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Vol. 100

JANUARY 1992

No. 1

Journal

of the

New York

Entomological Society

(ISSN 0028-7199)

Devoted to Entomology in General

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY

Editor: Janies K. Liebherr, Department of Entomology, Comstock Hall,

Cornell University. Ithaca, New York 14853
Assistant Editor: E^; Richard Hoebeke, Department of Entomology, Corn-

stock Hall, Cornell University, Ithaca, New York 14853
Book Review Editor: James S. Miller, Department of Entomology, Amer-

ican Museinn of Natural History, Central Park West at 79th Street,
v,‘l'^'New York, New York 10024

Publications Committee: Randall T. Schuh, American Museum of Natural

History, New York, Chairman; David L. Wagner, University of Con-
necticut, Storrs; Alfred G. Wheeler, Jr., Pennsylvania State Department
of Agriculture, Harrisburg.

The New York Entomological Society
Incorporating The Brooklyn Entomological Society

President: Durland Fish, Medical Entomology Laboratory, New York

Medical College, Valhalla, New York 10595

Vice President: Richard Falco, Westchester County Health Department,

White Plains, New York 10601

Secretary: Christine Falco, Westchester County Health Department, White

Plains, New York 10601

Treasurer: Louis Sorkin, Department of Entomology, American Museum

of Natural History, New York, New York 10024

Trustees: Class of 1990— Dennis J. Joslyn, Department of Biology, Rutgers

University, Camden, New Jersey; Bernard Furnival, New York, New
York. Class of 1991 — Sule Fischl, New York, New York; Theodore
Weisse, Northport, New York.

Annual dues are $23.00 for established professionals with journal, $10.00 without journal,
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mologica Americana can be purchased from Lubrecht and Cramer, RD 1, Box 244, Forestburgh,
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Meetings of the Society are held on the third Tuesday of each month (except June through
September) at 7 p.m. in the American Museum of Natural History, Central Park West at 79th
Street, New York, New York.

Mailed February 3, 1992

The Journal of the New York Entomological Society (ISSN 0028-7199) is published 4 times per year (January, April, July,
October) for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at
New York, New York and at additional mailing office. Postmaster: Send address changes to the New York Entomological
Society, % American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192.
Known office of publication: American Museum of Natural History, New York, New York 10024.

Journal of the New York Entomological Society, total copies printed 700, paid circulation 602, mail subscription 602, free
distribution by mail 19, total distribution 621, 79 copies left over each quarter.

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

J. New York Entomol. Soc. 100(1): 1-41, 1992

KEY AND DIAGNOSES FOR THE GENERA
(HEMIPTERA: PENTATOMIDAE: DISCO

L. H. Rolston

Department of Entomology, Louisiana Agricultural Experiment StauSfl?
Louisiana State University, Baton Rouge, Louisiana 70803

Abstract. — A key and diagnoses for 28 genera of Ochlerini are provided. New genera proposed
are: Barola, Catulona, Clypona, Coranda, Cromata, Forstona, Pseudadoxoplatys, Stalius and
Uvaldus. New species are Barola farfala, Catulona pensa, Catulona apaga, Clypona aerata,
Coranda castana, Cromata ornata, Forstona speciosa, Pseudadoxoplatys mendacis, Schaejferella
fusca, Stalius trisinuatus and Uvaldus concolor. New synonymy recognized is Melanodermus
StM, 1 867, as a junior synonym of Ochlerus Spinola, 1837, resulting in revival of the combination
Ochlerus circummaculatus St&l, 1867, and new combinations of Coranda picipes (Stcil, 1872)
and Stalius tartareus (St&l, 1862). Typhoeocoris Breddin, 1903, is placed in the synonymy of
Caracia St&l, 1872, and T. fulvifemur Breddin, 1903, in the synonymy of C. sexdens Stcil, 1872.
The holotype of Parochlerus latus Breddin, 1904, is redescribed. The male genitalia of this
species are described since only the female was known previously. A redescription of Tetro-
chlerus fissiceps Breddin, 1904, is given, and a voucher specimen is designated inasmuch as
type material of this species is unknown. A lectotype and paralectotype are designated for
Ochlerus tartareus Sled, 1862.

The previously established genera that are considered here were formerly included
in Halyini of the nominate pentatomid subfamily. A decade ago, these genera were
removed to Discocephalinae (Rolston and McDonald, 1979), and subsequently Och-
lerini was proposed to contain them apart from other genera in Discocephalinae
(Rolston, 1981).

Members of Ochlerini are exceedingly dull in appearance, being largely of funereal
hues. They have attracted little attention from taxonomists and as a consequence
their classification is rudimentary. The ochlerines reside in the tropics and subtropics
of the Western Hemisphere. Species of a few genera feed on herbaceous plants and
are commonly collected. However, most species apparently inhabit the forest canopy,
which explains their scarcity in collections and why most specimens were taken at
lights or on freshly felled trees.

The position of the rostrum and eyes relative to each other and to other morpho-
logical features are used extensively in the key and diagnoses. The anterior limit
(origin) of the labium has been determined with the venter of the insect uppermost
and the longitudinal axis of the thorax and abdomen at right angles to the line of
observation. The apparent relationship of the labium and eyes is altered when the
head is in an abnormal position. Likewise, the relationship of the apex of the rostrum

CHARACTERS INVOLVING THE ROSTRUM AND EYES

2

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

to the abdominal sternites and of the apex of the rostral segments to the coxae is
changed if the head is abnormally deflexed, reflexed or protruded. The proximity of
the eyes to the pronotum is, of course, decreased in the latter case. In using these
characters care must be exercised to ascertain that the head of the specimen is
normally positioned.

MEASUREMENTS

All measurements are in millimeters and, unless qualified, are rounded to the
nearest 0.05 mm. Dimensional lines on illustrations equal 0.5 mm.

EXCLUDED GENERA

Breddin (1903a) described Typhoeocoris fulvifemur as a new genus and new species
from Ecuador, indicating neither the disposition of the specimen upon which he
based his descriptions nor the classification other than “Rhynchoten.” Froeschner
(1981) ventured to catalog this insect as a halyine, but Breddin’s descriptions fit a
rather bizarre pentatomine described earlier by Stal (1872). I therefore propose the
following synonymy:

Caracia Stal, 1872:14-15.

Typhoeocoris Breddin, 1903a: 122. NEW SYNONYMY.

Caracia sexdens St&l, 1 872: 1 5.

Typhoeocoris fulvifemur Breddin, 1903a: 122-123. NEW SYNONYMY.

Breddin (1912) also described from Peru a new genus and new species, Melam-
byrsus hoplita, that he placed near Ochlerus and Melanodermus. Again, the disposition
of the type was not revealed. From the description, Melambyrsus seems similar to
Moncus St&l, but for the present it must remain a nomen dubium.

Spinola (1850a) proposed the generic name Audinetella in his synoptic table, par-
enthetically stating that the genus was based on an unpublished species from Brazil.
Description of the genus and species, Audinetella bipunctata Spinola, appeared the
same year (Spinola, 1850b). Here, however, the two female syntypes are said to have
come from Cayenne, and this is probably correct. These specimens pose a nomen-
clatural problem because one is Lincus croupius Rolston, 1983, and the other a
Paralincus species that is apparently unnamed. Audinetella Spinola, 1850, is senior
to both Lincus St&l, 1867, and Paralincus Distant, 1911. Use of Audinetella as the
senior synonym of Paralincus would be of small consequence since there is scant
primary literature on this little known genus and its junior synonym Vauriana Ruckes,
1958, but use of Audinetella as the senior synonym of Lincus would be very disruptive.
There is considerable primary literature on this large genus, much of it concerning
the role of Lincus species as vectors of diseases afflicting palms. Unfortunately, the
description of Audinetella bipunctata fits only Lincus croupius in several critical
aspects, so there is actually no choice in associating the binomen and specimen. For
these reasons I am petitioning the International Commission on Zoological Nomen-
clature to suppress the binomen Audinetella bipunctata Spinola, 1850, for purposes
of priority.

1992

GENERA OF OCHLERINI

3

Ochlerini Rolston, 1981

Diagnosis. Superior surface of third tarsal segment of hind legs shallowly excavated
in females (only flattened in Adoxoplatys) and sometimes in males.

Trichobothria on at least last stemite located laterad of adjacent spiracles, excepting
Pseudadoxoplatys with mesial trichobothrium of each pair on last stemite on imag-
inary line tangential to, and projecting caudad from, spiracular openings on last two
stemites (Fig. 5). Basal segment of rostrum projecting caudad of bucculae, terminating
on prostemum. Mesostemum thinly carinate mesially, metastemum usually so. Meta-
pleural ostioles each accompanied by auricle, this sometimes elongated but not drawn
out into ruga. Scutellum longer than wide at base (excepting brachypterous forms).
All tibiae broadly sulcate. Spiracles present on paratergite 8 of females, on sternite
8 of males.

Labium usually arising on or posterior to plane transecting head at right angles to
longitudinal axis of body and at anterior limit of eyes, but arising before this plane
in several genera ( Adoxoplatys , Neoadoxoplatys, Pseudadoxoplatys ; Ochlerus, Or-
batina, Schraderia\ and Phereclus).

Comments. Only the tarsal character appears unique among western hemisphere
pentatomids, all other characteristics of the tribe appearing in various combinations
elsewhere. The tribe’s greatest affinity outside of the subfamily is with Pentatomini.

KEY TO GENERA OF OCHLERINI

1 . Hemelytra fully developed 2

Brachypterous, membranes of hemelytra much reduced or absent 32

2(1). Clearly defined intercalary segment present at base of 2nd rostral segment; meta-
stemum broadly sulcate between lateral rims, with or without median, longitudinal
carina in sulcus 3

- Intercalary rostral segment absent; metastemum usually without lateral rims, usu-
ally carinate; flat or tectiform if lateral rims present 4

3(2). Antennae 4-segmented; inferior surface of femora armed with many small tuber-
cles of similar size, most of them in 2 rows Adoxoplatys Breddin (p. 6)

- Antennae 5 -segmented; inferior surface of femora armed with preapical pair of
spines, of which at least mesial member is stout, and usually with lesser spines

or tubercles Neoadoxoplatys Kormilev (p. 6)

4(2). Both trichobothria of each pair on stemite 7 laterad of imaginary line tangential
to outer margin of spiracular openings on stemites 6 and 7 and projecting caudad
(Fig. 9); length of head before ocelli usually less than 0.8 of width of head across
eyes; sides of abdomen usually convex from dorsal view 5

- Mesial trichobothrium of each pair on stemite 7 on such imaginary line (Fig. 5);

length of head before ocelli 0.8 or more of width of head across eyes; sides of
abdomen subparallel (Fig. 4) Pseudadoxoplatys, new genus (p. 7)

5(4). Width of scutellum at distal ends of frena about 0.6 or more of basal width; costal

angles of coria projecting little if any past apex of scutellum 6

- Width of scutellum at distal ends of frena less than 0.6 of basal width; costal angles

of coria projecting well beyond apex of scutellum 10

6(5). Scutellum reaching apex of abdomen Moncus St<U (p. 8)

- Scutellum shorter, an imaginary, transverse line at apex of scutellum crossing

penultimate connexival sclerites 7

4

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

7(6).

8(7).

9(8).

10(5).

11(10).

12(11).

13(12).

14(13).

15(10).

16(15).

17(16).

18(17).

19(15).

Anteapical tooth present on lateral margins of each jugum (Fig. 6)

Schraderia Ruckes (p. 9)

Lateral margins of juga unarmed 8

Humeral angles subacute, produced laterad of hemelytra by distance subequal to
one-half width of eye; elevated edge of each evaporative area sigmoid (Fig. 8) ...

Orbatina Ruckes (p. 10)

Humeral angles little produced; elevated edge of each evaporative area usually

arcuate (Fig. 10) 9

Anterolateral angles ofpronotum projecting anteriorly little if any beyond posterior
limit of eyes (Fig. 11); scutellum not constricted near posterior ends of frena;

metasternum with thin, weak carina Ochlerus Spinola (p. 10)

Anterolateral angles of pronotum projecting anteriorly well past posterior limit of
eyes, reaching near middle of eyes; scutellum usually slightly constricted near
posterior ends of frena; mestastemum flat, or tectiform, or thinly and weakly

carinate Stalius, new genus (p. 11)

Space between each eye and pronotum equal to or grea+er than 0.2 diameter of

eye (Figs. 21-24) 11

Eyes contiguous with pronotum or nearly so 15

Labium arising before imaginary plane bisecting head at anterior limit of eyes

Phereclus St&l (p. 1 4)

Labium arising behind such a plane 12

Inferior surface of femora armed distally with small, peg-like tubercles, these

reduced or absent on posterior femora Paralincus Distant (p. 14)

Inferior surface of femora smooth or granulate 13

Space between each eye and pronotum greater than one-half diameter of eye (Fig.

23); juga shorter than tylus Eritrachys Ruckes (p. 15)

Space between each eye and pronotum less than one-half diameter of eye (Figs.

24, 25); juga as long as or longer than tylus 14

Second rostral segment attaining mesocoxae, rostral apex lying on stemite 3 . . .

Herrichella Distant (p. 15)

Second rostral segment attaining metacoxae, rostral apex lying on stemite 6 ...

Barola, new genus (p. 1 6)

Second rostral segment terminating between procoxae and mesocoxae; rostrum

not projecting beyond stemite 3, usually not reaching onto abdomen 16

Second rostral segment attaining anterior limit of mesocoxae, or nearly so, or
reaching beyond this point; rostrum extending onto or beyond stemite 3 19

Anterolateral margins of pronotum convex; juga separated before tylus, pincer-

shaped (Fig. 29) Clypona, new genus (p. 1 7)

Anterolateral margins of pronotum concave or sinuous; juga contiguous before

tylus or nearly so 17

Anterolateral margins of pronotum slightly sinuous with small, triangular tooth
at anterolateral angles 18

Anterolateral margins of pronotum concave with anterolateral angles projecting
forward to anterior limit of eyes, this projection curving mesad and dorsad ....

Forstona, new genus (p. 20)

Basal plates somewhat tumescent near posterior margin, unevenly convex in pro-
file; pygophore strongly concave mesially ventrad of posterior margin; antennae

4-segmented Miopygium Breddin (part) (p. 1 9)

Basal plates evenly convex; pygophore convex mesially ventrad of posterior mar-
gin; antennae 5-segmented Uvaldus, new genus (part) (p. 25)

Humeri strongly emarginated (Fig. 35); veins of hemelytral membranes sometimes
reticulated Schaejferella Spinola (part) (p. 21)

1992

GENERA OF OCHLERINI

5

- Humeri weakly or not emarginated; veins of hemelytral membranes simple or

furcate, often with some basal cells 20

20(19). Anteocular processes well developed, acute (Figs. 43, 44, 47) 21

- Anteocular margins of head projecting at most as obtuse processes 23

21(20). Pronotum scarcely declivitous, anterolateral angles broadly rounded (Fig. 43) . .

Alathetus Dallas (part) (p. 23)

- Pronotum decidedly declivitous, anterolateral angles narrowly rounded or angular
22

22(2 1). Disk of basal plates swollen near posterior margin (Fig. 45); pygophore about one-
fourth as wide as maximum width of abdomen, posterior surface lacking subcon-
ical projection Alitocoris Sailer (part) (p. 23)

- Disk of basal plates almost evenly convex; pygophore about two-fifths as wide as

maximum width of abdomen, bearing large subconical projection on posterior
surface Macropygium Spinola (p. 24)

23(20). Second rostral segment ending between mesocoxae, rostral apex of female and

usually of male not reaching stemite 6 24

- Second rostral segment projecting past mesocoxae, rostral apex of both sexes

extending to or beyond stemite 6 29

24(23). Disk of basal plates swollen near posterior margin, or their posterior margin
standing well out from 9th paratergites, these profoundly concave basally, forming

pit; third antennal segment at least twice length of second 25

Disk of basal plates not conspicuously swollen, their posterior margin appressed
to 9th paratergites; length of third antennal segment much less than twice length
of second 26

25(24). Basal plates evenly convex, their posterior margin standing well out from 9th
paratergites (Fig. 53); 9th paratergites profoundly concave at base, each together
with adjacent margin of basal plate forming pit; dorsal margin of pygophore
projecting at bottom of mesial emargination as nearly horizontal projection . . .
Uvaldus, new genus (part) (p. 25)

- Disk of basal plates with swollen area, posterior margin appressed to 9th para-

tergites, these not forming basal pit; pygophore without projection at base of
emargination Alitocoris Sailer (part) (p. 23)

26(24). Lateral margins of juga anterior to anteocular concavity tapering sharply toward

apex (Fig. 65) 27

- Lateral margins of juga parallel or subparallel between anteocular concavity and

apex 28

27(26). Antennae 5-segmented Parochlerus Breddin (p. 27)

- Antennae 4-segmented Tetrochlerus Breddin (p. 29)

28(26). Second gonocoxae visible (Figs. 36, 39); humeri noticeably emarginated, produced

laterad of base of hemelytra by about two-fifths width of eye (Fig. 35)

Schajferella Spinola (part) (p. 21)

- Second gonocoxae hidden (Figs. 72, 78); humeri not produced

Catulona, new genus (p. 30)

29(23). Length of second antennal segment less than one-third length of third antennal

segment and less than width of eye Alitocoris Sailer (part) (p. 23)

- Second antennal segment at least one-half as long as third antennal segment and

longer than width of eye 30

30(29). Metastemum flat or slightly tectiform, sometimes weakly carinate for part of its

length Lincus Sill (p. 35)

- Metastemum clearly carinate mesially for entire length 31

6

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

31(30). Juga exceeding tylus, their lateral margins briefly subparallel between anteocular

concavity and apex (Fig. 79) Cromata, new genus (p. 33)

Tylus exceeding juga; lateral margins of juga before anteocular concavity tapering

toward apex (Figs. 84, 86) Coranda, new genus (p. 36)

32(1). Pronotum strongly explanate (Fig. 43) Alathetus Dallas (part) (p. 23)

Pronotum not explanate or with only anterolateral angles strongly produced 33

33(32). Anterolateral angles of pronotum extending anteriorly well beyond eyes; hemely-

tral membranes lacking Brachelytron Ruckes (p. 38)

- Anterolateral angles of pronotum not projecting anteriorly beyond eyes; hemelytral

membrane present but much reduced Miopygium Breddin (part) (p. 1 9)

Adoxoplatys Breddin, 1903

Adoxoplatys Breddin, 1903b:368; Kirkaldy, 1909:XXXIII, 238 (type species desig-
nated); Kormilev, 1 949:3 1 5-3 1 6, 324 (description, species keyed); Kormilev, 1 950:
343 (synonymy); Kormilev, 1955:12-15 (Breddin’s species redescribed, species
keyed); Ruckes, 1958:23 (systematic position). (Type species Adoxoplatys minax
Breddin, 1903, by subsequent designation, Kirkaldy, 1909.)

Diagnosis. Mesial tubercle at base of abdominal venter broad, flat, apposed by
sulcate metastemum. At least anterior femora armed on inferior surface with 2
irregular rows of small tubercles; superior surface of third tarsal segment of hind legs
flattened but not excavated. Labium arising well before imaginary plane bisecting
head at anterior limit of eyes; second rostral segment usually attaining mesocoxae,
with well defined intercalary segment at base; apex of rostrum usually on or beyond
stemite 5. Length of head before ocelli 0.6-0. 8 of width across eyes. Antennae
4-segmented. Juga surpassing tylus. Width of scutellum at distal end of frena nearly
one-half of basal width. Coria extending well past apex of scutellum. Body depressed.

Comments. This genus and Neoadoxoplatys are unusual among ochlerines in having
a sulcate metastemum, a prominent intercalary rostral segment, and usually a flat-
tened rather than excavated superior surface of the third tarsal segment of the hind
legs in both sexes.

The genus ranges from Panama to Argentina and contains 8 nominal species.

Neoadoxoplatys Kormilev, 1956

Neoadoxoplatys Kormilev, 1956:4; Ruckes, 1958:23 (systematic position). (Type
species Neoadoxoplatys saileri Kormilev, 1956, by original designation.)

Diagnosis. Metastemum broadly and shallowly sulcate, without median carina but
sometimes tectiform. Inferior surface of each femur armed with one or two pairs of
preapical spines, some spines stout and conspicuous; superior surface of third tarsal
segment of hind legs flattened in females, sometimes excavated. Base of abdominal
venter unarmed. Labium arising before imaginary plane bisecting head at anterior
limit of eyes; second rostral segment reaching mesocoxae, with intercalary segment
at base; apex of rostrum on or beyond sternite 5. Antennae 5-segmented. Juga longer
than or subequal to tylus. Length of head before ocelli 0.7 of width across eyes. Width

1992

GENERA OF OCHLERINI

7

Figs. 1-5. Pseudadoxoplatys mendacis. 1. Genital plates, caudoventral view. 2. Pygophore,
lateral view. 3. Pygophore, caudal view. 4. Habitus, with hemelytra slightly open. 5. Spiracles
and trichobothria on left side of stemites 6 and 7. Symbol: stemite 7 (S7).

of scutellum at distal end of frena 0.4-0. 5 of basal width. Coria projecting far past
apex of scutellum. Body depressed.

Comments. The genus ranges from Mexico to Argentina and contains 3 nominate
species.

Pseudadoxoplatys, new genus
Figs. 1-5

Type species Pseudadoxoplatys mendacis, new species.

Diagnosis. Mesial trichobothrium of pair on each side of stemite 7 located on line
tangential to lateral margin of spiracular openings on stemites 6 and 7 and projected
posteriorly (Fig. 5). Length of head before ocelli 0.8 of width across eyes. Metastemum
flat with thin, weak, mesial carina. Base of abdomen and femora unarmed. Labium
arising before an imaginary pane bisecting head at anterior limit of eyes, even before
antennifers; second rostral segment reaching mesocoxae; apex of rostrum on stemite
5. Antennae 5-segmented; basal segment projecting well beyond apex of head. Width
of reticulated part of one eye 0.2 width of head across eyes. Juga and tylus subequal
in length. Width of scutellum at distal end of frena about one-half of basal width;
costal angles of coria surpassing apex of scutellum. Sides of abdomen subparallel.

Comments. This genus bears a strong resemblance to Adoxoplatys and Neoadox-
oplatys because of the body shape (Fig. 4). However, it is unlike Adoxoplatys or
Neoadoxoplatys in having unarmed femora, in lacking an intercalary rostral segment
and in having the metastemum asulcate.

The excavated superior surface of the third tarsal segment of the hind legs of
females places this genus among the ochlerines.

8

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Pseudadoxoplatys mendacis, new species
Figs. 1-5

Description. Dark castaneous to fuscous with no pale markings.

Head 3. 0-3. 2 wide across eyes, 2. 9-3. 3 long. Width of eyes 0.6-0. 7, interocular
width 1. 7-2.0. Distance between ocelli 0.7-0. 8, across ocelli 1.15-1.3. Head almost
rectangular, scarcely concave before eyes; lateral margins ofjuga narrowly reflexed.
Segments I-V of antennae 1.6-1. 8, 0. 7-1.0, 2. 9-3. 4, 2. 5-2. 8, 2.4-2. 7 in length. Seg-
ments II-IV of labium 2. 6-2. 9, 2. 4-2. 9, 2. 3-2. 7 long.

Pronotum 6. 9-7. 8 wide at humeri, 3.0-3. 3 long mesially. Anterolateral margins
nearly straight; anterolateral angles obsoletely toothed; humeri not produced, rounded
(Fig. 4).

Scutellum 4. 5-5.4 wide at base, 5. 7-6. 5 long, without fovea in basal angles; frena
extending about two-thirds of distance from base to apex. Costal angles of coria
above penultimate tergum, far surpassing apex of scutellum; membranes fumose,
each with about 8 simple or furcate veins. Connexiva moderately exposed, with
lighter colored marginal spot in middle of each segment.

Metastemal carina thin, becoming evanescent posteriorly.

Genital plates as in Figure 1 . Emargination in posterior pygophoral margin deep,
U-shaped from caudal view (Fig. 3). Dorsal margin of inferior ridge V-shaped. Pos-
terolateral comers of pygophore incised, apparently at junction of lateral pygophoral
walls and inferior ridge. Parameres visible from caudal view, their apices enlarged,
ovoid. Posterior pygophoral margin sigmoid in profile (Fig. 2).

Body length 15.2-17.3.

Types. Holotype, <3 labeled “PERU: Madre de Dios, Rio Tambopata Reserve, 30
km SW of Puerto Maldonado, 290 m, Nov. 1-26, 1982, Edward S. Ross.” Deposited
in the California Academy of Sciences.

Paratypes, 499. 19 labeled same as holotype (LHR); 19 labeled “Frans Steinback
Coll. 1956. Todos Santos Prov. Chaparo Dep. 500 M. Cochabamba, Bolivia.” and
“J. C. Lutz Collection. 1961.” (NMNH); 19 labeled “Bolivia: Dept, of Santa Cruz,
1 120 ft. elevation” and “R. B. Cumming 12-X-59 black light trap” (RMB); and 19
labeled “Bolivia: Saavedra, Dept. Santa Cruz Agr. Exp. Sta.” and “R. B. Cumming.
27-XII-59. Black light trap” (AMNH).

Moncus Stal, 1867

Moncus Stal, 1867:524; Stal, 1872:12, 13 (keyed); Distant, 1911:245 (synonymy).

(Type species Ochlerus obscurus Dallas, 1851, by monotypy.)

Hemingius Distant, 1899:423-424 (synonymized by Distant, 1911). (Type species
Ochlerus scaber Walker, 1867 [=Ochlerus obscurus Dallas] by monotypy.)

Diagnosis. Scutellum extending to apex of abdomen, 0.8 as wide at distal end of
frena as at base, narrowest there, again widening before apex. Eyes large, reticulated
part of each 0.3 width of head across eyes. Labium arising on imaginary plane
bisecting head at anterior limit of eyes; second rostral segment reaching mesocoxae;
apex of rostrum on stemite 5 or 6. Antennae 5-segmented; basal segment projecting
past apex of head. Juga and tylus subequal in length, former compressed apically
and pinched around tylus. Length of head before ocelli 0.6 of width across eyes.

1992

GENERA OF OCHLERINI

9

Figs. 6-11. 6. Schraderia hughesae, head. 7, 8. Orbatina fulginia. 7. Head and part of

pronotum. 8. Evaporative area, left metapleuron. 9-1 1. Ochlerus handlirschi. 9. Spiracles and
trichobothria on left side of stemites 6 and 7. 10. Evaporative area, left metapleuron. 1 1. Head
and part of pronotum. Symbol: stemite 7 (S7).

Comments. The size of the scutellum set this genus apart from all other ochlerines.
There are 2 nominate species, both South American.

Schraderia Ruckes, 1959
Fig. 6

Schraderia Ruckes, 1959:3-5. (Type species Schraderia hughesae Ruckes, 1959, by
original designation.)

Diagnosis. Small preapical tooth present on lateral margin of each jugum (Fig. 6).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Similar to Ochlerus in other respects excepting male genitalia. Parameres divergent,
essentially linear (stem of parameres subparallel or convergent in Ochlerus', head of
parameres usually expanded laterad, forming right angle with stem).

Comments. The anteapical tooth on the lateral margin of each jugum distinguishes
members of Schraderia.

The 2 nominate species are from Central America.

Orbatina Ruckes, 1960
Figs. 7, 8

Orbatina Ruckes, 1960:228-229. (Type species Orbatina fuligina Ruckes, 1960, by
original designation.)

Diagnosis. Humeral angles moderately produced laterad, right angular (Fig. 7).
Width of reticulated part of one eye little more than 0.2 width of head across eyes,
0.7 of distance between ocelli. Labium arising well before imaginary plane bisecting
head at anterior limit of eyes; second rostral segment reaching mesocoxae or nearly
so; apex of rostrum on stemite 5. Antennae 5 -segmented; first segment clearly ex-
ceeding apex of head. Juga and tylus subequal in length. Length of head before ocelli
0.75 of width across eyes. Width of scutellum at distal end of frena 0.6 of basal width.
Costal angles of coria not or scarcely surpassing apex of scutellum; scutellum and
coria terminating above tergum 6. Elevated margin of evaporative areas sigmoid
(Fig. 8).

Comments. Among the ochlerine genera in which the scutellum is nearly as long
as or longer than the coria, and in which the scutellar width at the end of the frena
is about 0.6 or more of the basal width, Orbatina is distinguished by the laterally
produced humeri and relatively small eyes.

This genus is monotypic and known from Colombia, Bolivia and Panama.

Ochlerus Spinola, 1837
Figs. 9-1 1

Ochlerus Spinola, 1837:294; Herrich-Schaffer, 1844:63-64 (description, species keyed);
Dallas, 1851:1 50, 1 56-158 (keyed, synonymy, species keyed); Stal, 1 867:524 (keyed);
Stal, 1872:12, 13 (keyed, synonymy); Breddin, 1910:615-616 (genitalia). (Type
species Ochlerus cinctus Spinola, 1837, by monotypy.)

Menipha Amyot and Serville, 1843:1 12 (synonymized by Stal, 1867). (Type species
Menipha brunnea Amyot and Serville, 1843 [=Ochlerus cinctus Spinola, 1837], by
monotypy.)

Melanodermus Stal, 1867:524; Stal, 1872:13. (Type species Ochlerus circummacu-
latus Stal, 1860, by subsequent designation, Kirkaldy, 1909:XXXII.) NEW SYN-
ONYMY.

Diagnosis. Eyes large, width of reticulated part of an eye nearly 0.3 width of head
acoss eyes, subequal to distance between ocelli (Fig. 11). Labium arising before
imaginary plane bisecting head at anterior limit of eyes; second rostral segment
reaching mesocoxae; apex of rostrum on sternite 5, 6 or 7. Antennae 5-segmented;
basal segment projecting well beyond apex of head. Juga and tylus subequal in length;
juga unarmed, subacute or obliquely truncate at apex, there narrow. Length of head

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GENERA OF OCHLERINI

11

before ocelli 0.7 of width across eyes. Tooth at anterolateral angles of pronotum
small, usually directed laterad. Width of scutellum at distal end of frena 0.6-0.75 of
basal width; scutellum usually a little longer than coria, terminating above tergum
6 or 7. Metastemum thinly carinate mesially.

Comments. Ochlerus circummaculatus St&l, the type species selected by Kirkaldy
(1909) for Melanodermus, differs only trivially from other Ochlerus species, most
notably in the slightly larger tooth at the anterolateral angles of the pronotum and
the longer tooth on the bucculae. Melanodermus therefore becomes a junior synonym
of Ochlerus.

Ochlerus is one of 5 genera in which the scutellum is relatively long and broad
subapically. The other genera are: Moncus, Orbatina, Schraderia and Stalius.

There are 1 5 nominate species in the genus. The generic range is from Mexico to
Brazil.

Stalius, new genus
Figs. 12-20

Melanodermus Stal, 1867:524 (in part); Stal, 1872:13 (in part).

Stalius, new genus. (Type species Ochlerus tartareus Stal, 1862.)

Diagnosis. Metastemum flat or tectiform, or weakly and thinly carinate mesially.
Labrium arising on or near imaginary plane bisecting head at anterior limit of eyes;
second rostral segment reaching metacoxae; apex of rostrum on stemites 5 or 6.
Antennae 5-segmented; basal segment projecting past apex of head. Juga little or no
longer than tylus, each narrowly rounded apically. Length of head before ocelli 0.6-
0.7 width across eyes. Anterolateral angles of pronotum acute and directed forward,
or spinose and directed anterolaterad, in either case projecting well beyond posterior
limit of eyes (Figs. 12, 14). Scutellum and coria terminating above penultimate or
last abdominal segment; scutellum usually a little longer than coria and slightly
constricted near distal ends of frena; width of scutellum at distal ends of frena nearly
0.6 of basal width.

Comments. St&l (1867) originally placed two species in Melanodermus : Ochlerus
circummaculatus Stal, 1 860, and Ochlerus tartareus Stal, 1 862. Kirkaldy ( 1 909) chose
O. circummaculatus as the type species of Melanodermus. I have examined the type
material of these two species, which is conserved in the Naturhistoriska Riksmuseet,
Stockholm. In my opinion Ochlerus circummaculatus belongs in the genus Ochlerus,
which reduces Melanodermus St&l, 1867, to a junior synonym of Ochlerus Spinola,
1837. Ochlerus tartareus is not congeneric with O. circummaculatus and does not fit
into any established genus. Stalius is therefore proposed to accommodate O. tartareus
and the new species whose description follows. Of the 2 syntypes of Ochlerus tartareus,
both female, the one labeled (a) “Mexico,” (b) “Salle,’” (c) “Type,” (d) “300/80,”
(e) “Riksmuseum Stockholm” is selected as lectotype. The paralectotype is labeled
(a) “Columbia” (sic), (b) “Stal,” (c) “Type,” (d) “Paratypus,” (e) “301/80,” (f) “Riks-
museum Stockholm.”

Two other species were subsequently added in Melanodermus : M. picipes Stal,
1872, and M. dilutipes Breddin, 1904. I have examined the holotype of M. picipes,
also conserved in the Naturhistoriska Riksmuseet, but have been unable to locate

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 12-20. 12, 13. Stalius tartareus. 12. Head and anterior part of pronotum. 13. Genital

cup. 14-20. Stalius trisinuatus. 14. Habitus, composite drawing. 15. Left antennifer, lateral
view. 16. Genital plates, caudoventral view. 17. Pygophore, caudal view. 18. Genital cup. 19.
Aedeagus, right lateral view. 20. Right paramere, lateral view. Symbols: aedeagus (a); basal
plate (bp); capitate process (cp); paramere (pa); paratergite 9 (pt9); phallotheca (ph); phallothecal
process (php); proctiger (pr); secondary gonopore (sg); tubercle (t).

1992

GENERA OF OCHLERINI

13

the type of M. dilutipes. M. picipes is assigned to a new genus, Coranda, that is
proposed later in this paper.

Stalius trisinuatus, new species
Figs. 14-20

Description. Mostly fuscous; rostrum, legs, basal 0.2 of last antennal segment yel-
lowish brown or at least paler than rest of antennae; distal part of coxae and numerous
more or less confluent femoral spots, fuscous.

Antennifers cupped laterally about base of antennae (Figs. 14, 15). Segments I-V
of antennae 0.7-0. 9, 0.7-0. 8, 1.2-1. 4, 1.3-1. 6, 1.7-1. 9 in length. Width of head across
eyes 2.2-2. 5, length 1. 7-2.0; length of head before ocelli 0.6-0. 7 of width across eyes.
Interocular width 1. 1-1.2; distance across ocelli 0.9-1.05. Eyes slightly longer than
wide; width of an eye about one-half of interocular width. Juga and tylus subequal
in length; lateral margins of former subparallel between anteocular concavity and
narrowly rounded jugal apices (Fig. 14). Segments II-IV of labium 2. 2-2. 4, 1 .6-1.9,
1. 8-2.1 long.

Anterolateral angles of pronotum armed with oblique spine; anterolateral margins
of pronotum slightly sinuous, narrowly reflexed. Disk rough, rugose or coarsely gran-
ular. Width at humeri 5.2-6. 1, mesial length 2. 5-2. 8.

Scutellar margin usually broadly impressed along frena; disk somewhat rugose
basally, smooth caudad of frena. Width of scutellum at base 3. 3-4.0, width at distal
end of frena 0.6 of basal width; length 3. 9-5.0. Membranes of coria darkly fumose,
each with 9-10 simple veins.

Smooth, castaneous, submarginal band present on each side of abdominal venter.
Area about trichobothria similarly colored.

Ninth paratergite concave basally, abruptly bent in posterior direction near apex,
flattened and densely setose at apex (Fig. 1 6). Posterior margin of basal plates shal-
lowly concave beneath 9th paratergites from caudal view.

Posterior margin of pygophore sinuous from caudal view, broadly emarginate
mesially, less deeply and less widely emarginate laterally, entire emargination con-
sisting of 3 concave and 2 convex curves (Fig. 1 7). Membraneous tubercle present
on each side of genital cup arising from lateral wall, well removed from posterior
margin (Fig. 1 8). Parameres compressed, aduncate (Fig. 20). Aedeagus heavily scler-
otized and pigmented, phallotheca appearing monolithic (Fig. 1 9).

Types. Holotype, 5 labeled (a) “COSTA RICA: Puntarenas, Monte verde area. 6
June- 14 June 1973. 1400-1700 meters” and (b) “Erwin & Bevel Central American
Expedition, 1973.” Deposited in National Museum of Natural History, Washington.

Paratypes, 255, 922. 15 and 592 with same labeling as holotype (2 LHR, 4 NMNH);
15, 222 labeled “PANAMA: Chiriqui Prov. Santa Clara. May 23-25, 1980. E. Riley
& LeDoux” (1 LHR, 2 DBT); and 222 labeled “PANAMA, Chir. Renacimiento,
5000', Sta. Clara, May ’80. Ratcliffe” (1 AMNH, 1 HDE).

Comments. The lobed antennifers, spinose pronotal lobes, and the genitalia readily
distinguish this species from S. tartareus. The membraneous tubercles in the genital
cup of S. tartareus are located at the posterior margin of the genital cup, and the
mesial emargination in the posterior pygophoral margin is smoothly concave rather
than sinuously concave (Fig. 1 3).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 21-24. Head and anterior margin of pronotum. 21. Phereclus pluto. 22. Paralincus
terminalis. 23. Eritrachys bituberculata. 24. Herrichella thoracica.

Phereclus Stal, 1862
Fig. 21

Phereclus Stal, 1862:98; Stal, 1867:524 (keyed); Stal, 1872:14 (keyed). (Type species
Phereclus pluto Stal, 1862, by monotypy.)

Diagnosis. Eyes separated from pronotum by distance equal to 0.3 diameter of an
eye; width of reticulated part of one eye about one-fourth width of head across eyes.
Labium arising well before imaginary plane bisecting head at anterior limit of eyes;
second segment attaining mesocoxae. Anterior margin of pronotum moderately emar-
ginated, a line drawn across anterior limit of pronotum passing along or near anterior
margin of ocelli. Antennae 5-segmented; about one-half of basal segment protruding
beyond apex of head; second segment about one-half as long as third. Juga subequal
to tylus in length, their lateral margins tapering toward narrowly rounded apex of
head (Fig. 21). Scutellum nearly 0.5 as wide at distal end of frena as at base.

Comments. From the other 4 ochlerine genera in which the eyes are notably
separated from the pronotum, this genus may be distinguished by the origin of the
labium. In Phereclus the labium originates well before an imaginary plane bisecting
the head at the anterior limit of the eyes. In the other genera in which the eyes and
pronotum are markedly separated, the labium arises behind such a plane.

The 3 nominal species of this genus come from Colombia.

Paralincus Distant, 1911
Fig. 22

Paralincus Distant, 191 1 :246; Rolston, 1 983b: 183-1 84 (revision). (Type species Och-
lerus terminalis Walker, 1867, by monotypy.)

Vauriana Ruckes, 1958:10-12 (synonymized by Rolston, 1983b). (Type species Vau-
riana bimaculata Ruckes, 1958, by original designation.)

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GENERA OF OCHLERINI

15

Diagnosis. Femora armed distally on inferior surface with small, peg-like, seta-
bearing tubercles, these obscure or reduced in size and number on posterior femora.
Eyes separated from pronotum by about one-half the diameter of an eye (Fig. 22);
width of one eye about one-fourth width of head across eyes. Juga longer than tylus,
convergent apically, without anteocular process or preapical tooth. Antennae 5 -seg-
mented; basal segment of each slightly surpassing apex of head. Labium arising behind
imaginary plane bisecting head at anterior limit of eyes; second segment reaching
mesocoxae; apex of rostrum on stemites 4-6. Length of head from anterior limit of
ocelli to apex 0.7 of width across eyes. Anterior margins of pronotum shallowly
concave or sinuous. Scutellar width at distal end of frena 0.5 of basal width. Costal
angles of coria surpassing apex of scutellum. Metastemum carinate mesially.

Comments. There are 4 other genera in which the eyes are notably separated from
the pronotum: Barola, Eritrachys, Herrichella and Phereclus. Of these 5 genera, only
Paralincus has armed femora.

The 3 species of the genus are found in the Amazon basin and northern South
America.

Eritrachys Ruckes, 1959
Fig. 23

Eritrachys Ruckes, 1959:10-11. (Type species Eritrachys bituberculata Ruckes, 1959,
by original designation.)

Diagnosis. Eyes pedunculate, separated from pronotum by a little more than one-
half the diameter of an eye on anterior-posterior axis (Fig. 23). Antennae 5 -segmented;
first segment surpassing apex of head; second segment about half as long as third.
Juga shorter than tylus. Length of head before ocelli 0.6 of width across eyes. Labium
arising behind imaginary plane bisecting head at anterior limit of eyes; second rostral
segment attaining mesocoxae; apex of rostrum on penultimate stemite. Pronotal
emargination behind head and anterolateral margin on each side of head quite sin-
uous. Width of scutellum at distal end of frena 0.4 of basal width. Corium surpassing
apex of scutellum. Metastemum carinate mesially.

Comments. The pedunculate eyes, well separated from the pronotum, are distinc-
tive. This monotypic genus ranges from Costa Rica to Ecuador.

Herrichella Distant, 1911
Fig. 24

Herrichella Distant, 191 1:245-246. (Type species Herrichella thoracica Distant, 1911,
by monotypy.)

Diagnosis. Eyes separated from pronotum by distance equal to 0.2 diameter of eye
(Fig. 24). Antennae 5-segmented; first segment extending little past apex of head;
length of segment II less than 0.3 that of segment III. Juga exceeding tylus. Length
of head before ocelli 0.6-0. 7 width across eyes. Anteocular process poorly developed,
obtuse. Labium arising behind imaginary plane bisecting head at anterior limit of
eyes; rostral segment II attaining anterior limit of mesocoxae; apex of rostrum reach-
ing stemite 3. Anterior margin of pronotum moderately emarginated. Width of scu-

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 25-28. Barola farfala. 25. Head and anterior margin of pronotum. 26. Profile of left
buccula. 27. Left hemelytron. 28. Genital plates, caudoventral view.

tellum at distal end of frena 0.5 of basal width. Coria exceeding apex of scutellum.
Metastemum carinate mesially.

Comments. The original description is misleading with respect to relationships of
the rostrum to the coxae and sternites because the head of the holotype was appre-
ciably reflexed.

This is one of 5 ochlerine genera in which the eyes and pronotum are not contiguous
or nearly so. The other genera are Barola, Eritrachys, Paralincus and Phereclus.
Herrichella differs especially from Eritrachys is the length of the juga relative to the
tylus and in the form of the pronotum, from Paralincus in having unarmed femora,
from Phereclus in the origin of the labium and from Barola in having a shorter
rostrum, shorter second antennal segment and more developed metastemal carina.

Herrichella is monotypic and known only from Colombia.

Barola, new genus
Figs. 25-28

Type species Barola farfala, new species.

Diagnosis. Metastemum nearly flat, very shallowly sulcate on each side of weakly
developed, broad, median carina. Labium arising behind imaginary line traversing
head at anterior limit of eyes; bucculae strongly toothed (Fig. 26); segment II of
rostrum reaching metacoxae, apex attaining penultimate stemite. All trichobothria
well laterad of adjacent spiracle. Eyes separated from pronotum by about 0.4 max-
imum anterior-posterior diameter of eye (Fig. 25). Anteocular process lacking. An-
tennae 5 -segmented; length of segment II about two-thirds that of segment III. Length
of head before ocelli three-fourths of width across eyes. Anterior margin of pronotum
moderately emarginate behind head, an imaginary line across anterior limits of
pronotum passing near anterior limits of ocelli. Costal angles of coria surpassing apex

1992

GENERA OF OCHLERINI

17

of scutellum, lying above penultimate connexival segment. Width of scutellum at
distal end of frena about one-half of basal width. Femora unarmed.

Comments. This genus is similar to Herrichella but differs notably in having a less
developed metastemal carina, a longer rostrum and differently proportioned antennal
segments.

Barola farfala, new species
Figs. 25-28

Description. Dark brown above with ivory median macule at base of scutellum
and yellowish marginal macule on each connexvial segment, those macules on seg-
ments 5 and 6 covering most of segment; abdominal venter brownish yellow with
conspicuous dark punctation except mesially; legs light brown with slightly darker
subapical annulus on femora; last 2 antennal segments slightly lighter colored than
fuscous to dark brown basal segments.

Juga not surpassing tylus at apex, their lateral margins tapering from convexity
above antennifers to apex (Fig. 25). Vertex of head sharply delineated, strongly
convex, nearly tumescent. First antennal segment projecting well beyond apex of
head; segments I-V of antennae 0.9, 0.6, 0.9, 2.1, 3.0 long. Segments I-IV of labium
1.5, 2.9, 2.3, 2.3 in length. Head 2.6 wide across, 2.5 long. Interocular distance 1.4;
eyes 0.6 wide, 0.6-0.65 long from dorsal view; distance across ocelli 1.4. Base of
antennifers ventrally and line running from antennifers to base of head pale.

Pronotum 6.5 wide at humeri, 2.7 long at meson. Anterolateral margins sinuous,
reflexed. Punctation rugose.

Scutellum 4.0 wide at base, 5.1 long. Apex narrowly rounded, slightly elevated on
each side. Fovea in basal angles black, deeply punctured. Punctation of disk similar
to that of pronotum. Exocorium more strongly punctate than endocorium; mem-
branes dark, each with about 1 1 veins; costal angle acute, submarginal (Fig. 27).

Lateral borders of prothorax broadly calloused, pale. Pleural surfaces of thorax
otherwise densely punctate. Lateral borders of abdomen impunctate, shiny; sternites
3-6 with broad, impunctate sulcus; area between sulcus and lateral border uniformly
and darkly punctate; spiracles oval, black.

Basal plates submarginally impressed posteriorly (Fig. 28).

Length about 14 including membranes.

Types. Holotype, 9 labeled (a) “Barro Colo. Is., Panama C.Z. VII-28-1938.,” (b)
“Col. by E. C. Williams, Jr.” Deposited in the Field Museum of Natural History,
Chicago. Segment 3 of right antenna deformed.

Paratype, 19 labeled “PANAMA: Prov. Chiriqui, Rovira, 5-VII-64, A. Bruce, 2500
ft. mosquito light trap” (RMB).

Clypona, new genus
Figs. 29, 30

Type species Clypona aerata, new species.

Diagnosis. Second rostral segment ending between procoxae and mesocoxae; apex
of rostrum lying between metacoxae; labium arising well behind imaginary plane
bisecting head at anterior limit of eyes. Antennae 5 -segmented; diameter of 3 basal

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 29-34. 29, 30. Clypona aerata. 29. Habitus. 30. Genital plates, caudoventral view.

31. Miopygium cyclopeltoides, head and anterior pronotal margin. 32-34. Forstona speciosa.

32. Habitus. 33. Genital plates, caudoventral view. 34. Same, lateral view. Symbols: basal plate
(bp); 2nd gonocoxae (gx2); paratergite 9 (pt9).

segments appreciably greater than that of 2 distal segments; segment II short, one-
third or less length of segment III in females (males unknown). Eyes contiguous with
pronotum; width of reticulated part of one eye a little less than one-fourth width of
head across eyes. Anteocular process large, spinose (Fig. 29). Juga far surpassing tylus,
leaving deep incision in apex of head, pincer-shaped before tylus. Length of head
before ocelli 0.6 of width across eyes. Anterolateral margins of pronotum explanate;
anterolateral angles obtuse. Scutellum little longer than wide at base, length 0.05

1992

GENERA OF OCHLERINI

19

greater than basal width; width at distal end of frena 0.5 of basal width; scutellar
apex above tergum 5, surpassed by costal angle of coria. Femora and base of abdomen
unarmed. Metastemum thinly carinate mesially.

Comments. This genus differs from Miopygium in having 5 -segmented antennae,
apically dehiscent juga, a larger anteocular process and laterally explanate pronotum.
It is like Miopygium, Forstona and Uvaldus in having a relatively short rostrum.

Clypona aerata, new species
Figs. 29, 30

Description. Dorsum fuscous to dark castaneous with scattered flecks and vermi-
form lines brownish yellow, and general appearance of badly tarnished bronze. Venter
colored as dorsum but more extensively pale-marked on thorax. Antennae mostly
fuscous, first segment lighter toward base. Rostrum and legs brownish yellow; nu-
merous dark spots on femora and tibiae.

Juga projecting beyond tylus by distance nearly equal to three-fourths length of
tylus, narrowly rounded apically (Fig. 29); jugal margins forming incision in apex of
head, first diverging then converging. Diameter of eyes 0.6; distance between ocelli
1.1. Width of head across eyes 2. 7-2. 8, length 2.0-2. 1. Basal segment of antennae
largest in diameter, segments II and III larger in diameter than segments IV and V;
segments I-V of antennae 0.8, 0.5, 1.6-1. 9, 1.3-1. 4, 2. 2-2. 3 long. Bucculae obtusely
angled or toothed anteriorly, percurrent caudad of angle or tooth, then curving to
termination at base of head. Second through last labial segments 1.5, 0.8-0. 9, 0.7-
0.8 long.

Anterolateral margin of pronotum moderately convex from dorsal apsect; obtuse
anterolateral angles sometimes minutely toothed; humeri somewhat bossed before
junction of veins on coria; width of pronotum at humeri 6. 9-7. 4, length mesially
3.0-3. 3. Scutellum 4. 4-4. 8 wide at base, 4. 6-5.0 long; fovea in basal angles consisting
of a few large, shallow, punctate impressions. Connexiva broadly exposed; segments
forming weak serration, each with small, pale, marginal macule at middle. Mem-
branes dark, each with about 7 veins.

Basal plates rather evenly convex; posterior margin of each sigmoid from caudal
view; curving ventrad near meson, curving dorsad below 8th paratergite; posterior
margin from caudoventral view concave along mesial half (Fig. 30). Length 13.0-
13.5.

Types : Holotype, 2, labeled “Argentina, Ledeimo, July 14-48, P. Wygodzinsky.”
Deposited in National Museum of Natural History, Washington.

Paratypes, 222 labeled same as holotype (1 NMNH, 1 AMNH).

Miopygium Breddin, 1904
Fig. 31

Miopygium Breddin, 1904:153. (Type species Miopygium cyclopeltoides Breddin,

1904, by monotypy.)

Diagnosis. Rostrum relatively short, apex ranging from metacoxae onto third ab-
dominal stemite; second segment projecting little past procoxae; labium arising well
behind imaginary plane bisecting head at anterior limit of eyes. Antennae 4-seg-

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mented; basal segment projecting well beyond apex of head; diameter of basal 2
segments conspicuously greater than that of last 2 segments. Anteocular process small,
obtuse or acute (Fig. 31). Juga much longer than tylus, contiguous apically or nearly
so. Length of head before ocelli little more than one-half width across eyes. Antero-
lateral margins of pronotum carinate, a short, triangular tooth present at anterolateral
angles. Scutellar width at distal end of frena 0.5-0.55 of basal width; length of
scutellum usually 0. 1 longer than wide. Metastemum carinate mesially.

Comments. The rostrum of most ochlerines extends well onto the abdomen, often
nearly to the abdominal apex, but in Miopygium, Clypona, Forstona and Uvaldus
the rostrum does not project onto the abdomen or projects no farther than the third
stemite. Miopygium differs strikingly from the other 3 genera of this group in having
4-segmented antennae.

There are 2 nominate species, both from Brazil. M. cyclopeltoides has two forms,
one brachypterous.

Forstona, new genus
Figs. 32-34

Type species Forstona speciosa, new species.

Diagnosis. Anterolateral angles of pronotum expanded, curving around eyes, ex-
tending nearly to anterior limit of eyes (Fig. 32). Second rostral segment ending
between procoxae and mesocoxae; apex of rostrum lying between metacoxae; labium
arising well behind imaginary plane bisecting head at anterior limit of eyes. Antennae
5 -segmented; basal segment not reaching apex of head, this segment alone of markedly
greater diameter than last 2 segments; segment II three-fourths length of segment III
in female (male unknown). Eyes contiguous with pronotum; width of reticulated part
one-fourth width of head across eyes. Anteocular process present, acute. Juga con-
tiguous before tylus. Length of head before ocelli 0.7 of width across eyes. Scutellar
width at distal end of frena a little less than one-half basal width; length nearly 0.2
greater than basal width; apex of scutellum above tergum 5, surpassed by coastal
angle of coria. Femora and base of abdomen unarmed. Metastemum thinly carinate
mesially.

Comments. The antennae, pronotum and length of the scutellum distinguish this
genus from Miopygium, Clypona and Uvaldus.

Forstona speciosa, new species
Figs. 32-34

Description. Dull black above and below, but with shades of yellowish brown as
follows: fifth and distal part of fourth antennal segments, rostrum, tarsi, and small
macule at middle of scutellar base, on disc of each corium and marginally in middle
of connexival segments, the latter obscure.

Head rather flat above, densely punctate; margins before eyes narrowly reflexed,
width across eyes 2.7, length 2.4. Diameter of eyes 0.6; distance between ocelli 0.9.
Segments I-V of antennae 0.7, 0.8, 1.3, 1.5, 1.8 long. Bucculae subacutely toothed
anteriorly, curving posteriorly to termination a little before base of head. Second
through last labial segments 1.7, 1.3, 1.1 long.

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GENERA OF OCHLERINI

21

Pronotum behind cicatrices and scutellum strongly rugose; coria densely punctate,
weakly rugose. Pronotum 7.1 wide at humeri, 3.1 long mesially. Scutellum 4.5 wide
at base, 5.0 long; fovea in basal angles not prominent, consisting of a cluster of strong
punctures. Connexiva narrowly exposed. Membranes dark, each with about a dozen
veins.

Basal plates tumescent submarginally along posterior margin below 9th paratergite,
forming a rounded projection when viewed laterally (Figs. 33, 34).

Type. Holotype, 9 labeled “Brazil, Rondonia: Vilhena, Nov. 1973. M. Alvarenga.”
Deposited in the American Museum of Natural History. No paratypes.

Schaejferella Spinola, 1850
Figs. 35-42

Schaejferella Spinola, 1850b:88; Spinola, 1852:128 (reprint); Stal, 1867:523 (keyed);

Stal, 1872:12 (keyed, synonymy). (Type species Schaejferella litigiosa Spinola, 1850

[=Cimex incisus Herrich-Schaffer, 1839], by monotypy.)

Diagnosis. Humeri produced laterad of base of coria by distance subequal to one-
fourth width of eye and prominently emarginated (Fig. 35). Basal 3 segments of
antennae enlarged, with interspecific variation and sexual dimorphism in degree of
enlargement; length of segment II ranging from less than one-half of segment III to
subequal to segment III; basal segment scarcely surpassing apex of head. Anteocular
process present, weak and obtuse to moderately produced (Figs. 35, 38); juga con-
vergent or contiguous before tylus. Length of head before ocelli 0.7 of width across
eyes. Labium arising behind imaginary plane bisecting head at anterior limit of eyes;
segment II of rostrum reaching mesocoxae; rostrum ending on stemites 4 or 5. Width
of scutellum at distal end of frena about one-half of basal width. Coria extending
past apex of scutellum. Distal part of proctiger expanded laterally (Figs. 37, 40).
Metastemum mesially carinate. Basal plates as in Figures 36 and 39, their disk convex
with a low prominence near center.

Comment. Schaejferella resembles most closely Alitocoris and Macropygium in
characteristics other than those associated with the genitalia. There are two species
in the genus, the type species from Brazil and the other from the Andean region.

Schaefferella fusca, new species
Figs. 38-42

Description. Fuscous with following brown to brownish yellow: basal three-tenths
of last antennal segment, humeral emargination, rostrum, tarsi, and in part trochan-
ters and coxae.

Anteocular process reduced to obtuse tubercle (Fig. 38). Juga convergent before
tylus, their lateral margins subparallel between concavity before eyes and convexity
at apex. Disk of head strongly punctate, vertex rugose, juga transversely rugous
basally. Diameter of segments II and III of antennae (in female) only slightly greater
than that of segments IV and V; segments I-V of antennae 0.9, 1.3, 1.5, 1.6, 2.0
long. Apex of rostrum reaching middle of sternite 5; length of second through last
segments 2.4, 2.2, 1.8. Reticulated part of eye 0.7 wide, distance between eyes 1.6.
Distance across ocelli 1.5, between ocelli 1.1. Width of head across eyes 3.1, length
2.4.

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Figs. 35-42. 35-37. Schafferella incisa. 35. Head and pronotum. 36. Genital plates, cau-

doventral view. 37. Genital cup. 38-42. Schafferella fusca. 38. Head and anterior pronotal
margin. 39. Genital plates, caudoventral view. 40. Genital cup. 41. Pygophore caudal view. 42.
Distal part of pygophore, lateral view. Symbols: 2nd gonocoxae (gx2); inferior ridge (ir); par-
amere (pa); proctiger (pr).

Anterolateral margins of pronotum slightly sinuous; tooth at anterolateral angle
flat, subtriangular. Disk rugosely punctate. Pronotal width at humeri 7.5, length at
meson 3.3.

Scutellum rugose basally, finely punctured apically with a few yellowish brown
dots and vermiform marks. Basal width 4.6, length 5.2. Deep fovea present in each
basal angle. Frena extending along basal six-tenths of scutellum. Veins of hemelytral
membranes simple or furcate; each membrane with about 1 1 veins.

Genital plates as in Figure 39, with low tumescence near center of basal plates and
longitudinal furrows. From caudal view, posterior pygophoral margin with deep,
wide, U-shaped emargination half filled by inferior ridge (Fig. 41). Inferior ridge
incised mesially, bent posterolaterad on each side of incision. Paramere compressed;
posterior part of head convex on mesial face, concave on lateral face; convexities
reversed on anterior part of head. Proctiger expanded distally; enlargement reflexed,
concave ventrally, convex dorsally (Fig. 40). Extracting pygophore discloses in each
lateral wall a large emargination framing head of parameres (Fig. 42).

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GENERA OF OCHLERINI

23

Types. Holotype, 6 labeled “ECUADOR. Pastaza, Cuisimi, on Rio Cuisimi, 1 50
km SE Puyo, 350 m. July 19-23, 1971. B. Malkin.” Deposited in the American
Museum of Natural History.

Paratype, 9 labeled “PERU, Loreto. Headwaters of Loreto- Yacu, Yagua Indian
Vill. April 21-May 1, 1970. B. Malkin” (LHR).

Comment. This species is readily distinguished from the only other known species
of the genus, S. incisa, by the predominately fuscous color and lack of reticulation
in the hemelytral membranes.

Alathetus Dallas, 1851
Fig. 43

Alathetus Dallas, 1851:160; Rolston, 1982:156. (Type species Alathetus rufitarsus

Dallas, 1851, by monotypy.)

Diagnosis. Pronotum flat dorsally; anterolateral margins explanate, especially at
anterolateral angles (Fig. 43). Anteocular process present. Antennae 4-segmented;
basal segment reaching to or beyond apex of head. Labium arising behind imaginary
plane bisecting head at anterior limit of eyes; second rostral segment attaining or
surpassing mesocoxae; apex of rostrum on or beyond stemite 5. Juga surpassing tylus.
In alate species length of head before ocelli 0.7 width across eyes, and width of
scutellum at distal end of frena 0.4-0. 5 of basal width. Metastemum carinate mesially.
Brachypterous species lacking ocelli.

Comments. The genus contains 2 species, the type species from Jamaica and a
brachypterous species, A. haitiensis Rolston, 1982, from Hispaniola.

The form of the pronotum in Alathetus is distinctive among ochlerine genera.

Alitocoris Sailer, 1950
Figs. 44-46

Alitocoris Sailer, 1 950:69-7 1 . (Type species Alitocoris schraderi Sailer, 1 950, by orig-
inal designation.)

Diagnosis. Labium arising behind imaginary plane bisecting head at anterior limit
of eyes; second rostral segment nearly attaining to slightly surpassing mesocoxae;
apex of rostrum reaching stemites 3-5. Length of head before ocelli 0.6-0. 7 width
across eyes. Anteocular process present, usually a blunt tubercle, occasionally acute
and set apart by concave emargination in lateral margin of jugum (Figs. 44, 46).
Antennae usually 5 -segmented, 4-segmented in some species; first segment extending
little or not at all beyond apex of head; basal 3 segments of 5-segmented antennae
or basal 2 segments of 4-segmented antennae somewhat to much enlarged, their
diameter appreciably greater than that of following segments, particularly in males
of some species. Width of scutellum at distal end of frena 0.4-0. 5 basal width. Coria
surpassing apex of scutellum. Metastemum carinate mesially. Width of pygophore
one-fourth of maximum abdominal width. Basal plates with broad tumescence near
distal margin (Fig. 45). Posterior surface of pygophore concave, usually obscuring
genital cup from caudal view excepting portion visible through mesial emargination.

Comments. There are 5 described species, all from Central America. However,
undescribed species extend the genus range from Mexico to southern Brazil.

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Figs. 43-49. 43. A/athetus rufitarsis, head and pronotum. 44, 45. Alitocoris parvus. 44. Head.
45. Genital plates, caudoventral view. 46. Alitocoris manni, head. 47-49. Macropygium reticu-
lare. 47. Head and pronotum. 48. Pygophore and 7th abdominal stemite, ventral view. 49.
Genital plates, caudoventral view. Symbol: proctiger (pr).

Alitocoris shows considerable interspecific variation with respect to size, devel-
opment of anteocular processes, and antennae (relative diameter and length, and
number of segments). The genitalia, however, are interspecifically similar.

Macropygium Spinola, 1837
Figs. 47-79

Macropygium Spinola, 1837:287; Herrich-Schaffer, 1844:48 (description); Dallas,
1851:1 50, 158-159 (keyed, synonymy). (Type species Macropygium atrum Spinola,
1837 [=Cimex reticularis Fabricius, 1803], by monotypy.)

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GENERA OF OCHLERINI

25

Oxyrhinus Amyot and Serville, 1843:1 13 (synonymized by Dallas, 1851). (Type

species Oxyrhinus subsulcatus Amyot and Serville, 1843, by monotypy.)

Diagnosis. Anteocular process prominent, spinose (Fig. 47); juga convergent or
contiguous apically. Basal 3 of 5 antennal segments enlarged; basal segment usually
projecting beyond apex of head; segment II less than one-half length of III; segment
III in males nearly twice diameter of each of last 2 segments. Length of head before
ocelli 0.6 width across eyes. Labium arising behind imaginary plane bisecting head
at anterior limit of eyes; second rostral segment reaching mesocoxae or nearly so;
apex of rostrum on stemite 3 or 4. Anterolateral margins of pronotum explanate;
anterolateral angles projecting forward to middle of eyes. Scutellar width at distal
end of frena 0.5 basal width. Coria surpassing apex of scutellum. Many small tubercles
scattered over inferior surfaces of femora, most prominent on front legs. Sternite 7
in male longer than all preceding stemites combined. Pygophore large, 0.4 as wide
as greatest abdominal width, bearing large, mesial cone on disk (Fig. 48). Disk of
basal plates rather evenly convex, without prominent, swollen area (Fig. 49).

Comments. This is the most frequently collected of ochlerine genera and one of
the easiest to recognize. The genus is currently monotypic, although there are dif-
ferences among male genitalia that suggest the possibility of one or more additional
species.

Uvaldus, new genus
Figs. 50-59

Type species Uvaldus concolor, new species.

Diagnosis. Imaginary vertical plane at base of labium and at right angles to axis
of body passing through eyes; basal segment of rostrum projecting onto concave
prostemum, second segment approaching or reaching mesocoxae, apex of fourth
extending onto base of abdomen. Bucculae well developed, uniformly produced,
reaching base of head. Antennae 5-segmented; basal segment of each projecting
slightly past apex of head, third segment 2. 5-3.0 times length of second. Eyes con-
tiguous with pronotum. Anteocular process present on each side (Fig. 50). Length of
head before ocelli 0.6 of interocular width.

Anterolateral margins of pronotum weakly sinuous; small tooth at each antero-
lateral comer directed laterad; humeri scarcely produced, weakly emarginated. Width
of scutellum at distal end of frena slightly more than one-half basal width; apex of
scutellum reaching between imaginary line connecting anterolateral angles of fifth
abdominal segment and similar line connecting posterolateral angles of same segment.
Costal angles of coria projecting well past scutellar apex.

Mesostemum and metastemum carinate. Ostiolar sulci lacking; rugae short, each
reaching 0.15 of distance from ostiole to lateral margin of metapleuron; evaporative
areas small, confined to metapleura. Superior surface of last tarsal segments of hind
legs flattened or concavely depressed in both sexes. Abdomen without mesial tubercle
or spine at base. All trichobothria laterad of spiracular line. Basal plates rather evenly
convex, without swollen areas. Width of pygophore 0.3 of maximum abdominal
width.

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Figs. 50-59. Uvaldus concolor. 50. Habitus. 51. Genital plates, caudoventral view. 52. Same,
caudal view. 53. Same, lateral view. 54. Spermatheca. 55. Pygophore, caudal view. 56. Genital
cup. 57. Pygophore, ventral view. 58. Right paramere, lateral view. 59. Aedeagus, lateral view.
Symbols: basal plate (bp); paramere (pa); proctiger (pr); paratergite 9 (pt9).

Uvaldus concolor, new species
Figs. 50-59

Description. Light brown above with fuscous to black punctation, usually with
rufous to castaneous flecks on pronotum and scutellum and halos of the same color
around punctures; cicatrices dark brown; moderate size callus in each basal angle of
scutellum and elongated macule on disk of each corium usually ivory. Venter mostly
dark brown; coxae, trochanters and rostrum brownish yellow; femora, tibiae and
tarsi mottled in browns of varied intensity. Antennae uniformly light or dark brown.

Juga contiguous before tylus, leaving small indentation in apex of head, moderately
reflexed marginally, especially at anteocular concavities (Fig. 50); anterocular process

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GENERA OF OCHLERINI

27

on each side of juga short, obtuse. Punctation rather dense, irregular. Width of head
across eyes 2.4-2. 5, length 1.75-1.95; length before ocelli 1.5-1.55; interocular width
1.35-1.5; distance across ocelli 1.1-1. 2, between ocelli 0. 8-0.9, from each ocellus to
nearest eye 0.30-0.35. Segments I-V of antennae 0.65-0.7; 0.4-0. 6; 1.3-1.35; 1.2-
1.3; 1.6-1. 7 long. Fourth antennal segment of males slightly clavate, its greatest
diameter exceeding that of all other segments save first. Segments I-IV of labium
1.0; 1.6-1. 7; 1.0-1. 2; 1.0 long.

Anterolateral pronotal margins slightly sinuous, clearly reflexed. Punctation less
dense than on head, coarser, tending to cluster into short, transverse, streptococcal
lines. Pronotal width at humeri 5. 7-6. 2, medial length 2. 5-2. 8.

Fovea in each basal angle of scutellum shallow, triangular, densely punctate, black;
calloused macule contiguous with and about size of fovea, usually ivory. Punctation
similar to that on pronotum but denser. Scutellar width at base 3. 6-3. 9, length 4.0-
4.3. Distal margin of each cerium nearly straight, inner angle rounded; punctation
on endocorium and clavus as on pronotum, denser and finer on exocorium; mem-
branes heavily fumose, each with 6 or 7 simple veins. Connexiva exposed, finely
punctate, nearly uniform in color.

Posterior margins of basal plates sinuous from caudal view (Fig. 52), thickened,
appearing somewhat separated from other plates (Fig. 53). Paratergite 9 channelized
basally. Proximal part of spermathecal duct enlarged; spermathecal bulb small with
3 long diverticula (Fig. 54).

Sides of pygophore produced caudad, deeply sulcate; furrow dividing sides of
pygophore from convex area ventrad of genital cup (Fig. 55); ventral border of genital
cup convexly produced mesially, projecting horizontally. Distal half of proctiger
scleritized, granular (Fig. 56). Parameres bilobed; dorsal lobe globose, smooth; ventral
lobe thickly hirsute (Fig. 58).

Types. Holotype, 5 labeled “Brazil: Est. Rio de Janeiro, Araruama XL 1981. Coll.
M. Alvarenga.” Pygophore dissected. Deposited in the Carnegie Museum of Natural
History, Pittsburgh.

Paratypes 699 and 15 labeled same as holotype. (49$ CMNH; 19 DAR; 19 DBT;

15 LHR.)

Parochlerus Breddin, 1904
Figs. 60-66

Parochlerus Breddin, 1904:153. (Type species Parochlerus latus Breddin, 1904, by
monotypy.)

Diagnosis. Lateral jugal margins tapering sharply from anteocular concavities to
apex of head; juga convergent, projecting beyond tylus (Fig. 65); anteocular process
before each eye rudimentary, obtuse. Length of head before ocelli two-thirds of width
across eyes. Labium arising behind imaginary plane bisecting head at anterior limit
of eyes; second rostral segment reaching slightly past mesocoxae; apex of rostrum on
stemite 5. Antennae 5 -segmented; basal segment protruding past apex of head; seg-
ment II nearly as long as segment III. Scutellar width at distal end of frena about
one-half of basal width. Coria surpassing apex of scutellum. Metastemum carinate.
Comments. The generic name was an unfortunate choice because a close relation-

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Figs. 60-66. Parochlerus latus. 60. Pygophore, caudal view. 61. Genital cup, dorsal view.
62. Paramere. 63. Pygophore, lateral view. 64. Genital plates, caudoventral view. 65. Head.
66. Aedeagus. Symbols: inferior ridge (ir); membranous area of pygophore (m); proctiger (pr).

ship to Ochlerus is implied. Only Tetrochlerus is obviously near Parochlerus, and
these two genera appear to stand apart within the tribe.

Parochlerus is monotypic. Breddin’s meager description of the type species, based
on one female, is amplified by the following redescription, and the male is described.
The species is known only from Peru and Amazonas, Brazil.

Parochlerus latus Breddin, 1 904
Figs. 60-65

Parochlerus latus Breddin, 1904:153.

Holotype. Black to fuscous except as follows: edge of abdomen ventally, irregular
narrow border on lateral margins of juga, scattered calluses and macules on dorsum
and venter, basal 0.4 of last antennal segment, all yellowish; remainder of antennae,
coxae, tarsi brown to castaneous; evaporative areas slate colored.

Head 2.7 wide at eyes, 2.2 long; width across ocelli 1.25. Punctation rugose,
diagonally so on juga. Segments I-V of antennae 0.9, 1.1, 1.6, 1.8, 2.1 long.

Pronotum 7.0 wide at humeri, 3.2 long mesially; anterolateral margins nearly

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GENERA OF OCHLERINI

29

straight; humeri not produced laterally, obtusely angular; disk rugosely punctate
behind cicatrices.

Scutellum 4.5 wide at base, 5.1 long; frena extending 0.6 disance from base to
apex; disk somewhat rugose, especially basal part; apex moderately rounded, undif-
ferentiated by color or punctation. Coria punctured similarly to distal part of scu-
tellum; membranes brown, each with about 10 simple or furcate veins. Connexiva
moderately exposed, rather densely and strongly punctate.

Prostemum concave, mesostemum narrowly carinate mesially, both conspicuously
punctured; metastemum carinate mesially (mostly destroyed by pin). Spiracles oval.
Genital plates as in Figure 64.

The female holotype is labeled (a) “Peru, Amar,” (b) “Parochlerus latus Breddin,”
(c) “Coll. Breddin,” (d) “Typus,” (e) “Holotypus.” It is conserved in the Institut fur
Pflanzenschutzforschung, Eberswalde. The locality may not be interpreted correctly
from the handwritten label. The original description indicates the holotype to be a
male. Presumably the wrong sex symbol was used by Breddin or by the typesetter.

Males. Similar to female excepting genitalia.

Pygophore with apex V-shaped, mesial emargination of posterior margin truncate
from caudal view (Fig. 60). Inferior ridge with U-shaped, mesial emargination; area
between posterior pygophoral margin and inferior ridge not excavated. Pygophoral
appendage at each posterolateral comer separated by membrane; mesial face of
appendage concave; lateral face mostly membraneous, only anterior and dorsal mar-
gins sclerotized from lateral view (Fig. 63). Posterolateral pygophoral comers, ventrad
of appendages, acutely produced. Proctiger lacking tubercles; margin thickly fringed
with long hair, particularly subapically (Fig. 61). Parameres delicate, each bent at
right angle, somewhat flattened distally, bearing row of short hair proximad of bend
and peripheral fringe of long hair distad of bend (Fig. 62). Theca and structures
surrounding ejaculatory duct distad of theca sclerotized; membranous conjunctiva
minuscule (Fig. 66).

Tetrochlerus Breddin, 1904

Tetrochlerus Breddin, 1904:153. (Type species Tetrochlerus jissiceps Breddin, 1904,

by monotypy.)

Diagnosis. Differing from Parochlerus only in having 4-segmented antennae; seg-
ment III 0.6 length of segment II.

Comments. The type of T. fissiceps was not located. However, there is a specimen
in the National Museum of Natural History, Washington, DC, that fits B redd in’s
short description exactly, including a length of 1 1.5 from the apex of the head to the
tip of the hemelytral membranes, excepting that it is female. This specimen is de-
scribed below and designated voucher specimen. It bears the labels (a) “Balzapamba
(Ecuad). R. Haensch S. ,” (b) “Coll. Breddin,” (c) “J. C. Lutz Collection, 1961,” (d)
“Tetrochlerus fissiceps Breddin.”

The difference between Tetrochlerus fissiceps and Parochlerus latus in the number
of antennal segments is hardly sufficient grounds to place each species in a separate
genus. However, it seems best to leave this arrangement undisturbed until the male
of T. fissiceps is known, since the male genitalia of the two species may differ suffi-
ciently to justify retention of the present classification.

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Tetrochlerus fissiceps Breddin, 1 904
Tetrochlerus fissiceps Breddin, 1904:153.

Voucher. Dark brown except as follows: lateral margins of prothorax above and
below, of coria basally, of connexiva and abdominal venter, rufous; submarginal
macule in middle of connexival segments, flecks on abdominal venter and basal 0.4
of last antennal segment light brown to brownish yellow; anterior disk of pronotum,
head, antennae (except base of last segment), femora and tibiae, fuscous to black.

Head 2.2 wide across eyes, 1.8 long; length before ocelli 0.7 width across eyes;
dorsal punctation dense, strong, diagonally rugose on juga; interocular width 1.3,
each eye 0.45 wide, anterior-posterior diameter 0.55. Segments I-IV of antennae 0.7,
2.0, 1.3, 1.6 long. Segments II-IV of rostrum, 1.9, 1.3, 1.2 long; bucculae lacking
anterior tooth.

Pronotum 6.0 wide at humeri, 2.7 long mesially; anterolateral margins slightly
concave, narrowly reflexed; humeri not produced laterad; small tooth at anterolateral
angles slightly acute; disk rugosely punctate behind cicatrices.

Scutellum 3.8 wide at base, 4.2 long; frena extending 0.6 of distance from base of
scute! 1 uni to apex; basal disk rugosely punctate; apex slightly reflexed on each side,
black-edged where reflexed. Connexiva narrowly exposed, less densely punctate than
border of coria; punctures in macules black.

Prostemum concave; mesosternum and metastemum both carinate mesially and
conspicuously punctate. Spiracles oval. Outline of genital plates as in Parochlerus
latus (Fig. 64); basal plates convex, submarginally impressed at posterolateral angles.

Length 1 1.5

Catulona, new genus
Figs. 67-78

Type species Catulona pensa, new species.

Diagnosis. Small tooth at anterolateral angles of pronotum reflexed, directed ceph-
alad or anterolaterad. Juga as long as or slightly longer than tylus, their lateral margins
briefly parallel between anteocular concavities and apex of head; each jugum with
obtuse, rudimentary, anteocular process (Figs. 67, 73). Length of head before ocelli
0.7 of width across eyes. Eyes relatively small, width of each 0.2 of head width across
eyes. Antennae 5-segmented; basal segment reaching apex of head; segments II and
III subequal in length. Labium arising slightly behind imaginary plane bisecting head
at anterior limit of eyes; segment II reaching mesocoxae; rostral apex on stemites 4
or 5. Humeri weakly produced, scarcely emarginated. Width of scutellum at distal
end of frena slightly less than one-half of basal width. Costal angles of coria surpassing
scutellar apex, lying above penultimate abdominal segment. Metastemum thinly
carinate mesially. Second gonocoxae hidden (Figs. 72, 78).

Last abdominal tergum of male projecting mesially as thin, broad, convex process
partially covering genital cup. Proctiger with pair of long, basal processes (Figs. 70,
75). Mesial pit in pygophore at base of and ectad to inferior ridge (Figs. 68, 74).

Comments. The tergal process distinguishes this genus among ochlerines, but either
the pygophore must be removed or the hemelytra membranes displaced to see this
character.

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GENERA OF OCHLERINI

31

Figs. 67-72. Catulona pensa. 67. Head and pronotum. 68. Pygophore, caudal view. 69.
Distal part of pygophore, lateral view. 70. Genital cup. 7 1 . Pygophore, ventral view. 72. Genital
plates, caudoventral view. Symbols: inferior ridge (ir); pit (p); paramere (pa); proctiger (pr).

Catulona pensa, new species
Figs. 67-72

Description. Fuscous, usually with brownish dots and vermiform marks on dorsal
rugosities, and brownish yellow as follows: 3 basal spots on scutellum (1 mesial, 1
at each basal angle), small macule on disk of each corium, usually middle of each
visible connexival segment, basal one-fourth to all of last antennal segment, coxae
in part, extension of connexival macules as broken lateral border along abdominal
venter, cresent or spot bordering some or all of spiracles mesially. Length 10.7-1 1.0.

Vertex of head and base of tylus quite swollen (Fig. 67). Punctation on dorsum of
head fine to obscure. Juga and tylus of females subequal in length, each separately
rounded; juga exceeding tylus in male, leaving nearly square notch in apex of head.
Width of head across eyes 2. 2-2. 4, length 2.0-2. 5. Interocular distance 1.2- 1.4.
Distance across ocelli 1.25-1.4, between ocelli 1.0-1. 1. Segments I-V of antennae
0.6-0. 7, 0.8-0. 9, 0. 8-1.0, 1.2-1. 3, 1.8-2. 1 in length. Last 3 segments of rostrum 2.0-
2.3, 1.6-1. 9, 1. 5-2.0 in length.

Anterolateral pronotal margins slightly sinuous, slightly reflexed; triangular tooth
at each anterolateral angle directed cephalad, projecting past posterior margin of eye
(Fig. 67). Disk somewhat rugose. Width of pronotum at humeri 5. 8-6. 3, mesial length
2. 5-2. 8.

Basal width of scutellum 3. 7-4.0, length 4. 0-4. 6. Membranes of hemelytra dark,
with about 9 simple or furcate veins each, those toward costal margin usually faint.
Connexvia narrowly exposed.

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Evaporative area on each pleuron small, with ridges fanning out from orifice and
extending little past auricle. Mesial, longitudinal sulcus on abdominal venter broad,
shallow, extending from base over several or all stemites.

Genital plates as in Figure 72.

From caudal view, pygophoral emargination broad, semi-circular (Fig. 68). Dorsal
margin of inferior ridge evenly concave; tuft of hair projecting mesad located at each
lateral limit of inferior ridge and just entad of posterior pygophoral margin; area
between posterior pygophoral margin and inferior ridge sloping to mesial pit at base
of inferior ridge. From ventral view, pygophoral emargination shallow, slightly sin-
uous between mesial notch and posterolateral comers (Fig. 71). Proctiger bearing
long, depressed, basal process on each side, these curving posteroventrad, fringed
apically with long hairs (Fig. 69); apex of proctiger somewhat expanded and strongly
impressed; a pair of protuberances clothed in long hairs located between basal pro-
cesses and apical impression (Fig. 70). Parameres compressed, concave laterally,
convex mesially, bilobed apically; posterior lobe acute, anterior lobe broadly rounded.

Types. Holotype, 6 labeled “Brazil, Rio de Janeiro; Repressa Rio Grande, Decem-
ber 1977, M. Alvarenga.” Deposited in the Instituto de Biociencias, Universidade
Federal do Rio Grande do Sul, Porto Alegre, Brazil.

Paratypes, 499, 399 labeled “Brazil, Bahia: Encruzilhada. 960 m. Nov. 1972. M.
Alvarenga” (2 AMNH, 1 LHR) and 19 labeled “Fry Coll. 1905.100” (BMNH).

Comments. The swollen vertex of the head will separate this species from the one
other species of the genus. The swollen vertex also occurs in a few species of Lincus.

Catulona apaga, new species
Figs. 73-78

Description. Fuscous with brownish yellow as follows: at base of scutellum large
mesial spot and sometimes small spot at each basal angle, small spot on disk of each
corium and at mesial edge of each spiracle, short vermiform line on meson of pro-
notum and several such markings on abdominal venter, submarginal lateral line on
each stemite, fifth antennal segment. Length 14.0-14.3.

Neither vertex of head nor base of tylus tumescent. Dorsum of head rugosely
punctate, transversely to diagonally so on juga and tylus except apically. Juga sur-
passing tylus, somewhat convergent apically (Fig. 73). Width of head across eyes 3.0-
3. 1 , length 2. 5-2.6. Interocular distance 1 .6. Distance across ocelli 1 .45-1.5, between
ocelli 1.1. Segments I-V of antennae 0.9, 1.2-1. 3, 1 .2—1 .3-, 1.5-1. 6, 1.7 long. Last 3
rostral segments 2.5, 2.0, 1.9 long.

Anterolateral margins of pronotum shallowly concave, tooth at anterolateral angles
subtriangular, directed anterolaterad. Disk rugose, strongly punctate. Width of pro-
notum at humeri 7. 5-7. 6, length at meson 3.4-3. 5.

Scutellum 4.8 wide at base, 5. 5-5.6 long. Membranes of hemelytra dark, nearly
opaque, each with about 12 simple or furcate veins. Connexiva rather broadly ex-
posed.

Evaporative areas and basal plates similar to type species (Fig. 78). From caudal
view, posterior pygophoral margin forming obtuse, concave emargination; surface
entad of posterior margin sloping regularly to inferior ridge; deep, mesial, subtrian-
gular pit located at base of inferior ridge, a shallow sulcus extending from pit toward

1992

GENERA OF OCHLERINI

33

Figs. 73-78. Catulona apaga. 73. Head and anterior pronotal margin. 74. Pygophore, caudal
view. 75. Genital cup. 76. Posterior pygophoral margin, ventral view. 77. Distal part of py-
gophore, lateral view. 78. Genital plates, caudo ventral view. Symbols: inferior ridge (ir); pit
(p); paramere (pa); proctiger (pr).

posterior margin (Fig. 74). From ventral view, posterior pygophoral margin shallowly
and evenly concave (Fig. 76), from dorsal view slightly sinuous (Fig. 75). Proctiger
with pair of large, compressed, basal processes, each directed posterolaterad, fringed
apically with long hairs; pair of large, contiguous protuberances just distad of origin
of processes, each clothed in long hairs. Parameres compressed, lateral surface of
each concave, mesial surface unevenly convex, paramere-head entire, posterodorsal
edge forming acute tooth (Fig. 75).

Types. Holotype, 6 labeled (a) “Brasilia” and (b) “Coll. F. Salvator.” Deposited in
the National Museum of Natural History, Washington, DC.

Paratypes, 32$. 1$ labeled “Strum, Brazil” (LHR); 12 labeled (a) “Brasilia” and (b)
“Coll. F. Salvator” (SZCU); and 12 labeled “Beske Rio” (RNH).

Comments. This species differs from Catulona pensa in the moderately convex
rather than inflated vertex, in larger size and in details of the genitalia.

Cromata, new genus
Figs. 79, 80

Type species Cromata ornata, new species.

Diagnosis. Second rostral segment projecting past mesocoxae; apex of rostrum on

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

last sternite; labium arising near or on imaginary plane bisecting head at anterior
limit of eyes. Anteocular margins of head lacking well developed processes; lateral
jugal margins subparallel or parallel between anteocular concavity and apex of head;
juga as long as or longer than tylus (Fig. 79). Width of reticulated part of each eye
0.2 width of head across eyes. Length of head anterior to ocelli 0.7-0. 8 width of head
across eyes. Antennae 5 -segmented; basal segment not or scarcely exceeding apex of
head; segment II more than one-half length of segment III. Anterolateral margins of
pronotum strongly reflexed, especially toward apex; small tooth present at each
anterolateral angle. Width of scutellum at distal end of frena a little less than one-
half of basal width. Costal angles of coria surpassing apex of scutellum, lying above
penultimate abdominal segment. Metastemum thinly carinate mesially.

Comments. This genus appears most nearly related to Coranda, new genus, but
differs in the form of the head and pronotum.

Cromata ornata, new species
Figs. 79, 80

Description. Usually ochraceous with rufous borders along anterolateral margins
of pronotum, costal margins of coria basally, lateral margins of abdomen ventrally;
connexiva, femora distally and tibiae proximally also rufous, as is occasionally nearly
all of dorsum. Punctation, antennae, tibiae except at base, preapical femoral band,
large marginal spot at abdominal incisures, band on each side of transverse connexival
sutures and pair of spots on apex of scutellum all dark brown to black.

Head across eyes 2.2-2. 5 wide, 2.0-2. 5 long. Segments I-V of antennae 0.7-0. 8,
0. 7-1.0, 1. 1-1.3, 1.3-1. 6, 2. 0-2. 2 long. Segments II-IV of rostrum subequal in length,
2. 3-2. 7. Juga usually a little longer than tylus, their lateral margins sharply reflexed,
subparallel between anteocular concavities and apex of head (Fig. 79).

Anterolateral margins of pronotum slightly sinuous from dorsal view; pronotal
width at humeri 5. 7-6. 6, length mesially 2. 6-3. 2; disk divided along meson by im-
punctate line that continues for entire length of scutellum.

Scutellum 3.6-4. 1 wide at base, 4. 0-4. 6 long; fovae in basal angles small, consisting
of a few clustered punctures, isolated by an impunctate band; apex reflexed slightly
at large, dark, marginal spots. Costal margins of coria reflexed basally; large, im-
punctate lacuna present on disk of each corium; membranes lightly fumose, each
with about 7 veins. Connexiva broadly exposed.

Gonocoxae 2 tectiform where exposed (Fig. 80).

Types. Holotype, 2 labeled “Brazil, Espiritu Santos: Linhares, Sept. 1972, M. Al-
varenga.” Deposited in the American Museum of Natural History.

Paratypes, 829. 12 labeled same as holotype (AMNH); 12 labeled “Brazil, Mato
Grosso, Lat. 1 2°3 1 ' and Long. 55°37', Sinop, October, 1 974. M. Alvarenga” (AMNH);
422 labeled “Brazil, Bahia, Encruzilhada, 960 m. Nov. 1972, M. Alvarenga” (2
AMNH, 2 LHR); 12 labeled “BRAZIL: Sao Paulo, Sao Paulo, 3 February 1974, Coll.
V. N. Alin” (DAR); 12 labeled “Encruzilhada, Bahia, Brazil, XIII 1980” and “A.
Martinex e M. Alvarenga” (LHR).

Comments. Among ochlerines, this species is the most colorful. Even so it is rather
drab.

1992

GENERA OF OCHLERINI

35

Figs. 79-83. 79, 80. Cromata ornata. 79. Habitus. 80. Genital plates, caudoventral view.

8 1-83. Head and anterior part of pronotum. 8 1 . Lincus securiger. 82. L. styliger. 83. L. dentiger.
Symbol: 2nd gonocoxae (gx2).

Lincus Stal, 1867
Figs. 81-83

Lincus Stal, 1867:524; Stal, 1872:12, 14; Rolston, 1983a:2-7 (revision). (Type species
Pentatoma rufospilata Westwood, 1837, by original designation.)

Minilincus Ruckes, 1958:14-15 (synonymized by Rolston, 1983a). (Type species
Minilincus parvulus Ruckes, 1958, by original designation.)

Diagnosis. Labium originating behind imaginary plane bisecting head at anterior
limit of eyes; second rostral segment extending a little past mesocoxae; rostral apex
on or beyond penultimate stemite. Antennae 5 -segmented, basal segment reaching
or surpassing apex of head. Juga as long as or longer than tylus, sometimes contiguous
apically. Length of head before ocelli 0.6-0. 8 of width across eyes. Scutellum 0.4-
0.5 as wide at distal end of frena as at base. Coria extending past apex of scutellum.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 84-88. 84, 85. Coranda picipes. 84. Head and pronotum. 85. Pygophore, caudal view.

86-88. Coranda cast ana. 86. Head and pronotum. 87. Pygophore, caudal view. 88. Genital
cup. Symbols: inferior ridge (ir); paramere (pa); proctiger (pr).

Metastemum flat, or weakly tectiform, sometimes with weak, medial carina for only
part of its length.

Comments. Most species of Lincus are recognizable as members of the genus by
their well-developed pronotal lobes (Figs. 81, 82), although several species have
unremarkable pronotal lobes (Fig. 83). This is one of the largest ochlerine genera
with 35 described species (Rolston, 1983a, 1989; Dolling, 1984). Some of these species
are known to be of economic importance as vectors of plant diseases. The genus is
widely distributed in Central and South America.

Coranda, new genus
Figs. 84-88

Type species Melanodermus picipes Stal, 1872.

