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Home , Ablabesmyia, Ablabesmyia annulata, Ablabesmyia monilis, Anatopyniini, Boreoheptagyia, Chaetocladius

THE OF OTSEGO LAKE WITH KEYS TO THE IMMATURE STAGES OF THE SUBFAMILIES AND (DIPTERA)

Joseph P. Fagnani

Willard N. Harman

BIOLOGICAL FIELD STATION

COOPERSTOWN, NEW YORK

OCCASIONAL PAPER NO. 20 AUGUST, 1987

BIOLOGY DEPARTMENT

STATE UNIVERSITY COLLEGE AT ONEONTA THIS MANUSCRIPT IS NOT A FORMAL PUBLICATION. The information contained herein may not be cited or reproduced without permission of the author or the S.U.N.Y. Oneonta Biology Department ABSTRACT

The species of Chironomidae inhabiting Otsego Lake, New York, were studied from 1979 through 1982. This report presents the results of a variety of collecting methods used in a diversity of habitats over a considerable temporal period. The principle emphasis was on sound and rearing to associate immatures and adults. Over 4,000 individual rearings have provided the basis for description of general morphological stages that occur during the life cycles of these species.

Keys to the larvae and pupae of the 4 subfamilies and 10 tribes of

Chironomidae collected in Otsego Lake were compiled. Keys are also presented for the immature stages of 12 Tanypodinae and 2 Diamesinae species found in Otsego Lake. Labeled line drawings of the majority of structures and measurements used to identify the immature stages of most species of chironomids were adapted from the literature. Extensive photomicrographs are presented along with larval and pupal characteristics, taxonomic notes, synonymies, recent literature accounts and collection records for 17 species. These include: ­

Paratendipes albimanus (Meigen) (); Ortl10cladiinae ­

Psectrocladius () simulans (Johannsen) and I!ydrobaenus johannseni (Sublette); Diamesinae - ramosus Saether

(Protanypodini) and longimana Kieffer (); and

Tanypodinae - (Clinotanypus) pinguis (Loew)

(Coelotanypodini), Tanypu~ () punctipennis Meigen (),

Procladius (Psilotanypus) bellus (Loew); Var. 1 Roback,

i (Holotanypus) johnsoni Roback and Procladius (Holotanypus) denticulatus

Sublette (), and pilosella (Loew),

(Ablabesmyia) annulata (Say), Ablabesmyia (Ablabesmyia) basalis

(Walley), Ablabesmyia (Ablabesmyia) mallochi (Walley), Pentaneura

inconspicua (Malloch), (Conchapelopia) currani (Walley),

and (Thienemannimyia) norena Roback ().

Rearings provided as many as eight undescribed new associations of

immature stages with the respective adults (Ablabesmyia (Ablabesmyia)

basalis (Walley) (Tanypodinae: Pentaneurini)j (?)

scimitarus (Townes)j milleri (Townes), mirabile

(Townes) (Polypedilum) crassum Maschwitz, unguiculatus (Malloch) and protextus (Townes) (Chironominae:

Chironomini)j and ligulata ligulata Saether

(». Several unidentifiable and possibly undescribed new species were also associated.

One hundred twenty one species of Chironomidae in 4 subfamilies, 10

tribes and 51 genera were recorded from Otsego Lake. One hundred one of

the species reported were collected in one or more life stages in this study. Additional records were compiled from the literature. About

two-thirds (N=80) of the total species were in the Subfamily

Chironominae with about half (61) being in the Tribe Chironomini.

Orthocladiinae (22) and Tanypodinae (17) were much less diverse in

species, and only 2 species of Diamesinae are represented.

ii ADDENDUM

Since submission of this manuscript, two journal articles were published which change the names of two species presented here. The species identified as CryPtochironomus fulvus group (sensu Curry 1958, p. 435 and Darby 1962, p.'73) on page 13 was redescribed by Mason (1986, p. 404) as a new species and named Q. curryi Mason. Also, Ablabesmyia (b) basalis (Walley) reported on pages iii, vi, ix,. 12, 23, 28 and 60-63 was designated a synonym of !. (!.) monilis (Linnaeus) by

Roback (1985, p. 185). Mason, P. G. 1986. Four new species of the CryPtochironomus fulvus (Johannsen) species complex (Diptera. Chironomidae). Ent. scand. 161399-413. Roback, S. s. 1985. The immature chironomids of the eastern United States, VI. Pentaneurini- Ablabesmyia. Froc. Acad. Nat. Sci. Phila. 137(2)s153-212.

iii ACKNOWLEDGEMENTS

Our thanks are extended to Ron Embling and Charles

Winters of the SUNY College at Oneonta Graphic Arts

Department for information and assistance in photography and graphics and to June Lundon and staff at Milne Library for her help and cooperation in obtaining inter-library loans.

We wish to express special appreciation to Dr. Robert C.

MacWatters and Thomas Simmons for donation of material and

Dr. Selwyn S. Roback for kindly confirming most of the

Tanypodinae rearing identifications. We also thank Robert

Montione for loaning material for identification.

Acknowledgement is gratefully extended to Colleen Driscoll for her patience in typing the many versions of the manuscript and to Faye Hacker, Deborah Jicha and Victoria

Lamberton for additional typing.

iv TABLE OF CONTENTS

page l.~ ABSTRACT ••••.•••••.•••.••••.•••••..•••••••..•.••.•...••...• 0 •• 0 • • • ••

ACKNOWLEDGEMENTS. •••••.•••.••••••• ..••••.••••••••••..•.• . • . . . • • • • • • • i v

TABLE OF CONTENTS...... v

LIST OF TABLES •.••..•••• 0 ••••••••••••••••••••••••••••••••••••••••••• vii

LIST OF MAPS ••••.••••••••••••••••••••••••••••••.••••••• , • 0 •••••••••• vii

LIST OF PLATES AND FIGURES ••.•.••.•••••••••••••••• , .••... 0 •••••••••• viii

INTRODU CTION ••••.•.••••.••.• ..•••..•••••.••••••••••.•..•.•...•.. • • . • 1

MATERIALS AND HETHODS...... 1

Collection...... 1

Re a ring ., 0 •• III ••••••• " •••••• " • • • • • • • • • • • • • • • • • • • • • • • • 4

Slide preparation••••..••.••••••.•••••••••••••••••.••. 0...... 6

Illustrations...... 8

Material examined •••••••••••••• , •••.••••••••••••••••...•••••.••• 8

Structures and terminology •••• •..••••.••••••••••... 00 ••••• •••••• 9

Explanation of keys...... 10

RESU~1E OF SPECIES ••••••••.•••••..•••••.•••• o. ••••••••••••••••••••••• 11

KEY TO THE SUBFAMILIES OR TRIBES OF OTSEGO LAKE CHIRONONID LARVAE... 19

KEY TO THE TRIBES AND GENERA OF OTSEGO LAKE DIAMESINAE LARVAE...... 21

KEY TO THE TRIBES AND GENERA OR SUBGENERA AND SPECIES OF OTSEGO

LAKE TANYPODINAE LARVAE ••••••••••••••••••.••••••••... 0 •• • •• ••• • • 21

KEY TO THE SUBFAMILIES OR TRIBES OF OTSEGO LAKE CHIRONOHID PUPAE.... 24

KEY TO 1BE TRIBES AND GENERA OF OTSEGO LAKE DIAMESINAE PUPAE •••••••• 26

KEY TO THE TRIBES AND GENERA OR SUBGENERA AND SPECIES OF OTSEGO LAKE

TANYPODINAE PUPAE...... 26

NOTES ON SPECIES INCLUDED...... 29

Subfamily Chironominae (Tribe Chironomini) ••••••••••....•••.•..• 29

v TABLE OF CONTENTS (continued)

page NOTES ON SPECIES INCLUDED conintued

Paratendipes albimanus (Meigen) .....•...... 29

Subfamily Orthocladiinae...... 32

Psectrocladius (Psectrocladius) simulans (Johannsen) ...•... 32

Hydrobaenus johannseni (Sublette) •...... 34

Subfamily Diamesinae (Tribe Protanypodini) ••.•••...... ••••••• 37

Protanypus ramosus Saether ...... •.•...... •..... 37

Subfamily Diamesinae (Tribe Diamesini) .•...••.•...... 39

Potthastia longimana Kieffer ...... •..•...... •. 39

Subfamily Tanypodinae (Tribe Coelotanypodini) ...... •.•. 42

Clinotanypus (Clinotanypus) pinguis (Loew) ..•....•..••.•.•. 42

Subf amily Tanypodinae (Tri be Tanypodini)...... 44

Tanypus (Tanypus) punctipennis Meigen •...... •. 45

Subfamily Tanypodinae (Tribe Procladiini) ..•••••••...... •..•.. 47

Procladius (Psilotanypus) bellus (Loew); Var. 1 Roback ..... 47

P. (Holotanypus) johnsoni Roback ...... ••.••• 51

P. (!!.) denticulatus Sublet te...... 53

Subfamily Tanypodinae (Tribe Pentaneurini)...... 55

Labrundinia pilosella (Loew) ....•.•.....••.•..•..•...... •. 55

Ablabesmyia (Ablabesmyia) annulata (Say) ••...... 58

A. (~.) basalis (Walley) .....•..•.•.••.•...... •.•....•.• 60

A. (~.) mallochi (Walley)...... 63

Pentaneura inconspicua (Malloch) ....•..•.••...... •...•.••. 66

Conchapelopia (Conchapelopia) currani (Walley) .....•••••••• 67

Thienemannirnyia (Thienemannirnyia) norena Roback ...... ••• 69

LITERATURE CITED...... 71

vi LIST OF TABLES

page Table 1. A list of chironomids collected in Otsego Lake or

believed to be present in or about the lake...... 12

Table 2. Classification of Otsego Lake Chironomidae 18

LIST OF NAPS

page Map 1. Relevant landmarks and bathymetric map of Otsego Lake,

New york...... 2

LIST OF PLATES AND FIGURES

Plate Figure page 1. 1. Life cycle of a chironomid . 91

2. 2. Slide mountAd rearing showing arrangement

of par ts ...... •.....• ...... 93

3. 3-4. Larval head capsules and cephalic structures .... 95

4. 5-6. Larval head capsules and cephalic structures.... 97

5. 7-8. Larval head capsules and cephalic structures ..•. 99

6. 9-17. Generalized larval Tanypodinae head capsule

and cephalic structures..•...... •...• 101

7. 18-20. Generalized larval Chironominae morphology 103

8. 21-29. Generalized larval cephalic structures showing

measurements ...... ••...... •. 105

9. 30-34. Larval cephalic structures showing measurements. 107

10. 35-37. Larval cephalic premento-hypopharyngeal complex. 109

11. 38-40. Larval cephalic and anal segment morphology

showing measurements ...... •.. III

12. 41. Generalized pupal morphology...... •• 113

13. 42-45. Generalized pupal morphology and setation 115

vii LIST OF PLATES (continued)

Plates Figures page 14. 46-59. Thoracic horn morphology and measurements 117

15. 60-64. Paratendipes albimanus (Meigen) 119

16. 65-71. P. albimanus continued..••...... 121

17. 72-77 . P. a1bimanus continued 123

18. 78-85. P. a1bimanus continued...... •...... 125

19. 86-92. Psectrocladius (Psectrocladius) simu1ans

(Johannsen) 127

20. 93-100. P. simulans continued 129

21. 101-104. P. simu1ans continued 131

22. 105-112. Hydrobaenus johannseni (Sublette) 133

23. 113-119. H. johnannseni continued .•...... 135

24. 120-123. H. johnannseni continued 137

25. 124-130. Protanypus ramosus Saether 139

26. 131-136. P. ramosus continued 141

27. 137-143. P. ramosus continued 143

28. 144-151. P. ramosus continued 145

29. 152-159. P. ramosus continued 147

30. 160-166. Kieffer 149

31. 167-173.!. longimana continued 151

32. 174-179. Clinotanypus (C1inotanypus) pinguis (Loew) 153

33. 180-186. C. pinguis continued .....•.•...... •..... 155

34. 187-192. C. pinguis continued 157

35. 193-199. Tanypus (Tanypus) punctipennis Meigen...... •.... 159

36. 200-201.!. punctipennis continued•...... •...... 161

37. 202-205. Procladius (Psilotanypus) be1lus (Loew);

Var. 1 Roback ...... •...... 163

viii LIST OF PLATES (continued)

Plates Figures page 38. 206-212. P. bel1us continued ...... •...... 165

39. 213-215. P. bellus continued 167

40. 216-221. Procladius (Holotanypus) johnsoni Roback 169

41. 222-226 P. johnsoni continued...... •...... 171

42. 227-233. P. johnson! continued 173

43. 234-236. P. johnsoni continued 175

44. 237-244. P. (Ho1otanypus) denticulatus Sublette 177

45. 245-250. P. denticulatus continued 179

46. 251-258. P. denticulatus continued 181

47. 259-262. P. denticulatus continued 183

48. 263-270. P. denticulatus continued ...... •.... 185

49. 271-276. Labrundinia pilosella (Loew) 187

50. 277-284. L. pi10sella continued 189

51. 285-290. L. pilosella continued 191

52. 291-297. Ablabesmyia (Ablabesmyia) annulata (Say) 193

53. 298-300. A. annulata continued 195

54. 301-306. A. annulata continued ...... •. 197

55. 307-313. A. annulata continued 199

56. 314-321. Ablabesmyia (!.:.) basalis (Walley) 201

57. 322-328. A. basalis continued 203

58. 329-336. A. basalis continued ...... • 205

59. 337-339 A. basalis continued 207

60. 340-345. ~blabesmyia (~.) mallochi (Walley) 209

61. 346-354. A. mal10chi continued 211

62. 355-362. A. rnallochi continued ••...... •...... ••.. 213

63. 363-365. A. rna110chi continued 215 Ix LIST OF PLATES (continued)

Plates Figures page 64. 366-371. Pentaneura inconspicua (Malloch) 217

65. 372-379. P. inconspicua continued 219

66. 380-385. P. inconspicua continued 221

67. 386-390. Conchapelopia (Conchapelopia) curran! (Walley) .. 223

68. 391-397. C. curran! continued••...... 225

69. 398-405. C. currani continued 227

70. 406-407. C. currani continued 229

71. 408-414. Thienemannimyia (Thienemannimyia) norena

(Roback) 231

72. 415-421. T. norena continued 233

73. 422. T. norena continued 235

x INTRODUCTION

The species of Chironomidae inhabiting Otsego Lake, New York, were studied from 1979 through 1982. The principle emphasis of this work was on sound taxonomy and rearing to associate immature and adult stages.

It is hoped that the keys and information presented will be of general value to the beginning student as well as to the experienced chironomid taxonomist.

The field work was conducted from the State University of New York

College at Oneonta Biological Field Station. The Biological Field

Station is located on the west shore of Otsego Lake near the Village of

Cooperstown at Rat Cove. Most of the field work was concentrated in this area (Map 1).

Otsego Lake lies in Otsego County in New York State (42 0 40'N-700 00'W) in the glacially overdeepened headwaters of the Susquehanna River.

Harman and Sohacki (1976) and Harman et al. (1980) recently summarized the limnology and other characteristics of Otsego Lake. It is a finger lake with a maximum depth of 166 ft (50.5 m) and possessing characteristics typical of glaciated lakes (Map 1). It is dimictic, well-buffered and moderately productive, with diverse sediments and abundant chironomid microhabitats.

HATERIALS AND METHODS

CoHee ting

Field sampling usually occurred during June through September, from

1979 through 1982. Sampling was also conducted in April and May of 1981

1 2

CRIPPLE CREEK

OTSEGO LAKE (GLIMMERGLASS) OTSEGO COUNTY. NrW' "tOllllt

~ ....-.­~TOD... 'TATE UflrvUUUrv OF NEW YOII .. COU£Q~ AT OHtOIlfTA IUOLOGM::Al nu.D IYATKMlf COOI"EUTOWN. III Y.

THREEMILE POINT

LEATHERSTOCKING CREEK

BROOKWOOD POINT ~

BLACKBIRD BAY Map 1. Relevant landmarks and bathymetric map of Otsego Lake, New York. (Source: W.N. HarmRn, Department WILLOW BROOK of Biology, SUNY College at SUSQUEHANNA RIVER Oneonta, Oneonta, NY) 3

and 1982 and extended through October in 1981. In April of 1982,

sampling began within a week after ice out.

There are no reliable methods of sampling chironomid populations

that are not exceedingly time consuming due to the generally small size

and frequent great abundance of the organisms and large quantities of

sediment often required for adequate samples. Standard size (9 in x 9

in) Ekman grabs supplied the bulk of immature material in this study.

Live larvae and pupae were also obtained for rearing from dip net

samples, multiplate samplers, skims of the waters' surface with a fine mesh sieve or net and hand picking from rocks, wood and other debris and vegetation along the shoreline. Although Ekman grabs were taken in many

localities throughout the lake, most were collected in open water

sediments and were concentrated in the Rat Cove area, opposing shore and

intervening profundal zone (Map 1). The latter sampling methods were confined to this area of the lake.

Sample processing methods depended on the amount of sediment collected. Usually, grabs and other sediment samples were gf'ntly sieved

in a manner which also allowed collection of the smallest species. The

technique is a gentle swirling action that encourilges entrapment of

specimens in the meniscus at the air-water interface in the sieve. The

specimens were generally transferred with a probe or dropper to a white enamel pan or plastic tub with lake water that was kept in the shade until individual rearings were established. Any remaining specimens were preserved in 70% ethanol and retained for comparative purposes as whole larvae. Relatively clean samples, such as rocks, twigs, some

vegetation and multiplate samples, etc., were placed in a pan or tub 4

with lake water and rearings established with hand pick~d and swimming specimens found. Bulky but clean samples, such as vegetation, were often held in a pail or large tub of lake water with many swimming larvae and pupae easily siphoned off with a dropper or pipette. Small portions were then generally removed and inspected in a white enamel pan or plastic tub. Samples with mixed materials were treated accordingly.

Adult chironomids were collected as additional taxonomic specimens.

They were sampled by sweep net on and about the lake in many localities and by sweep net and aspirator on and near the Field Station facilities and preserved in 70% ethanol.

Rearing

Chironomid are holometabolous with four stages in the life cycle: egg, , pupa and adult (Fig. 1). The purpose of rearing is to obtain a direct association of immature stages with the respective adults. Although the immature stages of many species are described, rearing is still the favored method for obtaining the most accurate species level determinations. One reason for this is because the adult stage in most groups is much better known taxonomically, particularly the males. Also, in addition to the lack of species keys to larvae and pupae, the range of character variation in the immature stages is not known for most species and a large minority of life histories are not yet known, so that rearing larvae to adults is often still necessary to confirm species identity.

The method of rearing chironomids in this study followed Roback

(1976b). Individual larvae and pupae taken from samples with droppers or probes were placed in separate 6 dram vials about 1/4 full with 5

distilled water which were then capped with a foam plug. Six dram vials were selected because they were large enough containers for the largest species encountered and also fit conveniently into laboratory test tube racks. In the field, racks of rearing vials were generally kept in a cooler, with or without ice, depending on the season. In the laboratory, the rearings were kept in a cool (15-200 C) place whenever possible and examined daily for deaths, pupati0ns and adult emergences.

The majority of dead larvae were examined and discarded if successful rearings were obtained. Dead larvae retained, larvae which pupated, and those that subsequently emerged were preserved with 70% ethanol in the respective rearing vials. Emerged adults were allowed to fully harden and were then immobilized with a squirt of 70% ethanol and the vials then gently filled and capped.

Some multispecies or mass rearings were also undertaken and examined daily. Many pupae and fourth instar larvae, most noticeable as pre pupae were subsequently removed and individually reared. Pre pupae are fully developed larvae preparing to pupate with pupal and developing adult structures discernible. The larval thorax becomes noticeably swollen and the large, developing adult eyes appear as two dark spots in or just posterior to the larval head.

Roughly, about 4,000 individual larval rearings and several hundred pupal rearings were established. Approximately 10 percent (N=400) resulted in adult emergences witll an additional 7.5 percent (300) terminating after pupation. Slide preparation

When possible, some associations were retained in

Specimens should generally be examined under a dissecting scope for important characteristics prior to slide mounting. All individual associations that were slide mounted, were mounted on the same microscope slide, usually with thinness 1 circular coverslips 12 millimeters in diameter (Fig. 2). These rearing, preservation and slide mounting procedures greatly reduced the chance of a mis-association occurring.

Numerous slide making techniques have been described for chironomids. Beck (1968 and 1976) provided a thorough basic introduction to larval mounting, positioning and identification procedures. Hahn et al. (1977) also discussed basic equipment and procedures, while Russell and Soponis (1981) and Beckett and Lewis

(1982) discussed rapid and inexpensive larval processing methods.

Coffman and Ferrington (1984) briefly discussed preparation of larva,

pupa and adult specimens and \.Jirth (1961), Schlee (1966), Wirth and

Marston (1968) and Hansen and Cook (1976) described adult slide making

techniques.

There are obvious advantages and disadvantages that occur in

selecting a particular slide making procedure or technique. In this

study, specimens were mounted directly from alcohol or water into

polyvinyl lactophenol oriented as shown in Figure 2 and covered with a coverslip. Anterior and proximal aspects of specimens and structures 7

are usually mounted towards the observer because of the inverted image of the compound microscope. "Larvae and exuviae are mounted ventral side up with careful cleaning and positioning of the head, body segments and posterior structures. Slight manipulation of and gentle pressure on the coverslip, while viewed under a dissecting microscope, is usually required to properly orient the greatest number of taxonomic characteristics. Pupal exuviae are mounted do~sal side up after careful cleaning. Pupae are dissected with the abdomen usually mounted dorsal side up and the head and thorax mounted as in adults. Often, it is best to peel the pupal exuviae off the thorax and to gently extricate the adult abdomen from the exuviae if possible before mounting them. Adults are also dissected prior to mounting with the respective parts variously arranged as in Figure 2. Generally, male abdomens are mounted dorsal side up, female abdomens ventral side up and in both sexes, the thoraces are mounted laterally and heads dorsally.

Polyvinyl lactophenol is a clearing, semi-permanent medium. Though soluble in alcohol or water, alcohol retards clearing somewhat so larger, thick bodied specimens were usually rehydrated in distilled water prior to mounting. Experience showed that some of the largest and darkest specimens, particularly whole larvae and pupae and adult abdomens and thoraces required further clearing. After rehydration, these specimens were placed in a warmed 5 to 10% potassium hydroxide solution for several minutes to digest tissues and reduce coloration, soaked in distilled water for several minutes and then mounted.

Finished slides were placed in a light box for specimen clearing and media hardening. Gentle warming from an incandescent bulb in a small 8

slide cabinet enhanced clearing and increased the speed of hardening.

Slides were checked periodically for media shrinkage and filled when necessary. Clearing took from several hours to about 2 days depending on specimen thickness and tenacity. Hardening took from several days to about 2 weeks depending on mount thickness. When hardened, each coverslip was ringed with nail polish to retard further media shrinkage.

Illustrations

Line drawings used in this study were obtained from cited sources, usually with little modification. Photomicrographs were taken with ASA

100, Plus-X black and white print film in a Kodak Colorsnap 35 manual camera, mounted on a Bausch and Lomb 2X Dynazoom research compound microscope with apochromatic objectives. Standard photography and dark room procedures were used in shooting, developing and enlarging.

Magnification of each photomicrograph was determined as the product of objective magnification, camera focal length magnification (2.5X) provided by the manufacturer, zoom magnification if used and print enlargement magnification. The highest possible objective magnification was used to preserve image quality and resolution of structures photographed. All prints were negative enlargements of 2X or 3X accomplished by enlargement of a negative of a stage micrometer onto a millimeter rule.

Material examined

Specimens collected in this study provided the bulk of the material examined. However, other loaned and donated chironomids collected from

Otsego Lake were also examined.

