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JANUARY & FEBRUARY, 1993

USISSN0013-872X
NO. 1

ENTOMOLOGICAL NEWS

On Thomas Say's entomological
publications printed in
New Harmony, Indiana

The phylogenetic position of
Chloroniella peringueyi
(Megaloptera: Corydalidae)
and its zoological significance

Yves Bousquet 1

Norman D. Penny 17

A new species of Polycentropus
(Trichoptera: Polycentropodidae)
from Arkansas

D.E. Bowles, M.L. Mathis, S.W. Hamilton

A new species in the Polycentropus
cinereus group (Trichoptera:
Polycentropodidae) from
Arkansas and Texas S.R. Moulton, II, K.W. Stewart

Interactions of predaceous katydids
(Orthoptera: Tettigoniidae) with
neotropical social wasps
(Hymenoptera: Vespidae):
are wasps a defense
mechanism or prey?

31

35

Sean O'Donnell 39

Records of bat flies from Jordan,
Libya, and Algeria

A chamber for mass hatching
and early rearing of
praying mantids (Orthoptera:
Mantidae)

Elmidae of Taiwan, Part II:
Redescription of Leptelmis
formosana (Coleoptera:
Dryopoidea)

Z.S. Amr, M.B. Qumsiyeh 43

F.R. Prete, RJ. Mahaffey 47

BOOK REVIEWS

SOCIETY MEETING OF OCTOBER 28, 1992

SOCIETY MEETING OF NOVEMBER 18, 1992

M.-L. Jeng, P.-S. Yang 53

15, 16,60

34
46

THE AMERICAN ENTOMOLOGICAL SOCIETY

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Vol. 104. No. 1. January & February, 1993

ON THOMAS SAY'S ENTOMOLOGICAL
PUBLICATIONS PRINTED IN
NEW HARMONY, INDIANA 1

Yves Bousquet^

ABSTRACT: All entomological papers published by Thomas Say and printed in New
Harmony, Indiana, are briefly commented upon. The dates and citations of all new
taxa and new replacement names of Coleoptera proposed in these papers are presented in
a table. Four of Say's names, previously considered as junior synonyms, have priority:
Chlaenius circumcinctus Say, 1830. in place ofC. perplexus Dejean, 1831; Chlae niux soccatus
Say, 1 830, in place of C. melanarius Dejean, 1 83 1 ; Dytiscus confluens Say, 1 830, in place of D.
dauricus Gebler. 1832; Tenomerga cinerea (Say, 1831), in place of T. concolor (Westwood.
1835).

From January 1826 to October 10, 1834, the date of his death, Thomas
Say lived in New Harmony, a small community in southwestern Indiana
located along the Wabash River. Say published most of his entomologi-
cal and conchological observations while in New Harmony first in the
journal 'The Disseminator of useful knowledge" which became "The
Disseminator" on June 29, 1830. The journal was suspended on June 26.
1831 and reappeared only in 1834. During that interval. Say published
his descriptions in several pamphlets, printed in New Harmony, which
have become extremely rare. Some of these pamphlets were reprinted,
with minor editorial changes, in scientific journals and this action has
created confusion over the dates of many new species described by Say.

The purpose of this publication is to briefly comment on each of
Say's entomological papers published in New Harmony and give the
proper publication dates of all new taxa and new replacement names of
beetles published in these papers.

1. Say, T. 1830. Correspondence relative to the insect that destroys
the cotton plant. The Disseminator of useful knowlege; contain-
ing hints to the youth of the United States from the "School of
industry" 3: 19-21.

The journal, usually abbreviated as 'The Disseminator of useful
knowledge" was a semi-monthly publication, printed in octavo form.
The first issue appeared on January 1 6, 1 828 and the last one (vol. 3, no. 9)
on May 12, 1830. Three volumes were published (not two as noted in

1 Received March 19. 1992. Accepted May 18. 1992

2 Centre for Land and Biological Resources Research. Agriculture Canada. Ottawa,
Ontario, KIA OC6

ENT. NEWS 104(1): 1-14. January & February. 1993

ENTOMOLOGICAL NEWS

Titus, 1965): the first in 1828, the second in 1829, and the third, which was
not completed, in 1830. The third volume is extremely rare and the only
copy I am aware of is in The Public Library of Cincinnati & Hamilton
County, Department of Rare Books and Special Collections (Ian
MacPhail, personal communication).

This paper consists of a letter, dated January 1 827, from C.W. Capers,
University of Pennsylvania, to Say and Say's response, dated November

1. 1827, in which he describes the moth Noctua xylina (= Alabama
argillacea Hiibner, 1823). The paper was first published in 1828 in the
"Southern Agriculturist" (volume 1, page 203) (Horn and Schenkling
1928) and subsequently reproduced with minor changes by A. Fitch in
1858 in the Transactions of the N.Y. State Agricultural Society (volume
17, pages 813-814). The version reproduced by J.L. LeConte (in Say
18590: 369-371) is that published by Fitch.

9

2. Say, T. 1830. Descriptions of new species of North American
insects, and observations on some already described. The Dis-
seminator of useful knowledge; containing hints to the youth of
the United States from the "School of industry" 3: 67-69,
133-135.

This paper appeared in number 5 (March 17 issue, pages 67-69) and
number 9 May 12 issue, pages 133-135). It contains descriptions of some
taxa of Carabidae, including cicindelids, and observations on others.
The content of this paper was reprinted in Say (1830c) and (1 834/7).

3. Say, T. 1830. Descriptions of new species of North American
insects and observations on some already described. The Dis-
seminator 1(1):[3]; 1(3):[3]; 1(4):[3]; 1(5):[3]; 1(6):[3]; 1(7):[3].

"The Disseminator" continued "The Disseminator of useful
knowledge". It was published weekly, in folio, numbered separately, but
with the pages unnumbered. The first volume was published from June
29, 1830(number l)toJune25, 1831 (number52).Thejournal reappeared
on June 1 834, under the same title but as a new series, and ceased in April
1841. Seven volumes of the new series were published (Titus 1965).

The paper contains the descriptions of several species of Cara-
bidae which appeared on the third pages of numbers 1 and 3-7. Its con-
tent was reprinted in Say (1830c) and (1834/?).

4. Say, T. 1830-1834. Descriptions of new species of North
American insects, and observations on some of the species
already described. School Press, New Harmony [Indiana]. 81
pages (numbered [l]-73, 73 1/2-80).

Vol. 104, No. 1, January & February, 1993

The first 17 pages of this pamphlet, which are not numbered, are
unaltered reprints of Say's papers published in the third volume of "The
Disseminator of useful knowledge" (pages [1-4] in double columns) and
the first volume of "The Disseminator" (pages [5-17] in single columns).
The next pages are numbered 18 to 65 with an unnumbered blank page
before page 1 8, two unnumbered blank pages before pages 50 and 58, and
one unnumbered blank page before page 66 (numbered 46). The next
eight pages are incorrectly numbered 46 to 53 (instead of 66 to 73) and the
last eight pages are numbered 73 1/2 to 80.

The cover page of the pamphlet bears the date 1829-1833 and at the
bottom of page 1 8 is printed "August 20, 1 830." There are no other dates
indicated on the pamphlet. The publication dates of the pages, however,
were discussed by T.W. Harris in a letter addressed to J.L. LeConte dated
Nov. 29, 1852: "The last eight pages [of this pamphlet] were printed after
the title page, and only a short time before his [Thomas Say] last sickness
and death. About one half of the whole work, ending with the description
of Lathrobium dimidiatum, was actually printed and distributed in 1830.
Thence to Anthophagus verticalis inclusive, in 1831; thence loAleochara
semicarinata inclusive, in 1832; . . . Thence \oAgriluspolitus, in 1833; and
the remainder, to Elater exstriatus inclusive, in 1834" (Harris 1869: 222-
223). Therefore, pages [1]-41 were published in 1830 (with pages 18-41
likely printed on August 20), pages 42-49 in 1831, pages 50-57 in 1832,
pages 58-73 in 1833, and pages 73 1/2-80 in 1834 (before October 10). The
dates indicated on the cover page are incorrect and should have read
1830-1834.

I knowof five copies of this pamphlet. The copy in the Library of New
Harmony Workingmen's Institute contains only the pages 18-65
(Rosemary Alsop, personal communication). The Library of Congress
has a copy in its Rare Books Section which has the printed pages 18-65,
and the first 17 pages and a few of the remaining pages in a handwritten
form. The American Museum of Natural History Library possesses the
copy of S.H. Scudder which includes pages [l]-73. The copy was pre-
viously owned by J.L. LeConte and includes a few of his manuscript
notes on margins of pages [information written on front cover and
copied from Sherman Catalogue no. 6]. The American Museum of
Natural History Library also has a photostat of the pages 73 1/2-80
"made from Harvard University copy March 1948". The Houghton
Library, Harvard University, holds two copies of this pamphlet. One
copy has only the pages 1 8-65, and the second copy is complete. Byrd and
Peckham (1955) reported the presence of additional, incomplete copies
of this pamphlet in the libraries of the Indiana University, the University
of Minnesota, the Case Western Reserve University, and the Academy of
Natural Sciences.

ENTOMOLOGICAL NEWS

Pages [l]-57 of the pamphlet were republished with minor editorial
changes in 1834 in the New Series of the Transactions of the American
Philosophical Society (volume 4, pages 409-470) and pages 58-80 were
republished with minor editorial changes and scientific comments of
T.W. Harris in 1839 in the same journal (volume 6, pages 155-190). The
last 14 pages of the 1839 publication contain descriptions and notes not
previously published and were likely taken from manuscripts sent after
Say's death by his wife, Lucy W. Say, to T.W. Harris (Weiss and Ziegler
1931). The version reproduced by J.L. LeConte (in Say 18596: 521-629) is
that of the Transactions of the American Philosophical Society.

Carus and Engelmann (1861) correctly recorded the dates (1830-34)
and the number of pages (8 1 pages) of this pamphlet. Hagen ( 1 863) listed
it as published in 1829-1833 and containing 65 pages. Horn and Schen-
kling (1928) catalogued the pamphlet as "Disseminator of useful
knowledge. New Harmony, 1829-33, sep. p. 1-81". Summers (1982) re-
ported it as printed in 1834 and "included material originally published
in the Disseminator". I am not aware of any taxonomic works citing this
pamphlet except Blackwelder (1952) who listed it in the "Bibliography"
section as published in 1 830. The reproduction in the Transactions of the
American Philosophical Society of 1834 and 1839 is the reference cited
by most taxonomists and the publication dates of that journal (i.e. 1834,
1839) are those which are, erroneously, attached to Say's scientific
names.

5. Say, T. 1831. Descriptions of new species of North American
insects, found in Louisiana by Joseph Barabino. School Press,
New Harmony [Indiana]. 17 pp.

This pamphlet, published in March 1831, as indicated on the title
page, had 17 printed pages which were numbered 3-19. Scudder (1899)
called attention to this work which had been omitted by J.L. LeConte (in
Say 1859fl, b). As discussed by Bequaert (1950), Horn and Schenkling
(1928) incorrectly cited this pamphlet and the one published in January
1832 (item #8) as the same publication with different titles.

The publication contains the description of 22 species, 12 Coleop-
tera, four Hemiptera, five Hymenoptera, and one Diptera. All but two of
the descriptions were already published (Say 18306 and/or 1830<r), or
were republished later (Say 183 Id, 1835, 1836, or 1837).

6. Say, T. 1831. Descriptions of new species of curculionites of
North America, with observations on some of the species already
known. School Press, New Harmony [Indiana]. 30 pp.

Vol. 104. No. 1, January & February, 1993

This publication was published in July 1831, as indicated on the
front page. The title reported here appears on the front page, but the
second page bears another title "Descriptions of North American Cur-
culionides & an arrangement of some of our known species agreeably to
the method of Schoenherr". Pages 25-30 constitute the "supplement" and
could have been published later, possibly in 1832 (Anonymous 1861).
However, because there is no tangible evidence supporting that inter-
pretation, the year indicated on the pamphlet (i.e. 1831) should be ac-
cepted for nomenclatural purposes. The paper was republished, under
the title on the second page, with short comments by J.L. LeConte (in Say
1859a: 259-299).

7. Say, T. 1831. Descriptions of new species of heteropterous
Hemiptera of North America. School Press, New Harmony
[Indiana]. 39 pp.

This pamphlet was published in December 1831, as indicated on the
cover page. It was reproduced by A. Fitch, in 1 858, in the Transactions of
the N.Y. State Agricultural Society (volume 17, pages 755-812). Fitch
(1858) reported that although the title page bears the date 1831, only the
first four pages were printed that year, because Say mentioned, on page 5,
his pamphlet of January 1832 (item #8). However, since both pamph-
lets appeared nearly simultaneously (December 1831 and January 1832),
it is likely that both were printed concurrently. Say, who was involved in
the printing process (Weiss and Ziegler 1 93 1 ), had access to the plates of
the January 1832 pamphlet and could have modified the plates of the
December 1831 pamphlet. Obviously, it is also possible that the cover
date is incorrect but, in my opinion, it is unlikely that some pages were
published in 1831 and others later. Because of the uncertainty concern-
ing the publishing date, I believe that the year on the cover page (i.e. 1 83 1 )
should be accepted for nomenclatural purposes. The version rep-
roduced by J.L. LeConte (in Say 1859a: 310-368) is that published in the
Transactions of the N.Y. State Agricultural Society.

8. Say, T. 1832. New species of North American insects, found by
Joseph Barabino, chiefly in Louisiana. School Press, New Har-
mony [Indiana]. 16 pp.

This publication was published in January 1832, as indicated on the
cover page. It contains the descriptions of 19 species, seven Coleoptera,
nine Hemiptera, one Hymenoptera, and two Diptera. The descriptions
of some Hemiptera species were partly reprinted from the December

ENTOMOLOGICAL NEWS

1831 pamphlet (item #7). The paper was republished with short com-
ments by J.L. LeConte (in Say 1859a: 300-309).

Because of the scarcity of the New Harmony pamphlets and the sub-
sequent reproductions of some of them, the dates associated with many
species described by Say are often given incorrectly. This is mainly true
for species of Coleoptera. Table 1 lists all new taxa and new replacement
names of beetles proposed by Say and published in New Harmony, with
their references (including their subsequent reproductions, with the
exception of Say [18590, ,]) and valid names. By employing the correct
dates, four of Say's names, previously considered as junior synonyms,
have priority. They are: Chlaenius circumcinctus Say, 1830 [not 1834] in
place of Chlaenius perplexus Dejean, 1831; Chlaenius soccatus Say, 1830
[not 1834] in place of Chlaenius melanarius Dejean, 1831; Dytiscus con-
JluensSay, 1830, [not 1834] in place of Dytiscus dauricusGebler, 1832; and
Tenomerga cinerea (Say, 1831) [not 1834] in place ofTenomerga concolor
(Westwood, 1835) [not 1830].

Table 1. Alphabetical list of new taxa and new replacement names of Coleoptera pro-
posed by Say and printed in New Harmony, with references and valid names (i.e. correct
scientific names for the taxa).

Say's Taxa

Acanthocinus quadrigibbus
Acupalpus debilipes
Acupalpus ob e oletus
Agonum orbicollis

Agonum sururale

Agraphus
Agrilus atiosus
Agrilus fallax
Agrilus putillus
Aleochara bilobata
Aleochara exigua
Aleochara falsifica
Aleochara fasciata
Aleochara indentata
Aleochara lustrica
Aleochara minima
Aleochara obscuricollis
Aleochara propera
Aleochara semicarinata
Aleochara simplicicollis
Aleochara verna

References

1831a: 9; 1835: 195
1830c: 21; 18346: 435
1830c: 22; 18346: 435
18306: (4)[3]; 1830c: [11];

18346: 423
18306: (4)[3]; 1830c: [10);

18346: 422
18316: 13

1833: 68(48); 1839: 163
1833: 66(46); 1839: 163
1833: 67(47); 1839: 163
1833: 59; 1839: 156
1833: 59; 1839: 156
1833: 58; 1839: 155
18326: 56; 18346: 469
18326: 56; 18346: 469
18326: 55; 18346: 468
1833: 59; 1839: 156
18326: 55; 18346: 468
18326: 57; 18346: 470
18326: 57; 18346: 470
1833: 58; 1839: 155
1833: 58; 1839: 156

Valid Names

Acanthoderes quadrigibbus (Say, 1831)
Bradycellus rupestris (Say, 1823)
Bradycellus obsoletus (Say, 1830)
Agonum punctiforme (Say, 1823)

Agonum sururale Say, 1830

Agraphus Say, 1831
Agrilus atiosus Say, 1833 [not otiosus)
Agrilus fallax Say, 1833
Agrilus putillus Say, 1833
Aleodorus bilobarus (Say, 1833)
Oligota exigua (Say, 1833)
?Xenota falsifica (Say, 1833)
Phanerota fasciata (Say, 1 832)
Philhygra indentata (Say, 1832)
Aleochara lustrica Say, 1832
?Xenota minima (Say, 1833)
Aleochara obscuricollis Say, 1832
Amischa propera (Say, 1832)
?Funda semicarinata (Say, 1832)
Crataraea suturalis (Mannerheim, 1830)
Aleochara verna Say, 1833

Vol. 104, No. 1, January & February, 1993

Aleochara 4-punctata
Aleodorus
Altica exapta
Altica mellicollis
Altica ocreata
Amara dolosa

Amara furtiva
Amara grossa
Amara sera
Amblycheila
Anchomenus collaris

Anisodactylus dilatatus
Anthonomus calceatus
Anthonomus erythropterus
Anthonomus musculus
Anthonomus quadrigibbus
Anthonomus signatus
Anthophagus verticalis
Anthribus brevicornis
Anthribus cornutus
Aphrastus
Apion segnipes
Aracanthus
Aretharea

Aretharea helluonis

Ateuchus humectus
Bagous aereus
Bagous mamillatus
Bagous simplex
Balaninus nasicus
Balaninus nasutus
Balaninus rectus
Baridius acutipennis
Baridius nigrinus
Baridius scolopax
Baridius striatus
Baridius transversus
Baridius trinotatus
Baridius undatus

Barynotus erinaceus
Barynotus granulatus

18326:57; 18346:470

1833:60; 1839: 157

1832a: 6

1831a: 10; 1835: 199

1832a: 7

18306: (7)[3]; 1830c: [15];

18346: 429
18306: (7)[3]; 1830c: [16);

18346: 429
18306: (7)[3j; 1830c: |16];

18346: 430
18306: (7)[3]; 1830c: [16];

18346: 429
1830a:67; 1830c: [1|;

18346: 409
18306: (4)[3|; 1830c: [10];

18346: 421

1830c: 18; 18346: 431
18316: 15
18316: 25
18316: 15
18316: 15
18316: 25

1831c: 49; 18346: 463
18316:4
18316:4
18316:9
18316:6
18316:9
1830a:68; 1830c: [2];

18346:411
1830a: 69; 1830c: [2];

18346:411
1832a: 4
18316: 29
18316: 28
18316:29
18316: 16
18316: 16
18316: 16
18316:27
18316:26
18316: 26
18316: 17
18316: 18
18316: 17
18316: 17

18316: 12
18316: 12

?Xenota quadripunctata (Say, 1832)
Aleodorus Say, 1833
Altica carinata German 1824
?Disonycha mellicollis (Say, 1831)
Monomacra tibialis (Olivier, 1808)
Amara dolosa Say, 1830

Amara exarata Dejean, 1828
Euryderus grossus (Say, 1830)
Amara sera Say, 1830
Amblycheila Say, 1830
Agonum collare (Say, 1830)

Anisodactylus anthracinus (Dejean, 1829)
Odontopus calceatus (Say, 1831)
Anthonomus suturalis LeConte, 1824
Anthonomus musculus Say, 1831
Anthonomus quadrigibbus Say, 1831
Anthonomus signatus Say, 1831
Geodromicus verticalis (Say, 1831)
Phaenithon brevicome (Say, 1831)
Toxonotus cornutus (Say, 1831)
Aphrastus Say, 1831
Apion segnipes Say, 1831
Aracanthus Say, 1831
Aretharea Say. 1830

Aretharea helluonis Say, 1830

Canthon humectus (Say, 1832)
Tyloderma aereum (Say, 1831)
Bagous mamillatus Say, 1831
Lissorhoptrus simplex (Say, 1831)
Curculio nasicus (Say, 1831)
Curculio proboscideus (Fabricius, 1775)
Curculio proboscideus (Fabricius, 1775)
Craptus acutipennis (Say, 1831)
Pseudobaris nigrina (Say, 1831)
Aulobaris scolopax (Say, 1831)
Baris striata (Say, 1831)
Baris transversa (Say, 1831)
Trichobaris trinotata (Say, 1831)
Craptus undatus (Say, 1831)

[not undulatus]

Panscopus erinaceus (Say, 1831)
Anametis granulata (Say, 1830)

ENTOMOLOGICAL NEWS

Say's Taxa

Barynotus rigidus
Bembidium ephippiatum
Bembidium incurvum
Bembidium postremum
Bembidium semifasciatus
Bembidium sigillare
Bembidium tripunctatum
Brachinus stygicornis

Bruchus mimus
Bruchus musculus
Bruchus obsoletus
Bruchus obtectus
Bruchus oculatus
Bruchus transversus
Bruchus triangularis
Buprestis acornis
Buprestis impedita
Buprestis thureura
Buprestis ultramarina
Callopistus
Centrinus scutellum album

Ceutorhynchus curtus
Ceutorhynchus inaequalis
Ceutorhynchus triangularis
Chlaenius circumcinctus

Chlaenius soccatus
Chlaenius vigilans

Cleogonus sedentarius
Cleonus trivittatus
Colymbetes gutticollis
Coptotomus
Coptotomus serripalpus
Cossonus corticola
Cossonus multiforus
Cossonus platalea
Cryptorhynchus
anaglypticus

Cryptorhynchus bisignarus
Cryptorhynchus cribricollis
Cryptorhynchus elegans
Cryptorhynchus ferratus
Cryptorhynchus foveolatus
Cryptorhynchus obliquus
Cryptorhynchus palmacollis

References

18316: 11

1830c: 25; 18346: 439
1830c: 26; 18346: 440
1830c: 23; 18346: 437
1830c: 25; 18346: 438
1830c: 24; 18346: 437
1830f: 26; 18346: 439
18306: (1)[3); 1830c: (5);

18346: 415
18316: 2
18316:3
18316: 2
18316: 1
18316:2
18316: 3
18316: 1

1833: 63; 1839: 159
1833: 63; 1839: 160
1832a: 3

1833: 64; 1839: 160
18316:9
18316:21

18316: 29
18316: 20
18316: 20
18306: (3)[3j; 183ft-: [7];

1831a:3; 18346:418
18306: (3)[3j; 183ft-: [8];

18346: 419
18306: (3)[3); 183ft-: [8];

18346: 419
18316: 30
18316: 10

1830c:29; 18346:442
1830c: 29; 18346: 443
1830c: 30; 18346: 443
18316: 24;
18316: 30
18316: 24
18316: 18

18316: 19
18316: 28
18316: 18
18316: 28
18316: 19
18316: 28
18316: 27

Valid Names

Phyxelis rigidus (Say, 1831)
Pericompsus ephippiatus (Say, 1830)
Elaphropus incurvus (Say, 1831)
Bembidion postremum Say, 1830
Bembidion semifasciatum Say, 1830
Bembidion punctatostriatum Say, 1823
Elaphropus tripunctatus (Say, 1830)
Brachinus quadripennis Dejean, 1825

Gibbobruchus mimus (Say, 1831)
Meibomeus musculus (Say, 1831)
Acanthoscelides obsoletus (Say, 1831)
Acanthoscelides obtectus (Say, 1831)
Acanthoscelides oculatus (Say, 1831)
Althaeus hibisci (Olivier, 1795)
Acanthoscelides triangularis (Say, 1831)
Actenodes acornis (Say, 1833)
Cypriacis striata (Fabricius. 1775)
Descarpentriesina thureura (Say, 1832)
Buprestis salisburyensis Herbst, 1801
Compsus Schonherr, 1823
Odontocorynus scutellumalbum

(Say, 1831)

Acanthoscelidius curtus (Say, 1831)
Craponius inaequalis (Say, 1831)
Rhinoncus triangularis (Say, 1831)
Chlaenius circumcinctus Say, 1830

Chlaenius soccatus Say, 1830
Chlaenius tricolor Dejean, 1826

Pseudomus sedentarius (Say, 1831)
Cleonidius trivittatus (Say, 1831)
Rhantus gutticollis (Say, 1830)
Coptotomus Say. 1830
Coptotomus serripalpus Say, 1830
Cossonus corticola Say, 1831
Cossonus multiforus Say, 1831
Cossonus platalea Say, 1831
Conotrachelus anaglypticus (Say, 1831)

Eubulus bisignatus (Say, 1831)
Pheloconus cribricollis (Say, 1831)
Conotrachelus elegans (Say, 1831)
Apteromechus ferratus (Say, 1831)
Tyloderma foveolatum (Say, 1831)
Cryptorhynchus obliquus Say, 1831
Rhyssomatus palmacollis (Say, 1831)

Vol. 104, No. 1, January & February, 1993

Say's Taxa

Cryptorhynchus retentus
Cryptorhynchus tubulatus
Cupes cinerea
Cymindis laticollis

Deracanthus pallidus
Dermestes nubilus
Dryopthorus corticalis
Dytiscus bimarginatus

Dytiscus confluens
Dytiscus habilis
Elaphrus fuliginosus

Elaphrus ruscarius

Elater agonus
Elater apicatus
Elater armus
Elater attenuatus^
Elater baridius

Elater bilobatus
Elater cardisce
Elater choris
Elater curiatus
Elater discalceatus
Elater ectypus
Elater exstriarus
Elater fallax
Elater hamarus
Elater hieroglyphicus
Elater inquinatus
Elater mellillus
Elater oblessus'
Elater obliquus
Elater pectoralis
Elater soleatus
Elater sulcicollis
Erirhinus ephippiatus
Erirhinus mucidus
Erirhinus rufus [as rufous]
Erodiscus myrmecodes
Feronia deparca

Feronia oblongo-notata
Feronia obscura

References

18316: 27

18316: 20

1831a: 6; 1835: 167

1830a: 134; 1830c: |3];

18346: 413
18316:9
1832a: 3
18316: 24
1830c: 28; 1831a: 5;

18346: 442

1830c: 27; 18346: 440
1830c: 27; 18346: 441
18306: (1)(3|; 1830c: [6];

18346: 417
18306: (1)|3); 1830c: (6];

18346: 417
1834a: 75; 1839: 171
1834a: 74; 1839: 170
1834a: 75; 1839: 171
1833: 70(50); 1839: 166
1834a: 79; 1839: 176

1834a: 78; 1839: 174
1834a: 73^; 1839: 169
1834a: 76; 1839: 172
1834a: 77; 1839: 173
18340: 73Vi 1839: 169
1833: 72(52); 1839: 167
18340: 80; 1839: 177
1834a: 74; 1839: 170
18340: 74; 1839: 170
18340: 75; 1839: 172
18340: 78; 1839: 175
18340: 76; 1839: 173
1833: 70(50); 1839: 165
18340:77; 1839: 174
18340:76; 1839: 173
18340: 79; 1839: 176
1833: 70(50); 1839: 168
18316:25
18316: 14
18316: 25
18316: 15
18306: (6)|3]; 1830r: [13];

18346: 426
18306: (6)[3|; 1830c: [13];

18346: 425
18306: (5)[3j; 1830c: [12];

18346: 425

Valid Names

Conotrachelus retentus (Say, 1831)
Idiostethus tubulatus (Say, 1831)
Tenomerga cinerea (Say, 1831)
Cymindis laticollis Say, 1830

Aracanthus pallidus (Say, 1831)
Dermestes caninus Germar, 1824
Dryophthorus americanus Bedel, 1885
Hydaticus bimarginatus (Say, 1830)

Dytiscus confluens Say, 1830
Dytiscus habilis Say, 1830
Elaphrus fuliginosus Say, 1830

Elaphrus ruscarius Say, 1830

Limonius agonus (Say, 1834)
Ampedus apicatus (Say, 1834)
Limonius stigma (Herbst, 1806)
Parallelostethus attenuates (Say, 1825)
Hemicrepidius memnonius

(Herbst, 1806)

Hemicrepidius bilobatus (Say, 1834)
Cardiophorus cardisce (Say, 1834)
Negastrius choris (Say, 1834)
Horistonotus curiatus (Say, 1834)
Elathous discalceatus (Say, 1834)
Limonius ectypus (Say, 1833)
Drapetes exstriarus (Say, 1834)2
Ctenicera fallax (Say, 1834)
Ctenicera hamata (Say, 1834)
Ctenicera hieroglyphica (Say, 1834)
Glyphonyx inquinatus (Say. 1834)
Aeolus mellillus (Say, 1834)
Ampedus oblessus (Say, 1833)
Ampedus areolatus (Say, 1823)
Paradonus pectoralis (Say. 1834)
Ischiodontus soleatus (Say, 1834)
Ctenicera sulcicollis (Say, 1833)
Elleschus ephippiatus (Say, 1831)
Dorytomus mucidus (Say, 1831)
Elleschus ephippiatus (Say, 1831)
Myrmex myrmex (Herbst, 1797)
Amara deparca (Say. 1830)

Pterostichus adstrictus

Eschscholtz. 1823
Pterostichus sayanus Csiki. 1930

10

ENTOMOLOGICAL NEWS

Say's Taxa
Feronia obsoleta
Feronia permunda

Graphorhinus
Graphorhinus operculatus
Graphorhinus vadosus
Gyrinus obtusus
Gyrinus parcus
Harpalus amputatus
Harpalus ocreatus
Hydrocanthus atripennis
Hydrophilus castus
Hydroporus bifidus
Hydroporus discicollis
Hydroporus interruptus

Hydroporus nudatus

Hydroporus sericatus
Hypsonotus alternatus
Lamia crypta
Lathrobium armatum
Lathrobium cinctum
Lathrobium confluentum
Lathrobium dimidiatum
Lathrobium millepunctatum

Lathrobium similipenne
Lathrobium sphaericolle
Lepyrus geminatus
Listroderes lineatulus
Listroderes porcellus
Listroderes sparsus
Listroderes squamiger
Lixus concavus
Lixus lateralis
Lixus marginatus
Lixus musculus
Metonius

Metonius laevigatus
Metonius purpureus
Molosoma
Noterus bicolor

Nothiophilus porrecrus

Odontopus
Oedemera apicialis
Olisthopus cinctus

References

1830ft: (5)[3]; 1830o: [12];

1834ft: 424
18306: (6)[3); 1830c: [14];

1834ft: 426
1831ft: 8
1831ft: 9
1831ft: 8

1830r 34; 1834ft: 447
1830c: 34; 1834ft: 448
1830c: 19; 1834ft: 432
183Qr: 20; 1834ft: 433
1830c:33; 1834ft: 447
1831a: 7; 1835: 170
1830c: 31; 1834ft: 444
1830c: 32; 1834ft: 446
1830o: 32; 1834ft: 445

1830c: 31; 1834ft: 444

1830c: 31;
1831ft: 10
1832a: 5
1830c: 40;
1830c: 40;
1831r: 43;
1830c: 41;
1831c:42;

1834ft: 445

1834ft: 453
1834ft: 454
1834ft: 456
1834ft: 455
1834ft: 456

1830c: 40; 1834ft: 453

1831c: 42; 1834ft: 455

1831ft: 12

1831ft: 11

1831ft: 11

1831ft: 11

1831ft: 11

1831ft: 14

1831ft: 14

1831ft: 13

1831ft: 14

1833: 68(48); 1839: 164

1833: 69(49); 1839: 164

1833:69(49); 1839:. 164

1831c: 48; 1834ft: 462

1830c: 33; 1831a: 5;

1834ft: 446
1830ft: (3)[3]; 1830c: [7];

1834ft: 418
1831ft: 15

1831a: 9; 1835: 188
1830ft: (5)[3]; 1830c: [11];

1834ft: 424

Valid Names

Cyclotrachelus obsoletus (Say, 1830)
Pterostichus permundus (Say, 1830)

Graphorhinus Say, 1831
Epicaerus operculatus (Say, 1831)
Graphorhinus vadosus Say, 1831
Gyrinus obtusus Say, 1830
Gyrinus parcus Say, 1830
Harpalus amputatus Say, 1830
Notiobia terminata (Say, 1823)
Hydrocanthus atripennis Say, 1830
Hydrobiomorpha casta (Say, 1831)
Potamonectes bifidus (Say, 1830)
?Hydroporus dichrous Melsheimer, 1846
Potamonectes griseostriatus

(DeGeer, 1774)
?Potamonectes aequinoctialis

(Clark, 1862)

Hydroporus sericatus Say, 1830
Trichalophus alternatus (Say, 1831)
Ataxia crypta (Say, 1832)
Lathrobium armatum Say, 1830
Astenus cinctus (Say, 1830)
Sunius confluentus (Say, 1831)
Lobrathium dimidiatum (Say, 1830)
Achenomorphus corticinus

(Gravenhorst, 1802)
Ochthephilum similipenne (Say, 1830)
Apocellus sphaericollis (Say, 1831)
Lepyrus palustris (Scopoli, 1763)
Listronotus sparsus (Say, 1831)
Listronotus porcellus (Say, 1831)
Listronotus sparsus (Say, 1831)
Listronotus squamiger (Say, 1831)
Lixus concavus Say, 1831
?Lixus lateralis Say, 1831
?Lixus marginatus Say, 1831
?Lixus musculus Say. 1831
Pachyschelus Solier, 1833
Pachyschelus laevigatus (Say, 1833)
Pachyschelus purpuratus (Say, 1833)
Osorius Dejean, 1821
Hydrocanthus bicolor (Say, 1830)

Notiophilus aeneus (Herbst, 1806)

Odontopus Say, 1831

Nacerdes melanura (Linne, 1758)

Olisthopus parmatus (Say, 1823)

Vol. 104, No. 1, January & February, 1993

11

Say's Taxa

Omalium marginatus
Omalium rotundicolle
Oodes parallelus

Ophryastes
Orchestes ephippiatus
Orchestes pallicornis
Oxyporus stygicus
Oxytelus cordatus
Oxytelus emarginatus
Oxytelus rugulosus
Paederus cinctus
Paederus discopunctatus
Peritelus bellicus
Peritelus chrysorrhaeus
Phytonomus comptus
Phytonomus trivittatus
Pterocolus
Rhynchites aeratus
Rhyncholus latinasus
Rhynchophorus cicatricosus
Rhynchophorus immunis
Rhynchophorus inaequalis
Rhynchophorus interstitialis
Rhynchophorus placidus
Rhynchophorus praepotens
Rhynchophorus rectus
Rhynchophorus truncatus
Rhynchophorus venatus
Rugilus dentatus
Scymnus terminatus
Sitona indifferens
Sitona scissifrons
Spheracra

Staphylinus apicialis
Staphylinus connexus
Staphylinus dimidiatus
Staphylinus ephippiatus
Staphylinus inversus
Staphylinus iracundus
Staphylinus tachiniformis
Stenolophus cinctus
Stenus colon
Stenus femoratus
Stenus geniculatus
Stenus quadripunctatus
Stenus stygicus
Stomis granulatus

References

1832A: 50; 18346: 463
18326: 50; 18346: 464
18306: (3)|3j; 1830c: |8);

183 la: 4; 18346: 420
18316: 13
18316: 16
18316: 16

1831c: 46; 18346: 459
1831c:47; 18346:461
183k: 48; 18346:461
183k: 47; 18346:460
1831c: 44; 18346:457
183k: 43; 18346: 457
18316: 13
18316: 13
18316: 12
18316: 12
18316:5
18316: 5
18316: 30
18316:22
18316: 23
18316:23
18316:21
18316:23
18316:21
18316:22
18316: 22
18316: 22

183k: 44; 18346:457
183 la: 11; 1835:203
18316: 10
18316: 10
1830a: 133; 1830c: (3);

18346: 412

1830c: 37; 18346: 451
1830c: 35; 18346: 448
1830c: 37; 18346: 450
183(V: 35; 18346: 448
1830c: 36; 18346: 449
1830r 35; 18346: 449
1830c: 37; 18346: 450
1830c: 20; 18346: 434
183k: 45; 18346: 458
183k: 45; 18346:459
183k: 44; 18346:458
183k: 46; 18346: 459
183k: 45; 18346: 458
18306: (6)|3]; 1830c: |14]

18346:427

Valid Names

Eusphalerum marginatum (Say, 1832)
Olophrum obtectum Erichson, 1840
Lachnocrepis parallela (Say, 1830)

Ophryastes Say, 1831
Tachyerges ephippiatus (Say, 1831)
Rhynchaenus pallicornis (Say, 1831)
Oxyporus stygicus Say, 1831
Bledius cordatus (Say, 1831)
Bledius emarginatus (Say, 1831)
?Anotylus rugulosus (Say, 1831)
Homaeotarsus cinctus (Say, 1831)
Astenus discopunctatus (Say, 1831)
Agraphus bellicus (Say, 1831)
Cercopeus chrysorrhaeus (Say, 1831)
Hypera compta (Say, 1831)
Hypera trivittata (Say. 1831)
Pterocolus Say, 1831
Pselaphorhynchites aeratus (Say. 1831)
Cossonus corticola Say, 1831
Sphenophorus cariosus (Olivier. 1807)
Sphenophorus venatus (Say, 1831)
Sphenophorus inaequalis (Say, 1831)
?Sphenophorus interstitialis (Say, 1831)
Sphenophorus venatus (Say, 1831)
Cleonidius trivittatus (Say, 1831)
Sphenophorus rectus (Say, 1831)
Sphenophorus pertinax (Olivier, 1807)
Sphenophorus venatus (Say, 1831)
Rugilus dentatus Say, 1831
Diomus terminatus (Say, 1831)
Sitona indifferens Say, 1831
Sitona scissifrons Say, 1831
Leptotrachelus Latreille, 1829

Hesperus apicialis (Say, 1830)
Belonuchus connexus (Say, 1830)
Philonthus sericans (Gravenhorst, 1802)
Belonuchus ephippiatus (Say, 1830)
Quedius capucinus (Gravenhorst, 1806)
Quedius fulgidus (Fabricius, 1787)
Philonthus tachiniformis (Say, 1830)
?Bradycellus rupestris (Say, 1823)
Stenus colon Say, 1831
Stenus femoratus Say, 1831
Stenus flavicornis Erichson, 1840
Carpelimus quadripunctatus (Say, 1831)
Stenus stygicus Say, 1831
?Stomis granulatus Say. 1830

12

ENTOMOLOGICAL NEWS

Say's Taxa

Tachinus cincticollis

Tachinus humidus
Tachinus obsoletus
Tachinus trimaculatus

Tachyporus acaudus
Tachyporus faber
Tachyporus fumipennis
Tachyporus jocosus
Tachyporus moestus

Tachyporus opicus
Tachyporus ventriculus
Tanymecus confusus
Tenebrio rufinasus
Thamnophilus barbitus
Thamnophilus pallidus
Thamnophilus pandura
Thecesternus
Thylacites microps
Trox aequalis
Trox alternatus
Tychius amoenus
Tychius aratus
Tylodes clavarus
Xantholinus cephalus
Xantholinus hamatus
Zygops quercus

References

1832*: 51; 18346: 465

18326: 52; 18346: 465
18326:51; 18346:464
18326:51:18346:464

18326: 54; 18346: 467
18326: 55; 18346: 468
18326: 52; 18346: 466
18326: 53; 18346: 466
18326: 53; 18346: 466

18326: 54; 18346: 467

18326: 53; 18346: 466

18316:9

1831a: 8; 1835: 187

18316:6

18316:7

18316:7

18316: 8

18316:9

1832a: 5

1831a: 7; 1835: 179

18316: 26

18316: 26

18316: 29

183flt: 39; 18346: 452

1830c: 39; 18346: 453

18316: 20

Valid Names

Lordithon thoracicus thoracicus

(Fabricius, 1776)

?Mycetoporus humidus (Say, 1832)
Lordithon obsoletus (Say, 1832)
Lordithon thoracicus venustus

(Melsheimer, 1846)
Tachyporus jocosus Say, 1832
Tachyporus nitidulus (Fabricius, 1781)
Tachinus fumipennis (Say, 1832)
Tachyporus jocosus Say, 1832
Sepedophilus crassus

(Gravenhorst, 1802)
Sepedophilus opicus (Say, 1832)
Coproporus ventriculus (Say, 1832)
Tanymecus confusus Say, 1831
?Alphitobius rufinasus (Say, 1831)
Magdalis barbita (Say, 1831)
Magdalis armicollis (Say, 1824)
Magdalis pandura (Say, 1831)
Thecesternus Say, 1831
Minyomerus microps (Say, 1831)
Trox aequalis Say, 1832
Trox suberosus Fabricius, 1775
Smicronyx amoenus (Say, 1831)
Tychius aratus Say, 1831
Acalles clavarus (Say, 1831)
Nudobius cephalus (Say, 1830)
Neohypnus hamatus (Say, 1830)
Cylindrocopturus quercus (Say, 1831)

1 Obviously, Say (1833) intended, on page 70, to describe a new species under the binomen
Elater attenuatus. He apparently overlooked the fact that he already had described an Elater
attenuatus in 1825, on page 257. The descriptions are slightly different but seem to apply to
the same taxon (E. Becker, personal communication). Elater attenuatus Say, 1833 is a
primary homonym and probably also a junior synonym of Elater attenuatus Say, 1825.

2 This species is usually listed under the name Drapetes geminatus (Say, 1825). However,
Say's name, described under the binomen Elater geminatus, is a primary homonym of Elater
geminatus Germar, 1824 and so is invalid (ICZN, 1985, art. 57 [b]). Say (1834a) explicitly
proposed the name Elater exstriatus as a replacement name for E. geminatus Say, 1825.

3 Say explicitly proposed the name Elater oblessus as a new replacement name for Elater dis-
coideus Fabricius, 1801, a primary homonym of Elater discoideus Weber, 1801. Despite Say's
(1833: 70) comments which show that he simultaneously applied the new name to a dif-
ferent species, his name remains nevertheless the valid one for the species described as
Elater discoideus Fabricius, 1801 (ICZN, 1985, art. 72 [e]), not Elater sellatus Leng, 1918 as
given by some authors.

Vol. 104, No. 1, January & February, 1993 13

ACKNOWLEDGMENTS

I thank A. Wheeler, Common wealth of Pennsylvania, Harrisburg (PA). S.Granato, The
American Museum of Natural History Library, New York. A Kleine-Kreutzmann, Public
Library of Cincinnati & Hamilton County, Cincinnati, and R. Alsop. Library of New Har-
mony Workingmen's Institute. New Harmony, for providing xerox copies of some of Say's
publications and S. Sherman, Entomology Research Library, Agriculture Canada.
Ottawa, for her help locating various publications. My colleagues A Smetana and S.
Laplante of the Centre for Land and Biological Resources Research, Ottawa, and I.
MacPhail. The Morton Arboretum. Lisle (IL), reviewed the manuscript and offered
useful comments.

LITERATURE CITED

Anonymous. 1861. Catalogue of the Harris library. Proc. Bost. Soc. Nat. Hist. 7 (1859-

1861): 266-271.
Bequaert, J. 1950. A bibliographic note on Say's two tracts of March 1831, and January,

1832. Psyche 57: 146.

Blackwelder, R.E. 1952. The generic names of the beetle family Staphylinidae with an
essay on genotypy. Bulletin of the United States National Museum No. 200. 483 pp.
Byrd, C. K. and H.H. Peckham. 1955. A bibliography of Indiana imprints 1804-1853.

Indiana Historical Bureau, Indianapolis, xxi + 479 pp.

Carus, J.V. and W. Engelmann. 1861. Bibliotheca Zoologica. Verzeichniss der Schriften
uber Zoologie, welche in den periodischen Werken enthalten und vom Jahre 1846-1 860
selbsta'ndig erschienen sind mit Einschluss der allgemein-naturgeschichtlichen, peri-
odischen und palaeontologischen Schriften. Erster Band. Engelmann, Leipzig. 950 pp.
Fitch, A. 1858. Say's Heteropterous Hemiptera. Trans. N.Y. State Agri. Soc. 17 (1857):

754.
Hagen, H.A. 1863. Bibliotheca entomologica. Die Litteratur uber das ganze Gebiet der

Entomologie bis zum Jahre 1862. Zweiter Band. N-Z. Engelmann. Leipzig. 512 pp.
Harris, T.W. 1869. Entomological correspondence of Thaddeus William Harris. M.D.

edited by Samuel H. Scudder. Boston Society of Natural History. Boston. 375 pp.
Horn, W. and S. Schenkling. 1928. Index litteraturae entomologicae. Serie I: Die Welt-
Literatur uber die gesamte Entomologie bis inklusive 1863. Band III. W. Horn, Berlin.
Pp. 705-1056.

International Code of Zoological Nomenclature [ICZN]. 1985. International Code of
Zoological Nomenclature, third edition, adopted by the XX General Assembly of the
International Union of Biological Sciences. International Trust for Zoological Nomen-
clature, London, xx + 338 pp.
Say, T. 1825. Descriptions of new American species of the genera Buprestis. Trachys and

Elater. Ann. Lye. Nat. Hist. New York 1 (2): 249-268.
Say, T. 1 830a. Descriptions of new species of North American insects, and observations on

some already described. The Disseminator of useful knowledge 3: 67-69, 133-135.
Say, T. 18306. Descriptions of new species of North American insects and observations on
some already described. The Disseminator 1(1): [3]; 1(3): [3]; 1(4): [3]; 1(5): [3); 1(6): [3];

1(7): [3].

Say, T. 1830<?-1834a. Descriptions of new species of North American insects, and obser-
vations on some of the species already described. New Harmony [Indiana). 81 pp.
[1830c-: 1-41; 183k: 42-49; 18326: 50-57; 1833: 58-73; 1834a: 73 1/2-80].

Say,T. 1831a. Descriptions of new species of North American insects, found in Louisiana
by Joseph Barabino. New Harmony (Indiana). 17 pp.

14 ENTOMOLOGICAL NEWS

Say, T. 1831ft. Descriptions of new species of curculionites of North America, with obser-
vations on some of the species already known. New Harmony [Indiana]. 30 pp.
Say, T. 183 \d. Descriptions of new species of heteropterous Hemiptera of North

America. New Harmony [Indiana]. 39 pp.
Say, T. 1832a. New species of North American insects, found by Joseph Barabino, chiefly

in Louisiana. New Harmony [Indiana]. 16 pp.
Say, T. 18346. Descriptions of new North American insects, and observations on some

already described. Trans. Amer. Phil. Soc. (N.S.) 4: 409-470.
Say, T. 1 835. Descriptions of new North American Coleopterous insects, and observations

on some already described. Boston J. Nat. Hist. 1 (1834-37): 151-203.
Say, T. 1836. Descriptions of new North American Hymenoptera, and observations on

some already described. Boston J. Nat. Hist. 1 (1834-37): 210-305.
Say, T. 1837. Descriptions of new North American Hymenoptera, and observations on

some already described. Boston J. Nat. Hist. 1 (1834-37): 361-416.
Say, T. 1839. Descriptions of new North American Insects, and observations on some

already described. Trans. Amer. Phil. Soc. (N.S.) 6: 155-190.
Say, T. 1 859a. The complete writings of Thomas Say on the Entomology of North America.

Edited by John L. Le Conte, M.D. with a memoir of the author, by George Ord. Vol. 1.

Balliere Brothers, New York, xxiv + 412 pp (+ 54 plates).
Say, T. 1 859ft. The complete writings of Thomas Say on the Entomology of North America.

Edited by John L. Le Conte, M.D. with a memoir of the author, by George Ord. Vol. 2.

Balliere Brothers, New York, iv + 814 pp.
Scudder, S.H. 1899. An unknown tract on American insects by Thomas Say. Psyche 8:

306-308.
Summers, G. 1982. A bibliography of the scientific writings of Thomas Say (1787-1834).

Arch. Nat. Hist. 11:69-81.
Titus, E.B. (ed.). 1965. Union list of serials in libraries of the United States and Canada.

Third edition. Volume 2. Wilson Company, New York.
Weiss, H.B. and G.M. Ziegler. 1931. Thomas Say, early American naturalist. Thomas,

Springfield, xiv + 260 pp.

Vol. 104, No. 1, January & February, 1993 15

BOOK REVIEW

THOMAS SAY: NEW WORLD NATURALIST. Patricia Tyson Stroud.
1992. University of Pennsylvania Press, Philadelphia. 340 pp. $24.95.

Entomologists have often maligned Thomas Say for the inadequacies of his descrip-
tions and the loss of most of his specimens. George Ord's Memoir, read before the
American Philosophical Society two months after Say's death in 1 834, was not wholly flat-
tering, speaking of Say's "neglect of literature" and his "want of technical precision." Ord,
"a small man of sour impulses," (in Peter Mathiessen's words) did go on to praise Say's
"industry and zeal" and his "integrity" and "veracity". It remained for Weiss and Ziegler. in
their 1931 biography, to clarify Say's status as America's first systematic entomologist.

In a carefully researched and lavishly illustrated new biography, Patricia Tyson Stroud
paints Say in much broader strokes, not only as an entomologist but as a pioneer naturalist
of major stature. He did, after all, describe many birds, mammals, reptiles, and shells as
well as insects, and he was acquainted with most of the naturalists of his time. The Thomas
Say that emerges from these pages is indeed a "strong, self-determined, highly motivated,
even driving character," as Stroud promises in her Introduction.

Say did not attend the initial gathering of Philadelphia naturalists on June 25. 1812, but
he did attend the following meeting, when the group decided to call itself the Academy of
Natural Sciences of Philadelphia, with Say listed as a founding member. (Ord spoke of
these initial meetings as "a club of humorists" which met for "the purpose of amusement.")
Over the next few years Say served as the Academy's curator, librarian, and editor of the
Journal. In between his field trips to Florida and to the Rockies and the Red River of the
north he remained active in the Academy, and after his move to New Harmony he often
longed for the atmosphere of "the Athens of America." Say was never a success at earning
money, and because of his dependence on William Maclure's patronage he was never to
escape the isolation and drudgery that marked the final decade of his life.

Stroud's quotations from many of Say's previously unpublished letters reveal a more
forceful (though sometimes cranky and discouraged) Say than has been appreciated. He
had little use for Rafinesque's conceits, and was disturbed by Rafinesque's not wholly
favorable review of his American Entomology. He criticized John E. LeConte for dis-
regarding some of his species descriptions and for sending specimens abroad for identi-
fication for Say believed strongly in American taxonomic self-sufficiency. (Ironically, it
was LeConte's son, John L.. who in 1 859 collected Say's scattered publications in two lavish
volumes.) But Say did have fast friends, for example Philadelphian Reuben Haines and
naturalists Charles Lucien Bonaparte and Charles Lesueur.

Lucy Say also emerges as a stronger personality than previously apparent. After her
husband's death she saw to the disposition of his collections and effects and even learned
to engrave the plates for the seventh part of American Conchology. "I am looked upon as
being very singular [she wrote to a friend], particularly since I have commenced En-
graving a gentleman remarked 'Well! at what do you think the ladies will stop?'. I replied.
I hoped at nothing. . . that we were tired of cramping our genius over the needle and the dis-
taff." Lucy Say was the first woman to be elected to the Academy of Natural Sciences. She
outlived her husband by half a century.

There are a few minor points in Stroud's biography that bothered me. Dermestid
beetles are not really "microscopic predators." and the drink that Say noted on his Mexican
trip, pulque, is not the same as tequila, though both are derived from agave. As an
entomologist, I would have enjoyed being reminded of some of the exciting discoveries Say

16 ENTOMOLOGICAL NEWS

made on his various field trips, such as the remarkable tiger beetle Amblycheila cylin-
driformis and the fiery tarantula hawk Pepsis formosa. But Stroud's objective was to place
Say securely as a leading figure in the awakening of Americans to their biological riches,
and in this she has been most successful.

Howard E. Evans

Dept. of Entomology

Colorado State University

Fort Collins, CO

BOOK REVIEW

FLORISSANT BUTTERFLIES: A GUIDE TO THE FOSSIL AND
PRESENT DAY SPECIES OF CENTRAL COLORADO. Thomas C.
Emmel, Marc C. Minno, and Boyce A. Drummond. Stanford Univ.
Press, Ca. 1992. $14.95 paper; $35.00 cloth.

Having pleasant memories of two visits to the Florissant area, I was delighted when
asked to review this book. Boyce Drummond's stunning cover photograph of two Par-
nassius phoebus sayii immediately catches one's attention.

This book about an area renowned for the quantity of well preserved insect and plant
fossils begins with information on insect fossils and the Earth's geological record. This first
section contains descriptions and photographs of the twelve butterfly fossil species found
at Florissant. The next section covers the ecology of the Florissant region's extant butter-
flies and gives detailed habitat accounts of both plants and butterflies. Other topics include
butterfly diversity, biology, behavior and survival. The authors described the ninety-seven
species recorded in the area, and black and white photographs of the larvae, pupae, and
adults accompany many of these entries. In addition, ten color plates, showing both male
and female forms, illustrate all ninety-seven species. At the end of the book one finds
references, an excellent glossary, and a checklist of the present day butterflies of the Floris-
sant Region and Central Colorado. In addition, there are general and plant indexes.

Florissant Butterflies contains a great deal of information and is a pleasure to read.
Although a field guide version would be more practical than the current, 8x11 inch format,
the book will still appeal to the serious lepidopterist as well as the traveler and the naturalist
interested in Colorado.

Jane Ruffin, naturalist and
amateur lepidopterist.

Vol. 104. No. 1, January & February, 1993 17

THE PHYLOGENETIC POSITION OF

CHLORONIELLA PERINGUEYI

(MEGALOPTERA: CORYDALIDAE)

AND ITS ZOOGEOGRAPHIC SIGNIFICANCE 1

Norman D. Penny^

ABSTRACT: Male and female genitalia of Chloroniella peringueyi are illustrated for the
first time, and phylogenies of all genera of Corydalinae are presented, based both on
morphological analysis and plate tectonics. A possible general pattern of megalopteran
evolution is suggested.

When Glorioso( 1981) published his revision of the Corydalinae, he
did not include the South African genus Chloroniella in his phylogenetic
analysis because he was "unable to obtain specimens of the only known
species due to its scarcity in collections." Further attempts to locate
additional specimens by the present author have also been unsuccessful.
In November 1986 Drs. John T. Doyen and Charles E. Griswold col-
lected 14 specimens of a species that compares well with the original de-
scription of Chloroniella peringueyi Esben-Petersen at Algeria Forest
Camp in the Sederberg mountains of Cape Province, South Africa.
Thus, the time seems appropriate to revise the original cladistic analysis
to include all known genera.

METHODS

The cladistic analysis utilized the 70 character states from Glorioso's
study with a few modifications. His character 18, the stipal setae, was
found to be too variable for inclusion. Characters 30 and 41, the 1A
branches and male ninth sternal setae respectively, were not in-
cluded in his cladogram, but are included here. His first three character
states are common to all Corydalinae, and thus are uninformative about
relationships within the subfamily and have not been included. Five
additional characters have been found and are incorporated, four of
them exist in Chloroniella, two of them being unique to this genus (Table
1). Three of these new characters are autapomorphies 3 within the sub-
family, and two are shared with only one other genus. As in Glorioso's
study, the apomorphic state was derived from a comparison with genera
in Chauliodinae, these being considered the outgroup. In all, 7 1 charac-

1 Received February 5, 1990. Revision received and accepted September 12, 1992.

2 California Academy of Sciences, Golden Gate Park, San Francisco. California. USA,
94118.

3 Autapomorphies are also uninformative about relationships, but do provide evidence
about the monophyly of a group.

ENT. NEWS 104(1): 17-30. January & February. 1993

18

ENTOMOLOGICAL NEWS

3A

2A

CuP

CuA

Figures 1-2. Chloroniella peringueyi. 1. Head in dorsal view (10X). 2. Right forewing
(5X).

Vol. 104, No. 1, January & February, 1993

19

ters are considered in this analysis. To help construct the most par-
simonious cladogram, the PAUP computer program was used. Illus-
trations were made utilizing a Wild M6 binocular microscope with a
camera lucida attachment. Genitalia were studied after clearing in 10%
cold KOH for 24 hours and female genitalia were stained with Chlorazol
Black E. The terminology is the same as that used by Glorioso.

RESULTS

Individuals of Chloroniella have forewings with mostly plesio-
morphic character states, although a few apomorphic modifications
have arisen, such as reduced apical forks of R5 and MP (Fig. 2). The male
genitalia (Figs. 3, 4) provide the most distinctive characters, with
medially notched ninth sternum, medially fused papillae, a tuft of setae
at the base of the tenth tergites, and medially divided tenth gonocoxites.
Other useful characters are found in the female genitalia (characters 62
to 71) (Figs. 5, 6). The pale coloration with dark markings of Chloroniella
superficially resembles that of the New World Chloronia and the Asiatic
Neuromus (Fig. 1). Utilizing these and the other characters listed by
Glorioso, only one most parsimonious cladogram is generated by the

gonocoxjte

4

Figures 3-4. Chloroniella peringueyi apex of male abdomen (25X). 3. Dorsal view. 4.
Ventral view.

20

ENTOMOLOGICAL NEWS

PAUP program (Cladogram 1), with a consistency index of 0.688. This
cladogram has 109 steps.

CONCLUSIONS

The cladogram generated by PAUP is very similar to that presented
by Glorioso. Except for the addition of Chloroniella, there are no
other changes.

Megalopterans are today highly dependent on fresh water and there
is no reason to believe that previous ancestors behaved differently.
Adults are heavy-bodied, weak flyers and are seldom found far from
streams and lakes. Eggs are laid in large masses on land, and pupae are

ninth tergite
ninth gonocoxite

Figures 5-6. Chloroniella peringueyi apex of female abdomen (25X). 5. Lateral view. 6. Ven-
tral view.

Vol. 104, No. 1, January & February, 1993

21

found on land in the substrate near the margins of water courses. Larvae
of all megalopterans are aquatic, and with the exception of Pacific Coast
Neohermes and Protochauliodes are all confined to permanent bodies of
water. Thus, the ability to disperse is highly restricted, and present day
distribution patterns probably closely reflect past climatic and plate tec-
tonic events. Ancestral forms in Laurasia and Gondwanaland would
have suffered drastic distributional and richness reductions as "con-
tinental effects produced extensive aridity as the land-mass drifted

5
B 3 i ! g

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1

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1
1

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10',18.20.22.27r.28r.30,45.64.69

27,42.55.61,67
\

CLADOGRAM 1

Cladogram 1 . Phylogenetic position of corydaline genera using PAUP analysis. Numbers
refer to apomorphic states in Table 2.

