MARK F. O'BRIEN

Vol. 16, No. 2 Summer 1983 THE GREAT LAKES ENTOMOLOGIST

PUBLISHED BY THE MICHIGAN EN1"OMOLOGICAL SOCIErry THE GREAT LAKES ENTOMOLOGIST

Published by the Michigan Entomological Society

Volume 16 No.2

ISSN 0090-0222

TABLE OF CONTENTS

Seasonal Flight Patterns of in a Black Walnut Plantation. 7. . J. E. McPherson, B. C. Weber, and T. J. Henry ...... 35

Effects of Various Split Developmental Photophases and Constant Light During Each 24 Hour Period on Adult Morphology in Thyanta calceata (Hemiptera: Pentatomidae) J. E. McPherson, T. E. Vogt, and S. M. Paskewitz ...... 43

Buprestidae, Cerambycidae, and Scolytidae Associated with Successive Stages of Agrilus bilineatus (Coleoptera: Buprestidae) Infestation of Oaks in Wisconsin R. A. Haack, D. M. Benjamin, and K. D. Haack ...... 47

A Pyralid () as Pollinator of Blunt-leaf Orchid Edward G. Voss and Richard E. Riefner, Jr...... 57

Checklist of American (Arachnida: Araneae) Brent D. Ope II ...... 61

COVER ILLUSTRATION

Blister (Meloidae) feeding on Siberian pea-tree (Caragana arborescens). Photo­ graph by Louis F. Wilson, North Central Forest Experiment Station, USDA Forest Ser....ice. East Lansing, Michigan. THE MICHIGAN ENTOMOLOGICAL SOCIETY 1982-83 OFFICERS

President Ronald J. Priest President-Elect Gary A. Dunn Executive Secretary M. C. Nielsen Journal Editor D. C. L. Gosling Newsletter Editor Louis F. Wilson

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Copyright © 1983. The Michigan Entomological Society 1983 THE GREAT LAKES ENTOMOLOGIST 35

SEASONAL FLIGHT PATTERNS OF HEMIPTERA IN A NORTH CAROLINA BLACK WALNUT PLANTATION. 7. MIRIDAE J. E. McPherson, I B. C. Weber,2 and T. 1. Henry3

ABSTRACT

The seasonal flight patterns of 79 of Miridae collected in window traps in a North Carolina black walnut plantation are described. Flying height distributions and seasonal flight activities of Deraeocoris nebulosus (Uhler), Keltonia sulphurea (Reuter), Lygus lineo/aris (Palisot de Beauvois), Plagiognathus poUtus Uhler, and Reuteroscopus ornatus (Reuter) are considered in detail.

This is the last in a series of papers on seasonal flight patterns of Hemiptera in a black walnut (Jug/ans nigra L.) plantation near Asheville, North Carolina, and deals with the family Miridae; earlier papers dealt with the Pentatomoidea (1980), Coreoidea (l98Ia), Reduvioidea (l981b), Cimicoidea (l98Ic), Lygaeoidea (l98Id), and Tingidae and Aradidae (198Ie). The study was conducted from 24 March to 14 October in 1977, and from 24 March to 13 October in 1978. Specimens were collected weekly by window trapping; traps were suspended at 1, 2, 3, 4, 5, 6, and 7 m. The study site and trap construction are discussed in detail by McPherson and Weber (1980). All hemipteran specimens collected during this study are deposited in the Entomology Collection, Zoology Research Museum, Southern University, Carbondale.

RESULTS AND DISCUSSION

Seventy nine mirid species were collected during the two years of this study with the subfamily being best represented; numbers of specimens eollected for all taxa ranged from I to 2673 (Table I). Most of the species were collected in numbers too low to permit conclusions about seasonal flight patterns. However, Deraeocoris nebulosus (Uhler), Keltollia su/phurea (Reuter), Lygus lineolaris (Palisot de Beauvois), P/agiogllathus POlilUS Uhler, and Relitero­ scopus ornatus (Reuter) were collected in sufficient numbers (Table 1) to allow a more detailed discussion of flying height distributions and seasonal flight activities. D. Ilebulosus, a predaceous species, is generally associated with trees and shrubs (Wheeler et al. 1975). It has been reported to attack various species of mites and insects including, among others, European red mite, Panollychus ulmi (Koch); woolly apple aphid, Eriosoma lanigerum (Hausmann); clover aphid, Nearctaphis bakeri (Cowen); hop aphid, Phorodoll humuli (Schrank); cotton aphid, Aphis gossypii Glover; terrapin scale, Lecanium nigrofasciatum Pergande; eyespotted bud moth, Spiionota ocellana (Denis and Schiffer­ muller); and possibly codling moth, Laspeyresia pomollella (L.) (see literature survey of Wheeler et al. 1975). Wheeler et al. (1975), in their study of this , also reported its feeding on the mite Oliogonychus bicoior (Banks); the oak lace bug, Corythucha arcuata

I Department of Zoology, Southern Illinois University, Carbondale, IL 6290 L 2USDA Forest Service, North Central Forest EXperiment Station, Forestry Sciences Laboratory, Carbondale, IL 62901. 3Systematic Entomology Laboratory, lIBllI, Agricultuml Research Service. USDA, c/o National Mu­ seum of Natural History, Washington, DC 20560. w Table I. Seasonal flight activity of Miridae during 1977-78 in a North Carolina black walnut plantation. 0\

Collection Height (m) No. Range of Taxon Collected x ± SE Range Collection Dates MIRINAE Adelphocoris lineolatus (Goeze) 2 2.00 27 May Adelphocoris rapidus (Say) 11 1.82 ± 0.33 1-4 26 May-6 Oct. Agnocoris pulverulentus (Uhler) 2 4.50 ± 2.50 2-7 26 May-30 June ..., ater (L.) 4 4.25 ± 0.95 3-7 20 May-9 June ::r: elegans Heidemann 1 7.00 23 June tTl Dichrooscytus suspectus Reuter 1 4.00 2 June Cl Jusiformis (Say) 3 1.67 0.67 1-3 24 June g; Leptopterna dolabrata (L.) 5 3.20 ± 1.02 1-6 13 May-26 May ~ Lygocoris caryae (Knight) 11 3.82 0.52 2-7 13 May-7 July r Lygocoris geneseensis (Knight) 3 6.00 ± 1.00 4-7 13 May-17 June ;J;> iA Lygocoris sp. 1 4.00 27 May tTl Lygus lineolaris (Palisot de Beauvois) 929 2.60 ± 0.002 1-7 31 March-13 Oct. en recticornis (Geoffroy) 1 1.00 16 June tTl prob. americanus Carvalho 2 3.50 1.50 2-5 7 July ...,Z Phytocoris confluens Reuter I 5.00 30 June a Phytocoris conspurcatus Knight I 3.00 7 Oct. as:: Phytocoris ereclUs Van Duzee 2 4.50 ± 0.50 4-5 23 Jooe-5 Aug. ra Phytocoris Jenestratus Reuter I 7.00 20 May Cl Phytocoris intermedius Henry 3 5.33 ± 0.67 4-6 20 May-16 June en..., Phytocoris minutulus Knight 7 2.00 ± 0.53 1-5 30 J une-6 Oct. - Phytocoris penipectus Knight 1 6.00 13 May Phytocoris puella Reuter 2 5.50 0.50 5-6 23 June-30 June Phytocoris salicis Knight I 5.00 7 July Phytocoris tibialis Reuter 8 2.50 0.73 1-7 30 June-IS Sept. <: Polymerus basalis (Reuter) 21 1.90 0.33 1-7 1 April-13 Oct. P Prepops Jraterculus (Knight) 1 7.00 2 Sept. ~ Stenodema trispinosa Reuter 4 2.75 ± 1.44 1-7 1 April-6 May Z Stenotus binotatus (Fabricius) 6 4.00 ± 0.93 1-7 2 June-30 June ? Taylorilygus pallidulus (Blanchard) 36 2.11 :±: 0.21 1-6 30 June-7 Oct. N coelestiaUum (Kirkaldy) 37 4.59 ± 0.33 1-7 29 April-13 Oct. ::0 00 Trigonotylus doddi (Distant) 4 4.25 ± 1.03 2-7 30 June-6 Oct. v.> Tropidosteptes rufusculus (Knight) 2 6.00 ± 1.00 5-7 22 April-28 April

ORTHOTYLINAE Ceratocapsus setosus Reuter 2 2.50 0.50 2-3 29 Ju1y-5 Aug. Ceratocapsus sp. 1 6.00 15 July Diaphnocoris provancheri (Burque) 6 2.83 0.60 1-5 17 June-8 Sept. Halticus bractatus (Say) 24 2.75 ± 0.34 1-6 5 May-29 Sept. Halticus intermedius Uhler 3 3.33 :t 1.20 1-5 24 June-15 July >-l Heterocordylas maUnus Reuter 1 7.00 9 June ::r:: [/nacora stalii Reuter 82 2.09 :t 0.19 1-7 27 May-7 Oct. t'tI heidemanni Knight 33 2.06 0.33 1-7 19 May-18 Aug. Q ~ (Say) 3 2.67 ± 0.88 1-4 23 June-24 June t'tI Lopidea robiniae (Uhler) 1 7.00 7 July >-l>­ Orthotylus modestus Van IAlZee 7 5.14 0.59 2-7 20 May-9 June t'"' Orthotylus ornatas Van Duzee 3 5.00 ± 0.58 4-6 6 May-26 May >­ i": Orthotylus ramus Knight 5 4.80 ± 0.80 3-7 27 May-7 July t'tI Pilophorus crassipes Heidemann 4 6.00 ± 0.58 5-7 10 lune-6 Oct. (I:l Pseudoxenetus scutellatus (Uhler) 2 4.50 ± 2.50 2-7 13 May-27 May t'tI Z Sericophanes heidemanni Poppiu8 12 3.00 ± 0.51 1-6 5 May-13 Oct. >-l Slaterocoris atritibialis (Knight) 6 2.17 0.75 1-5 29 April-16 June 0a:: Slaterocoris stygicus (Say) 1 1.00 26 May 0 t'"' 0 Q Chlamydatus suavis (Reuter) 5 3.60 ± 1.25 1-7 12 May-6 Oct. til Criocoris sa liens (Reuter) 2 2.00 19 May-20 M"v >-l Icodema nigrolineatum (Knight) 2 3.50 ± 1.50 2-5 24 June-14 Keltonia sulphurea (Reuter) 194 1.32 ± 0.Q7 1-7 27 May-7 Oct. Lepidopsal/us c1aricomis Knight 14 3.36 0.27 2-5 6 May-3 June Lepidopsallus miniatus Knight 6 3.67 0.88 1-7 22 April-16 June Lepidopsal/us rostratus Knight 13 3.77 0.46 2-7 13 May-l July Microphylel/us modestus Reuter 2 5.00 1.00 4-6 27 May Plagiognathus albatus (Van Duzee) 1 1.00 15 July Plagiognathus cameolus Knight 4 2.50 ± 0.96 1-5 6 May-12 May v.> Plagiognathus dispar Knight 1 5.00 10 June -...j c...> Table 1. Continued. 00

