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PSYCHE

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George Salt.
Notes on the Strepsiptera and Their Hymenopterous Hosts.
Psyche 34(5):182-192, 1927.

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182 Psyche [October
NOTES ON THE STREPSIPTERA AND THEIR HYME- NOPTEROUS HOSTS.1
In the course of a study of the effects of stylopization on aculeate Hymenoptera, a number of new and interesting host records and various notes on parasitism by Strepsiptera have been accumulated.
Although their varied nature render it some- what disjointedJ it seems wise to bring these miscellaneous facts together in the present paper.
The following is an annoted list of the eighty-one species of Hymenoptera of which I have examined stylopized specimens. 'Two genera, Zethus and Monte~urnia~ and several species ap- parently constitute new host records.
Sphecid~.
Ammophila
abbreviata (Fabr.)? det. Banks. Virginia: Falls Church (N. Banks).
alberti Hald,, det. Banks.
Arizona: Palmerlee, Tucson (F.
H. Snow).
conij'era Arnold, det. Arnold. Belgian Congo : Kabare (Jos. Bequaert).
pictipennis Walsh and Riley, det. Fernald. Virginia: Falh Church and Great Falls (N. Banks).
vulgaris Cress., det. Robertson. Illinois: Carlinville (Chas. Robertson) ; Florida: Inverness (Chas. Robertson). Chlorion
(Isodontia)
aut-ipes Fern., det. Banks. Texas: Dallas (J. Boll). harri6 Fern.) det. Fox in Coll. M, C. 2. Connecticut:
Stamford (A. P. Morse).
(Proterosphex)
ichneumoneum (Linn.) det. Salt. Illinois: Carlinville (Chas. Robertson) ; Texas : Fedor.
pennsylvanicum (Linn.), det. Fernald. North Carolina : Southern Pines (H. Manee).
1Contributions from the Entomological Laboratory of the Bussey Insti- tution, Harvard University, No. 270.




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Notes on the Strepsiptera and their Hosts 183
Vespid~.
' Zethin~.
Zethus
pubescens Smith, det. Bequaert.
Zululand: Mfongosi (W. E*
Jones).
spinipes var. variegatus Sauss. ) det . Bequaert.. Texa~ : Dd- las (J. Boll).
Eumeninz
Eumenes
carnpaniformis var. marshalli Bequaert) det. Bequaert) Para- type. "Dr. Melle) Arcturus. 1916) Salisbury.') jraternus Say) det. Bequaert. Massachusetts: Forest Hills (George Salt).
maxillosus var. circinnalis Fabr. ) det . Bequaert. Moluccas : Amboine (Rouyer) .
maxiZZosus var. reginus Sauss., det. Bequaert. "Diego) Suarez.))
Montezwnia
sp. near brunnea Sm., det. Bequaert. Argentina: Chaco de Santiago (Del Estero) Rio Salado).
injernalis (Spin.)) det. Bequaert. Paraguay : Villarrica. Odynerus
annulatus Say) det. Bequaert. Texas: Austin) New Braun- fels; Dakota: Custer Co. (S. W. Garman). anormis Say, det. Robertson. Illinois: Carlinville (Chas. Robertson).
awensis Sauss. ) det.. Robertson. Illinois : Carlinville, (Chas. Robertson),
beZZulus Cress.) det. Bequaert. Texas: Lee Co. (Birkman). blandus Sauss.) det. Bequaert. California: Stanford Univ. bradleyi Robt.) det. Robertson. No locality label. foraminatus Sauss., det. Bequaert. Massachusetts : Woods Hole; Illinois: Carlinville (Chas. Robertson), Chicago; Pennsylvania : Philadelphia; Texas : Dallas (J. Boll), F'edor; Ohio : West Jefferson (Jos. Bequaert) ; Virginia : Vienna) Dyke) and Falls Church (N. Banks). fundatus Cress., det. Bequaert . Texas : Austin ; Illinois : Carlinville (Chas. Robertson).