About 3,000 presorted adult chironomids collected by Simmons (1983) from several locations on and about Otsego Lake in July and August of 9

1981 were identified and supplemented collection records (Tables 1

and 2).

Some larvae collected by Mr. Robert Montione in Goodyear Swamp at

the north end of Otsego Lake in 1983 were identified and also

supplemented collection records (Tables 1 and 2). In addition, some

immature specimens were obtained from the stomachs of profundal feeding

whitefish (Coregonu~ clupeaformis) collected by Dr. Robert W. MacWaters.

Food items were examined from nearly 100 stomachs obtained from bottom

gill net collections set in 20 to 30 meters of water off of Kingfisher

Tower and Peggs Point in September-October 1980 and May-June of 1981

(Map 1).

Structures and terminology

Anatomical and morphological terminology in Chironomidae has been much confused in the past. In this work, the terminology of Saether

(1971, 1976, 1977a, 1977b and 1980) was used where possible. Roback

(1976b, 1977, 1980 and 1981), Wilson and McGill (1982), Oliver and

Roussel (1983), Wiederholm (ed.) (1983) and Coffman and Ferrington

(1984) were additional sources for terminology used.

A large majority of the structures and measurements used to identify

the larvae and pupae of most species of chironomids are labeled in

Figures 3-59. As used here, lateral setae found on the abdominal

segments of many pupae (Fig. 41) refers to the large, generally

lamelliform or filamentous lateral setae also called macrosetae and swim

setae, and not to laterally placed hair setae or non-lame11iform or

filamentous setae. Counts given of paired structures such as lateral

setae are of only one side or one of the pair. Abbreviations used 10

for life stage and sex were larva (L), pupa (P), prepupa (PPL), male (M) and female (F). The sex was noted and an index number assigned to each specimen included in this work. This was done so that each photomicrograph refers to a specific specimen of a species of known sex. In the family Chironomidae, male individuals are generally smaller and more slender than the female of a species in all life stages.

Though sexually dimorphic in size, larvae could not be reliably determined as to sex. Pupal sex, however, is easily discernible. The antennal and genetal sheaths in male pupae are much enlarged compared with females to accommodate the adult structural differences. Rough designations of a species body size were given as small, medium and large. Generally, small species may range up to 4 or 6 mm in the immature stages, medium species to 10 or 12 mm, large species over 12 mm and very large over 18 mm.

Explanation of keys

The keys presented are traditional and were prepared for identifying whole larvae and cast skins or exuviae of larvae in the fourth (and usually third) instar and for pupae and pupal exuviae. The characters used are visible on slide mounted material of larvae mounted ventral side up and pupae mounted dorsal side up. Many characters are also visible on unmounted material.

The keys were intended for ease of identification and not to show taxonomic or phylogenetic relationships. Most characters were compiled mainly from the keys of Roback (1957, 1976b, 1977, 1980 and 1981), Beck

(1968 and 1976), Oliver et al. (1978), Wilson and McGill (1982), Oliver and Roussel (1983), Wiederholm (ed.) (1983) and Coffman and Ferrington

(1984). 11

Most specimens will key well using only the primary and secondary characters given in the keys, although additional characters are provided if they are missing or obscured on a specimen. It is stressed that only a small number of species are keyed in this work. Therefore, care must be taken in general use of the species keys.

RESUME OF SPECIES

One hundred twenty one species of chironornidae in 4 subfamilies, 10 tribes and 51 genera were recorded from Otsego Lake (Tables 1 and 2).

One hundred one of the species reported were collected in one or more life stages in this study. Additional records were compiled from the literature. About two-thirds (N=80) of the total species were in the

Subfamily Chironominae with about half (61) being in the Tribe

Chironomini. Orthocladiinae (22) and Tanypodinae (17) were much less diverse in species, and only 2 species of Diamesinae are represented. 12

Table 1. A list of chironomids collected in Otsego Lake or believed to be present in or about the lake. [An asterisk (*) denotes species collected only 8S adults on or about the lake in the present study. A plus (+) denotes species whose immature stages were collected in the lake with most being reared. Adult collection records reported by other nuthors are noted as l=Roback (1971), 2=Townes (1945), and 3=Nevin (1936).] 4,5

Subfamily Tanypodinae

Tribe Coelotanypodini + 1. 1Clinotany~ (~.) pinguis (Loew) 1861:308.

Tribe Pentaneurini + 2. 1Ab1abesmyia (~.) annulata (Say) 1823:15.

3. +~. (A_.J' basalis ( Walley ) 1925 : 273 .

4. • ~. (~.) ianta (Roback) 1959:131.

5 • +~. (~.) mallo~hi (Walley) 1925:273.

6. • ~. (Kare!ia) i.!linoe!!~is (Malloch) 1915:376.

7. 1~. q~.) peleensis (Walley) 1926:64.

8. +1Conchapelopia (~.) curran! (Walley) 1925:276.

9. 1!;;. (IIelopeloQi.!!) Qjlicauda1! (Walley) 1925 :277. + 10. Labrundinia QilQ~ell~ (Loew) 1866:5. + 11. Pentaneura incogspicul! (Mall oeb) 1915: 371. + 12. Thienemannimvia (T.) nor~na (Roback) 1957:38.

Tribe Procladiini + 13. Procladius (Psilotanypus) bellus (Loew) 1866:4; var. 1 Roback 1980:31.

14. +r. () Ro1otany~~ denticulatus Sublette 19 6 4:12 4 .

15. +r. (~.) iQhnso~i. Roback 1980:43.

Tribe Tanypodini + 16. Tan~ (T.) pun£1ip~gni~ Meigen 1818:61.

======~=====~======~======~======11

Table 1, continued.

Subfamily Diamesinae

Tribe Diamesini + 17. Pott!t!!tl.!!! longimana Kieffer 1922:362.

Tribe Protanypodini + 18. Protanypus ram~~us Saether 1975:368.

Subfamily Chironominae

Tribe Chironomini

19. 2Chironomus (Chaetolabis) atroviridis (Townes) 1945:114.

20. .2~. () anthfacinu~ Zetterstedt 1860:6499.

21. +3~. (~.) decQrus Johannsen 1905:239.

22. +2~. (~.) plumoslls (Linnaeus) 1758:587.

23. .~. (~.) ~ari~~ Meigen 1804:13.

24. 2~. (~.) tenta!!~ Fabricius 1805 :38.

25. +~. (~.) sp. 3.

26. +~. (~.) sp. 5. + 27. ~ladopelm~ amach!!~I~ (Townes) 1945:168. + 28 . ~. collator (Townes) 1945:169.

29. • ~. edwards.! (Kruseman) 1933:194.

30. +~. sp. 4. + 31. CrYQ1Qchironomus dig.!latus (Malloch) 1915:483. + 32. ~. f!!lvlls gpo (sensu Curry 1958, p. 435 and Darby 1962, p. 73).

33. +~. (?). ~cimitar!!~ ()Townes 1945:98. + 34. Cryptotendip~ £8su8rius (Townes) 1945:162. + 35. ~. emorsus (Townes) 1945:161.

======~======~=== 14

Table I, continued.

Subfamily Chironominae, continued

Tribe Chironomini, continued

36. 'Dierotendi~~ !Qbiger (Kieffer) 1921:71.

37. +~. mi!~El (Townes) 1945:110. + 38. 1~. modestus (Say) 1823:13.

39. 'Q. nervosus (Staeger) 1839:567. + 40. brunneipenni! (Johannsen) 1905:205.

41. 'g. !Qrs.!!.lis Oleigen) 1818:25.

42. '~. Q!!ganus (Meigen) 1838:7.

43. +' niK!ican~ (Johannsen) 1905:219. + 44. 19. subtenQen~ (Townes) 1945:65.

45. 'Glyptotendip~ (Q.) sen!!i! (Johannsen) 1937b:37.

46. 'Q. (Phytotendi~) !Qbiferus (Say) 1823:12.

47. 'Q. (~.) Q.!!.ri~ (Edwards) 1929:392. + 48. cur.!l!!!J!!~l!at!! (Malloch) 1915: 474.

49. .2Lauterborniella !&rayloiQ~ (Kieffer) 1911:51.

50. .~. varipennis (Coquillett) 1902:94. + 51. caQucus Townes 1945:24. + 52. 'M. pedellus var. pedellus (De Geer) 1776:378. + 53. Nilothauma mir!!bil~ (Townes) 1945:35. + 54. abort hus (Malloeh) 1915: 465.

55. '~. Qot!J!!Qgeti (Townes) 1945:159.

56. +~. tenui~da~us (Malloch) 1915:475.

57. · ~~dine (Townes) 1945:149.

58. +'Paratendip~! albiJ!!anus (Meigen) 1818:40.

======~=~======15

Table I, continued.

------.- .. - ._------­ Subfamily Chironominae, continued

Tribe Chironomini, continued + 59. () coracina (Zetterstedt) 1850:3508. + 60. zr.. (!:..) flavipes O!eigen) 1818:50. + 61. E. (!:..) obediens (Johsnnsep) 15)05:286.

62. ZE. (E.) punctipes (Wiedemann) 1817:65.

63. *ZFolypedilum (Pentapedilum) tritum (Walker) 1856:342. + 64. f. (Polypedilum) bergi ~'!aschwitz 1976:148.

65. 3E. (E.) convictum (~laU.er) 1856:161. + 66. r.. (E.) crassum Maschwitz 1976:99.

67. Zf. (f.) illinoense (Malloch) 1915:471.

68. Zf. (!:..) nubeculosum (Meigen) 1804:18.

69. ~E. (Tripodura) digitifer Townes 1945:45. + 70. 3£. halterale «('0(l1d111'11) 1~Olh:17. + 71. ,E. (T.) scalaenul11 (SchranU lR03:73. + 72. z£.. (!.) simulans Townes 1945:43.

73. t2Stenochironomus hilaris (Walker) 1848:17.

74. Z~. taeniapennis (Coquillett) 1901a:607. + 75. 2Stictochironomus devinctus (Say) 1829:150. + 76. ~. unguiculatus (Malloch) 1934:16. + 77. zTribelos iucundus var. dimorphus (Malloch) 1915:464. + 78. 21. protextus (Townes) 1945:69.

79. 2Xenochironomus xenolabis (Kieffer) in Tbienernann 1916:526.

Tribe Pseudochironol11ini + 80. banksi Townes 1945:17.

:: :: :. :- ; :- :-:.- ::: ::. -==::- =- ::". :.; =- =-::: =-=:::: =- ~ ~:-=- =-::- =- =":: =::: -:- =::- =~:: ;: ~ =- = =-- =- =-:. -=:: =: == =-:- =- ::- =- :: -:: =- -:: ': :- ::: : :- ::: :: :: =: == ==-.:::::::= = 16

Table 1, continued.

Subfamily Chironominae, continued

Tribe , continued

81. :I~. fulviventris (Johannsen) 1905:229.

82. :I~. netta Townes 1945:19.

Tribe Tanytarsini + , 83. C1~dotanyt!!.rsus sp. l.

84. +~. sp. 2.

85. +Mic ropsec t ra sp. 1.

86. .~. sp. 2. + 87. Par!!1anytarsus sp. l. + 88. ~. sp. 2.

89. ·Stempell ina sp . 1.

90. .~. sp. 2. + 91. sp. l.

92. .-.I. sp. 2 .

93. .!. sp. 3.

94. +!. sp. 4. + 95. !. sp. 5.

96. +1:. sp. 6.

97. +1:. sp. 7.

Subfamily Orthocladiinae

98. + sp. 1. + . 99. taris Roback 1957:61. + 100. (~.) sp. 1. + 101. ~. (Isocladius) myriophyll i 01 her 1984: 1287. ======17

Table 1, continued.

Subfamily Orthocladiiuae, continued + 102. ~. \1.) flnlpes Johannsen 1942:73.

103. +~. (!.) sylvestris (Fabricius) 1794:252. + 104. Heterotris~ocladins (1) hirtapex Saether 1975:39. + 105. Hydrobaenn~ j ohannseni (Subl cUe) 1967: 504. + 106. Orthocl&dius (g.) annectens Saether 1969:61.

107. + Q. (.g,) obumbratoa Johannsen 1905:281. + 108. Parakieffexiel1a sp. 1. + 109. f. sp. 2. + , 110. Parametrlocnemus lundQ~~!i (Johannsen) 1905:302. + 111. Psectrocladius

114. + 115. Orthocladiinae sp. 1.

116. *Orthocladiinac spp.

4Near the completion of this study, Mr. Robert Montione collected larvae at Goodyear Swamp in Otsego Lake of the following 5 species that were identified by the writer and are supplemental to this list: ~atarsia (sp. A of Roback 1978, p. 191) (Tanypodinae: Pentaneurini); sp. 1 (Chironominae: Tanytarsini); Parachaetocladius sp. 1, Paralimnophye~ sp. 1 and sp. 1 (Orthocladiinae).

SRearings provided as many as eight undescribed new associations of immature stages with the respective adults (Ablabesmyia (Ablabesmyia) basalis (Walley) (Tanypodinae: Pentaneurini); Cryptochironomus (7) scimitarus (Townes); Dicrotendipes miller! (Townes), Nilothauma mirabile (Townes) Polypedilum (Polypedilum) crassum Maschwitz, Stictochironomu8 unguiculatus (Malloch) and Tribelos protextus (Townes) (Chironorninae: Chironomini); and Zalutschia ligulata ligulata Saether (Orthocladiinae». Several unidentifIable and possibly undescribed new species were also associated. 18

Table 2. Classification of Otsego Lake Chironomidae.

+ Number of species

Subfamily Tribe· Found in this Study Total

Chironominae 62 80

Chironomini 45 61 Tanytarsini 16 16 Psendochironomini 1 3

Orthocladiinae 22 22

Tanypodinae 15 17

Pentaneurini 10 12 Proc1adiini 3 3 Coelotanypodini 1 1 Tanypodini 1 1

Diamesinae 2 2

Diamesini 1 1 Protanypodini 1 1

4 Subfamil ies 10 Tribe s 101 121

---

*Triba 1 status not universally agreed upon in subfamily Orthocladiinae. + Compiled from Table 1. 19

KEY TO THE SUBFAHILIES OR TRIBES Of OTSECn LA.~ CHIRONOHID LARVAE

1. Head with retractile, 4-segmented antennae (Figs. 3, 4, 9, 10, 21, 174, 222 and 271): premento-hypopharyn~e~lcomplex with a 4­ to 8-toothed ventral, median ligula and lateral, toothed parnligulae (Figs. 3, 4, 9, 10, 13, 250, 302 and 3~6): mentum weakly developed (Fig. 16) or with dorsomental teeth arranged in conspicuous plates (Figs. 3, 15. 194 and ~47) or in longitudinal rows (Fig. 177), SubfamHy Tanypodinae .. (p. 21)

Antennae not retractile and with 4 to 8 segments (Figs. ~-8, 18, 23-25, 60, 90, 106. 138 and 162; premento-hypcpharyngeal complex lacking toothed ligul a and paraligulae (Figs. 35-37): mentum usually a well develoned ventral toothed plate with usually more than 5 to numerous apical teeth (Figs. 5-8, 18, 31, 38, 67, 68, 95, 110 and 112)...... 2

2. Antennal segment 3 with areas of reduced sclerotization giving the appearance of annulations (Figs. 25 and 162); premento­ hvpopharyngeal complex '.d til pell developen median ligula and lateral para-ligulae resembling 3 seti'll hrushes (Fig. 36). Subfamily Diam€sinae (in part: Tribe Diamesini) ....(p. 21)

Antennal segment 3 never annulated (Figs. 23, 24, 71, 90, 113 and 138); premento-hYf'c'pharyngeal complex ,,!ith Bertles or modified setae (Figs. 35 and 37), rarelY with long setae or at most wi th 11 appendage d,~vel(\ped as a s iogle brush...... 3

3. Occipital margin of head carsule with a distinct posteriorly directed ventra-lateral projection on each side (Figs. 8, 124 and 129); labrum with an overlapDing 1'0'1,07 of serrate lamellae (Figs. 8, 132 and 136); tl appencLOige of premerto-hypopharyngeal complex strongly developed and deeplv divided into lamellae (Figs. 37, 133 and 135); head capsule with numerous closelY spaced long setae (Figs. 8 and 125). Subfamily Diamesinae

(in part: Tribe Protanypodini). ... < ••••••• (p. 21)

Occipital margin of head capsule without or with rl sTOJall posterior­ ly directed ventre-lateral projection on each side (Figs. 5-7, 18, 65, 86 and 105); labrum 'vlthout a Tffiv of lamellae or at most~ with a pair of lahral lamellae; M appendage of pre­ mento-hypopharyngesl complex usuallv weaklv developed (Fig. 35); head capsule rarely '\·rlth numerous setae...... 4

4. Mentum with ventral part expanded laterally to form distinct and radially striated ventromental plates (striations absent or greatly reduced in ) which never have setae or beard heneath (Figs-:-S;-6> 18, 60, 62 and 67); eye spots usually divided and lying in a vertical plane or with dorsal eye spot lying anterior to ventral eye spot (Figs. 5 and 6); antennae 5- or 6-segmented. Subfamily Chironominae..... 5 20

Hentum with ventromental plates present or absent: if present and distinct, then never striated and with or 'vithout setae or beard beneath (Figs. 7, 31, 93, 95, 108, 110 and 112); eye spots usually single, or if divided, then with dorsal eye spot lying posterior to ventral eye spot; antennae 4- to 8­ segmented. SubfamilY Orthocladiinae* (Psectro­ cladius (R..) !'limu}ans, p. 32 and l!L9robae!.1U~ j ohannseni, p. 3L~)

5. Antennae 5-segmented dud usuCllly long, arising from a tubercle (pedestal) at least as long as wide (Fig. 6); first antennal segment generally curved and with an antennal seta present (Fig. 24); Lauterborn's organs generally large and conspicuous or occurring at the apex of elongated pecioles (pedicels) (Figs. 6 and 24); ventromental plates usualJy strap-like and with median apices separated by less than the width of the median mental tooth (Fig. 6) or fan-shaped and more widely separated; abdominal segments usually with plumose setae...... • . • . . . Tribe Tanytarsini

Antennae usually shorter, 5- or 6-segmented and with antennal tubercle (pedestal) shorter than wide or absent (Figs. 5, 60 and 61); first antennal segment generally straight and lackin~ an antennal seta (Figs. 5 and 66); Lauterhorn's organs generally smaller and not conspicuous and never occurring at the apex of elongated petioles (pedicels) (Figs. 5 and 71); ventromental plates usually fan-shaped and "(dth median apices separated by more than the width of the median or median pair of mental teeth (Figs. 5, 18, 60, 62 and 67) or less often more narrowly separated or plates may be strap-like and contiguous to very narrowly separated; abdominal segments usually wi th only simple setae. . . , ...... 6

6. Pos terior parapod Hi th two rOFS of claY-Is arranged in a horse-shoe shape; ventrornental plates strap-like and with median 8pices corotiguous or very narrowly separated ...... Tribe Fseudochironomini; Genus Pseudochironomus

Posterior parapod with claws irregularly arranged (Figs. 63 and 64); ventromental plates with median apices u~~ally separated by more than the width of the median or median pair of mental teeth (Figs. 67 and 68) or if narrowlY separated or contigu­ ous, then plates not strap-like. Tribe Chironomini ...... • ...... • . . . .. (Paratendipes albimanus, p. 29)

*Tribal status not universally agreed upon. 21

KEY TO THE TRIBES AND GENERA OF OTSEGO LJ\KE Dlf\MESINAF. LARV/IF.

1. Occipital margin of head cRpsule with a distinct rORt~rior]v directed ventro-'ateral projection on each side (Figs. R, 124 and 129); head capsule with numerous closely spaced long setae (Figs. 8 and 25); labrum with an overlapping row of apically serrate lamellae (Figs. 8, 132 and 136); H appendage of pre­ mento-hypopharvngeal complex strongly developed and deeplv divided into lamellae (Figs. 37, 133 and 135): antennal segment 3 not annulate (Figs. 137-139); ;Y1andibL~ \·iith <1 long apical tooth and 5 inner teeth (Figs. 131, 137 and 139); mentum with­ out distinct teeth on middle two-thirds (Figs. 130, lLfO, 141 and 143), Tribe Protanypodini; Genus Protanypus ...... •(~. rClmosus, p. 37)

Occipital margin of head capsule without a posteriorly directed ventra-lateral projection on each side (Fig. 160); head capsule without numerous long setae; labrum without an overlapping row of serrate 1a'l1e11ae, but ~vith typical labral setae (Fig. 164); premento-hypopharyngeal complex with well developed median ligula and lateral paraligulae resembling 3 setal brushes (Fig. 36); antennal segment 3 \,;rith areas of reduced sc1erotiz­ ation giving the appearance of annulations (Figs. 161 and 162) (Tribe Diamesini); mandible with a lon~, attenuate apical tooth a large, hook-shaped median process and lacking seta interna (Fig. 166); mentum undulate and entirely without teeth (Fig. 164 and Oliver and Roussel 1983, Figure 432). r.enus Potthastia•••...•... (in part: longlmana group, p. 39)

KEY TO THE TRIBES t~uJ GENERA OR SUnGENEP~ N~D SPECIES OF OTSEGO LAKE T~~PODINAE LARVAE

1. Abdominal segments with a lateral hail- fringe (Figs. 187 and 188); mentum with dorsomental teeth fused iota distinct plates (Figs. 3, 15, 194, 203, 220, 247 and 250) or separatE' and loosely arranged in longitudinal rows (Fig. 177); head ratio (length/width) less than 1.5 , ...... • 2

Abdominal segments without a '.,-el1 defined lateral hair frint;e, although numerous scattered hairs may be present; dorsomental teeth absent (Fig. 16); head ratio (length/width 1.5 or

greater...... 0 •, • • • • • • • • • • • 5

2. Mentum with dorsomental teeth separate and loosely arranged in longitudinal rO\olS (Fig. 177); conical papilla present bee-ween bases of procerci (Oliver and Roussel 1983, Fig. 20) (Tribe Coelotanypodini); ligula usually with six teeth and with a 22

concave apical marBin (Fig. I7fi); mandible stronRly curved and with a large, pointed basal tooth (Figs. 175 and 177). Genus Clinotanypus...... (~. pinguis, p. 47. )

Mentum with dorsomental teeth fused into distinct plates (Figs. 3, 15, 194, 203, 220, 247 and 250); conical papilla not present between bases of procerci; ligula usually with 5 teeth and mandible variable but not as above. . • . • . . . . 3

3. Mentum with dorsomental teeth fused into distinct plates which are joined medially (Fig. 194); mandible with a short apical tooth, minute basal and accessory teeth and with basal two-thirds expanded (Fig. 195); ligula with 5 pale teeth and usually with a convex apical margin (Fig. 198); M appendage without a pseudoradula. Tribe Tanypodini; Genus Tanypus ...... • ...... (:£.. punc tipennis, p. 45)

Mentum with dorsomental teeth fused into distinct plates which are not joined medially (Figs. 3, 15, 202, 203, 205, 216, 220, 222, 223, 246, 247 and 249-251); mandible with apical tooth much longer, with a relatively large and bluntly rounded basal tooth and not expanded in basal two-thirds, but more or less smoothly curved from base to apex (Figs. 204, 205, 217, 218 and 248); ligula with 5 black teeth and with a concave apical margin (Figs. 202, 203, 205, 216, 220, 222, 223, 246, 247 and 249­ 251); M appendage with p pseudoradula (Fig. 15). Tribe Procladiini; Genus Procladius. . . • ...... 4

4. Paraligula with apical tooth short, generally at most 2 times as long as accessory teeth (Fig. 203); pecten hypopharvngis with less than 10 short teeth; posterior parapod with one or more small, strongly hooked clm'is with an expanded base (Figs. 106 and 207). some of which may have 3 to 4 large teeth on inner margin; smaller species. Subgenus Psilotanypus ...... (Frocladius (£..~.) bellus), p. 117)