Diagnosis. Anterolateral angles of pronotum acute, projecting forward beyond base
of eyes; anterolateral margins prominently reflexed, especially toward apex (Figs. 84,
86). Second rostral segment reaching anterior limit of metacoxae; rostral apex lying
on last abdominal stemite; labium arising on or near imaginary plane bisecting head
at anterior limit of eyes. Anteocular process absent; lateral jugal margins tapering
from anteocular concavity to apex of head; tylus longer than juga. Width of reticulated
part of each eye 0.25 as wide as head across eyes; length of head before ocelli 0.7-
0.8 of head width across eyes. Antennae 5 -segmented; basal segment surpassing apex

1992

GENERA OF OCHLERINI

37

of head; segment II one-half length of segment III; width of scutellum at distal end
of frena a little less than one-half of basal width. Costal angle of each corium sur-
passing apex of scutellum, lying above penultimate abdominal segment. Metasternum
thinly carinate mesially.

Comments. Members of this genus resemble Stalius tartareus in the form of the
pronotum and size. However, Stalius is among those genera with a relatively broad
and long scutellum. Coranda seems closely related to Cromata but differs in the form
of the head and pronotum.

Coranda picipes (Stal, 1872), New Combination
Figs. 84, 85

Melanodermus picipes Stal, 1872:1 4.

Description. Dark castaneous to fuscous dorsally with many interstitial rugae, basal
one-third of last antennal segment and marginal macule on each connexival segment
light castaneous to light brownish yellow. Venter castaneous to fuscous, generously
mottled with yellowish brown. Tibiae of dark yellowish brown legs annulated with
3 fuscous bands: 1 at base, 1 at apex, and 1 intermediate, these bands increasing
progressively in width from base to apex.

Head 1.9-2. 2 wide across eyes, 1.7-1. 9 long. Interocular width 1.1-1.25; distance
across ocelli 0.85-0.95. Anterior-posterior diameter of eyes slightly greater than
width, diameter 0.5, width 0.45. Segments I-V of antennae 0.8, 0.6, 1. 1-1.2, 1.1,
1.6-1. 8 long. Segments II-IV of rostrum about 2. 5-2.6, 2. 0-2. 2, 2.0-2. 2 in length.

Pronotum 4.8 wide at humeri, 1.9-2. 2 long mesially. Cicatrices subcircularly el-
evated laterally.

Scutellum 2.9-3. 1 wide at base, 3. 2-3. 5 long; apex slightly reflexed on each side.
Distal margin of each corium slightly convex; membranes heavily fumose, each with
9 simple veins. Connexiva broadly exposed.

Mesial emargination of pygophore deep from caudal view, sinuously V-shaped.
Parameres visible from caudal view, divergent, straight, their posterior surface sulcate
(Fig. 85). Inferior ridge divided mesially, with small, triangular tooth on each side
of division.

Length 9.8.

Distribution. Colombia (Bogota), Peru (La Libertad).

Comments. The holotype of Melanodermus picipes, which now lacks a pygophore,
was examined. It is conserved in the Naturhistoriska Riksmuseet, Stockholm.

This species is transferred from Ochlerus (= Melanodermus) because the short and
subapically narrow scutellum (relative to Ochlerus ), the long rostrum, and other
characteristics are contraindicative of Ochlerus. From the other species of the genus,
C. picipes is particularly distinguished by the form of the pygophore and fuscous
coloration.

Coranda castana, new species
Figs. 86-88

Description. Brown to light castaneous dorsally; cicatrices and densely punctate
band on each side of vertex fuscous; border anterior to and laterad of cicatrices and

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

much of head dark castaneous; connexiva alternated, with yellowish median macule
traversing all or part of each segment. Abdominal venter dully mottled in shades of
brown. Legs and rostrum brownish yellow; basal segment of antennae light brown,
segments II-IV light brown (V missing).

Head 2.0 wide across eyes, 1 .9 long; length of head before eyes 0.75 of width across
eyes. Interocular width 1.05, distance across ocelli 0.85. Each eye about 0.5 wide;
anterior-posterior diameter equal to width. Segments I-IV (V missing) of antennae
0.8, 0.5, 1.1, 1.2 long. Segments II-IV of rostrum 3.2, 2.1, 2.1 long.

Pronotum 5.1 wide at humeri, 2.2 long mesially.

Scutellum 3.0 wide at base, 3.3 long; apex slightly reflexed on each side. Distal
margin of each corium slightly convex; membranes heavily fumose, each with 9
simple veins. Connexiva broadly exposed.

Dorsal margin of pygophore tumescent on each side of mesial emargination (Figs.
87, 88). Parameres with stout apical hook on anterior edge. Inferior ridge flattened,
V-shaped from dorsal view.

Length 9.7.

Type. Holotype, 6 labeled (a) “Santa Jnez [sic] (Ecuad.) R. Haensch S.,” (b) “coll.
Breddin.” Deposited in the Akademie der Landwirtshaftswissenschaften, Eberswalde.
No paratypes.

Comment. The trisinuate posterior pygophoral margin and castaneous coloration
distinguish this species from C. picipes, which has a single medial emargination in
the posterior pygophoral margin and fuscous coloration.

Brachelytron Ruckes, 1958

Brachelytron Ruckes, 1958:17-19. (Type species Brachelytron angelicas Ruckes, 1958,

by original designation.)

Diagnosis. Brachypterous, coria truncate distally above tergum 4, not reaching
broadly rounded scutellar apex above tergum 5. Ocelli present but small. Antero-
lateral angles of pronotum curving around head, projecting anteriorly beyond eyes.
Anteocular processes short, acute. Juga surpassing tylus. Antennae 5 -segmented, first
segment extending only to apex of head. Labium arising behind imaginary plane
bisecting head at anterior limit of eyes; second rostral segment reaching mesocoxae.
Metasternum carinate.

Comments. The genus is monotypic, represented so far by a single specimen from
Brazil.

Brachypterous forms occur in two other genera, Alathetus and Miopygium. Of
these, only the latter genus is sympatric. In addition to the distinguishing character-
istics given in the key, Miopygium differs from Brachelytron in having 4-segmented
antennae, the basal segment of which projects beyond the apex of the head, and a
shorter rostrum, with the second segment reaching just beyond the procoxae.

CODENS FOR PARATYPE DEPOSITION

AMNH, American Museum of Natural History, New York
BMNH, British Museum (Natural History), London
CAS, California Academy of Sciences, San Francisco

1992

GENERA OF OCHLERINI

39

CMNH, Carnegie Museum of Natural History, Pittsburgh

DAR, D. A. Rider collection

DBT, D. B. Thomas, Jr. collection

HDE, H. D. Engleman collection

LHR, L. H. Rolston collection

NMNH, U.S. National Museum of Natural History, Washington
RMB, R. M. Baranowski collection
RNH, Riksmuseum van Natuurlijke Historie, Leiden
SZCU, Systematic Zoology, Charles University, Prague.

ACKNOWLEDGMENTS

Specimens for this study were kindly loaned by R. M. Baranowski (University of Florida),
H. Brailovsky (Universidad Nacional Autonoma de Mexico), W. R. Dolling (British Museum
[Natural History]), H. Dodge Engleman, Richard C. Froeschner (U.S. National Museum of
Natural History), Jocelia Grazia (Univ. Federal do Rio Grande do Sul), Donald B. Thomas,
Jr. (USDA), Randall T. Schuh (American Museum of Natural History), Pavel Stys (Charles
University) and P. H. van Doesburg (Riksmuseum van Natuurlijke Historie).

Type material was made available by Richard C. Froeschner, P. M. Giachino (Museo Re-
gionale de Scienze Naturali, Torino), A. Kaltenbach (Naturhistorisches Museum Wien), Per
Lindskog (Naturhistoriska Riksmuseet), G. Petersen (Akademie der Landwirtschaftswissen-
schaften der D.D.R.) and Randall T. Schuh.

Prof. Carlo Vidano (Universita Delgi Studi) attempted to examine the relevant material in
the Spinola collection before the collection was moved to the Museo Regionale di Scienze
Naturali, Torino. His efforts are appreciated.

Several individuals reviewed the manuscript at various stages of its development and offered
constructive criticism and suggestions: J. B. Chapin, J. E. Eger, Jr., and Donald B. Thomas, Jr.
Their comments were valuable. I am especially indebed to D. A. Rider for his considerable
contributions, and to J. P. Woodring for translation from German.

To all of these colleagues I am grateful. To the curators who unsuccessfully searched their
collections for Breddin type material, I am also grateful.

Approved for publication by the Director of the Louisiana Agricultural Experiment Station
as manuscript number 90-17-4437.

LITERATURE CITED

Amyot, C. J. B. and J. G. Audinet-Serville. 1843. Historie naturelle des insects. Hemiptera.
Paris.

Breddin, G. 1903a. Neue Rhynchoten aus den Anden. Soc. Ent. 18:122-124.

Breddin, G. 1 903b. Beitrage zur Hemipterenfauna der Anden. Sitz.-ber. Ges. naturf. Fr. Berlin
(19Q3):366-383.

Breddin, G. 1904. Neue Rhynchotenausbeute aus Siid-Amerika. Soc. Ent. 18(20): 153-1 54.
Breddin, G. 1910. Beitrage zur Systematik der Pentatomiden Sudamericas. Vierte Stuck. Sitz.-
ber. Ges. naturf. Fr. Berlin (1909):6 15-631.

Breddin, G. 1912. Zwei neue neotropsche Pentatomiden-Gattungen (Hem.) Arch. Naturges.
78 Abt. A(6):90-93.

Dallas, W. S. 1851. List of the Specimens of Hemipterous Iinsects in the Collection of the
British Museum. Part 1. London.

Distant, W. L. 1899. XLIX— Rhynchotal notes. III. Heteroptera: Discocephalinae and Pen-
tatominae (part). Ann. Mag. Nat. Hist. (7)4:421-445.

40

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Distant, W. L. 1911. Rhynchotal notes, liii. Neotropical Pentatomidae. Ann. Mag. Nat. Hist.
(8)7:242-258.

Dolling, W. R. 1984. Pentatomid bugs (Hemiptera) that transmit a flagellate disease of cul-
tivated palms in South America. Bull. Entomol. Res. 74:473-476.

Fabricius, J. C. 1803. Systema Rhyngotorum. Brunsvigae.

Froeschner, R. C. 1981. Heteroptera or true bugs of Ecuador: a partial catalog. Smithsonian
Contr. Zool. No. 322. Washington.

Herrich-Schaffer, G. A. W. 1837-1839, 1842-1844. Die Wanzenartigen Insecten. Vol. 4
(1837-1839), Vol. 7 (1842-1844). Numberg.

Kirkaldy, G. W. 1909. Catalogue of the Hemiptera (Heteroptera). Vol. 1. Cimicidae. Berlin.
Kormilev, N. A. 1 949. Dos especies neuvas del genero Adoxoplatys Breddin ( 1 903) de Bolivia
(Hemiptera, Pentatomidae). Notas Mus. La Plata 14(Zool. no. 1 28):3 1 3—324 + 2 figs.
Kormilev, N. A. 1950. Notes on neotropical Pentatomidae, with description of one new genus
and two new species (Hemiptera). Rev. Brasil. Biol. 10:339-346.

Kormilev, N. A. 1955. Notas sobre Pentatomoidea neotropicales II (Hemiptera). Acta Sci.

Instituos Investigacion San Miguel, Cuademo no. 1:3-16.

Kormilev, N. A. 1956. Notas sobre Pentatomoidea neotropicales V (Hemiptera). Acta Sci.

Institutes Investigacion San Miguel, Cuademo no. 4:3-7.

Rolston, L. H. 1981. Ochlerini, a new tribe in Discocephalinae (Hemiptera: Pentatomidae).
J. New York Entomol. Soc. 89(1):40^42.

Rolston, L. H. 1 982. A brachypterous species of Alathetus from Haiti (Hemiptera: Pentatom-
idae), J. Kansas Entomol. Soc. 55(1): 156-1 58.

Rolston, L. H. 1983a. A revision of the genus Lincus Stcil (Hemiptera: Pentatomidae: Dis-
cocephalinae: Ochlerini). J. New York Entomol. Soc. 9 1(1): 1-47.

Rolston, L. H. 1983b. The genus Paralincus (Hemiptera: Pentatomidae). J. New York En-
tomol. Soc. 9 1(2): 183-1 87.

Rolston, L. H. 1989. Three new species of Lincus (Hemiptera: Pentatomidae) from palms. J.
New York Entomol. Soc. 97(3):27 1-276.

Rolston, L. H. and F. J. D. McDonald. 1979. Keys and diagnoses for the families of Western
Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae
(Hemiptera). J. New York Entomol. Soc. 87(3): 189-207.

Ruckes, H. 1958. New genera and species of neotropical discocephaline and halyine penta-
tomids (Heteroptera, Pentatomidae). Am. Mus. Novitates no. 1868, 27 pp.

Ruckes, H. 1959. New genera and species of pentatomids from Panama and Costa Rica
(Heteroptera, Pentatomidae). Am. Mus. Novitates no. 1939, 18 pp.

Ruckes, H. 1960. Three new New World halyine pentatomids (Hemiptera: Pentatomidae). J.
New York Entomol. Soc. 68:225-231.

Sailer, R. I. 1950. Alitocoris, a new genus of Pentatomidae (Hemiptera). Proc. Entomol. Soc.
Washington 52(2):69-76.

Spinola, M. 1837. Essai sur genres d’insectes appartenants a l’ordre des Hemipteres L., ou
Rhyngotes Fabr. et a la section des Heteropteres. Paris.

Spinola, M. 1850a. Tavola sinottica dei genera spettanti alia classe degli insetti arthrodignati
Hemiptera Linn., Latr., Rhyngota Fabr., Rhynchota Burm. Modena 138 pp.; reprinted
1852 in Mem. Nat. Fis. Soc. Ital. Modena 25(part 1):43-100.

Spinola, M. 1850b. Di alcuni generi d’insetti arthrodignati nouvamente propositi. Pages 61-
138; reprinted 1852 in Mem. Mat. Fis. Soc. Ital. Modena 25(part 1 ): 1 0 1—1 78.

Stcil, C. 1860. Bidrag till Rio Janeiro-traktens Hemipter-fauna. K. Svenska Vet.-Ak. Handl.
2(7): 1-84.

Stcil, C. 1862. Bidrag till Rio Janeiro-traktens Hemipter-fauna. 2. K. Svenska Vet.-Ak. Handl.
3(6): 1-75.

StM, C. 1867. Bidrag till Hemiptererans systematik. Conspectus generum Pentatomidum
Americae. Ofv. K. Svenska Vet.-Ak. Forhl. 24(7):522-534.

1992

GENERA OF OCHLERINI

41

St&l, C. 1872. Enumeratio Hemipterorum. 2. Enumeratio Cimicinorum Americae. K. Svenska
Vet.-Ak. Handl. 10(4):3-65.

Walker, F. 1867. Catalogue of the Specimens of Hemipera Heteroptera in the Collection of
the British Museum. Vol. 1. London.

Westwood, J. O. 1837. In: F. W. Hope. A Catalogue of Hemiptera in the Collection of the
Rev. F. W. Hope, M.A. with Short Latin Diagnoses of the New Species. Part 1.

Received 30 October 1990; accepted 27 February 1991.

J. New York Entomol. Soc. 1 00( 1 ):42— 98, 1992

REVISION OF THE GENUS THYANTA STAL, 1862
(HETEROPTERA: PENTATOMIDAE)

II. NORTH AMERICA, CENTRAL AMERICA,

AND THE WEST INDIES

D. A. Rider1 and Joan B. Chapin

Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University Agricultural Center, Baton Rouge, Louisiana 70803;

1 present address, Department of Entomology,

North Dakota State University, Fargo, North Dakota 58105

Abstract.— The species of the pentatomid genus Thyanta St<U occurring north of South Amer-
ica are revised. All northern hemisphere species belong to the subgenera Thyanta and Argosoma
Rider. Diagnoses are given for the genus, subgenera, and the 14 included species and subspecies.
The subspecies accerra McAtee is transferred from T. pallidovirens (St&l) to T. custator (Fa-
bricius). The following new synonymy is recognized (junior synonym in parentheses): T. custator
accerra McAtee, 1919 (=T. pallidovirens spinosa Ruckes, 1957a), T. pallidovirens (Stdl, 1859)
(=T. pallidovirens setosa Ruckes, 1957a), and T. maculata (Fabricius, 1775) (=T. casta StM,
1862b). Lectotype designations are made for Cimex maculatus Fabricius, Pentatoma obsoleta
Dallas, P. pallidovirens, T. casta, and T. pseudocasta Blatchley. Thyanta planifrons Ruckes is
reported from the United States for the first time. A key is provided for the subgenera, species,
and subspecies that occur north of South America.

The genus Thyanta is restricted to the western hemisphere and is one of several
fairly large genera in the nominate tribe and subfamily of the Pentatomidae. His-
torically, identifications in this genus have been difficult. In fact, Jensen-Haarup
(1928) commented “The species are very variable, but at the same time inter se
much similar both as to colour and sculpture, and also regarding the genital segments;
a strict separation of most of the species is, therefore, rather difficult, if not rich and
fresh materials of species and specimens are at hand.” There has been no recent
treatment of the entire genus, the last being that of Jensen-Haarup in 1928. Malloch
(1919), Blatchley (1926), and Torre- Bueno (1939) all reviewed portions of the genus,
but their works have become outdated largely due to the addition of many new
species and subspecies (Ruckes, 1952, 1956, 1957a, b, c; Rider and Chapin, 1991).

The genus Thyanta is one of a group of pentatomine genera characterized by the
lack of a spine or tubercle on the third (second visible) abdominal stemite. Rolston
and McDonald (1984) provided a key to separate those genera occurring in the
Western Hemisphere north of South America. Because there are many species and
many of them are difficult to distinguish, the genus has been divided into two groups
according to geographical regions. The present paper reviews those species known
to occur north of South America. The South American species of Thyanta were
recently revised by Rider and Chapin (1991).

Care should be maintained when using the key to species. It is important to have
fresh, mature specimens. After death, specimens tend to become greasy and discol-
ored, making their determinations rather tentative. In the key, when certain characters

1992

THYANTA OF NORTH AND CENTRAL AMERICA

43

are described as black or piceous, the true structural color is black, not just darkened
due to discoloration. Teneral specimens and specimens of brown forms often have
darkened areas that may be misleading when working through the key. Most char-
acters of the genitalia can be seen without dissecting the specimen; however, accurate
determinations may require some dissection. In many cases there are no reliable
characters to separate females. The only way to identify females in these instances
is by association with males or sometimes by geographical distribution.

When label data is cited in the text, each letter in parentheses represents a different
label with (a) being closest to the specimen. Museum acronyms used in the text are
defined in the acknowledgements. All measurements are in millimeters. Measure-
ments in parentheses are of the holotype.

Thy ant a Stal 1862

Thyanta St&l, 1862a:58; Stal, 1867:529; Stal, 1872:34-35; Distant, 1880:65; Sum-
mers, 1898:45; Kirkaldy, 1909:94; VanDuzee, 1917:51; Blatchley, 1926:104, 112-
1 13; Jensen-Haarup, 1928: 185-186; Furth, 1974:21-22; Froeschner, 198 1:7 ^Mc-
Pherson, 1982:48, 76-77; Rolston and McDonald, 1984:74, 76; Froeschner, 1988:
592.

Type species. Cimex perditorF., 1794 (by subsequent designation, Kirkaldy, 1909:
XXX).

Diagnosis. Third (second visible) abdominal stemite lacking medial spine or tu-
bercle. Each ostiolar ruga sulcate proximally, reaching at least three-fourths distance
from mesial margin of ostiole to lateral margin of metapleuron. Bucculae evanescent
or arcuately truncate at posterior terminations. Juga and tylus usually subequal in
length; rostrum reaching at least to metacoxae. Femora unarmed; superior surface
of each tibia usually sulcate. Width of scutellum at distal end of frena two-fifths or
less basal scutellar width. Each paramere narrowly rounded to acute apically, lacking
denticles, usually lacking lateral lobe, rarely with spinose lateral lobe.

Comments. The genus Thyanta is closely related to two other pentatomine genera,
Cyptocephala Berg and Tepa Rolston and McDonald, from which it can be separated
reliably only by differences in the male genitalia. In all northern hemisphere species
of Thyanta, the head of each paramere lacks a lateral lobe. In Cyptocephala and
Tepa, the parameres have well-developed lateral lobes. Cyptocephala further differs
from Tepa and Thyanta by having minute denticles between the lateral lobe and the
apex of the paramere. The parameres of species of Tepa and Thyanta lack denticles.

Jensen-Haarup (1928) described the subgenus Parathyanta within Thyanta. Rol-
ston and McDonald (1984) placed Parathyanta in the synonymy of Cyptocephala.
At the same time, they transferred 4 species from Thyanta to Cyptocephala and 6
species from Thyanta to Tepa. The species of both Cyptocephala and Tepa have
been reviewed recently (Rolston, 1972, 1986; Rider, 1986a).

The genus Thyanta is divided into three subgenera: Argosoma Rider, Phacidium
Breddin, and Thyanta. All 8 species of the subgenus Phacidium are restricted to
South America and have been treated in an earlier paper (Rider and Chapin, 1991).
Argosoma contains 20 species, 6 of which are known to occur outside South America.
The nominate subgenus contains 9 species of which 7 are known to occur in the area
covered by the present paper.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

KEY TO SUBGENERA OF THYANTA STAl

1. Dorsal surface relatively shiny, glossy; punctures relatively large, sparse; pygophoral
opening relatively large; posteroventral surface of pygophore produced into small,
blunt, chin-like protuberance; ectal surface of each paramere concave; spermathecal

bulb globose Argosoma Rider

Dorsal surface appearing matte, not shiny; punctures relatively small, dense; pygo-
phoral opening relatively small, posteroventral surface of pygophore not produced
into blunt, chin-like protuberance; ectal surface of each paramere convex; sperma-
thecal bulb globose or digitiform 2

2(1). Pygophoral opening subtended by semicircular or rectangular impression; posterior
margin of pygophore with medially emarginate production in middle; theca large,
subtriangular, with small protuberance on each side near lateral dorsal margin;

proximal end of sclerotized rod cone-shaped; spermathecal bulb digitiform

Thyanta St&l

- Posteroventral surface of pygophore smoothly arcuate or with distinct sulcus, not
with semicircular or rectangular impression; posterior margin of pygophore nearly
transverse, lacking medial production; theca reniform, lacking protuberances on
lateral dorsal margin; proximal end of sclerotized rod may be slightly swollen, but
not cone-shaped; spermathecal bulb globose Phacidium Breddin

KEY TO SPECIES OF THYANTA OCCURRING NORTH OF SOUTH AMERICA

1 . Anterolateral margins of pronotum piceous; mesial angle of each pronotal cicatrice

black 2

Anterolateral margins of pronotum not piceous; coloration of mesial angle of each
pronotal cicatrice variable, often immaculate 3

2(1). Piceous markings along anterolateral pronotal margins relatively broad, usually
easily visible from dorsal view; pygophoral opening subtended by semicircular
impression; general form relatively broad (eastern U.S.) calceata (Say)

- Piceous markings along anterolateral pronotal margins relatively narrow, not easily

visible from dorsal view; pygophoral opening subtended by rectangular impres-
sion; general form relatively narrow (coastal plain from central Louisiana to Flor-
ida to New York) custator custator (Fabricius)

3(1). Humeral angles rounded to angulate, but never spinose 4

Humeral angles distinctly spinose 6

4(3). Dorsal punctation minute, dense, surface appearing matte; pygophoral opening
subtended by rectangular impression; ectal surface of each paramere convex, lack-
ing dorsomedial concave surface (Fig. 34); spermathecal bulb digitiform (Fig. 47)

5

Dorsal punctation coarse, sparse, surface glossy; posteroventral surface of pygo-
phore produced into blunt chin-like protuberance; each paramere with dorso-
medial surface concave (Fig. 126); spermathecal bulb globose (Fig. 136) 10

5(4). Postspiracular black spot usually present on each side of each abdominal stemite
and usually larger in diameter than adjacent spiracle; if absent or smaller than

spiracle, then humeral angles usually angulate (eastern U.S.)

custator accerra McAtee (part)

- Postspiracular area of each abdominal stemite immaculate; or, if black spot pres-

ent, then spot is usually smaller in diameter than adjacent spiracle; humeral angles
rounded, never angulate (western U.S.) pa/lidovirens (St&l)

6(3). Mesial angle of each pronotal cicatrice often marked with black; posterolateral
angle of each abdominal sternite usually black; pygophoral opening subtended by

1992

THYANTA OF NORTH AND CENTRAL AMERICA

45

semicircular impression, posterior margin with distinct medial slit (Fig. 71) .... 7

Without above black markings; pygophoral opening subtended by rectangular
impression, posterior margin straight, concave, or sinuous, without medial slit
(Fig. 41) (southwestern U.S.; Mexico; Guatemala) . . . custator accerra McAtee (part)

7(6). Complete or partial transhumeral reddish band usually present; both anterolateral

and posterolateral angles of each abdominal sternite usually piceous 8

- Reddish band between humeral angles usually absent; each abdominal sternite
with anterolateral angle immaculate, each posterolateral angle with at most a small

black spot 9

8(7). Humeral angles directed anterolaterad, usually approaching a 45-degree angle with
longitudinal axis of body (Fig. 63) (southern U.S. to northern Argentina)

perditor (Fabricius)

- Humeral angles directed primarily laterad and only slightly anterad (Fig. 78) (Baja

California, Mexico) spectabilis Ruckes

9(7). Yellow anterolateral pronotal margins contrasting with rest of pronotum; pronotal
cicatrices immaculate; usually smaller than 9.0 mm long by 6.0 mm wide (Cuba;
British West Indies) cubensis Barber & Bruner

- Anterolateral pronotal margins concolorous with rest of pronotum; mesial angle

of each pronotal cicatrice usually black; usually larger than 9.0 mm long by 6.0
mm wide (Revillagigedo Islands, Mexico) serratulata Ruckes

10(4). Southwestern U.S.; Mexico; Central America 11

Southern Florida; West Indies 12

11(10). Ostiolar canal narrowed at middle, becoming slightly wider towards apex (Fig.

137); posterolateral angle of each abdominal sternite usually immaculate; apex of
each paramere nearly spinose in medial view (Fig. 1 24) (southwestern U.S.; north-
western Mexico) planifrons Ruckes

- Ostiolar canal acuminate apically; posterolateral angle of each abdominal sternite

piceous, sometimes only a minute portion of the tip piceous; apex of each paramere
narrowly rounded in medial view (Fig. 140) (southern Texas; Mexico; Central
America) maculata (Fabricius)

12(10). Each paramere in medial view with apex straight or bending slightly ventrad (Fig.

155), concave surface oriented more mediad than dorsad; posteromesial angles
of basal plates distinctly excavated, resulting concavity broadly U-shaped (Fig.

165) (Trinidad and Tobago; Venezuela) vadosa Rider

- Each paramere in medial view with apex curving slightly dorsad, orientation of

dorsomedial concave surface variable; posteromesial angles of basal plates round-
ed or weakly emarginate 13

13(12). Concave surface of each paramere oriented more mediad than dorsad (southern

Florida) pseudocasta Blatchley

- Concave surface of each paramere oriented more dorsad than mediad 14

14(13). Apex of each paramere rounded in ectal view (Fig. 186); spermathecal duct with

relatively small amount of swelling and coiling below proximal flange (Fig. 197)
(Greater Antilles) obsoleta (Dallas)

- Apex of each paramere spinose in ectal view (Fig. 201); spermathecal duct with

large amount of swelling and coiling below proximal flange, swelling carrot-shaped
(Fig. 2 1 2) (Lesser Antilles; northern South America) testacea (Dallas)

Subgenus Thyanta Stal

Diagnosis. Punctures minute, dense. Posterior terminations of bucculae evanescent.
Anterolateral pronotal margins straight to concave, sometimes marked with piceous;

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

humeral angles rounded to angulate, often spinose; pronotal cicatrices sometimes
marked with piceous in mesial angles. Ostiolar canals acuminate apically. Superior
surface of each tibia sulcate.

Posterior margins of basal plates sinuous, posteromesial angles entire (Fig. 1 3).
Distal end of sclerotized rod cone-shaped (Fig. 14); spermathecal bulb digitiform;
cylindrical structure below proximal flange (Fig. 15). Pygophoral opening small,
subtended on posteroventral surface by a rectangular or semicircular impression;
posterior margin of pygophore straight to concave in caudal view, with medially
incised protuberance in middle (Fig. 9). Each paramere F-shaped, obtuse protuber-
ance on shaft usually prominent, apex spinose, ectal surface convex (Fig. 3), roughened
spiculate area on lateral surface linear (Fig. 5). Each lateral conjunctival lobe of
aedeagus with single spinose diverticulum (Fig. 6); dorsomedial conjunctival lobe
usually well-developed (Fig. 7), theca large; subtriangular in lateral view, with dor-
solateral protuberance on each side near caudal limit (Fig. 8); medial penial lobes
and penisfilum moderate in size.

Comments. Species of the subgenus Thyanta have the pygophoral opening sub-
tended by a semicircular or rectangular impression, and the posterior margin is
distinctly emarginate medially. Species of Phacidium have the posteroventral surface
of the pygophore arcuately rounded or sulcate, and the posterior margin is not
emarginate medially. The posteroventral surface of the pygophore in species of Ar-
gosoma is produced into a blunt, chin-like protuberance. The ectal surface of each
paramere is concave in Argosoma and convex in both Phacidium and Thyanta.

The female genitalia are also useful in separating species of Thyanta and Phacidium.
In Thyanta, the distal end of the sclerotized rod is cone-shaped, and the spermathecal
bulb is digitiform. In Phacidium, the distal end of the sclerotized rod is swollen
subapically and narrowed distally, and the spermathecal bulb is globose. The female
genitalia of both Phacidium and Argosoma are very similar, but females can usually
be separated by dorsal punctation which is relatively dense in Phacidium and less
dense and more coarse in Argosoma.

Thyanta ( Thyanta ) calceata (Say)

Figs. 1-15, Map 1

Pentatoma calceata Say, 1831:8.

Thyanta custator (of authors, not Fabricius): Herrich-Schaffer, 1844:96, 106, fig. 771;
Uhler, 1886:7 (part); Lethierry and Severin, 1893:148 (part); Kirkaldy, 1909:94
(part); Banks, 1910:90 (part).

Thyanta custator calceata: Uhler, 1872:399.

Thyanta calceata: Barber, 191 1:108-1 1 1; Van Duzee, 1917:53; Hart, 1919:184,217,
fig. 70; Blatchley, 1926:1 13, 1 17-1 18; Torre-Bueno, 1939:231; Ruckes, 1957a:21-
22; Hoffman, 1971:44; Furth, 1974:22, 23-24; McPherson, 1982:76-77, 79-80,
fig. 72; Rolston and McDonald, 1984:figs. 23, 27; Froeschner, 1988:593.

Diagnosis. General form broad, ovate. Transhumeral rubiginous band usually pres-
ent; vertex of head and tylus often reddish.

Lateral jugal margins nearly parallel for middle third of distance from eyes to apex
(Fig. 2). Humeral angles rounded to angulate; anterolateral pronotal margins piceous,
straight to weakly concave in dorsal view (Fig. 1); mesial angle of each pronotal

1992

THYANTA OF NORTH AND CENTRAL AMERICA

47

cicatrice piceous. Each abdominal stemite with postspiracular spot on each side and
posterolateral angles piceous.

Mesial margins of basal plates in caudoventral view slightly concave; posterior
margins sinuous; posteromesial angles rounded (Fig. 13). Posterolateral angles of
pygophore continuing onto posteroventral surface as vague carinae, forming semi-
circular impression in caudoventral view. Posterior margin of pygophore produced
posterodorsad, in ventral and dorsal views appearing convex with small, medial,
V-shaped emargination (Figs. 10, 1 1); slightly concave in lateral view (Fig. 12).

Types. The type specimen of Pentatoma calceata is apparently no longer in exis-
tence, and Say’s original description (1831) will lit equally well for either T. calceata
or T. custator. Previous usage has, however, fixed both species, and therefore des-
ignation of a neotype is not necessary.

Distribution. Eastern U.S. (Map 1).

Specimens examined. 1,105 specimens collected during every months of the year;
deposited in AMNH, ARH, AUA, CAS, CNC, CUC, DAR, DBT, EGER, FMNH,
FSCA, INHS, ISU, LRH, LSU, MSU, MSUE, NCSU, NDSF, OSUC, POLH, PUL,
SIUC, SMEK, TAMU, UAT, UCB, UCR, UCS, UGA, UIM, UMC, USNM, VPI,
WSU. UNITED STATES: Alabama: Autauga, Barbour, Bibb, Cherokee, Chilton,
Clay, Cleburne, Coosa, Covington, Dallas, De Kalb, Elmore, Houston, Lee, Lowndes,
Macon, Madison, Perry, Randolph, Shelby, Tallapoosa, Wilcox, Winston. Arkansas:
Benton, Boone, Carroll, Grant, Hempstead, Hot Spring, Logan, Marion, Nevada,
Newton, Polk, Sevier, Union, Yell. Connecticut: Litchfield, New Haven. District of
Columbia. Florida: Collier, Holmes, Taylor. Georgia: Bartow, Bibb, Bryan, Chatham,
Clark, Cobb, De Kalb, Lloyd, Lranklin, Lulton, Habersham, Hall, Harris, Johnson,
Madison, Marion, Oconee, Oglethorpe, Paulding, Peach, Rabun, Sumter, Talbot,
Thomas. Illinois: Champaign, Lranklin, Hancock, Hardin, Jackson, Johnson, Pope,
Union, Vermilion, Williamson. Indiana: Crawford, Howard, Porter, Tippecanoe. Iowa:
Henry, Van Buren. Kansas: Bourbon, Douglas. Kentucky: Barren, Carter, Green,
Laurel, Trigg. Louisiana: Allen, Acadia, Beauregard, Caddo, Calcasieu, Claiborne,
De Soto, East Baton Rouge, East Leliciana, Grant, Jefferson Davis, Natchitoches,
Rapides, Sabine, St. Landry, Tangipahoa, Vernon, Webster, West Leliciana, Winn.
Maryland: Allegany, Anne Arundel, Calvert, Charles, Lrederick, Montgomery, Prince
Georges. Massachusetts: Bristol. Michigan: Wayne. Mississippi: Adams, Amite, Co-
piah, Forrest, Franklin, Jones, Lauderdale, Leake, Lincoln, Monroe, Newton, Oktib-
beha, Panola, Perry, Pontotoc, Tallahatchie, Walthal. Missouri: Barry, Benton, Boone,
Butler, Callaway, Camden, Carter, Crawford, Dade, Douglas, Greene, Harrison, Jack-
son, Laclede, Marion, McDonald, Montgomery, Osage, Ozark, Pettis, Phelps, Ralls,
Randolph, St. Charles, St. Francois, Scotland, Scott, Shannon, Stone, Taney, Wright.
New Jersey: Bergen, Gloucester, Morris. New York: Albany, Erie, Nassau, Orange,
Suffolk, Ulster, Westchester. North Carolina: Buncombe, Columbus, Craven, Cum-
berland, Currituck, Harnett, Johnston, McDowell, Macon, Montgomery, Moore, Or-
ange, Pender, Scotland, Wake. Ohio: Adams, Fairfield, Gallia, Hocking, Licking,
Pickaway, Tuscarawas, Washington. Oklahoma: Craig, Latimer, McCurtain, Ottawa.
Pennsylvania: Franklin, Lancaster, Philadelphia, Westmorland. South Carolina: Ab-
beville, Charleston, Chesterfield, Dorchester, Florence, Greenville, Horry, Lexington,
Oconee, Pickens, Spartanburg, Williamsburg. Tennessee: Cumberland, Decatur, De
Kalb, Fentress, Knox, Lauderdale, Marion, Rhea, Scott, Sevier. Texas: Anderson,

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 1-15. T. calceata. 1. Habitus. 2. Head. 3-5. Right paramere. 3. Medial view. 4. Ectal
view. 5. Lateral view. 6-8. Theca and related structures. 6. Ventral view. 7. Dorsal view. 8.
Lateral view. 9-12. Pygophore. 9. Caudal view. 10. Ventral view. 1 1. Dorsal view. 12. Lateral

1992

THYANTA OF NORTH AND CENTRAL AMERICA

49

Angelina, Aransas, Bowie, Brazos, Camp, Cherokee, Dallas, Harris, Harrison, Jasper,
Lamar, Sabine, Smith, Walker. Virginia: Charolette, Chesterfield, Clifton Forge City,
Fairfax, Falls Church, Floyd, Gloucester, Hanover, King & Queen, Montgomery,
Nelson, New Kent, Norfolk, Pulaski, Virginia Beach, Wythe. West Virginia: Monroe,
Roane, Upshur.

Comments. Thyanta calceata can be separated from all other congeners except T.
custator custator by the distinctly black anterolateral pronotal margins. It can be
reliably distinguished from T. c. custator only by the male genitalia. In T. calceata,
the pygophoral opening is subtended by a semicircular impression; this impression

view. 1 3. Genital plates, caudoventral view. 14. Spermatheca. 1 5. Spermathecal pump. Symbols:
bp, basal plate; cyl, cylindrical structure below proximal flange; dfl, distal flange; dmc, dorso-
medial conjunctival lobe; dsp, dilation of spermatheca; gx2, second gonacoxae; jug, juga; lcl,
lateral conjunctival lobe; mpl, median penial lobe; pen, penisfilum; pfl, proximal flange; pla,
posterolateral angle of pygophore; pmp, posterior margin of pygophore; pt8, eighth paratergite;
pt9, ninth paratergite; rsa, roughened spiculate area on lateral surface of paramere; spb, sper-
mathecal bulb; sr, sclerotized rod; slO, tenth stemite; th, theca; tyl, tylus.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

is rectangular in T. c. custator. Specimens of T. calceata are generally shorter and
broader than specimens of T. c. custator, and the black markings on the anterolateral
pronotal margins are usually darker and more extensive in T. calceata. These last
two characters, however, may be apparent only when a series of specimens of each
species can be compared.

Most species of Thy ant a occur in two color forms, a green form in the warmer
months and a brown form in the cooler months. This is particularly well documented
in T. calceata (McPherson, 1977a, b, 1978a, b; McPherson and Paskewitz, 1982;
McPherson et al., 1983).

Thyanta ( Thyanta ) custator custator (Fabricius)

Figs. 16-30, Map 2

Cimex custator Fabricius, 1803:164.

Pentatoma custator: Dallas, 1851:251; Walker, 1867:288.

Thyanta custator: Stal, 1862a:58; Stal, 1872:34-35 (part); Uhler, 1872:399 (part);
Uhler, 1876:289-290 (part); Uhler, 1886:7 (part); Van Duzee, 1904:53 (part);
Kirkaldy, 1909:94 (part); Van Duzee, 1909: 155-1 56; Banks, 19 10:90 (part); Barber,
1911:108-111 (part); Barber, 1914:523; Van Duzee, 1917:52 (part); Blatchley,
1926:1 13, 1 15-1 16 (part); Torre-Bueno, 1939:231 (part); Ruckes, 1957a: 1-2, 4-
13, 20, 22, fig. 1; Ueshima, 1963:149, 152-153; Hoffman, 1971:44-45; Furth,
1974:22, 23, fig. 45; McPherson, 1982:76-77, 80-81, fig. 73; Rolston and Mc-
Donald, 1984:figs. 24, 28; Froeschner, 1988:593.

Thyanta accerra (of authors, not McAtee): Blatchley, 1926:1 14, 118.

Thyanta casta (of authors, not Stal): Blatchley, 1926:1 13, 1 16-1 17.

Diagnosis. Dorsal surface green to brown, often with varying amount of reddish
coloration between humeral angles, sometimes extending along anterolateral pronotal
margins and onto basal margin of each corium.

Lateral jugal margins sinuous, not parallel (Fig. 17). Anterolateral margins of
pronotum and mesial angle of each pronotal cicatrice piceous. Anterolateral pronotal
margins slightly concave in dorsal view; humeral angles rounded to angulate, never
spinose (Fig. 16). Ventral surface yellow-green to brown, posterolateral angles of
abdominal stemites black. Each abdominal stemite with postspiracular black spot
on each side.

Mesial margins of basal plates nearly straight in caudoventral view; posterior
margins sinuous; posteromesial angles narrowly rounded (Fig. 28). Each posterolat-
eral angle of pygophore continuing onto posteroventral surface as vague carina,
forming rectangular impression; mesial portion of posterior margin slightly convex
with small, V-shaped, emargination in middle in both ventral and dorsal views (Figs.
25, 26); pygophore slightly concave in lateral view (Fig. 27).

Figs. 16-30. T. custator custator. 16. Habitus. 17. Head. 18-20. Right paramere. 18. Medial
view. 19. Ectal view. 20. Lateral view. 21-23. Theca and related structures. 21. Ventral view.
22. Dorsal view. 23. Lateral view. 24-27. Pygophore. 24. Caudal view. 25. Ventral view. 26
Dorsal view. 27. Lateral view. 28. Genital plates, caudoventral view. 29. Spermatheca. 30.
Spermathecal pump.

21

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Types. The type specimen of Cimex custator is apparently no longer in existence
(Zimsen, 1964), and the original description does not adequately fix the species.
Fabricius’ description will fit either T. custator custator and T. calceata equally well.
Previous usage, however, has fixed both species, and therefore the designation of a
neotype is not necessary.

Distribution. Coastal plain from central Louisiana to Florida to New York, and
the Bahama Islands (Map 2).

Specimens examined. 1,243 specimens collected during every month of the year;
deposited in AMNH, ARH, AUA, BMNH, CAS, CNC, CUC, DAR, DBT, FMNH,
FSCA, INHS, ISU, LHR, LSU, MSU, MSUE, NCSU, OSUC, PUL, SIUC, SMEK,
TAMU, UAT, UCB, UCR, UCS, UGA, UIM, UMC, USNM, UUSL, WSU. UNIT-
ED STATES: Alabama: Baldwin, Barbour, Covington, Dallas, Escambia, Henry,
Houston, Lee, Macon, Madison, Mobile. Florida: Alachua, Bay, Brevard, Broward,
Charlotte, Clay, Collier, Columbia, Dade, De Soto, Dixie, Duvall, Franklin, Gadsden,
Glades, Gulf, Hamilton, Hardee, Hendry, Highlands, Hillsborough, Holmes, Indian
River, Jackson, Lake, Lee, Leon, Levy, Liberty, Madison, Manatee, Marion, Martin,
Nassau, Okaloosa, Orange, Osceola, Palm Beach, Pasco, Pinellas, Polk, Putnam, St.
Johns, St. Lucie, Santa Rosa, Sarasota, Seminole, Sumter, Suwanee, Taylor, Volusia,

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THYANTA OF NORTH AND CENTRAL AMERICA

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Wakulla, Washington. Georgia: Berrien, Brantley, Brooks, Charlton, Chatham, Clinch,
Decatur, Evans, Glynn, Grady, Jefferson, McIntosh, Peach, Pierce, Sumter, Thomas,
Tifton, Toombs, Ware. Louisiana: Acadia, Ascension, Avoyelles, East Baton Rouge,
East Feliciana, Iberville, Jefferson, Livingston, Orleans, St. Charles, St. James, St.
Mary, St. Tammany, Tangipahoa, Washington, West Baton Rouge. Mississippi:
Covington, Forrest, Hancock, Harrison, Jackson, Jefferson Davis, Pike. New Jersey:
Burlington, Cape May, Monmouth, Ocean. New York: Suffolk. North Carolina: Bla-
den, Brunswick, Columbus, Duplin, Hyde, Moore, New Hanover, Onslow, Pender.
South Carolina: Aiken, Bamburg, Barnwell, Beaufort, Charleston, Florence, Green-
ville, Horry, Jasper, Kershaw, Marion, Orangeburg, Richland. Virginia: Norfolk. Ba-
hama Islands: South Bimini Island.

Comments. This species has been the subject of much confusion in the past. At
one time the name T. custator was applied to nearly all specimens from the entire
United States. Ruckes (1957a) showed that the true custator form is confined to the
coastal plain from Louisiana to Florida to New York, but he felt that it was a separate
species distinct from the accerra form. The two forms have virtually identical gen-
italia. They cross-breed readily in the laboratory; and where their distributions over-
lap, specimens intermediate between the two forms are found fairly frequently. The
two forms have a very narrow overlap in their respective ranges, however, and
specimens taken from outside the area of overlap are usually easily identified. These
two forms probably are more correctly considered subspecies.

Thyanta c. custator and T. calceata are the only two species in the genus with
distinctly black anterolateral pronotal margins. They can be separated reliably from
each other only by the characters of the male genitalia. The pygophoral opening in
T. c. custator is subtended by a rectangular impression, while this impression is
semicircular in T. calceata. When large series are present, some separation can be
made based on the general shape of the body: T. custator is slightly longer and
narrower than T. calceata. Fortunately, there is very little overlap in their distri-
butions.

Thyanta ( Thyanta ) custator accerra McAtee
Figs. 31-47, Map 3

Thyanta custator var. accerra McAtee, 1919:16.

Thyanta custator: Stal, 1872:34-35 (part); Uhler, 1872:399 (part); Uhler, 1876:289-
290 (part); Uhler, 1877:404; Popenoe, 1884:62; Uhler, 1886:7 (part); Lethierry
and Severin, 1893:148 (part); Osborn, 1894:121; Uhler, 1894a:230-231; Van Du-
zee, 1894:171; Blatchley, 1895:269; Gillette and Baker, 1895: 16; Van Duzee, 1904:
53 (part); Barber, 1906:260; Kirkaldy, 1909:94 (part); Banks, 1910:90 (part); Bar-
ber, 1911:108-111 (part); Zimmer, 1912:232-233; Torre-Bueno, 1914:92; Van
Duzee, 1914:4-5 (part); Van Duzee, 1917:52 (part); Hart, 1919:184-185; Malloch,
1919:217, fig. 75; Blatchley, 1926:113, 115-1 16 (part); Torre-Bueno, 1939:231
(part).

Thyanta perditor (of authors, not Fabricius): Uhler, 1872:399 (part); Uhler, 1876:
289 (part); Uhler, 1877:404 (part); Popenoe, 1884:62; Uhler, 1886:7; Osborn, 1894:
121; Uhler, 1894a:230 (part); Gillette and Baker, 1895:16; Van Duzee, 1904:52-
53 (part); Kirkaldy, 1909:95 (part); Zimmer, 191 1:232 (part); Torre-Bueno, 1914:
92; Malloch, 1919:217, fig. 79; Blatchley, 1926:113, 114-115, fig. 24 (part).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

39

40

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THY ANT A OF NORTH AND CENTRAL AMERICA

55

Thyanta pallido-virens (of authors, not Stal): Banks, 1910:90 (part); Rolston and
McDonald, 1984:fig. 31.

Thyanta accerra: Deay and Gould, 1935:305-306; Torre-Bueno, 1939:233; Harris,
1943:150; Ward et al., 1977:40; McPherson, 1979b:94; McPherson, 1982:77, 78-
79, fig. 167; Froeschner, 1988:592-593.

Thyanta pallidovirens accerra: Ruckes, 1957a: 14-1 5, 19-22; Ueshima, 1963:152-
153; Hoffman, 1971:44-45; Furth, 1974:22-23; McPherson, 1979a:83-84.
Thyanta pallidovirens spinosa Ruckes, 1 957a: 1 8-19; Ward et al. 1 977:40; Froeschner,
1988:593. NEW SYNONYMY.

Diagnosis. Green to dark brown, sometimes with varying amounts of reddish
coloration between humeral angles, often extending onto basal margin of each corium.

Lateral jugal margins sinuous, not parallel (Fig. 33). Humeral angles rounded to
angulate (spinose in desert areas of southwestern United States and Mexico); an-
terolateral margins of pronotum straight to concave in dorsal view (Figs. 31, 32),
stramineous to green, sometimes reddish, but never piceous; pronotal cicatrices im-
maculate. Each abdominal stemite with postspiracular black spot present on each
side (eastern U.S.) or absent (western U.S.). Posterolateral angles of abdominal ster-
nites immaculate or minutely marked with piceous.

Mesial margins of basal plates in caudoventral view slightly concave; posterior
margins sinuous; posteromesial angles rounded (Fig. 45). Posterolateral angles of
pygophore continuing onto posteroventral surface of pygophore as vague carinae,
forming rectangular impression; mesial portion of posterior margin of pygophore
convex with slight mesial emargination in ventral and dorsal views (Figs. 42, 43);
pygophore only slightly concave in lateral view (Fig. 44).

Types. McAtee (1919) described accerra as a variety of T. custator, he examined
four specimens: three from Barachias, Alabama, and one from San Antonio, Texas.
Blatchley (1926) elevated accerra to full species rank, but his concept of the species
was incorrect. Blatchley’s T. accerra was actually T. c. custator. The type specimens,
which are housed in the U.S. National Museum of Natural History (Washington,
D.C.), were examined.

Ruckes (1957a) described T. pallidovirens spinosa from 75<3<3 and 5199 specimens.
The holotype, which was examined, is from Patagonia, Santa Cruz Co., Arizona, and
is deposited in the American Museum of Natural History (New York).

Distribution. Eastern two-thirds of U.S. west to Montana, Nevada, and southern
California; Mexico; and Guatemala (Map 3).

Specimens examined. 8,677 specimens collected during every month of the year;
deposited in AMNH, ARH, ASUT, AUA, BMNH, CAS, CNC, CUC, DAR, DBT,
EGER, ENGL, FMNH, FSCA, INHS, ISU, LACM, LHR, LSU, MSU, MSUB,
MSUE, NCSU, NDSF, NMSU, OSUC, POLH, PUL, SIUC, SMEK, TAMU, UAT,

Figs. 31-47. T. custator accerra. 31. Habitus. 32. Variation in humeral angle. 33. Head.
34-36. Right paramere. 34. Medial view. 35. Ectal view. 36. Lateral view. 37-40. Theca and
related structures. 37. Ventral view. 38. Dorsal view. 39. Lateral view. 40. Ectal view. 41-44.
Pygophore. 41. Caudal view. 42. Ventral view. 43. Dorsal view. 44. Lateral view. 45. Genital
plates, caudoventral view. 46. Spermatheca. 47. Spermathecal pump.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

UCB, UCR, UCS, UGA, UIM, UMAA, UMC, UNAM, UNL, USNM, UUSL, VPI,
WSU. UNITED STATES: Alabama: Chambers, Dallas, De Kalb, Elmore, Green,
Henry, Jefferson, Lee, Macon, Madison, Montgomery, Morgan, St. Clair, Tallapoosa.
Arizona: Apache, Cochise, Coconino, Gila, Graham, Greenlee, Maricopa, Mohave,
Navajo, Pima, Pinal, Santa Cruz, Yavapai, Yuma. Arkansas: Benton, Boone, Conway,
Graighead, Faulkner, Garland, Hempstead, Hot Spring, Howard, Lawrence, Lincoln,
Little River, Logan, Mississippi, Montgomery, Newton, Pike, Polk, Pulaski, Wash-
ington, Yell. California: Imperial, Inyo, Kern, Los Angeles, Riverside, San Bernardino,
San Diego, Santa Barbara, Ventura. Colorado: Boulder, Chafee, Clear Creek, Doug-
las, El Paso, Jefferson, Gunnison, Larimer, Las Animas, Lincoln, Logan, Montezuma,
Morgan, Otero, Prowers, Pueblo, Teller, Washington, Yuma. Delaware: Kent. District
of Columbia. Florida: Highlands. Georgia: Bartow, Clarke, Crawford, Dade, Decatur,
Fulton, Habersham, Henry, Lamar, Macon, Madison, Morgan, Oconee, Oglethorpe,
Peach, Spalding, Talbot, Twiggs, Whitfield, Wilkes, Worth. Hawaii: Honolulu. Illi-

1992

THYANTA OF NORTH AND CENTRAL AMERICA

57

nois: Adams, Alexander, Cass, Champaign, Clay, Coles, Cook, Cumberland, Edgar,
Effingham, Grundy, Hancock, Hardin, Henry, Jackson, Jo Daviess, Johnson, Kan-
kakee, Macon, Macoupin, Madison, Marion, Marshall, Mason, Massac, McDonough,
McHenry, Menard, Morgan, Ogle, Peoria, Piatt, Pope, Pulaski, St. Clair, Scott, Union,
Vermilion, Wabash, Warren, Washington, White, Will, Williamson. Indiana: Cass,
Clark, Dearborn, Floyd, Greene, Harrison, Howard, Jackson, Jefferson, Jennings,
Johnson, Knox, Kosciusko, Lagrange, Lake, Lawrence, Marion, Morgan, Noble, Owen,
Parke, Perry, Posey, Putnam, St. Joseph, Starke, Switzerland, Tippecanoe, Vander-
burgh, Washington, Wayne. Iowa: Audubon, Boone, Floyd, Fremont, Harrison, John-
son, Linn, Lyon, Monona, Polk, Story, Union, Woodbury. Kansas: Allen, Anderson,
Atchison, Barbor, Bourbon, Chautauqua, Cherokee, Cheyenne, Clark, Comanche,
Decatur, Dickenson, Doniphan, Douglas, Ellis, Ellsworth, Finney, Ford, Geary, Gove,
Graham, Gray, Greeley, Greenwood, Hamilton, Harper, Harvey, Hodgeman,
Johnson, Kearny, Kingman, Kiowa, Labette, Lane, Leavenworth, Linn, Logan, Meade,
Miami, Mitchell, Montgomery, Morton, Neosho, Norton, Osborne, Ottawa, Pawnee,
Phillips, Pottawatomie, Pratt, Rawlins, Reno, Riley, Rooks, Rush, Russell, Scott,
Sedgwick, Seward, Sheridan, Sherman, Smith, Stafford, Stanton, Stevens, Sumner,
Thomas, Wabaunsee, Wallace, Washington, Wichita, Wilson. Kentucky: Campbell,
Christian, Fayette, Green, Hardin, Jefferson, Oldham, Rowan, Scott, Trimble. Lou-
isiana: Acadia, Avoyelles, Bossier, Caddo, Calcasieu, Cameron, Catahoula, Claiborne,
Concordia, East Baton Rouge, East Feliciana, Evangeline, Franklin, Grant, Iberville,
Jefferson Davis, Livingston, Madison, Natchitoches, Ouachita, Rapides, Richland, St.
Landry, Tensas, Vermilion, Webster, West Baton Rouge. Maine: Oxford. Maryland:
Allegany, Anne Arundel, Baltimore, Calvert, Frederick, Montgomery, Prince Georges,
Queen Annes, Talbot, Washington. Michigan: Barry, Berrien, Calhoun, Ingham,
Ionia, Jackson, Kalamazoo, Kent, Lenawee, Livingston, Mason, Montcalm, Oakland,
Shiawassee, Van Buren, Washtenaw, Wayne. Minnesota: Grant, St. Louis. Missis-
sippi: Bolivar, Clay, Grenada, Hinds, Issaquena, Lauderdale, Lee, Lowndes, Marshall,
Monroe, Noxubee, Oktibbeha, Panola, Pontotoc, Warren, Yazoo. Missouri: Atchison,
Audrain, Barry, Barton, Benton, Boone, Buchanan, Caldwell, Callaway, Camden,
Cape Girardeau, Carroll, Carter, Cedar, Chariton, Clark, Clay, Clinton, Cole, Cooper,
Crawford, Dade, Dallas, Douglas, Dunklin, Franklin, Gasconade, Gentry, Greene,
Grundy, Henry, Howard, Howell, Jackson, Jasper, Johnson, Lafayette, Lawrence,
Lewis, Lincoln, Livingston, Madison, Maries, Marion, Miller, Mississippi, Moniteau,
Montgomery, Morgan, New Madrid, Newton, Nodaway, Oregon, Osage, Pemiscot,
Pettis, Phelps, Pike, Platte, Polk, Putnam, Ralls, Rankin, Ray, Reynolds, St. Charles,
Ste. Genevieve, St. Louis, Saline, Scott, Shannon, Stoddard, Stone, Sullivan, Taney,
Vernon, Warren, Washington, Wright. Montana: Custer. Nebraska: Adams, Cass,
Chase, Cherry, Cheyenne, Custer, Frontier, Garden, Hall, Keith, Lancaster, Lincoln,
Morrill, Nance, Otoe, Scotts Bluff, Sioux, Thomas, Thurston. Nevada: Clark, Es-
meralda, Lincoln, Mineral, White Pine. New Hampshire: Rockingham. New Jersey:
Atlantic, Burlington, Cape May, Cumberland, Gloucester, Salem. New Mexico: Ber-
nalillo, Catron, Chaves, Cibola, Colfax, De Baca, Dona Ana, Eddy, Grant, Guadalupe,
Hidalgo, Lincoln, Luna, McKinley, Otero, Quay, Rio Arriba, Roosevelt, Sandoval,
San Miguel, Santa Fe, Torrance, Union, Valencia. New York: Orange. North Car-
olina: Bertie, Bladen, Caldwell, Chowan, Cleveland, Craven, Dare, Duplin, Edge-
combe, Granville, Iredell, Johnston, McDowell, Mecklenberg, Montgomery, Moore,

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Pasquotauk, Pender, Person, Rowan, Scotland, Vance, Wake, Warren, Washington,
Wayne. North Dakota: Cass, Hettinger, Stark. Ohio: Adams, Athens, Clinton, Frank-
lin, Hamilton, Lawrence, Lucas, Monroe, Pickaway, Sandusky, Trumbull, Washing-
ton. Oklahoma: Alfalfa, Beckham, Caddo, Canadian, Carter, Cleveland, Comanche,
Custer, Garfield, Grant, Harper, Jackson, Kay, Kingfisher, Lincoln, Logan, Mayes,
McIntosh, Noble, Oklahoma, Osage, Pawnee, Payne, Sequoyah, Stephens, Texas,
Tulsa, Wagoner, Woods, Woodward. Pennsylvania: Adams, Berks, Bucks, Centre,
Franklin, Lancaster. South Carolina: Abbeville, Allendale, Anderson, Barnwell,
Charleston, Chesterfield, Florence, Horry, Laurens, Lee, Marlboro, Oconee, Richland,
Spartanburg, Sumter. South Dakota: Beadle, Bennett, Brookings, Brule, Buffalo, Clay,
Custer, Fall River, Jones, Lawrence, Minnehaha, Pennington, Stanley, Sully, Tripp,
Union. Tennessee: Benton, Cooke, Davidson, Hamilton, Knox, Roane, Sevier. Texas:
Anderson, Aransas, Atascosa, Bailey, Bandera, Bastrop, Bee, Bell, Bexar, Bosque,
Brazos, Brewster, Burleson, Burnet, Caldwell, Cameron, Camp, Collin, Colorado,
Comal, Comanche, Coryell, Crockett, Crosby, Culberson, Dallas, Deaf Smith, Denton,
Dimmit, Donley, Duval, Eastland, El Paso, Erath, Fayette, Flatonia, Floyd, Frio,
Galveston, Gillespie, Gonzales, Gray, Guadalupe, Hale, Harris, Hays, Hemphill, Hi-
dalgo, Hill, Houston, Howard, Hudspeth, Jackson, Jeff Davis, Jim Hogg, Jim Wells,
Kaufman, Kendall, Kennedy, Kerr, Kimble, King, Kinney, Kleberg, Knox, Leon, Live
Oak, Llano, Lubbock, Lynn, Madison, Marion, Maverick, McLennon, Medina, Mil-
am, Montague, Moore, Motley, Nueces, Oldham, Palo Pinto, Parker, Pecos, Potter,
Presidio, Randall, Real, Reeves, Robertson, Runnels, San Patricio, Shelby, Smith,
Starr, Sutton, Swisher, Tarrant, Taylor, Terrell, Travis, Upshur, Uvalde, Val Verde,
Victoria, Walker, Ward, Webb, Wichita, Willbarger, Willacy, Williamson, Wise,
Zapata. Utah: Iron, Juab, Kane, Piute, Salt Lake, Sanpete, Sevier, Washington.
Virginia: Accomack, Albemarle, Appamattox, Arlington, Augusta, Bedford, Bucking-
ham, Campbell, Caroline, Carroll, Charlotte, Charlottesville City, Chesterfield, Clifton
Forge City, Frederick, Hampton, James City, King George, King William, Loudoun,
Montgomery, Nelson, New Kent, Newport News City, Norfolk City, Patrick, Pittsyl-
vania, Petersburg City, Prince Edward, Richmond City, Roanoke, Rockbridge, Rock-
ingham, Staunton City, Suffolk, Virginia Beach City, Westmorland, Winchester City,
Wythe. West Virginia: Monroe, Morgan. Wisconsin. Wyoming: Goshen, Laramie,
Niobrara, Uinta.

MEXICO: Aguascalientes: Aguascalientes; Arroyo San Pedro, 38 mi N Aguasca-
lientes. Baja California Norte: Bahia de los Angeles; Canyon del Tajo; Diablo Dry
Lake, San Felipe Valley; Ensenada; Gonzaga Bay; Guadelupe Canyon; 4 mi SW La
Zopopita; L Camillas Canyon; 60 mi S Mexicali; Mission San Borja; Persebu; 13 mi
S Puertocitos; 24 mi N Punta Prieta; 9 mi SE Rancho Laguna Chapola; 7 mi NE
Ranch Rosarito; San Felipe; 1 5 mi S San Felipe; 2 1 km S San Quintin. Baja California
Sur: Bahia Concepcion; 40 mi S El Arco Mine; La Paz; 7 mi SW La Paz; 25 mi W
La Paz; Las Tinajitas; Loreto; 10.3 mi SW Los Mendanos; 8 mi SW Mission San
Javier; 15 mi N San Ignacio; San Jose de Comondu; 4 mi W San Jose del Cabo; 5
mi SE Santa Rosalia; 10 mi W Santa Rosalia; Santiago; 1 mi E Todos Santos; 4 mi
N Todos Santos; 30 mi E V. Insurgentes; 45.5 mi E V. Insurgentes. Chiapas: Cintalpa;
Comitan. Chihuahua: Ciudad Camargo; 20 mi SW Ciudad Camargo; 25 mi SW
Ciudad Camargo; Catarinas; Chihuahua; 3 mi NW Chihuahua; 16 mi SE Chihuahua;
46 mi N Chihuahua; 1.1 mi S Colonel Alvaro Obregon; Ciudad Delicias; 3 mi SE

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THYANTA OF NORTH AND CENTRAL AMERICA

59

Ciudad Delicias; 10 mi S Ciudad Delicias; El Herradero; Gallego; 8 mi S Gallego;
10 mi W Gallego; Garcia; Hidalgo del Parral; 8 mi NE Hidalgo del Parral; 25 mi W
Hidalgo del Parral; 44 mi S Hidalgo del Parral; Huejotitlan; 6 mi WSW Jimenez; 10
mi N Jimenez; 24 mi N Jimenez; La Bufa; 11.1 km S RR at La Junta; Madera;
Marqueno; Matachic; 6 mi NE Meoqui; Moctezuma; Nuevo Casas Grandes; 43 mi
SE Nuevo Casas Grandes; 45 mi NW Nuevo Casas Grandes; 2 km W Oginaga, Rio
Conchos; 7 mi E Pedemales; Salaices; Samalayuca; San Jose Babicora; Santa Barbara;
63 mi W Santa Barbara; Santa Clara; Santa Clara Canyon, 5 mi W Parrita; Valle de
Olivos; Valle de Zaragoza; 23 mi N V. Ahumada; 15 mi S V. Matamoras. Coahuila:
10 mi S Allende; 10 mi S Ciudad Acuna; Arroyo de la Zorra; 20 mi SE Arteaga;
Boquillas del Carmen; 1 4.3 mi S Castanos; Guadalupe; 1 2 mi N Hermanas; La Gloria,
S of Monclova; Nueva Rosita; Saltillo; 1 mi E Saltillo; 1 mi SE Saltillo; 4 mi S
Saltillo; 6 mi NE Saltillo; 7 mi SSW Saltillo; 12.4 mi S Saltillo; 15 mi N Saltillo; 15
mi S Saltillo; 1 6 mi SE Saltillo; 1 7 mi SE Saltillo; 20 mi SE Saltillo; 29 mi SE Saltillo;
39 mi W Saltillo; 66 km S Saltillo; 5 km N San Esteban; San Jose de la Nina; San
Pedro de las Colonias; 12 mi SE San Pedro de las Colonias; Serranas del Burro;
Torreon; 22 mi N Zoragoza. Durango: 8 mi S Canutillo; 30 mi N Cuencame; 14 mi
S Donata Guerra; Durango; 1 1 mi W Durango; 20 mi W Durango; 25 mi S Durango;
69 mi N Durango; Encino; Guadalupe Victoria; La Loma; 2 mi S Menores de Arriba;
Navajos, 20 mi E El Salto; Nombre de Dios; 18 mi SE Nombre de Dios; Pedricena;
3 mi NW Pedricena; Rodeo; San Juan del Rio; Santa Lucia; 5 mi NE Sauz. Gua-
najuato: Leon. Guerrero: Iguala. Hidalgo: Tasquillo. Jalisco: 2 mi S Ciudad Guzman;
Lagos de Moreno; 5.6 mi NE Lagos de Moreno; 13 mi SW Lagos de Moreno; San
Juan de los Lagos; 1 mi NE V. Hidalgo. Michoacan: 10.3 mi W Morelia; Zamora.
Morelos: Ruinas Xochicalco; Tepoztlan. Nayarit: San Bias. Nuevo Leon: Apodaca;
Aramberri; 3 mi E Galeana; Hacienda Vista Hermosa, V. Santiago; 9 mi W Iturbide;
Linares; 10 mi S Linares; 15 mi W Linares; 16 mi S Linares; 1.7 mi S Montemorelos;
15 mi SE Montemorelos; Monterrey; 4 mi S Monterrey; 5 mi S Monterrey; 6 mi S
Monterrey; 1 0 mi W Monterrey; Paso Mamulique; Pena Nevada Zaragoza; Pesqueria;
10 mi N Providencia; 41 mi S Sabinas Hidalgo; 22 mi S San Roberto; 40 mi S San
Roberto; 46 mi NW San Roberto; Santiago; Vallecillo. Oaxaca: 3 mi SE El Tule; 10
mi N Miltepec; Mitla; Oaxaca; 2 mi NW Oaxaca; Ruinas Zaachila; 13 km W Te-
huantepec; 22 mi SE Totalapan, San Jose Viejo. Puebla: Acatlan; 1 1 mi SE Acatlan;
19 mi NW Calcalcapan; Chilac; Puebla; 6 mi SW Tehuacan; 7 mi NE Tehuacan;
Valseguillo. Queretaro: Oro. San Luis Potosi: 3 mi W Cedral; 12 mi W Ciudad del
Maiz; 20 mi S Ciudad Valles; 19.6 mi N El Huizache; 28.5 mi S El Huizache; El
Salto; 34 mi S Salinas; 10 mi NE San Luis Potosi; 31 mi S San Luis Potosi; 46 mi
N San Luis Potosi; 84 mi NE San Luis Potosi; 1 23 mi NE San Luis Potosi; V Hidalgo;
12 mi NE V. Hidalgo. Sinaloa: Los Mochis; 5 mi N Mazatlan; 46 mi E Mazatlan.
Sonora: 20 mi SE Agua Prieta; 65 mi SE Agua Prieta; Alamos; 7 mi SE Alamos; 7
mi W Alamos; 10 mi W Alamos; Bahia Kino; 1 mi W Caborca; Cabullona; Campo
Utah; El Desemboque; El Fuerte; 5 mi E Esqueda; 20 mi S Estacion Llano; Guaymas;
26 mi SE Guaymas; Hermosillo; 40 mi N Hermosillo; 42 mi S Hermosillo; 9 mi
NNE Imuris; La Chiripa; La Choya; La Floresta Ranch, 8 mi E Tastiota; Magdalena;
Minas Nuevas; Mira Mar Beach; Navojoa; 5 mi E Navojoa; Nogales; Palm Canyon,
17 mi E Magdalena; Pitiquito; Pozo Coyote; Puerto Penasca; San Bernardino, Rio
Mayo; 8 mi S Santa Ana; Santa Rosa Ranch; Santa Teresa; Sonora; 20 mi S Sonoyta;

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

38 km S Sonoyta; Tastiota; Tiburon Island; V. de Sens, 5 mi SW Hermosillo; Yavaros.
Tamaulipas: Ciudad Victoria; 25 mi S Ciudad Victoria; 81 mi NW Ciudad Victoria;
35 km N El Limon; Gomez Farias; Guemes, 1 5 mi N Ciudad Victoria; Lago Republica
Espanola; Matamoros; San Fernando; San Jose. Veracruz: 1 mi NE Acultzingo;
Orizaba. Zacatecas: 4 mi NE Concepcion del Oro; Fresnillo; 1 mi N Fresnillo; 8 mi
S Fresnillo; 9 mi S Fresnillo; 25 mi W Fresnillo, Laguna Balderama; 14 mi N Luis
Moya; Sain Alto; 28 mi NE Sierra Viejo.

GUATEMALA: Jutiapa: 6 mi NE El Progreso.

Comments. This subspecies is extremely variable and has been the subject of much
confusion in the past. Its relationship with the nominate subspecies is discussed under
the comments section of that subspecies.

Ruckes (1957a) divided T. pallidovirens into four subspecies: pallidovirens from
the far western U.S.; setosa from the northwestern U.S.; spinosa from the south-
western U.S. and Mexico; and accerra from the eastern U.S. Ueshima (1963) showed
that specimens of pallidovirens from California have a different chromosome number
than specimens from the eastern U.S. (14 + XY in the male versus 16 + XY,
respectively). Because of this difference, he believed that the two populations were
genetically isolated and were probably two distinct species. The difference in chro-
mosome number has been confirmed in the present study. Cross-breeding and free-
choice mating experiments also have been conducted, the results of which support
the separation of the two taxa into distinct species (e.g., specimens from California
and Louisiana mated and laid eggs, but the eggs were infertile).

Ruckes (1957a) described T. pallidovirens setosa from Idaho and eastern areas of
Oregon and Washington. During this study, specimens from Nez Perce Co., Idaho
were karyotyped. They had a chromosome complement of 14 + XY in the male,
the same as T. pallidovirens from California.

Ruckes (1957a) described T. pallidovirens spinosa from the southwestern U.S.
Ueshima (1963) karyotyped specimens of this form collected from southeastern
California. He found that males had a chromosome complement of 14 + XY, and
he concluded that the designation of spinosa as a subspecies of T. pallidovirens was
probably correct. During the present study, however, specimens karyotyped from
several localities in southern Arizona were found to have a chromosome complement
of 16 + XY. Also, in cross-breeding experiments, specimens from Arizona and
Louisiana readily interbred with no apparent difficulties and produced viable young.
Therefore, it is probable that spinosa is T. custator accerra, not T. pallidovirens.

Ruckes (1957a) used the spinose humeral angles of T. pallidovirens spinosa to
separate it from T. custator accerra (=his T. p. accerra). This character is relatively
variable, however. Long series from many localities within the range of spinosa
usually include specimens with spinose humeral angles and specimens with rounded
humeral angles, as well as many intermediates. Geographical separation of the two
forms is not complete, and therefore spinosa is placed as a junior synonym of T. c.
accerra.

Distinguishing T. custator accerra and T. pallidovirens can be difficult. In general,
specimens of T. c. accerra (excluding the spinosa form) have a postspiracular black
spot on each side of each abdominal sternite, while specimens of T. pallidovirens do
not. However, some (< 10%) specimens of T. pallidovirens have postspiracular black
spots, but the spots are usually smaller than the adjacent spiracle. Conversely, some

1992

THY ANT A OF NORTH AND CENTRAL AMERICA

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(<10%) specimens of T. c. accerra lack postspiracular black spots. Fortunately, the
two species are geographically separated except in the southwestern U.S., where T.
pallidovirens overlaps considerably with the spinosa form of T. c. accerra. These two
forms can usually be separated by the degree of spinosity of the humeral angles.
Thyanta pallidovirens always has rounded humeral angles, while the spinosa form of
T. c. accerra usually has angulate to spinose humeral angles. Also, most specimens
of T. pallidovirens have at least a partial reddish transhumeral band, a condition that
is rare in the spinosa form of T. c. accerra.

Thyanta ( Thyanta ) pallidovirens (Stal)

Figs. 48-62, Map 4

Pentatoma pallidovirens St&l, 1859:227; Walker, 1867:288.

Thyanta pallidovirens: Stal, 1862a:58; Stal, 1872:35; Uhler, 1886:7; Lethierry and
Severin, 1893:148; Uhler, 1894a:231; Van Duzee, 1904:53-54; Banks, 1910:90
(part); Van Duzee, 1917:52; Torre-Bueno, 1939:232; Ruckes, 1 957a: 1-22, Froesch-
ner, 1988:593.

Thyanta custator (of authors, not Fabricius): Uhler, 1872:399 (part); Uhler, 1876:
289-290 (part); Kirkaldy, 1909:94 (part); Van Duzee, 1914:4-5 (part); Van Duzee,
1916:231; Van Duzee, 1917:52 (part).

Thyanta pallidovirens setosa Ruckes, 1957a: 17-1 8; Froeschner, 1988:593. NEW

SYNONYMY.

Diagnosis. Dorsal coloration often with varying amounts of reddish between hu-
meral angles, often extending onto basal margin of each corium and along antero-
lateral pronotal margins; sometimes apex of scutellum reddish.

Lateral jugal margins sinuous, not parallel (Fig. 49). Anterolateral margins of pro-
notum straight to slightly concave in dorsal view, green to reddish, never piceous;
humeral angles rounded (Fig. 48); pronotal cicatrices immaculate. Postspiracular
black spots usually lacking; if present, then usually smaller than adjacent spiracle.
Posterolateral angles of abdominal stemites immaculate.

Basal plates in caudoventral view with mesial margins straight to slightly concave;
posterior margins sinuous; posteromesial angles narrowly rounded (Fig. 60). Pos-
terolateral angles of pygophore continuing onto posteroventral surface of pygophore
as carinae, forming rectangular impression; posterior margin slightly concave in
caudal view, usually with small, medial, V-shaped emargination (Fig. 56); postero-
lateral angles prominent in ventral and dorsal views (Figs. 57, 58); pygophore slightly
concave in lateral view (Fig. 59).

Types. Stal (1859) described Pentatoma pallidovirens from 1<3 and 12 specimen
from California without designating a holotype or paratype. The 6 specimen labeled
(a) “California” (b) “Kinb.” (c) “Type” (d) “Typus” (e) “ Thyanta pallidovirens Stal”
(f) “109 51” (g) “349 84” (h) “Riksmuseum Stockholm” is designated the lectotype.
The 2 specimen labeled (a) “California” (b) “Kinb.” (c) “2” (d) “Type.” (e) “Allo-
typus” (f) “350 84” (g) “Riksmuseum Stockholm” is designated paralectotype. The
lectotype and the paralectotype, which are conserved in the Naturhistoriska Riks-
museet (Stockholm, Sweden), were examined.

Ruckes (1957a) described T. pallidovirens setosa from 1 8<5<5 and 1022 specimens.

Figs. 48-62. T. pallidovirens. 48. Habitus. 49. Head. 50-52. Right paramere. 50. Medial
view. 51 Ectal view. 52. Lateral view. 53-55. Theca and related structures. 53. Ventral view.
54. Dorsal view. 55. Lateral view. 56-59. Pygophore. 56. Caudal view. 57. Ventral view. 58.
Dorsal view. 59. Lateral view. 60. Genital plates, caudoventral view. 61. Spermatheca. 62.
Spermathecal pump.

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THYANTA OF NORTH AND CENTRAL AMERICA

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Map 4. T. pallidovirens.

The holotype, which was examined, is from Pullman, Whitman Co., Washington,
and is deposited in the American Museum of Natural History (New York).
Distribution. Western U.S. (Map 4).

Specimens examined. 3,606 specimens collected during every month of the year;
deposited in AMNH, CAS, DAR, EGER, FMNH, FSCA, INHS, ISU, LACM, LHR,
LSU, MSU, MSUB, MSUE, NCSU, ODAS, OSUC, PUL, SIUC, SMEK, TAMU,
UCB, UCR, UCS, UGA, UIM, UNAM, USNM, UUSL, WSU. CANADA: British

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Columbia: Cache Ck; Coldstream; Comox; Lytton; Malahat; Nanaimo; Vancouver
Isl; Royal Oak; Saanich Dist; Vernon; Victoria; Wellington.

UNITED STATES: California: Alameda, Alpine, Amador, Butte, Calaveras, Co-
lusa, Contra Costa, El Dorado, Fresno, Glenn, Humboldt, Inyo, Kern, Lake, Lassen,
Los Angeles, Madera, Marin, Mariposa, Mendocino, Merced, Modoc, Mono, Mon-
terey, Napa, Nevada, Orange, Placer, Plumas, Riverside, Sacramento, San Benito,
San Bernardino, San Diego, San Francisco, San Joaquin, San Luis Obispo, San
Mateo, Santa Barbara, Santa Clara, Santa Cruz, Shasta, Sierra, Siskiyou, Solano,
Sonoma, Stanislaus, Tehama, Trinity, Tulare, Tuolumne, Ventura, Yolo, Yuba. Ida-
ho: Ada, Benewah, Bingham, Blaine, Boise, Bonner, Butte, Cassia, Clearwater, El-
more, Franklin, Fremont, Gem, Gooding, Idaho, Kootenai, Lake, Latah, Lewis, Nez
Perce, Oneida, Owyhee, Shoshone, Twin Falls, Valley. Montana: Lake, Lewis & Clark,
Ravalli, Sanders. Nevada: Carson City, Douglas, Lyon, Washoe. Oregon: Baker,
Benton, Crook, Curry, Deschutes, Douglas, Grant, Harney, Hood River, Jackson,
Jejferson, Josephine, Klamath, Lake, Linn, Malheur, Marion, Multnomah, Polk,
Sherman, Tillamook, Umatilla, Union, Wasco, Washington, Wheeler, Yamhill. Utah:
Box Elder, Cache, Garfield, Morgan, Salt Lake, Sevier, Utah, Washington, Wheeler,
Yamhill. Utah: Box Elder, Cache, Garfield, Morgan, Salt Lake, Sevier, Utah, Wash-
ington, Weber. Washington: Asotin, Benton, Chelan, Clark, Columbia, Douglas, Grant,
King, Kittitas, Mason, Okanagon, Pierce, San Juan, Snohomish, Spokane, Thurston,
Walla Walla, Whitman, Yakima.

Comments. Ruckes (1957a) distinguished T. pallidovirens setosa from other U.S.
Thyanta by the increased amount of pilosity on the legs and rostrum. This is a
difficult character to discern. The brown autumnal-vernal forms of several other
species and subspecies are also characterized by more and longer hairs on the same
body structures (McPherson, 1979a). Because T. p. setosa differs from the nominate
subspecies in no other significant manner, T. p. setosa is placed as a junior synonym
of T. pallidovirens.

No reliable morphological character will consistently separate T. pallidovirens from
T. custator accerra, as discussed in detail under the comments section of T. c. accerra.
Basically, in the southwestern U.S. the two usually can be separated by the shape of
the humeral angles, which are rounded in T. pallidovirens and angulate to spinose
in T. c. accerra. In the northwestern U.S., the two species seem to be geographically
isolated (Maps 3, 4). Here, separation often can be made based on the presence (in
T. c. accerra) or absence (in T. pallidovirens ) of postspiracular black spots.

Thyanta ( Thyanta ) perditor (Fabricius)

Figs. 63-77, Map 5

Cimex perditor Fabricius, 1794:102; Fabricius, 1803:163.

Pentatoma fascifera Palisot de Beauvois, 1817:150, fig. 8 (syn. by Dallas, 1851:206).
Pentatoma collaris Westwood, 1837:40 (syn. by Dallas, 1851:206).

Cimex transversalis Herrich-Schaffer, 1841:66 (syn. by Dallas, 1851:206).

Cimex dimidiatus Herrich-Schaffer, 1 84 1 :fig. 629 (syn. by Dallas, 1851:206).
Pentatoma dimidiatum: Herrich-Schaffer, 1844:94 (syn. by Dallas, 1851:206).
Euschistus perditor: Dallas, 1851:206; Walker, 1867:247.

Pentatoma ( Mormidea ) perditor: Guerin-Meneville, 1857:367.

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THYANTA OF NORTH AND CENTRAL AMERICA

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Thyanta perditor: St&l, 1862a:58; St&l, 1862b: 104; Stal, 1868:29; Stal, 1872:34; Uhler,

1872:399 (part); Uhler, 1876:289; Uhler, 1877:404 (part); Distant, 1880:66; Berg,

1884:100; Distant, 1893:333; Lethierry and Severin, 1893:148; Uhler, 1893:705;

Uhler, 1894a:230 (part); Uhler, 1894b: 173; Distant, 1900:432; Van Duzee, 1904:

52-53 (part); Van Duzee, 1907:9; Kirkaldy, 1909:95; Banks, 1910:90; Zimmer,

1912:14 (part); Barber, 1914:523; Van Duzee, 1917:51-52; Barber, 1923:12;

Blatchley, 1926:113, 114-115 (part); Barber, 1939:292-293; Torre-Bueno, 1939:

230; Ruckes, 1957a:l, 20; Froeschner, 1988:593.

Euschistus fasciatus Walker, 1867:245 (syn. by Stal, 1872:34).

Euschistus adjunctor Walker, 1867:249 (syn. by Stal, 1872:34).

Thyanta ( Thyanta ) perditor: Rider and Chapin, 1991.

Diagnosis. Transhumeral rubiginous band usually present; tylus and vertex of head
often reddish.

Lateral jugal margins sinuous, not parallel (Fig. 64). Humeral angles spinose, each
spine directed anterolaterad; anterolateral pronotal margins not piceous, concave in
dorsal view (Fig. 63). Mesial comer of each pronotal cicatrice black. Each abdominal
stemite with postspiracular black spot on each side, both anterolateral and postero-
lateral angles usually piceous.

Basal plates with mesial margins straight to slightly convex in caudoventral view;
posterior margins sinuous (Fig. 75). Pygophoral opening subtended by semicircular
impression; posterior margin of pygophore produced posterodorsad, in ventral and
dorsal views convex medially with small, medial, V-shaped emargination (Figs. 72,
73), concave in lateral view (Fig. 74).

Types. Fabricius (1794) described Cimex perditor from 266 and 299 specimens
without designating a holotype or paratypes. Rider and Chapin (1991) made lectotype
and paralectotype designations. All four specimens, which are housed in the Univ-
ersetetes Zoologiske Museum (Copenhagen, Denmark), were examined.

Rider and Chapin (1991) confirmed the status of Pentatoma fascifera Palisot de
Beauvois, P. collaris Westwood, Cimex transversalis Herrich-Schaffer, Euschistus
fasciatus Walker, and E. adjunctor Walker as junior synonyms. They also designated
lectotypes for the latter two species. At one time Euschistus rubiginosus Dallas was
considered a synonym of T. perditor. Rider (1986b), however, examined the holotype
of E. rubiginosus and determined that it was a senior synonym of Euschistus incus
Rolston.

Distribution. This is the most widely distributed species in the genus, occurring
from southern Florida, Texas, and Arizona south through Central America, West
Indies, and South America to northern Argentina (Map 5).

Specimens examined. 844 specimens collected during every month of the year;
deposited in AMNH, AUA, BMNH, CAS, CNC, CU, DAR, DBT, FSCA, INHS,
ISU, LHR, LSU, MSU, MSUE, OSUC, PUL, SIUC, SMEK, TAMU, UAT, UCB,
UCR, UCS, UGA, UMAA, UMC, UNAM, USNM, WSU. UNITED STATES:
Arizona: Cochise: Southwest Research Station, 5 mi W Portal. Florida: Flamingo;
Mahogany Hammock; Paradise Key; Pine Castle; Windly. Broward: Deerfield Beach.
Collier: Immokalee; Royal Palm Park. Dade: Coral Gables; Everglades National Park;
Florida City; 5 mi SW Florida City; Goulds; Grossman Hammock; Hialeah; Home-
stead; Kendall; Miami; Miami Springs; Princeton. Highlands: Avon Park; Lake Plac-

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THY ANT A OF NORTH AND CENTRAL AMERICA

67

id; Sebring. Indian River: Vero Beach. Lee: Fort Myers. Manatee: Palmetto; Terra
Ceia. Martin: Stuart. Monroe: Big Pine Key; Key Largo; Key West; Marathon Key;
Upper Key Largo; Upper Matecumbe Key. Orange: Orlando. Palm Beach: Belle
Glade; Delray; Royal Palm Park. Polk: Lake Hamilton; Lakeland. Volusia: Daytona
Beach. Texas: Cameron: Brownsville; Sabal Palm Grove near Southmost.

MEXICO: Hochmilco; Lagos de Moreno; Paricutin. Aguascalientes: Aguascalien-
tes. Campeche: km 71 Carr. Campeche-Meizina. Chiapas: Arriaga; 4.4 mi N Bochil;
Bonampak Ruinas; Chicoasen; Chorreadera State Park; Cintalpa; 5 mi S Cintalpa;
13 mi W Cintalpa; Comitan; 31.5 mi SE Comitan; Dos Lagos; El Sumidero; 1.6 mi
S Jitotol; 13 mi S La Trinitario; 18 km S La Trinitario; 12 mi W Ocozocoautla;
Palenque; Palenque Ruinas; 23 mi S Palenque, 4 mi NE Pichucalco; 1 mi S Rayon;
2 mi SE Revolucion Mexicana; 3 mi W Rizo de Oro; Sanches Ranch Las Rosas; San
Cristobal de las Casas; 8 mi SE San Cristobal las Casas; 23 mi W San Cristobal de
las Casas; 39 mi E San Cristobal las Casas; Simojovel; 10 km WNW Soyalo; 2 mi
E Suchiapa; Sumidero Canyon; 9 mi SE Tapilula; Teopisca; 14 mi N Tuxtla Gutierrez;

3.5 km S Rio Tulija. Colima: Colima Volcano. Durango: 9 mi W La Ciudad; Peasa
Pena. Guanajuato: Leon. Guerrero: Acahuizotla; Acapulco de Juarez; 5 mi S, 2.5 mi
E Chilpancingo; km 8 Carr. Chilpancingo Omilteme; 17 mi N Mexcala; Mochitlan;

1.5 mi W Mochitlan; 13 mi SW Tierra Colorado. Hidalgo: Hwy 45, 17 mi NE
Huichapan. Jalisco: 16 km E Agua el Obispo; Ajijic; Guadalajara; 5 mi SE Plan de
Barrancas. Mexico: 4.3 mi NE Ixtapan de la Sal; Tejupilco; Temascaltepec, Real de
Arriba; Teotihuacan; 1 1 mi W Texcoco; 1 mi NE Tlamacas, P. N. Popocatepetl;
Valle de Bravo; 21 mi NW Valle de Bravo. Michoacan: Apatzingan; 1 1 mi E Apat-
zingan; Jiquilpan; Palo Alto; Tancitaro. Morelos: Cuautla; Cuernavaca; Jojutla; Mo-
relos; Pte de Itla; Xochicallo. Nayarit: 13 mi NW Ahuacatlan; Jesus Maria; Nayarit-
Jalisco line, Hwy 15; 15 km E San Bias; Tepic. Nuevo Leon: 9 mi S Monterrey; 3
mi S Pacheco. Oaxaca: 7.7 mi S Ejutla; El Camaron, 20 mi E Oaxaca; 2.7 mi NW
El Camaron; 10 mi S El Camaron; 27 mi E Juchitan; 3 km E La Ventosa; 3.4 mi SE
Matatlan; 1 1 mi N Matias Romero; SE Nejapa; 30 mi NW Oaxaca; 45 mi SE Oaxaca;
53 mi NE Oaxaca; Tehuantepec; Temazcal; 23 mi W Tequistlan; Totolapan; 18 mi
NW Totolapan; 23 km NW Totolapan; Tuxtepec. Puebla: 5.1 mi SW Tehuacan; 6
mi SW Tehuacan. Queretaro: Queretaro; 29 mi N Queretaro; 10 mi E San Juan del
Rio. San Luis Potosi: 5 mi E Ciudad del Maiz; 1 1 km N Ciudad Valles; 20 mi S
Ciudad Valles; El Pujal; El Salto Falls; Micos; 3 1 mi S San Luis Potosi; Tamazunchale;
30 mi S Tamazunchale; Valle Hidalgo; 2 mi E Xilita. Sinaloa: 22.6 mi S Culiacan;
Los Mochis; Mazatlan; 1 5 mi N Mazatlan; Valle Union; 5 mi E Valle Union. Tabasco:
30 mi W Cardenas; Chontalpa; Pajelagatero; Pico do Oro; 5 km S Villahermosa; 14
mi SE Villahermosa. Tamaulipas: Bocatoma: 1 1 mi SW Ciudad Victoria; 6 mi W
Rio Sabinas, near Encino; 19 mi NE Tula. Tlaxcala: 2 mi W Tlaxcala. Veracruz:
Alvarado; Catemaco; 7 mi SE Catemaco; Coatzacoalcos; Cordoba; 1 mi E Cordoba;

Figs. 63-77. T. perditor. 63. Habitus. 64. Head. 65-67 . Right paramere. 65. Medial view.
66. Ectal view. 67. Lateral view. 68-70. Theca and related structures. 68. Ventral view. 69.
Dorsal view. 70. Lateral view. 7 1-74. Pygophore. 7 1 . Caudal view. 72. Ventral view. 73. Dorsal
view. 74. Lateral view. 75. Genital plates, caudo ventral view. 76. Spermatheca. 77. Spermathecal
pump.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Cotaxtla; Extacion Mieron das Fortin; 1 mi W Fortin de las Flores; Jalapa; 10 mi E
Jalapa; Lake Catemaco; 5 mi N Lerdo de Tejada; Los Tuxtlas Biological Station; L.
Rivera. Atzagan; 4 mi NE Minatitlan; Nanchital; Orizaba; Papantla; Plan del Rio;
3 mi SW Sontecompapan; 15.8 mi S Tampico; Vega de Alatorre; Veracruz. Yucatan:
10 km N Piste.