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184 Psyche [October
histrio Lepel.) det. Robertson.
Florida: Inverness (Chas.
Robertson).
histrionalis Robt., det. Robertson. No locality label. manifestus Cress. ) det. Bequaert. Texas : (Fedor). perennis Sauss., det. Bequaert. Virginia: Great Falls) Falls Church and Dyke (N. Banks).
pratensis Sauss., det. Bequaert. Texas : El Paso. toas Cress.) det. Bequaert. ex as : Valentine, El Paso; New Mexico : Steins.
Pachodynerus
cubensis (Sauss.), det. Bequaert. Cuba: Soledad (Jos, Bequaert).
erynnis (Lepel.)) det. Bequaert.
Florida: Lakeland (Wm.
T. Davis)) Miami (Jos. Bequaert), Inverness (Chas. Robertson).
guadulpensis (Sauss.) , det. Bequaert . Grenada : Santeurs (Allen and Brues).
nasidens (Latr.)) det. Bequaert. Rep. Honduras: Pto. Cas- tilla (Jos. Bequaert) .
Ancistrocerus
albophaleratus (Sauss.)) det. Bequaert. Connecticut: Ca- naan (Jos. Bequaert) ; New Hampshire : Mount Wash- ington (Halfway House) (C. W. Johnson) ; Maine; Alberta: Bilby (George Salt).
birenimaculatus (Sauss.) ) det . Bequaert . New York : Flush- ing (Jos. Bequaert) ; New Jersey : Newfoundland. catskillensis (Sauss.) ) det. Bequaert. Massachusetts : Forest Hills (L. H. Taylor), Auburndale (C. W. Johnson); New York: Keene Valley (El. Notman) ; Connecticut : Colebrook (W. M. Wheeler).
julvipes (Sauss.) , det . Bequaert . Texas : Lee Co. (Birkman) ) New Braunfels.
parietum (Linn.), det. Bequaert.
New York: Ithaca.
digris (Sauss.)) det. Bequaert .
New Jersey : (Ramsey) , Fort
Lee (Jos. Bequaert) ; Virginia: Falls Church and Glen- carlyn (N. Banks) ; Pennsylvania: Melsh ; Connecticut : Colebrook ((W. M. Wheeler).




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19271 Notes on the Strepsiptera and their Hosts 185 waldenii (Vier.) det. Bequaert. Massachusetts : Forest Hills (0. E. Plath).
Belonogaster
filiventris (Sauss.) det. Bequaert. Belgian Congo : La Pan- da (Katanga) (Dr. Mich. Bequaert) .
junceus (Fabr.) , det . Bequaert . Victoria Nyanza : Masinga Archipelago (Dr. J. Carl).
Ro palidia
cincta (Lepel. ) det . Bequaert.
Belgian Congo : Gani (Li-
salto, Monveda) (Jos. Bequaert).
nobilis (Gerst.) det. Bequaert. Congo : Lisala. Polistes
canadensis var. annularis (Linn.), det. Bequaert. West Vir- ginia: Mongolia Co. (L. H. Taylor).
crinitus (Felton) ) det. Bequaert. Rep. Dominica : Sanchez (A. M. N. H.).
crinitus var. arnericanus (Fabr .)
det . Bequaert . Porto Rico :
Aibonito, Cayey) Coamo Springs7 Jayuya, Manati, and Mayaguey.
gallicus (Linn.)> deb. Bequaert. No locality label. pallipes Lepel., det. Bequaert. Connecticut : Colebrook, Robertsville, and Chapenville (W. M. Wheeler). mriatus Cress. det . Robertson. Illinois : Carlinville (Chas, Robertson).
Chloralictus
sp. Washington : Pullman (A. D.)
sp. (N. B. Several species are probably here grouped to- gether). Connecticut : Colebrook (W. M. Wheeler) ; Massachusetts : Forest Hills (George Salt). sparsus Robte7 det. Robertson.
Illinois: Carlinville (Chas.
Robertson).
versatus Robt., det. Robertson.
Illinois: Carlinville (Chas.
Robertson).
zephyrus Sm., det. Robertson.
Illinois : Carlinville (Chas.
Robertson).