Paraligula with apical tooth long, generally at least 3 times as long as accessory teeth (Figs. 223, 247 and 250); pecten hypo­ pharyngis with more than 10 teeth (generally around 14 teeth, most of which are of normal length and larger than in the sub­ genus Psilotanypus); posterior parapod with all claws forming a flat curve, slender and simr]e (Figs. 226, 243 and 244) (some claws may have very fine serrations on inner margin); larger species. Subgenus Holotanypus...... (Procladius (B..) johnsoni, P. 51 and Procladius (.!!..:) denticulatus, p. 53 )

5. Maxillary palp with basal segment subdivided into 2 to 5 segments (Figs. 296, 299, 322 and 346). Genus Ablabes~ 6

Maxillary palp with basal segment consisting of a single segment (Figs. 11, 366, 367, 387, 389 and 408) .•..•...... 8 23

6. Ligula with 5 black teeth and tdth median teeth paler, even in height and rounded distally (Figs. 300-303); maxillary palp with basal segment subdivided into 2 segments subequf11 in length (Figs. 296 and 299); posterior parapod lacking dark claws and wi th s[113.1l c] CIt-'S covering approximately the dis tal half (Fig. 294). Ablabesmyia (~.') .?nnula~.§:. (p. 58 )

Ligula with 5 black pointed teeth and with median tooth shorter than laterals (Figs. 317, 325, 326,aod 340-343); maxillary paIr with basal segment subdivided into 3 to 5 segments (Figs. 316, 322, 323, 343, 346, 348, 349 and 351); posterior pararod with 1 to 3 dark claws and a SpRrse number of sma]l subapical claws (Figs. 327,328,345 and 352)...... , . , ... 7

7. Maxillary palp with basal segment subdivided into 3 segments (Figs. 316, 322 and 323); the proximal segment is verv short and often difficult to see (Fig. 319) and the distal segment is slightlY longer and more slender than the middle segment. Ablahesmyia (~.) basalis .... , .....•.•...... (p. 60)

Maxillary palp with basal segment subdivided into 5 segments (Figs. 343, 346, 348, 349 and 351); the proximal 4 segments are sub­ equal in length and much shorter than the distal fifth segment. Ablabesmyia (~.) mallochi...... (p. 61)

8. Ligula with middle tooth longer than lateral teeth (Fig. 273); antennal segment 2 darker than basal segment (Figs. 271 and 275); posterior parapod with one bifid claw with outer tooth shorter and weaker than inner tooth (Figs. 280 and 283) and with basal seta modified as a compound spur (Figs. 278, 280 and 281), Genus Labrundinia. .. , .•.. (1. pilosell~. p. 55)

Ligula with middle tooth subequal to lateral teeth (Figs. 367 and 368) or shorter than lateral teeth (Figs. 396, 397, 411 and 413); antennal segment 2 pale in color and nct darker than basal segment (Figs. 367 and 387); posterior parapod with all claws simple (Figs. 369, 395 and 412); posterior parapod with basal seta smooth and unmodified (Figs, 369, 395 and 409) .. 9

9. Ligula with teeth even in height, a straight apical margin and with inner margins of first lateral teeth curved outward (Figs. 367 and 368); mandible T,dth large basal and accessory teeth (Figs. 367 and 372); procercus about 6 times as long as wide (Figs. 370 and 371). Genus Pentaneura. . (!. inconspicua, p. 66)

Ligula with a concave arical margin and with inner ID.:lrgins of first lateral teeth curved outward (Figs. 396, 397, 411 and 413); mandible with basal tooth reduced (Fig. 394), often appearing absent (Figs. 389 and 390): procercus 2 to 3 times as long as wide (Figs. 393, 395, 409 and 412). Jhienemannimyia group genera...... 10 2!~

10. Naxillary palpal apical appendage b, 3-segmented (Figs. 26 and 394); occipital margin pale (Fig. 386). Genus Conchapelopia.... • ...... • . (~ . (~.) currani, p. 67 )

Maxillary palpal apical appendage b, 2-segmented (Fig. 26); occipi­ tal margin dark (Fig. 410). Genus Thienemannimyia . • • • • • •• . . • • • •• • • • • • • (:!:-. (:!:-.) norena, p.69)

KEY TO THE SUBFAHILIES OR TRIBES OF OTSEGO LAKE CHIRONOHID PUPAE

1. Thoracic horn with from 2 lumened branches to a tuft of many fine thread-like filaments sometimes difficult to see in slide mounts (Figs. 53-56) and abdominal segment VIII usually with a sclerotized spur or comb (Figs. 41, 79, 80, 84 and 85), or thoracic horn simple (Fig. 52), often with long hairs and abdominal segment VIII always with a sclerotized spur or comb; anal lobe always with a hair fringe (Figs. 41, 80, 84 and 85). Subfamily Chironominae.....•..•.....••.... 2

Thoracic horn not branched, always simple and never with lo~g hairs (Figs. 41, 43, 46-48 and 57-59) or thoracic horn is absent; abdominal segment VIII almost never with a sclerotized spur or comb; hair fringe or fringe of small spines along margin of anal lobe may be present or absent., . . • • . . . . . • . . . 6

2. Thoracic horn simple (Fig. 52), often with long hairs (Fig. 51) and abdominal segments II or III through V or VI always with paired dorsal spine patches (Fig. 41); abdominal segment VIII always with a sclerotized spur or comb (Fig. 41) and wing sheath almost always with a subapical nose (Fig. 43) . • . • • ...... •• ,. .. Tribe Tanytarsini

Thoracic horn with from 2 lumened branches (Fig. 55) to a tuft of many fine thread-like filaments (Figs. 53 and 56), sometimes difficult to see in slide mounts and rarely with paired dorsal spine patches on abdominal segments; abdominal segment VIII usually with a sclerotized spur or comb (Figs. 41, 79, 80, 84 and 85), though sometimes reduced (absent in many lIarnischia complex genera of the tribe Chironomini) and wing sheatl! almost never with a subapical nose (Fig. 43)...... 3

3. Thoracic horn a tuft of many fine thread-like filaments (Figs. 53 and 56). ...• .... Tribe Chironomini (in p£lrt)

Thoracic horn with from 2 branches (Fig. 55) to 12 branches of lumened tubules (Fig. 54) which are sometimes relatively wide (Fig. 55). . • ...... • . . . . . • . . . • • . . • . 4 25

4. Thoracic horn with 4- to 12-branched lurnencd tubules (Fig. 54). Tribe Chironomini (in part) ... (Paratendipes albirnanus, p. 29)

Thoracic horn with 2-branched lumened tubules (Fig. 55) .. . 5

5. Small species; abdominAl segments II-VI ~'lith paired dorsi'll spine patches (similar to some Tanytarsini) (Fig. 41); thoracic horn 2-branched narrow-lumened tubules. •. Chironorrini (in part)

Meditun to large species; abdominal segments lacking paired dorsal spine patches, but with more uniform shagreen (Fig. 41) and with anterior band or median patch of stronger dorsal spinules; thoracic horn 2-branched wide-lumened tubules ...... Tribe Pseudochironomini, Genus Pseudochironornus

6. Thoracic horn linear (Figs. 57, 189 and 209) or club-shaped (Figs. 377 and 401) with a conspicuous plastron plate or large and ovoid or sac-like with a dark and usually reticulate respira­ tory atrium (Figs. SA, 59, 288, 305 and 306); anal lobe with 2 large lateral setae located in approxirnatelv the middle third of lobe outer margin (Figs. 262, 285 and 338); hair fringe (Fig. 192) or fringe of weak or strong spines (Figs. 212, 263, 286 and 310) may be present or absent (Figs. 383, 400 and 418). Subfamily Tanypodinae...... (p.26 )

Thoracic horn, when present, generally smaller and linear and never with a plastron plate (Figs. 49, 50, 99, 116 and 152-154); anal lobe with 3 (Figs. 41, 44, 45, 104, 123 and 170) or more (Figs. 155 and 156) apical or subapical macrosetae; hair fringe on anal lobe may be present (Figs. 41, 44, ~5, 104, 121 and 151) or absent (Figs. 172 and 173)...... 7

7. Generally large species; posterior pair of leg sheaths curved under the wing sheaths and anterior 2 pairs distallv straight (Coffman and Ferrington 1984, Fig. 25.263). Subfamily Diamesinae . (p.26)

Generally smaller species; all leg sheaths curved under wing sheaths (as in most Chironomidae) (Coffman and Ferrington 1984, Fig. 25.264) or, rarely, all leg sheaths distally straight (op.cit., Fig. 252.265). Subfamily Orthocladiinae*...... (Psectro­ _cladius (R..) simulans. p. 32 and !!,ydrobaenus j ohannsenl:.. r· 34 )

*Tribal status not universally agreed upon. 2G

KEY TO THE TRIBES AlID GENERA OF OTSEGO LAKE DIMfESINAE PUPAE

1. Abdominal segment VIII with a ventral caudal lobe in male (Figs. 155, 156 and 158) or p~ir of lobes in female (Fig. 151); anal lobe with 4 to 8 stout apical macrosetae and a sparse hair fringe of 4 to 6 setae anteriorly (Figs. 151, 155, 156 and 158); pupal head with 2 pairs of cephalic tubercJes (Figs. 148 and 150), the posterior pair appearing on the prothorax of exuviae (Fig. 1l19); thoracic horn tapering to a point. Tribe Protanypodini, Genus Protanypus..... , (~. ramosus, p. 37)

Abdominal segment VIII without a ventral caudal lobe (Tribe Diamesini); anal lobe truncate, with 3 8ubapical macrosetae and a minutely denticulate apico-lateral projection and lacking a hair fringe (Figs. 170-173); thoracic horn absent; abdominal segments with branched lateral setae, dark anterior transverse banding, uniform dorsal median shagreen and from segments 111­ VIII, small intersegmental spinules (Figs. 168 and 169). Genus Potthastia (in part: longimana group, p. 39)

KEY TO THE TRIBES AND GENERA OR SUBGENERA AND SPECIES Of OTSEGO LAKE TANYrOD1Np~ PUPAE

1. Anal lobe greatly reduced and rounded, shorter than abdominal segment VIII and male genital sheath; with 2 lateral setae located in posterior half (Roback 1977, Fig. 62). Tribe Tanypodini, Genus Tanypus ...... •... (!. punctipennis, p. 45)

Anal lobe larger and either plate-like (Figs. 192, 212, 236 and 239) or tr~angular and pointed (Figs. 285, 309, 339, 362, 383, 400 and 417) with lateral setae variously located...... 2

2. Anal lobe plate-like, with a uniseral hair fringe in addition to 2 lateral setae located in the anterior half and with medial setiferous projections apically (Fig. 192); thorax with a lateral thoracic tubercle (Fig. 189). Tribe Coelotanypodini, Genus Clinotanypu~. . . . . • . (~. pinguis, p. 42)

Anal lobe plate-like or triangular and pointed but lacking hair fringe and medial setiferous projections and with lateral setae variously located; thorax lacking lateral thoracic tubercle but IDay have thoracic comb (Figs. 41, 288, 308, 334, 357 and 379) .

•••••••••••••• e. "," ••••••• 3

3. Anal lobe plate-like, bearing storng spines on outer margin and with 2 lateral setae located in the anterior half (Figs. 212, 236, 262 and 269); thoracic horn usually linear to club-shaped and 27

with a conspicuous apical plastron plate (Figs. 209 and 253­ 257) (except Procladius johnsoni which has a large and globose thoracic horn with a small apical nipple and no clear pl;Jstron plate, Figs. 230-233). Tribe Procladiini, Genus Procladius .. •••••••• ••• (,l~""""G ~.," ..... 4

Anal lobe triangular and pointed with (Figs. 286 and 310) or without (Figs. 339, 362, 383, 400 and 420) weak spines on outer margin and with lateral setae variously located; thoracic horn either large and ovoid with a reticulate respiratory atrium and aeropyle entering a small subapical papilla (Figs. 288, 305, 331 and 356) or thoracic horn club-shaf:,ed with an ameboid respiratory atrium (Figs. 416 and 419), or if a conspicuous plastron plate is present, it is always lateral and respiratory atrium is not ameboid (Figs. 376-379 and 401-40~. Tribe Pentaneurini. • . . . • . , . . . . • . . . • ...... 6

4. Smaller species; anal lobe with inner corners generally rounded (Fig. 212); abdominal tergites generally clear; thoracic horn linear and narrow, about 4 times as long as wide and with a conspicuous apical plastron plate (Figs. 209 and 210). Sub­ genus Psilotanypus. . . . (Procladius (P~.) bell us; p. LI 7)

Larger species; anal lobe ~.,ri th inner corners generally produced into one or more points (Figs. 236 and 263-270); abdominal segments sometimes clear, but generally patterned (Fig. 252); thoracic horn either large and globose, lacking a clear plastron plate (Figs. 230-233) or linear to club-shaped and wide, about 3 times as long as Fide and with a conspicuous Rpical plastron plate (Figs. 253-257). Subgenus Holotanypus. , . , . . • . . 5

5. Large species; thoracic horn large and globose with a small apical nipple and no clear plastron plate (Figs. 230-233). Procladius (~.) johnson!. ... 0 • • • • • • • •• (p.51)

Hedium to large species; thoracic horn linear to club-shaped and wide, about 3 times as long as ,vide and wi th a conspicuous apical plastron plate (Figs. 253-257). ProcLldius_ (Holo

6. Thoracic horn club-sh8red, with an ameboid respiratory atrium (Figs. 416 and 419) or with a conspicuous lateral plastron plate (Figs. 376-379 and 404-405) ,. . 7

Thoracic horn ovoid, Kith a dark reticulate respiratorv atrium and with an aeropyle entering a subapical papilla (Figs. 287-289, 305, 329-331, 333, 355, 356, 358 and 359)...... 9

7. Thoracic horn with an ameboid respiratorv atrium and lacking a plastron plate surrounding clear area and ostia (Figs. 416 and 419); thoracic comb absent and thoracic shagreen heavy (Figs. 416 and 417); abdominal sp~ments I (or II)-VIII with lateral setae. Genus ThienemannimEa...•.. c.:!'_. (1:..) norena, p. 69) 28

Thoracic horn with a prominent lateral plastron plate and surround­ ing clear area partly circumscribed by ostia and with respir­ atory atrium not ameboid (Figs. 377-379 and 401-405); thoracic comb present (Figs. 376, 378 and 379) or absent to weakly developed (Figs. 401 and 402); abdominal segments lacking lateral setae anterior to segment III...... 8

8. Anal lobe about 1.5 times as long as wide and with 2 lateral setae located in the anterior half (Figs. 382 and 383); thoracic comb present (Figs. 376, 378 and 379); abdominal shagreen uniform and sparse (Figs. 380-383) and with all spinules simple (Figs. 384 and 385). Genus Pentaneura .•.. (I. inconspicua, p. 66)

Anal lobe about as long as wide and with 2 lateral setae located in the posterior half (Fig. 400); thoracic comb absent or weakly developed (Figs. 401 and 402); abdominal shagreen uniform and heavy (Figs. 399, 400 and 406) and with numerous multirayed spinules (Fig. 407). Genus Conchapelopia...... • . . • . . (~. (~.) currani, p. 67)

9. Small species; anal lobe about 1.5 times as long as wide with outer margin bearing weak spines and with 2 lateral setae located centrally (Figs. 282, 285 and 286); respiratory atrium of thoracic horn finely reticulate (Fig. 288). Genus Labrundinia. • . . • . • • • . • • • • • . . . • • • • (.!=.. pilosella, p. 55)

Larger species; anal lobe abont as long as wide and generally with outer margin smooth and 2 lateral setae located in the poster­ ior half (Figs. 338, 339, 361 and 362) (except which h<1s \.,reak spines on outer margin and lateral setae located approximately centrally, Figs. 309 and 310); respiratory atrium of thoracic horn coarsely reticulate (Fig. 331) . Genus Ablabesmyia...... • . . . . . •• 10

10. Large species; abdominal tergites with very heavy uniform shagreen with numerous multirayed spinules (Figs. 307, 309 and 311-313); anal lobe with weak spines on outer margin and 2 lateral setae located approximately centrally (Figs. 309 and 310). Ablabesmyia (!:...) annulata...... • (p. 53)

Medium sized species; abdominal tergites with very sparse shagreen and thorax and abdomen generally with some pigmontation (Figs. 336, 364 and 365); anal lobe with outer margin smooth and 2 lateral setae located in the posterior half (Figs. 338, 339, 361 and 362)...... 11

11. Wing sheath \.,rith major adult veins pigmented (Fig. 336); aeropyle of thoracic horn of uniform Iddth throughout its length (Fig. 333). Ablabesmyia (!:...) basalis (and others). . • • . . . .. (p. 60 )

Wing sheath and abdomen heavily pigmented (Figs. 364 and 365); aeropyle of thoracic horn bulbous distally (Figs. 358 and 359). Ablabesmyia (!:...) mallochi...... • • . . . .. (p. 63 ) 29

Subfamily Chironominae

Tribe Chironomini

Paratendi~~ ~lbima~~ (Meigen) 1818

(Plates 15-18; Figs. 60-85)

Nwnerous associations Rnd fourth instar larvae of £~ratendi~ albimanus were obtained from about one meter of water on multiplate samplers in Rat Cove (Map 1) and in littoral grab samples in Rat Cove and Hyde Bay in Otsego Lake in June and July of 1980. One fourth instar larva was also obtained from a bottom feeding whitefish stomach collected in 20-30 meters of water off of Peggs Point in June of 1981. Early instar larvae were particularly abundant in the deltiac fan of Shadow Brook in Hyde Bay during April and May of 1981.

Although adult males of seven species of Paratendip~~ are known from North America (Townes 1945 '1nd Sublette 1966), the immature stages of only ~. albimanus have been described. The larva and PUP3 I.... ere described by Johannsen (1937, p. 27 as Chironomu~ (Paratendi~) albimanu~) and Lenz 0954-1962). Chernovski (949) figured cephalic structures of P. albimanus (mandible, ;Intenna, mentum and ventro­ mental plate) ~nd~ncluded it in a key to four larval species (or "types") of Paratendip~~. Ward and Cummins (978) provided a thorough account on the biology of P. albimanus and on the morpho­ logical changes occurring with 1arv;-1 growth, as well as figuring the mandible, antenna, mentum and ventromental plates for all four larval stadia.

Larval characteristics:

Medium to moderately small Size.

Occipital margin dark and gula darkened posteriorly (Figs. 60, 61 and 65).

Antenna with 6 segments dnd with Lauterborn's organs alternate at apices of segments 2 and 3 (Figs. 62 and 71).

Labral setae I and LI plumose and labral seta I truncate apically and I¥ith bases fused (Figs. 69 and 70).

Pecten epipharyngis consisting of 3 simple plates apically attenuate (Fig. 69).

Premandible dark and with 2 apical teeth (Figs. 60 and 67-69). JO

Nandible with a short, dark and bluntly roundpd apical tooth, two dark inner teeth and a strong, pale dorsal tooth (Figs. 62, 66 and 67).

Mentum with 4 pale median teeth equal in height and 6 pairs of dark 1ate r a 1 tee t h; me d ian tee t h s harte r t han secand lateral teeth and first lateral teeth fused to and also shorter than second lateral teeth (Figs. 60, 62, 67 and 68) .

Ventromental plates widely separated and ;~ith coarse striae; the median apices are anteriorly produced and in contact with the median part of mentum (Figs. 67 and 68).

Procercus small, about as long as wide with 8 anal setae (Fig. 63) •

Pupal characteristics:

Medium to moderately small Slze.

Frontal apotome with prominant cephalic tubercles with long setae (Figs. 72, 73 and 75).

Thoracic horn with up to 12 lumened branches which are often difficult to see ln slide mounts.

Wing sheath with a dark pigmented border (Fig. 74).

Abdominal segment II with an uninterrupted dorsal, posterior row of recurved hooks (figs 78, 81 and 82); segments III-IV and IV-V with intersegmental spinules (Figs. 78 and 83); and segments II-VI with a dorsal, median shagreen which divides postero-medially on III (sometimes II) through VI (Figs. 78 and 83).

Pedes spurii A present on abdominal sternites III and IV (Fig. 77) •

Pedes spurii B present posteriorly on abdominal segment II and weakly indicated anteriorly on segment I (Figs. 74 and 76 ).

Abdominal segments V-VIII with 3, 4, 4 and 4 lateral setae, respectively, and lateral setae lacking on anterior segments (Fig. 79).

Abdominal segment VIII with a sclerotized comb with from 3 to 5 (or 6) teeth and some finer spinules (Figs. 79,80,84 and 85) .

Anal lobe with uniseral hair fringe (Figs. 80, 84 and 85). 31

Larvae of P. albimanus are distinguishable from other genera and species by hav(;:;-g ~ mandible with 2 dark inner teeth and a pale dorsal tooth, a mentum with 4 pale, equally sized median teeth which are always lower than the second lateral teeth and a 6-segmented antenna with alternate Lauterborn' s organs. The pupa is distinguished by the combination of most characteristics given above.

Paratendipes albimanus is common and widely distrl.buted in lotic and lentic habitats in North America (Townes 1945, Sublette 1960, Ward and Cummins 1978, Oliver and Roussel 1983 and Pinder and Reiss 1983). Townes (1945) reported examining specimens from an extensive number of collection localities in central New York, includlng Otsego Lake. Simpson and Bode (1980) reported collecting P. a1bimanus from mu1tip1ate samplers in streams in Ne\1I York Stat~~e writer has also collected larvae from Ashokan Reservoir on Esopus Creek, Esopus Creek, Schoharie and the Upper and Lower Blenheim-Gilboa Pumped Storage Reservoirs on Schoharie Creek and the Hudson River in New York.

Townes (1945) provided the following synonymies for Paratendipes a1bimanus:

"Chironomus annularis Meigen, 1804, Klass. Beschr. zweif1. Ins., p. 17; type 10cality:?France (?Paris museum). This reference not seen. Chironomus albimanu~ Meigen, 1818, Syst. Beschr. Europ. zweifi. Ins. 1:40. New name for Chironomus annularis Meigen, 1804, not (De Geer) Meige-;:;-I804 (~s ~-.--~- annulatus). 1l1e necessity for continuing this change in name is problematic. Chironomus albimanus Johannsen, 1905, Bull. N. Y. State Mus. 86:214~scription. Paratendipes albimanus Kieffer, 1911, Bull. Soc. d'Hist. Nat. Het~41-;-descript ion, generic pas i t ion. Chironomus (Par~~ndi~) albimanus Johannsen, 1937, Mem. Cornell Univ. Agr. Exp. Sta. 210:27; description of larva and pupa. Chironomus ~Eimanus Townes, 1938, Ann. Rpt. N. Y. State Conservation Dept. 27, suppI.: 165, 171; biology."

More recent literature records on P. albimanus are numerous. --~--- Ward and Cummins (1978, Table 1) listed the following additional references: Nietzke (1938), Brunelin (949), Thienemann (1950), Hall (1951), Curry (1954), Dittmar (1955), Roback (1957), Reiss (1968a and 1968b), Iovino and Minor (1970), Lehman (1971), and Learner and Potter (1974). Ward and Cummins (1979) subsequently reported on trophic relations in Paratendipes ~~imanu~ and Harper and Cloutier (1979) on emergence phenology. 32

Subfamily Orthocladiinae

Psectrocladius (Psectrocladius) simulans (Johannsen) 1937

(Plates 19-21; Figs. 86-104)

Several associations and fouth ins tar larvae of Psectrocladius simulans were obtained from littoral grab samples in Rat Cove (Map 1) and the opposing shore in Otsego Lake from mid-May through the end of July in 1980-1982. The immature stages were first described by Johannsen (1937, p. 67) and briefly by Roback (1957, p. 90) and Saether (1969, p. 83). Diagnosis and keys to subgenera and species groups of Psectrocladius larvae are presented in Cranston et al. (1983, p. 192).