GUATEMALA: Chocola. Chimaltenango: Yepocapa. Guatemala: Guatemala City.
Izabal: Livingston; Los Amates; Morales. Jutiapa: San Jeronimo. Sac.: Antigua Gua-
temala. Solola: Panajachel, Lago de Atitlan. Suchitepequez: Alta Vista; 8 mi N Santa
Barbara. BELIZE: Punta Gorda. EL SALVADOR: La Libertad; San Salvador. HON-
DURAS: 1 mi W Jicaro Galan Junction, Rio Nacaome; Tegucigalpa; Uyace Peak;
Yojoa Lake; Zomorano. NICARAGUA: km 4 to Masaya. Estetf: 1 3.4 mi NW Sebaco.
Leon: Los Zarzales; Malpaisillo. Managua: Los Brasiles; Managua; Tipitapa. Mu-
sawas: Waspuk R. Zelaya: Puerto Cabezas. COSTA RICA: Isla Bonita; Reventazon
Valley; San Carlos. Cartago: Santiago; Turrialba. Guanacaste: La Pacifica near Canas.
Heredia: Heredia; 2 mi E Puerto Viejo; Varablanca. Puntarenas: Monte verde. San
Jose: 5.5 km SE Desamparados; San Jose. PANAMA: New California. Bocas del
Toro: Chiriqui Grande. Canal Zone: Barro Colorado Island; Cristobal. Chiriqui:
Boquete; Cerro Punta; Porteillos. Code: Cerro Pena near El Valle. Panama: Cerro
Jefe; Madden Lake.

BAHAMA ISLANDS: Andros Island: Andros Town. Cat Island. Long Island:
Clarencetown. New Providence Island: Nassau. CUBA: El Cobre. Ciudad de la Ha-
bana: La Habana. Cienfuegos: Palmira; Soledad nr Cienfuegos. Granma: Cayamas.
Guantanamo: Mtns nr Guantanamo. Las Tunas: Jobabo. Pinar del Rio: San Vicente.
Santiago de Cuba: 12 mi N Santiago de Cuba. JAMAICA: Balaclava; Claremont
Baron Hill Trelawny; Hope Bay; Kingston; Liguanea Plain; Mandeville; Montego

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THYANTA OF NORTH AND CENTRAL AMERICA

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Bay. HAITI: Ouest: Kenscoff; Port-au-Prince. DOMINICAN REPUBLIC: 8 mi up
Macoris River. Distrito Nacional: Santo Domingo. La Romana: La Romana Center.
La Vega: Jarabacoa. Peravia : 2 km N Nizao; 21 km NW San Jose. Puerto Plata. San
Cristobal: San Cristobal. San Pedro de Macons: Boca Chica. PUERTO RICO: Cialitos
Cruces, El Verde, Mona Island. Aguadilla: Anasco, Isabela, Rincon, San Sebastian.
Arecibo: 7 km S Ciales, Dorado, Manati, Utuado, Vega Baja. Bayamon: Corozal.
Guayama: Arroyo. Humacao: Humacao, Loiza, 1 mi SE Luquillo, Naguabo. Ma-
yaguez: Boqueron, Guanica Forest, Guayanilla, Mayagiiez, San German. Ponce: Coa-
mo Springs, Juana Diaz, Ponce, 3 mi N Santa Isabel. San Juan: Rio Piedras. U.S.
VIRGIN ISLANDS: St. Croix: Frederiksted. St. Thomas. BRITISH WEST INDIES:
Antigua: Bello. Barbuda: Codrington. Dominica: Castle Bruce Road nr Savane David;
Clarke Hall Est; 4.7 mi E Pt Casse; 5 km SW Pt Casse; Roseau. Grand Cayman: 3
mi N Georgetown; Western Dist. Grenada: Grand Anse; Mount Gay Est; St. Georges.
St. Kitts: Basseterre; 4 mi W Basseterre. St. Lucia: Castries. St. Vincent. Trinidad:
Cumuto; St. Augustine. FRENCH WEST INDIES: Guadeloupe: Pointe-a-Pitre. Mar-
tinique: 2 km N St. Pierre. BARBADOS.

Comments. Thy ant a perditor is one of a group of very closely related species, all
of which have distinctly spinose humeral angles. The characters used to separate
these species are sometimes subtle and hard to diagnose unless a series of specimens
is available. Fortunately, most have very little overlap in their distributional ranges.

Thyanta {Thy ant a) spectabilis Ruckes
Figs. 78-92, Map 6

Thyanta spectabilis Ruckes, 1957c: 175-1 78, figs. 3-4.

Thyanta perditor (of authors, not Fabricius): Van Duzee, 1923:127.

Diagnosis. Rubiginous transhumeral band usually present; often tylus and vertex
of head reddish.

Lateral jugal margins sinuous, not parallel (Fig. 79). Anterolateral pronotal margins
concave, not piceous; humeral angles spinose, spines directed primarily laterad and
only slightly cephalad (Fig. 78). Mesial comer of each pronotal cicatrice usually
piceous. Each abdominal stemite with postspiracular black spot on each side, an-
terolateral and posterolateral angles piceous.

Basal plates in caudoventral view with mesial margins straight to slightly convex;
separated basally; posterior margins sinuous (Fig. 80). Spermathecal bulb digitiform,
but with numerous short protuberances (Fig. 82). Pygophoral opening subtended by
semicircular impression; posterior margin of pygophore produced posterodorsad, in
ventral and dorsal views convex medially with small, medial V-shaped emargination
(Figs. 84, 85); slightly concave in lateral view (Fig. 86).

Types. Ruckes (1957c) described Thyanta spectabilis from 266 and 692 specimens,
all from Baja California, Mexico. The 6 holotype was examined and is presently
conserved in the California Academy of Sciences (San Francisco).

Distribution. Baja California, Mexico (Map 6).

Specimens examined. 1 7 specimens collected during every month of the year except
February, April, September, and October; deposited in CAS, DBT, UCB, UIM,

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Figs. 78-82. T. spectabilis. 78. Habitus. 79. Head. 80. Genital plates, caudoventral view.
81. Spermatheca. 82. Spermathecal pump.

UNAM, USNM. MEXICO: Lower California; 2 km W El Centemario. Baja Cali-
fornia Norte: Bahia de los Angeles. Baja California Sur: Cabo San Lucas; 28 mi S
El Arco Mine, Rancho Santa Marguerita; 2 mi E El Coyote, NE of La Paz; 6 mi S,

1 mi E El Pescadero; La Paz; 6 km S San Pedro; Santa Anita; 7 mi N Santa Anita;

2 km SE, 3.5 km NE Santa Rita; 21.6 mi N Todos Santos; Venancio.

Comments. Thyanta spectabilis is very closely related to T. perditor, and may

actually be a subspecies of that species. The genitalia of the two species are virtually
identical; the only difference is that the parameres in T. spectabilis are somewhat
larger than those in T. perditor. This is expected, however, since specimens of T.

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Figs. 83-92. T. spectabilis. 83-86. Pygophore. 83. Caudal view. 84. Ventral view. 85. Dorsal
view. 86. Lateral view. 87-89. Right paramere. 87. Medial view. 88. Ectal view. 89. Lateral
view. 90-92. Theca and related structures. 90. Ventral view. 91. Dorsal view. 92. Lateral view.

spectabilis are larger than those of T. perditor. The only reliable character to separate
the two species is the orientation of the humeral spine. In T. perditor the humeral
angle is directed anterolaterad, while in T. spectabilis it is directed primarily laterad
and only slightly cephalad.

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Thy ant a {Thy ant a) cubensis Barber & Bruner
Figs. 93-108, Map 7

Thyanta cubensis Barber and Bruner, 1932:257-258, figs. 4-5; Bruner and Barber,
1949:158; Alayo, 1967:18, 20.

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THYANTA OF NORTH AND CENTRAL AMERICA

73

Figs. 93-108. T. cubensis. 93. Habitus. 94. Head. 95-97. Right paramere. 95. Medial view.
96. Ectal view. 97. Lateral view. 98-101. Theca and related structures. 98. Ventral view. 99.
Dorsal view. 100. Lateral view. 101. Ectal view. 102-105. Pygophore. 102. Caudal view. 103.
Ventral view. 104. Dorsal view. 105. Lateral view. 106. Genital plates, caudoventral view. 107.
Spermatheca. 108. Spermathecal pump.

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Map 7. T. cubensis (■); T. obsoleta (0); T. testacea (•).

Diagnosis. Dorsal coloration brown to green; transhumeral reddish markings lack-
ing.

Lateral jugal margins sinuous, not parallel (Fig. 94). Anterolateral margins of
pronotum slightly concave in dorsal view, usually contrastingly pale yellow; humeral
angles spinose with spines small and directed anterolaterad (Fig. 93); pronotal cica-
trices immaculate. Abdominal sternites with anterolateral angles immaculate, pos-
terolateral angles piceous. Postspiracular black spot present on each side of each
abdominal sternite.

Basal plates in caudoventral view with mesial margins straight to slightly convex;
posterior margins sinuous; posteromesial angles broadly rounded (Fig. 106). Pygo-
phoral opening subtended by semicircular impression in caudal view; posterior mar-
gin of pygophore produced posterodorsad, convex with small, medial, V-shaped
emargination in both ventral and dorsal views (Figs. 103, 104); pygophore slightly
concave in lateral view (Fig. 105).

Types. Barber and Bruner (1932) described Thyanta cubensis from 14 66 and 599
specimens, all from Cuba. The holotype and 1 1 paratypes were examined. The
holotype is housed in the U.S. National Museum of Natural History (Washington,
D.C.).

Distribution. Bahama Islands and Cuba (Map 7).

Specimens examined. 32 specimens collected during every month except January,
May, June, and November; deposited in AMNH, CAS, ISU, LHR, MSU, USNM.
BAHAMA ISLANDS: Andros Island: Mangrove Bay. Cat Island. CUBA: Boniato.
Archipielago de los Canarreos: Isla de Pines. Camagiiey: Camagiiey. Ciego de Avila:
Baragua. Cienfuegos: Soldad near Cienfuegos.- Granma: Cayamas. Pinar del Rio:

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THYANTA OF NORTH AND CENTRAL AMERICA

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Sierra Rangel. Sancti Spiritus: Zaza del Medio. Santiago de Cuba: Santiago de Cuba.
Villa Clara: Santa Clara.

Comments. Thyanta cubensis is often smaller than the other species related to T.
perditor, the humeral spines are shorter, the transhumeral reddish markings are
usually absent, and the pronotal and abdominal black markings are reduced or absent.

Thyanta ( Thyanta ) serratulata Ruckes
Figs. 109-121

Thyanta serratulata Ruckes, 1957c: 1 78-179, figs. 5-6.

Diagnosis. Body shape broad, stout.

Lateral jugal margins sinuous, not parallel (Fig. 1 10). Anterolateral pronotal mar-
gins immaculate, in dorsal view concave, serrate, especially anteriorly; humeral angles
spinose, spines relatively short, directed anterolaterad (Fig. 109). Pronotal cicatrices
immaculate or often marked with black in each mesial angle. Postspiracular black
spots absent. Anterolateral angles of abdominal stemites immaculate; posterolateral
abdominal angles concolorous with rest of segment or sometimes marked with black.

Basal plates in caudoventral view with mesial margins straight to slightly convex;
posterior margins sinuous (Fig. 1 16). Pygophoral opening subtended by semicircular
impression; posterior margin of pygophore produced posterodorsad, convex with
small, medial, V-shaped emargination in caudal view (Fig. 1 14); slightly concave in
lateral view (Fig. 1 1 5).

Types. Ruckes (1957c) described T. serratulata from 13<$<3 and 1322 specimens.
The holotype and 1 7 paratypes were examined. The holotype is housed in the Cal-
ifornia Academy of Sciences (San Francisco).

Distribution. Known only from the type locality: Clarion Island, Revillagigedo
Islands, Mexico.

Specimens examined. 21 specimens collected between 27 February and 8 May;
deposited in AMNH, CAS, LACM, USNM. MEXICO: Colima: Revillagigedo Is-
lands, Clarion Island.

Comments. Thyanta serratulata can be identified by the broad, stout shape, the
short humeral spine, the lack of post-spiracular black spots, and the reduction or
absence of black markings on the lateral abdominal angles.

Subgenus Argosoma Rider

Thyanta ( Argosoma ) Rider [in Rider and Chapin, 1991:33].

Type species. Pentatoma patruelis Stal, 1859 (by original designation, Rider and
Chapin, 1991).

Diagnosis. Punctation coarse, sparse, dorsal surface appearing glossy. Anterolateral
margins of pronotum straight to slightly concave, concolorous with surface of pro-
notum; humeral angles rounded to angulate, rarely spinose; pronotal cicatrices usually
immaculate, sometimes faintly marked with fuscous in mesial angles. Posterior ter-
mination of each buccula evanescent.

Distal end of sclerotized rod with or without subapical swelling, never cone-shaped;
spermathecal bulb globose; spermathecal duct below proximal flange slightly to great-
ly swollen and coiled, but never forming distinct cylindrical structure. Pygophoral

view. 1 15. Lateral view. 1 16. Genital plates, caudoventral view. 1 17-1 19. Right paramere. 1 17.
Medial view. 1 18. Ectal view. 1 19. Lateral view. 120. Spermatheca. 121. Spermathecal pump.

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THY ANT A OF NORTH AND CENTRAL AMERICA

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opening relatively large; posterior margin usually broadly and shallowly U-shaped;
postero ventral surface ofpygophore produced into blunt chin-like protuberance. Each
paramere acute to narrowly rounded apically, obtuse protuberance on shaft moderate
in size to absent, possessing distinct dorsomedial concave surface; roughened, spic-
ulate area on lateral surface of paramere usually circular, rarely linear ( T . boliviensis
Rider). Theca reniform, lacking dorsolateral protuberances; each lateral conjuctival
lobe usually with single diverticulum; median penial lobes and penisfilum usually
relatively small.

Comments. This is the largest subgenus, containing 20 species, and the included
species are also the most difficult to identify. It is often necessary to examine the
male genitalia in order to make accurate determinations. Within geographical areas,
the internal female genitalia are usually distinctive.

This subgenus can be divided into two groups based primarily on the structure of
the spermatheca. In T. boliviensis, T. brasiliensis Jensen-Haarup, T. emarginata
Rider, and T. hamulata Rider, the sclerotized rod is somewhat elongate and lacks
any subapical swelling. The remaining species have the sclerotized rod shorter and
distinctly swollen subapically, becoming narrowed apically.

Thyanta ( Argosoma ) planifrons Ruckes
Figs. 122-137, Map 6

Thyanta planifrons Ruckes, 1956:59-61, fig. 3; Rolston and McDonald, 1984:fig. 33.
Thyanta casta (of authors, not Stal): Uhler, 1894a:231 (part); Van Duzee, 1904:52,

54 (part); Kirkaldy, 1909:94 (part); Banks, 1910:90; Van Duzee, 1917:53 (part);

Van Duzee, 1923:127-128; Torre-Bueno, 1939:231 (part); Froeschner, 1988:593

(part).

Diagnosis. Vertex of head relatively flat; lateral jugal margins subparallel for middle
third of distance from eyes to apex (Fig. 123). Anterolateral margins of pronotum
straight to slightly concave in dorsal view; humeral angles rounded, not or only
slightly produced beyond base of adjacent corium (Fig. 122). Pronotal cicatrices
immaculate. Rarely with reddish markings on pronotum. Ostiolar canal widening
towards apex, wider distally than in middle (Fig. 137). Abdominal stemites lacking
black markings, rarely extreme tip of posterolateral angles of abdominal stemites
piceous.

Basal plates with mesial margins slightly convex in caudoventral view, separated
basally; posterior margins slightly convex; posteromesial angles rounded to slightly
emarginate (Fig. 134). Sclerotized rod slightly swollen subapically, narrowed apically
(Fig. 135); spermathecal duct only slightly swollen below proximal flange (Fig. 136).
Posterior margin of pygophore broadly and shallowly U-shaped in caudal view,
slightly sinuous medially (Fig. 1 30); posterior margin weakly concave in ventral and
dorsal views (Figs. 131, 132); posteroventral surface deeply emarginate in lateral
view (Fig. 133). Apex of each paramere nearly acute from both medial and ectal
views (Figs. 1 24, 125); concave surface oriented more dorsad than mediad; roughened
spiculate area on lateral surface elongate-circular (Fig. 126). Each lateral conjunctival
lobe of aedeagus with 1 or 2 non-sclerotized diverticula (Fig. 129); dorsomedial lobe
apparently absent (Fig. 128); median penial lobes spatulate (Fig. 127).

Types. Ruckes (1956) described T. planifrons from 6<3<3 and 729 specimens. The

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tyl

Medial view. 125. Ectal view. 126. Lateral view. 127-129. Theca and related structures. 127.
Ventral view. 128. Dorsal view. 129. Lateral view. 130-133. Pygophore. 130. Caudal view.
131. Ventral view. 132. Dorsal view. 133. Lateral view. 134. Genital plates, caudoventral view.

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THY ANT A OF NORTH AND CENTRAL AMERICA

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holotype, which is from 10 miles west of Alamos, Sonora, Mexico, and all 12 para-
types were examined. The holotype is housed in the American Museum of Natural
History (New York).

Distribution. Southwestern U.S. and northwestern Mexico (Map 6).

Specimens examined. 535 specimens collected during every month except Feb-
ruary; deposited in AMNH, ASUT, CAS, DAR, DBT, EGER, FSCA, LACM, LHR,
MSU, TAMU, UAT, UCB, UCR, UIM, UMC, UNAM, USNM. UNITED STATES:
Arizona: Cochise: Portal; 5 mi W Portal, SWRS. Pima: Baboquivari Mountains,
Browns Canyon; Organ Pipe Cactus Natl Monument. Santa Cruz: Madera Canyon.
California: Imperial. Riverside: Bautista Canyon; Deep Canyon; Palm Springs. San
Diego: Borego Valley.

MEXICO: Canipole; 10 mi SW Canipole; Carmen Island, Porto Ballandra; Punt-
bunda. Baja California Norte: Bahia de los Angeles; Catavina; 1 0 mi S Catavina; 8
km N Punta Prieta; 1 5 mi N Punta Prieta; San Felipe; 1 2 mi S San Felipe; 1 5 mi S
San Felipe; San Fernando. Baja California Sur: Bahia Concepcion; Cabo San Lucas;
3 mi W Caduano; Comondu; 10 mi SW Comondu; 20 mi N Comondu; 23 mi S
Comondu; 14 mi S El Arco Mine; 28 mi S El Arco Mine; 6.5 mi S, 1 mi E El
Pescadero; 15 mi N El Rufugio; El Sargento; El Triunfo; 2 mi NW El Triunfo; 6 mi
N El Triunfo; Escondido Bay; 3 mi N Guajademi; Hamilton Ranch; 1 km SW
Huatamote; Isla Annelvo; Isla Catalan; Isla Cerralvo; Isla Espirita Santo; La Paz; 2
mi S La Paz; 5 mi SW La Paz; 7 mi SW La Paz; 13 mi W La Paz; 14 mi W La Paz;
15 mi W La Paz; 20 mi NW La Paz; 21 mi W La Paz; 23 km W La Paz; 25 mi W
La Paz; 26 mi W La Paz; 33.5 km NW La Paz; La Purisima; Las Animas; Las
Barracas; Las Tinajitas; 2 mi SE Las Virgenes; 1 mi E Migriho; Miraflores; 5 mi S
Miraflores; 4 mi S Mission San Javier; Mulege; 1 mi S Mulege; 2 mi S Rancho de
la Ventana; 2.6 mi E San Antonio; 3 mi SW San Antonio; 5 mi S San Antonio; 5
mi W San Bartolo; San Domingo; 1 5 mi S San Domingo; 1 5 mi N San Ignacio; 27
mi W San Ignacio; San Jose del Cabo; 2 km W San Jose del Cabo; 10 mi SW San
Jose del Cabo; 1.3 mi N San Jose Viejo; 3 mi N San Jose Viejo; 5 mi S San Miguel;
3 km S, 1.3 km E San Pedro; 3.5 mi NE San Pedro; San Sebastian; 5 mi SE Santa
Rosalia; 12 mi S Santa Rosalia; Santiago; 6 mi SW Santiago; Sierra de la Laguna;
Todos Santos; 4 mi N Todos Santos; 28-29 km N Todos Santos; Venancio; 30 mi
E V. Insurgentes. Sinaloa: Mazatlan; 34 mi N Mazatlan. Sonora: 10 mi W Alamos;
Bahia de los San Carlos; Bahia Kino; 20 mi NNE Ciudad Obregon; El Desemboque;
Guaymas, Punta San Antonio; Hermosillo; La Choya; Minas Nuevas; 15 km S
Navojoa; 15 mi N Navojoa; San Bernardino, Rio Mayo; 20 mi S Sonoyta; Tecoripa;
Tiburon Island; Yavaros.

Comments. Thyanta planifrons can be separated from all other congeners by the
distal widening of the ostiolar canal. Also, the vertex of the head is relatively flat, a
character for which this species was named.

135. Spermatheca. 136. Spermathecal pump. 137. Ostiolar canal. Symbols: bp, basal plate; dfl,
distal flange; dsp, dilation of spermatheca: gx2, second gonacoxae; jug, juga; lcl, lateral con-
junctival lobe; mpl, median penial lobe; oc, ostiolar canal; pen, penisfilum; pfl, proximal flange;
pla, posterolateral angle of pygophore; pmp, posterior margin of pygophore; pt8, eighth para-
tergite; pt9, ninth paratergite; rsa, roughened spiculate area on lateral surface of paramere; spb,
spermathecal bulb; sr, sclerotized rod; slO, tenth stemite; th, theca; tyl, tylus.

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The widening of the ostiolar canal is unusual, but not unique. It also occurs in
Tepa jugosa Van Duzee, a species with essentially the same distribution as Thyanta
planifrons. The biological significance of this condition is not known, but would make
an interesting study. This is the first record of this species from the United States.

Thyanta ( Argosoma ) maculata (Fabricius)

Figs. 138-152, Map 8

Cimex maculatus Fabricius, 1775:704.

Thyanta casta Stal, 1862b: 104; Stal, 1872:35; Uhler, 1876:7; Distant, 1 880:66; Uhler,
1886:7; Uhler, 1894a:231 (part); Lethierry and Severin, 1893:147; Kirkaldy, 1909:
94 (part); Malloch, 1919:217, fig. 74; Torre-Bueno, 1939:231 (part); Froeschner,
1988:593 (part). NEW SYNONYMY.

Euschistus castus: Walker, 1867:244.

Thyanta maculata: Stal, 1872:35; Distant, 1893:334; Lethierry and Severin, 1893:
148; Kirkaldy, 1909:94; Rolston and McDonald, 1984:fig. 29.

Diagnosis. General color green to brown; often with varying amounts of reddish
on pronotum between humeral angles, often forming two oblong spots, one on each
side of middle. Apices of scutellum and coria occasionally rubiginous.

Lateral jugal margins sinuous, not quite parallel (Fig. 139). Anterolateral margins
of pronotum nearly straight in dorsal view; humeral angles rounded to angulate,
usually produced beyond base of adjacent corium by less than half width of eye (Fig.
138); pronotal cicatrices immaculate. Ostiolar canals acuminate apically. Postero-
lateral angles of abdominal stemites piceous, sometimes only minutely so. Postspi-
racular spots usually absent, though sometimes present in brown form.

Basal plates with mesial margins slightly convex in caudoventral view, separated
basally; posterior margins convex; posteromesial angles slightly emarginate (Fig. 1 50).
Sclerotized rod swollen subapically, narrowed apically (Fig. 151); spermathecal duct
moderately swollen and coiled below proximal flange (Fig. 1 52). Posterior margin of
pygophore broadly and shallowly U-shaped in caudal view (Fig. 146); lateral angles
prominent in ventral and dorsal views, chin-like protuberance relatively small (Figs.
147, 148); emarginate below middle in lateral view (Fig. 149). Apex of each paramere
obtusely rounded in ectal view (Fig. 141); concave surface oriented more mediad
than dorsad, shaft rather robust with small medial protuberance (Fig. 1 40); roughened
spiculate area on lateral surface circular (Fig. 142). Aedeagus with lateral and dor-
somedial conjunctival lobes large (Fig. 144), each lateral lobe with single, slightly
sclerotized diverticulum (Fig. 143); penisfilum and median penial lobes obscured by
conjunctiva (Fig. 145).

Types. Fabricius (1775) described Cimex maculatus from “America” without des-
ignating a holotype or paratypes. It is not possible to determine from his original
description how many specimens he examined, but evidently he had more than one,
as he mentions a variation. Only one syntype was located and examined. It is a 9 in
poor condition (apex of abdomen destroyed, left forewing missing, etc.), but it does
possess the characters that define this species. It has the following label data: (a)
“maculatus” (b) “Thyanta maculata F.,” and is here designated the lectotype. This
specimen is housed in the Universetetets Zoologiske Museum (Copehagen, Den-
mark).

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THY ANT A OF NORTH AND CENTRAL AMERICA

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Figs. 138-152. T. maculata. 138. Habitus. 139. Head. 140-142. Right paramere. 140.
Medial view. 141. Ectal view. 142. Lateral view. 143-145. Theca and related structures. 143.
Ventral view. 144. Dorsal view. 145. Lateral view. 146-149. Pygophore. 146. Caudal view.
147. Ventral view. 148. Dorsal view. 149. Lateral view. 1 50. Genital plates, caudo ventral view.
151. Spermatheca. 152. Spermathecal pump. Symbol: dmc, dorsomedial conjunctival lobe.

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Stal (1862b) described T. casta from Mexico without designating a holotype or
paratypes. One syntype, a 2, was located and examined. Although it lacks the reddish
markings that many specimens of T. maculata possess, it differs structurally in no
significant manner from T. maculata. Because it is not possible to determine the
number of specimens upon which St&Ts description was based, the syntype specimen
is designated the lectotype. It is labeled as follows: (a) “Mexico Coll. Signoret.” (b)
“Casta det. Stal” (c) “TYPE” (d) “Coll. Nat.-Mus. Wien” (e) “Thyanta casta STAL.”
The lectotype is housed in the Naturhistorisches Museum (Vienna, Austria).

Distribution. Southern Texas southward through Mexico and Central America to
southern Panama (Map 8).

Specimens examined. 499 specimens collected during every month of the year;
deposited in AMNH, ARH, BMNH, CAS, CNC, CUC, DAR, DBT, EGER, ENGL,
FSCA, INHS, LACM, LHR, LSU, MSU, MSUE, OSU, OSUC, SMEK, TAMU,
UAT, UCB, UCS, UGA, UMAA, UNAM, UNSM, UUSL. UNITED STATES:
Texas: Brazos. Burnet: Inks Lake State Park. Cameron: Boca Chica; 3 mi SW Boca
Chica; Brownsville; Harlingen; Sabal Palm Grove Sanctuary near Southmost. Hi-
dalgo: Bentsen Rio Grande Valley State Park; Edinburg; McAllen; Progresso; Santa
Ana Natl Wildlife Refuge. Kleburg: Kingsville. Presidio: Presidio. San Patricio: Cor-
pus Christi Lake State Park; Nueces River, 5 mi SW Mathis. Starr: Falcon Heights;
4-15 mi N Roma. Webb: Laredo. Zapata: 1 mi E Falcon Lake; Falcon State Park.

MEXICO: Campeche: Calkini; 1 2 mi E Campeche; El Remate; Escarcega; Ruinas
Edzna; km 54, Carr. Campeche-Merida. Chiapas: Aguacero, 16 km W Ocozocoautla;
Chicoasen Dam Area; 1 3 mi W Cintalpa; 2 mi N Ocozcoautla; Ruinas Bonampak;
San Cristobal de las Casas; Santo Domingo, 15 mi SE Simojovel; Simojovel; Su-
chiapa. Coahuila: Arroyo de la Zorra. Colima: 2 mi N Manzanillo. Guerrero: Aca-
pulco de Juarez; 3.7 mi E Marquelia; Tecpan de Galeana. Hidalgo: Otongo. Jalisco:

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Chamela; Estacion de Biologia Chamela. Michoacan: Acahuato. Morelos: 4.4 mi E
Cuernavaca; Villa de Ayala. Nayarit: 1 5 km E San Bias. Nuevo Leon: Apodaca; 3 mi
E Galeana; 16 mi S Linares, Anegade Arroyo; 4.1 mi S Montemorelos; Monterrey;

4 mi S Monterrey; 5 mi S Monterrey, Valle Alto; 6 mi S Monterrey. Oaxaca: 2.7
mi NW El Camaron; El Charquito; 6 mi W Jalapa del Marques; 8 mi N La Ventosa;
Puerto Escondido; Salina Cruz; Tehuantepec; 1 1 mi W Tehuantepec; 12 mi W Te-
huantepec; 13 km W Tehuantepec; 44 mi W Tehuantepec; 2.1 mi NW Totolapan.
Queretaro: 1 mi NW Ayutla. Quintana Roo: 1 km N Coba; 20 km N Felipe Carrillo
Puerto; 54 mi SE Peto; San Isidrio Puerto Morelos; 2 mi NE San Miguel; River El
Ramonal; Rancho El 24; Xcun Nuevo; km 146, Carr. Chetumal-Cancun; km 146,
Carr. Chetumal-Pto Juarez. San Luis Potosi: 12 mi S Ciudad Mante; Ciudad Valles;
1 1 km E Ciudad Valles; El Banito; El Salto Falls; Tamazunchale; 5 mi N Tamazun-
chale; 30 mi S Tamazunchale; Tamuin. Sinaloa: 1 mi NW El Venadillo; Mazatlan;

5 mi N Mazatlan; 10 mi S Mazatlan; Presidio River near Caton. Sonora: 7 mi W
Alamos. Tamaulipas: Abasolo; Antiguo Morelos; Bocatoma, Ciudad Victoria; 6 mi
S Ciudad Victoria; 25 mi S Ciudad Victoria; 5 mi SSE Gomez Farias; 6 mi S Gomez
Farias; Guemes; Hidalgo; 8 mi N Jimenez; La Pesca; Llera; 5 mi N Llera; 13 km E
Magiscatzin; Punta Piedras; Rio Corona, 30 km N Ciudad Victoria; San Fernando;
6.2 mi S San Fernando; 25 mi SE San Fernando; Tampico; Villagran. Veracruz:
Cordoba; Cotaxtla; Cotaxtla Expt Stn; Cuitlahuac; Lake Catemaco Area; Ojo de Agui;
Puente Nacionale; San Andres Tuxtla; 5 mi S Santiago Tuxtla; Tolome; 10 mi W
Veracruz. Yucatan: Chichen Itza; Piste; Progreso; 12 km N Quintana Roo, Hwy 295;
13 mi E Valladolid; 13.3 mi S Valladolid; 1 km S Xcalacoop; 10 km N Xcalacoop.

BELIZE: Belize: 12 mi NW Belize City. EL SALVADOR: Ruinas San Andres.
HONDURAS: Choluteca. Comayagua: 5 mi NW Comayagua. Cortes: Puerto Cortes.
NICARAGUA: Chinandega; N side Cosgoina Volcano, Gulf of Fonseca. Managua:
Jiloa; Masachapa. COSTA RICA: Limon. Guanacaste: La Pacifica, near Canas. PAN-
AMA: Chitre; La Chorrera. Canal Zone: Barro Colorado Island; Base of Cerro Galera.
Darien: Santa Fe.

Comments. Thyanta maculata was originally distinguished from other congeners
by the presence of two oblong reddish macules, one on each side of the middle of
the pronotum. Fewer than half the specimens of T. maculata actually exhibit this
character. Also, several other species of Thyanta are known to have the same type
of maculation in at least some specimens ( T . pseudocasta Blatchley, T. brasiliensis
Jensen-Haarup, T. curvata Rider).

Thyanta maculata can be separated from other congeners except T. pseudocasta
by the structure of the male genitalia. Each paramere is rather robust with the apex
rounded, and the dorsomedial concave surface is oriented more mediad than dorsad.
Thyanta pseudocasta has very similar male genitalia, but the aedeagus is slightly
different. The aedeagus of T. maculata has a dorsomedial conjunctival lobe, while
the same structure is apparently absent in T. pseudocasta. The two forms can usually
be separated based upon the prominence of the humeral angles. Each humeral angle
in T. maculata is usually produced beyond the base of the adjacent corium by less
than half the width of an eye, while in T. pseudocasta each humeral angle usually
protrudes beyond the corium by more than half the width of an eye.

Thyanta maculata is a variable species, especially with respect to size and color-
ation. For example, specimens from the Yucatan peninsula of Mexico average much

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smaller than specimens from other parts of the range, but they do not differ in any
other significant manner.

Thyanta ( Argosoma ) vadosa Rider
Figs. 153-167

Thyanta {Argosoma) vadosa Rider [in Rider and Chapin, 1991:55].

Diagnosis. Ovate; dorsal surface green to pale brown; some interstitial areas of
pronotum, scutellum, and elytra pale yellow; sometimes marked with reddish-purple
between humeral angles, on apex of scutellum, and on tylus and vertex of head.
Punctures green to pale brown.

Apex of head arcuately rounded; lateral jugal margins sinuous, subparallel for
middle third of distance from eyes to apex (Fig. 1 54); vertex convex. Anterolateral
margins of pronotum in dorsal view straight to slightly concave; humeral angles
rounded to angulate, often projecting beyond base of adjacent corium (Fig. 153).
Pronotal cicatrices immaculate. Punctation becoming sparse medially, central portion
of pronotal disc subcalloused. Posterior third of pronotum often darker than rest of
pronotum. Posterolateral angles of connexival segments piceous. Ostiolar canals acu-
minate apically. Postspiracular black spots usually absent (except in brown form);
posterolateral angles of abdominal stemites piceous, sometimes only minutely so.

Mesial margins of basal plates straight to slightly convex in caudoventral view;
posterior margins slightly convex; posteromesial angles broadly and shallowly emar-
ginate, lateral sides of concavity resulting from excavations in basal plates divergent,
not parallel (Fig. 165). Distal end of sclerotized rod swollen subapically, narrowed
apically (Fig. 166); spermathecal duct moderately swollen and coiled below proximal
flange (Fig. 167). Posterior margin of pygophore in caudal view broadly U-shaped,
medial portion straight to slightly convex (Fig. 161); chin-like protuberance appearing
relatively narrow in ventral and dorsal views (Figs. 162, 163); pygophore deeply
emarginate in lateral view (Fig. 1 64). Each paramere with concave surface oriented
mediad; from ectal view, parameral apex angling gently mesad (Fig. 1 56); from medial
view, apex acutely angulate, straight or bending slightly ventrad (Fig. 155); roughened
spiculate area on lateral surface circular (Fig. 157). Each lateral conjunctival lobe of
aedeagus without sclerotized diverticula (Fig. 160); dorsomedial conjunctival lobe
weakly developed (Fig. 159); median penial lobes spatulate (Fig. 158).

Types. Rider [in Rider and Chapin, 1991] described T. vadosa from 5 66 and 599
female specimens. The holotype 6 was examined and is deposited in the Canadian
National Collection, Ottawa, Canada.

Distribution. Trinidad and Tobago; Venezuela.

Specimens examined. 9 specimens collected in January, February, March, May,
July, September, and October, deposited in AMNH, ARH, CNC, USNM. BRITISH
WEST INDIES: Tobago. Trinidad: Bejucal; Curepe, Santa Margarita Circular Road;
Saint Augustine; Santa Margarita Hill.

Comments. In general appearance this is a typical species of the maculata group.
The shape of the emargination in the posteromesial angle of each basal plate of the
female is distinctive. Thyanta emarginata and T. excavata both have the postero-
mesial angles of the basal plates deeply emarginate, but the sides of the resulting
concavity are nearly parallel, not divergent as in T. vadosa. The male genitalia are

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THY ANT A OF NORTH AND CENTRAL AMERICA

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Figs. 153-167. T. vadosa. 153. Habitus. 154. Head. 155-157. Right paramere. 155. Medial
view. 156. Ectal view. 157. Lateral view. 158-160. Theca and related structures. 158. Ventral
view. 159. Dorsal view. 160. Lateral view. 161-164. Pygophore. 161. Caudal view. 162. Ventral
view. 163. Dorsal view. 164. Lateral view. 165. Genital plates, caudo ventral view. 166. Sper-
matheca. 167. Spermathecal pump.

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also distinctive. Thyanta vadosa is the only species with the apex of each paramere
not only acutely angulate (almost acuminate) but also straight or bending slightly
ventrad. In the maculata group, all other species having the apex of each paramere
acute to acuminate also have the apex bending dorsad.

Thyanta ( Argosoma ) pseudocasta Blatchley
Figs. 168-182, Map 9

Thyanta pseudocasta Blatchley, 1926:114, 120; Blatchley, 1930:64; Torre-Bueno,

1939:230; Rolston and McDonald, 1984:figs. 26, 34; Froeschner, 1988:594.
Thyanta casta (of authors, not Stal): Barber, 1914:523; Van Duzee, 1917:53 (part);

Torre-Bueno, 1939:231 (part); Froeschner, 1988:593 (part).

Diagnosis. General color pale green to pale brown, sometimes with anterior half
of pronotum reddish.

Lateral jugal margins subparallel for middle third of distance from eyes to apex
(Fig. 169). Anterolateral margins of pronotum nearly straight in dorsal view; humeral
angles prominent, produced beyond base of adjacent corium by more than half width
of eye, rounded to angulate (Fig. 1 68). Pronotal cicatrices immaculate. Ostiolar canals
acuminate apically. Posterolateral angles of abdominal stemites piceous, sometimes
only minutely so. Postspiracular black spots usually absent, sometimes present in
brown form.

Basal plates in caudoventral view with mesial margins straight to slightly convex;
posterior margins evenly convex, posteromesial angles rounded (Fig. 1 80). Sclerotized
rod slightly swollen subapically, narrowed apically (Fig. 181); spermathecal duct with
small amount of swelling and coiling below proximal flange (Fig. 182). Posterior
margin of pygophore in caudal view broadly and shallowly U-shaped, slightly sinuous
medially in caudal, ventral, and dorsal views (Figs. 176-178); pygophore emarginate
in lateral view (Fig. 179). Apex of each paramere from ectal view nearly acute (Fig.
171); from medial view narrowly rounded; concave surface facing mediad (Fig. 1 70);
roughened spiculate area on lateral surface circular (Fig. 172). Aedeagus with each
lateral conjunctival lobe apparently lacking sclerotized diverticula (Fig. 175); dor-
somedial lobe apparently absent (Fig. 174); median penial lobes relatively small (Fig.
173).

Types. Blatchley (1926) described T. pseudocasta without designating a holotype
or paratypes. In 1930, however, he designated 1 6 specimen as “type” (lectotype). He
did not mention any of the other syntypes. Although actual paralectotype designations
were not made, according to the International Code of Zoological Nomenclature
(Ride et al., 1985, sect. 74a[iv]), once the lectotype is designated the remaining
syntypes automatically become paralectotypes. Accordingly, no such designations
are needed, but labels have been added to the specimens to indicate their actual
status.

Regrettably, the $ lectotype is in deplorable condition; all that remains is the head
and pronotum. It has the following label data: (a) “Miami Fla. W. S. B. Coll. 3-11-
24” (b) “Purdue Blatchley collection” (c) “TYPE” (d) “Thyanta pseudocasta Blatch-
ley.” The lectotype is deposited in the Purdue University Collection (W. Lafayette,
IN).

Ten additional specimens that are believed to be part of the original syntype series

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THY ANT A OF NORTH AND CENTRAL AMERICA

87

Medial view. 171. Ectal view. 172. Lateral view. 173-175. Theca and related structures. 173.
Ventral view. 174. Dorsal view. 175. Lateral view. 176-179. Pygophore. 176. Caudal view.
177. Ventral view. 178. Dorsal view. 179. Lateral view. 180. Genital plates, caudoventral view.
181. Spermatheca. 182. Spermathecal pump.

have been located: 16 and 299 labeled (a) “Miami Fla. W. S. B. Coll. 3-1 1-24” (b)
“Purdue Blatchley collection” (AMNH, PUL), except 19 labeled (c) “Thyanta pseu-
docasta Blatchley” (LSU); 366 and 299 labeled (a) “C. Sable Fla. W. S. B. Coll. 2-23-
19” (b) “Purdue Blatchley collection” (PUL), except 16 labeled (b) “H G Barber
Colin 1950” (USNM), and 16 and 19 labeled “4-5-25” (AMNH); 16 labeled (a) “K.

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Map 9. T. pseudocasta.

West Fla. W. S. B. Coll. 3-2-19” (b) “Purdue Blatchley collection” (PUL); and 1<5
labeled (a) “Coxam Fla. W. S. B. Coll. 3-8-21” (b) “Purdue Blatchley collection”
(LSU). The lectotype and all but one paralectotype were examined.

Distribution. Southern Florida (Map 9).

Specimens examined. 153 specimens collected during every month of the year;
deposited in AMNH, ARH, CAS, CNC, CU, DAR, DBT, EGER, FSCA, LHR, LSU,

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THY ANT A OF NORTH AND CENTRAL AMERICA

89

MSU, PUL, SMEK, UCB, UCS, UGA, UMC, USNM. UNITED STATES: Florida:
Caxambus. Broward: Fort Lauderdale. Charlotte: Charlotte Harbor Area, Little Gas-
parillo Island; Punta Gorda. Collier: Marco Island; Royal Palm Park. Dade: Biscayne
Bay; Coral Gables; Homestead; Miami. Lee: Sannibel Island. Monroe: Big Pine Key;
Cape Sable; Everglades Natl Park; Key Largo; Key West; Marathon Key; Plantation
Key; Stock Island; Tavernier Key; 9 mi NW Key Largo.

Comments. Thyanta pseudocasta can be separated from all congeners except T.
maculata because the concave surface of its paramere is oriented more mediad than
dorsad. Thyanta pseudocasta may actually be a subspecies of T. maculata. The male
genitalia of the two species are very similar, but there are some differences in the
aedeagus. Thyanta maculata has a prominent dorsomedial conjunctival lobe, while
this structure is apparently absent in T. pseudocasta. Also, the humeral angles are
more prominent in T. pseudocasta. Because these species represent reproductively
isolated populations, a conservative approach is taken, and they are retained as full
species.

Thyanta ( Argosoma ) obsoleta (Dallas)

Figs. 183-197, Map 7

Pentatoma obsoleta Dallas, 185 1:25 1; Walker, 1867:289.

Thyanta obsoleta: Lethierry and Severin, 1893:148; Kirkaldy, 1909:94.

Thyanta casta (of authors, not Stal): Van Duzee, 1904:52, 54 (part); Kirkaldy, 1909:

94 (part); Barber, 1923:12; Barber, 1939:292-293.

Diagnosis. Coloration green to pale brown, often with transhumeral reddish mark-
ings that sometimes form two oblong spots, one each side of middle.

Lateral jugal margins subparallel for middle third of distance from eyes to apex
(Fig. 1 84). Anterolateral margins of pronotum straight to slightly concave in dorsal
view; humeral angles rounded to angulate, usually produced beyond base of adjacent
corium by about half width of eye (Fig. 183). Pronotal cicatrices immaculate. Ostiolar
canals acuminate apically. Posterolateral abdominal angles piceous; postspiracular
black spots usually absent, sometimes evident in brown form.

Mesial margins of basal plates in caudoventral view straight to slightly convex;
posterior margins slightly convex; posteromesial angles broadly rounded (Fig. 195).
Sclerotized rod swollen subapically, narrowed apically (Fig. 1 96); spermathecal duct
moderately swollen below proximal flange with only slight amount of coiling (Fig.
197). Posterior margin of pygophore broadly and shallowly U-shaped in caudal view
(Fig. 191); in lateral view concave (Fig. 194); posterior margin only slightly concave
in dorsal view, posterolateral angles not prominent (Fig. 193); postero ventral surface
distinctly produced into blunt chin-like protuberance in ventral view (Fig. 192). Each
paramere apically rounded in ectal view (Fig. 186); narrowly rounded from medial
view, curving dorsad; concave surface oriented more dorsad than mediad (Fig. 185);
roughened spiculate area on lateral surface circular (Fig. 187). Each lateral conjunc-
tival lobe of aedeagus with 1 or 2 subacute diverticula (Fig. 190), dorsomedial lobe
reduced (Fig. 1 89); median penial lobes large, spatulate (Fig. 1 88); penisfilum reduced.

Types. Dallas (1851) described Pentatoma obsoleta from Jamaica without desig-
nating a holotype or paratypes, and it is not possible to determine the number of
specimens upon which he based his description. Only 19 syntype was located, and

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Figs. 183-197. T. obsoleta. 183. Habitus. 184. Head. 185-187. Right paramere. 185. Medial
view. 186. Ectal view. 187. Lateral view. 188-190. Theca and related structures. 188. Ventral
view. 189. Dorsal view. 1 90. Lateral view. 191-194. Pygophore. 1 9 1 . Caudal view. 192. Ventral
view. 193. Dorsal view. 194. Lateral view. 195. Genital plates, caudoventral view. 196. Sper-
matheca. 197. Spermathecal pump.

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THY ANT A OF NORTH AND CENTRAL AMERICA

91

it is here designated the lectotype. It is labeled (a) “Jamaica” [dorsal surface], “45
1 1 1 1” [ventral surface] (b) “Type” (c) “35. Pentatoma obsoleta .” The lectotype, which
is housed in the British Museum of Natural History (London), was examined.

Distribution. Greater Antilles (Map 7).

Specimens examined. 60 specimens collected during every month of the year except
September; deposited in AMNH, ARH, BMNH, CAS, CNC, CU, DAR, DBT, ENGL,
LHR, OSU, SMEK, UAT, USNM. BAHAMA ISLANDS: San Salvador Island.
CUBA: Ciudad de la Habana: El Cano; Havana. Guantanamo: Guantanamo Bay
Naval Base, Caravella Point. Santiago de Cuba: Daiquiri; Jarahueca. JAMAICA:
Bluefields; Christiana; Mona, near Kingston; St. Andrew Ferry; Try. Duncans. St.
Andrew: Bamboo Lodge near Irish Town. Westmorland: Negril, Negrillo Cottages.
HAITI: Diquini. Ouest: Port-au-Prince. DOMINICAN REPUBLIC: Los Hidalgos;
8 mi up Macons River, Santo Domingo. Distrito Nacional: La Victoria; Santo Do-
mingo City. Peravia: 1 3 km NW Bam. Samana: Sanchez. San Cristobal. San Juan:
16 km SE San Juan; 28 km E San Juan. PUERTO RICO: Aguirre; Fortuna A.E.S.
Humacao: Vieques Island, Puerto Real. Mayaguez: Guanica Forest, Hwy 334 at
Ranger Station. Ponce: Coamo Springs; Ponce.

Comments. Examination of the male genitalia is necessary to separate this species
from other species in the maculata group. Thyanta obsoleta can be distinguished
from other Northern Hemisphere species by its apically rounded parameres, which
have the concave surface oriented more dorsad than mediad.

Thyanta ( Argosoma ) testacea (Dallas)

Figs. 198-212, Map 7

Pentatoma testacea Dallas, 1851:250; Walker, 1867:289.

Thyanta testacea: St&l, 1872:35; Berg, 1878:23, Lethierry and Severin, 1893:148;
Kirkaldy, 1909:95.

Thyanta casta (of authors, not Stal): Uhler, 1893:705; Uhler, 1894b: 174.

Thyanta signoreti Ruckes, 1956:65-66, fig. 7 (syn. by Rider and Chapin, 1991).
Thyanta {Argosoma) testacea: Rider and Chapin, 1991.

Diagnosis. General color green to brown, rarely with rubiginous transhumeral
markings.

Lateral jugal margins subparallel for middle third of distance from eyes to apex
(Fig. 1 99). Anterolateral pronotal margins straight to slightly concave; humeral angles
angulate to rounded, usually produced beyond base of adjacent corium by about half
width of eye (Fig. 198). Pronotal cicatrices immaculate. Ostiolar canals acuminate
apically. Posterolateral abdominal angles not marked with black or only minutely
so; postspiracular black spots usually absent, sometimes evident in brown form.

Basal plates in caudoventral view with mesial margins convex, separated basally;
posterior margins convex (Fig. 210). Distal end of sclerotized rod slightly swollen
subapically, narrowed apically (Fig. 211); spermathecal duct greatly swollen below
proximal flange, carrot-shaped (Fig. 212). Posterior margin of pygophore broadly
and shallowly U-shaped in caudal view (Fig. 206); slightly concave in lateral view
(Fig. 209). Each paramere apically acute in both medial and ectal views (Figs. 200,
201); concave surface oriented more dorsad than mediad; roughened spiculate area

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

view. 201. Ectal view. 202. Lateral view. 203-205. Theca and related structures. 203. Ventral
view. 204. Dorsal view. 205. Lateral view. 206-209. Pygophore. 206. Caudal view. 207. Ventral
view. 208. Dorsal view. 209. Lateral view. 210. Genital plates, caudoventral view. 211. Sper-
matheca. 212. Spermathecal pump.

on lateral surface circular (Fig. 202). Aedeagus with dorsomedial lobe apparently
absent (Fig. 204).

Types. Dallas (1851) described Pentotoma testacea from “S. America” without
designating a holotype or paratypes, and it is not possible to determine how many

1992

THY ANT A OF NORTH AND CENTRAL AMERICA

93

syntypes he had. Rider and Chapin (1991) designated the only known $ syntype as
lectotype. The lectotype, which is conserved in the British Museum of Natural History
(London), was examined.

Distribution. Lesser Antilles and northern South America (Map 7).

Specimens examined. 250 specimens collected during every month of the year,
deposited in AMNH, ARH, BMNH, CAS, CU, DBT, EGER, ENGL, INHS, LACM,
LHR, LSU, MSUE, SMEK, TAMU, USNM. BRITISH VIRGIN ISLANDS: Tortola
Island. U.S. VIRGIN ISLANDS: St. Croix: Canaan; Christiansted; E Hill; Experi-
ment Station Grounds; Hams Bluff. St. John: Estate Carolina, NW of Coral Bay;
Virgin Islands Natl Park. St. Thomas: Charlotte Amalie; Estate Lilliendahl; French-
man’s Bay. BRITISH WEST INDIES: Anguilla: N of Road Bay. Antigua: Coolidge;
Coolidge airport. Bequia. Dominica: Antrim; Cabrit Swamp; Clarke Hall; Grande
Savane; Macoucheri; Mero Beach; Salybia; Springfield Estate; S Chiltem. Grenada:
Caliveny Estate; Grand Anse, St. Georges Parish; Granville; Mt Gay Estate; St.
Georges; Santeurs. Montserrat: Galway’s Estate; Plymouth. St. Kitts: W Farm Gut.
St. Lucia: 1.5 mi N Canaries. Tobago: Bucco Bay; Grafton Estate. Trinidad: St.
George Co., Curepe C.I.B.C. Union. FRENCH WEST INDIES: Guadeloupe: Sur
Cotonnier. Martinique: Diamant; Sainte Anne. BARBADOS: Edge Hill; Freshwater
Bay; Groves St. George. CURASAO.

Comments. Thyanta testacea is very closely related to the other species in the
maculata group, and can be reliably identified only by examination of the male
genitalia. The apically acute parameres curving gently dorsad will separate it from
all congeners except T. patruelis, which it closely resembles. The chin-like protu-
berance on the posteroventral surface of the pygophore is somewhat less prominent
in T. testacea than in T. patruelis. There does seem to be a geographical separation
of the two forms with T. testacea restricted to northern South America and the Lesser
Antilles, and T. patruelis occurring from northeastern Brazil and southern Peru
southward.

ACKNOWLEDGMENTS

This study would not have been possible without the kind help of the many curators and
colleagues who generously lent specimens. The following is a list of institutions and their curators
who provided valuable assistance (acronyms are those used in the text; DAR is senior author’s
collection): AMNH— American Museum of Natural History, New York, R. T. Schuh; ARH—
University of Florida, Agricultural Research Center, Homestead, R. M. Baranowski; ASUT—
Arizona State University, Tempe, F. F. Hasbrouck; AUA— Auburn University, Auburn, AL,
W. E. Clark; BMNH— British Museum (Natural History), London, England, W. R. Dolling;
CAS— California Academy of Sciences, San Francisco, P. H. Amaud, Jr.; CNC— Canadian
National Collection, Ottawa, Ontario, R. Foottit; CU— Cornell University, Ithaca, NY, J. K.
Liebherr; CUC— Clemson University, Clemson, SC, J. C. Morse; DBT— D. B. Thomas personal
collection, Tuxtla Gutierrez, Mexico; EGER— J. E. Eger personal collection, Tampa, FL; ENGL—
H. D. Engleman personal collection, Coco Solo, Panama; FMNH— Field Museum of Natural
History, Chicago, IL, E. H. Smith; FSCA— Florida State Collection of Arthropods, Gainesville,
F. W. Mead; INHS— Illinois Natural History Survey, Champaign, D. Voegtlin; ISU— Iowa
State University, Ames, J. Laffoon; LACM— Los Angeles County Museum of Natural History,
CA, C. L. Hogue; LHR— L. H. Rolston personal collection, Baton Rouge, LA; LSU — Louisiana
State University, Baton Rouge, J. B. Chapin; MSU — Mississippi State University, Mississippi
State, R. L. Brown, P. R. Miller; MSUB— Montana State University, Bozeman, S. Rose; MSUE—
Michigan State University, East Lansing, R. L. Fischer; NCSU— North Carolina State Uni-

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versity, Raleigh, C. Parron; NDSF— North Dakota State University, Fargo, E. U. Balsbaugh,
Jr.; NMSU— New Mexico State University, Las Cruces, G. S. Forbes, J. R. Zimmerman;
ODAS— Oregon Department of Agriculture, Salem, R. L. Westcott; OSU— Ohio State Uni-
versity, Columbus, C. A. Triplehom; OSUC— Oregon State University, Corvallis, K. A. Phillips;
POLH — University of Colorado, Polhemus collection, Englewood, J. T. Polhemus; PUL—
Purdue University, West Lafayete, IN, A. V. Provonsha; SIUC— Southern Illinois University,
Carbondale, J. E. McPherson; SMEK— Snow Museum of Entomology, University of Kansas,
Lawrence, P. D. Ashlock; TAMU— Texas A&M University, College Station, J. C. Schaffner;
UAT— University of Arizona, Tucson, F. G. Werner; UCB— Essig Museum of Entomology,
University of California, Berkeley, J. A. Powell; UCR— University of California, Riverside, S.
I. Frommer; UCS— University of Connecticut, Storrs, J. E. O’Donnell; UGA— University of
Georgia, Athens, C. L. Smith; UIM — University of Idaho, Moscow, J. B. Johnson; UMAA—
University of Michigan, Ann Arbor, B. M. O’Conner; UMC— University of Missouri, Colum-
bia, R. L. Blinn, T. R. Yonke; UNAM— Instituto de Biologia, Universidad Nacional Autonomia
de Mexico, Mexico City, DF, H. Brailovsky; UNL— University of Nebraska, Lincoln, B. C.
Ratcliffe; USNM— U.S. National Museum ofNatural History, Washington, DC, R. C. Froesch-
ner, T. J. Henry; UUSL— University of Utah, Salt Lake City, J. Loye; VPI — Virginia Polytechnic
Institute and State University, Blacksburg, M. Kosztarab; WSU— Washington State University,
Pullman, R. S. Zack.

We would like to give special thanks to those who arranged the loan of pertinent type material:
N. M. Anderson, Universetetets Zoologiske Museum, Copenhagen, Denmark ( maculata , per-
ditor); P. H. Amaud, Jr., California Academy of Sciences, San Francisco ( serratulata , spectabilis);
W. R. Dolling, British Museum (Natural History), London ( obsoleta , test ace a); R. C. Froeschner
and T. J. Henry, National Museum ofNatural History, Washington, D.C. ( cubensis ); A. Kal-
tenbach, Naturhistorisches Museum, Vienna, Austria (casta); P. Lindskog, Naturhistoriska
Riksmuseet, Stockholm, Sweden (pallidovirens ); A. V. Provonsha, Purdue University, West
Lafayette, Indiana (pseudocasta ); R. T. Schuh, American Museum of Natural History, New
York (planifrons ). The holotypes for accerra (USNM), setosa (AMNH), and spinosa (AMNH)
were examined while the senior author visited the respective museums.

We also thank D. J. Schotzko and F. Merickel, University of Idaho, who generously sent
specimens of T. pallidovirens preserved in Camoy’s solution for some of the chromosome work
discussed in this paper. The chromosome study was also assisted by a Sigma Xi Grant-in-Aid
of Research and a Theodore Roosevelt Fund Field Research Grant.

We would also like to thank J. A. Moore, L. H. Rolston (Louisiana State University), and
D. B. Thomas, Jr. (USDA-ARS, Tuxtla, MX) for their critical reviews of the manuscript. We
are especially grateful to L. H. Rolston who provided important suggestions and encouragement
throughout this entire project.

This publication was approved by the Director of the Louisiana Agricultural Experiment
Station as manuscript number 90-17-4416.

LITERATURE CITED

Alayo, D. P. 1967. Catalogo de la fauna de Cuba— XVIII — Los Hemipteros de Cuba— II
Familia Pentatomidae. Mus. “Felipe Poey” Acad. Cienc. Cuba Trab. 43:1-47, 9 pis.
Banks, N. 1910. Catalogue of the Nearctic Hemiptera-Heteroptera. Am. Entomol. Soc., Phil-
adelphia, viii + 103 pp.

Barber, H. G. 1906. Hemiptera from southwestern Texas. Brooklyn Inst. Arts Sci. Mus., Sci.
Bull. 1 (9):25 5— 289.

Barber, H. G. 1911. The resurrection of Thyanta calceata Say from synonymy. J. New York
Entomol. Soc. 19(2): 108-1 11,2 figs.

Barber, H. G. 1914. Insects of Florida. II. Hemiptera. Bull. Am. Mus. Nat. Hist. 33:495-535.
Barber, H. G. 1923. A preliminary report on the Hemiptera-Heteroptera of Porto Rico col-
lected by the American Museum ofNatural History. Am. Museum Novitates 75:1-13.

1992

THY ANT A OF NORTH AND CENTRAL AMERICA

95

Barber, H. G. 1939. Scientific survey of Porto Rico and the Virgin Islands. Vol. XIV, Part
3. Insects of Porto Rico and the Virgin Islands— Hemiptera-Heteroptera (excepting the
Miridae and Corixidae). New York Acad. Sci. 1 4:263 — 44 1 .

Barber, H. G. and S. C. Bruner. 1932. The Cydnidae and Pentatomidae of Cuba. J. Dept.
Agric., Univ. Porto Rico 1 6(3):23 1—284, pis. 24-26.

Berg, C. 1878. Hemiptera Argentina: ensayo de una monographia de los Hemipteros Het-
eropteros y Homopteros de la Republica Argentina. An. Soc. Cient. Argent. 6( 1 ):23— 26.
[Reprinted as Berg, C. 1879. Hemiptera Argentina enumeravit speciesque novas des-
cripsit. Bonariae (and Hamburg), viii + 9-62 pp.]

Berg, C. 1884. Addenda et emendanda ad Hemiptera Argentina. An. Soc. Cient. Argent. 17(3):
97-118. [Reprinted as Berg, C. 1884. Addenda et emendanda ad Hemiptera Argentina.
Bonariae (and Hamburg), 2 1 3 pp.]

Blatchley, W. S. 1895. Notes on the winter insect fauna of Vigo County, Indiana— II. Psyche
7:267-270.

Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America with Special
Reference to the Faunas of Indiana and Florida. Nature Publ. Co., Indianapolis, 1,116 pp.

Blatchley, W. S. 1930. Blatchleyana. Nature Publ. Co., Indianapolis, 77 pp.

Bruner, S. C. and H. G. Barber. 1949. List of the Pentatomidae of Cuba with the description
of a new species (Hemiptera-Heteroptera). Mem. Soc. Cubana Hist. Nat. 19(2): 155-165.

Dallas, W. S. 1851. List of the specimens of hemipterous insects in the collection of the British
Museum. London. Part 1:1-368.

Deay, H. O. and G. E. Gould. 1935. Hemiptera unrecorded from Indiana. Proc. Indiana
Acad. Sci. 45:305-309.

Distant, W. L. 1880. Insecta. Rhynchota, Hemiptera-Heteroptera. In: F. D. Godman and O.
Salvin (eds.), Biologia Centrali-Americana, Vol. 1. London, 1893:329-462.

Distant, W. L. 1893. Insecta. Rhynchota, Hemiptera-Heteroptera. In: F. D. Godman and O.
Salvin (eds.), Biologia Centrali-Americana, Vol. 1. London, 1893:329-462.

Distant, W. L. 1900. Mr. W. L. Distant on Pentatominae. Ann. Mag. Nat. Hist. 5(7):386-
397, 420-435.

Fabricius, J. C. 1775. Systema entomologiae sistens insectorum classes, ordines, genera, spe-
cies, adjectis synonymis, locis, descriptionibus, observationibus. Flensburgi et Lipsiae,
xxxii + 832 pp.

Fabricius, J. C. 1794. Entomologia systematica emendata et aucta, secundum classes, ordines,
genera, species, adjectis synonymis, locis, observationibus, Vol. IV. Hafniae, viii +
472 pp.

Fabricius, J. C. 1803. Systema Rhyngotorum secundum ordines, genera, species adjectis
synonymis, locis, observationibus, descriptionibus. Brunsvigae, x + 314 pp.

Froeschner, R. C. 1981. Heteroptera or true bugs of Ecuador: a partial catalog. Smithson.
Contrib. Zool. 322:1-147.

Froeschner, R. C. 1988. Family Pentatomidae Leach, 1815. The stink bugs. In: T. J. Henry
and R. C. Froeschner (eds.), Catalog of the Heteroptera, or True Bugs of Canada and
the Continental United States. E. J. Brill, Leiden, New York, Kobenhavn, Koln, 958 pp.

Furth, D. G. 1974. The stink bugs of Ohio (Hemiptera: Pentatomidae). Bull. Ohio Biol. Surv.
(N.S.) 5(1): 1-60.

Gillette, C. P. and C. F. Baker. 1895. A preliminary list of the Hemiptera of Colorado. Colorado
Agric. Expt. Stn. Bull. 31, Tech. Ser. no. 1, 137 pp.

Guerin-Meneville, F. E. 1857. Ordre des Hemipteres, Latr. Premiere section. Heteropteres,
Latr. In: M. R. Sagra’s Historie Physique, Politique et Naturelle de Pile de Cuba. Arthus
Bertrand, Paris. 7:359-424.

Harris, H. M. 1943. Additions to the South Dakota list of Hemiptera. J. Kansas Entomol.
Soc. 16(4): 150-1 53.

96

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Hart, C. A. 1919. The Pentatomoidea of Illinois with keys to the nearctic genera. Illinois Nat.
Hist. Surv. Bull. 13(7): 157-223.

Herrich-Schaffer, G. A. 1841. Die Wanzenartigen Insecten, Vol. 6:37-72. Niimberg.

Herrich-Schaffer, G. A. 1844. Die Wanzenartigen Insecten, Vol. 7:41-134. Niimberg.

Hoffman, R. L. 1971. The insects of Virginia: no. 4. Shield bugs (Hemiptera: Scutelleroidea:
Scutelleridae, Corimelaenidae, Cydnidae, Pentatomidae). Virgina Polytechnic Inst, and
State Univ. Res. Div. Bull. 67:1-61.

Jensen-Haarup, A. C. 1928. Hemipterological notes and descriptions V. Entomol. Medd. 16:
185-202.

Kirkaldy, G. W. 1909. Catalogue of the Hemiptera (Heteroptera) with Biological and Ana-
tomical References, Lists of Foodplants and Parasites, Etc. Vol. I: Cimicidae. Felix L.
Dames, Berlin, 392 + XL pp.

Lethierry, L. and G. Severin. 1893. Catalogue General des Hemiptera, Vol. 1, Pentatomidae.
R. Friedlander et Fils, Brussels and Berlin, ix + 286 pp.

Malloch, J. R. 1919. Subfamily Thyreocorinae. [and] Addenda. Pages 206-219 in: C. A. Hart
(ed.), The Pentatomoidea of Illinois with Keys to the Nearctic Genera. Illinois Nat. Hist.
Surv. Bull. 13:157-223, 83 figs.

McAtee, W. L. 1919. Notes on nearctic Heteroptera. Bull. Brooklyn Entomol. Soc. 14(1):
8-16.

McPherson, J. E. 1977a. Notes on the Biology of Thyanta calceata (Hemiptera: Pentatomidae)
with information on adult seasonal dimorphism. Ann. Entomol. Soc. Am. 70(3):370-
372.

McPherson, J. E. 1977b. Effects of developmental photoperiod on adult color and pubescence
in Thyanta calceata (Hemiptera: Pentatomidae) with information on ability of adults to
change color. Ann. Entomol. Soc. Am. 70(3):373-376.

McPherson, J. E. 1978a. Sensitivity of immature Thyanta calceata (Hemiptera: Pentatomidae)
to photoperiod as reflected by adult color and pubescence. Great Lakes Entomol. 11(1):
71-76.

McPherson, J. E. 1 978b. Effects of various photoperiods on color and pubescence in Thyanta
calceata (Hemiptera: Pentatomidae). Great Lakes Entomol. 1 1(3): 1 55—1 58.

McPherson, J. E. 1979a. Effects of various photoperiods on color and pubescence in Thyanta
pallidovirens accerra (Hemiptera: Pentatomidae). Great Lakes Entomol. 12(2):83-84.

McPherson, J. E. 1979b. A revised list of the Pentatomoidea of Illinois (Hemiptera). Great
Lakes Entomol. 1 2(3):9 1—98.

McPherson, J. E. 1 982. The Pentatomoidea (Hemiptera) of Northeastern North America with
Emphasis on the Fauna of Illinois. Southern Illinois Univ. Press, Carbondale and Ed-
wardsville, 240 pp.

McPherson, J. E. and S. M. Paskewitz. 1982. Effects of continuous and split developmental
photophases during each 24 hour period on adult color and pubescence in Thyanta
calceata (Hemiptera: Pentatomidae). Great Lakes Entomol. 15(2):97-98.

McPherson, J. E., T. E. Vogt and S. M. Paskewitz. 1 983. Effects of various split developmental
photophases and constant light during each 24 hour period on adult morphology in
Thyanta calceata (Hemiptera: Pentatomidae). Great Lakes Entomol. 16(2):43-46.

Osborn, H. 1894. Notes on the distribution of Hemiptera. Proc. Iowa Acad. Sci. 1 (4): 1 20—
123.

Palisot de Beauvois, A. M. F. J. 181 7. Insectes recueillis en Afrique et en Amerique, dans les
royaumes d’Oware et de Benin, a Saint-Dominique et dans les Etats-Unis, pendant les
annees. 1786-1797. Paris, Parts 9-10:137-156, 157-172.

Popenoe, E. A. 1 884. Contributions to a knowledge of the Hemiptera-fauna of Kansas. Trans.
Kansas Acad. Sci. 9:62-64.

Ride, W. D. L., C. W. Sabrosky, G. Bemardi and R. V. Melville. 1985. International Code

1992

THY ANT A OF NORTH AND CENTRAL AMERICA

97

of Zoological Nomenclature, 3rd edition. Univ. of California Press, Berkeley and Los
Angeles, 338 pp.

Rider, D. A. 1986a. A new species and new synonymy in the genus Tepa Rolston and
McDonald (Hemiptera: Pentatomidae). J. New York Entomol. Soc. 94(4):552-558.
Rider, D. A. 1986b. The identity of Euschistus rubiginosus Dallas, 1851 (Hemiptera: Pen-
tatomidae). J. Kansas Entomol. Soc. 59(2):397-398.

Rider, D. A. and J. B. Chapin. 1991. Revision of the genus Thyanta Stcil, 1862 (Heteroptera:
Pentatomidae) I. South America. J. New York Entomol. Soc. 99:1-77.

Rolston, L. H. 1 972. The small Thyanta species of North America (Hemiptera: Pentatomidae).
J. Georgia Entomol. Soc. 7(4):278-285.

Rolston, L. H. 1986. The genus Cyptocephala Berg, 1883 (Hemiptera: Pentatomidae). J. New
York Entomol. Soc. 94(3):424-433.

Rolston, L. H. and F. J. D. McDonald. 1984. A conspectus of Pentatomini of the western
hemisphere. Part 3 (Hemiptera: Pentatomidae). J. New York Entomol. Soc. 92(1):
69-86.

Ruckes, H. 1952. Two new species of Thyanta Stcil (Pentatomidae, Heteroptera). Bull. Brook-
lyn Entomol. Soc. 47(3):65-68.

Ruckes, H. 1956. Six new species of Thyanta Stcil (Heteroptera, Pentatomidae). Bull. Brooklyn
Entomol. Soc. 5 1 (3):57— 68.

Ruckes, H. 1957a. The taxonomic status and distribution of Thyanta custator (Fabricius) and
Thyanta pallidovirens (St&l) (Heteroptera, Pentatomidae). Am. Mus. Nat. Hist. Novitates
1824:1-23.

Ruckes, H. 1957b. New species of Pentatomidae from North and South America (Heteroptera)
II. Bull. Brooklyn Entomol. Soc. 52:39-47.

Ruckes, H. 1957c. Three new species of Thyanta Stcil (Heteroptera: Pentatomidae). Pan-
Pacific Entomol. 33(4): 175-1 80.

Say, T. 1831. Descriptions of New Species of Heteropterous Hemiptera of North America.
New Harmony, Indiana, 39 pp.

Stcil, C. 1859. Hemiptera species novas. Pages 219-298, pis. 3, 4 in: Kongl. Svenska Vet.-
Akad., Kongl. Avenska Fregatten Eugenies Resa Omkring Jorden Under Befel Af C. A.
Virgin, Aren 1851-53 Zoologi IV Insekter (Pt. 27). P. A. Norsted & Soner, Stockholm.
StM, C. 1862a. Bidrag till Rio Janeiro-traktens Hemipter-fauna. Kongl. Svenska Vet.-Akad.
Handl. 3(6): 1-75.

Stcil, C. 1862b. Hemiptera mexicana enumeravit speciesque novas descripsit. Stett. Entomol.
Zeit. 23:81-118.

St&l, C. 1867. Bidrag till Hemipteremas systematik. Conspectus generum Pentatomidum
Americae. Ofv. Kongl. Vet.-Akad. Forh. 24(7):522-532.

Stcil, C. 1868. Hemiptera Fabriciana. Fabricianska Hemipterater efter de i Kopenhaven och
Kiel forvarade typexemplaren granskade och beskrifne. Kongl. Svenska Vet.-Akad. Handl.
7(1 1): 1-148.

Stcil, C. 1872. Enumeratio Hemipterorum. Bidrag till en forteckning ofver alia hittils kanda
Hemiptera, jemte systematiska meddelanden. Enumeratio Cimicinorum Americae. Kongl.
Svenska Vet.-Akad. Handl. 10(4):3-65.

Summers, H. E. 1 898. A general synopsis of the nearctic Pentatomidae. Iowa Acad. Sci. Proc.
6:40-46.

Torre-Bueno, J. R. de la. 1914. Phototropism in Heteroptera. Bull. Brooklyn Entomol. Soc.
9(5):90-96.

Torre-Bueno, J. R. de la. 1939. A synopsis of the Hemiptera-Heteroptera of America north
of Mexico, part I. Entomol. Am. 19:141-304.

Ueshima, N. 1963. Chromosome study of Thyanta pallido-virens (Stfil) in relation to tax-
onomy. Pan-Pacific Entomol. 39(3): 149-1 54.

Uhler, P. R. 1872. Notices of the Hemiptera of the western territories of the United States,

98

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

chiefly from the surveys of Dr. F. V. Hayden. Pages 392-423 in: F. V. Hayden, Prelim.
Rept. U.S. Geol. Surv. Montana and Portions of Adjacent Territories, 5th Annu. Rept.
Govt. Printing Office, Washington, D.C.

Uhler, P. R. 1876. List of Hemiptera of the region west of the Mississippi River, including
those collected during the Hayden explorations of 1873. Bull. U.S. Geol. Geogr. Surv.
Territories 1 (5):267— 36 1 , pis. 19-21.

Uhler, P. R. 1877. Report upon the insects collected by P. R. Uhler during the explorations
of 1875, including monographs of the families Cydnidae and Saldae, and the Hemiptera
collected by A. S. Packard, Jr., M. D. Bull. U.S. Geol. Geogr. Surv. Territories 3(2):355-
475, 765-801.

Uhler, P. R. 1886. Check-list of the Hemiptera Heteroptera of North America. Brooklyn
Entomol. Soc., Brooklyn, 32 pp.

Uhler, P. R. 1 893. 2. A list of the Hemiptera-Heteroptera collected on the island of St. Vincent
by Mr. Herbert H. Smith; with descriptions of new genera and species. Proc. Zool. Soc.
London ( 1 893):705-7 1 9.

Uhler, P. R. 1894a. Observations upon the heteropterous Hemiptera of Lower California,
with descriptions of new species. Proc. California Acad. Sci. (2nd series) 4:223-295.
Uhler, P. R. 1894b. On the Hemiptera-Heteroptera of the island of Grenada, West Indies.
Proc. Zool. Soc. London (1894): 167-227.

Van Duzee, E. P. 1894. A list of the Hemiptera of Buffalo and vicinity. Bull. Buffalo Soc.
Nat. Sci. 5:167-204.

Van Duzee, E. P. 1904. Annotated list of the Pentatomidae recorded from America north of
Mexico, with descriptions of some new species. Trans. Am. Entomol. Soc. 30(1): 1-80.
Van Duzee, E. P. 1907. Notes on Jamaican Hemiptera: a report on a collection of Hemiptera
made on the island of Jamaica in the spring of 1906. Bull. Buffalo Soc. Nat. Sci. 8:
1-79.

Van Duzee, E. P. 1909. Observations on some Hemiptera taken in Florida in the spring of
1908. Bull. Buffalo Soc. Nat. Sci. 9:149-163.

Van Duzee, E. P. 1914. A preliminary list of the Hemiptera of San Diego County, California.
Trans. San Diego Soc. Nat. Hist. (1)11, 57 pp.

Van Duzee, E. P. 1916. Check List of the Hemiptera (excepting the Aphididae, Aleurodidae
and Coccidae) of America, North of Mexico. New York Entomol. Soc., New York,

1 1 1 pp.

Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America north of Mexico, excepting
the Aphididae, Coccidae, and Aleurodidae. Univ. of California Publ., Tech. Bull. En-
tomol. 2:1-902.

Van Duzee, E. P. 1923. Expedition of the California Academy of Sciences to the Gulf of
California in 1921: the Hemiptera (true bugs, etc.). Proc. California Acad. Sci. (4th series)
12:123-200.

Walker, F. 1867. Catalogue of the specimens of Hemiptera Heteroptera in the collection of
the British Museum, Part 2, pp. 241-417, London.

Ward, C. R., C. W. O’Brien, L. B. O’Brien, D. E. Foster and E. W. Huddleston. 1977. An-
notated checklist of New World insects associated with Prosopis (mesquite). USDA Tech.
Bull. No. 1557, Washington, D.C., 115 pp.

Westwood, J. O. 1837. In F. W. Hope, A Catalogue of Hemiptera in the Collection of the
Rev. F. W. Hope, M.A. with Short Latin Descriptions of the New Species, Part 1. F. W.
Hope, London, 46 pp.

Zimmer, J. T. 1912. The Pentatomidae of Nebraska. Univ. Nebraska Contr. Dep. Entomol.
11(3):2 19-251. (1911).

Zimsen, E. 1964. The Type Material of I. C. Fabricius. Munksgaard, Copenhagen, 656 pp.
Received 27 September 1990; accepted 23 January 1991.

J. New York Entomol. Soc. 1 00( 1 ):99— 1 36, 1992

REVISION OF AROCERA SPINOLA, WITH THE
DESCRIPTION OF TWO NEW SPECIES
(HETEROPTERA: PENTATOMIDAE)1

D. A. Rider1

Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University Agricultural Center, Baton Rouge, Louisiana 70803;

1 present address, Department of Entomology,

North Dakota State University, Fargo, North Dakota 58105

Abstract.— Arocer a Spinola contains two subgenera, the nominate subgenus and Euopta St&l.
Diagnoses are provided for Arocera, Euopta, and all previously described species. Two new
species are described: A. {Euopta) variegata from Argentina and A. {Euopta) sexpunctata from
Ecuador. Arocera apta (Walker) and A. melanopyga (St&l) are resurrected from the synonymy
of A. spectabilis (Drury), A. placens (Walker) is resurrected from the synonymy of A. nigrorubra
(Dallas), A. aequinoxialis (Westwood) is resurrected from the synonymy of A. acroleuca (Perty),
and A. repleta is resurrected from the synonymy of A. elongata and placed as a subspecies of
that species. Arocera affinis Distant, 1880, A. protea Distant, 1880, A. altivola Distant, 1893,
A. chiriquensis Distant, 1893, and A. contralineata Piran, 1955 are all removed from the
synonymy of A nigrorubra (Dallas, 1851) and are placed as new synonyms of A. placens (Walker,
1 867). Arocera patibulata Distant, 1 880 and A. jalapensis Distant, 1 893 are both removed from
the synonymy of A. spectabilis (Drury, 1782) and are also placed as new synonyms of A. placens.
Arocera crucigera Haglund, 1868 and A. schumanni Distant, 1890 are both removed from the
synonymy of A. spectabilis ; A. crucigera is placed as a new synonym of A. apta (Walker, 1867),
and A. schumanni is placed as a new synonym of A. melanopyga (St<U, 1858). Lectotypes are
designated for A. affinis, A. schumanni, Pentatoma aequinoxialis, Strachia apta, and S. nigro-
rubra. A key is provided to aid in the identification of species of Arocera.

The genus Arocera Spinola sensu lato was revised by McDonald in 1984, but the
discovery of two undescribed species and several taxonomic problems have neces-
sitated a new revision of the genus. Rider (1991) divided Arocera into two genera
based primarily on differences in coloration, punctation, and the male and female
genitalia. He transferred seven species of Arocera [colombiana McDonald, inmaculata
(Piran), prinicipalis (Stal), rufolimbata Stal, rufonotata Stal, splendens (Blanchard),
and verdana McDonald] into his newly described genus Rhyssocephala. He provided
diagnoses for all known species and described three new species.

Arocera belongs in the nominate tribe and subfamily of the Pentatomidae and is
characterized by the lack of a spine or tubercle on the third (second visible) abdominal
stemite and by elongate ostiolar rugae, reaching more than half the distance from
the ostioles to the lateral metapleural margins. Rolston and McDonald (1984) pro-
vided a key to related western hemisphere genera occurring north of South America,
and Rolston (1987) provided a key to the South American genera that have both
elongate ostiolar rugae and the base of the abdomen unarmed. The species of Arocera
are some of the most brightly colored species of New World Pentatomidae. Rider
(1991) provided a key to the pentatomine genera, which have at least some brightly
colored species.

Measurements are in millimeters; measurements in parentheses are of the holotype.

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Measurements of the body were taken with the anterior and posterior margins of the
scutellum in approximately the same plane of focus. Total length was measured
slightly differently between sexes. In females, total length was simply the distance
from the apex of the head to the posterior apex of the body excluding hemelytral
membranes. Because in males the pygophore may be distended by varying amounts,
total length was measured from the apex of the head to the posterior-most part of
the last connexival segment. Total width was measured across the humeral angles,
although in some cases the width across the abdomen was greater. Head length was
measured from the apex of the head to an imaginary line drawn though the posterior
margins of the ocelli. Head width was measured across the eyes. When label data
are presented in the text, each letter in parentheses represents a separate label with
(a) being closest to the specimen. Type label data is cited as written and placed within
quotation marks. Determination labels and collection labels were sometimes omitted
from the label data citations for brevity. Acronyms are defined in the acknowledg-
ments.

Arocera Spinola, 1837

Arocera Spinola, 1837:316-318; Stal, 1861:139; Stal, 1867:529; Stal, 1872:37; Dis-
tant, 1880:72-73; Lethierry and Severin, 1893:158; Kirkaldy, 1909:109; Froesch-
ner, 1981:68; Rolston and McDonald, 1984:74, 75; McDonald, 1984:97-99; Rol-
ston, 1987:63, fig. 5.

Ooedosoma Amyot and Serville, 1843:128 [synonymized by Stal, 1872].

Estphoria Gistel, 1848:VIII (replacement name for Ooedosoma) [synonymized by
Kirkaldy, 1909].

Type species. Arocera : Arocera aurantiaca Spinola, 1837 ( =Pentatoma acroleucum
Perty, 1833), by monotypy. Ooedosoma : Pentatoma acroleucum Perty, 1833, by
monotypy.

Diagnosis. Dorsal coloration usually yellow, orange, or red, with black markings
(specimens of A elongata elongata are black with orange markings); dorsal punctation
minute, sparse, especially on pronotum.

Vertex and jugal surfaces glabrous with at most a few weak wrinkles on juga; lateral
jugal margins slightly reflexed, becoming inflated near apices; juga and tylus subequal
in length or juga slightly longer than tylus. First antennal segment reaching nearly to
apex of head. Rostrum reaching hind coxae or beyond; first rostral segment reaching
slightly beyond posterior terminations of bucculae; bucculae not lobed posteriorly.

Anterolateral pronotal margins reflexed; humeral angles rounded. Ostiolar rugae
elongate, curving slightly cephalad, each extending about two-thirds distance from
mesial margin of ostiole to lateral metapleural margin. Third (second visible) ab-
dominal segment unarmed. Tarsi three-segmented.

Posteroventral surface of pygophore forming obtuse, semicircular carina; surface
of pygophore between carina and inferior ridge lacking black spicules, but often with
irregular wrinkles or carinae (Fig. 40); dorsolateral angles of inferior ridge not pro-
duced into spiculate horns. Lateral wall of genital cup with large, minutely striated,
sclerotized tubercle (Fig. 4 1 , set); superior ridge prominent, usually expanding over
posterolateral angles of proctiger. Each paramere with basal process either obtuse
(Fig. 29, tub) or large and spatulate (Fig. 49, bsp). Posterior margin of proctiger entire,

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REVISION OF AROCERA

101

not produced caudally (Fig. 13). Ninth paratergites flat to slightly concave (Fig. 19).
Spermathecal bulb either constricted near base (Fig. 4) or armed with long digitiform
process (Fig. 6), not simply globose; dilation of spermatheca entirely membranous,
lacking slightly sclerotized area proximally (Fig. 1 ).

Comments. Stal (1872) divided the genus Arocera into two subgenera based pri-
marily on the shape of the head and the length of the first antennal segment. He
included only one species, A. acroleuca, in the nominate subgenus, but he considered
A. acroleuca and A. aequinoxialis to be synonyms. They are both valid species and
can be separated from the remaining species by the distinctive dorsal coloration, the
shape of the head, and the form of the female genital plates. The remaining eight
species known to Stal were placed in the subgenus Euopta Stal, but three of these,
A. rufolimbata, A. rufonotata, and A. splendens, Rider (199 1) transferred to the genus
Rhyssocephala. McDonald (1984) described A. colombiana and A. verdana, both of
which were also transferred to Rhyssocephala. Piran (1963) described Pharypia in-
maculata which Grazia (1986) transferred to Arocera. Rider (1991) then transferred
A. inmaculata to Rhyssocephala. Three valid species and one subspecies have been
added to Arocera since the work of Stal. Walker (1867) described Strachia apta and
S. placens which were both later transferred to Arocera. Showalter (1929) inadver-
tantly “described” A. elongata when he published a picture of the species with a
manuscript name of Uhler’s in an issue of National Geographic. In 1929, this was
sufficient for a valid species description. Finally, Van Duzee (1931) described A.
repleta which is here considered a subspecies of A. elongata.

One species has been placed in Arocera and was subsequently transferred to another
genus. Walker (1867) described Strachia nigropicta from at least two specimens from
Brazil. Distant (1900) transferred this species to Arocera. Later, Rolston (1976) trans-
ferred it to the genus Pharypia, placing it as a junior synonym of P. generosa Stal.
Rolston also made lectotype and paralectotype designations.

At present eleven species and one subspecies are recognized in Arocera.

KEY TO THE SUBGENERA AND SPECIES OF AROCERA SPINOLA

1. Dorsal surface pale yellow with large, U-shaped black marking (Figs. 25, 32);
dorsal surface of head transversely concave; ninth paratergites slender, width less
than half the length (Fig. 21) subgenus Arocera Spinola 2

- Dorsal surface yellow, orange, or red with dark brown or black macules, but not

forming U-shaped marking; dorsal surface of head flat to slightly concave; ninth
paratergites relatively broad, width greater than half the length (Figs. 19, 20)
subgenus Euopta St&l 3

2(1). Femora fuscous to black except, at most basal one-fourth yellow with margin
between yellow and black areas gradual, diffuse; lateral margins of black, U-shaped
marking on dorsum usually entire, extending laterad over distal end of R + M
vein (Fig. 25) (southern and eastern South America) acroleuca (Perty)

- Femora fuscous to black on distal one-half, basal one-half yellow with margin

between yellow and black areas sharp, well-defined; lateral margins of black,
U-shaped, dorsal marking usually emarginate near apex of corium, usually not
extending laterad over distal end of R + M vein (Fig. 32) (northern and western
South America) aequinoxialis (Westwood)

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

3(1). Antennal segment I fuscous or black, concolorous with rest of antennal segments

4

- Antennal segment I yellow, orange, or red, distinctly lighter in color than rest of

antennal segments 6

4(3). Posterolateral angles of pygophore armed with 1-3 minute, medially directed teeth

(Fig. 40); each paramere in ectal view uniformly elongate, apex curving only slightly
laterad (Fig. 43) (Colombia) nigrorubra (Dallas)

- Posterolateral angles of pygophore unarmed (Figs. 46, 55); each paramere in ectal

view sinuously elongate, apex curving distinctly laterad (Figs. 5 1 , 60) 5

5(4). Posteroventral surface of pygophore with relatively deep, semicircular depression
and sharply carinate ventral margin; in ectal view, lateral margin of each paramere
distinctly sinuous, subapical notch well-defined (Fig. 50), (Bolivia and northern
Argentina) variegata new species

- Posteroventral surface of pygophore with relatively shallow, semicircular depres-

sion, and obtusely carinate ventral margin; in ectal view, lateral margin of each
paramere slightly sinuous, subapical notch not distinct (Fig. 59) (Mexico and
Jamaica south to Argentina) placens (Walker)

6(3). Rostrum reaching onto posterior half of fifth (fourth visible) abdominal segment,
often reaching onto sixth abdominal segment; dorsal surface bright orange with
hemelytral membranes and ocellar areas of head black (Figs. 61, 69), or black

with most of head, apex of scutellum, and two spots on each corium orange (Fig.

62) 7

Rostrum at most reaching on to anterior half of fifth (fourth visible) abdominal
segment; dorsal surface yellow or orange with brown or black macules 9

7(6). Dorsal surface mostly black with six orange spots, one on head, one on basal third
of each corium, one on posterolateral angle of each corium, and one on apex of
scutellum (Fig. 62), spot on apex of scutellum sometimes continued cephalad as
thin orange line (northwestern South America) elongata elongata Showalter

- Dorsal surface mostly orange, sometimes with a few small black markings 8

8(7). Dorsal surface usually completely orange except for black hemelytral membranes

and black spot around each ocellus (Fig. 61), sometimes with other black macules,
but never in pattern described below; each paramere in ectal view not curving
laterad apically (Fig. 68) (Panama and Costa Rica) elongata repleta Van Duzee

- Dorsal surface orange except for base of head, hemelytral membranes and six

small spots black: one on each basal angle of scutellum, one on each side of
midline of scutellum about one-half distance from base to apex, and one on each
side of midline of pronotum (Fig. 69); each paramere in ectal view curving dis-
tinctly laterad apically (Fig. 75) (Ecuador) sexpunctata new species

9(6). Ocelli very small, distance from each ocellus to adjacent eye at least 3-4 times
diameter of ocellus; dorsal margin of each paramere in lateral and medial views
concave (Figs. 80, 81) (northwestern South America) rufifrons (Dallas)

- Ocelli of normal size, distance from each ocellus to adjacent eye no more than 1-

2 times diameter of ocellus; dorsal margin of each paramere in lateral and medial
views distinctly sinuous (Figs. 87, 88, 94, 95, 101, 102) 10

10(9). In lateral and medial views dorsal margin of each paramere with basal emargi-
nation very shallow (Figs. 87, 88) (Mexico and northern Central America) ....
melanopyga (Stcil)

- In lateral and medial views, dorsal margin of each paramere with basal emargi-

nation relatively deep (Figs. 94, 95, 101, 102) (Costa Rica, Panama, and South
America) 11

11(10). Humeral angles black (Fig. 90); connexiva alternately fuscous and pale yellow

1992

REVISION OF AROCERA

103

(Fig. 90); usually with black spots on abdominal venter (southern and eastern

Brazil) spectabilis (Drury)

- Humeral angles yellow (Fig. 97); connexiva uniformly pale yellow (Fig. 97); ab-
dominal venter uniformly pale yellow except for some black markings on basal
plates (northern and western South America) apt a (Walker)

Subgenus Arocera Spinola, 1837

Diagnosis. Dorsal coloration pale yellow with large, U-shaped, black marking,
open end on base of pronotum, closed end on hemelytral membranes (Figs. 25, 32).
Dorsal surface of head distinctly, transversely concave. Posterolateral angles of py-
gophore prominent in lateral view, apices ventral to middle of pygophore (Figs. 28,
35). Each paramere with obtuse basal process, lacking large spatulate process (Figs.
29, 36). Ninth paratergites elongate, slender, width less than half the length (Fig. 21);
spiracles on eighth paratergites exposed, clearly visible. Spermatheca with sclerotized
rod swollen near proximal ends (Figs. 1 , 3), spermathecal bulb constricted near base,
with or without digitiform process (Figs. 2, 4).