Collection Height (m) No. Range of Taxon Collected x ± SE Range Collection Dates

Plagiognathus guttulosus (Reuter) 1.00 13 May Plagiognathus nigronitens Knight I 1.00 23 June Plagiognathus politus Uhler 506 2.51 ± 0.004 1-7 5 May-13 Oct. Psallus strobicola Knight 5 5.40 ± 0.81 3-7 27 May-23 June ,.., Pseudatomoscelis seriatus (Reuter) 23 1.91 ± 0.33 1-7 2 June-29 ::r:: Reuteroscopus ornatus (Reuter) 2,673 4.58 ± 0.001 1-7 27 May-13 Oct. m Rhinocapsus rubricans (Provancher) 2 4.50 ± 2.50 2-7 16 June-28 July Cl Spanagonicus albofasciatus (Reuter) 30 4.23 ± 0.39 1-7 29 July-13 Oct. m ";p.,.., r Deraeocoris nebulosus (Uhler) 612 2.86 ± 0.002 1-7 I ;p. Deraeocoris nigrituius Knight 3 5.33 ± 0.88 4-7 13 June ~m Eustictus grossus (Uhler) 2 5.00 22 Sept. (/) harti Knight 7 2.43 ± 0.69 1-5 30 June-l Sept. m Z (Say) 5 1.60 ± 0.60 1-4 24 June-29 Sept. ,.., 0 ~ DICYPHINAE 0 Dicyphus famelicus (Uhler) 1.00 15 April r 0 Dicyphus prob. rhododendri Dolling 7.00 3 June Cl til ...., Sixeonotus tenebrosus (Distant) 3 3.33 ± 0.88 2-5 3 June-7 Aug.

CYLAPINAE -< Fulvius imbecilis (Say) 2 4.00 ± 3.00 1-7 29 July...s Sept. 0 Fulvius slateri Wheeler 8 4.50 ± 0.33 3-6 24 June-25 Aug. , :a; 'L ? IV 1983 THE GREAT LAKES E!\TOMOLOGIST 39

(Say); hawthorn lace bug, C. cydon[ae (Fitch); and greenhouse whitefly, Tria/eurodes .'aporariorum (Westwood). In the Harrisburg area of , it overwinters as adults and is trivoltine (Wheeler et al. 1975). In the present study, D. nebu/oslis adults were found from early April to late September (Table I). They were collected at all seven flying heights with about 60% captured at 2-3 m (Fig. I). This species overwintered as adults but the number of generations per year is unclear from the data available (Fig. 6). If, as Wheeler et al. (1975) reported for Harrisburg, this species is also trivoltine near Asheville, then the overwintered adults emerged in early April and reproduced during the spring. Flight activity of their offspring began in early June, generally increased from mid-June through July, peaked in eady August, and declined sharply during the second half of August; the length of this flight activity period (i.e., during much of the summer) suggests overlapping generations although it is possible it represents the activity of long-lived individuals. K. sillphurea has been collected on Ambrosia sp., Chenopodium a/bum L, Symphor­ icarpos orbicu/atlls Moench (Knight 1941), and Sida spinosa L. (Knight 1927, 1941, 1966). No information has been published on its life cycle other than its reported "breeding" on S. spinosa in early September in Georgia (Knight 1927). In the present study, K. su/phurea adults were found from late May to early October Cfable I). They were collected at all seven flying heights with about 85% collected at I m (Fig. 2). This species probably overwintered as eggs, this based on the large number of flying adults in the fall and the extended period of time in the spring before any adults were collected (i.e., late May) (Fig. 7). Also, if by "breeding" in early September Knight (1927) meant he observed copulating pairs at that time, then fertilized femlaes would have had sufficient time to oviposit before the end of the season. If this species does overwinter as eggs, then the flight data suggest three generations; adults of the first generation were present from late May to about mid-July, those of the second from mid-July to late August, and those of the third from early September to the end of the season. L. linea/aris (tarnished plant bug) feeds on numerous plants including alfalfa, apple, apricot, aster, bean, beet, blackbeny, cabbage, carnation, carrot, celery, cherry, chrysan­ themum, clover, cotton, cucumber, currant, dahlia, grape, lettuce, marigold, pea, peach, pear, peony, plum, potato, quince, raspberry, rose, strawberry, tobacco, and turnip (Kelton 1975). However, it also feeds on certain including alfalfa plant bug, Ade/phocoris lin eo/allis (Goeze); potato leafhopper, Empoasca fabae (Harris); pea aphid, Acynhasiphon pisum (Harris); Colorado potato , Leptinotarsa decemJineata (Say); alfalfa weevil, Hypera postica (Gyllenhal); species of Noctuidae; P/euraprllcha insu/saria (Guem6e); alfalfa blotch leafminer, Agramyza frantella (Rondani); Aphidius ervi pulcher Baker; A, smithi Sharma and Subba Rao; Praan sp.; species of Formicidae; and the harvestman Pha/angillm opilio L. (Wheeler 1976). This species overwinters as adults (Guppy 1958, Kelton 1975, Ridgway and Gyrisco 1960, Stewart and Khoury 1976) and has two (Guppy 1958, Kelton 1975) or perhaps three (Ridg­ way and Gyrisco 1960, Stewart and Khoury 1976) generations per year. Ridgway and Gyrisco (1960) used tanglefoot traps to determine flying height patterns between 0 and 18 feet. They found that L. lineo/aris flew fairly close to the ground; of 323 adults collected, 300 were captured within 6 ft of the ground and only one as high as 15-18 feet. In the present study, L. lineo/aris adults were found from late March to mid-October (Table I). They were collected at all seven flying heights with about 60% captured at 1-2 m and about 3% at 7 m (Fig. 3); this roughly corresponded to the flying height pattern reported by Ridgway and Gyrisco (1960). This species overwintered as adults and is apparently bivoltine near Asheville (Fig. 8). Adults began to emerge from overwintering sites in late March. Their adult offspring (summer generation) occurred from about late May to mid-August. Adults of the second (overwintering) generation occurred from about late August or early September to the end of the season, It is possible that an additional generation occurred during the summer but this could not be deternrined from the data available. 40 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No.2

100,------,- 100,------r

.. 1 !! ~ 2 ~ 80 ~ 80 .;" .;'0 '6 60 .5 ~ 60 '0 40 ~ 40 C ..c: .. () ~ 20 0; 20 n. Q.

2 3 4 561 2 3 4 561 Meters Meters

100~------, 100.------~ .. 3 ., 4 ~ 80 ~ 80 II '0 > ;; ~ 60 ~ 60 '0 40 o 40 C.. C., () "i 20 a; 20 n. Q.

2 3 4 587 234 5 e 7 Meters Meters 100 5 .... 60 ::I ";; '6 60 .5 '0 40 C.. ~ 20 n.

2 3 456 7 Meters

Figs. 1-5 Flying height distributions offive mirid species during 1977-78 in a North Carolina black walnut plantation: (I) Deraeocoris nebulosus, (2) Keltonia sulphurea, (3) Lygus lineoluris. (4) Plagiognathus politus, (5) Reuteroscopus ornatus.

p, poUtus has been collected from several plants including Pyrus (Leonard 1915, Knight 1941), Ambrosia sp., Betula sp" Carya sp., Corylus sp., Juniperus virginiana L., Pinus strobus L., Quercus sp., Robinia pseudoacacia L., Salix: sp., Solidago sp., Symphoricarpos orbiculatus Moench, Taxodium distichum (L.) (Knight 1941), Erigeron, and Verbascum (Froeschner 1949). It overwinters as eggs (Leonard 1915) and apparently has two broods per year (is bivoltine?) (Froeschner 1949, Knight 1941). In , adults of the first genera­ tion have been collected between 10 June and 20 August and those of the second between g August and 31 September (Froeschner 1949). In the present study, P. paUlUS adults were found from early May to mid-October (Table I). They were collected at all seven flying heights with almost 50% captured at 1 m (Fig. 4). This species apparently overwintered as eggs, thus agreeing with the findings of Leonard (1915). Supporting this conclusion were the large number of flying adults in the fall and the 1983 THE GREAT LAKES ENTOMOLOGIST 41

25 25,------, .. 6 .. 7 § 20 § 20 ".;: ".;: '5 15 '0 15 .E .E '0 /1 "0 ~ 10 ~ 10 c: . \ c: II> '"(J / I (J ~ 5 l 5 0.. ) ~ M A M' J J A S 0 M A M J J A S 0

25 25,------, ., 8 9 '"iii 20 ::> ;;;" '5 15 .E "0 A c 10 1\ '"(J r\~ ;;; 5 0.. ;\)'\1\ J \ M A M J J A S 0 M A M J J A S 0

'" 10 ~ § 20 1\ ".;: Ii :;:; 15 .5 ~ 10 (\ c: I I (J ;;;'" 5 I ! 0.. I \ \ M A M J J A S'O

Figs. 6-10. Seasonal flight activities offive mirid species during 1977-78 in a r-iorth Carolina black walnut plantation: (6) Deraeocoris nebulosus, (7) Keltonia su/phurea, (8) Lygus linealaris, (9) Plagiognathus politus, (10) Reulerascopus omalus. extended period in the spring before the first adults were collected (i.e., early May) (Fig. 9). It is apparently bivoltine near Asheville; adults ofthe first (summer) generation were present from early May to late July, and those ofthe second (overwintering) generation from early or mid-August to the end of the season. R. ornatus has been collected from Ambrosia (Kelton 1964, Knight 1941) and Cheno­ podium album L. (Knight 1941), No information has been published on its life cycle. In the present study, R. ornatus adults were found from late May to mid-October (Table 1). They were collected at all seven flying heights with more than 55% captured at 5-7 m (Fig. 5). This species probably overwintered as eggs, this based on the same reasons as those given above for P. politus. The number of generations per year is unclear from the data available (Fig. 10). However, it is likely that at least one generation is represented by the few adult 42 THE GREAT LAKES ENTOMOLOGIST Vo!. 16, No.2 specimens collected between late May and mid-August, and a later generation by the dra­ matic increase in numbers (i.e., increase in flight activity) in September. Interestingly, of the 2673 adults collected during this study, almost 93% were captured in September and almost 50% during the third week of September (Fig. 10).