Larval characteristics:

Medium to moderately small S1ze.

Occipital margin pale.

Antenna 5-segrnented, with each segment consecutively smaller and with Lauterborn's organs small, distal and opposite on segment 2 (Fig. 90).

Labral seta I palmate, with from 5 to 7 (or 8) branches (Figs. 89, 92 and 94).

Pecten epipharyngis consisting of 3 simple spines subequal 1n length (Fig. 89).

Premandible simple, dark, broad and moderately curved and attenuated apically (Figs. 89 and 90).

Mandible with an attenuate apical tooth which 1S longer than the combined width of the 3 inner teeth; tip of apical tooth and the 3 inner teeth are dark (Figs. 86, 87, 90 and 94) .

Mentum with 5 pairs of d~rk lateral teeth and with a single median tooth sometimes appearing bifid apically, which is mostly pale except for an apical median dark spot (Figs. 86, 93 and 95).

Ventromental plates present and distinct, triangular and with a sparse beard beneath (Figs. 93 and 95).

Procercus lightly sclerotized with 4 or 5 small basal spurs and 5 anal setae (Figs. 88 and 91). 33

Pupal characteristics:

Medium to moderately small Size.

Frontal apotome with distinct cephalic tubercles.

Thoracic horn linear, slightly wid~~ned distally and with about 12 major spines on outer margin (Figs. 97 and 99).

Abdominal segment II with posterior rows of dorsal recurved spines; segments Ill-VI with posterior rows of dorsal spines; and tergites IV-VI with transversely oval spine patches centrally (Figs. 96, 98, 101 and 103).

Pedes spurii A and pedes spurii B lacking on abdominal segments.

Abdominal segments VII and VIII with 4 and 5 lateral setae, respectively, and lateral setae lacking on anterior segments (Figs. 102 and 103).

Anal lobe with 3 apical m~crosetae and a uniseral hair fringe with setae longer than 1/2 the length of the lobe (Figs. 100 and 104).

Both the larva and pupa presented here key to Psectroc~c1ius elatus Roback rather than to P. simulans in Roback (1957, p. 86). Despite some discrepancies, these-associated specimens undoubtedly belong to P. simulans. According to Saether 0969, p. 84), the smaller body size and fewer spines in the central patches of the pupal tergites exhibited in the Otsego Lake specimens, fall within the limits of variation in other Psectrocladius species and should not be used as characteristics to separate the pupa? of P. elatus and P. simulans. The larva seems to agree with descriptIons of P. simulans fairly well except that the premandible (Figs. 89 and 90)-is dark, broader and slightly less curved and attenuaU~d apically (see Saether 1969, Fig. 41 E and Oliver et a1. 1978, Fig. 137).

The larvae of Psectrocladius species are quite distinctive. The palmate labral seta I is unique. The combination of the large ventromental plate and beard beneatll, the simple premandible and spurs on the procercus are also characteristic (Cranston et al. 1983). T11e pupae are distinguished by an anal lobe with a hair fringe of long setae and 3 apical macrosetae, the abdominal tergite central spine groups and posterior spine rows and the prominant cephalic tubercles.

Johannsen (1937, p. 67) collected the holotype male and allotype fema Ie reared from larvae in C:Ula 1 Pond, New York. Simpson and Bode (1980, p. 57) reported a £sil~~terus group species, appearing to be this species, as being found in moderate to large rivers in New York. The writer has also obtained associations in New York State from Ashokan Reservoir on Esopus Creek and Schoharie and the Upper Blenheim-Gilboa Pumped Storage Reservoirs on Schoharie Creek. 34

Sublette (1967) provided the following synonymies for P. simulans:

"Spaniotoma (Psectrocladius) simulans Johannsen, 1937, 205:67. Description of larva, pupa, male and female. Hydrobaenus (Psectrocladius) ~imulans (Johannsen); Johannsen in Johannsen and Townes, 1952, 80: 22. In key. Psectrocladius simulans (Johannsen); Roback, 1957, 9:90. Further description of larva and pupa. Psectrocladius simulans (Johannsen); Sublette and Sublette, 1965, 276:159-.-0Istribution. "

Psectrocladius (P.) vernalis f1alloch (described in the immature stages by Sublet~64b, p. 119) was also frequently collected in littoral areas of Otsego Lake.

Hydrobaenus johannseni (Sublette) 1967

(Plates 22-24; Figs. 105-123)

One specimen of Hydrobaenus johannseni was collected and associated from Otsego Lake in this study. A skim of the waters' surface with a fine sieve near the shoreline of Rat Cove (Map 1) on 23 April 1982 contained a pupal exuviae with the larval skin attached and encompassing the abdomen on the pupal skin. Though common in the Chironominae, this is the first association obtained by the writer of this type in another subfamily over many years of collecting. Usually, the larval skins are quickly shed by newly formed pupae. Both immature stages are fairly easily identified using the keys of Saether (1976, p. 75) who also provided the first immature descriptions (op. cit., p. 138). Diagnosis and key to species groups of ~robaenus larvae are presented in Cranston et a1. (1983, p. 175).

Larval characteristics:

Medium size.

Occipital margin dark.

Antenna 6-segmented, with each segment consecutively smaller and segment 6 hair-like and often difficult to see; Lauterborn's organs are distal and opposite on segment 2 and slightly shorter than segment 3 (Figs. 106 and 113).

Labral seta I plumose (Fig. 111).

Pecten epipharyngis consisting of 3 simple spines subequal in length (Fig. Ill). ]')

Premandible dark and ,,lith 2 apical teeth.

Mandible dark in distal half and with apical tooth bluntly rounded and much shorter than the combined width of the 3 inner tee th (Figs. 106 and 107).

Mentum dark, ,,,ith 2 median teeth and 6 pairs of lateral teeth (Figs. 108, 110 and 112).

Ventromental plates present and distinct, rounded posteriorly and lacking a beard beneath (Figs. 108, 110 and 112).

Procercus heavily sclerotized with 7 aLaI setae (Fig. 109).

Pupal characteristics:

Medium size.

Frontal apotome with small cephalic tubercles (Fig. 114).

Thoracic horn linpar, of uniform width and spinulose, with stronger spines on outer margin (Figs. 115 and 116).

Abdominal segment II with a weak posterior, median shagreen and posterior rows of weak dorsal recurved spines (Fig. 118); segments II-VI with a dorsal, median shagreen with spinules increasing in size posteriorly (Figs. 117-120).

Pedes spurll A present on abdominal sternites IV-VIII (Fig. 122) .

Pedes spurii B present posteriorly on abdominal segment II.

Abdominal segments VII and VIII with 4 and usually 5 lateral setae, respectively, and lateral setae lacking on anterior segments.

Anal lobe with an apical rugosity at base of 3 apical macrosetae, a uniseral hair fringe mostly concentrated in basal half with setae shorter than 1/2 the length of the lobe, and genital sheaths of male hearly reaching the end of anal lobe and bearing short apical projections (Figs. 121 and 123).

Hydrobaenus iohannseni is the second most common Hydrobaenus species in temperate North Ameri.ca (Saether 1976). The larva is extremely similar to the more common !!. pi.lipes and is generally distinguishable by the ring organ of antennal segment 1 being 5-10 micrometers (x=8) from the base as compared to 10-20 micrometers (x=14) in!!. .£ilipes and by other characters gi.ven by Saether (1976). The pupa is fairly distincti.ve and easily distinguised from!!. Eilipe~ by the absence of filamentous lateral setae on abdominal segment VI and 36

by the presence of an apical rugosity at the base of the macrosetae on the anal lobe (Figs. 121 and 123).

Larvae of Hydrobaenus sppcies are morphologically simil~r to several genera including Zalutschia and (Cranston et al.1983). Zalutschia is~-Iso found in Otsego Lake (Table 1).

Saether (1976) reported the holotype male and para types from Ithaca, New York, another male collected at Tonawanda Creek, Genesee Co. in New York State and the following synonymies:

" nivoriundus Johnson 1900: 627, nec Fitch (misidentification of Chironomus ( = ) nivoriundus Fitch); Johannsen 1905: 274 (male, female, pupa, larva, pupa and larva probably only in part), 1934: 348 (synonomy); Kieffer 1906b: 28; Potthast 1915: 361 (pupa, larva) Orthocladiu~ nigritus Malloch 1915: 525, pro parte (see Sublette 1966: 594 and below) Trissocladius nivoriundus (Joh.) Thienemann 19"f4: 585,588, 534 (pupa, larva) Hydrobaenus (Chaetocladius) nivoriundus (Joh.) Johannsen 1947: 173 (male, female) Hydrobaenus (~obaenus) nivoriundus (Fitch) Johannsen 1952: 23 (male, female) Hydrobaenus nivoriul1dus (Joh.) Roback 1957: 76, 81 (pupa, larva) Orthocladius domus Sublette 1966: 594 (male, syn.n. (based on a paratyp~ male of Q. nigritus Mall.) Orthocladius (Orthocladius) johannseni Sublette 1967: 504 (male, female), new species for Q. nivoriundus Joh. nec Fitch Trissocladius domus (Subl.) Saether 1969: 42 (generic placement~n.n. Trissocladius hamiltoni Saether 1969: 42 (male, pupa, larva), syn.n. nec Orthocladius nivoriundus Fitch Malloch 1915: 525 (see Sublette 1967: 504) nec Dac ty loc lad ius !.:i vor iundus Kie f fer 1917: 359 nec Fi tch , probably a synonym of ~. pilipes Mall. nec Spaniotoma (Orthocladius) nivoriunda Joh. nec Fitch Johannsen 1937: 64 (pupa, larva). At least the pupae belong to ~. pilipes (see p. 115). nec Trissocladius joha~nseni Saether 1969: 45 (generic placement) synonym of H. pilipes Mall. (see p. 115)." 37

Subfamily Diamesinae

Tribe Protanypodini

Protanypus ramosus Saether 1975

(Plates 25-29; Figs. 124-159)

Although a common constituent of the Otsego Lake profundal community based on whitefish stomach analyses, the larvae of Prota~ ramosu~ were rarely collected in grab samples. None were reared or associated in this study. Numerous fourth instar larvae and mature, pharate male and female pupae were obtained from bottom feeding whitefish stomachs collected in 20 to 30 meters of water off of Kingfisher Tower and Peggs Point (Map 1) in September-October 1980 and ~n May of 1981. Several adult males and females were also collected in May of 1981 and early June ~n 1979.

Protanypus is a small genus with only 5 species described from North America and one possibly Holarctic species (Saether 1975, Wiederholm 1975 and Doughman 1983). Two species are known only as larvae (Saether 1975). P. ramosus was first described by Saether (1975, p. 368) in all stages.

Larval characteristics:

Large size.

Antenna with 4 segments (Figs. 137-139),

Occipital marg~n and gula region dark (Figs. 124 and 142).

Occipital margin with a distinct, posteriorly directed ventro­ lateral projection on each side (Figs. 8, 124 and 129).

Head capsule with numerous closely spaced long setae (Fig. 125) .

Labrum with an overlapping row of from 14 to 16 apically serrate lamellae (Figs. 132 and 136).

M appendage of premento-hypopharyngeal complex strongly developed and deeply divided into lamellae (Figs. 133 and 135 ).

Mandible with a long and dark, attenuate apical tooth and 5 dark inner teeth (Figs. 131, 137 and 139). 38

Premandible narrow, straight and with weak lnner teeth (Figs. 133 and 135).

Maxilla with apically serrate lamellae present dorsal to palp (Fig. 134).

Mentum without distinct teeth on middle two-thirds, with a palr of larger, extreme lateral teeth and with one or two smaller, more distal lateral teeth on each side (Figs. 130, 140, 141 and 143).

Posterior parapod with some (2 to 3) smaller, hooked claws with denticles on inner margin (Figs. 144-147).

Pupal characteristics:

Large Slze.

Pupal head with 2 palrs of cephalic tubercles discernable (Figs. 148 and 150); the posterior pair of tubercles appear on the pro thorax and the anterior pair on the frontal apotome of exuviae (Fig. 149).

Thoracic horn linear, tapering to a point with generally very fine spinules for most of its length (Figs. 152-154, 157 and 159).

Abdominal segment I with ventral shagreen consisting of thin spinules 5 to 6 microns long on posterior 1/3 to 1/5 of sternite.

Abdominal segments II-IV and V-VIII with 2 and 3 lateral setae, respectively.

Abdominal segment VIII with a ventral caudal lobe ln male (Figs. 155, 156 and 158) and a pair of lobes ln female (Fig. 151).

Anal lobe with 4 to 8 stout apical macrosetae and a sparse uniseral hair fringe anteriorly of 4 to 6 setae (Figs. 151, 155, 156 and 158).

Pedes spurii A and B lacking on abdominal segments.

Protanypus is the only genus in the tribe Protanypodini (Oliver and Roussel 1983, p. 158). The larva is easily recognized and very distinctive in that the head capsule bears numerous long setae (Fig. 125), a pair of large, posteriorly directed ventro-lateral projections (Figs. 8, 124 and 129), mental teeth that are confined to the extreme lateral margins (Figs. 130, 140, 141 and 143) and in other characters given above. The pupa is apparently also unique in several characteristics including abdominal segment VIII bearing a ventral caudal lobe in males (Figs. 155, 156 and 158) and a pair of lobes in females 39

(Fig. 151). Also, the anal lobe bears 4 to 8 stout apical macrosetae and a sparse uniseral hair fringe anteriorly of 4 to 6 setae (Figs. 151, 155, 156 and 158) and the prothorax bears a pair of anterior thoracic tubercles (Fig. 149).

Prota~E~s larvae presented here key to Protanypus sp. B in Saether 0975, p. 385) and to P. saetheri Wiederholm in DoughmAn (1983, p. 50). However, adult-malesand females collected at Otsego Lake key to and agree very well with Saether's (1975, p. 368) descriptions of P. ramosus and the immature stages are undoubtedly this species. Saethe~---rc;p. cit.) used the number of labral lamellae (scales) as a principle character for separating P~otanypus larvae. Based on 2 fourth instar larvae in that study, a count of 11 to 12 lAbral lamellae was obtained for ~. ramosus and used in the key. Individual lamellae are often difficult to discriminate and count, but a larger series of carefully mounted Otsego Lake specimens indicate 14 to 16 (or 17) labral lamellae are usually discernable (Figs. 132 and 136). The pupae collected agree very well with Saether's (op. cit.) description except that the ventral caudal lobe of ahdominal segment VIII in the male is apically rounded (Figs. 155, 156 and 158) rather than truncated and the thoracic horn is occassionally without distinct spinules (Figs. 152 and 153) .

This is apparently the first collection record of Protanypus in New York State. North American cn1lection records reported by Saether (975) included: Miller (941), Oliver 0963, 1964 and (968), Hamilton (1965 and 1971), Brinkhurst et a1. (1968), Hiltunen (1969), Saether (1970) and Saether and McLean (1972). Additional records include: Wiederholm (1975), Oliver et a1. (1978), Doughman (1983) and Oliver and Roussel (1983). Saether (1975, p. 375) examined specimens of P. ~amosus from numerous 01 igotrophic lakes and mentioned (p. 386) that the Pro tan~12 t a xa dis c usse d by l-Ji 11 e r (l941 ), Br inkh ur s t eta1. (1968), Hiltunen (1969) and Hamilton (1971) are P. ra~osus. Johannsen (1937, p. 45) described the larva of B~il1i;~ (7) from Perch Lake, Ontario which is a Protanypus species differing little from ramosus.

Tribe Diamesini

Potthastia longimana Kieffer 1922

(Plates 30-31; Figs. 160-173)

Only 3 specimens of Potthastia longimana from the profundal zone of Otsego Lake were obtained in this study. All three were from bottom feeding whitefish stomachs collected in 20 to 30 meters of water off of Kingfisher To''''er and Peggs Point (Map 1) in May and early June of 1981. 40

Two specimens were pupae, each containing a mature male and female within and the third was a larva, ~lich because of its small size was considered to be a third instar.

The immature stages of Potthastia longimana were described by Pagast (1933, larva as Diamesa and 1947, p. 536, pupa) and Pankratova (1970). The larva was figured by Roback (1953) and keyed and figured in Mason (1973), Oliver et al. (1978), Simpson and Bode (1980) and Oliver and Roussel (1983). Oliver (1983, p. 120) also discussed the group designation and diagnoses in larvae of Potthastia.

Larval characteristics:

Large size.

Occipital margin dark (Fig. 160).

Antenna with 5 segments; antennal segment 3 with areas of reduced sclerotization giving the appearance of annulations; antennal segment 2 longer than segment 1 and a 1so longer than the comb ined length s of the th i rd to fifth segments (Figs. 161 and 162).

Labral seta I bifid (Fig. 164).

Premandible with 11 to 16 teeth and usually fan-shaped (Fig. 165 and Oliver 1983, Fig. 7.8).

Mandible with a long, attenuate apical tooth, three small inner teeth, a large hook-shaped median process and lacking a seta interna (Fig. 166 and Oliver 1983, Fig. 7.8).

Mentum undulate, entirely without teeth and ventromental plates vestigial (Figs., 163, 164, 166 and Oliver 1983, Fig. 7.8).

Pupal characteristics:

Large size.

Thoracic horn absent.

Abdominal segments with branched lateral setae, dark anterior transverse banding, uniform dorsal median shagreen, and from segments III-VIII, small intersegmental spinules (Figs. 168 and 169).

Anal lobe truncate, with 3 subapical macrosetae, a minutely denticulate apico-lateral projection and lacking a hair fringe (Figs. 170-173).

Pedes spurii A and B lacking on abdominal segments. 41

Although additional species may be involved in the longimana group in North America, the char,qcteristics presented of .!"'ots.!:J~stia 19n9imana larva and pupa agree for the most part wi th the limited amount of descriptive information available. 'lne larva of the P. longimana group are unique among the Diamesini in several characteristics including the toothless mentum (Figs. 163, 164 and 166), the apically attenuated mandible (Fig. 166) and the long second antennal segment (Figs. 161 and 162) (Oliver 1983, p. 121). TIle singlp, early instar larva collected appears to differ from the fourth instar mainly in the smaller body size and in the premandibles being some\"hat club-shaped (Fig. 165) rather than fan-shaped (e.g., Oliver 1983, Fig. 7.8).

Potthastia longimana is apparently widely distributed in North America (Coffman and Ferrington 1984, the above cited works and many accounts in the literature). Simpson and Bode (1980, p. 29) reported .!"'.. longiman~ as occurring occassionally on multiplate samplers at 4 unspecified sites in New York. The writer has also collected larvae from the Susquehanna River at Oneonta, and in Esopus Creek in New York State. 42

Subfamily Tanypodinae

Tribe Coe1otanypodini

Genus C1inotanypus Kieffer 1913

Subgenus C1inotanypus Kieffer 1913

"Ta~: Loew 1861 (Part):308, 309. ~oc1adius: Johannsen 1905 (Part):129-133. Psi1otanypus: Kieffer 1906b (Part):40. Clinotanypus Kieffer 1913b:157. Proc1adius: Malloch 1915a (Part):391. Clinotanypus: Edwards 1929:302. Clinotanypus: Johannsen 1934:347. C1inotanypus: Johannsen 1952:11. C1inotanypus: Roback 1957b:46.

Clinotanypus: Sublette and Sublette 1965:145. II

C1inotanypus (C1inotanypus) ~U1S (Loew) 1861

(Plates 32-34; Figs. 174-192)

C1inotanypus ringuis is ubiquitous in the northeast (Roback, 1971) and was found frequently in the shallow water sediments of Rat Cove in Otsego Lake (Map 1). ~. pinguis was not reared to the adult, but numerous larval and pharate adult male and female associations were obtained. The immature stages are quite distinctive and easily identified using Boesel (1974) or Roback (1976b). Two point four meters was the maximum depth that this species was collected in grab samples in Otsego Lake which is also (and probably coincidently) the maximum depth that Roback (1976b) found it.

Larval characteristics:

Large size.

Occipital margin and head capsule reddish-brown (Figs. 174 and 180) .

Mandible strongly curved and with a large, pointed basal tooth (Figs. 175 and 177). 43

Ligula usually with six pale teeth (rarely 5 or 7) and a concave apical margin (Fig. 176).

Maxillary palp with a large basal segment and protracted distal segments on a membranous apex (Figs. 177, 181 and 182).

Mentum with 6-8 separate, loosely arranged dorsomental teeth in a longitudinal row which are not on a distinct plate (Fig. 177).

Abdomen with a lateral hair fringe (Figs. 187 and 188) and a conical papilla between bases of procerci (Boesel 1974, Fig. 18 and Oliver and Roussel 1983, Fig. 20).

Procercus about 2.5 to 3 times as long as wide with about 14 anal setae (Figs. 178 and 179).

Pupal characteristics:

Large size.

Thoracic horn large and linear with a dark respiratory atrium and prominant apical plastron plate (Fig. 189).

Lateral thoracic tubercle present (Fig. 189).

Abdominal segments VII and VIII with 6 (sometimes 5) and 5 lateral setae, respectively, and lateral setae lacking on anterior segments (Fig. 190).

Anal lobe plate-like and slightly wider than long, with a uniseral hair fringe in addition to 2 lateral setae located in the anterior half and with medial setiferous projections apically (Fig. 192).

Roback (1971) reported adults collected at Buffalo, Babylon, Bemus Pt., Ithaca, Lewiston, Mayville, Mud Pond at McLean Res., and Otsego Lake in New York State, other extensive accounts in the northeast and also provided the following synonymies for this distinctive species:

"Ta~ pinguis Loew 1861:308 (adult description). Tanypus flavicinctus Loew 1861:309 (adult description). New Synonym. Tanypus thoracicus Loew 1866:4 (adult description). New Synonym. Procladius thoracicus: Johannsen 1905:129 (adult male). Procladius caliginosus Johannsen 1905:131 (adult female). New Synonym. Procladius pinguis: Johannsen 1905:133 (adult and larva). Procladius flavicinctus: Johannsen 1905:132 (adult male). Procladius pinguis: Kieffer 1906b:39 (adult, generic position). Psilotanypus flavicinctus: Kieffer 1906b:39 (adult, generic position). Psilotanypus thora£icus: Kieffer 1906b:40 (adult, generic position). Procladius caliginosus: Kieffer 1906b:39 (adult, generic position). Proc1adius or .!:s il~~~~~ flavic inc tus: Johannsen 1908:270 (generic position). Procladius thoracicus: Johannsen 1908:270 (generic position). Procladius thoracicus: Malloch 1915a:391 (adults). ----~-- C1 inotanypus flavic inc tus, ca 1iginosus, thoracicu~, pin~uis: Johannsen 1934:347 (generic position). Clinotanypus pinguis: Johannsen 1937:25 (pUpil). Clinotanypus thoracicus: Morrissey 1950:89 (distribution and immature stages). Clinotanypus thoracicus, flavicinctus, pinguis, caliginosus: Johannsen 1952: 11 (adults in key). C1inotanypus flavicinctus?: Neff 1955:7 (immature stages and adult biology). Clinotanypus thoracicus?: Roback 1957b:46 (immature stages). Clinotanypus ~is: Roback 1957b:46 (pupa in key). Cl inotanypus thorac icus: Beck and Beck 1959: 91 (distribution). Clinotanypus caliginosus, thoracicus, flavicinctus, pinguis: Sublette and Sublette 1965:145, 146 (generic placement and distribution). Clinotanypus prob. pinguis: Roback 1969a:16 (food of larva)."

Roback (1976a) provided a subsequent note on feeding habits.