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186 Psyche [October
Andrena
andrenoides Cress., det. Robertson. Illinois: Carlinville (Chas. Robertson).
asteris Robt,. det . Robertson. Illinois : Carlinville (Chas. Robertson).
bradteyi Vier.? det. Salt. Massachusetts : Forest Hills (George Salt).
canadensis D. T., det. Viereck. Connecticut: Colebrook (W. M. Wheeler).
carolina Vier,? det, Viereck.
Massachusetts: Billerica (P.
J. Darlington Jr.).
cressoni Robt., det. Viereck.
Connecticut: Colebrook (W.
M. Wheeler).
erigenia? Robt., det. Robertson. Illinois : Carlinville (Chas. Robertson).
jorbesi Robt., det. Viereck.
Connecticut: Colebrook (W.
M . Wheeler).
jragilis Sm.? det. Viereck. New York: Oliverea (Catskills) (Jos. Bequaert) .
hippotes Robt., det. Robertson. Illinois : Carlinville (Chas. Robertson).
hirticincta Prov.? det. Salt, Connecticut : Colebrook, Rob- ertsville, and Doolittle Pond (Lichfield Co.) (W. M. Wheeler) ; Vermont : Chittenden (Rutland). itlinoiensis Robt., det. Robertson. Illinois : Carlinville (Chas. Robertson).
irnithtrix var. claytoni~ Robt., det. Robertson. Illinois: Carlinville (Chas. Robertson).
rnandibularis Robt. det. Robertson. Illinois : Carlinville (Chas. Robertson).
miranda Sm., det. Viereck. Connecticut: Colebrook (W. M. Wheeler) ; Wisconsin.
nasoni Robt., det. Viereck. Massachusetts : Forest Hills (L. H. Taylor, George Salt).
nuda Robt., det. Robertson. Illinois : Carlinville (Chas. Robertson).
salictaria Robt,, det . Robertson. Illinois : Carlinville (Chas. Robertson).




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19271 Notes on the Stripsiptera and their Hosts 187 solidaginis Robt., det. Viereck. Nebraska: Malcolm (C. R. Oertel); New Jersey: Alpine and Fort Lee (Jos. Be- quaert) .
vicina Sm., det. Viereck. Massachusetts: Petersham (C. T. Brues) .
Panurginus
innuptus Ckll., det. Steven-s. North Dakota: Minot (0. A. Stevens).
Pseudo panurgus
cethiops Cress., Colorado: Wray.
rudbeckice Robt., det. Robertson. Illinois: Carlinville (Chas. Robert son).
solidaginis Robt ., det. Robertson. Illinois : Carlinville (Chas. Robertson).
Various writers have given data on the extent of parasitism by Strepsiptera, and in his monograph and its supplements, Pierce (1909, 1911, 1918) has brought together these records and added others. Most of the published notes, however, deal with Polistes and Andrena, whereas the following have to do largely with other genera.
In 1922, Dr. W. M. Wheeler collected a total of 373 speci- mens of Andrena hirticincta Prov, Forty-four of these or 11 .9% were stylopized, 6 of 23 males or 26.1%, and 38 of 350 females or 10.9%. His collecting records for the same year of species of Chloralictus, mostly albipennis, show 98 stylopized of a total of 470 specimens collected, an infestation of 20.9%; 88 of 378 males or 23.3% were stylopized, and 10 of 92 females or 10.9% No collecting records are available for the Vespidse other than Polistes but some indication of the extent to which they suffer from strepsipterous attack may be obtained from the fre- quency of the occurrence of stylopized individuals in collections. Of 4525 specimens of miscellaneous Eumeninse examined by the writer, 74, or 1.64%, were stylopized. Collections of particular genera contained stylopized specimens as follows: Masarinse of 96 specimens, 0 stylopized, or 0 % . Zethus of 53 " 2 i' ' 3.8%.
Eurnenes of 214 " 3 i b - - ' 1.4 %.