ACKNOWLEDGMENTS

We wish to thank Mr. D. Brenneman and the staff of the North Carolina Division of Forestry, Morganton, for their help in collecting data and maintaining the window traps. This research was partially supported by the USDA Forest Service, North Central Forest Experiment Station.

LITERATURE CITED

Froeschner, R. C. 1949. Contributions to a synopsis of the Hemiptera of Missouri, Pt. IV. Hebridae, Mesoveliidae, Cimicidae, Anthocoridae, Cryptostemmatidae, Isometopidae, Meridae (sic). Amer. Midland Natur. 42: 123-188. Guppy, J. C. 1958. Insect surveys ofclovers, alfalfa, and birdsfoot trefoil in eastern Ontario. Canadian Entomo!. 90:523-531. Kelton, L. A. 1964. Revision of the Reuteroscopus Kirkaldy 1905 with descriptions of eleven new species (Hemiptera:Miridae). Canadian Entomo\. 96:1421-1433. ----,. 1975. The lygus bugs (genus Lyglls Hahn) of North America (: Miridae). Mem. Entomo!. Soc. Canada 95: 1-101. Knight, H. H. 1927. Notes on the distribution and host plants of some North American Miridae (Hemiptera). Canadian Entomo!' 59:34-44. ----. 1941. The plant bugs, or Miridae, of Illinois. Illinois Natur. Hist. Surv. Bull. 22:1-234. ---. 1966. Keltonia, a new genus near Reuteroscopus Kirk., with descriptions of new species (Hemiptera:Miridae). Canadian Entomo!' 98:590-591. Leonard, M. D. 1915. The immature stages of Plagiognathus politus Uhler and Campyl­ omma verbasci Herrick-Schaeffer (sic) (Capsidae, Hemiptera). J. Entomo!. Soc. 23:193-197. McPherson, J. E. and B. C. Weber. 1980. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 1. Pentatomoidea. Great Lakes Entomo!' 13:177-183. ----. 1981a. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 2. Coreoidea. Great Lakes Entomo\. 14: 11-13. ----'. 1981b. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 3. Reduvioidea. Great Lakes Entomo!. 14: 15-17. 1981c. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 4. Cimicoidea. Great Lakes Entomo!. 14:19--22. 1981d. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 5. Lygaeoidea. Great Lakes Entomo!. 14: 133-136. ----. 1981e. Seasonal flight patterns of Hemiptera in a North Carolina black walnut plantation. 6. Tingidae and Aradidae. Great Lakes Entomol. 14: 137-140. Ridgway, R. L. and G. G. Gyrisco. 1960. Studies of the biology of the tarnished plant bug, Lygus lineolaris. J. Econ. Entomo!. 53:1063-1065. Stewart, R. K. and H. Khoury. 1976. The biology of Lygus lineolaris (Palisot de Beauvois) (Hemiptera:Miridae) in Quebec. Ann. Entomo!. Soc. Quebec. 21:52-63. Wheeler, A. G., Jr. 1976. Lygus bugs as facultative predators. p. 28-35, in D. R. Scott and L. E. O'Keefe (eds.). Lygus bug: host-plant interactions. Unlv. Press Idaho, Moscow. Wheeler, A. G., Jr., B. R. Stinner, and T. J. Henry. 1975. Biology and nymphal stages of Deraeocoris nebulosus (Hemiptera:Miridae), a predator of pests on ornamen­ tals. Ann. Entomo!' Soc. Amer. 68:1063-1068. 19S3 THE GREAT LAKES ENTOMOLOGIST 43

EFFECTS OF VARIOUS SPLIT DEVELOPMENTAL PHOTOPHASES AND CONSTANT LIGHT DURING EACH 24 HOUR PERIOD ON ADULT MORPHOLOGY IN THYANTA CALCEATA (HEMIPTERA: PENTATOMIDAE) J. E. McPherson, IT. E. Vogt, I and S. M. Paskewitz 2

ABSTRACT

Rearing immatures of Thyanta calceata in a range of split photophases during each 24 h period and in constant light showed that the adult dimorphic response in color and pu­ bescence could be produced; individuals reared in photoperiods in which each scotophase was at least 2 h in length generally developed into the fall/spring morpho

Thyanta calceata (Say) ranges from New England south to , and west to Illinois (Blatchley 1926) and Missouri (Oetting and Yonke 1971). This phytophagous stink bug exhibits adult dimorphism. McPherson (l977a) has shown it to be bivoltine and seasonally dimorphic; green adults with short pubescence (shorter than diameter of tibia) are found during the summer months, and brown adults with long pubescence during the fall and spring. Adult dimorphism results from developmental photoperiod (McPherson 1977b, 1975a) with a threshold photoperiod ofabout 12.5L: 11.5D involved in the dimorphic response (McPherson 1975b); reared in photophases above and below the threshold develop into the summer and fall/spring morphs, respectively. To determine if the photophase during each 24 h period had to be continuous (e.g., 16 h) or could be split (e.g., S h, S h) and still produce the same morph, McPherson and Paskewitz (l9S2) reared animals under SL: 16D, 8L:4D:8L:4D, and J6L:SD photoperiods. The SL:4D:8L:4D photoperiod exposed the animals to only 8 h of continuous light but to a total of 16 h oflight/24 h. They found that those reared under SL: 16D and SL:4D:8L:4D became the fall/spring morph, and those in 16L:8D the summer morpho Thus, during each 24 h period, it is the length of each photophase, rather than the combined lengths of all photo­ phases, that determine the adult morpho Also, since seotophases of 16 h and 4 h were involved in the production of the fall/spring morph, and 8 h the production of the summer morph, it appeared that the scotophase was functioning only to break the photophase and the length of the scotophase was unimportant down to 4 h. This raised another question. What was the length of the scotophase below which the animals would no longer respond but, instead, develop into the summer morph? The results of an experiment to determine this are presented here.

METHODS AND MATERIALS

Fifty males and 50 females from F I generation laboratory stock were placed in an incu­ bator (23.9 1.1"C) under a 24L:OD photoperiod; the stock was established with indi­ viduals collected July 1982 in Poinsett County, . They were maintained in mason jars (five of each sex/jar) provided with cheesecloth as an oviposition site, a paper toweling strip, and filter paper, and fed green snap beans (Phaseolus vulgaris L.), as described by McPherson (1971).

IDepartment of Zoology, Southern Illinois University, Carbondale, IL 62901. 2Department of Entomology, University of Georgia, Athens, GA 30602. "'"'

>-l ::r:: t'I1 a ;:0 t'I1 Table I. Comparison of color and pubescence between Thyanta calceata adults reared in various split photophases and constant light. >-l> t""' Color > t'I1 '"r:n Dorsal Ventral Pubescence t'I1 Z Photoperiod Sex Brown Green Prob. Brown Green Prob. Short Long Prob. >-l 0 s::: 8L:4D:8L:4D 0 49 49 I 0 50 0 a a a t""' 9L:3D:9L:3D 49 0.76 48 2 0.50 I 49 0.50 0

aH 9L:3D:9L:3D 0 49 I 48 2 I 49 r:n 10L:2D: 1OL:2D 48 2 0.50a 48 2 0.69a 2 48 0.50a >-l

1OL:2D: 10L:2D 0 48 2 48 2 2 48 IlL: ID: IlL: 1D 19 31 35.46b* 19 31 35.46 b* 18 32 14.06b * -< !2.. IlL: ID: IlL: 1D 0 19 31 19 31 18 32 I1.5L:0.5D: 11.5L:0.5D I 49 18.06 b* I 49 18.06 b* 50 0 O.ooa* ~ Z 11.5L:0.5D: 11.5L:0.5D 0 I 49 I 49 50 0 ? 24L:OD 3 47 0.31a 3 47 0.31a 49 I 0.50a N 8L:4D:8L:4D 0 49 I 49 I 0 50 \:i; 00 24L:OD 3 47 8l.J3b* 3 47 81.13 b* 49 I o.ooa* """ 8L:4D:8L:4D ¥ 43 7 43 7 3 47 9L:3D:9L:3D 46 4 OAlb 45 5 O.09b 3 47 O.OOb

9L:3D:9L:3D S' 46 4 45 5 3 47 IOL:2D: lor .:2D 44 6 O. lOb 44 6 O.OOb 3 47 O.OOb

10L:2D: JOL:2D ¥ 44 6 44 6 3 47 >-l ilL: 1D: ilL: 1D 8 42 49.0gb* S 42 49.0S b* 36 14 43.04b* ::r: til Cl IlL: 1D: IlL: 1D ¥ 8 42 8 42 36 14 :;:l 11.5L:0.5D: 11.5L:0.5D I 49 4AOb* I 49 4AOb* 50 0 O.ooa* til ~ 11.5L:0.5D: 11.5L:0.5D ¥ I 49 I 49 50 0 24L:OD 0 50 0.50a 0 50 0.50a 49 1 0.50a ~ ::-:: til 8L:4D:8L:4D ¥ 43 7 43 7 3 47 r.J'l O.ooa* O.ooa* b til 24L:OD 0 50 0 50 49 I 81.13 * Z exact probability test. as:: x 2 x' test for independent samples corrected for continuity. 0 *Significant at the 0.05 level of probability. § '""r.J'l >-l

..,.. v. 46 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No.2

Each resulting egg cluster was placed in one of the following six photoperiods and reared to adults as described by McPherson (1971): 8L:4D:8L:4D, 9L:3D:9L:3D, 10L:2D: IOL:2D, IlL: ID: IlL: ID, II.5L:0.5D: 11.5L:0.5D, and 24L:OD. All individuals were reared In23.9 I. 1°C and in about 260 ft-c during the light phases (Sylvania, 15W Daylight, FI5T8/D). Adult characters compared were color (green or brown) and pubescence (long or short); short hairs were defined as those shorter than the diameter of the tibia. The 0.05 level of significance was chosen for all comparisons.