Tri be Tanypod ini

Genus Tanypus Meigen 1803

Subgerrus Tanypus Meigen 1803

"Pelopia Meigen 1800:18 (name suppressed, ICZN 1963b). Ta~ Meigen 1803:261; Type species Tanypus pun£!i£ennis Meigen 1818 (New name for T. cincta Meig. 1803 nee. Fabr.). Tanypus: Johannsen 1905 (part):121-125. Protenthes Johannsen 1907b:400; Genotype T. punctipennis ----Me i g--: Kieffer 1913a:13 (nee. Kieffer 1906a:319)." 4S

Tanypus (.!anypus)punctipennis ~leigen 1818

(Plates 35-36; Figs. 193-201)

Only one larva of in the fourth ins tar was collected in a grab sample at the base of Nupha~ vari~~atum in about one meter of water in Rat Cove (Map 1) during mid-July, 1981. This individual did not pupate and no pupa were otherwise obtained .

.!. ~nctipennis is the type species for the genus TanYE~~ (Roback 1971). The immature stages were describ~d by Grodhaus (1967, p. 3), Roback (1969b, p. 414 and p. 415 as carinat~ var.) and Roback 0977, p. 75).

Larval characteristics:

Moderately large Size.

Occipital margin dark (Fig. 193).

Mandible with a short apical tooth, minute basal and accessory teeth and with basal two-thirds expanded (Fig. 195).

Ligula with five pale teeth and a convex apical margin (Fig. 198).

Paraligula with main fork about half-way from base (Fig. 198).

Mentum with dorsomental teeth fused into distinct plates which are joined medially (Fig. 194).

Dorsomental teeth dark and middle tooth darkly pigmented along apical margin (Fig. 194).

Anal tubules in three pairs (rather than the usual two pairs) (Fig. 199).

Abdominal segments with a lateral hair fringe (as in Clinotanypus pinguis, Figs. 187 and 188).

Procercus about 4.5 to 5 times as long as wide with about 14 pale anal setae (Figs. 197 and 200).

Unfortunately, the pupal stage was unavailable for photographing in this study. Characteristics used by Roback (1977) for distinguishing the pupa included:

Moderately large Size.

Thoracic horn large and ovoid, asymetrical1y biconvex with a dark reticulate respiratory atrium, a small apical nipple 46

and no clear plastron plate (e.g., Fig. 59).

Abdominal segments II m: III tilJ:ough VIII with lateral setae; generaily 5 lat G f21 setae on segment VIII And num­ erous but iees tnan 50 leteral setae on segments VI and VII and on ant2clor segmeuts (op. cit., p. 66 and Table 3).

Anal lobe greatly reduced and rounded, shorter than abdominal segment \.~l and male genital sheath Rnd with 2 lateral setae located in the posterior half (op. cit., Fig. 62).

Roback (1977) reported larvae collected at Ste~ling Forest, New York and Roback (1971) reported adult males collected from Lechworth State Park in New York State" The "Triter has reared .:!:. punctipennis from Iroquois Lake in Central Park, Schenectady, New York. Roback (1971) also presented (~xt2n8ive accounts d' this holaretic species with a wide distribution in North America.

Roback (1971 as amended by Roback 1977) provided the following synonymies for 1. pu~ctipenni~:

"Tanypus puncti:.E_e11l'-~~ Me~F2n 1818:61 (adult description). Protenthes ~~~tipenu~~: Maltoch 1915a(Part):383 (description of pupa and adljlts)~ £~~c t~_12-ennis Protenthes ViE 0 americanus? Kieffer 1923a:297 (SC2tUS ~ncert~in, probably female of stellatus) ~ Tanypus p~nsti~enni~: Edwards 1929:299 (brief description of aaults)o nee. T5!pus 12u,,':.tipen~::is: Johannsen 1937;19. Janypus stei-~t\J3: Johar;r,se" 1937:20 (irmnatures). neco Ta.ny~us EI}ct~L.nnis: Hortissey 1950:90 nee .. Pelopia pu.ncci2en~.~~~ Johannsen 1952:10. nee. Pelopia Eunctj-E_~_~~LJ.is: Neff 1955:4" Pe10pia stellata: Rob,s.cki957b:48 (irm~1atures).. nee. Tanypus ~~~~E":~,,i;3: Beck and Beck 1959:91. nee. Pelopia PU-'2::J:.~?e[\nis; Dendy Bnd Sublette 1959:509. Tanypus puncti,:;e:mi:2 : Sublette 19648.: 132--133 (description of Edwards i m2terial)~ Tanypus carinatus SubleLte 1964·a:HO--1l2 (description of a.dults)" ?\12,",," Synonym~ Tanypus new species j Sublette 1.9648:110,157, 159, 161 (distribijtion, ecology and phenology). Tanypus ~~.~IHl_is; Si:blette an.d Sublette 1965 (Part):lSO (generic placement). Tanypus punctipennis: Roback 1966a:114-116 (designation of Lectotype and description). Tanypus cari~!.d;:U:~: Grodhaus J967:3 (immatures). Tanypus ~~!i~a~~~ var.: Roback 1969b:41S (immatures). "

Usinger and Kellen (1955 , Paine and Gaufin (1956), Fagan and Enns (1966) and Kimerle and Enns ( 968) are additional references on larval biology listed by Beck \ 977 p~ 23:)~ 47

Tribe Procladiini

Genus Procladius Skuse 1889

Subgenus Psilotanypus Kieffer 1906

"Procladius: Johannsen 1905(Part):125, 126. Psilotanypus: Kieffer 1906a:318. Psilotanypus: Kieffer 1906b:39. nee. Psilotanypus: Johannsen 1908:271. Psilotanypus: Coquillett 1910a:597. nee. Psilotanypus: Coquillett 19l0b:375, 378. nee. Psilotanypus: Malloch 19l5a:395. Procladius: Malloch 19l5a(Part):390. Psilotanypus: Edwards 1929:301. Psilotanypus: Goetghebuer 1936:4. Procladius: Johannsen 1937(Part):20. Psilotanypus: Brundin 1947:8. Psilotan.Y£us: Cae 1950:133. Procladius: Johannsen 1952(Part):lO. Psilotanypus: Freeman 1955:56,61."

Procladius (Psilotanypus) bellus (Loew) 1866; Var. 1 Roback 1980

(Plates 37--39; Figs. 202-215)

One specimen or Procladius (?~.) bellus was associated from Otsego Lake in this study. A larva collected in a grab sample from the shallow water sediments in Rat Cove (Map 1) on 7 July 1980, was reared to an adult male. The immature stages were first described by Johannsen (1905, p. 132 as ~. adumbratus), followed by Malloch (1915, p. 388 as Protenthes bel1us) and Johannsen (1937, p. 22 as P. adumbratus and p. 23 as P. bellus after Malloch, op. cit.).- Roback (1980, p. 31) provided the mo~ecent immature descriptions and keys.

Larval characteristics:

Moderately small Size.

Occipital margin dark.

Mentum with dorsomental teeth fused into distinct plates which are not joined medially (Figs. 202, 203 and 205).

Ligula with apical half or more black, with 5 black teeth and a concave apical margin (Figs. 202, 203 and 205). 48

Paraligula with apical tooth short, generally at most 2 times as long as accessory teeth (Fig. 203).

Pecten hypopharyngis with less than 10 short teeth.

Mandible with a long apical tooth and a relatively large and bluntly rounded basal tooth (Figs. 204 and 205).

Abdominal segments with a lateral hair fringe (as 1n Clinotanypus .£i.E!£~' Figs, 187 and 188).

Anterior parapod with serrated claws (as 1n P. (Holotanypus) johnsoni, Fig. 221).

Posterior parapod with one or more small and strongly hooked simple claws with an expanded base (Figs. 206 and 207).

Procercus about 3 times as long as wide with around 18 pale anal setae (Fig. 208).

Pupal characteristics:

Moderately small size.

Thoracic horn linear, narrow and short, about 4 times as long as wide, with a dark respiratory atrium and an apical plastron plate (Figs. 209 and 210).

Abdominal tergites generally clear.

Abdominal segments VII and VIII with 4 and 5 lateral setae, respectively, and lateral setae lacking on anterior segments (Fig. 213).

Anal lobe plate-like, slightly wider than long, with inner corners rounded and outer margin bearing strong spines and 2 lateral setae located in anterior half (Fig. 212).

Procl_adius species immatures are generally very similar morphologically with few distinguishing characteristics. Although Roback (980) provided keys to the immature stages, inunatures associated with adult males are still necessary to confirm identification in most species.

Larvae in the subgenus Psilot~"~_ may be distinguished from the subgenus Holotanypu~ (formerly Procladius sensu stricto but amended by Roback 1982 and Fittkau and Roback 1983) by their smaller size, generally smaller apical tooth of the paraligula (Fig. 203), the reduced size and number of teeth of the hypopharyngeal pecten and by the one to three small, strongly curved claws with expanded bases of the posterior parapods (Figs. 206 and 207) (Moore and Moore 1978, Roback 1980 and Fittkau and Roback 1983L Psilotanypus pupae may also be distinguished by their smaller size, smaller and relatively narrower thoracic horns 49

(Figs. 209 and 210), the rounded inner corners of the anal lobe (Fig. 212) and the generally clear or unpatterned abdomen (Roback 1980).

f. bellus is the only North American PsilotanlEus species in which the immatures are knO\·Jr1.R.oD.9.ck (1980) separated this species into three varieties based on adult and immature characteristics. A principal character used for separating the larvae is whether one or more of the smallest posterior parapod claws have intermediate teeth on the inner margin. The smallest claws are simple in variety 1 (Figs. 206 and 207) while one and two claws have intermediate teeth in varieties 2 and 3, respectively. Variety 1 pupae are distinguished by the small size of the anal lobes which are less than 413 micrometers in length as compared to greater than 430 micrometers in varieties 2 and 3. Variety 2 pupae are distinguished by the relative narrowness of the plastron plate which is less than 0.87 of the thoracic horn width as compared to greater than 0.87 in variety 3.

Roback (1980) reported examining a variety 1 female reared from a larva from Warwick Creek in Sterling Forest, New York, a variety 2 male reared from a pupa, a pupa, and an adult male from a lake also in Sterling Forest, New York. 1ne writer has also reared variety 1 males and females from the Hudson River in New York State. Roback (1971) examined specimens widely distributed in ;'iorth America and provided the following synonymies for P. bel1us:

"Tanypu~ bellus Loew 1866:4 (adult). Tanypus pusillus Loew 1866:5 (adult description), : Johannsen 1905:128 (redescription of ---;dult) . Rrocladius pusillus: Johannsen 1905:128 (adults), Procladius adumbratus Johannsen 1905:132 (adult female and immature stages). New Synonym. Psilotanypus E~llu~ Kieffer 1906b:39 (adult placement). Proc1adiu~ Eusil1us: Kieffer 1906b:39 (adult placement). Procladius adumbratus: Kieffer 1906b:39 (adult placement). ~~ius ~~us: Johannsen 1908:270 (generic placement) . Procladius or~siiontanY~iJs bellus: Johannsen 1908: 270 (generic placement. Protenthes ?el)u~: Malloch 19158.:388 (immature stages and adultj. Protenthe~ riparius Malloch 1915a: (paratypes):389 (adults). Ne\v Synonym. Procladius flavidus Kieffer 1923a:297 (adult female). ----- Procladius ~ali'::~ro Garret 1925:10 (adults). New Synonym. Protenthe~ _~::.£~E!"::-\_:::': Frison 1927: 176 (listing of types). Procladius pusillus: Johannsen 1937:22 (generic placement). Procladius ripariUS: Johannsen 1937:22 (generic placement). ProCladw-5 rna 1i fero: Johannsen 1937: 22 (gener ic placement). Proc lad ius flavi_~1!~: Johannsen 1937: 22 (gener ic placement). Procladius hellus: Johannsen 1937:23 (immatures, after Ma 110c 1-1-)~- 50

Procladius (Psilotanyp~~) adumhratus Johannsen 1937:22 (immatures) . Procladius bellus: Johannsen 1952:113 (adult in key). Proc1adius ~dumb~atus Johannsen 1952:102 (adults in key). ProcladiL~~ 1~.~.li)lus: Johannsen 1952: 10 (adult in key). Procladius riparius: Johannsen 1952: 11 (adult in key). Procladius adumbratus?: Wurtz and Roback 1955:199 (distribution and ecology). Procladius be11us: Roback 1957b:48 (immatures in key). Procladiu; ad~mbratus: Roback 1957b:49 (adult female). Procladius beJ.lus:-·B~ck and Beck 1959:91 (distribution). Procladius pusilitis: Beck and Beck 1959:91 (distribution). Procladius bellus: Sublette 1964a:126 (adult description). Procladius (Ps{l;!~~pus) bellus: Sublette 1964b:lll (adult and distribution). Proc1adius (Ps.) adumbratus: Sublette and Sublette 1965:14g-(ge~eric placement). Procladius (Ps.) belhls: Sublette and Sublette 1965:149 (generi~plac2we~t). Proc1adius (Ps.) malifero: Sublette and Sublette 1965:149 (generic placement). Procladius (~.) riparius: (Part) Sublette and Sublette 1965:149 (generic placement).

Procladius bellus: Roback 1969a:16 (food of larva). II

I 'Procladius Skuse 1889:283 (2 included species, type not designated). Tanypus: Johannsen 1905(Part):155. nee Trichotanypus Kieffer 1906a:319. Procladius: Coquiilett 1910a:594 (designation of P. paluJicola as type). nee Procladius: Malloch 1915a:390. Prothenthes: Malloch 1915a(Part):381. Trichotan~~~ Kieffer 1918b:62. Procladius: Edwards 1929:300. Procladius: Goetghebuer 1936:8. Procladius: Johannsen 1937(Part):20. Procladius: Cee 1950:133. Procladius: Johannsen 1952(Part):lO. Proc ladius: Freeman ) 955: 56. Ii Ho1otanypus: Roback 1982:98. 51

Proc1adius (Holotany~) johnsoni Roback 1980

(Plates 40-43; Figs. 216-236)

This species was extremely abundant in and restricted to the profundal zone of Otsego Lake. Nu,nerous larvae and pupae were associated and reared to adult males and femaies from late April through mid-June in 1981 and 1982. Most specimens were collected just east of Rat Cove in about 20 m of water (Map 1). Procladius (H.) 1ohnsoni was also a dominant food item (especially as pupae and pr~~pal larvae) of many bottom feeding whitefish collected in 20-30 m of water off of Kingfisher Tower and Peggs Point (Map 1) between early May and mid-June in 1981.

Roback 0980, p. 43) first described the immature stages of ~. johnsoni from a single larva reared to an adult male from the profunda1 of Muskoka Lake, Ontario, Canada. The present study apparent ly represents the first state and national record of occurrence for P. johnsoni.

Larval characteristics:

Moderately large Slze.

Occipital margin dark (Fig. 229).

Mentum with dorsomental teeth fused into distinct plates which are not joined medially (Figo 216, 220, 222 and 223).

Ligula with apical halE or more black, with 5 black teeth and a concave apical ITk'H:gin (Fig, 216, 220, 222 and 223).

Paraligula with apical tooth long, generally at least 3 times as long as accessory teeth (Fig. 223).

Pecten hypopharyngis with more than 10 teeth, generally around 14 teeth, most of ''''hich ar.e of normal length and larger than in the subgenus £silot:anY.£us.

Mandible with a long apical tooth and a relatively large and bluntly rounded basal tooth (Fig. 217 and 218).

Abdominal segments with a lateral hair fringe (as in Clinotanypus pinguis, Figs. 187 and 188).

Anterior para pod with serrated claws (Fig. 221).

Posterior parapod with all claws forming a flat curve, slender and simple (Fig. 226) (some claws may have very fine serrations on inner margin). 52

Procercus about 3 times as long as wide with around 18 pale anal setae (Fig. 225).

Pupal characteristics:

Moderately large size.

Thoracic horn large and globose with a dark respiratory atrium, a small apical nipple and no clear plastron plate (Figs. 230-233) .

Abdominal tergites generally patterned.

Abdominal segments VII and VIII with 4 and 5 lateral setae, respectively, and lateral setae lacking on anterior segments (Fig. 235).

Anal lobe plate-like, slightly wider than long, with inner corners generally produced as one or more spines and outer margin bearing strong spines and 2 lateral setae located in anterior half (Fig. 236).

Procladius species inm1atures are generally very similar morphologically with few distinguishing characteristics. Although Roback (1980) provided keys to the iw~ature stages, immatures associated with adult males are still necessary to confirm identification in most species. Characteristics for distinguishing the immature stages of the subgenera Holo~~ from Psilotanypus are given under ~. bellus. Roback (1982) erected the subgenus Holotany£us to include the Holarctic species formerly placed in the subgenus Procladius (Fittkau and Roback 1933).

Roback (1980) found the larva of E' johnsoni not readily separable from the larger sized Procladius (H.) species known. In the key to larvae (op. cit., p. 29), R. jOhnsoni specimens are separated from P. sublettei group species (which includes P. denticulatus) a;d other similar larvae by their larger rel;tive size based on the lengths of the mandible (greAter than 200 micrometers) (Figs. 216-218, 22, 227 and 228), antennal segment 1 (greater than 190 micrometers) (Figs. 216, 222, 227 and 228) and the ligula (greater than 140 micrometers) (Figs. 220 and 223). In the final couplet, P. johnsoni is distinguished from ~. vesus (thus far reported only from Alaska and the Northwest Territories of Canada in Roback 1971 and 1980, respectively) by the shorter apical paraligular tooth of the one specimen examined (about 12 micrometers versus about 18 micrometers long, respectively) (Fig. 223). As discussed under ~. denticulatus, reared profundal specimens of P. denticulatus are larger than littoral specimens in this taxon-and~;~indistinguishablefrom~. johnsoni based on size. Although littoral specimens of P. denticulatus key properly, profundal larvae key to R. johnsoni o~ vesus in Roback's (1980) key and are thus inseparable from R. johnsoni in Otsego Lake and probably other northern localities as well. 53

The large, globose thoracic horn with small apical nipple and no clear plastron plat~ (Figs. 230-233) will distinguish the pupa from all known Procladius species. In other characters, the pupa is very similar to P. denticulatus and other sublettei group species (Roback 1980) .------

Procladius (HolotanY2u~) d~~ticul~tus Sublette 1964

(Plates 44-48; Figs. 237-270)

Procladius (R.) denticulatus was one of the most abundant and ------widely distributed Tanypodinae species in Otsego Lake. Numerous larval-adult associated rearings were obtained from littoral grab samples at the north end of the lake near Cripple and Hayden Creeks, in Hyde Bay and at the south end in Rat Cove (Hap 1) throughout the sunnners in 1980 through 1982. In addition, larval and pupal rearings from profundal samples east of Rat Cove were quite common. Discussed below and photographed, these profundal reared specimens were considerabley larger than littoral rearings examined.

Roback (1957, p. 49) presented a b::-ief description of the immatures under ~. riparius and Roback (1930, p. SO) more extensively illus­ trated and described the immature stages or this primarily northern species.

Larval characteristics:

Medium to moderately large SIze.

Occipital margin dark (Figs. 238, 241 and 245).

Mentum with dOJ:somental tepth fused into distinct plates which are not joined medial (Figs. 246, 247 and 249-251).

Ligula with apical half or more black, with 5 black teeth and a concave apical margin (Figs. 246, 247 and 249-251).

Paraligula with apical tooth long, generally at least 3 times as long as accessory teeth (Figs. 247 and 250).

Pecten hypopharyngis with more than 10 teeth, generally around 14 teeth, most of which are of normal length and larger than in the subgenus Psilotanypus.

Mandible with a long apical tooth and a relatively large and bluntly rounded basal tooth (Fig. 248).

Abdominal segments with a lateral hair fringe (as in Clinotanypus plnguls, figs 187 and 188). 54

Anterior parapod with serrated claws (as 1n P. (H.) johnsoni, Fig. 221).

Posterior parapod with all claws forming a flat curve, slender and simple (Figs. 243 and 244) (some claws may have very fine serrations on inner margin).

Procercus about 3 times as long as wide with around 18 pale anal setae (Figs. 240 and 242).

Pupal characteristics:

Medium to moderately large Slze.

Thoracic horn linear to club-shaped, wide and long, about 3 times as long as wide, with a dark respiratory atrium and an apical plastron plate (Figs. 253-257).

Abdominal tergites generally patterned (Fig. 258).

Abdominal segments VII and VIII with 4 and 5 lateral setae, respectively, and lateral setae lacking an anterior segments (Fig. 261).

Anal lobe plate-like, slightly wider than long, with inner corners generally produced as one or more spines and outer margin bearing strong spines and 2 lateral setae located in anterior half (Figs. 262-270).

Procladius species immatures are generally very similar morphologically with few distinguishing characteristics. Although Roback (1980) provided keys to the immRture stages, inunatures associated with adult males are still necessary to confirm identification in most species. Characteristics for distinguishing the immature stages of the subgenera Holotanypus from Psilotanypus are given under ~. bellus. Roback (1982) erected the subgenus Holotanypus to include the Holarctic species formerly placed in the sugbenus Procladius (Fittkau and Roback 1983).

After studying numerous rearings, it became apparent that specimens of f. denticulatus reared from the profundal zone of Otsego Lake were consistently larger than those from littoral sediments. Moreover, they were large enough for the larvae to key to !'. iohnsoni or vesus (see discussion under ~. johnsoni) in Roback's (1980, p. 29) revision, while littoral specimens keyed properly to 5 inseparable species. For the most part, profundal pupae keyed properly to i. denticulatus and sublettei which are as yet inseparable, although some individuals exceeded the anal lobe maximum length given for these spec1es.

TIle effort was made 1n this presentation to demonstrate the apparent habitat related size differences by the specimens and photographs selected. Specimens indexed as M-l and M-2 were reared from the profundal zone east of Rat Cove (Map 1), while specimens M-3, M-4, M-5 55

and F-l were reared from several locations in the littoral zone. Exam­ ination of the slide material corroborate the photographic enlargements.

Roback (1980) reported an adult male collected at Sterling Forest in New York State and in 1971 and 1980 examined a small number of specimens mainly from across northern North America,

Roback (1980) provided the following synonymies for Procladius denticulatus:

I 'Procladius denticulatus Sublette 1964: J.23-126" (descrip­ tion of male type).

I 'Procladius riparius (Part):Roback 1957:49(Sta. 22B)" (brief description of immatures).

and also:

Procladius (Procladius) denticulatus Roback 1971:189, 190 (adult male description and distribution).

Tribe Pentaneurini

Genus Labrundinia Fittkau 1962

l'Ablabesmyia: Johannsen 1905 (Part):152. Tanypus: Malloch 19L5a (Part):372. Tanypus: Walley 1928 (Part):583). Pentaneura: Edwards 1929 (Group F, Part):294. Pentaneura: Johannsen 1946 (Group E, Part):283. Labrundinia Fittkau 1962: 372. iabrundinia: Beck and Beck 1966: 337. ' ,

Labrundinia filosella (Loew) 1866

(p la tes 49-51; Figs. 271-290)

Only one specimen of this small Pentaneurini genus was collected and associated in this study. A prepupal larva was obtained from a grab sample collected from 15 feet of water in Rat Cove (Map 1) on 20 August 1981 and reared to an adult female. TI-iOugh the females in this genus are poorly known, several adult males collected in July or August in 1981 by Simmons (983) conformed to the descriptions of Labrundinia pilosella in Roback (1962 and 1971). However, Roback (personal communication) stated the need for revision in the poorly known genus. Apparently their 5~

small size has hampered collection and thus availability for study. Roback (1962) provided detailed descriptions of the immature stages.

Larval characteristics:

Small size.

Occipital margin and head capsule pale (Figs. 271 and 277).

Antennal segment 2 heavily sclerotized and darker than basal segment (Figs. 271 and 275).

Mandible with a large quadrate basal tooth and prominant accessory tooth (Fig. 273).

Ligula with 5 dark teeth and with median tooth longer than lateral teeth (Fig. 273).

Head capsule with several lateral tubercles or spines just posterior to eye spots (Figs. 271 and 276).