Comments. The difference between Arocera and Euopta are distinct and involve
both the male and female genitalia, and the two subgenera could be elevated to
genera. They do, however, have a number of characters in common, and so I have
chosen to recognize them as subgenera. The subgenus Arocera contains only two very
closely related species, A. acroleuca and A. aequinoxialis.

Arocera ( Arocera ) acroleuca (Perty, 1833)

Figs. 1, 2, 25-31, Map 1

Pentatoma acroleucum Perty, 1833:168, pi. 33, fig. 11.

Cimex acroleucus: Burmeister, 1835:366.

Arocera aurantiaca Spinola, 1837:318; Herrich-Schaffer, 1844:90 [synonymized by
Signoret, 1851].

Ooedosoma acroleucum: Amyot and Serville, 1843:128.

Strachia acroleuca: Dallas, 1851:263; Signoret, 1851:338; Stal, 1860:22; Walker,
1867:318.

Arocera acroleuca: Stal, 1861:140; Distant, 1880:73 (part); Lethierry and Severin,
1893:158-159 (part); Piran, 1966:86; McDonald, 1984:102-103, figs. 9-16 (part).
Arocera ( Arocera ) acroleuca : Stal, 1872:38 (part); Kirkaldy, 1909:109 (part).

Diagnosis. Dorsal surface yellow with large, U-shaped, black marking; lateral mar-
gins of U-shaped mark usually entire and extending laterad across distal end of R +
M vein (Fig. 25). Dorsal surface of head black with a few transverse wrinkles; antennae
black. Anterolateral pronotal margins slightly convex (Fig. 25), slightly re flexed.
Ventral surface yellow except most of head black; rostrum usually extending onto
third (second visible) abdominal segment, yellow except segment IV black. Legs
fuscous to black except coxae and trochanters yellow, each femur gradually becoming
yellow near base. Extreme distal apices of juga often pale brown, this color sometimes
continuing along reflexion of juga.

Posterolateral angles of pygophore continuing onto pygophoral surface as weak,
obtuse, semicircular carina; inferior ridge in caudal view distinctly sinuous (Fig. 27).
Each paramere in lateral and medial views F-shaped, with small, obtuse protuberance

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Figs. 1-18. 1, 2. Arocera (A) acroleuca. 1. Spermatheca. 2. Spermathecal pump. 3, 4. A. (A.)

aequinoxialis. 3. Spermatheca. 4. Spermathecal pump. 5, 6. A. ( Euopta ) sexpunctata. 5. Sper-
matheca. 6. Spermathecal pump. 7, 8. Rhyssocephala splendens. 7. Spermatheca. 8. Sperma-
thecal pump. 9, 10. R. infuscata. 9. Spermatheca. 10. Spermathecal pump. 11-14. A. ( E .)
spectabilis. 1 1. Spermatheca. 12. Spermathecal pump. 13. Proctiger, dorsal view. 14. Proctiger,
lateral view. 15, 16. R. macdonaldi. 15. Proctiger, dorsal view. 16. Proctiger, lateral view. 17,
18. R. ecuadoriensis. 17. Proctiger, dorsal view. 18. Proctiger, lateral view. Symbols: csb,

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105

on shaft (Figs. 29, 30); in ectal view paramere-head slightly produced both medially
and laterally, distal margin nearly straight (Fig. 31). Sclerotized rod of spermatheca
swollen and then somewhat elongate proximally (Fig. 1); spermathecal bulb con-
stricted near base, with one long digitiform process connecting to top of bulb, and
one short digitiform process connecting to side of bulb just above constriction
(Fig. 2).

Types. Perty (1833) described this species from a single S specimen. The type
specimen of Pentatoma acroleuca, which is housed in the Zoologische Museum,
Munich, was examined. The label data for the holotype specimen is (a) “Type. Perty.”
(b) “Brasilia a: acroleuca Pert” (c) “AROCERA ACROLEUCA Perty”.

Spinola (1837) described Arocera aurantiaca from Brazil. The type specimen is
not in the Spinola collection and was not located, but the description matches either
A. acroleuca or A. aequinoxialis very closely. The type locality of Brazil puts it within
the known range of A. acroleuca.

Specimens examined. 45 specimens collected from 2 October to 9 April; deposited
in AMNH, BMNH, DAR, MACN, USNM, ZMB, ZSM. BOLIVIA: Santa Cruz:
Tchito, Buena Vista. BRAZIL: Bahia: Encruzilhada; Nova Conquista. Goias: Ana-
polis; Chapada dos Veadeiros, 18-24 km N Alta Paraiso; Jatai. Mato Grosso: Campo
Grande; Nivac. Minas Geraes: Barroso; Diamantina; Faz. dos Campos; Pedra Azul;
Sete Lagoas; Visosa. Rio de Janeiro: Rio de Janeiro. Sao Paulo: Campinas; Cipo;
Sao Paulo.

Distribution. Eastern and central South America (Map 1).

Comments. This species is nearly identical in coloration with A aequinoxialis, and
the two were considered conspecific until now. There are distinct differences in both
the male and female genitalia as well as a couple of fairly consistent somatic characters.
Arocera acroleuca has the femora fuscous to black except for at most the basal one-
fourth yellow, and the transition from fuscous to yellow is gradual. Arocera aequi-
noxialis has the basal one-half of the femora yellow and the transition from fuscous
to yellow is sharp. Both of these species are almost identical in coloration to Pharypia
gracilirostris Stal but may be separated from that species by the unarmed abdominal
venter and the shorter first rostral segment.

Arocera ( Arocera ) aequinoxialis (Westwood, 1837)

Figs. 3, 4, 21, 32-38, Map 1

Pentatoma aequinoxialis Westwood, 1837:33.

Arocera acroleuca (of authors, not Perty): Distant, 1880:73 (part); Lethierry and
Severin, 1893:158-159 (part); McDonald, 1984:102-103, figs. 9-16 (part).
Arocera (Arocera) acroleuca (of authors, not Perty): Stal, 1872:38 (part); Kirkaldy,
1909:109 (part).

constriction of spermathecal bulb; dfl, distal flange; dil, dilation of spermatheca; dp, digitiform
process; pesr, proximal end of sclerotized rod; pfl, proximal flange; ppp, posterior projection
of proctiger; sdil, sclerotized portion of dilation of spermathecal duct; spd, spermathecal duct;
sr, sclerotized rod; spb, spermathecal bulb.

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Figs. 19-24. Genital plates. 19. Arocera ( Euopta ) sexpunctata. 20. A. ( E .) variegata. 21. A.
{A.) aequinoxialis. 22. Rhyssocephala macdonaldi. 23. R. ecuadoriensis. 24. R. infuscata. Sym-
bols: bp, basal plate; gx2, gonacoxae 2; sp, spiracle; slO, stemite 10; 8pt, eighth paratergite; 9
pt, ninth paratergite.

Diagnosis. Coloration essentially the same as A. acroleuca except for the following
differences: basal half of femora yellow, transition from fuscous to yellow sharp;
lateral margins of dorsal, U-shaped, black mark usually indented medially just an-
terior to hemelytral membranes, black extending laterally sometimes to distal end
of each R + M vein, but usually not extending across R + M vein (Fig. 32); apices
and reflexion of juga always black.

Posterolateral angles of pygophore continuing onto pygophoral surface as weak,
obtuse, semicircular carina; inferior ridge in caudal view distinctly sinuous medially
(Fig. 34). Each paramere in lateral and medial views F-shaped, with obtuse protu-
berance on shaft (Figs. 36, 37); in ectal view paramere-head distinctly produced both
medially and laterally, apical margin distinctly concave (Fig. 38). Spermatheca with
apex of sclerotized rod swollen, but not elongate proximally (Fig. 3); spermathecal
bulb globose with basal constriction, lacking digitiform processes (Fig. 4).

Types. Westwood (1837) described Pentatoma aequinoxialis without designating
a holotype or paratypes. One <5 syntype and 1 2 syntype were located. The 6 specimen
labeled (a) “TYPE.= =WESTW. (HOPE) C. Hemipt. 1837 Part I, page 33 Distant,
P. Z. S., 1900, p. 807-825.” (b) “Type” (c) “America aq.” (d) “aquinoxialis Hope”

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Figs. 25-38. 25-31. Arocera (A.) acroleuca. 25. Habitus. 26-28. Pygophore. 26. Ventral

view. 27. Caudal view. 28. Lateral view. 29-31. Right paramere. 29. Lateral view. 30. Medial
view. 31. Ectal view. 32-38. A. (A.) aequinoxialis. 32. Habitus. 33-35. Pygophore. 33. Ventral
view. 34. Caudal view. 35. Lateral view. 36-38. Right paramere. 36. Lateral view. 37. Medial
view. 38. Ectal view. Symbols: emg, lateral emargination in dorsal U-shaped marking; ir, inferior
ridge; par, paramere; pla, posterolateral angle of pygophore; prt, proctiger; rm, R + M vein;
tub, tubercle near base of paramere; vm, ventral margin of pygophore.

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Map 1. Arocera {A.) acroleuca (•); A. (A.) aequitloxialis (O).

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(e) “Pentat. marginalis H Sch: Hahn. 320” (f) “Arocera acroleuca Perty” (g) “TYPE
HEM: 167 Vi PENTATOMA AEQUINOXIALIS WESTWOOD HOPE DEPT. OX-
FORD” is designated the lectotype. The 9 specimen labeled (a) “Rio” (b) same as
(g) for lectotype, is designated paralectotype. Both specimens were examined and are
housed in the Oxford Entomological Museum, England.

Specimens examined. 86 specimens collected during every month of the year except
February; deposited in AMNH, BMNH, CAS, DAR, DBT, HCO, LHR, MACN,
USNM, ZMB. GUATEMALA: Morales. COSTA RICA: Guanacaste: Guanacaste.
PANAMA: Darien: Santa Fe. COLOMBIA: Amazonas: Leticia. Antioquia: Valley of
Medellin. Cauca: Cauca Valle. GUYANA: East Demerara-West Coast Berbice:
Rockstone, Essequebok R. SURINAM: Marowijne: Nassau Mts, Marowijne R.
FRENCH GUIANA: Cayenne: Cayenne; Crique Boulanger; Hwy D6 to Kaw, 33.5
km SE Roura; Hwy N2 to Regina, 67 km S Cayenne. ECUADOR: Pastaza: Cuisimi,
on Rio Cuisimi, 1 50 km SE Puyo. PERU: Archinamiza. Cusco: Quince Mil. Huanuco:
Tingo Maria, Rio Huallaga. Loreto: Iquitos, Amazon R; Rio Tapiche; middle Rio
Ucayali. Madre de Dios: Rio Tabopata Reserve, 30 km SW Puerto Maldonado. San
Martin: El Boqueron Pass; Palmas del Espino Uchiza. BOLIVIA: Las Juntas; Sara.
El Beni: Reyes. La Paz: Tumupasa. Santa Cruz: Buena Vista; Saavedra Expt. Stn.;
Santa Cruz. BRAZIL: Amazonas: Tefe. Mato Grosso: Corumba Para: Santarem.

Distribution. Central America and northern and western South America (Map 1).

Comments. This species is very closely related to A. acroleuca but may be separated
from that species by differences in both the male and female genitalia and usually
by several color characters. The basal half of each femur is yellow, and the boundary
between the fuscous and yellow areas is sharp; the same boundary in A. acroleuca is
gradual, and only the basal one-fourth of the femur is yellow.

Subgenus Euopta Stal, 1872
Arocera ( Euopta ) Stal, 1872:38; Kirkaldy, 1909:109.

Type species. Cimex spectabilis Drury, 1782, by subsequent designation (Kirkaldy,
1909:XXX).

Diagnosis. Dorsal coloration yellow, orange, or red, with black markings, but not
forming distinct U-shaped marking. Dorsal surface of head flat to only slightly con-
cave. Pygophore with posterolateral angles prominent in lateral view, apices dorsal
to middle of pygophore (Fig. 42). Each paramere C-shaped, with large, spatulate
process near base (Fig. 49). Ninth paratergites relatively broad, width greater than
half length (Figs. 19, 20); spiracles on eighth paratergites small, entirely or partially
concealed under seventh abdominal tergites. Sclerotized rod of spermatheca acu-
minate and curved proximally, not swollen (Figs. 5, 11); spermathecal bulb not
constricted basally, possessing single, long, digitiform process (Figs. 6, 12).

Comments. The above characters will easily separate the species of this subgenus
from the species of the nominate subgenus.

The species of this subgenus may be grouped into three species groups. Arocera
elongata and A sexpunctata, the largest species, have the first antennal segment pale,
the hemelytral membranes completely black, and the parameres similarly shaped.
Arocera nigrorubra, A. placens, and A. variegata, the smallest species, have the first
antennal segment black, the hemelytral membranes black with the posterior margins

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pale, and the paramere-head and basal, spatulate process nearly parallel. Arocera
apta, A. melanopyga, A. rufifrons, and A. spectabilis are medium-sized and have the
first antennal segment pale, the hemelytral membranes black with the posterior
margins pale, and the paramere-head and basal spatulate process nearly perpendic-
ular.

Arocera ( Euopta ) nigrorubra (Dallas, 1851)

Figs. 39-43, Map 2

Strachia nigrorubra Dallas, 1851:267; Walker, 1867:318.

Strachia quadripunctata Signoret, 1851:337, pi. 10, fig. 4 [synonymized by Stal, 1872].
Arocera nigrorubra: Stal, 1861:140; Lethierry and Severin, 1893:159; Rolston, 1976:

3; McDonald, 1984:105-108, figs. 26-36 (part).

Arocera ( Euopta ) nigrorubra: Stal, 1872:38; Kirkaldy, 1909:109.

Diagnosis. Dorsal surface of head entirely black, with weak, transverse striations
on bases ofjuga; apex of head rounded. Lateral jugal margins reflexed, sinuous; apices
of juga somewhat inflated. Antennal segment I black (remainder of antennae missing
in only known specimen). Dorsal surface of pronotum orange-red with anterior
(except medially) and anterolateral margins and humeral angles black; and with small,
black spot on each side of middle (Fig. 39). Scutellum black with orange-red, cru-
ciform marking, posterior part of cross not reaching apex of scutellum; small, orange-
red spot laterally on each side near inner corial angle. Basal one-third to one-half of
each corium black except orange intruding anteriorly for short distance on clavus;
small, round, black spot just mesial to apex of each R + M vein; embolium much
wider apically than basally; posterior margins of coria convex (Fig. 39). Hemelytral
membranes black with posterior margins pale (Fig. 39) Connexiva alternately black
and orange-red.

Ventral surface of head and thorax completely black; rostrum black, extending to
base of abdomen; legs black. Abdominal stemites orange-red laterally, black medially,
except last segment orange-red with medial, chordate, black spot; posterolateral angles
black, and small, black spot surrounding each spiracle.

Posteroventral surface of pygophore with semicircular impression, and weak sub-
marginal carina parallel to posterior margin of pygophore becoming prominent me-
dially; posterior margin in ventral view arcuately rounded, posterolateral angles
armed with two to three minute, medially-directed teeth (Fig. 40). Parameres uni-
formly elongate, each lacking subapical emargination, narrowly rounded apically in
ectal view, curving only slightly laterad (Fig. 43). Female unknown.

Types. Dallas (1 85 1) described Strachia nigrorubra from an indeterminate number
of specimens from Colombia without designating a holotype or paratypes. Only one
syntype specimen, a <3, was located and is here designated lectotype. It has the fol-
lowing label data: (a) “Type” (b) “Colombia” (c) “fusagasuga Rolu.” (d) “33. Strach-
ia nigrorubra.” This specimen was examined and is conserved in the British Mu-
seum (Natural History), London.

The type specimen of Strachia quadripunctata Signoret, 1851 was not examined,
but the figure in the original description is of a specimen with a nearly identical color
pattern as that of the holotype of S. nigrorubra. Until further evidence is discovered,
it seems best to leave this species as a junior synonym of A. nigrorubra.

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111

Figs. 39-51. 39-43. Arocera ( Euopta ) nigrorubra. 39. Habitus. 40-42. Pygophore. 40. Ven-

tral view. 41. Caudal view. 42. Lateral view. 43. Left paramere, ectal view. 44-51. A ( E .)
variegata. 44, 45. Habitus. 46-48. Pygophore. 46. Ventral view. 47. Caudal view. 48. Lateral
view. 49-51. Right paramere. 49. Lateral view. 50. Medial view. 51. Ectal view. Symbols: bsp,
basal spatulate process; ir, inferior ridge; par, paramere; pla, posterolateral angle of pygophore;
prt, proctiger; san subapical notch; set, sclerotized tubercle.

Oo

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= A. nigrorubra
= A. variegata
O = A. placens

Map 2. Arocera ( Euopta ) nigrorubra (#); A. ( E .) placens (O); A. ( E .) variegata (A).

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113

Distribution. Colombia (Map 2).

Specimens examined. This species is known only from the type specimen.

Comments. Although the only known specimen is distinctively marked, the mark-
ings are not unique. I have examined several specimens of A. p/acens with very
similar markings. The only reliable method to separate these two species and also
the closely related A. variegata is by the male genitalia. In both A. placens and A.
variegata, the lateral surface of each paramere in ectal view is distinctly sinuous with
the apex curving distinctly laterad. Each paramere of A. nigrorubra is more uniform
in shape with the apex curving only slightly laterad. Also, the two to three medially
directed teeth on each posterolateral angle of the pygophore of A. nigrorubra are
diagnostic.

Arocera (Euopta) variegata, new species
Figs. 20, 44-51, Map 2

Description. Broadly oval, somewhat convex above, distinctly convex below; dorsal
coloration variable, but usually yellow to orange with black markings; dorsal punc-
tuation very weak, indistinct, especially on pronotum and basal disc of scutellum.

Dorsal surface of head black, usually with apices of juga becoming brown; jugal
surfaces weakly wrinkled; lateral jugal margins sinuous, nearly parallel, narrowly but
distinctly reflexed, becoming slightly inflated distally; juga slightly longer than tylus.
Antennae entirely fuscous to black. Ocelli relatively small, distance from each ocellus
to adjacent eye at least one and one-half times diameter of ocellus.

Anterolateral pronotal margins slightly convex, distinctly reflexed; pronotal disc
yellow to orange with anterior margin and sometimes anterolateral margins black,
often forming black spot on each humeral angle, and on each side of midline of
pronotal disc (Fig. 44). Scutellum usually yellow or orange with six black macules;
a small spot in each basal angle, two larger spots medially along base, one large,
subtriangular spot near distal end of each frenum (Fig. 44), and sometimes one small
spot on apex; sometimes scutellum entirely pale, lacking black markings (Fig. 45).
Coria yellow to orange with basal one-fourth to one-third black, sometimes with pale
coloration continuing cephalad on each clavus and onto basal portions of each corium
(Fig. 44); each corium with small black spot near distal end of R + M vein, sometimes
lacking black markings except for this spot (Fig. 45); hemelytral membranes fumose
with margins transparent. Connexiva yellow or orange with posterior one-third of
each segment fuscous to black (Fig. 45).

Ventral surface of head and thoracic segments fuscous to black. Rostrum fuscous
to black, extending onto base of abdomen. Legs entirely fuscous to black, except
sometimes coxae brown. Ventral surface of abdomen yellow to orange with transverse
black spots medially, those near base widest and becoming gradually narrower on
distal segments, there forming triangular black area medially; also with black spot
in each posterolateral angle of abdominal stemites and black spot just mesad and
cephalad of each spiracle; spiracle usually just in edge of black spot.

Posterior margin of pygophore in ventral view concave, posterolateral angles un-
armed (Fig. 46); weak submarginal carina parallel to posterior margin becoming
wrinkled tumescence medially (Figs. 46, 47); posteroventral surface with relatively
deep, semicircular depression; ventral margin sharply carinate (Fig. 47). Each par-

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amere C-shaped, distal part sinuously elongate in ectal view with relatively deep
subapical notch, apex curving distinctly laterad (Fig. 51); basal, spatulate process
projecting in approximately same plane as paramere-head. Basal plates widely sep-
arated basally, converging distally, mesial margins somewhat convex, posteromesial
angles rounded, posterior margins slightly sinuous mesially (Fig. 20).

Measurements. Total length 9.78-1 1.83 (9.78); total width 5.99-7.10(5.99); medial
length of pronotum 2.05-2.37 (2.05). Medial length of scutellum 4.31-5.19 (4.31);
basal width 4.36-4.53 (4.36); width at distal end of frena 1.66-2.21 (1.66). Length
of head 1.71-1.93 (1.71); width 2.51-2.84 (2.51); intraocular width 1.44-1.60 (1.44);
intraocellar width 0.85-0.98 (0.85); ocellar diameter 0.15; distance from ocellus to
adjacent eye 0.24-0.29 (0.26). Length of segments I-V of antennae 0.40-0.55 (0.55),
0.85-1.07 (0.85), 1.18-1.25 (1.21), 1.56-1.64 (1.56), and 1.69-1.71 (1.71), respec-
tively. Length of segments II-IV of rostrum 1.44-1.62 (1.44), 1.14-1.36 (1.14), and
1.03-1.12 (1.07), respectively.

Holotype. 6 labeled “ARGENTINA, Horca Molle, nearTucuman X- 18- 1968 C.W.&
L.B. O’Brien.” Deposited in the Florida State Collection of Arthropods, Gainesville.

Paratypes. 3 9 specimens, (a) “ARGENTINA: Prov. Tucuman, Acheral, 350 m,
9. XII. 1966 leg. W. Weyrauch” (b) “ex-col Weyrauch” (9 LHR); “S. P. Cololao Tu-
cuman 1950 Leg. J. M. Amau” (9 MACN); “SUR-YUNGAS chulumani-1-48 bri-
darolli” [location in Bolivia] (9 MACN).

Distribution. Bolivia and northern Argentina (Map 2).

Comments. This species is most closely related to A. nigrorubra and A. placens,
and may be reliably separated from those species only by the male genitalia. In A.
nigrorubra each paramere in ectal view is uniformly elongate with the apex curving
only slightly laterad, and the posterolateral angles of the pygophore are armed with
several small teeth. In both A. placens and A. variegata each paramere in ectal view
is sinuously elongate with the apex curving distinctly laterad, and the posterolateral
angles of the pygophore are unarmed. Arocera placens has the ventral margin of the
pygophore obtusely carinate, whereas A. variegata has the ventral margin of the
pygophore sharply carinate.

Etymology. This species is named variegata to reflect its variable coloration.

Arocera ( Euopta ) placens (Walker, 1867)

Figs. 52-60, Maps 2, 4

Strachia placens Walker, 1867:316-317.

Arocera protea Distant, 1880:73-74, pi. 7, fig. 18; Distant, 1893:336; Lethierry and
Severin, 1893:159; Rolston, 1976:3; Froeschner, 198 1:68; Brailovsky and Barrera,
1982:237. NEW SYNONYMY.

Arocera ajfmis Distant, 1880:74, pi. 7, fig. 19; Distant, 1893:336; Lethierry and
Severin, 1893:159; Rolston, 1976:3; Brailovsky and Barrera, 1982:236-237. NEW
SYNONYMY.

Arocera patibulata Distant, 1880:74, pi. 7, fig. 17. NEW SYNONYMY.

Arocera altivola Distant, 1893:337, pi. 31, fig. 18; Lethierry and Severin, 1893:159;
Rolston, 1976:3. NEW SYNONYMY.

Arocera chiriquensis Distant, 1893:337, pi. 30, fig. 19; Lethierry and Severin, 1893:
159; Becker and Grazia-Vieira, 1971:1 1; Rolston;, 1976:3; Grazia, 1984:73. NEW

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115

Figs. 52-60. Arocera { Euopta ) placens. 52-54. Habitus. 55-57. Pygophore. 55. Ventral view.
56. Caudal view. 57. Lateral view. 58-60. Right paramere. 58. Lateral view. 59. Medial view.
60. Ectal view.

SYNONYMY.

Arocera jalapensis Distant, 1893:337, pi. 30, fig. 18; Lethierry and Severin, 1893:
159; Rolston, 1976:3. NEW SYNONYMY.

Arocera placens: Distant, 1900:391; Rolston, 1976:3.

Arocera ( Euopta ) affinis: Kirkaldy, 1909:109.

Arocera {Euopta) altivola: Kirkaldy, 1909:109.

Arocera {Euopta) chiriquensis: Kirkaldy, 1909:109.

Arocera {Euopta) jalapensis: Kirkaldy, 1909:109.

Arocera {Euopta) patibulata: Kirkaldy, 1909:109.

Arocera {Euopta) placens: Kirkaldy, 1909:1 10.

Arocera {Euopta) proteus: Kirkaldy, 1909:1 10.

Arocera contralineata Piran, 1955:82; Becker and Grazia-Vieira, 1971:11; Grazia,
1984:73. NEW SYNONYMY.

Arocera spectabilis (of author, not Drury): McDonald, 1984:100 (part).

Arocera nigrorubra (of author, not Dallas): McDonald, 1984:105-106 (part).

Diagnosis. Dorsal coloration extremely variable, usually yellow or orange with

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= A. spectabilis
O — A. apta
= A. rufifrons

Map 3. Arocera ( Euopta ) apta (O); A. (E.) rufifrons (▲); A. (E.) spectabilis (#).

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117

brown or black macules, sometimes mostly black with yellow to red markings. Dorsal
surface of head entirely black or variously marked with yellow or orange; lateral jugal
margins slightly reflexed, nearly parallel, inflated apically and converging slightly
over apex of tylus; jugal surfaces weakly wrinkled near base. Ocelli relatively large,
each separated from adjacent eye by about its own diameter. Antennal segment I
black. Ventral coloration also extremely variable. Rostral segments II-IV fuscous to
black, apex extending onto base of abdomen.

Posterior margin of pygophore concave in ventral view; posterolateral angles un-
armed; postero ventral surface with relatively shallow semicircular depression, ventral
margin obtusely carinate (Fig. 56); weak submarginal carina parallel with posterior
pygophoral margin. Each paramere in ectal view sinuously elongate, apex curving
distinctly laterad with relatively shallow subapical emargination (Figs. 58, 59). Basal
plates subtriangular, mesial margins nearly straight, widely separated basally, con-
verging apically to narrowly rounded posteromesial angles, posterior margins nearly
straight, transverse.

Types. Walker (1867) described Strachia placens from at least two specimens, one
of which was from Santo Domingo, Hispaniola. The type specimen should be con-
served at the British Museum (Natural History), but it was not located. The descrip-
tion, however, is adequate to fix this species.

Distant (1880, 1893) described a large number of Arocera species, including the
following six which are examples of A. placens. Arocera protea was described from
one specimen from San Geronimo, Guatemala in 1880. The holotype was not ex-
amined, but the description with accompanying illustration and location are adequate

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to fix this species. Also in 1880, he described A. affinis from at least four specimens
from Mexico, Guatemala (San Geronimo), and Nicaragua (Chontales) without des-
ignating a holotype or paratypes. Two 2; syntypes were located. The specimen labeled
(a) “affinis Dist.” (b) “S. Geronimo, Guatemala. Champion.” (c) “Distant Coll. 1911-
383.” (d) “SYNTYPE” is here designated lectotype. The other specimen is here
designated paralectotype and is labeled the same as the lectotype except that it lacks
label (a). Both of these specimens were examined. Distant described A. patibulata in
1880 from Irazu, Costa Rica. McDonald (1984) considered it a junior synonym of
A. spectabilis. However, both Distant’s description and illustration indicate that the
first antennal segment is black, and since this specimen is from Costa Rica it is
undoubtedly A. placens. The holotype was not examined. In 1893, Distant described
A. altivola from one 2 specimen from Volcan de Chiriqui, Panama; A. chiriquensis
from one 6 specimen from Volcan de Chiriqui, Panama; and A. jalapensis from one
2 specimen from Jalapa, Mexico. The holotype specimens for the latter species were
examined. All of Distant’s type material is conserved in the British Museum (Natural
History), London.

Piran (1955) described A. contralineata from 22 specimens with the holotype 6
from Paso de los Fibres, Corrientes, Argentina. The holotype was not located, but
20 paratypes were examined from the Museo Argentino de Ciencias Naturales “Ber-
nardino Rivadavia,” Buenos Aires. All 20 specimens are typical specimens of A.
placens.

Specimens examined. 194 specimens collected during every month of the year;
deposited in AMNH, BMNH, CNC, DAR, DBT, EGER, ENGL, FSCA, LHR, MACN,
OSUC, UNAM, USNM, ZMB. MEXICO: Chiapas: 30 m W San Cristobal de las
Casas. Nayarit: Tepic. Oaxaca: Oaxaca. San Luis Potosi: 24.7 m E Lauda de Ma-
ta micros. Qro; Tamazunchale; 2 m S Tamazunchale; 3 m W Xilitla. Veracruz: 25 m
S Acayucan; Cordoba; El Limon; Jalapa. GUATEMALA: San Geronimo. BELIZE:
Rio Grande. HONDURAS: Lago Yojoa. COSTA RICA: Guanacaste: La Pacifica nr.
Canas. PANAMA: Boquete; Maje Stn. Bocas del Toro: Bocas del Toro; Miramar.
Canal Zone: Cardenas Village; Coco Solo Hospital; Madden Dam; Pipeline Rd.
Chiriqui: Fortuna; N. Candelas; Volcan de Chiriqui. Panama: Cerro Azul; Cerro
Campana; Las Cumbres.

CUBA: S. Rangel; Taco Taco. JAMAICA: Hill Gardens. Clarendon: Milk River
Bath. St. Andrew: Irishtown. PUERTO RICO: dales: 7 km S Ciales. HAITI: Laxilo.
DOMINICAN REPUBLIC: Pedernales: 24 km N Cabo Rijo. ST. LUCIA. TRINI-
DAD: Arima Valley; Simla. Curepe: Santa Margarita Circular Rd.

COLOMBIA: Magdalena: Rio Magdalena. Putomayo: El Pueblito, Parque Na-
cional Tairona Sierra Nevada de S. Marta; Sta. Rosa de Sucumbios Kofan Indian
Village, Rio San Miguel. Valle del Cauca: Palmira. VENEZUELA: Distrito Federal:
Caracas. FRENCH GUIANA: Cayenne: Cayenne. ECUADOR: Oriente, Rio Bu-
rn ay acu; Tres Cerritos. Esmeraldas: Esmeraldas. Los Rios: Rio Palenque Biol. Stn.,
N of Quevedo at km 56. Napo: Coca; Shushufindi fields, 25 m N Limoncocha.
Pastaza: Cuisimi, on Rio Cuisimi, 1 50 km SE Puyo. Pichincha: Los Banos; Quevede;
1 m W Sto. Domingo de los Colorados; Rio Palenque Stn., 47 km S Sto. Domingo
de los Colorados; Tinalandia, 12 km E Santo Domingo de los Colorados. PERU:
Cusco: Macchu Picchu Pueblo. Hudnuco: Cueva de las Pavas Cyn. 8 km S Tingo
Maria; Tingo Maria. Junin: Satipo. Loreto: Iquitos, Amazon R; Explorama Lodge,

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119

50 m NE Iquitos on Amazon R; Lake Yarinacocha, 10 km NW Pucallpa; 34 km W
Pucallpa, Toumavista Rd, km 3; Rio Maranon; Yurimaguas. Madra de Dios: Puerto
Maldonado. San Martin: Moyobamba region. BRAZIL: Amazonas: Benjamin Con-
stant, Rio Javary; Manaus; 5 km N Manaus; Parintins; Tefe. Bahia: Encruzilhada.
Distrito Federal: 30 km SW Brasilia. Mato Grosso: Corumba; Sinop. Para: Amazon
R nr Obidos; Cameta. Sao Paulo: Est. Biol. Boracea, Salesopolis. PARAGUAY: Alto
Paraguay. Central: Colonia Nueva Italia. Concepcion: Horqueta, 44 km E Paraguay
R. Cordillera: San Bernardino. ARGENTINA: Catamarca. Corrientes: Paso de Los
Libres; Santo Tome. Entre Rios: Liebig. Salta: Aguas Blancas.

Distribution. Mexico and Cuba south to Argentina (Maps 2, 4).

Comments. This species is extremely variable in coloration, and some color forms
are quite similar to A. spectabilis, A. melanopyga, and A. rufifrons. Arocera placens
can be separated from these three species by the black antennal segment I. This
species is most closely related to A. nigrorubra and A. variegata from which it can
reliably be distinguished only by the male genitalia. Each paramere in ectal view is
sinuously elongate with a relatively shallow subapical notch, the apex curves distinctly
laterad, and the ventral margin of the pygophore is obtusely carinate. In A. nigrorubra,
each paramere in ectal view is uniformly elongate and the apex curves only slightly
laterad; while in A. variegata, each paramere is sinuously elongate with a relatively
deep subapical notch, the apex curving distinctly laterad, and the ventral margin of
the pygophore is sharply carinate.

Arocera ( Euopta ) elongata elongata Showalter, 1929
Figs. 62-68, Map 6

Arocera elongata Showalter, 1929:33, 40, pi. V; Van Duzee, 1937:25-27; Piran, 1962:

5, fig. 4; Becker and Grazia- Vieira, 1971:11; Sabrosky, 1972:87-88; McDonald,

1984:114, figs. 56-66 (part); Grazia, 1984:73.

Diagnosis. Head orange except triangular area behind each ocellus black; antennae
black except first segment orange. Dorsal surface black except five orange spots as
follows: one occupying basal one-third of each corium; one on posterolateral angle
of each corium; and one on apex of scute! lum (Fig. 62), sometimes continuing ceph-
alad as thin orange line. Anterolateral margins ofpronotum straight to slightly convex.

Rostrum black except segment I orange, extending to middle of fifth (fourth visible)
abdominal segment or more; legs black. Propleura completely black; ventrolateral
margins of coria orange. Ventral surface of abdomen orange except segment seven
and lateral portions of segment six black; genital segments black except some orange
markings on genital plates of female.

Posteroventral surface of pygophore forming obtuse, semicircular carina; second,
more-clearly defined submarginal carina becoming series of small wrinkles laterally;
surface between carinae depressed (Fig. 63). Inferior ridge in caudal view sinuous
(Fig. 64). Each paramere C-shaped with large, spatulate process near base of shaft
and small tubercle just dorsal to spatulate process (Figs. 66, 67), in ectal view not
curving laterad apically (Fig. 68).

Types. The lectotype specimen (designated by Sabrosky, 1972) was examined, and
is conserved in National Museum of Natural History, Washington, D.C. The lectotype

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 6 1-68. 6 1 . Arocera ( Euopta ) elongata repleta, habitus. 62-68. A. ( E .) elongata elongata.

62. Habitus. 63-65. Pygophore. 63. Ventral view. 64. Caudal view. 65. Lateral view. 66-68.
Right paramere. 66. Lateral view. 67. Medial view. 68. Ectal view.

has the following label data: (a) “Oct.” (b) “Chapada” and (c) “Type No. 52107
U.S.N.M.”

Specimens examined. 88 specimens collected during every month of the year;
deposited in AMNH, BMNH, CAS, CNC, DAR, DBT, EGER, ENGL, FSCA, LHR,
MACN, UNAM, USNM. PANAMA: San Bias: Punta Escoses. COLOMBIA: Aguae-
laro; Buena Vista. Antioquia: Jerico. Meta: Restrepo. VENEZUELA: Merida: Merida.
ECUADOR: Oriente Zasayacu. Guayas: La Libertad. Napa: Coca; Jatun Sacha Biol.

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REVISION OF AROCERA

121

Stn, 23 km E Puerto Napo; La Selva, E of Limoncocha; Limoncocha; Lumbaqui;
Rd between El Puyo and Puerto Napo; Shushufindi Fids, 25 m N Limoncocha.
Pastaza: Puyo. Zamora-Chinchipe: Cumbaratza; Zamora, 35 m ESE Loja. PERU:
Montoya. Cusco: Macchu Picchu; Pilcopata; Quince Mil; Santa Isabel, Valley of R.
Ccosnipata. Huanuco: Pozuzo; Tingo Maria. Jumn: Prov. Jauja, Satipo; Prov. Tarma,
Utcuyacu. La Libertad: Cumpang. above Uctubamba. Loreto: Middle Rio Ucayali;
Pucallpa. BOLIVIA: Lara. Cochabamba: Cochabamba; Crista! Mayu, Chapare; Pal-
mar, Prov. Chapare; Todos Santos, Prov. Chapare Santa Cruz: nr. Santa Cruz.
BRAZIL: Chapada. Amazonas: Benjamin Constant Rio Javary. Mato Grosso do Sul:
Corumba.

Distribution. Eastern Panama and northwestern South America (Map 6).
Comments. Both subspecies of A. elongata can be readily identified by their char-
acteristic coloration alone. Arocera e. elongata is black with five large orange-red
spots on the dorsum, and A. e. repleta usually has the dorsum uniformly orange-red
(except for black markings on the head and hemelytral membranes). These color

122 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Map 6. Arocera ( Euopta ) elongata elongata (O); A. ( E .) sexpunctata (•).

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REVISION OF AROCERA

123

patterns are remarkably constant and are unique within the genus. The male and
female genitalia of the two subspecies are identical, however, and since each seems
to have distinct geographical distributions, they are best considered subspecies. Ar-
ocera e. elongata occurs throughout northwestern South America and eastern Pan-
ama. Arocera e. repleta occurs in Panama and Costa Rica.

Arocera ( Euopta ) elongata repleta Van Duzee, 1931

Fig. 6 1 , Map 4

Arocera repleta Van Duzee, 1931:94.

Arocera elongata (of authors, not Showalter): McDonald, 1984: 1 14, figs. 56-66 (part).

Diagnosis. Dorsal surface bright orange except hemelytral membranes and trian-
gular area behind each ocellus black (Fig. 61). Sometimes small, variable, black spots
present on basal areas of pronotum and scutellum, and rarely on central area of each
corium (one specimen has anterior two-thirds of scutellum and most of medial area
of pronotum black). Antennae black except segment I orange. Anterolateral margins
of pronotum straight to slightly convex.

Segment I of rostrum orange, remaining segments black, rostrum extending at least
to middle of fifth (fourth visible) abdominal segment. Thoracic pleura black except
anterolateral margins of propleura and lateral margins of meso- and metapleura
orange. Legs black. Ventral surface of abdomen and genital segments as in nominate
subspecies.

Types. The holotype specimen 2 of Arocera repleta was examined. It is housed in
the California Academy of Sciences, San Francisco. The holotype is type no. 3602
and is labeled from the Panama Canal Zone.

Specimens examined. 28 specimens collected from 1 3 May to 1 3 August; deposited
in AMNH, CAS, DAR, DBT, EGER, ENGL, LHR, USNM, COSTA RICA: Pun-
tar enas: Monteverde. PANAMA: Bocas del Toro: 2 m N divide on hwy to Chiriqui
Grande; 10 km NE Fortuna Dam. Canal Zone: Barro Colorado Island; Madden
Dam; Margarita. Chiriqui: Fortuna. Code: Cerro Gaital. Colon: Sta. Rita Ridge.

Distribution. Panama and Costa Rica (Map 4).

Comments. This subspecies can be identified by its distinctive coloration. No other
species or subspecies has the dorsum uniformly orange-red as in this subspecies.

Arocera (Euopta) sexpunctata, new species
Figs. 5, 6, 19, 69-75, Map 6

Arocera elongata (of authors, not Showalter): McDonald, 1984: 1 14, figs. 56-66 (part).

Description. Dorsal surface of head orange except triangular area behind each
ocellus black. Segment I of each antenna orange, remaining segments black. Dorsal
surface of body bright orange except six small black spots as follows: one in each
basal angle of scutellum, one on each side of midline of pronotum just posterior to
cicatrices, and one on each side of midline of scutellum about one-half distance
between base and apex (Fig. 69). Anterolateral pronotal margins straight to slightly
convex. Hemelytral membranes black. Connexival segments uniformly black.

Rostrum black except segment I orange, extending to middle of fifth (fourth visible)
abdominal segment. Thoracic pleura black except anterolateral margins of propleura

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 69-75. Arocera ( Euopta ) sexpunctata. 69. Habitus. 70-72. Pygophore. 70. Ventral
view. 71. Caudal view. 72. Lateral view. 73-75. Right paramere. 73. Lateral view. 74. Medial
view. 75. Ectal view.

and lateral margins of meso- and metapleura orange; ventrolateral margins of coria
orange. Legs black. Abdomen completely black except some orange markings on
genital plates of females.

Pygophore in ventral view with inferior ridge concave, slightly emarginate medially
(Fig. 70), with submarginal semicircular depressed area wrinkled, especially laterally
(Fig. 71); in caudal view inferior ridge sinuous (Fig. 71). Each paramere C-shaped
with large, spatulate process near base of shaft and small tubercle just dorsad of
spatulate process (Figs. 73, 74); in ectal view, paramere-head distinctly curving laterad
(Fig. 75). Basal plates widely separated basally, converging distally, mesial margins
convex, posteromesial angles broadly rounded, posterior margins sinuous (Fig. 19).

Measurements. Total length 13.72-17.35 (15.30); total width 7.89-9.30 (8.67);
medial length of pronotum 2.52-3.39 (2.92). Medial length of scutellum 6.39-7.57
(6.78); basal width 5.05-5.91 (5.68); width at distal end of frena 1.89-2.37 (2.05).
Length of head 2.32-2.65 (2.54); width 3.06-3.46 (3.31); intraocular width 1.60-
1.88 (1.80); intraocellar width 0.91-1.1 1 (1.07); ocellar diameter 0.29-0.31 (0.29);
distance from ocellus to adjacent eye 0.29-0.31 (0.29). Length of segments I-V of
antennae 0.74-0.83 (0.83), 1.36-1.69 (1.55), 2.17-2.50 (2.32), 2.54-2.80 (2.70), and
2.37-2.58 (2.48), respectively. Length of segments II-IV of rostrum 2.70-3.24 (3.02),
3.02-3.68 (3.39), and 1.91-2.43 (2.13), respectively.

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REVISION OF AROCERA

125

Holotype. 6 labeled “ECUADOR Pichingha, 15 km E Sto. Domingo 1.XI.74 Col.
J. Baier at lights.” Deposited in the National Museum of Natural History, Wash-
ington, D.C.

Paratypes. 666, 1 292. Labeled as holotype ( 6 299 ENGL; 2 LHR); labeled as holotype
except “31-X-74” (2 ENGL); (a) “Tropical rainforest general collecting” (b) “EC-
UADOR, Pichincha, Puerto Quito, 750m. x-xi. 1982 G. Onore” (c) “Brit. Mus.
1982-246” (4 66 2 BMNH); “ECUADOR Santo Domingo de los Colorados, Pichincha
7 February 1958 R. W. Hodges, 2000'” (6 LHR); (a) “Ecuador Puerto Ore-llanae”
(b) “Col. R. de Lafebre” (2 UNAM); “ECUADOR: Pichincha Prov., Tinalandia
2800' elev. 1 2 km E of Santo Domingo June 28-30, 1 980 Coll. Dan Bogar” (2 EGER);
“ECUADOR: Pichincha: Santo Domingo January 19, 1968 B. Heineman” (222
AMNH); and “ECUADOR: Pichincha Sto. Domingo de los Colorados Sept. 22, 1 970
R. E. Dietz IV, leg.” (222 USNM).

Distribution. All known specimens are from the province of Pichincha, Ecuador
(Map 6).

Comments. Specimens of this species are very distinctively and consistently marked.
No other species has a bright orange dorsal surface with small black spots as described
above.

Arocera ( Euopta ) rufifrons (Dallas, 1851)

Figs. 76-82, Map 3

Strachia rufifrons Dallas, 1851:267; Walker, 1867:318.

Arocera rufifrons: Lethierry and Severin, 1893:159; Rolston, 1976:3; McDonald,

1984:108-109, figs. 37-43 (part).

Arocera {Euopta) rufifrons : Stal, 1872:38; Kirkaldy, 1909:110.

Diagnosis. Broadly oval, dorsal coloration yellow to orange with the following
black areas: basal half of head, pronotal cicatrices and area just anterior to each
cicatrice, sometimes anterolateral margins of pronotum, large basal spot on each side
of middle of pronotum, large spot on each basal angle of scutellum, lateral semicircular
spot near apex of frenum on each side of scutellum, large spot on each hemelytron
extending from scutellum to lateral margin, and hemelytral membranes except pale
apices (Fig. 76).

Apex of head broadly rounded, nearly arcuate; lateral jugal margins sinuous, re-
flexed for most of their length; jugal apices not distinctly inflated, jugal surfaces
distinctly striated. Ocelli very small, distance from each ocellus to adjacent eye at
least 3-4 times diameter of ocellus. Antennal segment I pale except extreme distal
end fuscous, remaining segments black. Embolium becoming distinctly wider api-
cally, width at distal end of R + M vein at least twice basal width (Fig. 76). Corium
usually appearing distinctly wrinkled in area of black spot. Connexiva uniformly
yellow or orange or sometimes with black spot on each posterolateral angle.

Ventral surface of head uniformly yellow or orange. Rostrum fuscous to black,
except most of segment I yellow or orange, reaching to base of fourth (third visible)
abdominal segment. Ventral surface of thorax yellow to orange with large black spot
on central portion of each pleuron, and usually medial areas black. Legs fuscous to
black except coxae pale. Ventral surface of abdomen yellow to orange, sometimes

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 76-89. 76-82. Arocera ( Euopta ) rufifrons. 76. Habitus. 77-79. Pygophore. 77. Ventral

view. 78. Caudal view. 79. Lateral view. 80-82. Right paramere. 80. Lateral view. 81. Medial
view. 82. Ectal view. 83-89. A. ( E .) melanopyga. 83. Habitus. 84-86. Pygophore. 84. Ventral
view. 85. Caudal view. 86. Lateral view. 87-89. Right paramere. 87. Lateral view. 88. Medial
view. 89. Ectal view.

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REVISION OF AROCERA

127

with medial black spots, especially on posterior segments, sometimes with a few
irregular black spots just mesial to spiracles.

Posterior margin of pygophore in ventral view concave, sharply carinate laterally,
becoming thickened and somewhat rugulose medially; posteroventral surface deeply
impressed, somewhat V-shaped ventrally; ventral margin carinate laterally becoming
more rounded medially (Fig. 77); inferior ridge in caudal view slightly concave with
broad medial area produced slightly dorsad (Fig. 78). Parameres in medial and lateral
views C-shaped, each with shaft relatively thick dorsal to basal spatulate process
(Figs. 80, 81), dorsal margin broadly and shallowly concave, apex curving gently
dorsad (Figs. 80, 81), basal spatulate process in nearly same plane as paramere-head
(Fig. 82). Basal plates widely separated basally, converging to narrowly rounded
posteromesial angles; posterior margins nearly straight with mesial areas more caudal
than lateral areas.

Types. Dallas (1851) described Strachia rufifrons from an indeterminate number
of specimens without designating a holotype or paratypes. McDonald (1984) located
29 syntypes and designated the specimen labeled 85a as lectotype, and the specimen
labeled 85b as paralectotype. Both specimens were examined and are conserved in
the British Museum (Natural History), London. The lectotype has the following label
data: (a) “Braz.” (b) “85a.”

Specimens examined. 22 specimens collected from September to April; deposited
in AMNH, BMNH, EGER, USNM, ZMB. COLOMBIA: Rio Dagus. ECUADOR:
Napo: Coca; Tena PERU: Achinamiza; Chanchamayo. Huanuco: Cueva de las Pavas
Cyn, 8 km S Tingo Maria. Loreto: Upper Rio Tapiche. BRAZIL: Guanabara, Cor-
coyado. Amazonas: Tefe.

Distribution. Colombia, Ecuador, Peru, and Brazil (Map 3).

Comments. McDonald (1984) partially misidentified this species, including with
it specimens of A. melanopyga from Mexico. Arocera rufifrons, however, is relatively
rare and is restricted to northwestern South America. It may be recognized most
easily by the very small ocelli and the male genitalia.

Arocera ( Euopta ) melanopyga (Stal, 1858)

Figs. 83-89, Map 5

Strachia melanopyga St&l, 1858:437; Walker, 1867:316.

Arocera melanopyga: St&l, 1861:140; Stal, 1862:107; Distant, 1880:73, pi. 7, fig. 1 1;

Lethierry and Severin, 1893:159; Brailovsky and Barrera, 1982:237.

Arocera {Euopta) melanopyga: Stal, 1872:38; Kirkaldy, 1909:109.

Arocera schumanni Distant, 1893:336, pi. 31, fig. 17; Lethierry and Severin, 1893:
159; Rolston, 1976:3; Brailovsky and Barrerra, 1982:238. NEW SYNONYMY.
Arocera ( Euopta ) schumanni: Kirkaldy, 1909:1 10.

Arocera spectabilis (of authors, not Drury): McDonald, 1984:1 00-102, figs. 1-8 (part).

Diagnosis. Dorsal surface of head smooth, yellow, except basal half fuscous to
black, black not extending beyond anterior margins of eyes, often reduced with
medial, narrow, longitudinal, pale line. Lateral jugal margins slightly reflexed, nearly
parallel, apices of juga slightly inflated. Each ocellus relatively large, located about

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

its own diameter from adjacent eye. Antennae black except segment I and sometimes
basal portions of segment II yellow to orange.

Dorsal surface of pronotum often with anterior margins fuscous to black, marking
sometimes extending along anterolateral margins to humeral angles. Posterior disk
of pronotum usually with large black spot on each side of midline (Fig. 83). Black
markings on pronotum may be reduced or completely absent; black spots on posterior
disk may be diffuse brown. Scutellum with black spot in each basal angle (Fig. 83).
Black or fuscous trans-corioscutellar band usually present (Fig. 83), but may be
reduced and interrupted by thin, pale, line on middle of scutellum and sometimes
on coria. Hemelytral membranes fumose except apical margins pale (Fig. 83). Con-
nexival segments yellow to orange, usually with fuscous to black spots in posterolateral
angles (Fig. 83)

Ventral surface of head yellow to orange. Rostrum usually fuscous to black except
segment I yellow to orange, sometimes orange markings also on segments II— III,
extending to anterior margin of fourth (third visible) abdominal stemite. Coloration
of thoracic pleura variable ranging from yellow or orange to fuscous to black on
extensive areas of meso- and metapleura, propleura rarely with black markings. Legs
with varying amounts of fuscous and pale areas. Abdominal stemites sometimes
uniformly pale, but usually with black spots laterally and sometimes along midline.

Posterior margin of pygophore in ventral view concave, sinuous medially (Fig. 84);
in caudal view inferior ridge sinuously V-shaped; postero ventral surface wrinkled,
with V-shaped depressed area (Fig. 85). Each paramere flat, thin; apex undulating,
appearing tri-lobed; in lateral and medial views, lobes not as distinct as in A. spec-
tabilis (Figs. 87, 88); basal spatulate process nearly perpendicular to paramere-head
(Fig. 89). The female genitalia are typical for the subgenus.

Types. Stal (1858) described Strachia me/anopyga from at least two female spec-
imens from Mexico. The type material is probably deposited in the Naturhistoriska
Riksmuseet, Stockholm, Sweden, but the specimens were not located. This species
can, however, be fixed by Stal’s original description and its type locality. Arocera
me/anopyga is the only known species occurring in Mexico that has the first antennal
segment pale.

Distant (1893) described A. schumanni from 26 and 2$ specimens without desig-
nating a holotype or paratypes. The <5 specimen labeled (a) “Tepic, Mex. July, Schu-
mann.” (b) “B. C. A., Hem. 1. Arocera schumanni” (c) “SYNTYPE” is designated
lectotype. The remaining specimens are designated paralectotypes and have the fol-
lowing label data: (a) “schumanni Dist.” (b) “Tepic, Mex. July. Schumann.” (c)
“Distant Coll. 1911-383.” (d) “SYNTYPE” (6); (a) “Tepic, Mex. July. Schumann.”
(b) “Distant Coll. 191 1-383.” (c) “SYNTYPE” ($); (a) “Tepic, Mex. July. Schumann.”
(b) “B. C. A., Hem. 1. Arocera schumanni” (c) “SYNTYPE” (2). All four specimens
were examined; in important characters of the male and female genitalia they do not
differ significantly from A. me/anopyga. The specimens are conserved in the British
Museum (Natural History), London.

Specimens examined. 50 specimens collected from 1 9 April to 9 October; deposited
in BMNH, CAS, DAR, DBT, EGER, LHR, UNAM. MEXICO: Chiapas: Aguacero;
Chicoasen Dam Area; Parque Laguna Belgica, 12 m N Ocozocuatla; 20 km NE
Tuxtla Gutierrez. Guerrero. Jalisco: Estacion de Biologia “Chamela”, 8 km S Cha-
mela. Nayarit: 8.7 m E San Bias; Tepic. Oaxaca: Palomeres. Quintana Roo: Bacalar;

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129

Nuevo, X-Can. Tamaulipas: Bocatoma, 6 m S Gomez Farias; 8 m W El Limon; 73. 1
m N Manuel. Veracruz: Los Tuxtlas; 15.8 m S Tampico. BELIZE: Orange Walk
Town. GUATEMALA: Cayuga; Escuintla Penon de Siquinala.

Distribution. This species occurs in Mexico and northern Central America (Map

5).

Comments. This species has recently been considered a junior synonym of A.
spectabilis, from which it is geographically isolated. Specimens of A. spectabilis are
consistently marked, whereas A. melanopyga is extremely variable.

Arocera melanopyga is most similar in coloration to A. rufifrons and A. apta, both
of which occur only in South America. It differs from both of these species by the
lack of black markings on the propleura and by the male genitalia. It further differs
from A. rufifrons by the large ocelli. The ocelli are very small in A. rufifrons.

The only similarly colored sympatric species is A. placens, which can be separated
by the male genitalia and the color of antennal segment I. The first antennal segment
is pale in A. melanopyga but black in A. placens.

Arocera ( Euopta ) spectabilis (Drury, 1782)

Figs. 11-14, 90-96, Map 3

Cimex spectabilis Drury, 1782: index, 66, pi. 46, fig. 4.

Strachia spectabilis: Dallas, 1851:266; Stal, 1860:22; Walker, 1867:318.

Strachia generosa St&l, 1854:233; Walker, 1867:317 [synonymized by Stal, 1872],
Arocera spectabilis: Stal, 1861:140; Lethierry and Severin, 1893:159; Grazia, 1977:

163; McDonald, 1984:100-102; figs. 1-8 (part).

Arocera ( Euopta ) spectabilis: Stal, 1872:38; Kirkaldy, 1909:1 10.

Diagnosis. Dorsal surface of head weakly wrinkled near base of juga, with apical
portion orange-red, basal portion black; black usually extending onto basal areas of
juga and tylus (Fig. 90). Juga distinctly inflated apically; lateral margins reflexed,
sinuous, tapering to truncately rounded apex. Antennae black except segment I red-
orange; ocelli relatively large, each located about its own diameter from adjacent eye.

Anterior margin of pronotum black except yellow intruding medially to anterior
reflexion; anterior black areas usually extending along anterolateral margins, then
forming distinct black spot on each humeral angle (Fig. 90). Large black spot on each
side of midline of posterior pronotal disk, extending onto base of scutellum. Broad
black band traversing coria and scutellum near middle of scutellum (Fig. 90), some-
times divided on midline of scutellum by thin yellow line (Fig. 90). Hemelvtral
membranes black, apical margins transparent (Fig. 90). Connexia red-orange with
large black spot in each posterolateral angle. Anterolateral pronotal margins nearly
straight; hemelytra lacking wrinkles. Embolium somewhat widened apically.

In ventral view, posterior margin of pygophore concave, slightly sinuous (Fig. 91);
weak submarginal carina parallel to posterior margin; posteroventral surface of py-
gophore with broad V-shaped depression, ventral margin obtusely carinate (Fig. 92).
Each paramere undulating dorsally, apex curving dorsad slightly more than in A.
apta, with large spatulate process near base protruding laterad, perpendicular to
paramere-head (Fig. 96) with small rectilinear tubercle apically (Fig. 94); subtrian-
gular, flattened, dorsal margin appearing tri-lobed in lateral and medial views (Figs.
94, 95). Basal plates widely separated basally, converging apically; mesial margins

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 90-103. 90-96. Arocera (Euopta) spectabilis. 90. Habitus. 9 1-93. Pygophore. 9 1 . Ven-
tral view. 92. Caudal view. 93. Lateral view. 94-96. Right paramere. 94. Lateral view. 95.
Medial view. 96. Ectal view. 97-103. A. ( E .) apta. 97. Habitus. 98-100. Pygophore. 98. Ventral
view. 99. Caudal view. 100. Lateral view. 101-103. Right paramere. 101. Lateral view. 102.
Medial view. 103. Ectal view.

1992

REVISION OF AROCERA

131

convex, posterior margins nearly straight, posteromesial angles rounded slightly
prominent.

Types. The type of Cimex spectabilis was not located, but this species is easily
fixed by the illustrations of Drury (1782).

I was not able to examine any of the type material of Strachia generosa, but since
Stal (1872) himself placed his species as a junior synonym of A. spectabilis I have
followed this synonymy.

Specimens examined. 72 specimens collected from 6 August to 1 2 March; deposited
in AMNH, EGER, UNAM, USNM, ZMB. BRAZIL: E. Bresslau; Guapi; Laguna
Aztezca, Holango, Hidalgo. Bahia: Encruzilhada; Rio, v. olfers, Sello. Esperito Santo:
Concei^ao da Barra. Minas Geraes: Diamantina; Pedra Azul; Vigosa. Pernambuco:
Tapera. Rio Grande do Norte: Natal. Rio de Janeiro: Petropolis; Rio de Janeiro. Rio
Grande do Sul: Pelotas; Porto Alegre. Santa Catarina: Corupa; Rio Natal. Sao Paulo:
Casa Grande, Boraceia Field Stn.

Distribution. Southern and eastern Brazil (Map 3).

Comments. This species resembles some specimens of A. placens from which it
may be separated by the pale first antennal segment; the first antennal segment is
black in A. placens. It is most closely related to A. apt a, which in fact may be a
subspecies of this species. The genitalia of the two are very similar, but the differences
in coloration are distinct and constant. Arocera spectabilis always has black markings
on the humeral angles, the connexiva, and the abdominal venter. These markings
are never present in A. apt a.

Arocera ( Euopta ) apta (Walker, 1867)

Figs. 97-103, Maps 3, 5

Strachia apta Walker, 1867:323.

Arocera crucigera Haglund, 1868:157; Lethierry and Severin, 1893:159; Froeschner,
1981:68. NEW SYNONYMY.

Arocera {Euopta) crucigera: St&l, 1872:38; Kirkaldy, 1909:109.

Arocera apta: Distant, 1880:73, pi. 7, fig. 12; Distant, 1893:336; Lethierry and Sev-
erin, 1893:159; Piran, 1963:107; Rolston, 1976:3; Becker and Grazia- Vieira, 1977:
55; Froeschner, 1981:68; Grazia, 1984:73.

Arocera ( Euopta ) apta: Kirkaldy, 1909:109.

Arocera malonopyga [sic] (of authors, not Stal): Piran, 1956:29.

Arocera melanopyga (of authors, not Stal): Piran, 1962:6.

Diagnosis. Dorsal coloration very similar to A. spectabilis, ground color yellow
with six large black macules: one on basal half of head extending onto anterior margin
of pronotum, one on each side of midline of posterior pronotal disc continuing onto
basal areas of scutellum, one on each corium near middle extending onto scutellum
near distal end of frenum, and one consisting of black hemelytral membranes (Fig.
97). Black on basal half of head extends at most only slightly onto basal areas of
juga; black markings entirely absent from humeral angles (Fig. 97). Connexiva entirely
pale yellow. Dorsal punctation minute and very sparse.

Lateral jugal margins slightly reflexed, sinuous, nearly parallel, extending slightly
beyond apex of tylus; jugal surfaces weakly wrinkled. Antennae black except segment
I and sometimes basal portion of segment II pale. Each ocellus relatively large,

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

separated from adjacent eye by about its own diameter. Ventral surface yellow except
for one black macule on middle of each pleurite. Rostrum yellow except segment IV
black, usually extending onto fourth (third visible) abdominal stemite; legs black
except basal half of each femur yellow.

Pygophore with posterior margin in ventral view concave, slightly emarginate
medially (Fig. 98), posteroventral surface with shallow, submarginal, triangular, de-
pressed area (Fig. 99); inferior ridge in caudal view sinuous (Fig. 99). Each paramere
undulating, trilobed dorsally, apex not curving dorsad as much as in A. spectabilis,
with large spatulate process near base of paramere protruding laterad (Fig. 101),
nearly perpendicular to paramere-head (Fig. 103). Female genitalia typical for sub-
genus.

Types. Walker (1867) described Strachia apta from an indeterminate number of
specimens from Brazil without designating a holotype or paratypes. Only 1 9 syntype
was located and is here designated the lectotype. It has the following label data: (a)
“62 57“ [dorsal surface], “Braz“ [ventral surface] (b) “Type” (c) “4 1 . Strachia apta.”
[dorsal surface] “nereous, with black Ve“ [ventral surface] (d) “68.” This specimen
was examined and is housed in the British Museum (Natural History), London.

The type material of Arocera crucigera Haglund should be deposited in the Na-
turhistoriska Riksmuseet, Stockholm, Sweden, but no specimens were located.
McDonald (1984) examined the holotype and considered A. crucigera to be a junior
synonym of A. spectabilis. The type locality of the Brazilian Amazon more likely
places it in the known distribution of A. apta.

Specimens examined. 173 specimens collected from every month of the year;
deposited in AMNH, BMNH, CAS, CIAT, CNC, DAR, DBT, EGER, ENGL, FSCA,
LHR, MACN, OSUC, UNAM, USNM, ZMB. COSTA RICA: Esparta; Turrialba.
Heredia: 1 km S Pt. Viejo. Puntarenas: Finca Las Cruces, 6 km S San Vito de Java;
Las Cruces, 7 km S San Vito; Palmar. PANAMA: Gamboa; Punta Vacamonte, 8°52'N
79°4Q'W. Bocas del Toro: 2 mi N Divide on hwy to Chiriqui Grande; Miramar, 9°N
82°15'W. Canal Zone: Barro Colorado Island; Coco Solo Hospital; Ft. Gulch; Ft.
Gulion. Chiriqui: Renacimiento Santa Clara; Volcan de Chiriqui. Cocle: Cerro Gaital.
Colon: Santa Rita Ridge. Darien: 23 km E Cananzas. Panama: Altos de Maje, Chepo;
Cerro Campana; ~ 10 km N El Llano; Las Cumbres.

COLOMBIA: Amazonas: Leticia. Antioquia: Medellin. Atlantico. Caqueta: Rio
Orteguaza nr Rio Peneya. Meta: Pto. Gaiton-Carimagua; Restrepo. Valle de Cauca:
Buenaventura. VENEZUELA: Rio Chaurito. Amazonas: San Carlos de Rios Negro.
Aragua: Rancho Grande. Carabobo: Valencia. Distrito Federal: Caracas. Merida: Hwy
to El Vigia. Yaracuy: Aroa. GUYANA: Essequibo R., Moraballi Crk; East Berbice-
Corentyne: Oronoque & New River heads. Mazaruni-Potaro: Bartica District, Penal
Settlement; Kartabo Point; Takutu Mtns. FRENCH GUIANA: Cayenne: Cayenne;
Hwy D6 to Kaw, 34 km SE Roura; Hwy N2 to Regina, 45 km S Cayenne; Hwy N2
to Regina, 67 km S Cayenne. St. Laurent du Maroni: St. Jean; Hwy ext. Nl, 20 km
SE St. Laurent; 13 km SSE St. Laurent. ECUADOR: Salidero. Azuay: Sta. Isabel, 65
km SW Cuenco. El Oro: Santa Rosa; W of Santa Rosa, just S of Machala. Esmeraldas:
San Mateo. Guayas: Guayaquil. Los Rios: Pichilingue Expt Stn, Quevedo; Rio Pa-
lenque. Manabi: Prov. Penchancha, 8 km N Tinalandia, Sto. Domingo de los Co-
lorados. Napo: Coca; La Selva; Limoncocha, on Rio Napo; Lumbaqui Stn, 70 km
W Lago Agrio. Pastaza: Puyo; 1 6 km W Puyo. Pichincha: Puerto Quito; Sto. Domingo

1992

REVISION OF AROCERA

133

de los Colorados; Rio Palenque Stn, 47 km S Sto. Domingo de los Colorados;
Tinalandia, 12 km E Sto Domingo de los Colorados; 15 km E Sto. Domingo de los
Colorados. PERU: Achinamiza. Amazonas: Rio Santiago. Cusco: Pilcopata; Prov.
Paucartambo, Haciendo Maria, Shores of Rio Ccosnipata. Huanuca: Tingo Maria.
Junin: Prov. Tarma, Utcuyacu. La Libertad: Cumpang. above Uctubamba. Loreto:
Callicebus Res. Stn. Mishana, Rio Nanay, 25 km SW Iquitos; Guyabamba, 3 kms
from Iquitos; Iquitos, Platanal; middle Rio Ucayali; Requena. Madre de Dios: mouth
La Torre River; Rio Tambopata Reserve, 30 km SW Puerto Maldonado. San Martin:
El Boqueron Pass; Rio Huallaga. Ucayali: Pucallpa; 34 km W Pucallpa; 3 km NE
Abra Diuisoria on Tingo-Maria-Pucallpa hwy. BOLIVIA: Sara; S. F. Chipiriri. Co-
chabamba: Chapare, Alto Palmar; Chapare, Villa Tunari. El Beni: Blancaflor; Ri-
beralta; Rurrenabaque. La Paz: Ixiamas. Santa Cruz: Buena Vista, Proc. Ichilo; Colpa
Gas P. Stn, 9 mi W Warnes; 10 mi W Portachuelo; Santa Cruz. BRAZIL: Amazonas:
Amazons between Para and Manaus; Benjamin Constant, Rio Javary; Hyutanahan,
Rio Purus; Manaus; 5 km N Manaus; 244 km Manaus; Uypiranga Rio Negro, 14
km from Manaus; Puraquequara; Sao Paulo de Olivenga; Tefe. Bahia: Rio, v. olfers,
Sello. Mato Grosso: 10°25'S, 59°28'W; Sinop. Mato Grosso do Sul: Corumba. Para:
Almeirim, Sao Raimundo; 330 m S Belem; Jacareacanga; Santarem; Urua, 65 km
SW Itaituba on BR 230. Rondonia: Porto Velho. Santa Catarina: Rio Vermelho.
ARGENTINA: Salta: San Martin Palitos.

Distribution. Costa Rica to Brazil (Maps 3, 5).

Comments. This species is similar in appearance to some specimens of A. placens
but differs from that species by the pale first antennal segment. It is also closely
related to A. melanopyga, but A. apta is very consistently marked, whereas A. me-
lanopyga is extremely variable. The two are also geographically isolated, with A.
melanopyga restricted to Mexico and northern Central America, and A. apta occurring
from Costa Rica southward into much of South America. Arocera apta is also similar
to A. rufifrons but can be separated from that species by its larger ocelli and usually
by the yellow ground color. Arocera rufifrons has very small ocelli, and the ground
color is usually orange.

This species is most closely related to A. spectabilis and may only be a subspecies
of that species. The genitalia of the two species are very similar, but the differences
in coloration are distinctive and consistent. Arocera spectabilis always has black
markings on the humeral angles, the connexiva, and the abdominal stemites; these
markings are always absent in A. apta.

NOMINA DUBIA

Arocera fasciativentris Breddin, 1901

Arocera fasciativentris Breddin, 1901:139; Kirkaldy, 1909:367; McDonald, 1984:118

[nomen dubium\.

Arocera fasciativentris Breddin was described from Rio Grande do Sul, Brazil.
McDonald (1984) listed this species as a nomen dubium as he was not able to locate
any type material and it is not possible to fix the species from the description. He
speculated that it was probably a junior synonym of A. spectabilis. The description
does match A. spectabilis fairly well, except that Breddin indicates that the antennae
are black. This suggests that A. fasciativentris may be a junior synonym of A. placens.

134

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Arocera capitata Breddin, 1901

Arocera capitata Breddin, 1901:139; Kirkaldy, 1909:367; McDonald, 1984:118 [no-
men dubium ].

McDonald (1984) also listed this species as a nomen dubium and suggested that
it was a junior synonym of A spectabilis. Once again, its description fits A spectabilis
fairly well, but its type locality of Bolivia places it well outside the range of A.
spectabilis. Breddin mentions that the eyes are unusually small. I have not seen any
specimens in which the compound eyes are small, but A. rufifrons has very small
ocelli, and if Breddin was actually referring to the ocelli, then A. capitata is probably
a junior synonym of A. rufifrons.

ACKNOWLEDGMENTS

I would like to thank the following individuals for their help in lending specimens pertinent
to this project (codens in parentheses; DAR is author’s collection): AMNH— American Museum
of Natural History, New York, R. T. Schuh; BMNH— British Museum (Natural History),
London, W. R. Dolling; CAS— California Academy of Sciences, San Francisco, P. H. Amaud;
CIAT— Centro Intemacional de Agricultura Tropical, Cali, Colombia, A. Pantoja; CNC—
Canadian National Collections, Ottawa, Canada, M. D. Schwartz; DBT — D. B. Thomas personal
collection, Tuxtla Gutierrez, Mexico; EGER— J. E. Eger personal collection, Tampa, Florida;
ENGL— H. D. Engleman personal collection, Coco Solo, Panama; FSCA— Florida State Col-
lection of Arthropods, Gainesville, F. Mead; HCO— Hope Entomological Collections, Oxford,
England, I. Lansbury; LHR— L. H. Rolston personal collection, Baton Rouge, Louisiana; MACN —
Museo Argentino de Ciencias Naturales “Bernardino Rivadavia,” Buenos Aires, A. O. Bach-
man; OSUC— Ohio State University, Columbus, C. A. Triplehom; UNAM— Universidad Na-
cional Autonomia de Mexico, Mexico City, H. Brailovsky; USNM — United States National
Museum of Natural History, Washington, D.C., T. J. Henry; ZMB— Zoologisches Museum,
Berlin, J. Deckert; ZSM— Zoologische Staatssammlung, Munich, M. Baehr.

I would like to give special thanks to P. H. Amaud, M. Baehr, W. R. Dolling, T. J. Henry,
and I. Lansbury for the loan of relevant type material. I would also like to express my appre-
ciation to J. B. Chapin, J. A. Moore, and L. H. Rolston (Louisiana State University), J. E. Eger
(DowElanco, Tampa), and D. B. Thomas (USDA-AG/SEA, Tuxtla, Mexico) for their helpful
review of an early version of the manuscript.

This publication was approved by the Director of the Louisiana Agricultural Experiment
Station as manuscript number 90-17-4513.

LITERATURE CITED

Amyot, C. J. B. and J. G. A. Serville. 1843. Histoire naturelle des insectes Hemipteres. In:
“Suites a Buffon.” Fain et Thunot, Paris, LXXVI + 675 + 8 pp., 12 pis.

Becker, M. and J. Grazia-Vieira. 1971. Contribu^ao ao conhecimento da superfamilia Pen-
tatomoidea na Venezuela (Heteroptera). Iheringia (Zool.) 40:3-26.

Becker, M. and J. Grazia-Vieira. 1977. The Pentatomoidea [Heteroptera] collected in French
Guiana by the expedition of the Museum National d’Histoire Naturelle. Ann. Soc. Ent.
Fr. (N.S.) 13(1 ):53— 67.

Brailovsky, H. and E. Barrera. 1982. Hemiptera-Heteroptera de Mexico. XXII. Nuevos re-
gistros de la tribu Pentatomini y descripcion de una nueva especie del genero Pseude-
voplitus Ruckes (Pentatomidae). An. Inst. Biol., Univ. Nal. Auton. de Mex. 52 (1981),
Ser. Zool. (1):23 1-246, 15 figs.

1992

REVISION OF AROCERA

135

Breddin, G. 1901. Neue neotropische Wanzen und Zirpen. Soc. Entomol. 16:41-42, 52-53,
59-60, 82-84, 90-91, 100-101, 107-108, 123-124, 138-139.

Burmeister, H. C. C. 1835. Handbuch der Entomologie, Tome 2. T. Enslin, Berlin. Abtheil
1 :i— xii, 1-400, 2 pis.

Dallas, W. S. 1851. List of the specimens of Hemipterous Insects in the Collection of the
British Museum. Taylor & Francis Incorporated, London 1:1-368, pis 1-1 1.

Distant, W. L. 1880. Insecta. Rhynchota, Hemiptera-Heteroptera. In: F. D. Godman and O.
Salvin (eds.), Biologia Centrali-Americana, Vol. 1. London, 1880:1-88.

Distant, W. L. 1893. Insecta. Rhynchota, Hemiptera-Heteroptera. In: F. D. Godman and O.
Salvin (eds.), Biologia Centrali-Americana, Vol. 1. London, 1893:329-462.

Distant, W. L. 1900. Rhynchotal notes. — IV. Heteroptera: Pentatomidae. Ann. Mag. Natur.
Hist. (7)5:386-397, 420-435.

Drury, D. 1782. Illustrations of Natural History Wherein Are Exhibited Upwards of Two
Hundred and Forty Figures of Exotic Insects According to Their Different Genera, Vol.
2. B. White, London 3:I-XII, 1-76, pis. 1-50, index.

Froeschner, R. C. 1981. Heteroptera or true bugs of Ecuador: A partial catalog. Smithson.
Contrib. Zool. 322:iv + 147 pp.

Gistel, J. N. F. X. 1848. Naturgeschichte des Thierreichs. Fiir hohere Schulen. Hoffman,
Stuttgart, XVI + 216 + 4 pp., 32 pis.

Grazia, J. 1977. Revisao dos Pentatomineos citados no “Quarto catalogo dos insetos que
vivem nas Plantas do Brasil.” (Hemiptera-Pentatomidae-Pentatomini). Dusenia 10(3):
161-174.

Grazia, J. 1984. Pentatomini da Venezuela (Heteroptera, Pentatomidae). An. Soc. Ent. Brasil
1 3( 1 ):7 1—8 1 .

Haglund, C. J. E. 1868. Hemiptera nova. Stett. Entomol. 29:150-163.

Herrich-Schaffer, G. A. W. 1 844. Die Wanzenartigen Insecten. C. H. Zeh’schen Buchhandlung,
Niimberg. 7:41-134.

Kirkaldy, G. W. 1909. Catalogue of the Hemiptera (Heteroptera) with Biological and Ana-
tomical References, Lists of Foodplants and Parasites, etc. Felix L. Dames, Berlin, 1:1-
XL, 1-392.

Lethierry, L. F. and G. Severin. 1893. Catalogue General des Hemipteres. Pentatomides. R.
Friedlander and Fils, Bruxelles and Berlin, 1:I-X, 1-286.

McDonald, F. J. D. 1984. Revision of Arocera Spinola (Hemiptera: Pentatomidae). J. New
York Ent. Soc. 92(2):97-120.

Perty, J. A. M. 1833. Delectus animalium articulatorium quae in itinere per Brasiliam annis
1817-1820, jussu et auspiciis Maximiliani Josephi Bavariae regis augustissimi peracto,
collegerunt Dr. J. B. de Spix et Dr. C. F. Ph. de Martius. Fasc. 3. Monachii. Pp. 1 25—
224, pis. 25-40.

Piran, A. A. 1955. Arocera contralineata, especie neuva de la fauna Argentina (Hemiptera,
Pentatomidae). Neotropica 1:81-84.

Piran, A. A. 1956. Hemipteros raros o poco conocidos y no mencionados para las faunas de
Brasil, Uruguay, Argentina, Paraguay y Bolivia. Rev. Soc. Uruguaya Entomol. 1(1 ):29—
35, 2 figs.

Piran, A. A. 1962. Hemiptera neotropica V. Notas sobre sistematica y zoogeografia de Pen-
tatomidae. Acta Zool. Lilloana 18:5-10.

Piran, A. A. 1963. Hemiptera neotropica VIII. Especies neuvas o poco conocidas de las jauans
de Colombia, Peru, Brazil, Bolivia y Paraguay. Physis 24(67): 107-1 12. [“Brazil” in error
for Ecuador]

Piran, A. A. 1966. Hemiptera Neotropica X. Rev. Soc. Ent. Arg. 28:85-89.

Rider, D. A. 1991. Rhyssocephala, new genus, with the description of three new species
(Heteroptera: Pentatomidae). J. New York Ent. Soc. 99:582-609.

136

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Rolston, L. H. 1976. An evaluation of the generic assignment of some American Pentatomini
(Hemiptera: Pentatomidae). J. New York Ent. Soc. 84( 1 ):2— 8.

Rolston, L. H. 1987. Two new genera and species of Pentatomini from Peru and Brazil
(Hemiptera: Pentatomidae). J. New York Ent. Soc. 95(1 ):62— 68.

Rolston, L. H. and F. J. D. McDonald. 1984. A conspectus of Pentatomini of the western
hemisphere. Part 3 (Hemiptera: Pentatomidae). J. New York Ent. Soc. 92(l):69-86.

Sabrosky, C. W. 1972. The case of the elusive author. Proc. Ent. Soc. Wash. 74:87-88.

Showalter, W. J. 1929. Insects: exploring the wonders of the insect world. Nat. Geog. Mag.
(July): 1-90.

Signoret, V. A. 1851. Description de nouvelles especes d’Hemipters. Ann. Soc. Ent. Fr. 9(2):
329-348.

Spinola, M. M. 1837. Essai sur les genres d’lnsectes appartenants a l’ordre des Hemipteres
Lin. ou Rhyngotes, Fab. et a la section Heteropteres, Dufour. Chez Yves Gravier, Genes,
383 + 16 pp.

Stcil, C. 1854. Nya Hemiptera. Ofv. Kongl. Sven. Vet.-Akad. Forh. 1 1 (8):23 1—255.

StM, C. 1858. Hemipterologiska bidrag. Ofv. Kongl. Sven. Vet.-Akad. Forh. 1 5(9— 1 0):433—
454.

St&l, C. 1 860. Bidrag till Rio Janeiro-traktens Hemipter-fauna. Kongl. Sven. Vet-Akad. Handl.
2(7): 1-84.

StM, C. 1861. Miscellanea Hemipterologica. Stett. Ent. Zeit. 22(4-6): 129-1 53.

Stcil, C. 1862. Hemiptera Mexicana enumeravit speciesque novas descripsit. Stett. Ent. Zeit.
23( 1— 3):8 1 — 1 18.

Stcil, C. 1867. Bidrag till Hemipteremas systematik. Conspectus generum Pentatomidum
Americae. Ofv. Kongl. Vet.-Akad. Forh. 24(7):522-532.

StM, C. 1872. Enumeratio Hemipterorum: Bidrag till en foreteckning ofver alia hitills kaanda
Hemiptera, jemte systematiska meddelanden. Kongl. Sven. Vet.-Akad. Handl. 1 0(4): 1—
159.

Van Duzee, E. P. 1931. Four new neotropical Heteroptera. Pan-Pacific Ent. 8(2):93-95.

Van Duzee, E. P. 1937. A few new Hemiptera. Pan-Pacific Ent. 13:25-27.

Walker, F. 1867. Catalogue of the specimens of Heteropterous-Hemiptera in the collection
of the British Museum. British Museum, London 1:1-240.

Westwood, J. O. 1837. A Catalogue of Hemiptera in the Collection of the Rev. F. W. Hope,
with short Latin descriptions of new species. J. Bridgewater, London 1 : 1 — 46.

Received 18 December 1990; accepted 17 April 1991.

J. New York Entomol. Soc. 100(1): 137-141, 1992

DESCRIPTION OF THE MALE OF OXYPORUS BALLI
CAMPBELL, WITH NOTES ON DISTRIBUTION OF
OXYPORUS IN MEXICO
(COLEOPTERA: STAPHYLINIDAE)

Jose Luis Navarrete-Heredia

Lab. de Morfofisiologia Animal, Fac. de Ciencias, UNAM,

Apdo. Postal 21-518, 04000 Mexico, D.F., and
Instituto Gran Union, Kramer 74, Col. Atlantida,

04370 Mexico, D.F.

Abstract. — The male of Oxyporus balli Campbell is described for the first time,
based on specimens collected in Morelos, Mexico. Conspecific females collected in
the same locality differed in color. The host mushrooms for O. balli are recorded for
the first time: Amanita caesarea and A. rubescens. New records and host mushroom
for O. mexicanus Fauvel and O. lawrencei Campbell are included.

The original description of Oxyporus balli by Campbell (1969) was based on two
females collected in the Mexican states of Michoacan and Morelos. The purpose of
this paper is to describe the male of O. balli and comment on color pattern variation
in females, all collected recently in Morelos.

Oxyporus balli Campbell, 1969
Can. Entomol. 101:254

Description of male. Black but labrum and tarsi light yellow, elytra (except broad
vitta on extreme outer apical angles) bright yellow.

Head quadrate to slightly transverse, its width (including eyes) slightly broader
than width of pronotum; temples shorter than eyes. Frons near to clypeus with two
depressions between antennae. Anterior margin of clypeus slightly emarginate me-
dially. Mandibles shorter than length of head, with deep depression close to base;
left mandible with two teeth near middle to receive right mandible, one directed
ventrally and one medially; each mandible slightly curved with acute apex.

Thorax with pronotum slightly wider than long, widest near anterior margin, grad-
ually narrowed to base. Suture between mesepisternum and mesepimeron slightly
carinate. Tarsi slender, lacking fine setae ventrally; first tarsomere shorter than second,
fifth longest.

Abdomen with tergite VIII and sternites VII and VIII moderately densely covered
with long setae near posterior margin; other sternites with row of long setae sparsely
placed near posterior margin. Aedeagus (Fig. 5) with broad median lobe, its apex
broadly rounded; parameres without apical setae.

Material examined. MEXICO: Morelos, Tlayacapan, San Jose de los Laureles,
3. VI. 1990, cols. J. L. Navarrete y G. A. Quiroz, ex Amanita caesarea, #283, Oxyporus
balli Campbell, Det. J. L. Navarrete 1990, (1$); same data except for: ex Amanita
rubescens, 05.VIII.1990, #440 (1 6, 1 9); same data except for: 06.VIII.1990, #453,

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1 2

Figs. 1, 2. Dorsal habitus. 1. Oxyporus balli Campbell, male. 2. O. balli, female.

( 1 <3). Specimens are deposited in my personal collection, and in the Canadian National
Collection.

Distribution. Known only from Michoacan and Morelos at high altitude. The
specimens examined were collected at 1,700 m.

Habitat and habits. The specimens examined were collected from two species of
Amanita in a cloud forest, in June and August.

Remarks. The color pattern of the two females examined differs from that described
by Campbell (1969) as follows: the metastemum, mesepimeron, metepisternum,
metepimeron and sternites I-V are yellow; tergites III-VI are yellow with black spots
at lateral margins; in one female, the hind legs are mostly yellow except for a small
black spot near the apex of the femur. Male and female habitus and color pattern
are illustrated in Figures 1-2. This species resembles O. rufipennis LeConte from
eastern North America, however it differs primarily in having the extreme outer
apical angles of the elytra black, the elytral epipleura inconspicuous, the mesepi-
sternum broader (Figs. 3, 4), and by differences in the aedeagus (Figs. 5, 6).

NEW MEXICAN DISTRIBUTIONAL RECORDS FOR OXYPORUS

O. lawrencei Campbell is known from Eslava (Distrito Federal), Sierra de Taxco
(Guerrero), and Derrame del Chichinautzin (Morelos) (Campbell, 1974; Navarrete-
Heredia and Novelo-Gutierrez, 1990). The following label data provide new records

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OXYPORUS BALL I

139

5 6

Figs. 3-6,Oxyporus spp. 3, 4: Pterothorx, lateral view. 3. O. balli. 4. O. rufipennis LeConte.
5, 6: Aedeagus, lateral and ventral views, respectively. 5. O. balli. 6. O. rufipennis.

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for an additional state: MEXICO: Mexico, San Rafael, cerca “caseta,” 09.IX.1989,
col. J. L. Navarrete, ex hojarasca, cerca de hongos, Oxyporus lawrencei Campbell,
Det. J. L. Navarrete 1989, (1); km 14-17 carr. Cuernavaca-Ocuilan, 27. VII. 1990,
ex hojarasca, cerca de Tricholoma sp., #408, Oxyporus lawrencei Campbell, Det. J.
L. Navarrete 1990, (1).

O. mexicanus Fauvel is known from Durango, Guerrero, Oaxaca and Morelos
(Campbell, 1969; Navarrete-Heredia and Novelo-Gutierrez, 1990). The following
label data provide additional habitat information: MEXICO: Morelos, Tlayacapan,
San Jose de los Laureles, 3. VI. 1990, cols. J. L. Navarrete y G. A. Quiroz, ex Amanita
caesarea #277, Oxyporus mexicanus Fvl., Det. J. M. Campbell (4); same data except
for, #283, Det. J. L. Navarrete (3).

REMARKS

Scheerpeltz (1960) described O. bolivianus from Bolivia; the female type is de-
posited in Munich (A. F. Newton, pers. comm.). This species was excluded from
Campbell’s ( 1 969) revision. Scheerpeltz ( 1 960) also cited questionable locality records
for various Mexican species: for O. smithi Bemhauer and O. guerreroanus Bemhauer,
Iguala and Tapantitlan (also cited as Tepantitlan) (the first locality is in Guerrero
and the latter is without precise geographic locality); for O. mexicanus Fauvel and
O. flohri Sharp, Juquila in Oaxaca; and for O. fenyesi Bemhauer (=0. mexicanus),
Toluca, the capital of the Estado de Mexico.

The type series of O. flohri was originally collected by Flohr at “Alto del Tizar,”
a questionable locality in Mexico (see Campbell, 1990). Campbell suggested that this
locality is probably in the mountainous areas of Chiapas. Recently, Biol. L. Delgado
(Instituto de Ecologia) discovered a locality named “Alto del Tizar” in the Mexican
state of Veracruz, near to Naolinco. It is possible this “Alto del Tizar” is the true
type locality of O. flohri. If confirmed as such, this geographic locality will represent
the first record of an Oxyporus from eastern Mexico.

ACKNOWLEDGMENTS

The author acknowledges Biol. G. A. Quiroz-Rocha for her help with the line drawings and
her company during collecting trips. Drs. J. H. Frank (University of Florida, Gainesville), J.
M. Campbell (Biosystematics Research Centre, Ottawa), and A. F. Newton, Jr. and M. K.
Thayer (Field Museum of Natural History, Chicago) kindly reviewed an earlier draft of the
manuscript. Mr. L. Delgado (Instituto de Ecologia) discovered and informed me of the “Alto
del Tizar” locality in Veracruz. I also thank M. J. A. Hernandez G. (Lab. de Microcine, Fac.
de Ciencias) for preparation of the habitus photographs.

LITERATURE CITED

Campbell, J. M. 1969. A revision of the New World Oxyporinae (Coleoptera: Staphylinidae).
Can. Entomol. 101:225-268.

Campbell, J. M. 1974. A new species of Oxyporus (Coleoptera: Staphylinidae) from Mexico
with comments on Oxyporus elegans LeC. Coleopts. Bull. 28:155-157.

Campbell, J. M. 1 990. A new species of Oxyporus (Coleoptera: Staphylinidae) and rediscovery
of O. flohri from Guatemala. Coleopts Bull. 44:21 1-215.

Navarrete-Heredia, J. L. and R. Novelo-Gutierrez. 1990. Contributions to the knowledge of

1992

OXYPORUS BALLI

141

Oxyporinae (Coleoptera: Staphylinidae) associated with mushrooms (Fungi: Basidio-
mycetes) in Mexico. Coleopts Bull. 44:229-232.

Scheerpeltz, O. 1 960. Zur Kenntnis neotropischer Staphyliniden (Col.). Btr. Z. Neotrop. Fauna
2:65-138.

Received 9 November 1990; accepted 16 August 1991.

J. New York Entomol. Soc. 100(1): 142-1 54, 1992

REVIEW OF THE SPECIES OF TELEPHANUS ERICHSON
FROM THE MALAGASY REGION, WITH
DESCRIPTION OF A NEW SPECIES
(COLEOPTERA: SILVANIDAE)

M. C. Thomas

Florida State Collection of Arthropods, P.O. Box 147100,

Gainesville, Florida 32614-7100

Abstract. — Six species of Te/ephanus Erichson are recorded from the Malagasy Region, in-
cluding one new species, Te/ephanus gomyi, from Reunion Island. Te/ephanus spinosus Schau-
fuss is considered a junior synonym and junior homonym of Te/ephanus spinosus Grouvelle.
A key to the species and illustrations of the adults are provided and the affinities of this group
of species are briefly discussed.

While examining a collection of Cucujidae (sens, lat.) from the Mascarene Islands
(Reunion and Mauritius) in the Museum d’Histoire Naturelle, Geneva I discovered
three specimens of an apparently undescribed apterous, microphthalmic species of
Te/ephanus. This led to an examination of the types and other specimens of all five
species described from the Malagasy Region (Madagascar and the Mascarene Islands).
This study revealed three distinctive groups of species and raised questions over the
generic placement of the Old World members of Telephanus.