RESULTS AND DISCUSSION

Rearing males or females in 8L:4D:8L:4D, 9L:3D:9L:3D, and IOL:2D: IOL:2D produced similar results; most individuals were brown (males, 96--98%; females, 86--92%) with long pubescence (males, 96--100%; females, 94%) (Table I). Rearing the two sexes in llL:ID: IlL: I D produced a marked increase in the percentage ofgreen adults (males, 62%; females, 84%) and adults with short pubescence (males, 36%; females, 72%). Rearing individuals in 11.5L:0.5D: 11.5L:0.5D produced a second increase (98% green adults, 100% short pu­ bescence; both sexes) and these percentages were similar to those for individuals reared in constant light. These results show that the developmental photophase generally must be longer than 2 h for the fall/spring morph (brown with long pubescence) to be produced. Between 2 hand 1/2 h, most individuals no longer respond but appear as though reared in constant light (i.e., develop into green adults with short pubescence). As the length of the scotophase is de­ creased, females generally fail to respond before males (recall that at IlL: ID: ilL: ID, 84% of the females were green compared to only 62% of the males). Thus, scotophase, as shown in the earlier experiment (McPherson and Paskewitz 1982), does function to break the photophase but can be overridden ifthe scotophase is not of sufficient duration.

ACKNOWLEDGMENT

We wish to thank Dr. H. E. Barton, Department of Biological Sciences, Arkansas State University, State University, for sending us the specimens used in establishing the labora­ tory culture.

LffERATURE CITED

Blatchley, W. S. 1926. Heteroptera or true bugs of eastern North America with especial reference to the faunas of Indiana and Florida. Nature Pub!. Co., Indianapolis. McPherson, J. E. 1971. Laboratory rearing of Euschistus tristigmus tristigmus. J. Econ. Entomo!. 64: 1339-1340. ----. 1977a. Notes on the biology of Thyanta calceata (Hemiptem:Pentatomidae) with information on adult seasonal dimorphism. Ann. Entomol. Soc. Amer. 70:370-372. 1977b. Effects of developmental photoperiod on adult color and pubescence in Thyanta calceala (Hemiptera:Pentatomidae) with information on ability of adults to change color. Ann. Entomo!. Soc. Arner. 70:373-376. ----. 1978a. Sensitivity of immature TJryanta calceata (Hemiptera:Pentatomidae) to photoperiod as reflected by adult color and pubescence. Great Lakes Entomo!. 11:71-76. 1978b. Effects of various photoperiods on color and pubescence in Thyanta calceata (Hemiptera:Pentatomidae). Great Lakes Entomo!' II: 155-158. McPherson, 1. E. and S. M. Paskewitz. 1982. Effects of continuous and split developmental photophases during each 24 hour period on adult color and pubescence in Thyanta calceata (Hemiptera:Pentatomidae). Great Lakes Entomo!' 15:97-98. Oetting, R. D. and T. R. Yonke. 1971. Biology of some Missouri stink bugs. 1. Entomo!. Soc. 44:446--459. 1983 THE GREAT LAKES ENTOMOLOGIST 47

BUPRESTIDAE, CERAMBVCIDAE, AND SCOlVTIDAE ASSOCIATED WITH SUCCESSIVE STAGES OF AGRILUS BILINEATUS (COlEOP"rERA: BUPRESTIDAE) INFESTATION OF OAKS IN WISCONSIN 1 Roben A. Haack2, Daniel M. Benjamin3, and Kevin D. Haack4

ABSTRACT

The species of Buprestidae, Cerambycidae, and Scolytidae found in association with Agrilus bilineatus (Weber) in declining oaks, Quercus spp., in Wisconsin, were Chryso­ bothris femorata (Olivier) and Dicerca sp. (Buprestidae); Amniscus macula (Say), Cyrta­ phorus verrucosus (Olivier), picipes (Fabricius), Graphisurusfasciatus (DeGeer), Neodytus acuminatus (Fabricius), Sarosesthes fulminans (Fabricius), and Xylotrechus colonus (Fabricius) (Cerambycidae); and Monarthrum fasciatum (Say), Monarthrum mali (Fitch), Pseudopityophthorus minutissimus (Zimmerman), and Xylaterinus paUlUS (Say) (Scolytidae). In general, weakened oaks were first attacked by A. bilineatus. and at times that same year by C. femorata. G. fasdalus, and P. minutissimus. Infestation by M.fasci­ atum, M. mali, and X. poUtus began the season following first attack by A. bilineatus. With the exception of A. bilineatus, the above mentioned Buprestidae and Cerambycidae ap­ peared to preferentially infest dead wood, often those portions that had died the previous season.

The twolined chestnut borer, Agrilus bilineatus (Weber), (Coleoptera: Buprestidae) is a major pest of weakened oaks (Quercus spp.) throughout eastern Nonh America. Recent outbreaks (1976-1980) in southern Wisconsin occurred where oaks had been stressed by drought, ice-storm damage, and fall cankerworm, Alsophila pomelaria (Harris), (Lepidop­ tera: Geometridae) defoliation. The biology of A. bilineatus has recently been studied by Dunbar and Stephens (1975, [976) in Connecticut; Cote and Allen (1980) in New York and Pennsylvania; and Haack, Benjamin, and Schuh (1981) and Haack and Benjamin (1982) in Wisconsin. In declining oaks, A. bilineatus is normally the first borer to attack (Dunbar and Stephens 1975). Initial attack usually occurs first in the live crown and then proceeds downward along the trunk in succeeding years. Attacked trees usually die during August and September of the second or third year of infestation. Occasionally, tree death occurs in the first year of attack when girdling is complete below the first major branches (Haack and Benjamin 1982). Once an oak is attacked or killed by A. bilineatu$, it becomes a suitable host to several other wood borers, inner bark (phloem) borers, bark beetles, and ambrosia beetles in the families Buprestidae, Cerambycidae, and Scolytidae. Study of borer succession is of practi­ cal imponance when attempting to assess the impact of several insect species on a host. Nevertheless, few such studies have been done. Savely (1939) reponed on the succession of animals in oak logs over a four year period in North Carolina, and Cote and Allen (1980)

I Research supported by the School of Natural Resources, College of Agricultural and Life Sciences, University of Wisconsin-Madison, and tne Wisconsin Department of Natural Resources. 2Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611. 3Department of Entomology, University of Wisconsin, Madison, WI 53706. 4Department of Entomology, A&M University. College Station, TX 77843. 48 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No.2 listed the associated borers they had reared from A. bilineatus-infested oak bolts. We present here observations on the Buprestidae, Cerambycidae, and Scolytidae associated with A. bilineatus in oaks in various stages of decline, recorded in Wisconsin in 1979.

MATERIALS AND METHODS

The period of adult emergence was investigated in 1979 by capturing adults in traps (0.4-().7 m2l made of nylon mosquito netting stapled at ca. breast height (1.3 m) on 51 oaks (20-70 cm db h) in four host-condition categories. Trees were assigned to a category based on external symptoms and signs of A. bilineatus attack during fall 1978 and spring 1979. Cate­ gories were (I) apparently healthy, i.e., having no evidence of 1978 A. bilineatus attack; (2) some 1978 crown attack; (3) tree death in 1978; and (4) tree death in 1977. Traps were installed in late May 1979 on 10 white oaks (Q. alba 1..), 7 black oaks (Q. velutina Lam.), 2 red oaks (Q. rubra L.), and I bur oak (Q. macrocarpa Michx.) in an oak woodlot near Madison, Dane County, Wisconsin, and on 10 white oaks, 7 black oaks, 2 red oaks, and I bur oak in a natural oak-hickory forest in the Kettle Moraine State Forest, Jefferson County, Wisconsin. Adults were removed and counted twice each week from 20 May through 5 July and then weekly through 14 September. Adults were identified at the University of Wiscon­ sin and at the Insect Identitication and Beneficial Introduction Institute, USDA, Beltsville, . In an earlier paper (Haack and Benjamin 1982) the within-tree distributions of A. biline­ atus larvae, larval galleries, and adult exit holes were presented for five host-condition categories of oaks from the Kettle Moraine State Forest site. During that study we also recorded the number and location of all other borers encountered. Briefly, 25 red and black oaks (5/category, 23-46 cm dbh) were sampled in December 1979 after having been assigned to a host-condition category in September 1979, when symptoms of current-year A. biline­ atus attack were most evident. Categories were (I) apparently healthy, i.e., having no symptoms of 1979 A. bilineatus attack; (II) 25-50% crown death in 1979 after one season of attack; (III) tree death in 1979 after one season of attack; (IV) tree death in 1979 after two seasons of attack; and (V) tree death in 1978 after at least two seasons of attack. After felling, bolts 30-cm-Iong were cut at 2-m intervals from tree base out along one major branch to a final diameter of ca. 5 cm. We recorded the length and diam. (inside the bark) of each bolt. All borers were recovered from the bark, cambial region, and wood by carefully splitting with hammer and chisel; numbers were recorded per square metre of bolt area. The Scolytidae were identified to species. However, the Buprestidae and Cerambycidae (larvae) were only identified to the family and genus level, using the keys of Burke (1917), Craighead (1923), and Peterson (1960); some larvae were reared to adults.

RESULTS AND DISCUSSION

Two species of Buprestidae and seven species of Cerambycidae were collected from the emergence traps. The Buprestidae were A. bilineatus and Chrysobothris femorata (Olivier). The Cerambycidae were Amniscus macula (Say), Cyrtophorus verrucosus (Olivier), Euderces picipes Fabricius), fasciatus (DeGeer), Neoclytus acuminatus (Fabricius), Sarosesthes fulminans (Fabricius), and Xylotrechus colonus (Fabricius). Table 1 presents number collected, collection period, host trees, and host tree condition for each borer species. Borers were collected only from traps on dead oaks (categories 3 and 4); none were collected from the healthy or crown-attacked oaks. However, later inspection of the crown­ attacked oaks revealed A. bilineatus exit holes in crown branches and along the upper bole of each, but none along the lower trunks where the traps had been placed. The most commonly collected cerambycid in our study was G. fasciatus; it had a similar ranking in the studies of Savely (1939) and Cote and Allen (1980). At times, A. bilineatus, C.jemorata, and G. fasciatus were all found in the same traps suggesting concurrent attack since each is considered to be univoltine (Haack and Benjamin 1982, Chittenden 1905, and Craighead 1983 THE GREAT LAKES ENTOMOLOGIST 49

Table 1. Adult Buprestidae and Cerambycidae collected from emergence traps at breast height (1.3 m) on Agrilus bilineatus-infested oaks in an oak woodlot (Madison, Dane Co., Wisconsin) and in a natural oak-hickory forest (Kettle Moraine State Forest, Jefferson Co., Wisconsin) from 20 May through 14 September 1979.