22 ENTOMOLOGICAL NEWS

through middle latitudes in the late Permian and Triassic" (Riek, 1970).
There would have been a polarizing effect during the Triassic as "the
lower-latitude zone featured high aridity to about 45N and 55S, except
in western Europe, where somewhat greater humidity characterized the
higher latitude areas" (Frakes, 1979). At the same time, where aquatic
habitats continued to exist, the ameliorating local environment may
have allowed primitive forms to survive long after terrestrial forms
became extinct. Subsequent improving climatic conditions in lower
latitudes during the Paleogene (65 to 22.5 mya) would have allowed
plates, such as South America and India, drifting into the tropical zones,
to retain and even expand their high latitudinal faunas, while these same
faunas were being eliminated closer to the poles "an abundance of
atmospheric moisture at high latitudes seems to have retreated equator-
ward through the interval, and the first glaciers since the Paleozoic
formed and expanded in Antarctica" (Frakes, loc. cit.).

The three suprageneric groups of Megaloptera (Sialidae, Chaulio-
dinae and Corydalinae) all show elements of a previous austral distribu-
tion (Maps 1-3). Each of the three groups has at least one endemic species
in South Africa, and Sialidae and Chauliodinae each have endemic
species in Madagascar. No megalopterans are known from other parts of
sub-Saharan Africa. An endemic sialid species is known from southern
Asia, as well as five genera of Corydalinae and four genera of
Chauliodinae. In the U.S.S.R. and Europe only Sialis occurs today,
although fossil evidence indicates that Corydalidae was previously
present.

In the Australian Region, two endemic groups of Sialidae occur in
Australia. One additional genus of C\\au\iodinae,Archichaulidoes is con-
fined to Australia, New Zealand and Chile.

The Neotropical Region contains endemic sialid species formerly
placed in Protosialis, as well as a very distinctive Chilean species, Pro-
tosialis chilensis. This region also shares two endemic genera of Cory-
dalinae and two genera of Chauliodinae with the Australian Region.

The only other genus of Corydalinae found in the Neotropical
Region, Corydalus, as well as the chauliodine Protochauliodes, may pro-
vide clues to the largest anomaly of the Gondwanian distribution pat-
tern. Corydalus is not only found throughout South and Central
America, but also occurs throughout most of North America as well.
However, as Glorioso pointed out, "Because of the diversity and pres-
ence of the most primitive species of Corydalus and Playneuromus in the
Amazonian region, I hypothesize this as the ancestral range of the com-
mon ancestor of these genera, with subsequent range extension into
Central America." (p. 273). To this could also be added the genus

Vol. 104, No. 1, January & February. 1993 23

Chloronia (Penny & Flint, 1982). Protochauliodes has a broad distribu-
tion, which may be explained in a similar manner. In addition to being
found in Australia, New Zealand, and Chile, it is also present in the
western United States. This genus probably has moved northward from
its previous Gondwanian distribution. The Chauliodinae also contain
fourother genera that are endemic to North America. As indicated by the
distribution ofCorydalus and Protochauliodes, there appears to have been
a pathway for northward movement from South America into the
western United States. However, this pathway was probably of early
origin, as primitive chauliodine eggs have been found in Tertiary
deposits of the Laramie Formation. Thus, the western Nearctic Region
may have become a secondary center of evolution within the Chaulio-
dinae at a relatively early date.

This theory of Gondwanian distribution would have to hypothesize
a few other factors. Corydalinae would have had to become extinct in
Madagascar and Australia. The geographical pattern ofSialis would be
best explained by a Laurasian, rather than Gondwanian evolution,
perhaps from an earlier vicariant event. It should also be noted that the
higher classification of Sialidae is badly in need of revision. Such a study
could affect the zoogeographical analysis of this family.

The cladogram generated in this study provides another anomaly to
a Gondwanian distribution. Although the majority of genera fit well into
such a pattern, such as the three South American genera and two primi-
tive lineages in southern Asia, the presence of Chloroniella in South
Africa in this scenario necessitates either a later evolution of this genus
or a movement of an ancestral Chloroniella from Asia and into southern
Africa. As mentioned earlier, there are no megalopterans today in tropi-
cal Africa, and an early extinction with subsequent reintroduction from
Asia seems doubtful. A non-parsimonious cladogram, with Chloroniella
evolving from a position completely compatible with what we know of
plate tectonics, is presented in Cladogram 2. In this case, all five Asiatic
genera are together at the base of the cladogram. Some of these five
genera in three clades could have either been present in southern Asia
from a previous vicariant event (perhaps the split of Laurasia and
Gondwanaland), or been present as a single or multiple ancestor on the
Indian subcontinent as it drifted northward, later forming the genera
Protohermes, Neurhermes, Neoneuromus, Neuromus, andAcanthacorydalis.
As the continents of Africa and South America separated, they took with
them the ancestors of Chloroniella, and Platyneuromus, Chloronia. and
Corydalus, respectively. Although not the most parsimonious clado-
gram, there are only four more steps (113 steps; consistency index 0.664)
than the most parsimonious cladogram, and in view of the general

24

ENTOMOLOGICAL NEWS

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2

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Vol. 104, No. 1, January & February, 1993

25

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B9

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26

ENTOMOLOGICAL NEWS

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13

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Vol. 104, No. 1, January & February, 1993

27

indications of a Gondwanian distribution, it is felt that this non-
parsimonious modification is justified.

Because of similar biologies and restrictions to fresh water environ-
ments, the three main groups of megalopterans may have had similar
histories of speciation and vicariant events. Much of this early evolution
may have been caused by plate tectonics, leading to the similar world-
wide distribution patterns evident today. Further studies of sialid and
chauliodine phylogeny may reveal whether geographically associated

c
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8

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3

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si

f

7.1 OM 8.20,22,45,52.54.64.66.68.69

1.3,4.6,13.15,29,63

CLADOGRAM 2

Cladogram 2. Phylogenetic position of corydaline genera reflecting geographical dis-
tributions.

28

ENTOMOLOGICAL NEWS

genera of the three groups had similar cladistic histories, or not. They
may also reveal which of the two cladograms presented best fits a general
megalopteran evolution pattern.

Table 1. Characters of the Corydalinae with their plesiomorphic and apomorphic
states.

Character

Plesiomorphic

Apomorphic

1. Head shape

2. Postocular flange

3. Postocular spine

4. Posterior tentorial pits

5. Cranial disk

6. Antennae

7. Clypeal margin

8. Male mandibles

9. Female mandibles

10. Labrum shape
10'.

11. Labral position

12. Labral anterior margin

13. Maxilla shape

14. Lacinia apex

15. Galeal setae

16. Galeal sensory peg
16'.

17. Maxillary palp

18. Maxillary palp apex

19. Maxillary palp setae

20. Maxillary palp sensory

areas

21. Labial palp

22. Labial palp sensory

areas

23. Last branch of radial

sector

24. Ml + 2 branches

25. M3 + 4 branches

26. 1A branches

27. Rl-Rs crossveins

28. Medial crossveins

29. M-Cu crossveins
29'.

30. Cubital accessory

crossveins

31. Costal crossveins

32. Male ninth sternum

33. Male ninth sternum

Robust

Absent

Feebly developed

Linear

Without spines

Feebly subserrate

Entire

Not enlarged

Three-fourths head length

or shorter
Triangular

Over mandibles
Sparsely setose
Short and broad
With three elongate setae
Flattened, lanceolate
Well developed

Five-segmented

Conical

Long

One at apex

Four-segmented
one at apex

Bifurcate

Two

Two

Two

Three

Two

Three

Absent

Vertical or oblique
Hind margin not notched
More or less quadrate

Flattened

Present

Moderately to well developed

Arcuate

With spines

Filiform

Incised medially

Enlarged

Length of head

Ovoid

Broadly truncate

Between mandibles

Fimbriate

Relatively elongate

Without elongate setae

Bristlelike

Poorly developed

Absent

Four-segmented

Broadly rounded

Short

Two at apex

Three-segmented
Two at apex

Not bifurcate

Four or more

One

Three

Four or more

Three or more

Four

Six or more

Present

Reticulate

Hind margin notched

Attenuate

Vol. 104, No. 1. January & February, 1993

29

Table 1. Character stales (continued)

Character

Plesiomorphic

Apomorphic

34. Male ninth sternum

35. Male ninth sternum

36. Male ninth sternum

37. Male ninth sternum
37'.

38. Male ninth sternum

39. Male ninth sternum

40. Male ninth sternum

41. Male ninth tergum

42. Male ninth tergum

43. Ninth tergal internal

inflection

44. Ninth tergal internal

inflection

45. Membrane between 9th

& 10th sternites
45'.

46. Genital papillae

47. Genital papillae

medially

48. Tenth gonocoxites

49. Tenth gonocoxites

50. Tenth gonocoxites

51. Tenth gonosrylus
51'.

52. Ninth gonostylus

53. Ninth gonosrylus

54. Ninth gonostylus

55. Ninth gonostylus

apodeme

56. Tenth tergites

57. Tenth tergites

58. Tenth tergites

59. Tenth tergites

60. Tenth tergites

61. Eleventh tergum

62. Lateral sclerite of

ovipositor

Without median projection
Without posterolateral lobes
Without setiferous lateral

protuberances
With normal setae

Not sclerotized dorsally
Without internal ridges
Not locking with tenth

sternite
More or less quadrate

With dorso-lateral

incisions
Without median fossa

Arched
Thin

Present
Separated

Without median projection
Antero-lateral corner

without acute projection
Medially joined
Digitiform

Unguiform
Relatively short
Sparsely setose
Parallels ninth tergum

Short

Without sensory field on

apical surface
Rounded
Uniramous
Not bearing basal tuft

of hairs
Present
Well sclerotized

With median projection
With posterolateral lobes
With setiferous lateral

protuberances
With short, stout setae
With fine setae
Sclerotized dorsally
With internal ridges
Locking with tenth sternite

Short, with broad median

incision
Without dorso-lateral

incisions
With median fossa

Inverted V-shaped
Thickened, bilobate

Thickened, regularly

convoluted
Absent
Fused

With median projection
Antero-lateral corner with

acute projection
Medially separated
Short, broad
Papilliform
Clavate
Long, bent
Densely setose
Directed medially

Long, thin

With sensory field on

apical surface
Laterally compressed
Biramous
Bearing basal tuft of hairs

Absent

Weakly sclerotized

30

ENTOMOLOGICAL NEWS

Table 1. Character states (continued)

Character

Plesiomorphic

Apomorphic

63. Gonostylus

64. Sternal pouch

65. Sclerites between 8th

sternum & gonopore

66. Bursa copulatrix

67. Number of

spermathecae

68. Spermathecal duct

69. Accessory glands

70. Accessory glands

71. Accessory glands

Articulated with gonocoxite Fused with gonocoxite

Absent
Absent

Not saclike
Two

Continuous with bursa

Present

Short

Sigmoid

Present
Present

Saclike
Ore

Separate from bursa,

T-shaped
Absent
Long
Linear

LITERATURE CITED

Esben-Petersen, P. 1924. South African Megaloptera. Annals of the South African
Museum, 19:151-158. (Esben-Petersen's name was erroneously spelled "Ebsen-Peter-
sen" in this article.)

Frakes, L.A. 1980 (1979). Climates throughout Geologic Time. Elsevier Scientific Pub-
lishing Company. New York.

Glorioso, M J. 1981. Systematics of the dobsonfly subfamily Corydalinae (Megaloptera:
Corydalidae). Systematic Entomol., 6:253-290.

Penny, N.D. and Flint, O.S., Jr. 1982. a revision of the genus Chloronia (Neuroptera: Cor-
ydalidae). Smithsonian Contributions to Zoology, No. 348. 29 pp.

Riek, E.F. 1970. Composition and distribution of the fauna, pp. 187-204, In C.S.I.R.O., The
Insects of Australia. Melbourne University Press.

Vol. 104, No. 1, January & February, 1993 31

A NEW SPECIES OF POLYCENTROPUS
(TRICHOPTERA: POLYCENTROPODIDAE) FROM

ARKANSAS 1

David E. Bowles^, Michael L. Mathis^, Steven W. Hamilton^

ABSTRACT: Polycentropus stephani, a new species from Arkansas, is described and illus-
trated. This species is a member of the confusus species-group and is most closely related to
P. chelatus. P.floridensis and P. neiswanderi. but differs primarily in having a prominent spur
on the basoventral swelling of the phallus. Known only from the interior highlands. P.,
stephani may be endemic to that region.

The Polycentropus confusus species-group (Trichoptera: Polycen-
tropodidae) consists of 16 previously described species with all occur-
ring in eastern North America (Hamilton el al 1990). During a survey of
the Trichoptera of the interior highlands of Arkansas (Bowles and
Mathis 1989), some undescribed adult caddisflies belonging to the
Polycentropus confusus species-group were collected with ultraviolet-light
traps. These caddisflies were initially identified as Polycentropus species
B and C (Bowles and Mathis 1989), but were subsequently determined to
be conspecific. Herein, we describe that species. Morphological ter-
minology follows that of Hamilton (1986) and Hamilton et al (1990).

The holotype and allotype are deposited at the National Museum of
Natural History (NMNH), Washington, DC. Paratypes are deposited at
the Royal Ontario Museum (ROM), Florida State Collection of Arthro-
pods (FSCA), Illinois Natural History Survey (INHS), University of
North Texas (UNT), and the NMNH. All material is preserved in 70%
ethanol.

Polycentropus stephani, new species

(Figures 1-4)
Polycentropus species B and C, Bowles and Mathis, 1989:237

Adult. Length of forewing: Male, 5. 1 mm; female, 6. 1 mm. Body and wing color light
brown. Setae on dorsum of head and thorax tan.

1 Received May 27, 1992. Accepted June 21, 1992.

* United States Air Force Armstrong Laboratory, Occupational and Environmental
Medicine Branch, Brooks AFB, Texas, 78235

3 Department of Biological Sciences. University of Arkansas, Fayetteville, Arkansas.
72701

4 Center for Field Biology and Department of Biology. Austin Peay State University,
Clarksville, Tennessee 37044

ENT. NEWS 104(1): 31-34. Januarv & February. 1993

32 ENTOMOLOGICAL NEWS

Male genitalia. Abdominal sternite IX large, semicircular in lateral view, anterior
margin rounded, posterior margin sinuate. Intermediate appendages apically free, slightly
decurved, proximally fused to membranous dorsum of segment IX. Body of each preanal
appendage short, with broad emargination of posterior margin; dorsal process long and
decurved. Each inferior appendage with elongate ventral portion, in lateral view only
slightly narrowed distad, in ventral aspect narrowing gradually distad; dorsobasal arm of
inferior appendage large, curving posterad, in lateral view narrowing abruptly into ventral
portion, with turned-in blade-like portion at base, in caudal view this part broadly tri-
angular with apex rounded. Phallobase tubular, moderately decurved, basoventral swell-
ing bearing a prominent caudally directed spur; phallic sclerite elongate.

Female genitalia. Sternite VIII broad, membranous; lateral lobes elongate, expanded
at mid-point, tapering posteriorly. Vaginal sclerites forming vase-shaped sac; vulvar
sclerite circular, with rimmed opening posteriorly.

Type Material: United States, Arkansas. Holotype. <f, Logan Co., Mt. Magazine,
Green Beach, Gutter Rock Creek, 1 May 1987, R.Leschen,UV-light(NMNH). Allotype. 9,
same data as holotype (NMNH). Paratypes. 1 cf, 1 9, same data as holotype (FSCA); 1 cf, 1
9, same data as holotype (ROM); 2 d 1 cf. Independence Co., unnamed intermittent stream,
18 April 1987, P. Harp (NMNH, UNT); 2 c? cf, 2 9 9, Washington Co., Devil's Den State
Park, 22 April 1989, C. E. Carlton, blacklight (NMNH); 1 cf, Washington Co., 2 mi. N
Bugscuffle, dirt road at Hwy 265, near Strickler, 20 April 1987, R. Leschen, at light
(INHS).

Etymology. We name this species in honor of Karl Stephan whose unfailing en-
thusiasm for collecting insects has been invaluable in the development of the interior
Highlands caddisfly inventory.

DISCUSSION

This species belongs in the confusus species-group and exhibits
characters similar to several species within that group. The broadly
emarginate basal portion of the preanal appendage is similar to that
observed for several members of the confusus species-group including P.
floridensis Lago and Harris, P. neiswanderi Ross, and P. thaxtoni Hamilton
and Holzenthal. The phallus of P. stephani is most similar to that off.
chelatus Ross and Yamamoto from which it differs in the presence of a
prominent spur situated on the basoventral swelling of the phallobase.
This spur is unique among the species of the confusus species-group. The
female of/ 3 , stephani bears resemblance to several other species in the
confusus species-group. However, the females off. chelatus, P. floridensis
and P. thaxtoni are unknown and detailed comparisons among these
species can not be completed at this time.

This species has been collected only from the interior highlands of
Arkansas and may be endemic to the region. Polycentropus centralis
Banks was the only other member of the confusus species-group collected
in conjunction with P. stephani. Small intermittent streams are the prob-
able habitat of the immatures based on collection of the adults. However,
nothing is known about the biology of this species. A description of the
type locality was presented by Mathis and Bowles (1989), and a descrip-
tion of a paratype locality by Flint and Harp (1990).

Vol. 104, No. 1, January & February, 1993

33

1

int.app.

inf.app.

Figs. 1-4: Polycentropus stephani n. sp. 1. Male genitalia, lateral. 2. Male phallus, lateral. 3.
Dorsobasal arm of inferior appendage, caudal. 4. Female genitalia. ventral. Abbreviations:
b.v.s., basoventral spur; d.p., dorsal process of preanal appendage; 1.1. s. VII, lateral lobe of
eighth sternite; pre. app., preanal appendage; s. IX. ninth sternite; VIII. eighth tergite.

34 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

We thank Chris Carlton and Richard Leschen for providing us specimens of this
species. Oliver Flint kindly provided some additional information about paratypes and
reviewed this manuscript. Chad McHugh and Stephen R. Moulton II gave commentary
on earlier versions of this manuscript.

LITERATURE CITED

Bowles, D. E., and M. L. Mathis. 1 989. Caddisflies(Insecta:Trichoptera) of mountainous

regions in Arkansas, with new state records for the order. J. Kansas Entomol. Soc.

62:234-244.
Flint, O. S., Jr., and P. A. Harp. 1990. Lepidostoma (Nosopus) ozarkense (Trichoptera:

Lepidostomatidae), a new species from Arkansas. Entomol. News 101:81-87.
Hamilton, S. W. 1986. A new species of Polvcentropus (Trichoptera: Polycentropodidae)

from Cuba. Proc. Entomol. Soc. Wash. 88:731-733.
Hamilton, S. W. 1986. A new species of Polvcentropus (Trichoptera: Polycentropodidae)

from Cuba. Proc. Entomol. Soc. Wash. 88:731-733.
Hamilton, S. W., S. C. Harris, and P. K. Lago. 1990. Description of two new species of

Polvcentropus from Alabama with a checklist of the Polvcentropus confusus-group

(Trichoptera: Polycentropodidae). J. New York Entomol. Soc. 98:362-368.
Mathis, M. L., and D. E. Bowles, 1989. A new microcaddisfly genus (Trichoptera:

Hydroptilidae) from the interior Highlands of Arkansas. J. New York Entomol. Soc.

97:187-191.

SOCIETY MEETING OF OCTOBER 28, 1992

INSECTS IN THE CLASSROOM-
TURNING KIDS ON TO SCIENCE WITH BUGS

Dr. Harold B. White, organizer

University of Delaware

A diverse set of speakers and styles highlighted a presentation on teaching entomology
in schools. This meeting, more in line with a workshop than a lecture, featured presen-
tations by five entomologists drawn from the Science Alliance's Speakers Bureau, a
resource "pool" of professionals available for speaking to students in Delaware schools.
The objectives of this meeting, as outlined by Dr. White, were twofold: to give teachers ideas
on presenting science (and particularly entomology) to their students, and to provide
encouragement to entomologists to make themselves available for presentations in the
schools. Dr. White also highlighted the Society's educational activities, including the Field
Day and the Calvert Prize.

Dr. Douglas Tallamy. University of Delaware, talked on "Introducing Insect Behavior"
and featured an excellent series of slides related to the subject. A major point he attempts to
bring out to students is to consider how an insect's appearance is related to its behavior, e.g.,
defensive, mimicry, crypsis, etc. and in so doing attempts to get beyond many students'
impressions that insects are grotesque. An impressive set of slides focused on the monarch
butterfly and other milkweed insects and illustrated how this complex of insects is able to
avoid the latex flow when feeding on milkweed leaves. Another interesting series of slides

Continued on page 38

Vol. 104, No. 1, January & February, 1993 35

A NEW SPECIES IN THE

POLYCENTROPUS CINE RE US GROUP

(TRICHOPTERA: POLYCENTROPODIDAE)

FROM ARKANSAS AND TEXAS 1

Stephen R. Moulton II, Kenneth W. Stewart 2

ABSTRACT: Polycentropus harpi. a new species belonging to the P. cinereus group, is de-
scribed and illustrated. The species was collected from the Edwards Plateau physiographic
subregion of southcentral Texas and the Ouachita Mountains of Arkansas. This addition
brings the total number of species belonging to the P. cinereus group to three.

The diversity of caddisflies in Texas is still poorly known despite
earlier attempts to document the fauna (Edwards 1 973). In order to better
understand the species composition and distribution of caddisflies in
Texas, a long term study was undertaken to intensively sample all major
physiographic subregions in the state. The caddisfly fauna of the Interior
Highlands (Ozark and Ouachita Mountains), though better known than
Texas (Unzicker etal. 1970, Bowles and Mathis 1989, Mathis and Bowles
1992), still needs a thorough analysis of its biogeographic affinities.
Current research of the senior author is addressing those needs. In the
course of examining recent collections from the Edwards Plateau
physiographic subregion of central Texas and the Ouachita Mountains
in Arkansas, we have discovered an undescribed species belonging to
the Polycentropus cinereus group. Herein we describe and illustrate the
new species, and discuss its relationship to other members of the
group.

Morphological terminology follows that of Hamilton et al. (1990).
Type material is deposited in the collections of Clemson University
(CU), Illinois Natural History Survey (INHS), National Museum of
Natural History (NMNH), and the University of North Texas (UNT). All
material is preserved in 70% ethanol.

Polycentropus harpi, new species

Male. Length of forewing 4.0-7.0 mm. Thoracic pleura, sterna and legs yellow; nota and
dorsum of head darker brown with numerous clear-yellow, erect setae. Forewings covered
with a mixture of yellow and brown setae; membranes interspersed with small indistinct
pale spots; stigmal region with spots coalesced to form a large pale area (not evident in

1 Received June 29, 1992. Accepted August 27, 1992.

2 Department of Biological Sciences, University of North Texas, Denton, Texas 76203

ENT. NEWS 104(1): 35-38. January & February. 1993

36

ENTOMOLOGICAL NEWS

material older than one year). Fork 1 of hind wing present; discoidal cell closed. Male
genitalia (Figs. 1-4): Abdominal sternite IX (s.IX) in lateral aspect broad basally, tapering
dorsally; ventrally concave. Preanal appendages (pre. app.) broad, short and rounded
apically. Membranous terga IX and X fused (IX+X). inferior appendage (inf. app.) roughly
quadrate in lateral aspect; posterior margin deeply emarginate; dorsomesal processes
(dm.pr.) in lateral view acute and projecting posteriorly, their mesal surfaces with two to
three clear, stout setae projecting mesally; ventromesal lobes of each inferior appendage
broadly rounded with dark, truncate peg-like setae covering the apicodorsal and apicoven-
tral surfaces. Phallobase broad and deeply concave in lateral view; apicodorsal area of
phallus membranous; in dorsal aspect, left phallic rod (1. ph. r.) lanceolate and curving

pre.app.

dm.pr.

inf.app.

dm.pr.

ph.scl.

r.ph.rd:

Fig. 1-4. Polycentropus harpi n.sp. 1, male genitalia, lateral. 2, sternite IX and inferior
appendages, ventral. 3. phallus, lateral. 4, phallus, dorsal. VIII = abdominal segment 8,
s.IX = abdominal sternite 9, IX+X = fused, membranous abdominal segments 9 and 10,
pre.app. = preanal appendage, inf.app. = inferior appendage, dm.pr. = dorsomesal pro-
cess, ph.scl. = phallic sclerite, r.ph.rd. = right phallic rod, l.ph.rd. = left phallic rod.

Vol. 104, No. 1, January & February, 1993 37

gradually to left; right phallic rod (r. ph. r.) shorter and straight; phallic sclerite (ph. scl.)
straight and gradually tapering to a point in dorsal view; in lateral view, phallic sclerite
slightly bowed with anterior end emarginate.

Female. Unknown.

Immatures. Unknown.

Material. Holotype, male: U.S.A., Texas, Kendall Co., Cibolo Creek below confluence
with Ranger Creek, Boerne, 17-111-1992, K. W. Stewart UV light (NMNH). Paratypes.:
same data as holotype, 1 male, (NMNH); same as holotype but 18-IV-1992, D. E. Bowles, 1
male, (INHS); Arkansas: Garland Co., Bear Creek at Camp Clear Fork, 27-IX- 1986, D. E.
Bowles, 1 male (NMNH); Montgomery Co., Strawn Spring. 0.5 mi E Caddo Gap. 12-IX-
1980, H. W. Robison, 4 males, (INHS); same as previous but Jones's Creek at AR Hwy 8,
Caddo Gap, 2 males (CU); Fourche Mt., E Mena District Rifle Range, ca. 10 mi NW Mena,
1 1-V1-1991, B. Ewing, 2 males, (CU); Polk Co.. Ewing Farm, 7 mi W Mena, 26-V1-1991, B.
Ewing, 2 males, (UNT); same data as previous but 27-VI-1991, 1 male, (NMNH).

Etymology: We name this species in honor of George L. Harp (Arkansas State Uni-
versity) who has made extensive contributions to our understanding of aquatic insects
in Arkansas.

DISCUSSION

Polycentropus harpi is most closely related to P. cinereus Hagen. It is
readily distinguished from the latter by the following combination of
male characters: ( 1 ) much deeper posterior emargination of the inferior
appendage, (2) left dorsal phallic rod curved to the left; this structure is
straight in P. cinereus. and (3) general convex outline of the dorsomesal
processes of the inferior appendages when viewed ventrally; this outline
is straight in P. cinereus. Polycentropus harpi and P. cinereus are easily dis-
tinguished from P. sabulosus Leonard and Leonard on the basis of the
posterior emargination of the inferior appendage. Specimens of P. harpi
from Arkansas are much smaller than those from Texas and may have
the left phallic rod angled more sharply to the left.

The P. cinereus group contains two species recorded from North
America (Armitage and Hamilton 1990), in addition to P. harpi. The
nominate species, P. cinereus, is transcontinental (Armitage and
Hamilton 1990). Polycentropus sabulosus is known only from Michigan.
Polycentropus harpi is known only from the type series. In Texas P. harpi
has been collected with P. picana Ross while in Arkansas it has been
collected with P. centralis Banks.

ACKNOWLEDGMENTS

We thank D. E. Bowles. B. Ewing. and H. W. Robison for collecting additional
specimens of P. harpi. M. L. Mathis allowed the senior author to examine material in the
University of Arkansas Arthropod Collection. We thank S. W. Hamilton and J Kennedy
for reviewing an early draft of the manuscript. The comments of three anonymous
reviewers were appreciated. This study was supported in part by the National Science
Foundation (DEB-9200895) and a UNT Faculty Research Grant to KWS.

38 ENTOMOLOGICAL NEWS

LITERATURE CITED

Armitage, B. J. and S. W. Hamilton, 1990. Diagnostic atlas of North American caddisfly

adults. II. Ecnomidae, Polycentropodidae, Psychomyiidae, and Xiphocentronidae.

The Caddis Press. Athens, AL. 152 pp.
Bowles, D. E. and M. L. Mathis. 1989. Caddisflies (Insecta: Trichoptera) of mountainous

regions in Arkansas, with new state records for the order. Jour. Kans. Entomol. Soc.

62:234-244.

Edwards, S. W. 1973. Texas caddisflies. Texas Jour, of Science 24:491-516.
Hamilton, S. W., S. C. Harris, and P. K. Lago. 1990. Description of two new Polycentropus

from Alabama with a checklist ofthePolycentropusconfusus species-group (Trichoptera:

Polycentropodidae). Jour. New York Entomol. Soc. 98:362-368.
Mathis, M. L. and D. E. Bowles. 1992. A preliminary survey of the Trichoptera of the

Ozark Mountains, Missouri, U.S.A. Entomol. News 103:19-29.
Unzicker, J. D., L. Aggus, and L. O. Warren. 1970. A preliminary list of the Arkansas

Trichoptera. Jour. Georgia Entomol. Soc. 5:167-174.

Continued from page 34

showed the entire process of silphid beetles burying a mouse carcass and the development
of the beetle brood. He also showed how the carrion beetle life cycle could be replicated in
the classroom. A related presentation, by Dr. Judith Hough-Goldstein, entitled "Amazing
Insects", also used slides to illustrate many basic ideas concerning insects such as body
characteristics, major orders, metamorphosis, etc. She also stresses to students that insects
can be competitors to humans, and ends her presentation with live insects, including a
hissing cockroach. In addition, she brings to the classroom a collection of striking and
interesting specimens.

Dr. Elzie McCord Jr., of the Stine Haskell DuPont Labs, presented "Insect Show and
Tell". He goes to the classroom to promote insects as interesting, not "yucky", to encourage
participation by the students, introduce students to what scientists do and show that scien-
tists can come from diverse backgrounds. He attempts to develop with the students lots of
interactions and much exposure to live insects, aptly demonstrated by his talk to the
society, and he tries to leave the students with a setup of live insects for their classroom. Dr.
Susan Whitney, University of Delaware, in "Bug Out", presented an overview of a wonder-
ful series of lessons she has developed for 6-10 year olds entitled Bug Out. This series incor-
porates interaction, observation, projects and short lectures to illustrate many aspects of
insect biology (available at low cost through North Carolina State University). She ended
by having the entire audience participate in a sample lesson of drawing a "mystery" bug
and then identifying it; the trick here is that the children receive much information about
how an insect is constructed while participating in a fun game. Dr. Harold White, Uni-
versity of Delaware, ended the session with "Aquatic Insects." As an amateur entomologist
(he is a biochemistry professor) he stresses this aspect of the science to kids, and sees his
presentations as a way to get children interested in science perse, not just entomology. His
demonstrations were simple but effective, and relied only on a petri dish, overhead pro-
jector and a few live aquatic insects. By simply placing the insects in water in the dish on
the projector, locomotion, morphology and respiration of aquatic insects all can be illus-
trated. He stressed, above all, that the best experience for the kids is to take them out-
side to a pond!

Over 50 society members, guests and teachers attended this meeting at the Uni-
versity of Delaware.

Jon K. Gelhaus,
Corresponding Secretary

Vol. 104, No. 1, January & February, 1993 39

INTERACTIONS OF PREDACEOUS KATYDIDS

(ORTHOPTERA: TETTIGONIIDAE)

WITH NEOTROPICAL SOCIAL WASPS

(HYMENOPTERA: VESPIDAE): ARE WASPS

A DEFENSE MECHANISM OR PREY? 1

Sean O'Donnell 2

ABSTRACT: Field observations of interactions between predaceous tettigoniids, Phlugis
poecila and Ancistrocercus inficitus, and neotropical eusocial wasps, Polistes instabilis and P.
versicolor, show that tettigoniids prey on unguarded wasp nests and may employ chemical
or tactile cues to distinguish the nests. Tettigoniids prey on brood in P. instabilis nests from
which adult wasps are experimentally removed. Polistes instabilis and Polybia occidentalis
wasp workers are not tolerant of tettigoniids roosting near their nests, and tettigoniids near
wasp nests behave as if avoiding detection by adult wasps. These observations suggest that
predation on brood is an important feature of tettigoniid associations with neotropical
wasp nests.

Associations of tettigoniids with active colonies of social wasps in the
neotropics have been interpreted as a commensalistic or mutualistic
relationship, wherein the tettigoniids gain protection from predators for
the duration of their diurnal inactive period and the wasps gain
additional warning of predator approach (Downhower and Wilson
1973; Richards 1978). Here I report a case of predation on a wasp colony
by the katydid Phlugis poecila Hebard (Orthoptera: Tettigoniidae). Com-
bined with observations of interactions of social wasps and green-faced
katydids, Ancistrocercus inficitus (Walker), roosting near their nests, and
with the results of experimental removal of adult Polistes wasps from
nests with katydids roosting nearby, these observations suggest that an
important component of the association is predation on wasp broods
by katydids.

Nest predation observations

On 3 August 1990 at 1200h, I discovered a small, newly-founded (pre-
adult emergence) nest of Polistes versicolor (Olivier) (Hymenoptera: Ves-
pidae) 2 m above the ground under a Heliconia sp. leaf at the edge of a
clearing nearGamboa, Republic of Panama. The nest was attended by a
single female wasp, presumably the foundress, during observations. A
female tettigoniid, Phlugis poecila, was resting on the same leaf at a dis-

1 Received May 16, 1992. Accepted July 27, 1992

~ Department of Entomology, University of Wisconsin. Madison, WI 53706

ENT. NEWS 1(M{1): 39-42. January & February. 1993

40 ENTOMOLOGICAL NEWS

tance of 5 cm when the nest was discovered. Phlugis poecila is predaceous,
though little else is known of its behavior (D. Nickle, pers. comm.).

The P. poecila female approached the nest at approximately 5 min
intervals over the course of 45 min following nest discovery. On each
approach the tettigoniid remained at a distance roughly equal to her
antennal length and gently touched both the nest and adult wasp with
the distal ends of her antennae; whenever her antennae contacted the
wasp the katydid retreated to a distance of between 10 and 30 cm from the
nest and remained motionless until the next approach. The wasp made
no obvious response to the tettigoniid throughout this period. The wasp
departed for 4 minutes on a foraging trip, but the katydid did not
approach during this time.

After 50 min of observation I removed the wasp from the nest and
placed her in a sealed container. The tettigoniid approached and anten-
nated the unoccupied nest 5 min later, then retreated 3 cm. After 4 min
the tettigoniid returned, antennated the nest, then approached and
touched the nest petiole and the bases of the cells with her labial and
maxillary palpi. The tettigoniid climbed onto the nest and touched the
brood cell entrances with her palpi before climbing back to the base of
the nest. At this point she held the nest with her forelegs and chewed at
the bottom of one brood cell near its attachment to the nest petiole. This
chewing continued for 10 min, at the end of which I collected the nest
and the tettigoniid.

The damage to the wasp nest was clearly visible at 8X magnification
as holes chewed into brood cell bases. A total of three cells bore similar
damage, suggesting that the nest had been attacked before my obser-
vations. All cells contained larvae, however.

Nest predation experiment

I surveyed 45 active P. instabilis de Saussure colonies for the presence
of katydids roosting within 1 m of their nests on 8 July 1992 at Palo Verde
National Park, Costa Rica. Green-faced katydids were found near 7 1 % of
the nests surveyed (Xnumber katydids present = 2.96, sd = 3.45, range =
Oto 17); over 90% of the katydids were roosting within 10 cm of the nests.
The number of adult wasps occupying the nests at the time of the survey
ranged from 2 to 1 1 (X = 5.2, sd = 2.28). None of the wasp nests bore
damage, for example holes or tears in brood cell walls, which might sug-
gest attempted katydid predation.

Downhower and Wilson (1973) noted that a green-faced katydid
roosting near a Polistes sp. nest in Costa Rica fed on wasp pupae after the
nest had been moved and the adult wasps had been removed. I removed
all adult wasps from five P. instabilis colonies on 13 July 1992 between

Vol. 104, No. 1, January & February, 1993 41

1930h and 2030h, after nearby katydids had left to forage, to test the
hypothesis that brood in unguarded wasp nests would be preyed upon by
katydids. I did not otherwise manipulate the nests or katydids. At 600h
the following morning 4 to 12 green-faced katydids were present at four
of the treated nests. Brood cells in these four nests were apparently
chewed open on the sides and their contents partially removed (number
of cells damaged ranged from 6 to 15). No katydids were present at the
remaining nest, which was undamaged. Four surveys of the manipulated
nests were conducted over the ensuing 36 hours. The number of cells
damaged increased on the four nests with katydids present, and katydids
were observed chewing through cells and consuming brood on each of
these nests. The nest without katydids present was not damaged.

Downhower and Wilson (1973) found that A. inficitus were able to
relocate two Polistes sp. nests in the morning after the nests were dis-
placed 1 m from their original site the previous night, and that the nest
petioles were especially attractive to the tettigoniids; the adult wasps
placed in a nearby cage were not attractive. These results, combined with
my observations, suggest that some tettigoniid species are capable of dis-
tinguishing wasp nests using chemical or tactile cues, and that the tet-
tigoniids can discriminate between the nest and the adult wasps using
these cues. In the case of P. poecila, substrate-borne cues were apparently
employed in discriminating between the nest and the adult wasp.

Wasp responses to tettigoniids

Ancistrocercus inficitus often perch on twigs within 10 cm of wasp
nests, adopting the cryptic posture described for other neotropical
orthopterans (Robinson 1969). My observations in Guanacaste, Costa
Rica in 1988 and 1991 showed \ha\.A. inficitus roosting near eusocial wasp
(Polistes instabilis and Polybia occidentalis [Olivier]) colonies were at-
tacked and occasionally driven away after being detected by wasp
workers. Though roosting tettigoniids often remained motionless for
several hours, initial detection was probably visual, since the wasps often
oriented to and approached the tettigoniids following movement. Wasp
workers that contacted tettigoniids with their antennae raised their
wings, elevated their bodies, and bit the intruders repeatedly (observed
on several occasions at one Polistes instabilis nest in 1988 and at two
Polybia occidentalis nests in 1991). When attacked by wasps, A. inficitus
slowly raised the body part being bitten and only moved away if the wasp
worker was especially persistent. A similar response to tettigoniids was
noted at a Synoeca sp. colony in Costa Rica (Downhower and Wilson
1973). Social wasps are not tolerant of roosting tettigoniids, though this
may be a general response by workers to foreign insects near the nest.

42 ENTOMOLOGICAL NEWS

Roosting near wasp colonies may afford protection to predaceous tet-
tigoniids, but it is unclear which potential predators of tettigoniids would
be repelled by the wasps (Downhower and Wilson 1973). Both wasp
nests and green-faced katydids are fed to nestling trogons (Trogon sp.) by
their parents in Guanacaste (F. Joyce, pers. comm.).

Association with wasp nests during the daylight hours maximizes the
probability that the nest will be unoccupied or occupied by few wasps,
since social wasp foragers return to and cluster on their nests at night.
This is especially true in relatively young colonies of independent-
founding wasps such as Polistes spp., however, even a small number of
adult wasps present on the nest can apparently deter katydid predation.
Further observations of tettigoniids roosting near small wasp colonies
are needed to establish the frequency of predation on unmanipulated
nests and the impact of tettigoniid predation on wasp colony success.

ACKNOWLEDGMENTS

Thanks are extended to Donald Windsor who served as my Smithsonian research
advisor during field work in Panama. Ann Eraser, Bruce Howlett, John Lill, Karen Lon-
don, and Adrienne Nicotra of Organization for Tropical Studies course 92-3 provided valu-
able suggestions and assisted ably with the wasp nest survey in Costa Rica. David Nickle
identified the tettigoniid from Panama and supplied useful background information on its
habits. Robert Jeanne, Tom Phillips, and two anonymous reviewers read and commented
on an earlier version of the manuscript. Financial support was provided by a Short Term
Fellowship from the Smithsonian Tropical Research Institute, a grant from the University
of Wisconsin-Madison Graduate School, and an Organization for Tropical Studies/Pew
Charitable Trust Tropical Fellowship.

LITERATURE CITED

Downhower, J.F. and D.E. Wilson. 1973. Wasps as a defense mechanism of katydids.

Am. Midi. Nat. 89:451-455.
Richards, O.W. 1978. The social wasps of the Americas. Brit. Mus. (Nat. Hist.), London.

580 pp.
Robinson, M.H. 1969. Orthopteroid defensive behaviour in Panama. Trans. R. Ent. Soc.

Lond. 121:281-303.

Vol. 104, No. 1, January & February, 1993 43

RECORDS OF BAT FLIES FROM JORDAN,
LIBYA AND ALGERIA 1

Z. S. Amr, M. B. Qumsiyeh^

ABSTRACT: Eight species of bat flies (Insecta: Diptera: Streblidae and Nycteribiidae)
collected from bats from Jordan, Libya and Algeria are listed along with their respective
hosts. Brachytarsina flavipennis, Stylidia biarticulata, Stylidia Integra and Basilia nana are
new records for Jordan.

In the course of extensive collecting throughout Jordan, Libya and
Algeria in 1981, several species of ectoparasites were removed from bats.
Although the Chiroptera of Jordan and North Africa have been studied
(Qumsiyeh 1980, Qumsiyeh el al. In press, Hufnagl 1972, Hayman and
Hill 1971), little information is available on their associated bat flies.

The only record of a bat fly from Jordan was reported by Kock and
Nader (1979). Anciaux de Faveaux (1976) provided a list of parasitic
insects from Algerian bats and Hurka (1982) reported on the bat flies of
coastal Libya.

This paper provides additional records of bat flies from Libya,
Algeria and Jordan.

MATERIALS AND METHODS

Bats were collected by mistnet or while roosting in caves or crevices.
A total of 63 bats were collected: 12, 15 and 36 from Jordan, Libya and
Algeria respectively. Each bat was individually examined and their
ectoparasites were stored in vials containing 75% alcohol, "n" designates
the number of bats examined.

Systematic List

Family Streblidae

Brachytarsina flavipennis Macquart 1851

Material examined:

Jordan: Wadi Khanzaireh( W.Araba). 9.2. 1981, ex 2M,Rhinolophusblasii

(n = 7).

Libya: Kuf National Park, 13.3.1981, ex 6M, Khinolophus mehelyi (n == 9).
Algeria: Misserghin Cave, 7.5.1981, ex IF, Myotis blythi (n = : 10).

1 Received May 4. 1992. Accepted August 26, 1992.

2 Department of Biology, Jordan University of Science and Technology. Irbid. Jordan.

ENT. NEWS 104(1): 43-46. January & February. 1993

44 ENTOMOLOGICAL NEWS

Remarks: This species has been reported from Miniopterus schreibersi
and several species of the genus Rhinolophus (Theodor 1967). Hurka
(1982) indicated its presence on R. mehelyi.

Family Nycteribiidae
Stylidia biarticulata (Harmann 1864)

Material examined:

Jordan: Wadi Khanzaireh (W. Araba), 9.2.1981, ex 2M, Rhinolophus

blasii (n = 7).

Libya: Kuf National Park, 13.3.1991, ex 1M, R. mehelyi (n = 9).
Algeria: Misserghin Cave, 7.5.1981, ex 1M, 2F, Myotis blythi (n = 10).

Tipasa, 5.7.1981, ex 1M, IF, Miniopterus schreibersi (n = 2).

Tipasa, 5.7.1981, ex 1M, R. mehelyi (n = 1).

Remarks: Theodor (1967) reported Stylidia biarticulata from Rhinolophus
euryale, R.ferrumequinum, R. hipposideros minimus, R. blasii, Myotis myotis
and Miniopterus schreibersi. Hurka ( 1 982) considered Stylidia biarticulata
to be a westpalearctic species.

Stylidia integra (Theodor and Moscona 1954)

Material examined:

Jordan: Dibbin National Park, 27.2.1981, ex 1M, Rhinilophus hip-
posideros (n = 1).

Remarks: Kock and Nader (1979) commented on the distribution of 5 1 .
integra and they suggested this species represents a saharosindian
faunal element. It is known from Rhinolophus blasii and R. acrotis
(Theodor 1965).

Basilia nana Theodor and Moscona 1954

Material examined:

Jordan: Dibbin National Park, 27.2.1981 and 9.8.1981, ex 1M, 3F,
Myotis nattereri (n = 1 ).

Remarks: It seems that Basilia nana is host-specific for the genus Myotis.
Theodor (1965) reported Myotis nattereri and M. myotis as hosts for
Basilia nana.

Basilia daganiae Theodor and Moscona 1954

Material examined:

Libya: KufNational Park, 15-16.3. 1981, ex, \M,3,Ptpistrelluspipistrellus

(n ---- 5).

8 km SSEHaniya, 3.4.1981, ex 1M, P. pipistrellus (n = 1).

Vol. 104, No. 1, January & February, 1993 45

Remarks: Theodor (1965) reported that Basilia daganiae parasitized
Pipistrellus Kuhlii in Deganya.

Penicillidia dufouri (Westwood 1835)

Material examined:

Algeria: Misserghin Cave, 7.5.1981, ex 4M, 3F Myotis blythi (n = 10).
Cap Aokas, 23.6.1981, ex 1M, 3F, M. blythi (n = 3). "
Tipasa, 5.7.1981, ex 1M, Miniopterus schreibersi (n = 2).
Tipasa, 5.7.1981, ex 1M Rhinolophus euryale (n = 11).

Remarks: Rhinolophus hipposideros was reported as host of this bat fly
(Theodor 1967).

Nycteribia pedicularia Latreille 1796

Material examined:

Algeria: Misserghin Cave, 7.5.1981, ex IF, Myotis blythi (n = 10).

Remarks: Rhinolophus hippisideros minimus, R. euryale and Miniopterus
schreibersi are known hosts for Nycteribia pedicularia (Vermeil 1960,
Theodor 1967).

Nycteribia schmidlii Schiner 1853

Material examined:

Algeria: Misserghin Cave, 7.5.1981, ex 1M, 3F, Miniopterus schreibersi

(n = 8).

Sig cave, 9.5.1981, ex 1M, 3F, M. blythi (n = 1).

Tipasa, 5.7.1981, ex 2M, IF, R. euryale (n = 1).

Tipasa, 5.7.1981, ex 2M, 6F, Miniopterus schreibersi (n = 2).

Remarks: In addition to the above species, Nycteribia schmidlii has been
taken from Rhinolophus ferrumequinum and Mvotis mehelvi (Fa\coz 1923,
Theodor 1967).

ACKNOWLEDGMENTS

We are grateful to T. C. Maa for his help in species determination and D. Kock for pro-
viding reprints on bat flies. This work was supported in part by a grant (No. 26/92)
from the Deanship of Academic Research (J. U. S. T.)

LITERATURE CITED

Anciaux de Faveaux, M. 1976. Distribution de chiropteres en AJgerie avec notes
ecologiques et parasitologiques. du Bulletin de la Societete d'Histoire Naturelle de
1'Afrique du Nord, 67: 69-80.

Falcoz, L. 1923. Biospeologica n 49. Diptera Pupipara ( 1 Serie). Arch.. Zool. Exper. Gen.,
61: 521-552.

46 ENTOMOLOGICAL NEWS

Hayman, R. and Hill, J. 1971. The Mammals of Africa, an Identification Manual.

Part 2. Order Chiroptera. Washington, Smithsonian Inst. Press. 73 pp.
Hufnagl, E. 1972. Libyan-Mammals. The Oleander Press. London. 88 pp.
Hurka, K. 1982. On the insect bat ectoparasites of coastal Libya. Cimicidae, Nycteribiidae,

Strebiidae and Ischnopsylidae. Vestn. Cesk. Spol. Zool. 46: 85-91.
Rock, D. and Nader, I. 1979. Two bat flies from the kingdom of Saudi Arabia, their

nomenclature, host specificity and zoogeography (Insecta: Diptera: Nycteribiidae).

Senckenbergiana bio., 60: 65-73.
Qumsiyeh, M. B. 1980. New records of bats from Jordan. Saugetier Kdl. Mitt.. 28:

36-39.
Qumsiyeh, M. B., Disi, A. M. and Amr, Z. In Press. Systematics and distribution of the

bats (Mammalia: Chiroptera) of Jordan. Dirasat.
Theodor, 0. 1967. An Illustrated Catalogue of the Rothschild Collection of Nycteribiidae.

The British Museum. Publication No. 655: 1-506. London.
Theodor, O. and Moscona, A. 1954. On bat parasites in Palestine. I. Nycteribiidae.

Strebiidae, Hemiptera, Siphonaptera. Parasitology, 44: 157-245.
Vermill, C. 1960. Contribution a 1 etude des Nycteribiidae et des Strebiidae de Tunisie.

Ann. Parasit. Hum. Comp., 35: 737-743.

SOCIETY MEETING OF NOVEMBER 18, 1992
ECOLOGY OF PAVEMENT ANTS

Mr. Thomas King,

Peerless Pest Control

Philadelphia, PA

We are all well aware of the detrimental effects of increasing urbanization and develop-
ment on the natural environment, including the loss of diverse habitats and the numerous
insect denizens which occur there. The informative and humorous presentation by Mr.
Thomas King, drawing upon his own observations and those of others, reminds us that
there are numerous insects, among these the pavement ant, Tetramorium caespitum (Lin-
naeus), who find in urbanization an opportunity for range expansion and population
growth.

The pavement ant, although probably evolved in Europe or Africa, is now found scat-
tered throughout the world, including diverse places as Belize, Chile, North America and
Australia. Although possibly arriving in North America with the early European colonists,
it continues its rapid spread at a local level even today. For example, a monograph on the
ants of Colorado in the early 1960's stated that Tetramorium caespitum was absent from
Denver; during a recent trip to Denver, Mr. King found the ant abundant everywhere he
looked. The success of the pavement ant in cities is most probably related to the habitat in
which it originally evolved, i.e., open areas of scarce, low vegetation. It is equally at home in
the wall-to-wall pavement of any major city (thus its apt common name), or in the lawns of
the suburbs.

Mr. King's interest in ants in general extends back to age 7. More recently, his ant
research has included one year studying the fire ant at Texas Tech University, and con-
tinues with observations on interactions between the introduced Tetramorium caespitum
and other ant species in the Philadelphia area. Surprisingly, although the pavement ant is
well adapted to the urban and disturbed environment, it does face severe competition from

Continued on page 52

Vol. 104, No. 1, January & February, 1993 47

A CHAMBER FOR MASS HATCHING AND

EARLY REARING OF PRAYING MANTIDS

(ORTHOPTERA: MANTIDAE) 1

Frederick R. Prete, Robert J. Mahaffey^

ABSTRACT: A rugged, easily maintained polycarbonate and lucite chamber for the mass
hatching and early rearing of praying mantises is described.The chamber is sealed after
eggs are introduced so that even the smallest prey (e.g., Drosophila) cannot escape. An
aquarium pump forces fresh air through a water bottle into the sealed chamber. Gas
exchange and the introduction of prey, food, and water occur through several holes that are
plugged with foam rubber. If necessary, the temperature of individual chambers can be
raised above ambient by placing an incandescent light bulb at the appropriate distance.

The praying mantis' dramatic method of prey capture and the rela-
tive ease with which the predatory strike can be elicited in the laboratory
have made this insect an important investigatory tool for the study of
visually guided behaviors (e.g., Barnes, 1979; Barnes and Mote, 1980;
Collet, 1987; Liske and Mohren, 1984; Kirmse, 1985; Horridge, 1986;
Rossel, 1986; Prete, 1991; Prete, 1992a, b; Prete, et al, 1992a, b).

In spite of the popularity of mantises among professional and
amateur entomologists, information regarding methods of mass rearing
and maintaining these insects remains primarily anecdotal. A few infor-
mative reports on small scale rearing have appeared (e.g.. Heath, 1980),
but problems unique to those rearing large numbers of mantises remain
unaddressed in the literature. Here, we describe a chamber in which egg
cases can be incubated and young (e.g., up to fourth or fifth instar
Tenodera or Sphodromantis) mantises can be easily raised prior to being
placed in individual containers. The chamber solves several of the prob-
lems that mantis breeders face for instance: maintaining high
humidity, maintaining high prey densities, feeding the introduced prey,
and preventing the escape of small prey.

MATERIALS AND METHODS

The chambers are easily built and, with reasonable care, have an
indefinite life. The materials needed for the construction of one chamber
are these (measurements are given in inches for appropriate items): i) a
polycarbonate rodent cage (51 x 41 x 22 cm; Fisher Scientific); ii) one
piece of clear lucite (53 x 42 x 0.64 cm); iii) eleven foam rubber plugs ( 3.5

1 Received June 27, 1992. Accepted July 18, 1992.

2 Department of Psychology, Youngstown State University, Youngstown, Ohio 44555.

ENT. NEWS 104(1): 47-52. January & February. 1993

48 ENTOMOLOGICAL NEWS

x 4 cm; e.g., standard Drosophila vial plugs); iv) 42 cm long x 3.8 cm (1.5
inches) wide aluminum piano hinge; v) 42 cm long x 1 .3 cm (.5 inch) wide
aluminum angle; 2.5 cm long ( 1 inch) 6-32 round head bolts, 1 .6 cm long
(.625 inch) 10-32 Hat head bolts, 1.6 cm long (.625 inch) 6-32 Oat head
bolts, appropriate washers and nuts. A finished chamber is pictured in
Fig. 1.

The first step is to cut the lucite into two sections across its width. One
piece will serve as a door, the other as the permanently attached portion
of the top. The width of the door can be varied to suit individual prefer-
ences but we suggest cutting the lucite into two approximately equal sec-
tions. This allows sufficient access to the chamber interior when re-
moving mantises and cleaning without having to remove the entire top.
The two pieces of lucite are connected with the piano hinge on the out-
side of the chamber. If the hinge is not pre-drilled, the 10-32 fiat head
screws are spaced 10cm between centers with the heads recessed into the
lucite. The aluminum angle is attached to the lip of the door with the 6-32
flat head screws, also spaced 10cm between centers and recessed into the
lucite. Fewer screws should not be used in that one purpose of the hinge
and angle is to prevent the lucite from warping. If the two top pieces do
not fit snugly under the hinge, the gap can be filled with a strip of .64 cm
(.25 inch) wide adhesive foam rubber weather stripping (e.g., Door &
Window Weather Strip, Macklanburg-Duncan, Oklahoma City, OK).
The top is affixed to the bottom with the 6-32 round head screws. Holes
should be drilled through the top such that the screws pass through the
center of the rolled lip of the box; The 6-32 nuts will fit snugly within the
lip. Holes should be drilled for four bolts on each side (two on each side
of the top and two on each side of the door) and for three along the back
edge of the top. This is the minimum number sufficient to prevent the top
from warping.

Seven 3.2 cm (1.25 inch) holes are drilled in the sides of the box, and
four in the top (see Fig. 1 for placement). Foam rubber plugs are inserted
into each hole. Standard Drosophila vial plugs work well; however; larger
holes may be drilled as long as the plugs used fit snugly. Prey, and food
for the prey are introduced through these holes.

Fresh humid air is supplied to the chamber by a standard aquarium
pump attached to an 8 oz (237 ml) polyethylene wash bottle (Carolina
Biological) containing water. After clipping off the thin tip to increase air
flow, the nozzle of the bottle is inserted into the chamber through the
center of one of the foam rubber plugs. Any mid-sized pump (x;2500 cc/
min at 4 PSI) is sufficient to aerate two chambers. If chambers become
too humid, any number of foam rubber plugs can be replaced with
square pieces of fine screen taped over the holes and/ or the wash bottle

Vol. 104, No. 1, January & February, 1993

49

<u

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50 ENTOMOLOGICAL NEWS

can be emptied. Internal chamber temperature can be monitored by an
aquarium thermometer and, if necessary, the temperature can be raised
above ambient by placing an incandescent light bulb at the appro-
priate distance.

Perch sites for the mantises are supplied by a continuous (2.5 meter)
length of 15 cm high heavy (_> 5 x mm) plastic mesh folded back and
forth inside of the chamber (e.g., Co-Polymer Gutter Guard, Allumax
Home Products, Lancaster, PA).

RESULTS AND DISCUSSION

Prior to hatching, egg cases are hung from the plastic mesh with wire
hooks and the door is bolted closed. Generally, we incubate simul-
taneously four to eight egg cases of about the same age in one chamber.
Although the chamber is relatively easy to use, precautions are
necessary.

For over two hundred years, those who have written about keeping
mantises have continually reminded their readers that cannibalism can
be held to a minimum by supplying the mantises with sufficient prey
(Prete and Wolfe, 1992). This is easily done with a chamber from which
prey cannot escape and, if fed, survive well until eaten. For early instars,
we suggest the following procedure: just before the mantises hatch and
prior to bolting the chamber lid closed, place a jar (approximately 50 x
1 10 cm) half filled with commercial Drosophila food on its side under one
of the foam rubber plugs. Placing the jar on its side prevents mantises
from becoming entrapped in the food. Shortly after the first hatching
(and then as needed) anesthetized flies can be added by pouring them
through a funnel inserted into one of the holes in the top of the chamber.
If the foam rubber plug is kept in place during the procedure and is just
pushed aside by the funnel's spout, flies and mantises cannot escape.
When the jar needs refilling, it can be righted easily by means of a long
sturdy wire with 1.5cm of the tip bent at a right angle. The wire is inserted
through a hole in the top of the chamber (with the plug kept in place), and
hooked under the lip of the jar. Then, with the jar's base pushed against
the side of the chamber, it is pulled upright and slid, if necessary, directly
under the foam rubber plug. With practice, this procedure takes only
seconds. Once upright, the jar can be refilled with Drosophila food
through a funnel and returned to its side with the wire hook.

Obviously, other prey, such as crickets of any size, can be introduced
into the chamber through a funnel of appropriate size. Prey can be sup-
plied with slices of vegetables impaled on a thin stainless steel wire that is
bent at the tip and inserted through one of the holes in the top. The wire

Vol. 104, No. 1, January & February, 1993 51

should extend sufficiently far beyond the top to prevent it from falling
into the chamber. Although crickets can be maintained on just vege-
tables with high water content, such as potatoes, if they become thirsty or
hungry they will prey on the mantises. To avoid this problem, we also
supply the crickets with powdered laboratory rodent food and fresh
water. The former is simply poured through a funnel into the chamber.
Water is supplied in a slice of wet sponge impaled on a thin stainless steel
wire as is done with vegetables. The sponge should not be so wet that it
loses water into the chamber. We have found it best to place a small plas-
tic dish into which the crickets can climb (such as a small jar lid) under
the sponge to keep excess water off the chamber floor. This can be done
by placing the dish under the hole through which the sponge will be
inserted before the lid is bolted down or by first threading the wire on
which the sponge is impaled through the center of the dish. Obviously, if
the latter method is used, a hole large enough to accept the dish has to be
cut in the lid.

Once mantises reach approximately the sixth instar (depending on
species), we transfer them into aquaria with screen tops; immediately
after their final molt, they are placed in individual containers.