Posterior parapod with one bifid claw with the outer tooth shorter and weaker than the inner tooth (Figs. 280 and 283).

Posterior parapod with basal seta modified as a compound spur (Figs. 278, 280 and 281).

Procercus about 4 times as long as wide with 7 dark anal setae (Fig. 280).

Pupal characteristics:

Small size.

Thoracic horn large and ovoid or reniform, with a dark, very finely reticulate respiratory atrium (Figs. 279, 287 and 288) and a gently curved, distally bulbous aeropyle entering a small subapical papilla which has a reduced pastron plate (Fig. 289).

Thoracic comb present (Fig. 279, 287, 288 and 290).

Thorax and abdomen clear and abdominal tergites with a very sparse dorsal shagreen of minute simple spinules (Figs. 279, 282 and 285).

Abdominal segments VII and VIII with 4 and 5 lateral setae, respectively, and lateral setae lacking on anterior segments (Figs. 282 and 285).

Anal lobe triangular and pointed, about 1.5 times as long as wide, with outer margin bearing weak spines and 2 lateral setae located centrally (Figs. 282, 285 and 286). 57

Roback (1971) tentatively synonymized Labrundinia floridana, described in all stages by Beck and Beck (1966, p. 339)~with 1. pilo~l~. Although adult males were not readily separable in Roback's (971) study, the L. florid<:n2 larva did not agree with earlier descriptions of ~.-piloseila~y Malloch (1915) and Roback (1962). Those larvae were not reared through to the adult ['rage, however, and could have represented incorrect associations (Roback 1971). The material presented here agrees quite closely with those earlier descriptions of 1::0 .EiJ.osellcL In la]~vae of the .!::' flori~na type, the occipital margin and posterior fourth of the hpad capsule is darkened, the head capsule texture is granular or nodulose and the posterior parapod spur is cather slender with several fine basal spines. Oliver et al. (1978, p. 64), Simpson and Bode (·~Q80, p. 22) and Oliver and Roussel (1983, p. 43) discussed and photographed the L. floridana type larva.

Roback (1962) reported the larval posterior parapods to be fused for about three-quarters of their length in~. pilosella, Figure 280 shows clearly separated posterior psrapods. They were, however, fused for three-quarters of their iength up'jn mounting. ~;Ihen pressure \¥as placed on the coverslip [0 flatten the head capsule and to properly display the mouthparts and sensory structures, the parapods were observed to separate and a narrow adheSive surface became visible as an off-hue, running three-quarters of the length of each parapod.

The characteristics given for the pupa will distinguish Labrundinia species from all non-Pentaneurini genera and from other Pentaneurini except the genus Ablcp~smyia. Small body size Rnd a long and narrow anal lobe with weak spines on the outer margin and lateral setae located centrally will distinguish the pupa from most Ablabesmyia species.

Roback (1971) reported adUlt males collected at Ithaca and Pine Grove, New York, a wide distribution in North America and provided the following synonymies for ~. E-ilosel!~:

W'Ta.nypus pi~9_se_;_l~~ Loe'Vl 1866:5:.6 (adult female)" Ablabesmyia ~los~l~~: Johannsen 1905:152, 153 (redescription of adult female). Tanypus Eilosellus: l'1alloch 1915a: 372 (adults and immatures). Tanypus Eilo~~ll~s: Walley 1928:581, 583 (adult in key, brief notes). Pentaneura E)losellc:: Bauber 1945:502 (phenology). Pentaneura pilosell~: Johannsen 1946: 283 (descri.ption of adult male and female). Pentaneura pilosella: Johannsen 1952:9 (distribution, brief

description of rnale) 0 Pentaneura pilosella: Beck and Beck 1959:91 (distribution). Pentaneura i~losell~: Roback 1962:253-258 (description of adult male and irnmatures). Pentaneura pilosella: Sublette 1964a:lOO (adult male description) . 58

Pentaneura (K.) pilosella: Sublette 1964b:112, 113 (brief description, notes, male). Pentaneura (K.) pilosella: Sublette and Sublette 1965:147 (generic placement). ?Labrundinia floridana Beck and Beck 1966: 339 (description of adults and immatures). New Synonym. nec. Labrundinia pilose11a: Beck and Beck 1966:340 (misdetermination of Eilosella Loew, adults and imma tures). I ,

Subgenus Ablabesmyia Johannsen 1905

"Tan~: Say 1823:15. Ablabesmyia: Johannsen 1905:135. Tanypus: Johannsen 1907b:400. Tanypus: Malloch 19l5a (Part):375-377. Ta~: Walley 1928 (Part):589-59l. Pentaneura: Edwards 1929 (Group A):288. Pentaneura: Johannsen 1946 (Group A, Part):269. Pentaneura: Freeman 1955 (Subgenus Ablabesmyia, Part):35. Pentaneura: Roback 1957 (monilis group):39. Pentaneura: Roback 1959 (Subgenus Ablabesmyia, monilis group): 113-115. Ablabesmyia: Fittkau 1962:417. Ablabesmyia: Beck and Beck 1966 (Part):38l. "

Ablabesmyia (Ablabesymia) annulata (Say) 1923

(Plates 52-55; Figs. 291-313)

One specimen of this very large sized Ablabesmyia species was associated in this study. This individual was collected in the littoral zone in 20 feet of water on the east shore of Otsego Lake across from Rat Cove on 19 July 1981 (Map 1). It reared to an adult female which keyed in Roback (1971) because of very distinctive leg band coloration. Boesel (1972) provided the original descriptions of the immature stages.

Many adult males and females of ~blabesmyia annulata were also collected in this study, although much fewer larvae were found in grab samples. Simmons (1983) reported this species to comprise 83.5 percent of 3,239 adult insects collected on and about the lake in July and August 1981. Thus, 2,705 individuals were A. annulata, suggesting a great 59

numerical and relative abundance should be found in the lake. Perhaps, larvae had the facility to avoid the grab sampler in the present study.

Larval characteristics:

Very large size,

Occipital margln and head capsule light brown (Fig. 293).

Maxillary palp with basal segment subdivided into 2 segments subequal in length (Figs, 296 and 299).

Head capsule with prominent antero-lateral notches at eye spots (Figs. 293 and 298).

Ligula with 5 black teeth with the m(~dian 3 teeth paler, even in height and rounded distally (Figs. 300-303).

Procercus long and narrow, about 4 to 5 times as long as wide with 7 pale anal setae (Fig, 295).

Posterior parapod lacking dark claws and with small claws covering approximately the distal half (Fig. 294).

Pupal characteristics:

Very large size.

Thoracic horn large and ovoid with a dark, very coarsly reticulate cespiratory atrium, an extremely reduced or absent subapicaL papilla and plastron plate and a short, dark, linear &eropyIe (Fig. 305). Tne aeropyle is not visible in figure 305 because of a mounting artifact but is clearly illustrated by Boesel (1972, Figs, 10-12).

Thorac ic comb present: (Fig, 308).

Abdominal tergites with a very heavy and uniform shagreen (Figs. 306, 307, 309 and 312).

Abdominal shagreen with numerous multirayed spinul~s (Figs. 311 and 313) which is a unique character among Ablabesmyia species, though present in other Pentaneurini genera (Roback, personal co~~unication).

Abdominal segments VII and VIII with 4 and 5 lateral setae, respectively, and lateral setae lacking on anterior segments (Fig. 307).

Anal lobe triangular and pointed, slightly longer than wide, with outer margin bearing weak spines and 2 lateral setae located approximately centr.ally (Fig. 309). 60

Roback (1971) reported adult males collected at Ithaca, Cranberry Lake and Otsego Lake in New York State. This species, though seldom collected, appears to be widely distributed in North America (Roback 1971 and Boesel 1972). The "lriter haS collected larvae of A. annulata in Schoharie Reservoil- on Schohacie Creek and in the Hoha~k River .-

Roback (1971) provided the following synonymies for this large and easily recognizable species:

I 'Tanypus annulatus Say 1823:15 (description of male). Tanypu~ annulatus: Johannsen 1905:144 (repetition of Say's original description, possible synonymy with monilis). Ablabesymia annulatus: Kieffer 1906b:41 (distribution). Tanypus monilis: Malloch 1915a:375 (misid~ntification of monilis (1.), description of adults and pupa). Tanypus illinoensis Malloch 1915a(Part):376 (Paratype male 24016, misdetermined). Tanypus rno~il~~: Walley 1928:272 (in key, misidentification o f mo nil i s (L.». Tanypus monilis: Walley 1928:588 (general description, distribution, misidentification of monilis (L. ». Pentaneura rnoni1is: Hauber 1945:496 (in -ke-y-,-­ misidentification of ----monilis (L.). Pentaneura annulata: Johannsen 1946:272 (adult description) ~~~ ~~ Johannsen 1946(Part):274 (specimen misdetermined as monilis). Pentaneura sunulata: Morrissey 1950:88 (notes, misidentification of annulata as monilis, brief pheno logy) , Pentaneura annulate: Johannsen 1952:6 (in key). Pentaneura annulata: Sublette 1957:381 (ecology and pheno l~gl) . Pentaneura (A.) annulata: Roback 1959:130 (description of adult male)-.------Pentaneura (A.) annulata: Dendy and Sublette 1959:508 (in key) . Pentaneura annulate:: Beck ~961:125 (distribution). Ablabesmlia~-;iata: Sublette and Sublette 1965:148 ( pIac erne n t ) " Ablabesmyia an~ulata: Beck and Beck 1966:320 (in key)."

Ablabesmyia (Ablabesmyia) basalis (Walley) 1925

(Plates 56-59; Figs. 314-339)

Ablabesmyia basalis was quite abundant and widely distributed in grab samples collected from littoral sediments in Otsego Lake during the summer. Larval-adult associations were obtained from early June through early August in 1979 through 1981, Though not a complete description, 61

this is the first account of characteristics of the immature stages of A. basalis. However, after examination of associated material from this study and a series of other adults and A. monilis (Linnaeus) associations, Roback (personal communication) indicated that A. basalis will be placed in synonymy with the Holarctic and cos;;opolitan type species for the genus, A. moniiis (Roback, in preparation). The immature stages of A, monil(;- have been described under the genus name of Pentaneura by Johannsen (1937, p. 12), Roback (1957, p. 39), Wurtz and Roback (1955, p. 199) and by others under the genera Pentaneura and Ablabesmyia.

Larval characteristics:

Medium size.

Occipital margin dark and dark ventral head spot present (Fig. 318) .

Maxillary palp with basal segment subdivided into 3 segments (Figs. 316, 322 and 323); the proximal segment is very short and often difficult to see (Fig. 319) and the distal segment is slightly longer and more slender than the middle segment.

Ligula with 5 black, pointed teeth and with the median tooth shorter than the lateral teeth (Figs. 317, 325 and 326).

Posterior parapod with 1 or 2 (sometimes 3) dark claws (Figs. 327 and 328).

Procercus about twice as long as wide with 7 pale anal setae (Figs. 321 and 324).

Pupal characteristics:

Medium size.

Thoracic horn large and ovoid with a dark, COArsely reticulate respiratory atrium (Figs. 329 to 331) and a gently curved aeropyle of uniform width entering a small subapical papilla which has a reduced plastron plate (Fig. 333).

Thoracic comb present (Figs. 329 and 334).

Wing sheath with major adult veins pigmented (Fig. 336).

Abdominal tergites with very sparse shagreen.

Abdominal segments VII and VIII with 4 and 5 lateral setae, respectively, and lateral setae lacking on anterior segments (Fig. 337). 62

Anal lobe triangular and pointed, slightly longer than wide and with 2 lateral setae located in the posterior half (Figs. 338 and 339).

Roback (1971) provided the following synonymies for A. basalis:

"Tanypus basalis Walley 1925:273 (description male adult). Tanypus basalis Walley 1928:591 (distribution and collection data). nee. Pentaneura basalis: Hauber 1945:496, 498 (in key, pupal description). Pentaneura basalis: Johannsen 1946:272 (brief description rna 1e ge nitalia ) . nee. Pentaneura basalis?: Roback 1957b:40 (misidenti­ fication of basalis Walley). nee. Pentaneura basalis?: Sublette 1957:380 (misidenti­ fication of~asalis Walley). Pentaneura basali~Beck and Beck 1959:91 (distribution). Pentaneura (A.) basalis: Roback 1959:127 (description adult m:;;le). Pentaneura (A.) basalis: Dendy and Sublette 1959:508 (in key) . Ablabesmyia basalis: Sublette and Sublette 1965:148 (placement). ' ,

Roback (1971) reported a northern North American d lstribution for Ablabesmyia basalis and examined adult males collected at Pine Grove and Watson in New York State. The writer has collected larvae of A. basalis in Schoharie Reservoir on Schoharie Creek and in the Hudso~ River in New York. However, as A. basalis will be synonymized with ~. monilis, Roback (op. cit.) reported adult males of ~, monilis collected at Millwood, Lake George and in two localities at Ithaca in New York State and found it to be one or the most widely distributed Ablabesmyia species in North America. Also, Simpson and Bode (1980, p. 21) discussed and photographed a species called A. parajanta which is very probably ~. basalis (= monilis),

Roback (1971) provided the following synonymies for A. monilis:

"Tipula monilis Linnaeus 1758:587 (adult description). : Johannsen 1905(Part):142 (brief description of adults and immatures), Ablabesmyia monilis: Kieffer 1906b:42 (distribution). Tanypus monilis: Johannsen 1907b:400 (proposal of monilis as type of Tanypus). Tanypus Monilis?: Malloch 19l5a:375 (description of adults and irmnaturesl. nec. Tanypus monilis: Walley 1925:272 (in key). nee. Tanypus monilis: Walley 1928:588 (distribution and general description). Tanypus monilis?: Johnson 1929:131 (distribution). Pentaneura monilis: Edwards 1929:289 (brief description adults). Pentaneura monilis: Johannsen 1937:12 (description of larva and pupa). 63

nee. ~entaneura moni)is: Hauber 1945:196 (in key, misdetermination of monilis Linnaeus). Pentaneura monili s: Johannsen1946 (Part): 27/+ (description ---- and~omparative notes on adult male). Pentaneura monilis: Johannsen 1952:6 (in key). Ablabesmyia monilis?: Judd 1953:813 (ecology and pheno logy). Pentaneura monilis?: Wurtz and Roback 1955:199 (immature stages and distribution), Pentaneura moni1is: Roback 1957b:39 (brief description of immature stages). Pentaneura (A.) monilis: Dendy and Sublette 1959:508 (in key) . Pentaneura (A.) moni1is: Roback 1959:114-117 (description of adult male and comparisons with European material). Ablabesmyia americana Fittkau 1972:430 (proposed new name for moni1is Joh. nee. Linnaeus). Ablabesrnyia monilis: Fittkau 1962:437 (description of adults and immatures). Pentaneura (A.) monilis: Sublette 1964a:86 (brief description and California distribution). Ab1abesmyia americana: Sublette and Sublette 1965: 148 (placement), Ablabesmyia monilis: Roback 1969a:17, 18 (food of larva)."

Ablabe~myia (Ab1abesmyia) mallochi (Walley) 1925

(Plates 60-63; Figs. 340-365)

This regionally common and widely distributed species was not particularly abundant in sediment grab samples in Otsego Lake. Only one specimen was associated in this scudy. It was collected on 19 July 1981 in three feet of water on the east shore of Otsego Lake across from Rat Cove (Map 1) and reared to an adult male. Roback (1957, p. 39 as Pentaneura auriensis) and Beck and Beck (1966, p. 325 as Ablabesmyia~~02hIand p. 328 as A. ornata?) described the immature stages.

Larval characteristics:

Medium size.

Occipital margIn dark and dark ventral head spot present (Figs. 340, 353 and 354).

Maxillary palp with basal segment subdivided into 5 segments (Figs. 343, 346, 348, 349 and 351); the proximal 4 segments are subequa1 in length and much shorter than the distal fifth segment. 64

Ligula with 5 black, pointed teeth and with the median tooth shorter than the lateral teeth (Figs. 340-343).

Posterior parapod with 2 or 3 (sometimes 1) dark claws (Fig. 345 and 352).

Procercus about twice as long as wide with 7 pale Anal setae (Fig. 344).

Pupal characteristics:

Medium size.

Thoracic horn large and ovoid with a dark, coarsely reticulate respiratory atrium (Figs. 355 and 356) and a gently curved, distally bulbous aeropyle entering a small subapical papilla which has a reduced plastron plate (Figs. 358 and 359).

Thoracic comb present (Figs. 356 and 357).

Abdomen and wing sheath distinctively pigmented (Figs. 364 and 365).

Abdominal tergites with very sparse shagreen.

Abdominal segments VIr and VIII with 4 and 5 lateral setae, respectively, and lateral setae lacking on anterior segments (Figs. 360 and 361).

Anal lobe triangular and pointed, slightly longer than wide and with 2 lateral setae located in the posterior half (Figs. 361 and 362).

Roback (1971) reported adult males collected at Albany, New York and in 1959 reported adult males collected at Mud Pond in McLeAn Res. in New York State. The writer has reared several specimens from the Upper Blenheim-Gilboa Pumped Storage Reservoir on Schoharie Creek and the Hudson River in New York and Simpson and Bode (1980, p. 20) reported the larvae of ~. mallochi commonly collected in sparse numbers over a wide range of physical and chemical conditions in rivers and streams of the State. Roback (1977) examined specimens from a wide distribution in North America and provided the following synonymies:

"Tanypus mallochi Walley 1925:273 (description of adult male). Tanypus mallochi Walley 1928:585, 590 (in key, relation­ ship). Tanypus illinoensis Malloch 19l5a(Part):376 (Paratype 13705). Pentaneura mallochi: Hauber 1945:497 (in key, brief pupal description). Pentaneura mallochi: Johannsen 1946:275 (adult male description). 65

Pentaneura illinoensls: Johannsen 1946(Part, var. 6):273, 274) , Pentaneura monilis: Johannsen 1946CPart):274. Pentaneura ;;aTY;~hi: Johannsen 1952:7 (in key). Pentaneura monilis (L.) var: Rohack 1957b:39 (pupa). Pentaneura -;'uriensis Roback 1957b:39 (description of male and immat~;;~s»)- New Synonym, Pentaneura (A.) mallochi: Roback 1959:122 (adult male description), Pentaneura (A.) auriensis Roback 1959:123 (brief adult de se ription-)-,--- Pentaneura (:2') ae~:L~ifasci3t"!: Dendy ar:d Sublette 1959: 507 (description of adult male an2 female, New Synonym). Pentaneura CA.) mallochi: Dendy and Sublette 1959:508 (in key). - PentaE-eura (~,) aequifasdata: Darby 1962: 116-121 (description adults, ecology), Pentaneura (A.) mallochi: Sublette 1964a:86 (description of adults), Ablabesmyia aequifasciat~: Sublette 1964b:113 (brief male description), Ablabesmyia mailochi: Sublette and Sublette 1965:148 (placement) . Ablabesmyia maliochi: Beck and Beck 1966:325 (description adults and immatures), AblabeE~~ ornata? Beek and Beck 1966:328 (description adults and immatures), New Synonym. Ablab5=smyia malloehi Roback 1969a: 17 (food of larva)."

Genus PentBn'2.Ura Phi 1ipp i 1865

"~ntane\lra Phil£ppi 1865:629. Tanypus: Malloch 19158 (Part):3?l. Tany~~: Walley 1928 (Part):584. nec. Pentaneura: ~2wards 1929:287. Penta;e~: 30hannser. 19L~6 (Group D Part):279) Pentane~~~: Johannsen 1952 (Part):5. nec. Pentaneura: Freeman 1955: 20. nee. Pe~taneura: Roback 1957:28. Pentaneura~Fittkau 1962:364. Pental~eura (Pentaneura): Sublette 1964a (Part):93, 98. P~~eura: Beck and Beck 1966:334." 66

Pentaneura inconspicua (Malloch) 1915

(Plates 64-66; Figs. 366-385)

Pentaneura inconspicua was not found in sediment grab samples. Several adult males and females were associated with larvae collected from Rat Cove (Map 1) in 0.1 to 0.5 m of water on sticks, branches, rocks and multiplate samplers in late June and early July, 1980 and early June in 1979. Beck and Beck (1966, p. 334) described the immature stages of P. inconspicua as P. inculta.

Larval characteristics:

Medium size.

Occipital margin dark (Fig. 366).

Ligula with 5 dark teeth even in height and with the first lateral teeth with inner margins curved outward (Figs. 367 and 368).

Mandible with large basal and accessory teeth (Figs. 367 and 372).

Posterior parapod with one dark claw (Fig. 369).

Procercus long and narrow, about 6 times as long as wide, with a distinctive sclerotized band and 7 dark anal setae (Figs. 370 and 371).

Supra-anal seta long, longer than anal setae and heavily sclerotized (Figs. 369-371).

Anal tubules almost twice as long as posterior parapods.

Pupal characteristics:

Medium size.

Thoracic horn large and club-shaped, with a prominant lateral plastron plate and surrounding clear area partly circumscribed by ostia (Figs. 377-379).

Thoracic comb present (Figs. 376, 378 and 379).

Abdominal shagreen uniform and sparse (Figs. 380-383) with spinules simple (Figs. 384 and 385).

Abdominal segments VII and VIII with 4 and 5 lateral setae, respectively, and lateral setae lacking on anterior segments (Fig. 382) . 67

Anal lobe triangular and pointed, about 1.5 times as long as wide and with 2 lateral setae located in the anterior half (Figs. 382 and 383).

This is apparently the first state collection record of this named species in Pentaneura although Simpson and Bode (1980, p. 25) discussed Pentaneura sp. which is very probably this species. Roback (1971) examined specimens with a wide distribution in North America and provided the following synonymies for _~' inconspicua:

"~~ inconspicuus Malloch 1915a:371 (adult). Tanypus inconspicuus: Frison 1917:177 (designation of Lectotype). Tan~ inconspicuus: Walley 1928:584 (in key). Pentaneura incons~icua: Johannsen 1946: 279 (in key). Pentaneura inconspicua: Johannsen 1952:5 (in key). Pentaneura (Pentaneura) comosa Sublette and Sublette 1965:93~%(adult)-, New Synonym. Pentaneura (~') inconspicua: Sublette and Sublette 1965:147 (generic placement). Pentaneura inculta Beck and Beck 1966:334, 335 (adults and immatures), New Synonym. I I

Genus Conchapelo~ Fittkau 1962

Subgenus Conchapelopia Fittkau 1962

ranypus: Walley 1925:276. "Tanypus: Walley 1928 (Part):583. Pentaneura: Edwards 1929 (Group D Part):292. Pentaneura: Johannsen 1946 (Group D Part):279. Thienemannimyia Fittkau 1957 (Subgenus Conchapelopia):317. Conchapelopia Fittkau 1962:223. Conchapelopia: Beck and Beck 1966:346."

Conchapelopia (Conchapelopia) currant (Walley) 1925

(Plates 67-70; Figs. 386-407)

Associations of this common species in Otsego Lake were obtained from littoral sediments and multiplate samplers in Rat Cove (Map 1) from late May through mid-July in 1980. Fourth instar larvae of Conchapelo~ currani were obtained during the same period in 1981. Roback (1981, p. 98) presented the first descriptions of the immature stages of C. currani and found it to be an uncommon species in that h8

study. Only one larval specimen was obtained and reared to an adult male in four years of collecting and rearing. Adults of C. currani were also uncommonly collected in previous studies (Roback 1971).

Larval characteristics:

Medium size.

Occipital margin pale (Fig. 386).

Mandible with basal tooth reduced (Fig. 394), often appearIng absent (Figs. 389 and 390).

Maxillary palpal apical appendage b, 3-segmented (Figs. 26 and 394).

Ligula with 5 dark teeth and a concave apical margin with the first lateral teeth with inner margins curved outward (Figs. 396 and 397).

Posterior parapod with several darkened claws (Figs. 393 and 395).