Of about 100 described species of Telephanus, Hetschko (1930) listed only eight
from the Old World, five from the Malagasy Region and three from southeastern
Asia. The remainder are confined to the New World. Adults of Telephanus are
recognized by the combination of the following character states: securiform maxillary
palpi; elongate antennal scapes, frons without longitudinal grooves; closed procoxal
cavities; tarsomeres II— III simply lobed; prothorax and often elytra laterally spinose;
aedeagus inverted, parameres on ventral aspect of median lobe.

Three species, Telephanus antennatus Waterhouse, described from Dory, New
Guinea; T. pictus Waterhouse, described from Borneo; and T. spinicollis Waterhouse,
described from Aru, Indonesia, were transferred from Telephanus to Psammoecus
Latreille by Nevermann (193 1: 198). I have examined the type specimens, housed in
the British Museum (Natural History), of those species and agree with Nevermann’s
conclusion as to their assignment.

It also seems unlikely that the Malagasy members of Telephanus are strictly con-
generic with the New World members of the genus. The Reunion and Mauritius
species share some similarities, especially antennal proportions, with Indophanus
Pal, a monotypic genus from South India placed near Psammoecus and Telephanus
(Pal, 1981). The four Madagascar species share some character states that are closer
to Uleiota Latreille than to New World Telephanus. These include antennomere III
more than twice as long as II and possession of a longitudinally raised lateral margin
of the frons. In approximately 30 species examined of New World Telephanus,
antennomere III is no more than 1.5 times the length of II and the frons is not

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REVIEW OF TELEPHANUS

143

Fig. 1. Telephanus alluaudi Grouvelle. Line = 1.0 mm.

laterally margined. The hindwing venation of T. armatus (Fig. 1 4) differs considerably
from both Uleiota dubius (Fabricius) (Fig. 13) and Telephanus velox (Haldemann)
(Fig. 15). However, the two species from Mauritius and Reunion resemble New
World Telephanus in antennal and frontal characters, and it may be that the Uleiota-
like structures in the Madagascar species represent retention of ancestral character

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Fig. 2. Telephanus gomyi, n. sp. Line = 1.0 mm.

states. For that reason, I am proposing no generic changes at present. The distribution
of the species of Telephanus, as presently constituted, also is anomalous and suggests
that the composition of this genus merits closer study.

The number and disposition of socketed prothoracic spines are important diag-
nostic features of New World Telephanus. Unfortunately, many of the spines are
missing from most of the Madagascar specimens examined during this study and

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REVIEW OF TELEPHANUS

145

Fig. 3. Telephanus armatus Grouvelle, habitus of male. Inset is dorsal view of right hind
leg. Line = 1.0 mm.

their presence has been inferred from the empty sockets. The Madagascar specimens
examined that bear specific locality information were collected in Antananarivo,
spelled either “Annanarivo” or “Antanarivo” on the specimen labels. Members of
the type series of three species in the Museum National d’Histoire Naturelle, Paris
were examined by S. A. Slipinski, who affixed lectotype and paralectotype labels to
some of the specimens. Since these designations have not been published they are
not valid under the International Code of Zoological Nomenclature.

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Fig. 4. Telephanus spinosus Grouvelle, male. Line = 1.0 mm.

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147

Figs. 5-7. 5. Telephanus armatus Grouvelle, left elytron of female. 6. Telephanus spinosus

Grouvelle, same. 7. Telephanus cedius Schaufuss, pronotum. Lines = 1.0 mm.

Specimens were borrowed from the following institutions:

BMNH = British Museum of Natural History, London. Mr. A. D. Kerley.

MCSN = Museo Civico di Storia Naturale “Giacomo Doria,” Genova. Dr. Roberto
Poggi.

MNHN = Museum National d’Histoire Naturelle, Paris. Mile. Nicole Berti.
MHNG = Museum d’Histoire Naturelle, Geneva. D. Ivan Lobl.

ZMHB = Museum fur Naturkunde der Humboldt Universitat zu Berlin, Bereich
Zoologisches Museum, Berlin. Dr. Fritz Hieke.

KEY TO THE SPECIES OF TELEPHANUS OF THE MALAGASY REGION

1. Terminal antennomeres dark (Figs. 1, 2). Mascarene Islands 2

Terminal antennomeres pale (Figs. 3, 4, 8). Madagascar 3

2( 1 ). Eyes reduced (Fig. 2), elytral humeri rounded; hind wings absent; median lobe sharply

constricted medially, flagellum thick (Fig. 10); Reunion Is gomyi, n. sp.

- Eyes not reduced (Fig. 1), elytral humeri normal; hind wings present; median lobe

not sharply constricted medially, flagellum slender (Fig. 9); Mauritius

alluaudi Grouvelle

3(1). Elytra paler than head and pronotum 4

- Elytra same color as head and pronotum 5

4(3). Pronotum with conspicuous tubercles at anterio-lateral angles (Fig. 7)

cedius Schaufuss

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- Pronotum without conspicuous tubercles at anterio-lateral angles (Fig. 8)

gracilis Schaufuss

5(3). Trochanter and basal part of femur pale; hind trochanter in male prolonged poste-
riorly as acute spine (Fig. 1 2); hind femur in male armed medially with a blunt spine
and a carina (Fig. 4), hind tibia armed with a large spine medially on ventral edge;
elytral suture in male elevated at about apical third, appearing “pinched”; lateral
margin of elytra in female not explanate (Fig. 6) spinosus Grouvelle

- Only trochanter pale; hind trochanter in male slightly prolonged posteriorly, truncate;
hind femur of male with carina medially and acute spine at about apical third (Fig.

3); hind tibia in male with small tubercle located dorsally near apex (Fig. 3); elytral
suture not elevated (Fig. 3); lateral margin of elytra in female explanate (Fig. 5) . .
armatus Grouvelle

Telephanus alluaudi Grouvelle
Figs. 1, 9

Telephanus Alluaudi Grouvelle, 1 899: 172

Types. Lectotype, here designated, in MNHN with label data as follows: “I. Maurice
Curepipe Ch. Alluaud”/“MUSEUM PARIS 1917 Coll. GROU VELLE”/“T elephan-
us alluaudi Grouv”/“LECTOTYPE Left sp”/“PARALECTOTYPE Right”/“Tele-
phanus alluaudi Gru. det S.A. Slipinski 85” [two specimens glued to the same card].
I select the specimen on the left as lectotype and the specimen on the right as
paralectotype; 2, “I. Maurice Curepipe Ch. Alluaud 1897”/“TYPE‘’7“MUSEUM
PARIS coll. Alluaud’7“Telephanus alluaudi Grouv”[in Grouvelle’s hand] [two spec-
imens glued to same card; both are paralectotypes].

Type locality. Described from Curepipe, Mauritius, from an unstated number of
specimens.

Diagnosis. Adults of this species are most similar to those of the following species,
but can be distinguished by their normal humerus (Fig. 1) and metastemum, presence
of kind wings, more basally narrowed pronotum (Fig. 1), and structure of the male
genitalia (Fig. 9). The elytral maculae (Fig. 1) may be absent. Length 3.0 mm-3.5
mm.

Biology. The only information was provided with the original description: “Cap-
tured between the leaves of a small species of Pandanus. ”

Discussion. This species and the new species described below constitute an easily
recognized group within the Malagasy Region components of Telephanus, charac-
terized by their small size, coloration, antennal proportions, and male genital struc-
ture.

Telephanus gomyi, new species
Figs. 2, 10

Types. Holotype male in MHNG with data as follows: “La Reunion Cirque de
Salazie Piton Marmite 1 ,800 m. 4- 1 - 1 972 Tamisage tronc pourri Y. Gomy.” Paratype
females, 2, with same data [MHNG].

Diagnosis. Most similar to adults of T. alluaudi, adults of this species can be
distinguished by their small eyes, lack of hind wings and consequent reduction of
the humerus and metasternum, more parallel-sided pronotum (Fig. 2) and structure
of the male genitalia (Fig. 10).

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Description. Elongate, narrow. Pale testaceus, appendages somewhat paler, dark
markings as follows: antennomeres IX-XI and apex of VIII, elytral maculae at about
apical third. Length 2.8 mm. Head: Transverse (1:1.18), broadest just behind eyes.
Punctation absent or obscured by coarse turberculate microsculpture; pubescence
composed of pale, stout, recumbent to suberect setae, directed anteriorly; not ob-
scuring surface. Eyes small, in dorsal view about 5 facets wide, length about 0.18 of
head capsule length measured from apex of clypeus to basal transverse groove.
Antennae elongate, attaining about midpoint of elytra; scape elongate and stout;
antennomere proportions as follows: 5:1:1.25:2.75:2.75:2.5:2.25:2.25:2:2:2.25; an-
tennomeres VIII-XI broader than flagellar portion of antenna. Thorax: Pronotum
subquadrate, slightly longer than broad (1:0.88); broadest across apical angles, grad-
ually, evenly narrowed basally. Anterior angles right, posterior obtuse. Disk broadly,
vaguely longitudinally impressed laterally. Surface sculpture and pubescence as head.
Lateral margin, beginning at apical angle, with 1 1 tubercles armed with spines; two
additional tubercles and spines are present on each side of the apical margin and one

1 50 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

Figs. 9-12. 9. Telephanus alluaudi Grouvelle, male.genitalia. 10. Telephanus gomyi Thomas,
n. sp., male genitalia. 11, 12. Telephanus spinosus Grouvelle. 11. Male genitalia. 12. Male
anterior trochanter. Lines = 0.5 mm.

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is present on each side below and between the 1 st and 2nd tubercles. Elytra somewhat
oval in shape (length/width 1:0.48), depressed across basal third, humeral angle
absent; lateral margin explanate and with spined tubercles; spines alternating slightly
in length; striae composed of large, very shallow punctures; pubescence composed
of stout, pale setae; rows of setae alternate suberect and directed posteriorly with
recumbent and directed laterally; a few much longer setae near apex. Male genitalia:
Similar to that of T. alluaudi, differing in the more constricted median lobe, less
apically acute parameres, and broader flagellum (Fig. 10).

Variation. There are no obvious sexual differences. The two female paratypes are
2.8 mm and 3.0 mm in length. One has immaculate elytra.

Discussion. There are only two other known apterous species of Telephanus: T.
darlingtoni Nevermann (1937) and T. acrolophus Thomas (1984), both from the
mountains of eastern Jamaica. There do not seem to be any special affinities between
the two Jamaican species and T. gomyi. This species is named after its collector,
Ives Gomy, who collected many rare silvanids and cucujids on Reunion and Maur-
itius islands.

Telephanus armatus Grouvelle
Figs. 3, 5, 14

Telephanus armatus Grouvelle, 1890:127

Types. Lectotype, here designated, in MNHN with data as follows: “Madagas.”/
“exemplaire typique”/“LECTOTYPE”/“MUSEUM PARIS 1917 Coll. GROU-
VELLE”/“T. armatus A. Grouv”/“Telephanus armatus Grouv det. S.A. Slipinski
85”. Paralectotypes, 7, with label data as follows: 1, “Madagas.”/“exemplaire ty-
pique”/“MUSEUM PARIS 1917 Coll. GROU VELL’ 7“ P ARALECT OT YPE” ; 1,
“Antananarivo 1889 Sikora”/“Telephanus armatus Grouv. (An. M. C. xxix)”/“Mu-
seo Civico di Genova”; 5, “Antananarivo 1889 Sikora”/“Museo Civico di Genova”
[MNHN, MCSN]. Although specimens from the type series are housed both in Paris
and Genova, Grouvelle’s “exemplaire typique” labels on the Paris specimens leave
little choice in designating a lectotype.

Type locality. Madagascar.

Diagnosis: Adults of this species could only be confused with adults of spinosus.
In both sexes of armatus the femora are unicolorous, while the femora are bicolored
in both sexes of spinosus. Males of both species are easily distinguished by the
secondary sexual characters of the hind legs (Figs. 3, 4), and females by the differently
shaped elytra (Figs. 5, 6). Length 5.5 mm-6.0 mm.

Discussion. The type labels on several specimens listed below are explained by the
footnote on the page this species is described (Schaufuss, 1893:598): “While at the
printers, Mr. Grouvelle tells me that he has described the species under the above
name.” For this reason, I do not consider that Schaufuss (1893:598) intended to
describe this species as new at the time of publication, and the type labels are spurious.
Schaufuss (1893:599) described but did not name a variety in which the last two
antennomeres are pale; this is merely individual variation as intermediates occur.
Nevermann (1931:198) maintained that the date of Schaufuss’ publication was 1890
rather than 1893.

Distribution. Madagascar.

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Figs. 13-15. Hind wings. 13. U/eiota dubius (Fabricius). 14. Telephanus armatus Grouvelle.
15. Telephanus velox (Haldeman). Line = 1.0 mm.

Specimens examined. 20, as follows: 1, “Type’7“Madagascar”/“Coll. L. W. Schau-
fuss”/“Telephanus armatus Grouv”/“Zool. Mus. Berlin”; 2, “Coll. L. W. Schaufuss”/
“Telephanus armatus var. Type Annanarivo”/“Zool. Mus. Berlin” [both on same
pin]; 1, “Type”/“Telephanus armatus var. Type Annanarivo”/“Zool. Mus. Berlin”;
1, “Type”/“Telephanus ar=matus Grouv. Antanarivo”/“Zool. Mus. Berlin”; 7,
“Madagascar”/“Zool. Mus. Berlin” [live on one pin, two on a second pin]; 1, “Mad-

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153

agaskar”/“Zool. Mus. Berlin” [there are three specimens on the same pin, two are
female spinosus, one is a female armatus ]; 6, “Madagaskar” [one spelled “Mada-
gascar”]/“Zool. Mus. Berlin”; 1, “Madagaskar Antanarivo Sikora S.”/“Telephanus
spinosus Gr. det. S. A. Slipinski” [misidentified]. [ZMHB]

Telephanus spinosus Grouvelle
Figs. 4, 6, 11, 12

Telephanus spinosus Grouvelle, 1890:xcii

Telephanus spinosus Schaufuss, 1893:599, NEW SYNONYM, NEW HOMONYM.

Types. Of T. spinosus Grouvelle, lectotype male, here designated, in MNHN with
label data as follows: “Madagascar”/“Type”/“MUSEUM PARIS 1917 Coll. GROU-
VELLE”/“Telephanus spinosus A. Grouv” [in Grouvelle’s hand]/“LECTOTYPE”/
“Telephanus spinosus A. Grou. det S. A. Slipinski 81.” Of T. spinosus Schaufuss,
lectotype female, here designated, in ZMHB with label data as follows: “Type”/
“Telephan. spinosus Schauf. II. [female symbol] Annanarivo”/“Zool. Mus. Berlin.”
Three other specimens of the type series are in the ZMHB with data as follows: 1 ,
“Type”/“Telephan. spino=sus Schauf. II. [male symbol] Antanarivo Mag”/“Coll. L.
W. Schaufuss”/“ZooL Mus. Berlin” [head and pronotum missing]; 1, “Type”/“Te-
lephan. spino=sus Schauf. II. [female symbol] Antanarivo”/“Coll. L. W. Schaufuss”/
“Zoo!. Mus. Berlin” [pronotum and base of elytra damaged]; \ “Type”/“Coll. L. W.
Schaufuss”/“Telephan, spin-sus Schauf. II. [male symbol] Annanarivo Mag”/“Zool.
Mus. Berlin” [fragments only].

Type locality. Madagascar.

Diagnosis. See diagnosis of armatus, above. Length 6.0 mm-6.9 mm.

Discussion. Schaufuss (1 893:599) clearly felt he was describing this as a new species,
preceding his description with: “Teleph. spinosus nov. spec,” even though it had
been described by Grouvelle (189Q:xcii) three years earlier. Schaufuss (1893:600)
noted that he had received specimens of this species from Grouvelle. Hetschko ( 1 930)
made no mention of this situation in his catalog.

Specimens examined: 6, as follows: 2, “Madagascar”/“Zool. Mus. Berlin” [three
cards on pin, one specimen missing]; 1 , “Madagascar”/“Zool. Mus. Berlin”; 1 , “Mad-
agaskar”7“Telephanus spinosus A. Grouv”/“Zool. Mus. Berlin”; 2, “Madagaskar”/
”Zool. Mus. Berlin” [there are three specimens on the same pin, two are female
spinosus, one is a female armatus ]. [ZMHB]

Telephanus cedius Schaufuss
Fig. 7

Telephanus cedius Schaufuss, 1893:601

Type. Holotype, in ZMHB with label data as follows: “Telephanus cedius m Type
Annanarivo”/“Coll. L. W. Schaufuss”/“Zool. Mus. Berlin”. Head missing, sex un-
known.

Type locality. Antananarivo, Madagascar.

Diagnosis. In addition to the key character mentioned above, in the only known
specimen of this species the punctation of the pronotum is coarser and the micro-
sculpture more pronounced than in any of the specimens of gracilis examined. Un-

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fortunately, the head of the holotype is missing. Schaufuss (1893:601) stated that the
antennae of gracilis and cedius were similar. Schaufuss (1893:601) gave the length
of the holotype as 5 mm.

Te/ephanus gracilis Schaufuss
Fig. 8

Telephanus gracilis Schaufuss, 1893:600

Types. Lectotype, here designated, in ZMHB with following data: ’’Telephanus
gracilis m. Annanarivo Type“/”Coll. L. W. Schaufuss”/“Zool. Mus. Berlin.” Sex
unknown. Paralectotypes, 6, with label data as follows: 1, “Type”/“Telephanus grac-
ilis m. var. Type Annanarivo”/“Zool. Mus. Berlin”; 1 , “Type”/“Telephanus graci=lis
Schauf. II. Annanariva M.”/“Zool. Mus. Berlin” [damaged]; 4, “Coll. L. W. Schau-
fuss”/“Zool. Mus. Berlin.” [ZMHB]

Type locality. Antananarivo, Madagascar.

Diagnosis. Individuals of this species are superficially similar to cedius in sharing,
according to Schaufuss (1893:601), tricolored antennae. They can be distinguished
from those of cedius by their lack of prominent tubercles on the anterolateral angles
of the pronotum (Fig. 8) and the somewhat sparser pronotal macro- and micro-
sculpture. Length 4.6 mm-5.0 mm.

ACKNOWLEDGMENTS

I thank D. G. H. Halstead, S. A. Slipinski, and two anonymous reviewers for their critical
reading of the manuscript, and Sandra Davis for her help in translating the German. I am
especially grateful to the museum curators listed above for the loan of specimens that made
this study possible. This is Contribution No. 753, Bureau of Entomology, Division of Plant
Industry, Florida Department of Agriculture & Consumer Services.

LITERATURE CITED

Grouvelle, A. 1890. [Descriptions of Hyliota atrata and Telephanus spinosus]. Bull. Soc.
Entomol. France 1890:xcii.

Grouvelle, A. 1899. Descriptions de Clavicomes d’Afrique et de la region Malgache, 2e
memoire. Ann. Soc. Entomol. France 68:137-185.

Hetschko, A. 1930. Cucujidae. Coleopterorum Catalogus 1 5(109): 1-93.

Nevermann, F. 1931. Berichtigung zum Cucujidenkatalog. Entomol. Blatt. 32:195-199.
Nevermann, F. 1937. Beitrag zur Kenntnis der Telephanus (Col. Cucujidae). Stett. Entomol.
Zeit. 98:1-22.

Pal, T. K. 1981. A new genus and a species of Psammoecinae (Coleoptera: Silvanidae) from
South India. Oriental Insects 15:257-261.

Schaufuss, L. W. 1893. Beitrag zur Kaferfauna Madagascars. Nunquam otiosus 3:587-624.
Thomas, M. C. 1984. A new apterous species of Telephanus (Coleoptera: Silvanidae) with a
discussion of phylogenetic relationships of the Silvanidae. Coleopts. Bull. 38:43-55.

Received 19 February 1991; accepted 20 June 1991.

J. New York Entomol. Soc. 100(1): 155-1 77, 1992

HYOROPTILIDAE (TRICHOPTERA) OF COSTA RICA:

THE GENUS OXYETHIRA EATON

Ralph W. Holzenthal and Steven C. Harris

Department of Entomology, University of Minnesota,

St. Paul, Minnesota 55108, and

Aquatic Biology Program, Department of Biology, University of Alabama,

Tuscaloosa, Alabama 35487

Abstract.— Nine new species of Oxyethira (Trichoptera: Hydroptilidae) are described from
Costa Rica: O. apinolada, O. cuernuda, O. culebra, O. espinada, O. hilosa, O. rareza, O. sencilla,
O. sierruca, and O. tica. Males of each new species are described and figured. In addition,
distribution records are presented for eight previously described species occurring in Costa Rica:
O. arizona Ross, O. azteca (Mosely), O. costaricensis Kelley, O. glasa (Ross), O.janella Denning,
O. parazteca Kelley, O. parce (Edwards and Arnold), and O. simulatrix Flint. A key is provided
to males of the Costa Rican species.

This paper on the genus Oxyethira Eaton is the latest addition to our series dealing
with the taxonomy and systematics of the microcaddisflies of Costa Rica (Trichoptera:
Hydroptilidae). In this paper, we describe nine new species of Oxyethira from Costa
Rica (one is also known from Panama) and provide distribution records for an
additional eight Costa Rican species. These new species and new records are the
result of an ongoing field inventory of Costa Rican microcaddisfly biodiversity (Harris
and Holzenthal, 1990).

Kelley (1984) provided the most recent review of the genus, its subgenera and
species groups, and he included a detailed discussion of male and female morphology.
His checklist of the world species included 43 from the Neotropics. The occurrence
of 17 species, including nine new species, in Costa Rica alone, suggests that many
more Neotropical Oxyethira await discovery. Important papers describing portions
of the Neotropical Oxyethira fauna include those of Botosaneanu (1979) and Flint
1964, 1968a, b) for the Antilles, Mosely (1937) for Mexico, Mosely (1939) for Brazil,
Flint (1974) for Surinam, Flint (1980) for Argentina, and Kelley (1983) for the
Neotropics in general. Previously described Neotropical Oxyethira and their general
distribution include aculea Ross 1941 (Mexico), alaluz Botosaneanu 1979 (Cuba),
andina Kelley 1983 (S Andes), arctodactyla Kelley 1983 (Venezuela), argentinensis
Flint 1982 (Argentina), arizona Ross 1948 (SW USA; Central America; Greater
Antilles) ( cirrifera Flint 1964 is a synonym), azteca (Mosely) 1937 (SW USA to
Argentina1), bicornuta Kelley 1983 (N Brazil), bidentata (Mosely) 1934 (S Andes),
brasiliensis Kelley 1983 (Brazil), campesina Botosaneanu 1977 (Cuba), circaverna
Kelley 1983 (Ecuador; Panama), colombiensis Kelley 1983 (Colombia), costaricensis
Kelley 1983 (Costa Rica), dactylonedys Kelley 1983 (Paraguay), dalmeria (Mosely)
1937 (Mexico), discaelata Kelley 1983 (N Brazil), glasa (Ross) 1941 (Cuba; SE USA),
hyalina Muller 1879 (Brazil), jamaicensis Flint 1968 (Jamaica), janella Denning 1948
(SE USA to Amazon1), ( neglecta Flint 1964 is a synonyn), lagunita Hint 1981 (N
Argentina to S Brazil), longissima Hint 1974 (Surinam), macrosterna Hint 1974

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(Surinam; N Brazil), maryae Kelley 1983 (Colombia), maya Denning 1947 (SE USA;
Mexico), merga Kelley 1983 (Venezuela), obscura Flint 1974 (Surinam), parazteca
Kelley 1983 (Costa Rica; Ecuador), parce (Edwards and Arnold) 1961 (SW USA to
Peru1), paritentacu/a Kelley 1983 (Belize), puertoricensis Flint 1968 (Puerto Rico),
quelinda (Botosaneanu) 1979 (Cuba), quinquaginta Kelley 1983 (Ecuador), santi-
agensis Flint 1982 (N Argentina to S Brazil), scaeodactyla Kelley 1983 (Ecuador),
simulatrix Flint 1968 (Jamaica; Central America), spirogyrae Muller 1879 (Brazil),
spissa Kelley 1983 (Brazil), tega Flint 1968 (Cuba; Jamaica), ulmeri Mosely 1937
(SW USA; N Mexico), unispina Flint 1974 (Surinam), vipera Kelley 1983 (S Andes),
zilaba (Mosely) 1939 (Brazil; Paraguay; Argentina). Complete citations of papers
containing original descriptions of these species can be found in Kelley’s ( 1984) paper.

Terminology for this paper follows that of Marshall (1979). In Figures 1-9, letters
A, B, and C refer to lateral, dorsal, and ventral views, respectively, of the genitalia;
E and F are lateral and dorsal (or ventral where indicated) views of the phallus or
internal sclerites of the female genitalia. Type material will be deposited in the
National Museum of Natural History, Smithsonian Institution, Washington, D.C.
(NMNH), the Instituto Nacional de Biodiversidad, Costa Rica (INBIO), the Uni-
versity of Minnesota Insect Collection, St. Paul (UMSP), and the University of
Alabama, Tuscaloosa (UAL), as indicated below. All specimens of previously de-
scribed species upon which new records are based are from UMSP, except where
indicated.

Oxyethira Eaton

Oxyethira Eaton, 1873:143. Type species by original designation, Hydroptila costalis
Curtis, is a species of Orthotrichia and should be replaced by Hydroptila flavicornis
Pictet, accroding to Neboiss 1963.

Adult Oxyethira can be recognized by the presence of three ocelli, a 0-3-4 tibial
spur formula, and a mesoscutellum with its anterior margin convex and its posterior
margins slightly concave. General characteristics of the male genitalia include ster-
num VII with a short, apicomesal process; tergum and sternum VIII excised apically;
segment IX withdrawn into segment VIII, with upper half often reduced to a trans-
verse band and venter pointed or broadly rounded anteriorly; segment X membra-
nous and indistinct; the presence of a subgenital plate with a bilobed process origi-
nating at its basoventral margin; and a pair of short, fused inferior appendages. The
phallus is typically long and slender with a variety of modifications including scler-
otized processes, spines, and membranous lobes.

Characteristics of the female genitalia include sternum VII with a short process;
reduction of segments VIII, IX, and X; segment VIII narrowed laterally into anteriorly
directed apodemes, and X terminating in a pair of cerci. Several internal processes
and sclerites associated with the reproductive structures are diagnostic in the females.
These include the spermathecal process, which orginates anteroventrally, and the
associated spermathecal sclerite; the sclerotized dorsum of the oviduct; the posterior

1 The discovery of O. tica and the resurrection of O. parce from synonymy with O. azteca
requires a reassessment of the distributions of these species.

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HYDROPTILIDAE OF COSTA RICA

157

ring sclerite which encircles the oviduct in some species; and in species of the Azteca
Group a pair of ventral processes which support the oviduct.

Wiggins (1977) provided a detailed description of the fifth instar larva of the genus.
Larvae typically live in standing water or the slower areas of rivers. Fifth instar larvae
build transparent, flattened, bottle-shaped cases entirely of silk. Larvae generally feed
by puncturing filamentous algal cells and sucking out the cytoplasm.

Oxyethira apinolada, new species
Fig. 1

This species appears to be a member of Kelley’s ( 1 984) Aeola Group of the subgenus
Oxytrichia. It is somewhat similar to O. viper a Kelley in the dorsolateral projections
of segment IX and the dorsolateral incision of segment VIII, although this incision
is much shorter in the new species. It differs from O. vipera in the longer inferior
appendages, trilobed subgenital plate, and absence of bilobed processes.

Male. Length 2. 4-2. 6 mm. Brown in alcohol. Antennae with 27 segments. Genitalia
as in Figure 1. Abdominal sternum VII with short, apicomesal process. Segment VIII
with lightly sclerotized, setose, dorsolateral lobes; tergum VIII with broad, shallow,
mesal excavation; sternum VIII with deep, V-shaped, mesal excavation. Segment IX
extending anteriorly to middle of segment VII; in dorsal view, posterior portion wide
and rounded; in lateral view, posterior portion narrow, tapered; segment IX bearing
elongate, narrow, apically acute, sclerotized, dorsolateral processes. Segment X mem-
branous, indistinct. Inferior appendages sclerotized, elongate, slightly convex in lat-
eral view; tapering to acute apex; in ventral view, rectangular. Subgenital plate ta-
pering posteriorly; in ventral view, wide, trilobed, the lateral lobes bearing numerous
stout setae. Bilobed processes absent. Phallus somewhat rectangular; bearing scler-
otized subapical process nearly encircling distal membranous lobe and second, short,
acute, subapical, sclerotized process.

Holotype. Male, COSTA RICA: GUANACASTE: Parque Nacional Rincon de la
Vieja, Quebrada Agua Apinolada, 10.759°N, 85.292°W, el. 795 m, 25. vi. 1986, Hol-
zenthal, Heyn, Armitage (NMNH).

Paratype. COSTA RICA: ALAJUELA: Rio Pizote, 5 km (air) S Brasilia, 10.972°N,
85.345°W, el. 390 m, 12.iii. 1986, Holzenthal and Fasth, 1 male (UMSP).

Etymology. Named for the type locality, Quebrada Agua Apinolada.

Oxyethira cuernuda, new species
Fig. 2

This species appears to be a member of subgenus Tanytrichia Kelley based on the
elongate nature of segment IX and general structure of the phallus. The elongate
dorsal processes of segment VIII and the bifid lateral processes of the phallus separate
the new species from others in the subgenus.

Male. Length 2. 0-2. 4 mm. Brown in alcohol. Antennae with 26 segments. Genitalia
as in Figure 2. Abdominal sternum VII with short, apicomesal process. Segment VIII
with dorsolateral margin produced into a long, thin process; tergum and sternum
VIII deeply excised. Segment IX elongate, narrow; extending anteriorly beyond mid-
dle of segment VI. Segment X membranous, indistinct. Inferior appendages in lateral
view short, sclerotized, acute posteriorly; in ventral view widely separated, apices

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Fig. 1. Oxyethira apinolada, new species, male genitalia. A. Lateral. B. Dorsal. C. Ventral.
D. Phallus, lateral; inset, details of apical sclerites. Abbreviations: VII, VIII, IX - abdominal
segments VII, VIII, IX, respectively; ap pro VII = apicomesal process of segment VII; dl pro
IX = dorsolateral process of segment IX; dl pro VIII = dorsolateral process of segment VIII;
inf app = inferior appendage; sg = subgenital plate.

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HYDROPTILIDAE OF COSTA RICA

159

Fig. 2. Oxyethira cuemuda, new species, male genitalia. A. Lateral. B. Dorsal. C. Ventral.
D. Phallus, lateral. E. Phallus, ventral. Abbreviations: bil pro = bilobed process; others as in
Figure 1.

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narrow. Subgenital plate in lateral view elongate, narrow, and ventrally curved,
apicoventral margin with shallow notch; in ventral view wide, with pair of small
mesal projections. Bilobed processes elongate, bearing stout apical setae. Phallus
elongate, apex bulbous, membranous, bearing pair of apicolateral sclerites, each
divided distally into thin processes.

Holotype. Male, COSTA RICA: ALAJUELA: Rio Pizote, 5 km (air) S Brasilia,
10.972°N, 85.345°W, el. 390 m, 12.iii.1986, Holzenthal and Fasth (NMNH).

Paratypes. COSTA RICA: same data as holotype, 26 males (NMNH, UMSP,
INBIO, UAL); same, except 5 km N Dos Rios, 10.948°N, 85.29 1°W, 9.iii. 1986,
Holzenthal and Fasth, 9 males (UMSP).

Etymology. Spanish: “with horns,” referring to the hornlike processes of segment

VIII.

Oxyethira culebra, new species
Fig. 3

Oxyethira culebra is a member of the Aeola Group, most similar to O. bidentata
(Mosely). Both species have elongate, dorsolateral, rodlike processes associated with
abdominal segment IX, but O. culebra differs from O. bidentata and all others in the
group in the structure of the inferior appendages.

Male. 2.8 mm. Brown in alcohol. Antennae broken. Genitalia as in Figure 3.
Abdominal sternum VII with short, apicomesal process. Segment VIII with shallow
dorsolateral emargination; tergum VIII with broad, shallow, mesal excavation and
rounded lateral lobes; sternum VIII with deep, V-shaped, mesal excavation. Segment
IX narrow, extending anteriorly to middle segment VI; bearing narrow, elongate,
sinuate dorsolateral processes, each with single, long apical seta. Segment X mem-
branous, apex rounded in dorsal view. Inferior appendages in lateral view elongate,
narrow, their apices acute, upturned; each with pair of small subapicoventral lobes;
in ventral view, inferior appendages apically divergent, each with subapicomesal
sclerotized point and setose lobe. Subgenital plate and bilobed processes absent.
Phallus narrow, sinuate; ejaculatory duct protruding from apex, hairlike.

Holotype. Male, COSTA RICA: ALAJUELA: Rio Pizote, 5 km (air) S Brasilia,
10.972°N, 85.345°W, el. 390 m, 12.iii.1986, Holzenthal and Fasth (NMNH).

Etymology. Spanish: “snake,” referring to the sinuate phallus and dorsolateral
processes of segment IX.

Oxyethira espinada, new species
Fig. 4

This species belongs to the subgenus Tanytrichia Kelley and is most closely related
to O. paritentacula Kelley. It shares with that species the widely separated dorsal
processes of abdominal segment VIII. However, in O. espinada these processes are
short, do not converge apically, and have a series of large, dorsal, apical and subapical
spinelike setae.

Male. Length 2. 5-2. 7 mm. Brown in alcohol. Antennae with 34 segments. Genitalia
as in Figure 4. Abdominal sternum VII with short, apicomesal process. Segment VIII
in lateral view very acute posteriorly, apex upturned; sternum VIII with deep, narrow,
U-shaped, mesal excavation; tergum VIII with paired, narrow, sclerotized, dorsobasal

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HYDROPTILIDAE OF COSTA RICA

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Fig. 3. Oxyethira culebra, new species, male genitalia. A. Lateral. B. Dorsal. C. Ventral. D.
Phallus, lateral. Abbreviations as in Figures 1 and 2.

processes; each process bearing several stout, apicodorsal and subapicodorsal, spine-
like setae. Segment IX elongate, narrow, extending anteriorly to middle of segment
VI. Segment X membranous, indistinct. Inferior appendages in lateral view short,
apically acute; in ventral view, widely separated, apically truncate. Subgenital plate

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Fig. 4. Oxyethira espinada, new species, male genitalia. A. Lateral. B. Dorsal. C. Ventral.
D. Phallus, ventral. Abbreviations as in Figures 1 and 2.

in lateral view short, base rounded, apex acute; in ventral view, triangular. Bilobed
processes elongate, each bearing stout apical seta. Phallus with basal portion elongate,
tubular; distal portion bearing two sclerotized lateral processes, each with acute,
outwardly directed apex.

Holotype. Male, COSTA RICA: ALAJUELA: Rio Pizote, 5 km N Dos Rios,
10.948°N, 85.29 1°W, 9.iii. 1986, Holzenthal and Fasth (NMNH).

Paratypes. Same data as holotype, 2 males (UMSP, UAL); same, except 5 km (air)

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S Brasilia, 10.972°N, 85.345°W, el. 390 m, 12.iii.1986, Holzenthal and Fasth, 1 male
(INBIO).

Etymology. Spanish: “having spines,” referring to the spine-laden dorsal processes
of segment VIII.

Oxyethira hilosa, new species
Fig. 5

In most aspects this species is similar to O. paritentacula Kelley (subgenus Tany-
trichia ), but the distinctive phallus of the new species, with its threadlike lateral
projections, clearly separates the new species from O. paritentacula.

Male. Length 1.7-2. 2 mm. Brown in alcohol. Antennae broken. Genitalia as in
Figure 5. Abdominal sternum VII with short, apicomesal process. Segment VIII in
lateral view acute posteriorly; sternum VIII with deep mesal excavation; tergum VIII
with paired, sclerotized, basodorsal processes, their bases narrowly separated, their
apices narrow, acute, slightly convergent. Segment IX elongate, narrow; extending
anteriorly well into segment VI. Segment X membranous; truncate in lateral view,
rounded in dorsal view. Inferior appendages short in lateral view, their apices round -
ed; in ventral view, widely separated with apices slightly triangular. Subgenital plate
in lateral view wide; in ventral view wide, with rounded apicomesal protuberance.
Bilobed processes elongate, each bearing stout, apical seta. Phallus with pair of long
lateral arms arising in mesal membranous area; each arm thin, strongly recurved
dorsally, with dorsal, threadlike, subapical extension.

Holotype. Male, COSTA RICA: ALAJUELA: Rio Pizote, 5 km (air) S Brasilia,
10.972°N, 85.345°W, el. 390 m, 12.iii.1986, Holzenthal and Fasth (NMNH).

Paratypes. Same data as holotype, 13 males (UMSP, UAL, INBIO); PANAMA:
International Canal Zone, Lion Hill, 26.vii. 1981, R. B. Kimsey, 3 males (NMNH).

Etymology. Spanish: “having threads,” referring to the distinctive phallus.

Oxyethira rareza, new species
Fig. 6

This species is aberrant within the genus in most respects. It shares in common
with O. quinquaginta Kelley ( incertae sedis ) vestigial inferior appendages and an
asymmetrical subgenital plate, but it also shares similar elongate, dorsal processes
of segment IX with O. andina Kelley (Aeola Group of subgenus Oxytrichia ).

Male. Length 1.8-2. 1 mm. Brown in alcohol. Antennae with 34 segments. Genitalia
as in Figure 6. Abdominal sternum VII with broad, V-shaped mesal excavation;
lacking apicomesal process. Segment VIII in lateral view acute posteriorly; sternum

VIII with very deep, narrow, V-shaped excavation; tergum VIII with broad, U-shaped
excavation. Segment IX narrow, extending anteriorly well into segment VI; tergum

IX a narrow, sclerotized ribbonlike band extending posteriorly through segment VIII,
its apex recurved. Inferior appendages vestigial. Subgenital plate asymmetrical, de-
veloped only on left side, its apex bearing long, narrow, sclerotized arm. Left bilobed
process long, thin, apex with stout seta; right bilobed process absent. Phallus short,
stout, with narrow, subapicolateral sclerotized strip; apex of phallus membranous,
ejaculatory duct protruding.

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Fig. 5. Oxyethira hilosa, new species, male genitalia. A. Lateral. B. Dorsal. C. Ventral. D.
Phallus, lateral. E. Phallus, ventral.

Holotype. Male, COSTA RICA: ALAJUELA: Rio Pizote, 5 km N Dos Rios,
10.948°N, 85.291°W, el. 470 m, 9.iii.l986, Holzenthal and Fasth (NMNH).

Paratypes. Same data as holotype, 1 male (INBIO); same, except 5 km (air) S
Brasilia, 10.972°N, 85.345°W, el. 390 m, 12.iii.1986, Holzenthal and Fasth, 4 males

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(UMSP, UAL); HEREDIA: Quebrada Sura, Est. Biol. La Selva, 10.437°N, 84.010°W,
el. 50 m, 20-2 l.vi. 1986, Holzenthal, Heyn, Armitage, 1 male (UMSP).

Etymology. Spanish: “oddity,” referring to the unique genitalic structures of this
species.

Oxyethira sencilla, new species
Fig. 7

This new species shares similarities with O. macrosterna Flint and related species
(subgenus Tanytrichia ), but can be separated from other species in the subgenus by
the absence of the lateral phallic processes in the new species.

Male. Length 2.0-2. 2 mm. Antennae with 27 segments. Brown in alcohol. Genitalia
as in Fig. 7. Abdominal sternum VII with short, apicomesal process. Segment VIII
narrow; in lateral view with posterodorsal emarginations; tergum VIII with truncate,
mesal excavation and pair of lateral lobes; sternum VIII with deep, U-shaped ex-
cavation. Segment IX narrow, tapered, extending anteriorly into segment V. Segment
X indistinct, membranous. Inferior appendages short, apices acute; in ventral view
widely separated, apices appearing rounded. Subgenital plate in dorsal and ventral
views narrow with small apical excision; in lateral view thin, apex narrow. Bilobed
processes thin, elongate, each bearing stout apical seta. Phallus short, stout; with pair
of sclerotized, distal processes— dorsal one thin, apically acute, median one crescent
shaped; phallus when distended truncate distally, with prominent sickle-shaped dor-
solateral process.

Holotype. Male, COSTA RICA: ALAJUELA: Rio Pizote, 5 km N Dos Rios,
10.948°N, 85.291°W, el. 470 m, 9.iii.l986, Holzenthal and Fasth (NMNH).

Paratypes. Same data as holotype, 14 males (UMSP, INBIO); same, except 5 km
(air) S Brasilia, 10.972°N, 85.345°W, el. 390 m, 12.iii.1986, Holzenthal and Fasth,
2 males (UAL).

Etymology. Spanish: “simple, unadorned,” referring to this species’ lack of distinct
genitalic structures.

Oxyethira sierruca, new species
Fig. 8

This species is most similar to O. quinquaginta Kelley ( incertae sedis, according
to Kelley, 1984), with which it shares several atypical characters. These include the
many antennal segments, the vestigial inferior appendages, and the asymmetrical
dorsal arms of the subgenital plate. The new species is distinguished from its congeners
by the serrate margin of segment VIII and the structure of the phallus.

Male. Length 2.6-3. 3 mm. Brown in alcohol. Antennae with 45 segments. Ab-
dominal sternum VII with long apicomesal process. Segment VIII with posterodorsal
margin serrate, ragged; sternum VIII extended posteriorly; in ventral view with deep,
quadrate excavation and rounded lateral lobes; tergum VIII with small, shallow,
rounded excavation. Segment IX narrow in lateral view, rounded anteriorly in dorsal
and ventral views; extending anteriorly into segment VII; tergum IX reduced to thin
band. Inferior appendages vestigial. Subgenital plate reduced to rounded mesal lobe,
but with pair of prominent, asymmetrical dorsal arms— left arm with wide apex

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Fig. 6. Oxyethira rareza, new species, male genitalia. A. Lateral. B. Dorsal. C. Ventral. D.
Phallus, lateral. Abbreviations: bil pro = bilobed process; pro sg = process of subgenital plate;
sg = subgenital plate; trg IX = tergum IX.

bearing mesal excavation, right arm rectangular, sinuate along mesal margin. Bilobed
processes lacking. Phallus short, stout; with pair of lateral, sclerotized rods— left rod
thick, curving dorsally; right rod angled at midlength, subapico ventral margin serrate.

Holotype. Male, COSTA RICA: GUANACASTE: Quebrada Garcia, 10.6 km ENE
Quebrada Grande, 10.862°N, 85.428°W, el. 470 m, 8.iii.l986, Holzenthal and Fasth
(NMNH).

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Fig. 7. Oxyethira sencilla, new species, male genitalia. A. Lateral. B. Dorsal. C. Ventral. D.
Phallus, lateral. E. Phallus, ventral. Abbreviations as in Figures 1 and 2.

Paratypes. Same data as holotype, 96 males (UMSP); ALAJUELA: Reserva Fores-
tal San Ramon, Rio San Lorencito and tribs., 10.216°N, 84.607°W, el. 980 m, 30.iii-
l.iv. 1987, Holzenthal, Hamilton, Heyn, 7 males (INBIO); CART AGO: Quebrada
Plantanillo, 5 km E Moravia de Chirripo, 9.82 1°N, 83.407°W, el. 1,130 m, 7. viii. 1 987,

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Holzenthal, Morse, Clausen, 1 male (UMSP); PUNT ARENAS: Rio Singri, 2 km S
Finca Helechales, 9.057°N, 83.082°W, el. 720 m, 21.ii. 1986, Holzenthal, Morse,
Fasth, 2 males (UAL); Rio Guineal, 1 km S Finca Helechales, 9.076°N, 83.092°W,
el. 840 m, Holzenthal, Morse, Fasth, 3 males (UMSP).

Etymology. Spanish: “little saw,” referring to the serrate posterior margin of seg-
ment VIII of this species.

Oxyethira tica, new species
Figs. 9, 10

This species, a member of the Azteca Group of the subgenus Loxotrichia, is very
similar in overall appearance to O.janella Denning and O. puertoricensis Flint. Males
of the new species are distinguished by the acute, elongate process on the ventrolateral
margin of segment VIII which is absent in O.janella and enlarged in O. puertoricensis.
In the females, the ventral processes supporting the oviduct (Fig. 1 0C, D) are short
in the new species and elongate in both O.janella (Fig. 10E, F) and O. puertoricensis.

Previously published records of O.janella from Costa Rica (Bueno-Soria and Flint,
1978; Holzenthal, 1988) probably represent misidentifications of O. tica. To date,
we have not seen O. janella from mainland Costa Rica, although it is known from
Cocos Island in the Pacific Ocean, about 500 km southwest of the mainland.

Male. Length 2.2-3. 1 mm. Brown in alcohol. Antennae with 34 segments. Genitalia
as in Figure 9. Abdominal sternum VII with short, apicomesal process. Segment VIII
produced slightly midlaterally into segment VII, anterior margin with ventrolateral
process, extending almost to length of dorsolateral lobe and narrowing to acute apex;
VIII deeply excised ventrally with acute lateral lobes flaring slightly outward; dorsally
with U-shaped mesal excision. Segment IX elongate posteriorly, bearing setose pro-
cess, with shallow notch at apex. Inferior appendages indistinct and fused with seg-
ment IX. Subgenital plate elongate, narrowing posteriorly to acute apex; bilobed
processes closely appressed and about half length of subgenital plate. Phallus tubular
with several indistinct folds at apex. Female. Length 2. 2-2. 4 mm. Antennae with 25
segments. Similar in overall appearance to male. Genitalia as in Figure 10. Apodemes
of segment VIII short. Segments IX and X short and apparently fused dorsally, bearing
pair of cerci at apex. Oviduct supported by paired posteroventrally extended rods
which are about twice the length of the spermathecal sclerite; posterior ring sclerite
thin, curved in lateral view.

Holotype. Male, COSTA RICA: GUANACASTE: Parque Nacional Santa Rosa,
Quebrada El Duende, near La Casona, 10.838°N, 85.6 14°W, el. 280 m, 29. vi. 1986,
Holzenthal, Heyn, Armitage (NMNH).

Paratypes. COSTA RICA: ALAJUELA: Rio Pizote, 5 km N Dos Rios, 10.948°N,
85.29 1°W, el. 470 m, 9.iii. 1 986, Holzenthal and Fasth, 2 males (UMSP); Rio Pizote,
5 km (air) S Brasilia, 10.972°N, 85.345°W, el. 390 m, 12.iii.1986, Holzenthal and
Fasth, 17 males (UMSP). GUANACASTE: Parque Nacional Rincon de la Vieja,
Quebrada Agua Apinolada, 10.759°N, 85.292°W, el. 795 m, 25.vi.1986, Holzenthal,
Heyn, Armitage, 2 males (INBIO); Parque Nacional Guanacaste, Estacion Pitilla,
Rio Orosi, 10.991°N, 85.428°W, el. 700 m, 1 9-20. vi. 1988, C. M. and O. S. Flint,
R. Holzenthal, 1 male (INBIO); Parque Nacional Guanacaste, El Hacha, Quebrada
Alcomoque, 1 1.009°N, 85.577°W, el. 250 m, 26.vii.1987, Holzenthal, Morse, Clau-

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HYDROPTILIDAE OF COSTA RICA

169

Fig. 8. Oxyethira sierruca, new species, male genitalia. A. Lateral. B. Dorsal. C. Ventral.
D. Phallus, lateral. Abbreviations as in Figures 1 and 2.

sen, 1 male (UAL); Quebrada Garcia, 10.6 km ENE Quebrada Grande, 10.862°N,
85.428°W, el. 470 m, 8.iii. 1986, Holzenthal and Fasth, 1 male (UAL); Rio Los
Ahogados, 1 1 .3 km ENE Quebrada Grande, 10.865°N, 85.423°W, el. 470 m, 7.iii. 1 986,
Holzenthal and Fasth, 2 males (UMSP); Rio Mena, 4.2 km W Santa Cecilia, 1 1 .059°N,
85.448°W, el. 260 m, 1 l.iii. 1986, Holzenthal and Fasth, 1 male (UMSP); Rio Tem-
pisquito, 3 km S route 1, 10.790°N, 85.552°W, el. 75 m, 6.iii. 1986, Holzenthal and
Fasth, 1 male (UMSP); Rio Tizate, 7.2 km NE Canas Dulces, 10.773°N, 85.449°W,
el. 275 m, 28.vi.1986, Holzenthal, Heyn, Armitage, 1 male (UMSP). HEREDIA:

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Fig. 9. Oxyethira tica, new species, male genitalia. A. Lateral. B. Dorsal. C. Ventral. D.
Phallus, lateral. Abbreviations: dl lo VIII = dorsolateral lobe of segment VIII; vl lo VIII =
ventrolateral lobe of segment VIII; others as in Figure 2.

Rio Bijagual on road to Magsasay, 10.408°N, 84.076°W, el. 140 m, 12.ii.1986, Hol-
zenthal, Morse, Fasth, 1 male (UMSP). LIMON: Rio Telire and small tribs., SE
Suretka, 9.554°N, 82.892°W, el. 48 m, l.ii.1986, Holzenthal, Morse, Fasth, 1 male
(UMSP); Rio Uatsi, 8 km (air) W Bribri, 9.62°N, 82.90°W, el. 60 m, 25.iii.1987,
Holzenthal, Hamilton, Heyn, 5 males (UMSP). PUNT ARENAS: 2.8 mi E of Golfito,

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HYDROPTILIDAE OF COSTA RICA

171

Fig. 10. Oxyethira spp., female genitalia. A-D. Oxyethira tica, new species. A. Lateral. B.
Ventral. C. Internal sclerites, lateral. D. Same, ventral. E-F. Oxyethira janella Denning. E.
Internal sclerites, lateral. F. Same, ventral. Abbreviations: ov su rd = oviduct support rod; post
r scl = posterior ring sclerite; sp scl = spermathecal sclerite.

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1 8-1 9.vii. 1 967, 0. S. Flint, Jr., 1 male (NMNH); Quebrada Pita, 3 km (air) W Golfito,
8.642°N, 83.193°W, el. 15 m, 1 5.ii. 1 986, Holzenthal, Morse, Fasth, 1 male (UMSP);
Rio Ceibo, route 2, 6 km W rd. to Buenos Aires, 9.149°N, 83.377°W, el. 250 m,
20. ii. 1986, Holzenthal, Morse, Fasth, 1 male (INBIO); Rio Singri, 2 km (air) S Finca
Helechales, 9.057°N, 83.082°W, el. 720 m, 2 1 .ii. 1986, Holzenthal and Fasth, 2 males
(UMSP). SAN JOSE: Parque Nacional Braulio Carrillo, Estacion Carrillo, Quebrada
Sanguijuela, 10.160°N, 83.963°W, el. 800 m, 27.iii. 1987, Holzenthal, Hamilton,
Heyn, 1 male (UMSP); Rio General, 1 mi S San Isidro, l.vii. 1967, P. J. Spangler, 1
male (NMNH); Rio Negro, 3.5 km SE jet. route 239, 9.680°N, 84.394°W, el. 230 m,
21.iii. 1986, Holzenthal and Fasth, 1 male (UMSP).

Etymology. Named for the Costa Ricans, or “Ticos,” in recognition of the warm
hospitality they have extended to the senior author and his associates during their
research in the country.

DISTRIBUTION RECORDS FOR OXYETHIRA IN COSTA RICA

Oxyethira arizona Ross, 1948:202, fig. 4, male, United States: Arizona (INHS);

Bueno-Soria and Flint, 1978:204; Blickle, 1979:54, fig. 158; Kelley, 1984:439;

Holzenthal, 1988:62 ( arizonica , lapsus ).

Records. ALAJUELA: Reserva Forestal San Ramon, Rio San Lorencito and tribs.,
10.216°N, 84.607°W, el. 980 m, 1 3-1 6.vi. 1988, C. M. and O. S. Flint, R. Holzenthal,
1 male. GUANACASTE: Rio Tempisquito, 3 km S route 1, 10.790°N, 85.552°W,
el. 75 m, 6.iii.l986, Holzenthal and Fasth, 2 males; Parque Nacional Guanacaste,
Maritza, Rio Tempisquito, 10.958°N, 85.497°W, el. 550 m, 19-20.vii.1987, Hol-
zenthal, Morse, Clausen, 1 male.

Oxyethira azteca (Mosely) 1937:165, fig. 13, male, Mexico ( Loxotrichia ) (BMNH);

Flint, 1968:54, fig. 138-139, 149; Bueno-Soria and Flint, 1978:205; Holzenthal,

1988:62.

Records. ALAJUELA: Rio Pizote, 5 km (air) S Brasilia, 10.972°N, 85.345°W, el.
390 m, 12.iii.1986, Holzenthal and Fasth, 184 males; Rio Pizote, 5 km N Dos Rios,
10.948°N, 85.29 1°W, el. 470 m, 9.iii. 1986, Holzenthal and Fasth, 446 males. CAR-
TAGO: Rio Platanillo, 2.2 km E Tayutic, 9.82°N, 83.55°W, el. 730 m, 30.U986,
Holzenthal, Morse, Fasth, 1 male. GUANACASTE: Rio Mena, 4.2 km W Santa
Cecilia, 1 1.059°N, 85.448°W, el. 260 m, 1 l.iii. 1986, Holzenthal and Fasth, 46 males.
HEREDIA: Est. Biol. La Selva, Quebrada Sura, 10.437°N, 84.010°W, el. 50 m, 20-
21.vi. 1986, Holzenthal, Heyn, Armitage, 7 males; Est. Biol. La Selva, Rio Puerto
Viejo, 10.440°N, 84.0 12°W, el. 30 m, 19.vi.1986, Holzenthal, Heyn, Armitage, 4
males; Rio Bijagual, road to Magsasay, 10.408°N, 84.076°W, el. 140 m, 1 2.ii. 1 986,
Holzenthal, Morse, Fasth, 536 males; Rio Sarapiqui, 7 km W Puerto Viejo, 10.452°N,
84.067°W, el. 50 m, 1 1 .ii. 1986, Morse and Fasth, 18 males. PUNTARENAS: Rio
Ceibo, route 2, 6 km W rd. to Buenos Aires, 9. 149°N, 83.377°W, el. 250 m, 20.ii. 1986,
Holzenthal, Morse, Fasth, 76 males; Rio Singri, 2 km (air) S Finca Helechales,
9.057°N, 83.082°W, el. 720 m, 2 1 .ii. 1986, Holzenthal, Morse, Fasth, 3 males; SAN
JOSE: Rio General, 1 mi S San Isidro, l.vii. 1967, P. J. Spangler, 5 males (NMNH).

Oxyethira costaricensis Kelley, 1983:44, fig. 4, male, Costa Rica (NMNH); Holzen-
thal, 1988:62.

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HYDROPTILIDAE OF COSTA RICA

173

Record. HEREDIA: Los Cartagos, 24.vi.1967, P. J. Spangler, male holotype
(NMNH). Note: This species is known only from the male holotype.

Oxyethira glasa (Ross) 1 94 1 :70, fig. 28, male, United States: Oklahoma ( Loxotrichia )
(INHS); Blickle, 1979:54, fig. 160; Holzenthal, 1988:63.

Records. ALAJUELA: Rio Pizote, 5 km N Dos Rios, 10.948°N, 85.29 1°W, el. 470
m, 9.iii. 1 986, Holzenthal and Fasth, 2 males.

Oxyethira janella Denning, 1948:397, fig. 2A-B, male, United States: Florida (D. G.
Denning collection); Flint, 1968a:52, figs. 136-137, 148; Bueno-Soria and Flint,
1978:205 (record suspect); Holzenthal, 1988:63 (record suspect).

Record. COSTA RICA: Isla del Coco: 500 km SW [mainland] Costa Rica, Chatham
Bay, at light, 8-10. iv. 1979, R. Silberglied, 1 male (LACM).

Oxyethira parazteca Kelley, 1983:53, fig. 16, male, Ecuador (NMNH); Kelley, 1984:
442; Holzenthal, 1988:63.

Records. ALAJUELA: Rio Pizote, 5 km N Dos Rios, 10.948°N, 85.29 1°W, el. 470
m, 9.iii.l986, Holzenthal and Fasth, 20 males; Rio Pizote, 5 km (air) S Brasilia,
10.972°N, 85.345°W, el. 390 m, 12.iii.1986, Holzenthal and Fasth, 3 males. CAR-
TAGO: Quebrada Platanillo, 5 km E Moravia de Chirripo, 9.82 1°N, 83.407°W, el.

l, 130 m, 7.viii. 1987, Holzenthal, Morse, Clausen, 2 males; Rio Platanillo, 2.2 km
ETayutic, 9.82°N, 83.55°W, el. 730 m, 30.i. 1986, Holzenthal, Morse, Fasth, 2 males.
GUANACASTE: P.N. Rincon de la Vieja, Quebrada Agua Apinolada, 10.759°N,
85.292°W, el. 795 m, 25.vi.1986, Holzenthal, Heyn, Armitage, 38 males; Rio Gon-
gora (sulfur mine), 4 km (air) NE Quebrada Grande, 10.887°N, 85.470°W, el. 590

m, 21.vii. 1987, Holzenthal, Morse, Clausen, 4 males.

Oxyethira parce (Edwards and Arnold) 1961:405, figs. 23-25, United States: Texas
( Protoptila ) (S. W. Edwards collection).

Records. ALAJUELA: P.N. Rincon de la Vieja, Quebrada Provision, 10.769°N,
85.28 1°W, el. 810 m, 4.iii. 1986, Holzenthal and Fasth, 3 males; Reserva Forestal
San Ramon, Rio San Lorencito and tribs., 10.216°N, 84.607°W, el. 980 m, 30.iii-

l. iv.1987, Holzenthal, Hamilton, Heyn, 2 males; same, except 13-1 6. vi. 1988, C. M.
and O. S. Flint, Holzenthal, 1 male; Rio Pizote, 5 km (air) S Brasilia, 10.972°N,
85.345°W, el. 390 m, 1 2.iii. 1986, Holzenthal and Fasth, 2 males. CART AGO: Chi-
tana, 1 9.vi. 1 967, Flint and Ortiz, 4 males, 1 0 females (NMNH); Quebrada Platanillo,
5 km E Moravia de Chirripo, 9.82 1°N, 83.407°W, el. 1,130 m, 7.viii.l987, Holzen-
thal, Morse, Clausen, 5 males; Reserva Tapanti, Rio Grande de Orosi, 9.686°N,
83.756°W, el. 1,650 m, 1 8-2 l.iii. 1987, Holzenthal, Hamilton, Heyn, 1 male; Tur-
rialba, 26.viii.1972, G. F. and S. Hevel, 1 male, 1 female (NMNH). GUANACASTE:
Parque Nacional Guanacaste, Estacion Pitilla, Rio Orosi, 10.99°N, 85.428°W, el. 700

m, 1 9-20. vi. 1988, C. M. and O. S. Flint, Holzenthal, 16 males, 8 females; P.N.
Rincon de la Vieja, Quebrada Agua Apinolada, 10.759°N, 85.292°W, el. 795 m,
25.vi.1986, Holzenthal, Heyn, Armitage, 2 males; P.N. Santa Rosa, Quebrada El
Duende, nr La Casona, 10.838°N, 85.614°W, el. 280 m, 29.vi.1986, Holzenthal,
Heyn, Armitage, 1 male; Quebrada Garcia, 10.6 km ENE Quebrada Grande, 10.862°N,
85.428°W, el. 470 m, 8.iii. 1986, Holzenthal and Fasth, 2 males; Rio Gongora (sulfur

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mine), 4 km (air) NE Quebrada Grande, 10.887°N, 85.470°W, 21.vii.1987, Holzen-
thal, Morse, Clausen, 2 males; Rio Tizate, 7.2 km NE Canas Dulces, 10.773°N,
85.449°W, el. 275 m, 28. vi. 1986, Holzenthal, Heyn, Armitage, 1 1 males. HEREDIA:
Rio Bijagual, road to Magsasay, 10.408°N, 84.076°W, el. 140 m, 12. ii. 1986, Hol-
zenthal, Morse, Fasth, 17 males. LIMON: La Lola near Matina, 1 l.iii. 1965, S. and
W. D. Duckworth, 1 male (NMNH); Reserva Biol. Hitoy-Cerere, Rio Cerere, 9.67 1°N,
83.028°W, el. 90 m, 23-24.iii. 1987, Holzenthal, Hamilton, Heyn, 27 males; Rio
Banano, 1 6 km WSW Bomba, 9.888°N, 83.1 67°W, el. 1 50 m, 26.iii. 1 987, Holzenthal,
Hamilton, Heyn, 3 males; Rio Telire and small tribs., SE Suretka, 9.554°N, 82.892°W,
el. 48 m, l.ii. 1 986, Holzenthal, Morse, Fasth, 1 1 males. PUNTARENAS: 2.8 mi E
Golfito, 18-1 9. vii. 1967, O.S. Flint, Jr., 3 males, 7 females (NMNH); Quebrada Pita,
3 km (air) W Golfito, 8.642°N, 83.193°W, el. 15 m, 15.ii.1986, Holzenthal, Morse,
Fasth, 28 males; Rio Bellavista, 1.5 km NW Las Alturas, 8.95 1°N, 82.846°W, el.
1,400 m, 8-9. iv. 1987, Holzenthal, Hamilton, Heyn, 31 males; Rio Ceibo, route 2,
6 km W rd. to Buenos Aires, 9.149°N, 83.377°W, el. 250 m, 20.ii. 1986, Holzenthal,
Morse, Fasth, 27 males; Rio Coton in Las Alturas, 8.938°N, 82.826°W, el. 1,360 m,
16.ii.1986, Holzenthal, Morse, Fasth, 1 male; Rio Guineal, 1 km (air) E Finca He-
lechales, 9.076°N, 83.092°W, el. 840 m, 22. ii. 1986, Holzenthal, Morse, Fasth, 14
males; Rio Jaba at rock quarry, 1.4 km (air) W Las Cruces, 8.79°N, 82.97° W, el.
1,150 m, 14.vi. 1986, Holzenthal, Heyn, Armitage, 23 males; Rio Jaba, 2.4 km (air)
NW San Vito, 8.832°N, 82.99 1°W, el. 970 m, 13.vi.1986, Holzenthal, Heyn, Ar-
mitage, 5 males; Rio Singri, 2 km (air) S Finca Helechales, 9.057°N, 83.082°W, el.
720 m, 2 l.ii 1986, Holzenthal, Morse, Fasth, 73 males. SAN JOSE: Pacuare, Rio
General, l.vii. 1967, Flint and Ortiz, 2 males, 5 females (NMNH); Rio General, 1
mi S San Isidro, l.vii. 1967, P. J. Spangler, 9 males, 27 females (NMNH); Rio Negro,
3.5 km SE jet. route 239, 9.68°N, 84.394°W, el. 230 m, 2 l.iii. 1986, Holzenthal and
Fasth, 1 male.

Oxyethira simulatrix Flint, 1968:43, figs. 89-91, 99, male, Jamaica (NMNH);
Holzenthal, 1988:63.

Records. GUANACASTE: Parque Nacional Guanacaste, Maritza, Rio Tempis-
quito, 10.958°N, 85.497°W, el. 550 m, 19-20.vii.1987, Holzenthal, Morse, Clausen,
1 male; Rio Tempisquito, 3 km S route 1, 10.790°N, 85.522°W, el. 75 m, 6.iii. 1 986,
Holzenthal and Fasth, 1 male. LIMON: Rio Telire and small tribs., SE Suretka,
9.554°N, 82.892°W, el. 48 m, l.ii. 1986, Holzenthal, Morse, Fasth, 11 males. PUN-
TARENAS: Rio Ceibo, route 2, 6 km W rd. to Buenos Aires, 9.149°N, 83.377°W,
el. 250 m, 20.ii. 1986, Holzenthal, Morse, Fasth, 1 male; Rio Singri, 2 km (air) S
Finca Helechales, 9.057°N, 83.082°W, el. 720 m, 2 l.ii. 1986, Holzenthal, Morse,
Fasth, 1 male. SAN JOSE: P.N. Braulio Carrillo, Quebrada Sanguijuela, Est. Carrillo,
10.160°N, 83.963°W, el. 800 m, 27.iii.1987, Holzenthal, Hamilton, Heyn, 1 male;
Rio General, 1 mi S San Isidro, l.vii. 1967, P. J. Spangler, 1 male (NMNH).

KEY TO MALES OF COSTA RICAN OXYETHIRA

1 . Inferior appendages absent or vestigial (Figs. 6A, C); subgenital plate asymmetrical

(Figs. 6C, 8B) 2

- Inferior appendages present (Figs. 5A, C, 3A, C); subgenital plate symmetrical

(Fig. 2C) or absent 3

2(1). Antennae with less than 40 segments; subgenital plate asymmetrical and well

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HYDROPTILIDAE OF COSTA RICA

175

developed only on left side (Fig. 6C); tergum IX a long, narrow, ribbonlike band

(Fig. 6B); segment VIII smooth dorsally (Fig. 6B) O. rareza, n. sp.

- Antennae with more than 40 segments; subgenital plate asymmetrical and well
developed on both sides (Fig. 8B); tergum IX reduced to thin band (Figs. 8A, B);

segment VIII serrate dorsally (Fig. 8B) O. sierruca, n. sp.

3(1). Inferior appendages elongate, thin and fingerlike (Kelley, 1983, figs. 4a, b)

O. costaricensis Kelley

- Inferior appendages variable, but not thin and fingerlike (Figs. 3 A, 5 A, 7 A, 9 A) 4

4(3). Segment IX short, extending anteroventrally into segment VII (Figs. 1A, 9 A),

often broadly rounded posteriorly in ventral view (Fig. 1C) 5

- Segment IX elongate, extending anteroventrally into segment VI (Fig. 5A) or V

(Fig. 7 A), usually narrowing posteriorly in ventral view (Fig. 4C) 13

5(4). Subgenital plate short in lateral view (Ross, 1948, figs. 4, 4a; Flint, 1968a, fig. 89),
wide in ventral view (Fig. 1 C); inferior appendages short in lateral view (Fig. 1 A;
Ross, 1948, figs. 4, 4a) 6

- Subgenital plate elongate in lateral view (Fig. 9A; Flint, 1991, figs. 1 34, 135, 1 38),

triangular in dorsal view (Fig. 9B); inferior appendages elongate in lateral and
ventral views (Figs. 9A, C) 9

6(5). Segment VIII with deep, V-shaped mesal incision ventrally (Fig. 2C); inferior
appendages in ventral view elongate, rectangular and separate (Fig. 2C)

O. apinolada, n. sp.

- Segment VIII with shallow, rounded mesal incision ventrally; inferior appendages

in ventral view fused, short (Flint, 1968a, fig. 91; Ross, 1948, fig. 4a) 7

7(6). Dorsolateral process of segment VIII slender, narrowing posteriorly (Ross, 1948,

fig. 4) 8

- Dorsolateral process of segment VIII broad basally, upcurved and bearing stout

spine at apex (Ross, 1941, fig. 28) O. glasa (Ross)

8(7). Phallus tubular with ejaculatory duct protruding and distally recurved (Flint,

1968a, fig. 90) O. simulatrix Flint

- Phallus divided into central tubular lobe bearing ejaculatory duct and lateral lobe

encircling the shaft (Ross, 1948, figs. 4b, c) O. arizona Ross

9(5). Segment VIII with prominent dorsolateral lobe, incised medially (Figs. 9A, B;
Mosely, 1937, fig. 3b); subgenital plate elongate and triangular in dorsal view, in
lateral view curving ventrad distally (Figs. 9A, B) 10

- Segment VIII without dorsolateral lobe, no incision medially; subgenital plate

short and less triangular in dorsal view, in lateral view curving dorsad distally
(Kelley, 1983, fig. 16) O. parazteca Kelly

1 0(9). Inferior appendages in ventral view with wide, V-shaped excision at apex (Mosely,

1937, fig. 13); bilobed processes of subgenital plate prominent 11

Inferior appendages in ventral view with shallow, rounded excision at apex (Fig.

9C); bilobed processes of subgenital plate indistinct (Fig. 9B) 12

11(10). Dorsal process of segment IX originating at midlength; bilobed process about half
length of subgenital plate; phallus with apico ventral lip short (Flint, 1991, figs.
138-140) O. azteca (Mosely)

- Dorsal process of segment IX originating anteriorly of midlength; bilobed process

greater than half length of subgenital plate; phallus with apicoventral lip elongate
(Flint, 1991, figs. 134-137) O. parce (Edwards and Arnold)

12(10). Segment VIII with ventrolateral lobe small and rounded posteriorly (Hint, 1968a,

fig. 81) O. janella Denning2

2 Not seen from mainland Costa Rica during this study. Literature records of O. janella from
the mainland need to be reexamined in light of the discovery of O. tica.

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Segment VIII with ventrolateral lobe elongate and narrowing to an acute tip

posteriorly (Fig. 9A) O. tica, n. sp.

13(4). Segment VIII with dorsolateral processes (Figs. 2A, 4A, 5 A) 14

- Segment VIII without dorsolateral processes (Figs. 3 A, 7 A) 16

14(13). Processes of segment VIII arising dorsobasally (Figs. 4A, 5 A) 15

- Processes of segment VIII dorsolateral in position, not basal or anterior (Fig. 2A)

O. cuernuda, n. sp.

1 5( 1 4). Dorsobasal processes of segment VIII bearing numerous thick spines, widely sepa-
rated basally, then converging distally (Fig. 4B); phallus bearing two simple, scler-

otized lateral processes (Fig. 4D) O. espinada, n. sp.

Dorsobasal processes of segment VIII lacking spines, narrowly separated basally
and only slightly convergent distally (Fig. 5B); phallus with sclerotized lateral
processes divided at apex, with dorsal arm thin and elongate (Figs. 5D, E) ....

O. hilosa, n. sp.

1 6( 1 3). Segment IX extending anteriorly to middle of segment VI, bearing narrow, elongate
dorsolateral processes (Fig. 3 A); inferior appendages elongate (Fig. 3C); subgenital

plate absent; phallus tubular, lacking sclerotized processes (Fig. 3D)

O. culebra, n. sp.

Segment IX extending anteriorly into segment V, lacking dorsolateral processes
(Fig. 7 A); inferior appendages short (Figs. 7 A, C); subgenital plate prominent (Fig.

7 A); phallus with pair of sclerotized distal processes (Figs. 7D, E) O. sencilla, n. sp.

ACKNOWLEDGMENTS

We are very grateful to the Instituto Nacional de Biodiversidad and the Servicio de Parques
Nacionales of Costa Rica and their personnel for facilitating this research. Appreciation is
extended to Jacqueline Larson for typing an early draft of the manuscript. Oliver S. Flint, Jr.,
Smithsonian Institution, is thanked for loaning specimens and providing comments on the
manuscript. This material is based upon work supported by the National Science Foundation,
grants BSR-85 12368 and BSR-89 17684. Paper No. 19,014, Scientific Journal Series, Minnesota
Agricultural Experiment Station, St. Paul, Minnesota.

LITERATURE CITED

Blickle, R. L. 1 979. Hydroptilidae (Trichoptera) of America North of Mexico. New Hampshire
Agr. Exp. Sta. Bull. 509:1-97.

Botosaneanu, L. 1979. The caddis-flies (Trichoptera) of Cuba and of Isla de Pinos: a synthesis.

Stud. Fauna Curacao and other Caribb. Isl. 59(1 85):33— 62.

Bueno-Soria, J. and O. S. Flint, Jr. 1978. Catalogo sistematico de los tricopteros de Mexico
(Insecta: Trichoptera), con algunas registros de norte, centro y sudamerica. An. Inst.
Biol. Nal. Auton. Mexico, Ser. Zoologia (1): 189-2 18.

Denning, D. G. 1948. New species of Trichoptera. Ann. Entomol. Soc. Am. 41:397-401.
Eaton, A. E. 1873. On the Hydroptilidae, a family of Trichoptera. Trans. Entomol. Soc.
London. 1873:125-150.

Edwards, S. W. and C. R. Arnold. 1961. The caddis flies of the San Marcos River. Tex. J.
Sci. 13:398-415.

Flint, O. S., Jr. 1964. The caddisflies (Trichoptera) of Puerto Rico. Tech. Pap. Univ. Puerto
Rico Agric. Exp. Sta. 40:1-80.

Flint, O. S., Jr. 1968a. The caddisflies of Jamaica (Trichoptera). Bull. Inst. Jamaica Sci. Ser.
19:1-68.

Flint, O. S., Jr. 1968b. Bredin-Archbold-Smithsonian Biological Survey of Dominica 9. The
Trichoptera (caddisflies) of the Lesser Antilles. Proc. U.S. Nat. Mus. 125(3665): 1-86.

1992

HYDROPTILIDAE OF COSTA RICA

177

Hint, O. S., Jr. 1974. The Trichoptera of Surinam. Studies on the Fauna of Suriname and
other Guy anas 14:1-151.

Flint, O.S., Jr. 1980. Studies on Neotropical caddisflies XXVI: new species from Argentina
(Trichoptera). Rev. Soc. Entomol. Argentina 39:137-142.

Flint, O. S., Jr. 1991. Studies of Neotropical caddisflies, XLV: the taxonomy, phenology, and
faunistics of the Trichoptera of Antioquia, Colombia. Smithsonian Contrib. Zool. 520:
1-113.

Harris, S. C. and R. W. Holzenthal. 1990. Hydroptilidae (Trichoptera) of Costa Rica: the
genus Mayatrichia Mosley. J. New York Entomol. Soc. 98:453-460.

Holzenthal, R. W. 1988. Catalogo sistematico do los tricopteros de Costa Rica. (Insecta:
Trichoptera). Brenesia 29:51-82.

Kelley, R. W. 1 983. New Neotropical species of Oxyethira (Trichoptera: Hydroptilidae). Proc.
Entomol. Soc. Washington 85:41-54.

Kelley, R. W. 1984. Phylogeny, morphology and classification of the micro-caddisfly genus
Oxyethira Eaton (Trichoptera: Hydroptilidae). Trans. Am. Entomol. Soc. 1 10:435-463.
Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bull. Brit.
Mus. Nat. Hist. (Entomol.) 39:135-239.

Mosely, M. E. 1 937. Mexican Hydroptilidae (Trichoptera). Trans. Roy. Entomol. Soc. London
86:151-190.

Mosely, M. E. 1939. The Brazilian Hydroptilidae (Trichoptera). Novit. Zool. 41:217-239.
Neboiss, A. 1963. The Trichoptera types of species described by J. Curtis. Beitr. Entomol.
13:582-635.

Ross, H. H. 1941. Descriptions and records of North American Trichoptera. Trans. Am.
Entomol. Soc. 67:35-126.

Ross, H. H. 1948. Notes and descriptions of Nearctic Hydroptilidae (Trichoptera). J. Wash-
ington Acad. Sci. 38:201-206.

Wiggins, G. B. 1977. Larvae of the North American Caddisfly Genera (Trichoptera). Univ.
Toronto Press, Toronto. 401 pp.

Received 23 January 1991; accepted 25 April 1991.

J. New York Entomol. Soc. 100(1): 1 78—1 79, 1992

ON THE SPIDER SUBFAMILY Z A V ATT ARICIN AE
(ARANEAE, GNAPHOSIDAE)

Norman I. Platnick

Department of Entomology, American Museum of Natural History,

Central Park West at 79th Street, New York, New York 10024

Abstract. — The spider genus Zavattarica Caporiacco, and the monogeneric subfamily Za -
vattaricinae, were based on the supposed presence of a reduced number of spinnerets. However,
both described species actually have the normal gnaphosid complement of six spinnerets, and
Zavattarica Caporiacco is placed as a junior synonym of Zelotes Gistel.

Caporiacco (1941) established a new subfamily of gnaphosid spiders, the Za vat-
taricinae, for a new genus and species, Zavattarica bimamillata, based on a single
female from Ethiopia. The new genus and subfamily were said to differ from all other
gnaphosids in having only two spinnerets (the “inferiori” or anterior lateral pair).
Subsequently, Caporiacco (1947) expanded the diagnosis of the group, describing a
second species, Zavattarica tetramamillata, based on two females from Tanzania,
supposedly differing from the type species and all other gnaphosids in having only
four spinnerets (including the “superiores” or posterior lateral pair).

The existence of this subfamily was recognized in the catalog of Roewer (1955),
and maintained in the supplement to that catalog by Brignoli (1983). Aside from
those catalog citations, however, there seems to have been no significant mention of
these taxa in the subsequent literature.

Through the courtesy of Ms. Sarah Mascherini of the Museo Zoologico de “La
Specola” in Firenze, Italy, I have recently had the opportunity to examine the holotype
of Z. bimamillata. During a recent visit to the laboratory of Dr. Vladimir I. Ovtsha-
renko at the Zoological Institute, USSR Academy of Sciences, Leningrad, I also had
the opportunity to examine type material of Z. tetramamillata, which had been
borrowed by Dr. Ovtsharenko from the Hungarian National Museum in Budapest.

In both cases, Caporiacco’s observations on the spinnerets are simply erroneous.
Six spinnerets are present in these specimens. The posterior median and posterior
lateral pairs are inconspicuous, because they are partially contracted into the abdo-
men, but they are clearly present. The spiders have the metatarsal preening combs
typical of zelotine gnaphosids (Platnick and Shadab, 1982:figs. 1, 2), and epigyna
typical of the genus Zelotes Gistel (see figs. 1, 2, prepared with the help of Dr.
Mohammad U. Shadab of the American Museum of Natural History). Zavattarica
is therefore here placed as a junior synonym of Zelotes, resulting in the new com-
binations Zelotes bimamillatus (Caporiacco) and Zelotes tetramamillatus (Capo-
riacco).

Although no generally accepted subfamilial classification of gnaphosids exists (Plat-
nick, 1990), the worldwide and extremely speciose complex of genera sharing the
metatarsal preening comb (including at least Zelotes, Drassyllus Chamberlin, Cam-
illina Berland, Trachyzelotes Lohmander, Urozelotes Mello-Leitao, and Setaphis

1992

SUBFAMILY ZAVATTARICINAE

179

Figs. 1, 2. Zelotes bimamillatus (Caporiacco), epigynum. 1. Ventral view. 2. Dorsal view.

Simon) has long been informally recognized as a group. Ironically, Caporiacco’s
completely baseless subfamilial name Zavattaricinae may live on, for there seems to
have been no earlier family-group name based on any of these genera!

LITERATURE CITED

Brignoli, P. M. 1983. A catalogue of the Araneae described between 1940 and 1981. Man-
chester, 755 pp.

Caporiacco, L. di. 1941. Arachnida. In: Missione Biologica Sagan-Omo 12(Zool. 6): 1-1 59.
Caporiacco, L. di. 1947. Arachnida Africae Orientalis, a dominibus Kittenberger, Kovacs et
Bomemisza lecta, in Museo Nationali Hungarico servata. Ann. Hist.-Nat. Mus. Natl.
Hungarici 40:97-257.

Platnick, N. I. 1990. Spinneret morphology and the phylogeny of ground spiders (Araneae,
Gnaphosoidea). Am. Mus. Novit., No 2978:1-42.

Platnick, N. I. and M. U. Shadab. 1982. A revision of the American spiders of the genus
Drassyllus (Araneae, Gnaphosidae). Bull. Am. Mus. Nat. Hist. 173:1-97.

Roewer, C. F. 1955. Katalog der Araneae von 1758 bis 1940, bzw. 1954. Brussels, 2a-b:l-
1751.

Received 21 February 1991; accepted 25 March 1991.

NOTES AND COMMENTS

J. New York Entomol. Soc. 100(1): 180-1 81, 1992

THE IDENTITY AND SYNONYMY OF HYDROESSA FUSCA
GERMAR (VELIIDAE: HETEROPTERA)

Hydroessa fusca Germar (1838) has been an enigma in modem times, not recog-
nized by any worker since Mayr (1 868) who transferred it to the genus Velia Latreille
without comment. In the Drake Collection (USNM) I found one female, with a Mayr
determination label in his handwriting, collected during the Novara Expedition at
the Cape of Good Hope, South Africa. This specimen belongs to the species currently
known as Ocellovelia germari Distant, which must therefore fall as a junior synonym.
In the discussion below I substantiate the synonymy.

Ocellovelia fusca Germar, New Combination

Hydroessa fusca Germar, 1838:123 (Location of types unknown.)

Velia fusca. Mayr, 1868:180.

Ocellovelia germari Distant, 1 904:436 NEW SYNONYMY. (Type in BMNH.)

Discussion. The location or existence of Germar’s types is unknown. Horn and
Kahle (1935, p. 89) state that the Hemiptera part of his collection is in “ Zool. Univ.
Mus., Lemberg.” Presumably Mayr saw Germar’s types or at least knew the species,
but unfortunately he gives no discussion. Ocellovelia and Angilia are the only two
South African genera with general facies resembling Velia as construed at that time.

The description of Hydroessa (now Microvelia Westwood) fusca by Germar (1838)
is very brief, however it contains several characters that provide clues as to its identity
even without the determination of Mayr (1868): 1. Each elytra with a basal oblique
white streak (the only ones visible in old or greasy specimens of Ocellovelia germari
Distant), 2. The first two antennal segments basally light, distal two hairy, with dark
hairs, 3. Head exserted, 4. Hemelytra a little shorter than abdomen, 4. Venter black,
margins fuscous, 5. Legs pallid, knees darker, and 6. Twice the size of Hydroessa
reticulata Burmeister.

The description matches specimens of Ocellovelia germari in the Polhemus Col-
lection, with the exception of the rather rough size indication; however, all other
veliids known to occur in the Cape region were also compared to test the possibility
that Mayr made a mistake in his determination.

Character 6 immediately eliminates all African Microvelia species as they are all
smaller or about the same size as reticulata, i.e., males 1.5 mm, females 1.6 mm.
This character also eliminates Rhagovelia and Angilia species, all 4.8 mm or larger.
All three of these genera match the description poorly in other respects as well. Other
veliid species occurring in the Cape region belong to the genera Ocellovelia China
& Usinger (3. 7-4. 2 mm), Pseudovelia Hoberlandt (2.5 mm), and Xiphoveloidea Hob-
erlandt (2.5 mm). As can be seen, the size of fusca given by Germar does not match

1992

NOTES AND COMMENTS

181

any of the latter three genera very closely so we must examine other characters for
the latter two genera.

Pseudovelia species agree in the following: Venter black with fuscous margins; first
antennal segment basally pallid. They disagree in the following: Hemelytra with four
(2 + 2) oblique diffuse sordid white streaks; head not exserted; distal antennal seg-
ments with light colored hairs; hemelytra longer than abdomen.

Xiphoveloidea species agree in the following: Hemelytra with oblique basal white
streaks. They disagree in the following: Venter black, without fuscous margins; first
antennal segment uniformly dark; head not exserted; distal antennal segments with
light colored hairs; hemelytra longer than abdomen (macropterous form rare).

The possibility remains that fusca could be synonymous with Ocellovelia distanti
China and Usinger, but this species is rare, and is darker, which does not match
Germar’s description stating that fusca is dorsally fuscous.

The conclusion is that no other veliid matches the description of fusca as well as
Ocellovelia germari Distant, and this species is common in the Cape region; thus,
Mayr’s determination is supported, and germari must fall as a synonym.

Because no type-material of Hydroessa fusca Germar is known to exist, and in
order to insure stability, I here designate a neotype, to be placed in the U.S. National
Museum of Natural History, Washington, D.C. The neotype data is as follows:
macropterous male, South Africa, Cape Province, Blinkwater Falls, Table Mt., Cape-
town, XI-21-1949, B. Malkin.— John T. Polhemus, University of Colorado Museum,
3115 South York St., Englewood, Colorado, USA 80110.

ACKNOWLEDGMENTS

I am indebted to R. C. Froeschner, Smithsonian Intitution, U.S. National Museum of Natural
History (USNM), Washington, D.C. for the opportunity to study material in the Drake Col-
lection under his care.

LITERATURE CITED

Distant, W. L. 1904. On South African Tingidae and other heteropterous Rhynchota. Trans.
S. Afr. Phil. Soc. 14:425-436, pi. 8.

Germar, E. F. 1838. Hemiptera Heteroptera promontorii bonae spei nundum descripta, quae
collegit C. F. Drege. Silbermann’s Rev. Entomol. 5:121-192.

Horn, W. and E. Kahle. 1935. Uber entomologische Sammlungen, Entomologen & Entomo-
Museologie. Entomol. Beih. 2:1-160, pis. 1-16.

Mayr, G. 1868. Hemiptera. In: Reise der Osterreichischen Freggate Novara um die Erde in
den Jahren 1857, 1858, 1859. Teil, II, Abt. I, Wien, 204 pp., 5 pis.

J. New York Entomol. Soc. 1 00( 1 ): 1 82-1 84, 1992

DISPERSAL OF THE ERGOT FUNGUS
CLA VICEPS PURPUREA BY THE
LAUXANIID FLY MINETTIA LUPULINA

Insect visitors of diseased plants may be important in the dispersal of many plant
pathogens, and their behavior may be important in the distribution of pathogens in
plant populations (A’Brook, 1973; Burdon, 1987; DeNoog, 1988; Roche, pers. comm.).
Just as not all insect visitors of flowers are effective pollinators (Schemske and
Horvitz, 1984) it is possible that not all visitors of diseased plants are effective
dispersers of pathogens. In this study I assess the ability of two insect visitors of the
pathogen Claviceps purpurea (Fr.) Tul. (Clavicipitaceae) to disperse it to the host
plant Festuca arundinacea Schreb. (Poaceae).

Although the ascomycetous fungus Claviceps purpurea is a well studied, polycyclic
pathogen of many species of grasses (Loveless, 1971), the secondary dispersal vectors
of this pathogen have not been previously well described. At the time of primary
infection, the wind disperses ascospores after sclerotia germinate (Lutrell, 1 980). Once
ascospores successfully infect the ovaries of grasses, the fungus induces the grasses
to produce a sweet liquid called honeydew in which conidia are formed. It has been
noted that the honeydew is attractive to insects, and that the fungus is secondarily
dispersed by flies and beetles (Mower and Hancock, 1975). I known of no study,
however, that quantifies the dispersal abilities of the insect visitors of grasses infected
by C. purpurea.

At the Mountain Lake Biological Station in western Virginia, I found that indi-
viduals of F. arundinacea that are infected by C. purpurea and producing honeydew
are predominantly visited by two insects, a mycophagous fly, Minettia lupulina (F.)
(Diptera: Lauxaniidae) and a mycophagous beetle, Acylomus sp. (Coleoptera: Phal-
acridae). Because healthy panicles of the grass do not produce nectar or other rewards,
it is unknown what causes insects to disperse the pathogen to uninfected individuals.

The beetle visits the infected florets when honeydew is present and sclerotia are
just beginning to form. The beetles lay eggs at that time (Lemon, unpubl. data). Beetle
larvae develop within the sclerotia. During the early stages of sclerotia development,
I found developing beetle larvae, obvious feeding damage, and frass. At the end of
the growing season, I found pupae and adult beetles within mature sclerotia as well.

The purpose of this study was to determine if the beetles or flies could be responsible
for secondary dispersal of the pathogen. I investigated whether or not these two
species carried spores on their bodies, and if so whether or not they transferred viable
spores from their bodies to the surrounding environment.

MATERIALS AND METHODS

To assess whether the flies or beetles were carrying spores, 12 individuals of each
insect were collected in the vicinity of infected florets of F. arundinacea during the
time of honeydew production. To determine whether the flies were carrying spores

1992

NOTES AND COMMENTS

183

on their bodies and/or in their guts, four flies were selected and each placed in a
separate sterile vial to which 1 ml of sterile distilled water was added and then shaken
for 1 minute using a vortex shaker. The number of C. purpurea spores per ml of
water was counted by using a hemacytometer. To assess whether spores were present
in fly guts, the abdomens were pierced, contents were shaken for an additional minute,
and the spores counted. Gut spore content was calculated as the difference in spore
counts before and after piercing. The same procedure was used for beetles. Because
no fungal spores were found on the first four beetles, the remaining eight were also
surveyed.

Since spores were found on the first four flies, the remaining eight were used to
determine whether or not the flies carried and deposited viable spores onto objects
on which they landed. The eight live flies were kept in separate sterilized glass vials
for twenty four hours where they walked about and defecated. After the removal of
the flies, each vial was rinsed with one ml of sterile distilled water. The number of
spores in the rinse water was counted by using a hemacytometer, and one drop of
the rinse water was plated onto each of two potato dextrose agar plates to count
resultant colonies. Voucher specimens of the beetle are deposited at the Florida State
Collection of Arthropods, Gainesville.

RESULTS

None of the twelve beetles surveyed carried C. purpurea spores internally or ex-
ternally. In contrast, of four flies, three carried spores of C. purpurea externally, and
all carried spores internally. The four individuals had a mean of 5.1 x 105 spores/
ml (SD 4.1 x 105 range 0-1.04 x 106) on their body, and 9.1 x 106 spores/ml (SD
2.4 x 106, range 5.3 x 106 — 1.24 x 1 07) in their gut.

When kept in vials, the flies deposited a mean of 2.76 x 106 spores/ml on the vial
walls (SD 2.75 x 106, range 0-8.0 x 1 06). Some of the deposited spores were viable.
Plating of approximately 0.1 ml of the spore suspension on agar yielded a mean of
36.57 colonies/plate (SD 45.81, range 0-100). The number of spores/ml of solution
was not correlated with the number of colonies that developed (Spearmans rank
correlation: r = 0.19 P < 0.33).