Collection Species (n) Period Host and Host Condition a

BUPRESTIDAE

Agrilus bilineatus 126 8 June-26 July Q. alba (1), Q. rubra (1), Q. velutina (1) Chrysobothris femorata 8 13 June- 4 July Q. alba (2), Q. rubra (2), Q. velutina (1,2)

CERAMBYCIDAE

Amniscus macula 2 1June- 8 June Q. velutina (2) Cyrtophorus verrucosus 6 8 June- 4 July Q. alba (2), Q. velutina (2) Euderces picipes 3 3 June-ll June Q. velutina (2) Graphisurus fasciatus 28 8 J une-29 June Q. alba (2), Q. rubra (2), Q. velutina (1,2) Neoclytus acuminatus 4 17 June-26 June Q. rubra (2), Q. velutina (2) Sarosesthes fulminans 7 20 June- 4 July Q. velutina (2) Xylotrechus colonus 13 8 June- 4 July Q. rubra (2), Q. velutina (2)

aparenthetical numbers represent the host tree condition in 1979 during the period ofadult emergence: 1 ~ oaks dead for ca. 1 year (died fall 1978). 2 ~ oaks dead for ca. 2 years (died fall 1977).

1923, respectively). On no occasion did we collect C. femorata or G. fasciatus from cate­ gory 3 oaks without also collecting A. bilineatus, but A. bilineatus was at times the only borer recovered from a given trap. Savely (1939) also found the above three borers, as well as X. colonus, together in oak logs sampled within a year of felling. Craighead (1923) reported a one-year life cycle for X. colonus. However, Gardiner (1960) stated that two years were required for X. colon us to complete development in Quebec. A one-year life cycle has also been reported for N. acuminatus (Baker 1972), A. macula (Craighead 1923), and C. verrucosus (Duffy 1953). To our knowledge, voltinism data are not available for E. picipes and S. fulminans. However, Craighead (1923) and Knull (1946) reported that these two species normally attack dead wood, often the season following tree death. If this was the case in our study, then E. picipes and S. fulminans would be univoltine in Wisconsin since they emerged from oaks the second summer following tree death. Although category I oaks had appeared unattacked, two of the five oaks had been at­ tacked by A. bilineatus in 1979. Nevertheless, as Haack and Benjamin (1982) reported, all recovered A. bilineatus larvae (n = 83) had died as first or second instars. Only one other borer was found, a Chrysobothris larva. This larva was found alive in the cambial region of a branch sample (7 cm dia.) where three dead A. bilineatus larvae were recovered. Ap­ parently, A. bilineatus seldom attacks relatively vigorous oaks, but when it does, larval development is usually not successful. Borers other than A. bilineatus seem to attack vigor­ ous oaks even less often; the above Chrysobothris larva probably arrived once A. bilineatus larvae were already present based on the apparent precedence ofthe latter's galleries. Category II oaks had 25-50% crown death after one season of A. bilineatus attack. We recovered larvae of A. bilineatus and Chrysobothris, and pupae and teneral adults of Pseudopityophthorus minutissimus (Zimmerman) (Scolytidae); the percentage of samples containing each borer and the host tissues from which they were collected are given in Table II, and their mean densities are presented in Figure I. Over 90% of the A. bilineatus larvae were in pupal cells constructed in outer bark if thick or in sapwood if the bark was thin. However, A. bilineatus larvae feed and develop primarily in phloem (inner bark). Two of the 50 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No. 2

80 CATEGORY II

N ~ 60 BILINEATUS /' B - BUPRE STI OA E A_A

a:: 40 s ­ SCOLYTIOAE / S W ID ~ ,-,-' ::;) 20 / z ,_,-,-!-, 0 AI

0.5 4.5 8.5 12.5 HEIGHT ABOVE GROUND (M)

Fig. 1. Mean Agrillls bilineatus, Buprestidae (Chrysobothris), and Scolytidae (Pselldopityophthorlls) densities at various heights above ground for five oaks sampled in December 1979 that had 25-50<% crown death in September 1979 after one season of A. bilineatus attack, Kettle Moraine State Forest, Jefferson Co., Wisconsin.

Chrysobothris larvae were found constructing pupal cells in the sapwood; the others were in the cambial region. P. minutissimus was recovered from a single gallery system in a 4-cm­ dia. branch sample. Also working in Wisconsin, McMullen et al. (1955) recorded two P. minutissimus generations per year with flight periods in May and August, and with last­ instar larvae being the overwintering form. In our study, the relatively warm fall tempera­ tures of 1979 may have allowed for more advanced development (pupae and adults) relative to the McMullen et aI. (1955) study (larvae). In southern , P. minutissimus successfully overwinters in every stage but the pupal stage (Rexrode 1969). No empty P. minutissimus galleries (representing the first generation of 1979) were observed. This is not surprising because this beetle normally infests only dead or dying oaks (Rexrode 1969). Therefore, the category II oaks were probably not suitable for P. minutissimus colonization during its May flight period. However, by August, these oaks were suitable for P. minutissimus, as a result of the A. bilineatus branch-girdling that had taken place that summer. Category III oaks had died the year of sampling after one season of A. bilineatus attack. We recovered larvae of A. bilineatus, Chrysabothris, and Graphisllrus, and brood of P. minutissimus (Table II, Fig. 2). Larval densities of A. bilineatus were highest in the upper clear bole; major branching began between 7 and 9 m in the oaks of this study. Over 90% of the A. bilineatlls were recovered from pupal cells. We found 24 Chlysobothris larvae con­ structing pupal cells in sapwood and 37 in the cambial region. All Graphisurus larvae were collected from the same tree; two of the 17 larvae were constructing pupal cells in the cambial region. We collected last-instar larvae, pupae, and callow adults of P. minutissimus from one branch sample (5 cm dia.). Category IV oaks had died in the year of sampling after two seasons of A. bi/ineatus attack. We collected larvae of A. bilineatus, Chrysobothris, Graphisurus, Neaciytus, and Xylotrechus, and brood of P. minutissimus and the ambrosia beetles (Scolytidae) Monar­ thrum Jasciatum (Say), Monarthrum mali (Fitch), and Xyloterinus paUtus (Say) (Table II, Fig. 3). Note that the values given for the numbers of ambrosia beetles in Table II represent numbers of active galleries, not numbers of individuals. No living A. bilineatus larvae were found above the 6.5-m sampling height where it had attacked the year before; the average lowest extent of A. bilineatus exit holes was ca. 7.5 m. Current A. bilineatus larval densities 1983 THE GREAT LAKES ENTOMOLOGIST 51

CATEGORY III

120 A

100

(\j :::;; /\1\ SO

0:: UJ 60 S al """"'"' :::!! /:-~B- BUPRESTIDAE \ :::> C- CERAMBYCIDAE A/'1 z 40 5 - SCOLYTIDAE

20 B__B_B -- c 0 B c- --C-!:l:::;:::::e:::::::::~ .::::::@

0,5 4,5 S.5 12,5 HEIGHT ABOVE GROUND (M)

Fig. 2. Mean Agrilus bilineatus, Buprestidae (Chrysobothris), Cerambycidae (GraphisllruS), and Scoly­ tidae (Pseudopityophthorus) densities at various heights above ground for five oaks sampled in Decem­ ber 1979 that had died in September 1979 after one season of A. bilineatus attack, Kettle Moraine State Forest, Jefferson Co" Wisconsin.

100 CATEGORY IV

80 A 1:.. BlllHEATUS OJ ::e 60

0: W 40 '"::e :::> z 20

0

4,5 8,5 12,5 HEIGHT ABOVE GROUND 1M J

Fig. 3, Mean Agrillis bilineatlts, Buprestidae (Chrysobothris), Cerambycidae (Graphisurus, Neoc!ytus, XyJotrechlls), and Scolytidae (Monarthrllm, Pselldopityophthorlls, Xyloterinus) densities at various heights above ground for five oaks sampled in December 1979 that had died in September 1979 after two seasons of A. hilineatlls attack, Kettle Moraine State Forest. Jefferson Co" Wisconsin, 52 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No.2 were highest in the 2.5-m samples; lower densities at 4.5 and 6.5 m may be related to the relati vely drier condition of the cambial region through either differential larval mortality (if egg density was uniform) or differential oviposition (if parent females oviposited preferen­ tially), or a combination of both. We found 35 Chrysobothris larvae constructing pupal cells in sapwood and 112 in the cambial region. Chrysobothris densities were highest in the upper trunk of category IV oaks where A. bilineatus had attacked the previous year. Graphisurus larval densities were highest in the region (6.5 and 8.5 m) that represented the lowest extent of A. bilineatus infestation during the previous year. In the cambial region, a few Graph­ isurus larvae but no Xylotrechus larvae were preparing pupal cells. Xylotrechus and Neo­ elytus larvae were recovered primarily in samples from the zone between previous-year and current-year A. bilineatus attack. The active galleries of M. Jasciatum and X. POlilUS con­ tained mostly callow adults and some pupae; M. mali galleries had callow adults only. In Indiana, X. politus overwinters as adults and is univoltine (MacLean and Giese 1967). In Missouri, M. Jasciatum overwinters as adults and completes 2-3 generations per year with the initial flight period occurring between late March and mid-May (Roling and Kearby 1974). In Wisconsin, X. POlilUS appears to be univoltine with adult emergence occurring mostly from early April to mid-May (J. O. Haanstad, pers. comm.). 5 Flight period data are not available for the Monarthrum species in Wisconsin, but they are probably similar to that of X. politus because we found galleries ofX. poUtus and Monarthrum spp. ofsimilar length and having similar life stages in the same oaks sampled in June 1979. Therefore, since the A. bilineatus flight period does not occur until early June in Wisconsin (Haack and Benjamin 1982), the lower trunk of these oaks was probably attacked first in 1979 by ambrosia beetles. We also noticed in these category IV oaks that A. bilineatus larval mines seldom were found in the stained region of phloem and sapwood surrounding the entrance hole of each am­ brosia-beetle gallery, suggesting that the staining (and thus the ambrosia beetles) preceded A. bilineatus. The relatively early flight period of the ambrosia beetles may help to explain why none were collected in our adult-emergence study; traps were installed the third week of May which was after their peak flight. The P. minutissimus brood (adults and pupae) was recovered from two galleries in the same branch sample (8 cm dia.) that had been infested the previous year by A. bilineatus, indicating that this scolytid attacks material both con­ currently with and the year follOWing A. bilineatus attack. Category V oaks had died the year prior to sampling after at least two seasons of A. bilineatus attack. We found larvae of Chrysobothris. Dicerca (Buprestidae), Cyrlophorus. Graphisurus, Neoclytus, Sarosesthes, and Xyiotrechlls, and brood of X. poUtus (Table 2, Fig. 4). Chrysobothris larvae were found only in lower-trunk samples, indicating that they infest material killed the previous year but not material killed two seasons earlier; seven larvae were in sapwood and 17 in the cambial region. However, the Dicerca larvae were only found in samples from branches that had died two seasons earlier. Savely (1939) collected Dicerca larvae from oak logs sampled ca. two years after felling. The cerambycid larvae were collected primarily from the lower trunk where death had occurred the previous year. Nevertheless, a few Graphisurus, Neoclytus, and Xylotrechus larvae (n 18) were collected from branches that had died two seasons earlier. It is uncertain ifthese larvae had developed from eggs laid in 1978 or 1979. However, since (I) most larvae appeared fully grown and were in or constructing pupal cells, (2) cerambycid exit holes were already present in some of these same samples, and (3) the phloem and xylem in these samples appeared very dry relative to the lower-trunk samples, it is probable that these larvae began development in 1978 and thus would have required two years to complete their life cycle. Sarosesthes larvae were most common at 2.5 m; this species appeared to prefer the lower trunk of oaks having died the previous season. Active X. politlls galleries were found mostly in lower trunk samples. Apparently, this scolytid attacks oaks the year after death, but not portions having died two seasons earlier. We found several old galleries ofM.fasciatum. M. mali, and X. poUtus that were active the previous season; old P. mifllltissimus galleries were also present.