Procercus about twice as long as wide with 7 pale anal setae (Figs. 393 and 395).

Pupal characteristics:

Medium size.

Thoracic horn large and club-shaped, with a prominant lateral plastron plate and surrounding clear area partly circumscribed by ostia (Figs. 401-405).

Thoracic shagreen light to moderate and thoracic comb absent or weakly developed (Figs. 401 and 402).

Abdominal shagreen uniform and heavy (Figs. 399, 400 and 406) with numerous multirayed spinules (Fig. 407).

Abdominal segments III-VII with four lateral setae and segment VIII with 5 lateral setae (Fig. 399).

Anal lobe triangular and pointed, about as long as wide and with 2 lateral setae located in the posterior half (Fig. 400).

Roback (1971) reported adults of C. currani collected from Otsego Lake and Bemus Pt. in New York State and from several other localities in northern North America. The writer has also reared this species from water milfoil (Myriophyllum spicatum) in Saratoga Lake, New York. 69

Roback (1971) provided the following synonym1es for C. curran1:

'ITanypus curran1 Walley 1925:276 (description of male). Ta~ currani Walley 1928:585 (distribution). Tanypus currani Curran 1930:29 (distribution), Pentaneura currani: Johannsen 1946:279 (in key). ~~ currani: Johannsen 1952:8 (in key). nec. Pentan~~c;rrani: Roback 1957b:33 (immature stages). Pentaneura (P.) currani: Sublette and Sublette 1965:147

(generi~ placeme~). II

Genus Thienemannimyia Fittkau 1957

Subgenus Thienemannimyia Fittkau 1957

"Tanypus: Malloch 1915a (Part):369. Tanypus: Walley 1928 (Part):586. Pentaneura: Edwards 1929 (Group C Part):289, 290. Pentaneura: Johannsen 1946 (Group D Part):179, 280. Thienemannimyia: Fittkau 1957:315. Pentaneura: Roback 1957 (Part):38. Thienemannimyia Fittkau 1962:164. II

Thienemannimyia (Thienemannimyia) norena Roback 1957

(Plates 71-73; Figs. 408-422)

This common species was not found in sediment grab samples in Otsego Lake. Male and female associations of Thienemannimyia norena were obtained from larvae reared from multiplate samplers and from larvae found quite abundantly in the silted-over tread of a tire in Rat Cove (Map 1) in about 0.5 m of water during July, 1980. Roback (1957, p. 38) presented the original descriptions of the holotype male and immature stages and Roback (1981, p. 109) provided additional descriptions of the immature stages.

Larval characteristics:

Medium size.

Occipital margin dark (fig. 410).

Mandible with basal tooth reduced (as 1n Conchapelopia currani, fig. 394). 70

Maxillary palpal apical appendage b, 2-segmented (see fig. 26 for palpal appendage lettering).

Ligula with 5 dark teeth and a concave apical margin with the first lateral teeth with inner margins curved outward (figs. 411 and 413).

Posterior parapod with several darkened claws (figs. 409 and 412 ).

Procercus about 3 times as long as wide with 7 pale anal setae (figs. 409 and 412).

Pupal characteristics:

Medium size.

Thoracic horn large and club--shaped, with an ameboid respiratory atrium and lacking a plastron plate, surrounding clear area and ostia (Figs. 416 and 419).

Thoracic shagreen heavy and thoracic comb absent (Figs. 416 and 419) .

Abdominal shagreen uniform and heavy (Figs. 415, 417, 418 and 421) with mostly simple and bifid spinules (Fig. 422).

Abdominal segments I, VII and VIII with 5 lateral setae and abdominal segments II-VI with 6 lateral setae (Fig. 417).

Anal lobe triangular and pointed, about as long as wide and with 2 lateral setae located in the posterior half (Fig. 417, 418 and 420).

Thienemannimyia norena is a common, primarily eastern species according to Roback (1981). Although the pupae are quite distinctive, the larva of T. norena is presently inseparable from the other common ThienemannimyIa species in the east,]:. senata.

Roback (1971) reported adult males collected at Canadarago Lake, North Fairhaven and at Ithaca, New York. Roback (1981) reported one male at Cayuga Lake and another collection record of pupal skins in New York S ta t e .

Roback (1971) provided the following synonymies for T. norena:

I 'Pentaneura norena Roback 1957b:38 (adult wEle and imma t ures ) . Thienemannimyia noreana: (lapsus) Fittkau 1962: 174 (generic placement). Pentaneura carnea: Johannsen 1946 nee. Fabr. 1805 {Part):278 (misdetermination of carnes Fabr.). Pentaneura (P.) norena: Sublette and Sublette 1965:147 ---~neri~ placement). II 71

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Plate 1

Figure 1. Life cycle of a chironomid midge (unmodified from Jamnback 1969, fig. 34). Eggs

..-- Adult Larva

/ Pupa

1 93

Plate 2

Slide monnted rearing showing arrangement of parts

Figure 2. Ablabesmyia annulata female with associated immatures. )( -c 0 0 '­ Q) 0 .c. ..c. +- +­ +- (J) - C't ::J '"C :::J C '"C c ~·3~ « c

c -Q) Q) 00 0.-- +­ E :::J > ::J 0 a..~ '"C -0 .c Q) « c

- Q) / ~_2/ '- > o ::J -Ix Q) 95

Plate 3

Larval head capsules and cephalic structures

Figure 3. frocladius (Tanypodinae: Procladiini).

4. Ablabesmyi~ (Tanypodinse: Pentaneurini).

(Figs. 3 and 4 were slightly modified from Oliver and Roussel 1983. figs. 4 and S. respectively). ANTENNA------~

MANDIBLE ~ ~~r------~ttXpl&§ARY ~ I""A.-'-'~ LIGULA ~ PECTEN tJ: ,.o,;;;.~+-I---;",;~---- HYPOPHARYNGIS ~ PARALIGUU ~ ;5"7'-----"------DORSOHENTAL B TEF:I'H ~; ::;~ w"0:0 ~u

3

APICAL TOOTH

PECTEN :.:~---+-+l------L1 GU LA HYPOPHARYNGIS -----f-:---...'---" PARALIGUlA

4 97

Plate 4

Larval head capsules and cephalic structures

Figure 5. ~h!ronomus (Chironominae: Chironomini).

6. Mi~ropsectr~ (Chironominae: Tanytarsini).

(Figs. 5 and 6 were slightly modified from Oliver and Roussel 1983, figs. 6 and 7, respectively). APICAL TOOTH APICODORSAL TOOTH INNER TOOTH

!i~~l~rBoRN·s---.....,

....1lf---BLADE

y<----,------ANTENNA

/~"

\,--,,4,--c1Ac-;~.....,--PREMANDt8LE

STR IATION ---\------j~-:z:<4@F'7-f! '_"-;r;!---EYE SPOT

'+---~r_-- VENTROMENTAL PLATE

ventral view

5

"'(;7------BLADE

-+-+------~~b~~~f~6'NL (APICAL SPUR)

-I,------ANTENNAL TUBE RCLE (PEDESTAL)

MANDl BLE ---+--l,+­ ----"H---+---s I SETA INTERNA---\---¥'-;-T-4'1~ PECTEN EPI PHA R YNG IS----I-I:+i-r-""7.'""'''''''~ (n'lT-'T~~ITJIf----IPREMANDIBLE

EYE SPOTS---","",,(II'~II.

ventral view

6 99

Plate 5

Larval head capsules and cephalic structures

Figure 7. Cricotopu8 (Orthocladiinae).

8. ~rot~~ (Diamesinae: Protanypodini).

(Figs. 7 and 8 were slightly modified from Oliver and Roussel 1983, figs. 8 and 9, respectively). MAXI LLARY----'=-\-----'"' y,::J'..,....A"'~'A<~~~l_----PECTEN PALPUS EPIPHARYNGIS <:-~~~4.i,~:A-----PREMANDIBLE CHAETULAE----~A0'----"

yF----C\---VElITROHElITAL PUrE

I (

'V':;!----OCCIPITAL MARGIN

\'entral view

7

LABRAL LAMELLAE

/ '", ., ! '\ (' ",.:. "

\c\-----POSTERIOR PROJECTION

ventral view

8 101

Plate 6

Generalized larval Tanypodinae head capsule and cephalic structures

Figure 9. Head capsule.

10. Head capsule.

11. Maxillary palp.

12. Antenna.

13. Premento-hypopharyngeal complex lacking M appendage.

14. Mandible.

15. Mentum and M appendage of premento-hypopharyngeal complex (Macropelopiinil.

16. Mentum and M appendage of premento-hypopharyngeal complex (Pentaneurini).

17. Palatal surface of labrum.

(Figs. 9-16 were unmodified from Cranston and Reiss 1983. fig. 2.1 A-n, respectively; fig. 17 was slightly modified from Saether 1980, fig. 68 with lettering as in Roback 1974. 1976 and 1977 and probable homologies from Saether (op. cit.). Lauterborn organs ventral view blade antenna: b seto basal ""/: / ring acCos sory blade

maxillary palp--­ ring organ ring orgon 11 12

premento-hypopharyngeal "'--- submentum _hypopharynx ~"~~ /. ~ ~ .\ 13 sensillum minusc.ulum- ventrolateral setae dorsal tooth 2 /"'\'\;/ /" 3 antennal ratio (AR) \ 'i \ ,A..~\ ~~, ~ ;::k" '_, basal segment accessory - '\. ,(' ,t flagellum tooth 14 (.'0" ,.",~i apical toot h ­ eye ~ ~ A ~I~ v labial veSIcle pecten hypopharyngls dar somental tooth dorsomentu~ / ~_.c::-----,~---~ 15 IT'l(lntum \Jentromentum

', , dorsal view , ' ~, \ 16 10 , " chaeta

ventral view 17 103

Plate 7

Generalized larval Chironominae morphology

Figure 18. Head capsule.

19. Habitus of whole larva.

20. Labro-epipharyngeal region.

(Figs. 18-20 were slightly modified from Cranston and Reiss 1983. figs. 2.3 and 2.2 A and B, respectively). tooth ventral view

organ

lobrol sclente

clypeol seta

frontal opalame ------

dorsal view 18

triangulum occipitale --- foramen occlpitole

dorsal view -:~~ fronlal apotame _ / sehgment.,' C,"'" segments anterior parapads ,r"\ \"'" lateral viewI -, \IIJ \ //\,~ sclertle 3 abdominal I'~ segments i ) '\ Di~ bar .1 venlral tubules "

pl' \\ I ( as ena~rparapods : >"'"

anal tubules ~ / /", ~ / lateral tubules

procercus ~ premondibular brush anal setae 19 / , 20 105

Plate 8

Generalized larval cephalic structures showing measurements

Figure 21. Antenna (Tanypodinae).

22. Detail of antennal flagell urn (Tanypodinae) .

23. Antenna (Orthocladiinae).

24. Antenna of Tanytarsus (Chironominae: Tanytarsini> •

25. Antenna (Diamesinae: Diamesini> .

26. Lettering of apical maxillary palpal appendages in !~!en~~annimyia group species (Tanypodinae: Pentaneurini> •

27. Mandible of ~rilli~ (Orthocladiinae).

28. Mandible (Chironominae).

29. Mandible (Tanypodinae).

(Figs. 21-24. 28 and 29 were slightly modified from Saether 1980. figs. 81 - 86. respectively; fig. 25 was slightly modified from Oliver and Roussel 1983. fig. 10; fig. 26 was unmodified from Roback 1981. fig. 19; and fig. 27 was slightly modified from Cranston 1982. fig. 4). peg sensillum Lauterborn organ --t////11Jll.­ blade blade accessory blade _-II--A7 / peg sensillum nng. organ I accessory blade

segment I length

I', " ring organ " 22 I:" :'1/ I.: .. _.\ ~ 21 \ , 23

1 I . annulated Co-==- Lauterborn organ ~ ~. third segment .~ . ' \ 1\ peg sensllJum blade ~\ -ttf., a style ,/o~ 9 blade accessory bladel 25

... ~ngth of apical tooth antennal seta

combined width [ 26 of inner teeth \ r:; ring organ -} ~') ~1 ~~ 27 seta submenti pecten mandibularis seta subdentalis

seta subdentalis

seta interna seta submenti

ring organ 29

28 107

Plate 9

Larval cephalic structures showing measurements

Figure 30. Mandible of ~edilUI!I (Chironominae: Chironomini> •

31. Mentum and ventromental plates of RheocricotQP~~ (Orthocladiinae).

32. Maxilla of ~bl~lle~i!! (Tanypodinae: Pentaneurini> .

33. Maxilla of an Orthocladiinae.

34. Maxilla of Chironomus (Chironominae: Chi ronomini> .

(Fig. 30 was slightly modified from Maschwitz 1975, fig. 15; fig. 31 was slightly modified from Cranston 1982. fig. 6; figs. 32 and 34 were slightly modified from Saether 1980. figs. 76 and 75. respectively; and fig. 33 was unmodified from Cranston and Reiss 1983, fig. 2.2 C). /mentum " \" ~fl \~: 4' ventromental I( -/-. /.' beard width \~?~ fir m","m It~ seta submenti ventromental plate ventral view ventral view 31 30

paraxial seta ontaxiol seta multilobale sensillum ::'::::':~/~~~·;; I~/,_;oo::,::~ '''' d"'",',"~ lacinia "''";A~l maxillary palp ,/ L:J antaxial ""' /!, I t:J \seta 0«,""' ~,Ii! ~. A";~~'f 9( ".~ setae maxillaris, setae maxillaris 2 A" baslconlcum -~!blse~slllum ~ '_, seta maxllla"s 1+2 paraxial seta appendix seta I ca~,nat beard pecten galea"s choetulae of palpiger ventral view ventral view 32 33

lacinial chaeta

a seta paraxial seta sensilla basiconica bisensillum multilobate sensillum antaxial seta maxillary palp chaetulae of palpiger

...... :

dorsal view 34 109

Plate 10

Larval cephalic premento-hypopharyngeal complex

Figure 35. Hydrob~enus (Orthocladiinae).

36. Diamesini (Diamesinae).

37. f!~a~~~ (Diamesinae: Protanypodini).

(Figs. 35 and 37 were slightly modified from Saether 1980, figs. 80 and 79, respectively, and fig. 36 was unmodified from Cranston and Reiss 1983, fig. 2.2 D). M appendage 7"1J-~-t-- median lamellae labial palp paramedian lamellae ---\-----\~yr.;u paraligula chaetulae of prementum pecten hypopharyngis ~-TI-m:t--=::-l1

dorsal view ventral view 35

M appi?ndage

ventral view 36

labial palp pecten hypopharyngis \

ventral view 37 III

Plate 11

Larval cephalic and anal segment morphology showing measurements

Figure 38. Cephalic structures of Polypedilum (Chironominae: Cbironomini) .

39. Anal segment of (Orthocladiinae).

40. Procercus of Psectrocladius (Orthocladiinae).

(Fig. 38 was slightly modified from Maschwitz 1975, fig. 10 and figs. 39 and 40 were slightly modified from Cranston 1982, figs. 8 and 9, respectively). mentum ventromental plate a-gular length b-mentum width c-distance between ventromental plates d-ventromental plate width e-ventromental plate length

occipital margin

ventral view 38

"" "'" lenglh of anal seta lenglh of anallUbuly'""" ,>\ \ f "'" " ". '~"~ \'

procercus -----.J,-'.,-----'~~::3~

supra-anal seta -----:.....,-~,J. anal tubule ------=...--1.

posterior parapod --,-~-/-- claws of posterior parapod ----~l"!.

of posterior parapod lateral view 39

r------anal seta

lower lateral seta of procercus upper lateral seta of procercus procercus

height of procercus

spur or tubercle on base of procercus

lateral view 40 11J

Plate 12

Generalized pupal morphology

Figure 41. Diagrammatic dorsal view of a chironomid pupal exuviae showing major diagnostic features (slightly modified from Wilson and McGill 1982, fig. 1). cePhalic tubercle and seta \ I I M~ ~ennal sheath ~--;::.---.~ ~~~ ;~::;;~; ~::~ ~/~~ '.~., ':'. . .,"" respl.ratory atrium - • .,.;:,..<",. . . I plastron plate ~ !--thorax \ ~::: ::':Ch \~~ recurved hooks on II \ pedes spurii B

longitudinal spine patch pearl row spinules

IV band abdominal segments \ band \ \ \ \

41 115

Plate 13

Generalized pupal morphology and setation

Figure 42. Frontal apotome and ocular field.

43. Thorax.

44. Abdomen (Orthocladiinae).

45. Abdomen (Orthocladiinae).

(Figs. 42-45 were slightly modified from Saether 1980, figs. 52, 51, 53, and 54, respectively). thoracic horn

supraaJar median antepronotal(s)

if nose lateral antepronotal(s) pearls frontal apotome ~ cephalic tubercle lateral view frontal seta --+---''---l.J )'"vertical ocular field _ 43 IJI " P02 \ " Vv- postorbital I 42

tergites sternites

hooklets

conju nc ti ve IIJlIT -}.c======:c::======C'\ .~ pedes spurii B j \. ~ I ~

pedes spurii A muscle marks. ventral 2

o seta VI muscle marks. dorsal.; v.; v5-~""'­ V4

anal lobe

anal macrosetae ~'--~ dorsal view ventral view 44 45 117

Plate 14

Thoracic horn morphology and measurements

Figure 46. Tanypodinae, generalized.

47. ~rQ~la~i~~ (Tanypodinae: Procladiini).

48. ~rocla~i~~ (Tanypodinae: Procladiini).

49. Q~tbQQla4i~~ (Orthocladiinae).

50. ~~~~~ota~~~~~~! (Orthocladiinae).

51. ~la4Qta~tar!~~ (Chironominae: Tanytarsini).

52. RbeQta~ta~!us (Chironominae: Tanytarsini).

53. ~ryptQt~~4!pe~ (Chironominae: Chironomini).

54. ~Ql~e4il~ (Chironominae: Chironomini).

55. Ps~~do~b!~Q~omus (Chironominae: Pseudochironomini).

56. ~hirQ~Q~us (Chironominae: Chironomini).

57. ~~t~~~!a (Tanypodinae: ).

58. ~blah~smy!a (Tanypodinae: Pentaneurini).

59. Tanypus (Tanypodinae: Tanypodini).

(Fig. 46 was slightly modified from Saether 1980, fig. 55; figs. 47 and 48 were slightly modified from Roback 1980, fig. 39 A and n, respectively; fig. 49 was unmodified from Soponis 1977, fig. 85 a; figs. 50 and 55 were unmodified from Saether 1980, figs. 56-58, 61 60 and 59, respectively; figs, 56-59 were unmodified from Coffman and Ferrington 1984, figs. 25.618, 25.292, 25.321 and 25.324, respectively). plastron plate

horn sac aeropyle

d thoracic hom respiratory / atrium a - plastron plate width b - thoracic horn width felt chamber c- plastron plate length d - thoracic horn length

lateral view dorsal view lateral view 46 47 48

~ \

49 50 51 52

53 54 55

.... ; l",:.,.: - ..;. --~."' . :. ~, ~'~ _~ - ....'~ ,r: ~~ 57 58 59 119

Plate 15

Larva and pupa (Plates 15-18; Figs. 60-85)

Larva:

Figure 60.* Head capsule. F-l, 150X.

61. Head capsule, M-3, 1001.

62. Anterior of head capsule, M-3. 237.5X.

63. Procerci and posterior parapod cl aws. M-3, 237.5X.

64. Posterior parapod claws. M-3. 4851 (Oil).

*Abbreviations used for life stage and sex were larva (L), pupa (P), prepupa (PPL), male (M) and female (F). 60

61 121

Plate 16

~. albim~~~! continued

Larva continued:

Figure 65. Head capsule, F-1, 100 X.

66. Antenna, mandible and maxillary palp, F-1, 237.5X.

67. Mentum and ventromental plates, F-1, 237.5X.

68. Mentum, F-1, 485X (Oil).

69. Premandible, pecten epipharyngis and labral setae I, F-2, 237.5X.

70. Plumose labral setae I, F-2, 485X (Oil).

71. Antenna, F-1, 485X (Oil). 65 66

67 68

69

71

70 123

Plate 17

f. albimanus continued

Pupa:

Figure 72. Frontal apotome and ocular field, M-3. 100X.

73. Frontal apotome and ocular field, F-2, 100X.

74. Pedes spurii B on abdominal segment II, M-3, 237.5X.

75. Cephalic tubercles and frontal setae, F-2, 237.5X.

76. Pedes spurii B on abdominal segment II, F-2, 237.5X.

77. Pedes spur11 A ventral on abdominal segment IV. M-3. 237.5X. 72 73

74 75

76 77 125

Plate 18

~. albimanus continued

Pupa continued:

Figure 78. Abdominal segments II and III with dorsal shagreen, recurved hooks on posterior margin of II and intersegmental spinules between III and IV, F-2, 1001.

79. Abdominal segments VI-VIII and anal lobe, M-3, SOX.

80. Details of combs and lateral setae of abdominal segment VIII and anal lobe with hair fringe, M-3, 1001.

81. Recurved hooks on posterior margin of abdominal segment II, F-2, 237.51.

82. Recurved hooks on posterior margin of abdominal segment II, F-2, 485X (Oil).

83. Abdominal segments IV and V with dorsal shagreen and intersegmental spinules, M-3, 1001.

84. Details of combs and lateral setae of abdominal segment VIII and anal lobe with hair fringe, F-l, 1001.

85. Details of combs and lateral setae of abdominal segment VIII and anal lobe with hair fringe, F-2, 1001. 78 79 80

81 82

83 8 127

Plate 19

Larva and pupa (Plates 19-21; Figs. 86-104)

Larva:

Figure 86. Head capsule, L-1, 100X.

87. Head capsule, F-2, 100X.

88. Procerci and posterior parapods , L-1, 100X.

89. Labrum. L-1, 237.5X.

90. Antennae, L-1, 237.5X

91. Spurs at base of procerci, L-1, 237.5X.

92. Palmate labral setae I, L-1. 485X (Oil). J

87

86

88

89 90

91 92 129

Plate 20

~. !im~l~ns continued

Larva continued:

Figure 93. Mentum and ventromental plates. L-1. 237.51.

94. Palmate labral setae I and mandibular teeth. F-2. 4851 (Oil).

95. Mentum and beard of ventromental plates. L-l. 237.51.

Pupa:

Figure 96. Abdominal segment II with posterior rows of dorsal recurved spines and segment III with posterior rows of dorsal spines. F-2. 1001.

97. Thoracic horn. F-2. 1001.

98. Abdominal segments IV and V with posterior rows of dorsal spines and central spine patch on V. F-2. 1001.

99. Thoracic horn. F-l. 1001.

100. Anal lobe with hair fringe and macrosetae. F-2. 1001. 93 94

95 96

97 98

i

99 100 131

Plate 21

~. simulan~ continued

Pupa continued:

Figure 101. Abdominal segments II-V with posterior rows of dorsal spines and central spine patches on IV and V, F-2, 501.

102. Abdominal segments VII and VIII lateral setae and dorsal shagreen, F-2, 1001.

103. Abdominal segments VI-VIII with central spine patch and posterior spine rows on VI, lateral setae on VII and VIII and anal lobe. F-2, SOX.

104. Anal lobe with hair fringe and macrosetae, F-2, 100X. 101 102

103 04 133

Plate 22

Larva and pupa (Plates 22-24; Figs. 105-123)

Larva:

Figure 105. Head capsule and anterior parapods. M-1. SOX.

106. Antenna and mandible, ~~1, 237.5X.

107. Head capsule, M-1, 100X.

108. Mentum with warn median teeth and ventromental plates. M-1. 237.5X.

109. Procerci and posterior parapods. M-1. 100X.

110. Warn median teeth of mentum with notches indicating first laterals, M-1, 237.5X.

111. Pecten epipharyngis. plumose labral setae I and simple labral setae II. M-1, 485X (Oil).

112. Mentum and ventromental plates (unmodified from Saether 1976, fig. 24 G). 105 106

107 108

109 no

111 112 135

Plate 23

H. johannseni continued

Larva continued:

Figure 113. Antenna, M-1, 237.SX.