DISCUSSION

At Mountain Lake, the fly, M. lupulina may be a spore dispersal vector for C.
purpurea, but the mycophagic beetle, Acylomus sp. was not. This is unusual since
the beetles, but not flies, were breeding in the fungus and, presumably, might be
more specific to the fungus than are the flies. The flies carried spores on their body,
and in their gut, and transferred the spores to their surroundings. It seems likely that
flies transmit spores to receptive grass stigmas by touching them with their bodies
or by detecating on them. It was not unusual for some florets on the panicle to be
producing honeydew at the same time that other florets on the same panicle were
producing stigmas. I observed flies walking around on grass panicles which were
producing honeydew and receptive stigmas at the same time. It is likely that the
severity of infection on an individual panicle is caused by flies visiting diseased florets
and moving spores to healthy florets on the same panicle. I also observed flies moving
from diseased panicles to different healthy panicles with receptive stigmas. Flies are

184

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(1)

probably moving spores from diseased plants to healthy plants by landing on them
when searching for spore-filled honey dew in the diseased plant population.— Kathleen
M. Lemon, The Nature Conservancy, 1815 North Lynn St., Arlington, Virginia 22209.

ACKNOWLEDGMENTS

Thanks to Carol Horvitz, David Janos, and Helen Alexander who made useful comments
on the manuscript. D. A. Grimaldi identified the fly and M. C. Thomas identified the beetle.
This work was supported by the Pratt Predoctoral Fellowship from Mountain Lake Biological
Station, University of Virginia, and by the Jerry Curtis Plant Sciences Scholarship from the
Department of Biology of the University of Miami.

LITERATURE CITED

A’ Brook, J. 1973. The effect of plant spacing on the number of aphids trapped over cocksfoot
and kale crops. Ann. Appl. Biol. 74:279-285.

Burdon, J. J. 1987. Disease and Plant Population Biology. Cambridge University Press,
Cambridge.

DeNoog, M. P. 1988. The role of weevils in the infection process of the fungus Phomopsis
subordinaria in Plantago lanceolata. Oikos 52:51-58.

Loveless, A. R. 1971. Conidial evidence for host restriction in Claviceps purpurea. Trans.
Brit. Mycol. Soc. 56:419-434.

Lutrell, E. S. 1980. Host parasite relationships and development of the ergot sclerotium in
Claviceps purpurea. Can. J. Bot. 58:942-958.

Mower, R. L. and J. G. Hancock. 1975. Mechanisms of honeydew formation by Claviceps
species. Can. J. Bot. 53:2828-2834.

Schemske, D. W. and C. C. Horvitz. 1984. Variation among visitors in pollination ability: a
precondition for mutualism specialization. Science 225:519-521.

J. New York Entomol. Soc. 1 00( 1 ): 1 84, 1992

A NEW HOMONYM IN THE MEZIRINAE
(HEMIPTERA: ARADIDAE)

In 1967 1 established a new genus, Argocoris Kormilev, for a new described species,
Argoeoris grossi Kormilev, from Queensland, Australia (Kormilev, 1967, Rec. S.
Austral. Mus. 15:519). I have been advised that Argocoris Kormilev, 1967, is a junior
homonym of Argocoris Mayr, 1 864, Hemiptera: Pentatomidae (Verh. Zool. Bot. Ges.,
Wien, 14:905) which was created for Argocoris redtenbacheri Mayr, 1864.

To rectify this homonym, I therefore now propose the replacement name of Pseu-
doargocoris Kormilev, new generic name, for Argocoris Kormilev, 19 67 .—Nicholas
A. Kormilev, 211 Pasadena Avenue N., Apt. 312, St. Petersburg, Florida 33710.

ACKNOWLEDGMENT

I thank Dr. R. C. Froeschner (Smithsonian Institution, Washington, D.C.) for bringing this
situation to my attention.

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Journal of the

New York Entomological Society

VOLUME 100 JANUARY 1992 NO. 1

CONTENTS

Key and diagnoses for the genera of Ochlerini (Hemiptera: Pentatomidae: Dis-
cocephalinae) L. H. Rolston 1—41

Revision of the genus Thvanta St&l, 1862 (Heteroptera: Pentatomidae) II. North
America, Central America, and the West Indies

D. A. Rider and Joan B. Chapin 42-98

Revision of Arocera Spinola, with the description of two new species (Heteroptera:

Pentatomidae) D. A. Rider 99-136

Description of the male of Oxyporus bal/i Campbell, with notes on distribution
of Oxyporus in Mexico (Coleoptera: Staphylinidae)

Jose Luis Navarrete-Heredia 137-141

Review of the species of Telephanus Erichson from the Malagasy region, with
description of a new species (Coleoptera: Silvanidae) M. C. Thomas 142-154

Hydroptilidae (Trichoptera) of Costa Rica: The genus Oxyethira Eaton

Ralph W. Holzenthal and Steven C. Harris 155-177

On the spider subfamily Zavattaricinae (Araneae, Gnaphosidae)

Norman I. Plat nick 178-179

Notes and Comments

The identity and synonymy of Hydroessa fusca Germar (Veliidae: Heteroptera)

John T. Polhemus 180-181

Dispersal of the ergot fungus Claviceps purpurea by the lauxaniid fly Minettia
lupulina Kathleen M. Lemon 182-184

A new homonym in the Mezirinae (Hemiptera: Aradidae)

Nicholas A. Kormilev 184

APRIL 1992

No. 2

Vol. 100

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY

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CONTRIBUTIONS FROM FORMER STUDENTS
IN HONOR OF THE 80TH BIRTHDAY OF

PROFESSOR JOHN G. FRANCLEMONT

Organized by
James K. Liebherr
and

E. Richard Hoebeke

John G. Franclemont, Angie and Duffy (NY: Tompkins Co., Ithaca, Snyder Heights, 13-X-
1991, D. C. Ferguson).

J. New York Entomol. Soc. 1 00(2): 1 85—1 90, 1992

A REMEMBRANCE OF JOHN G. FRANCLEMONT

Ronald W. Hodges

Systematic Entomology Laboratory, ARS, USDA,

% National Museum of Natural History, MRC-168,
Washington, D.C. 20560

John G. Franclemont, Jack to friends and colleagues, is a keen student of the
Lepidoptera and a superb field naturalist. He was bom on 1 5 April 1912 and raised
in Buffalo, New York. He began collecting moths and butterflies at a very early age
and was well instructed in the fine points of spreading specimens by an uncle. Summer
vacations were spent on a farm where Jack’s natural history interests and skills were
further developed. Then, for several summers he was a nature counsellor at a Boy
Scout camp in the Adirondack Mountains. By the time Jack entered Cornell Uni-
versity he knew that he wanted to become a specialist in the Lepidoptera, but he
studied broadly and became well grounded in history, literature, and natural history
generally with special emphasis on insects, plants, birds, and mammals. Soon, he
met professor William T. M. Forbes, then the dean of Lepidopterists in North Amer-
ica, and the two became close associates. Jack Franclemont, Karl Krombein (also
from Buffalo and now a Senior Scientist of the Department of Entomology, Smith-
sonian Institution), Laurence Rupert (a high school mathematics teacher in Horse-
heads, NY), and Ross Arnett were close friends during his undergraduate years. Many
of Jack’s fellow students also have became well-known entomologists including Ross
Arnett, Jr., John Belkin, William Bodenstein, Eugene Gerberg, Karl Krombein,
Woodrow Middlekauff, Eugene Munroe, L. L. Pechuman, Harvey Scudder, and
Henry Townes. He received a BS degree in 1935 and was awarded the Ph.D. degree
in 1953, both from Cornell University.

When WWII broke out, Jack entered the Army Medical Corps as an enlisted man.
Subsequently, he went through officers’ candidate school to become a commissioned
officer. He served as a mosquito specialist in the Solomon Islands in the South Pacific
and in the Philippines. Sometimes his unit arrived on islands while the Marines were
still endeavoring to secure them. He collected intensively on Guadalcanal, Bougain-
ville, New Georgia, and at Baguio in the Philippine Islands. He was moved westward
from island to island and was stationed briefly in Japan at the end of the war. He
was honorably discharged in San Francisco with the rank of Captain.

For a brief time Jack was an assistant entomologist ( 1 946-47) at Cornell University.
Then, he was an entomologist with the Bureau of Entomology and Plant Quarantine
(the predecessor of the Systematic Entomology Laboratory), U.S. Department of
Agriculture at the Smithsonian Institution from 1947 to 1953 where he was respon-
sible for research, curation, and identification of noctuoid and geometroid moths. In
1953 he became an associate professor of entomology at Cornell University, in 1959
full professor, and in 1982 emeritus. He continued to teach the advanced insect
systematics course in Lepidoptera and to advise graduate students for several years
after retirement.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

During Jack’s professional career he was the acknowledged authority on noctuoid
moths of the Western Hemisphere. His extensive bibliography demonstrates the
breadth of his interests, particularly the clarification of many difficult species-group
problems and recognition of closely similar taxa.

His personal library was developed as a hobby and an investment. It has become
one of the best privately held libraries in systematic entomology, and it is far superior
to those in most universities. Naturally, the major emphasis is the Lepidoptera, but
it is strong in several other areas. He was actively accumulating rare books and
journals while many were still available and relatively affordable. It has been given
to Cornell University, but he has lifetime use of it.

Building a personal collection of moths, particularly of the macrolepidoptera, has
been a driving force during most of Jack’s life. Although collecting the macrolepi-
doptera of the United States became a monumental endeavor, he never neglected
the local fauna. This interest became ever more encompassing over the years and
has resulted in the amassing of probably the finest collection of moths ever assembled
through personal fieldwork, spreading, and curation. The level of quality in spread
specimens and genital preparations set new standards, which, through his students,
will influence North American museum collections well into the future. The material
he provided for the teaching collection for the introductory insect systematics course
was far more extensive and better prepared than the main collections of most uni-
versities, and typically was destroyed systematically by budding systematists. While
he was a student and for the first few years in the 1950’s he collected assiduously a
locality on upper Six Mile Creek in the Ithaca watershed area, almost always accom-
panied by his dog. This area may be the best sampled locality for macrolepidoptera
in the United States. He collected for several years at Arlington, Virginia and full
summers in the following areas: Lakehurst, New Jersey; Highlands, North Carolina;
Madera Canyon, Santa Rita Mountains (four years), Flagstaff (three years), and Chir-
icahua Mountains (three years), Arizona; and Big Timber, Montana. He has given
most of his estimated 350,000 specimen collection to the Department of Entomology,
Cornell University where it is well housed and readily accessible to workers. He gave
the Pyraloidea and microlepidoptera to the National Museum of Natural History,
Smithsonian Institution.

As well as collecting adults he has reared many broods of larvae. He established
or verified hosts and preserved larvae of several hundred species. During collecting
seasons one or more glass containers each with a single gravid adult female moth
might be seen in his lab. Often viable eggs were laid to be followed with the provision
of an array of suspected plant host leaves for the newly hatched larvae to try. The
guessing game always began with knowledge of associated species’ larval food plants.
Success in finding an acceptable host plant was reasonably frequent, but the inclem-
ency and relatively short growing season of the Ithaca area sometimes cut off the
food supply before a brood had finished eating or the reverse when a southern species
was captured on a spring field trip. Rearing larvae was a prominent part of all field
work and often was one of the duties of a series of graduate student-field assistants.
Many students have been field assistants and have gained immeasurably from their
experiences.

Throughout his career Jack’s interest in moths and many other insects, excellent
memory, knowledge of the literature, true interest in students, and fine sense of

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JOHN G. FRANCLEMONT

187

humor have served as a foil for many developing scientists. For most of his years at
Cornell his schedule varied little: at the office shortly after eight, lunch in the office
dinner at home, then back to the office for the evening— an exception being the
inevitable collecting on good evenings. The routine changed somewhat on Saturday
and Sunday, but he was at the office even then. Any of his students and many others
felt free to enter his office at any time to discuss a problem, seek advice, or talk.
When a prominent visitor was in town. Jack usually arranged to have a social evening
to which the graduate students and others were invited. Among many, I vividly
remember one evening when Vladimir Nabokov was the guest. We students were
and remain Jack’s extended family. Among his Ph.D. students are George Ball (1954),
Charles Triplehom (1957), Donald Anderson (1958), Ronald Hodges (1961), Carroll
Knowlton (1961), Feliciano Calora (1962), Donald Davis (1962), James Marshall
(1964), Robert Dagleish (1967), Douglas Ferguson (1967), Jay Shaffer (1967), Robert
Poole (1969), George Godfrey (1970), Ring Carde (1971), David Furth (1976), Ali
A. Ahmadi (1977), Timothy McCabe (1978), Frank Ramburg (1979), Richard Brown
(1980), and John Rawlins (1982). Those for whom he chaired a master’s degree
committee or was a minor advisor for the doctoral degree are William Stockton,
Mercedes Delfinado, Oliver Flint, Stuart Neff, Benjamin Foote, Lloyd Knutson,
Albertus Bratt, Siu-Lam Lee, Thomas Pliske, Thomas Wood, Robert Dietz, Jay
Abercrombie, Arthur Shapiro, Andre Comeau, Jorma Tahvanainen, Kuo-Ching Feng,
John Burton, James Liebherr, Frank Slansky, Stanley Tuckwell, J. Mark Scriber,
Ronald Kugler, Mark Lacey, Judith Abrams, Daniel Bickel, Stuart Krasnoff, and
James Miller.

Jack was a true mentor to his students and thus provided guidance, challenges,
and encouragement. He never expected them to agree with him but rather relished
the differences and anticipated independence of thought. Much of his teaching and
perhaps that for which he is best remembered was on a one-on-one basis. He worked
best with self-motivated individuals. He effectively developed a sense of self confi-
dence in his students that has helped them immensely. Because there was so much
personal interaction between Jack and students, they have many cherished memories
that illumine his personality. Above all, Jack is highly ethical, honest, positively
forthright, helpful, humorous in a gentle, nondestructive way, and caring. All his
students benefited from these qualities.

BIBLIOGRAPHY

1. 1936. A new generic name (Lepidoptera, Euchromiidae). Entomol. News 47:275.

2. 1937. Descriptions of new genera (Lepidoptera, Noctuidae, Cuculliinae). Can. Entomol.
69:127-130.

3. 1938. Descriptions of new melanic forms (Lepidoptera, Geometridae, Noctuidae, and
Arctiidae). Entomol. News 49:108-114.

4. 1939a. The usage of the names Epizeuxis Hiibner and Zanclognatha Lederer (Lepidoptera,
Phalaenidae, Herminiinae). Can. Entomol. 71:113-116.

5. 1939b. A revision of the American species of the genus Enargia Hiibner (Lepidoptera,
Phalaenidae, Amphipyrinae). Can. Entomol. 71:113-116.

6. 1939c. Comachara cadburyi, a new genus and species (Lepidoptera, Phalaenidae, Sar-
rothripinae). Entomol. News 50:216-218.

188

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

7. 1941a. Gluphisia lintneri Grote and related species (Lepidoptera, Notodontidae). Can.
Entomol. 73:92-94.

8. 1941b. Some new noctuids from New York State with notes on other well known species
(Lepidoptera, Phalaenidae, Amphipyrinae). Can. Entomol. 73:1 1 1-1 14.

9. 1941 c. The pulverulenta group of the genus Pseudanarta Grote (Lepidoptera, Phalaenidae,
Cucculliinae). Can. Entomol. 73:132-136.

10. 194 Id. Notes on some Cuculliinae (Lepidoptera, Phalaenidae) I. Entomol. News 52:201-
205.

1 1. 1942a. Notes on some Cuculliinae (Lepidoptera, Phalaenidae) II. Entomol. News 53:31 —
35, 63-66.

12. 1942b. [The genus Antaea.] In: W. T. M. Forbes (ed.), Lepidoptera of Barro Colorado
Island. Bull. Mus. Comp. Zool. 90:281-285.

1 3. 1 943. Notes on some Cuculliinae (Lepidoptera, Phalaenidae) III. Entomol. News 54:92-97.

14. [1947] 1946. A revision of the species of Symmerista Hiibner known to occur north of
the Mexican border (Lepidoptera, Notodontidae). Can. Entomol. 78:96-103.

15. 1948. [The genus] Symmerista. In: W. T. M. Forbes (ed.), The Lepidoptera of New York
and neighboring states, part II. Cornell Univ. Agric. Exp. Sta. Mem. 274:222-224.

16. 1949a. The Lepidoptera of New York and neighboring states, part II. [Book review.]
Lepidopterists’ News 3:5-6.

17. 1949b. Synonymical notes relating to Menopsimus caducus Dyar (Lepidoptera, Phalaen-
idae). Proc. Entomol. Soc. Washington 51:74-75.

18. 1 949c. The occurrence of Anomis commoda Butler in the United States and its life history.
Bull. Brooklyn Entomol. Soc. 44:69-71.

19. 1949c. A new moth on coconut from Cuba, with descriptions of new genera for related
species (Lepidoptera, Phalaenidae). Proc. Entomol. Soc. Washington 51:279-285.

20. 1950a. A new moth from Patagonia (Lepidoptera, Phalaenidae). Proc. Entomol. Soc.
Washington 52:40-41.

21. 1950b. The Linnaen subgeneric names of Phalaena (Lepidoptera, Heterocera). J. New
York Entomol. Soc. 58:41-53.

22. 1950c. Proposed use of the Plenary Powers to validate as subgeneric names as from
Linnaeus, 1758, certain terms published for groups of species within the genus “ Phalaena ”
Linnaeus, 1758 (Class Insecta, order Lepidoptera). Bull. Zool. Nomencl. 6:304-312.

23. 1 950d. On the identity of Therina fervidaria Hiibner (Lepidoptera, Geometridae, Ennom-
inae). Bull. Brooklyn Entomol. Soc. 45:290.

24. 1950e. On the types of two genera in the Lepidoptera (Arctiidae and Drepaniidae). The
Entomologist 83:199-200.

25. 1950f. A new generic name (Lepidoptera, Phalaenidae, Acontiinae). Proc. Entomol. Soc.
Washington 52:271-272.

26. 1 950g. Notes on some genera and species of eastern moths with descriptions of new species
(Lepidoptera, Phalaenidae). Bull. Brooklyn Entomol. Soc. 45:144-155.

27. 1951a. Correction on Hiibner’s Florida. Lepidopterists’ News 5:6.

28. 1951b. The species of the Leucania unipuncta group, with a discussion of the generic
names for the various segregates of Leucania in North America (Lepidoptera, Phalaenidae,
Hadeninae). Proc. Entomol. Soc. Washington 53:57-85.

29. Field, W. D., J. F. G. Clarke and J. G. Franclemont. 1951. On a recent proposal to correct
an error committed by the International Commission on Zoological Nomenclature at the
Paris 1948 meeting. Science 113:68-70.

30. 1952. Four undescribed moths from eastern North America, with notes on closely related
species (Lepidoptera, Phalaenidae olim Noctuidae). Bull. Brooklyn Entomol. Soc. 47:123-
137.

3 1 . 1954a. A new species of Copablepharon (Lepidoptera, Noctuidae, Noctuinae). Bull. Brook-
lyn Entomol. Soc. 49:25-27.

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JOHN G. FRANCLEMONT

189

32. 1954b. The identify of Homoptera lineosa Walk. (Lepidoptera, Noctuidae, Catocalinae).
Bull. Brooklyn Entomol. Soc. 49:120.

33. Forbes, W. T. M. and J. G. Franclemont. 1954. Subfamily 3. Cuculliinae. In: W. T. M.
Forbes (ed.), Lepidoptera of New York and neighboring states, part III. Cornell Univ.
Agric. Exp. Sta. Mem. 329:1 16-164.

34. 1955. A new species of Abagrotis (Lepidoptera, Noctuidae). Bull. Brooklyn Entomol. Soc.
50:44-46.

35. 1956. Grant no. 1891 (1955). Taxonomic and biological studies of the noctuoid and
geometroid moths of the New Jersey pine barrens. Year Book Amer. Phil. Soc. 1956:206-
207.

36. 1957. The genus Euclidia, with the description of a new species (Lepidoptera, Noctuidae,
Catocalinae). Bull. Brooklyn Entomol. Soc. 52:5-15.

37. Forbes, W. T. M. and J. G. Franclemont. (1958) 1957. The striated band (Lepidoptera,
chiefly Arctiidae). Lepidopterists’ News 1 1:147-150.

38. 1959. Grant no. 2391 (1958). Taxonomic and biological studies on the moths of the
southern Appalachians. Year Book Amer. Phil. Soc. 1959:231-233.

39. Amadon, D. and J. G. Franclemont. 1960a. Proposals concerning homonymous family
group names .... Bull. Zool. Nomencl. 17:220-223.

40. 1960b. Family 52. Nolidae. In: W. T. M. Forbes (ed.), The Lepidoptera of New York and
neighboring states, part IV. Cornell Univ. Agric. Exp. Sta. Mem. 371:50-56.

41. 1961. Grant no. 303— Johnson Fund (1960). Lepidoptera Heterocera (moths) of southern
Arizona. Year Book Amer. Phil. Soc. 1961:292-293.

42. 1964a. Zale perculta species nov. (Insecta: Lepidoptera: Noctuidae: Catocalinae). Pilot
Register of Zoology, card 1 .

43. 1964b. Notes and two new genera (Lepidoptera: Noctuidae). Proc. Entomol. Soc. Wash-
ington 66:71-73.

44. 1964c. Grant no. 3339 — Penrose Fund (1963). Lepidoptera Heterocera (moths) of south-
eastern Arizona. Year Book Amer. Phil. Soc. 1964:266-267.

45. 1966a. Two new species of Arctiidae from southern Arizona (Lepidoptera, Arctiidae,
Arctiinae). Proc. Entomol. Soc. Washington 68:250-257.

46. 1966b. A new genus, Nychioptera, with descriptions of two new species and notes on the
genus Hemeroplanis (Lepidoptera: Noctuidae). Proc. Entomol. Soc. Washington 68:295-
306.

47. 1967a. Identity of Ceratonyx satanaria, and the larva and pupa of C. arizonensis (Geo-
metridae, Ennominae). J. Lepidopterists’ Soc. 21:11-14.

48. 1 967b. Two new species of noctuids (Lepidoptera, Noctuidae, Noctuinae). Proc. Entomol.
Soc. Washington 69:97-102.

49. 1968. A new species of Metaxaglaea (Lepidoptera, Noctuidae, Cuculliinae). Entomol.
News 79:57-63.

50. 1969a. Two new species of Lithophane from California (Noctuidae, Cuculliinae). J. Lep-
idopterists’ Soc. 23:10-14.

51. 1969b. William Trowbridge Merrifield Forbes [obituary]. Entomol. News 80:88.

52. 1969c. The generic name Callarctia and its type (Lepidoptera: Arctiidae). Proc. Entomol.
Soc. Washington 71:256.

53. Eisner, T. et al. 1970. Defense mechanisms of arthropods. XXVII. Osmeterial secretions
ofpapilionid caterpillars ( Baronia , Papilio, Eurytides). Ann. Entomol. Soc. Amer. 63:914-
915.

54. 1972a. The generic names Salia and Colobochyla of Hiibner (Lepidoptera: Noctuidae).
Proc. Entomol. Soc. Washington 74:75-77.

55. 1972b. Notes on species of North American Leucania with the description of a new species
(Lepidoptera: Noctuidae: Hadeninae). Proc. Entomol. Soc. Washington 74:141-147.

190

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

56. Franclemont, J. G. and R. W. Poole. 1972. A new genus, Stenaspilatodes (Lepidoptera:
Geometridae: Ennominae). Proc. Entomol. Soc. Washington 74:31-34.

57. 1973a. A new noctuid from Arizona (Lepidoptera: Noctuidae: Cuculliinae). Proc. Entomol.
Soc. Washington 75:172-176.

58. 1973b. Mimallonoidea, Mimallonidae and Bombycoidea, Apatelodidae, Bombycidae, and
Lasiocampidae. In: R. B. Dominick et al. (eds.), The Moths of America North of Mexico.
20. 1 : [i]— viii + 1-86, pi. 1-11.

59. 1976. New species of Arizona noctuids, I. (Lepidoptera: Noctuidae: Hadeninae). Proc.
Entomol. Soc. Washington 78:409-416.

60. 1980. “ Noctua c-nigrum ” in eastern North America, the description of two new species
of Xestia Hiibner (Lepidoptera: Noctuidae: Noctuinae). Proc. Entomol. Soc. Washington
82:576-586.

61. 1981a. The identity of Mamestra passa and Morrisonia peracuta of Morrison (Lepidoptera:
Noctuidae: Hadeninae). Proc. Entomol. Soc. Washington 83:133-136.

62. 1981b. The identity of Noctua marginata Fabricius, 1775, with a note on Schinia mar-
ginata (Haworth, 1809) (Lepidoptera: Noctuidae: Catocalinae and Heliothinae). Proc. En-
tomol. Soc. Washington 83:175-176.

63. Blanchard, A. and J. G. Franclemont. 1981. Charadra ingenua Smith in West Texas
(Lepidoptera: Noctuidae: Pantheinae). Proc. Entomol. Soc. Washington 83:797-798.

64. Blanchard, A. and J. G. Franclemont. 1982a. A new species of Zale Hiibner from Texas
and New Mexico (Lepidoptera: Noctuidae: Catocalinae). Proc. Entomol. Soc. Washington
84:134-137.

65. Blanchard, A. and J. G. Franclemont. 1982b. Marilopteryx carancahua a new genus and
new species from east Texas (Lepidoptera: Noctuidae: Hadeninae). Proc. Entomol. Soc.
Washington 84:270-276.

66. Lafontaine, J. D., J. G. Franclemont and D. C. Ferguson. 1982. Classification and life
history of Acsala anomala (Arctiidae: Lithosiinae). J. Lepidopterists’ Soc. 36:218-226.

67. 1983. Mimallonidae, Apatelodidae, Bombycidae, Notodontidae, Dioptidae, Arctiidae. In:
R. W. Hodges, et al. (ed.), Check List of the Lepidoptera of America North of Mexico. E.
W. Classey Ltd. and the Wedge Entomological Research Foundation, London, pp. 107-
108, 112-119.

68. Franclemont, J. G. and E. L. Todd. 1983. Noctuidae (exclusive of Papaipema and Euxoa).
In: R. W. Hodges, et al. (ed.), Check List of the Lepidoptera of America North of Mexico.
E. W. Classey Ltd. and the Wedge Entomological Research Foundation, London, pp. 120-
159.

69. 1 985a. A new species of Psammathodoxa Dyar from Costa Rica (Lepidoptera: Noctuidae:
Catocalinae). Proc. Entomol. Soc. Washington 87:746-750.

70. 1 985b. A new species of Parascotia with notes on the genera Mycterophora and Parascotia
(Lepidoptera: Noctuidae: incertae sedis). Proc. Entomol. Soc. Washington 87:826-833.

7 1 . 1985c. A new species of Meganola Dyar from eastern North America (Lepidoptera: Noc-
tuidae: Nolinae). Proc. Entomol. Soc. Washington 87:871-874.

72. 1986a. New species of tortricid moths from eastern North America (Lepidoptera: Tor-
tricidae). Proc. Entomol. Soc. Washington 88:56-62.

73. 1986b. New species and new genera of noctuids from Arizona, Texas, and Mexico: the
genera Matigramma and Acritogramrpa (Lepidoptera: Noctuidae: Catocalinae). J. Kansas
Entomol. Soc. 59:143-172.

74. Metzler, E. H. and J. G. Franclemont. 1991. A review of four species names of Paectes
from North America (Noctuidae: Euteliinae). J. Lepidopterists’ Soc. 45:34-41.

J. New York Entomol. Soc. 100(2): 19 1-202, 1992

NEOTROPICAL MICROLEPIDOPTERA XXV, A NEW
ARBOREAL SPECIES OF ACROLOPHUS FROM
PERU (LEPIDOPTERA: ACROLOPHIDAE)

Donald R. Davis and Charles L. Hogue

Department of Entomology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, and
Entomology Section, Natural History Museum of Los Angeles County,

Los Angeles, California 90007

Abstract. — A new species, Acrolophus arboreus, of the endemic American family Acrolophidae
is described from the upper Amazon of Peru. Similar larval material from Manaus, Brazil,
suggests the species to be widespread through the Amazon Basin. The larvae are arboreal and
construct elongate tubes of silk and wood fragments on the bark of several unrelated tree species.
The larvae are believed to be scavengers on lichens and similar plant material.

Little is known about the larval biology for the approximately 265 described species
of the endemic American moth family Acrolophidae. Consequently, one can only
speculate as to how diverse their life histories might be (Davis et al., 1986; Davis,
1 987; Davis and Milstrey, 1 988; Davis, 1 990). It is generally believed that acrolophid
larvae develop predominantly within a terrestrial habitat. The larvae of Acrolophus,
for example, typically construct long silken, subterranean tubes amongst plant debris
and roots upon which they feed (Hasbrouck, 1964). The few published records of
arboreal Acrolophus refer to species living in the root system of epiphytic bromeliads
(Picado, 1913; Beutelspacher, 1969).

Field work over the past decade by the junior author (Hogue) has resulted in the
discovery of an even “more arboreal” Acrolophus, whose larvae are believed to be
scavengers on plant material growing on the bark of certain trees. A second arboreal
and undescribed Acrolophus has also been observed in Costa Rica (D. Janzen, pers.
comm.). A description of the Peruvian species is provided below.

Deposition of specimens referred to in this paper are: LACM for the Los Angeles
County Museum of Natural History, Los Angeles, California and USNM for the
National Museum of Natural History, Smithsonian Institution, Washington, D.C.
Abbreviations of morphological terms used for larval description defined in Davis,
1987.

Acrolophus arboreus, new species

ADULT (Figs. 1, 2). Length of forewing: o 12-13 mm; o, 12 mm. Moderately
large moth with uniformly fuscous hindwings and variably marked dark fuscous to
light brown forewings which are paler in the female. Labial palpi elongate in male,
curving, over head to metanotum; much shorter and porrect in female.

Head: Vestiture short, slightly rough over middle (largely obscured by labial palpi
in male); scales moderately slender to piliform with bidentate to acute apices, gen-
erally longer and more slender laterally along eye margin; fuscous with white scale

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Figs. 1-6. Adult morphology, Acrolophus arboreus. 1. Male, length of forewing 11 mm. 2.
Female, length of forewing 12 mm. 3. Basal third of male antenna, ventral view (200 n m). 4.
Dorsal view of Figure 3 (150 /urn). 5. Sensilla coeloconica enlarged from Figure 4 (27 /urn). 6.
Detail of single sensillum coeloconicum (5 yum). (Scale lengths in parentheses; bar scale for all
photographs = Fig. 3.)

1992

NEW ARBOREAL ACROLOPHUS

193

bases in male, mostly white in female. Eye relatively large; interocular index ca. 1.1
mm; cornea with dense covering of interfaceted setae, setae gradually becoming longer
over posterior surface of eye; eyelash absent. Antenna short, less than 0.25 the length
of forewing, 34-35 segmented; male with basal % strongly bipectinate (Figs. 3, 4);
dorsal surface of rami densely covered with sensilla coeloconica (Figs. 5, 6), scape
and pedicel fuscous irroriated with buff and dorsum of flagellum covered with dull
white scales; female antenna simple with dorsum of basal % covered with scattered
fuscous to brown scales, pedicel and scape white. Male with labial palpus elongate,
curving over head to metanotum, fuscous laterally, pale buff to white mesally; labial
palpus of female porrect, much shorter, ca. 1.5 x eye diameter, mostly light brown
laterally, white mesally, with a prominent tuft projecting anteriorly from venter of
second segment.

Thorax: Pronotum with dense mixture of short, moderately broad scales overlaid
by elongate piliform scales, dull white irrorated with gray in male, mostly white
irrorated with buff in female; venter densely covered with mostly white piliform
scales. Forewing of male mostly light to medium fuscous with numerous fine, dark
reticulate spots, a dark fuscous discal streak to base and a small rhomboidal anal
spot; base of anal area distinctly lighter, mostly dull white; cilia brownish fuscous
with a darker terminal line; forewing of female much lighter, light brown with darker
brown reticulations and a much smaller, more slender brownish discal streak; sub-
terminal area distinctly whiter. Hindwing uniformly fuscous in both sexes. Foreleg
dark fuscous dorsally in male and dull white ventrally; light brown dorsally in female,
white ventrally; epiphysis present, elongate, equal in length to foretibia. Midleg and
hindleg progressively paler in both sexes; midtibia with dense brush of white piliform
scales dorsally.

Abdomen: Almost entirely fuscous in male with caudal margins edged with white
ventrally, especially over caudal three segments; generally paler in female, mostly
light brown.

Male genitalia: As shown in Figures 35-38. Uncus moderately short, broad, and
divided about half its length. Gnathos fused, mostly rounded with a slight, median
cleft. Juxta absent. Valva broad, of complex outline, with a prominent, acute, apical
lobe and an elongate, curved, acute saccular lobe closely associated with a membra-
nous inner lobe from mesal membrane of valva. Aedoeagus elongate exceeding length
of genital capsule, of complex form, with a broad median, uncinate comutus and ca.
9 slender comuti along apex of vesica.

Female genitalia: As shown in Figure 39. A single pair of short posterior apophyses
present. Caudal margin of lamella antevaginalis irregularly serrate, well sclerotized.
Ostium broad in diameter, width ca. equal to length of apophyses. Ductus bursae
broad, short, sclerotized. Corpus bursae bilobed, relatively small, length only slightly
exceeding that of ductus bursae; walls completely membranous.

LARVA (Figs. 10-34). Length of largest larva 32 mm; maximum diameter 3 mm.
Body mostly gray dorsally over anterior half, lighter and predominantly cream to
white over remainder of body; thoracic and anal plates and pinacula dark fuscous
to gray.

Head: Dark reddish brown anteriorly, dark fuscous posteriorly. Maximum width
2 mm. AF2 arising at level of apex of frons. P2 more distant from PI than PI is to
ecdysial line. Six pairs of stemmata present (Figs. 15, 28); 3-5 contiguous anteriorly

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;K\

Figs. 7-10. Larval biology of Acrolophus arboreus. 7. Larval cases (see arrows) on tree trunk,
Yanamono, Peru. 8. Underside view of chambered node of larval tube. 9. Larval tube, length
ca. 35 cm (scale = 2 cm). 10. Larva, length ca. 30 mm.

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NEW ARBOREAL ACROLOPHUS

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between SI and SSI; 6 reduced. Antennal sensilla as shown in Figures 29, 30, similar
to that of Acrolophus pholeter Davis (1988) except segment 4 lacking apical extension.
Labrum similar to A. pholeter. Mandible with 2 distinct cusps and two much reduced
cusps; lateral (first) cusp elongate, acute (Fig. 18). Maxilla as shown in Figures 25-
26; all sensilla basiconica (3 + 3 + 2) shorter than those of A. pholeter. ; A2 styloconic;
sensillum digitiformium (SD) greatly reduced, subapical. Spinneret and labial palpus
(Figs. 20, 24) similar to A. pholeter. Apex of mentum with a pair of minute secondary
setae (Fig. 22).

Thorax: Pronotal and mesonotal plates dark reddish brown to nearly black. Spi-
racular plate almost completely separated from pronotal plate by a narrow fissure;
all 3 lateral setae together on spiracular plate. Coxal plates narrowly separated (Fig.
31). Tarsal claw (Fig. 32) with minute, slender axillary seta.

Abdomen: Al-6 with 1 1 pairs of primary setae, SV trisetose. A7-8 with SV3 absent.
A9 with 9 setal pairs, SV unisetose. Ventral crochets in a uniserial ellipse of ap-
proximately 50-53 hooks; anterior and posterior sides of prolegs densely covered
with small scattered spines, more numerous anteriorly. Anal proleg with ca. 28 hooks
in a half ellipse open posteriorly and covered with dense spines anteriorly. A8 with
spiracle greatly enlarged, equalling size of prothoracic spiracle. Anal plate with 4
pairs of elongate setae.

LARVAL CASE (Figs. 7-9). An elongate tube up to 35 cm in length and 5 mm
in diameter, constructed of minute wood fragments held together by silk; a 3 to 6
chambered, oval, abruptly enlarged dome-like section variably situated somewhere
along midlength of tube, up to 40 mm long, 30 mm wide and 15 mm high. Inner
surfaces of larval tube and chamber smoothly lined with thin layer of whitish silk;
cavity devoid of frass or debris. Larval tube extends through chamber intact, without
any openings; partitions of central chamber possibly due to successive enlargement
of chamber. Color light to dark brown.

PUPA (Figs. 40-48). Length of largest pupa: o, 12 mm; o, 12.8 mm. Dark reddish
brown in color. Vertex smooth except for a pair of small setae near middorsal line.
Frons with a small pair of setae near inner margin of eye (Figs. 40, 4 1 , 44). Antenna
of male relatively longer and much broader than in female; labial palpus ca. twice
as long in male than female. Wings extend to middle of A3. Mesonotum with two
pairs of dorsal setae, D1 more anterior and separated further apart than D2. Dorsum
of A3-8 with a low, transverse, minutely and densely spined (Figs. 45, 46) ridge
parallel to anterior margin. A10 a relatively small, acutely furcate cremaster dorsally
(Figs. 47, 48).

HOLOTYPE. 6, Yanamono, 80 km east of Iquitos, Loreto Department, Peru; 3
Jul 1990, emerged 28 Jul 1990, C. L. Hogue (LACM).

PARATYPES. PERU. Loreto Department: Amazon Safari Camp, Rio Momon,
near Iquitos: 1 9, 25 Jun 1980, emerged 12 Aug 1980, C. Hogue, CLH 268.1, genitalia
slide 3669 (LACM); 1 6 pupa, 25 Jun 1980 (LACM). Yanamono, 80 km E Iquitos:
1 6, emerged Jul 1982, C. Hogue (LACM); 1 6, emerged 20 Jul 1983, pupal slide
3653-54 (LACM); 1 <3, emerged 23 Jul 1983, C. Hogue (LACM); 1 <3, emerged 23
Jul 1983, antenna slide 28638 (USNM); 1 <5, emerged 28 Jul 1983, genitalia slide
30926, C. Hogue (USNM). Same data as holotype, 1 <3 (LACM); 20 larvae, 3 Jul
1990, C. Hogue slides 30696-98 (USNM, LACM).

HOST. Unknown; most probably various lichens and other plant debris on the

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Figs. 11-18. Acro/ophus arboreus, larval morphology. 11. Chaetotaxy of pro- and meso-
thorax, abdominal segments, 1, 5, 8, 9. 12. Head, dorsal view (0.5 mm). 13. Head, ventral
view. 14. Abdominal segments 8-10, dorsal. 15. Head, lateral view. 16. Labrum, dorsal (0.5
mm). 17. Labrum, ventral. 18. Mandible (0.5 mm). (Scale lengths in parentheses.)

Figs. 19-26. Acrolophus arboreus, larval morphology. 19. Head, dorsal view (0.6 mm). 20.
Labrum and maxilla (231 Aim). 21. Head, ventral view (0.6 mm). 22. Labium and maxilla,
ventral view (200 /urn). 23. Head, anterior view (0.6 mm). 24. Labial palpi and spinneret (60
Aim). 25. Maxilla (75 nm). 26. Sensilla of maxillary palpus (1 5 /urn). (Scale lengths in parentheses;
bar scale for all photographs = Fig. 19.)

Figs. 27-34. Acrolophus arboreus, larval morphology. 27. Head, lateral view (0.5 mm). 28.
Stemmatal area (231 pm). 29. Antenna, lateral view (23.1 pm). 30. Antenna, distal view (23.1
pm). 31. Thorax, ventral view of T1 and 2 (0.67 mm). 32. Pretarsus, lateral view of T1 (50
jum). 33. Abdominal proleg, A3 (150 pm, A = anterior, L = lateral). 34. Anal proleg (150 pm).
(Scale lengths in parentheses; bar scale for all photographs = Fig. 27.)

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NEW ARBOREAL ACROLOPHUS

199

luM:

Ki W

Figs. 35-42. Acrolophus arboreus, genital morphology and pupae. 35. Male genitalia, ventral
view (0.5 mm). 36. Lateral view. 37. Valva, mesal view. 38. Aedoeagus. 39. Female genitalia,
ventral view (0.5 mm). 40. Male pupa, head. 41. Female pupae, ventral view (0.5 mm). 42.
Dorsal view of Figure 4 1 . (Scale lengths in parentheses.)

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Figs. 43-48. Acrolophus arboreus, pupal morphology. 43. Head, ventral view (0.6 mm). 44.
Frons, with paired setae (see arrows) (0.33 mm). 45. Dorsal view of A6-10 (0.6 mm). 46. Detail
of serrated dorsal ridge of A7 (see r in Fig. 45) (60 jum). 47. Lateral view of A6-10 (0.6 mm).
48. Caudal view of A 10 (0.6 mm). (Scale lengths in parentheses; bar scale for all photographs
- Fig. 43.)

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NEW ARBOREAL ACROLOPHUS

201

bark of several trees including Chimarrhis glabriflora Ducke (Rubiaceae) and Ficus
insipida Willd. (Moraceae).

FLIGHT PERIOD. June-August. Because collecting was conducted only during
this period, which occurs in the middle of the dry season, the maximum flight period
and seasonality of this species are uncertain.

DISTRIBUTION. Adults known only from type locality, an area in the Peruvian
Amazon in the vicinity of Explorama Lodge (=Yanamono), ca. 80 km E Iquitos,
Department of Loreto. Perhaps widely distributed in the Amazon Basin as indicated
by larval cases observed in Manaus, Brazil (see ethnological note).

ETYMOLOGY. The specific name is derived from the Latin arboreus (of trees)
in reference to the unusual, arboreal habit of the larva.

TAXONOMIC DISCUSSION. Currently, Acrolophus arboreus does not appear
closely related to any described species in this large genus. The unusual structure of
the male genitalia, particularly the rather complex form of the valva and thickened
gnathos, show no close affinities to a similarly arboreal and undescribed Acrolophus
from Costa Rica.

Larval chaetotaxy appears to be very conservative within Acrolophus. Possibly the
reduced sixth stemma is the most diagnostic larval character for this species. Among
the few known larval cases, the swollen, centrally located chamber is unique. The
similar case observed in Manaus, Brazil (see ethnological note) probably represents
this species.

BIOLOGICAL OBSERVATIONS. Larval tubes have been observed on the trunks
of several tree species, including two identified by A. Gentry (pers. comm., letter of
31 Aug. 1982 to Hogue) as Chimarrhis glabriflora Ducke and Ficus insipida Willd.
The tubes are located from 3 to 7 m above the ground and normally oriented
vertically. They are easy to see, usually being of slightly different color than the tree
bark. The tubes are weakly attached to the bark surface along their entire length but
adhere more tightly at the upper end, their apparent origin.

The function of the chambered, swollen node located along the middle of the larval
tube was not observed. Possibly the larva uses this as a retreat even though no
openings in the tube were observed within the cup-like chamber. An outer shell
around the tube might provide some added protection. Likewise, larval feeding was
not observed. Possibly the larva gleans lichens or other plant material from either
end of the tube.

ETHNOLOGICAL NOTE. During a visit to the Municipal Market in Manaus,
Brazil, on 25 April 1990, Hogue noticed the larval tubes of A arboreus (or a very
close relative) in a stall where basketry and sundries were sold. Hogue was told by
the vendor that the tubes were called (in Portuguese) “rede de tamaquare,” meaning
“net of the tamaquare lizard.” He believed that this lizard, which is frequently found
climbing on tree trunks in the nearby forest, was responsible for making the tubes.

The tubes were described by the vendor as useful in “black magic,” to influence
the humor of one’s spouse or sweetheart, as follows: “Colocar o nome dentro da
rede, e colocar embaixo do traveseiro.” Put the name [of one’s lover, on a piece of
paper] inside the net and place it under [her] pillow. In the morning she will wake
up in a perfectly friendly mood.

The common name “tamaquare” (syn. “taquare”) in Brazil applies to Enya/ia spp.
(von Ihering 1968:675). A poorly preserved specimen of the alleged net maker shown

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to Hogue by the vendor appeared not to be of this genus but the very common, often
arboreal, Tropidurus torquatus Van Lidth de Jeude, 1917, known in Brazil as “ta-
raquira.” In any case, identification of the lizard is unimportant to the story, the
locals being imprecise taxonomists.

ACKNOWLEDGMENTS

We wish foremost to express our gratitude to John G. Franclemont for the very competent
advice and assistance which he generously provided over the years and to congratulate him for
the numerous accomplishments he achieved throughout his distinguished career, both as en-
tomologist and teacher. We are indebted to Young Sohn of the Department of Entomology,
National Museum of Natural History, for the line drawings and to Susann Braden and Walt
Brown of the Smithsonian SEM Lab and Victor Kranz of the Smithsonian Office of Photographic
Services for photographic assistance. Acknowledgment is made to the University of California
at Los Angeles Extension and Society Expeditions, Seattle, Washington, for financing the trips
on which the material was collected. Thanks also to Tina Ross for preparing the final draft of
the manuscript.

The senior author (Davis) is responsible for the systematic and morphological portions of
this paper and the biological and ethnological observations are by the junior author (Hogue)
who discovered the species. This report constitutes contribution XXV in the Smithsonian
Neotropical Microlepidoptera series.

LITERATURE CITED

Beutelspacher, C. 1969. Una especie nueva de Acrolophus Poey, 18832, de Bromeliaceas
(Lepidoptera: Acrolophidae). An. Inst. Biol. Univ. Nat. Auton. Mexico. 4° Ser. Zool. (1):
43-48.

Davis, D. R. 1987. Tineidae. Pages 362-365 in: F. W. Stehr (ed.), Immature Insects, Vol. 1.
Kendall/Hunt Publishing Co., Dubuque, Iowa.

Davis, D. R. 1 990. Three new species of Acrolophus from the southeastern United States with
remarks on the status of the family Acrolophidae (Lepidoptera: Tineoidea). Proc. En-
tomol. Soc. Wash. 92(4):694-704.

Davis, D. R., D. H. Clayton, D. H. Janzen and A. P. Brooke. 1986. Neotropical Tineidae,
II: Biological notes and descriptions of two new moths phoretic on spiny pocket mice
in Costa Rica (Lepidoptera: Tineoidea). Proc. Entomol. Soc. Wash. 88(1):98-109.

Davis, D. R. and E. G. Milstrey. 1988. Description and biology of Acrolophus pholeter,
(Lepidoptera: Tineidae), a new moth commensal from gopher tortoise burrows in Florida.
Proc. Entomol. Soc. Wash. 90(2): 164-1 78.

Hasbrouck, F. S. 1964. Moths of the family Acrolophidae in America north of Mexico
(Microlepidoptera). Proc. U.S. Nat. Mus. 1 14(3475):487-706.

Picado, C. 1913. Les Bromeliacees epiphytes, considerees comme milieu biologique. Bull.
Scient. Fr. Belg. 47:215-360.

von Ihering, R. 1968. Dicionario dos animais do Brasil (rev. ed.). University of Brasilia, Sao
Paulo, 790 p.

Received 3 July 1991; accepted 8 October 1991.

J. New York Entomol. Soc. 100(2):203-20.8, 1992

TWO NEW SPECIES OF MOMPHA FROM CALIFORNIA
(LEPIDOPTERA: MOMPHIDAE)

Ronald W. Hodges

Systematic Entomology Laboratory, ARS, USD A,

%National Museum of Natural History, MRC-168,

Washington, D.C. 20560

Abstract.— Two new species of Mompha, franclemonti and powelli, from California are de-
scribed to provide names that support long-term biodiversity studies of natural areas in Cali-
fornia. M. powelli was reared from Zauschneria californica (Onagraceae).

Specimens of an undescribed species of Mompha were collected during a long-
term survey of Lepidoptera of Big Creek Reserve, Monterey County (one of the
University of California’s natural land reserves) by Jerry A. Powell (UC Berkeley).
Subsequently, a closely similar, but slightly smaller species of Mompha was collected
in the hills immediately east of San Leandro, Alameda County. The second species
later was reared from Zauschneria californica Presl, Onagraceae. To make a name
available for his planned report and to document some of the diversity within Mom-
pha both species are described.

The new species are black marked with white and are most similar to Mompha
metallifera (Walsingham). In the National Museum of Natural History’s (USNM)
insect collection several, mainly eastern, specimens also are very similar to metal-
lifera, including a series reared from Cuphea viscosissima Jacquin, Lythraceae (as C.
petiolata) by Annette F. Braun and incorrectly determined as metallifera. This species
proved to be undescribed. I have not found any other male specimen whose genitalia
match those of the holotype of metallifera. After dissecting males (females are not
well represented) of most of the miscellaneous specimens, I find that no less than 1 2
species are present. Only three of them have names: metallifera, argentimaculella
(Murtfeldt), and annulata (Braun). Careful study of color pattern leads me to believe
that separation of these species on this basis may be impossible or extremely difficult.
Sexual dichromatism occurs in the series reared by Braun: males have the distal xh
of the antenna uniformly black; females have the distal three segments white preceded
by black segments.

My initial intent was to analyze these similar-appearing species, prepare an iden-
tification key, and illustrate the genitalia of each entity. This proved to be too large
a project for the present purpose and thus is deferred until a thorough study can be
made of nearctic Mompha. My preliminary study of this genus leads me to suspect
that the fauna may be more than 100 species and have several cryptic species’
complexes.

Mompha franclemonti, new species
Figs. 1, 3, 5

Diagnosis. A superficially very dark-gray moth having shining silver transverse
fasciae, one at 2/5, 4/5, and V6 forewing length. All surfaces, except ventral surface of

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Figs. 1-2. Photographs of Mompha species. 1. M. franclemonti, n. sp., holotype <3. 2. M.
powelli, n. sp., holotype <3.

fore- and hindwings and white areas, with shining reflections, usually mainly silvery
but sometimes with yellow or yellow-gray cast. Mompha franclemonti is most similar
to M. powelli but differs as follows: 1) The mesial surface of the labial palpus is paler
than the lateral surface and the pale-gray tipped scales are less contrasting on the
mesial surface than the lateral surface {franclemonti ); the mesial and lateral surface
of the labial palpus are similarly shaded and hued, and the pale-gray to off-white
areas of the scales are large {powelli ). 2) The forewing has many, small white dots
0 franclemonti ); the forewing lacks white dots {powelli). 3) The aedoeagus has a nearly
parallel-margined, very broadly rounded cornutus and a sclerotized band with tooth-
like projections (, franclemonti ); the lateral margins of the cornutus taper within the
distal Vi to a slender, rounded apex {powelli ). 4) The distal lobes of the juxta are
separated mesially by a U-shaped area and appear like a pair of nipples {franclemonti );
the distal lobes of the juxta are separated by a very narrow, V-shaped area, are broad
basally and taper laterally within the distal Vi to a rounded apex {powelli ).

Description. Head: Haustellum very dark gray with a few pale-gray tipped scales
basally, becoming naked and yellowish distally. Labial palpus medium gray mesially,
dark gray laterally, many scales tipped with pale gray on second segment and off-
white on third segment. Frons nearly black with shining metallic silver reflections;
vertex and occiput dark gray with shining metallic silver reflections; scales behind
eye nearly black. Antenna very dark gray, distal segment gray preceded by 1 Vi very
dark-gray segments (more pronounced on ventral surface), then somewhat irregularly
alternate Vi segments gray and very dark gray. Thorax: Tegula very dark gray basally,
gray distally; meso- and metathorax very dark gray. Foreleg mainly very dark gray
becoming darker distally, apex of tibia with a few slightly paler scales, apexes of first

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Figs. 3-4. Male genitalia of Mompha species. 3. M. franclemonti, n. sp., paratype, USNM
genitalia slide 86613. 4. M. powe/li, n. sp., paratype, USNM genitalia slide 8661 1.

four tarsomeres white, apex of fifth tarsomere gray. Midleg as for foreleg but tibia
with a faint, gray transverse fascia at Vi length and prominently white at apex. Hindleg
as for midleg but with prominent, transverse, white fascia at Vi length and spurs
mainly white with some gray areas. Forewing mainly very dark gray with prominent
salt-and-pepper pattern caused by a small white area just before the apex of many
individual scales or at the apex of scales on the distal part of the wing; a patch of
raised scales on the fold at Vi wing length and one on the posterior Vi of the wing
just beyond % wing length, a diffuse zone of silver-colored scales at base of wing, a
silver, transverse fascia at % wing length, another at 4/5 wing length, and a curved

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Figs. 5-6. Female genitalia of Mompha species. 5. M. franclemonti, n. sp., paratype, USNM
genitalia slide 15992. 6. M. powelli, n. sp., paratype, USNM genitalia slide 86612.

silver band on the outer margin of the wing; fringe at apex of wing gray preceded by
a narrow black line, then white dots, then a black area with scattered white dots,
fringe in tomal area medium to pale gray; undersurface different shades of gray;
length 3. 0-4.0 mm. Hindwing medium to pale gray on dorsal and ventral surfaces.
Abdomen: dorsal surface mainly very dark gray, prominently shining silvery gray
laterally and on posterior margin of several segments, mesial parts of terga 3-5 with
yellowish gray to orange gray; ventral surface very dark gray, apex of sterna broadly
shining white or silver white, eighth sternum mainly dark with a few white scales
mesially. Male genitalia (Fig. 3): anterior margin of costal lobe of valva arising from
base of valva, continuing as heavily sclerotized transtilla, maximum width of costal

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207

lobe 0.75 x width of valva at base; transtilla incomplete, not meeting mesially; ae-
doeagus with very broad, distal comutus that is broadly rounded distally and with
a diffuse sclerotized band bearing many denticular projections; juxta with pair of
short lobes distally. Female genitalia (Fig. 5).

Etymology. Named in honor of my long-time friend and mentor, John G. Fran-
clemont.

Holotype. 6, USA. California: Monterey Co.: Big Creek Reserve, UCNLWRS,
mining ridge, 80-180 m, sage chapparal, 21/23 III 1989, F. Hsu, J. Powell; deposited
in the University of California, Berkeley, Entomology Museum.

Paratypes. Same data as holotype, USNM genital slides 15991, 15992, 86613,
RWHodges slide 5140, 13 66, 2 99 (UCB, USNM). Same locality as holotype, 21/22
II 1988, J. A. Powell, 1 3 (UCB).

Mompha powelli, new species
Figs. 2, 4, 6

Diagnosis. Superficially, powelli is similar to franclemonti but differs as indicated
under franclemonti.

Description. Head: Haustellum shining pale gray becoming yellowish gray distally.
Labial palpus medium gray, most scales of second and third segments broadly tipped
with pale gray. Frons and vertex shining silver gray, occiput very dark gray with
shining yellow reflections, row of scales behind eye very dark gray to shining gray.
Antenna very dark gray, most scale bases slightly paler than apexes. Thorax: Tegula,
meso- and metathorax very dark gray. Foreleg mainly very dark gray; coxa promi-
nently shining silver gray; apexes of many scales on femur and tibia pale gray, apex
of epiphysis with a few off-white scales and apex of tibia white; apex of tarsomeres
one, two, and four white. Mid- and hindlegs much as for foreleg but scales of femur
and tibia without pale apexes, tibia with a prominent white fascia at 3/5 length.
Forewing mainly very dark gray, scale bases often slightly paler gray, a diffuse zone
of translucent silver-colored scales at base, a prominent shining silver fascia at 2/5
length and one at % length, a shining silver spot on posterior margin at xh length, a
few silver scales on anterior margin at V2 length, and a band of shining silver scales
paralleling distal margin; a patch of raised scales on the fold at V2 the wing length
and another on the posterior margin at 3/5 length; a few pale-gray tipped scales on
posterior V3 of wing before first silver fascia; fringe at apex of wing gray preceded by
a narrow black line, then white dots, then a black area with scattered white dots,
fringe in tomal area medium to pale gray; undersurface mainly medium to pale gray;
length 2. 7-3. 7 mm. Hindwing medium to pale gray on dorsal and ventral surfaces.
Abdomen: as for franclemonti. Male genitalia (Fig. 4): anterior margin of costal lobe
of valva arising from base of valva, continuing as heavily sclerotized transtilla,
maximum width of costal lobe approximately 0.50 x width of valva at base; transtilla
incomplete, not meeting mesially; aedoeagus with large distal comutus, lateral mar-
gins tapering within distal V2 to slender, rounded apex; lobes of juxta broad, mesial
margins slightly diverging, lateral margins angled mesially at V2 length to rounded
apex. Female genitalia (Fig. 6).

Etymology. Named in honor of the collector and colleague, Jerry A. Powell.

Holotype. 6, USA. California: Alameda Co.: Fairmont Ridge, SE of San Leandro,

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3 V 1989, J. Powell; J. Powell No. 89E6, emgd. 28 v 1989, reared from Zauschneria
calif [ornica\, deposited in the University of California, Berkeley, Entomology Mu-
seum.

Paratypes. Same data as holotype but emerged 24-29 V 1989, USNM genitalia
slide 15990, 4 <3<3, 4 99. Same locality as holotype, 19 iv 1989, J. Powell; J. Powell
No. 89D33, emgd. 9-22 V 1989, reared from Zauschneria calif [ornica], USNM
genitalia slides 15989, 86611, 86612, 5 66, 4 99. Same locality as holotype, 3 V 1 989,
31 VIII 1989, J. Powell, 10 86, 6 99 (UCB, USNM).

Distribution. An additional specimen of powelli was collected at Whittier, Los
Angeles Co., California on 4 III 191 1 by P. H. Timberlake. It is not included in the
type series, but study of the male genitalia (USNM genitalia slide 5324) shows it to
be conspecific.

ACKNOWLEDGMENTS

I thank Jerry A. Powell (University of California, Berkeley; UCB) for allowing me to study
material collected by him; my wife, Elaine R. S. Hodges, for the line drawings of genitalia;
Douglas C. Ferguson for the photographic prints and review of the manuscript; and Don R.
Davis, Richard C. Froeschner, and Arnold S. Menke for review of the manuscript.

Received 11 July 1991; accepted 1 November 1991.

J. New York Entomol. Soc. 100(2):209-227, 1992

SIX NEW SPECIES OF CATASTEGA
(LEPIDOPTERA: TORTRICIDAE: EUCOSMINI) FROM
MEXICO AND SOUTHWESTERN UNITED STATES

Richard L. Brown

Department of Entomology, Drawer EM, Mississippi State, Mississippi 39762

Abstract. — In celebration of the eightieth birthday of Dr. John G. Franclemont, six new
species of Catastega (Lepidoptera: Olethreutinae: Eucosmini) are described as follows: C. spec-
tra, from Texas, C. nebula, from Arizona, C. strigatella, from New Mexico, Texas, and Mexico,
C. triangulana, from Arizona and Mexico, C. adobe, from Arizona, New Mexico, and Texas,
and C. plicata, from Arizona and Mexico.

This paper is dedicated to Dr. John G. Franclemont, because of his direct and
indirect contributions in documenting the Lepidoptera fauna of the southwestern
United States. Dr. Franclemont, assisted by a number of graduate students, spent a
total of 31 months in Arizona during nine extended periods between 1959 and 1974;
these field trips were notable for the intense collecting, e.g., 99 nights out of a possible
100 in 1960, and the resulting large number of undescribed taxa that were collected
(Franclemont, 1961, 1964, 1976). During these trips to Arizona, Dr. Franclemont
collected several species of Olethreutinae in various genera that remain undescribed,
and his graduate students collected three of the following six new species of Catastega
Clemens. Species of Catastega from Texas, New Mexico, and Mexico similar to those
in Arizona also are treated here.

The North American genus Catastega was resurrected from synonymy with Epi-
notia Hubner and characterized by apomorphies of the male anellus and female
sterigma by Brown (1986). At present the genus includes two eastern species, C.
timidella Clemens and C. aceriella Clemens, which have larvae forming serpentine
frass and silk tubes, and one western species, C. marmoreana (Heinrich), for which
hosts and larval habits are unknown. The following six new species indicate that
morphological diversity in this genus is greatest in the southwestern United States
and Mexico.

METHODS AND MATERIAL

A stereomicroscope with an ocular micrometer was used to examine and measure
specimens. The forewing length was measured from the outer edge of the tegula at
wing base to the outermost edge of the fringe scales at apex. Setae and comuti were
counted by examining their sockets with a compound microscope. The “Methuen
Handbook of Colour” (Konerup and Wanscher, 1983) was used as a standard for
describing colors of imagos as observed under an incandescent light source.

Descriptions of the forewing are based on a comparative study of pattern elements
in more than 600 tortricid species representing all tribes as defined by Horak and
Brown (1991). The terminology for pattern elements is based largely on Nijhout

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Fig. 1. Catastega nebula, n. sp., holotype, forewing with pairs of costal strigulae labeled
1-10.

(1978), as modified and discussed by Brown and Powell (1991). The term umbra is
used for narrow, transverse lines of scales in fascial or interfascial areas that contrast
with the paler ground color; the term spot is used for contrasting pigmentation that
is confined to a restricted portion of a fascia (e.g., pretomal spot) or an interfascial
area (e.g., white interfascial spot between subbasal and median fasciae in Figs. 2 and
3). The term patch is used to describe a color field that differs from the ground color
and that covers at least one fascia and one interfascial area. The term stria is used
for narrow, transverse lines of gray (silver when viewed at an appropriate angle to
light) scales that originate at the costal strigulae and extend towards the inner or
outer margin. Costal strigulae, which mark the margins of fasciae and usually occur
in pairs, are numbered 1 to 10 from base to apex (Fig. 1).

Unidentified Olethreutinae and specimens identified as Epinotia, with which Ca-
tastega has been confused, have been borrowed from 27 institutions and several
private collections. Specimens of the following new species of Catastega were bor-
rowed from or are deposited in the collections of the following: Andre Blanchard
(AB, deposited in USNM), American Museum of Natural History, New York (AMNH),
Cornell University Insect Collection (CUIC), Edward C. Knudson, Houston, TX
(ECK), Richard L. Brown, Mississippi State, MS (RLB), University of California,
Berkeley (UCB), University of Connecticut, Storrs (UCONN), U.S. National Museum
of Natural History (USNM).

KEY TO THE SPECIES OF CATASTEGA

1 . Forewing with pale yellow ground color, subbasal fascia forming longitudinally elongate
spot between middle of discal cell and CuP plicata, n. sp.

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- Forewing ground color grayish brown, brownish gray, or brownish red, subbasal fascia

extending to costa or inner margin or not forming contrasting spot 2

2. Forewing with brownish red ground color, without contrasting fasciae adobe, n. sp.

- Forewing ground color not brownish red, with or without contrasting fasciae 3

3. Forewing with termen slanted towards tomus, apex acute, interfascial areas distinctly

strigate strigatella, n. sp.

- Forewing with termen not slanted towards tomus, apex not acute, interfascial areas

not distinctly strigate 4

4. Forewing with brown, triangular, pretomal spot, without other contrasting fasciae . .

triangulana, n. sp.

- Forewing with or without brown, pretornal spot, with other contrasting fasciae 5

5. Forewing with subbasal fascia and pretomal spot not distinct on inner margin, pale

inner marginal half and dark costal half separated by sinuate border . . timidella Clemens

- Forewing with subbasal fascia and pretomal spot distinct on inner margin, inner mar-
ginal half not contrastingly paler than costal half nor separated by sinuate border 6

6. Ocellar area of forewing with well separated silver striae enclosing large group of
peppered scales; male costal fold present or absent; occurring west of a line between
San Antonio, Texas, and Winnipeg, Manitoba 7

- Ocellar area of forewing with approximate silver striae not enclosing large group of

peppered scales, male costal fold absent; occurring east of a line between San Antonio,
Texas, and Winnipeg, Manitoba aceriella Clemens

7. Forewing with pretomal spot contrastingly darker than median fascia on costa, without

white interfascial spot on inner margin, male costal fold present; female sterigma
surrounded by sternum VII marmoreana (Heinrich)

- Forewing with pretomal spot usually not darker than median fascia on costa, with or

without white interfascial spot, male costal fold absent; female sterigma posterior to
sternum VII 8

8. Silver stria between pretomal spot and postmedian fascia with irregular distal margin,

ocellar area with many white scales; male cucullus rounded ventrally; occurring in
Arizona nebula, n. sp.

- Silver stria between pretomal spot and postmedian fascia with even distal margin,
ocellar area with few white scales; male cucullus angled ventrally; occurring in Texas

spectra, n. sp.

Catastega spectra, new species
Figs. 2, 8, 9, 15

Description. Head: vertex grayish brown anteriorly, light grayish brown posteriorly,
posterior scales with apices white to orange gray; labial palpus light grayish brown,
second segment with orange-white or orange-gray scales on ventral base and with
concolorous band on dorsal surface near two-thirds length, band becoming diffuse
laterally and extending towards apex of ventral scale tuft. Thorax: mesonotum light
grayish brown with white-tipped scales, with dark grayish-brown, transverse band
at middle and concolorous semicircular band on posterior lobe, semicircular band
approximate to transverse band at midline, bands diffuse and less distinct in some
specimens; tegula usually dark grayish brown basally, light grayish brown or light
grayish orange intermixed with yellowish white apically, most scales with white tips,
some specimens with small, brownish orange or grayish orange spot surrounded by
dark grayish brown at base. Forewing (Fig. 2): length 7.0-7. 7 mm in males, 6. 7-7. 7
mm in females; male costal fold absent; termen concave, with indistinct white strigula

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Figs. 2-7. Catastega n. spp. imagos, all photographed at same magnification. 2. C. spectra,
male Mt. Locke, Texas. 3. C. nebula, male, Chiricahua Mts., Arizona. 4. C. strigatella, female,
Guadalupe Mts., Texas. 5. C. triangulana, female, Santa Rita Mts., Arizona. 6. C. adobe,
holotype, male, Chiricahua Mts., Arizona. 7. C. p/icata, male, Santa Rita Mts., Arizona.

between R5 and M,; costa with eight pairs of white to yellowish white strigulae
apparent, pairs 1-4, 9, and 10 separate, 5 approximate to 6 and 7 approximate to 8
with each of two pairs appearing as single pair, strigulae separated by dark grayish-
brown fasciae and umbrae; ground color light grayish brown, most scales with white
to yellowish white tips, some specimens with large, white (as in Fig. 2) or small,
brownish orange interfascial spot on inner margin; fasciae incomplete, dark grayish
brown, broken transversely to form umbrae or longitudinally to form spots; basal
fascia represented by narrow umbrae and isolated, small groups of scales; subbasal
fascia represented by narrow, isolated umbra on costa and large, subquadrate spot
on inner margin, outer margin of spot slanted toward wing apex, some specimens
with subbasal spot confluent with median fascia at midwing; median fascia narrow
on costa between strigulae 4 and 5, becoming broader at midwing, broken by ground
color or white and one or two, small, transverse, gray (silver) bars at CuA2 to form
triangular pretornal spot, spot enclosing 2-3, small, transverse, gray (silver) bars near
inner margin and 2-3, small, longitudinal dark brown bars near midwing, median
fascia with scales more peppered with yellow white apices than scales in subbasal

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fascia; postmedian fascia narrow on costa between strigulae 6 and 7-8 and extending
toward termen, angled towards tornus and widened near R5 to form spot enclosing
2-3 small groups of dark-brown scales, fascia enclosing small, gray (silver) bar at
tornus, scales between R5 and tornus more peppered with white apices than scales
in median fascia, postmedian fascia bordered distally and proximally by wide, gray
(silver) striae, proximal stria with distal margin even and not broken by scales of
postmedian fascia; grayish brown scales in costal and terminal areas apical to strigulae
7-8 with orange color when viewed at angle to light; outer margin with bases of
attenuate fringe scales white to orange white, forming terminal line from apex to
midway between CuA, and CuA2; fringe between apex and tornus with proximal
scales dark grayish brown with white apices, distal scales light brownish gray, without
white apices except at terminal strigula between R5 and Mt. Underside brown except
for yellowish white costal strigulae 3-10, scales in terminal area between strigulae 9
and tornus with white tips. Hindwing: upperside uniformly light brown; underside
light grayish brown with distinct ripples of yellowish white on costa and indistinct
ripples on terminal area between M, and CuA^

Male genitalia (Figs. 8, 9): tegumen moderately broad, densely setose posterolateral-
ly, moderately setose dorsomedially; uncus with forks well separated at their bases,
forks sharply bent posteriorly at near midlength, each fork densely setose dorsally
from near midlength to below its base; socius subquadrate, dorsoapically rounded,
ventroapically fusing with base ofgnathos, densely setose laterally from dorsal margin
to base of gnathos except for smooth, rimlike, apical margin, vertical length of setose
area at apical margin 0.50-0.53 x the length of straight line distance from ventral
base of tegumen to base of uncus forks; aedeagus with 18 cornuti; anellus cuplike,
not closely surrounding aedeagus, dorsal side elongate, length subequal to that of
aedeagus, medially cleft; valva with elongate cluster of spiniform setae on sacculus,
costa not indented distal to costal hook, dorsal and ventral margins of valva sub-
parallel at middle third, cucullus elongate, apical margin forming obtuse angle with
ventral margin of valva (two preparations examined).

Female genitalia (Fig. 15): sternum VII with scales sparse laterally, dense postero-
medially, with a few rugae posterolaterally; tergum VIII with sparse setae and scales
scattered on lateral extensions, each extension lateral to bases of apophyses with less
than 20 scales and setae, dorsum with sparse setae; papillae anales with lateral and
medial margins of each ventral surface subparallel, not expanded posteriorly, densely
setose, some lateral setae with well-developed, papillose bases, rugae absent; sterigma
conical, lamella antevaginalis well developed, without rugae, lamella postvaginalis
reduced to narrow rim, posterolateral angles acute, setae absent; ductus bursae en-
circled by colliculum, moderately long on one side, sinewy, ductus expanded ante-
riorly before corpus bursae; signa subequal in size or with one signum smaller (three
preparations examined).

Holotype. Male, labeled “McDonald Obser., Mt. Locke, Tex., 1 9-VII- 1971, A.
Blanchard.” The type locality at McDonald Observatory is located 6,828 ft above
sea level in Jeff Davis Co. The holotype is in excellent condition and is deposited in
the U.S. National Museum of Natural History. Of the variation within the species,
the holotype has an interfascial area on the inner margin between the subbasal and
median fasciae that is light brownish gray, enclosing a small brownish orange spot
(as in Fig. 1).

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Fig. 8. Catastega spectra, n. sp., male genitalia, USNM slide 16026, Mt. Locke, Texas. Scale
= 1 mm.

Paratypes. United States. Texas: same data as holotype, 1 1 males, genitalia slides
USNM 16026, 17771, 4 females, genitalia slides USNM 17649, 17753, 17754 (RLB,
USNM).

Diagnosis. Among previously described Catastega, C. spectra is most similar in
genitalia to C. aceriella and C. timidella. The new species is superficially distinct by
the light brown hindwing, the more contrasting subbasal and median fasciae, and
the contrasting lightly pigmented area between these two fasciae on the inner margin.
Distinctive characters of male genitalia include an uncus with forks that are well
separated at their bases and that are bent posteriorly, an anellus with the dorsal plate
subequal in length with the aedeagus, and a valva with the medial third having
subparallel dorsal and ventral sides and with the apical margin of the cucullus forming
an obtuse angle with the ventral margin. The female genitalia of the three species
are similar in form, but the new species differs in having a wider tergum VIII, narrower
papillae anales, a longer, smooth lamella antevaginalis that ventrally covers the
ostium, and narrower signa. Catastega spectra is very similar in forewing pattern and
genitalia to a new species from Arizona, which is differentiated in the following
description.

Catastega nebula, new species
Figs. 1, 3, 10, 16

Description. Head and thorax: as in C. spectra. Forewing (Figs. 1 , 3): as in C. spectra
except length 7. 7-8. 5 mm in males, 7. 8-8. 6 mm in females; costa with 9 pairs of
strigulae apparent, with pairs 5 and 6 separate and distinct; median fascia more
narrow and less defined between costa and midwing; postmedian fascia between
midwing and tornus with white scales intermixed with brown and dark grayish brown

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Figs. 9-14. Catastega n. spp., male genitalia, all photographed at same magnification. 9. C.
spectra, USNM slide 16026. 10. C. nebula, USNM slide 17738. 11. C. strigatella, AB slide
3765 (in USNM). 12. C. triangulana, holotype, USNM slide 28866. 13. C. adobe, USNM slide
17778. 14. C. plicata, holotype, USNM slide 17769.

scales with white tips, proximal margin of fascia and distal margin of gray stria
irregular, with gray stria often broken by white or white-tipped scales of fascia.
Hindwing: underside light grayish brown with indistinct ripples of yellowish white
on costa and without ripples on terminal area between M, and CuA,.

Male genitalia (Fig. 1 0): as in C. spectra except uncus with forks curved posteriorly,
not strongly bent, dorsum of each fork setose on basal one-fourth; socius with apical
margin of lateral surface densely setose, vertical length of setose area at apical margin
of socius 0.37-0.40 x the length of straight line distance from ventral base of tegumen
to base of uncus forks, medial surface with smooth rim on apical margin; aedeagus
with 22-30 comuti; valva with base of costa deeply indented distal to costal hook,
cucullus rounded ventrally (five preparations examined).

Female genitalia (Fig. 16): as in C. spectra except sternum VII of some specimens

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with rugae lateral to midline from anterior to posterior margins; tergum VIII with
setae and scales concentrated on posterior margins of lateral extensions, each exten-
sion lateral to bases of apophyses with more than 30 scales and setae; papillae anales
with each ventral face narrow anteriorly, expanding posteriorly, lateral margin rugose;
lamella antevaginalis with longitudinal rugae (three preparations examined).

Holotype. Male, labeled “Rustler Park 8500', Chiricahua Mts., Ariz. VII- 14-72,
at light, J. Powell, male genitalia slide 310 R.L. Brown.” The type locality is in
Cochise Co., Arizona. The holotype is deposited at University of California, Berkeley.

Paratypes. United States. Arizona: Cochise Co.: same data as holotype, 7 males,
genitalia slides JAP 3629, RLB 1768, 3 females, genitalia slide RLB 311; same data
except 3 July 1972, 1 male, RLB photo 159 on file, 12 July 1972, 2 females, genitalia
slide RLB 621; Cave Creek, 1 mi SW Portal, 24-29 July 1972, J. Powell, 1 male,
genitalia slide JAP 3626; Huachuca Mts., Carr Canyon, 5,300 ft, 29 July 1986, D.
L. Wagner, 1 female. Santa Cruz Co.: Santa Rita Mts., Madera Canyon, 5,100 ft,
Bog Springs Campground, July 10-26 1964, D. R. Davis, 2 males, genitalia slides
USNM 17738, 28867; Madera Canyon, 4,880 ft, 25 Jul. 1959, R. W. Hodges, 1
female, genitalia slide RLB 382, 13 Aug. 1959, 1 female, genitalia slide RLB 383
(deposited in CU, RLB, UCB, UCONN, USNM).

Diagnosis. Differences in wing pattern between C. spectra and C. nebula are subtle
but are consistent among specimens examined. The separation of costal strigulae 5
and 6 and the irregular distal margin of the gray (silver) stria between the pretornal
spot and postmedian fascia are the most distinctive forewing characters for distin-
guishing C. nebula. The postmedian fascia in C. nebula has many white scales in the
area of the ocellus, and these are lacking in C. spectra ; however, the degree of
pigmentation of the fasciae is a variable character in some species of Catastega. The
setation of the apical margin of the lateral surface of the socius, in contrast to a
smooth margin in related species, is a result of the socius being apically inflected
inwardly, such that the medial surface is also setose apically. In addition, the setose
area of the socius is vertically shorter in C. nebula than in C. spectra, and this is
independent of the difference in the size of body, as indicated by forewing lengths.

Catastega strigatella, new species
Figs. 4, 11, 17

Description. Head: vertex and labial palpus brownish gray, scales with white to
light gray apices. Thorax: mesonotum brownish gray with two, transverse, dark
brownish gray bands on posterior half, with area between bands and posterior end
light gray to white; tegulae basally dark brownish gray, apically light brownish gray
with increasing amount of white to light gray on apices of scales. Forewing (Fig. 4):
length 8. 2-9. 3 mm in males, 8. 3-9. 7 mm in females; male costal fold present, length
of fold 0.39-0.41 x forewing length; termen straight, slanted towards tomus to pro-
duce acute angle at apex, with light gray strigula between R5 and M,; costa with nine
pairs of light gray strigulae apparent, pairs 1 and 2 separate, poorly expressed, 3 and
4 approximate, appearing as two pairs, 5 and 6 separate, 7 and 8 confluent, appearing
as one pair, 9 and 10 separate; ground color light brownish gray, some specimens
light gray on costal half of wing, apices of scales varying in amount of light gray;
fasciae usually incomplete, dark grayish brown; basal fascia indistinct except for

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Figs. 15-16. Catastega n. spp., female genitalia. 15. C. spectra, USNM slide 17753. 16. C.
nebula, RLB slide 621 (in UCB). Scale = 1 mm.

distal margin, extending as umbra from near costa to near inner margin; subbasal
fascia complete from costa to inner margin in some specimens, outwardly angled
near midwing, usually broken transversely at midwing or near CuP to form con-
trasting subquadrate spot on inner margin, proximal and distal margins of spot often
dark grayish brown and enclosing brownish gray scales with gray apices, proximal

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margin of spot extending as umbra from inner margin to CuP, distal margin extending
as umbra from inner margin to CuP or midwing, margins of fascia usually distinct
from costa to radius; median fascia broken into costal spot extending from costa to
radius, medial spot extending from midwing to CuA2, and triangular pretomal spot
extending from inner margin to CuA, when most expressed, proximal and distal
margins and central areas of spots concolorous with those of subbasal inner marginal
spot, costal spot usually indistinct, often with only margins expressed, medial spot
wide at midwing, becoming narrow toward CuA2, spot crossed by oblique, dark,
brownish gray line, usually with one or two umbrae between CuA2 and inner margin;
postmedial fascia represented by narrow umbra extending from costa to M3, often
broken near R3 and appearing connected with interfascial umbrae between strigulae
7 and 8 or with umbra of subterminal fascia between strigulae 8 and 9, ocellus absent,
area between M3 and CuA2 (area of ocellus in related species) with two, short, ap-
proximate, gray striae appearing only slightly silver when viewed at angle to light;
subterminal fascia reduced to umbra, broken near M3, appearing to originate from
small spot between costal strigulae 9 and 10, extending to near CuA,, subparallel
with postmedial umbra, area between these umbrae and between radius and midwing
forming large spot (or patch from elements of postmedial and subterminal fasciae),
concolorous with median and subbasal spots; terminal fascia represented at wing
apex as small spot concolorous with other spots; outer margin without terminal line;
fringe with each scale white to light gray at extreme apex, sharply contrasting with
dark brownish gray at subapex, gradually becoming lighter brownish gray to gray
from subapex toward base, proximal scales with irregular length and with white
apices of scales not forming lines. Underside light grayish brown, costa with strigulae
4-10 distinct. Hindwing: upperside uniformly light grayish brown; underside light
brownish gray, costal area with ripples of orange gray.

Male genitalia (Fig. 1 1): tegumen moderately broad, moderately setose postero-
laterally, bare or with sparse setae dorsomedially; uncus with forks asetose, approx-
imate and slender basally, gradually curved posteriorly, base of uncus with group of
dense setae below each fork; socius strongly rounded dorsoapically, gently rounded
ventrally to base of gnathos, lateral surface with apical third densely setose from
dorsal margin to near base of gnathos, medial surface with smooth rim on apical
margin; gnathos with base prolonged, arising from ventral one-third of vinculum;
aedeagus with more than 3 1 cornuti; anellus cuplike, not closely surrounding aede-
agus, dorsal side elongate, length about two-thirds that of aedeagus, dorsomedial
margin concave; caulis short; valvae with small, elongate cluster of spiniform setae
on ventroapical margin of sacculus, neck between cucullus and sacculus moderately
constricted (four preparations examined).

Female genitalia (Fig. 1 7): sternum VII moderately setose laterally and anteriorly,
densely setose posteromedially, rugose laterally; tergum VIII with apices of lateral
extensions moderately scaled (20-25 each side) and sparsely setose (3-5 each side),
dorsum sparsely setose; papillae anales with lateral margins evenly rounded, ventral
face slightly broadest at one-third distance from posterior end, densely setose, lateral
setae with moderately large, papillose bases, rugae absent; sterigma reduced, lamella
antevaginalis forming a smooth, narrow rim around ostium, subequal in length with
and superimposed on sclerotized posteroventral margin of ostium, lamella postvag-
inalis absent or reduced to very narrow portion of posterodorsal sclerotized margin

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Figs. 17-18. Catastega n. spp., female genitalia. 17. C. strigatella, RLB slide 902 (in RLB).
18. C. triangulana, USNM slide 17688. Scale = 1 mm.

of ostium, setae absent; ductus bursae encircled by long, sinewy colliculum; signa
subequal in size (three preparations examined).

Holotype. Male, labeled “Brewster Co., Tex., Big Bend Nat. Park, Chisos Basin,
29-III-82, Leg. E.C. Knudson.” The specimen is in excellent condition and is de-
posited in the U.S. National Museum of Natural History.

Paratypes. Mexico. Coahuila: 17 mi SE Saltillo, 6,800 ft, 23 Sept. 1976, J. A.
Chemsak & J. Powell, 1 male, genitalia slide JAP 5013, 1 female. United States. New
Mexico: Otero Co.: 2 mi NE Cloudcroft, 8,600 ft, 1 Jul. 1964, F., P., & M. Rindge,
1 male. Socorro Co.: 28 mi SW Magdalena, 8,500 ft, 21 Jul. 1964, F., P., & M.

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Rindge, 1 female, 22 Jul. 1964, 3 females, genitalia slide RLB 903. Texas: Brewster
Co.: same data as holotype, 3 males, genitalia slides RLB 1 147, ECK 287, 6 females,
genitalia slide RLB 1 148; Big Bend Nat. Park, Green Gulch, 31 Mar. 1970, A.&M.E.
Blanchard, 1 male, genitalia slide AB 3765, 1 female, genitalia slide RLB 902, 3 Apr.
1984, E. C. Knudson, 1 male, 5,400', 2-4 Apr. 1986, E. C. Knudson, 1 male. Cul-
berson Co.: Guadalupe Mts., McKittrick Canyon, 5,000 ft, 23 May 1973, R. W.
Hodges, 1 male, genitalia slide USNM 17654; Guadalupe Mts, Smith Canyon, 22
May 1973, A.&M.E. Blanchard, 1 female, RLB photo 51 on file; Guadalupe Mts.
N.P., Frijole, 29-31 Mar. 1990, E. C. Knudson, 1 male, same data except Ship on
the Desert, 1 male, 1 female. Jeff Davis Co.: Davis Mts. St. Pk., 5 Jun. 1986, E. C.
Knudson, 1 female (deposited in AMNH, ECK, RLB, UCB, USNM).

Diagnosis. The strigate interfascial areas, fascial pattern, and the slanting termen
of the forewing differentiate this species from other Catastega. This species also
differs from others in having a dark longitudinal line crossing the medial spot of the
median fascia and a subterminal umbra that is broken from its origin between
strigulae 7 and 8 and is reconnected with the interfascial umbra originating from
between strigulae 9 and 10. The male genitalia are unique in having a socius that is
ventrally lengthened and tapered into the long base of the gnathos. The valva has a
well-defined neck separating the cucullus from the sacculus, similar to C. marmoreana
and the following three new species.

Catastega triangulana, new species
Figs. 5, 12, 18

Description. Head: vertex and labial palpus light grayish brown, posterior scales
of vertex with light gray apices. Thorax: mesonotum grayish brown to light brown,
scales with light gray apices, light gray increasing posteriorly; tegulae light brown,
some specimens with apices of posterior scales light gray. Forewing (Fig. 5): length
6. 7-7. 3 mm in males, 7. 3-7.9 mm in females; male costal fold present, length of fold
0.38-0.41 x forewing length; termen straight, not slanted toward tornus, without
distinct strigulae, some specimens with lighter fringe scales between apex and R5,
between R5 and M„ and on each side of CuA2; costa with five pairs of light brownish
gray strigulae apparent, strigulae less distinct in some specimens, pairs 5 and 6
separate, 7 and 8 confluent, appearing as one pair, 9 and 10 separate; ground color
brownish gray, specimens varying in number of scales with light-gray apices; fasciae
as in C. strigatella except only pretomal triangle on inner margin well expressed,
brown with dark brown margins or uniformly dark brown; costal two thirds from
wing base to outer margin of median fascia forming indistinct grayish brown patch,
slightly darker than ground color; subbasal fascia expressed as two dark brown umbrae
from CuP to inner margin; postmedian fascia represented by dark brown umbra
extending from between costal strigulae 6 and 7 to tornus, dentate and broken in
area of ocellus between two, short gray (silver) striae; subterminal fascia extending
from between costal strigulae 8 and 9 to near CuA1? subparallel with postmedian
umbra, some specimens with subterminal and postmedian umbrae connected by
short, longitudinal line near M2; terminal fascia indistinct. Underside grayish brown,
costa with strigulae 5-10 distinct in some specimens. Hindwing: upperside uniformly

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light grayish brown; underside light brownish gray, some specimens with costa orange
gray, without ripples.

Male genitalia (Fig. 12): as in C. strigatella except uncus with forks less slender
basally, setose to midlength; socius strongly rounded dorsoapically and ventroapi-
cally, apical margin concave; gnathos arising from ventral base of socius at middle
third of tegumen; aedeagus with approximately 20 comuti; valva with sacculus gently
rounded apically, neck and cucullus broad (two preparations examined).

Female genitalia (Fig. 1 8): as in C. strigatella except sternum with rugae very fine,
becoming stronger anteriorly; tergum VIII with lateral extensions reduced to narrow
bars, densely scaled and sparsely setose, dorsum without setae, anterior margin with
two triangular extensions; sterigma less reduced, lamella antevaginalis forming a
smooth, moderately broad rim around ostium, longer than sclerotized posteroventral
margin of ostium, lamella postvaginalis moderately long, posterolateral angles with
or without single seta; colliculum of ductus bursae long, not sinewy (two preparations
examined).

Holotype. Male, labeled “4 mi ESE Pine, Gila Co. Ariz. 5400 ft, 1 September 1961,
Ronald W. Hodges. Genitalia slide by R.L. Brown, USNM 28866.” The holotype,
which is missing the left mesothoracic leg, is deposited in the U.S. National Museum
of Natural History.

Paratypes. United States. Arizona: Cochise Co.: Huachuca Mtns., Ash Canyon,
27-31 Jul. 1986, D. L. Wagner, 1 male; 2 mi SW Portal, 2-5 Aug. 1974, J. Powell,

1 male, genitalia slide RLB 1 324. Gila Co.: same data as holotype, 1 female, genitalia
slide USNM 17688. Pima Co.: Santa Rita Mtns., Madera Canyon, 4,400 ft, 10 Oct.
1959, R. W. Hodges, 1 female, genitalia slide RLB 297. Santa Cruz Co.: Santa Rita
Mtns., Madera Canyon, 4,880 ft, 30 Jul. 1959, R. W. Hodges, 1 female, 5,600 ft, 17
Oct. 1959, 1 female (deposited in CU, RLB, UCB, UCONN, USNM).

Other material examined. The following specimens are tentatively identified as C.
triangulana because of their similar forewing pattern and male genitalia: Mexico.
Durango: 28 mi E El Salto, 8,000 ft, 22 Jul. 1964, [beating] Arctostaphylos sp., J.
Powell, 3 males, same data except J. A. Chemsak & J. Powell, black & white lights,
8 males, genitalia slides JAP 1787, RLB 1423; 30 mi W Durango, 8,500 ft, 31 Jul.
1964, J. Powell, 2 males, 8,400 ft, 3-7 Aug. 1972, D. Veirs, J. Powell, C. D. MacNeill,

2 males, 1 female, 3-8 Aug. 1972, J. Powell & D. Viers, 1 male, 8,400 ft, Tepalcates,
4-8 Aug. 1972, 1 male, genitalia slide RLB 1334; 9 mi E La Ciudad, 9,000 ft, 23
Jul. 1964, J. Powell, 1 male. Oaxaca: 7 mi SE Nochixtlan, 7,000 ft, 7 Oct. 1975, J.
Powell & J. Chemsak, 1 male.

These specimens (deposited in UCB) are not included in the type series nor in the
description because they represent either a wide range of variation in C. triangulana
or a distinct species. Unfortunately, both series are small, and the series from Mexico
does not include females. The specimens from Mexico are larger, having a male
forewing length of 7. 8-9. 2 mm, and the fore wings appear slightly darker than spec-
imens from Arizona. The forewing pattern of specimens from Mexico differs in having
a more sinuate line of separation between the costal patch, extending from wing base
to median fascia, and the lighter inner margin. This pattern difference, which falls
within the range of variation of some Catastega species, is a result of the subbasal
and median fasciae extending closer to the inner margin in specimens from Mexico
than in those from Arizona. Male genitalia of specimens from Mexico, based on

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examination of three slide preparations, differ in having a slightly more narrow,
elongate, and apically constricted cucullus than those from Arizona; however, these
may be allometric differences.

Diagnosis. This species and the following new species, C. adobe, both have the
triangular pretornal spot as the most contrasting fascial element of the forewing
pattern, which is unique among North American Catastega. The color and pattern
of C. triangulana, however, is most similar to that of the C. strigatel/a. Catastega
triangulana differs in its lack of expression of strigulae and fasciae and in having a
suffusion of pigment in the interfascial areas between the basal, subbasal, and median
fasciae; both subbasal and median fasciae are broken near the same vein in both
species but are more expressed on the inner margin in C. strigatella. The male and
female genitalia of C. triangulana are most similar to those of C. adobe; males of
the two species differ slightly in shape of the valvae, and females differ in development
of the sterigma, being more reduced in the latter species.