ACKNOWLEDGMENTS

This research was supported in part by a University Research Council Grant from
Youngstown State University. We thank S. P. Grossman of The University of Chicago, and
B. Greenberg of the University of Illinois at Chicago forcritically reading an earlier version
of this manuscript. We also thank our referees and editor for their thoughtful criticisms and
suggestions.

LITERATURE CITED

Barnes, S. N. 1979. The visual system of the mantis Tenodera aridifolia sinensis. Invest.

Ophthalmol. Vis. Sci. Suppl.: 277-278.
Barnes, S. N., and Mote, M. I. 1980. Lamina mono polar cells of the praying mantis

Tenodera-aridifolia: Response pattern and receptive fields. Invest. Ophthalmol. Vis. Sci.

Suppl.: 88.

Collett, T. S. 1987. Binocular depth vision in arthropods. TINS. 10: 1-2.
Copeland, J., and Carlson, A, D. 1977. Prey capture in mantids: Prothoracic tibial flexion

reflex. Jour. Insect. Physiol. 23: 1151-1156.
Corrette, B. J. 1980. Motor control of prey capture in the preying mantis, Tenodera aridifolia

sinensis. Ph.D. dissertation, Univ. of Oregon 217 pp.
Heath, G. H. 1980. Rearing and studying the praying mantids. Amat. Entomol. Soc. Leaf.

36: 1-15.
Horridge, G. A. 1986. A theory of insect vision: velocity parallax. Proc. R. Soc. Lond. B.

229: 13-27.

Kirmse, W. 1985, Short communication. Visual position information controlling smooth-
tracking in the praying mantis. Jour. Exp. Biol. 1 19: 365-367.
Liske, E., and Mohren, W. 1984. Saccadic head movements of the praying mantis, with

52 ENTOMOLOGICAL NEWS

particular reference to visual and proprioceptive information. Physiol. Entomol. 9:

29-38.
Prete, F. R. 1990. Prey catching in mantids: The role of the prothoracic tibial flexion reflex.

J. Insect Physiology, 36: 335-338.
Prete, F. R. 1991. Configurational prey selection by the praying mantis., Sphodromantis

lineola (Burr.): The effects of size and direction of movement. Brain, Beh. Evol. 36:

300-6.
Prete, F. R. 1992a. The effects of background pattern and contrast on prey discrimination

by the praying mantis Sphodromantis lineola (Burr). Brain. Beh. Evol. (in press).
Prete, F. R. 1992b. The discrimination of visual stimuli representing prey versus non-prey

by the praying mantis, Sphodromantis lineola (Burr). Brain, Beh. Evol. (in press).
Prete, F. R. and M. M. Wolfe 1992. Religious supplicant, seductive cannibal, or reflex

machine? In search of the praying mantis. J. Hist. Biol., 25: 91-136.
Prete, F. R., H. Lum, and S. P. Grossman 1992a. Non-predatory ingestive behaviors of the

praying mantis Sphodromantis lineola (Burr.) Brain. Beh. Evol. 39: 124-32.
Prete, F. R., C. A. Klimek, and S. P. Grossman 1990a. The predatory strike of the mantis,

Tenodera aridifolia sinensis (Sauss.). J. Insect Physiol. 36: 561-565.
Prete, F. R., P. J. Placek, M. A. Wilson, R. J. Mahaffey and R. R. Nemcek 1992b. The

Effects of Stimulus Speed and Order of Presentation on the Discrimination of Visual

Stimuli Representing Prey by the Praying Mantis Sphodromantis lineola (Burr.), Brain,

Behav. Evol. (in press).
Rossel, S. 1986. Binocular spatial localization in the praying mantis. Jour. Exp. Biol. 120:

265-281.

Continued from page 46

many species it did not initially evolve with, including native species of Pheidole and
Monomorium ants, and even small rodents like mice and rats. For example, T. caespitum
won't forage in bright sunlight and the resultant heat, therefore limiting its foraging times
in warm areas to night and morning hours; some species of ants can forage at much higher
temperatures during the day, and mice probably compete with the pavement ant for food
during the nocturnal periods. Other factors which may impact the competitiveness of
T.caespitum are the large nest sizes and foraging areas it maintains (nests may be up to 7
square meters, foraging areas may be up to 40 square meters), its tolerance of other ant
species in its area and its high investment in reproductives (up to 50% of the colony's energy
may go toward reproductives).

There were several notes of local entomological interest preceding Mr. King's talk. Con-
cerning the widely publicized decline in monarch butterfly populations due to severe cold
and logging in their overwintering forests in Mexico, Dale Schweitzer suggested that local
factors may have also played an additional factor. He reported that population levels in
Cumberland Co., New Jersey appeared to be building in July, but never appeared as a flush
of adults in August as expected, possibly due to cooler summer temperatures and/or dis-
ease. Mildred Morgan stated that numbers of monarchs tagged at Cape May. New Jersey
by Jane Ruffin and herself was one-tenth that of the previous year. Barbara Kirschenstein
reported on small flies (family Phoridae) attracted to iodized salt. Society president Joe
Sheldon urged everyone to attend the traveling insect exhibition, "Backyard Monsters" at
the Franklin Institute, Philadelphia, which features monstrous robotic insects, a mar-
velous collection of OH MY! insects from around the world, interactive exhibits and an
operational scanning electron microscope. The meeting at the Academy of Natural
Sciences was attended by 27 members and their guests.

Jon K. Gelhaus,
Corresponding Secretary

Vol. 104, No. 1, January & February. 1993 53

ELMIDAE OF TAIWAN PART II:

REDESCRIPTION OF LEPTELMIS FORMOSANA

(COLEOPTERA: DRYOPOIDEA) 1

Ming-Luen Jeng, Ping-Shih Yang^ 3

ABSTRACT: Leptelmisformosana is the only member of the genus known from Taiwan. It
is redescribed and the male genitalia and other characters are illustrated. Because of
similar male genitalia but somewhat different external morphology, we regard Leptelmis
vietnamensis from Vietnam as a subspecies of L. formosana. A key is modified from Brown
and Thobias (1984) to include all known Leptelmis species of Asia.

The genus Leptelmis Sharp was reviewed recently by Brown and
Thobias (1984). More than twenty species are known from Asia and
Africa. Leptelmis formosana Nomura is the only species known from
Taiwan. Nomura described this species in 1962 based on two adults
collected by Yano in 1938. The descriptions only pointed out the dif-
ferences between L. formosana and L. parallela Nomura from Japan.
Brown and Thobias omitted these two species in their key to Asian
species of Leptelmis since the original diagnoses were too ambiguous to
separate them from L. gracilis Sharp from Japan. Actually, L. gracilis is
quite distinct in elytral shape. Because its humeri are not prominent and
the elytra are very broad at their apical 1/3, the body looks expanded pos-
teriorly. Both L. formosana and L. parallela have prominent humeri and
their elytra are subparallel-sided (Fig. 1 ). However, it is necessary to note
that the wing polymorphism may accompany morphological change of
pronotum and elytra (Deleve 1945; Brown, personal communication).

While examining the insect collections of Taiwan Agricultural
Research Institute (TARI), three specimens of Leptelmis formosana
were found. We redescribe and illustrate the species here. In addition, we
regard Leptelmis vietnamensis Deleve from Vietnam as a subspecies of
L. formosana due to its similar male genitalia but somewhat different
external morphology. A key to all known Asian species, modified from
Brown and Thobias (1984), is provided to include L. formosana and
L. parallela .

The following description of coloration is based on alcoholic
specimens viewed under a white light source. Body length is measured
from apex of pronotum to apex of elytra.

1 Received July 3, 1992. Accepted July 30, 1992.

2 Laboratory of Insect Conservation, Department of Plant Pathology and Entomology,
National Taiwan University, Taipei, Taiwan 10764, R.O.C.

3 This study was supported by the National Science Council, Republic of China, Grant
No. NSC 82-0409-B-002-053.

ENT. NEWS 104(1): 53-59. January & February. 1993

54

ENTOMOLOGICAL NEWS

1 mm

Fig. 1. Leptelmis formosana formosana Nomura, dorsal aspect.

Vol. 104, No. 1, January & February, 1993

55

Leptelmis formosana formosana Nomura

Leptelmis formosana Nomura, 1962, Toho Gakuho 12:48.

: Brown and Thobias, 1984, Pan-Pacific Entomol. 60(1 ):28.

Length 2.3 2.4 mm, width 0.8 0.9 mm. Body elongate, subparallel-sided, convex
dorsally. General coloration brown, with elytra feebly shining. Epicranium darker than
other portions; venter lighter than dorsum; antennae, palpi, tarsi and genitalia translucen-
tly testaceous.

Head retractable within prothoracic collar; visible portion finely granulate and pubes-
cent. Vertex concave at middle, impressed in a band on each side toward antennal base.
Frons convex at middle, about 3/5 as broad as width across eyes. Eyes rather large. Fronto-
clypeal suture indistinct. Labrum transverse, anterior margin feebly truncate with frontal
angle round. Antennae 11-segmented, barely reaching pronotal base; apex of distal seg-
ment acute.

Pronotum longer than broad by about 1 . 1 times; widest at basal 2/5, thence subparallel
posteriorly to base; narrowest at middle transverse impression. Anterior pronotal margin
arcuate and projecting over the vertex; anterior angles subacute, slightly protruding out-
wards; sides conspicuously bisinuate, not crenate; posterior margin feebly sinuate; basal
angles subquadrate. Surface finely and sparsely granulate anteriorly, coarsely and deeply
punctate at transverse impression and posterior portion. A subtriangular elevation located
behind the transverse impression; two upper tubercles of the elevation very prominent; the
lower tubercle smaller, with an indistinct ridge posteriorly; an indistinct impression com-
posed of some punctures extending from transverse impression to near lower tubercle of
the elevation. Two oblique, convergent grooves behind the subtriangular elevation deep,
with two oblique elevations posteriorly. Base with two small, round feeble impressions.

Scutellum fiat very sparsely granulate.

Elytra 2.4 times as long as pronotum; humeri prominent; sides subparallel in anterior
2/3, thence tapering posteriorly to a rounded apex; feebly depressed at base, but convex at
humeri. Each elytron bearing 9 punctate-striae; the 3rd and 4th striae merge on apical
declivity. Strial punctures on disk rather large, subquadrate, separated from one another
by less than half their diameters; punctures on apical declivity smaller and shallower.
Strial intervals on disk narrower than half diameter of punctures; the third interval (be-
tween 2nd and 3rd striae) elevated at base. Lateral borders feebly margined and finely
serrate. Epipleura narrowed gradually towards apex.

Fig. 2. Hind wing of L. formosana formosana.

56

ENTOMOLOGICAL NEWS

0.5mm

Fig. 3. Prosternum and mesosternum of L. formosana formosana.

Hind wing with venation as in Fig. 2. Venation essentially like that of the Cylloepus
(Hinton 1940, Figs. 251, 252); Veins 3Aj and 3A2 separated near base.

Prosternum coarsely punctate posteriorly. Prosternal process with sides subparallel or
slightly expanded posteriorly; apex truncate (Fig. 3).

Mesosternum with inconspicuous, blunt hind angles.

Metasternum with large, deep punctures; punctures separated at most by 1/2 their
diameters. Anterior position bare between mesosternum and longitudinal sulcus.

Abdomen with first two visible sterna with large, deep punctures similar to those on
metasternum; punctures on the last three sterna finer and sparser. Apex of last sternum
feebly truncate in males and round in females; males with two tufts of hairs and a semicir-
cular depression at the apex of 5th sternum.

Legs long and slender, with fine pubescence and sparse granules. Tibiae with in-
conspicuous rows of small setae present along innerdistal margins. Tarsi 5-segmented, that
of foreleg shortest and hindleg longest; segments progressively longer from base to apex;
apical segment as long as segments 1-4 combined, without ventroapical tuft of setae; claws
large, each with a basal tooth.

Male genitalia as shown in Fig. 4. It is noteworthy that while the genitalia are connected
ventrally with sternum IX the parameres are parallel-sided, but when sternum IX is
removed, the parameres expand outward as illustrated.

Variation: In one specimen the pronotum has its broadest width at base.

Specimens examined: Icf, Tamsui, Taihoku (Taipei Hsien), 24-VIII-1941, S. Miyamoto
leg.; 2 99, Heito (Pintong Hsien), V-1933, Y. Miwa leg. These specimens are deposited in
Department of Applied Zoology, TARI.

Distribution: The type locality of this species is Takezaki (Chuchi,
Chiayi Hsien). When more specimens become available we expect that
this species may be distributed from northern to southern Taiwan. At
present, the only known specimens are the two type specimens (in
National Science Museum, Tokyo) and the three specimens reported
here.

Vol. 104, No. 1, January & February, 1993

57

0.1mm

Fig. 4. male genitalia of L. formosana formosana.

Leptelmis formosana vietnamensis Deleve

Leptelmis vietnamensis Deleve, 1968, Ann. Hist.-Nat. Mus. Nat. Hung., Pars Zool. 60:154.
: Brown and Thobias, 1984, Pan-Pacific Entomol. 60(1):27.

Since the male genitalia of this taxon is so similar to that of L. for-
mosana, we regard it as a subspecies of the later. Compared with the
nominate subspecies, the pronotum of this subspecies has (1) a smaller
elevation; (2) more indistinct upper tubercles; (3) a longer ridge behind
the lower tubercle and (4) a conspicuous longitudinal impression at
anteromiddle of the elevation. Its body size is a little smaller than the
nominate subspecies. These differences are shown in Fig. 5a and b.

E
to

O'

a

Fig. 5a. Pronotum of L. formosana formosana; b. of L. formosana vietnamensis.

58 ENTOMOLOGICAL NEWS

Specimens examined: Holotype male, labelled "Vietnam, Prov. Ha-
Tinh, forestiere Huong-son, 1 50 m, foret trop. pluv./ a la lumiere, 1 5, VIII,
1963, T. Pocs./ Prepar. genit. No. 21266.2". Paratypes, Icf, 19, with identi-
cal data; Prepar. genit. No. 21266.1 and 21266.3 respectively.

The following key is modified from Brown and Thobias, 1984, to
include L. formosana and L. parallela .

Key to Asiatic Species and Subspecies of Leptelmis*

1. Elytra maculate 2

Elytra essentially uniform in color or with the humeri lighter 5

2. Each elytron with 5 yellow spots (Vietnam) L. signata

Elytra with fewer than 5 spots or markings 3

3. Elytra without vitta in apical half on intervals, with only humeral and apical markings;

larger (3.0 mm) (Sumatra) L. stricticollis

Elytra with both vitta, humeral and apical markings; smaller (less than 2.5 mm)
4

4. Elytra with strial interval 3 raised from base to apex; smaller (1.8 mm) (Philip-
pines) L. tawitawiensis

Elytra with strial interval 3 raised only at base; larger (2.3 mm) (Vietnam) . . .L. basalis

5. Elytra without prominent humeri 6

Elytra with prominent humeri 7

6. Elytra expanded posteriorly; venter punctate; larger (2.5-2.8 mm) (Japan). . . .Lgracilis
Elytra not expanded posteriorly; venter granulate; smaller (2.15 mm) (South India)

L. philomina

1. Tarsi 4-segmented (South China) L. flavicollis

Tarsi 5-segmented 8

8. Elytra with humeri paler 9

Elytra uniform in coloration; or if the elytra with humeral spot and /or paler 3rd strial
interval, the body size less than 2.5 mm 10

9. Larger (2.5-2.6 mm) (Japan) L. parallela

Smaller (2.0 mm) (North India) L. fracticollis

10. Elytra with strial interval 3 raised from base to apex 11

Elytra with strial interval 3 raised at base only or extending to apical 1/3 13

11. Male genitalia with parameressubparallel in apical half(Sumatra, Java) . . . .L.sulcata
Male genitalia with parameres tapering from base to apex 12

12. Penis subparallel laterally, longer than basal piece by 1.7 times (Sri Lanka)

L. cederholmi

Penis dilated at apical 1/4 and thence tapering basally, longer than basal piece by 1.5
times (Vietnam) L. obscura

13. Two upper tubercles of triangiriar elevation on pronotum very prominent; longitudinal
impression at anteromiddle of the pronotal elevation inconspicuous (Taiwan)

L. formosana formosana

Upper tubercles of triangular elevation on pronotum not very prominent; longitudinal

impression at anteromiddle of the pronotal elevation distinct (Vietnam)

L. formosana vietnamensis

* The species L. nietneri was transferred to the genus Podelmis by Jach (1984).

Vol. 104, No. 1, January & February, 1993 59

ACKNOWLEDGMENTS

We thank Department of Applied Zoology (Liang-Yih CHOU), Taiwan Agricultural
Research Institute, R.O.C. and Termeszetrudomanyi Miizeum ( Otto MERKL), Hungary,
for lending us the precious specimens. We also express gratitude to Harry G. NELSON
(Field Museum of Natural history, Chicago), Harley P. BROWN (Dept. of Zoology. Univ.
of Oklahoma, Norman, U.S.A.) and M. A JACH (Naturhistorisches Museum Wien, Aus-
tria) for revising the English manuscript.

LITERATURE CITED

Brown, H. P. and M. P. Thobias. 1984. World synopsis of the riffle beetle genus Leptelmis
Sharp, 1888, with a key to Asian species and description of a new species from India
(Coleoptera, Dryopoidea, Elmidae). Pan-Pacific Entomol. 60(1): 23-29.

Deleve, J. 1945. Contribution a 1'etude des Dryopidae. III. Le genre Pseudomacronychus
Grouvelle et le dimorphism alaire de ses especes. Bull. Mus. R. Hist. Nat. Belg.

Hinton, H. E. 1940. A monographic revision of the Mexican water beetles of the family

Elmidae. Novit. Zool. 42:217-396.
Jach, M. A. 1984. Die Koleopterenfauna der Bergbache von Sudwest-Ceylon. Arch.

Hydrobiol. Suppl. 69 (2): 228-332.
Nomura, S. 1962. Some new and remarkable species of the Coleoptera from Japan and its

adjacent regions. Toho Gakuho 12:35-51.

60 ENTOMOLOGICAL NEWS

BOOK REVIEW

A SYNTHESIS OF THE HOLARCTIC MIRIDAE (HETEROP-
TERA): DISTRIBUTION, BIOLOGY, AND ORIGIN, WITH EM-
PHASIS ON NORTH AMERICA. A. G. Wheeler, Jr. and T. J. Henry.
1992. Thomas Say Foundation, Vol. 15. Entomological Society of
America, Lanham, MD. 282 pp. $30 (members), $50 (non-members).

The authors have determined that about 5% (98 spp.) of New World mirids also occur in
the Old World, and thus appear to have Holarctic distributions. However, they present
good evidence that 61 of these species (3%) have been introduced into North America
through commerce, so actually only 37 spp. (2%) are likely Holarctic. Distinguishing these
two types of origins are important to those in the biological control, quarantine, and
biogeography fields.

Although the adults and nymphs of most mirid species are not hardy or long-lived,
diapausing mirid eggs imbedded in plant tissue can easily survive long ocean voyages, and
were the likely means of dispersion. Many mirids were not detected until years after their
initial establishment, as commonly happens with immigrant species. A number of them
were first found by one or both of the authors.

A total of 98 species are included in this book, arranged by subfamily. For each species,
the known distribution (with a map), host plant and habits, and zoography is discussed,
with references cited. Most species are phytophagous, but some are predators. Several are
economically important pests. Ten additional species formerly thought to be Holarctic are
discussed briefly.

Five tables follow the text. These list indigenous mirids (species present since the
Beringian land connection), and species which were introduced through eastern, north-
western southern, and multiple ports of entry. The tables place the species in functional
groups, but the contents (or index) must be used to locate the discussion on each species in
the text.

There are two indices, listing the common and Latin names of the mirid species and of
their host plants. There are also ca. 500 references, for those who wish further information.
However, this list is the most comprehensive for taxonomic papers, and is less complete for
economic and biocontrol citations, especially after 1986. There was a refreshing ab-
sence of misspellings and other errors throughout the book.

This small volume contains a wealth of information, it will be useful to taxonomists,
economic entomologists, and quarantine and biological control specialists.

W. H. Day
USDA-ARS-BIRL
Newark, DE

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USISSN0013-872X

NEWS

A new species of Aleodorus (Coleoptera:
Staphylinidae) from Costa Rica and
generic reassignment of Falagria

costariccnsis to Aleodorus

Egg surface ultrastructure in

Mantispa interrupta (Neuroptera:
Mantispidae)

E. Richard Hoebeke 61

Bruce Cutler 68

Reclarification of males of Alloperla
concolor and A. neglecta (Plecoptera:
Chloroperlidae), with new distribution
records for both species

B.C. Kondratieff, R.F. Kirchner

New records of spiders (Araneae)

from Cape Cod, MA, including two
possible European immigrants

First karyotypic data on a cupedid beetle
(Coleoptera: Archostemata) showing
achiasmatic meiosis J. Galian, J.F. Lawrence

An ecotonal study of carrion beetles
(Coleoptera: Silphidae) in the Great
Swamp N.W.R., NJ

Techniques for obtaining adult-associated

immature stages of predacious tachydromiine

flies (Diptera: Empidoidea), with implications

for rearing and biocontrol J.M. Cumming, B.E. Cooper

Establishment of Hippodamia variegata
and new records of Prop y/ea
quatuordecimpunctata (Coleoptera:
Coccinellidae) in the eastern U.S.

Epilachna vigintioctopunctata (Coleoptera:
Coccinellidae), new record for
Western Hemisphere, with a review
of host plants

R.F.W. Schroder. M.M. Athanas, C. Pavan

73

Robert L. Edwards 79

83

Paul P. Shu beck 88

93

A.G. Wheeler, Jr. 102

BOOKS RECEIVED & BRIEFLY NOTED

111

67, 72, 78

THE AMERICAN ENTOMOLOGICAL SOCIETY

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Vol. 104, No. 2, March & April, 1993 61

A NEW SPECIES OF ALEODORUS
(COLEOPTERA: STAPHYLINIDAE) FROM COSTA

RICA, AND GENERIC REASSIGNMENT OF
FALAGRIA COSTARICENSIS TO ALEODORUS l

E. Richard Hoebeke^

ABSTRACT: Aleodorus maureenae. new species of falagriine Staphylinidae, is described
from Costa Rica. Falagria costaricensis Bemhauer is redescribed and transferred to Aleo-
dorus. A lectorype is designated.

The genus Aleodorus was established by Say (1833) for the North
American species Aleochara bilobata, also described by Say that same
year. Chitalia was proposed by Sharp (1883) to accommodate four new
species (crenata, granigera, debilis, and dubius) collected at various
localities in Mexico and Central America. The latter genus was later
determined to be a junior synonym of Aleodorus (Fenyes, 1912).

Members of Aleodorus are restricted to the Western Hemisphere. At
present, four species are known to occur in America north of Mexico,
with the Nearctic species having been revised by Hoebeke ( 1985). Black-
welder (1944) lists five species from Mexico and Central America, and
one species from South America. Pace (1989, 1990) added 4 taxa to the
existing South American fauna by describing 3 new species from Argen-
tina, Peru, and Brazil, and by reassigning Falagria discisa Erichson
(Brazil) to Aleodorus. The Neotropical species have not been revised.

In March and April 1973, specimens of a distinctive, yet unrecog-
nized, species of Aleodorus were collected from Berlese samples of leaf
mold and leaf litter in virgin forest in Puntarenas and Guanacaste prov-
inces, Costa Rica, by J. Wagner and J. Kethley of the Field Museum of
Natural History (Chicago). Specimens of this new species, which I dis-
covered among recently prepared and unidentified Staphylinidae in the
Field Museum collection (FMNH), are described below.

Furthermore, after examination and dissection of syntypes of
Falagria costaricensis Bernhauer. I have found these to belong to the
genus Aleodorus and herein propose this reassignment.

MAY ? 1993

1 Received September 12. 1992. Accepted October 14. 1992

- Department of Entomology, Cornell University, Ithaca. New York

ENT. NEWS 104(2): 61-67, March & April. 1993

62 ENTOMOLOGICAL NEWS

Aleodorus maureenae, new species
(Figs. 1-6)

Diagnosis. In overall adult body size and in pronotal shape, Aleodorus maureenae
is similar to A. granigerus (Sharp), but differs most noticeably by the very roughened
surface of the head, thorax, and elytra caused by a dense, uniform covering of setiferous
asperities.

Description. Agreeing with generic characters given by Hoebeke (1985).

Length 2. 8-3.4 mm (n = 9,x = 3.2mm). Body color uniformly dark rufo-brunneous, with
distal antennal articles, mouthparts, and, in some specimens, last two abdominal segments
rufo-testaceous. Habitus as in Figs. 1-2.

Head (Fig. 1) quadrate, slightly longer than wide, posterior angles broadly rounded,
posterior margin slightly arcuate to truncate. Eyes moderately large, prominent, their long-
est diameter nearly subequal to length of temple. Dorsal surface with a dense, uniform
covering of asperities, each bearing a short, erect microseta; dorsum with narrow, median
area between posterior margins of eyes, and median frontal prominence between antennal
bases devoid of asperities smooth and glossy (see Fig. 1) (some specimens with smooth,
glossy area between eyes appearing as small dimple, or absent altogether); cuticular sur-
face between asperities smooth, glossy. Gena and ventral surface of head without as-
perities, smooth and glossy. Antennae moderately long, reaching anterior 0.4 of elytra;
distal articles beyond article IV compactly organized; scape somewhat incrassate, nearly
equal to length of article II; article II and III elongate, II slightly shorter than III; article IV
somewhat quadrate, but slightly longer than wide; articles V-X becoming gradually more
transverse; article XI obovate, slightly shorter than IX + X,

Pronotum (Fig. 1) slightly wider than head, broadest across anterior third, strongly
narrowed and converging behind towards base; posterior angles nearly acute; posterior
margin broadly truncate. Disc narrowly and deeply channeled along median line, channel
terminating in deep, subbasal fovea; surface on either side of channel densely and uni-
formly covered with setiferous asperities; cuticular surface between asperities smooth and
glossy. Scutellum large, flat, densely punctured (punctures minutely asperate), on either
side of a broad smooth, slightly impressed, median channel.

Elytra (Figs. 1-2) about as long as prothorax, humeri well developed, lateral margins
broadly arcuate posteriorly, posterior angles sinuate, posterior margin truncate; surface
with a dense, uniform covering of setiferous asperities; in some specimens, asperities tend-
ing to be arranged in longitudinal series, and thus appearing somewhat costate; cuticular
surface between asperities smooth, glossy.

Abdomen (Fig. 2) broad at base, but narrower than elytra. Terga III-V (first three visible
tergites) broadly, transversely impressed at base; impressions coarsely foveate, each fovea
limited laterally by distinct, flattened ridge; basins of foveae smooth and glossy, without
microsculpture. Tergite VI slightly impressed at base with several obsolete ridges and
foveae. Tergal surfaces posterior to basal impressions moderately densely punctured and
pubescent, some punctures at most minutely asperate; cuticle smooth and glossy. Sterna
III-V strongly constricted at base; basal constriction coarsely foveate (often visible in
lateral view).

Male. Eighth tergite with apical margin broadly arcuate at middle, with comb of
minute denticles. Median lobe of aedeagus as in Figs. 3-4. Paramere and apical lobe of
paramerite as in Fig. 5.

Female. Eighth tergite with apical margin as in male. Spermatheca as in Fig. 6.

Secondary sexual characteristics. None apparent.

Material examined. Holotype: male, COSTA RICA: Puntarenas; OTS Sta. finca Las
Cruces, 4000 ft.; San Vito; 111:18:1973. 8258'W-846'N, leg. J. Wagner. J. Kethley/
FM(HD)#73-322, 73CRIII-18d FLC Berlese ISOOcc. leaf litter in stream bed. away from

Vol. 104, No. 2, March & April, 1993

63

flowing water, steep banks, virgin forest cover. Terminalia. aedeagus and parameres mounted
(in Euparal) on microslide and affixed below specimen. The holotype is deposited in the
Field Museum of Natural History, Chicago (FMNH).

Paratypes. 9: Same data as holotype. 1; same data, except 111:16:1973, 4; same data,
except 11:19:1973, 2; COSTA RICA: Guanacaste; Canas. Miravalles Volcano, 1042'N-
857'W;IV:8:1973. leg. J. Wagner, J. Kethley/FM(HD) #73-385, 73CRIV-8e: Berlese 2 liters
cone, leaf litter + soil in dry rivulet #1. 1; same data, except FM(HD) #73-386, 73CRIV-8f.
1. Eight paratypes deposited in the FMNH; 1 paratype (female), with same data as
holotype. except with the date 111:16:1973. is deposited in the Cornell University insect
Collection (CUIC).

Etymology. This elegant species is named for my wife, Maureen, who, over the years,
has graciously provided encouragement and constant support of my work on the
Staphylinidae.

Geographic distribution. Known only from the type localities in Puntarenas and
Guanacaste provinces, Costa Rica.

Bionomics. Little is known about the habitat of this species, but specimens at hand
have been collected from Berlese samples of leaf litter in stream beds, in dry rivulets, and
on slopes above stream banks in virgin forests of Costa Rica.

Remarks. Only slight external morphological variation exists be-
tween the populations from Puntarenas and Guanacaste provinces. The

1

Figs. 1-2. Aleodorus maureenae n. sp. (OTS Sta. finca Las Cruces, Puntarenas province,
Costa Rica), scanning electron photomicrographs. 1, head, thorax, scutellum, and upper
one-third of elytra, dorsal aspect. 2, lower two-thirds of elytra, and abdominal segments III-
VI, dorsal aspect.

64

ENTOMOLOGICAL NEWS

Figs. 3-6. Aleodorus maureenae n. sp. 3, Median lobe of aedeagus, lateral aspect. 4, Median
lobe of aedeagus, dorsal aspect. 5, Paramere. 6, Spermatheca. Scale line, 0.1 mm.

ten specimens from Puntarenas province are slightly larger than those
from Guanacaste province (2 specimens) (cf. 3.1-3.4 mm vs. 2.8 mm, re-
spectively). Furthermore, the setiferous asperities on the heads of the
Guanacaste specimens are slightly less dense (asperities separated by
slightly more than their diameters) than on the heads of the Puntarenas
specimens (asperities separated by less than or equal to their diameters).
The density of the asperities on the thorax and elytra of the Puntarenas
and Guanacaste specimens is similar. For all other external characters,
specimens from the two Costa Rican localities are identical.

Generic Reassignment and Redescription of
Falagria costaricensis Bernhauer

Bernhauer ( 1 940) described Falagria costaricensis from Costa Rica.
All subsequent authors and cataloguers have followed this original
generic placement. The diagnostic morphological features of Falagria
species [type species Falagria caesa Erichson, 1837 = sulcata (Paykull,
1789) nee (Miiller, O. F., 1776)] include a bicarinate scutellum, comb of
minute denticles on the apical margin of tergum VIII, margined hypo-
mera, deep pronotal sulcus, and uniform elytral punctation.

I have carefully examined specimens of the synrype series of F. cos-
taricensis and found them to belong to the genus Aleodorns Say [type
speciesAleochara bilobata Say]. Members of this genus are characterized

Vol. 104, No. 2, March & April, 1993 65

by the unique mesosternum which is on a different plane from that of the
metasternum (mesosternum appears elevated), the short, abbreviated
mesosternal process which does not extend between the coxae, and the
long, generally coiled flagellum of the male aedeagus.

Aleodorus costaricensis (Bernhauer), new combination

(Figs. 7-9)

Falagria costaricensis Bernhauer, 1940:159. Lectotype here designated: La Caja: 8 kil-
[ometers]. w(est]. San Jose, C|osta].R[ica)., Schmidt 1934/H and written "costaricensis
Brnh. Typ" (white label)/Handwrirten "costaricensis Brh. Typus Falagria" (red label)/
Chicago NHMus, M. Bernhauer Collection/ LECTOTYPE Falagria costaricensis
Bernhauer, desig. E. R. Hoebeke 1992 (red label) (FMNH).

Paralectotypes, 5, here designated: Same data as lectotype. with the additional
labels: Syntypus (red label)/Bernhauer det./Coll. DEI Eberswalde/PARALECTO-
TYPE Falagria costaricensis Bernhauer, desig. E. R. Hoebeke 1992 (red label) (IPFE).

Redescription. In agreement with generic characters given by
Hoebeke (1985).

Length 2.1-2.6 mm. (n = 5, x = 2.4 mm). Body color rufo-brunneous. with antennae
(especially toward apices), mouthparts, and legs generally rufo-testaceous; in some speci-
mens, basal three abdominal segments light rufo-brunneous.

Head somewhat quadrate, nearly as long as wide, posterior angles somewhat obtuse,
posterior margin truncate to slightly arcuate; neck very slender, about 0.3 x head width
across eyes. Eyes moderate in size, longest diameter slightly greater than temple length.
Dorsal surface smooth and glossy, moderately sparsely, but uniformly punctured and
pubescent, except for broad median area; punctures very fine, non-asperate. Antennae
moderately long, reaching posteriorly to near 0.5 elytral length; articles I-III elongate; arti-
cle III slightly longer than II; article IV somewhat quadrate, slightly longer than wide;
articles V-X becoming gradually broader, more transverse; article XI obconical, pointed
apically, slightly shorter than IX + X.

Pronofum subequal to width of head, broadest across anterior third, gradually narrow-
ing and converging behind toward base; posterior angles acute; posterior margin truncate.
Disc narrowly and deeply sulcate along median line, terminating in deep, subbasal fovea;
surface on either side of sulcus sparsely punctured and pubescent; punctures very fine,
non-asperate; surface between punctures smooth, glossy. Scutellum large, flattened, with
fine, asperate punctures on either side of a broad, smooth, median channel.

Elytra approximately 1.2 x longer than pronotum, humeri well developed, lateral
margins broadly arcuate in posterior half, posterior angles sinuate, posterior margin trun-
cate; surface with moderately dense and uniform covering of very fine punctures and micro-
setae; area adjacent to scutellum with slightly more dense punctures; cuticular surface
between punctures smooth and glossy.

Abdomen slightly narrower than elytra, parallel-sided, tapering to apex; terga III-V
transversely impressed at base; impressions of terga III + IV with large, rather coarse,
foveae, each bordered laterally by fine, slightly elevated, ridges (less so on tergum V); basins
of large foveae obscurely granulate (with imbricate microsculpture); tergal surface pos-
terior to basal impressions of terga III-V moderately sparsely punctured, pubescent; cuticle
smooth and glossy.

66

ENTOMOLOGICAL NEWS

Male. Eighth tergite with apical margin broadly arcuate at middle, with comb of
minute denticles. Median lobe of aedeagus as in Fig. 7. Paramere and apical lobe of
paramedic as in Fig. 8.

Female. Eighth tergite with apical margin as in male. Spermatheca as in Fig. 9.

Secondary sexual characteristics. None apparent.

Figs. 7-9. Aleodorus costaricensis (Bernhauer). 7, Median lobe of aedeagus, lateral aspect. 8,
Paramere. 9, Spermatheca. Scale line, 0.1 mm.

Remarks. The syntype series of F. costaricensis bears a striking resem-
blance to specimens identified &s Aleodorus dubius (Sharp) from Mexico,
Guatemala, and Costa Rica. There is close agreement in body length,
coloration, pronotal configuration, punctation, and, more importantly,
the genitalic characters of both sexes (shape of median lobe and coiled
flagellum of aedeagus, apical lobe of paramerite, and Spermatheca);
these latter genitalic characters are virtually identical for the two species.
Based on this evidence, I strongly suspect that these species are con-
specific. However, I have not, as yet, examined the type series of A. dubius,
and have studied only a limited number of identified specimens.
Therefore, this hypothesis must remain tentative until a comprehensive
revision of the Neotropical Aleodorus is completed.

ACKNOWLEDGMENTS

I am thankful to the following institutions and individuals for providing all specimens
studied (codens identify the collections in the text): (FMNH) Field Museum of Natural
History, Chicago, Illinois, A. F. Newton, Jr. and M. K. Thayer; and (IPFE) Institut fur
Pflanzenschutzforschung, Kleinmachnow der Akademie der Landwirtschaftswissen-
schaftern, Eberswalde-Finow, Federal Republic of Germany, L. Zerche. J. Howard Frank
(University of Florida, Gainesville) and James K. Liebherr (Cornell University) each pro-
vided helpful suggestions and critical review of the manuscript.

Vol. 104, No. 2, March & April, 1993 67

LITERATURE CITED

Bernhauer, M. 1940. Neue Staphyliniden aus Costa Rica. Arb. Morphol. Taxon. Entomol.

Berlin Dahlem 7:158-161.
Blackwelder, R. E. 1944. Checklist of the coleopterous insects of Mexico, Central

America, the West Indies, and South America. Part I. Bull. U.S. Natl. Mus. 185:1-188.
Erichson, W. F. 1837. Die Kafer der Mark Brandenburg, vol. 1, pt. 1, pp. 1-384. Berlin.
Fenyes, A. 1912. Falagria Mannh. and its relatives. J. New York Entomol. Soc. 20:20-27.
Hoebeke, E. R. 1985. A revision of the rove beetle tribe Falagriini of America north of

Mexico (Coleoptera: Staphylinidae: Aleocharinae). J. New York Entomol. Soc. 93:913-

1018.
Miiller, O. F. 1776. Zoologiae Danicae prodromus, seu animalium daniae et norvegiae

indigenarum characteres, nomina, et synonyma imprimis popularum. Copenhagen.

274 pp.
Pace, R. 1989. Aleocharinae neotropiche del Museo Ungherese di Storia Naturale

(Coleoptera, Staphylinidae). Ann. Hist.-Nat. Mus. Natl. Hung. 81:53-107.
Pace, R. 1990. Nuovi Falagriini. Hoplandriini ed Aleocharini della Regione Neotropica

(Coleoptera. Staphylinidae) (LXXX contribute alia conoscenza della Aleocharinae).

Gion Ital. Entomol. 5:157-180.

Paykull, G. 1789. Monographia staphylinorum Sueciae. Upsala, 81 pp.
Say, T. 1830-1834. Descriptions of new species of North American insects and obser-
vations on some of the species already described. New Harmony, Indiana. 81 pp.

(1833:58-73]
Sharp, D. 1883. Biologia Centrali-Americana: Insecta, Coleoptera. 1 (pt. 2): 145-3 12.

London.

BOOKS RECEIVED AND BRIEFLY NOTED

SYSTEMATICS AND ECOLOGY OF THE SUBGENUS IXODIOPSIS (ACARI: IXO-
DIDAE: IXODES). R.G. Robbins & J.E. Keirans. 1992. Thomas Say Fd., Entomol. Soc.
Amer. 159 pp. $25.00 ESA member. S40.00 other.

The publisher states this is the first cladistic analysis within the Ixodidae and the first
quantitative investigation of/xocles. For the seven species ofLcodiopsis, all known host and
distributional data are summarized, and dichotomous identification keys, accompanied
by scanning electron photomicrographs, are provided.

CLASSIFICATION. CLADISTICS. AND NATURAL HISTORY OF NATIVE NORTH
AMERICAN HARPALUS LATREILLE (INSECTA: COLEOPTERA: CARABIDAE:
HARPALINI). EXCLUDING SUBGENERA GLANODES AND PSEUDOPHONUS.
G.R. Noonan. 1991. Thomas Say Fd., Entomol. Soc. Amer. 310 pp. $30.00 ESA member.
$50.00 other.

This work revises the native North American species ofHarpalus and includes keys to
separate all North American members of the genus from those of othergenera of No. Amer.
Harpalini.

68 ENTOMOLOGICAL NEWS

EGG SURFACE ULTRASTRUCTURE IN

MANTISPA INTERRUPTA
(NEUROPTERA: MANTIS PI DAE) 1

Bruce Cutler^

ABSTRACT: The eggchorion ofMantispa interrupta was examined by scanning and trans-
mission electron microscopy. The egg surface consists of reticulations connected by
bridges that rise from the inner chorion surface. The egg stalk surface is featureless even at
high magnifications. This same morphology is seen in the Chrysopidae, as reported by
Hinton(1981).

The eggs of mantispids are white ovals on short stalks and are found
on the undersides of exposed surfaces, such as leaves and anthropogenic
structures (Kuroko 1961; Redborg and MacLeod 1983, 1984, 1985; Rice
1986). They greatly resemble those of Chrysopidae, except that the stalks
are rarely more than 2-3 times the length of the egg and usually less. Illus-
trations of gross morphology are found in Hungerford (1936), Kuroko
(1961), and Merti ( 1940).

METHODS

A female Mantispa interrupta Say was found on West Campus,
University of Kansas, Lawrence, Douglas County, Kansas, on October
13, 1990, laying eggs on a slightly curled red leaf of a 3 m tall planted sugar
maple (Acer saccharum Marshall). Accessible leaves were examined for
90 minutes, but no additional Mantispa were found. The Mantispa female
was enclosed in a glass container with the sides lined with chroma-
tography paper. Eggs were laid in the laboratory on October 1 5, 20, 27, 30,
November 3, 9, 1 3. The female died November 14. Larval emergence was
from 13-17 days after laying. Approximate egg counts were 200-300 per
batch, with a total of about 1500.

Eggs to be fixed were cut out of the main batch of eggs along with the
chromatography paper to which the stalks were fastened. Fixation was
in 2.5% glutaraldehyde in 0.1 M, 7.3 pH sodium cacodylate buffer at
room temperature for 1 hour, followed by fresh fixative at 4C for 2-3
days. For scanning electron microscopy, eggs were then rinsed in buffer,
run through a dehydration series of ethanol to 100% ethanol, then placed
in two 10 minute changes of HMDS (hexamethyldisilazane) and air

1 Received August 24, 1992. Accepted October 19, 1992

2 Electron Microscopy Laboratory and Department of Entomology, University of Kansas,
Lawrence, Kansas 66045-2106

ENT. NEWS 104(2): 68-72, March & April, 1993

Vol. 104, No. 2, March & April, 1993 69

dried from the HMDS. Many eggs, regardless of developmental stage,
were subsequently seen to be collapsed. A few remained intact, and these
were the ones studied. In retrospect, critical point drying would probably
have produced greater numbers of non-collapsed eggs. The paper con-
taining the eggs was glued to a stub and sputter-coated with 200-250 A of
gold-palladium alloy. Specimens were examined with a Philips 501
scanning electron microscope.

For transmission electron microscopy, eggs were rinsed in buffer,
postfixed in 1% osmium tetroxide in the same buffer at 4C for 2.5 hours.
After rinsing in buffer, eggs were dehydrated in an ethanol series to 100%
ethanol, followed by 100% acetone and ultimately embedded in the
epoxy resin, EM-BED 812 (Electron Microscopy Sciences). Silver to
silver-gold sections were cut with a diamond knife, stained with uranyl
acetate and lead citrate, and examined with a JEOL 1 200 EX II transmis-
sion electron microscope.

RESULTS AND DISCUSSION

The surface of the eggs of Mantispa interrupta was virtually identical
to that of Chrysopa species illustrated by Hinton (1981, figs. 50A-E).
Figure 1 shows a whole egg in side view. The micropyle is also very
similar to the micropyle of Chrysopa, and the reticulations of the mic-
ropyle are identical to those of the general egg surface. The only pub-
lished electron micrograph of a Mantispa egg is Hinton's fig. 50F,
showing a view of the side surface ofM interrupta. The reticular surface is
clearly identical to that of figure 3 here. Figure 2 shows the base of the
stalk and the posterior pole of the egg. As in the Chrysopa egg in Hinton's
fig. 50E, the reticulations disappear at the base of the stalk, which is
featureless at magnification up to 20,OOOX. Figure 4 shows the details of
chorionic structure in cross section. The outer reticulations and inner
chorion give no indication of an aeropyle. The reticulations are essen-
tially solid and arise from a solid layer sitting on the inner chorion. The
outer portion of the reticulations is more electron lucent than the base,
and this less dense appearing material also forms the bridges. The inner
chorion consists of two layers of chorionin bridged by pillars. Nowhere
in examined sections are there connections between the spaces between
pillars and the outside. Similar inner (but not outer) chorionic structure
is found in the stick insect Carausius (Hinton, 1981, fig. 180A). In Car-
ausius the pillars arise from the inner layer and are capped at the outer
end. In Mantispa the pillars appear to arise from both layers and join in
the middle. Hinton ( 198 1 ) refers to transmission electron micrographs of
Chrysopa, but does not illustrate them. His statement that "there is no

70

ENTOMOLOGICAL NEWS

space for a film of air" implies that the outer chorionic layer sits on the
inner as inMantispa. To reiterate, the resemblance to the eggs ofChrysopa
is great. Certainly, on the basis of the ultrastructure of the egg surface of
those species of Chrysopidae and Mantispidae examined, one could not
distinguish one family from the other.

Figures 1-2. Mantispa interrupta egg. 1. Side view of egg, S = stalk, M = micropyle, scale
line = 100 |jm. 2. Base of egg stalk(s), scale line = 10pm.

Vol. 104, No. 2, March & April, 1993

71

Figures 3-4. Mantispa intemtpta egg: chorion structure. 3. Surface view, R = reticulation, B
= bridges, scale line - 1 u. 4. Section through chorion, see text for detailed explanation.
R = reticulation, B = bridge, I = inner chorion, P = pillars, scale = 0.2 urn.

ACKNOWLEDGMENTS

I would like to thank Kevin Hoffman, Clemson University, South Carolina, for iden-
tifying the Mantispa; and Hank Guarisco, Kansas Biological Survey, for sharing informa-
tion and reviewing the manuscript. Byron Alexander, University of Kansas, also reviewed
the manuscript. The adult female specimen and some eggs and larvae are deposited in the
Snow Entomological Museum, University of Kansas, voucher number BC 101390.

72 ENTOMOLOGICAL NEWS

LITERATURE CITED

Hinton, H. E. 1981. Biology of Insect Eggs, v. I, II, III. Pergamon Press, Oxford, 1125

pp.
Hungerford, H. B. 1936. The Mantispidae of the Douglas Lake, Michigan Region, with

some biological observations (Neurop.). Entomol. News 47:69-72; 85-87, plate 1.
Kuroko, H. 1961. On the eggs and first instar larvae of two species of Mantispidae. Esakia

3: 25-32, plates 10-14.
Merti, C. 1940. Contribucion al estudio de Mantispa decorata Erd. (Hemip. Cor.). Revista

de la Sociedad Entomologica Argentina 10: 304-307.

Redborg, K. E. and E. G. MacLeod 1983. Climaciella brunnea (Neuroptera: Mantis-
pidae): a mantispid that obligately boards spiders. Jour. Natural History 17: 63-73.
. 1984. Maintenance feeding of first instar mantispid larvae (Neuroptera,

Mantispidae) on spider (Arachnida, Araneae) hemolymph. Jour. Arachnology 13:

139-140.

_. 1985. The developmental ecology of Mantispa uhleri Banks (Neuroptera:

Mantispidae). 111. Biol. Monograph 53, 130 pp.
Rice, M. E. 1986. Communal oviposition by Mantispa fuscicon. is (Say) (Neuroptera: Man-
tispidae) and subsequent larval parasitism on spiders (Arachnidae: Araneida) in south
Texas. Jour. Kans. Entomol. Soc. 59: 121-126.

BOOKS RECEIVED AND BRIEFLY NOTED

THE HOT-BLOODED INSECTS. B. Heinrich. 1993. Harvard Univ. Press. 597 pp.
$75.00

This author of Bumblebee Economics and Ravens in Winter presents here what is now
known about thermoregulation in all of the major insect groups, offering new insights into
physiology, ecology, and evolution. By describing the environmental opportunites and
challenges faced by a wide range of insect life, Heinrich attempts to explain their great
variety of physiological and behavioral adaptations for survival in their world.

INSECT PATHOLOGY. Y. Tanada & H.K. Kaya. 1993. Academic Press. 666 pp.

Originally intended as an update of Steinhaus' 1949 text, "Principles of Insect Pathol-
ogy", this book developed into a new text on insect pathology, on the different types of dis-
eases in insects, and on their biological control. Each of the 16 chapters ends with a very
comprehensive listing of additional references.

THE BIOLOGY OF MOSQUITOES. Volume 1. DEVELOPMENT, NUTRITION, AND
REPRODUCTION. A.N. Clements. 1992. Chapman & Hall. 509 pp. $99.50

This is the first of two volumes arising from the rewriting of "The Physiology of Mos-
quitoes", published in 1963, so it is written from the viewpoint of a physiologist. This first
volume covers subjects such as genetics, embryology, larval biology, growth & develop-
ment, metamorphosis, adult physiology, and nutrition of adults and larvae.

Vol. 104, No. 2, March & April, 1993 73

A RECLARIFICATION OF THE MALES OF

ALLOPERLA CONCOLOR AND A. NEGLECTA

(PLECOPTERA : CHLOROPERLIDAE), WITH NEW

DISTRIBUTION RECORDS FOR BOTH SPECIES 1

Boris C. Kondratieff, 2 Ralph F. Kirchner 3

ABSTRACT: Historically, there has been confusion concerning the identification of
Alloperla concolor Ricker and Alloperla neglecta Prison. Studies of types and specimens
determined by S.W. Hitchcock (United States National Museum of Natural History), T.H.
Prison (Illinois Natural History Survey), and material in the author's collections revealed
new characters allowing for accurate separation. Figures of the male epiproct of both
species showing specific details are presented. New distribution records are also noted.

Adults of the genus Alloperla are generally recognized by their deli-
cate habitus and lime green or yellow coloration in vivo. Presently, this
genus includes 28 Nearctic species. Many of these species are regionally
endemic and often are only locally abundant.

Ricker (1935) described Alloperla concolor from Homing's Mills,
Ontario, Canada based on a male and two females. Prison, also in 1935
described Alloperla neglecta from North Carolina near Newfound Gap
based on three males. In 1942, Prison synonymized A. concolor, after con-
cluding that this species was "specifically identical with neglecta." Ricker
apparently concurred with this opinion. However, Hitchcock (1968) pre-
sented morphological evidence that Alloperla concolor was a valid
species, based on the shape of the male epiproct. He again in 1974 men-
tioned that both species were specifically distinct, based on the charac-
ters of the epiproct, Surdick (1985) listed A. concolor as being north-
eastern in distribution and A. neglecta restricted to the southern Appa-
lachian Mountains of North Carolina and Tennessee.

After we collected specimens of A. concolor from West Virginia in
1990, a review of the taxonomic status of both species was undertaken.
Types of both species were examined, but unfortunately the male
terminalia of the holotype of A. concolor had been lost (G. W. Wiggins,
Royal Ontario Museum, personal communication). Additionally,

1 Received September 15, 1992. Accepted September 16. 1992

2 Department of Entomology, Colorado State University, Fort Collins, Colorado 80523

3 U.S. Army Corps of Engineers, Water Quality Section (ED-HW), 502 8th Street. Hunt-
ington. West Virginia 25701

4 The views of the second author do not purport to reflect the position of the Depart-
ment of the Army or the Department of Defense.

ENT. NEWS 104(2): 73-78. March & April. 1993

74

ENTOMOLOGICAL NEWS

material determined by Hitchcock (United States National Museum of
Natural History) and Prison (Illinois Natural History Survey) were
also examined.

RESULTS AND CONCLUSIONS

Figs. 1-6 illustrate the apex of the epiproct of both species. Contrary to
Hitchcock's (1968) observations that the epiproct of A. neglecta lacked
distal serration, both species have these serrations (Fig. 1 ). In lateral view
the epiproct of A. concolor resembles the head of a duck, flattened and
forming a distinct serrated edge at the top (Figs. 2A, 3), whereas A.
neglecta has subparallel margins with the tip rounded (Figs. 2B, 5). In
dorsal aspect the epiproct of A. concolor is nearly as wide as long, but in A.
neglecta the epiproct is two times as long as wide (Figs. IB, 5, 6). Speci-
mens collected from southwestern Virginia, which were thought to be
Alloperla concolor by Kondratieff and Kirchner (1987), are A. neglecta.
This represents a northern range extension. The records of A. concolor
from West Virginia represent a new state record and a southern range
extension for this species. Map 1 indicates the recorded distribution of
both species, including the records listed below.

1 A

1 B

Fig. I. Apex of epiproct, dorsal view. A. A. concolor: B.. A. neglecta.

Vol. 104, No. 2, March & April, 1993

75

B

Fig. 2. Epiproct, lateral view. A., A. concolor; B.,A. neglecta.

Material Examined: Alloperla concolor: Ontario, Canada: Homing's Mills, 1 m pinned
(holotype), 2 f pinned. Connecticut: Hartland, 14-VM966, SW Hitchcock, 1 m, 1 f.
Barkhamstead, 1 -VI- 1967, SW Hitchcock, 4 m 3 f. New Hampshire: North Woodstock,
13-V1-1964, SW Hitchcock, 3 m, 8 f. West Virginia: Nicholas Co., near Richwood. North
Fork Cherry River, 14- V- 1990, B. C. KondratiefT, J. L. Welch & R. F. Kirchner, 2 m. 1 f; same
but 2-V1- 1992, 1 m.

Alloperla neglecta: North Carolina: Swain Co., (near) Newfound Gap. 3560 ft, 28-V-
1934, TH Prison, 1 m (pararype); Haywood Co., Shining Rock Natl. Rec. Area, East Fork
Little Pigeon River, 17-V-1983, BC Kondratieff & RF Kirchner, 18 m; same but 23- V- 1990,
BC Kondratieff, RF Kirchner & JL Welch, 7 m, 6 f. Tennessee: Sevier Co., (West Prong)
Little Pigeon River, (near) Newfound Gap, 14-V-1939, TH Prison & HH Ross, 4 m; Sevier
Co., Gatlinburg, 14- VI- 1940, TH Prison etal. 1 m 5 f. Virginia: Grayson Co., Lewis Fork.
2-VII-1978, RF Kirchner & GT Voreh, 4 m; same but 18-V-1990, BC Kondratieff. RF
Kirchner & JL Welch, 58 m, 35 f.

76

ENTOMOLOGICAL NEWS

*.&

Figs. 3-6. Epiproct,/! concolor; 3. Lateral (680X), 4. apex, dorsal view (81 OX). A. neglecta; 5.
lateral (600X), 6. apex, dorsal view (925X).

Vol. 104, No. 2, March & April, 1993

77

Map 1. Distribution of /I. concolor (), A. neglecta ().

ACKNOWLEDGMENTS

We thank Oliver S. Flint, Jr. (United States Museum of Natural History), Glenn B.
Wiggins (Royal Ontario Museum), and Kathryn C. McGiffen (Illinois Natural History
Survey) for providing critical material for study. Additionally, we appreciate the comments
on this project by the participants of the XI International Symposium on Plecoptera
(August 17-20), Tomahawk, Wisconsin.

78 ENTOMOLOGICAL NEWS

LITERATURE CITED

Prison, T. H. 1935. New North American species of the genus Alloperla (Plecoptera:

Chloroperlidae). Trans. Amer. Entomol. Soc. 61: 331-344.
Prison, T. H. 1942. Studies of North American Plecoptera with special reference to the

fauna of Illinois. Bull. Illinois Nat. Hist. Sur. 22: 235-355.
Hitchcock, S. W. \968.Alloperla (Chloroperlidae: Plecoptera) of the Northeast with a key

to species. J. New York Entomol. Soc. 76: 39-46.
Hitchcock, S. W. 1974. Guide to the Insects of Connecticut. Part VII. The Plecoptera or

stoneflies of Connecticut. State Geological and Natural History Survey of Connecticut,

Department of Environmental Protection. Bull. 107. 262 pp.
Kondratieff, B. C. and R. P. Kirchner. 1987. Additions, taxonomic corrections, and

faunal affinities of the stoneflies (Plecoptera) of Virginia, USA. Proc. Entomol. Soc.

Wash. 89: 24-30.

Ricker, W. E. 1935. New Canadian perlids (Part II). Can. Entomol. 67: 256-264.
Surdick, R. P. 1985. Nearctic genera of Chloroperlinae (Plecoptera: Choroperlidae).

Illinois Biol. Monogr. 54: 146 pp.

BOOKS RECEIVED AND BRIEFLY NOTED

CATERPILLARS. ECOLOGICAL AND EVOLUTIONARY CONSTRAINTS ON
FORAGING. N.E. Stamp and T.M. Casey, eds. 1993. Chapman & Hall. 587 pp. $75.00

This volume provides a framework for a unified theory of foraging, taking into account
how two or more constraints influence caterpillars. Ranging from behavioral and nutri-
tional ecology to insect physiology and biophysics, it will serve as a valuable resource for
courses in insect-plant interactions, insect ecology, population and community ecology,
and biological control.

INSECT LEARNING. ECOLOGICAL AND EVOLUTIONARY PERSPECTIVES.
D.R. Papaj and A.C. Lewis, eds. 1993. Chapman & Hall. 398 pp. $54.95

This volume challenges the widespread view that insect behavior is rigidly pro-
grammed and inflexible. There is increasing recognition that there is plasticity in the
behavior of insects that can be explained only by learning. This book surveys a number of
studies on the ecology, evolution, and mechanisms of learning in a number of insect
species, both social and non-social.

Vol. 104, No. 2, March & April, 1993 79

NEW RECORDS OF SPIDERS (ARANEAE) FROM
CAPE COD, MASSACHUSETTS, INCLUDING
TWO POSSIBLE EUROPEAN IMMIGRANTS 1

Robert L. Edwards 2

ABSTRACT: Trochosa ruricola (Lycosidae) and Lepthyphantes tenuis (Linyphiidae), the for-
mer previously known only from Europe and Asia, the latter only from Europe and the west
coast of North America, occur and appear to be well established on Cape Cod, Massa-
chusetts, USA. Six southerly distributed species, Gladicosa pulchra, Lycosa acompa,
Drassylus dixinus. Thymoites expulsa, Grammonota vitatta. and Dictyne pixi also are com-
monly found here.

This paper is presented as a reference to use in connection with Kas-
ton's Spiders of Connecticut, revised in 1981, which is still a widely used
volume and the only one of its kind. Kaston recorded 478 species of
spiders in Connecticut. So far I have recorded 465 species from Cape
Cod (Edwards, unpubl.). There are some uncertainties in the list, some
are yet to be identified, and a few are almost certainly undescribed. After
five years of intensive sampling, additions to the list continue to show up
with regularity. The study area is on the southwestern tip of Cape Cod,
township of Falmouth, and extends for 15 km north from the village of
Woods Hole to Hatchville. The area is dominated by suburban develop-
ments, salt and brackish marshes and a few extensive tracts of second
growth pine and deciduous woodland.

The widely distributed palearctic species, Trochosa ruricola (De Geer)
is common here, with mature specimens found throughout the year. In
the British Isles this species is regularly taken in the same habitats with
Trochosa terricola Thorell (Roberts 1985). Trochosa ruricola and T. terricola
are the most common of the four Trochosa species found in the British
Isles. These two species are common on Cape Cod, occupying somewhat
separate habitats. Trochosa ruricola and T. terricola are most abundant in
open areas such as fields, lawns and gardens, marsh environments, and
edges of woods. Trochosa ruricola is more abundant in wetter areas and
closer to the shore.