5Department of Entomology. University of Wisconsin, Madison, WI 53706. 1983 THE GREAT LAKES ENTOMOLOGIST 53

Table 2. Percent of oak samples containing Buprestidae, Cerambycidae, and Scolytidae described by host condition category and height above ground (5 oaks/category; Kettle Moraine State Forest, Jefferson Co., Wisconsin; December 1979).

Sampling Heights (m)

Borer 0.5 2.5 4.5 6.5 8.5 10.5 12.5 14.5 (Na) Siteb

Category II Oaks A. bilineatus 80 80 100 100 100 100 (186) B,C,SW Chrysobothris 60 40 20 (6) C,SW P. minutissimus 20 (8) C

Category III Oaks A. bilineatus 100 100 100 100 100 100 100 100 (638) B,C,SW Chrysobothris 60 40 40 20 20 20 60 20 (61) C,SW Graphisurus 20 20 20 20 20 20 (17) C P. minutissimus 20 (14) C

Category IV Oaks A. bilineatus 100 100 100 60 (230) B,C,SW Chrysobothris 20 80 80 100 100 80 40 20 (147) C,SW Graphisurus 60 80 80 60 (137) C Neoclytus 60 60 20 (11) SW Xylotrechus 80 80 (14) C M. fasciatum 20 40 (3) SW M. mali 20 20 (3) SW P. minutissimus 20 (14) C X. poUtus 60 60 80 60 20 20 (42) SW

Category V Oaks Chrysobothris 60 80 40 (24) C,SW Dicerca 20 20 (2) C Cyrtophorus 20 20 20 20 (4) SW Graphisurus 100 100 100 100 20 20 (99) C Neoclytus 20 60 40 40 40 (11) SW Sarosesthes 60 100 60 20 (28) C,SW Xylotrechus 80 100 100 60 80 (48) C X. poUtus 80 80 80 20 (40) SW aN the number of individuals (or gallery systems for tile ambrosia beetles: M. jasciatum, M. mali, X. politus) recovered from all samples for the indicated host condition category of oaks. bSite refers to the area(s) in which each borer was recovered: B outer bark, C cambial region, SW sapwood.

SUMMARY

A successional pattern of Buprestidae, Cerambycidae, and Scolytidae can now be ap­ proximated for a typical declining oak in Wisconsin. In general, A. bilineatus begins the assault by first attacking the branches and upper trunk during the summer of year one. At times, A. bilineatus is joined that first summer by small numbers of Chrysobothris, 54 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No.2

50 CATEGORY V

40 N ::Ii C",c 8'

"- / ~C C~ CERAMSYCIDAE 30 C a: \ S- SCOLn.C" w ro 20 ::Ii :::l C Z 10 5-i::::-..... ~ a/" '-'5a:::::-"'" C "­ 0 ~A>-C~---B~B

0.5 4,5 8,5 12.5 HEIGHT ABOVE GROUND (M)

Fig, 4. Mean Buprestidae (Chrysobolhris, Dicerca), Cerambycidae (Cyrlophorus, Graphisums, Neo­ ely/us, Saroseslhes, Xylotrechus), and Scolytidae (Xy/oteriflus) densities at various heights above ground for five oaks sampled in December 1979 that had died in fall 1978 after at least two seasons ofA. bilinea/us attack, Kettle Moraine State Forest, Jefferson Co" Wisconsin,

Graphisurus, and P. minutissimus. During spring of year two, the lower trunk is colonized by ambrosia beetles such as M. fasciatum, M. mali, and X. poUtus. In the summer of year two, A bilineatus moves its attack to the lower trunk where ambrosia beetles have already begun construction of their galleries. Also at this time, Chrysobothris and Graphisurus may join A. bilineatus in the lower trunk, and also reattack the upper trunk and crown along with P. minutissimus. Joining Chrysobothris and Graphisurus in the upper trunk during the summer of year two are Neoclytus and Xylotrechus. In spring of year three, X. poUtus continues to colonize the lower trunk which had died in the fall of year two. In the summer of year three, following the exit of A. bilineatus, the lower trunk is now infested by Graph­ isurus, Neociytus, and Xylotrechus as well as Cyrtoplwrus and Sarosesthes. The upper trunk and major branches which had died two seasons earlier are now suitable for Dicerca infestation. In this way the insect fauna in the crown branches, upper trunk, and lower trunk changes from year to year during the decline of an oak tree and also after its death.

ACKNOWLEDGMENTS

We thank Dr. Dave J. Hall and Paul E. Pingrey, Wisconsin Department of Natural Re­ sources, for assistance in locating forest stands; Tom Byrnes for access to his oak woodlot; John O. Haanstad for use of his Xyloterinus politus emergence data; Dr. D. M. Anderson, Insect Identification and Beneficial Insect Introduction Institute, USDA, Beltsville, MD, for identification of the Scolytidae; and Dr. Douglas C. Allen, Dr. David G. Nielsen, Dr. James D. Solomon, and John O. Haanstad for critical review ofthis manuscript.

LITERATURE CITED

Baker, W. L. 1972. Eastern forest insects. USDA Misc. Pub!. 1175. Burke, H. E. 1917. Flat-headed borers affecting forest trees in the United States. USDA Bull. 437. Chittenden, F. H. 1905. The flat-headed apple tree borer (Chrysobothris femorata). USDA Bur. Entomo!' 32 rev. 1983 THE GREAT LAKES ENTOMOLOGIST 55

Cote, W. A., III and D. C. Allen. 1980. Biology of two-lined chestnut borer, Agrilus bilineatus. in Pennsylvania and New York. Ann. Entomol. Soc. Amer. 73:409-413. Craighead, F. C. 1923. North american cerambycid larvae. Canadian Dept. Agric. Entomol. Branch Bull. 27. Duffy, E. A. J. 1953. A monograph of the immature stages of British and imported timber beetles (Ceram bycidae). British Mus. (Natur. Hist.) London. Dunbar, D. M. and G. R. Stephens. 1975. Association of twolined chestnut borer and shoestring fungus with mortality of defoliated oak in Connecticut. For. Sci. 21: 169-174. ----. 1976. The bionomics of the two lined chestnut borer. p. 73--83, in I. F. Anderson and H. K. Kaya (eds.). Perspectives in forest entomology. Academic Press, New York. Gardiner, L. M. 1960. Description of immature forms and biology of Xylntrechus colonus (Fab.) (Coleoptera: Cerambycidae). Canadian Entomo!' 92:820-S25. Haack, R. A. and D. M. Benjamin. 1982. The biology and ecology of the twolined chestnut borer. Agrilus bilineatus (Coleoptera: Buprestidae), on oaks, Quercus spp., in Wisconsin. Canadian Entomo!. 114:385-396. Haack, R. A., D. M. Benjamin, and B. A. Schuh. 1981. Observations on the biology of Phasgonophora sulcata (Hymenoptera: Chalcididae), a larval parasitoid of the twolined chestnut borer, Agrilus bilineatus (Coleoptera: Buprestidae). in Wisconsin. Great Lakes Entomo!' 14: 113-116. Knul!, J. N. 1946. The longhorned beetles of Ohio (Coleoptera: Cerambycidae). Ohio BioI. Surv. Bull. 39: 133-354. MacLean, D. B. and R. L. Giese. 1967. The life history of the ambrosia beetle Xyloterinus poUtus (Coleoptera: Scolytidae). Canadian Entomo!' 99:285-299. McMullen, L. H .• E. W. King, and R. D. Shenefelt. 1955. The oak bark beetle, Pseudo­ pityophthorus minutissimus (Zimm.) (Coleoptera: Scolytidae) and its biology in Wiscon­ sin. Canadian Enlomol. 87:755-757. Peterson, A. 1960. Larvae of insects. Edwards Brothers, Ann Arbor. Rexrode, C. O. 1969. Seasonal development and habits of Pseudopityophthorus spp. (Coleoptera: Scolytidae) in southern Ohio. Canadian Entomol. 101:306-313. Roling, M. P. and W. H. Kearby. 1974. Life stages and development of Monarthrum fasciatum (Coleoptera: Scolytidae) in dying and dead oak trees. Canadian Entomo!. 106: 1301-1308. Savely, H. E., If. 1939. Ecological relations of certain animals in dead pine and oak logs. Ecol. Monog. 9:321-385. 1983 THE GREAT LAKES ENTOMOLOGIST 57