Catastega adobe, new species
Figs. 6, 13, 19

Description. Head: vertex and frons yellowish white, becoming yellowish gray
posterolaterally; labial palpus yellowish white with light brownish-gray band near
apex of second segment. Thorax: mesonotum yellowish red anteriorly with tips of
scales yellowish white, becoming brownish red posteriorly; tegulae brownish red.
Forewing (Fig. 6): length 5. 7-6. 5 mm in males, 6. 3-6.9 mm in females; male costal
fold present, length of fold 0.33-0.35 x forewing length; termen slightly concave,
strigulae absent; costa with five pairs of orange white strigulae apparent, strigulae 5,
6, 9, and 10 separate, strigulae 7 and 8 confluent, appearing as single pair; ground
color brownish red; fasciae absent except for pretornal spot; outer two-thirds peppered
by variable number of brown and grayish brown scales with white tips, some spec-
imens more peppered with brown scales on median fold, pretornal area, area between
ocellus and R4, and wing margins; pretornal spot brownish red suffused with variable
number of peppered brown scales; ocellus weakly expressed, orange white, bordered
by gray (silver) striae proximally and distally; outer margin with orange white terminal
line formed from apices of distal row of disc scales and bases of attenuate fringe
scales; fringe scales between apex and tomus with white apices, appearing peppered,
proximal fringe brown, distal fringe grayish brown. Underside brown except for
orange white costa and strigulae. Hindwing: light brown dorsally, light brown to
grayish brown ventrally.

Male genitalia (Fig. 13): tegumen narrow ventrally, becoming moderately wide
dorsally, sparsely setose laterally, bare dorsomedially; uncus moderately setose from
base to midlength of forks; socius sub-triangular, expanded and bulbous dorsobasally,
apical margin relatively straight from base of uncus to projecting ventral angle,
irregularly serrate at high magnification, lateroapical half and medioapical third
setose; gnathos arising from ventral bases of socii, well sclerotized basally; aedeagus
with 10-18 comuti; anellus not closely surrounding aedeagus, forming moderately
shallow cup with long dorsal side, length less than half that of aedeagus; valva with
sacculus and cucullus separated by neck without setae, sacculus with elongate, poorly

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NEW CATASTEGA

223

Figs. 19-20. Catastega n. spp., female genitalia. 19. C. adobe, USNM slide 17658. 20. C.
plicata, USNM slide 17739. Scale = 1 mm.

defined cluster of spiniform setae on ventral margin, ventroapical comer strongly
rounded, not overlapping base of neck (three preparations examined).

Female genitalia (Fig. 19): sternum VII with moderately dense scales throughout,
more dense posteromedially, slightly rugose anteriorly; tergum VIII with lateral ex-
tensions reduced to narrow triangles, apices of lateral extensions setose and scaled,
bare dorsally, anterior margin with two, short triangular projections; papillae anales
densely setose, setae with small, papillose bases, rugae absent; sterigma with cone
reduced, lamella antevaginalis narrow, lamella postvaginalis with 0-2 setae poster-

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

olaterally; colliculum of ductus bursae moderately sclerotized, long on one side, not
encircling ductus; signa subequal in size and shape (three preparations examined).

Holotype. Male, labeled “ Ariz . Cochise Co., Portal, 4 mi. W., 5300'; Chiricahua
Mts., Aug. 3-6, 1964, D.R. Davis. Genitalia slide by R.L. Brown, USNM 17639.
Photograph on file, 135 R.L. Brown.” The holotype is in excellent condition and is
deposited in the U.S. National Museum of Natural History.

Paratypes. United States. Arizona: Cochise Co.: same data as holotype, 3 males,
genitalia slides USNM 17770, 17778, 1 female, genitalia slide USNM 17658. New
Mexico: Lincoln Co.: Capitan Mts., near Nogal, 1 Aug. 1989, Wagner & Epstein, 1
male, 1 female. McKinley Co.: Ft. Wingate, Jul. 16-23 [year and collector unknown],
1 female, July 24-31, 1 female. Socorro Co.: 28 mi SW Magdalena, Bear Trap Camp,
6,500', July 6 1965, F., P. & M. Rindge, 1 female, July 7 1965, 1 male. Texas:
Culberson Co.: Guadalupe Mts. N. P., McKittrick Canyon, 30-3 1 July 1989, Wagner,
Epstein, & Knudson, 1 male, Dog Canyon, 6,400', 19-20 Aug. 1990, E. C. Knudson,
4 females; Sierra Diablo W.M.A., 27 Jun. 1981, E. C. Knudson, 1 male. Jeff Davis
Co.: Mt. Locke, 27 May 1979, E.C. Knudson, 1 male (deposited in AMNH, ECK,
RLB, UCONN, USNM).

Diagnosis. Catastega adobe differs from other Catastega and most other Oleth-
reutinae in having brownish red forewings without contrasting fasciae (except for the
triangular pretomal spot, which is derived from median fascia). The form of the
socius in the male and the form of sterigma with reduced lamella antevaginalis and
developed lamella postvaginalis in the female is diagnostic for this species.

Catastega plicata, new species
Figs. 7, 14, 20

Description. Head: vertex pale yellow to yellowish white; labial palpus yellowish
white, intermixed with grayish brown except ventrobasally and at two-thirds length.
Thorax: mesonotum yellowish white to pale yellow anteriorly, becoming pale orange
to grayish orange posteriorly; tegulae pale orange intermixed with grayish orange
basally, becoming yellowish white apically. Forewing (Fig. 7): length 5. 8-7. 6 mm in
males (5. 8-6. 8 mm in Arizona specimens; 6. 0-7. 6 mm in Mexico specimens), 5.8-
8.3 mm in females (5. 8-7.0 mm in Arizona specimens; 6. 7-8. 3 mm in Mexico
specimens); male costal fold present, length of fold 0.56-0.62 x forewing length;
termen slightly concave, strigulae absent; costa in female with six pairs of yellowish
white to orange white strigulae apparent, pairs 3, 4, 9 and 10 separate, pairs 5 and
6 and pairs 7 and 8 confluent, each confluent pair appearing as single pair, males as
in females except with pairs 1-4 yellowish gray, with individual strigulae widely
spaced on costal fold, strigulae 5-8 in both sexes with striae concolorous with ground
color, indistinct except shiny when viewed at angle to light, strigulae 9 with stria
gray (silver) from R4 to R5, strigulae 10 with short gray (silver) stria extending to R5
or near termen; ground color pale yellow intermixed with variable amounts of yel-
lowish white and pale orange, suffused with reddish orange to brownish orange on
costal area between median fascia and apex and terminal area between apex and
tornus; fasciae brown intermixed with variable amounts of dark brown and grayish
orange; basal fascia absent; subbasal fascia forming longitudinally elongate spot be-
tween middle of discal cell and CuP, some specimens with fascia expressed between

1992

NEW CATASTEGA

225

costa and radius and/or with suffusion of grayish orange between CuP and inner
margin; median fascia brown or brown intermixed with grayish orange, usually ex-
tending from costa to between CuA, and CuA2, rarely complete, some specimens
with separate, sub-triangular pretornal spot, some specimens with dark brown, lon-
gitudinal streak on accessory cell, inner margin of fascia extending basally near costa
beyond apices of strigulae 4; postmedian fascia expressed as large, rounded spot,
dark brown intermixed with varying amounts of brown and/or grayish orange, sep-
arated from median fascia by ground color or suffusion of grayish orange; subterminal
and terminal fascia not expressed; outer margin bordered by white to yellow white
terminal line formed from bases of proximal attenuate fringe scales, pre-apical area
of proximal fringe scales forming reddish orange line from apex to tornus, becoming
more irregular towards latter, distal fringe scales uniformly grayish orange. Underside
light grayish brown to grayish orange, costal area rippled by yellowish white to orange
white strigulae. Hindwing: upperside yellowish gray to yellowish white, underside
yellowish white.

Male genitalia (Fig. 14): tegumen narrow ventrally, becoming expanded at bases
of socii, moderately setose laterally, sparsely setose dorsomedially; uncus setose from
base to one-third length of forks, apices of forks curved medially; socius dorsobasally
expanded, bulbous, apical margin gently rounded dorsoapically and tapered ventrally
into bases of gnathos, irregularly serrate at high magnification, lateroapical half and
medioapical third setose; gnathos arising from base of ventral socius; aedeagus with
28-32 comuti; anellus appearing relatively reduced, sclerotized portion forming shal-
low cup, not closely surrounding aedeagus, dorsal and ventral sides of sclerotized
portion subequal in length, dorsal side with lightly sclerotized extension reaching to
two-thirds length of aedeagus; valva with sacculus and cucullus separated by neck
without setae, sacculus with poorly defined cluster of spiniform setae on ventral
margin, ventroapical corner strongly rounded, slightly overlapping base of neck (four
preparations examined).

Female genitalia (Fig. 20): sternum VII with moderately dense scales throughout,
more dense posteromedially, rugose medially, except for midline, from posterior to
anterior margins; ovipositor (segment VIII and posteriorly) modified for insertion;
tergum VIII with setae much longer than setae on papillae anales, sparse and random
on well developed, triangular, lateral extensions, moderately dense in irregular row
on dorsum, scales absent, anterior margin with long, rounded projection; papillae
anales fused and attenuate posteriorly, setae moderately dense, bases not on papillae,
setose surface wide laterally, narrow ventrally; posterior and anterior apophyses thick
and wide with expanded bases; sterigma skewed, lamella antevaginalis well devel-
oped, smooth, lamella postvaginalis reduced to narrow rim, without posterolateral
angles and setae; colliculum of ductus bursae moderately sclerotized, sinewy, long
on one side; signa subequal in size and shape (four preparations examined).

Holotype. Female, labeled “ Ariz . Santa Cruz Co., Madera Canyon 5100', Santa
Rita Mts., July 10-26, 1964, D.R. Davis; Bog Spring Camp Ground; Genitalia Slide
by R.L. Brown USNM 17769.” The holotype is in excellent condition, except for
missing the right prothoracic leg, and is deposited in the U.S. National Museum of
Natural History.

Paratypes. Mexico. Durango: 3 mi E Revolcaderos, 11 Aug. 1972, J. Powell, 1
male; Durango, Arroyo los Mimbres, 2,200 m, 19 Aug. 1984, E. Welling, 1 male, 3

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

females; 10 mi W El Salto, 8,800 ft, 23 Jul. 1964, J. A. Chemsak & J. Powell, 1
female; 31 mi E El Salto, 18 Jul. 1964, J. A. Chemsak, 1 female, genitalia slide RLB
1413; 30 mi W Durango, 5 Aug. 1972, J. Powell, 1 male, 8,500 ft, 31 Jul. 1964, J.
A. Chemsak & J. Powell, 1 male, 2 females, 8,400 ft, 3-8 Aug. 1972, J. Powell, D.
Veirs, & C. D. MacNeill, 25 males, genitalia slide JAP 3734, 7 females, Tepalcates,
8,400 ft, 4-8 Jul. 1972, 6 males, genitalia slide RLB 1412, 1 female, same data
without elevation, 11 Aug. 1986, Brown & Powell, 2 males, 2 females. Sinaloa: 8
mi W El Palmito 6,400 ft, 8-12 Aug. 1972, J. Powell, D. Veirs, & C. D. MacNeill,
1 male, 2 females, 6 Aug. 1986, J. Brown & J. Powell, 1 male. United States. Arizona:
Cochise Co.: Huachuca Mts., Ash Canyon, 27-31 Jul. 1986, D. L. Wagner, 1 male,
Carr Canyon, 5,300 ft, 29 Jul. 1986, D. L. Wagner or Wagner and Powell, 4 females.
Pima Co.: Madera Canyon, Bog Springs, 30 Jul. 1973, J. Powell, 1 male. Santa Cruz
Co.: same data as holotype, 1 male, 2 females, genitalia slide USNM 17739, same
data plus “from flowers Yucca schottii 1 male; Madera Canyon, 4,880 ft, 19 Jul.
1959, R. W. Hodges, 1 male, genitalia slide RLB 328, 25 Jul. 1959, 1 female, genitalia
slide RLB 327, 5,600 ft, 1 Aug. 1959, 1 female, genitalia slide RLB 494, 4,880 ft, 2
Aug. 1959, 1 male, 4 Aug. 1959, 2 males, 1 female, 4,880 ft, 18 Aug. 1959, 1 male
(deposited in CU, RLB, UCB, UCONN, USNM).

Diagnosis. The pale yellowish color combined with the forewing pattern of the
subbasal and medial fasciae and postmedian spot in this species is unique among
known North American Eucosmini. The length of the male costal fold in C. plicata
and C. marmoreana is more than half the forewing length, whereas the costal fold
is absent or much shorter in other Catastega species. Male genitalia differ from those
of other known species by the combined forms of the socius and valva; female
genitalia differ from other known species by the combined forms of the ovipositor
and sterigma. The form of any one structure is similar to that in one or more other
species of Catastega.

Comments. The female of C. plicata is considered to have a form of ovipositor
for inserting eggs in crevices because of developed setae and loss of scales on the
eighth tergum, reduced ventral faces of the papillae anales, and developed apophyses,
which are characteristic of other species known to insert eggs in buds, bark crevices,
and other constricted areas. In contrast to many species with inserting ovipositors,
the ovipositor is not greatly lengthened and the moths fly during late summer, rather
than early spring. The ovipositor of C. plicata is similar to that of an undescribed
species of Catastega that occurs in southeastern United States; the latter species has
a forewing pattern and male genitalia very similar to C. timidella. Although a single
male has been collected from flowers of Yucca, the host remains unknown.

ACKNOWLEDGMENTS

I thank Dr. John G. Franclemont for his guidance during the period when I first began
gleaning new species of Catastega from among unidentified Olethreutinae. I gratefully acknowl-
edge Frederick H. Rindge (AMNH), L. L. Pechuman (CUIC), Edward C. Knudson, Jerry A.
Powell (UCB), David L. Wagner (UCONN), Don R. Davis (USNM), and the late Andre
Blanchard for the loan of specimens. Photographs of the imagos, except Figure 1, were made
by Howard Lyons; drawings were made by Amy Trabka, and plates were made by Joe MacGown.

1992

NEW CATASTEGA

227

LITERATURE CITED

Brown, R. L. 1986. Resurrection of Catastega Clemens and revision of the Epinotia vertum-
nana (Zeller) species-group (Tortricidae: Olethreutinae). J. Lepid. Soc. 40:327-346.

Brown, R. L. and J. A. Powell. 1991. Description of a new species of Epiblema (Lepidoptera:
Tortricidae: Olethreutinae) from coastal redwood forests in California with an analysis
of the forewing pattern. Pan-Pacific Entomol. 67:107-1 14.

Franclemont, J. G. 1961. Grant no. 303 — Johnson Fund ( 1 960), $800. Lepidoptera Heterocera
(moths) of southern Arizona. Year Book Amer. Philos. Soc., 1961:292-293.

Franclemont, J. G. 1964. Grant no. 3339 — Penrose Fund (1963), $800. Lepidoptera Heter-
ocera (moths) of southeastern Arizona. Year Book Amer. Philos. Soc., 1964:266-267.

Franclemont, J. G. 1976. New species of Arizona noctuids, I. (Lepidoptera: Noctuidae: Haden-
inae). Proc. Entomol. Soc. Wash. 78:409-416.

Horak, M. and R. L. Brown. 1991. Taxonomy and phylogeny. Pages 23-50 in: L. P. S. van
der Geest and H. H. Evenhuis (eds.), Tortricid Pests, Their Biology, Natural Enemies
and Control. Elsevier Science Publishers, Amsterdam, 808 pp.

Komerup, A. and J. H. Wanscher. 1983. Methuen Handbook of Colour (3rd ed.). Methuen
and Co., London, 252 pp.

Nijhout, H. F. 1978. Wing pattern formation in Lepidoptera: a model. J. Exp. Zool. 206:1 19-
136.

Received 18 July 1991; accepted 22 November 1991.

J. New York Entomol. Soc. 100(2):228-256, 1992

PUPAL MORPHOLOGY AND THE SUBFAMILY
CLASSIFICATION OF THE NOTODONTIDAE
(LEPIDOPTERA: NOCTUOIDEA)

James S. Miller

Kalbfleisch Curatorial Fellow, American Museum of Natural History,
Central Park West at 79th Street, New York, New York 10024

Abstract. — Pupal morphology is surveyed among 46 species of Notodontidae representing all
currently-recognized subfamilies, and 24 phylogenetically informative characters are identified.
Comparison of a previous notodontid subfamily phylogeny based on larval and adult mor-
phology with one constructed from pupal data shows close concordance. Although it was not
possible to identify a synapomorphy for the Notodontidae from pupae, this life stage provides
a long list of potential synapomorphies useful at the subfamily level.

The immature stages of Lepidoptera provide crucial character information for
studies in taxonomy and phylogeny. Unfortunately, our knowledge of pupal mor-
phology is lagging behind the growing body of data on larvae. This can be attributed,
at least in part, to a lack of available material. For example, in the North American
Notodontidae, where larvae are known for the majority of species, preserved pupae
are relatively uncommon in collections. Among tropical groups, the situation is
dismal; in the Neotropical notodontid subfamily Dioptinae, larvae are known
for about 4% of the fauna (Miller, 1992), and pupal material is available for less
than 2%.

Use of pupal characters has a long history in Lepidoptera systematics. Among the
earliest comparative studies are those of Jackson (1890), Poulton (1891), and Chap-
man (1892, 1893), all of whom stressed the potential taxonomic value of pupae.
Early attempts at a phylogeny for the lepidopteran families (Packard, 1895; Chapman,
1 896) relied heavily on pupal characters. Hinton’s ( 1 946) classification of insect pupae
drew attention to the major subdivisions of the Lepidoptera. Pupae figure prominently
in modern treatments as well. Minet (1991), in his recent phylogeny for the Ditrysia—
by far the largest and most homogeneous part of the order (Nielsen, 1 989)— identified
several important pupal characters.

The preeminent worker on Lepidoptera pupae was Edna Mosher, whose papers
form the basis for study even today. Her research uncovered characters useful in
elucidating problematic family interrelationships that were intractable using adult or
larval morphology. In addition to providing a general framework for pupal mor-
phology and classification of the Lepidoptera (Mosher, 1 9 1 6b), her work focused on
the families Saturniidae (1914, 1916a) and Sphingidae (1918a). She also published
the most comprehensive study available on pupae of the Notodontidae (Mosher,
1918b).

Pupae are taxonomically important in other families of Lepidoptera, an example
being the Geometridae (Rindge, 1952; McGufhn, 1987; Bolte, 1990). Nakamura’s

1992

NOTODONTID PUPAE

229

(1976) research on pupae of Japanese Lymantriidae treated 33 species in 16 genera.
After identifying a set of useful characters, he subdivided the genera and provided
a phylogeny. Common (1990), in his recent book on Australian moths, relied exten-
sively on pupae for family-level diagnostic characters, providing pupal illustrations
for representatives of all the Australian moth families.

There are distinct differences among lepidopteran groups in the form of the cocoon,
in the mode of pupation, and in the method of escape by the adult from the cocoon
and pupa (Hinton, 1 946; Common, 1 975). A well-known example from the Noctuidae
is the unusual boat-shaped cocoon found in Sarrothripinae and Nolinae, which pro-
vides evidence of relationship between the two subfamilies (Forbes, 1954; Francle-
mont, 1960; Kitching, 1984).

Differences in cocoon structure and in the mode of pupation can also be found
within the Notodontidae (Packard, 1895; Griinberg, 1912). For example, Cerura
larvae excavate a shallow depression in a twig (usually on the foodplant) and construct
an extremely dense cocoon formed from chewed wood particles and salivary secre-
tions (Dyar, 1891; Chapman, 1892). When the adult moth emerges it produces a
secretion from the head, possibly containing an enzyme (Kafatos and Williams, 1 964),
which dissolves the tough cocoon (Chapman, 1 892; Hinton, 1 946). Dolinskaya ( 1989)
divided notodontids into two groups: those— such as Cerura, Furcula, Gluphisia, and
Clostera— that pupate on the food plant, and those— such as Phalera, Stauropus,
Notodonta, Pheosia, and Spatalia — that leave the host to pupate in the soil or litter.
She further pointed out that in the first group of genera, all except Clostera exhibit
reduction of the cremaster (see Character 20 below). Although characters relating to
cocoon structure and pupation behavior will undoubtedly provide valuable infor-
mation for future work on notodontid systematics, they were not used here simply
because available data are scarce.

This paper is intended to compliment the pioneering works of Packard (1895),
Mosher (191 6b, 1 9 1 8b), and Dolinskaya ( 1 989) on notodontid pupae. Those authors
demonstrated extensive variation in a variety of characters, and all used pupal traits
in their subfamilial and generic diagnoses. However, their findings have not been
examined in the context of a world-level classification. In a recent paper (Miller,
1991), I recognized nine subfamilies for the world Notodontidae. My cladistic hy-
pothesis for those is shown in Figure 1 . This paper is not a comprehensive treatment
of pupae, but it provides an overview with two major aims: to identify potentially
useful characters for notodontid phylogeny and classification, and more generally, to
stimulate further interest in the morphology of this life stage. In particular, I stress
the implications of pupal data for understanding subfamily relationships.

MATERIALS AND METHODS

Morphological terminology (see Fig. 2) follows Mosher (1916b) and Common
(1990). Most of the pupae I studied had been preserved in alcohol. Intact pupae were
examined in almost all cases, but where these were not available, exuviae were used.
Line drawings were made using a camera-lucida attached to a Zeiss SV8 stereomi-
croscope. Scanning electron micrographs were taken with a Zeiss DSM950 micro-
scope. Specimens for SEM were air-dried, mounted on stubs in silver paste, and then
dried again overnight in an oven at 47°C before being coated with gold-palladium.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

— — DOIDAE

— — — — THAUMETOPOEINAE

— j— — — PYGAERINAE

— — r— — — — — — — — PLATYCHASMATINAE

— NOTODONTINAE

— — PHALERINAE

— — DUDUSINAE

— HETEROCAMPINAE

NYSTALEINAE

— DIOPTINAE

Fig. 1. Cladogram of subfamilies in the Notodontidae based on an analysis of 174 larval

and adult characters (from Miller, 1991).

The list of species examined is shown in Table 1 . Comprehensive study of outgroup
taxa was beyond the scope of this paper. The Doidae— containing two relatively small
genera, Doa Neumoegen and Dyar and Leuculodes Dyar (Franclemont, 1983; Don-
ahue and Brown, 1987)— and Notodontidae are thought to be sister groups (Miller,
1991). I used pupae of Doa ampla (Grote) (Fig. 2) for outgroup comparison. The
pupa of another doid, D. dora Neumoegen and Dyar, was figured by Brown (1990).
I have generally relied on Mosher (1916b) in my assessment of pupal morphology
among quadrifid Noctuoidea, but needless to say, this is a cursory approach; the
group contains tremendous morphological variation, and Mosher’s treatment is rel-
atively brief.

I performed cladistic analyses using pupal characters scored for 16 notodontid
species. These 16 taxa were chosen to represent the nine subfamilies recognized in
Miller (1991). Twenty-four characters comprising 61 character states (Table 2) were
identified. The resulting data matrix (Table 3) was then analyzed using the Hennig86
parsimony program of Farris (1988), with Doa ampla as an outgroup. Shortest trees
were produced by implicit enumeration. The analysis provides a test of how pupae
fit with my previous hypothesis of subfamily relationships based on adult and larval
characters (see Fig. 1).

PUPAL MORPHOLOGY

Below is the list of characters for which I found useful variation. My treatment is
preliminary; more characters certainly remain to be discovered, and it is impossible
to fully assess morphological variation using such a small species sample. However,
all the traits I discuss deserve consideration in future taxonomic research on the
Notodontidae. Characters with numbers in parentheses were used in the cladistic
analyses (see Table 2).

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Fig. 2. Pupa of Doa ampla (Grote) (Doidae). A. Ventral view. B. Lateral view. C. Dorsal
view. A = abdominal segment; an = antenna; ao = anal opening; cr = cremaster; e = eye; fr
= front; go = genital opening; lp = labial palpi; ml = mesothoracic leg; ms = mesothorax; msp
= mesothoracic spiracle; mt = metathorax; mw = mesothoracic wing; pb = proboscis; pi =
prothoracic leg; pt = prothorax; sp = spiracle; tl = metathoracic leg. Scale line = 2 mm.

Head:

Eyes. ( 1 ) Mosher (191 6b) recognized two parts of the eye in Lepidoptera pupae:
one part she termed the “sculptured eye-piece,” and the other the “glazed eye-piece.”
I found this division difficult to see, at least in notodontids, and the morphological
distinction may be unnecessary for this group.

Mosher (1918b) identified an interesting character in her study of Maine noto-
dontids. She noticed that in Clostera species, the proximo-lateral angles of the pro-
boscis do not reach the eye-pieces (Fig. 4A). This is also illustrated by the pupae of
Doa ampla (Fig. 2A) and Thaumetopoea pinivora (Fig. 3A). In most notodontids,
the proximo-lateral angles of the proboscis reach the eye-pieces (Figs. 5A, 6A, 8A-
12A). In Cerura and Furcula they do not (Fig. 7 A), but in other Notodontinae they
do (Fig. 8A). Thus, according to my survey only the Doidae, the Thaumetopoeinae,
the Pygaerinae, as well as Cerura and Furcula of the Notodontinae, exhibit the first

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Table 1. List of species for which pupae were examined arranged according to the classi-
fication of Miller (1991).

Taxon

Locality

Collection

Doidae

Doa ampla (Grote)*

Arizona

CUIC

Notodontidae

Thaumetopoeinae

Thaumetopoea pinivora (Treitschke)*

Germany

CUIC

Pygaerinae

Clostera albosigma Fitch*

New York

CUIC, NYSM

C. inclusa Hiibner

Illinois

INHS

Platychasmatinae

Platychasma virgo Butler*

Japan

SSC

Notodontinae

Notodontini

Notodonta simplaria Graef*

New York

NYSM

Pheosia portlandia Hy. Edwards

Oregon

NYSM

Hyperaeschra georgica Herrich-Schaffer

New York

CUIC

Dicranurini

Centra scitiscripta Walker

Florida

AMNH

Furcula occidentalis (Lintner)*

New York

AMNH

F. scolopendrina (Boisduval)

New York

NYSM

F. modesta (Hudson)

New York

NYSM

Gluphisia septentrionis Walker*

New York

AMNH, CUIC

Phalerinae

D at ana robust a Strecker

Texas

CUIC, USNM

D. integerrima Grote and Robinson

New York

CUIC, NYSM

D. ministra (Drury)*

New York

CUIC

D. drexelii Hy. Edwards

New York

CUIC

D. contract a Walker

New York

CUIC

D. angusii Grote and Robinson

New York

CUIC

Phalera sp.**

—

—

Ellida caniplaga Walker

New York

CUIC

Nadata gibbosa Smith

Illinois

INHS, NYSM

Dudusinae

Crinodes besckei Hiibner*

Costa Rica

INHS

Goacampa variabilis Schaus*

Costa Rica

INHS

Heterocampinae

Heterocampa umbrata Walker

Illinois

INHS

H. bill neat a Packard

Illinois

INHS

H. guttivitta Walker*

New York

NYSM

Hyparpax aurora (Smith)

New Jersey

AMNH, USNM

Lochmaeus manteo Doubleday*

New York

NYSM

Misogada unicolor Packard

New York

NYSM

1992 NOTODONTID PUPAE 233

Table 1. Continued.

Taxon

Locality

Collection

Schizura ipomoeae Doubleday

Connecticut

AMNH

S. badia (Packard)

New York

AMNH

S. unicornis (Smith)

New Jersey

AMNH

S. concinna (Smith)

Illinois

INHS

Litodonta hydromeli Harvey

Texas

SJW

Spat alia argent ina (Schiffermuller)

Europe

CUIC

Nystaleinae

Didugua argentilinea Druce*

Texas

USNM

Symmerista albifrons Smith*

Illinois

INHS

S. leucitys Franclemont

New York

NYSM

Dioptinae

Cyanotricha necyria Felder

Colombia

AMNH

Josia ligula Hiibner*

Ecuador

SJW

Phaeochlaena gyon Fabricius

Brazil

BMNH

Phryganidia californica Packard

California

USNM

Zunacetha annulata Guerin*

Panama

AMNH

Incertae Sedis

Nerice bidentata Walker

New York

NYSM

Lophopteryx cuculla Esper

Germany

CUIC

Drymonia trimacula Esper

Germany

CUIC

*, Species used for the cladistic analyses (see text).

**, Characters for Phalera pupae were taken from descriptions and figures in the literature.
Acronyms for collections: AMNH, American Museum of Natural History, New York, NY;
BMNH, The Natural History Museum, London, GB; CUIC, Cornell University Insect
Collections, Ithaca, NY; INHS, Illinois Natural History Survey, Champaign, IL; NYSM,
New York State Museum, Albany, NY; SJW, Susan J. Weller collection, LSU, Baton Rouge,
LA; SSC, Shigero Sugi collection, Tokyo, Japan; USNM, United States National Museum,
Washington, DC.

character state, while all other taxa show the second. The “primitive” character state
in Cerura/Furcula may not be homologous with others; in both genera, the eye-piece
itself is relatively small (see Fig. 7A, B).

Front shape. The front in pupae is defined as the sclerite to which the antennae
are attached (Mosher, 1 9 1 6b). There are various shapes of the front. It is smoothly
rounded in most species, whereas it protrudes to varying degrees in others, Zunacetha
annulata (Dioptinae) (Fig. 12) being an example. The front in Datana and Phalera
species is highly sculpted (Fig. 13; Dolinskaya, 1989).

Some primitive Lepidoptera pupae have a “cocoon-cutter” on the head used to
cut through the fabric of the cocoon, but there is no comparable structure in Ditrysia
(Hinton, 1946). Various notodontids have modifications on the adult head (Miller,
1991) that assist in digging through the substrate upon emergence from the pupa.
However, there seems to be no correlation between adult and pupal stages regarding
modified heads. For example, notodontids with specializations in adults, such as
Thaumetopoea and Goacampa, show nothing in pupae (Figs. 3, 9). In contrast, the

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Table 2. Pupal characters used in a cladistic analysis of sixteen representative notodontid
species. Autapomorphic characters were not used, and are omitted from the list.

1. Proximo-lateral angles of the proboscis: not reaching eye-pieces (0); reaching eye-pieces

(1).

2. Labial palpus sclerite: present (0); absent (1).

3. Length of proboscis: approximately xh that of wings, prothoracic legs and mesothoracic legs
meeting at midline (0); approximately Vi that of wings, only mesothoracic legs meeting at
midline (1); equal to or greater than that of wings, prothoracic and mesothoracic legs not
meeting at midline (2).

4. Anterior suture of proboscis: strongly curved, arching forward toward vertex of head (1);
not strongly curved and arching forward (0).

5. Antennae: shorter than mesothoracic legs (0); extending beyond mesothoracic legs (1);
greatly elongate, extending to A5 (2).

6. Thorax length (ventral view): less than % total body length (0); equal to % total body length

(1).

7. Secondary setae: absent (0); covering thorax and abdomen (1).

8. Dorsum of thorax: smooth or with a small ridge along midline (0); with a distinct crest
along midline (1).

9. Sculpturing on caudal margin of mesothorax: not present (0); with a series of shallow
depressions separated by raised ridges (1); with a series of small, quadrangular plates
separated by deep pits (2).

10. Shape of mesothorax: smoothly rounded dorsally, posterior margin gradual (0); with a
dorsal hump, posterior margin projecting at body midline (1).

1 1. Lateral portions of metathorax: roughly parallel-sided (0); sinuate (1).

1 2. Thoracic cuticle: smooth or with a few scattered, shallow pits (0); covered with many deep
pits (1).

13. Anterior margin of each movable segment (A5-A7): unmodified (0); with a diffuse band
of small pits (1); with a single row of lunate pits (2); with a distinct carina (3).

14. Abdominal cuticle: smooth or with fine wrinkles (0); with small pits scattered over the
surface (1); covered with large, deep pits (2); covered with large, shallow pits and raised
ridges (3).

15. Abdominal spiracles: small (0); large (1).

16. Abdominal spiracle shape: spiracle margins equal in size, spiracle opening laterally (0);
anterior spiracular margin more robust and raised, so that spiracle opens posteriorly (1).

17. Segments A8-A10: tapered posteriorly (0); broadly rounded (1).

18. Anterio-dorsal margin of A 10: unmodified (0); with a deep furrow (1); sharply elevated,
with a lip-shaped edge (2).

19. Dorsum of A 10: smooth (0); with a raised, reticulate area anterior to cremaster (1); with
numerous, small longitudinal ridges (2).

20. Cremaster: variously shaped (0); bifurcate (1); long and spine-like (2); broad, compressed
(3); absent (4).

21. Base of cremaster: smooth (0); fluted (1).

22. Number of cremaster setae: 20 or more (0); between eight and ten (1); setae absent (2);
cremaster absent (?).

23. Body: cylindrical (0); dorso-ventrally compressed (1).

24. Body: variously shaped, but roughly elongate (0); wide relative to its length (1).

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Table 3. Data matrix of 24 pupal characters (Table 2) for 16 species of Notodontidae (see
Table 1) with Doa ampla used as an outgroup.

5

10

15

20

24

Doa

00000

00000

00000

00000

0000

Thaumetopoea

01000

01000

00000

00001

0201

Clostera

00000

01000

10011

00002

0000

Platychasma

10201

00000

10010

00020

0000

Notodonta

11100

00000

10011

01001

0100

Furcula

11000

00000

10011

01004

0711

Gluphisia

11000

00000

10011

01004

0711

Datana

11100

01000

10021

00101

0200

Phalera

11100

01000

10021

00101

0200

Goacampa

11210

00110

10331

00000

0100

Cr inodes

11210

00110

10311

00201

0100

Heterocampa

11211

00020

10111

00011

0200

Lochmaeus

11211

00020

10111

00011

0200

Symmerista

10211

00000

11211

10001

1100

Didugua

10211

00000

11211

10001

1100

Josia

11212

10001

10010

10003

1100

Zunacetha

10212

10001

10010

10003

1100

head of Z. annulata has a tubercle in the pupal stage (Fig. 12) but is unmodified in
adults.

Labial palpi. (2) Although the labial palpi can be relatively large in taxa outside
the Notodontidae (e.g., Doa ampla, Fig. 2 A), they are either highly reduced or absent
among notodontids (Mosher, 1916b, 1918b; Common, 1990; Minet, 1991). Most
commonly they are absent, but a small, triangular labial palpus sclerite occurs among
various notodontid subfamilies (Dolinskaya, 1989) including the Platychasmatinae
(Fig. 5 A), Nystaleinae (Fig. 1 1A), and Dioptinae (Fig. 12 A).

It is interesting that, contrary to modem ideas, Mosher’s concept of the Noctuoidea
excluded the Notodontidae. She placed notodontids in a separate superfamily, the
Notodontoidea, along with the Geometridae. Unfortunately, there is still no pupal
character known, comparable to the metathoracic tympanum in adults, that can be
used to define the Noctuoidea (as the term is currently applied). Mosher (1916b,
1918b) grouped the Notodontidae and Geometridae together because in both the
pupae lack labial palpi, but instead have only “a very small triangular or polygonal
area” (191 6b: 1 25) remaining. Adult and larval data certainly do not support Mosher’s
phylogenetic hypothesis (see Miller, 1991; Minet, 1991).

Length of proboscis. (3) Mosher (191 6b) used the term maxillae for the proboscis
of Common (1990); I follow the latter. Both authors noted that, although highly
variable in length, the proboscis is always visible in Lepidoptera. In notodontids,
extremes range from the thaumetopoeines, where the proboscis is less than xh the
length of the mesothoracic wings (Fig. 3A), to the Dioptinae, where it can extend
beyond the caudal margin of the wings, actually touching A5 (Fig. 12A). I divided
the character into three states (Table 2). In species with an extremely short proboscis,

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

the prothoracic and mesothoracic legs meet at the ventral midline (Figs. 3A, 4A, 7A,
8A). In taxa such as Datana and Notodonta, with a tongue only slightly shorter than
the thorax, the mesothoracic legs meet at the midline but the prothoracic ones do
not (Fig. 6A; Mosher, 1918b). Finally, in long-tongued species (Figs. 5A, 9A-12A)
neither pairs of legs meet along the midline (Mosher, 1918b; Dolinskaya, 1989).

This character has been noted by all authors since Packard (1895). However,
proboscis length in pupae can vary greatly even within notodontid subfamilies. For
example, there are heterocampine species in which the tongue is as long as the wings
(Fig. 10A), and species where the proboscis is much shorter than the wings. Similar
variation occurs in the Nystaleinae. It is not surprising that tongue length in pupae
is correlated with that in adults (Poulton, 1891).

Anterior sutures of proboscis. (4) I here define the “anterior sutures” of the proboscis
as the sutures where the base of the proboscis and the head join (see Figs. 10, 11).
In most species, these either arch slightly forward at their midpoint (Fig. 3A), or are
straight with an orientation essentially perpendicular to the body midline (Fig. 8A).
Four subfamilies of the Notodontidae— the Dudusinae, Heterocampinae, Nysta-
leinae, and Dioptinae — exhibit a different configuration. In these, the anterior sutures
of the proboscis are strongly curved, bending toward the vertex of the head, with the
proximo-medial angles of the proboscis being narrow (Figs. 9 A- 12 A).

Length of antennae. (5) Like the proboscis, the antennae of pupae vary in length.
In addition, the antennae can either be shorter than the mesothoracic legs or they
can extend beyond the mesothoracic legs. The former is typical of most notodontids
(e.g., Fig. 9 A) as well as their sister group, Doa (Fig. 2 A), while the latter appears to
be restricted to the Platychasmatinae (Fig. 5 A), Heterocampinae (Fig. 10 A), Nys-
taleinae (Fig. 11 A) and Dioptinae (Fig. 12A). Mosher (1916b), based on study of
Phryganidia californica, listed long antennae, touching A5, as a diagnostic trait for
the Dioptinae. This character state occurs in all the dioptines I examined.

Thorax:

Thorax length. (6) Dioptinae are unusual in that the thorax is elongate. In the
species I studied, the thorax is 2h the total body length (Fig. 12), whereas in other
notodontids the thorax is less than 2A the body length (e.g., Figs. 3, 5, 10).

Secondary setae. (7) There are primary setae on the body in pupae analogous to
the primary setae in larvae (Common, 1990). Those of pupae are small and difficult
to locate (Mosher, 1916b). The primary setae of Zunacetha annulata (Fig. 12) are
large compared to the other species I examined; usually they are miscroscopic. I did

Figs. 3, 4. Pupae of Notodontidae. 3. Thaumetopoea pinivora (Treitschke). 4. Clostera
albosigma Fitch. A. Ventral view. B. Lateral view. C. Dorsal view. Scale lines = 3 mm.

Figs. 5, 6. Pupae of Notodontidae. 5. Platychasma virgo Butler. 6. Datana robusta Strecker.
A. Ventral view. B. Lateral view. C. Dorsal view. Scale lines = 3 mm.

Figs. 7, 8. Pupae of Notodontidae. 7. Furcula scolopendrina (Boisduval). Scale line = 3 mm.
8. Gluphisia septentrionis Walker. Scale line = 2 mm. A. Ventral view. B. Lateral view. C.
Dorsal view.

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9

Figs. 9, 10. Pupae of Notodontidae. 9. Goacampa variabilis Schaus. 10. Lochmaeus manteo
Doubleday. A. Ventral view. B. Lateral view. C. Dorsal view, as = anterior suture; pf = femur
of prothoracic leg. Scale lines = 3 mm.

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NOTODONTID PUPAE

241

not examine primary setal patterns in pupae, but such research might be valuable.
They have rarely been used for taxonomic purposes.

Secondary setae can also occur. For example, larvae and pupae of Lymantriidae
are covered with long secondary setae (Poulton, 1891; Nakamura, 1976). The rela-
tionship between presence or absence of secondary setae in the larval and pupal stages
of Notodontidae is interesting. This character shows a high level of homoplasy in
larvae; secondary setae occur in caterpillars of the Thaumetopoeinae, Pygaerinae,
Phalerinae, Nystaleinae, Dioptinae, and Dudusinae, and each of these cases is thought
to have evolved separately (Miller, 1991). Presence of secondary setae in the pupal
stage appears to exhibit a more restricted taxonomic distribution. Genera such as
Cyanotricha (Dioptinae) and Dasylophia (Nystaleinae) have secondary setae in larvae
but not in pupae. Based on the notodontid pupae I examined, secondary setae occur
only in the subfamilies Thaumetopoeinae, Pygaerinae, and Phalerinae. Undoubtedly,
examples in other subfamilies will eventually be found. Secondary setae in notodontid
pupae are extremely short, almost microscopic (Mosher, 1918b).

Femur of prothoracic legs. Both Common (1990) and Mosher (1918b), in their
diagnoses for the family, stated that the femur of the prothoracic leg is never visible
in pupae of Notodontidae. The structure is frequently visible in various other families.
I found that the profemur can be seen in at least one notodontid species, Goacampa
variabilis (Dudusinae) (Fig. 9A). This is obviously a derived character state for the
Notodontidae, but without examining additional species its usefulness for phylogeny
is unknown.

Prothoracic crest. (8) In the two dudusines I studied, Crinodes besckei and Goa-
campa variabilis, there is a distinct crest along the midline on the dorsum of the
mesothorax (Fig. 1 4). It is present, but less obvious, on the prothorax as well. Although
a few other taxa show a slightly raised area along the midline (e.g., Figs. 3C, 10C),
it is not pronounced.

Mesothoracic spiracle. The shape of the mesothoracic spiracle varies among species
(Mosher, 1 9 1 6b), with sculpturing around the spiracle showing the most significant
differences. Unfortunately, thorough study would require SEM for all taxa, something
beyond the scope of this paper.

Pits on mesothoracic caudal margin. (9) On the dorsum of the pupa some taxa
exhibit modifications of the mesothoracic caudal margin. The most remarkable of
these occurs in the Heterocampinae (Fig. 10C). Here, there is a row of from six to
twelve small, quadrangular plates separated by deep pits (Figs. 15-17). The plates
are slightly convex (Fig. 1 6) and their surfaces are rugose under high magnification
(Fig. 1 8). This apomorphy, first mentioned by Dyar ( 1889) and described by Packard
(1895) in species of North American Heterocampinae, has been noted by most
subsequent authors. Based on my study and on reports in the literature, the trait is
restricted to Heterocampinae but is quite widespread within the subfamily, occurring
in at least nine genera (Dyar, 1889; Packard, 1895; Mosher, 1916b, 1918b; Weller,
1987; Dolinskaya, 1989). Its function is unknown.

A similar, but less spectacular, modification can be found in Goacampa variabilis
and Crinodes besckei (both Dudusinae). In these species, the caudal margin of the
mesothorax bears a row of shallow depressions separated by raised longitudinal ridges
(Figs. 9C, 14). Although the character in Heterocampinae and Dudusinae may be
homologous, the character states in the two groups are clearly separable.

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Figs. 13, 14. Scanning electron micrographs of notodontid pupae. 13. Front of Datana
ministra (Drury), ventral view (500 yum). 14. Meso- and metathorax of Goacampa variabilis
Schaus, dorsal view with anterior at top (500 yum). Scale lengths in parentheses.

Shape of mesothoracic caudal margin. (10) A modification of the mesothoracic
caudal margin, not related to sculpturing, occurs in pupae of Dioptinae. Here, the
mesothorax is humped posteriorly and its caudal margin projects sharply posteriad
at the body midline (Fig. 12C; Mosher, 1916b). In other notodontids, the caudal
margin of the mesothorax forms a shallow curve posteriorly (e.g., Fig. 11C). The
derived state seems to provide yet another synapomorphy for the Dioptinae.

Lateral extensions of metathorax sinuate. (11) In all pupae I studied except Doa
ampla and Thaumetopoea pinivora, the lateral extensions of the metathorax have a
constriction opposite the A2 spiracle and an expanded distal portion (e.g., Figs. 9B,
1 IB). This configuration is more marked in some taxa than in others, but it seems
to define a monophyletic group that includes all notodontid subfamilies except the
Thaumetopoeinae.

The length of the lateral portions of the metathorax also varies among Notodon-
tidae. In some species they extend as far posteriad as A4, either touching the anterior
margin of that segment (e.g., Thaumetopoea pinivora, Fig. 3B) or extending beyond
it (e.g., Platychasma virgo, Fig. 5B). In other taxa, they extend only as far as the
middle of A3 (e.g., Goacampa variabilis, Fig. 9B). However, this character shows so
much homoplasy that its value is questionable.

Forewing venation. In most Ditrysia, the only wings visible in the pupa are the
forewings (Mosher, 1916b). The trifid Noctuoidea, to which the Doidae and Noto-
dontidae belong (Miller, 1991), have the forewing cubital vein three-branched (com-
prising veins CuA,, CuA2, and M3), a plesiomorphic trait. The derived state is the
quadrifid condition, where M2 branches from the cubital as well. These veins are
usually difficult to see in pupae, except during later development when the pharate

Figs. 11, 12. Pupae of Notodontidae. 11. Didugua argentilinea Druce. Scale line = 3 mm.
12. Zunacetha annulata Guerin. Scale line = 2 mm. as = anterior suture; lp = labial palpi. A.
Ventral view. B. Lateral view. C. Dorsal view.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Figs. 15-20. Scanning electron micrographs of notodontid pupae. 15. Posterior margin of
mesothorax in Heterocampa guttivitta Walker, dorsal view (400 /urn). 16. A single plate from
the structure in Figure 15 (50 pm). 17. A pit between plates (40 pm). 18. Surface structure of
a single plate (20 pm). 19. Symmerista leucitys Franclemont ventral view, showing bottom of
wing and segments A4-A5 on right side; tips of antennae in upper right (500 pm). 20. Similar
view of Ellida canip/aga (500 pm). Scale lengths in parentheses.

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NOTODONTID PUPAE

245

adult can be seen through the pupal cuticle. Nevertheless, the trifid condition can be
used to separate pupae of the Doidae and Notodontidae from other Noctuoidea.
There is one exception; the subfamily Platychasmatinae, containing the monobasic
genus Platychasma, is the only notodontid group in which the forewing is quadrifid
(Nakamura, 1956; Miller, 1991).

Cuticle surface structure. (12) Thoracic surface structure seems to vary indepen-
dently of that on the abdomen. The nystaleine pupae I examined are characterized
by numerous, deep pits covering the thorax (Fig. 1 9). The thorax of other notodontids
either has a few shallow pits scattered over its surface (e.g., Ellida caniplaga; Fig.
20), or is smooth.

Appendages standing out in relief. In most notodontid pupae the appendages are
fused together to form a relatively flat surface. In Cerura and Furcula, the proboscis,
legs, and antennae stand out in relief from the body surface (Mosher, 1916b), a
character state also noted by Dolinskaya (1989) for the Palearctic genera Harpy ia
and Ptilophora. I placed Ptilophora in the subfamily Notodontinae along with Cerura
and Furcula, but I put Harpyia in the Heterocampinae (Miller, 1991). The derived
state is not found in other Notodontinae, such as Gluphisia and Notodonta (Dolin-
skaya, 1989). This trait needs further study.

Abdomen:

The movable segments (A5-A7). (1 3) In primitive Lepidoptera, abdominal segments
A1-A7 are movable, while A8-A10 are always fused (Mosher, 1916b). In more
advanced families, such as the Notodontidae, the only movable segments of the
abdomen are A5-A7 (Mosher, 1918b). Segments A1-A4 show little morphological
variation among notodontid species. The movable segments, on the other hand, often
exhibit unique structural modifications not found on other abdominal segments.
Segments A9 and A 10, bearing the cremaster, show extreme variability (see Character
20).

Various types of specializations can be found along the anterior margin of the
movable segments in Notodontidae. The traits appear to provide valuable phylo-
genetic information, especially useful at the subfamily level. Among more primitive
notodontid subfamilies, the movable segments are not modified (e.g., Figs. 3, 7, 8).
In advanced groups, however, at least four discrete character states occur: Hetero-
campine pupae have a diffuse band of small pits ringing the anterior margin of each
movable segment (Figs. 10, 21). Nystaleine pupae have a single row of lunate pits
(Figs. 1 1, 22). Mosher (1918b) first described this trait for Symmerista, but it occurs
in other Nystaleinae as well. Finally, in Dudusinae there is a sharp carina near the
anterior margin of each movable segment (Fig. 9). A similar carina can be found in
Hyperaeschra georgica (Fig. 24). This species has long been placed in the subfamily
Notodontinae (Packard, 1895). If that placement is correct, the carina has evolved
there independently.

Cuticle structure. (14) Various authors have characterized the abdomen in Noto-
dontidae as being punctate (Mosher, 1918b; Marumo, 1920; Common, 1990). I did
not survey abdominal surface structure in detail, but there is considerable variation
among species. The cuticle surface of Doa ampla and Thaumetopoea pinivora is
smooth or somewhat wrinkled, without punctations. In other groups, small punc-
tations are the general rule (Figs. 1 9-22), but in Datana and Phalera (both Phalerinae),

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Figs. 21-26. Scanning electron micrographs of notodontid pupae. 21. Segment A4 of Het-
erocampa guttivitta Walker, dorsal view with anterior at top (500 fim). 22. Segment A5 of
Symmerista leucitys Franclemont, ventral view with anterior at top (200 gm). 23. Terminal
segments of Goacampa variabilis Schaus, dorsal view with posterior at top (500 gm). 24.
Segments A4-A7 of Hyperaeschra georgica Herrich-Schaffer, dorsal view with anterior at top,
showing paired pits (1 mm). 25. A single pit on A6 of H. georgica (100 gm). 26. Segments A8-
A10 of Datana ministra (Drury), dorsal view with posterior at top, showing furrow on A10
(500 gm). Scale lengths in parentheses.

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247

the abdomen is covered with large pits (Fig. 26; Dyar, 1891; Dolinskaya, 1989). The
abdomen of Goacampa variabilis exhibits complex surface sculpturing, with shallow
pits and furrows (Fig. 23). This character shows variation analogous to that on the
surface of the larval head (see Miller, 1991). Like that character, SEM study across
a wide variety of species would be valuable.

Paired pits on A5-A7. In Hyperaeschra georgica, there is a pair of large dorsal pits
on the anterior margin of A5-A7 (Figs. 24, 25). The character is distinctive, but I
observed it only in this species. I have not seen reference to these structures in the
literature, and their function is unknown.

Spiracle size. (15) The abdominal spiracles are found laterally on segments A2-
A8 (Fig. 2B). The one on A8 is always smaller than others, and according to Mosher
(1916b) it is not functional. The abdominal spiracles show size variation across taxa.
I simply scored them as being either large or small; a more detailed analysis is needed.
Small spiracles occur in Doidae, Thaumetopoeinae, Platychasmatinae, and Dioptinae
(Figs. 2B, 3B, 5B, 12B).

Spiracle shape. (16) I used one character involving spiracle shape. In Dioptinae
and Nystaleinae, the anterior margin of each abdominal spiracle is more robust and
heavily sclerotized. Furthermore, this margin is raised so that the spiracular opening
faces somewhat posteriorly rather than laterally, the typical orientation.

There are also differences in the sculpturing around the spiracle (Mosher, 1918b).
For example, some species have the entire spiracle raised from the body surface (Fig.
2A). This is yet another character that would benefit from comprehensive SEM study.

Shape of segments A8-A10. (17) Pupae in the subfamily Notodontinae show a
great deal of morphological variation. One potential synapomorphy involves the
shape of the terminal abdominal segments (A8-A10), which are broadly rounded
(Figs. 7, 8) rather than being tapered as in other Notodontidae. Many notodontines
have lost the cremaster, and these exhibit the rounded abdomen. However, the trait
also occurs in taxa, such as Notodonta, where the cremaster is present (Ferguson,
1963).

Male and female pupae exhibit different external morphologies on the venter of
the terminal segments, allowing one to distinguish the sexes (Jackson, 1890; Poulton,
1891; Mosher, 1918b). However, I did not examine this character complex from a
taxonomic standpoint because often, specimens of only a single sex were available
for study.

Anterior margin of A10. (18) In Datana and Phalera pupae, the anterior margin
of A 1 0 on the dorsal side bears a deep furrow (Poulton, 1891; Nagano, 1916; Mosher,
1918b; Dolinskaya, 1989). The posterior edge of this furrow is often sculpted (Fig.
26). Judging from the species I examined and figures in the literature, there is inter-
specific variation in shape. Datana species are extremely difficult to separate based
on adult characters, so it would be interesting to see whether differences in furrow
shape provide valuable species-specific characters.

There is an analogous modification in pupae of Cri nodes besckei, where instead of
a deep furrow, the anterior margin of A10 is sharply elevated and lip-shaped. This
character does not occur in the other dudusine I examined, Goacampa variabilis
(Figs. 9C, 23).

Reticulate area on A 10. (19) Some pupae have a sharply raised, reticulate area on
the dorsal surface of segment A 10 just anterior to the cremaster. This structure was

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Figs. 27-32. Scanning electron micrographs of notodontid pupae. 27. Cremaster and A10
of Heterocampa guttivitta Walker, dorsal view (200 yu m). 28. Cremaster and A10 of Symmerista
leucitys Franclemont, ventral view (200 yum). 29. Cremaster and A 10 of Ellida caniplaga Walker,
ventral view (200 pm). 30. Cremaster of Zunacetha annulata Guerin, ventral view (100 yum).
31. Cremaster of Josia ligula Hiibner, ventral view (100 yum). 32. Cremaster setae of J. ligula,
ventral view (50 yum). Scale lengths in parentheses.

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249

discussed for two Palearctic species, Fentonia ocypete Bremer and Spatalia argent ina,
by Dolinskaya (1 989). It also occurs in Hyparpax perophoroides (Strecker) (Ferguson,
1963) and Misogada unicolor. All these taxa belong in the subfamily Heterocampinae
(Forbes, 1948; Miller, 1991). In Heterocampa guttivitta, this reticulate area consists
of a few laterally oriented ridges (Fig. 27). In some heterocampines it is absent
altogether. Platychasma has numerous, small longitudinal ridges on segments 9 and
10 (Fig. 5C), but these are probably not homologous with the structure in Hetero-
campinae.

Cremaster shape. (20) There is almost endless variation in cremaster morphology
among Notodontidae; my study does not begin to tap the structure’s taxonomic
potential. Proper treatment of this character complex would require a monograph in
itself. My goal here is to describe some of the variation, and to begin outlining ways
in which the various character states might be useful for phylogenetic purposes.

In species belonging to at least six subfamilies— the Thaumetopoeinae, Notodon-
tinae, Phalerinae, Dudusinae, Heterocampinae, and Nystaleinae— the cremaster is
bifurcate (e.g., Figs. 3, 27, 28). There may be examples in other subfamilies as well.
I apply a broad definition to this term, scoring any cremaster as bifurcate whose
parts are divergent. A bifurcate cremaster may be the ground-plan state for the entire
Notodontidae. The distinctive fork-shaped cremaster of Heterocampine (Figs. 10,
27) has been noted by various authors (Packard, 1895; Mosher, 1918b; Marumo,
1920; Ferguson, 1963), and it seems to be a definitive subfamily trait, at least for
Holarctic species. A bifid cremaster with spiny projections (Fig. 26) is characteristic
of Datana and Phalera (Dyar, 1891; Poulton, 1891; Mosher, 1918b).

Other cremaster types include the long, spine-like cremaster of Clostera (Packard,
1895; Nagano, 1916; Mosher, 1 9 1 8b) and Ellida (Fig. 29), and the broad, compressed
cremaster of Dioptinae (Figs. 30, 31).

Finally, there are notodontid pupae where the cremaster is entirely absent. This
was described over a hundred years ago by Poulton (1891) and Dyar (1891) for
species of Cerura and Furcula (Notodontinae), and has subsequently been discussed
by other workers (e.g., Packard, 1895; Dolinskaya, 1989). In these species, the ninth
and tenth abdominal segments are broadly rounded and the abdomen is completely
smooth posteriorly (Fig. 7). An almost identical condition occurs in the genus Glu-
phisia (Packard, 1895; Ferguson, 1963; Dolinskaya, 1989; Fig. 8). I previously made
the suggestion (based on larvae and adults) that Cerura/ Furcula and Gluphisia not
only belong together in the same subfamily, the Notodontinae, but that they belong
in the same tribe, the Dicranurini (see Table 1). Cremaster morphology provides
another synapomorphy in support of my hypothesis.

Base of cremaster fluted . (21) The base of the cremaster in all dioptine pupae I
examined has a characteristic series of longitudinal ridges on its ventral surface. A
possibly homologous condition occurs in Nystaleinae, where the cremaster is a dif-
ferent shape than in Dioptinae but also has longitudinal ridges at its base. Adult and
larval data show that these two subfamilies are sister groups (Weller, 1989; Miller,
1991), and I draw attention to this character as another potential synapomorphy.

Cremaster setae. (22) In many families of Lepidoptera there are hook-shaped setae
on the cremaster (Fig. 32). These setae can serve either to anchor the pupa to the
lining of the cocoon, or in species lacking a cocoon, to attach it to a silk pad on the
substrate spun previously by the caterpillar (Common, 1990). There are four pairs
of cremaster setae in most Geometridae, and a nomenclature (“Dl, D2, LI, SD1”)

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has been applied to them (e.g., see Bolte, 1 990). Presumably, these terms were derived
from the primary setal names on the anal plate of larvae (McGuffin, 1987). Although
eight is also a typical number of cremaster setae for Notodontidae (see below), I
could not homologize those in notodontids with the ones in Geometridae, and so
have not applied setal names.

Mosher claimed that presence of hooked setae “is the exception in the Notodon-
tidae, as most of them pupate in the ground” (19 16b: 5 8), implying that pupation in
the ground obviates the need for setae. I found that hook-shaped cremaster setae
occur in six of the nine notodontid subfamilies. Based on my survey, the groups in
which they are absent are the Thaumetopoeinae (Fig. 3), Phalerinae (Fig. 26), and
Heterocampinae (Fig. 27). Mosher (1918b) used this as diagnostic for Heterocam-
pinae. Among members of the subfamily Notodontinae, setae occur in Notodonta
(Ferguson, 1963) and other more primitive members of the group, but they are
obviously absent in taxa that lack the cremaster altogether (see above).

Where setae do occur, they vary in number from only eight in Symmerista and
Notodonta (Ferguson, 1 963) to over 20 in P/atychasma (Fig. 5). The outgroup number
is about 20 as well ( Doa ampla, Fig. 2). There is also variation within subfamilies.
For example, among the Dioptinae numbers range from eight or ten setae in Josia
and Zunacetha (Figs. 30-32) to about 25 in Cyanotricha necyria. Phryganidia cali-
fornica (Dioptinae) is unique in having hook-shaped setae on the dorsum of segments
A7-A9, as well as on the cremaster itself (Miller, 1987). The long cremaster of Clostera
species (Pygaerinae) has a few stout terminal hooks, each bearing tiny setae that
Mosher (191 8b) noted are easily destroyed and difficult to see.

Body shape:

Body dorso-ventr ally compressed. (23) Both Dyar (1891) and Packard (1895) noted
that the body of Cerura pupae is slightly flattened dorso-ventrally. Packard (1895)
also applied that description to the pupae of Gluphisia species. He did not, however,
use the character as evidence of relationship between the two genera. Although not
particularly well-marked in F. scolopendrina (Fig. 7B), the trait is readily apparent
in other Furcula species (Dyar, 1891) as well as in Cerura, and it can easily be seen
in Gluphisia septentrionis (Fig. 8B).

Body wide relative to its length. (24) There is considerable variation in body pro-
portion among the species I studied. I recognized two character states for this trait:
in the majority of taxa the body is elongate, while in three taxa — Thaumetopoea,
Furcula and Gluphisia (Figs. 3, 7, 8)— the body is wide relative to its length.

CLADISTICS OF PUPAL DATA

Analysis of the data matrix in Table 3, with all nine multistate characters except
proboscis length and antenna length (Characters 3 and 5) treated as non-additive,
produced a single tree (Fig. 33). There are several important features of that clado-
gram. First, taxa that I claimed are related based on adult and larval morphology
(Miller, 1991) also group together when pupal characters are used. This is a significant
result; some of my previous conclusions— such as placement of Gluphisia with Fur-
cula, and Crinodes with Goacampa — might be considered controversial.

Second, the subfamily tree based on pupae (Fig. 34) corresponds well with previous
hypotheses. For example, two pupal synapomorphies (Characters 16 and 21) unite

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NOTODONTID PUPAE

251

ij= Doa

| — .[= Thaumetopoea

n= Clostera
■ n= Phalera

— Datana

■=\ [pGluphisia

I. — Furcula

— Notodonta

i— n= Crinodes

Goacampa

=4 [j= Lochmaeus

I, IL~ Heterocampa

- ij= Platychasma

— r?= Didugua

I, IL~ Symmerista

~ ir= Josia

Zunacetha

Fig. 33. Cladogram produced by a Hennig86 analysis of 24 pupal characters for 16 species
of Notodontidae (see Tables 1, 2) with Doa ampla as an outgroup. Length = 52, Cl = 71, RI
= 80.

the Dioptinae and Nystaleinae, a theory supported by adult and larval data (Weller,
1989; Miller, 1991). Comparison of my adult/larval tree (Fig. 1) with the one from
pupae in fact shows only two differences: In the pupal cladogram (Fig. 34), the
Phalerinae appear two nodes lower than before, while the subfamily Platychasmatinae
has moved higher, appearing as the sister group to the Dioptinae + Nystaleinae.

To further test these comparisons, I combined the 174 characters from my adult/
larval matrix (Miller, 1991: appendix IV) for the 17 test species with the 24 pupal
characters (Table 3). This produced a single data matrix, containing 198 characters,
derived from all three life stages. Analysis of that matrix using Hennig86 and implicit
enumeration again gave a single tree (Fig. 35). According to this cladogram, the
position of the Phalerinae suggested by pupal morphology (Fig. 34) may be the correct
one. On the other hand, the combined analysis supports placement of the Platy-
chasmatinae as suggested by adult and larval characters (Fig. 1); rather than being a

THAUMETOPOEINAE

PYGAERINAE

PHALERINAE

NOTODONTINAE

DUDUSINAE

HETEROCAMPINAE

PLATYCHASMATINAE

NYSTALEINAE

DIOPTINAE

Fig. 34.
Figure 33.

Relationships among notodontid subfamilies as implied by the cladogram in

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

j= Doa

{= Thaumetopoea
rp Clostera

n= Platychasma

ij= Phalera
[f~~~ ■ Datana

" ff= Gluphisia

ll L Furcula

I. IL^ Notodonta
1 ■■ . n= Crinodes

IL- Goacampa

rp Lochmaeus

h Heterocampa

p Didugua
II ==^= Symmerista
[j= Josia
■■■ - Zunacetha

Fig. 35. Cladogram produced by a combined matrix of adult, larval, and pupal characters
(total = 198; see text) for 16 species of Notodontidae with Doa ampla as an outgroup. Length
= 367, Cl = 62, RI = 62.

derived group, the Platychasmatinae is the sister-group of all notodontid subfamilies
exclusive of the Thaumetopoeinae and Pygaerinae.

Perhaps the most important result of my research on pupal morphology is the
discovery of potential synapomorphies for notodontid subfamilies, summarized in
Table 4. Clearly, all of my conclusions will be subject to change as pupae from
additional notodontid taxa are studied and as more is learned concerning morphology
of this life stage.

KEY TO THE PUPAE OF NOTODONTID SUBFAMILIES

Mosher (1918b: 58-60) gives a key to pupae for six North American notodontid
genera representing five subfamilies. The key below treats all nine of the currently
recognized subfamilies (see Miller, 1991). However, the user should be reminded
that it is based on relatively few taxa. In particular, pupae of Neotropical and Old
World species may not key out successfully.

1. Proximo-lateral angles of proboscis extending to eye-pieces (Figs. 5, 6, 8-12) 4

Proximo-lateral angles of proboscis not extending to eye-pieces (Figs. 3, 4, 7) 2

2(1). Cremaster present; eye-piece not small 3

Cremaster absent; eye-piece small Furcula, Cerura (Notodontinae; Fig. 7)

3(2). Labial palpi absent; abdominal cuticle covered with fine wrinkles; abdominal spir-
acles small; body wide; cremaster composed of two horns . . Thaumetopoeinae (Fig. 3)
- Labial palpi present; abdominal cuticle covered with small pits; abdominal spiracles

large; body elongate; cremaster a single elongate spine with hooks at tip

Pygaerinae (Fig. 4)

4(1). Anterior suture of proboscis not strongly curved, arching only slightly forward at

midline, proximo-medial angles of proboscis not acute (Figs. 5-8) 5

- Anterior suture of proboscis strongly curved, arching sharply forward at midline,

proximo-medial angles of proboscis acute (Figs. 9-12) 7

5(4). Antennae extending beyond mesothoracic legs; proboscis as long as wings; forewing
venation quadrifid; cremaster with more than 20 hook-shaped setae

Platychasmatinae (Fig. 5)

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253

Table 4. Summary of potential subfamily synapomorphies from pupal morphology. Char-
acters are based on study of the species listed in Table 1.

Thaumetopoeinae: (1) Labial palpus sclerite absent; (2) cremaster setae absent; (3) body wide
relative to its length.

Pygaerinae: (1) Cremaster long and spine-shaped, with stout hooks at tip.

Platychasmatinae: (1) Forewing venation quadrifid; (2) Antennae extending beyond mesotho-
racic legs; (2) Dorsum of A 1 0 with numerous, fine longitudinal ridges.

Notodontinae: (1) Terminal segments of abdomen broadly rounded.

Phalerinae: (1) Abdomen covered with large, deep pits; (2) Anterio-dorsal margin of A10 with
a deep furrow; (3) Cremaster setae absent.

Dudusinae: (1) Dorsum of thorax with a distinct crest along midline; (2) Caudal margin of
mesothorax with a series of shallow depressions separated by longitudinal striations; (3)
Anterior margin of each movable segment with a distinct carina.

Heterocampinae: (1) Caudal margin of mesothorax with a series of small, quadrangular plates
separated by deep pits; (2) Anterior margin of each movable abdominal segment ringed with
a diffuse band of small pits; (3) Dorsum of A10 with a reticulate area anterior to cremaster;
(4) Cremaster fork-shaped, lacking setae.

Nystaleinae: (1) Labial palpus sclerite present; (2) Thoracic cuticle covered with deep pits; (3)
Anterior margin of each movable abdominal segment ringed by a single row of lunate pits.

Dioptinae: (1) Thorax long, equal to % total body length; (2) Caudal margin of mesothorax
projecting posteriad at body midline; (3) Antennae extremely long, often touching A5; (4)
Abdominal spiracles small; (5) Cremaster broad, compressed.

- Antennae not extending beyond mesothoracic legs; proboscis shorter than wings;

forewing venation trifid; cremaster with fewer than 10 setae or with setae absent
(Figs. 6-12) 6

6(5). Terminal abdominal segments (A8-A10) broadly rounded; abdomen covered with
small pits; dorsum of A 10 unmodified; cremaster often reduced or absent, but when
present setae occur Notodonta, Gluphisia (Notodontinae; Fig. 8)

- Terminal abdominal segments tapered; abdomen covered with large, deep pits; dor-
sum of A 10 with a deep furrow; cremaster a series of spines, setae absent

Phalerinae (Figs. 6, 26)

7(4). Caudal margin of mesothorax strongly sculpted; abdominal spiracles facing laterally;
base of cremaster lacking flutes on ventral surface; labial palpi absent (Figs. 9, 10)
8

- Caudal margin of mesothorax not strongly sculpted; abdominal spiracles facing

posteriorly ; base of cremaster with longitudinal flutes on ventral surface; labial palpi
often present (Figs. 11,12) 9

8(7). Caudal margin of mesothorax with a series of shallow depressions separated by
raised ridges; thorax with a dorsal crest along midline; anterior margin of each
movable abdominal segment (A5-A7) with a carina Dudusinae (Figs. 9, 14, 23)

- Caudal margin of mesothorax with a series of small, quadrangular plates separated

by deep pits; thorax without a dorsal crest; anterior margin of each movable ab-
dominal segment with a diffuse band of small pits, no carina

Heterocampinae (Figs. 10, 15-18, 21, 27)

9(7). Surface of thorax deeply pitted; thorax not elongate; mesothorax unmodified; anterior
margin of each movable segment with a single row of lunate pits; proboscis only as

long as wings; cremaster bifid, not dorso-ventrally compressed

Nystaleinae (Figs. 11, 19, 22, 28)

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Surface of thorax with a few small pits or smooth; thorax elongate, % the length of
abdomen (ventral view); anterior margins of movable segments unmodified; pro-
boscis often longer than wings, touching A5; cremaster dorso-ventrally compressed,
not bifid Dioptinae (Figs. 12, 30-32)

DISCUSSION

According to Mosher, “the pupae of [Notodontidae] vary considerably, and there
is no one character which will serve to separate them from those of the nearly related
families” (1918b:58). None of the diagnostic characters she listed for the Notodon-
tidae (Mosher, 1 9 1 6b, 1 9 1 8b) adequately define the group; most are plesiomorphic
traits. Both Mosher and Common (1990) emphasized reduction or loss of labial palpi
as definitive (see Character 2). Unfortunately, this character is extremely variable
among families of Lepidoptera. For example, within the Noctuoidea labial palpi are
present in Noctuidae but absent in Arctiidae (Mosher, 1916b). Minet (1991) used
reduction of labial palpi in pupae to define a large ditrysian clade that includes the
Geometroidea, Hedyloidea, Hesperioidea, and Papilionoidea.

Another pupal trait that Mosher (1916b, 1918b) and Common (1990) stressed in
their family diagnoses is presence of punctures on the abdomen. I found, however,
that these punctures are absent in Thaumetopoea pinivora (representing the subfamily
Thaumetopoeinae; see Character 14), so the trait cannot be used to define the No-
todontidae. None of the other characters listed by either Mosher or Common can
be used as potential family-level synapomorphies.

My own study does little to alleviate this problem; I was unable to find a pupal
character unique to the Notodontidae. I can only recommend that future researchers
pay attention to morphological detail in the search for a synapomorphy in pupae. If
one is to be found, it will not likely involve an obvious feature of the external anatomy.
However, lack of a synapomorphy from pupae does not call monophyly of the group
into question; there are numerous synapomorphies from adults and larvae (Miller,
1991).

Notwithstanding the problems outlined above, research on pupae makes clear the
potential of this life stage for understanding the phylogeny and systematics of the
Notodontidae. I have identified pupal traits that elucidate relationships among sub-
families (e.g., Characters 1, 3, 4), as well as traits that help define subfamilies (Table
4). Future research should focus on the pupae of notodontid groups, such as the
Hemiceratini, whose positions in the family classification remain enigmatic (see
Miller, 1991). Perhaps characters from pupae will provide answers that have not
been forthcoming from the study of adults and larvae.

ACKNOWLEDGMENTS

I thank the following people for the loan of pupal material: George Godfrey (Illinois Natural
History Survey), Rick Hoebeke (Cornell University Insect Collections), Ian Kitching (The
Natural History Museum, London), Tim McCabe (New York State Museum), Alma Solis
(United States National Museum), Shigero Sugi (Tokyo, Japan), and Susan Weller (Louisiana
State University, Baton Rouge). Andre Sharkov (AMNH) kindly translated the papers of Do-
linskaya from Russian into English, Peling Fong (AMNH) assisted with scanning electron
microscopy, and Alima Edwards (AMNH) helped with manuscript preparation. I am grateful

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NOTODONTID PUPAE

255

to Caroline Michalski (AMNH) and Bob Poole (USNM) for comments on the manuscript. This
research was supported by a Kalbfleisch Curatorial Fellowship from the AMNH. I would
especially like to acknowledge J. G. Franclemont, who has been an inspiration to me since my
first days as a graduate student.

LITERATURE CITED

Bolte, K. B. 1990. Guide to the Geometridae of Canada (Lepidoptera). VI. Subfamily Laren-
tiinae. I. Revision of the genus Eupithecia. Mem. Entomol. Soc. Can. 151:1-253.

Brown, J. W. 1990. The early stages of Doa dora Neumoegen and Dyar (Lepidoptera: Noc-
tuoidea: Doidae) in Baja California, Mexico. J. Res. Lepid. 28:26-36.

Chapman, T. A. 1892. The emergence of the Ceruras. Entomologist 25:302-304.

Chapman, T. A. 1893. Some neglected points on the pupae of the heterocerous Lepidoptera.
Trans. Entomol. Soc. London 1893:97-119.

Chapman, T. A. 1896. On the phylogeny and evolution of the Lepidoptera from a pupal and
oval standpoint. Trans. Entomol. Soc. London 1896:567-587.

Common, I. F. B. 1975. Evolution and classification of the Lepidoptera. Annu. Rev. Entomol.
20:183-203.

Common, I. F. B. 1990. Moths of Australia. E. J. Brill, Leiden.

Dolinskaya, I. V. 1989. Chrysalid morphology of certain Far Eastern notodontid moth species
(Lepidoptera: Notodontidae). Vestn. Zool. 1989:60-67. [In Russian; transl. by Andre
Sharkov, AMNH]

Donahue, J. P. and J. W. Brown. 1987. Doidae (Noctuoidea). Pages 534-536 in: F. W. Stehr
(ed.), Immature Insects. Kendall/Hunt, Dubuque, Iowa.

Dyar, H. G. 1889. Preparatory stages of Janassa lignicolor, Walker. Entomol. Amer.
5:91-92.

Dyar, H. G. 1891. Preparatory stages of Datana perspicua, G. & R., and Cerura borealis,
Boisd. Canadian Entomol. 23:82-87.

Farris, J. S. 1988. Hennig86 reference. Version 1.5. Port Jefferson Station, New York.

Ferguson, D. C. 1 963. Immature stages of four Nearctic Notodontidae (Lepidoptera). Canadian
Entomol. 95:946-953.

Forbes, W. T. M. 1948. Lepidoptera of New York and neighboring states. Part 2. Notodon-
tidae. Cornell Agric. Exp. Stn. Mem. 274:203-237.

Forbes, W. T. M. 1954. Lepidoptera of New York and neighboring states. Part 3. Noctuidae.
Cornell Agric. Exp. Stn. Mem. 329:433 pp.

Franclemont, J. G. 1960. Nolidae. Pages 50-55 in: W. T. M. Forbes (ed.), Lepidoptera of
New York and neighboring states. Part 4. Agaristidae through Nymphalidae including
Butterflies. Cornell Agric. Exp. Stn. Mem. 371:188 pp.

Franclemont, J. G. 1983. Arctiidae. Pages 1 14-1 19 in: R. W. Hodges et al. (eds.), Check List
of the Lepidoptera of America North of Mexico. E. W. Classey Ltd. and The Wedge
Entomological Research Foundation, London.

Griinberg, K. 1912. Notodontidae. Pages 284-319 in: A. Seitz (ed.), Macrolepidoptera of the
World. Vol. 2. Alfred Kemen, Stuttgart.

Hinton, H. E. 1946. A new classification of insect pupae. Proc. Zool. Soc. London 1 1 6:282—
328.

Jackson, W. H. 1890. Studies in the morphology of the Lepidoptera. Part I. Trans. Linn. Soc.
London, Zool. Ser. 2, 5:143-186.

Kafatos, T. C. and C. M. Williams. 1964. Enzymatic mechanism for the escape of certain
moths from their cocoons. Science N.Y. 146:538-540.

Kitching, I. J. 1984. An historical review of the higher classification of the Noctuidae (Lep-
idoptera). Bull. Br. Mus. (Nat. Hist.) Entomol. 49:153-234.

Marumo, N. 1920. A revision of the Notodontidae of Japan, Corea and Formosa with de-

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scriptions of five new genera and five new species. J. Coll. Agric. Imp. Univ. Tokyo 4:
273-359.

McGuffin, W. C. 1987. Guide to the Geometridae of Canada (Lepidoptera). II. Subfamily
Ennominae. 4. Mem. Entomol. Soc. Can. 138:182 pp.

Miller, J. S. 1987. A revision of the genus Phryganidia Packard, with description of a new
species (Lepidoptera: Dioptidae). Proc. Entomol. Soc. Washington 89:303-321.

Miller, J. S. 1991. Cladistics and classification of the Notodontidae (Lepidoptera: Noctuoidea)
based on larval and adult morphology. Bull. Am. Mus. Nat. Hist. 204:230 pp.

Miller, J. S. 1992. Host-plant associations among prominent moths. BioScience 42:50-57.
Minet, j. 1991. Tentative reconstruction of the ditrysian phylogeny (Lepidoptera: Glossata).
Entomol. Scand. 22:69-75.

Mosher, E. 1914. The classification of the pupae of the Ceratocampidae and Hemileucidae.
Ann. Entomol. Soc. Am. 7:277-300.

Mosher, E. 191 6a. The classification of the pupae of the Satumiidae. Ann. Entomol. Soc. Am.
9:136-156.

Mosher, E. 1916b. A classification of the Lepidoptera based on characters of the pupa. Bull.
111. St. Lab. Nat. Hist. 12:14-159.

Mosher, E. 1918a. Pupae of common Sphingidae of Eastern North America. Ann. Entomol.
Soc. Am. 1 1:403-442.

Mosher, E. 1918b. Pupae of some Maine species of Notodontoidea. Maine Agr. Exp. Sta.
Bull. 259:29-84.

Nagano, K. 1916. Life-history of some Japanese Lepidoptera containing new genera and
species. Bull. Nawa Entomol. Lab. 1:1-27.

Nakamura, M. 1956. Contribution to the knowledge of some Japanese notodontid moths.
Revisional notes X. Tinea 3:142-143.

Nakamura, M. 1976. Pupae of Japanese Lymantriidae (Lepidoptera). Kontyu (Tokyo) 44:
411-434.

Nielsen, E. S. 1 989. Phylogeny of major lepidopteran groups. Pages 28 1-294 in: B. Femholm,
K. Bremer and H. Jomvall (eds.), The Hierarchy of Life. Elsevier, Amsterdam.
Packard, A. S. 1895. Monograph of the bombycine moths of America north of Mexico
including their transformations and origin of the larval markings and armature, part I:
family 1, Notodontidae. Mem. Nat. Acad. Sci. 7:390 pp.

Poulton, E. B. 1891. The external morphology of the lepidopterous pupa: its relation to that
of the other stages and to the origin and history of metamorphosis. Parts I-V. Trans.
Linn. Soc. London, Zool. Ser. 2, 5:187-212; 245-263.

Rindge, F. H. 1952. Taxonomic and life history notes on North American Eupithecia (Lep-
idoptera, Geometridae). Am. Mus. Novitates 1569:27 pp.

Weller, S. J. 1987. Litodonta hydromeli Harvey (Notodontidae): description of life stages. J.
Lepid. Soc. 41:187-194.

Weller, S. J. 1989. Phylogeny of the Nystaleini (Lepidoptera: Noctuoidea: Notodontidae).
Ph.D. thesis, Univ. Texas, Austin, 396 pp.

Received 23 August 1991; accepted 22 November 1991.

J. New York Entomol. Soc. 100(2):257-266, 1992

FRANCLEMONTIA INTERROGANS (WALKER), A
NEW GENUS FOR AN OLD SPECIES
(LEPIDOPTERA: NOCTUIDAE: AMPHIPYRINAE)

Douglas C. Ferguson

Systematic Entomology Laboratory, Agricultural Research Service, USD A,
%National Museum of Natural History, NHB-168, Washington, D.C. 20560

Abstract.— The identity of Leucania interrogans Walker, 1856, described from an unknown
type locality, is confirmed, and a lectotype is designated for it. It is a distinctive and rarely
collected species of amphipyrine noctuid of the southeastern U.S. that belongs to neither Leu-
cania nor Phragmatiphila, where placed by previous authors. A comprehensive search failed
to reveal any genus to which interrogans might be referred, and Franclemontia, new genus, is
proposed for it. The possible sister-group relationship of this new genus to atypical North
American elements of Chortodes is suggested. All members of this large group of amphipyrine
genera are believed to be host-specific stem borers in marsh grasses.

Walker (1856:1 14) described Leucania interrogans from two specimens from the
Milne collection in the British Museum but gave no type locality. Hampson (1910:
270) transferred interrogans to his newly described genus Phragmatiphila and pro-
vided an illustration of one of the syntypes, but their country of origin remained
unknown for about a century. John G. Franclemont, while stationed at Fort Rucker,
Ozark, Dale Co., Alabama, in the early 1940’s, collected a specimen that he subse-
quently recognized as P. interrogans. After Hampson, however, no mention of the
name is also in the 1983 check list (Franclemont and Todd, in Hodges et al., 1983).
The Milne collection is known to have contained material purchased from John
Abbot (Doubleday [1839], in Scudder, 1869:123), and Abbot was about the only
source of moths from the southeastern United States prior to Doubleday’s stay at
St. Johns Bluff, Duval County, Florida, in 1838. Georgia is therefore the probable
type locality.

Few have been collected, probably no more than eight or ten specimens, including
the two types in the Natural History Museum, London (BMNH). A male and female
(Figs. 3, 4) from The Wedge Plantation, McClellan ville, Charleston County, South
Carolina, are deposited in the National Museum of Natural History (USNM). Others
include the specimen collected by Franclemont at Fort Rucker, Alabama and another
from McClellanville, South Carolina (Cornell Univ. Coll.); and one from near Pen-
sacola, Escambia County, Florida that was reported by Kimball (1965: 103) from the
Shirley M. Hills collection, and probably identified for Kimball by Franclemont. The
present location of the Hills collection could not be determined. I have been advised
of two other specimens in private collections but have not seen them. The species
remained in the genus Phragmatiphila from 1910 to the present, although it bears
little resemblance to the type species of that genus, the European Phragmatiphila
nexa (Hiibner) (Figs. 5-7, 12).

Phragmatiphila nexa appears to be more closely related to species of the North
American genus Meropleon Dyar, which I revised earlier (1982). Although the male

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

genitalia of the two genera agree closely, the female genitalia differ enough to dissuade
me from synonymizing them. Also, P. nexa has a conspicuous, s-shaped, white discal
spot on the forewing not present in any species of Meropleon. A similarly shaped
but less conspicuous s-shaped discal spot is present in P. interrogans, however, and
this may be why Hampson associated the species with Phragmatiphila.

A review of similar moths worldwide, including other species of Phragmatiphila
as now understood (Poole, 1989, vol. 2:800) failed to reveal anything that could be
considered congeneric with interrogans. In the event that such a species exists, it to
would need a generic name. I establish the following genus to accommodate this rare
and little-known species.

Franclemontia, new genus

Type species: Leucania interrogans Walker, 1856. Monotypic.

The features that distinguish this genus are summarized in the accompanying list
(Fig. 1), and the following notes will explain them:

1. Franclemontia interrogans (Figs. 2-4, 8-11) is unique among the genera to
which it seems most closely related in having wide, bipectinate male antennae.
Although I mentioned (1982:231) that Meropleon titan has bipectinate male
antennae with short branches, they are really more lamellate than pectinate.
Franclemontia has true bipectinate male antennae with branches whose length
may equal nearly 4 times the thickness of the antennal shaft.

2. Neither F. interrogans nor species of Meropleon, or any other species examined,
have the sharp, bladelike, transverse frontal process that distinguishes P. nexa,
and which is visible even without removing the frontal vestiture. Species of
most genera have a somewhat convex, rounded front, although the “ Chor -
todes ” species that I listed as Group B have an obtusely conical front with a
decumbent, pointed apex.

3. Phragmatiphila nexa (Fig. 5) has a conspicuous, s-shaped or z-shaped reniform
spot somewhat similar in shape to the less distinct reniform of F. interrogans.

4. The male of interrogans has a pair of eversible scent glands with coremata or
long hair pencils, each contained in a lateral pocket at the base of the abdomen
(Fig. 10). These recessed bundles of hair extend for the length of the first three
abdominal segments. No similar structures were found to be present in any
of the related grass-feeding amphipyrine genera investigated, although they
appear elsewhere in the Amphipyrinae.

5. The uncus is flattened and widest in the middle, tapering both toward the apex
and base. Phragmatiphila nexa is the only other species examined that has
this type of uncus. That of the Australian Bathytricha truncata Turner is also
wide but not tapered.

6. The anellus is thick and leathery and finely and densely scobinate over much
of its surface in Phragmatiphila and Meropleon, but not so in Franclemontia,
Chortodes, or other genera studied except Bathytricha from Australia.

7. The juxta is deeply notched or v-shaped in Franclemontia and Phragmatiphila
but not in other genera. “ Chortodes ” Group B is unusual in having the juxta
shaped like a spear point or arrowhead.

1992

FRANCLEMONTIA INTERROGANS

259

Chortodes

Chortodes

Francl

Phrag.

Meropleon

Group A

Group B

1

Bipectinate
c? antenna

+

-

-

-

-

2

Frontal process
absent

+

-

+

+

-

3

S-shaped

reniform

+

+

-

-

-

4

Hair pencils at
base of abdomen

+

-

-

-

-

5

Uncus broad
in middle

+

+

-

-

-

6

Anellus not leathery
or scobinate

4-

-

-

+

+

7

Juxta notched
or V-shaped

+

4-

-

-

8

Cucullus developed
(may be rudimentary)

+

-

-

+

+

9

Ampulla

+

+

-

+

+

10

Digitus

+

-

-

-

+

11

Large costal sclerite
and costal process

+

+

+

-

+

12

Sacculus swollen, with
large costal lobe

+

-

-

-

+

13

Clavus a long,
pointed process

+

-

-

-

-

14

Aedoeagus with ventro-
distal serrate ridge

+

+

-

-

+

15

£ genitalia short,
stout

+

-

-

-

+

16

Ovipositor lobes
straight, pointed

+

-

-

-

-

Fig. 1 . Selected characters of Franclemontia and related genera. Plus and minus signs in-
dicate presence or absence. Francl. = Franclemontia ; Phrag. = Phragmatiphila (others spelled
out).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Figs. 2-5. Adult moths. 2. Franclemontia interrogans (Wlk.) 9. Lectotype (BMNH). 3. F.
interrogans 9. Wedge Plantation, McClellanville, S.C., 29 April 1973, R. B. Dominick (USNM).
4. F. interrogans 6, same locality, 7 April 1977, D. C. Ferguson (USNM). 5. Phragmatiphila
nexa $. Germany (USNM). Magnification: 2x.

8. The valve of the male genitalia of Franclemontia has a fairly well-formed
cucullus, although no coronal spines are present. However, every stage from
no cucullus to a fully developed triangular one may elsewhere be found within
one genus (e.g., Chortodes).

9. A small ampulla is present in F. interrogans and P. nexa, although that of
nexa is minute.

10. A digitus, or digitus-like process, occurs in Franclemontia, although elsewhere
in this immediate group a possible homologue may be seen only in one of the
two species of “Chortodes” Group B (“C.” enervata (Guenee)).

1 1 . The large, bowed, concave, costal sclerite of the valve, which is an especially
conspicuous feature of all species of Meropleon, is similar in P. nexa. It is
present in modified form in F. interrogans and Chortodes enervata. The costal
sclerite, when present, has a pointed or scoop-shaped free end Vi to % of the
way out the costa. In F. interrogans this free process is about halfway out and
takes the form of a triangular costal process of unique form, as may be seen

1992

FRANCLEMONTIA INTERROGANS

261

Figs. 6-10. Male genitalia. 6. Phragmatiphila nexa (Germany). 7. Same specimen, aedoea-
gus. 8. Franclemontia interrogans, genitalia of specimen shown in Figure 4. 9. Same specimen,
aedoeagus. 10. Same specimen, abdominal sterna 1-4, showing eversible structures with hair
pencils (Character 4).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

in Figure 8. The j -shaped binary sclerite at this site in “C.” enervata is probably
derived from a fusion of the pointed costal sclerite and an enlarged digitus.

12. In Franclemontia the sacculus is swollen and rounded, forming a costal lobe
almost reaching the costobasal angle of the valve. The sacculus of “Chortodes”
enervata is similar, but without the clavus.

1 3. Franclemontia interrogans is unusual in having a clavus in the form of a long,
sharp, thornlike process arising from the costal lobe of the sacculus. These
processes lie in the same plane as the juxta and flank it, one on each side. No
other American members of the group have a clavus.

14. The aedoeagus of Franclemontia has 3 large, acute, dentate processes (comuti)
on the vesica, and 5 smaller ones on the distal end of the aedoeagus, clustered
in two groups subventrally. Although I equated those on the aedoeagus with
the similarly situated, longitudinally serrate ridge of Phragmatiphila nexa, it
is doubtful whether the two structures are homologous. Those of Francle-
montia suggest Meropleon and the arrangement in P. nexa is found otherwise
only in the species of “ Chortodes ” Group B.

15. The female genitalia are unusually shortened longitudinally, from ovipositor
to and including the corpus bursae, and have a short, thick, rugose “neck” on
the bursa copulatrix. The female genitalia of “Chortodes” defecta are surpris-
ingly similar in almost every detail.

1 6. The ovipositor lobes of the female genitalia are straight and pointed, not blunt
and bent somewhat ventrad as in most other genera of the group. In P. nexa
they are rounded at the tips but not downcurved.

The genus Chortodes Tutt is a holarctic group treated in North American check
lists as Hypocoena Hampson, but recently regarded as a senior synonym of Hypocoena
(Kononenko, Lafontaine and Mikkola, 1 989). However, the American species include
at least two genera, which explains why I list them as Groups A, Chortodes sensu
stricto, and B, “ Chortodes .” Group B includes C. enervata (Guenee) and C. defecta
of authors (not Grote). Group A consists of all others, including the type species, C.
morrisii Dale.