Both sexes and all instars of Trochosa ruricola have a claw on their
palp, including the cymbium; T. terricola does not. In addition, males of
T. ruricola have a unique ridge on the fang, nearer the proximal end of the
anterior margin. The male palp of T. terricola has a loop near the ter-

1 Received August 27, 1992. Accepted September 22, 1992.

2 Research Associate, U.S.N.M. Home address: Box 505, Woods Hole, MA 02543.

ENT. NEWS 104(2): 79-82, March & April, 1993

80 ENTOMOLOGICAL NEWS

minus of the embolus (Brady 1979, fig. 30); whereas that of ruricola is
barely curved. The epigyna of T. ruricola and T. terricola are very much
alike, but the presence or absence of a claw on the female palp suffices to
separate the two species.

On Cape Cod, T. ruricola outnumbers T. terricola in pitfall traps by a
factor of two or more. Adults of T. ruricola have been taken year round,
with females carrying egg sacs taken May through July. During the day,
females can be found in shallow, often silk-lined burrows in dry, matted
grass, old mouse nests, and under boards and rocks. Four sacs of T.
ruricola contained from 72 (July) to 17 1 (May) eggs. For further details on
the diagnosis and ecology of Trochosa terricola see Brady 1979; for T.
ruricola, see Roberts 1985.

Lepthyphantes tenuis (Blackwall) (Linyphiidae) is commonly taken in
tall grass in wet boggy areas and around pools. It has been found in
association with small ( 10 cm), horizontal sheet-like webs, usually
well above ground level. Mature specimens have been taken July
through October. Immatures could easily be mistaken for those of
Lepthyphantes zebra (Emerton). The genitalia of the adults are well illus-
trated in Roberts 1985. This species also occurs on the west coast of North
America. It has been recorded from the state of Washington by Crawford
1988, and from British Columbia, Canada by West et al 1984.

There is a distinct southern element in the other species newly found
on Cape Cod. The following six species illustrate this. B. J. Kaston's
treatise (op. cit.) included all known New England species, but did not
include these. All six species are well established on Cape Cod.

Gladicosa pulchra (Keyserling) (Lycosidae) is a handsome, strikingly
marked lycosid found on tree trunks, especially the rough barked trunk
of pitch pine (Pinus rigida). It is also found in holes and other protective
shelters on the trunks of more smoothly barked trees, such as the scarlet
oak. This lycosid matures late in the summer and descends to the ground
in the fall where it is taken in pitfall traps during a brief period in October
and early November. Young spiders appear on tree trunks in the spring,
usually before the end of April. Gladicosa pulchra has been taken as far
north as Long Island, but is generally found south of the Mason-Dixon
line (Brady 1986).

Lycosa acompa Chamberlin is common on Cape Cod in open
grassland, deciduous forest litter and old gravel pits where it is taken in
pitfall traps. The few records of L. acompa in the literature suggest that it
is typically a southern species. Young et al. 1989, note that it is a common
species in Washington County, Misssissippi.

Drassyllus dixinus Chamberlin (Gnaphosidae) is taken from June to
August as adults in pitfall traps in old fields. The northernmost pub-

Vol. 104, No. 2, March & April, 1993 81

lished record is for Patrole, Maryland, (Platnick and Shadab, 1982).

Tymoites expulsa Gertsch and Mulaik(Theridiidae) is found on dune
grass in the salt marsh near the ocean. Adults have been found in the
spring and fall. When Levi (1957) revised the genus, the northernmost
record was from North Carolina.

Grammonota vitatta Barrows (Linyphiidae) has been regularly col-
lected as immatures and adults along the edges of brackish and fresh-
water marshes during the colder months of the year. Previously, the
northernmost record was the type locality. Cape May, New Jersey,
( Bishop etal 1932).

Dictynapixi Chamberlin and Gertsch (Dictynidae) adults have been
taken in June by sweeping old fields dominated by such forbs as false
indigo and sweet fern. Immatures and subadults have been taken in pit-
fall traps in October. The type locality is Washington Crossing, New
Jersey; and it has been recorded from North Carolina, Arkansas, and
Michigan (Chamberlin and Gertsch, 1958).

The erigonines constitute about 20% of the total spider fauna in the
study area (Edwards, unpubl.). One species Kaston considered ubi-
quitous and extremely common in Connecticut, Ceraticelusfissiceps (O.
P.-Cambridge), has yet to be taken in the study area. On Cape Cod,
apparently C. fissiceps has been replaced by C. alticeps (Fox). Sixteen
species Kaston did not find in Connecticut but from Massachusetts or
farther north have been taken on Cape Cod. And among the erigonines
he recorded from Long Island and/or further south but not Connecticut
or elsewhere in New England are Grammonota maculata Banks, G.
pallipes Banks and Ceraticelus laticeps (Emerton). All are found on Cape
Cod and are common. These data also suggest that there may be a ten-
dency for more southerly distributed species to extend their range
northward along the coast.

With regard to the possible immigrants from Europe, it is a matter of
record that this region has been studied and collected by individuals
interested in spiders for over 100 years. Earlier investigators might have
missed Lepthyphantes tennis. It is a small spider and could easily have
been misidentified as one of the other two common Lepthyphantes
species. The relatively large size and abundance today of Trochosa
ruricola, as well as the lack of records elsewhere in New England, sug-
gests a more recent arrival. Since the 1 880's, Woods Hole. Cape Cod, has
been an area of considerable activity, principally focused on the marine
environment. The several institutions are often visited by scientists and
research essels from various parts of the world, especially from Europe,
thus providing many opportunities for stowaways, including spiders.

82 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

This study was funded in part by a grant from the natural Heritage & Endangered
Species Program of the State of Massachusetts.

Allen Brady (Hope College) kindly provided me with the initial identifications of
Trochosa ruricola and Lycosa acompa. He was most helpful in discussions of lycosid species,
their diagnosis and ecology and with comments on this manuscript. Daniel Jennings
(Univ. of Maine) thoroughly critiqued the final draft with his usual care and thoughtful-
ness. Charles Dondale and James Redner (BRC, Canada) assisted with helpful comments,
and information on the distribution ofLepthyphantestenuis. I am grateful to Vincent Roth,
Portal, Arizona, who provided my first assist in coping with a number of taxonomic prob-
lems concerning lycosids, and to Jonathan Coddington (USNM) for his comments and
suggestions on an early draft of the paper.

LITERATURE CITED

Bishop, S. C. and C. R. Crosby. 1932. Studies in American spiders: the Genus Gram-

monota. J. New York Ent. Soc., 40:393-421.
Brady, A. R. 1979. Nearctic species of the wolf spider Genus Trochosa (Araneae: Lyco-

sidae). Psyche, 86(2-3): 167-2 12.
1986. Nearctic species of the new wolf spider Genus Gladicosa

(Araneae:Lycosidae). Psyche, 93(3-4):285-319
Chamberlin, R. V. and W. J. Gertsch. 1958. The spider family Dictynidae in America

north of Mexico. Bull. American Mus. Nat. Hist., 116(1):1-152.
Crawford, R. L. 1988. An annotated checklist of the spiders of Washington. Burke Mus.

Cont. in Anthropology and Nat. Hist. No. 5:1-48.
Kaston, B. J. 1981. Spiders of Connecticut (rev. ed.). Connecticut St. Geol. Nat. Hist. Surv.

Bull., 70:1-1020.
Levi, H. W. 1957. The spider Genera Enoplognatha, Theridion, and Paidisca in America

north of Mexico (Araneae: Theridiidae). Bull. American Mus. NaL Hist., 112(1):1-

124.
Platnick, N. L. and M. U. Shadab 1982. A revision of the American spiders of the genus

Drassylus (Araneae, Gnaphosidae). Bull. American Mus. Nat. Hist., 173(l):l-97.
Roberts, M. J. 1985-1987. The Spiders of Great Britain and Ireland. 3 volumes, 1 : 1-229; 2: 1-

204; 3:1-256. Harley Books, Essex, England.
West, R., C. D. Dondale and R. A. Ring. 1984. A revised checklist of the spiders (Araneae)

of British Columbia. J. Ent. Soc. Brit. Columbia, 81:80-98.
Young, O. P., T. C. Lockley and G. B. Edwards. 1989. Spiders of Washington County,

Mississippi. J. Arachnol. 17(l):27-42.

Vol. 104, No. 2, March & April, 1993 83

FIRST KARYOTYPIC DATA ON A CUPEDID

BEETLE (COLEOPTERA: ARCHOSTEMATA)

SHOWING ACHIASMATIC MEIOSIS 1

J. Galian^^ j.p. Lawrence^

ABSTRACT: The chromosomes of a species of Cupedidae, Distocupes varians (Lea) were
studied for the first time. The male diploid chromosome number of the species is 2n = 19
and the male sex chromosome system is of the XO type. The presence of 9 pairs of
autosomes agrees with the hypothesis that suggests that this number is the ancestral condi-
tion for the whole Order Coleoptera. On the other hand the analysis of the spermatogenesis
reveals an achiasmatic meiosis pattern. The occurrence of this kind of meiosis in
Cupedidae (never recorded in Polyphaga, but present in some groups of Adephaga) is in
agreement with hypotheses that relate Archostemata with Adephaga.

The suborder Archostemata is a primitive group of beetles which
includes three families, Cupedidae, Micromalthidae and Ommatidae
(Lawrence etal. 1987). The only species of Micromalthidae so far karyo-
typically studied is Micromalthus debilis LeConte (Scott, 1936, 1941)
which has haplodiploidy, males with n = 10 and females with 2n = 20.
The family Cupedidae has 25 species worldwide and is represented in
Australia by the monotypic genus Distocupes and four species ofAdino-
lepis (Neboiss, 1984). To date nothing has been published on the karyo-
types of these species.

The relationships between Archostemata and the other three subor-
ders of Coleoptera are still in dispute. Crowson (1955, 1960) considers
that there are three ancestral stocks: Archostemata, Adephaga and Myx-
ophaga plus Polyphaga. Lawrence and Newton (1982) and Kukalova-
Peck and Lawrence (in press) suggest that Archostemata, Myxophaga
and Adephaga may form a monophyletic group based on wing venation
and folding.

In the present paper information on the chromosomes of the species
Distocupes varians (Lea) is reported and the relationships of its karyotype
with those of the other suborders is discussed.

MATERIALS AND METHODS

Five individuals of Distocupes varians were collected in December

1 Received June 1 1992. Accepted September 26, 1992

2 CSIRO Division of Entomology, GPO Box 1700, Canberra, ACT 2601, Australia

3 Departamento de Biologia Animal y Ecologia, Facultad de Veterinaria, Universidad de
Murcia, Apdo. 4021, Murcia 30071, Spain

ENT. NEWS 104(2): 83-87, March & April, 1993

84 ENTOMOLOGICAL NEWS

1990 from a permanent colony living in a garden in O'Connor, Can-
berra, Australia. The beetles are deposited in the Australian National
Insect Collection, Canberra. Male specimens wre injected with a 0.04 M
sodium acetate plus 0.05% colchicine solution for ten minutes and then
anesthetized. The testes were dissected out, fixed in 3:1 ethanol: acetic
acid solution, and then squashed in a drop of 1% lacto-propionic
orcein.

RESULTS

The male diploid chromosome number of Distocupes varians is 2n =
19 with nine pairs of autosomes plus X. The karyogram made from
metaphase II cells (Fig. 1 ) shows 9 pairs of metacentric and submetacen-
tric chromosomes gradually decreasing in size. The X chromosome
seems to be a metacentric element about the size of the second pair.

In meiosis during early prophase I the homologues condense (Fig. 2,
3) and at zygotene the central regions of the bivalents, probably of
heterochromatic nature, are heavily stained while the rest is weakly
stained (Fig. 3). During pachytene (Fig. 4) to metaphase I (Fig. 5) the
homologues remain in parallel alignment without showing any trace of
chiasmata. Homologous centromeres appear more deeply separated in
some bivalents (Fig. 5). The onset of anaphase I is denoted by a parallel
separation of homologues, which is delayed in the telomeric regions of
some bivalents (Fig. 6). At prophase II chromosomes appear as single
structures (Fig. 7). The two chromatids are seen again at metaphase II,
but they remain parallel instead of the typical cruciform figure due to
chromatid repulsion (Fig. 8). The X chromosome condenses pre-
cociously (Fig. 4) and moves undivided to one pole at anaphase I, and
divides its chromatids during anaphase II.

During specimens preparation, the testes of this species were found
to be of the normal follicular type, in which a number of small sperm
tubes are attached individually to the vas deferens (Snodgrass, 1935),
rather than the tubular type (Jeannel, 1941), which consists of a single,
coiled tube. This feature is of phylogenetic importance (see below).

DISCUSSION

Male meiosis of Distocupes varians resembles the achiasmatic pattern
that was previously described in Adephaga (Carabidae:Bembidiini,
Pogonini and perhaps Harpalini) by Serrano ( 1 98 1 a). Since achiasmatic
meiosis is considered to have evolved secondarily (John, 1990), this
character represents a specialized condition within the Cupedidae, an

Vol. 104, No. 2, March & April. 1993 85

unexpected result in view of the supposed archaic nature of the group.
However, it may also be considered as a latent tendency of the first
coleopterans that may appear in particular groups or species. Given the
lack of reports of achiasmatic meiosis in the suborder Polyphaga, in
spite of the large number of species studied (2000 in Smith and Virkki,
1978, and many more since then), the occurrence of this kind of meiosis
is in agreement with the hypotheses that relate the suborder Archo-
stemata with the suborder Adephaga (Lawrence and Newton, 1982;
Kukalova-Peck and Lawrence, in press).

I *
'
*

P 3 f U 3 * *r :

4

* / + + i"

\ f <:v>

w

f 8

Chromosomes of Distocupes varians. Figure 1. Karyogram made from two metaphase II
cells with n = 9 + X (above) and n = 9 (below). Figure 2. Early zygotene. Figure 3.
Zygotene. Figure 4. Postpachytene stage. Note the precocious condensation of the X
chromosome. Figure 5. Metaphase I. Figure 6. Anaphase I. Figure 7. Prophase II with n = 9
(left) and n = 9 + X (right). Figure 8 Metaphase II. Arrows show the X chromosome.
The bar equals 5 um.

86

ENTOMOLOGICAL NEWS

On the other hand, the chromosome number of Distocupes varians 2n
= 19, is very close to that found in other primitive coleopteran groups
(Table 1), thus supporting the hypothesis that the 2n = 20 karyotype is
ancestral for the Order Coleoptera, and that higher numbers developed
thereafter in the adephagan and polyphagan stocks, with disappearance
of the ancestral karyotype in modern Adephaga.

If the lack of the typical polyphagan Xyp sex-chromosome mecha-
nism in D. varians is corroborated in other species of Archostemata, it
will indicate that this system is characteristic of the suborder Polyphaga
but is not found in the other suborders of Coleoptera (Table 1 ). Thus the
Xyp system evolved when Polyphaga became separated from the other
suborders and represents an apomorphy for the suborder.

Finally, the occurrence of follicular testes in D. varians, as well as in
two other Cupedidae, Priacma serrata LeConte and Prolixocupes lobiceps

Table 1. Chromosome numbers in Coleoptera

Suborder

Species

References(*)

ARCHOSTEMATA
CUPEDIDAE

MICROMALTHIDAE

MYXOPHAGA

ADEPHAGA

POLYPHAGA

Distocupes varians

n (cf ) = 9 + X

2n (cf) = 19

Male achiasmatic meiosis

Micromalthus debilis

n (cf) = 10

2n (9) = 20

Sex-chromosome mechanism by

haplo-diploidy (arrhenotoky)

Ytu zeus

n(cf) = 9 + XY

2n (cf ) = 20

Ancestral karyotype
n(cf) = 18 + X

2n (cf ) = 37

Ancestral karyotype

n (cf ) = 9 + Xyp (parachute)

2n (cf ) = 20

(*) 1, Present study; 2, Scott (1936); 3, Mesa and Fontanetti (1985); 4, Serrano (1981b); 5,
Smith and Virkki ( 1978).

Vol. 104, No. 2, March & April, 1993 87

(LeConte) (K.W. Cooper, pers. comm.) conflicts with the reports of
tubular testes in Prolixocupes latreillei (Solier) and also in Tetraphalerus
wagneri Waterhouse (family Ommatidae) (Vidal-Sarmiento, 1969). It
appears that tubular and follicular testes, which characterize Adephaga
and Polyphaga, respectively, both occur within the suborder Archo-
stemata. If this is the case, it suggests that modern Archostemata are not
monophyletic or that this character has undergone reversal. However,
more data is needed to confirm the presence of tubular testes in either
Ommatidae or Cupedidae.

ACKNOWLEDGMENTS

We are grateful to K. Houston for providing the material and J. Serrano, M. Horak and T. A
Weir for their valuable comments on the manuscript. J. Galian was supported by a post-
doctoral grant (PF91-22936091) of the Spanish Government for staying in Australia.

LITERATURE CITED

Crowson, R.A. 1955. The natural classification of the families of Coleoptera. London.

Crowson, R.A. 1960. The phylogeny of Coleoptera. Ann. Rev. Entomol. 5: 1 1 1-134.

John, B. 1990. Meiosis. Cambridge Univ. Press. Cambridge.

Jeannel, R. 1941. Faune de France. 39. Coleopteres Carabiques. Premiere Parte. Le-
chevalier. Paris.

Kukalova-Peck, J. and J.F. Lawrence, (in press). Evolution of the hind wing in Coleop-
tera. Can. J. Zool.

Lawrence, J.F. and A.F., Jr. Newton. 1982. Evolution and classification of beetles. Ann.
Rev. Ecol. Syst. 13: 261-290.

Lawrence, J.F., T.A. Weir, and J.E. Pyke. 1987. Archostemata and Myxophaga. In:
Zoological catalogue of Australia. Volume 4. Coleoptera: Archostemata, Myxophaga
and Adephaga. D.W. Walton (ed.). Pp. 6-16. Australian Government Publishing Ser-
vice. Canberra.

Mesa, A., and C.S. Fontanetti. 1985. The chromosomes of a primitive species of beetle:
Ytu zeus (Coleoptera, Myxophaga, Torridincolidae). Proc. Acad. Natl. Sci. Philadelphia
137: 102-105.

Neboiss, A. 1984. Reclassification of Cupes Fabricius (s. lat.), with description of new
genera and species (Cupedidae: Coleoptera). Syst. Entomol. 9: 443-447.

Scott, A.C. 1936. Haploidy and aberrant spermatogenesis in a Coleopteran, Micromalthus
debilis Leconte. J. Morphol. 59: 485-515.

Scott, A.C. 1941. Reversal of sex production in Micromalthus. Biol. Bull. 81: 420-431.

Serrano, J. 198 la. Male achiasmatic meiosis in Caraboidea (Coleoptera, Adephaga).
Genetica57: 121-137.

Serrano, J. 1981b. Chromosome numbers and karyotypic evolution of Caraboidea.
Genetica 55: 51-60.

Smith, S.G. and N. Virkki. 1978. Animal cytogenetics 3, Insecta 5, Coleoptera. Born-
traeger. Berlin.

Snodgrass, R.E. 1935. Principles of Insect Morphology. McGraw-Hill. New York.

Vidal-Sarmiento, J.A. 1969. El sistema reproductor masculino y organos de copulacion
en Cupesidae (Coleoptera-Archostemata). Rev. Soc. Entomol. Argent. 31: 43-48.

88

ENTOMOLOGICAL NEWS

AN ECOTONAL STUDY OF CARRION BEETLES

(COLEOPTERA: SILPHIDAE) IN THE GREAT

SWAMP NATIONAL WILDLIFE REFUGE,

NEW JERSEY 1

Paul P. Shubeck 2

ABSTRACT: Carrion beetles were collected in chicken breast baited traps situated at 15
meter intervals, along a 60 m base line intersecting a woodland and field in Great Swamp
National Wildlife Refuge, NJ. There was no evidence, for any species, of a declining linear
gradient in numbers of carrion beetles from their preferred to their secondary habitat. An
abrupt edge effect was noted for one species, Nicrophonts orbicollis. which was taken in the
woodland and at the woodland/field interface, but not in the field proper. On the other
hand, Necrophila americana, Oiceoptoma noveboracense, and Oiceoptema inaequale were
each approximately equally abundant on either side of the actual interface of their pre-
ferred and secondary habitat. Furthermore, this junction zone (ecotone) may end between
15 m and 30 m into the field for Necrophila americana since it was found to be significantly
more abundant at 30 m into the field (its preferred habitat).

Two previous studies have shown that carrion beetles (Silphidae)
manifest slight to strong preferences for particular habitats. Anderson
(1982) observed species preferences, in Canada, for coniferous forests,
deciduous forests, fields and marshes. Shubeck (1983) observed species
preferences when trapping carrion beetles in woodland, field, and marsh
habitats in NJ. These findings are especially interesting in view of an
earlier study which showed there is much random flight when carrion
beetles search for carrion (Shubeck, 1968). In the New Jersey study it had
been found that eight species of carrion beetles had slight or strong pre-
ferences for woodland or field habitats. Only 5% of the individuals were
taken in a marsh.

Neither of the above habitat studies discussed ecotonal aspects of
boundaries between habitats. According to Odum ( 197 1 ), "An ecotone is
a transition between two or more diverse communities as, for example,
between forest and grassland ..." He further stated, "It is a junction zone
or tension belt which may have considerable linear extent but is
narrower than the adjoining community areas themselves."

In this current study an attempt was made to determine what
influence, if any, the ecotone might have on carrion beetle populations
in terms of numbers. Specific objectives included an attempt to deter-
mine if there might be (1) a declining linear gradient in numbers of car-

1 Received September 11, 1992. Accepted September 16, 1992.

Biology Department, Montclair State College, Upper Montclair, New Jersey 07043.
Present Address: 65 Pleasantview Ave., New Providence, NJ 07974.

ENT. NEWS 104(2): 88-92. March & April. 1993

Vol. 104, No. 2, March & April, 1993 89

rion beetles, along a base line, from the preferred habitat to the sec-
ondary habitat, and (2) an abrupt drop in numbers at the interface be-
tween the preferred habitat and the secondary habitat. In other words,
does the change occur at the interface itself or at 15 or 30 m inside of
either habitat.

The study was conducted in the Great Swamp National Wildlife
Refuge, Basking Ridge, NJ. Collections were made during June, July and
August 1984, and April and May 1985.

MATERIALS AND METHODS

Carrion beetles were trapped in five No. 10 food cans (3.78 1 ), each of
which was concealed in a wooden box having 1.27 cm wire mesh at the
top and a rain cover 5 cm over the opening. These have been described
elsewhere (Shubeck, 1976). A base line of 60 meters length was estab-
lished which intersected the boundary at which a deciduous woodland
and an old field were adjacent to each other. These habitats have been
described in a previous paper (Shubeck, 1983).

The five traps were situated along the base line so that one trap was 30
m into the woodland, a second was 1 5 m into the woods, a third was at the
edge of the woods where the field began, a fourth 1 5 m into the field, and a
fifth was 30 m into the field. One chicken breast was placed in each trap a
week before the first collection, a second added when collections began,
and each week thereafter the older in each trap was replaced with a fresh
chicken breast.

Traps were examined each week, carrion beetles were removed, iden-
tified, and the data recorded. There was a total of 2 1 weekly collections
12 during June, July, and August in 1984, and 9 during April and May in
1985. A previous study (Shubeck et a/., 1981), which ran from April
through November, had shown that over 98% of silphids collected in
Great Swamp were taken from early April through August.

RESULTS AND DISCUSSION

A total of 1,173 carrion beetles (Silphidae) was collected during this
study (Table 1). In rank order of abundance they were: 1. Necrophila
americana (525); 2. Oiceoptoma noveboracense (349); 3. Oiceoptoma inae-
quale (206); 4. Nicrophorus orbicollis (66); 5. Nicrophorus tomentosus ( 1 2); 6.
Nicrophorus pustulatus (6); 7. Necrodes surinamensis (5); 8. Nicrophorus
marginatus (4). In view of the small numbers for the last 4 species this
analysis and discussion will be limited to the four most abundant species
whose larger numbers may be treated statistically.

Table 1 shows total numbers, for 1984 and 1985, of species trapped

90 ENTOMOLOGICAL NEWS

along the base line intersecting the woodland and field interface. The
data, for each species, show how many beetles were trapped 30 m into the
woodland habitat, 15 m into the woodland, at the edge of woodland/
field, 15 m into the field, and 30 m into the field.

Table 1 shows that no species exhibits a linear gradient in numbers
collected from their preferred to their secondary habitat. Even the 4
species found in low numbers seem to bear this out (Table 1).

Only for Nicrophorus orbicollis was an abrupt edge effect noted. This
species was trapped in approximately equal numbers at the edge of the
woodland, and at 1 5, as well as 30 m into the wooded area. No individual
of this species was taken within the field proper. These data seem to indi-
cate that for jV. orbicollis, the interface of its woodland habitat with a field
can be a very real "barrier," or edge, beyond which this species seldom
ventures in search for food. Although in a previous habitat study
(Shubeck, 1983) 10 of a total of 73 individuals (13.7%) were taken in the
field, the great majority (86.3%) was collected in the forest their pre-
ferred habitat.

On the other hand, the 3 most abundant species behaved very dif-
ferently. Necrophila americana, Oiceoptoma noveboracense, and Oiceop-
toma inaequale were captured in somewhat similar numbers along the
base line (with the possible exception of TV. americana since 30% of its
numbers were taken 30 m into the field). It seems clear for these species
there is a definite ecotone, or zone, which extends from within one
habitat, across the edge into its adjacent habitat. Within this zone it
appears these beetles were approximately equally abundant on either
side of the actual interface of the preferred and the secondary habitat. In
order to support this observation the chi square statistic was used to
determine if the numbers of individuals from each of these 3 species,
taken on either side of the interface (i.e. forest vs. field) were significantly
different. The total number of individuals collected at 15 m and at 30 m
into the forest was compared with the total number collected at 1 5 m and
30 m into the field for each of these species respectively. It was found
there was no significant difference in the total number taken on either
side of the actual interface (but within the 60 m wide margin tested) in the
case of 2 species; Oiceoptoma noveboracense (X2 = 1.39, df = 1, P > .2)
Table 1, Oiceoptoma inaequale (X2 = 1.16, df = 1, P > .2) Table 1. Chi
square analysis of Necrophila americana , on the other hand, indicated
there is a significant difference in the numbers on either side of the inter-
face (X2 - 9.43, df = 1, P > .01) Table 1. However, the data in Table 1 may
indicate the junction zone, or tension belt (Odum, 1971), may end bet-
ween 15 and 30 m into the field for this species, since it has been noted
that N. americana manifested somewhat larger numbers 30 m into the

Vol. 104. No. 2, March & April, 1993 91

field. Chi square treatment of the numbers when all five collections for
this species are included indicate a significant difference in the numbers
(X2 = 34.26, df = 4, P > 0.00 1 ). On the other hand, when the latter collec-
tion numbers (30 m into the field) are excluded, and the remaining four
collections are subjected to chi square analysis, no significant difference
in collection numbers is noted (X 2 = 0.945, df = 3, P > .8). The very
obvious difference is the large number of beetles taken 30 m into the
field. I believe this can be accepted as evidence that this species may have
been clear of the ecotone at that point, since the field is its preferred
habitat (Shubeck, 1983), and larger numbers of individuals would be
expected to be there.

This study appears to indicate that there is a zone (ecotone), at least
1 5-30 m wide on either side of the field/woodland interface, within which
there are no significant differences in the numbers collected (on either
side of the interface) of Oiceoptoma noveboracense, Oiceoptoma inaequale,
and Necrophila americana. Nicrophorus orbicollis, however, seemed to
manifest an abrupt edge effect in that its numbers were taken in the
woods and at the interface but not in the field. It appears for this species
that the habitat preference for the woods is so strong that the concept of
"ecotone" may be meaningless.

Table 1. Totals of carrion beetles collected (1984 & 1985) at 15 meter intervals along a base
line which intersected a woodland and an adjacent field in Great Swamp National
Wildlife Refuge, N.J.

WOODS/
SILPHIDAE WOODS WOODS FIELD FIELD FIELD TOTAL

30m 15m Om 15m 30m

Necrophila americana

99

86

91

91

158

(525)

Oiceoptoma noveboracense

63

76

90

47

73

(349)

Oiceoptoma inaequale

58

21

61

24

42

(206)

Nicrophorus orbicollis

24

20

22

(66)

Nicrophorus tomentosus

2

5

1

4

(12)

Nicrophorus pustulatus

1

1

2

2

(6)

Necrodes surinamensis

1

4

(5)

Nicrophorus marginatus

2

2

(4)

92 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

I would like to Thank John Korky, Biology Department, Montclair State College, for read-
ing the manuscript and offering many suggestions, and also Helen Roberts, Mathematics
and Computer Science Department, Montclair State College, who reviewed the statistics
and corrected some of the P values and interpretations. Their assistance is much
appreciated.

LITERATURE CITED

Anderson, R. S. 1982. Resource partitioning in the carrion beetle (Coleoptera: Silphidae)
fauna of southern Ontario: ecological and evolutionary considerations. Can. J. Zool.
60:1314-1325.

Odum, E. P. 1971. Fundamentals of Ecology, 3rd Edition. Saunders, Philadelphia, PA.

Shubeck, P. P. 1968. Orientation of carrion beetles to carrion: random or non-random?
J. New York Entomol. Soc. 76:253-265.

Shubeck, P. P. 1976. An alternative to pitfall traps in carrion beetle studies (Coleoptera).
Entomol. News 87:176-178

Shubeck, P. P., N. M. Downie, R. L. Wenzeland S. B. Peck. 1981. Species composition
and seasonal abundance of carrion beetles (Coleoptera) in an oak-beech forest in
Great Swamp National Wildlife Refuge, N.J. Entomol. News 92:7-16.

Shubeck, P. P. 1983. Habitat preferences of carrion beetles in The Great Swamp National
Wildlife Refuge, New Jersey (Coleoptera: Silphidae, Dermestidae, Nitiduli-
dae, Histeridae, Scarabaeidae). J. New York Entomol. Soc. 91:333-341.

Vol. 104, No. 2, March & April, 1993 93

TECHNIQUES FOR OBTAINING

ADULT-ASSOCIATED IMMATURE STAGES

OF PREDACIOUS TACHYDROMIINE FLIES

(DIPTERA: EMPIDOIDEA), WITH IMPLICATIONS

FOR REARING AND BIOCONTROL 1

J. M. Gumming, B. E. Cooper^

ABSTRACT: Gravid females of four species of tachydromiine flies, namely Mega-
grapha exquisita, Platypalpus holosericus. P. aequalis. and P. melleus were induced to oviposit
by decapitation. Eggs were placed on a saline nutrient agar medium prior to hatching.
Ovaries containing fully mature eggs were additionally dissected from females of P.
holosericus. These eggs were transferred in situ in each ovary to agar plates, where they
embryonated and hatched, demonstrating parthenogenesis in this species. First instar lar-
vae of all four species were held in agar medium for several weeks, and were presented with
various prey organisms and other food materials. Only those that fed on Drosophila
melanogaster larvae, or on each other, developed to later larval instars. In one instance, a
fully mature larva of M. exquisita pupated after diapausing, and developed as far as the
teneral adult stage. Implications of the results of this study are discussed in terms of the
potential for obtaining taxonomic and phylogenetic information on previously unknown
immature stages, and for rearing Tachydromiinae as biological control agents of agri-
cultural pests.

The beneficial nature of empidoid flies as predators of insect pests
has long been recognized (reviewed for example by Smith, 1969, p. 18),
with the potential economic importance of one subfamily, the
Tachydromiinae, recently attracting considerable interest 3 . For exam-
ple, adult tachydromiines have been identified as important regu-
lators of small Diptera, Thysanoptera and aphid pests in cereal and oil
seed crops (Berest, 1987; Brunei et al, 1989; Chvala, 1975; Crook and
Sunderland, 1984; Jones, 1965, 1969, 1976a, 1976b; Potts and Vickerman,
1974; Stark, 1990; Stark and Wetzel, 1987; Sunderland et al. 1985), leaf-
mining flies in greenhouse and field situations (Kovalev, 1966; Rotheray,
1989; Whitfield, 1925), as well as psyllids and phytophagous mites in
orchards (Chvala, 1975; Fleschner and Ricker, 1953).

Despite this interest, effective use of tachydromiines as biological
control agents of agricultural pests has been severely hindered by a lack

1 Received July 3, 1992. Accepted September 26, 1992.

2 Biological Resources Division, Centre for Land and Biological Resources Research.
Agriculture Canada, Research Branch, Central Experimental Farm, Ottawa, Ontario,
Canada K1AOC6.

3 Classification of the Tachydromiinae within either the Empididae or Hybotidae is dis-
cussed in Cumming and Cooper (1992).

ENT. NEWS 104(2): 93-101. March & April. 1993

94 ENTOMOLOGICAL NEWS

of information about immature stages and life histories of empidoid flies
in general. For example, no empidid species has ever been successfully
reared through all life stages in the laboratory. Even the informative
outline of the life cycle ofEmpis tessellata Fabricius, described by Hobby
and Smith (1961), was pieced together from collections of mature larvae
taken from leaf litter (reared to adults) and from eggs obtained from
mated females (hatched to first instar larvae). The entire literature on the
structure and habits of immature stages of the Tachydromiinae, essen-
tially amounts to brief descriptions of the larva of a Platypalpus species
by Beling (1888), the larva and pupa of a Drapetis species by Malloch
(1917), and the larva of Crossopalpus curvipes (Meigen) by Smith (1989).
Smith (1989) additionally lists rearing records of adults of Crossopalpus
sp. from dung, C. nigritellus (zetterstedt) from fungi, Elaphropeza ephip-
piata (Fallen) from woodland soil, and Tachypeza nubila (Meigen) from
fungi and under bark, indicating that the immatures of Tachydromiinae
appear to occupy various terrestrial habitats.

The primary purpose of this paper is to add to our knowledge of the
Tachydromiinae by reporting on techniques used to obtain adult-
associated immature stages and on preliminary investigations into the
establishment of a rearing method for members of the subfamily, in
addition, a procedure is described for determining female reproduc-
tive mode.

PROCEDURE AND RESULTS

Rearing Method. Adult females of megagrapha exquisita (Malloch),
Platypalpus holosericus Melander, P. aequalis Loew, and P. melleus Melan-
der were netted locally during the summer months and placed indi-
vidually in plastic tubes containing moistened tissue paper, for transport
back to the laboratory 4 . Gravid females with fully mature eggs were
induced to oviposit by decapitation, following the method described by
Linley (1965) for the ceratopogonid fly species Leptoconops bequaerti
(Kieffer). This was most easily achieved after slightly anaesthetizing a
specimen with CC>2 and transferring it to moistened filter paper before
removing the head with fine dissecting scissors. Decapitated females
generally started ovipositing immediately and sometimes continued to
lay eggs for up to an hour after oviposition commenced.

Following oviposition, eggs from each female were transferred with a
fine brush to the surface of a saline nutrient agar medium that had been
allowed to set to a depth of approximately 4mmina50X9mm seal-tight

4 Voucher specimens are deposited in the Canadian National Collection of Insects and
Arachnids (CNC).

Vol. 104, No. 2, March & April, 1993 95

petri dish (Fig. 1). The saline nutrient agar medium used to rear
specimens of Tachydromiinae was originally developed for rearing pre-
dacious larvae of the ceratopogonid fly species Culicoides melleus
(Coquillett) and is fully described by Linley (1985). The advantages dis-
cussed by Kettle etal. (1975) in using agar to rear Culcoides larvae, and the
necessity demonstrated by Linley (1985) of adding supplementary
vitamins for rearing predacious larvae such as Culicoides (see below
under "Food Requirements"), appear generally applicable to the rearing
procedure employed here. Seal-tight petri dishes, which are designed
with tight-fitting lids (Fig. 1), are a necessary modification, essential in
containing the small highly motile tachydromiine larvae in the agar

medium.

The agar-filled petri dishes were maintained in the dark, in an
environmental chamber at a constant 20 C. Darkening of the chorion of
the egg on the first day denoted initiation of embryogenesis in all four
species, and indicated that the eggs were either fertilized during oviposi-
tion, or were developing parthenogenetically. In all four species, larval
cephalic structures and segmentation usually became visible through
the semi-transparent chorion towards the end of the first week after
oviposition (Fig. 2), with the eggs generally hatching by the end of the
second week. In some eggs of P. holosericus, hatching was delayed up to at
least four weeks after oviposition without noticeable developmental
effects, by keeping the eggs at 11 C.

First instar larvae of all four species appeared to burrow through the
agar medium easily (Figs. 3-4), and could be kept alive in the tightly
sealed petri dishes with little maintainence for several weeks. Larvae
seemed unaffected by fungus and bacteria, even when some older
cultures became heavily contaminated with these microorganisms, and
often appeared somewhat attracted to these contaminants (Fig. 6). Those
that fed (see below under "Food Requirements") appeared to progress
through three larval instars, as described for the distantly related empi-
doid, Liancalus virens Scopoli (Vaillant, 1948). The larval growth rate for
all species varied considerably as experimentation to determine food
requirements progressed, although in one batch of eggs of M. exquisita,
final instar larvae (Fig. 5) developed relatively rapidly, within ap-
proximately five weeks from the time of hatching. Final instar larvae of
all species that were still alive towards the end of the summer were cooled
down and allowed to diapause at 1 C for three months. Most larvae sur-
vived the diapause period, and in one instance, a single larva of M.
exquisita pupated (without forming a cocoon) in the agar medium
approximately five weeks after the temperature was increased to 15 C.
Within three weeks the pupa developed to the teneral adult stage (Fig. 8),

96 ENTOMOLOGICAL NEWS

but died before eclosion occurred.

Food Requirements. Attempts were made to determine the food
requirements of the larvae, since the diet of larval tachydromiines is
unknown. Although never directly established, tachydromiine larvae
have been assumed to be predacious (Chvala and Kovalev, 1989), based
on limited observations of other empidoid species (reviewed by Smith,
1969, p. 6). First instar larvae of all four tachydromiine species were
offered various small soil organisms, such as nematodes belonging to
the genus Panagrellus, all stages of the oribatid mite species Oppia nitens
C.L. Koch, and an inoculum of protozoans, but feeding on these mic-
roorganisms was not observed. In addition, early as well as later instar
larvae did not appear to scavenge on dead organisms, or on moistened
pieces of highly proteinaceous dried puppy meal, which were also added
to some cultures.

Finally, small larvae of the pomace fly, Drosophila melanogaster
Meigen, were presented as prey. Larvae of all four species of tachy-
dromiines readily fed on the Drosophila larvae within the agar medium
(Fig. 7). The smaller first instar tachydromiine larvae however, were only
able to overpower the smallest (first instar) Drosophila larvae. Later
tachydromiine instars fed on Drosophila larvae of various sizes, and
occasionally these older predacious larvae also cannibalized smaller
sibling larvae in the same culture. Since Drosophila larvae could survive
in the agar medium for one or two days before starving, periodic replace-
ment of prey larvae was required to sustain the tachydromiine larval
cultures.

Reproductive Mode. Females of P. holosericus are suspected of re-
producing parthogenetically (as has been suggested by Tuomikoski,
1935 and Chvala, 1975 for some Palearctic species of Platypalpus), since
males of this common Nearctic species are not represented in the main
North American empidoid collections, and have never been collected
locally. To determine the reproductive mode in P. holosericus, ovaries
were dissected from four gravid females and transferred individually to
eight agar plates for observation. During dissection each ovary, contain-
ing an average of approximately 30 fully mature eggs, was surgically
removed from the lateral oviduct to prevent any possibility of accidental
fertilization. Despite heavy fungal contamination of the ovariole tissue,
the occurrence of parthenogenesis in P. holosericus was convincingly
demonstrated when most of the eggs in all eight of the dissected
ovaries hatched.

DISCUSSION

The results obtained to date provide a first step in the development of
a general procedure for rearing tachydromiine flies, which will aid in the
accumulation of valuable taxonomic and life history data. Most impor-

Vol. 104, No. 2, March & April, 1993

97

Figs. 1-8. 1, Seal-tight petri dish filled with saline nutrient agar medium for rearing Tachy-
dromiinae (0.6 X); 2, eggs ofPlatypalpu.s holosericus Melander containing developing larvae
(33 X); 3, first instar larvae of P. holosericus in agar medium ( 18 X); 4, first instar larva of P.
holosericus in agar medium (36 X); 5, late instar larva of Megagrapha exquisita ( Malloch ) on
top of agar medium (12 X); 6, first instar larvae of P. holosericus amongst fungal contamina-
tion of agar medium (27 X); 7, late instar larva of P. holosericus feeding on early instar larva
of Drosophila melanogaster Meigen (18 X); 8. lateral view of pupa of M exquisita containing
teneral adult (14 X).

98

ENTOMOLOGICAL NEWS

tantly, induction of oviposition behavior by decapitation, in conjunc-
tion with maintenance of the eggs in agar-filled petri dishes to avoid
desiccation, appears to be a useful technique for obtaining adult-
associated first instar larvae (and possibly later stages) of Tachy-
dromiinae 5 The technique may work with, and should be attempted on,
other poorly known Empidoidea (e.g. Atelestinae, Brachystomatinae,
Ceratomerinae, Microphorinae, and Nemidind) and taxonomically
problematic Cyclorrhapha (e.g. Opetidae), for which immature stages
are not known (see for example Sinclair, 1992). Even if the first instar lar-
vae obtained are not reared successfully to a further stage, the taxonomic
information gained from having properly associated immatures of any
stage for such groups, would be valuable for testing previously proposed
classifications and formulating new phylogenetic hypotheses.

Dissection of ovaries from parthenogenetic species, such as P. holo-
sericus, can also yield large numbers of first instar larvae. Of perhaps
greater significance however, is the use of this dissection procedure for
determining whether certain females within a species, or all females, are
able to reproduce parthenogenetically rather than bisexually. Partheno-
genesis is presumed to occur in certain species of the genus Platypalpus,
where males have been rarely collected, or remain unknown. In the
Palearctic Region, some species are thought to be entirely partheno-
genetic [e.g. P. major (Zetterstedt) (Chvala, 1975, 1989)], or partially
parthenogenetic [e.g. P. ecalceatus (Zetterstedt) (Chvala, 1989; Tuomi-
koski, 1935)] throughout their range, whereas others [e.g. P. candicans
(Fallen) and P. cursitans (Fabricius) (Chvala, 1975; Frey, 1943; Tuomi-
koski, 1935)] appear to exhibit geographic parthenogenesis. Conclusive
determination of reproductive mode for species of Platypalpus at the
population level could be important for future screening of possible
biological control agents. This is because increased reproductive poten-
tial associated with parthenogenesis can be a desirable attribute for
beneficial insects being considered for release programs (Aeschlimann,
1990; Doutt et al, 1976), and mating requirements for parthenogenetic
females can be effectively ignored.

Larval food requirements of empidoids in general are poorly
understood, and no information is available for Tachydromiinae. The
procedures outlined here however, allow for experimentation with
various prey organisms and other food materials to determine which
broad categories of food types can be consumed by larvae. The results

Morphological study of the immature stages of the Tachydromiinae will he dealt with in
subsequent papers.

Vol. 104, No. 2, March & April, 1993 99

obtained on the four species studied here, indicate that larvae of many if
not all Tachydromiinae are predacious, probably on small soil or litter
inhabiting organisms such as other Diptera larvae, rather than being
saprophagous or microorganism feeders. The use of larval Diptera as the
major source of prey for the larvae of some other empidoid groups, has
been noted by Smith (1969). The apparent attraction of the larvae of all
four tachydromiine species towards areas of heavy fungal and bacterial
contamination in older cultures, suggests that natural organisms cap-
tured by these predacious larvae probably include small mycetophagous
or saprophagous Diptera larvae, or other soft-bodied prey.

A general procedure for rearing tachydromiine flies could have
important implications for biological control programs targeted against
a variety of small-sized insect pests. For example, the predatory activity
of many adult tachydromiines appears to be both intense and of long
duration (Chvala, 1975; Stark and Wetzel, 1989; Whitfield, 1925), and
adults are considered to occupy small-sized predator niches not gen-
erally shared by other (usually larger-sized) predators (Chvala, 1975).
This, in conjunction with the ability of several species to reach very high
population densities (e.g. recorded as high as 40 to 60 individuals of
Platypalpus per meter 2 in cereal crops by Stark, 1990), and the apparent
lack of a fixed diapause stage in at least some species of Crossopalpus,
Platypalpus, and Stilpon (Chvala, 1975) suggests potential benefits for the
development of future mass-rearing programs for this group of preda-
cious flies. Common parthenogenetic species like the relatively large,
voracious P. holosericus in North America, or the very similar European
P. major, appear to be ideal candidates for further research.

ACKNOWLEDGMENTS

We would like to thank A Borkent (Salmon Arm, B.C.) for his discussions on
ceratopogonid rearing techniques and J. R. Linley (University of Florida, Vero Beach) for
graciously supplying supplementary vitamins for the saline nutrient agar. J. E. O'Hara
(Biological Resources Division CLBRR) and B. J. Sinclair(Carleton University, Ottawa)
kindly reviewed the manuscript and suggested many improvements.

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Beling, T. 1888. Beitrag zur Metamorphose einiger zweiflugeliger Insecten aus den

Familien Tabanidae, Empidae, und Syrphidae. Verh. Zool. Bot. Ges. Wien 38: 1-4.
Berest, Z. L. 1987. Trophic relations of natural enemies of cereal leaf aphids. Vestnik Zool.

21:45-48. [In Russian.]

!00 ENTOMOLOGICAL NEWS

Brunei, E., P. Grootaert and J. Mesquida. 1989. Entomofaune associee a la floraison du

colza (Brassica napus L.): Note preliminaire sur les Dolichopodidae et les Empididae

(Insectes: Diptera). Med. Fac. Landbouww. Rijksuniv. Gent 54(3a): 727-737.
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Fauna Entomol. Scand. 3: 1-336.
Chvala, M. 1989. Monograph of northern and central European species of Platypalpus

(Diptera, Hybotidae), with data on the occurrence in Czechoslovokia. Acta Univ.

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Chvala, M. and V. G. Kovalev. 1989. Family Hybotidae, pp. 174-227. In. Soos, A. and L.

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insects and spiders by ELISA. Ann. Appl. Biol. 105: 413-422.
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951-998.
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Fleschner, C. A. and D. W. Ricker. 1953. An empidid fly predacious on citrus red mites. J.

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102 ENTOMOLOGICAL NEWS

ESTABLISHMENT OF HIPPODAMIA VARIEGATA

AND NEW RECORDS OF PROPYLEA

QUATUORDECIMPUNCTATA (COLEOPTERA:

COCCINELLIDAE) IN THE EASTERN

UNITED STATES 1

A. G. Wheeler, Jr. 2

ABSTRACT: Hippodamia variegata is a Palearctic coccinellid known previously in North
America from a few areas of eastern Canada. It has been released in eastern and western
states for biological control of aphids, but its establishment in the United States has not
been documented. On the basis of late-season surveys in the northeast in 1992, H. variegata
is reported from 38 counties in eight states from northern New England to eastern
Pennsylvania and northern New Jersey. All localities surveyed are mapped. Its abundance
relative to other coccinellines collected on weeds in disturbed habitats and its plant
associations are indicated; the origin of U.S. populations is discussed. Records forPropylea
quaiuordecimpunctata. another Old World coccinellid, are given for Massachesetts, New
Hampshire, New York, and Vermont.

Hippodamia (Adonia) variegata (Goeze) is an Old World coccinellid
first recorded from North America by Gordon (1987). He reported its
establishment in the vicinity of Montreal, Quebec, noting that Nearctic
populations may be adventive rather than the result of intentional
releases. This aphid predator was released in the United States (Arizona,
California, Florida, and Georgia) beginning in 1957-1958, but no record
of Canadian releases is available (Gordon 1985, 1987).

Hippodamia variegata (South African strain) was evaluated in the
laboratory (and eventually released) as a potential biological control
agent of the greenbug, Schizaphis gramium (Rondani), that could in-
crease the diversity of coccinellid predators in Texas sorghum fields
(Michels and Bateman 1986). Invasion of the western United States by
the Russian wheat aphid, Diuraphis noxia (Mordvilko), in 1986 (Stoetzel
1987), led to foreign exploration for natural enemies of this introduced
pest and the introduction of various strains of//, variegata from Eurasia.
In 1987, it was released in several eastern and western states by the
USDA's Animal and Plant Health Inspection Service (APHIS) (Obrycki
and Orr 1990, Flanders et al. 1991). Several biological studies on this
introduced aphidophagous coccinellid have been conducted in North
America, including its developmental rates at several constant tem-
peratures (Michels and Bateman 1986) and an evaluation of several

1 Received December 3, 1992. Accepted December 28. 1992.

2 Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA
17110-9408.

ENT. NEWS 104(2): 102-1 10, March & April. 1993

Vol. 104, No. 2, March & April, 1993 103

aphid species as suitable prey (Obrycki and Orr 1990).

Despite numerous releases in western and eastern states since 1987
(including Colorado, Kansas, Maryland, Massachusetts, New Mexico,
Pennsylvania (Flanders el al. 1991 ), and New Jersey (R. Chianese, per-
sonal communication),//, variegata has been recorded in North America
only from Quebec and Ontario (Gordon and Vandenberg 1991,
McNamara 1991). Since 1990, participants in the USDA's Cooperative
Agricultural Pest Survey (CAPS) have been asked to look for//, variegata
in the northeast.

The purpose of this paper is to document the establishment of//.
variegata in eight northeastern states, map the known U. S. range, and
provide information on its abundance relative to other coccinellids
occurring in the same habitats. In addition, new records are given for
Propylea quatuordecimpunctata (L.), another coccinellid that CAPS par-
ticipants have been asked to search for in northeastern states.

METHODS

After I discovered //. variegata in New York and Vermont in late
August 1992, surveys were begun to help determine the extent of its
northeastern range. The habitats surveyed (11-13,1 8-20, and 27 Septem-
ber, and 6 October) were those that seemed likely to support a diverse
coccinellid fauna, particularly disturbed, weedy sites such as railroad
yards and urban vacant lots. Mowed roadside vegetation generally
yielded few coccinellids, and cropland was not surveyed because of the
time that would have been needed to obtain permission for sampling.

Herbaceous vegetation was swept with a standard insect net, and all
adult Coccinellini and numbers of each species (except in late August
and early October) were recorded. At some sites, particular plant species
were examined to determine host associations of//, variegata. Totals of
all species at a site include adults collected by both techniques. Even
though the duration of sampling (usually 10-15 minutes), number of
sweeps, and vegetation varied among the sites, the numbers of coc-
cinellines recorded at a given locality allow comparisons of relative den-
sity between //. variegata and other coccinellids present during Sep-
tember.

Specimens thought to represent H. variegata (and those of several
other species that could not be identified accurately in the field) were
collected for subsequent determination. Voucher material of//, variegata
and Propylea quatuordecimpunctata has been deposited in collections at
Cornell University, Ithaca, NY (CUIC); National Museum of Natural
History, Washington, DC (USNM); and Pennsylvania Department of
Agriculture, Harrisburg (PAD A).

104

ENTOMOLOGICAL NEWS

RESULTS

Eleven coccinelline species in six genera were encountered during
surveys for Hippodamia variegata (Table 1). Nearly 250 specimens of//.
variegata were obtained at 48 localities in eight states: Connecticut (5
counties), Massachusetts (6), New Hampshire (2), New Jersey (1), New
York (16), Pennsylvania (3), Rhode island (1), and Vermont (4). Positive
and negative sites for H. variegata, as well as recent release sites in
Massachusetts, New Jersey, and Pennsylvania, are shown in Fig. 1.

Figure 1 . Known U.S. distribution of Hippodamia variegata. Dots = detection sites; circles =
sites where the coccinellid was not found; stars = recent release sites in Massachusetts,
New Jersey, and Pennsylvania (not shown are 1957-1958/1987 release sites in Delaware,
Maine, and Maryland).

Vol. 104, No. 2, March & April, 1993 105

The following new records document the establishment of//, varie-
gata in the eastern United States; all collections were made by the author
from 28 August to 6 October 1992.

CONNECTICUT: Fairfield Co., Danbury; Hartford Co., New Brittain; New Haven
Co., Waterbury; Tolland Co., Mansfield; Windham Co., Abington and Dayville.
MASSACHUSETTS: Berkshire Co., Rt.41 S.of Housatonic; Franklin Co., Ervingand
Greenfield; Hampden Co., Springfield; Hampshire Co., Northampton; Middlesex Co.,
Marlborough; Worcester Co., Athol, Gardner. Millbury, and Winchendon. NEW
HAMPSHIRE: Cheshire Co., North Walpole and Troy; Sullivan Co., Claremont. NEW
JERSEY: Sussex Co., McAfee and Rt. 23 E. of Montague. NEW YORK: Albany Co., Rt.
90E, Service Plaza, Town of Rotterdam nr. Albany; Clinton Co., June. rts. 87 & 456 E. of
Beekmantown; Columbia Co., Hillsdale; Dutchess Co., Fishkill; Greene Co., Leeds;
Orange Co., Newburgh and Port Jervis; Otsego Co., Cooperstown Junction; Putnam
Co., Brewster; Rensselaer Co., Brunswick; Saratoga Co., Ballston Spa; Schenectady
Co., Duanesburg; Schoharie Co., Cobleskill; Sullivan Co., Wurtsboro; Ulster Co.,
Kerhonkson; Warren Co., Glens Falls; Washington Co., Whitehall. PENNSYLVANIA:
Philadelphia Co., Philadelphia; Pike Co., Matamoras; Wayne Co., Hawley and
Waymart. RHODE ISLAND: Providence Co., Chepachet and Nasonville. VERMONT:
Addison Co., Vergennes; Chittenden Co., Colchester; Rutland Co., Fair Haven;
Washington Co., Montpelier.

In northern New England and northeastern New York, H. variegata
was found at 20 of 29 sites sampled during 1 1 - 1 3 September. It was either
the only coccinellid species or the most numerous one at 1 3 of those sites.
During 18-20 September, it was present at 22 of 24 sites in southern New
England, southeastern New York, and eastern Pennsylvania and was
most abundant at 1 1 sites. It was taken at 4 of 9 sites in northeastern
Pennsylvania and northern New Jersey on 27 September and was most
numerous at 2 sites. It was found at one location in Philadelphia during
limited surveys in southeastern Pennsylvania on 6 October.

Hippodamia variegata was frequently taken by sweeping legumes
such as red clover (Trifolium pratense L.) and sweet clover (Melilotus spp.)
that were infested with pea aphids, Acyrthosiphon pisum (Harris), or by
beating inflorescences of horseweed (Conyza canadensis (L.) Cronquist).
It was also observed on volunteer alfalfa (Medicago sativa L.) and on com-
posites such as aster (Aster spp.), chicory (Cichorium intybus L.), golden-
rod (Solidago spp.), mugwort (Artemisia vulgaris L.), ragweed (Ambrosia
artemisiifolia L.), spotted knapweed (Centaurea maculosa Lam.), and
tansy (Tanacetum vulgare L.). A mating pair was found under a mat of
knotweed (Polygonum aviculare L.).

An adult H. variegata collected 20 September at Marlborough,
Massachusetts, was not killed right away. A few days later a parasitoid
cocoon was observed beneath its body, and the braconid Dinocampus
coccinellae (Schrank) emerged on 3 October. This Holarctic species is a

10 6 ENTOMOLOGICAL NEWS

known parasitoid of//, variegata (and other coccinellids) in Europe. The
Massachusetts record from H. variegata is noteworthy because this coc-
cinellid had proved unsuitable as a host (100% mortality) in laboratory
studies using North American D. coccinellae (Obrycki 1989). Successful
parasitism of Canadian populations of//, variegata has since been re-
ported (Orr et al. 1992).

Propylea quatuordecimpunctata was collected in 16 counties in four
states: Massachusetts (1 county), New Hampshire (1), New York (9), and
Vermont ( 5). It occurred at 1 1 of 29 sites during 11-13 September (and was
most abundant at 1 site) and 3 of 24 sites the following week. It was not
found during surveys of northeastern Pennsylvania and northern New
Jersey on 27 September or in the limited southeastern Pennsylvania sur-
veys on 6 October. The largest number of specimens (>20; not shown in
Table 1) was observed in late August at the Clinton Co., New York, site
listed below. The following records of P. quatuordecimpunctata were
obtained from 28 August to 27 September 1992.

MASSACHUSETTS: Franklin Co., Greenfield. NEW HAMPSHIRE: Grafton Co.,
West Lebanon. NEW YORK: Albany Co., Rt. 90E, Service Plaza, Town of Rotterdam nr.
Albany; Clinton Co., June. rts. 87 & 456 E. of Beekmantown; Dutchess Co., Fishkill;
Orange Co., Newburgh; Putnam Co., Brewster; Rensselaer Co., Brunswick; Saratoga
Co., Ballston Spa; Warren Co., Glens Falls; Washington Co., Whitehall. VERMONT:
Addison Co., Vergennes; Chittenden Co., Colchester; Rutland Co., Fair Haven; Wash-
ington Co., Montpelier; Windham Co., Brattleboro.

DISCUSSION

Hippodamia variegata should be considered a common and wide-
spread coccinellid in the northeastern states; the localities reported
herein can be regarded as the first records of establishment in the United
States. It was generally present in the areas surveyed except in more
western portions of eastern New York and in parts of northern New Jer-
sey and eastern Pennsylvania.

The current U.S. range of H. variegata could reflect expansion of
Canadian populations discovered in 1984 (Gordon 1987). As noted ear-
lier, it cannot be determined if this coccinellid's occurrence in Quebec is
the result of a fortuitous importation with commerce or deliberate
introduction associated with biological control work. Its extensive
northeastern distribution suggests H. variegata was present in the United
States when Gordon (1987) gave Quebec as the first North American
record.

Rather than having spread rapidly from the Montreal area, this coc-
cinellid may be present in the eastern states as a result of earlier U.S.
releases (probably those since 1987 rather than ones during 1957-1958)

Vol. 104, No. 2, March & April, 1993 107

that led to its establishment, which is only now being documented.
Schaefer <?/ a/. (1987) offered a similar hypothesis as one explanation of
the North American origin oiCoccinella septempunctata: that establish-
ment from earlier biocontrol releases went undetected for several years.
Northeastern and southeastern Pennsylvania populations of H. varie-
gata may be discontinuous, and its occurrence in Philadelphia may be
the result of 1987 releases there, even though recovery attempts have
been unsuccessful (Flanders et al. 1991 ). Similarly, the New Jersey pop-
ulations may be the result of establishment from 1991 releases (see
Fig. 1).