Pupa:

Figure 114. Frontal apotome and ocular fie ld , M-1, SOX.

115. Thoracic horn, M-'-1 , SOX

116. Thoracic horn, M-1, 100X.

117. Abdominal segments IV and V dorsal shagreen and setation, M-1, SOX.

118. Abdominal segment II dorsal shagreen and posterior rows of recurved spines, M-1, 100X.

119. Abdominal segment V dorsal shagreen and setation, M-1, 100X. 113 114

115 116 117

118 119 137

Plate 24

~. johannseni continued

Pupa continued:

Figure 120. Minute anterior dorsal spinules on abdominal segments II-VI and anal lobe, M-1, 485X (Oil).

121. Anal lobe with hair fringe, macrosetae and rugose base of macrosetae, M-l, 1001.

122. Pedes spurii A ventral on abdominal segments IV-VIII (VII is figured), M-l, 4851 (Oil).

123. Anal lobe with hair fringe, macrosetae and rugose base of macrosetae, M-l, 1001. 120 12 1

122

123 139

Plate 25

Protany~ ramosus Saether

Larva and pupa (Plates 25-29; Figs. 124-159)

Larva:

Figure 124. Head capsule, L-3, SOX.

125. Head capsule setae, L-3, 237.SX.

126. Anal tubules (two pairs), L-3, lOOX.

127 . Procerci and posterior parapod, L-3, SOX.

128. Procerci, L-3, lOOX.

129. Posteriorly directed ventral projections of occipital margin of head capsule, L-2, lOOX.

130. Lateral teeth of mentum, L-l, 237.5X. 124 125 126

127 128

129 130 141

Plate 26

P. ramosus continued

Larva continued:

Figure 131. Anterior of head capsule, L-3, lOOX

132. Labral lamellae and setae, L-3, 237.5X.

133. Premandible and epipharynx (upper) and medioventral appendix of prementum (lower), L-3, 237.5X.

134. Maxillary palp, L-3, 485X (Oil).

135. Premandible and epipharynx (upper) and medioventral appendix of prementum (lower), L-3, 485X (Oil).

136. Labral lamellae, L-3, 485X (Oil). 131 2

133 134

36

135 143

Plate 27

P. ramosus continued

Larva continued:

Figure 137 . Mandible and antenna, L-3, 237.5X.

138. Antenna, L-3, 485X (Oil).

139. Mandible and antenna, L-3, 237.5X.

140 . Anterior of head capsule, L-1, 100X.

141. Toothless middle portion of mentum, L-1, 237.5X.

142. Head capsule, L-1, SOX.

143. Toothless middle portion of mentum, L-1, 485X (Oil). 137 138 139

140 141

]43 145

Plate 28

f. ramosuS continued

Larva continued:

Figure 144. Posterior parapod claws (left), L-3, lOOX.

l4S. Posterior parapod claws (right) , L-3, 100X.

146. Detail of denticles on posterior parapod claws (left) , L-3, 237.5X.

147. Detail of denticles on posterior parapod claws (right), L-3, 237.5X.

Pupa:

Figure 148: Cephalic tubercles of pupal head 1n dorsal view, F-4, SOX.

149. Cephalic tubercles of exuviae, F-S, SOX.

150. Cephalic tubercles of pupal head in dorsal view, F-4, 100X.

151. Caudal lobes on venter of abdominal segment VIII and anal lobe, F-7, SOX. 144 145

146 147

148 149

150 151 147

Plate 29

f. ramosus continued

Pupa continued:

Figure 152. Thoracic horn, M-3, 100X.

153. Thoracic horn, M-l, lOOX.

154. Thoracic horn, F-4, 100X.

155. Caudal lobe ventral on abdominal segment VIII and anal lobe, M-l, SOX.

156. Caudal lobe ventral on abdominal segment VIII and anal lobe, M-6, SOX.

157. Thoracic horn spinules, F-4, 237.5X.

158. Caudal lobe ventral on abdominal segment VIII and anal lobe, M-2, SOX.

159. Thoracic horn spinules, F-5, 237.5X. 153 154

156 157

159 149

Plate 30

Potthastia longimana Kieffer

Larva and pupa (Plates 30-31j Figs. 160-173)

Larva, (7) third instar:

Figure 160. Head capsule, L-1, 100X.

161. Antenna, L-1, 237.5X.

162. Antenna with annulated third segment, L-1, 485X (Oil).

163. Anterior of head capsule, L-1, 237.5X.

164. Toothless mentum and bifid labral setae I, L-1, 485X (Oil).

165. Premandib1es, L-1, 485X (Oil).

166. Mandible, L-1, 485X (Oil). 161

160

162

163 164

165 166 151

Plate 31

P. lo~gimana continued

Larva continued:

Figure 167. Procerci and posterior parapods, L-l, IOOX.

Pupa:

Figure 168. Abdominal segment VII branched lateral setae and dorsal shagreen, M-I, IOOX.

169. Abdominal segments VII and VIII dorsal shagreen and maculation and dorsal and lateral setae, F-l, SOX.

170. Anal lobe with macrosetae, F-l, SOX.

171. Anal lobe with macrosetae, M-l, SOX.

172. Detail of anal lobe, F-I, IOOX.

173. Detail of anal lobe, M-I, lOOX. 167 168 169

/ ~. \

170 171

173 153

Plate 32

Clinotanypus (Clinotanypus) pinguis (Loew)

Larva and pupa (Plates 32-34; Figs. 174-192)

Larva:

Figure 174. Head capsule, L-1, 75X.

17S. Mandibles, L-l, 237.5X.

176. Ligula, paraligula and pecten hypopharyngis, M-2, 237.SX.

177. Anterior of head capsule, L-l, 150X.

178. Procerci, posterior parapods and anal tubules, L-l, SOX.

179. Procerci, L-1, 100X. 175

174 176

178

179 155

Plate 33

~. pinguis continued

Larva continued:

Figure 180. Head capsule, L-l, 50X.

181. Anterior of head capsule, L-l, 100X.

182. Maxillary palp, L-l, 237.5X.

183. Antenna, M-2, lOOX.

184. Antenna, M-2, 237.5X.

185. Antenna, L-l, 237.5X.

186. Posterior parapod claws, L-1, lOOX. I I I I I I I I I I I I I I I I • I 182 I \ 81 I 180 I I I I I I I I I

185 I 184 I 183 I I I

J I I I I

186 I I I I I I I 157

Plate 34

~. pinguis continued

Larva continued:

Figure 187. Lateral hair fringe of abdominal segments, L-l, SOX.

188. Lateral hair fringe of abdominal segments, LI-l, lOOX.

Pupa:

Figure 189. Thoracic horn and lateral thoracic tubercle ~n dorsal view, M-l,lOOX.

190. Abdominal segments VII and VIII and anal lobe lateral setae, M-l, SOX.

191. Dorsal mark on abdominal segment I, M-l, SOX.

192. Anal lobe with hair fringe, lateral setae and medial setiferous projections, M-I, SOx. 187 188 189

191

190

192 159

Plate 35

Tanypus (Tanypus) punctipennis Meigen

Larva (Plates 35-30; Figs. 193-201)

Larva:

Figure 193. Head capsule, L-1, 100X.

194. Dorsomentum and maxillary palps, L-l, 237.5X.

195. Mandibles, L-1, 237.5X.

196. Antenna, L-l, 237.5X.

197 . Procerci and posterior parapods, L-l, SOX.

198. Ligula and paraligula, L-1, 485X (Oil).

199. Anal tubules, L-1, lOOX. 194

193

195

196

199 197 Plate 36

T. punctipennia continued

Larva continued:

Figure 200. Procerci, L-1, lOOX.

201. Posterior parapod claws, L-1, lOOX. 201 163

Plate 37

Procladius (P8ilota~) bellus (Loew); var. 1 Roback

Larva and pupa (Plates 37-39; Figs. 202-215)

Larva:

Figure 202. Anterior of head capsule, M-l, lOOX.

203. Ligula, paraligulae and dorsomentum, M-l, 237.5X.

204. Antenna, mandible and maxillary palp, M-1, 237.5X.

205. Anterior of head capsule, M-l, 150X. 204

205 165

Plate 38

£. bellus continued

Larva continued:

Figure 206. Posterior parapod and claws, M-l, SOX.

207. Detail of posterior parapod claws, M-l, lOOX.

208. Procerci, M-i, lOOX.

Pupa:

Figure 209. Thoracic horn, M-l, lOOX.

210. Detail of plastron plate and thoracic horn, M-l, 237.SX.

211. Dorsal mark on abdominal segment I, M-l, lOOX.

212. Anal lobe, M-l, lOOX. 208

207

209

2lJ

210 167

Plate 39

K. ~ellus continued

Pupa continued:

Figure 213. Abdominal segments VII and VIII lateral setae and dorsal shagreen, M-l, lOOX.

214. Frontal apotome and ocular field, M-l, lOOX.

215. Anterior of thorax, M-l, SOX. /( 213 214

215 169

Plate 40

Procladius (Holotanypus) johnsoni Roback

Larva and pupa (Plates 40-43; Figs~ 216-236)

Larva:

Figure 216. Anterior of head capsule, M-l, 150X.

217. Mandible and maxillary palp, M-l, 237.5X.

218. Mandible and antenna, M-l 237w5X g

219. Autenn&-, 1'1-1, 485X (0:1.1).

220. Ligula, pEJ:'aligulae and dOrS0l112nturn, M-l, 237.5X.

221. Serrated anterior parnpod claws, PPL-2, 485X (Oil). 216

\ I I I I I 217 218 219

220 221 171

Plate 41

P. _iohnsoni continued

Larva continued:

Figure 222. Anterior of head :apSULe, PPL-l, lSOX.

223. Ligula and paraligulae, PPL-l, 237.5X.

224. Maxillary palps, PiL-l, 237.51.

225. Procerci, PPL-1. lOOX.

226. Posterior parapod c18~s, M-l, 237.5X. 225 226 173

Plate 42

~ ~ jQhnsoni continued

Larva continued:

Figure 227 Anterior of head capsule , PP1-1 , lOOK,

.~., ..."­ ~l-l 228 . Anterior ~.i .i head .:::apsu!.e , , lOOX"

229~ Head capsule, PPL-l 5 50X~

Pupa:

Figure 230. Thoracic horTI, iF, 100X.

232. Thoracic. horns .'l".j anterior parapod claws, PPl.-l, SOX.

233. Detail of apex of thoracic horn, FPL-I, 485X (Oil). 227 228

229 230 231

232 233 175

Plate 43

~i?'i.1pa continued:

,~ Q ~, Figure 234. 8. !:;. c::. C[(~:t T'i.E i IGOX~

235. Abdominal segm€~:8 VII and VIII ana anal lobe lateral setae and dorsal. shag~eenJ M-l~ SOX. 234

235 236 177

I,arva:

239~ Procerci and pOGteri~~r 237 238 239

l

240 241 242

243 244 79

249,

". :'~ .,.., , ­ .l)V.J 1..; J (=;t :ii 246

245

247

249

248

250 :: -\\ \.,. v- :"J .f"\. "

:;: ~.

258~ <:..,:.,~, 252

253 254 255

257 258 '-,:,. 259 260

261 262 .-'':'0 " ...: 263 264

265 266

267 268

269 270

272

273 271

276 275 274 189

Plate 50

Larva continued:

Figure 277. Head capsule, F-l, 100X.

278. Posterior parapod spurs, F-l, 237.SX.

280. Procerci and posterior parapods, F-l, lOOX.

281. Posterior parapod spurs, F-l, 485X (Oil).

283. Posterior parapod claws with detail of bifid claw, F-l, 48SX (Oil).

Pupa:

Figure 279. Thorax and abdominal segments I-III, F-l, SOX.

282. Abdominal segments IV-VIII and anal lobe, F-l, SOX.

284. Dorsal mark on abdominal segment I, F-l, lOOX. 277 278 279

280 281 282

283 284 191

Plate 51

L. pilosella continued

Pupa continued:

Figure 285. Abdominal segments VII and VIII and anal lobe lateral setae and shagreen, F-l, 100X.

286. Anal lobe, F-l, lOOX.

287. Thoracic horn and thoracic comb, F-l, lOOX.

288. Thoracic horn and thoracic comb, F-l, 237.5X.

289. Detail of aeropyle and apex of throacic horn, F-l, 485X (Oil).

290. Anterior of thorax, F-l, lOOX. 285 286 287

289

288

290 193

Plate 52

Larva and pupa (Plates 52-55; Figs. 291-313)

Larva:

Figure 291. Anterior of head capsule, L-1, 100X.

292. De ta i1 of antennal fl age11 um, L-1, 237.5X

293. Head capsule, L-l, SOX.

294. Posterior parapod and cl aw s, F-1, SOX.

295. Procerci, F-l, 100X.

296. Maxillary palp and mandibular tee th, L-l, 237.5X.

297. Posterior parapod claws, M-l, 100X. 291

293 4 295

296 195

Plate 53

A. annulata continued

Larva continued:

Figure 298. Anterior of head capsule showing lateral notches at eye spots, L-l, 75X.

299. Anterior of head capsule, L-l, 150X.

300. Premento-hypopharyngeal complex, F-l, l50X. 298

300 197

Plate 54

A. annulata continued

Larva continued:

Figure 301. Premento-hypopharyngeal complex, F-l, 100X.

302. Ligula, paraligulae and pectin hypopharyngis, F-l, 237.SX.

303. Detai of distally rounded middle ligular teeth, F-l, 485X (Oil).

Pupa:

Figure 304. Frontal apotome and ocular field, F-l, SOX.

305. Thoracic horn and thoracic comb, F-l, SOX.

306. Dorsal mark and shagreen on abdominal segment I, F-l, SOX. 301 302

303 304

305 306 199

Plate 55

A. annulata continued

Pupa continued:

Figure 307. Abdominal segments VII and VIII lateral setae and dorsal shagreen, F-1, SOX.

308. Thoracic comb, F-1, 100X.

309. Abdominal segment VIII and anal lobe, F-1, SOX.

310. Detail of anal lobe, F-1, 100X.

311. Multirayed spinules of dorsal shagreen on abdominal segment VII, F-1, 48SX (Oil).

312. Dorsal shagreen and muscle marks on abdominal segment VII, F-1, 100X.

313. Multirayed spinules of dorsal shagreen on posterior of abdominal segment III, F-I. 48SX (Oil). 307 308

309 310 311

312 313 201

Plate 56

Ablabesmyia (Ablabesmyia) basalis (Walley)

Larva and pupa (Plates 56-59; Figs. 314-339)

Larva:

Figure 314. Detail of antennal flagellum, L-l, 485X (Oil).

315. Anterior of head capsule, L-1, 100X.

316. Maxilary palp and mandibular teeth. L-l. 485X (Oil).

317. Anterior of head capsule. M-2. 100X.

318. Head capsule, M-l. 50X.

319. Maxillary palp. mandible and antenna, M-l, 237.5X.

320. Mandibular teeth and antennal flagellum. M-2. 485X (Oil).

321. Procerci, M-l. 100X. 314 315 316

317 318 319

320 321 203

Plate 57

A. basalis continued

Larva continued:

Figure 322. Maxillary palp, M-2, 485X (Oil) .

323. Maxillary palp, M-2, 485X (Oil) .

324. Procerci, M-l, lOOX.

325. Premento-hypopharyngeal complex, M-l, 237.5X.

326. Ligula, paraligulae and pecten hypopharyngis, M-l, 485X (Oil).

327. Posterior parapod and claws (note two dark claws), M-2, lOOX.

328. Posterior parapods and claws (note one dark claw on each parapod), M-l, lOOX. 322 323 324

325 326

127 328 205

Plate 58

A. basalis continued

Pupa:

Figure 329. Thoracic horn and thoracic comb, M-2, 100X.

330. Thoracic horn, M-l, 100X.

331. Thoracic horn, M-I, 237.5X.

332. Dorsal mark on abdominal segment I, M-I, 100X.

333. Detail of aeropyle and apex of thoracic horn, M-I, 485X (Oil).

334. Thoracic comb, M-2, 237.5X.

335. Frontal apotome and ocular field, M-2, 100X.

336. Maculation of wing sheath, M-2, SOX. 329 330 331

332 333

334 335 336 207

Plate 59

A. basalis continued

Pupa continued:

Figure 337. Abdominal segments VII and VIII lateral setae and dorsal shagreen, M-2, 100X.

338. Anal lobe, M-2, lOOX.

339. Anal lobe, M-l, 100X. 337 338 339 209

Plate 60

Ablabesmy!~ (Ablabesmyi~) ~allochi (Walley)

Larva and pupa (Plates 60-63; Figs. 340-365)

Larva:

Figure 340. Head capsule, F-l, 150X.

341. Premento-hypopharyngeal complex, M-l, 237.5X.

342. Ligula, paraligulae and pecten hypopharyngis, M-2, 485X (Oil).

343. Anterior of head capsule, M-2, 150X.

344. Procerci, M-l, 100X.

345. Procercus and posterior parapod (note two dark parapod claws), M-l, 100X. 341

340 342

344

143 345 211

Plate 61

A. mallochi continued

Larva continued:

Figure 346. Maxillary palp and mandibular teeth, M-2, 485X (Oil).

347. Anterior of head capsule, M-2, 100X.

348. Maxillary palp and mandibular teeth, M-2, 485X (Oil) .

349. Maxillary palp and mandibular teeth, M-l, 485X (Oil) .

350. Anterior of head capsule, M-l, 100X.

351. Maxillary palp and mandibular teeth, M-I, 485X (Oil).

352. Posterior parapod and claws (note three dark claws), M-2, 100X.

353. Head capsule, M-l, SOX.

354. Head capsule, F-l, 100X. \ "\ 346 347 348

\ .

i I / L' l...... _­ i 349 350 351

352 353 354 213

Plate 62

A. mallochi continued

Pupa:

Figure 355. Thoracic horn, M-1, 100X.

356. Thoracic horn and thoracic comb, M-2, lOOX.

357. Thoracic comb, M-2, 237.5X.

358. Detail of aeropyle and apex of thoracic horn, M-1, 237.5X.

359. Detail of aeropyle and apex of thoracic horn, M-1, 485X(Oil).

360. Abdominal segments VII and VIII lateral setae and dorsal shagreen, M-1, 100X.

361. Abdominal segments of VII and VIII and anal lobe lateral setae, M-1, SOX.

362. Anal lobe, M-l, 100X. 355 356 357

358 359

1!!lI5:isii+!%", 360 361 362 215

Plate 63

A. mallochi continued

Pupa continued:

Figure 363. Frontal apotome and ocular field, M-I, IOOX.

364. Abdominal segments I-III and wing sheath maculation, M-I, SOX.

365. Dorsal mark and maculation on abdominal segment I, M-I, IOOX. 363 364

365 217

Plate 64

Pentaneura inconspicua (Malloch)

Larva and pupa (Plates 64-66; Figs. 366-385)

Larva:

Figure 366. Head capsule, F-1, 150X.

367. Anterior of head capsule, F-1, 237.5X.

368. Ligula, F-1, 485X (Oil).

369. Posterior parapod and claws, F-l, lOOX.

370. Procerci and posterior parapods, F-l, SOX.

371. Procerci, F-1, lOOX. 367

366 368

369 370 371 219

Plate 65

P. inconspicua continued

Larva continued:

Figure 372. Anterior of head capsule, F-l, 237.5X.

373. Procerci and posterior parapods, M-l, SOX.

374. Head capsule, F-l, lOOX.

Pupa:

Figure 375. Frontal apotome and ocular field, M-2, lOOX.

376. Thoracic horn and thoracic comb, M-l, lOOX.

377. Thoracic horn, F-l, 237.5X.

378. Thoracic horn, K-2, 237.5X.

379. Thoracic horn and thoracic comb, M-l, 237.5X. 372 373

374 375 376

377 378 379 221

Plate 66

K. inconspicua continued

Pupa continued:

Figure 380. Dorsal mark and shagreen on abdominal segment I, F-l, 100X.

381. Dorsal mark and shagreen on abdominal segment I, M-1, 100X.

382. Abdominal segments VII and VIII and anal lobe lateral setae, M-2, 100X.

383. Anal lobe, M-2, 100X.

384. Abdominal segment VI dorsal shagreen, M-1, 237.5X.

385. Abdominal segment VII dorsal shagreen, M-1, 485X (Oil) • 380 381

382 383

384 385 223

Plate 67

Conchapelopia (Conchapelopia) currani (Walley)

Larva and pupa (Plates 67-70; Figs. 386-407)

Larva:

Figure 386. Head capsule, L-I, 75X.

387. Anterior of head capsule, L-I, 150X.

388. Anterior of head capsule, M-l, 100X.

389. Maxillary palp and mandible, M-1, 237.5X

390. Maxillary palp and mandible, M-1, 237.51. I I I I I I I I I 386 387 I I I I 388 I I I 390 .,• 389 225

Plate 68

f. currani continued

Larva continued:

Figure 391. Head capsule, L-l, SOX.

392. Anterior of head capsule, L-l, lOOX.

393. Procerc i and posterior parapods, M-l, SOX.

394. Maxillary pa Ips and mandibular teeth, L-l, 237.SX.

395. Procerci and posterior parapod claws, L-I, lOOX.

396. Ligula, L-l, 237.SX.

397 . Ligula, M-l, 237.SX. 391 392 393

394 395

396 397 227

Plate 69

c. currani continued

Pupa:

Figure 398. Frontal apotome and ocular field, M-I, 100X.

399. Abdominal segments VII and VIII lateral setae and dorsal shagreen, F-I, SOX.

400. Anal lobe, H-I, 100X.

401. Thoracic horn and thoracic shagreen, F-l, 100X.

402. Thoracic horn and thoracic shagreen, M-l, 100X.

403. Detail of plastron plate and apex of thoracic horn, F-l, 237.5X.

404. Ostia and plastron plate of thoracic horn, M-l, 237.5X.

405. Ostia of thoracic horn, M-I, 485X (Oil). 398 399 400

401 402

403 404 405 229

Plate 70

f. currani continued

Pupa continued:

Figure 406. Dorsal mark and shagreen on abdominal segment I, F-I, 100X.

407. Multirayed spinules of dorsal shagreen on abdominal segment VIII, F-I, 48SX (Oil). 406 407 231

Plate 71

Thienemannimyia (Thienemannimyia) norena (Roback)

Larva and pupa (Plates 71-73; Figs. 408-422)

Larva:

Figure 408. Anterior of head capsule, F-2, 100X.

409. Procerci and posterior parapods, F-2, SOX.

410. Head capsule, F-2, SOX.

411 • Ligula and pecten hypopharyngis, F-l, 237.SX.

412. Procerci and posterior parapod claws, F-2, 100X.

413. Ligula and paraligulae, F-l, 237.SX.

Pupa:

Figure 414. Frontal apotome and ocular field, M-l, 100X. 408

410

409

412

411

413 414 233

Plate 72

I. norena continued

Pupa continued:

Figure 415. Dorsal mark and shagreen on abdominal segment I (note adult setae within), F-3, 100X.

416. Thoracic horn and thoracic shagreen, M-l, 100X.

417. Abdominal segments VII and VIII and anal lobe lateral setae, M-l, SOX.

418. Anal lobe, M-1, 100X.

419. Detail of ameboid respiratory atrium of thoracic horn, M-l,237.SX.

420. Anal lobe, F-l, 100X.

421. Spinules of dorsal shagreen on abdominal segment VI, M-l, 237.SX. 415 416

417 418 419

420 421 235

Plate 73

I. norena continued

Pupa continued:

Figure 422. Mostly bifid spinules of dorsal shagreen on abdominal segment VI, M-l, 485X (Oil). 422