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188 Psyche [October
Nortonia
Pachymenes
Monobia
A lastor
Pterocheilus
Stenogaster
Belonogaster
Ro palidia
In addition, 217 specimens of Synagris were examined rather summarily but none was observed to be stylopized. To give detail of the sex, number, and position of the para- sites in the various species of hosts would require more space than is at our disposal, but the general conditions are indicated by the following analyses of the parasitism in four of the better- known genera.
Odynerus (sens. lat.) .
51 specimens (29 d, 22 Q ) carrying I parasite (15 3, 35 Q ,I?)
13 '
(63, 79)
" 2parasites (53,219).
3 "
( 39)
3 ' (38, 69).
2 ' ( 28)
(' 4 '
(33, 59).
1 ' (18)
" 6 L6
(23, 49).
Average infestation :-I .43 parasites per host. 99 parasites (28 8, 7 1 Q ) situated as follows :-42 on the right, 56 on the left, 1 median ;
under the 2nd. 3rd. 4th. 5th. abdominal segments. 8 parasites 1 21 6
9 parasites 6 45 19 1
In this genus of hosts the female parasites prefer a position anterior to that of the male.




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19271 Notes on the Strepsiptera and their Hosts 189 29 specimens (3 a", 26 Q )carrying 1
parasite
11 (1 d\ 10 Q ) 2 d" parasites
2 " (l&lQ)." 29 14
6 " ( 69) " lc?,lQ ::
' 307
:' ;:{ :: 23,IQ 1:
1c?,2Q "
" ( <' 40n
1 " (
2 " ( 2Q) " 53

1 " ( 19) " 7 3
(6
3 " ( 3 Q) I' larval "
-
65 " (53,609) " 118..larval"
Average infestation of wasps carrying adult parasites :-I. 9 para- sites per wasp.
118 parasites (943, 24 9 ) situated as follows:-54 on the right, 63 on the left, 1 median; 105 dorsal, 3 lateral, 10 ventral; under the 2nd. 3rd. 4th. 5th. abdominal segments. 3 parasites 1 44 41 8
Q parasites 1 2 21
The normal position for the male is here obviously under the 3rd. and 4th. segments. In the eight cases in which a male was found under the 5th. segment, only once was it the only parasite; twice it was one of two, twice one of three, once one of four, and twice one of five parasites, and may be considered to have been crowded from its normal position. On the other hand, the female usually occurs under the 5th. segment; where it was under the 3rd. it was one of three parasites, where it was under the fourth it was one of two or one of three parasites. This con- dition, in which the males normally lie anterior to the females in the body of the host, is exactly the opposite of that in Odynerus where the females lie anteriorly, the males behind. Chloralictus
313 specimens (276 3, 37 9 ) carrying I parasite. 161 " (1423, 19 9) " 2 parasites.
45 " (446 19) " 3

1 " (13)
11
4
6 1
-
-
520 " (4633,579) 774 LL
- --
Average infestation:-1 .5 parasites per host.