Evidence for evaluating the status of//, variegata in the New World
adventive or indigenous is meager, emphasizing a general need for
more field work and documentation of insect distributions. It also points
to the desirability of recording all sites where nonindigenous organisms
are released for biological control purposes and of conducting thorough
recovery surveys to determine whether establishment has taken place.
Areas well removed from release sites should be surveyed to allow for
dispersal by highly mobile species.

That H. variegata was the most abundant coccinellid occurring at
several sites was surprising. The comments to be made on other species
collected during the survey are speculative because numerous factors
affect the composition and abundance of coccinellid communities
(Hagen 1962, Hodek 1973, Honek 1985), and widely distributed species
can vary intraspecifically in various diapause characteristics (Hodek
1973; Obrycki and Tauber 1981, 1982; Tauberer a/. 1986).

The small numbers ofAdalia bipunctata taken on herbaceous plants
during the survey were expected because it prefers arboreal habitats
(Hodek 1973, Honek 1985). The generalist P. quatuordecimpunctata,
which was not abundant in the survey, tends to occur at relatively low
densities (Honek 1985). This is a distinctive, easily recognized coc-
cinellid, and, in contrast to //. variegata, its establishment and spread in
the United States have been documented (Dysart 1988, Wheeler 1990).
Coccinella septempunctata, a Palearctic species that has spread rapidly to
become the most common member of the genus east of the Rocky Moun-
tains (Gordon and Vandenberg 1991 ), was found in relatively low num-
bers during the September survey. The convergent lady beetle, //. con-
vergens, superficially resembles //. variegata in dorsal color pattern, and
the abundance of this native species was thought to be hindering detec-
tion of//, variegata in the East. Only one adult //. convergent, however,
was collected during the survey.

The small numbers of C. septempunctata and H. convergent seen in
September do not necessarily indicate low densities for the entire season.

108 ENTOMOLOGICAL NEWS

In a New Jersey study, the number of both species that were swept from
herbaceous weeds declined rapidly through August from much higher
June-July levels and were low in September. Coccinella septempunctata
shows a prolonged aestival-autumnal-hibernal diapause (Obrycki and
Tauber 1 98 1 ), and in New Jersey it began to aggregate as early as mid-July
(Angalet et al. 1979). Hippodamia convergens does not enter aestival
diapause in the Ithaca, New York, area (Obrycki and Tauber 198 1 ), but it
probably also would have been more abundant in the present study if
sites had been sampled earlier in the season.

In contrast, the failure to collect Coccinella novemnotata during the
survey possibly reflects its current scarcity. Biological control releases of
the Palearctic C. septempunctata may be contributing to the declining
numbers observed recently in populations of the native C. novemnotata. 3
In Virginia, the dominance of C. septempunctata is thought responsible
for the disappearance of both C. novemnotata and H. convergens from
alfalfa fields/*

Detection of H. variegata at additional northeastern localities is
favored by the absence or low density of//, convergens in late season.
Where these species co-occur in the East, //. variegata adults generally
can be recognized in the field by their smaller size compared to the con-
vergent lady beetle. Field identifications must be substantiated by mic-
roscopically observing a fine raised margin or bead at the pronotal base
(absent in //. convergens and similar-appearing eastern species of the
genus) and the white anterior coxae (black in H. convergens). Gordon
(1987) and Gordon and Vandenberg (1991) provide additional charac-
ters for recognizing this quite variable coccinellid.

Further survey work undoubtedly will show that H. variegata is even
more widely distributed in eastern North America. Its continued spread
in the East, establishment in agroecosystems, association with various
aphid prey and impact on their densities, and possible effects on pop-
ulations of native coccinellids such as H. convergens require further
study.

"Biological Control, Predators, and Strategy," a paper presented by R. D. Gordon, 24
September 1991, at annual meeting of the Eastern Branch, Entomological Society of
America, Richmond, Virginia.

"Apparent Displacement of the Convergent Lady Beetle by the Sevensported Lady
Beetle," a paper presented by R. L. Pienkowski, 24 September 1991 , at the annual meeting
of the Eastern Branch, Entomological Society of America, Richmond. Virginia.

Vol. 104. No. 2, March & April, 1993 109

Table 1. Adult Coccinellini collected during surveys for Hippodamia variegata in north-
eastern United States, September 1992. a

Species

No. of
Specimens

No. of Maximum No.
Sites Collected

Adalia bipunctata L.

3

2

2

Coccinella septempunctata L.

66

23

12

C. transversogutlata richardsoni Brown

1

1

1

C. trifasciata perplexa Mulsant

8

5

3

Coleomegilla maculata lengi Timberlake

67

29

12

Cycloneda munda (Say)

92

24

23

Hippodamia convergens Guerin

1

1

1

H. glacialis glacialis (F.)

7

4

3

H. parenthesis (Say)

102

30

22

H. variegata (Goeze)

210

45

27

Propylea quatuordecimpunctata L.

24

14

5

a Numbers of H. variegata, Propylea quatuordecimpunctata, and other species collected
during 28-29 Aug. and 6 Oct. 1992 are not included.

ACKNOWLEDGMENTS

I thank E. R. Hoebeke (Department of Entomology, Cornell University, Ithaca, NY) for
confirming my identification of//, variegata and reviewing the manuscript; R. V. Flanders
(USDA, APHIS, National Biological Control Institute, Hyattsville, MD) for reviewing the
manuscript; T. L. Burger and D. J. Nelson (USDA, APHIS, National Biological Control
Laboratory, Niles. MI), P. W. Schaefer (USDA, ARS, Beneficial Insects Research
Laboratory, Newark, DE), and R. Chianese (Division of Plant Industry, New Jersey
Department of Agriculture, Trenton) for information on eastern U. S. releases of H.
variegata; and C. Jung (Denver, CO) for providing Fig. 1.

LITERATURE CITED

Angalet, G.W., J.M. Tropp, and A.N. Eggert. 1979. Coccinella septempunctata in the

United States: recolonizations and notes on its ecology. Environ. Entomol. 8:896-

901.
Dysart, RJ. 1988. The European lady beetle Propylea quatuordecimpunctata: new locality

records for North America (Coleoptera: Coccinellidae). J. New York Entomol. Soc. 96:

119-121.
Flanders, R.V., DJ. Nelson, R. Deerburg, and CJ. Copeland. 1991. Aphid biological

control project: FY 1990 project report. USDA-APHIS Biological Control Laboratory,

Niles, Mich. 20 pp. + 26 tables.
Gordon, R.D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. J. New

York Entomol. Soc. 93:1-912.
Gordon, R.D. 1987. The first North American records of Hippodamia variegata (Goeze)

(Coleoptera:Coccinellidae). J. New York Entomol. Soc. 95:307-309.

1 10 ENTOMOLOGICAL NEWS

Gordon, R.D. and N. Vandenberg. 1991. Field guide to recently introduced species of

Coccinellidae (Coleoptera) in North America, with a revised key to North American

genera of Coccinellini. Proc. Entomol. Soc. Wash. 93:845-864.
Hagen, K.S. 1962. Biology and ecology of predaceous Coccinellidae. Annu. Rev. Entomol.

7:289-326.

Hodek, I. 1973. Biology of Coccinellidae. Junk, The Hague; Academia, Prague. 260pp.
Honek, A. 1985. Habitat preferences of aphidophagous coccinellids (Coleoptera). Ento-

mophaga 30:253-264.
McNamara, J. 1991. Family Coccinellidae, ladybird beetles, pp. 229-237. In : Y. Bousquet

(ed.). Checklist of beetles of Canada and Alaska. Res. Branch Agric. Can. Publ.

1861/E.
Michels, GJ., Jr. and A.C. Bateman. 1986. Larval biology of two imported predators of

the greenbug, Hippodamia variegata Goetz (sic) and Adalia flavomaculata DeGeer,

under constant temperatures, Southw. Entomol. 11:23-30.
Obrycki, JJ. 1989. Parasitization of native and exotic coccinellids by Dinocampus coc-

cinellae (Schrank) (Hymenoptera:Braconidae). J. Kans. Entomol. Soc. 62:21 1-218.
Obrycki, J J. and C J. Orr. 1990. Suitability of three prey species for Nearctic populations

ofCoccinellaseptempunctata. Hippodamia variegata. and Propylea quatuordecimpunctata

(Coleoptera:Coccinellidae). J. Econ. Entomol. 83:1292-1297.
Obrycki, JJ. and MJ. Tauber. 1981. Phenology of three coccinellid species: thermal

requirements for development. Ann. Entomol. Soc. Am. 74:31-36.

Obrycki, JJ. and MJ. Tauber. 1982. Thermal requirements for development of Hip-
podamia convergens (Coleoptera:Coccinellidae). Ann. Entomol. Soc. Am. 75:678-683.
Orr, C J., J J. Obrycki, and R.V. Flanders. 1992. Host-acceptance behavior of Dinocam-
pus coccinellae (Hymenoptera: Braconidae). Ann. Entomol. Soc. Am. 85:722-730.
Schaefer, P.W., R J. Dysart, and H.B. Specht. 1987. North American distribution ofCoc-

cinella septempunctata (Coleoptera:Coccinellidae) and its mass appearance in coastal

Delaware. Environ. Entomol. 16:368-373.
Stoetzel, M.B. 1987. Information on and identification of Diuraphis noxia (Homop-

tera:Aphididae) and other aphid species colonizing leaves of wheat and barley in the

United States. J. Econ. Entomol. 80:696-704.
Tauber, M J., C.A. Tauber, and S. Masaki. 1986. Seasonal adaptations of insects. Oxford

University Press, New York. 41 1pp.
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adventive lady beetle (Coleoptera:Coccinellidae). Entomol. News 101:164-166.

Vol. 104, No. 2. March & April. 1993 111

EPILACHNA VIGINTIOCTOPUNCTATA

(COLEOPTERA: COCCINELLIDAE),

NEW RECORD FOR WESTERN HEMISPHERE,

WITH A REVIEW OF HOST PLANTS l

Robert F. W. Schroder, 2 Michael M. Athanas,2 Crodowaldo Pavan^

ABSTRACT: The first discovery of the phytophagous coccinellid Epilachna vigintioc-
topunctata (F.) in the Western Hemisphere was made October 8, 1990 on wild cucurbit
plants near Curitiba, Parana, Brazil, new records since that discovery are given for
Paranagua, Parana and Itajai, Santa Catarina. host plants of agricultural importance
are given.

Epilachna vigintioctopunctata (F.) is reported for the first time in the
Western Hemisphere, it was not known to occur in the Western Hemi-
sphere prior to this report (Anonymous 1992). Gordon (1975) makes no
mention of this pest in the Western Hemisphere.

Nine adult beetles were collected on October 8, 1990 feeding on wild
cucurbit vines (species unknown) found along Highway 101 about 10
kilometers south of Curitiba, Parana, Brazil. No eggs or larvae were
found at the site. The adults were similar in appearance and size to
Epilachna varivestis Mulsant, having a brownish yellow color with dark
brown spots on the elytron. Four specimens were identified by Robert D.
Gordon, USDA, Systematic Entomology Laboratory, as the phyto-
phagous coccinellid E. vigintioctopunctata (E. 28-punctata). During a trip
to Brazil in April 1 992, we visited with Sonia M. N. Lazzari, a specialist in
taxonomy of Coleoptera, Department of Zoology, Federal University of
Parana, Curitiba. She indicated that on December 31. 1991, they col-
lected one E. 28-punctata adult from a pepper plant at Paranagua, a coas-
tal town east of Curitiba. on April 26, 1992, 25-30 adults were found
defoliating Solanum americanum along the coastal highway near Itajai,
Santa Catarina, Brazil. Adults sent to R. D. Gordon were also identified
as E. 28-punctata. During the period from April 26 to May 5, 1992, we did
not find any other specimens in the regions surveyed near Passo Fundo
(RS), Ponta Grossa (Parana) and Campinas (SP). Voucher specimens
are deposited in the Federal University of Parana. Department of Zool-
ogy Insect Collection.

Schaefer ( 1 983) indicates that this coccinellid is a serious pest in Asia

1 Received October 24, 1992. Accepted November 23. 1992.

- USDA. Agricultural Research Service. Plant Sciences Institute. Insect Biocontrol

Laboratory. Beltsville, MD 20705
3 Department of Genetics and Evolution, Institute of Biology. UNICAMP. Campinas.

S.P.. Brazil.

ENT. NEWS 104(2): 1 1 1-1 12. March & April. 1993

112 ENTOMOLOGICAL NEWS

and is reported on a wide range of host plants of agricultural importance
including potato, eggplant, tomato, tobacco, bitter gourd, sweet gourd,
ribbed gourd, snake gourd, cucumber, pumpkin, zucchini, beet, sugar-
beet, marrow, cotton melon, rock melon, squash, cowpea, peanut, okra,
alfalfa, vetch, clover, cotton and banana. This list does not include weeds
and grasses that Schaefer mentions.

According to Richards and Filewood (1990), there were 3 different
Australian species of Epilachna that were incorrectly referred to as E. 28-
punctata. He identified them as subspecies in the. 28-punctata complex,
all occurring in Australia. Two of the subspecies, E. 28-punctata pardalis
(Boisduval) and E. vigintisexpunctata vigintisexpunctata (Boisduval) feed
on solanaceous plants, and E. cucurbitae Richards feeds on cucurbits.
Another subspecies in the complex, Epilachna 28-punctata 28-punctata
(Fabricius) is not known to occur in Australia, but is found in India,
Pakistan, Japan, SE Asia and Oceania. All are important pests of
agricultural crops. Richards at least clarified the situation for Australia.
Because of the existing confusion over identification, or species limits,
much greater uncertainty exists over the host plants list for E. 28-punctata
compiled by Schaefer. This is only one example of a greater problem that
exists throughout the range of this species.

There is reason to be concerned about this new pest in the Western
Hemisphere, because it is reported as a serious pest on many eco-
nomically important crops in Asia. Since it looks very similar to E.
varivestis, its presence may go undetected in regions of the Western
Hemisphere where E. vrivestis is known to occur. Epilachna 28-punctata is
a new introduction, restricted to the coastal region of Brazil. Other than
the defoliation of the S. americanum plants from which we collected the
beetle, there are no other reports of it causing any damage to plants.
Therefore, the opportunity exists to delineate its distribution, restrict its
spread and maximize the use of a wide range of biocontrol agents
(Schaefer 1983) to reduce the potential threat of this pest to agricul-
ture in Brazil.

LITERATURE CITED

Anonymous. 1992. CAB, International Institute of Entomology, Distribution Maps of
Pests, Series A; map No. 532.

Gordon, R. D. 1975. A revision of the Epilachninae of the Western Hemisphere (Coleop-
tera: Coccinellidae). USDA Tech. Bull. 1943, 409 pp.

Richards, A. M. and L. W. Filewood. 1990. Feeding behavior and food preferences of
pest species comprising the Epilachna vigintioctopunctata (F.) complex (Col., Coc-
cinellidae). J. Appl. Ent. 1 10:501-515.

Schaefer, P. W. 1983. Natural enemies and host plants of species in the Epilachninae
(Coleoptera: Coccinellidae) a world list. University of Delaware, Agricultural
Experiment Station Bulletin 445, 42 pp.

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MAY & JUNE, 1993

USISSN0013-872X
NO. 3

NEWS

Identification of Osmia kenoyeri and O.
virga (Hymenoptera: Megachilidae),
two blueberry pollinators R.W. Rust, E.A. Osgood 113

Tabanidae as dietary items of Rafinesque's
big-eared bat: implications for its
foraging behavior

Susan E.Ellis 118

A huge nest of the common yellowjacket,
Paravespula vulgaris (Hymenoptera:
Vespidae), in California

R.D. Akre, E.A. Myhre, Y. Chen 123

Seasonal flight activity of Lipoptena

mazamae (Diptera: Hippoboscidae) in

South Carolina L.D. Cline, J.E. Throne

Overwintering aggregations of female
Brachymeria intermedia
(Hymenoptera: Chalcididae)

First records of parasitoids for slime mold
beetles of the family Sphindidae
(Coleoptera: Cucujoidea) Joseph V. McHugh

First report of a twisted-wing insect
(Strepsiptera) larva in a caddisfly
(Trichoptera) K.L. Manuel, R.M. Bohart

A method for sex determination of
Colorado potato beetle pupa,
Leptinotarsa decemlineata
(Coleoptera: Chrysomelidae)

129

Paul W. Schaefer 133

136

139

Yvan Pel letter 140

Establishment of Urophoraquadrifasciata
(Diptera: Tephritidae) and
Chrysolina quadrigemina
(Coleoptera: Chrysomelidae) in
portions of eastern U.S. E. Richard Hoebeke 143

BOOKS RECEIVED AND BRIEFLY NOTED 117, 128, 135

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Vol. 104, No. 3. May & June, 1993 113

IDENTIFICATION OF OSMIA KENOYERI AND

O. VIRGA (HYMENOPTERA: MEGACHILIDAE),

TWO BLUEBERRY POLLINATORS 1 ' 2

Richard W. Rust 3 , Eben A. Osgood 4

ABSTRACT: The sexes of Osmia kenoyeri and O. virga are associated and the females of
both are described. Both species visit lowbush blueberry, Vaccinium spp. for pollen and
nectar. The nesting biology of O. kenoyeri is described.

The genus Osmia contains over 130 species in North America
(Sandhouse 1939, Hurd 1979). Many of the species are only known by
one sex, often the male, because of distinctive abdominal sternal and
genitalic characters. Here we associate the sexes and describe for the first
time the females of Osmia kenoyeri Cockerell and O. virga Cockerell. Both
species have been collected on Vaccinium angustifolium Art. and V. myr-
tilloides Michx. in the lowbush blueberry complex in Maine (Boulanger
et al. 1967, Stubbs et al. 1992).

Osmia (Acanthosmioides) kenoyeri Cockerell

Female: Length 10 mm; fore wing 6.5 mm; head width 3.5 mm. Color dark greenish-
blue; antennae, mandibles, legs dark redbrown to black; tegulae black with greenish-blue
apical edge. Pubescence of vertex, pronotum. scutum, scutellum white; metasomal tergum
1 white with some black hairs; tarsi with white to reddish brown hairs; rest of body includ-
ing scopa black. Head wider than long, densely covered with punctures, all punctures of
approximaely similar diameter; compound eyes slightly convergent below; dorsal 1/3 of
gena visible beyond compound eye, gena wider than compound eye; clypeal margin
broadly concave, apical margin narrowly impunctate, clypeus in profile slightly convex,
densely punctate; hypostomal carina moderately raised, abruptly reduced before angle to
half its height; ocelli equally spaced between compound eyes and each other; length of
flagellomere 1 only slightly less than 2 and 3 together; mandible with 4 teeth (Fig. 1 ), apical
distance (between teeth) twice width of median constriction, carinae parallel, lower twice
as wide as upper, upper tooth oblique, next tooth smallest of the 4 with ventral margin trun-
cate, third tooth triangular, fourth tooth (lowest) longest; maxillary palpal segments 2 and 3
subequal, equalling length of 4 and 5 together; labial palpal segments 1 shorter than 2, 3
equal to4; ventral margin of galea with dense, short hairs. Thorax densely covered with fine
punctures, equalling those on head; propodeal triangle minutely rugose; propodeal pit
oval shining; hind tibial spurs straight, only apical 1/5 bent; strigilis with velum shallow
concave, amlus long, acutely pointed; fore wing with apical papillae small, cells covered

1 Received July 21, 1992. Accepted December 18, 1992.

2 Contribution No. 1656 from the Maine Agricultural Experiment Station.

3 Department of Biology and Program in Ecology. Evolution and Conservation Biology,
University of Nevada, Reno, NV 89557.

4 Department of Entomology, University of Maine, Orono. ME 04469-01 18.

ENT. NEWS 104(3): 113-117. May & June. 1993

14 ENTOMOLOGICAL NEWS

with few, short, light hairs, vein A 3 times B. Metasomal terga 1-4 with wide apical
impunctate bands, bands 1/4 to 1/3 punctate portion.

Male: The male of O. kenoyeri is extremely distinctive and can be distinguished from all
other Osmia by the structure of mid tarsus 1, the shape of the genitalia and metasomal ster-
num 2 (Sandhouse 1939).

Type: The holotype oWsmia kenoyeri, a male, is from Nebraska Hill, Colorado and was
collected on Trifolium. It is located at the University of Colorado Museum, Boulder,
Colorado. Males from Maine were identical with the holotype.

Distribution: Osmia kenoyeri is known from Deblois, Washington County, Maine;
Grand Sable Dunes, Alger County, Michigan (M. Arduser, pers. comm.); and Sandhouse
(1939) reported it from Colorado, California, Alberta, and the Yukon.

Remarks: In White (1952), the female of O. kenoyeri keys out with O.
Integra Cresson (couplet 18) but differs from O. Integra in the small size of
the fore wing papillae and the white pubescence on the dorsum of the
thorax. In Mitchell (1962), the female of O. kenoyeri also keys to O. Integra
(couplet 1 2) and the male keys with Osmia felti Cockerell (couplet 9) but is
easily separated by the globose first segment of the mid tarsus.

Sandhouse (1939) placed O. kenoyeri in the subgenus Acanthos-
mioides. White (1952) excluded O. kenoyeri from Acanthosmioides and
placed it in the subgenus Melanosmia (equals Centrosmia). His decision
was based primarily on character similarities with Osmia bucephala
Cresson. Hurd (1979) placed O. kenoyeri in the subgenus Acanthos-
mioides. He provided no explanation for the placement. However, with
the correct association of the female, O. kenoyeri appears to be more
closely associated with Acanthosmioides. In the female, the broad apical
width of the mandible, general body and pubescence coloration, and
apical margins of terga and in the male, the body coloration, mandible,
metasomal sterna 2 to 7, genitalia, and leg characters strongly suggest
Acanthosmioides.

Sandhouse ( 1939) suggested that Osmia hendersoni Cockerell may be
the female of O. kenoyeri. The type of O. hendersoni (U.S. National History
Museum #27891) was examined and found not to be the female of O.
kenoyeri. The shape of the mandible and hypostomal carina, and im-
punctate bands on abdominal terga 2 and 3 were different. The type of O.
hendersoni is extremely worn with badly torn wings and missing
flagellomeres.

Biology: A nesting site of O. kenoyeri was found on the "blueberry
barrens" in Deblois, Washington County, Maine (EAO). Females were
observed to excavate burrows in the sandy loam soil in late May and
adult activity continued to mid June. Burrows varied from 40 to 75 mm in
diameter and were not symmetrical in construction. They entered the
ground at about a 30 angle and terminated in a series of linear cells
within 2 to 3 cm of the soil surface. One nest contained 3 urn-shaped
cells, 2 with developing larvae while the third was open and not pro-

Vol. 104, No. 3, May & June, 1993

115

visioned. Cells were approximately 80 x 1 10 mm and composed of a mix-
ture of fine plant fibers, not leaf or leaf pieces, and soil. Cell walls varied
from 0.75 to 1 .5 mm thick. Cells were easily separated from the surround-
ing soil and remained intact when excavated. Larvae were mature by
early July and in cocoons. The cocoon was composed of two layers. The
outer layer was a dense mat of silk fibers that adhered to the soil-fiber cell
wall and the inner layer was thinner. A highly polished red brown matrix
with individual silk threads was visible and easily separated from the
outer layer. The outer anterior surface of the cocoon was formed by a
large, flat white nipple which separated the cocoon from the top of the
soil-fiber cell. Nineteen males and 9 females were collected the following
May from a 30 x 30 cm screened area of the nesting site. No parasites or
predators were recovered from the cages.

Examination of the pollen grains remaining in the fecal pellets
showed both the tetrad grain structure of Ericaceae (Vaccinium) and a
smaller, tricolporate grain, perhaps a Fabaceae.

Osmia (Chenosmia) virga Sandhouse

Female: Length 10 mm; fore wing 7.0 mm; head width 3.0 mm. Color olive-greenish
blue; flagellomeres, legs reddish brown, scape black, mandible red brown with black edges.
Pubescence white; mandibles, tarsi with golden to red brown hairs, scopa black. Head
slightly wider than long, punctures continuous and of similar size; gena with upper 1/3 visi-
ble beyond compound eye, twice as wide as compound eye; inner margin of compound eye
convergent below; lateral ocelli 2'/2 diameters distant from compound eye, less than 1
diameter from median ocellus; clypeal apical margin truncate, slightly wavy, narrowly
impunctate; hypostomal carina low, uniform, not toothed; mandible with 4 teeth (Figure
1 ), apical width only slightly greater than mid width, carinae divergent, lower twice as wide
as upper; flagellomere 1 longer than 2 or 3; maxillary palpal segment 3 longer than 2. twice 3

Figure 1. Female mandibles of Osmia kenoyeri Cockerel! (left) and Osmia virga Cockerell
(right), scale line is 1.0 mm.

16 ENTOMOLOGICAL NEWS

and 4 together, galea ventral surface with numerous short straight hairs, dorsal surface with
few scattered, straight hairs; labial palpal segment 2 longer than 1. Thorax densely and
evenly punctate, puncture size similar to vertex; propodeal triangle rugose above, granular
below; propodeal pit narrow, parallel sided; tegulae almost impunctate; wings minutely
and evenly haired, vein A twice B; strigilis with malus margin truncate, velum very short.
Abdomen with impunctate bands of metasomal terga 1 to 3Vi as wide as punctate portion, 4
to 6 narrower.

Male. The male of O. virga is easily identified by the wide impunctate bands on
metasomal terga 1 to 4, the truncate margin of metasomal sternum 4, and the structure of
the genitalia, especially the broad penis valves (Sandhouse 1939, Mitchell 1962).

Type. The type specimen is from Water Tank, Pennsylvania and is located in the United
States Natural History Museum. Washington, D.C. (Holorype #52883). Maine males were
identical to the holotype.

Distribution: We have seen specimens from Deblois and Orono, Maine. Sandhouse
(1939) reported O. virga from Massachusetts, New Jersey, Connecticut, and Virginia and
Mitchell (1962) added Wisconsin and Pennsylvania.

Remarks: In Mitchell (1962), the female of O. virga keys out as
O. atriventris Cresson (couplet 17) from which it differs in body color,
hypostomal carina, mandibular carinae, impunctate bands of meta-
somal terga 4 and 5, structure of the strigilis, and scopal color.

Stubbs et al. (1992) reported 99% Vaccinium pollen from the pollen
load of O. virga from Maine.

DISCUSSION

The distributional pattern seen in O. kenoyeri. Rocky Mountains and
northeastern U.S. and Canada, is not uncommon within the genus.
Osmia Integra, O. bucephala, O. subaustralis Cresson, O. inermis (Zet-
terstedt), and O. nigriventris (Zetterstedt) show similar distributions with
the northeastern and Rocky Mountain distributions connected through
northern Canada and Alaska (Sandhouse 1939, Mitchell 1962, Rust
1974). The latter two species are Holarctic. Biologically, O. kenoyeri
groups with two other Acanthosmioides, O. nigrobarbata Cockerell and
O. unca Michener, in excavating burrows in the ground where the cells
are lined with plant materials and soil (Rust et al. 1974). Also, the two
layered construction of the cocoon and the large, flat white nipple area
are characteristics observed in other Acanthosmioides species.

ACKNOWLEDGMENTS

We would like to thank D. Bowers (University of Colorado, Boulder) and R. J.
McGinley (U.S. National Museum) for the loan of type specimens, M. Arduser(St. Louis,
MO) for first suggesting that the Maine material might be O. kenoyeri and for the Michigan
distributional records, T. Griswold (U.S. DA Bee Biology and Systematics Laboratory,
Logan, UT) and F. A. Drummond (University of Maine, Orono) for reviewing the
manuscript.

Vol. 104. No. 3. May & June, 1993 117

LITERATURE CITED

Boulanger, L.W., G.W. Wood, E.A. Osgood, and C.O. Dirks. 1967. Native bees

associated with the low-hush blueberry in Maine and eastern Canada. Maine Agr. Exp.

Stat. Bull. T26: 22pp.
Hurd, P.D. Jr. 1979. Apoidea. In: Catalog of Hymenoptera in America north of Mexico.

eds: K.V. Krombein, P. D. Hurd Jr.. D.R. Smith, and B.D. Banks. Smithsonian Press.

Washington, D.C.
Mitchell, T.B. 1962. Bees of the eastern United States. Vol. 2. North Carolina Agr. Exp.

Stat. Tech. Bull 152: 557pp.
Rust, R.W. 1974. The systematics and biology of the genus Osmia. subgenera Osmia.

Chalcosmia, and Cephalosmia. Wasmann J. biol. 32: 1-93.
Rust, R.W., R.W. Thorp, and P.F. Torchio. 1974. The ecology of Osmia nigrifrons with a

comparison to other Acanthosmioides. J. Nat. Hist. 8: 29-47.
Sandhouse, G.A. 1939. The North American bees of the genus Osmia. Mem. Entomol.Soc.

Washington 1: 1-167.
Stubbs, C.S., H.A. Jacobson, E.A. Osgood, and F.A. Drummond. 1992. Alternative

forage plants for native (wild) bees associated with lowbush blueberry. Vaccinum

Acanthosmioides spp.. in Maine. Maine Agr. Exp. Stat. Bull. T148:54pp.
White, J.R. 1952. A revision of the genus Osmia. submenus Acanthosmioides ( Hymenoptera:

Megachilidae). Univ. Kansas Sci. Bull. 35: 219-307.

BOOKS RECEIVED AND BRIEFLY NOTED

CATALOGUE OF THE CYPHOPALPATORES AND BIBLIOG-
RAPHY OF THE HARVESTMEN ( ARACHNIDA, OPILIONES) OF
GREENLAND, CANADA, U.S.A., AND MEXICO. J.C. Cokendolpher
& V.F. Lee. 1993. Distributor: The Wishing Well, 1200 Clover Drive,
Burkburnett, TX 76354. 82 pp. paperback. $9.50 incl. shipping.

The goal of this publication is twofold: a catalogue and geographical checklist are pro-
vided to the 225 species and 50 genera of Cyphopalpatores and second, a complete litera-
ture survey, including newspapers, theses, dissertations, and government reports, is
provided for both fossil and recent species. All topics are covered, including folklore.

NYMPHS OF NORTH AMERICAN STONEFLY GENERA (PLE-
COPTERA). K.W. Stewart and B.P. Stark. 1993. Univ. of North Texas
Press. 464 pp. 244 illus. paperback. $34.50.

This book is a reprint of the 1988 edition published by the Entomological Society of
America as Volume XII of the Thomas Say Foundation. It is a baseline reference for serious
study of North American Plecoptera and for stream ecological studies.

118 ENTOMOLOGICAL NEWS

TABANIDAE AS DIETARY ITEMS

OF RAFINESQUE'S BIG-EARED BAT:

IMPLICATIONS FOR ITS FORAGING BEHAVIOR 1

Susan E. Ellis 2 * 3

ABSTRACT: Lepidopterans and dipterans formed the entire diet of a maternity aggrega-
tion of Rafinesque's big-eared bat (Plecotus rafinesquii) in a single-sample analysis of fecal
pellets. Tabanidae made up 31% of the total sample volume. The presence of male
Tabanidae as prey items may be significant, because few natural predators have been iden-
tified for this insect. Literature concerning the behavior of tabanids is reviewed and com-
pared to the sparse information concerning P. rafinesquii foraging. The foraging habits of
this bat species are apparently more varied than previously reported. The unusual pre-
sence of tabanids as prey items suggests that this bat could be of use in tabanid
control.

Knowledge of the life histories of insects in the diets of insectivorous
vertebrates may give insight into the foraging habits of these predators.
From an economic perspective, identifying the prey of insectivorous bats
could have important implications for biological control. Predation on
flying insects, usually reproductively competent adults, can have a much
greater impact on insect populations than predation on immature stages
(Buckner, 1967).

Rafinesque's big-eared bat, Plecotus rafinesquii Lesson has been de-
scribed as "the least known" bat species of the eastern United States
(Harvey, 1992; Barbour and Davis, 1969). Information about the natural
history of this insectivorous bat is fragmentary. Most studies have
focused on its distribution and roosting ecology (e.g., Jones, 1977;
Whitaker and Winter, 1977; Hall, 1963), and there is but one study of its
feeding ecology (Clark, 1991). Harvey (1992) reports that "almost noth-
ing is known about their feeding behavior." Information about this bat's
diet comes from an examination of the stomach contents and fecal
material collected from a single animal which had eaten moths
(Hamilton, 1933).

The purpose of this study is to report on items included in the diet of a
North Carolina aggregation of P. rafinesquii as determined by fecal
analysis, and discuss the foraging habits of the bats in relation to the
habits of their insect prey.

1 Received December 12, 1992. Accepted February 25, 1993.

Delaware Agricultural Experiment Station, Department of Entomology and Applied
Ecology, College of Agricultural Sciences, University of Delaware, Newark, DE 19717.

3 Current address: Department of Biology, York University, 4700 Keele Street, North York,
Ontario M3J 1P3 Canada.

ENT. NEWS 104(3): 1 18-122. May & June, 1993

Vol. 104, No. 3, May & June, 1993 1 19

MATERIALS AND METHODS

A sample of sixty fecal pellets was collected from a maternity roost of P.
rafmesquii from Chowan County, North Carolina in late July, 1992. This
material was analyzed by: 1) percentage frequency and 2) percentage
volume of insect taxa in the pellets, as described by Whitaker (1988).
After the fecal material was softened in 70% isopropyl alcohol for at least
two hours, individual fecal pellets were teased apart under a binocular
microscope at 45X, and the volume of each taxa in each pellet was
visually estimated. Fragments were identified to order and family using
standard insect identification keys (Borror, et al, 1989; Arnett, 1985;
Borror and White, 1970) and by comparing structures of wet mounted
fragments with those of pinned specimens in the museum collection at
the University of Delaware.

RESULTS

Lepidoptera occurred in 78.3% of the samples and composed 67% of
the sample volume. Three types of lepidopteran eggs were present in the
samples, occurring in 27.6% of the pellets containing lepidopterous
fragments. Dipterans occurred in 60% of the samples, composed 33% of
the sample volume, and included TabanicJae (horse flies, or Tabanus
spp.) and Tipulidae (crane flies). Tabanids constituted 31% of the total
sample volume and occurred in 46.6% of the pellets. The balance of the
dipterans consisted of tipulids (less than 2% volume, 8% frequency) and
other fragments, unassignable to family. Three of the tabanids were
identified as males: one by the aedeagus; two by the holoptic head cap-
sule characteristic of males. No tabanid eggs were found in the pellets,
and no recognizable female structures were found among the tabanid
fragments.

DISCUSSION

The North Carolina maternity aggregation of P. rafmesquii preys
heavily on lepidopterans and dipterans in at least part of its active
season. Extrapolating prey size from the fragments of legs in the pellets
and by comparison with museum specimens, I estimate that these prey
were all within a 20-25 mm range in body length.

The presence of male tabanids provides clues about the foraging
habits of the bats. Tabanid flies are considered economically-important
pests because the females are obligate blood-feeders, capable of both
inflicting a "vicious bite" and of transmitting pathogens (Pedigo, 1989).
In at least 34 Tabanus spp., males hover in groups as a prelude to mating

120 ENTOMOLOGICAL NEWS

(Gaugler and Schutz, 1989). These authors described the mating be-
havior of tabanid flies as a "potential weak link" in their life cycles and
propose that disrupting the equilibrium of tabanid populations might be
achieved through controls exerted on hovering males. Corbet and Had-
dow (1962) suggest that hovering swarms of these flies should be vulner-
able to predation, however they cite only one report (Blickle, 1959), in
which an adult Bembex belfragei Cresson (Hymenoptera: Sphecidae)
captured a hovering male Tabanus bishoppi Stone in Florida. Corbet and
Haddow (1962) postulate that swarming by males at low light levels may
reduce their detectability by diurnal predators. Plecotus rafinesquii ap-
pears to be one of the few predators exploiting this weak link, and may
exert some measure of population control on these flies.

Males of Tabanus species usually gather at dawn and dusk around a
landmark, where they hover in a fairly stationary group "a few feet above
ground level" (Gaugler and Schutz, 1989; Bailey, 1948). Landmarks vary,
but share the common feature of being relatively open and free of ob-
struction. Some reported landmarks are: forest-meadow margins, roads
(Gaugler and Schutz, 1989), shrubs, and a driveway (Bailey, 1948). Taba-
nids are strong omni-directional fliers capable of holding a defined ter-
ritorial position over a landmark and within a swarm (Downes, 1969).

The timing of swarming probably depends on temperature, light or
other environmental cues (Gaugler and Schutz, 1989; Bailey, 1948),
however a response to low light intensity appears to be the primary
initiator of swarming by tabanids (Corbet and Haddow, 1962). Low light
intensity may act as a species-specific cue, coordinating the activity
times of males and females (Gaugler and Schutz, 1989; Corbet and Had-
dow, 1962). The onset of hovering is closely synchronized and may con-
tinue for 10-23 minutes; cessation is more variable, probably because
factors such as fatigue and thoracic heating affect individual males dif-
ferently (Corbet and Haddow, 1962). However, thresholds of light inten-
sity probably also provide the cue for cessation of hovering (Corbet and
Haddow, 1962). All studies report a diurnal to crepuscular time window
for hovering by these flies (Schutz and Gaugler, 1992; Gaugler and
Schutz, 1989; Downes, 1969; Corbet and Haddow, 1962; Bailey, 1948).

Two conclusions may be drawn from this. First, foraging/ 1 , rafinesquii
shares space with these hovering flies. This indicates that P. rafinesquii
forages near ground level and bears out observations made by Clark
( 1 99 1 ) of light-tagged P. rafinesquii flying "about 1 m above ground level".
Barbour and Davis (1969) observed P. rafinesquii confined in a small
room and reported that it was capable of hovering, and of swift and
maneuverable flight. The presences of male tabanids in the diet of P.
rafinesquii indicates that these bats forage to some extent in open areas
containing tabanid landmarks. However, the flying agility ascribed to

Vol. 104, No. 3. May & June, 1993 121

them would allow them to navigate within the clutter of forested areas
where they have been observed foraging (Clark, 1991).

Second, P. rafinesquii shares some activity time with its prey, indicat-
ing that these bats forage in early morning or early evening, an observa-
tion that contrasts with reports that the species does not forage at twilight
(Harvey, 1991; Jones, 1977; Barbour and Davis, 1969).

Although Tipulidae accounted for less than 2% of the volume and
occurred in only 8% of the samples, these flies form "discrete assembly
stations" for mating, comparable in form if not behavioral complexity, to
those otTabanus species (Downes, 1969). An additional inference con-
cerning P. rafinesquii foraging habits is that it may sometimes exploit
clumped resources. Since I was unable to identify lepidopterans below
the ordinal level, I cannot cite specific behaviors of these insects that
might further support this inference. It is possible, however, that in-
dividuals of P. rafinesquii prey on swarms of hovering flies while enroute
to forested areas, then switch to moths after entering the forest. In the
absence of site-specific data this speculation seems reasonable, because
P. rafinesquii has often been associated with forested areas (Clark, 1991;
Whitaker and Winter, 1977; Barbour and Davis, 1969).

SUMMARY

The limited data presented in this single-sample prey analysis from
one maternity roost of P. rafinesquii provides evidence that these bats preyed
on 20-25 mm lepidopterans (67% volume) and dipterans (33% volume),
including tabanids (31% volume) and tipulids (less than 2% volume).
Male Tabanus spp. exhibit diurnal to crepuscular group hovering be-
havior in open areas, indicating that the spatial and temporal foraging
repertoire of P. rafinesquii may be more varied than previously
assumed.

Plecotus rafinesquii exploits what some tabanid researchers (i.e.,
Gaugler and Schutz, 1989) have identified as a potential weak link in the
life cycles of Tabanus species. Other tabanid researchers (i.e., Corbet and
Haddow, 1962), have indicated that this weak link is apparently rarely
exploited by other predators. Thus, P. rafinesquii may have some impact
on populations of these economically important insects by preying on
hovering males. It seems likely, from the percentage of lepidopteran eggs
occurring in the sample (27.6%), that P. rafmesauii predation also has an
impact on moth populations. Specifically, by limiting oviposition by
moths, P. rafinesquii affects proliferation of the destructive larval stages
of these insects.

Insectivorous bats extend their impact beyond individual prey num-
bers by removing reproductive adult insects, thus limiting mating and

122 ENTOMOLOGICAL NEWS

oviposition. These are effective methods of regulating insect populations
(Pedigo, 1989; Buckner, 1967). Further investigation of the foraging
habits off. rafinesquii and other insectivorous bats could yield important
information about the biological control potential of these predators.

ACKNOWLEDGMENTS

P. Trail provided me with the fecal material from a P. rafinesquii maternity roost he has
studied for several years. M. Clark generously shared observations from her field studies of
P. rafinesquii in North Carolina. R. Weber, R. Raimist, M. Fenton and two anonymous
reviewers made helpful suggestions on the manuscript. R. Weber provided invaluable
guidance and insight throughout.

LITERATURE CITED

Arnett, R.H., Jr. 1985. American insects. Van Nostrand Reinhold Co., New York. 640-
708 pp.

Bailey, N.S. 1948. The hovering and mating of Tabanidae: a review of the literature with
some original observations. Ann. Entomol. Soc. Am. 41:403-412.

Barbour, R.W. and W.H. Davis. 1969. Bats of America. Univ. of Kentucky Press, Lex-
ington, pp. 179-182.

Borror, DJ., C.A. Triplehorn and N.F. Johnson. 1989. Introduction to the study of
insects, 6th ed. Saunders College Publishing, Philadelphia. 499-572 pp.

Borror, D.J., Jr. and R.E. White. 1970. A field guide to the insects of America north of
Mexico. Houghton Mifflin, Boston. 262-275 pp.

Buckner, C.H. 1967. Avian and mammalian predators of forest insects. Entomophaga
12:491-501.

Clark, M.K. 1991. Foraging ecology of Rafinesque's big-eared bat, Plecotus rafinesquii, in
North Carolina (abstract). Bat Research News 33:68.

Corbet, P.S. and AJ. Haddow. 1962. Diptera swarming high above the forest canopy in
Uganda, with special reference to Tabanidae. Trans. R. Ent. Soc. Lond. 1 14:267-284.

Downes, J.A. 1969. The swarming and mating flight of Diptera. Ann. Rev. Entomol.
14:271-298.

Gaugler, R. and S. Schutz. 1989. Environmental influences on hovering behavior of
Tabanus nigrovittatus and T. conterminus (Diptera: Tabanidae). J. Insect Behavior
2:775-786.

Hall, J.S. 1963. Notes on Plecotus rafinesquii in central Kentucky. J. Mammal. 44:119-
120.

Hamilton, WJ., Jr. 1933. The insect food of the big brown bat. J. Mammal. 14:155-156.

Harvey, M J. 1992. Bats of the eastern United States. Arkansas Game and Fish Commis-
sion. 36-37 pp.

Jones, C. 1977. Plecotus rafinesquii. Mamm. Species 69:1-4.

Pedigo, L.P. 1989. Entomology and pest management. MacMillan Publishing Co., New
York. pp. 127-128.

Schutz, S. and R. Gaugler. 1992. Thermoregulation and hovering behavior of salt marsh
horse flies (Diptera: Tabanidae). Ann. Entomol. Soc. Am. 85:431-436.

Whitaker, J.O., Jr. 1988. Food habits analysis of insectivorous bats. In Ecological and
behavioral methods for the study of bats, T.H. Kunz,ed. Smithsonian Ins. Press, Wash-
ington, pp. 171-179.

Whitaker, J.O., Jr. and F.A. Winter. 1977. Bats of the caves and mines of the Shawnee
National Forest, Southern Illinois, USA. Trans. 111. State Acad. Sci. 70:301-313.

Vol. 104, No. 3. May & June, 1993 123

A HUGE NEST OF THE COMMON

YELLOWJACKET, PARAVESPULA VULGARIS

(HYMENOPTERA: VESPIDAE), IN CALIFORNIA 1

Roger D. Akre, Elizabeth A. Myhre, Yi Chen 2

ABSTRACT: A large, perennial colony of Paravespula vulgaris was collected in California
in July 1992. The envelope was ca 97 diam and had multiple entrances. The colony con-
sisted of 220 nonfunctional queens, 2 developing queens, 10 functional queens, ca 84,000
workers, and ca 32,000 males. The 22 combs consisted of ca 220,000 worker and 9,000 re-
productive cells; 64% were empty. The colony was estimated to be 2-3 years old.

Yellowjackets typically have a one year life cycle, and the only mem-
bers of the colony to overwinter are inseminated queens. They start new
colonies in the spring. However, some colonies continue for more than
one season (Akre and Reed 1981) and these colonies are always poly-
gynous (Carpenter 1989). Large, perennial colonies have previously
been reported for Paravespula germanica (F.) in New Zealand (Thomas
1960, Plunkett et al. 1989, Clapperton et al. 1989), Tasmania (Spradbery
1973a), Algeria and Morocco (Vuillaume<?f al. 1969), and Chile (Jeanne
1980, Chiappa et al. 1987). Perennial colonies are unusually large, and
Spradbery (1973a) estimated one nest weighed 1,000 pounds. Vespula
squamosa (Drury), the southern yellowjacket, has perennial colonies in
the southern United States, especially in Florida (Tissot and Robinson
1954, Akre etal. 1981, Ross and Matthews 1982) and in Texas. Perennial
colonies of the common wasp occur in New Zealand (Plunketter al. 1989)
and a perennial colony of P. vulgaris (L.) was recorded in California years
ago (Duncan 1939). Unconfirmed reports indicate many more perennial
colonies are being found in southern California where winter tem-
peratures are mild, and prey populations are high enough to sustain a
colony through the winter months. One P. vulgaris colony persisted from
its discovery in 1984 until at least 1986 in Berkeley, CA (Alameda Co.)
(Gambino 1986). A two year colony of P. pensylvanica (Saussure) was
reported in British Columbia (Spencer 1960), and Nakahara (1980)
recorded a huge perennial colony of P. pensylvanica (Saussure) in Hawaii
(see also Gambino et al. 1990). The purpose of this paper is to present a
detailed analysis of a nest off*, vulgaris collected in Burlingame (San
Mateo Co.), California.

1 Received November 21, 1992. Accepted January 2, 1993.

2 Department of Entomology, Washington State University. Pullman, WA 99164-6382.

ENT. NEWS 104(3): 123-128. May & June, 1993

124

ENTOMOLOGICAL NEWS

MATERIALS AND METHODS

A large nest of Paravespula vulgaris was discovered in Burlingame,
California, and a local pest control firm was called to kill the colony (Fig.
1). The nest was built into a dense mass of vines hanging 1.2 to 1.5 m

Fig. 1 . The nest was suspended in a large mass of vines. Although the entrance holes are not
readily apparent from this angle, several were at least 4 cm diam (R. Schoeppner photo-
graph).

Vol. 104, No. 3, May & June, 1993 125

above a creek bank. The nest was ca 62 cm wide X 76 cm tall X 35 cm
thick, with a circumference of 244 cm, a diameter of 97 cm (measure-
ments were estimated from a photograph), somewhat heart-shaped, and
had multiple entrances (ca 120 on each side). The colony was killed the
week of 1 2 July 1992, and all nest material and occupants were shipped to
Washington State University at Pullman for study. The nest, as collected,
complete with severed vines, weighed ca 64 kg. The nest was stored in a
freezer at 5.0 F until the occupants and combs were counted and
analyzed. Males were individually counted first, and it was determined
that 1,385 fit into a 250 ml graduate cylinder when it was vigor-
ously tapped on the substrate to settle and compact them. The count was
stopped when they no longer settled. Similarly, 2,01 1 workers were
needed to fill a 250 ml cylinder. All remaining individuals, except
queens, were measured volumetrically to estimate number of workers
and males in the colony. Queens were individually tallied, injected with
Kahle's to preserve them, and placed into 70% ETOH until they were dis-
sected for ovarian development.

All combs of the nest were analyzed for the presence of pupal caps,
larvae, and eggs. A 12 in grid divided into 1 in squares facilitated the
count (MacDonald et al 1974). In some sections the cells were indi-
vidually counted. The nest was sectioned in California to fit into the
shipping containers, and we tried to reconstruct comb placement before
the analysis began. Sections were numbered and flagged as they were
unpacked to aid us in this endeavor.

RESULTS

The colony consisted of ca 84,000 workers, 32,000 males, and 232
queens (Table 1). The total number of cells in the 22 comb layers of the

Table 1. Numbers and types of adults collected with the colony. Functional, developing,
and nonfunctional queens (parens) add up to "Queens."

Adult Type

Number Counted

Queens

232

Functional

(10)

Developing

(2)

Nonfunctional

(220)

Males

31,811

Workers

83784

Total Adults

115.827

126

ENTOMOLOGICAL NEWS

nest was ca 230,000, but 146,000 of these cells were empty (Table 2). Dis-
sections of the queens revealed that 10 had fully functional ovaries that
filled the gaster (eggs are 1 .0- 1 .5 mm when ready to lay). Two had a slight
development of the ovarioles but no eggs were >1 mm. All functional
queens had multiple age spots or dark discolorations on the gaster (Ross
1984). The remaining 220 queens showed no ovarian development.

DISCUSSION

Although large, this colony was probably entering a period of decline
as evidenced by the great number of empty cells. Ten functional queens
should have been able to lay eggs in many of these empty cells unless
there was conflict and fighting among workers and/or queens in the nest.
We could not determine any spheres of influence for any of the queens
because of the state of the nest when received.

The large number of males in the colony suggests that workers were
laying eggs and producing at least some or perhaps most of these
males.

Questions asked of neighbors suggested that this nest was observed
for 30 years. We have no evidence to support this claim and indeed, the
nest analysis, including the construction of reproductive cells on the
periphery of some combs, suggested that the nest was probably 2 years
old, at most.

Photoperiod is probably the stimulus that allows new queens to cir-
cumvent reproductive diapause and to start egg development after they
are inseminated (Spradbery 1973b, Ross and Matthews 1982). Queens
frequently rejoin the parent colony if the photoperiod is still increasing

Table 2. Occupants of cells from the 22 combs. Small, medium, and large larval counts
(parens) add up to "Larvae."

Worker

Reproductive

Total

Eggs . .

^884

82

~>966

Larvae

26918

417

~>7 17S

Small

(4 089)

(91)

(4 182)

Medium

(11 9">M

(170)

(P09S)

Large. .

(109">4)

(174)

(11 098)

Pupae .

SI SS3

97-7

V S">S

Empty cells

1189S4

7779

146711

Total

-no 1~>9

9">70

->->9 S99

Vol. 104, No. 3, May & June, 1993 127

(up to 21 June) in that geographical area, and if the winter temperatures
are mild. They can also rejoin the colony as nondiapausing, functional
queens when the daylight is very short as the "window" of receptiveness
seems to be 10-14 hrs (Spradbery 1973b). Photoperiods longer or shorter
do not induce reproductive diapause, and the ovaries in these new
queens are able to develop.

ACKNOWLEDGMENTS

We are indebted to George Bristol, Pescadero, and Marc Maas, Burlingame, for sending
us the colony. We are especially thankful to Robert Schoeppner, San Mateo Mosquito
Abatement District, Burlingame, for data on nest location and for many photographs of
the nest, including one that appears in this paper, that aided us in our analysis. R. Zack and
R. Schoeppner reviewed the article.

LITERATURE CITED

Akre R. D., A. Greene, J. F. MacDonald, P. J. Landolt, and H. G. Davis. 1981. Yellow-
jackets of America North of Mexico. USDA Agric. Handbook 552. 102 p.

Akre, R. D., and H. C. Reed. 1981. A polygynous colony of Vespula pensylvanica
(Saussure) (Hymenoptera: Vespidae). Ent News 92: 27-31.

Carpenter, J. M. 1989. Testing scenarios: wasp social behavior. Cladistics 5: 131-144.

Clapperton, B. K., P. A. Alspach, H. Moller, and A. G. Matheson. 1989. Impact of com-
mon and German wasps (Hymenoptera: Vespidae) on the New Zealand beekeeping
industry. N. Z. J. Zool. 16: 325-332.

Chiappa, T. E., R. H. Jopia, L. C. Morales, and L. J. Cook. 1987. Overwintering nests of
Vespula germanica (F.) (Hymenoptera: Vespidae) in Central Chile. Acta Ent. Chilena 14:
171-182.

Duncan, C. D. 1939. A contribution to the biology of North American vespine wasps.
Stanford Univ. Publ. Biol. Sci. 8: 1-271.

Gambino, P. 1986. Winter prey collection at a perennial colony of Paravespula vulgaris (L.)
(Hymenoptera: Vespidae). Psyche 93: 331-340.

Gambino, P., A. C. Medeiros, and L. L. Loope. 1990. Invasion and colonization of upper
elevations on east Maui (Hawaii) by Vespula pensylvanica (Hymenoptera: Vespidae).
Ann. Entomol. Soc. Am. 83: 1088-1095.

Jeanne, R. L. 1980. Evolution of social behavior in the Vespidae. Ann. Rev. Ent. 23: 371-
396.

MacDonald, J. F., R. D. Akre and W. B. Hill. 1 974. Comparative biology and behavior of
Vespula atropilosa and V. pensylvanica (Hymenoptera: Vespidae). Melanderia 18: 1-66.

Nakahara, L. M. 1980. Western yellowjacket (Vespula pensvlvanica) first record of aerial
nest in state. Coop. PI. Pest Rept. USDA, APHIS. 5(14): 270.

Plunkett, G. M., H. Moller, C. Hamilton, B. K. Clapperton, and C. D. Thomas. 1989
Overwhelming colonies of German (Vespula germanica) and common wasps (Vespula
vulgaris) (Hymenoptera: Vespidae) in New Zealand. N. Z. J. Zool. 16: 345-353.

Ross, K. G. 1984. Cuticular pigment changes in worker yellowjackets (Hymenoptera: Ves-
pidae). J. N. Y. Ent. Soc. 91: 394-404.

Ross, K. G. and R. W. Matthews. 1982. Two polygynous overwintered Vespula squamosa
colonies from the southeastern U. S. (Hymenoptera: Vespidae). Fla. Ent. 65: 176-184.

Spencer, G. J. 1960. On the nests and populations of some vespid wasps. Proc. Ent. Sex;.
British Columbia 57: 13-15.

128 ENTOMOLOGICAL NEWS

Spradbery, J. P. 1973a. Wasps: an account of the biology and natural history of solitary

and social wasps. Univ. Washington Press, Seattle. 408 p.
Spradbery, J. P. 1973b. The European social wasp, Paravespula germanica (F.) (Hymenop-

tera: Vespidae) in Tasmania, Australia. IUSSI Proc. VII Internal. Congr. pp. 375-380.
Thomas, C. R. 1960. The European wasp (Vespula germanica Fab.) in New Zealand, inf.

Ser. Dept. Sci. Ind. Res. New Zealand. 27: 1-74.
Tissot, A. N., and F. A. Robinson. 1954. Some unusual insect nests. Florida Ent. 37:

73-92.
Vuillaume, M., J. Schwander, and C. Roland. 1969. Note preliminaire sur 1'existence de

colonies perennes et polygynes de Paravespula germanica C. R. Acad. Sci. Ser. D. 269:

2371-2372.

BOOKS RECEIVED AND BRIEFLY NOTED

BIOLOGY AND CONSERVATION OF THE MONARCH BUTTER-
FLY. S.B. Malcolm and M.P. Zalucki, eds. 1993. No. 38 Science Series,
Natural History Museum of Los Angeles County. 219 pp. 139 figs. 100
tables. Cloth. $90.00.

This volume presents an interdisciplinary approach to understanding how Danaus
plexippus functions, providing information for an objective approach to conservation of
the species. Forty-four papers written by 50 authors present a diverse array of research on
biological and conservation topics. These papers embrace four main themes of monarch
biology: communication and mating, host plant exploitation and chemical defense,
migration, and overwintering.

THE SCIENCE OF ENTOMOLOGY. 3rd ed. W.S. Romoser and J.G.
Stoffolano. 1994. Wm. C. Brown, Publ. 532 pp. Cloth. $60.90.

The authors' stated objective for this third edition is to provide a broad, balanced
introductory text to the science of entomology, from both basic and applied points of view,
for use in a one-quarter or one semester general course.

Vol. 104. No. 3, May & June. 1993 129

SEASONAL FLIGHT ACTIVITY OF LIPOPTENA

MAZAMAE (DIPTERA: HIPPOBOSCIDAE)

IN SOUTH CAROLINA 1

L. Daniel Cline, James E. Throne 2

ABSTRACT: Flying winged adults (volants )ol'Lipoptenama:anwe were collected in sticky
traps at two of three grain storage sites in southeastern South Carolina that were being sam-
pled for seasonal occurrence of stored-product insects. Of the 42 volants captured during
the 55-week test, at least one was caught in every month from April through November.
None were caught from December through March. Most were caught in traps placed near
wooded areas or heavy shade where deer were likely to travel or feed. Sticky traps provide
an alternative to conventional trapping methods for hippoboscids. but have the disadvan-
tage of not providing positive host information.

Lipoptena mazamae Rondani is a parasite of all species and sub-
species ofdeer(Odocoileus spp.) and brocket (Mazama spp.) wherever the
hosts occur in the Neotropical region (Bequaert 1957). It also has been
found infrequently and accidentally on domestic cattle. Lipoptena
mazamae has been found as far south as Argentina and as far north as the
states bordering the Gulf of Mexico and up the Atlantic coast into South
Carolina (Bequaert 1957). Little is known of its life history and behavior.
Because it is essentially a tropical insect, its populations in the United
States may fluctuate with the severity of the winter.

Current techniques for the detection and collection of Hippobos-
cidae can be difficult (Pfadt & Roberts 1978). On live domestic animals,
an uncooperative attitude coupled with low numbers of parasites may
make them hard to find and the quickness of their movements may make
them difficult to catch. Placing newly killed hosts in a bag or screened
enclosure may facilitate collections. Collections on large wild animals
nearly always requires killing the host and searching a standardized
area of its body where the parasites are most likely to occur (Samuel &
Trainer 1972). Volants (winged adults) can be collected with sweep nets
or by collecting specimens from skin and clothing as the collector walks
through a selected area. Both of thestj methods require a great deal of
time and care (Hare 1945).