A PYRALID MOTH (LEPIDOPTERA) AS POLLINATOR OF BLUNT-LEAF ORCHID

Edward G. Voss i and Richard E. Riefner, Jr. 2

As early as 1912, mosquitoes were observed bearing the pollen masses (pollinia) of the blunt-leaf orchid, Habenaria obtusata (Pursh) Richardson, in Reese's Bog, a cedar swamp at the north end of Burt Lake, Cheboygan County, Michigan, near the campus of the University of Michigan Biological Station (Dexter 1913). A number of subsequent observers in Canada and the northern United States have reported pollinia of this orchid on mosqui­ toes, always females of the genus Aedes. Details of mosquito behavior and the pollination process have recently been described, with excellent pictures, by Stoutamire (1968), Thien (1969), and Thien & Utech (1970). These authors also noted three species of Xanthorhoe (Geometridae) as removing pollinia; two other geometrids were listed by Thien and Utech (1970) as very rarely bearing pollinia, but no other or insects of other orders have been reported as pollinators. The blunt-leaf orchid is a circumpolar species, ranging in North America from the tundra south into Michigan (Voss 1972) as far as the latitude of Saginaw Bay (with an old outlying record in northern St. Clair County). Toward this southern limit of its range, it favors coniferous swamps and forests, where the soil is cold and often moist. The inconspicuous plants are rarely over 20 cm tall, with a single leaf and a few greenish flowers about 5---{) mm long, scarcely larger than the mosquitoes which pollinate them. While it is comforting to contemplate that mosquitoes have some important function in the cedar swamps where this little orchid thrives, they have no monopoly on pollination. We now report, as an apparent pollinator, a small moth not much larger than the orchid flower (and smaller than the geometrid species previously noted). Anageshna primordialis (Dyar) was originally placed (1906) in Geshna, but was trans­ ferred by Munroe (1956) to a monotypic new genus. It is now classified in the Pyralidae, subfamily Pyraustinae (not Nymphulinae as originally thought). Forbes (1923, p. 581), who had first questioned its taxonomic placement, stated its occurrence as "Very common in damp places in June and early July." Indeed, it seems to be a rather common moth in the cedar swamps of northern Michigan, although (like most microlepidoptera) not previously reported from the state in any literature we have seen. The wingspread is barely 15 mm, and the wings are brownish with transverse bands of rather angular pale spots (Fig. I). Speci­ mens in the collections of the University of Michigan Museum of Zoology and the Ento­ mology Museum of the Department of Entomology, Michigan State University, indicate a range throughout the Upper Peninsula and northern half of the Lower Peninsula, i.e., the entire range of the orchid in the state; the moth has also been collected farther south, in Berrien, Livingston, Oakland, and Washtenaw counties. Dates in the northern part of the state are 16 June-21 July. None of the museum specimens bear pollinia, perhaps because they were probably not collected in the habitat of this orchid. On 24 June 1976, the specimen shown in Figure I was noted as it was stuck on a flower of Habenaria obtusata in Reese's Bog, where mosquitoes continue to be major pollinators. Eventually the moth pulled itself free without removing the pollinium. However, on 25 June 1981, a moth similarly stuck removed the pollinium, firmly attached to an eye, the usual location for pollinia of this species to become cemented on insect visitors (Fig. 2). One specimen captured in free flight bore two pollinia, both attached to the right eye (Fig. 3). This little pyralid moth has been repeatedly observed resting on the lateral sepals of the orchid flowers; after probing a flower, it pushes its proboscis into the spur and feeds upon

iHerbarium, The University of Michigan. Ann Arbor. MI 48109. 27439 La Palma Ave., Suite 206, Buena Park, CA 90620. 58 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No.2

Fig. I. Anageshna primordialis from a flower of Habenuriu ob/usa/a, Reese's Bog, Cheboygan Co., Michigan. 24 June 1976. Wingspan is 15 mm. (Photo by D. Bay.)

Fig. ' Hnium of Habenaria obtusata attached to right eye of Anageshna primordialis collected on the or"'Hu .n Reese's Bog. 25 June 1981. Kote the stalk by which the pollinium is connected to the sticky pad or viscidium which is firmly cemented to the lower front of the eye. (Photo by D. Bay,) 1983 THE GREAT LAKES ENTOMOLOGIST 59

Fig. 3. Anageshna primordialis with two pollinia attached to the right eye, collected in flight in Reese's Bog, 25 June 1981. (Photo by R. Riefner.)

nectar before taking flight. Moths bearing pollinia have been captured into the evening hours. The moths are apparently not attracted to the flowers from a distance, but as sug­ gested by Stoutamire (1968) for mosquitoes, move toward the flowers after coming to rest on the inflorescence. This behavior seems to suggest attractants perceived close at hand. The flowers tested UV- and have no scent perceptible to us. Thien and Utech (1970) suggested that the longer proboscis in Xanthorhoe (about twice as long as in the mosquitoes) might confer some advantage in obtaining nectar, especially from spurs only partly filled. Further observations are required to confirm that moths do indeed deposit pollen on visits to flowers and to determine any means by which they compete for nectar with mosquitoes, Anageshna, like the mosquitoes, ranges well to the south of the orchid (into Florida; Kimball 1965, p. 219) and clearly is not closely dependent upon it. In fact, it appears to be a moth of quite catholic tastes, having been reported to visit pig carrion in a state of advanced decay (Payne & King 1969).

ACKNOWLEDGMENTS

We are grateful to Dr. Eugene Munroe (Ottawa, Ontario) for identifying the moths we collected; to David Bay for photographic assistance; to curators of the insect collections at Michigan State University and The University of Michigan for the opportunity to examine specimens. Field observations were made in conjunction with the Boreal Flora course at the University of Michigan Biological Station. The specimens shown in Figures 1 and 2 are in the collection of the senior author; that in Figure 3 will be placed in the collection of the Biological Station. 60 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No.2

LITERATURE CITED

Dexter, John Smith. 1913. Mosquitoes pollinating orchids. Science 37:867. Dyar, Harrison G. 1906. The North American Nymphu1inae and Scopariinae. J. New York Entomol. Soc. 14:77-107. Forbes, William T. M. 1923. The Lepidoptera of New York and neighboring states. Cornell Vniv. Agric. Exp. Sta. Mem. 68. 729 p. Kimball, Charles P. 1965. The Lepidoptera of Florida an annotated checklist. Arthropods of Florida and neighboring land areas Vol. 1. Florida Dept. Agric., Gainesville. 363 p. + 26 pI. Munroe, Eugene. 1956. Geshna primordialis Dyar, with descriptions of two new genera and two new subspecies (Lepidoptera: Pyralidae). Canadian Entomol. 88:126-128. Payne, Jerry A. and Edwin W. King. 1969. Lepidoptera associated with pig carrion. J. Lepid. Soc. 23: 191-195. Stoutamire, Warren P. 1968. Mosquito pollination of Habenaria obtusata (Orchidaceae). Michigan Bot. 7:203-212. Thien, Leonard B. 1969. Mosquito pollination of Habenaria obtusata (Orchidaceae). Amer. J. Bot. 56:232-237. Thien, Leonard B. and Frederick Utech. 1970. The mode of pollination in Habenaria obtusata (Orchidaceae). Amer. J. Bot. 57: 1031-1035. Voss, Edward G. 1972. Michigan Flora Part I Gymnosperms and Monocots. Cranbrook Inst. Sci. Bull. 55. 488 p. 1983 THE GREAT LAKES ENTOMOLOGIST 61

CHECKLIST OF AMERICAN ULOBORIDAE (ARACHNIDA: ARANEAE) Brent D. Opell l

ABSTRACT

Names, synonyms, and distributions ofu10borid known from North, Central, and South America are provided.

The first comprehensive revision of American Uloboridae was Muma and Gertsch's (1964) treatment ofthe family's United States and Canadian members. This was followed by Lehtinen's (1967) reevaluation of the family's generic division, Chickering's (1968) study of Panamanian and West Indian Miagrammopes and Opell's (1979) revision of all Mexican and Central and South American Uloboridae except members of the genus Miagrammopes. Studies by Lubin et al. (1982) and Opell (1981, 1982) broughtthe number ofknown American uloborid genera to 14 and the number of recognized species to 71. The purpose ofthis list is to provide ready access to the names, synonyms, nomenclatural changes, and general distribution of the American Uloboridae. As a revision of Miagram­ mopes sensu lato (Huanacauria, Miagrammopes, Mumaia. Miagramopsidis. and Ranguma, the latter two not represented in the New World) nearing completion will render obsolete any listing of these species, they are only summarized. Included are all other known taxa except the following which, largely due to the unavailability oftype material, are of uncertain status: Uloborus minutus Mello-Leimo, U. tetramaculatus Mello-Leimo, U. formosus Marx (in Banks), U. ater Mello-Leimo, Petrunkevitchia venusta Mello-Leitao, and P. pusilla Mello-Leitao. Genera are arranged according to Opell's (1979) phylogenetic hypothesis. Within each genus, species are arranged alphabetically, the only exception being in where species are listed under three groups. The type species of each genus, if not included in a generic synonym or listed below the genus name, is marked by an asterisk. A colon following a name denotes the first transfer ofthe species to the cited genus. Five genera (Uloborus, Zosis, and Philoponella. Miagrammopes and the closely related Mumaia) and four species (U. campestratus, U. segregatus, Z. geniculatus and P. semiplumosa) are found in North, Central, and South America. Uloborus trilineatus, P. republicana and P. tingena are found in both Central and South America and Mumaia mexican a in both North and Central America. As Figure 1 shows, the greatest number and percent of endemic genera and species are found in South America and the least in North America. Because this evaluation may be biased by the number of named Miagrammopes species, totals exclusive of these species are also provided.

ACKNOWLEDGMENTS

Collections made by William G. Eberhard in Colombia and Costa Rica extended the known ranges of several Central and South American species. Preparation of this list was supported in part by National Science Foundation grant DEB-8011713.

1Department of Biology, Polytechnic Institute and State University, Blacksburg, V A 24061. 62 THE GREAT LAKES ENTOMOLOGIST Vol. 16. No.2

NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA (Canada, Unite<;l (5, Mexico through (South Qf Panama) States, ,~, Mexico) Panama [, I" Indies)

Coni faber, Sybota [IDEMIC ~ENERA Ariston I-k.ianacauria. Orinomana, Ponella

TOTAL NLMlER 6 10 OF GENERA

EI\()(MIC SPECIES 11 (11) 23 (6) 29 (16)

TO TAL NLMlER 16 (15) 31 (12) 36 (22) Of SPECIES

Fig. I. Distribution and number of known American uloborid taxa. Numbers in parentheses inicate totals exclusive of members of the closely related genem Hlianacallria. Miagrammopes. and Mumaia.

ARISTON O. Pickard-Cambridge 1896. 3 species.