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190 Psyche [~cto ber
774 parasites, all females, situated as follows-386 on the right, 358 on the left, 30 median; 763 dorsal, 4 lateral, 7 ventral; 40 under the 3rd., 712 under the 4th., and 22 under the 5th. ab- dominal segments.
In seven of the eleven cabes in which the parasite was lateral or ventral, it was one of three parasites; in three cases, one of two; and in only one case was it the only parasite. The normal position of the parasite in Chloralictus, therefore, is dorsal, under the 4th. abdominal segment. It will be noticed that here, as also in the case of the sty- '
lopized Halicti studied by Perkins (1892, 1918), all of the para- sites found were females; no males or male exuvia were seen in Halictus. This fact at once raises the question of parthenogenesis among the females of Halictoxenos already suggested by Perkins. Mrs. Schrader (1924), to be sure, has shown that the females of Acroschismus must be fertilized to produce their young; but in that genus males are of common occurrence and, indeed, in the present study were found in much larger numbers than the females. Pierce (1918) claims to be the only one who has ever captured an Halictus with a male parasite. In view of the very rare occurrence of males of Halictoxenos, then, or their almost complete absence, parthenogenesis is strongly suggested in this group notwithstanding Mrs. Schrader's contrary observations on A croschismus.
Andrena.
147specimens (41 c(\ 106 Q )carrying 1
parasite (40 c? 107 Q ).
18 ( 4 6 149) 2 Q parasites ( 36 9 ).
3 " ( ' 23 ' (63 1.
3 " ( 3Q)
1 " ( 19) " 39 ( 39).
2 " ( 2 9) " 1 larval parasite
-
- --
174 " (45 d\ 129 Q ) " 200 parasites (49 3, 149 9 , 2 larval). -
- -----
Average infestation:-1 .1 parasites per host. 200 parasites (493, 149 9 , 2 larval) situated as follows:-97 on the right, 93 on the left, 1 median, 3 une~t~ruded, 6 unrecorded; 187 under the 4th.) 9 under the 3rd., 1 under the 5th. abdominal segments, 3 unextruded. The normal position for the parasites of Andrena is obviously under the 4th. abdominal segment. '




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19271 Notes on the Strepsiptera and their Hosts 191 Finally, attent,ion must be drawn to a matter of nomen- clature. It appears to have been overlooked that von Heyden (1868, p. 398) used the name Xenos srnithii for two male para- sites of Sphex ichneumoneus L. from North America. Pierce (1909, p. 185) notes the specimens but apparently does not recognize the description. There can be no doubt of the validity of the name, however, for not only is the insect partly described, but also the name of tlhe host, the essential part of many a more recent description of a strepsipterous parasite, is given. Homi- lops bishoppi Pierce (1909, p. 146)) parasitic on Sphex ichneu- rnoneus L. at Waco, Texas, is known only in the female sex, while Xenos srnithii was described by von Heyden from two males only. It is therefore possible to consider the former the female of the latter, especially as it is a common practice to couple male and female strepsiptera simply by their infestation of a single species of host.
For loan of specimens I am very grateful to Mr. N. Banks of the Museum of Comparative Zoology, Mr. C. W. Johnson of the Boston Society of Natural History, Mr. Chas. Robertson of Carlinville, Illinois, Prof. W. M. Wheeler and Prof. C. T. Brues of the Bussey Institution, Harvard University, and Prof. Joseph Bequaert of the Harvard Medical School. My thanks are due to the various authorities to whom the host determinations are individually credited. To Dr. Joseph Bequaert I am especially grateful, not only for determination of the Vespidse and for the gift of specimens, but also for generously allowing me to search for stylopized individuals in his large and valuable collection. References.
Heyden, L. von 1868, Exotische Xenos-Arten. Berliner Entom. Zeitschr., 11, 398.
Perkins, R. C.L. 1892, Stylopized bees. Ent. Mon. Mag., 28, 1-4. 1918, Further notes on Stylops and stylopized bees. Ent. Mon. Mag., 54, 115-129.
Pierce, W. D. 1909, Monographic revision of the twisted winged insects comprising the order
Strepsiptera Kirby. Bull. U. S. Nut.
Mus., 66, pp. xii and 232, 15 pis.




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Psyche [October
1911, Notes on insects of the order Strepsiptera, with descriptions of new species. Proc.
U. S. Nut. Mus., 40, 487-511.
1918, The comparative morphology of the
order Strepsiptera together with records and descriptions of insects. Proc. U. S. Nut. Mus., 54, 391-501, pis. 64-78.
Schrader, S. H. 1924, Reproduction in Acroschismus wheeler2 Pierce. Journ. Morph. Physiol., 39, 157- 197, 4 pis.




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