During a study of the flight activity of stored-product insects around
grain bins, we captured L. mazamae on sticky traps. Given the scarcity of
information on their biology and the complete lack of information on

1 Received December 7, 1992. Accepted December 28, 1992.

2 USDA, ARS Stored-Product Insects Res. & Dev. Lab.. P. O. Box 22909, Savannah,
Georgia 31403.

ENT. NEWS 104(3): 129-132. May & June. 1993

130 ENTOMOLOGICAL NEWS

volants' seasonal occurrence, we report our findings of this insect at the
northern end of its range. In addition, we present a collection method
that passively catches the parasites with no trauma to their hosts and
minimal effort by the collector.

MATERIALS AND METHODS

Sticky traps were used to monitor three sites in southeastern South
Carolina for flying insects from 18 March 1987 to 6 April 1988 (55 con-
tinuous weeks). Traps consisted of clear flat plexiglass (30.5 cm by 30.5
cm) coated on each side with sticky substance (Tangle-Trap, Tangle-
foot Co., Grand Rapids, Michigan) 3 and held in a vertical position on a
wooden stake. Details of trap design and preparation are given in
Throne and Cline (1989).

The traps were deployed in two nearly concentric rings around grain
storage bins. The inner traps were generally about 0.5 m from the bins
and the outer traps varied from 5 to 45 m from the bins to accommodate
the cooperator's need to use the space. Four inner and four outer traps
were deployed at two sites (Bamberg and Hampton Counties) while five
inner and five outer traps encircled the larger third site (Barnwell
County). A map detailing the placement of the traps and the surround-
ing area is given in Throne and Cline (1989). In general, the Bamberg and
Hampton County sites were surrounded by cultivated fields with
wooded areas within 25 m of the grain bins. The Barnwell County site
was surrounded by pasture.

Each trap was exposed for one week after which it was replaced and
the exposed surfaces returned to the laboratory for examination. The
entire sticky surface (both sides) was examined at a magnification of at
least 10X. Hippoboscids were removed, recorded, and stored in vials of
alcohol. Identification was determined from characters detailed in
Peterson & Maa (1970). Voucher specimens were placed in the U.S.
National Museum Collection and in the Florida State Collection of
Arthropods.

RESULTS AND DISCUSSION

A total of 42 hippoboscids were collected and all were identified as L.
mazamae. Most (78.6%) were collected at the Hampton Co. site while
21.4% were collected at the Bamberg Co. site. None were caught at the
Barnwell Co. site. At least one volant was caught in every month from

3 Names of products are included for the benefit of the reader and do not imply endorse-
ment or preferential treatment by USDA.

Vol. 104, No. 3, May & June, 1993

131

April through November (Figure 1). None were caught in the four months
from December through March. This is similar to the finding of Hare
(1945) who found flying adults of a closely related species, L. depressa
(Say), from late March to early December in California with a peak in
July. The deer hosts are present in the area year-round, however, it is
unclear whether volants cease to emerge or cease to fly during cold
weather.

At the two sites where L. mazamae were caught, most were found in
the outer traps placed to the south of the grain storage area (44.4% at site 1
and 66.7% at site 2). At both sites, these were the areas either close to
woods or near heavy shade. At the Bamberg County site, the south outer
trap was placed where deer frequently traveled from a heavily wooded
area into a field where either corn or soybeans are usually grown. At the
Hampton County site, the south outer trap was placed at the edge of a
neglected grove of mature pecan trees where deer presumably come to
search for food. The third site was not a very suitable habitat for deer,
therefore, it is not surprising that no hippoboscids were caught there.
Hare (1945) found that most L. depressa (66%) were collected in or near

C7>

13
D
O

k_
(U

4
3
2
1

Bamberg Co.
(n = 9)

Hampton Co.
(n = 33)

May 20 Jul 29 Oct 7 Dec 16 Feb 25

Figure 1. Number ofLipoptena mazamae Rondani caught per week on sticky traps at two
sites in South Carolina, 1987-88.

132 ENTOMOLOGICAL NEWS

the tree shadows bordering wooded areas. A concentrated effort to place
traps in areas where hosts are likely to feed and rest would most likely
increase the number of volants that are caught. Although sticky traps
provide a relatively easy method for collecting hippoboscids, a disad-
vantage of the method is that they do not provide positive host
identification.

ACKNOWLEDGMENTS

We thank Mark Culik and Pat Lang for technical assistance; Messrs. Bates, Peeples,
and Rentz for allowing us to conduct this study on their farms; and W. W. Wirth and R. V.
Peterson (Cooperating Scientist and Scientist, respectively, USDA-ARS, Systematic Ento-
mology Laboratory) for confirming the identifications and reviewing the manuscript.

LITERATURE CITED

Bequaert, J.C. 1957. The Hippoboscidae or louse-flies (Diptera) of mammals and birds.

Part II. Taxonomy, evolution and revision of American genera and species. Entomol.

Amer. 36:4 17-611.
Hare, J.E. 1945. Flying stages of the deer lousefly, Lipoptena depressa (Say), in California

(Diptera, Hippoboscidae). Pan-Pac. Entomol. 21:48-57.
Peterson, B.V. and Maa, T.C. 1970. A new Lipoptena from Chile, with a key to the new

world species (Diptera: Hippoboscidae). Can. Entomol. 102:1 1 17-1 122.
Pfadt, R.E. and Roberts, I.H. 1978. X. Louse flies (Family Hippoboscidae). In: R.A. Bram

[ed.]. Surveillance and collection of arthropods of veterinary importance. USDA Agric.

Hbk. 518. pp. 60-71.

Samuel, W.M. and Trainer, D.O. \972.LipoptenamazamaeRondam, 1878 (Diptera: Hip-
poboscidae) on white-tailed deer in southern Texas. J. Med. Entomol. 9:104-106.
Throne, J.E. and Cline, L.D. 1989. Seasonal flight activity of the maize weevil, Sitophilus

zeamais Motschulsky (Coleoptera: Curculionidae), and the rice weevil, S. oryzae (L.), in

South Carolina. J. Agric. Entomol. 6:183-192.

Vol. 104, No. 3. May & June, 1993 133

OVERWINTERING AGGREGATIONS OF

FEMALE BRACHYMERIA INTERMEDIA

(HYMENOPTERA: CHALCIDIDAE). 1

Paul W. Schaefer 2

ABSTRACT: Discovery of 24 overwintering female Brachymeria intermedia in a single
aggregation in a window sash in an unheated building; a single female in an attic window
7.4 m above ground; a cluster under felt (tar) paper in a dog house; and in naturally occur-
ring dead stumps or trees all suggest that any dry, well protected site is suitable for
overwintering.

The introduced polyphagous pupal parasite Brachymeria intermedia
has spread throughout much of the northeastern United States and
Canada in close association with its principle host, the gypsy moth,
Lymantria dispar (L.) (Lepidoptera: Lymantriidae). Two, possibly three,
generations may occur during a season (Leonard 1 98 1 ). Adult females of
both generations overwinter while males die off before winter (Dowden
1935). Dowden (1935) speculated that overwintering adults "probably
hibernate under the bark of dead trees or in similar places". Only
decades later are we beginning to fully understand where overwinter-
ing occurs.

In Japan, Gyotoku (1957) found overwintering Brachymeria lasus
(Walker) (as obscurata) under the bark of a Chamaechyparis obtusa Endl.
(Cupressaceae) tree with two aggregations totaling ca. 30 wasps on the
lower 1.8 m of the south facing side of the trunk.

In laboratory experiments, Simser and Coppel (1980) showed the
presence of a chemical that led both B. lasus and B. intermedia to choose
previously used or conditioned overnight resting sites versus new or
unconditioned sites. They speculated that this chemically mediated
aggregating behavior might extend to overwintering individuals. Pro-
duced by adults of both sexes, this aggregation pheromone from B. inter-
media was later identified as 3-hexanone (Mohamed and Coppel 1987).

Waldvogel and Brown (1978) first reported the discovery of overwin-
tering Brachymeria intermedia. Groups of 5 to 15 females where found in
tunnels made by wood borers in a dead Quercus prinus L. tree in central
Pennsylvania. A similar discovery was made 6 January 1983 by Robert
Grebeck (USDA, BUR, Newark, DE) when ca. 1 5 females were found ca.
60 cm. off the ground in a dead Pinus rigida Mill, stump in Belleplain,
Cape May Co., New Jersey (B. Grebeck, pers. comm.).

1 Received December 23, 1992. Accepted March 15, 1993.

2 USDA, ARS, Beneficial Insects Introduction Research Unit Newark, Delaware, 19713.

ENT. NEWS 104(3): 133-135. May & June, 1993

134 ENTOMOLOGICAL NEWS

Overwintering B. intermedia also use non-natural sites. I report here
on a Dec. 30, 1981 discovery of a single aggregation of 24 female B. inter-
media found in a window frame/casing in a residence in Wethersfield,
Hartford Co., Connecticut. The window was located on an east wall of an
unheated shed attached to a family residence. The aggregating females
were clustered together in a vertical groove used to receive and hold a cot-
ton sash cord in an old-fashioned counter-balanced window. The groove
was in the side of a lower frame in a two-frame window. The cotton cord
had worn out and parted, however the end fragment still filled the groove
the entire 23 cm length. There was ample space between the cord and the
13x13 mm groove in the edge of the window frame so that wasps could
move about freely. At the time of discovery, the lower window was raised
almost daily but this did not appear to interfere with the overwintering B.
intermedia as they tended to cluster together against the cotton cord,
thereby avoiding being rubbed against the window casing as the sash
moved. Wasps were observable only after removing the window jams
and removing the sash from the frame. There was no indication of the
approach route used to enter this site. The window fit the frame so loosely
that it was possible the wasps entered the site simply by passing through
the crack between the sash and frame.

In late December 1982, another overwintering female B. intermedia
was found in a similar location in the same Wethersfield residence. One
female was found simply between a loose fitting window sash and its
frame. The window was a north facing unheated attic window at a height
of 7.4 m above ground level and about 2 m above the roof of the above
mentioned shed. This would suggest that overwintering sites are not
limited to lower strata.

Others have observed B. intermedia overwintering in different artifi-
cial situations. Bill Metterhouse (NJ Department of Agriculture, Tren-
ton), in the fall of 1982, found at least 25 female B. intermedia over-
wintering under felt (tar) paper covering a dog house at his residence in
Monmouth Co., New Jersey. Furthermore, that spring he found over 50
live B. intermedia females on the inside of his cellar windows and many
others dead on the basement floor (B. Metterhouse, pers. comm.).

All evidence reconfirms that only B. intermedia females overwinter.
Overwintering, singly or in aggregations, appears to occur only in con-
cealed, well protected, relatively dry sites, such as in dead trees that have
been excavated by other insects, and in man-made objects.

If the pheromone-mediated aggregation behavior of B. intermedia
females is involved in overwintering, as in over-night site selection, as
Simser & Coppel (1980) have speculated, then an artificial aggregation
site could be constructed, baited with aggregation pheromone, and

Vol. 104, No. 3. May & June. 1993 135

deployed as a survey tool to assess population levels or to measure over-
wintering survival.

ACKNOWLEDGMENTS

I thank Ronald Weseloh, Conn. Agric. Exp. Stn, New Haven; William Mctterhouse,
N.J. Dept. Agric.. retired; and Philip Taylor. USDA. BUR. Newark. Del., and two anony-
mous reviewers for suggestions on the manuscript.

LITERATURE CITED

Dowden, P.B. 1935. Brachymeria intermedia (Nees), a primary parasite, and B. compsilurae
(Cwfd.), a secondary parasite, of the gypsy moth. J. Agric. Res. 50(6):495-523.

Gyotoku, N. 1957. An example of hibernation of Brachymeria obscurata Walker. Shin-
konchu 10(7):52 (in Japanese).

Leonard, D. E. 1981. Brachymeria intermedia (Nees). (Hymenoptera: Chalcididae). pp.
394-398. In Doane, C.C. & M.L. McManus (eds.). The gypsy moth: Research toward
integrated pest management. U.S. Dept. Agric., Expanded Gypsy Moth Res. & Dev.
Program, Tech. Bull. 1584. 757 pp.

Mohamed, M.A. and H.C. Coppel. 1987. Pheromonal basis for aggregation behavior of
parasitoids of the gypsy moth: Brachymeria intermedia (Nees) and Brachvmeria lasus
(Walker) (Hymenoptera: Chalcididae). J. Chem. Ecol. 13(6): 1385-1393.

Simser, D.H., and H.C. Coppel 1980. Aggregation behavior of Brachymeria lasus
(Walker) in the laboratory. Environ. Entomol. 9:486-488.

Waldvogel, M.G. and M.W. Brown. 1978. An overwintering site of the gypsy moth
parasite. Brachymeria intermedia. Environ. Entomol. 7:782.

BOOKS RECEIVED AND BRIEFLY NOTED

BUGS OF THE WORLD. G.C. McGavin. 1993. Facts on File. 192 pp.

Another in the "Of the World" series, this is a popular guide to the classification and
biology of bugs, by Oxford entomologist. Dr. McGavin.

L'ABDOM EN ET LES GENITALI A DES FEMELLES DE COLEOP-
TERES ADEPH AG A. T. Deuve. 1 993. Tome 1 55, Memoires du Museum
National D'Histoire Naturelle.

With minor differences, this work is reproduced from a doctoral thesis of the University
of Paris 6, submitted 25 November 1988 under the title "Morphological and phylogenetic
studieson the abdomen and the female ectodermicgenitalia of the Coleoptera Adephaga".
Present text entirely in French language. Paperback.

136 ENTOMOLOGICAL NEWS

FIRST RECORDS OF PARASITOIDS FOR SLIME

MOLD BEETLES OF THE FAMILY SPHINDIDAE

(COLEOPTERA: CUCUJOIDEA)!

Joseph V. McHugh 2

ABSTRACT: The first three records of parasitoids for the family Sphindidae (Coleoptera:
Cucujoidea) are provided: (1) Sphindus americanus parasitized by Pentelicus sp. (varicornis
or near) (Hymenoptera: Encyrtidae) in New York, (2) Carinisphindus sp. by another Pen-
telicus sp. (probably sp. nov.) in Puerto Rico, and (3) Eurysphindus comatulus by Blacus
koenigi (Hymenoptera: Braconidae) in New York. These observations also represent the
first host information for the genus Pentelicus and a new host record for Blacus koenigi.

Sphindidae is a small family (9 genera and 51 species) of myxo-
mycophagous (slime mold eating) beetles that is represented in every
major biogeographical region in the world. Little has been published on
the biology of sjjhindids other than life history notes of one species
(Burakowski & Slipinski, 1987), host food records (see McHugh, 1993 for
citations) and a discussion of the possibility of assistance in slime mold
spore dispersal (see Blackwell, 1984; McHugh, 1993). During the sum-
mer of 1990, simple attempts to rear species of sphindids resulted in the
identification of the first parasitoids known for the family.

In June, a few specimens of an undescribed species of Carinisphindus
were collected from the sporocarp of a myxomycete (Stemonitis sp.) in the
Caribbean National Forest at El Verde Field Station, elev. 300 M., in
Puerto Rico. After the beetles were extracted and the slime mold was
examined closely, three Carinisphindus pupae were found. Within two
weeks, a single parasitoid wasp emerged from each pupa. One specimen
was caught and identified as an apparently undescribed species of the
encyrtid genus Pentelicus Howard (= Hemaenasius Ashmead) (J. S.
Noyes, pers. comm.).

In late July, a laboratory culture of Sphindus americanus LeConte also
was found to be parasitized by encyrtid wasps. The beetle culture was
started about one month earlier with field-collected sporocarps ofFuligo
septica (L.) Wiggers from Ithaca, New York. The parasitoid was deter-
mined as another species of Pentelicus, closely resembling Pentelicus
varicornis (Girault), but possessing an unusually long first funicle seg-
ment. This antennal feature may support recognition as a new species
(J. S. Noyes, pers. comm.). Collecting in two subsequent years suggests

1 Received December 5, 1992. Accepted January 5, 1993.

2 Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York

14853.

ENT. NEWS 104(3): 136-138. May & June, 1993

Vol. 104, No. 3, May & June, 1993 137

that this Pentelicus is most abundant in New York during the latter part of
the summer (late July- August), although sporocarps of the slime mold
are found colonized by S. americanus as early as May.

A culture of Eurysphindus comatulus McHugh was established in
August from fruiting bodies ofMucilago Crustacea Wiggers collected in
Brooktondale, Tompkins Co., New York. This culture was thought to be
free of arthropods other than mites and various life stages of E. coma-
tulus, but after two weeks it produced many specimens of the braconid
Blacus koenigi Fischer. The remains of many parasitized last-instar
larvae were found in a characteristic pose. Each larval skin was draped
over the top of a white, silky cocoon with the legs wrapped around the
cocoon as though holding it.

DISCUSSION

The Pentelicus species parasitizing Carinisphindus sp. in Puerto Rico
represents the first known parasitoid for a sphindid as well as the first
host record for a species of this encyrtid genus. The occurrence of a
second Pentelicus species parasitizing a species of Sphindus, a genus
closely related to Carinisphindus (see McHugh, 1993), suggests that the
association in Puerto Rico was not incidental.

The discovery that Blacus koenigi is a parasitoid of Eurysphindus coma-
tulus sheds light on the biology of this poorly understood braconid
genus. apek (1969) states that the tribe Blacini is made up mostly of
parasitoids of the larvae of curculionid beetles and related groups, but
adds that the taxonomic position of the genus Blacus "may be doubted as
very little is known about its biology, host relations, etc." later, Capek
(1970) suggests that the biology of Blacus involves parasitism of wood
boring Coleoptera larvae. Some species of Blacus are known to parasitize
mycophagous ("true fungus" feeding) beetles (Achterberg, 1975). In a list
of label data, Achterberg (1975) also reports thatfi. koenigi was collected
from a sporocarp ofStemonitisfusca Roth, a myxomycete known to be a
host of E. comatulus (as well as several other sphindid species). Achter-
berg adds that this species is "mainly collected in August and first half of
October". The parasitized culture ofE. comatulus was started with slime
mold sporocarps collected in August.

All specimens are deposited in the Cornell University Insect Collec-
tion with the exception of four specimens of Pentelicus Ivaricornis which
are in the reference collection of J. S. Noyes (The Natural History
Museum, London) and 16 specimens of Blacus koenigi at the Biosys-
tematics Research Centre (Agriculture Canada, Ottawa).

13 g ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

I thank P. R. Fraissinet for his help with field work in Puerto Rico and New York. E. R.
Hoebeke made original determinations of parasitoids and helped locate additional infor-
mation. J. S. Noyes provided the final determinations and information about the encyrtids.
The determination of specimens of B. koenigi was confirmed by M. J. Sharkey. P. R.
Fraissinet and E. R. Hoebeke also read and commented on an early version of this note.
Funds for the work in Puerto Rico were provided by the Center for International Studies at
Cornell University. Other support for this study was provided by NSF Grant No. BSR-87-
17401 and Hatch Project No. NY(C)- 139426 (both to Q. D. Wheeler).

LITERATURE CITED

Achterberg, C. van. 1975. A revision of the tribus Blacini (Hymenoptera, Bracondiae,

Helconinae). Tijdschr. Entomol. 118: 159-323.
Blackwell, M. 1984. Myxomycetes and Their Arthropod Associates. In Q. Wheeler and M.

Blackwell, eds., Fungus-Insect Relationships: Perspectives in Ecology and Evolution.

514 pp. Columbia Univ. Press, New York.
Burakowski, B. and S. A. Slipiriski, 1987. A New Species of Protosphindus (Coleoptera:

Sphindidae) From Chile With Notes and Descriptions of Immature Stages of Related

Forms. Annali Mus. Civ. Stor. Nat., Geneva, 86: 605-625.
Capek, M. 1969. An attempt at a natural classification of the family Braconidae based on

various unconventional characters (Hymenoptera). Proc. Entomol. Soc. Wash. 71(3):

304-312.
Capek, M. 1970. A new classification of the Braconidae (Hymenoptera) based on the

cephalic structures of the final instar larva and biological evidence. Can. Entomol. 102:

846-875.
McHugh, J. V. 1993. A revision ofEurysphindus LeConte (Coleoptera: Sphindidae) and a

review of sphindid classification and phylogeny. Syst. Entomol. 18: 57-92.

Vol. 104, No. 3. May & June, 1993 139

FIRST REPORT OF A TWISTED-WING INSECT

(STREPSIPTERA) LARVA IN A CADDISFLY

(TRICHOPTERA) 1

Kenneth L. Manuel, 2 Richard M. Bohart^

ABSTRACT: A microcaddisfly Oxyethira janella (Trichoptera: Hydroptilidae) was obser-
ved with a first stage strepsipteran triungulin larva in its abdominal cavity. The triungu-
linid belongs to the genus Xenos or Pseudoxenos (Strepsiptera: Stylopidae). This is the first
report associating Strepsiptera with Trichoptera.

While identifying adult aquatic insects, the senior author removed a
strepsipteran triungulin larva from the abdominal cavity of a female
microcaddisfly, Oxyethira janella. The O. janella specimen was collected
by light trap on the South Fork Edisto River, a Coastal Plain sand bot-
tomed blackwater river near Cope, Orangeburg County, South Carolina,
on October 22, 1990. Subsequently, the junior author identified the strep-
sipteran as a first stage larva of either Xenos, a Polistes wasp parasitoid, or
Pseudoxenos, a parasitoid of sphecid and eumenid wasps. The mounted
triungulin specimen is in the University of California at Davis Insect
Museum.

Polistes wasp colonies are extremely common in the dense riparian
vegetation overhanging the South Fork Edisto River. Roving triungulin
larvae released from strepsipteran parasitized wasps may come into con-
tact with caddisflies and other insects seeking daytime shelter in the low
light, high humidity environment of the riparian vegetation.

The O. janella specimen containing the triungulinid in its abdomen
was probably incidentally "parasitized." Due to the relative short life
span of most caddisfly adults, a strepsipteran may not be able to com-
plete its life cycle in a caddisfly even if it were physiologically adapted to
the host's body. In addition, the small ( 1 mm long) abdomen of O. janella
may not allow the complete development of the triungulin larva to an
adult.

To our knowledge, no aquatic insect has been reported as a normal or
incidental strepsipteran host. Aquatic entomologists, however, may
wish to look for additional examples of strepsipteran "parasitism" while
they are involved in adult aquatic insect identification.

ACKNOWLEDGMENTS

We thank Robert Kelley (Greenville, South Carolina, USA) and Alice Wells (Museum
and Art Galleries of The Northern Territory, Darwin, Australia) for their critical review
of the manuscript.

1 Received September 29, 1922. Accepted November 7, 1992

2 Duke Power Company, Applied Science Center. 1 3339 Hagers Ferry Road, Huntersville,
North Carolina 28078

3 Department of Entomology, University of California, Davis, California 95615-8584

ENT. NEWS 104(3): 139. May & June, 1993

140 ENTOMOLOGICAL NEWS

A METHOD FOR SEX DETERMINATION OF THE
COLORADO POTATO BEETLE PUPA,

LEPTINOTARSA DECEMLINEATA
(COLEOPTERA: CHRYSOMELIDAE) 1

Yvan Pelletier 2

ABSTRACT: A method for the determination of the sex of the Colorado potato beetle Lep-
tinotarsa decemlineata. at pupal stage is presented. This method is based on sexual differ-
ences of the external morphology of pupae. External morphology differences between
sexes at adult stage are also depicted.

The sex of the adult Colorado potato beetle can be determined by
examining morphological differences of the last abdominal sternite
(Rivnay 1928). The available graphic representations (Busvine 1980;
Rivnay 1928) do not clearly show the distinctive characters. Rivnay
(1928) provided a drawing of the ventral view of the tip of the abdomen of
the female only. Busvine ( 1 980) depicted the tip of the abdomen for both
male and female but his schematic drawings are difficult to interpret.
Pictures (Fig. Ic, d) show more visibly that the distal end of the last ster-
nite is depressed with a somewhat truncated border in the male; whereas
the depression is absent and the posterior border rounder in the
female.

The determination of the sex of Colorado potato beetle pupae would
be useful in situations where sexual dimorphism influenced larval
parameters such as larval weight (Pelletier and Smilowitz 1991). This
would reduce the time delay before sex determination and, more impor-
tantly, would allow sex determination of a larger proportion of beetles,
otherwise reduced by mortality during the pupal stage. A method for the
determination of the sex of Colorado potato beetle pupae is described for
the first time.

I observed that the 7th visible sternite of males is complete and de-
pressed in its center (Fig. 1 a) and the posterior margins of the 6th visible
sternite is somewhat truncated. In females, the 7th visible sternite is
divided in its center by a suture that is usually dark in color (Fig. 1 b). The
center of the 6th visible segment extends slightly posteriorly. To validate
this method, 100 fully grown larvae collected from the field were indi-
vidually caged in 1 oz cups filled with soil and allowed to pupate. The sex

1 Received September 24, 1992. Accepted January 16, 1993.

2 Agriculture Canada, Research Branch, P.O. Box 20280, Fredericton, N.B. E3B 4Z7

Canada.

ENT. NEWS 104(3): 140-142. May & June, 1993

Vol. 104. No. 3. May & June. 1993

141

142 ENTOMOLOGICAL NEWS

was then determined using the characters described above and each
pupa put back in its container. After emergence, each adult was sexed
again using the descriptions reported by Ravnay (1928) and Busvine
(1980). Sex determinations conducted on pupal and adult stages corres-
ponded in all cases.

ACKNOWLEDGMENTS

I thank G. Boiteau, Agriculture Canada, Fredericton and Z. Smilowitz, Pennsylvania
State University, for their constructive comments offered in review of the manuscript.

LITERATURE CITED

Busvine, J.R. 1980. Recommended methods for measurement of pest resistance to pesti-
cides, pp. 59-63. Food and Agriculture Organization of the United Nations. Rome.

Pelletier, Y., and Smilowitz, Z. 1991. Biological and genetic study on the utilization of
Solanum berthaultii Hawkes by the Colorado potato beetle (Leptinotarsa decemlineata
(Say). Can. J. Zool. 69(5): 1280-1288.

Rivnay, E. 1928. External morphology of the Colorado potato beetle (Leptinotarsa
decemlineata Say). J. New York Ent. Soc. 36(2): 125-141.

Figure 1. Ventral view of the tip of the abdomen of male (A) and female (B) pupa and male
(C) and female (D) adult of the Colorado potato beetle.

Vol. 104, No. 3, May & June, 1993 143

ESTABLISHMENT OF UROPHORA

QUADRIFASCIATA (DIPTERA: TEPHRITIDAE)

AND CHRYSOLINA QUADRIGEMINA

(COLEOPTERA: CHRYSOMELIDAE) IN

PORTIONS OF EASTERN UNITED STATES 1

E. Richard Hoebeke 2

ABSTRACT: This paper presents the first recorded distribution in the eastern United
States of the introduced weed biocontrol agents Urophora quadrifasciata (Diptera: Tephri-
tidae) and Chrysolina quadrigemina (Coleoptera: Chrysomelidae). The seed-head fly U.
quadrifasciata, released in North America in the early 1970's for the control of spotted and
diffuse knapweed (Centaurea spp.), is recorded from numerous localities in New York,
Pennsylvania, Vermont, New Hampshire, Massachusetts, Rhode Island, Connecticut, and
New Jersey. The leaf beetle C. quadrigemina, released in the mid- 1940's for the control of St.
Johnswort (Hypericum perforatum), is also documented from the northeastern U.S. with
records from New York, Pennsylvania, Ohio, West Virginia, and Maryland. A brief review
of the history and background information on the success of these two weed biocontrol
agents in North America are presented. Each of the biocontrol agents are also briefly
described.

This paper provides a brief review and background information on
two classical weed biocontrol projects that continue to attain moderate
success in managing two of North America's most dominant and abun-
dant introduced weeds on uncultivated land: spotted knapweed (Cen-
taurea maculosa Lamarck) and St. Johnswort (Hypericum perforatum L.).
The principal focus of this paper emphasizes new distributional data in
the eastern United States for the introduced fruit fly Urophora quadrifas-
ciata (Meigen) and the leaf beetle Chrysolina quadrigemina (Suffrian).
Both biocontrol agents were primarily released in western North
America for control of spotted knapweed and St. Johnswort, respec-
tively. These distributional records provide the first evidence of estab-
lishment of these introduced biocontrol agents in the eastern United
States.

I. The weed: Centaurea maculosa (Asteraceae)

Spotted knapweed is a herbaceous composite and short-lived peren-
nial introduced from Europe to the dry rangelands of western North
America (Harris, 1980; Harris & Myers, 1984). Because of its allelopathic

Received November 7, 1992. Accepted February 25, 1993.

Department of Entomology, Cornell University, Ithaca, New York 14853-0999.

ENT. NEWS 104(3): 143-152. May & June, 1993

ENTOMOLOGICAL NEWS

properties, low forage value, and drought adaptations, this knapweed
species has been able to displace and outcompete most other herbaceous
plants over vast areas of its new homeland (Harris & Myers, 1 984). It was
first collected in North America at Victoria, British Columbia in 1893
(Groh, 1943). In western Canada, it is distributed in British Columbia
and Alberta (Watson & Renney, 1974). The main areas of infestation
( > one million ha) are confined to the western United States in Montana,
Idaho, Washington, and Oregon (Maddox, 1982). In eastern North
America, this weed is common in Ontario, Quebec, and the Maritimes in
Canada (Frankton & Mulligan, 1970), and along roadsides and in fields
and waste areas in the northeastern and northcentral United States
(Cox, 1985).

The biocontrol agent: Urophora quadrifasciata 1 (Tephritidae)

Members of Urophora Robineau-Desvoidy (Diptera: Tephritidae)
have been widely employed as biocontrol agents of thistles and knap-
weeds of the composite family Asteraceae ( = Compositae). The genus
contains almost 100 known species distributed in Europe, temperate
Asia, Africa and the New World (White & Elson-Harris, 1992). Two
Urophora species, both of European origin and collectively referred to as
the seed-head flies, have been successfully introduced into western
North America for biocontrol of spotted knapweed (C. maculosa) and
diffuse knapweed (C diffusa Lamarck). Urophora a/finis (Frauenfeld)
and U. quadrifasciata (Meigen) oviposit into the developing inflorescen-
ces ofCentaurea species, induce gall formation, and ultimately reduce
seed production (Harris, 1980; Harris & Myers, 1984).

The usually univoltine U. affmis was originally released in British
Columbia (Ned's Creek) in 1970 and 5 western states in 1973 (Harris,
1980; Maddox, 1979; Story & Anderson, 1978; Story, 1985). By 1992, U.
affmis had been confirmed as established and increasing in abundance
in British Columbia, Idaho, Oregon, Montana, Washington and Wyom-
ing (Julien, 1992). Releases of U. affmis occurred in the east in 1971
(Ontario), 1979-1980 (Quebec), and 1983 (Maryland and New York)
(Harris & Myers, 1984; pers. commun., Stephen D. Hight, USDA-ARS,
Insect Biocontrol Laboratory, Beltsville, MD). Eastern establishment
had been confirmed as of 1992 in New York, Quebec, and Virginia
(Julien, 1992).

In the Palearctic, Urophora quadrifasciata may actually represent a species complex (see
White & Clement. 1987). with more than one species confused under the name quaclri-
fasciata.

Vol. 104. No. 3, May & June, 1993 145

The bivoltine U. quadrifasciata was first introduced in 1970 at Ned's
Creek, British Columbia, but was not released in the western United
States (Story, 1985). By May 1981, larvae oft/, quadrifasciata had been
found in spotted knapweed seed heads examined at a site in extreme
northwest Montana, ca. 400 km from the Ned's Creek original release
site (Story, 1985). By 1982, U. quadrifasciata was known to be established
in northwest and westcentral Montana (Story, 1985). Specimens of U.
quadrifasciata had been released and become established in Quebec in
1979(Julien, 1982) and 1980 (Harris & Myers, 1984). Between 25-31 May
1983, seed heads of field-collected spotted knapweed from British
Columbia were released by USDA-ARS personnel at 3 sites in New York
(Warren, Essex, and Tompkins counties) and at Beltsville, Maryland.
This material contained larval stages of both U. quadrifasciata and U.
affinis. A 1985 follow-up survey recovered only U. affinis at the Warren
and Essex County release sites located in the Adirondack region of
northern New York (pers. commun., S. D. Hight).

In Tompkins Co. (Trumansburg), New York, during July 1990,
numerous small tephritid flies were collected from the immature flower
heads of tyrol knapweed, C. dubia Suter, a common knapweed of the
fields and roadsides of southeastern Canada and northeastern U.S.
(Gleason & Cronquist, 1991). The flies were identified as U. quadrifas-
ciata (by the author and later confirmed), and a survey was initiated to
determine the geographic range in the northeast of this introduced seed-
head fly. The survey was conducted throughout portions of the north-
eastern states during June-September 1990-1992. During the survey, no
specimens of U. affinis were collected from knapweed.

In the following list of distributional data for U. quadrifasciata. the
abbreviations ERH (for the author) and AGW (for A. G. Wheeler, Jr.) for
collectors are used; dates of collection are expressed as "day-month
(Roman numeral)-year"; and hosts are abbreviated as follows: C.
maculosa, spotted knapweed (SK); C. dubia (=C. nigrescens & C. vochinen-
sis\ short-fringed knapweed (SFK); and C.jacea, brown knapweed (BK).
Treatment and usage of scientific and common names of Centaurea
follow Gleason and Cronquist (1991). The data below are also mapped
in Figure 1. All specimens, unless stated otherwise, are deposited in the
Cornell University Insect Collection. The author and A. G. Wheeler, Jr.
take responsibility for the host plant identifications, with the exception
of C dubia (see acknowledgments).

UNITED STATES: CONNECTICUT: Tolland Co., 1-84 West, N. of E. Wellington. 1 1-
MII-90, AGW, SK. MASSACHUSETTS: Plymouth Co., Rte. 58. nr. South Carver. 5-VIII-
90, AGW, SK. Worcester Co., Gardner, 4-V1II-90, AGW, SK. NEW HAMPSHIRE:
Hillsboro Co., Nashua, 5-VIII-90. AGW. SK. NEW JERSEY: Hunterdon Co., Exit 1 1 on

146

ENTOMOLOGICAL NEWS

Rte. 78, 12 mi. E. Phillipsburg, 10-VIII-91, ERH, SK. Sussex Co., High Point StPk., 28-VII-
90, AGW, SK; Rte. 23, nr. High Point St. Pk., 28-VII-90, AGW, SK. NEW YORK:
Allegany Co.: Rte. 1 7, Exit 33 (to Alfred), 7-VIII-92, ERH, SK; Rte. 17, 0.5 mi. E. Exit 37, 7-
VIII-92, ERH, SK; Alfred, 7-VIII-92, ERH, BK. Broome Co.: Binghamton, Junct. Rte. 81
and Rte. 12, 17-VIII-90, ERH, SK. Chemung Co., West Elmira, 30-VI-92, ERH. SK; Pine
City, 30-VI-92, ERH, SK; Elmira Heights, 7-IX-92, ERH, SK. Chenango Co., Brisben, 17-
VIII-90, ERH, SK; Norwich, 17-VIII-90, ERH, SK. Clinton Co., Rte. 456 @ junct. Rte. 87 E.
of Beekmantown, 29-VIII-92, AGW, SK; 1-87, Exit 36, S. of Pittsburgh, 2-VIII-92, AGW,
SK. Dutchess Co., 1-84 West, rest area nr. Storm ville, 1 l-VIII-90, AGW, BK. Greene Co.,
1-87 North, nr. Catskill, 3-VIII-90, AGW, SK. Jefferson Co., Plesis, 16-VIII-92, AGW, SK;
Wellesley Island St.Pk., 16-VIII-92, AGW, SK. Madison Co., DeRuyter, 17-VIII-90, ERH,
BK?; New Woodstock, 14-VII-91, ERH, SK. Ontario Co., nr. E. Victor (Farmington), 5-VII-
92, ERH, SK. Orange Co., 1-84 East, nr. Middletown, 3-VIII-90, AGW, SK. Putnam Co.,
Rte. 202, nr. Brewster, 1 l-VIII-90, AGW, BK. Rensselaer Co., Johnsonville, 3-VIII-90,

Figure 1. Northeastern United States. Distribution of Urophora quadrifasciata based on
examined specimens (dots). Known release sites (NY: Essex, Tompkins, and Warren coun-
ties; MD: Beltsville) (stars).

Vol. 104. No. 3, May & June. 1993 147

AGW, SK. Saratoga Co., 1-87 North, nr. Ushers. 3-V1II-90, AGW. SK. Schuyler Co.,
Alpine Junction, 6-V1II-92. ERH. SK; Watkins Glen. 31-V1II-90, 15-VIII-92. ERH. SK.
Steuben Co., Bath, 31-VIII-90, ERH, SK. Tioga Co., Owego, 17-V1II-90. ERH, SK;
Waverly, 17-V1II-90, ERH, SK. Tompkins Co., Ithaca, 14-VI-91, ERH. SK; Tnimansburg,
2 1- VI I -9 1,27- VI -92, ERH, SK; Town of Ulysses, N. of Jacksonville, 15-VIII-89.1 1.1 3. 16- VII-
90, 15-V1II-90, 12.15.17.20.27.VI-91.25-V1I-91, 15-V1II-91, 22-V1-92, 10. 12.29- VII-92, ERH,
SFK. Ulster Co., Mohonk Preserve, nr. New Paltz, 3-VIII-90, AGW, SK. Warren Co.,
Peggy Ann Rd., W. of Glens Falls, 3-V1II-90, AGW, SK. Yates Co., 10 mi. N. of Watkins
Glen, junct. Rte.42 and 14.4-IX-90, ERH, SK. PENNSYLVANIA: Bradford Co., Sayre.
17-VIII-90, ERH, SK; Wysox, 17-VIII-90, ERH, SK. Carbon Co., Rte. 534,0.3 mi. S. of junct.
Rte. 940 nr. East Side, 15-VII-90, AGW, SK. Cumberland Co., Rte. 1 14 @ junct. 1-81, nr.
Hogestown, 18-VII-90, AGW, SK. Dauphin Co., Rte. 39. nr. junct. Rte. 322 N. of Harris-
burg, 16-VII-90, AGW, SK. Lackawanna Co., 1-84 East, Mt. Cobb exit, 25-VII-90, AGW.
SK. Lancaster Co., Rte. 272, N. of Buck, 24-VII-090. AGW, SK. Lebanon Co., Rte. 934.
Indiantown Natl. Cem., 15-VII-90, AGW, SK. Luzerne Co., Rte. 93 @ 1-81, nr. West
Hazleton, 15-VII-90, AGW, SK. Lycoming Co., nr. Loyalsock, 28-VII-91, ERH. SK;
Muncy. 20-VII-90, K. Valley, SK. Monroe Co., Tobyhanna, 2 1 -VII-90, AGW. SK. Pike Co.,
Rte. 402, nr. Blooming Grove, 28-VII-90, AGW. SK. Schuylkill Co., Rte. 443. nr. New
Ringgold, 23-VII-90, T. Price. SK: Frackville. 15- VII-90, AGW, SK. Sullivan Co., Sones-
town, 20- VII-90, K. Valley, SK. Susquehanna Co., Montrose, 17-VIII-90. ERH, SK. Wayne
Co., Angels, 21-VII-90, AGW, SK. RHODE ISLAND: Kent Co., West Warwick. 5-VIII-90,
AGW, SK. VERMONT: Chirtenden Co., Camp Johnson, Colchester, 28-VIII-92, AGW,
SK. Franklin Co., Missisquoi Natl. Wildlife Refuge, 28-VIII-92, AGW, SK.

Comments. Adults oW. quadrifasciata and U. affinis are superficially
similar. Females of both species can be easily separated using the key in
White & Clement (1987:575). Among the chief characters that dis-
tinguish U. quadrifasciata are the four complete, transverse, black bands
and yellow base of the wing, with the first (or basal) and second
transverse bands broadly united at the costal margin (see Figure 8 of
Plate II in Freidberg & Kugler, 1989). In contrast, adults of U. affinis are
recognized by having 3 or 4 transverse bands (variable in intensity and
completeness), with the first band, when present, less pronounced than
the other three and separated from the second transverse band (as in
Figures 2-3 of Plate II in Freidberg & Kugler, 1989).

The Animal and Plant Health Inspection Service, Plant Protection
and Quarantine (APHIS-PPQ) of the USDA is initiating a biocontrol
program for diffuse and spotted knapweed in the eastern states (states
invited to participate in 1992 include Pennsylvania, New York, Michi-
gan, and Virginia). The locality records reported herein represent a
"prerelease inventory", establishing important base line data on existing
U. quadrifasciata populations in eight northeastern states.

II. The weed: Hypericum perforation (Hypericaceae)

St. Johnswort (also klamath weed or goatweed), Hypericum per-
foratum, a weed native to Europe, northern Africa, and large portions of

148 ENTOMOLOGICAL NEWS

Asia to China and Japan, was introduced into Australia and North
America. On the latter continent, it has become a serious weed on
rangelands in dry areas (Johansson, 1962). The first known introduction
of St. Johnswort into the United States was reported in 1 793 near Lancas-
ter, Pennsylvania. By 1900 it had spread westward and was reported in
California around the Klamath River, which provides the basis for one
of the plant's common names (Rosenthal et al, 1984). It is a hardy, deep-
rooted, perennial herb occurring in neglected meadows, fields, and pas-
tures and along roadsides from Newfoundland to Manitoba, south to
Florida and Texas, and in the far West from British Columbia to central
California (Cox, 1985). It remains a noxious rangeland weed only in the
western United States (Johansson, 1962).

In heavily infested areas, this weed is especially injurious by displac-
ing valuable and desirable forage plants. It is also toxic to livestock when
ingested in considerable quantities, causing a photodermatitis on un-
pigmented areas of grazing livestock exposed to direct sunlight (Jo-
hansson, 1962).

The biocontrol agent: Chrysolina quadrigemina (Chrysomelidae)

The first use of insects as a means of weed control in North America
was initially attempted in the mid-1940's with two European species of
leaf beetle (Chrysolina) that feed on St. Johnswort,//. perforation (Hollo-
way & Huffaker, 195 1). Chrysolina hyperici (Forster) and C. quadrigemina
(Suffrian) [= gemellata auct. and geminata auct] have become estab-
lished in release areas in western and eastern North America, with the
latter species exhibiting a greater ability to increase its distributional
range, particularly in California (Holloway & Huffaker, 1951).

Chrysolina hyperici was introduced into California (via Australia) in
1945 and is now established in many localities in the West, including
Oregon, Washington, Idaho, Montana, Colorado, and British Colum-
bia. Similarly, C. quadrigemina was introduced into California (via Aus-
tralia) in 1946, and is also established in the same areas as C. hyperici.
Another species, C. varians (Schaller), was introduced for the biocontrol
of Hypericum in British Columbia where it remains established and
apparently restricted (Johansson, 1962; Brown, 1962).

Populations of//, perforatum were dramatically reduced when both
C. hyperici and C. quadrigemina were introduced into Australia (Clark,
1953), the United States (Holloway & Huffaker, 1951; Holloway, 1957),
and western (British Columbia) and eastern Canada (Ontario and Nova
Scotia) (Smith, 1958; Harris & Maw, 1984). A relatively small number of
specimens of C hyperici (252) and C. quadrigemina (182) were originally
released in eastern Ontario in 1969 and 1970, respectively, for control of

Vol. 104, No. 3, May & June, 1993

149

St. Johnswort (Harris & Maw, 1984). Fields et al. (1988) demonstrated
that 18 years after their initial release both species of leaf beetle had
spread nearly 90 km from the original release site near Picton, Ontario.
They also concluded that the present distribution ofChrysolina spp. in
eastern Canada was probably due to natural dispersal, and that both
species were capable of finding widely separated stands of the host
plant.

No systematic release of either Chrysolina species has occurred in the
eastern United States. However, C. quadrigemina has been collected from
several widespread localities in New York, Pennsylvania, Maryland,
West Virginia, and Ohio, since 1989. Several states in the east have not
been surveyed; therefore, no statement can be made about the presence
or absence of this species in these regions. The known distributional
records for C. quadrigemina document its establishment and range
expansion in the eastern United States, apparently resulting from a
natural dispersal of populations from eastern Ontario. These data are
recorded below and mapped in Figure 2. All specimens were collected
from H. perforatum, unless stated otherwise. The host plant iden-

Figure 2. Northeastern United States. Distribution of Chrysolina quadrigemina based on
examined specimens (dots). Known release site (near Picton, Ontario) (star).

150 ENTOMOLOGICAL NEWS

tifications, where noted, are provided by the author and other collectors.
The leaf beetle determination is the responsibility of the author.

The New York and Pennsylvania specimens are deposited in the Cor-
nell University Insect Collection; Ohio and West Virginia specimens in
the collection of the West Virginia Department of Agriculture, Charles-
ton, WV; and Maryland specimens in the collection of the Maryland
Department of Agriculture, Annapolis, MD.

UNITED STATES: MARYLAND: Prince Georges Co., Brandywine, l-V-91, C. L.
Staines. NEW YORK: Cattaraugus Co., Allegany St. Pk., 24 July 1985. A. E. Hajek.
Chemung Co., no specific locality, 1 -VII-90, C. Klass. Erie Co., Tonawanda, 3-VII-92, E. R.
Hoebeke. Ontario Co., Geneva, 16- VI-91, ERH; along Rte. 90, W. of Geneva exit 16- Vl-91,
ERH. Tompkins Co., Ithaca, 10-VII-89, 12-IX-90, C. Klass; Ithaca, Sept.-Oct. 1990, R.
Campbell (student collection); Ithaca, Forest Home Wildflower Garden, l-X-89, ERH;
Town of Ulysses, N. of Jacksonville, 15. 17.20- VI -91, ERH; Trumansburg, fairgrounds, 1-
VII-90, ERH; Trumansburg, Falls Rd. nr. H. A Smith Woods, 6-VII-89, ERH; Trumans-
burg, Taughannock Falls St. Pk., 6-VII-89, ERH. OHIO: Ashland Co., Mohican Mem. St.
For., 30-V-87, S. M. Clark. PENNSYLVANIA: Berks Co., Slote Nurseries, nr. Angelica, 25-
X-90, AGW. Centre Co., Scotia Barrens, 29-V1-91, A. G. Wheeler, Jr. WEST VIRGINIA:
Greenbrier Co., Anthony, 9-VI-92, SMC. Pocahontas Co., Cass, 8-VII-92. SMC, ex.. H.
punctatum. Randolph Co., Cheat Mountain, 1 mi. n. Barton Knob, 3800 ft. elev., 8-VII-92,
SMC. Tucker Co., Dolly Sods Scenic Area, 16-IX-92, SMC.

Comments. Of the sixteen Chrysolina species recorded in North
America (Brown, 1962), C. quadrigemina can be generally distinguished
from its congeners in eastern North America by the following charac-
teristics: its color (blue, blue-green, brassy green, or bronze individuals
with venter and legs dark blue or blue-green); its distinctly larger, more
robust size (6.0-7.1 mm); and, in the male, by the presence of a saucer-
shaped impression on abdominal sternite V, and by the size and details
of the aedeagus (Brown, 1962; Wilcox, 1972; Frazer & Emberson, 1987).

ACKNOWLEDGMENTS

I am especially grateful to A G. Wheeler, Jr. (Bureau of Plant Industry, Pennsylvania
Dept. of Agriculture, Harrisburg) for providing both specimens and locality records for
northeastern U.S. populations oft/, quadrifasciata and C. quadrigemina, and for reviewing
the manuscript; and to Shawn M. Clark (West Virginia Department of Agriculture,
Charleston) and Charles L. Staines, Jr. (Maryland Department of Agriculture, Annapolis)
for allowing me to publish locality records for C. quadrigemina collected in West Virginia,
Ohio, and Maryland. I also thank Robert Richard (USDA, Biocontrol of Weeds Facility,
Bozeman, MT) and two other anonymous reviewers for commenting on the manuscript.
William J. Dress (L. H. Bailey Hortorium, Cornell University) identified the short-fringed
knapweed, C. dubia (identified as C. nigrescens), surrounding my property upon which
specimens off/, quadrifasciata were first collected in July 1990. Allen L. Norrbom (Sys-
tematic Entomology Laboratory, USDA, Washington, D.C.) kindly confirmed the iden-
tification off/, quadrifasciata.

Vol. 104, No. 3, May & June, 1993 151

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Clark, L. R. 1953. The ecology of Chrysomela gemellata Rossi and C. hyperici ForsL, and
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Cox, D. D. 1985. Common flowering plants of the Northeast: their natural history and
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Fields, P. G., J. T. Arnason, and B. J. R. Philogene. 1988. Distribution ofChrysolina spp.
(Coleoptera: Chrysomelidae) in eastern Ontario, 1 8 years after their initial release. Can.
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Frankton, C. and G. A. Mulligan. 1970. Weeds of Canada. Can. Dept. Agric. Publ. 948.
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Frazer, B. and R. Emberson. 1987. Rediscovery of Chrysolina quadrigemina (Suffrian)
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Freidberg, A. and J. Kugler. 1989. Diptera: Tephritidae. Fauna Palaestina, Insecta IV.
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Rosenthal, S. S., D. M. Maddox, and K. Brunetti. 1984. Biological methods of weed con-
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Smith, J. M. 1958. Biological control of Klamath weed. Hypericum perforatum L., in British
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ENTOMOLOGICAL NEWS

Story, J. M. 1985. First report of the dispersal into Montana of Urophora quadrifasciata
(Diptera: Tephritidae), a fly released in Canada for biological control of spotted and
diffuse knapweed. Can. Entomol. 117:1061-1062.

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and C. maculosa. Can. J. Plant Sci. 54:687-701

White, I. M. and S. L. Clement. 1987. Systematic notes on Urophora (Diptera, Tephri-
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White, I. M. and M. M. Elson-Harris. 1992. Fruit flies of economic significance: their
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OL.104

USISSN0013-872X
SEPTEMBER & OCTOBER, 1 993 NO. 4

INTO

scriptions of immature stages of

Nemafus desan^isi (Hymenoptera:

Tenthredinidae), a pest of Salicaceae

in Argentina and Chile S.M. Ovruski', D.R. Smith

Carpenter ant (Hymenoptera: Formicidae)
tunnels visualized by computed

tomography

An inexpensive vacuum collector for insect

sampling S. W. Wilson, J.L. Smith, A.M. Purcell, ///

Insect removal from sticky traps using a citrus oil

solvent R.S. Miller, S. Passoa, R.D. Waltz, V. Mastro

BOOK REVIEWS

CALVE RT AWARDS FOR 1993

SOCIETY MEETING OF FEBRUARY 24, 1993

153

Amblycerus teutoniensis (Coleoptera:
Bruchidae), a new species of seed
beetle C.S. Ribeiro-Costa, J.M. Kingsolver 161

Dragonflies and damselflies (Odonata) of Buck

Creek, Pulaski County, KY R.G. Payne, G.A. Schuster 165

Caddisflies (Trichoptera) of Wildcat

Creek, Pickens County, SC M.A. Floyd, J.C. Morse 171

Abundance and seasonal activity of
Eucinetoidea (Coleoptera) in a
raspberry plantation and adjacent
sites in southern Quebec C. Levesque, G-Y. Levesque 180

New distribution record for Ischnoptera bilunata

(Dictyoptera: Blattellidae) E.P. Benson, A.G. Appel 187

First record of Sepedophilus coronadensis
(Staphylinidae) from Mexico

Jose Luis Navarrete-Heredia 191

Genera of Baetidae (Ephemeroptera) from

Central America C.R. Lugo-Ortiz, W. P. McCafferty 193

RifS.El-Mallakh 198

203

209
214
197
215

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Vol. 104, No. 4, September & October, 1993 153

DESCRIPTIONS OF IMMATURE STAGES OF

NEMATUS DESANTISI (HYMENOPTERA:

TENTHREDINIDAE), A PEST OF SALICACEAE

IN ARGENTINA AND CHILE*

Sergio M. Ovruski 2 , David R. Smith 3

ABSTRACT: The egg, first-and last-instar larvae, prepupa, and pupa ofNematus desaniisi
Smith are described and illustrated. This species is a serious pest otSalix spp. and Populus
spp. in Argentina and Chile.

Only three species of the tenthredinid subfamily Nematinae are
known in South America: Pristiphora brasiliensis Malaise, P. plaumanni
Wong, and Nematus desantisi Smith. Host plants of the Pristiphora species
are unknown. Nematus desantisi is a serious pest ofSalix spp. in much of
Argentina and Chile, but it has also been recorded from Populus spp. in
Argentina (Gianti and Dapoto 1990). Reproduction of N. desantisi is by
thelytokus parthenogenesis (De Sands and Gallego de Sureda 1984),
which facilitates its development as a pest.

The earliest record of W. desantisi is from Chubut, Argentina, in 1980
(De Santis 1981 ), and shortly after it was described by Smith (1983). Later
it was collected in Chile (Gonzalez^ al. 1986, Gonzalez 1989)andinthe
Argentina provinces of Rio Negro, Neuquen, Mendoza, San Juan, San
Luis, Buenos Aires (De Santis and Gallego de Sureda 1984), Catamarca
(Vattuone 1989), and Tucuman, Salta, and Jujuy (Ovruski 1991, Ovruski
and Fidalgo, 1991 ). De Santis and Gallego de Sureda (1984), Mallea et al.
(1985), Gonzalez et al. ( 1986), Gonzalez ( 1989), and Gianti and Dapoto
(1990) provided some biological data and gave brief descriptions of the
egg, late-instar larva, cocoon, pupa, and adults.

This paper, part of the graduation thesis of the senior author (Ovruski
1991 ) includes a detailed description of the egg, external morphology of
the first- and last-instar larvae, prepupa, and pupa of this serious pest.
The descriptions are from series of specimens collected on willow in Tafi
del Valle, Tucuman Province, Argentina, in January, March, and Octo-
ber of 1990, and reared in the laboratory. Larval terminology is based
largely on that of Wong (1963).

1 Received February 19. 1993. Accepted March 26. 1993.

- Centre de Investigaciones para la Regulation de Poblaciones de Organismos Nocivos
(CIRPON). Pasaje Caseros 1050. C.C. 90. S.M. de Tucuman. Argentina.

3 Systematic Entomology Laboratory. ARS. PSI. U.S. Department of Agriculture, c/o
National Museum of Natural History NHB 168. Washington. D.C. 20560.

ENT. NEWS 104(4): 153-160. September & October. 1993

154 ENTOMOLOGICAL NEWS

Description of Immature Stages

Egg (Fig. 1). Length, 1.0-1.2 mm. Entirely green; elongated kidney-shaped with one end
narrow and slightly curved, averaging from 0.26-0.28 mm in diameter; other end broader,
averaging from 0.37-0.39 mm in diameter, with narrowly rounded apex; chorion smooth.
Described from numerous series of eggs laid in leaves on first day of oviposition, and from
mature ovarial eggs from females reared in laboratory.

Last-instar larva (Figs. 2-11,18). Length, 1 7-20 mm. Head capsule pale yellow with black
eyespots; longitudinal dark brown to black band present along coronal suture, and a light
brown band from vertex laterally to each ocularium; dorsal half of frons, antenna, clypeus,
maxillary palpus, labial palpus, and basal part of mandible light brown; apex of mandible
black.

Body entirely green when alive, with two darkened dorsolateral lines; thoracic legs pale
yellowish with dark brown tarsal claws; apex of epiproct and caudal protuberances
(pseudocerci of Middleton 1921) light brown.

Head capsule (Fig. 2) circular in front view, with few scattered setae that are about 0.09
mm long and slightly longer setae on genae. Antenna (Fig. 3) with 4 segments, each reduced
to short sclerotized pieces included in oval antacoria; 1st segment very small and subcir-
cular, its diameter at least half diameter of apical segment and bearing light colored sen-
sory pit; 2nd and 3rd segments crescent-shaped and almost equal in size, each with two
light-colored sensory pits; 4th (apical) segment subcircular and bearing four light-colored
sensory pits. Clypeus wider than long and bearing 4-6 setae. Inner surface of labrum (Fig. 4)
with apical margin emarginate and sinuous at the middle, lateral margins rounded, with
10-12 long setae on each side. Mandibles asymmetrical, strongly sclerotized; each with two
large sharp teeth and five smaller lateral teeth (Fig. 5). Maxillary palpus 4-segmented, 2nd
segment with 1 seta, longer than 3rd and 4th segments combined; lacinia with 10-12 short
spines; palpifer with 4 setae. Labial palpus 3-segmented.

Prothoracic, mesothoracic, and metathoracic terga (Fig. 7) each apparently divided into
4 annulets and with few setae. Prothorax with a large spiracle and a single pore slightly
below spiracle. Second prothoracic annulet bearing one pore positioned dorsolaterally.
Mesothoracic and metathoracic segments similar to each other; preepipleurite with 5-6
setae and a single pore; postepipleurite with 2 setae. Middle thoracic leg (Fig. 9) with sub-
rectangular coxa, 1.25X longer than wide, with 13-16 scattered setae each about 0.09 mm
long; trochanter subquadrate, slightly longer than broad, with 10-12 setae that are 0.05 mm
in length; femur subcylindrical, with 6-7 setae similar to the former and 1 seta on femoral
process; tibia subcylindrical, 2X longer than wide, with 7-8 scattered setae that are 0.10 mm
in length and 3-4 shorter setae and one pore on apical margin; tarsus with a simple
claw.

Abdominal segments 2-7 and 10 with prolegs (Fig. 18); segments 1-8 each with 5 dorsal
annulets; 9th segment divided into 4 annulets, and 10th without annulets. Third ab-
dominal segment (typical abdominal segment) as follows (Fig. 6): 1st and 5th annulets
glabrous; 2nd annulet with vertical row of 4 setae dorsolaterally; 3rd annulet with vertical
row of 4 setae and 2 pores on dorsum and pleuron; 4th annulet with 2 setae located subdor-
sally; with small spiracle; postspiracular area with 1 pore and 1 seta above pore (sometimes
absent); preepipleurite nearly always with 5 setae (occasionally 4) and a single pore; post-
epipleurite usually with longitudinal row of 5 setae and 1 pore; an emarginate lobe bearing
2 setae posterior to postspiracular area; proleg usually with 3 to 4 setae. Ninth segment
apparently with 4 dorsal annulets, distribution of setae and pores as in Fig. 8. Epiproct with
a pair of caudal protuberances, several short setae and 1 pore. Numerous setae about 0.10
mm in length on subanal and suranal lobes (Fig. 8). Spiracles of abdominal segments 1-7

Vol. 104, No. 4, September & October, 1993

155

11

Figs. 1-11. Nematus desantisi. 1, Egg. 2-1 1, Last-instar larva. 2, Head capsule, front view. 3,
Antenna. 4. Inner surface of labrum. 5. Left and right mandibles, dorsal view. 6. Third
abdominal segment. 7. Head and thorax. 8, Apical abdominal segments. 9. Middle thoracic
leg. 10, Spiracle of third abdominal segment. 1 1, Pore.