*'A. albicans O. Pickard-Cambridge 1896. SE Mexico and Honduras. A. aris/us OpeII 1979. Panama. A. mazolus Opell1979. SW Mexico.

HYPTIOTES Walckenaer 1837. 4 species.

Mithras (preoccupied) C. L. Koch 1834. Type-M. paradoxlls C. Koch 1834. Vptiotes WaIckenaer 1837. Type-V. anceps Walckenaer 1837. Hyptiotes: Erickson 1845, emendation. Cylfopodia Hentz 1847. Type-C. cava/us Hentz 1847.

H. cava/us (Hentz) 1847. E half of U.S. and Canada. H. ger/schi Chamberlin & Ivie 1935. W U.S. and Canada. H. pllebla Muma & Gertsch 1964. SW U.S. H. tehama Muma & Gertsch 1964. Central W U.S.

SIRATOBA Opell 1979. 2 species.

Ads/on (in part) O. Pickard-Cambridge 1896.

*S. referena (Muma & Gertsch) 1964. SW U.S. and N Mexico. S. sira Opell 1979. Central Mexico.

HUANACAURIA Lehtinen 1967. Type-Miagrammopes bambusicola Simon 1892.

Miagrammopes (in part) O. Pickard-Cambridge 1869.

3 species. Brazil (2), Venezuela (I). 1983 THE GREAT LAKES ENTOMOLOGIST 63

MIAGRAMMOPES O. Pickard-Cambridge 1869. Type-M. thwaitesii O. Pickard-Cambridge 1869.

22 species. Antilles (I), Brazil (6), Cuba (2), Guiana (I), Jamaica (3), Panama (7), Puerto Rico (I), Venezuela (I), Virgin Islands (I).

MUMAIA Lehtinen 1967. Type-Miagrammopes corticeus Simon 1892.

Miagrammopes (in part) O. Pickard-Cambridge, 1869.

6 species. Antiles (I), Argentina (I), Mexico (I), Panama (I), Puerto Rico (2), St. Vincent (I), U.S. (I), Venezuela (I), Virgin Islands (I).

SYBOTA Simon 1892. 3 species.

Sylvia (preoccupied) Nicolet 1849. Sybota Simon 1892, replacement name for Sylvia.

S. abdominalis (Nicolet) 1849. Chile. *Sylvia abdominalis Nicolet 1849. Sylvia ater Nicolet 1849. Sylvia similis Nicolet 1849. Sylvia rubiginosa Nicolet 1849. Sylvia vittata Nicolet 1849. Uloborus abdominalis: Simon 1887. Sybota abdominalis: Simon 1892. S. mendozae Opell 1979. Argentina. S. osornis Opell 1979. Chile.

ORINOMANA Strand 1934. 2 species.

Orinomus (preoccupied) Chamberlin 1916. Orinomana Strand 1934, replacement name for Orinomus.

O. bituberculata (Keyserling) 1882. Peru. Uloborus bituberculatus Keyserling 1882. *Orinomus lamprus Chamberlin 1916. Orinomana lampra: Strand 1934. O. mana Opell 1979. Chile.

ULOBORUS Latreille 1806. 8 species.

Uloborus Latreille 1806. Type-Uloborus walckenaerius Latreille 1806. Phillyra Hentz 1850. Type-Phillyra mammeata Hentz 1850. Veleda Blackwall 1859. Type-Veleda lineata Blackwall 1859. Philoponus Thorell 1887. Type-Philoponus pteropus Thorell 1887.

U. campestratus Simon 1893. SE U.S. through N South America. U. cinereus O. Pickard-Cambridge 1898. U. spernax O. Pickard-Cambridge 1898. U. diversus Marx, in Banks 1898. W U.S. and NW Mexico. U. albineus Marx, in Banks 1898. U. californicus Banks 1904. U. utahensis Chamberlin 1919. U. saphes Chamberlin 1924. U. crepedinis Chamberlin 1924. 64 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No.2

U. eberhardi Opel! 1981. Costa Rica. U. e/ongatus Open 1982. Argentina. U. glomosus (Walckenaer) 1847 (1842). E half of U.S. & Canada, N. Mexico. Epeire g/omosa Walckenaer 1837 (1842). Phillyra mammeata Hentz 1850. Phillyra riparia Hentz 1850. Uloboms plumipes: Emerton 1888. Uloborus americanus: Comstock 1912 (1913). Uloborus glomosus: Chamberlin & Ivie 1944. U. metae Opell 1981. E central Colombia. U. segregatus Gertsch 1936. Texas through N South America. U. trilineatus Keyserling 1882. S. Mexico through South America. U. penicillatus Simon 1891. U. aegrotus Simon 1893. U. maniculatus Simon 1893. U. bucki Mello-Leitao 1943 U. plumipes Mello-Leitao 1947. U. plumipedatus Roewer, 1951 replacement name for U. plumipes MelJo-Leitao, preoccupied by U. plumipes Lucas 1846.

ZOSIS Walckenaer 1837. 2 species.

Uloborus (in part) Latreille 1806. Zosis Walckenaer 1837. Type-Zosis caraibe Walckenaer 1837. Orithyia Blackwall 1858. Type-Orithyia williamsii Blackwall 1858.

Z. geniculatus (Olivier) 1789. pantropical. Aranea geniculata Olivier 1789. Zosis caraibe Walckenaer 1837. Uloborus zosis Walckenaer 1841. Uloborus latreillei Thorell 1858. Orithyia williamsii B1ackwall 1858. Uloborus domesticus Doleschall 1859. Orithyia luteola Blackwell, 1865. Uloborus williamsii: O. Pickard-Cambridge 1871. Uloborus geniculatus: Thorell 1890. Ulobarus luteolus: Roewer, 1954. Zosis geniculatus: Lehtinen 1967. Z. peruvianus (Keyserling) 1882. E central Colombia to N central Brazil to N central Argentina. U/obrus peruanus Keyserling 1882. Uloborus peruvianus: Bonnet 1959, emendation. Zosis peruvianus: Opell 1981.

OCTONOBA Opell 1979. 1 species.

Uloborus (in part) Latreille 1806.

*0. octonaria (Muma) 1945. Central and SE U.S. Uloborus octonarius Muma 1945. Octonoba octonaria: OpeU 1979.

CONIFABER Opel!, in Lubin et al. 1982. 1 species.

"Conifaber pan'us Opell, in Lubin et al. 1982. E central Colombia. 1983 THE GREAT LAKES ENTOMOLOGIST 65

PONELLA Opel! 1979. 1 species.

Uloborus (in part) Latreille 1806.

*P. lactescena (Mel!o-Leitao). 1947. Paraguay and S. BraziL Uloborus lactescens Mello-Leitao 1947. Panella lactescena: OpeU 1979.

PHILOPONELLA Mel!o·Leitao 1917. 15 species.

Uloborus (in part) Latreille 1806. Philopanella Mel!o·Leitao 1917.

Philoponella republicana species group Opell 1979.

P. divisa Opell 1979. N South America. *P. republicana (Simon) 1891. Panama & South America. Uloborus republicana Simon, 1891. Uloborus cuminamensis Mello·Leitiio 1930. Uloborus mundior Chamberlin & Ivie 1936. Zosis mundior: Lehtinen 1967. Philopenella republicana: Opell 1979. P. signatella (Roewer) 1951. SE Mexico through Honduras. Uloboms signatus O. Pickard·Cambridge 1898. Uloborus signatellus Roewer 1951, replacement name for preoccupied U. signatus O. Pickard·Cambridge 1898. Philoponella signatella: Opel! 1979. P. tingena (Chamberlin & Ivie) 1936. Costa Rica through Colombia. Uloborus tingnes Chamberlin & Ivie 1936. Philoponella tingena: Opell 1979. P. viGina (0. Pickard-Cambridge) 1898. S Mexico through Central America. Uloborus vicinus O. Pickard·Cambridge 1898. Philoponella vicina: Opell 1979.

Philoponella semiplumosa species group Opel! 1979.

P. arizonica (Gertsch) 1936. . Uloborus arizonicus Gertsch 1936. Philoponella arizonica: Opell 1979. P. oweni (Chamberlin) 1924. SW U.S. Uloboms oweni Chamberlin 1924. Philoponella oweni: Opell 1979. P. semiplumosa (Simon) 1893. Texas through N South America. Uloborus semiplumosus Simon 1893. Uloborus variegatus O. Pickard·Cambridge 1898. Uloborus abstrusus Gertsch & Davis 1942. Philoponella semiplumosa: OpeU 1979. P. subvittata Opell 1981. Guyana. P. viltala (Keyserling) 1882. South America. Uloborus vittatus Keyserling 1882. Uloborus servulus Simon 1892. Uloborus semiargenteus Simon 1893. Uloborus amazonicus Mello-Leitao 1949. Philoponella vittata: Opell 1979. 66 THE GREAT LAKES ENTOMOLOGIST Vol. 16, No, 2

Philoponella Jasciata species group Opel! 1979.

P, bella Opell 1979. NE Colombia. P. collina (Keyserling) 1882. Peru. Uloborus collin us Keyserling 1882. Philoponella collina: Opel! 1981. P. Jasciata Mello-Leitao 1917. SE Brazil & Paraguay. Uloborus Jasciatus Mello-Leimo 1917. Philoponella Jasciata: Opel! 1979. P. para Opell 1979. Paraguay.

LITERATURE CITED

Chickering, A. M. 1968. The genus Miagrammopes (Araneae, Uloboridae) in Panama and the West Indies. Breviora 289:1-28. Lehtinen, P. T. 1%7. Classification of the cribellate spiders and some allied families, with notes on the evolution of the suborder Araneomorpha. Ann. Zool., Fennici 4: 199-468. Lubin, Y. D., B. D. Open, W. G. Eberhard, and H. W. Levi. 1982. Orb plus conewebs in Uloboridae (Araneae), with a description of a new genus and four new species. Psyche 89:29-M. Muma, M. M. and W. 1. Gertsch. 1964. The family Uloboridae in North America north of Mexico. Amer. Mus. Novitates 21%:1-43. Opell, B. D. 1979. Revision of the genera and tropical American species of the spider family Uloboridae. Bull. Mus. Compo Zool. 148:443-549. ----. 1981. New Central and South American Uloboridae (Arachnida, Araneac). Bull. Amer. Mus. Nat. Hist., 170:219-228. ----. 1982. A new Uloborus Latreille species from Argentina (Arachnida: Araneae: Uloboridae). Proc. BioI. Soc. Washington, 95:552-554. INFORMATION FOR AUTHORS

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