156

ENTOMOLOGICAL NEWS

vertical, oval-shaped, and 2.25X longer than wide (Fig. 10), 8th abdominal spiracle longer
and very similar to prothoracic spiracle. Pores with 3 dark sharp protuberances on edge of
opening (Fig. 11). Eversible ventral glands present on abdominal segments 1-7, situated
between and slightly anterior to prolegs (not evident unless fully extended). Integument
with very small grayish granules.

Described from a series of 10 larvae, collected from Salix humboldtiana Willdn., and 5.
babylonica L. in Tafi del Valle.

First-instar larva (Figs. 12-17). Length, 1.5-2.0 mm at hatching, and reaching to 3.5-
4.0mm at beginning of second instar. First-instar (Fig. 17) similar to last-instar, except for
coloration of head capsule, mouthparts, thoracic legs, epiproct and caudal protuberances,
morphology of antenna, mandibles, thoracic legs, and abdominal spiracles, and distribu-
tion, number, and types of setae on head capsule, thorax, and abdomen.

Head capsule and mouthparts brownish black, same color as thoracic legs, except
trochanters which are greenish brown. Body, at hatching, translucent with grayish tone,
but pale green after feeding. Apex of epiproct and caudal protuberances dark brown.

Head capsule (Fig. 12) with numerous small setae each about 0.02 mm in length.
Antenna (Fig. 1 3 ) elongate, cone shaped; basal 3 segments ring-shaped and apical segment
subconical; diameter of 1st segment subequal to length of antenna, bearing 2 light-colored
sensory pits; 2nd and 3rd segments bearing a single light-colored sensory pit, and 4th seg-
ment bearing 4 light-colored sensory pits. Left mandible (Fig. 14A) with 7 sharp teeth, 2 of
which are very large and sharp, and a single, large, saw-shaped tooth; right mandible (Fig.
14B) with 6 sharp teeth and 1 large saw-shaped tooth.

16

Figs. 12-16. Nematus desantisi, first instar larva. 12, Head capsule, front view. 13. Antenna.
14A, Dorsal view of left mandible. 14B, Dorsal view of right mandible. 1 5, Middle thoracic-
leg. 16, Spiracle of third abdominal segment.

Vol. 104, No. 4, September & October, 1993 157

Middle thoracic leg (Fig. 1 5) with coxa subquadrate, bearing 20-22 scattered short setae
each about 0.02 mm in length; trochanters subtriangular, 2X broader than long, with 6
short setae; femur subquadrate, 1.25X broader than long, with 9-10 short setae; tibia sub-
quadrate, with 5 setae each about 0.03 mm in length, and 6-7 setae 0.015 mm in length, 2
large setae each about 0.05 mm in length and 4 pores near apical margin; tarsus with a large,
curved tarsal claw.

Third abdominal segment with vertical row of 3 setae on both 1st annulet and 2nd
annulets; vertical row of 4 setae on 4th annulet; 3rd and 5th annulets glabrous; spiracle sub-
circular (Fig. 16), about as long as wide, with brownish spots on anterior and posterior
sides, and grayish granules surrounding it on all sides; postspiracular area with 2 setae;
preepiplurite with 5-6 setae; postepiplurite with longitudinal row of 6-7 setae; 3 setae on
lobe posterior to postspiracular area; proleg with 6. occasionally 7, setae. Pores not found.

Described from 10 larvae reared in the laboratory.

Prepupa and pupa (Figs. 19, 20). Initially (first day), prepupa very similar to last-
instar larva. On second day, body curved and reduced in size, averaging 7.5 mm in length
(Fig. 19). Head capsule pale green, but with light brown lateral bands, longitudinal black-
ish band on coronal suture, and light brown spot on frons; thoracic legs greenish brown
and body dark green.

Pupa initially somewhat tough, entirely green with light brown eyes, averaging 7.8 mm
in length; antenna curved posteriorly, exterior to wings and extended to 4th abdominal seg-
ment; prothoracic and mesothoracic legs and wings curved toward ventral surface of
thorax; metathoracic legs extend to 5th abdominal segment. Mature pupa (Fig. 20) more
sclerotized; head, antenna, thorax, and thoracic legs yellowish brown; mandibles dark
brown, eyes black, and abdomen light green.

Fifteen prepupae and pupae, reared in the laboratory from larvae collected in Tafi del
Valle, were studied. The prepupa transforms to the pupa and adult stage inside a yellowish
brown oval cocoon from 7-8 mm long and from 3-4 mm in diameter.

DISCUSSION

The first descriptions of the immature stages of N. desantisi by De
Santis and Gallego de Sureda (1984), Mallea el al. (1985), Gonzalez
(1989), and Gianti and Dapoto (1990) were based principally on colora-
tion and measurements. De Santis and Gallego de Sureda (1984) pub-
lished the first figures of the mature larva, eggs in willow leaves, first-
instar larva, and prepupa inside the cocoon. These same authors men-
tioned that the eggs of N. desantisi are initially oval-shaped, flat, and tran-
slucent. Conversely, Mallea et al. (1985), Gonzalez etal. (1986) and Gon-
zalez (1989) described the eggs as kidney-shaped. This latter observation
agrees with our studies.

Gonzalez (1989) stated that the larva of N. desantisi has prolegs on
abdominal segments 2-8. Our studies show that the larva is typical of the
Nematinae and Nematus, with prolegs on abdominal segments 2-7 and
10, as defined by Smith and Middlekauff (1987).

Very few sawfly larvae are known from southern South America, but
the following combination of characters should aid in the recognition of
the larva of Nematus desantisi: feeding on Salix spp. and Populus spp.; pre-

158

ENTOMOLOGICAL NEWS

17

Figs. 17-18. Nematus desantisi. 17, First-instar larva. 18, Last-instar larva.

Vol. 104. No. 4. September & October, 1993

159

19

20

Figs. 19-20. Nematus desantisi. 19, Cocoon and prepupa. 20, Mature pupa.

160 ENTOMOLOGICAL NEWS

sence of prolegs on abdominal segments 2-7 and 10; flat, 4-segmented
antenna; mandibular dentition as described above; 5-annulate ab-
dominal segments (segments 1-8) with setae on annulets 2-4; and pre-
sence of a pair of caudal protuberances on the epiproct.

ACKNOWLEDGMENTS

We thank Dr. Patricio Fidalgo and Ing. Agr. Arturo L. Teran (CIRPON, S.M. de Tucu-
man, Argentina) for suggestions and critical reviews of the manuscript. Lie. Nora E.
Ovruski (EEAOC, S.M. de Tucuman, Argentina) for helping collect the specimens, and
Ing. Hugo Lazaro (CIRPON) for the photographs of the larvae, prepupa, and pupa. We
also thank W.W. Middlekauf, University of California, Berkeley; H. Goulet, Agriculture
Canada, Ottawa; and R.V. Peterson and D.A Nickle, Systematic Entomology Laboratory,
USDA Washington, D.C., for review of the manuscript.

LITERATURE CITED

De Santis, L. 1981. Estudiodeuna nueva plaga defoliadora delsaucecriolloenla provin-

cia de Chubut. Novedades del Museo de la Plata 1(1): 9.
De Santis, L. and A. Gallego de Sureda. 1984. La falsa oruga de los sauces y alamos

(Nematus desantisi). Academia Nacional de Agronomia y Veterinaria, Buenos Aires

38(7): 1-22.
Gianti, H.E. and G.L. Dapoto. 1990. Biologia, daftos y posibilidades de control de la

"Falsa oruga o cuncuna del sauce." Presencia 4(20/21): 11-12.
Gonzalez, R.H. 1989. Insectos y Acaros de importancia agricola y cuarentenaria de Chile.

Universidad de Chile. 310 pp.
Gonzalez, R.H., G. Barria, and M.A. Guennero. 1986. Nematus desantisi Smith, nueva

especie de importancia forestal en Chile (Hymenoptera: Tenthreidnidae). Revista

Chilena de Entomologia 14: 13-15.
Mallea, A.R., G.S. Macola, J.G. Garcia Saez, and S.J. Lanati. 1985. Observaciones

bioetoecologicas sobre Nematus desantisi Smith, 1983 (Hymenoptera: Tenthredinidae),

en Mendoza. Intersectuum 17(1-3): 1-14.
Middleton, W. 1921. Some notes on the terminal abdominal structures of sawflies. Proc.

Entomol. Soc. Wash. 23: 139-144.
Ovruski, S.M. 1991. Estudios biologicos morfologicos de la falsa oruga de los sauces y

alamos. (Nematus desantisi) y consideraciones sobre su distribucion. Tesis de gradua-

cion (Seminario), Fac. de Cs. Naturales e instituto M. Lillo, Universidad Nacional de

Tucuman, Argentina, 94 pp.
Ovruski, S.M. and P. Fidalgo. 1991. Distribucion geografica de Nematus desantisi Smith

(Hymenoptera: Tenthredinidae), plaga de salicaceas. Ciencia e Cultura, Sao Paulo

(suplemento) 43(7): 36-37.
Smith, D.R. The first record of Nematus Panzer from South America: a new species from

Argentina (Hymenoptera: Tenthredinidae). Proc. Entomol. Soc. Wash. 85: 260-262.
Smith D.R. and W.W. Middlekauf. 1987. Symphyta, pp. 618-649. In Stehr, F.W., ed.,

Immatuire Insects. Kendall Hunt, Dubuque, Iowa. 754 pp.
Vattuone, E.M. 1989. La falsa oruga de los sauces y alamos (Nematus desantisi Smith) en la

provincia de Catamarca. CIRPON, Revista de Investigacion 7(1-4): 85.
Wong, H.R. 1963. The external morphology of the adult and ultimate larval instar of the

larch sawfly, Pristiphora erichsonii (Hymenoptera: Tenthredinidae). Canadian Ento-
mol. 95: 897-921.

Vol. 104, No. 4, September & October, 1993 161

AMBLYCER US TE UTONIENSIS

(COLEOPTERA: BRUCHIDAE),

A NEW SPECIES OF SEED BEETLE 1 ' 2

Cibele S. Ribeiro-Costa, 2 John M. Kingsolver^
ABSTRACT: Amblycerus teutoniensis is described and illustrated.

To provide a specific name to be used in future species group ar-
rangements, this description is presented.

Amblycerus teutoniensis, new species

(figs. 1-8)

Dimensions: Medium body length 5.14 mm; width 3.36 mm. Pronotum length 1.38-
1.80 mm (mean = 1.62 mm); width 1.96-2.63 mm (mean =2.44 mm). Elytra length 2.48-4.20
mm (mean =3.52 mm); width 2.84-3.80 mm (mean =3.36 mm).

Integument: Black except four basal antennal segments, tarsi and calcaria reddish;
entire pygidium and abdomen reddish yellow.

Vestiture: Head, pronotum, elytra, venter of thorax and appendages with brown and
bluish gray hairs in irregular mottled pattern (fig. 1 ). Pronotum sometimes with four small
rounded bluish gray spots (fig.2). Scutellum densely pubescent with light yellow hairs
(fig.4). Pygidium (fig.5) and abdomen covered with light yellow and golden yellow hairs in
a mottled pattern and with some scattered small brown patches.

Body: Subquadrate (fig.l). Vertex micropunctate; frons and clypeus more coarsely
punctate than vertex except granulose in narrow apical band; labrum punctate basally.
Frons gently flattened, frontal carina evanescent in lower half, sometimes absent; fron-
toclypeal suture indistinct. Mesal margin of eye with fine carina and umbilicate punctures;
eye finely faceted, moderately protruding laterally; ocular sinus (emargination) 1/3 length
of eye and ocular index (width across eyes/width between eyes) 2.8: 1 ; postocular lobe long.
Antenna subserrate from fifth to tenth segments, eighth to tenth segments slightly wider
than long; terminal segment subelliptical (fig.6). Pronotum (fig.2) subconical. lateral
margins gently arcuate; disk evenly convex; basal lobe broadly angulate, usually not sul-
cate; surface densely punctulate, lateral one-third of disk on either side also coarsely
punctate, middle also with some punctures smaller than those on lateral areas; basal and
apical margins without sulci; lateral carina (fig.3) divided near base and gently divergent
toward apex, delimited by fine shallowdorsal and fine deep ventral sulci; cervical boss with
two fine setae (fig. 3), posterior angle of pronotum with one seta. Prosternum moderately
narrow with sulcate lateral margins and slightly expanded beyond procoxae. Scutellum 1.3

1 Received October 16, 1992. Accepted April 22, 1993

- Contribution no.738 from Departamento de Zoologia of the Universidade Federal do
Parana, C.P.: 19030; CEP:81531-970, Curitiba, Parana, Brasil.

3 Florida State Collections of Arthropods, Division of Plant Industry, PO Box 147100.
Gainesville. Florida 32614-7 UK), U.S.A.

ENT. NFWS 104(4): 160-164. September & October. 1993

162 ENTOMOLOGICAL NEWS

times as long as wide, apex trilobed (fig.4). Elytra as long as wide, evenly convex except
slightly depressed around scutellum; surural, third, fifth, seventh and ninth intervals gently
elevated on middle apical portion; striae strongly impressed, strial punctures fine; elytral
apices rounded. Mesosternum elevated, rounded apically. Postmesocoxal sulci meeting
mesally at acute angle, then extending laterally and connecting to parasutural sulci, the lat-
ter extending beyond middle length of metasternum. Metepisternum punctulate, also with
many coarser punctures, lacking striate file; metepisternal sulcus nearly right angled, verti-
cal arm reaching apex and longitudinal arm very short, not reaching middle length of
metepisternum. Metasternum between middle coxae not bulging. Face of hind coxa in dis-
tal two-thirds and along posterior border of proximal one-third setose and densely punctu-
late; many scattered larger punctures on distal two-thirds; proximal one-third glabrous in
part and punctate except in a reduced area. Metafemur slender, 2.9 times as long as wide
(fig.7); ventral face slightly sulcate in distal three-fourths; mesoventral carina complete but
lacking blunt, angulate process near apex. Ventral face of hind tibia slightly concave, each
margin with row of punctures and short, stiff setae; mesal face lacking tumidity at distal
two-thirds; apex with a few, short coronal denticles. Mesal tibial spur one-fifth as long as
lateral spur and one-third as long as first metatarsal segment (fig.7). Pygidium (fig.5) verti-
cal, subtruncate apically; surface punctulate and with many coarser punctures. Fifth vis-
ible abdominal sternum slightly emarginate in male and rounded in female. Eighth tergite
rounded in male.

Male terminalia (figs.8,9): Median lobe with ventral valve acute apically, lateral
margins incurvate, base broad; dorsal valve subtriangular, lateral margins nearly straight,
apex rounded. Internal sac armature (fig.8) consisting of two basal subconcave, slipper-
shaped sclerites with short irregular protuberances near apex (figs.8D,8d); two subbasal
sclerites, four times as long as wide, subrectangular, slightly sinuate and with serration
directed apicad along a margin ( fig.Sb) near the place of precedent pair and in part overlap-
ping it, two subbasal sclerites, 2.1 times as long as wide at base, subtriangular, very sinuate
and armed with denticles directed basad (fig.8e); two long, laminar, median sclerites,
slightly angulate toward basal and median portions, with one or two rows of basally directed
denticles along middle apical portion; unpaired, median wishbone-shaped sclerite, shorter
than the laminars (1.3 as long as the length of laminars), with strongly incurvate lateral
margins on middle apical portion, rounded apex in lateral view (fig.SC) and distinctly
separate stems; apical sclerite with broad lateral areas and long stems (fig.8). Internal sac
membrane with spines on basal and median portions (figs.8A.8F). Lateral lobes with mod-
erately deep rounded cleft between them (fig.9).

Etymology: The species name refers to the place where the holotype was
collected: Nova Teutonia, Santa Catarina, Brazil.

Type Material: Holotype, male: BRAZIL: Sta.Catarina, Nova Teutonia; 14.11.1944; F.
Plaumann; deposited in the National Museum of Natural History, Washington (NMNH).
Allotype, BRAZIL: Sao Paulo, Ilha da Vitoria, 16-27.111.1964, Exp.Dep.Zool, deposited in
the Museu de Zoologia de Sao Paulo (MZSP); one male paratype with same label as
holotype, deposited in the Museu de Entomologia do Departamento de Zoologia da
Universidade Federal do Parana (DZUP); additional three paratypes. BRAZIL: Mato
Grosso, Chapada dos Guimaraes, April, Acc.No.2966, deposited in the Carnegie Museum
of Natural History, Pittsburg (CARN); Rio Grande do Sul, Sao Leopoldo, 15.X.1982,
CJ.Becker, 60.602, deposited in the Fundacao Zoobotanica do Rio Grande do Sul
(FZB.MCN); PARAGUAY: Sao Bernardino, 27.XII, Amaranthaceae, K. Fiebrig
(NMNH); Depto. Alto Parana, Centra For. Alto Parana, 25 30'S, 54,44'W, 14-16- V-1986.
Pogue & Solis (NMNH).

Vol. 104, No. 4, September & October. 1993

163

BLACK INTEGUMENT
BROWN HAIRS

BLACK INTEGUMENT
BLUISH GRAY HAIRS

BLACK INTEGUMENT
LKSHT YELLOW HAIRS

REDDISH YELLOW INTEGUMENT
LIGHT YELLOW HAIRS

REDDISH YELLOW INTEGUMENT
GOLDEN YELLOW HAIRS

Figs.l-9.Amblycemsteutoniensis, new species. 1. dorsal hahitus; 2. pronotum; 3. lateral view
of pronotum; 4. scutellum; 5, pygidium; 6, antenna; 7, hind trochanter. femur, tibia and first
metatarsal segment; 8. malegenitalia. median lobe: A-spineson basal portion enlarged, b-
subbasal serrate sclerites, c- lateral view of wishbone-shaped sclerite, d- ventral view of
basal sclerite. D- same enlarged, e-subbasalspinoussclerites, F-spines on median portion,
enlarged; 9. legmen.

164 ENTOMOLOGICAL NEWS

DISCUSSION

This species is most closely related to A. canescens (Boheman). Both
share many characters such as body, except pygidium and abdomen,
mottled with brown and bluish gray hairs, postocular lobe long, eyes
finely faceted, lateral carina of pronotum divided, scutellum trilobed,
longitudinal arm of metepisternal sulcus very short, metepisternum and
hind coxa evenly punctured, mesal mesotibial spur about middle length
of lateral spur and one-third length of the first hind tarsal segment.

Amblycerus teutoniensis can be distinguished from ,4. canescens by the
reddish yellow integument of the first four antennal segments, tarsi,
pygidium and abdomen; these parts are entirely black in A. canescens.

The characters in the internal sac of male genitalia are comparable in
these two species. Differences are found in the shape of both pairs of sub-
basal sclerites (short in teutoniensis, longer in canescens; the other sub-
triangular with denticles directed basad in teutoniensis (fig.Se), sub-
rectangular with denticles directed apicad in canescens; in the shape of
the long laminar sclerites (slightly angulate in teutoniensis, strongly
angulate in canescens; and in the wishbone-shaped sclerite (with incur-
vate lateral margins and distinctly separate stems in teutoniensis, nearly
straight and moderately separate stems in canescens).

ACKNOWLEDGMENTS

We would like to thank Renato C. Marinoni for comments on this paper.

LITERATURE CITED

Boheman, C.H. 1833. In Schoenherr, C.J. Genera et species curculionidum cum syn-
onymia hujus familiae: species novae aut hactenus minus cognitae, descriptionibus a
Dom. Leonardo Gyllenhal. C.H. Boheman. et entomologis aliis. Vol. 1(1): 1-385. Paris.

Vol. 104, No. 4, September & October, 1993 165

THE DRAGONFLIES AND DAMSELFLIES

(ODONATA) OF BUCK CREEK,
PULASKI COUNTY, KENTUCKY 1

Randall G. Payne^, Guenter A. Schuster^

ABSTRACT: Seven families of Odonata representing 32 species were found to inhabit
Buck Creek, a fifth-order tributary of the upper Cumberland River. Of these 32 species, 17
were new records for Pulaski County. Buck Creek was found to be relatively undisturbed
and of high water quality as indicated by its diverse community of Odonata.

There have been relatively few published surveys of the Odo-
nata from Kentucky. Resner (1970) compiled a list of all known odonate
species occurring in the Commonwealth, adding three species to the list.
The last additions to Kentucky's species list were by Crowley and Wilson
(1979). The current total number of odonate species known for Kentucky
is 138.

Buck Creek, a fifth-order tributary of the upper Cumberland River,
had exceptionally good water quality and a diverse aquatic fauna
(Marker, # al. 1979). Recent studies of fishes (Cicerello and Butler, 1985),
freshwater Unionidae ( Schuster, et al. 1 989), and Trichoptera ( Floyd and
Schuster, 1990) have reported large numbers of species for each of these
groups of organisms. Because Buck Creek was relatively undisturbed, as
reflected in previous studies, it was thought to potentially support a
diverse community of odonates.

STUDY AREA

Buck Creek is located in southcentral Kentucky (37 10' N, 84 30'
W). This stream drains approximately 767 km2 in Lincoln, Pulaski and
Rockcastle counties. It flows southward for 107.2 km and discharges into
the Cumberland River, near Cumberland River km 859. Buck Creek
flows entirely within the Eastern Highland Rim subsection of the Inte-
rior Low Plateau Physiographic Province (Quarterman and Powell,
1978). The surface geology is composed primarily of Mississippian Age
limestone (Schuster, et al. 1989).

The land use in the watershed is primarily agricultural. The upper

1 Received February 25, 1993. Accepted March 8. 1993

2 219 Cherokee Trail, Somerset, Kentucky 42501

3 Department Of Biological Sciences, Eastern Kentucky University. Richmond. Ken-
tucky 40475

ENT. NEWS 104(4): 165-170, September & October. 1993

166

ENTOMOLOGICAL NEWS

NORTH

SCALE

BRUSHY CREEK

KM 5 10 15 20

CUMBERLAND RIVER

Figure 1 . Location of collecting sites along the mainstem of
Buck Creek, Pulaski County, Kentucky (after Butler, 1985).

one-half of the watershed is utilized for crop production and the remain-
der of the watershed lies within the boundary of Daniel Boone National
Forest. The stream has numerous braids that become isolated pools dur-
ing the drier times of the year.

Vol. 104. No. 4, September & October, 1993 167

MATERIALS AND METHODS

Six collecting sites were chosen on the mainstem of Buck Creek
(Figure 1). Sites one and two were located on the fourth-order section of
the stream, and sites three through six were on the fifth-order segment.
Exact localities of each collection site are given in Table 1 . Two sites were

Table 1. Location of collecting sites on Buck Creek, Pulaski County, Kentucky.
Site Location

1 State Route (SR) VOcrossing, approximately 4.9 km WNW of Woodstock and 19.4 km
N of Somerset.

2 SR 39 crossing, approximately 3.2 km S of Woodstock and 15.9 km NE of
Somerset.

3 SR 1677 crossing, approximately 2.2 km W of Dahl and 13.4 km ENE of
Somerset.

4 SR 1675 (old SR 80) crossing at Stab, approximately 14.6 km ENE of
Somerset.

5 SR 1003 crossing, approximately 7.2 km S of Stab and 14.6 km E of
Somerset.

6 SR 192 crossing, approximately 4.9 km NW of Mt. Victory and 15.8 km
ESE of Somerset.

visited per week, and a collecting circuit of all sites was completed every
three weeks. Collection of adults began in June and continued through
October 1991 and from April to mid-September 1992. Adults were col-
lected using a D-frame net and a .22 caliber rifle loaded with 1/15 oz,
number 12 shot shotshell. During the second collecting season a light
weight aerial net was used for collection. Representatives of each species
collected were placed in the Natural History Museum of Eastern Ken-
tucky University or in the first author's collection.

RESULTS

Five families of Anisoptera (Aeshnidae, Corduliidae, Gomphidae,
Libellulidae and Macromiidae) and two families of Zygoptera (Calop-
terygidae and Coenagrionidae) were found at Buck Creek (Tables 2 and
3 respectively), including 19 species of Anisoptera (Table 2) and 13
species of Zygoptera (Table 3). This is approximately 25% of the 138
species of Odonata known to occur in Kentucky. Of these 32 species, 17
were new records for Pulaski County, 14 anisopterans (Table 2) and three
zygopterans (Table 3).

168 ENTOMOLOGICAL NEWS

Table 2. Anisoptera collected at Buck Creek, Pulaski County, Kentucky, (June -November
1991; April - mid-September 1992): * = new county record; A=adult; N = nymph.

Aeshnidae

Basiaeschna Janata (Say) A

Boyeria vinosa (Say) A

Corduliidae

Epitheca princeps (Hagen) * A

Neurocordulia yamaskanensis Provancher * A

Somatochlora linearis (Hagen) * N

Gomphidae

Dromogomphus spinosus Selys A

Gomphus (Gomphus) lividus Selys * A

Gomphus (Gomphurus) lineatifrons Calvert * A

Gomphus (Hylogomphus) viridifrons Hine * A

Hagenius brevistylus Selys * A

Stylogomphus albistylus (Hagen) * A

Libellulidae

Libellula luctosa Burmeister * A
Libellula lydia (Drury) A

Libellula pulchella Drury A

Erythemis simplicicollis (Say) * A
Pachydiplax longipennis (Burmeister) A

Sympetrum vicinum (Hagen) A

Macromiidae

Didymops transversa (Say) * A

Macromia alleghaniensis (Williamson) * A

Table 3. Zygoptera collected at Buck Creek, Pulaski County, Kentucky, (June - November
1991; April - mid-September 1992): * = new county record; A= adult.

Calopterygidae

Calopteryx maculata (Beauvois) A

Hetaerina americana (Fabricius) A

Coenagrionidae

Argia fumipennis violacea (Hagen) A

Argia moesta (Hagen) A

Argia sedula (Hagen) A

Argia tibialis Rambur * A

Vol. 104, No. 4, September & October, 1993 169

Table 3. (Continued)

Argia translata Hagen A

Enallagma basidens Calvert * A

Enallagma civile (Hagen) A

Enallagma divagans Selys * A

Enallagma exsulans (Hagen) A

Ischnura posita (Hagen) A

Ischnura verticalis (Say) A

DISCUSSION

Kentucky is near the northern or southern limits of distribution of
many odonates. Montgomery (1967) used Peterson's Resemblance
Equation to indicate the degree of similarity or difference of odonates in
the North Central States. The similarity of the species of Enallagma of
Kentucky and southern Indiana was 0.35, while between Kentucky and
Tennessee it was 0.79, where a value of 1.0 indicates total similarity.

Buck Creek was found to possess a very diverse odonate community.
The assemblage of a diverse community of Gomphidae was indicative of
a relatively undisturbed habitat. One gomphid, Gomphus lineatifrons
Calvert, has been found to inhabit only streams of high water quality (S.
W. Dunkle, pers. comm.; Roback and Westfall, 1967), Carle (1979) re-
ported that of the Anisoptera in Virginia, 75% of those that were rare
inhabited relatively undisturbed lotic environments. While Buck Creek
is relatively undisturbed, certain perturbations such as clear cutting and
gravel removal had occurred during the course of this study. Continued
monitoring of Buck Creek is encouraged. Water quality assays and
periodic surveys of macroinvertebrates should continue in order to
detect any deleterious practices in and around this stream.

ACKNOWLEDGMENTS

We thank D. L. Batch and J. R. Maki of Eastern Kentucky University, and two
anonymous reviewers for their comments on this manuscript. We thank S. W. Dunkle for
verification of many species, and for helpful suggestions. Thanks go to M. J. Westfall. Jr. for
suggestions concerning curation methods. We also thank S. L. Jones for providing equip-
ment and financial support during this study.

LITERATURE CITED

Butler, R. S. 1985. Comparative feeding ecology of darters (Percidae: Etheosioma) in Buck
Creek, Pulaski County, Kentucky. M. S. Thesis. East. Ky. Univ.. Richmond. 247 pp.

Carle, F. L. 1979. Environmental monitoring potential of the Odonata, with a list of rare
and endangered Anisoptera of Virginia, United States. Odonatologica 8(4): 319-323.

Cicerello, R. R. and R. S. Butler. 1985. Fishes of Buck Creek, Cumberland River
Drainage, Kentucky. Brimleyana 11: 133-159.

170 ENTOMOLOGICAL NEWS

Crowley, P. H. and A. D. Wilson. 1979. New species records of damselflies (Odonata:

Zygoptera) in Kentucky. Trans. Ky. Acad. Sci. 40: 52.
Floyd, M. A. and G. A. Schuster. 1990. The caddisflies (Insecta: Trichoptera) of the Buck

Creek System, Pulaski County, Kentucky. Trans. Ky. Acad. Sci. 51: 127-134.
Marker, D. F., Jr., S. M. Call, M. L. Warren, Jr., K. E. Camburn, and P. Wigley. 1979.

Aquatic biota and water quality survey of the Appalachian Province, eastern Kentucky.

Tech. Rep. Ky. Nat. Pres. Comm., Vol. 1, Frankfort, KY 1152 pp.
, M. L. Warren, Jr., K. E. Camburn, S. M. Call, G. J. Fallo, and P.

Wigley. 1980. Aquatic biota and water quality survey of the upper Cumberland River

basin. Tech. Rep. Ky. Nat. Pres. Comm., Frankfort, KY. 683 pp.
Montgomery, B. E. 1967. Geographical distribution of the Odonata of the North Central

States. Proc. N. Central Branch Ent. Soc. Amer. 22: 121-129.

Quarterman, E. and R. L. Powell. 1978. Potential ecological/geological natural land-
marks on the Interior Low Plateau. U. S. Dept. Interior, Washington, D. C. 738 pp.
Resner, P. L. 1970. An annotated check list of the dragonflies and damselflies (Odonata)

of Kentucky. Trans. Ky. Acad. Sci. 31: 32-44.
Roback, S. S. and M. J. Westfall, Jr. 1967. New records of Odonata nymphs from the

United States and Canada with water quality data. Trans. Am. Ent. Soc. 93: 101-124.
Schuster, G. A., R. S. Butler and D. H. Stansbery. 1989. A survey of the unionids

(Bivalvia: Unionidae) of Buck Creek, Pulaski County, Kentucky. Trans. Ky. Acad. Sci.

50: 79-85.

Vol. 104, No. 4, September & October, 1993 171

CADDISFLIES (TRICHOPTERA)

OF WILDCAT CREEK,
PICKENS COUNTY, SOUTH CAROLINAi

Michael A. Floyd, John C. Morse^

ABSTRACT: Sixty-two species of caddisflies (Trichoptera) were identified from collec-
tions made from Wildcat Creek over a period of 33 years. A new distributional record for
South Carolina was obtained for Diplectrona metaqui. Eight species, Polycentropus carlsoni,
Wormaldia thyria, Neotrichia collata, Stactobiella delira, Neophylax atlanta. Goera fuscula,
Pseudogoera singularis. and Agarodes griseus, are considered to be threatened in South
Carolina. Psilotretafrontalis should be removed from the list of threatened species in South
Carolina. The diverse caddisfly fauna of Wildcat Creek adds further support to the recom-
mendation by other authors for its use as a biodiversity reference stream.

The aquatic insect fauna of Wildcat Creek has been well documented
for mayflies (Ephemeroptera), stoneflies (Plecoptera), and other signifi-
cant biota (Westfall 1947; McCaskill 1967, unpub. thesis, Clemson Univ.,
1973, unpub. dissertation, Clemson Univ.; McCaskill and Prins 1968;
Carlson 1971, unpub. thesis, Clemson Univ.; White et al. 1979; Stark
1983; Adler 1987; and Daniels and Morse 1992). Because of its potential
use as a biodiversity reference stream, as indicated by the high diversity
of Ephemeroptera and Plecoptera and the occurrence of several other
rare or unique plant and animal species (Daniels and Morse 1992), an
additional faunistic study was undertaken to document the caddisfly
(Trichoptera) fauna.

Wildcat Creek is a second order tributary of Six Mile Creek in
southwestern Pickens County, South Carolina (Fig. 1). It is a clear
stream with a gravel and sand substrate and a riparian zone composed of
mixed hardwoods. Wildcat Creek lies within the Piedmont Physio-
graphic Region and drains approximately 204 ha (504 ac), 47% (96 ha or
236 ac) of which lies within the Clemson University Experimental Forest
(CUEF). The portion of the watershed within the CUEF has been pro-
posed as a Registered Heritage Site as part of the Heritage Trust Program
of the South Carolina Wildlife and Marine Resources Department (Fig.
1). This designation would minimize the effects of anthropogenic distur-
bances such as clearcutting, private development, or sedimentation. A
more detailed description of the watershed, including its management

1 Received February 13, 1993. Accepted March 29, 1993

2 Clemson University, Department of Entomology, Long Hall, Box 340365, Clemson, SC
29634-0365

ENT. NEWS 104(4): 171-179, September & October, 1993

172

ENTOMOLOGICAL NEWS

Pleasant Hill
Bell-Hupp Ln. \. Church Rd.

Clemson

University

Experimental

Forest

Clemson

University

Experimental

Forest

(CUEF)

To Six Mile

To Clemson

_ _ _ Proposed Registered Heritage Site
(SC Heritage Trust Program)

1 Km

1 Mi

Fig. 1. Map of Wildcat Creek and surrounding area.

Vol. 104, No. 4, September & October, 1993 173

by the Clemson Univesity Forestry Department and its biological,
educational, and recreational importance, was provided by Sorrells
(1984) and Daniels and Morse (1992).

MATERIALS AND METHODS

The 3980 specimens of Trichoptera examined in this study are
housed within the Clemson University Arthropod Collection (CUAC).
Many of these specimens have been collected since 1956 by students
from the Clemson University Aquatic Insects course (ENT 469/669) or
Taxonomy of Immature Insects course (ENT 808). Other collections
have been made by Carlson ( 1 97 1 , unpub. thesis, Clemson Univ.), Hoff-
man and Morse (1990), and by the authors. Collections of larvae have
been made by qualitative benthic collecting, while adults have been
obtained by a variety of methods. Carlson ( 197 1 , unpub. thesis, Clemson
Univ.) used a modified emergence trap composed of parachute-netting
which was stretched across the stream, touching either the water or
ground on all four sides. Malaise traps and ultraviolet light traps were
used by Hoffman and Morse (1990) and the authors. The authors per-
formed or verified all identifications.

RESULTS AND DISCUSSION

Sixty-two species of caddisflies, representing 17 families, were iden-
tified from collections obtained from Wildcat Creek (Table 1, Page 176).
A new distributional record for South Carolina was obtained for Diplec-
trona metaqui Ross. Eight species of Wildcat Creek Trichoptera, Polycen-
tropus carlsoni Morse, Wormaldia thria Denning, Neotrichia collata
Morton, Stactobiella delira (Ross), Goera fuscula Banks, Pseudogoera
singularis Carpenter, Psilotreta frontalis Banks, and Agarodes griseus
Banks, were designated as threatened by the Invertebrate Taxa Review
Committee of the South Carolina Heritage Trust Program (Morse et al.
1979, and unpublished data). Unless otherwise noted, distributional and
habitat information listed below were taken from Morse et al. (1979).

Polycentropus carlsoni is known from only two states, Alabama and
South Carolina. In Alabama it has been reported from two locations in
Calhoun County (Harris et al. 1 99 1 ). South Carolina records include two
locations in the CUEF (Wildcat Creek, the type locality for this species,
and Indian Creek, a first order tributary of Six Mile Creek, Lake Issa-
queena [Adler 1992, unpub. dissertation, Clemson Univ.]) and a spring-
brook 10 km south of Clemson in Pendleton, Anderson County (Hoff-
man and Morse 1990). Each of these collections is from first order, clear.

174 ENTOMOLOGICAL NEWS

cold streams (Hoffman and Morse 1990). The United States Department
of Interior (1984) listed P. carlsoni as Category 2 (possibly endangered or
threatened, but lacking conclusive data). With the exception of two
males captured by ultraviolet light traps in Alabama, this species has
been captured solely with the use of modified emergence traps or
Malaise traps (Hoffman and Morse 1990).

Wormaldia thyria has been found at no other locality in South Car-
olina except Wildcat Creek. It has been reported from four other states,
Alabama (Frazer etal 1991), North Carolina (Denning 1950), Tennessee
(Etnier and Schuster 1 979), and Virginia (Parker and Voshell 1 98 1 ). It is
found in small, clear, cold streams in the Mountains and Upper
Piedmont.

Neotrichia collata is known from only one other locality in South Car-
olina, a tributary of Brasstown Creek in Oconee County. It is re-
ported to inhabit small, cold, rocky, rapidly flowing streams and has
been found in seven other states: Alabama (Harris et al. 1983), Illinois
and Kentucky (Ross 1944), Maine (Blickle 1979), New York (Morton
1905), Utah (Baumann and Unzicker 1981), and Vermont (Harris et al.
1991).

Stactobiella delira has been reported from 26 states and one Canadian
Province (a distribution including British Columbia and California
extending east to Maine and South Carolina). However, it is known from
only two localities in South Carolina, Wildcat Creek and South Fork of
the Saluda River, both of which are clear, cold, rocky streams.

Goera fuscula is known from only two other South Carolina streams,
the Chattooga River and a tributary of Brasstown Creek (both in Oconee
County). It requires cold, rocky, upland streams and has been reported
from Georgia (Schmid 1983), Massachusetts, New York, Tennessee, and
Virginia (Flint 1960), Maine (Mingo and Gibbs 1980), North Carolina
(Banks 1905), and Quebec (Roy and Harper 1979).

Pseudogoera singularis has been reported in South Carolina from only
two other streams, Toxaway Creek and Yellow Branch, both of which lie
in Oconee County. The larvae and pupae inhabit moss-covered water-
falls in small, clear, relatively unpolluted streams at elevations of 245 to
825 m (800 to 2700 ft.) (Wallace and Ross 197 1 ). It has been reported from
Georgia (Wallace and Ross 1971), North Carolina (Carpenter 1933), and
Tennessee (Etnier and Schuster 1979).

Psilotreta frontalis should no longer be considered threatened in
South Carolina because its distribution in the state now has increased to
six counties: Aiken, Greenwood, Lexington, Oconee, Pickens, and
Saluda (Floyd, unpub. data). It has been reported from 1 8 other states in
the eastern United States (Parker and Wiggins 1987).

Vol. 104, No. 4. September & October. 1993 175

Argores griseus has been reported from 20 states in the eastern United
States (Harris et al 1983, Harris et al. 1991, Parker and Wiggins 1987,
Schmid 1983). It is found in depositional areas of small spring seepages
and spring streams in three localities in the upper Piedmont of South
Carolina. These include Wildcat Creek, Indian Creek (Adler 1992,
unpub. dissertation, Clemson Univ.), and a springbrook in Pendleton,
Anderson County.

Although not designated by Morse et al. ( 1 979) as threatened in South
Carolina, one additional species, Neophylax atlanta Ross, undoubtedly
deserves such a designation. It has been reported from four states,
Alabama, Georgia, Virginia, and South Carolina (Ross 1947,Harrise/a/.
1991) but has been found at only two sites, Indian Creek (Adler 1992,
unpub. dissertation, Clemson Univ.) and Wildcat Creek, in South
Carolina.

Wildcat Creek also is a paratype locality for the subspecies Cheu-
matopsyche harwoodi enigma Ross, Morse, and Gordon ( 197 1 ). In South
Carolina it is known additionally from South Saluda River and Eastatoe
Creek in Pickens County, as well as Thompson River, Coley Creek, and
Bearcamp Creek in the mountains of Oconee County (Morse etal. 1 989).
Outside South Carolina it has been reported from cold, rocky Piedmont
and Mountain streams in Arkansas (Bowles and Mathis 1989), Georgia
(Ross, Morse, and Gordon 1971), North Carolina (Gordon 1974), and
Virginia (Parker and Voshell).

Because of the high diversity of such orders as mayflies and stone-
flies, as well as the presence of other rare and unique aquatic insects,
Daniels and Morse (1992) suggested that Wildcat Creek be used as a
biodiversity reference stream. The diverse caddisfly fauna detailed in
this study serves to reinforce this recommendation. Furthermore, the
entire watershed should be protected and managed to maintain its high
diversity of plants and animals and thus preserve its potential use for
education, recreation, and research.

ACKNOWLEDGMENTS

We thank the many former students of Clemson University who made collections from
Wildcat Creek over the past 33 years. Special thanks go to Paul H. Carlson and Kevin M.
Hoffman for supplying a large quantity of adult specimens from their respective studies.
Additional adult and larval specimens were provided by Cindy R. L. Adler. David Scar-
borough helped prepare Figure 1. Useful reviews of the manuscript were provided by Peter
H. Adler and John A. Durant. This is Technical Contribution no. 3370 of the South Car-
olina Agricultural Experiment Station, Clemson University.

17 6 ENTOMOLOGICAL NEWS

Table 1. Caddisflies (Trichoptera) of Wildcat Creek, Pickens County, South
Carolina.

Classification follows that of Weaver and Morse (1986). Stages of development (larva,
pupa, and adult) by which species are represented are indicated by L, P, and A, respectively.
Stages of development are followed by number of specimens in that particular stage. Dates
refer to capture times of adult specimens. New distributional records for South Carolina
are indicated by the symbol (#). Threatened species as listed by the invertebrate Taxa
Review Committee of the South Carolina Heritage Trust Program (Morse et al. 1979) are
indicated by an asterisk (*). An additional species not listed as threatened by Morse et al.
(1979), but deserving such designation, is indicated by a plus symbol (+)

Suborder Annulipalpia
Infraorder Curvipalpia
Superfamily Hydropsychoidea
Family Hydropsychidae

Ceratopsyche sparna (Ross), L (4), A (14 F, 6 M), 10 Apr.-29 June.

Cheumatopsyche harwoodi enigma Ross, Morse, & Gordon, A (87 F, 48 M),
8 Apr.-26 Sep.

Cheumatopsyche pettiti (Banks), A (17 F, 6 M), 18 Apr.-22 Aug.

Cheumatopsyche pinaca Ross, A (39 F, 9 M), 13 Apr.-22 Aug.
#Diplectrona metaqui Ross, L (1).

Diplectrona modesta Banks, L (25), A (22 F, 15 M), 4 Apr.-8 Sep.

Hydropysche betteni Ross, A (17 F, 4 M), 17 Apr.-30 Jul

Parapsyche cardis Ross, L (14), A (2), 21 May.

Potamyia flava (Hagen), L (2).
Family Polycentropodidae

Cymellus marginalis (Banks), A (5 F), 19-21 Jun.

Nyctiophylax nephophilus Flint, A (2 M), 21 May.

Phylocentropus lucidus (Hagen), A (1 F), 22 Aug.

Phylocentropus placidus (Banks), A (1 F), 21 May.

Polycentropus blicklei Ross & Yammamota, A (14 F, 27 M), 14 Apr. -25 Sep.

* Polycentropus carlsoni Morse, A (5 F, 3 M), 15 Apr.-4 Jul.
Polycentropus cinereus Hagen, A (22 F, 26 M), 22 Apr.-4 Sep.
Polycentropus confusus Hagen, A (3 M), 20 Apr.
Polycentropus maculatus Banks, A (3 F), 15 Apr.-12 May.

Family Psychomyiidae

Lype diversa (Banks), A (212 F, 1220 M), 10 Apr.-14 Oct.
Psychomyia flavida Hagen, A (460 F), 20 Apr. -4 Sep.

Superfamily Philopotamoidea
Family Philopotamidae

Chimarra aterrima (Hagen), A (13 F, 10 M), 4 Apr.-20 Jun., 6-13 Oct.

Chimarra obscura (Walker), A (1 F), 21 May.

Dolophilodes distinctus (Walker), L (9), P (4), A (64 F, 68 M), 10 May-8 Nov.

Dolophilodes major (Banks), A (1 F), 21 May.

Wormaldia moesta (Banks), A (1 1 F, 18 M), 20 Apr.-7 Oct.

* Wormaldia thyria Denning, A (1 M), 22 Jul.

Infraorder Spicipalpia
Superfamily Hydroptiloidea
Family Glossosomatidae

Agapetus iridis Ross, A (328 F, 144 M), 20 Apr.-13 Aug.

Glossosoma nigrior Banks, L (1), A (51 F, 25 M), 19 Mar.-23 Sep.
Family Hydroptilidae

Hydroptila amoena Ross, A (17 M), 19 Apr.-18 Sep.

Vol. 104, No. 4, September & October, 1993 177

Hydroptila gunda Milne, A (27 M), 10 Apr.-21 May.
Hydroptila quinola Ross, A (6 M), 19 Apr.-27 Jun.

* Neotrichia collata Morton, A (2 M), 12-20 Jun.

* Stactobiella delira (Ross), A (2 F, 1 M), 10-20 Apr.

Superfamily Rhyacophiloidea
Family Rhyacophilidae

Rhyacophila Carolina Banks, L (4), P (1), A (74 M), 10 Apr- 19 Sep.

Rhyacophila juscula (Walker), L (5), A (2 F, 5 M), 18 May-21 May, 31 Aug.- 18 Sep.

Rhyacophila glaberrima Ulmer, L (1), A (35 F, 59 M), 24 Apr.-4 May, 21 Aug.-26
Oct.

Rhyacophila minor Banks, P (1), A (1 M), 15-22 May.

Rhyacophila nigrita Banks, A (21 M), 23 Apr. -26 May, 6 OcL

Rhyacophila teddyi Ross, A ( 1 M), 22 May.

Rhyacophila torva Hagen, L (6), A (25 M), 4 Apr.-24 Sep.

Suborder Integripalpia
Infraorder Plenitentoria
Superfamily Limnephiloidea
Family Limnephilidae

* Goerafuscula Banks, L (3), A (6 F, 3M), 20 Apr.-27 Jun.
Pycnopsyche antica (Walker), L (15), A (5 F, 4 M), 4 Sep.-8 Nov.
Pycnopsyche gentilis (MacLachlan), L (45), P (8), A (13 F, 13 M), 12 Sep.-14 Oct.
Pycnopsyche guttifer (Walker), L (7).

Pycnopsyche luculenta (Betten), L (23), A (8 F, 4 M), 26 Sep.-8 Nov.
Family Uenoidae

+Neophylax atlanta Ross, A (2 M), 1 1-14 Oct.

Neophylax mitchelli Carpenter, L (4), A (6 M). 1 1-14 Oct.

Neophylax oligius Ross, A (13 M), 1-14 Oct.
Family Brachycentridae

Brachycentrus nigrosoma (Banks), L (1).

Micrasema sp., A (1 F), 8-15 May
Family Lepidostomatidae

Lepidostoma latipenne (Banks), A (83 F, 61 M), 15 Apr.-8 Nov.

Lepidostoma Ontario Ross. A (1 F. 1 M). 20 Apr, 17 Jun.

Infraorder Brevitentoria
Superfamily Leptoceroidea
Family Odontoceridae

* Pseudogoera singularis Carpenter, P (2), A (2 F, 2 M), 31 Aug.- 18 Sep.

* Psilotreta frontalis Banks. L (17). A (57 F. 93 M). 19 Apr.-21 Jun.. 8-15 Aug.
Family Calamoceratidae

Anisocentropus pyraloides (Walker), L (8), P (1). A (12 F. 31 M). 17 May-8 Aug.

Heteroplectron americanum (Walker), L (1), A (5 F), 20 Apr-22 May.
Family Leptoceridae

Ceraclea protonepha Morse & Ross, A (3 F, 3 M). 20 Apr. 21 May.

Oecetis inconspicua (Walker). A (10 F. 4 M), 20 Apr.-27 Jun. 22 Aug.-18 Sep.

Triaenodes ignitus (Walker). A (4 F. 7 M). 19 Apr.-22 Aug.
Family Molannidae

Molanna blenda Sibley. A (1 1 F. 10 M), 19 Apr.-24 May. 22 Jun.-23 Sep.

Superfamily Sericostomatoidea
Family Helicopsychidae

Helicopsyche paralimnella. L (2).
Family Sericostomatidae

* Agarodes griseus Banks. A (2 F, 4 M), 5-24 May.

178 ENTOMOLOGICAL NEWS

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(Trichoptera). Ann. Entomol. Soc. Amer. 64: 890-894.
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180 ENTOMOLOGICAL NEWS

ABUNDANCE AND SEASONAL ACTIVITY

OF EUCINETOIDEA (COLEOPTERA)

IN A RASPBERRY PLANTATION

AND ADJACENT SITES
IN SOUTHERN QUEBEC (CANADA) 1

Claire Levesque, Gilles-Yvon Levesque^

ABSTRACT: In a raspberry plantation and adjacent sites in southern Quebec, we col-
lected adult Eucinetoidea with unbaited pitfall traps and flight interception traps, from
early May through late October in 1987, 1988 and 1989. A total of 350 Clambidae (three
species), 33 Eucinetidae (four species) and 48 Scirtidae (five species) were collected. The
most common species at the soil surface of the raspberry plantation was Clambus arma-
dillo. The most abundant species in flight traps were Clambus howdeni, C. armadillo,
Eucinetus testaceus and Cvphon variabilis. Adults of C. armadillo, C. howdeni and C. variabilis
were active mainly in spring and/or early summer and probably overwintered as adults;
whereas Eucinetus testaceus flew mainly in August and probably overwintered as larvae.

Although world production of raspberries has increased about 5%
each year over the past decade (Pritts 1991 ), little is known on variations
in beetle communities associated with raspberry (Rubus idaeus L.) plan-
tations, particularly for mycophagus insects. These ecosystems appear
as rather stable sites because plants of many raspberry cultivars, with
perennial root systems and biennial canes, may be kept over about ten
years. Over a three-year period (1987-1989), we collected nearly 60,000
beetles in a raspberry plantation and adjacent sites in southern Quebec.
We have already presented results for the Nitidulidae and Elateroidea
(Levesque and Levesque 1992, 1993). We now present results on the
abundance and seasonal activity of adult Clambidae, Eucinetidae and
Scirtidae ( = Helodidae). These three families of Eucinetoidea are of no
known economic importance and little is known of their life history,
ecology and behavior. Wheeler and Hoebeke (1984) reviewed published
records of hosts and microhabitats of species of Clambidae and Euci-
netidae, families considered mainly mycophagous. However, no scirtids
are known to be associated with fungi (Wheeler and Hoebeke 1984). All
known larvae of Scirtidae are aquatic, phytophagous or detritivorous,
and are common in small lentic habitats and shallow areas of peatland
pools and marches. Scirtid adults are terrestrial and common along the
margins of pools or on emergent plants (Larson 1987).

1 Received February 6, 1993. Accepted March 26, 1993.

2 291 rue des Diamants, Fleurimont, Quebec, Canada JIG 4A1.

ENT. NEWS 104(4): 180-186. September & October, 1993

Vol. 104, No. 4, September & October, 1993 181

MATERIALS AND METHODS

Beetles were collected from early May through late October on a
monocultural raspberry farm at Johnville, near Sherbrooke, in south-
ern Quebec. We sampled from the Boyne cultivar in this conventionally
cultivated plantation (about 7 ha, on sandy soil).

Ground surface-active beetles were caught with pitfall traps at the
following sites: 1) a raspberry row planted in 1978 (old plants), 2) a
raspberry row planted in 1985 (young plants), 3) a woods-field boun-
dary (boundary), and 4) an adjacent wooded site dominated by eastern
white pine, Pinus strobus L. (pine woods). Pitfall traps consisted of glass
jam jars (450 ml, 6.5 cm diameter at the top) partially filled with 100 ml of
4% formalin. Traps were inserted into the soil beneath the canopy as
close to the cane of raspberry plants as possible at the first two sites. At
each site, 20 traps were set in a row (5 m apart) and were emptied
weekly.

In addition, we studied beetles flying close to the ground with flight
interception traps at four sites: 1 ) an open site near the center of the plan-
tation (A), about 20 m from old plants; 2) an open site near a pond (B),
about 5 m from young raspberry plants; 3) a woods-field boundary (C);
and 4) a pine woods (D). These traps were not located between rows of
raspberry plants because of grower's activities and public access during
harvest. Flight traps were modified from the large-area "window" trap
design promoted by Peck and Davies ( 1980). Each consisted of a gray 1 .5
mm mesh window screen (1.22 m height, 1.52 m width, about 1.85 m 2 of
surface) fastened to a wooden frame. The frame itself was suspended by
two lateral triangular wooden supports (1.83 m at the base, 1.25 m
height), 2-4 cm over a set of two galvanized metal pans (25 by 61 cm at the
top, 7.5 cm deep) which were placed directly on the ground. The insects
were caught in the pans partially filled with 2% formalin into which a few
drops of detergent were added. We installed one flight trap at each site;
however, the trap was operated in 1988 and 1989 only in the pine woods
(D). Samples were collected twice a week and the formalin solution was
replaced at each collection; the trap counts from individual sampling
periods were combined on a weekly basis.

Levesqueand Levesque( 1992) presented more detailed information
about the sampling methods and study sites, including a sketch-map of
the raspberry farm.

RESULTS AND DISCUSSION

Abundance of Eucinetoidea Catches. We collected a total of 350
Clambidae( three species), 33 Eucinetidae (four species) and 48 Scirtidae

182

ENTOMOLOGICAL NEWS

(five species). Voucher specimens of some species are deposited in the
Canadian National Collection (Ottawa).

Pitfall trapping resulted in the collection of 262 Clambidae (two
species), 3 Eucinetidae (two species) and 1 Scirtidae (Table 1 ). Almost all
eucinetoids represented Clambus armadillo (DeGeer) (97% of catches).
Adults of this species were collected in similar numbers in rows of old
and young raspberry plants; the explanation of this result and possible
association of these adults with fungi on R. idaeus are unknown. Clambus
armadillo was also caught in large numbers at the ground surface of a wet
meadow in southern Quebec on clay soil (Levesque and Levesque 1987).

Table 1. Total catches of Eucinetoidea species in pitfall traps (1987-1989) at Johnville,
Quebec.

Family and species

Old Young Boundary Pine Total
plants plants woods

Clambidae

Clambus armadillo (DeGeer)
Clambus howdeni Endrody-Younga

Eucinetidae

Eucinetus morio LeC.
Eucinetus oviformis LeC.

Scirtidae

Cyphon variabilis (Thunberg)

131 122

2 -

5

1

1

2

258
1 4

1

2

Total catches of eucinetoids in flight traps comprised 88 Clambidae
(three species), 30 Eucinetidae (four species) and 47 Scirtidae (five
species) (Table 2); all the species monitored at Johnville were collected in
flight traps.

Adults of Clambus howdeni Endrody-Younga flew chiefly at the
woods-field boundary and in the pine woods, while Clambus armadillo
was mainly captured in the three flight traps (A, B and C) near the
raspberry plants (Table 2). Eucinetus testaceus LeC. was the most abun-
dant eucinetid species and flew chiefly in the pine woods (Table 2). In two
mixed deciduous-coniferous forest sites in New Hampshire, Chandler
(1991) observed the flight activity of Clambus howdeni and four species of
Eucinetus (chiefly E. morio LeC.). Eucinetus morio was a slime-mold asso-
ciate, whereas C. howdeni and the other three species of Eucinetus (E.
oviformis LeC., E. punctulatus LeC., and E. testaceus) were associated with
Basidiomycete fruiting bodies (Chandler 1991).

Vol. 104, No. 4, September & October, 1993

183

Table 2. Total catches of Eucinetoidea species in flight traps (1987-1989) at Johnville,
Quebec.

Family and species

Open Open
site site

near near

center pond

(A) (B)

Boundary

(C)

Pine Total
woods a

(D)

Clambidae

Clambus armadillo (DeGeer)

Clambus gibbulus (LeC.)

Clambus howdeni Endrody-Younga

Eucinetidae
Eucinetus mono LeC.
Eucinetus oviformis LeC.
Eucinetus testaceus LeC.
Eucinetus sp.

12

10

1

13

2
22

21

3

1

19

36

2
50

4

2

23

1

Scirtidae

Cvphon obscurus (Guerin)

1

4

3

- 8

Cvphon prob. ruficollis (Say)

1

1

Cyphon pusillus (LeC.)

1

1

Cyphon variabilis (Thunberg)

4

18

8

3 33

Prionocvphon limbatus LeC.

4

4

a not sampled in 1987.

The five species of Scirtidae flew in open sites (A and B) and the
boundary site (C). The most abundant species was Cyphon variabilis
(Thunberg), apparently eurytopic, but mainly collected at the open site B
which was near a pond (Table 2). Nilsson (1980) studied the flight activity
of nine scirtid species in the vicinity of a river in northern Sweden, and
found Cypon variabilis to be one of the three most abundant species.

Seasonal Abundance of Four Species of Eucinetoidea. Adults of
Clambus armadillo were caught by pitfall traps from May through Octo-
ber during the three-year study, mainly in May and the first half of June
(Fig. 1 ). They flew from May until early September, but chiefly in May. In
a wet meadow of southern Quebec, the soil-surface activity was observed
in late spring and summer, mainly in June and July (Levesque and
Levesque 1987).

We observed the flight of Clambus howdeni only in May, June and
July during the three-year study (Fig. 2). In New Hampshire, flight
occurred from May through July, but a few adults were also collected in
late summer (Chandler 1991).

184

ENTOMOLOGICAL NEWS

CO
LU

LU
LJJ
CD

1_L

o

QC
LU
CD

Clambus armadillo

1987

10

PUFAUTRAPS

FLIGHT TRAPS

120 150 180 210 240 270 300

| MAY | JUN | JUL | AUG | SEP | OCT |

JULIAN DATE (days)

Fig. 1. Seasonal abundance of Clambus armadillo in pitfall and flight traps at Johnville,
Quebec.

Adults of Cyphon variabilis flew from May through October, but
mainly in May (Fig. 2). In northern Sweden, Nilsson (1980) observed two
flight periods, the first from mid-May until early July, the second in late
summer and autumn; the peak of captures occurred in June. According
to Larson (1987), adults of this species overwinter and reproduce in
spring and early summer.

Adults ofEucinetus testaceus flew from June through September dur-
ing the three-year study, and mainly in August (Fig. 2). We caught a
teneral adult on June 25, 1989. In New Hampshire, adults of E. morio flew
mainly in spring and early summer, and one adult was also caught in
October (Chandler 1991). In the Finger Lakes region of New York, a
breeding population (adults and larvae) of E. oviformis was found in late
summer, in association with a wood-rotting fungus; in addition, mature
larvae and some adults of this species were still present as late as October
18(Hoebekeera/. 1987).

Vol. 104, No. 4, September & October, 1993

185

CO

LLJ

LU
LU
CD

O

or

LU
CD

20-

10-

20-

10

20-

10

Clam bus
howdeni

Cyphon variabilis

V///X

v///\

Eucinetus
testaceus

MAY JUN JUL AUG SEP OCT

Fig. 2. Seasonal abundance ofClambus howdeni, Cyphon variabilis and Eucinetus testaceus,
in flight traps (1987-1989) at Johnvillc Quebec.

Two life-history patterns are observed among northern European
species of Scirtidae: 1) larvae overwinter and reproduction occurs in
midsummer, and 2) adults overwinter and reproduce in spring and early
summer (Larson 1987). According to Nilsson ( 1980), the seasonal flight
pattern of Scirtidae coincide with these two life-history patterns. If such
is the case in other eucinetoids, we suspect that Clambus armadillo and C.
howdeni overwinter as adults, whereas Eucinetus testaceus overwinters
as larvae.

ACKNOWLEDGMENTS

We appreciate the help of J.M. Campbell and J. McNamara (Centre for Land and
Biological Resources Research, Agriculture Canada. Ottawa, Ontario) for identifications

186 ENTOMOLOGICAL NEWS

and confirmations of most species collected in this study. We are grateful to Donald S.
Chandler (University of New Hampshire, Durham), J. McNamara and two anonymous
reviewers for their helpful comments on this manuscript. Also, we thank Michel Couture
and Lucie Labrecque, owners of "La Framboisiere de 1'Estrie, Enr." at Jonville (Quebec).
This study was partially supported by the Fonds F.C.A.R. (Quebec).

LITERATURE CITED

Chandler, D.S. 1991. Comparison of some slime-mold and fungus feeding beetles

(Coleoptera: Eucinetoidea, Cucujoidea) in an old-growth and 40-year-old forest in

New Hampshire. Coleopts. Bull. 45: 239-256.
Hoebeke, E.R., Q.D. Wheeler and R.L. Gilbertson. 1987. Second Eucinetidae-Conio-

phoraceae association (Coleoptera; Basidiomycetes), with notes on the biology of

Eucinetus oviformis LeConte (Eucinetidae) and on two species of Endomychidae. Proc.

Ent. Soc. Wash. 89: 215-218.
Larson, DJ. 1987. Aquatic Coleoptera of peatlands and marshes in Canada. Mem. Ent.

Soc. Can. 140: 99-132.
Levesque, C. and G.-Y. Levesque. 1987. Activite, succession saisonniere et taille de

coleopteres epiges d'un pre du sud du Quebec. Naturaliste Can. 114: 495-506.
Levesque, C. and G.-Y. Levesque. 1992. Epigeal and flight activity of Coleoptera in a

commercial raspberry plantation and adjacent sites in southern Quebec (Canada):

Introduction and Nitidulidae. Great Lakes Ent. 25: 271-285.
Levesque, C. and G.-Y. Levesque. 1993. Abundance and seasonal activity of Coleoptera

Elateroidea in a raspberry plantation and adjacent sites in southern Quebec (Canada).

Coleopts. Bull. 47. (in press)
Nilsson, A.N. 1980. Distribution and seasonal flight-patterns of Helodidae (Coleoptera)

at the northern coast of the province Angermanland, northern Sweden. Ent. Tidskr.

101: 151-153. ( in Swedish with English abstract)
Peck, S.B. and A.E. Davies. 1980. Collecting small beetles with large-area "window"

traps. Coleopts. Bull. 34: 237-239.
Pritts, M.P. 1991. The genus Rubus, pp. 1-2, In M.A. Ellis, R.H. Converse, R.N. Williams

and B. Williamson (eds.), Compendium of raspberry and blackberry diseases and

insects. APS Press, The American Phytopathological Society, St. Paul, Minnesota.
Wheeler, Q.D. and E.R. Hoebeke. 1984. A review of mycophagy in the Eucinetoidea

(Coleoptera), with notes on an association of the eucinetid beetle, Eucinetus oviformis,

with a Coniophoraceae fungus (Basisiomycetes: Aphyllophorales). Proc. Ent. Soc.

Wash. 86: 274-277.

Vol. 104. No. 4. September & October, 1993 187

NEW DISTRIBUTION RECORD

FOR ISCHNOPTERA BILUNATA

(DICTYOPTERA: BLATTELLIDAE) 1

Eric P. Benson, Arthur G. Appel^

ABSTRACT: Occurrence of Ischnoptera bilunata in Alabama is reported for the first time.
This record extends the known distribution of the species in the United States outside of
Florida. Discovery of/, bilunata can be credited, in part, to confusion with the Asian cock-
roach, Blatlella asahinai.

Ischnoptera bilunata Saussure, was first reported in the United States
from Florida in 1990 (Atkinson et al 1990: treated as the Nicaraguan
cockroach, /. bergrothi [Griffini], due to an error in identification). Pre-
viously, the species was known from Brazil, Bolivia, Paraguay and
Argentina (Princis 1969). Recent collections of/, bilunata in Alabama
(Figure 1) extend the known range of this cockroach in the United States
outside of Florida.

In July, 1991, five female and five male cockroaches collected in
Mobile, Mobile County, Alabama were identified as /. bilunata. Indi-
viduals were collected in the evening in a residential yard on grass and
on a patio around lights. In July, 1992, two females and one male /.
bilunata were collected in the evening on a house porch around lights in
Fair Hope, Baldwin County, Alabama. A third collection was made in
the evening of August, 1992, of two female and two male /. bilunata in
Dothan, Houston County, Alabama near outdoor lights of a super-
market. Specimens from these collections are deposited in the Auburn
University Entomological Museum, the Museum of Comparative Zool-
ogy at Harvard University, Cambridge, Massachusetts and with the Ter-
minix Insect Collection in Mobile, Alabama.

Before the collections of/, bilunata, 21 species of cockroaches were
considered established in Alabama (Dakin and Hays 1970, Pratt 1988,
Atkinson et al. 1991). An additional species, Blaberus craniifer Burmeis-
ter, has been listed from Lee County as an adventive (Dakin and Hays
1970). The only representative from the genus Ischnoptera previously
recorded in Alabama is the dark wood cockroach, /. deropehiformis
(Brunner). This species is distributed throughout the state, often in
wooded areas (Dakin and Hays 1970). All specimens of/, deropehiformis

1 Received December 31, 1992. Accepted February 25, 1993.

2 Department of Entomology, Auburn University, Auburn. Alabama 36849-5629.

ENT. NEWS 104(4): 187-190, September & October. 1993

188

ENTOMOLOGICAL NEWS

Figure 1. Collection sites of adult Ischnoptera bilunata Saussure, in Alabama. A: Mobile
Co., Mobile, VII-27-1991 (5 males, 5 females); B: Baldwin Co., Fair Hope, V1I-27-1992 (1
male, 2 females); C: Houston Co., Dothan, VIII- 14-1992 (2 males, 2 females).

Vol. 104, No. 4, September & October, 1993 189

in the Auburn University Entomological Museum were collected be-
tween the dates of April 24 and August 9. Specimens of/, bilunata were
collected between July 27 and August 14.

Both sexes of/, bilunata are active at night and attracted to light
(Atkinson et al. 1990). This description agrees with our observations.
Ischnoptera bilunata also has been recorded as being abundant in open,
moist, grassy vegetation on the margins of streams, rivers and ponds
(Atkinson et al. 1990, 1992). The Alabama collection sites were all adja-
cent to areas that could be considered open and grassy. Further, these
areas had thatch and leaf litter that could retain moisture. However, only
the Mobile site had a water source nearby. This consisted of a drainage
ditch that, depending on rainfall, was intermittently filled with water.

In 1986, the Asian cockroach, Blattella asahinai Mizukubo was re-
ported from the Tampa area of Florida (Brenner et al. 1986). The Asian
cockroach is very similar in appearance to the serious indoor pest, the
German cockroach, Blattella germanica (L.). The adults of both Blattella
species are 10 to 13 mm in length, brownish both ventrally and dorsally,
with two narrow pronotal spots and wings covering the body (Atkinson et
al. 1990). Unlike B. germanica, B. asahinai lives outdoors, often in leaf lit-
ter and grassy areas and readily flies to lighted areas at night during warmer
months (Brenner et al. 1986, 1988). Adult Asian and German cock-
roaches are similar in appearance to /. bilunata. Adult /. bilunata are 1 5 to
18 mm in length, brownish dorsally and black ventrally, with two tri-
angular pronotal spots and wings covering the body (Atkinson et al.
1990). Reports of B. asahinai and corresponding media coverage appar-
ently heightened the awareness of some pest control operators and
homeowners about cockroaches around lighted areas in the summer. In
the past few years, the Alabama Cooperative Extension Service has
received calls each summer about "Asian cockroaches" around porch
lights. These reported cockroaches have turned out to be one of the Par-
coblatta species and, now,/, bilunata. With the similarities \oB. germanica
and B. asahinai, we anticipate additional reports of/, bilunata in the
future.

ACKNOWLEDGMENTS

The authors are indebted to Fred Strickland and Donnie Byrne for being observant and
interested in collecting many of the specimens. We thank Louis Roth and Tom Atkinson
for their assistance in identification of the specimens and review of this manuscript. We
also appreciate the comments and assistance from Wheeler Foshee, Wayne Clark and
Michael Williams in preparing this manuscript.

190 ENTOMOLOGICAL NEWS

LITERATURE CITED

Atkinson, T.H., P.O. Koehler, and R.S. Patterson. 1990. The Nicaraguan cockroach,
our newest immigrant. Pest Management. 9(10): 12-13.

Atkinson, T.H., P.G. Koehler, and R.S. Patterson. 1991. Catalog and atlas of the cock-
roaches (Dictyoptera) of North America north of Mexico. Misc. Publ. Entomol. Soc.
Amer. 78: 1-85.

Atkinson, T.H., J.R. Mangold, and P.G. Koehler. 1992. Two neotropical cockroaches of
the genus Ischnoptera (Dictyoptera: Blattellidae) established in Florida. Florida En-
tomol. 75(1): 109-115.

Brenner, R.J. P.G. Koehler, and R.S. Patterson. 1986. A profile of America's newest
import, the Asian cockroach. Pest Management 5(Nov/Dec): 17-19.

Brenner, R J., R.S. Patterson, and P.G. Koehler. 1988. Ecology, behavior, and distribu-
tion of Blattella asahinai (Orthoptera: Blattellidae) in central Florida. Ann. Ent. Soc.
Amer. 81(3): 432-436.

Dakin, M.E. and K.L. Hays. 1970. A synopsis of Orthoptera (sensu lato) of Alabama. Ala.
Agric. Exp. Stn. Auburn Univ. Bull. 404.

Pratt, H.D. 1988. Annotated checklist of the cockroaches (Dictyoptera) of North America.
Ann. Entomol. Soc. Am. 81(6): 882-885.

Princis, K. 1969. Blattaria: subordo Epilamproidea: fam. Blattellidae, pars 13: pp. 713-
1038. In M. Beier [ed.], Orthopterorum catalogus. Junk, The Hague.

Vol. 104, No. 4, September & October, 1993 191

FIRST RECORD OF

SEPEDOPHILUS CORONADENSIS

(STAPHYLINIDAE) FROM MEXICO. 1

Jose Luis Navarrete-Heredia 2

ABSTRACT: The staphylinid beetle, Sepedophilus coronadensis is reported from Guerrero,
Mexico.

Sepedophilus Gistel is a genus of Tachyporinae (Staphylinidae) and is
considered to be primarily mycophagous because most of the species are
usually found in habitats rich in fungal mycelia (e.g. forest litter, decay-
ing logs, piles of decaying organic matter, etc.) (Hammond 1973;
Campbell 1976). A recent summary and discussion on Sepedophilus'
biology, including information on unidentified Mexican species, is pro-
vided by Newton (1984).

Sepedophilus is represented in Mexico by three named species from
Veracruz: Sepedophilus bicolor (Bnhr.). S. ferrugineus (Bnhr.) and S. sub-
pubescens (Schub.) (Blackwelder 1944: 151, under Conosomus genus).
This number probably is just a small proportion of the species that exist
in Mexico because the genus is widely distributed in temperate and
tropical regions and is usually collected in several kinds of habitats (per-
sonal observations from Guerrero, Hidalgo, Mexico, Morelos, Puebla,
Oaxaca, Veracruz, Yucatan, and from the examination of several speci-
mens deposited in the Field Museum of Natural History, Chicago, IL.

S. coronadensis Campbell is a species included in Campbell's revision
of species of America north of Mexico in the bisignatus group, and is
known from the Santa Rita and the Huachuca mountains of southeas-
tern Arizona. One specimen from Ramsey Canyon was collected by sift-
ing deep layers of wet, oak leaf litter (Campbell 1976).

The purpose of this note is to record for the first time S. coronadensis
from Mexico, and to provide host fungal records for this species.

During several field trips to the mountains of Taxco, Guerrero, Mex-
ico, six specimens (one male and five females) of this species were col-
lected associated with mushrooms ofAmanita rubescens, Cortinarius sp.,
Ganoderma applanatum and Tylopilus sp. (FIRST HOST RECORDS). In
this area, the specimens were found in cloud forest and in oak-pine

1 Received April 13, 1993. Accepted May 3, 1993.

2 Lab. de Morfofisiologia Animal, Fac. de Ciencias, UN AM. Apdo. Postal 21-518
(Coyoacan), 04000 Mexico. D.F.. MEXICO.

ENT. NEWS 104(4): 191-192. September & October, 1993

192 ENTOMOLOGICAL NEWS

forest, between 2 180-2 300 m. Adults of the species have been collected
from May through August (Campbell 1976), but in Guerrero, one male
and one female were collected in October.

MATERIAL EXAMINED: MEXICO: Guerrero, Desviacion a Pto. Oscuro, Bosque de
Pino-Encino, 2 260 m, 3.VII. 1986, J.L. Navarrete col., exAmanita rubescens (19); same data,
except: ex Cortinarius sp. (29); same data, except: Parque El Huizteco, Bosque Mesofilo de
Montana, 2 300 m, 25.VII. 1986, ex Tylopilus sp. (19); same data, except: km lOcarr. a Tetipac,
El Peral, Bosque Mesofilo de Montana, 2 180 m, 25. X. 1986, ex Ganoderma applanatum ( 1 tf ,
19). Specimens are deposited in the Institute de Biologia, UNAM (IBUNAM) and in
my collection.

ACKNOWLEDGMENTS

I would like to thank J.M. Campbell for the specific identification of the species; and
A.F. Newton Jr. and M.K. Thayer for their kind help and hospitality during a recent visit to
the Field Museum of Natural History.

LITERATURE CITED

Blackwelder, R.E. 1944. Checklist of the coleopterous insects of Mexico, Central

America, the West Indies, and South America. Part. 1. U.S. Nat. Mus. Bull. 185: 1-

188.
Campbell, J.M. 1976. A revision of the genus Sepedophilus Gistel (Coleoptera: Staphy-

linidae) of America north of Mexico. Mem. Ent. Soc. Can. 99: 1-99.
Hammond, P.M. 1973. Notes on British Staphylinidae. Part III. The British species of

Sepedophilus Gistel (Conosomus Auctt.). Entomologist's mon. Mag. 108: 130-165.
Newton, A.F., Jr. 1984. Mycophagy in Staphylinoidea (Coleoptera) [pp. 302-353]. In:

Fungus-insect relationships. Wheeler, Q.D. and M. Blackwell (editors). Columbia

University Press, New York.

Vol. 104. No. 4, September & October, 1993 193

GENERA OF BAETIDAE (EPHEMEROPTERA)
FROM CENTRAL AMERICA 1

C. R. Lugo-Ortiz, W. P. McCafferty 2

ABSTRACT: In addition to eight genera of Baetidae(Ephemeroptera) previously authen-
ticated as occurring in Central America, we report or confirm for the first time the
occurrence of Acerpenna Waltz and McCafferty and Baetis Leach. New records of Acer-
penna from Belize, Costa Rica, Honduras, Nicaragua, and Panama indicate the genus is
well established in the Neotropics. Baetis magnus McCafferty and Waltz (rhoduni group) is
newly reported from Costa Rica and Guatemala. The generic placement of Baetis sinuosus
Navas (probably/w.vca/w.v group) is provisionally accepted and thereby confirms the pres-
ence of a second species of Baetis in Central America. Whereas most baetid genera in Central
American have Neotropical affinities, the Baetis spp. appear to have Nearctic affinities.

Eight genera of the mayfly family Baetidae have been correctly repor-
ted from Central America. The biogeography of these genera was treated
by McCafferty et al. (1992). Baetodes Needham and Murphy was first
reported from Central America (Honduras) by Packer (1966). Subse-
quent records include Cohen and Allen ( 1972) (El Salvador and Guate-
mala), Cohen and Allen ( 1978) (Panama), and McCafferty ( 1985) (Belize
and Costa Rica). Callibaetis Eaton was first reported from an unspecified
Central American country by Eaton (1881). Subsequent records include
Eaton (1885) (Guatemala), Packer (1966) (Honduras), and McCafferty
and Lugo-Ortiz (1992) (Nicaragua). Camelobaetidius Demoulin was re-
ported from Costa Rica, El Salvador, Guatemala, Honduras, and Pan-
ama by Traver and Edmunds (1968). Cloeodes Traver was reported from
Panama by Flowers ( 1991 ). Fallceon Waltz and McCafferty was reported
(as Baetis s. lato) from Belize and Costa Rica by McCafferty (1985). Gua-
jirolus Flowers was described from Panama by Flowers ( 1 985). Moribaetis
Waltz and McCafferty was reported from Costa Rica, Guatemala, and
Panama by Waltz and McCafferty (1985). Paradoeodes Day was reported
from Panama by Flowers (1991).

All of the above genera, with the exception of Guajirolus and Mori-
baetis, which are strictly Neotropical, are Panamerican (Nearctic plus
Neotropical). In addition to the above reported Central American rec-
ords, we have seen Cloeodes from Costa Rica and Honduras; Fallceon
from Honduras; Guajirolus from Costa Rica; and Paradoeodes from
Costa Rica and Honduras.

1 Received March 26, 1993. Accepted April 10, 1993.

2 Department of Entomology, Purdue University, West Lafayette. IN 47907.

ENT. NEWS 104(4): 193-197, September & October. 1993

194 ENTOMOLOGICAL NEWS

In their analysis of Mesoamerican mayflies, McCafferty et al. (1992)
showed the Central American baetid fauna generally to have Neotropi-
cal affinities, rather than Nearctic. Although McCafferty et al. ( 1 992) did
not know Paracloeodes from Central America at that time, they predicted
its presence there, which would corroborate a recent Neotropical center
of dispersal for the genus. McCafferty et al. (1992) considered the dis-
tributional and cladistic data regarding Callibaetis to be too inconclusive
for deciphering a biogeographic history for the genus. Neotropical ele-
ments of this Western Hemisphere genus must be better understood
prior to hypothesizing its evolutionary history.

We have discovered one additional genus of Baetidae from Central
America and confirm the presence of another genus previously reported
on the basis of less restrictive generic concepts. Below, we provide these
records and discuss their significance.

Acerpenna Waltz and McCafferty

TheAcerpenna we report from Central America is represented by lar-
vae of two new species to be described elsewhere.

NEW RECORDS. Acerpenna spp. BELIZE. Cayo Prov.: Roaring Creek, VI/20/74, V. Resh,
larvae, PERC (Purdue Entomological Research Collection, West Lafayette, Indiana); (?):
Sibun River, Gracy, 19/VI/74, V. Resh, larvae, PERC. COSTA RICA: Guanacaste prov.:
Rio Tenorio at Finca La Pacifica, E of Panamerican H wy., II/8- 1 1/69, W. P. McCafferty, lar-
vae, PERC; Limon Prov.: Rio Banano, 16km. WSW of Bomba,9.888N/83.167W, 150m.,
111/26/87, Holzenthal, Hamilton, and Heyn, larvae, FAMU (Florida A&M University,
Tallahassee, Florida); San Jose Prov.: Rio parmita Chiquito, Rt. 12, 6.5 km. SW jet. Rt. 2.,
9.703N/83.970W, IV/10/87, HolzenthaL Hamilton, and Heyn. larvae, FAMU. HONDURAS.
Cortes Prov.: Chamalech, Rio Chamel, X/18/64, J. S. Packer, larvae, PERC. NICARA-
GUA: Presa El Clavo, IX/10/80, larvae, PERC.

REMARKS. The genus Acerpenna was erected by Waltz and McCafferty
(\981b) to include A. macdunnoughi(Ide) and A. pygmaea(Hagen). Later,
McCafferty and Waltz (1990) added A. akataleptos (Traver) and A. harti
McDunnough). All of these species occur in North America north of
Mexico, the southernmost range of any being that of A. pygmaea in the
Edwards Plateau in central Texas (McCafferty and Davis 1992; Lugo-
Ortiz and McCafferty, 1993).

The discovery of Acerpenna in the Neotropics is significant, indicat-
ing an additional Panamerican baetid genus. Waltz (pers. comm.) has
seen female adults of Acerpenna from Panama, and we have also found
the genus in South America and. the Greater Antilles, and we expect to
find it in Mexico. Further analysis of the group may indicate that Acer-
penna is yet another Panamerican baetid genus of Neotropical origin.

Vol. 104, No. 4, September & October, 1993 195

Baetis Leach

Our Central American record of Baetis is based on larvae of B.
magnus McCafferty and Waltz. This species is a member of the rhodani
group of species, which in the Nearctic is particularly well represented in
the southwestern United States (Morihara and McCafferty 1979). Baetis
magnus is known as far north as western Nebraska (McCafferty and
Waltz 1986), and we recently have found it throughout Mexico.

NEW RECORDS. Baetis magnus. COSTA RICA. Cartago Prov.: Reserva Tapanti, Rio
Badilla, 9.688N/83.757W, 1640 m., 111/21/87, Holzenthal and Hamilton, larvae, FAMU;
Rio Birris, Rt. 8 ca. 2.5 km (air) NE of San Juan de Chicoa, 9.960N/83.844 W, 2850 m., II/4/
86, Holzenthal, Morse, and Fasth, larvae, FAMU; Puntarenas Prov.: Rio Bellavista, ca. 1 .5
km NW of Las Alturas, 8.951N/82.846W, 1400 m., Wl 5- 17/86. Holzenthal, Heyn, and
Armitage, larvae, FAMU; Rio Sinigri, ca. 2 km (air) S of Finca Helechales, 9.057N/
83.082 W, 720m., 11/21/86, Holzenthal, Morse, and Fasth, larvae. FAMU; Rio Jaba at rock
quarry. 1.4 km (air) W of Las Cruces, 8.79N/82.97W. 1150 m., Vl/14/86, Holzenthal.
Heyn. and Armitage, larvae, FAMU; San Jose Prov.: Parque Nacional Braulio Castillo,
park headquarters. 10.059N/84.017W, 1650 m.. VII/7/86, Holzenthal. Heyn, and Armi-
tage. larvae, FAMU; 12 mi. N of San Isidro del General, Panamerican Hwy., 5200 ft. V1I/20/
62. G. G. Musser, larvae, PERC. GUATEMALA. Solala Prov.: River at Panajachel. 1/16/89.
B. C. Kondratieff, larvae, PERC.

REMARKS. The genus Baetis has been the subject of considerable
recent revisionary work that has resulted in many synonymies and
recombination of species and the recognition of new genera (Morihara
and McCafferty 1979; Waltz and McCafferty 1985, 1987b, 1987c;
McCafferty and Waltz 1990). As pointed out by McCafferty et al. (1992),
no previous records of Baetis s. str. in Central America have been con-
firmed. Navas (1924) described Baetis sinuosus from Costa Rica on the
basis of a male adult. We have not been able to secure Navas' material for
evaluation. Our examination of Navas' description and figure of the
hindwing does, however, indicate a strong probability that it is a true
Baetis. most likely of the fuscatus group. Moreover, it has an apparent
similarity to B. flavistriga McDunnough, a widespread North American
species that we have seen in northern Mexico. We are therefore pro-
visionally confirming the present generic placement of this species.

Although South American species have been described in the genus
Baetis (see Hubbard and Peters 1981; Hubbarde/a/. 1992), those generic
placements are doubtful and have yet to be verified. If the Neotropical
element of Baetis proves to be limited to a sparse representation in Cen-
tral America as indicated, then it apparently is one of the fewexamples of
a Panamerican group whose Neotropical representatives are derived
from the Nearctic. This has been shown to be the case for a relatively
small number of mayfly genera, such as Isonychia Eaton (Isonychiidae)

19 6 ENTOMOLOGICAL NEWS

and Stenonema Traver, Rhithrogena Eaton, and Iron Eaton (Hep-
tageniidae) (see McCafferty et al. 1992).

ACKNOWLEDGMENTS

We thank the following for the loan or donation of Central American specimens: R. W.
Flowers and M. L. Pescador (Florida A&M University, Tallahassee), R. W. Holzenthal
(University of Minnesota, St. Paul), and B. C. Kondratieff (Colorado State University, Fort
Collins). We also thank R. D. Waltz (Indiana Department of Natural Resources, Indian-
apolis) for reading the manuscript. This paper has been assigned Purdue Experiment Sta-
tion Journal No. 13741.

LITERATURE CITED

Cohen, S. D. and R. K. Allen. 1972. New species ofBaetodes from Mexico and Central

America. Pan-Pac. Entomol. 48: 123-135.
Cohen, S. D. and R. K. Allen. 1978. Generic revisions of mayfly nymphs. III. Baetodes in

North and Central America (Baetidae). J. Kans. Entomol. Soc. 51: 253-269.
Eaton, A. E. 1881. An announcement of new genera of the Ephemeridae. Entomol. Mon.

Mag. 17: 191-197.
Eaton, A. E. 1883-88. A revisional monograph of recent Ephemeridae or mayflies. Trans.

Linn. Soc. London. Zool. Ser. 3: 1-352.
Flowers, R. W. 1985. Guajirolus, a new genus of Neotropical Baetidae. Stud. Neotrop.

Fauna Environ. 20: 27-31.
Flowers, R. W. 1991. Diversity of stream-living insects in northwestern Panama. J. N. Am.

Benthol. Soc. 10: 322-334.
Hubbard, M. D. and W. L. Peters. 1981. Ephemeroptera. pp. 55-63 In: S. H. Hulbert, G.

Rodriguez, and N. D. Santos [eds.]. Aquatic biota of tropical South America, Part 1:

Arthropoda. San Diego State University, San Diego.
Hubbard, M. D., E. Dominguez, and M. L. Pescador. 1992. Los Ephemeroptera de la

Repiiblica Argentina: un catalogo. Revta. Soc. Ent. Argent. 50: 201-240.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1933. The mayflies (Ephemeroptera) of Texas

and their biogeographic affinities. In: L. Corkum and J. Ciborowski [eds.]. Proceedings

of the seventh international conference on Ephemeroptera. Sandhill Crane Press,

Gainesville. In press.
McCafferty, W. P. 1985. New records of Ephemeroptera from Middle America. Intern.

Quart. Entomol. 1:9-11.
McCafferty, W. P. and J. R. Davis. 1992. New and additional records of small minnow

mayflies (Ephemeroptera: Baetidae) from Texas. Entomol. News 103: 199-209.
McCafferty, W. P. and C. R. Lugo-Ortiz. 1992. Registros nuevos y notas sobre los

Ephemeroptera de Nicaragua. Rev. Nica. Entomol. 19: 1-7.
McCafferty. W P. and R. D. Waltz. \986.Baetismagnus. new species, formal new name for

Baetis sp. B of Morihara and McCafferty (Ephemeroptera: Baetidae). Proc. Entomol.

Soc. Wash. 88: 604.
McCafferty, W. P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae

(Ephemeroptera) of North and Middle America. Trans. Am. Entomol. Soc. 1 16: 769-

799.
McCafferty, W. P., R. W. Flowers, and R. D. Waltz. 1992. The biogeography of

Mesoamerican mayflies, pp. 173-193 In: S. P. Darwin and A. L. Welden [eds.], Bioge-
ography of Mesoamerica: proceedings of a symposium. Tulane Univ. Stud. Zool. Bot..

Suppl. Publ. 1.

Vol. 104, No. 4, September & October, 1993 197

Morihara, D. K. and W. P. McCafferty. 1979. The Baetis larvae of North America

(Ephemeroptera: Baetidae). Trans. Am. Entomol. Soc. 105: 139-221.
Navas, L. 1924. Insectos de la America Central. Broteria. 21: 55-86.
Packer, J. S. 1966. A preliminary study of the mayflies of Honduras. Ceiba 12: 1-10.
Traver, J. R. and G. F. Edmunds, Jr. 1968. A revision of the Baetidae with spatulate-

clawed nymphs (Ephemeroptera). Pac. Insects 10: 629-677.
Waltz, R. D. and W. P. McCafferty. \985.Moribaetis: a new genus of Neotropical Baetidae

(Ephemeroptera). Proc. Entomol. Soc. Wash. 87: 239-251.
Waltz, R. D. and W. P. McCafferty. 1987a. Revision of the genus Cloeodes Traver

(Ephemeroptera: Baetidae). Ann. Entomol. Soc. Am. 80: 191-207.
Waltz, R. D. and W. P. McCafferty. 1987b. New genera of Baetidae previously included

in Baetis Leach (Ephemeroptera). Ann. Entomol. Soc. Am. 80: 667-670.
Waltz, R. D. and W. P. McCafferty. 1987c. Systematics of Pseudocloeon. Acentrella,

Baetiella, and Liebebiella, new genus (Ephemeroptera: Baetidae). J. N. Y. Entomol. Soc.

95: 553-568.

1933 CALVERT AWARD PRESENTED FOR STUDY OF
STREAM INVERTEBRATES

Justin Schilling grew up near Wissahickon Creek in southeastern Pennsylvania. He
noticed that a small tributary flowing near a limestone quarry had a lot of chalky sediment.
He hypothesized that the sediment was limestone dust (calcium carbonate) and that
mollusks whose shells are made of calcium carbonate would be more abundant on
Wissahickon Creek downstream from the tributary. After a thorough study of water quality
and an analysis of stream invertebrates at three locations, he refuted his hypothesis. In the
process he learned a lot about the identification of local aquatic insects. Among the
approximately 25 students who entered insect-related projects at the Delaware Valley
Science Fairs, Justin was one of the few who could discuss insect biology and taxonomy.
The American Entomological Society awarded Justin Schilling its annual Calvert Award
for his project entitled "The Effects of a Limestone Tributary on Invertebrate Life in the
Wissahickon Creek." Justin is a sophomore at Abington Friends School in Jenkintown.
Pennsylvania.

The Calvert Award recognizes outstanding insect-related work by a young person in
the Delaware Valley. It honors Dr. Philip P. Calvert who joined the Society as a teenager,
later became its president, and was a member for 74 years until his death in 1961. As a pro-
fessor of biology at the University of Pennsylvania and as an associate of the Academy of
Natural Sciences of Philadelphia. Dr. Calvert stimulated an interest in insects among
many young people. The award includes membership in The American Entomological
Society and the Young Entomologists' Society and a check for $50.

The Calvert Award was presented April 28 at the Society's membership meeting at the
Academy of Natural Sciences of Philadelphia. Also honored were Benjamin Martin for his
runner-up project, "The Extraction and Testing of Natural Pesticides", and Ben Blanchard

for his honorable mention project, "A Quantitative Study of Water Quality in Local

Streams using Macroinvertebrates."

Harold B. White. Vice President. A.E.S.
and Chair. Education Committee

198 ENTOMOLOGICAL NEWS

CARPENTER ANT (HYMENOPTERA:

FORMICIDAE) TUNNELS VISUALIZED

BY COMPUTED TOMOGRAPHYi

Rif S. El-Mallakh2

ABSTRACT: Carpenter ant (Camponotus sp.) tunnels within a wooden beam of Douglas
fir (Pseudotsugo menziesii) were studied by X-ray computed tomography. A pattern charac-
terized by the main, or lead, tunnel within the pith and giving rise to an ever expanding set
of tunnels was noted. This innovative method of examination of wood specimens provides
accurate information about the interior tunnel systems of the ants without the need for sec-
tioning the wood and destroying the specimen.

Carpenter ants, Camponotus sp., are common, economically impor-
tant animals with a world wide distribution. Within the United States the
nine most common species are found primarily in the heavily wooded
moist northeast and northwest (Olkowski, Daar, and Olkowski, 1991).
Most species seek protection within tunnels excavated in dead or decay-
ing wood, but are capable of chewing through undamaged wood as well.
They may be attracted to man made structures, particularly if the wood is
weathered or if the structure contains small hollow areas within wooden
walls (Olkowski, Daar, and Olkowski, 1991). Since they do not feed on
the wood and since a colony may require 3-6 years to grow to its mature
size of roughly 2000 individuals, they are often slow to cause extensive
damage (Ebling, 1978).

The economic importance of carpenter ants derives primarily from
their tunnel excavations, consequently tunnel structure has been exten-
sively studied. It has long been known that they preferentially chew
through the softer heart and spring wood to produce a pattern of ever
enlarging concentric tunnels (Goetsch, 1953). in the past, making these
observations has required the destruction of the wooden structure that
holds the nest; this paper presents the results of an alternative method
which uses X-ray computed tomography to map the 3-dimensional
architecture of Camponotus tunnels without destroying the original
specimen.

1 Received December 3. 1992. Accepted April 7. 1993

- Department of Psychiatry and Behavioral Sciences. University of Louisville School of
Medicine, Louisville, Kentucky 40292

F.NT. NEWS 104(4): 198-202. September & October. 1993

Vol. 104, No. 4, September & October, 1993 199

MATERIALS AND METHODS

A severely damaged wood beam was obtained from the Field
Museum of Natural History in Chicago, Illinois (Figures 1 & 2). The
specimen was donated by a sculptor who had bought it for his work. Its
origins prior to that are unknown. The wood was identified by vascular
anatomy to be Douglas fir, Pseudotsuga menziesii (Mirb.) Franco. It
measured 9cm X 8.5 cm X 63.5 cm.

The wood was scanned with a 1005 type head EMI computerized
axial tomography (C.A.T.) scanner (EMItronics Inc., X-ray Systems
Division, Northbrook, Illinois), an early generation computed tomog-
raphy machine manufactured in the early 1970s. It is designed to obtain
cross-sectional X-ray images of the human brain and to reconstruct
them digitally. Each cross-sectional plane is scanned several times from
a variety of angles over 240 while corresponding X-ray attenuation data
are gathered. Since the degree of X-ray attenuation is dependent on the
density of the object, the image is a computer reconstruction of the den-
sity of the material. In this study, an 8 mm thick x-ray beam was used for
each slice, with a energy of 120 kV and 33mA.

RESULTS

A representative sample of the images is shown in Figures 3a-f. In
these photographs white areas are wood of sufficient density that it can
be resolved by the computer (usually summer wood). The dark areas are
either empty spaces or wood of such low density (usually spring wood)
that it cannot be resolved by the machine at the energy settings used. The
leading end of the ant tunnels is seen in the pith of the beam in Figure
3a. This section corresponds to the right arrow in Figure 1. As one
travels toward the outer end of wood beam (i.e., toward the end pictured
in Figure 2), the extent of the tunnels becomes progressively enlarged
(Figures 3a-3f)- While the density of the spring wood was less than the
resolution of the machine at the energy settings used, selective destruc-
tion of the spring wood can be ascertained by the clear presence of rem-
nants of summer wood within the major portion of the tunnels (Figures
3c, 3d, and 3e).

200

ENTOMOLOGICAL NEWS

^ Ir. .t

*

*VS' 4 '-

vm

.<* * 1. * ' ft ;A

^v?* ^

Figure 1 . The Douglas fir specimen used in this study. Only the portion between the arrows
was scanned. The right arrow corresponds to Figure 3a, while the left arrow corresponds to
Figure 3f.

Figure 2. Typical macroscopic pattern of severe Componotus damage. This photo corres-
ponds to Figure 3f.

Vol. 104, No. 4, September & October, 1993

201

Figure 3. Six representative photographs from a total of 24 taken. Each image is the average
density of 8 mm thick slice of wood, (a) The leading end of the runnel. Note that the ants
start at the soft pith, (b-e) A series revealing the progressive widening of the ant excavations.
Remnants of denser spring wood can occasionally be seen within the main body of the tun-
nel, (e) At the outer edge of the tunnel, the entire thickness of the beam is damaged.

DISCUSSION

Computerized tomography is a medical diagnostic instrument that
visualizes the internal soft tissue anatomy of the human body and brain
(Huckman, 1975; Weisberg, 1979). Its use in clinical medicine is now
quite extensive.

The application of this technique to the study of tunnels created by
Camponotus is proposed, and our study verifies that the results obtained
by this method are similar to previously reported patterns of tunnel
architecture. Specifically, the X-ray data is compatible with the primary
or leading end of carpenter ant tunnels being through the softer pith and
with the ants expanding their tunnels laterally with some sparing of the
denser summer wood (Goetsch, 1953).

202 ENTOMOLOGICAL NEWS

However, there are various limitations to the technique. First, the
great size and intricacy of the machine prevent any level of portabil-
ity. This reduces the potential application of this technique to relatively
small specimens that must be brought to the instrument. Second, at
current prices, computerized tomography time is very expensive (human
diagnostic charges average $600.00 per hour, and one hour would be
needed for most specimens). Third, while overall tunnel structure can be
visualized, small structural details, or the insects themselves are below
the level of resolution. The thickness of the X-ray beam, 8 mm in the
model used in this study, and software limitations of this and more
advanced models, make maximal resolution approximately 5 mm.
Finally, in this study full appreciation of the tunnel system was limited
due to inability to visualize the less dense spring wood. This problem can
be partly alleviated by decreasing the energy of the X-ray beam, and is
not a problem when scanning more dense material.

Nonetheless, this technique may be useful for ascertaining the extent
of insect damage to valuable wooden specimens (e.g., sculptures, an-
tiques, etc.). More importantly, since 3-dimensional reconstructions can
be made from the digitalized data, additional perfection of this tech-
nique can lead to the creation of manipulatable perspective images that
can be used for teaching or comparing 3-dimensional nest structure of
various species.

ACKNOWLEDGMENTS

lam indebted to George Wilder for help in the identification of the wood; Eric Smith for
the loan of the wood specimen; Donald Baumgartner for his insightful comments; and Saf-
wat El-Mallakh for the photographic work. Rush Presbyterian - St. Luke's Hospital,
Chicago, Illinois, graciously donated the use of their CAT. scanner.

LITERATURE CITED

Ebeling, W. 1978. Urban Entomology. Univ. of Calif. Press, Berkeley, CA, 695 pp.

Goetsch, W. 1953. The Ants. The Univ. of Mich. Press, Ann Arbor, MI, pp. 52-58.

Huckman, M.S. 1975. Clinical experience with the intravenous infusion of iodinated con-
trast material as an adjunct to computed tomography. Surgical Neurology 4: 297-
318.

Olkowski, W., Daar, S., and Olkowski, H. 1991. Common-Sense Pest Control. Taunton
Press, Newton, CT, 715 pp.

Weisberg, L.A. 1979. Computed tomography in the diagnosis of intracranial disease.
Annals of Internal Medicine 91: 87-105.

Vol. 104, No. 4, September & October, 1993 203

AN INEXPENSIVE VACUUM COLLECTOR
FOR INSECT SAMPLING 1

S. W. Wilson2, J. L. Smith 2 , A. H. Purcell, III 3

ABSTRACT: An inexpensive suction sampler was constructed by modifying a gasoline
powered leaf blower/vacuum (Weed Eater) Comparison of the sampler with use of an
aquatic dip net to collect a planthopper species from emergent aquatic plants showed that
the sampler was more efficient in collecting adults and was especially useful for capturing
early instars. The Weed Eater sampler was comparable or superior to a D-Vac for collect-
ing grassland arthropods (Wright and Stewart 1992). The total cost of this light (<6 kg.)
sampler, including modifications, is ca. $130.

The commonly used methods of collecting or sampling insects by
sweeping, beating, aspirating, or hand-picking undercollect very small
insects and mites disproportionately and miss arthropods at the base of
low vegetation (Herms et al, 1990; Perfect and Cook, 1983; Schotzko
and O'Keeffe, 1989; South wood, 1978; Summers et al., 1984). Suction
collectors such as the D-Vac (Dietrick, 1961) overcome these inade-
quacies at the costs of large size and weight (18.1 kg), expense
(US$1 100+), portability, and reliability. In this paper we describe a
relatively small (5.7 kg), inexpensive ($130) and portable vacuum col-
lector (hereafter referred to as the "bug-vac") easily converted from a
gasoline-powered leaf blower.

MATERIALS AND METHODS

Modifications were made to a Weed Eater hand held leaf blower
(Model 1920, 920, or 960) and Weed Eater vacuum attachment kit
(Model 952-701613 (VA 1905)) purchased at a discount store (the manu-
facturer's address is: Poulan/Weed Eater, Division White Consolidated
Industries, Inc., Shreveport, Louisiana 71139-9329, USA). Materials for
modification include duct tape (heavy duty cloth tape), eight 1 .2 cm x 0.3
cm machine screws, eight lock washers and nuts, one piece of 0.95 cm
metal circular screen, and two U-clamps (to attach the strap to the body
of the leaf blower). Detailed instructions for assembly and use as a leaf
vacuum are supplied by the manufacturer with the vacuum attachment
kit; modifications to these instructions include the following:

1 Received January 30, 1993, Accepted March 20, 1993

- Department of Biology, Central Missouri State University, Warrensburg, MO 64093

3 Department of Entomological Science. University of California. Berkeley. CA 94720

ENT. NEWS 104(4): 203-208. September & October. 1993

204 ENTOMOLOGICAL NEWS

1 ) The strap on the vacuum bag (part of the vacuum attachment kit)
was cut off where it connects to the bag, the bag disposed of (the
machine can be operated with the vacuum bag attached), and a
clamp sewn onto each end of the strap. The clamps on the strap
were attached to the ring on the leaf blower handle.

2) The basal end of the vacuum tube (from the vacuum attachment
kit) was fastened to its fitting sleeve by four bolts and nuts rather
than just by the large, circular hose clamp supplied for this pur-
pose. Four holes were drilled through the plastic sleeve on the
machine, the base of the plastic vacuum tube, and the hose clamp.
The exposed ends of the bolts were wrapped with duct tape around
the outside of the vacuum tube ("a" in Fig. 1).

3) We used two methods to prevent the insect-collecting bag from
being accidentally sucked into the vacuum fan. Either ( 1) a flat cir-
cular piece of wire screen (0.95 cm mesh) can be bolted into the
interior of the vacuum tube about 30 cm from the distal end of the
tube or (2) the screen can be fashioned into a cup-shaped basket
that fits snugly within the tube with the rim of the basket bent flush
and taped along the outside 2 cm of the vacuum tube. Loose ends
of wire in the screen are soldered. In either method, any protrud-
ing bolts or the wire rim of the basket are tightly wrapped with
heavy tape to cover these sharp protrusions ("b" in Fig. 1).

Orienting the leaf blower in a reverse (or backward) position facili-
tates starting.

RESULTS AND DISCUSSION

The bug vac proved equal or superior to sampling with a D-Vac or
net. Wright and Stewart (1992) converted an Atco "Blow- Vac" leaf-
blower using our directions and compared its sampling efficiency to a D-
Vac in three grassland sites in Great Britain. They found that the
"Blow- Vac" collected comparable numbers of Diptera, Hymenoptera,
and most Homoptera (Auchenorrhyncha). The D-Vac caught signifi-
cantly greater numbers of two species of leafhoppers (Cicadellidae) at
one of the three sites. However, their "Blow- Vac" proved much more effi-
cient than the D-Vac for capturing Coleoptera and Araneae.

The improved collection of immature planthoppers such as the
delphacid Pissonotus piceus (Van Duzee) (Homoptera: Fulgoroidea)
using the bug-vac was demonstrated in collections from its host, the
emergent aquatic plant "mild water pepper" (Polygonum hydropiperoides
Michx.) Sweeping this plant proved to be an inadequate method of
collecting this delphacid. A more efficient method, used once per week

Vol. 104. No. 4. September & October, 1993

205

Fig. 1 . Using the gasoline-powered leaf blower adapted as an insect vacuum ("bug-vac") to
collect insects from an aquatic emergent plant. Four bolts attach the basal end of the
vacuum tube to a sleeve on the machine; exposed ends of the bolts are covered with cloth
tape (a). A wire screen is bolted to the inside of the collector tube; exposed ends of the bolts
are covered with cloth tape(b). A net bag is inserted in the vacuum tube and attached with
two large rubber bands (c).

206

ENTOMOLOGICAL NEWS

during 1989, was to strike the plants with an aquatic dip net and aspirate
the insects from the net. During 1990, the bug- vac was used to sample
insects from plants as the collector waded through the vegetation (Fig. 1 ).
The bug-vac reduced average sampling times from 45 to 5 minutes and
collected a greater mean number of planthoppers (Fig. 2; t = 2.87, p <
0.05). It was especially useful for collecting the small (length ca 1 .0 mm.)
first instar nymphs (Fig. 3; t = 3.15; p < 0.05), few of which were
damaged.

As with other vacuum collectors, the bug-vac develops higher intake
velocities with the collecting tube near the ground. Despite this, we
collected significantly higher numbers of the leafhopper Fieberiellaflorii
Stal (Homoptera: Cicadellidae) from its typical shrub or hedge host
plants in Berkeley, California with the bug-vac, than with a standard

600

100 -

\

.III

1

1

1

;;

i

\

;

]

II

CMCNO CM O O CNCVJ

I I I I I I I I I I I I

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^^J^ZJr3Z3<Da><DO><U

<<<<mcncniocn

O

Fig. 2. Number of Pissonotus piceus collected per week with a dip net (dark stippling) and
bug-vac (light stippling); N dip net = 1587, N bug vac = 3732.

Vol. 104. No. 4. September & October. 1993

207

sweep net (Purcell, unpuhl. data). In these sampling experiments, equal-
length halves of boxwood (Buxus sp.) or privet (Ligustrum sp.) hedges
were sampled by thoroughly sweeping or vacuuming horizontally with
the net or bug-vac. Typical of other suction collectors, the bug-vac was
more efficient than sweeping, especially in collecting early instar
nymphs of F. florii.

The light weight and size of the bug-vac allow it to be transported, dis-
assembled, in a large suitcase if the exhaust tube and terminal end of the
vacuum tube are removed. Also, it can be carried in a carton that fits
easily in the luggage compartment of small automobiles. One disadvan-
tage shared with the D-Vac is the high noise level of the bug-vac; ear pro-
tection should be worn during operation. The engine has proven to be
reliable and durable over four seasons of use.

450

300 -

50 -

J_

o ' r-

CN CM

I I

to

O
I

o

cr> ' vo

CN
I I

co

O
I

in

I

CN
I

co ' T

OslO
I I

Q.
<D

Fig. 3. Number of Pissonotus piceus first instars collected per week with a dip net (dark stip-
pling) and bug-vac (light stippling); Njjp net = 92, N^ ug vac =

208 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

We thank L. R. Nault for sponsoring our demonstration of the bug vac at the Inter-
national Auchenorrhyncha Conference in Wooster, Ohio, in August 1990. We appreciate
the review of the manuscript by R. F. Denno, constructive comments of J. H.Tsaiand M. R.
Wilson, and data and enthusiasm of A. J. A. Stewart.

LITERATURE CITED

Dietrick, E. J. 1961. An improved back pack motor fan for suction sampling of insect pop-
ulations. J. Econ. Entomol. 54:394-395.

Herms, D. A., D. G. Nielsen, and T. D. Syndor. 1990. Comparison of two methods for
sampling arboreal insect populations. J. Econ. Entomol. 83:869-874.

Perfect, T. J. and A. G. Cook. 1983. Population sampling for planthoppers, leafhoppers
(Hemiptera: Delphacidae & Cicadellidae) and their predators in flooded rice. Bull.
Entomol. Res. 73:345-355.

Schotzko, D. J. and L. E. O'Keeffe. 1989. Comparison of sweepnet, D-Vac, and absolute
sampling, and diel variation of sweep net sampling estimates in lentils for pea
aphid (Homoptera: Aphididae), nabids (Hemiptera: Nabidae), lady beetles (Coleop-
tera: Coccinellidae), and lacewings (Neuroptera: Chrysopidae). J. Econ. Entomol.
82:491-506.

Southwood, T. R. E. 1978. Ecological Methods. Halstead Press, New York. 524 pp.

Summers, C. G., R. E. Garrett, and F. G. Zalom. 1984. New suction device for sampling
arthropod populations. J. Econ. Entomol. 77:817-823.

Wright, A. F. and A. J. A. Stewart. 1992. A study of the efficiency of a new inexpensive type
of suction apparatus in quantitative sampling of grassland invertebrate populations.
Bull. British Ecol. Soc. : 116-120.

Vol. 104. No. 4, September & October. 1993 209

INSECT REMOVAL FROM STICKY TRAPS USING
A CITRUS OIL SOLVENT

Richard S. Miller 2 , Steven Passoa 3 , Robert D. Waltz 4 , Victor Mastro 5

ABSTRACT: A new procedure using citrus oil was developed for removing both heavily
sclerotized and soft-bodied insect specimens from sticky traps. The scales of adult
Lepidoptera are usually left intact. Procedures for using standard techniques such as pin-
ning, slide preparation, and genitalic dissection are also discussed.

Although sticky traps are used extensively for studying and monitor-
ing insect populations (Peterson 1964; Murphy 1985), removal and iden-
tification of trapped specimens is difficult (Lindgren et al. 1983; Murphy
1985; Knodel and Agnello 1990). The sticky material in the traps, usually
polyisobutylene (PIB), often obscures or distorts critical characters
needed for accurate determination. Murphy (1985) tested the use of
various solvents to remove sclerotized insects, such as Coleoptera and
Hymenoptera, from sticky traps. He found that polar solvents are unsuit-
able and suggested several alternative nonpolar solvents including
toluene, heptane, hexane, and xylene. Ethyl acetate, methychloroform,
petroleum spirits, gasoline, and kerosene proved less effective. Because
all these solvents are, to some extent, toxic to humans and flammable,
they must be used under a fume hood in the laboratory away from flames
or electric equipment. Besides the laboratory hazards, these solvents are
ineffective for extracting adult Lepidoptera and soft-bodied insects
(Murphy 1985).

Because some of the targets of the USDA exotic pest detection pro-
gram are microlepidoptera, we sought alternative solvents and pro-
cedures for sticky trap insect identification. One potential alternative
solvent is citrus oil, which has the initial advantage that it is on the GRAS
(Generally Regarded As Safe) list of the Food and Drug Administra-
tion.

1 Received November 4, 1992. Accepted February 25, 1993.

2 Department of Entomology, Leon Johnson Hall, Montana State University, Bozeman.
Montana 59717. Formerly USDA/APHIS/PPQ, 8895 E. Main St., Reynoldsburg. OH
43068.

3 USDA/APHIS/PPQ, 8895 E. Main St.. Building 3, Room 109, Reynoldsburg. OH
43068.

4 Division of Entomology and Plant Pathology, Indiana Department of Natural Re-
sources, 402 West Washington Street, Room W290, Indianapolis, Indiana 46204.

5 Methods Development Lab., Building 1368, Otis Air National Guard Base, Massa-
chusetts 02542-5008.

ENT. NEWS 104(4): 209-213, September & October. 1993

210 ENTOMOLOGICAL NEWS

METHODS

We tested citrus oil marketed by three sources for effectiveness in the
extraction of various orders of insects imbedded in PIB. Formulations
included Durkee lemon extract used as a food additive and purchased
at a local supermarket; a histological clearing agent marketed as Histo-
Clear by National Diagnostics; and Livos thinning agent #7222 sold
by Livos Plant Chemistry Inc.* Insect specimens used in the tests were
those submitted on delta, Rebell, and wing traps to the USDA Pest
Identification Laboratory in Reynoldsburg, Ohio. The effectiveness of
the formulations was observed for most orders of insects.

RESULTS

All citrus oil formulations examined were viable alternatives to other
solvents listed by Murphy (1985), although cost per unit volume varies
widely. Our tests of various handling techniques suggest the following
procedures for removing and identifying specimens from PIB.

Screening Samples. Sticky trap bottoms are most effectively
screened with an illuminated magnifying glass mounted on a stable
base. It is also advantageous to cover the work area with scraps of card-
board to protect the work surface from being fouled with PIB or
scratched with a scalpel blade. If a stereomicroscope is used, protect the
objective lens with a neutral density or polarizing filter to prevent con-
tact with PIB.

Insect Removal. If the specimen is fresh and heavily sclerotized, for
example a beetle, it may be lifted directly from the trap bottom and
placed into solvent. A few drops of citrus oil on the trap bottom will
loosen the specimen and ease removal. However, if the specimen is dry
and brittle or soft bodied, it should be left untouched on the trap. The
extraneous portion of the trap (and any excess PIB) surrounding the
insect should be cut away with a scalpel before it is placed in the solvent
bath. Movement should be minimized, because any distortion of the PIB
will probably damage the specimen by pulling it apart. The volume of
the solvent bath should be at least sufficient to cover the specimen. After
a few hours the insect will float clear of the trap and PIB. Most specimens
can be left in the solvent overnight until any residual PIB has dissolved.
To prevent saturation of the citrus oil, the cut portion of the trap bottom
should be removed after the insect has been freed. The length of time

*FOOTNOTE. Mention of commercial products in this paper does not constitute a recom-
mendation by the United States Department of Agriculture.

Vol. 104, No. 4, September & October, 1993 211

required in the solvent varies with the amount of PIB to be dissolved and
the condition of the solvent. Solvent effectiveness will eventually decline
when it becomes saturated and, consequently, the time required to
remove the PIB will increase. Glassware (e.g., petri dishes) must be used
because citrus oil will react with plastic.

One way to speed removal of the PIB is to use an ultrasonic cleaner.
The insect is put in a small vial filled with solvent and placed in the
ultrasonic cleaner with water. The vial is required for two reasons. It
dampens the sound waves protecting fragile insects from excessive
movement and potential damage, and it conserves solvent because it is
not necessary to fill the whole tank. Most hard-bodied insects will be
cleaned in 5-10 seconds. Wings of Lepidoptera can also be descaled in
this manner for morphometric and venational studies.

Occasionally, a film of dried PIB will adhere to the specimen when it
is removed from spent solvent. This residue can usually be removed by
rinsing the insect in xylene and/or absolute ethanol. Leaving material
overnight in fresh citrus oil is another option. Although specimens may
be left in the citrus oil for extended periods without apparent damage,
they do become more brittle after 24 hours.

Pinning. After allowing the specimen to air-dry for a few minutes, it
may be pinned. If the insect must be relaxed before pinning, it can be
immersed in water for a few hours (or in subboiling water for a few
minutes).

Alcoholic Specimens. Insects which normally are stored in alcohol
may be rehydrated by placing them in subboiling water for a few minutes
before permanent preservation in 80% alcohol.

Slide-mounted Specimens. Very small specimens, e.g., springtails,
thrips, mites, scale crawlers, some nematoceran Diptera, and Hymenop-
tera for which the preparation of slide mounts may be necessary, can be
transferred directly from PIB into Euparal. Specimens that need to be
cleared before mounting should be handled as in the above section, thus
significantly reducing clearing time in KOH. Warming the Euparol prior
to mounting is helpful.

Preparation of Genitalia. The following procedure can be used to
prepare genitalia of moths trapped in PIB. Standard techniques dis-
cussed by Holloway et al (1987) have been modified and shortened to
save time in screening large samples.

1. Pull the abdomen from the trap substrate.

2. Immerse in citrus oil to clean specimen.

3. If still not free of PIB, return specimen to solvent for another 12
hours.

212 ENTOMOLOGICAL NEWS

4. When cleaned of PIB and if time permits, place abdomen in 10%
potassium or sodium hydroxide (KOH or NaOH) for 12-24 hours at
room temperature. Alternatively, wear safety goggles and boil the ab-
domen in hot hydroxide until it is soft.

5. Wash abdomen in water, or preferably, a 5% solution of glacial
acetic acid and water to neutralize the KOH or NaOH.

6. Place abdomen in 50% alcohol and mechanically brush scales
from it.

7. Stain with mercurochrome or chlorozal E black if desired.

8. Either place specimen in vial of 70% alcohol, mount in Hoyer's
solution, or clear and dehydrate the specimen for mounting in a resin
such as Euparol or Canada balsam.

The time required to prepare lepidopteran genitalia varies exten-
sively. Large moths generally require a longer KOH or NaOH bath than
smaller moths. Typically, a large moth may require almost a day at room
temperature, whereas smaller moths may need only a few hours. There-
fore, we recommend monitoring the progress of maceration. For those
unfamiliar with this technique, we suggest making trial runs with moths
of various sizes before attempting to use this technique on actual un-
knowns. Specimens left in citrus oil too long will be brittle, while those
left in KOH too long will be over-cleared and difficult to see.

In many cases where quick determination is required, "valve-rip-
ping" may be utilized. In this procedure the genitalic valva is grasped at
the base, pulled off the abdomen, placed in citrus oil, and then cleaned in
alcohol. Identification of many genera of Tortricidae and Noctuidae can
be confirmed by examining only the shape of the valve.

CONCLUSION

The success of each preparation depends largely on the condition of
the specimen when it is removed from the trap bottom. Fresh specimens
that are shallowly embedded in the PIB provide the best results, but even
those totally immersed can usually be recovered if they have not decom-
posed. Larger insects usually fare better, because small insects are more
likely to become immersed and decay. Extensive struggle by the insect on
the trap after capture often results in loss of setae and' scales. Addi-
tionally, scales, setae, and wings are often dislodged if the specimens are
manipulated before removal of the polyisobutylene. Specimens re-
moved with citrus oil using the above procedures have been maintained
for more than two years with no adverse effects.

Citrus oil offers distinct advantages over previously used solvents for
removing PIB, and most orders of insects have been extracted sue-

Vol. 104. No. 4, September & October, 1993 213

cessfully from sticky traps using this procedure. Generally, the tech-
nique works well for all taxa tested, but a higher percentage of success
occurs in fresh and more sclerotized specimens. Unlike solvents listed by
Murphy (1985), citrus oil does not leave specimens unduly brittle, and
subsequent laboratory and curatorial techniques can be easily accom-
plished after removal. Solvent toxicity is reduced or absent, although a
fume hood is still recommended to avoid breathing the fumes. Most
important, soft-bodied insects and Lepidoptera can be treated without
damage, if properly handled. Several problems remain: citrus oil is flam-
mable, the process remains time consuming, all specimens are not
recoverable, trap bottoms are often not reusable, and no specimens are
perfect "display quality." Other dry-trapping methods with screens to
remove unwanted nontargets are recommended if specimen quality is
critical.

ACKNOWLEDGMENTS

We thank Michael A. Ivie, Charles A. Triplehorn, and Raymond E. Hite, two anony-
mous reviewers, and the editor for critical comment on an early draft of this manuscript.

LITERATURE CITED

Holloway, J.D., J.D. Bradley, and D J. Carter. 1987. Lepi