Franclemontia interrogans (Walker), New Combination

Figs. 2-4, 8-10

Leucania interrogans Walker, 1856:1 14.

Phragmatiphila interrogans Hampson, 1910:270, pi. 143, fig. 3; Kimball, 1965:103;
Franclemont and Todd, in Hodges et al., 1983:138.

Type locality: Not given [probably Georgia— see earlier comment].

Types: Described from a male and female, of which the female was illustrated by
Hampson (1910, pi. 143, fig. 3) and is figured again in this paper (Fig. 1). I hereby
designate this female syntype as the lectotype. The male syntype cannot now be
found, although its genitalia are preserved in a vial in the collection of The Natural
History Museum, London.

Description: Although the generic description serves also to characterize the species,
inasmuch as there is only one, I offer the following supplementary information to
eliminate any doubt that might remain concerning the identity of the species. Male

1992

FRANCLEMONTIA INTERROGANS

263

Figs. 11, 12. Female genitalia. 1 1. Franclemontia interrogans, genitalia of specimen shown
in Figure 3. 12. Phragmatiphila nexa (Sweden).

antenna with longest pectinations equal to more than 3 times diameter of antennal
shaft; female antenna simple. Eyes large, very slightly elliptical, protuberant, equal
to full hemisphere in female, slightly more than hemisphere in male. Palpi long,
porrect or somewhat upturned; second segment as long as eye, third segment Vi as
long as second in both sexes (palpal vestiture mostly worn off in specimens examined).
Frons rounded and bulging, but not conical or ridged. Proboscis only slightly reduced,
about as well developed as those of Chortodes species (described as rudimentary in
potentially related African genera). Legs long and normally developed, with long
hindtibial spurs, the longest being slightly longer than the second palpal segment;
hind leg slightly shorter and with tibia more swollen in female; legs roughly and
loosely scaled, with thin posterior fringe of longer, hairlike scales; epiphysis reaching
almost to end of foretibia. Vestiture of vertex and thorax consisting of long hairlike
scales, but with some broad scales concealed beneath; thorax and abdomen without
special scale tufts or crests.

Hindwing with vein M2 obsolescent. Shape of male and female forewings differ;
that of male normal except that outer margin is noticeably angled at vein M3; female
forewing not angulate on outer margin but with costa unusually convex or arched.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Forewings pale brown, the usual dead-grass color of many grass-feeding moths, with
slightly darker, predominantly longitudinal shades, and the following markings: a
pale, somewhat indefinitely s-shaped or z-shaped reniform with an elongated dark
center, bending distad in direction of outer margin at end near costa and basad at
end nearest inner margin; the latter end tapering off very acutely as a thin, pale-
edged, dark streak directed toward wing base; a dark-brown dot, ringed by a paler
shade, at position of orbicular; a small, black, basal dot; and a weak, sinuous, dark,
postmedial line broken into a series of dots or dashes (antemedial line obsolete).
Hindwing almost unicolorous gray brown in male; slightly paler, more variegated,
and with a distinct, dusky, sinuous, transverse band in female. Undersurfaces with
forewing darker than hindwing but with contrastingly paler costa and outer margin,
at least in female; diffuse, dusky, discal spots on both wings, that on hindwing of
female preceded by a thin, short, longitudinal dark streak; hindwing also with weak,
dark, convex postmedial line, especially apparent in female. Terminal line a broken
series of dark crescents or dashes on both upper- and undersurfaces of both wings.
Length of forewing: <5, 15 mm (N = 1); $, 15-16 mm (N = 2).

Genitalia. Discussed under genus and illustrated. The male (Figs. 8, 9) is unusual
in many respects, including the possession of what appears to be a small digitus, a
structure normally absent in members of this group. The digitus in this case, however,
may be an ampulla that has migrated to a different position. The female genitalia
(Fig. 1 1), with their short, stout overall aspect and wide, straight, rigid, rugose neck
region of the bursa copulatrix, would be unique were they not almost identical to
the female genitalia of “Chortodes” defecta (Grote), previously known as Hypocoena
defecta (Grote).

Distribution. SOUTH CAROLINA: Wedge Plantation, near McClellanville,
Charleston Co. FLORIDA: near Sarasota, Escambia Co. (Kimball, 1965). ALA-
BAMA: Fort Rucker, near Ozark, Dale Co. (Franclemont Coll., Cornell Univ.). The
types are probably from Georgia, but this is speculative.

Flight period. The only collection dates available are 7, 15 and 21 April for South
Carolina; 6 April for Florida; and 5 April for Alabama. The species is undoubtedly
univoltine, as are probably all grass-boring Amphipyrinae in North America.

Habitat. The South Carolina specimens were taken at light near old, abandoned,
plantation rice fields (now brackish) at the edge of the vast Santee Delta marshes,
on the south side of the South Santee River about 8 km (5 miles) upstream from the
coast. I would therefore have guessed that the species is associated with some select
microhabitat in salt or brackish estuarine marshland; the Florida site could have
been similar. However, the locality in Alabama is not coastal.

DISCUSSION

Each of the genera herein used for comparison (Fig. 1) has other distinguishing
features not mentioned. For example, Phragmatiphila nexa is the only member of
the complex with a signum, which is small and depicted edgewise on the right side
of the bursa in Figure 12. Species of Meropleon are consistent in having a unique
fringe of persistent (hard-to-remove) scales on the posterior margin of the seventh
sternum of the female abdomen. Males of both species of “Chortodes” Group B have
a distinctive juxta that is shaped like an arrowhead or spear point.

1992

FRANCLEMONTIA INTERROGANS

265

I also examined genitalia, or published illustrations of genitalia, representing a
wide array of other genera of amphipyrine noctuids whose species bore in grass stems,
indeed, all that could be identified as containing species that might be related to F.
interrogans. Species of the following genera were included in this survey, in most
cases the type species: Archanara Walker, Chortodes Tutt ( =Hypocoena Hampson),
Spartiniphaga McDunnough, Ommatostola Grote, Benjaminiola Strand, Mammi-
frontia Barnes and Lindsey (North America); Nonagria Ochsenheimer, Photedes
Lederer, Rhizedra Warren (Europe); Acrapex Hampson, Busseola Thurau, Carelis
Bowden, Conicofrontia Hampson, Poecopa Bowden, Poeonoma Tams & Bowden,
Sciomesa Tams & Bowden, Sesamia Guenee, Speia Tams & Bowden (Africa, Asia),
and Bathytricha Turner (Australia).

I examined the genitalia of all species of Meropleon Dyar (Ferguson, 1982) and
Chortodes in related revisionary projects. Nonagria, widespread in the Old World,
is clearly more different. Many of the above-mentioned genera are African, and I
had thought that the sister group of Franclemontia might be found among them. For
example, Conicofrontia and Sciomesa show certain similarities to Franclemontia in
valve shape, and Sciomesa has pectinate male antennae. However, Franclemontia
does not fit any of those genera because of other important differences, such as the
hair pencils at the base of the male abdomen, the long thomlike clavus, and the short,
stout female genitalia. Keys and illustrations for the African genera were given by
Janse (1939), Tams and Bowden (1953) and Bowden (1956). The Australian genus
Bathytricha, of which I saw the male genitalia of a specimen believed to be B. truncata
(Walker) (the type species), kindly sent by T. L. McCabe, is also related to this group.
The valve (including character 1 1) is remarkably similar to that of Phragmatiphila
and Franclemontia, although the sclerites it bears are more similar to those of Chor-
todes enervata. It has bipectinate male antennae (character 1 ), a broad uncus (character
5), a scobinate anellus (character 6), a rudimentary cucullus (character 8), and a
cockscomb-shaped sclerite on the aedoeagus (character 14). However, characters 3,
4, 7, 9, 1 2, and 1 3 appear to be absent. In the end I concluded that the closest relatives
of Franclemontia are also North American, and almost certainly the species of Chor-
todes Group B.

Curiously, this whole complex of genera appears to be unrepresented in the neo-
tropics. For example, the only New World species of Acrapex, an otherwise Old
World tropical genus, is also found in the warm-temperate zone of eastern North
America, not in the American tropics (Ferguson, 1991).

My inspiration for the generic name Franclemontia needs little explanation. I was
one of Dr. Franclemont’s earlier graduate students, and one whose career in Lepi-
doptera systematics might never have happened had our paths not crossed. It is an
honor to me to be the one to name a genus after him, and I have tried to fit the
name to an appropriate moth.

ACKNOWLEDGMENTS

I thank R. W. Poole, a colleague in the Systematic Entomology Laboratory, for answering
queries concerning the world fauna and the possible relationships of Franclemontia ; and Linda
Pitkin and Martin Honey, the Natural History Museum, London, for sending on loan the type
material of Leucania interrogans Walker. I am indebted to the late R. B. Dominick of The
Wedge Plantation, whose hospitality and interest in Lepidoptera gave me access to one of the

266 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

few known habitats of this species and led to this research on it. I thank Richard L. Brown,

Ray Gagne, Ronald W. Hodges, and Timothy L. McCabe for reviewing the paper.

LITERATURE CITED

Bowden, J. 1956. New species of African stem-boring Agrotidae (Lepidoptera). Bull. Entomol.
Res. 47:415-428.

Ferguson, D. C. 1982. A revision of the genus Meropleon Dyar, with descriptions of a new
species and subspecies (Lepidoptera; Noctuidae). Entomography 1:223-235.

Ferguson, D. C. 1991. First report of a species of the genus Acrapex Hampson from the New
World, with description of a new species from the Carolinas and Virginia (Noctuidae:
Amphipyrinae). J. Lepid. Soc. 45:209-214.

Hampson, G. F. 1910. Catalogue of the Lepidoptera Phalaenae in the British Museum. Volume
9. Taylor & Francis, London, xv + 552 pp., pis. 137-147.

Hodges, R. W., et. al. 1983. Check List of the Lepidoptera of America North of Mexico. E.
W. Classey and The Wedge Entomological Research Foundation, London, xxiv +
284 pp.

Janse, A. J. T. 1937-40. The Moths of South Africa. Volume 3. Cymatophoridae, Callidulidae,
and Noctuidae (partim). E.P. & Commercial Printing Co., Ltd., Durban, xv + 435 pp.,
46 pis., 102 text figs.

Kimball, C. P. 1965. The Lepidoptera of Florida. Division of Plant Industry, Florida Dept,
of Agriculture, Gainesville, v + 363 pp., 26 pis.

Kononenko, V. S., J. D. Lafontaine and K. E. Mikkola. 1989. An annotated checklist of
noctuid moths (Lepidoptera, Noctuidae) of Beringia. Entomologicheskoye Obozreniye,
No. 3, pp. 549-567. Also published (in English) in 1990, Entomological Review 69:1 17-
138.

Poole, R. W. 1989. Noctuidae. Lepidopterorum Catalogus Fascicle 1 18. E. J. Brill/Flora and
Fauna Publications, Leiden, New York, etc. 1314 pp. in 3 vols.

Scudder, S. H. (ed.). 1869. Entomological correspondence ofThaddeus William Harris, M.D.
Occasional Papers Boston Soc. Nat. Hist. 1: 375 pp., illus.

Tams, W. H. T. and J. Bowden. 1953. A revision of the African species of Sesamia Guenee
and related genera (Agrotidae-Lepidoptera). Bull. Entomol. Res. 43:645-678.

Walker, F. 1856. List of the Specimens of Lepidopterous Insects in the Collection of the
British Museum. Volume 9. Published by order of the Trustees of the British Museum
by Edward Newman, London, 1-252 pp.

Received 3 July 1991; accepted 21 October 1991.

J. New York Entomol. Soc. 100(2):267-272, 1992

AN ANNOTATED KEY AND ILLUSTRATIONS TO THE
SPECIES OF THE GENUS CUCULLIA
(LEPIDOPTERA: NOCTUIDAE: CUCULLIINAE) FROM
MEXICO AND CENTRAL AMERICA

Robert W. Poole

Systematic Entomology Laboratory, PSI, ARS, USD A
% National Museum of Natural History, NHB-168,

Washington, D.C. 20560

Abstract.— A key to the Mexican and Central American species of the genus Cucullia is
presented. The key is annotated with the geographical distribution of each species and supple-
mented with illustrations of the adults of each species.

Jack Franclemont, among his many and diverse interests, has always had a special
affection for the species of the genus Cucullia Schrank. The species in Cucullia have
a characteristic long, pointed forewing and the patagia are raised into a pointed hood.
Although these two features are not unique to Cucullia, the species of the genus are
usually easily recognizable. The genus is primarily Holoarctic with the bulk of its
diversity in Europe, North America, and Asia. Species are relatively less well rep-
resented in Africa. The genus is strongly developed in Canada and the United States,
but much more poorly represented south of the U.S.-Mexican border. The south-
ernmost known extension of the genus is in Costa Rica. Only a single species occurs
in Mexico and Central America that is not found in the United States, Cucullia
costaricensis Hampson. I have recently completed a revision of the genus Cucullia
which will soon appear as part of a fascicle in the Moths of America North of Mexico
series (Poole, 1992). The purpose of this paper is to complete the treatment of the
New World species of the genus by presenting an annotated key and illustrations to
the species known or strongly suspected to occur in Mexico and Central America.
This paper is meant to supplement the Moths of America North of Mexico fascicle.
Color illustrations of all of the species (except costaricensis ) along with full discussions
and descriptions of the species, and photographs of the male and female genitalia
may be found there. References to original descriptions and dates of publication may
also be found in that publication or alternatively may be retrieved from Poole ( 1 989).

Many of the species treated are U.S. species which have not yet been collected
south of the border but probably occur in northern Mexico. I have not included those
species that are part of the Mohave Desert region fauna of southern California that
also occur in northern Baja California. The reader will want to refer to the Moths of
America North of Mexico fasicle for the identification of species of Cucullia from
this region of Mexico.

The following is a list of the species treated in this paper arranged by species groups
following the arrangement in the Moths of America North of Mexico fascicle.

asteris species group

Cucullia montanae Grote

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Cucu/lia postera Guenee
Cucullia obscurior Smith
convexipennis species group
Cucullia lilacina Schaus
Cucullia eucaena Dyar
Cucullia costaricensis Hampson
Cucullia oribac Barnes
speyeri species group
Cucullia new species #1
Cucullia laetifica Lintner
Cucullia new species #2
Cucullia new species #3
Cucullia eccissica Dyar
antipoda species group
Cucullia antipoda Strecker
strigata species group

Cucullia serraticornis Lintner

KEY TO THE SPECIES OF CUCULLIA

1 . Fore tibia with a distinct claw; front of head with large, triangular frontal process

antipoda

Cucullia antipoda Strecker (Fig. 1 6) has not been seen from Mexico, but is abundant
in the drier areas of the western United States and has been collected in southern
New Mexico and Arizona. It probably will be found in Chihuahua and Sonora.

- Fore tibia without a distinct claw; front of head without frontal process 2

2. Palpi studded with thin brown hairs giving palpi a bristly look; each antennal segment

of male ventrally expanded into a laterally projecting plate of variable development,
each plate with distinct hairs; early spring fliers serraticornis

Cucullia serraticornis Lintner (Figs. 17, 18) has not yet been collected in Mexico. It
occurs commonly in southern California and in southeastern Arizona. It undoubtedly
will be found in Sonora as well.

- Palpi not studded with thin brown hairs, not appearing bristly; antennal segments of

male simple 3

3. Lower third of postmedial line of forewing a conspicuous light line edged with black
on both sides; orbicular and reniform usually obviously present; forewing not gray,

but if gray usually with a distinct violet or blue tint 4

- Lower third of postmedial line of forewing not conspicuous although often vaguely

present; orbicular and reniform usually obscure or absent; forewing some shade of
gray with or without a bluish tint, never purplish 11

4. A dark lateral streak through the middle of the forewing along the cubital vein 5

- No dark lateral streak through the middle of the forewing along the cubital vein 6

5. Larger species (forewing length from base to apex approximately 25 mm)

costaricensis (9)

- Smaller species (forewing length from base to apex approximately 20 mm) oribac

Cucullia oribac Bames (Figs. 5, 6) has been seen from Veracruz, Morelos, Chiapas,
and the Federal District in Mexico. Its northern limit is southern and central Arizona

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CUCULLIA

269

Figs. 1-12. Cucullia species. 1. C. montanae Grote, 3, Portal, Cochise Co., Arizona, USA.
2. C. obscurior Smith, 3, Alta Lakes, San Miguel Co., Colorado, USA. 3. C. postera Guenee, 3,
Hill City, Pennington Co., South Dakota, USA. 4. C. lilacina Schaus, 3, Lomas de Chepultepec,
Distrito Federal, Mexico. 5. C. oribac Barnes, 3, Zacualpan, Mexico, Mexico. 6. C. oribac
Barnes, 9, Ciudad de Mexico, Distrito Federal, Mexico. 7. C. eucaena Dyar, 3, Mount Locke,
Jeff Davis Co., Texas, USA. 8. C. eucaena Dyar, 9, Lomas de Chepultepec, Distrito Federal,
Mexico. 9. C. costaricensis Hampson, 3, Mount Poas, Alajuela, Costa Rica. 10. C. costaricensis
Hampson, 9, Orizaba, Veracruz, Mexico. 11. C. new species #2, 3, Lomas de Chepultepec,
Distrito Federal, Mexico. 12. C. new species #1,3, Hondo, Medina Co., Texas, USA.

to western New Mexico. It probably occurs generally in Mexico, possibly extending
further south into Guatemala.

6. Forewing lilac or violet with a distinct fawn-brown patch between the reniform and
outer margin lilacina

The violet forewing color of Cucullia lilacina Schaus (Fig. 4) is very distinctive. The
species has been collected in Veracruz and the Federal District in Mexico and the
state of Cartago in Costa Rica. The northern limit of its range is in Arizona, New
Mexico, and western Texas.

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Figs. 13-18. Cucullia species. 13. C. laetifica Lintner, <5, Irving, Dallas Co., Texas, USA.
14. C. new species #3, 9, Panther Pass, Brewster Co., Texas, USA. 15. C. eccissica Dyar, <3,
Lomas de Chepultepec, Distrito Federal, Mexico. 16. C. antipoda Strecker, <3, Ephraim, Sanpete
Co., Utah, USA. 17. C. serraticornis Lintner, <3, Loma Linda, San Bernardino Co., California,
USA. 18. C. serraticornis Lintner, 9, Loma Linda, San Bernardino Co., California, USA.

Figs. 19-20. Opsigalea species. 19. O. ocel/ata Walker, <3, Lomas de Chepultepec, Distrito
Federal, Mexico. 20. O. blanchardi Todd, <3, Fort Davis, Jell' Davis Co., Texas, USA.

- Forewing not as above 7

7. Hindwing pure white or with a white base and dark brown outer margin 8

- Hindwing dull brown and if base lighter than outer margin, not conspicuously so

obscurior

Cucullia obscurior Smith (Fig. 2) has not yet been collected in Mexico, but it has
been collected in extreme southern Arizona and probably occurs in northern Sonora
as well.

8. Basal area of forewing with a large area of light brown; hindwing of male with broad

band of dark brown; median and middle subterminal area of forewing contrastingly
lighter than rest of wing montanae

Cucullia montanae Grote (Fig. 1) has not yet been collected in Mexico, but has been
seen from southern Arizona, New Mexico, and western Texas. It probably occurs in
northern Sonora and Chihuahua.

- Basal area of forewing without large patches of light brown; hindwing of male (but

not of female) without broad band of dark brown; median and middle subterminal
area of forewing not significantly lighter than remainder of forewing 9

9. Forewing with postmedial line distinct near costal margin; median area of male not

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CUCULLIA

271

uniform violet-gray; male without rusty patch between reniform and apex; female
with dark streak on median vein of forewing costaricensis (9 and <$)

Cucullia costaricensis Hampson (Figs. 9, 10) is the largest and most southerly species
of the genus in the New World. The only species costaricensis might be confused with
is eucaena. Its larger size and superficial differences are illustrated in the plate. In the
male genitalia the smaller spine of the vesica is very short, about as long as wide,
but in eucaena the spine is at least twice as long as wide. The clasper of the valve
barely exceeds the costa of the valve in costaricensis but greatly surpasses the costa
in eucaena. It has been collected in the state of Veracruz in Mexico, Baja Verapaz
and Quezaltenango in Guatemala, and San Jose, Cartago, and Alajuela in Costa Rica.

- Forewing with postmedial line indistinct near costal margin; median area of male

uniform purple-gray; male with rusty patch between reniform and apex; female without
dark streak on median vein of forewing 10

10. Forewing with small patch of white scales at base of forewing; male genitalia with

clasper medially placed in valve; ductus bursae of female genitalia without knob at
caudal end eucaena

Cucullia eucaena Dyar (Figs. 7, 8) is known from Puebla, Veracruz, Guerrero, and
the Federal District in Mexico. As in costaricensis and oribac the sexes are strongly
dimorphic. The male of eucaena is almost identical to postera from southern Arizona.
The white spot at the base of the forewing should identify it, but the male genitalia
may need to be checked to be sure. The female is suffused with dark brown, but the
female of postera is similar to the male.

- Forewing without a small patch of white scales at base of forewing; male genitalia

with clasper distally placed in valve; ductus bursae of female genitalia with knob at
caudal end postera

Cucullia postera Guenee (Fig. 3) is primarily a species of the northern parts of the
United States and most of Canada. It also occurs throughout the Rocky Mountain
region and has been taken near the Mexican border in southern New Mexico. It has
not yet been collected in Mexico, but probably will be. The male is almost identical
to the male of eucaena except for the small white patch mentioned in the key. For
certainty the male genitalia must be checked. Females resemble the males unlike the
female of eucaena.

1 1 . Forewing very light gray with central median area almost white; black dash on lower

fourth of postmedial line contrasting with light coloration of median area

new species # 1

Cucullia new species #1 (Fig. 12) has not yet been collected in Mexico, but occurs
throughout eastern and central Texas south to Brownsville and is commonest along
the coast. It probably occurs in Tamaulipas and possibly Nuevo Leon as well. The
light gray tinted with white coloration of the forewing will separate it from laetifica.

- Forewing not as above 12

1 1 . Forewing uniform dark black-gray; claspers of right and left valves of male genitalia

equal in size; vesica of aedoeagus with two spines new species #2

Cucullia new species #2 (Fig. 1 1 ) is the largest and darkest species of the speyeri
species group. It has been collected in the Federal District of Mexico. Elsewhere it is
known only from southeastern Arizona. Its largest size and dark coloration will
distinguish it from the other species in the speyeri species group.

Not as above

13

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

1 3. Forewing with a light patch in the median area; forewing with a slightly metallic blue

tint laetifica

Cucullia laetifica Lintner (Fig. 1 3) has not yet been collected in Mexico. The species
is common in southern Arizona, New Mexico, and Texas and undoubtedly occurs
throughout northern Mexico as well.

- Forewing without a light patch in the median area; forewing uniformly light gray

without a metallic blue tint 14

13. Vesica of male genitalia with two spines eccissica

Cucullia eccissica Dyar (Fig. 1 5) has been collected in the states of Puebla and Mexico
in Mexico. In the United States it occurs in southern California and southern Arizona.

- Vesica of male genitalia with three spines new species #3

Cucullia new species #3 (Fig. 14) has not been collected in Mexico as yet, but occurs
in western Texas and southern Arizona and probably occurs in Coahuila, Chihuahua
and Sonora as well. Its smaller size and the three spines in the vesica will distinguish
it from eccissica.

In addition to the species of Cucullia keyed above, there are two species of the
genus Opsigalea Hampson occurring in Mexico that may be mistaken for species of
Cucullia. These two species are Opsigalea ocellata (Walker) (Fig. 19) and Opsigalea
blanchardi Todd (Fig. 20).

LITERATURE CITED

Poole, R. W. 1989. Lepidopterorum Catalogus. Fascicle 118. Three Volumes. E. J. Brill,
Leiden, 1314 pp.

Poole, R. W. 1 992. Noctuidae. Fascicle 26. 1 . Cuculliinae, Stiriinae, and Psaphidinae (in part).
Moths of America North of Mexico. Wedge Entomological Foundation, Washington (in
press).

Received 12 August 1991; accepted 18 September 1991.

J. New York Entomol. Soc. 100(2):273-285, 1992

A REVISION OF THE GENUS HYPSOROPHA HUBNER
(LEPIDOPTERA: NOCTUIDAE: OPHIDERINAE)

Tim L. McCabe

Biological Survey, New York State Museum, The State Education Department,

Albany, New York 12230

Abstract. — The genus Hypsoropha (Lepidoptera: Noctuidae) is revised and two new species,
H. franclemonti, and H. baja are described. Male and female genitalia and adult habitus are
illustrated for all known species in the genus. Character states and resulting cladogram are
presented.

Hypsoropha Hiibner (Lepidoptera: Noctuidae: Ophiderinae) is a genus known from
Middle and North America. Two of the included species, H. monilis Frabicius (1 777)
and H. hormos Hiibner (1818) are common in the southeastern United States. Hyp-
soropha adeona Druce (1889) was described from Mexico and has recently been
recorded from Costa Rica by Daniel H. Janzen (in collection of J. G. Franclemont
and USNM). A fourth species is herein recorded from Cuba and the Bahamas and
a fifth from Mexico and Arizona. The only other species previously associated with
the genus, Tiauspa [ Hypsoropha ] argyria Butler (1879), was transferred from the
Noctuidae and made the type of a notodontid genus, Ctianopha, by Schaus (1901).

Both Hypsoropha monilis and H. hormos range over most of the southeastern
United States with H. hormos being the most widespread, having been reported as
far west as Arizona (Forbes, 1954), and as far north as New Hampshire and New
York (Crumb, 1956). Hypsoropha hormos larvae feed on Persimmon ( Diospyros
virginiana L.) [Ebenaceae] (Dyar, 1899; Glover, 1878; Guenee, 1852) and Sassafras
( Sassafras albidum (Nutt.)) Nees [Lauraceae] (Forbes, 1954). H. monilis larvae have
been reported from Diospyros (Crumb, 1956) and I have seen larvae (Fig. 29) from
New Jersey on D. virginiana.

Hypsoropha Hiibner, 1818

Type species: Noctua monilis Fabricius by subsequent designation by Grote, 1874.
Gloee Hiibner, 1808, suppressed (ICZN Op. 789).

Type species: Gloee monilis sensu Hiibner by subsequent designation by Berio,
1957.

Euphais Hiibner, 1818.

Type species: Noctua monilis Fabricius by subsequent designation by Berio, 1957.
Monogona G uenee, 1852.

Type species: Hypsoropha hormos, by monotypy.

Tiauspa Walker, 1858.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Type species: Hypsoropha hormos, by monotypy.

Gloce Neave, 1939, mispelling.

Description: Palpus upturned beyond vertex in males; male antenna pectinate to
subpectinate, female antenna filiform; eyes naked; reniform inconspicuous or absent;
orbicular a black point; forewing apex slightly falcate; female possessing appendix
bursa.

Hypsoropha' s monophyly is established by the following synapomorphies: forewing
with a transverse series of about four circular white markings in the fold along the
postmedian line; 8th abdominal sternum in the male with a peculiar pincer-like
development.

The nearest relatives appear to belong to the genus Psammathodoxa as suggested
by Franclemont (1985). Psammathodoxa and Hypsoropha share a chitinized devel-
opment of the eighth abdominal sternum, although this has become much more
elaborate in the latter.

Hypsoropha franclemonti, new species

Description. Antenna pectinate in male, simple in female; palpus upturned beyond
vertex in male, to middle of front in female; head, tegula and thorax violet-gray;
forewing with even, oblique postmedian line starting as four white markings in fold
and terminating at costa near apex; antemedian line, when traceable, bowed out at
middle; reniform obscure, orbicular a black point; wings silvery gray with diffuse
whitish scaling along posterior margin and at apex; hindwing buff from base to middle,
gradually darkening toward margin; fringe white. Male genitalia and eighth stemite
as in Figures 10 and 12. Female genitalia as in Figure 9. Expanse 25-34 mm.

Diagnosis. H. franclemonti can be distinguished from H. monilis and H. hormos
by the even, oblique postmedian line; from H. adeona and H. baja by the buff-gray
hindwing and less mottled forewing. The female bursa copulatrix has one band of
signa in H. franclemonti versus two bands in H. adeona. Other structural differences
in genitalia are evident from the illustrations (compare Figs. 9-12, 13-16, and 25-
28). Hypsoropha franclemonti is presently known only from Cuba and Great Exuma
Island in the Bahamas. Diospyros crassinervis (Krug & Urb.) Standi, has been reported
in Cuba as well as in the Bahamas (Correll and Correll, 1982) and may prove to be
an acceptable food plant for the new species.

Type material. BAHAMAS, Holotype male: Great Exuma Island, Simon’s Point,
lat 23.31.50 long. 75.47.30, 11 April 1986, T. L. McCabe [NYSM]; Paratypes, 17:
CUBA (2) Santiago (no additional data) [USNM, BMNH]; and the BAHAMAS (14)
(same data as Holotype except dates run from 10-16 April 1986) [NYSM, LACM,
JGF]; Bahamas: J. M. St. J. Yates (no additional data) (1) [BMNH]. Paratypes will
be deposited in the USNM, the BMNH, the LACM, and the collection of J. G.
Franclemont.

It gives me great pleasure to name this species after Prof. John G. Franclemont. I
have benefitted greatly from his frequent counsel. It seems especially fitting to com-
memorate him with a species in a group for which a connection had been made
through his knowledge and insight.

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REVISION OF HYPSOROPHA

275

Figs. 1-8. 1. Hypsoropha franclemonti, Holotype male, Bahamas, 15 April 1986. 2. H.

franclemonti, Paratype female, same data. 3. H. adeona, male, Guanacoste, Costa Rica. 4. H.
adeona, female, Veracruz, Mexico. 5. H. baja, Holotype male, Baja California, 10-11 Sept.
1984. 6. H. baja, Paratype female, same locale, 9 Sept. 1984. 7. H. monilis, male, Liberty Co.,
Florida, 25 March 1983. 8. H. hormos, male, Alachua Co., Florida, 3 April 1986. Scale line
10 mm.

Hypsoropha baja, new species
Figs. 5, 6, 25-28

Description. Antenna pectinate in male, simple in female; palpus upturned beyond
vertex in male, to middle of front in female; head, tegula and thorax violet-gray;
forewing with even, oblique postmedian line starting as four white markings in fold
and terminating at costa near apex; antemedian line, when traceable, bowed out at

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Figs. 9-12. Hypsoropha franclemonti. 9. Female genitalia (ventral view), McCabe slide 1499,
Bahamas, 10 April 1986. 10. Male genitalia, McCabe slide 1497, Bahamas. 1 1. Aedoeagus with
vesica uneverted, McCabe slide 1497, Bahamas. 12. Seventh and eighth abdominal sterna,
McCabe slide 1497, Bahamas. Scale line 1 mm.

middle; reniform obscure, orbicular a black point; wings brownish gray with diffuse
whitish scaling at apex, heavily mottled; hindwing buff from base to middle, slight
darkening toward margin; fringe white. Male genitalia and eighth sternum as in
Figures 26-28. Female genitalia as in Figure 25. Expanse 29-35 mm.

Diagnosis. H. baja can be distinguished from H. franclemonti by the more irrorated
forewings and male eighth stemites that lack teeth, and from H. adeona by the smaller
size and gray hindwings (creamy in H. adeona). H. baja has a variable am line (absent
to well represented whereupon it is dentate across from cell) and a variable amount
of freckling (almost smooth in one specimen). It is presently known only from the

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REVISION OF HYPSOROPHA

277

Figs. 13-16. Hypsoropha adeona. 13. Seventh and eighth abdominal sterna, McCabe slide
1515, Guanacaste, Costa Rica. 14. Female genitalia (ventral view), McCabe slide 1514, Ve-
racruz, Mexico. 15. Male genitalia, same data as Figure 13. 16. Aedoeagus with vesica everted,
same data as Figure 13. Scale line 1 mm.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Figs. 17-20. Hypsoropha monilis. 17. Seventh and eighth abdominal sterna, McCabe slide
1513, Alachua Co., Florida, 28 March 1986. 18. Male genitalia, same data as Figure 17. 19.
Aedoeagus with vesica everted, same data as Figure 17. 20. Female genitalia (ventral view),
McCabe slide 1405, Alachua Co., Florida, 30 March 1986. Scale line 1 mm.

Baja and Arizona. Diospyros californica (Brandegee) I. M. Jhtn. has been reported
in the Baja (Wiggins, 1980) and may prove to be an acceptable food plant for this
new species.

Type material. MEXICO, Holotype male: Baja Calif. Sur Cabo Pulmo, sea level,
10-1 1 September 1984, J. P. & K. E. Donahue [LACM]. Paratypes, 10: MEXICO:
same data as holotype [5 specimens— LACM, NYSM]; Mexico: Baja Ca Sur, 12 mi
S P. Colorada, Arroyo Las Barracae, 15-18 Oct 1989, N. Bloomfield [1-LACM];
Mexico: Colima, 13 mi N Manzanillo, nicr. Toro 24-26 May 1989 N. Bloomfield
[1-LACM] Aguaje, Sinaloa, Mex., VI-18 1956, R. P. Allen [1-CAS]; Baja Calif., San

1992

REVISION OF HYPSOROPHA

279

Figs. 21-24. Hypsoropha hormos. 21. Seventh and eighth abdominal stemites, McCabe
slide 1491, Alachua Co., Florida, 3 April 1986. 22. Aedoeagus with vesica everted, same data
as Figure 21. 23. Male genitalia, same data as Figure 21. 24. Female genitalia (dorsal view),
McCabe slide 1404, Alachua Co., Florida, 27 March 1986. Scale line 1 mm.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Figs. 25-28. Hypsoropha baja. 25. Female genitalia (ventral view), McCabe slide 1769,
Baja, California, 24 July 1981. 26. Male genitalia, McCabe slide 1787, Baja, California, 24 July
1981. 27. Aedoeagus, same data as Figure 26. 28. Seventh and eighth abdominal stemites,
same data as Figure 26.

Bartola Microwave Tower, 2000 ft., July 24, 1981, R. Holland [1-AMNH]. UNITED
STATES: Arizona: August 22, 1954, Madera Canyon, Santa Rita Mts., Southern
Arizona, Lloyd M. Martin [1-LACM].

Hypsoropha adeona Druce
Figs. 3-4, 13-16

Hypsoropha adeona Druce, 1889:338, pi. 30, fig. 28.
Hypsoropha adeona', Poole, 1989:546.

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REVISION OF HYPSOROPHA

281

Table 1. Character states.

1. Male eighth sternum: narrow (0), flattened (1); expanded (2).

2. Male eighth sternum with supporting struts: absent (0); present (1).

3. Male antennae: heavily pectinate (0); subpectinate (1).

4. Valves with fold: central (0); marginal (1).

5. Male eighth sternum with teeth: absent (0); present (1).

6. Pm line: straight (0); indented (1).

7. Forewing white markings: absent (0); present (1).

8. Saccus a “V” (0); a “U” (1).

9. Female ductus bullae broad and chitinized (1) or not (0).

10. Female bursal bands/signa: two bands (0); one band (1); one signum (2).

1 1. Female appendix bursae: absent (0); weakly developed (1); well developed (2).

12. Female with mid- ventral chitinized pocket between ovipositor lobes: absent (0); present

(1).

13. Eight stemite directed: posteriorly (0); anteriorly (1).

14. Aedoeagus a narrow, curved tube (0) or broad and straight (1).

15. Uncus: simple (0); slightly modified (1); keel-shaped (2).

16. Cucullus: absent (0); present (1).

17. Forewing with circular markings: absent (0); present (1).

18. Postmedian line: broken (0); entire (1).

Description. Antenna subpectinate in male, simple in female; palpus upturned
beyond vertex in both sexes; head, tegula and thorax buff with yellowish tint; forewing
with even, oblique postmedian line starting as four white markings in fold and
terminating at costa near apex; antemedian line, when traceable, bowed out opposite
cell; reniform obscure, orbicular a black blotch; wings yellow-brown and heavily
mottled; hindwing with basal two thirds yellow, gradually darkening toward margin;
fringe brown. Male genitalia and eighth sternum as in Figures 15 and 13. Female
genitalia as in Figure 14. Expanse 35-40 mm.

Diagnosis. Both H. adeona and H. franclemonti possess the even, oblique post-
median lines, but H. franclemonti has a buff-gray hindwing (see diagnosis under H.
franclemonti). The moth occurs from Mexico to Costa Rica and the food plant is
unknown.

Type locality. Mexico [Veracruz], Jalapa [BMNH].

Hypsoropha monilis (Fabricius)

Figs. 7, 17-20

Noctua monilis Fabricius, 1777:283.

Hypsoropha monilis ; Hiibner, 1816:249; Hypsoropha monilis ; Walker, 1857:994;
Hypsoropha monilis ; Dyar, 1903:219; Hypsoropha monilis ; McDunnough, 1938:
125; Hypsoropha monilis ; Franclemont and Todd, 1983:123; Hypsoropha monilis ;
Poole, 1989:546.

Description. Antenna pectinate in male, simple in female; palpus upturned beyond
vertex in both sexes; head, tegula and thorax orange-brown; forewing with postmedian

282 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Table 2.

Character matrix.

Characters

Taxa

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18

H. adeona

2

1

1

1

1

0

1

0

0

0

2

1

1

0

1

1

1

0

H. baja

2

1

1

1

0

0

1

0

1

0

2

1

1

0

0

1

1

0

H. franclemonti

2

1

0

0

1

0

1

1

0

1

2

0

1

1

0

0

1

0

H. hormos

1

0

0

0

0

1

1

1

0

0

2

1

1

1

0

0

1

1

H. monilis

1

0

0

1

0

1

1

1

1

2

1

0

1

0

2

0

1

1

P. natadoides

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

line indented below reniform, starting as four white markings in fold and terminating
at costa near apex; antemedian line rather evenly excurved when traceable; reniform
obscure, orbicular a black point; wings orange-brown; hindwing orange-brown, slight-
ly darker toward margin; fringe concolorous with ground. Male genitalia and eighth
sternum as in Figures 18 and 17. Female genitalia as in Figure 20. Expanse 33-44
mm.

Diagnosis. H. monilis is distinguished from H. adeona and H. franclemonti by the
indented postmedian line. A spade-shaped juxta represents an autapomorphy for H.
monilis. The species occurs from northern Florida north to North Carolina, Illinois,
and Missouri (Forbes, 1954). The larva feeds on Diospyros virginiana and has been
described by Dyar (1903a). The head has a conspicuous, rounded, black spot which
extends from the posterior margin to along the adfrontals for half their length. The
dorsum is black with a pale middorsal stripe, then a subdorsal pale stripe, this followed
by a black stripe (Fig. 29). H. monilis larvae are sawfly mimics, but spend the daytime
hidden at the base of the tree, and large numbers of larvae climb at dusk onto usually
one or two limbs (D. F. Schweitzer, pers. comm.). The larva diapaused as a prepupa
in a weak cocoon.

Type locality. “Anglia” [in error for North America]. The type has not been located,
but it was illustrated by Westwood (1854) on plate 54, figure 61b (plate labelled
“doubtful British species”).

Hypsoropha hormos Hiibner
Figs. 8, 21-24

Hypsoropha hormos Hiibner, 1818:10, figs. 27, 28.

Monogona hormos: Guenee, in Boisduval and Guenee, 1852:402.

Tiauspa hormos ; Walker, 1857:995.

Hypsoropha hormos; Dyar, 1903b:219; Hypsoropha hormos; McDunnough, 1938:

1 25; Hypsoropha hormos ; Franclemont and Todd, 1983:1 23; Hypsoropha hormos;

Poole, 1989:546.

Description. Antenna pectinate in male, simple in female; palpus upturned beyond
vertex in both sexes; head, tegula and thorax pearl-gray; forewing with postmedian
line indented below reniform, starting as four white markings in fold and terminating
at costa near apex; antemedian line bowed out at middle; reniform obscure, orbicular
a black point; wings pearl gray with black scaling along postmedian line; median area

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REVISION OF HYPSOROPHA

283

Fig. 29. Last instar larva of Hypsoropha hormos, Bear Swamp East, Cumberland Co., New
Jersey (shown on leaf of Diospyros virginiana).

and apex lighter gray; hindwing uniform gray-buff gradually darkening toward margin;
fringe concolorous with ground. Male genitalia and eighth sternum as in Figures 23
and 21. Female genitalia as in Figure 24. Expanse 22-32 mm.

Diagnosis. H. hormos is readily distinguished from H. adeona and H. franclemont i
by the indented postmedian line; from H. monilis by the gray color and black scaling
along the postmedian line and much smaller size. A pointed male eighth sternum
represents an autapomorphy for H. hormos. The species occurs from the Florida
Keys west to Arizona (Forbes, 1954) and north as far as New Hampshire and New
York (Crumb, 1956). The larva was described and the food plant, Diospyros, was
recorded by Crumb (1956). The larva has a reticulate head pattern. A green and a
brown color phase occur; the green phase has purplish spots subdorsally, these being
fused on segment five and eight. The mandible has a mesal tooth. Adults show much
more brown overall in the northern parts of their range.

Type locality. Georgia [type deposition unknown].

PHYLOGENETIC ANALYSIS

Polarization of character states was accomplished by outgroup comparison with
Psammothodoxa natadoides Franclemont, which has been recognized as a near rel-
ative of Hypsoropha (Franclemont, 1985). The Hennig 86 computer program (Farris,
1988) was employed. The cladogram in Figure 30 is the most parsimonious tree
possible for the data presented. The tree exhibited a consistency of 0.786.

Not immediately evident from the bi- and tri-state partitioning of characters is a

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Psammothodoxa

■ H. hormos
- H. monilis

I H. franclemonti

I H. adeona

1 H. baja

Fig. 30. Cladogram.

phyletic series: the eighth abdominal stemite in the male progresses from a simple,
thin, modified pincer in H. hormos, to a slightly heavier pincer in H. monilis, to an
even heavier and expanded pincer in H. franclemonti and H. baja and finally to a
very heavy, expanded structure in H. adeona. Hypsoropha hormos, which has the
largest known distribution, is hypothesized to be the least derived species in the
genus.

ACKNOWLEDGMENTS

I thank John Franclemont of Cornell University, Robert Poole of the United State National
Museum, James Miller of the American Museum of Natural History, Julian Donahue of the
Los Angeles County Museum, Paul Amaud and Norman Penny of the California Academy of
Science, S. Morton Adams of West Shokan, John Rawlins of the Carnegie Museum, and Martin
Honey of the British Museum (Natural History) for searching for Caribbean, Middle and North
American specimens for me. I am also indebted to Ole Karsholt of the Zoologisk Museum,
Glasgow, who kindly searched for Hypsoropha and confirmed that the type of H. monilis was
not deposited in that museum and who sent related genera collected in Argentina. Mr. Sidney
Russell, on behalf of the Bahamian government, granted permission to collect on Exuma Island.
Christopher Supkis printed the photographs. Patricia Reman provided the illustrations. Dale
Schweitzer kindly sent me immatures of H. monilis and gave his observations on their habits.
Contribution number 656 of the New York State Science Service.

LITERATURE CITED

Berio, E. 1957. Ulteriori modifiche e Cambiamenti nella nomenclatura dei generi di noctuidae
del globo. Memorie Soc. Entomol. Ital. 36:14.

Butler, A. G. 1879. On the Lepidoptera of the Amazons, collected by Dr. James W. H. Trail,
during the years 1873 to 1875. Part III.— Noctuites. Trans. Entomol. Soc. Lond. 1879:
1 60-166.

Correll, D. S. and H. B. Correll. 1982. Flora of the Bahama Archipelago. J. Cramer, Vaduz,
1692 pp.

1992

REVISION OF HYPSOROPHA

285

Crumb, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agric. Tech. Bull. 1135,
356 pp.

Druce, H. 1889. Heterocera. In Godman and Salvin, Biologia Centrali Americana. Insecta.
Taylor and Francis, London. P. 338, PI. 30, Fig. 28.

Dyar, H. G. 1899. Life history of Hypsoropha hormos, Hiibn. Can. Entomol. 31:289-290.

Dyar, H. G. 1903a. New North American Lepidoptera with notes on larvae. Proc. Wash.
Entomol. Soc. 5:291.

Dyar, H. G. 1903b. A list of North American Lepidoptera and key to the literature of this
order of insects. Government Printing Office, Washington, D.C., 723 pp.

Fabricius, J. C. [1777]. Genera Insectorum. M. F. Bartschii, Chilonii [Kiel], p. 283.

Farris, J. S. 1988. Hennig 86 (a phylogenetic computer program), version 1.5. Stony Brook,
N.Y.

Forbes, W. T. M. 1954. Lepidoptera of New York and neighboring states. Cornell Expt. Sta.
Mem. 329, 433 pp.

Franclemont, J. G. 1985. A new species of Psammathodoxa Dyar from Costa Rica (Lepi-
doptera: Noctuidae: Catocalinae). Proc. Entomol. Soc. Wash. 87(4):746-750.

Franclemont, J. G. and E. L. Todd. 1983. Noctuidae, pp. 120-159. In: R. W. Hodges et al.
(eds.), 1983 Check List of the Lepidoptera of America North of Mexico. E. W. Classey
Limited and the Wedge Entomological Research Foundation, London, 284 pp.

Glover, T. 1878. Illustrations of North American Entomology. J. S. Tomlinson, Washington,
D.C., plate 13, figs. 1 & 2.

Grote, 1874. List of the Noctuidae of North America. Bull. Buffalo Soc. Nat. Sci. 2:29.

Guenee, A. 1852. In J. B. A. D. de Boisduval & A. Guenee, Histoire Naturelle des Insectes.
Species General des Lepidopteres. Roret, Paris, Pt. 6, p.402.

Hiibner, J. 1808. Sammlung exotischer Schmetterlinge. Augsburg 1:14, figs. 27, 28.

Hiibner, J. 1816. Verzeichniss bekannter Schmetterlinge. Augsburg P. 249.

Hiibner, J. 1818. Zutrage zur Sammlung exotischer Schmetterlinge. Augsburg 1:10.

McDunnough, J. 1938. Checklist of the Lepidoptera of Canada and the United States of
America. Part 1, Macrolepidoptera. Mem. S. Calif. Acad. Sci. 1:1-272.

Neave, S. A. 1939. Nomenclator Zoologicus. 2:477. Richard Clay & Co., Ltd., Suffolk, En-
gland.

Poole, R. W. 1989. Lepidopterorum Catalogus (N.S.). Fascicle 118. Noctuidae (3 parts). E.
J. Brill/Flora & Fauna Publications, New York, 1314 pp.

Schaus, W. 1901. A revision of the American Notodontidae. Trans. Entomol. Soc. London
1901:272.

Walker, F. [1858] 1857. List of the Specimens of Lepidopterous Insects in the Collection of
the British Museum. E. Newman, London. Vol. 13, p. 984, 995.

Westwood, J. O. 1854. Wood’s Index Entomologicus; or, a complete illustrated catalogue,
consisting of upwards of two thousand accurately coloured figures of the Lepidopterous
insects of Great Britain. A new and revised edition, with supplement, containing figures
and notices of nearly two hundred newly discovered species, synoptic lists, etc. London.
G. Willis, Great Piazza, Covent Garden. Pp. 298.

Wiggins, I. 1980. Flora of Baja California. Stanford University Press, Stanford, California,
1025 pp.

Received 1 February 1991; accepted 21 October 1991.

J. New York Entomol. Soc. 100(2):286-310, 1992

LIFE HISTORY AND SYSTEMATICS OF THE
WEST ANDEAN MOTH AUCULA FRANCLEMONTI WITH
DESCRIPTION OF A NEW SPECIES FROM ECUADOR
(LEPIDOPTERA: NOCTUIDAE: AGARISTINAE)

John E. Rawlins

Section of Invertebrate Zoology, Carnegie Museum of Natural History,

4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213

Abstract. — Systematics, biology, and morphology of the rare South American agaristine moth,
Aucula franclemonti Todd and Poole (Lepidoptera: Noctuidae), are presented in the context of
other agaristine genera. A. kimsa Todd and Poole is placed in synonymy with A. franclemonti,
and A. ivia Todd and Pool is synonymized with A. tusora Todd and Poole; A. franclemontoides,
new species, is described from east Andean Ecuador. A brief phylogenetic analysis of the five
species in the A. franclemonti species-group is offered, with a key to the species-group based
on male genitalia. Female genitalia of A. franclemonti are described, and an external mating
plug (sphragis) noted, perhaps functionally related to a permanently everted and sclerotized
endophallus in males. Larvae were found feeding on Vitis tiliifolia Humb. & Bonpl. (Vitaceae),
and all immature stages reared from the egg are briefly described. A complete chaetotaxy of
the last instar larva is illustrated in detail. Modifications of the pupal cremaster and the presence
of subventral seta SV2 on the seventh abdominal segment are apomorphic features shared only
with other agaristine species.

Species of Aucula are infrequently encountered agaristine moths found in semiarid
to mesic habitats in Panama and tropical South America. Kiriakoff (1977) sum-
marized the taxonomic status of species described by earlier workers. In 1981, Todd
and Poole revised Aucula, thoroughly describing adult males and providing a key to
all species based on the male genitalia. They recognized as valid three previously
described species, transferred two species to related monobasic genera ( Arpia Schaus
and Darcetina Felder), and excluded three species from Aucula, leaving them tem-
porarily in generic combinations as originally described. They also described 2 1 new
species based primarily on genitalic differences after examining slightly more than
100 male specimens in the following collections: National Museum of Natural His-
tory, Washington, D.C. (NMNH); The Natural History Museum, London (BMNH);
Universidad Central de Venezuela; Zoologische Sammlung des Bayerischen Staates,
Munich; and Carnegie Museum of Natural History, Pittsburgh (CMNH). Of the 24
species currently considered valid (Poole, 1989), nine are known from unique types,
and only seven species are known from more than five specimens. The longest series
for any species is 18 specimens (Aucula ivia Todd and Poole), 13 of which are from
the same locality. Todd and Poole mentioned only 12 female specimens associated
with six species but did not figure adult females or their genitalia.

A series of 23 males and two females of A. franclemonti Todd and Poole were
collected at a single locality in western Ecuador during the summer of 1983. A.
franclemonti was previously known only from the holotype. This material constitutes
the largest known series of any Aucula species, providing information on character

1992

AUCULA FRANCLEMONTI

287

Figs. 1-6. Adults of Aucula, dorsal (left) and ventral views (right). 1-2. A. franclemonti,
male; forewing length 25 mm; Ecuador, Carchi (CMNH). 3-4. A. franclemonti, female; forewing
length 27 mm; Ecuador, Carchi (CMNH). 5-6. A. franclemontoides, holotype male; forewing
length 24 mm; Ecuador, Morona-Santiago (CMNH).

variation not available to Todd and Poole. Eggs were also obtained and the immature
stages reared; a larva was found feeding in the field and an adult female was reared
from it, confirming the identity of the foodplant. The following paper reviews the
systematics of A. franclemonti and closely related species, describes as new a related
species from eastern Ecuador, describes the female genitalia, provides basic life
history information, and presents a detailed morphological description of the larva
in the context of other agaristine and noctuid larvae.

SYSTEMATICS OF THE AUCULA FRANCLEMONTI GROUP

Todd and Poole (1981) informally recognized a group of six superficially identical
species of Aucula from the Andes “characterized by an enlarged broad uncus and a

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Figs. 7-16. Outlines of unci from male genitalia of species in the Aucula franclemonti group
(Figs. 7-10 and 12-13 taken from Todd and Poole (1981)). 7. A. jenia, holotype. 8. A. otasa,
holotype. 9. A. tusora, holotype. 10. A. ivia, holotype. 1 1. A. franclemontoides, holotype (Gen-
italia slide: Rawlins 860 (CMNH)). 12. A. kimsa, holotype. 13. A. franclemonti, holotype. 14.
A. franclemonti (Genitalia slide: Rawlins 849 (CMNH)). 15. A. franclemonti (Genitalia slide:
Rawlins 858 (CMNH)). 16. A. franclemonti (Genitalia slide: Rawlins 857 (CMNH)).

characteristically shaped valve,” the latter referring to the abruptly angled ventral
margin at the distal end of the sacculus and a slender, setose, and smoothly arched
valve apex. These genitalic features are unique to this group, both in the context of
other species in the genus, as well as other species of Agaristinae (Kiriakoff, 1977).
Their apomorphic condition supports the monophyly of a taxon here christened the
A. franclemonti group. All specimens in this group examined by Todd and Poole
were from Colombia and Ecuador west of the Andes, or from southern Peru and
adjacent Bolivia east of the Andes.

Todd and Poole (1981) described the largest individuals in the group (FWL = 26-
27 mm) as two species, A. franclemonti based on a single male (NMNH), and A.
kimsa based on two males (BMNH). All type specimens of both species are from
the same locality in western Colombia, San Antonio near Cali. The two species were
distinguished by the shape of the distal half of the uncus, smoothly lobed in A.
franclemonti (Fig. 13) and contrastingly concave in A. kimsa (Fig. 12). Similarly,
Todd and Poole (1981) described two medium-sized species (FWL = 19-22) from
northwestern Bolivia and adjacent southeastern Peru, A. ivia based on 18 males from
both countries (NMNH, BMNH, CMNH), and A. tusora based on a single male from
Bolivia. A. tusora was distinguished from A. ivia by the smoothly tapered base of
the uncus, contrasting with a distinctive, angulate, or “shouldered” condition in A.
ivia. Other than this condition, the habitus and male genitalia of the two species are
virtually identical despite the somewhat worn condition of the unique type specimen
of A. tusora.

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AUCULA FRANCLEMONTI

289

Figs. 17-20. Male genitalia of Aucula. 17. External genitalia of Aucula franclemonti (Gen-
italia slide: Rawlins 848 (CMNH)). 18. Aedeagus of Aucula franclemonti (Genitalia slide:
Rawlins 849 (CMNH)). 19. External genitalia of male holotype of Aucula franclemontoides
(Genitalia slide: Rawlins 860 (CMNH)). 20. Aedeagus of male holotype of Aucula franclemon-
toides.

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species-group.

The A. franclemonti species-group contains two other distinctive and easily rec-
ognized species. The smallest in the group (length of forewing, FWL = 17 mm), A.
otasa Todd and Poole, was described from a single Bolivian male (BMNH) and
characterized by an autapomorphic incision of the dorsal margin of the sacculus, a
very sharply angled emargination of the ventral outline of the valve apex distal to
the saccular angulation, and a spatulate uncus (Fig. 8). Another medium-sized species
from western Ecuador, A. jenia Todd and Poole (FWL = 23-24 mm), is easily
recognized by an unique black bar across the yellow basal region on the ventral side
of the hindwing (Fig. 25), and by male genitalia with a nearly straight outer margin
beyond the saccular angulation and a broadened, strap-like uncus without lateral
lobes (Fig. 7). The black wing bar is autapomorphic for A. jenia, but the genitalic
features are considered plesiomorphic as for most species of Aucula other than those
in the A. franclemonti group. One additional specimen of A. jenia has been examined
and found to agree with all diagnostic features of the type series (Ecuador, Chim-
borazo, 1 1 km NE Pallatanga, 2,800 m, 1 1 November 1987, C. Young, R. Davidson,
J. Rawlins (CMNH)).

Capture of 23 males in the A. franclemonti group at a single locality in northwestern
Ecuador near the Colombian border (Ecuador, Carchi, Chical, 00°56'N, 78°11'W,
1,250 m, J. Rawlins, R. Davidson (CMNH)) provide new data on the nature and
extent of variation in adult characters, including those purportedly diagnostic for A.
franclemonti and A. kimsa. Specimens were taken during a period of intensive col-
lecting extending from June 21 to August 16, 1983, and include the following dates
(number of male specimens in parentheses): June 29 (1), July 1 (2), July 2 (6), July
5(1), July 6(1), July 8(1), July 1 1 (3), July 15(1), July 17 (4), July 18 (1), August
1 (1), August 7 (1). All genitalic features were assessed in six males by dissection,
and the shape of the uncus and the distal half of the valve was examined in all
remaining males by brushing abdominal apices.

All males were virtually identical in color and wing pattern (Figs. 1-2). There is
slight variation in forewing size: FWL, 23-27 mm; mean = 24.8 mm; mode = 25
mm; N = 23. Several genitalic features are highly variable, especially the degree of
setation and sclerotization of the valves and the shape and degree of sclerotization

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AUCULA FRANCLEMONTI

291

of the uncus. In particular, the shape of the distal half of the uncus varies from a
condition more deeply concave than that of the type specimen of A. kimsa to indi-
viduals with lateral lobes larger than those on the type specimen of A. franclemonti
(Figs. 12-17). Collection of these specimens at a single locality over a short time
period suggests they are conspecific members of a single population exhibiting con-
siderable intrapopulation variation in genitalic characters, especially the shape of the
uncus. Type specimens of A. kimsa and A. franclemonti fall within the range of
character variation observed in this series, and their recognition as separate species
is not supported. In contrast, some features of the male genitalia do not vary, notably
the shape and sclerotization of the juxta, and the shape and size of the aedeagus (see
discussion below).

The degree and extent of sclerotization at the base of the uncus in this conspecific
series of A. franclemonti also varies greatly, ranging from a weakly sclerotized con-
dition with smoothly arched outline to a more strongly sclerotized condition resulting
in angled “shoulders” at the uncal base. The latter condition is very similar to the
only diagnostic feature separating the single male type of A. tusora from the relatively
extensive series of A. ivia, suggesting, albeit indirectly, that those two names represent
but a single species as well.

In addition to the above specimens from western Ecuador, a single, slightly dam-
aged specimen of Aucula was captured in Ecuador in a wet forest on the eastern slope
of the Andes. Because this specimen was the first Aucula specimen known to me
from the eastern Andean region north of southern Peru, special attention was given
to its study. Externally the specimen closely resembles A. franclemonti and A. tusora
(Fig. 3), but genitalic dissection revealed a distinctive new species of Aucula of the
A. franclemonti group (Figs. 19-20).

Aucula franclemontoides, new species
Figs. 3-4, 19-20

DIAGNOSIS. This tropical American species is placed in the genus Aucula by the
following combination of characters: rounded (not falcate) forewings, characteristic
wing pattern (Fig. 3-4), lack of a yellow basal patch on the ventral side of the forewing,
and male genitalia lacking a distinct median process on the sacculus. It differs from
all described species of Aucula by features of the male genitalia, especially the strongly
sclerotized, enlarged lobes on the dorsal margin of the juxta and the greatly enlarged
and elongated lateral lobes of the uncus (Fig. 1 9).

DESCRIPTION (male only, female unknown). Forewing: length, 24 mm; maxi-
mum width, 12 mm. Hindwing: length, 18 mm. Extremely similar in all external
features to A. franclemonti Todd and Poole, including antennal pectination, shape
and size of frontal process, color pattern of wings and body; differs from A. francle-
monti as follows: ground color of forewing brownish black, slightly darker and less
reddish brown than A. franclemonti and similar in color to A. jenia\ outer margin of
yellow basal patch on hindwing smoothly and evenly curved as in A. tusora and A.
otasa, not crenulate due to faint inward extension of black scales along veins as in
A. franclemonti and A. jenia.

Male abdomen. Abdominal terga 3-7 with antecostae evanescent on midline; Ter-

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gum 8 subquadrate, unmodified; Sternum 8 with anterior margin slightly thickened
and sinuate. Abdominal Segment 2 with lateral hair pencils resting in scale-lined,
elongate pleural invaginations on segments 2-4.

Male genitalia. Figures 19-20. Tegumen slightly wider than long, strongly scler-
otized dorsally, membranous ventrally. Vinculum narrow; saccus not developed.
Juxta a smooth flattened plate, subacute ventrally, prolonged dorsally into two round-
ed and posteriorly extended lobes separated by a deep notch; lobes of juxta heavily
sclerotized, densely covered with microscopic spinules, Anellifer membranous except
in region immediately adjacent to aedeagus. Uncus inflated at base to more than half
width of tegumen, narrowly constricted at middle, then abruptly dilated into two
long lateral lobes and terminated in a very short, broadly truncated, and slightly
asymmetrical apex. Valves symmetrical, subtrapezoidal, with elongate, curved apices;
costa with large triangular process at middle subtended by flat, coarsely setose editum;
sacculus broad, setose, with ventral margin smoothly curved, forming abrupt obtuse
angle with apex of valve; dorsal margin of sacculus distinct, raised into rounded
prominence near base of valve; apex of valve constricted distally with outline strongly
concave ventrally and dorsally; valve with distal end slightly flattened and widened,
dorsally deflected, and densely setose on mesal surface.

Aedeagus greatly modified (Fig. 20); basiphallus broadly expanded anteriorly into
a weakly sclerotized coecum; distiphallus shorter than its diameter, but bearing
dorsally a strongly sclerotized, flattened, apical process which curves smoothly ven-
trad; endophallus (vesica) permanently everted, very heavily sclerotized, forming a
hollow tube curved dorsad, terminated in a subcircular meatus (primary gonopore)
with a minute spinule approximate to its ventral lip.

BIOLOGY AND IMMATURE STAGES. Unknown, but are expected to be very
similar to those of A. franclemonti, described below. Expected foodplants for A.
franclemont aides and other species of Aucula are species of Vitis and related genera
of Vitaceae.

DISTRIBUTION. Known only from the type locality where the habitat is sub-
montane rainforest of exceptional richness and floristic diversity.

DISCUSSION. Diagnostic modifications of the juxta and uncus leave little doubt
that this species is distinct from other members of the A. franclemonti group.

Of special interest is the aedeagus, which is similar to those of other species in the
A. franclemonti group, none of which was figured by Todd and Poole (1981). They
interpreted the aedeagus to have a “vesica without spines or comuti,” but careful
study and dismemberment of one aedeagus did not reveal any eversible structure
beyond the external meatus of the strongly sclerotized, tubelike apex of the phallus.
Comparative study of the aedeagus reveals that the basiphallus (portion of aedeagus
internal to junction with anellus (manica)) is inflated and weakly sclerotized. The
distiphallus (sclerotized external portion of aedeagus from manica to base of endo-
phallus) is very short with an elongate dorsal process. The endophallus is not mem-
branous, but rather very strongly sclerotized into a permanently everted tube so that
the apparent apical opening of the phallus is the primary gonopore, not the secondary
gonopore (phallotreme) as is the usual case in most Lepidoptera.

PRIMARY TYPE DATA. Holotype male, CMNH. Verbatim text of four pin
labels: ECUADOR: Morona-Santiago. Rio Culebrillas, 34 km SE Gualaceo. 2,200
m. 22-23 Oct 1987 / J. Rawlins, C. Young, R. Davidson. Wet forest. / Genitalia

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Slide No. 860 John E. Rawlins / HOLOTYPE Aucula franclemontoides Rawlins [red
paper].

Etymology: The name franclemontoides is an adjective referring to the great re-
semblance of this species to A. franclemonti.

ALPHABETICAL CHECKLIST OF SPECIES IN THE AUCULA FRANCLEMONTI GROUP

Aucula franclemonti Todd and Poole, 1981.

Aucula kimsa Todd and Poole, 1981. NEW SYNONYMY.

Aucula franclemontoides Rawlins, new species.

Aucula jenia Todd and Poole, 1981.

Aucula otasa Todd and Poole, 1981.

Aucula tusora Todd and Poole, 1981.

Aucula ivia Todd and Poole, 1981. NEW SYNONYMY.

KEY TO ADULTS OF SPECIES IN THE AUCULA FRANCLEMONTI GROUP
BASED PRIMARILY ON THE MALE GENITALIA

(Derived in part from key in Todd and Poole, 1981)

1. Uncus widest near middle (Figs. 8-17, 19); yellow region on lower surface of HW

unmarked (Figs. 2, 4, 6) 2

- Uncus widest at apex, strap-like (Fig. 7); yellow region on lower surface of HW with

black stripe across middle (Fig. 25) jenia

2. Dorsal margin of sacculus with triangular lobe near middle; apical part of valve nearly

straight otasa

- Dorsal margin of sacculus not lobed; apical part of valve distinctly curved 3

3. Sclerotized dorsal edge ofjuxta v-shaped, the convergent sides nearly straight in outline

(Fig. 1 7); uncus with lateral outline slightly concave, straight, or evenly convex (Figs.
12-17) franclemonti

- Sclerotized dorsal edge ofjuxta with two rounded lobes protruded posteriorly, in outline

forming two rounded lips separated by a distinct notch (Fig. 19); uncus with lateral
lobes 4

4. Uncus with distal lateral outline evenly convex (similar to Figs. 9-10) tusora

- Uncus with greatly enlarged lateral lobes (Figs. 11, 19) franclemontoides, n. sp.

PHYLOGENETIC RELATIONSHIPS

There are few characters shared among the five species of the A. franclemonti group
that permit phylogenetic analysis relative to an outgroup consisting of all other species
of Aucula. Five informative characters were observed as follows, the state occurring
in the outgroup scored 0:

Character 1. Lateral lobes of uncus: 0, present; 1, absent.

Character 2. Lateroventral outline of apical half of valve: 0, straight; 1, concave,
smoothly curved; 2, concave, deeply angled.

Character 3. Distinct, rounded lobes on dorsal edge ofjuxta: 0, absent; 1, present.
Character 4. Ventral outline of valve at distal end of sacculus: 0, smoothly curved;
1 , sharply angled.

Character 5. Length of free, apical extension of valve: 0, apex of valve not con-
spicuously elongated; 1 , apex of valve elongated and constricted.

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Characters were distributed as follows, with the number of each character followed
by its state in parentheses: A. jenia, 7(0), 2(0), 2(0), 4{ 1), 5(1); A. otasa, 7(1), 2(2),
2(1), "7(1), 5(1); A. franclemonti, 7(1), 2(1), 2(0), 4{ 1), 5(1); A. franclemontoides, 7(1),
2(2), 2(1), "7(1), 5(1); A. tusora, 7(1), 2(1), 2(1), "7(1), 5(1); other Aucula species
(outgroup), 7(0), 2(0), 2(0), "7(0), 5(0). Characters autapomorphic for single species
were not considered, such as small body size and a notch in the dorsal margin of
sacculus in A. otasa, the black HW stripe in A. jenia, huge uncal lobes in A. francle-
montoides, and a relatively short apical elongation of the valve in A. tusora.

A phylogenetic analysis produced a minimally supported but fully resolved hy-
pothesis of relationship. All possible phylogenetic trees (105) were examined with
respect to the above five characters, and a single tree of minimum length (Fig. 21)
was found without invoking hypotheses of homoplasy (Consistency = 1 .00). East
Andean species from Ecuador, Peru, and Bolivia (A. tusora, A. otasa, and A. fran-
clemontoides) are hypothesized to form a monophyletic subgroup based on distinctive
lobing of the juxta. This subgroup and A. franclemonti form a monophyletic group
supported by two synapomorphies, uncal lobes and an inwardly curved or angled
ventral margin along the outer half of the valve.

The monophyly of the A. franclemonti group is supported by two derived char-
acters, apical elongation of the valves and a sharply angled ventral margin at the
distal end of the sacculus. On the other hand, there is no evidence that other species
of Aucula form a monophyletic group, and genitalic features suggest that the im-
mediate sister-lineage to the A. franclemonti group may be one including Aucula byla
Todd and Poole. There is also serious question whether the genus Aucula is mono-
phyletic as treated by Todd and Poole (1981), because diagnostic features currently
used to define the genus, such as wing shape, coloration, and the absence of a median
process of the sacculus, are not convincingly apomorphic at the level of Agaristinae.

DESCRIPTION OF THE ADULT FEMALE OF AUCULA FRANCLEMONTI

Todd and Poole only briefly described females of Aucula and their genitalia. A
more complete description of female A. franclemonti is given here based on three
specimens, two mated females caught in the field (one is the female parent of CMNH
Culture 83-186), and a single virgin female eclosing in the laboratory (CMNH Culture
83-351).

General. Maculation, color, and shape as in males with the following differences:
size slightly larger than males (FWL = 27-28 mm); costa of forewing slightly but
distinctly angled just beyond middle; apical segment of labial palp more than twice
as long as wide (length and width subequal in males).

Female genitalia. Figure 22. Lobes of papillae anales subtriangular due to slight
elongation of posterior dorsal angle; lobes sparsely setose, more densely so near
posterior margins; posterior apophyses compressed and widened near base, cylin-
drical anteriorly, subequal in length to papillae anales; Tergum 8 unmodified; anterior
apophyses slightly shorter than posterior apophyses; ostium bursae small, circular,
sclerotized, located in membranous protuberance between ventral edges of Tergum
8; ductus bursae straight, slender, membranous, sclerotized only at extreme base by
ostium bursae; corpus bursae membranous, ovoid, tapered smoothly into ductus
bursae; anterior enlargement of corpus bursae (fundus bursae) without signa but with

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£

£

Figs. 22-23. Female genitalia of Aucula franclemonti (Genitalia slide: Rawlins 859 (CMNH)).
22. Ventral view without sphragis. 23. Detail of ostial region with sphragis in position.

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Figs. 24-25. Ventral view of Aucula species. 24. Female of A. franclemonti (CMNH Culture
No. 83-351) during reflex immobilization following a slight disturbance. 25. Ventral view of
A. jenia, male; forewing length 23 mm; Ecuador, Chimborazo (CMNH).

sparsely scattered acanthae; small medial zone of irregular sclerotization anterior to
ostium bursae, reflexed under Sternum 7; posterior margin of Sternum 7 sclerotized,
broadly concave. Mated female without discemable internal spermatophore; plug of
amorphous sclerotized material in ostium bursae (Fig. 23), extended inward along
short sclerotized base of ductus bursae, and outward over protuberance encircling
ostium.

Discussion. The external female genitalia are structurally correlated in size and
shape with the male genitalia. The small ostium and adjacent sclerotized portion of
the ductus bursae match precisely in diameter and length the permanently everted
and strongly sclerotized endophalli of males.

A sclerotized plug was present in the ostia bursarum of both mated females, and
absent in the single female known to be virgin by virtue of its having eclosed in
isolation. The chitinous nature of this copulatory obstruction suggests that it is
probably homologous to the collum of the unmodified noctuid spermatophore. Its
presence in both mated females available for study, and its absence in a virgin female,
suggests that is is functioning as a sphragis preventing multiple matings. I am not
aware of sphragides elsewhere in the Noctuidae.

LIFE HISTORY NOTES ON AUCULA FRANCLEMONTI

A female of A. franclemonti was collected on 5 July 1983 at the same Ecuadorian
locality as the series of 23 males discussed above. A search was made for likely
foodplants in the Onagraceae, Vitaceae, and Portulacaceae in anticipation of obtain-
ing another gravid female. Another female was taken on the night of July 1 1. When

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Figs. 26-27. Immature stages of Aucula franclemonti (CMNH Culture No. 83-351). 26.
Last larval instar resting on under surface of leaves of foodplant, Vitis tiliifolia. 27. Pupa exposed
in pupal chamber formed by boring in soft wood; pupal position reversed from that found in
nature.

placed in a small container with leaves of a wild grape {Vitis, Vitaceae), it deposited
28 eggs by the morning of July 12. These hatched after an incubation period of
slightly more than four days. Five eggs and five first instars were preserved (CMNH
Culture 83-186), and the remaining larvae began feeding at once on the dorsal surface
of wild grape leaves. First instar larvae remained relatively immobile on the leaves
while feeding, but when disturbed moved about rapidly as “semiloopers” with prolegs
advancing the length of two or more segments at each step (multisegmental loco-

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motion). Detailed developmental observations were not made, but three late third
instars were preserved on July 26, and two last larval instars (fifth) were preserved
on August 3, 18 days after hatching. The last three instars moved by advancing the
prolegs a single segment per step without semilooping (unisegmental locomotion),
and fed by biting leaf edges.

During the rapid development of this isofemale culture, larvae were sought in the
field on foliage of the same species of Vitis. During the night of 26 July 1983, a last
instar larva was found feeding by Mark Smyers (Fig. 26). On July 3 1 , the larva began
boring into rotting wood, constructing a pupation chamber which was rendered
observable by carefully splitting the piece of wood into which it had bored. The
chamber was completed by August 1, and the prepupa remained inactive for four
days, pupating on August 5 (Fig. 27). The pupal chamber was transported back to a
laboratory at the University of Texas, Austin. By rare good fortune, eclosion of a
female moth was directly observed on 1 1 September 1983 at 10:00 hours CST. The
freshly eclosed moth climbed up to an elevated position in the laboratory enclosure
and remained motionless for nearly 40 minutes before expansion of the wings. When
disturbed, this virgin female defended itself by reflex immobilization with the wings
appressed stiffly above the abdomen and the abdomen expanded and strongly curved
under the thorax (Fig. 24), similar to the behavior of wild-caught males and other
aposematic Lepidoptera, including a diversity of sympatric Arctiidae (Watson, 1975).

Foodplant: Vitis tiliifolia Humb. & Bonpl. ex. Roem. & Schultes; voucher specimen
in herbarium of Carnegie Museum, Thompson and Rawlins No. 811, collected 10
July 1983. Determination by Michael O. Moore, 1990, The University of Georgia.

The immature stages of A. franclemonti are described briefly below.

Egg: Deposited singly or in clumps of 2-3 against major veins on underside of
leaf of foodplant, partially or completely inserted under lanate pubescence. Pale
yellow when laid, turning reddish brown prior to hatch. Egg slightly variable in size
and shape, moderately flattened; diameter, 0.96-1.09 mm; height, 0.60-0.74 mm.
Microsculpture consisting of 22-25 vertical costulae only slightly raised above outline
of egg; all vertical costulae begin near outer edge of flattened base of egg, but only
half extend to micropylar rosette, the other half evanescing at about % the distance
to rosette; long and short vertical costulae alternating; vertical costulae connected by
diffuse transverse costulae, separating columns of 10-13 indistinct areoles. Micro-

Figs. 28-29. Chaetotaxy of the head of last instar larva of Aucula franclemonti. 28. Dor-
sofrontal view of head. 29. Lateral view of head. Symbols used: A1-A3, anterior setae; Aa,
anterior pore; acl, anteclypeus; af, adfrontal sclerite; AF1-AF2, adfrontal setae; AFa, adfrontal
pore; afs, adfrontal suture (=lateral adfrontal suture); cl, clypeus (=postclypeus of authors);
CL1-CL2, clypeal setae; el, ecdysial line; ep, epicranium; eps, epicranial suture (=coronal suture
or medial adfrontal suture); f, front; F 1 , frontal seta; Fa, frontal pore; fcls, frontoclypeal suture;
LI, lateral seta; La, lateral pore; lbr, labrum; LR1-LR6, labral setae; MD1-MD3, dorsal pro-
prioceptor setae; MDa, dorsal pore; MG1, genal proprioceptor seta; MGa, genal pore; P1-P2,
posterior setae; Pa-Pb, posterior pores of head; poc, postocciput; pocs, postoccipital suture;
S1-S3, stemmatal setae; Sa-Sb, posterior stemmatal pores; SS1-SS3, substemmatal setae; SSa,
substemmatal pore.

1 mm

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pylar rosette less than 0. 10 mm in diameter, surrounded by a slight depression joining
upper ends of vertical costulae.

First instar: Length, 2. 8-3. 2 mm; width of head, 0.52 mm. Head yellowish brown,
body pale grayish white with long white setae arising from contrasting brown pinac-
ulae. Tactile setae as in last instar with the following exceptions: only one SV seta
present on A1 and A7, two SV setae present on A2; five coxal setae (DC1-DC2
missing, Figs. 36-37). Prolegs on A5 and A6 subequal, those on A4 slightly smaller,
those on A3 still smaller.

Intermediate instars: Head yellow, with setal bases minutely but contrastingly dark
brown. Body color pale reddish gray, densely spotted with dark spots vaguely forming
broken transverse stripes. Two SV setae on A7, the anteriormost seta (SV2) much
smaller than the posterior and on a separate pinaculum.

Last instar: Length, 48 mm; head width, 4.5 mm; width of mandible, 1.16 mm.
Head blocky, strongly hypognathous, width greater than prothorax. Body elongate,
enlarged gradually posteriorly, distinctly swollen on Ab8; most setae long, conspic-
uous, erect (especially on segment Ab8), set in distinct, slightly elevated pinaculae
which are enlarged dorsally (especially dorsal pinaculae on segment Ab8). Detailed
chaetotaxy and external morphology presented below.

Head pale brownish yellow, bases of setae brownish black. Thorax and abdomen
with ground color light gray tinted with olivaceous to yellowish brown on middle of
segments; pattern of dark gray to black spots, forming irregular transverse bands on
posterior portion of segments; transverse pattern less distinct anteriorly on each
segment; segments T2 to Ab9 with large blotches of white laterally, most extensive
on sides of Ab7 in subdorsal and lateral regions; ventral region on all segments pale
yellowish white, contrasting with adjacent subventral color. Prothoracic and anal
shields concolorous with head; thoracic legs brownish yellow on ventral and lateral
surfaces, darker brown on dorsal surface; subventral pinaculum above prolegs yel-
lowish brown dorsally, darker brown ventrally; spiracles black; most setae pale gray
to white.

Pupa (terminology follows Mosher, 1916): Length, 22 mm. Pupa very dark reddish
brown, dull in lustre due to strong microsculpture which is finely spinulose dorsally.
Labial palpus visible externally; base of maxilla extended laterally to sculptured
eyepiece; prothoracic femur not visible; mesothoracic leg not extended anteriorly to
sculptured eyepiece (externally not contiguous with eyepiece); terga 3 and 4 separated
by flexible cuticle, not fused; cremaster truncate, with flattened lateral processes
bearing reduced setae along posterior margins.

Discussion: Although the egg does not differ in a distinctive way from many other
non-agaristine noctuids, the larval and pupal stages of A. franclemonti are charac-
teristic of those found in a diverse representation of other agaristines that I have
reared and studied in detail (e.g., Alypia octomaculata (Fabricius), Brephos nyassana
(Bartel), Chelonomorpha formosana Miyake, Euscirrhopterus gloveri Grote and Rob-
inson, Eudryas grata (Fabricius), Eudryas unio (Hiibner), Mimeusemia vilemani
Hampson, Psychomorpha epimenis (Drury), Tuerta chrysochlora Walker, Xerociris
wi/sonii (Grote)). In particular, mature larvae are characterized by pale, hypognathous
head capsules with setal insertions often accentuated with dark spots; body segments
with contrasting transverse patterns of 1-8 bands, sometimes broken into spots
( Aucula ) or obscured by a dark ground ( Xerociris ), often with patches of orange or
white laterally. These larval generalizations hold for many species described in some

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Figs. 30-32. Mouthparts of last instar larva of Aucula franclemonti. 30. Medial surface of
left mandible. 3 1 . Left lateral view of hypopharyngeal complex, labial and maxillary palp;
removed. 32. Left antenna. Symbols used: amc, anterior mandibular condyle; AN 1-AN2, setae
of an2; anl-an3, antennal segments; ANa, pore of an2; anf, antennifer; elf, transverse cleft of
hypopharynx; dlsp, distolateral spinules; dst, dististipes (=palpifer of authors); mrO, outer man-
dibular ridge; mrl-mr3, inner mandibular ridges; mt, inner tooth; MX1-MX2, setae of max-
illary stipes; MXa, pore of basistipes; pig, palpiger (=stipes labii of Crumb (1929)); pm, post-
mentum; PM1, postmental seta; pmc, posterior mandibular condyle; prlsp, proximolateral
spinules; prm, prementum; sat, subapical cleft; 1 1— 13, apical teeth.

detail by Gardner (1946, 1948; Aegocera venulia (Cramer), Sarbanissa albifascia
(Walker)), by Crumb (1956; Alypia langtonii Couper, Alypia ridingsii Grote, Andro-
loma maccullochii (Kirby) (but body pattern longitudinal), Eudryas brevipennis Stretch,
Euscirrhopterus cosyra (Druce)), those figured by Sugi (1987; Asteropetes noctuina
(Butler), Chelonomorpha japana Motschulsky, Maikona jezoensis Matsumura, Sar-
banissa subflava (Moore), S. venusta (Leech)) and those figured by Common (1990;
Agarista agricola (Donovan), Argyrolepidia subaspersa (Walker), Cruria donowani
(Boisduval), Comocrus behri (Angas), Periscepta polysticta (Butler), Phalaenoides
tristifica (Hiibner)). Of all agaristines known to this author, the immature stages of
A. franclemonti most closely resemble those of Mimeusemia vilemani from Taiwan.

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Fig. 33. Maxillolabial complex of last instar larva of Aucula franclemonti (semischematic).
Symbols used: bst, basistipes; cd, cardo; dst, dististipes; fs, fusiliger; LB1, stipular seta of labium;
LBa-LBc, pores of palpiger; lg, laciniogalea; LGa, pore of laciniogalea (=Mxe of Gerasimov);
LP1, apical seta of lpl; Ip 1 , basal segment of labial palpus; lp2, distal segment of labial palpus;
LSS, lateral sensillum styloconicum of laciniogalea; mpl-mp3, segments of maxillary palpus;
MPa, pore of basal segment of maxillary palpus (=Mxd of Gerasimov); MPbl, lateral pore of
mp2 (=Mxf of Gerasimov); MPb2, medial sensillum of mp2; MPc, pore of mp3 (=Mxj of
Gerasimov (1952)); =SD of Grimes and Neunzig (1986a); MSS, medial sensillum styloconicum
of laciniogalea; MX1-MX3, setae of maxillary stipes; MX4, seta of first segment of maxillary
palpus; MX5-MX7, apical setae of maxilla (=STIII-STI of Grimes and Neunzig, 1 986b)); MXa,
pore of basistipes; pig, palpiger; pm, postmentum; PM1, postmental seta; pmsc, posterior
sclerotization of postmentum; prm, prementum; sp, spinneret.

Figs. 34-35. Chaetotaxy of the body segments of last larval instar of Aucula franclemonti
(semischematic views). 34. Left side of thorax and first two abdominal segments. 35. Left side
of abdominal segments 6-10. Symbols used (broken outlines indicate features hidden from
view by other structures): Abl-AblO, abdominal segments; ad, adenosma (ventral cervical

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gland); D1-D2, dorsal setae; L1-L3, lateral setae; LAa, lateral pore of AblO; MD1, dorsal
proprioceptor seta; MSD1-MSD2, subdorsal proprioceptor setae; MV 1-MV3, ventral proprio-
ceptor setae; MXD1, posterior proprioceptor seta of Tl; PP1, paraproct seta; SD1-SD2, sub-
dorsal setae; spr, spiracle; SV1-SV4, subventral setae; T1-T3, thoracic segments; VI, ventral
seta; XD1-XD2, anterior dorsal setae of Tl; XDa-XDc, anterodorsal pores of Tl.

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Figs. 36-37. Chaetotaxy of the prothoracic leg of last instar larva of Aucula franclemonti.
36. Anterior view of left prothoracic leg. 37. Posterior view of left prothoracic leg. Symbols
used: Cl, seta of anterior coxal sclerite; C2-C5, setae of posterior coxal sclerite; clw, claw of
pretarsus; cxa, anterior sclerite of coxa; exp, posterior sclerite of coxa; exs, coxal suture; DC1-

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The pupation chamber and pupa of A. franclemonti are also very similar to those
of agaristine species I have studied (see list above). Pupation occurs in chambers
bored in soft wood. Pupae characteristically have heavy microsculpture, are often
spinulose dorsally, and the truncate cremaster bears flattened lateral flanges with
setae greatly reduced or absent. The labial palpi are visible, the prothoracic femora
are not, and the mesothoracic leg is not contiguous with the eyepiece. Terga 3 and
4 are not fused but are closely joined by flexible intersegmental cuticle.

The vitaceous host of A. franclemonti is expected, as most of the known foodplants
for agaristine species worldwide are in the Vitaceae, including Leeaceae (Rhamnales),
and the remainder in the following unrelated families of plants: Dilleniaceae (Dil-
leniales); Lauraceae (Laurales); Onagraceae (Myrtales); Haloragaceae (Haloragales);
Portulacaceae, Nyctaginaceae and Cactaceae (Caryophyllales); Loranthaceae (San-
talales); Boraginaceae (Lamiales); Rubiaceae (Rubiales); Araceae (Arales); Smilaca-
ceae and Dioscoreaceae (Liliales).

CHAETOTAXY OF THE MATURE LARVA OF AUCULA FRANCLEMONTI

Chaetotaxy and general morphological features are presented in considerable detail
in Figure 28-37. No previous author has presented exhaustively the chaetotaxy of
an entire larval noctuoid, although a consistently applied terminology for most setae
has emerged in recent years as best summarized by Stehr (1987). Current setal
nomenclature for Lepidoptera larvae results from a long and confusing history of
comparative study including Fracker (1915), Schierbeek (1917), Hinton (1946), Ger-
asimov (1952), and Hasenfuss (1963). Descriptive work with noctuoid larvae has
become increasingly refined since early work by Ripley (1923) and Crumb (1929),
through comprehensive, regional treatments by Gardner ( 1 946, 1 948), Crumb (1956),
Beck (1960), and Godfrey (1972, 1987), to an exceptionally detailed and well-illus-
trated work by Miller (1991) who illustrated the partial chaetotaxy of the agaristine
Alypia octomaculata. Beck (1960) and Godfrey (1972) provide nomenclature for the
hypopharyngeal complex, and Godfrey (1972) supplies a general rationale for de-
scription of the mandible, similar in principle to that proposed by Gerasimov (1952).
A review of chaetotaxy for abdominal segment 10 was given by Stehr (1987), but
locations for the various setae were not clearly illustrated and application of his
terminology remains problematic for lateral and subventral setae of the last abdom-
inal segment.

Two systems, the maxillolabial complex and the thoracic legs, have not been
consistently illustrated or labelled in larval descriptions, although Gerasimov (1952)
offered a system naming most setae. Grimes and Neunzig (1986a, 1986b) provided
a clear nomenclature for apical sensillae of the terminal segment of the maxillary
palpus and the laciniogaleae, but did not offer terminology for more basal setae or
sensillae of the maxilla. The setae of the thoracic legs were labelled in part by

DC2, dorsal subprimary seta of coxa; fe, femur; FE1-FE2, femoral setae; MCI, proprioceptor
seta of anterior coxal sclerite; MTR1-MTR3, proprioceptor setae of trochanter; MTRa, antero-
dorsal pore of trochanter; ti, tibia; T11-T16, tibial setae; tr, trochanter; ts, tarsus; TS1-TS4,
tarsal setae.

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Gerasimov (1952), but his system was not closely or consistently followed by recent
treatments (Nielsen and Kristensen, 1989; Miller, 1991).

Chaetotaxal nomenclature adopted here follows the Hinton system (1946) as sum-
marized by Stehr (1987) but agrees with Singh and Goel’s (1987) interpretation of
subdorsal abdominal setae. Nomenclature of setae and pores on the prothorax follows
Hinton (1946), not Hasenfuss (1963), and the names of those on abdominal segment
10 follows Stehr (1987). Terminology for the mandibles follows Gerasimov (1952)
and Godfrey (1972); that for the hypopharyngeal complex follows Godfrey (1972).
Names for setae and pores of the antenna, maxilla, and labium follow Gerasimov
(1952) where possible, conflicting with Grimes and Neunzig (1986b) only with respect
to three setae (sensillae) of the laciniogalea. Finally nomenclature for the setae and
pores of the thoracic legs follows Gerasimov (1 952) in part, differing from the systems
proposed by Miller (1991) and Nielsen and Kristensen (1989).

Names and symbols for setae newly applied in this paper are in agreement with
general principles of a system first introduced by Hinton (1946). His nomenclature
based on relative morphological position may be rendered useful, informative, and
straightforward by attention to synonymy, homonymy, and priority of usage by earlier
authors. Numbering of setae within groups follows general trends and facilitates rapid
and accurate recognition of setae: ontogenetic appearance, anterior to posterior, dorsal
to ventral, medial to lateral, and sequential continuity around circular or cylindrical
structures. Where possible, symbols for setae and pores are chosen as mnemonic
abbreviations of the names they represent. Abbreviations for a few head setae have
been changed from those of Hinton to avoid homonymy with those on other body
parts, but the L and MD setae of the head and body are still homonymous. Finally,
symbols for setae and pores are entirely upper case followed by numbers for setae
and lower case letters for pores; symbols for sutures, sclerites, and other structures
and regions are lower case with the exception of traditional symbols for segments,
T1-T3, Abl-AblO.

Head: Figures 28-29. Only a single substemmatal pore and a single genal proprio-
ceptor seta on each side of head; six labral setae (here symbolized LR 1-LR6 following
a peripheral sequence, medial to lateral, around each labral lobe) but no obvious
labral pores; three stout but otherwise unmodified epipharyngeal setae.

Antenna: Figure 32.

Mandible: Figure 30. Three distinct apical teeth are subtended by three inner ridges,
the first ridge as numbered here originates from the posterior side of a truncate, inner
mandibular tooth; a tiny subapical cleft present where outer (posterior) mandibular
ridge joins distal margin of mandible.

Maxilla: Figure 33. Nomenclature for apical setae follows Gerasimov (1952), MX5-
MX7, equalling STIII to STI of Grimes and Neunzig (1986b); pore MPc is the same
as Mxj of Gerasimov (1952) and the sensillum digitiformium (SD) of Grimes and
Neunzig (1986a).

Hypopharyngeal complex: Figure 3 1 . Proximolateral spinules long, well developed;
medial surface of hypopharynx smooth; transverse cleft indistinct, not sharply incised.

Labium: Figure 31, 33. Nomenclature for divisions of the mentum follows Snod-
grass (1935). Spinneret almost as wide as long, membranous, broadly tubular.

Thorax: Figure 34. Two SV setae on segment Tl, and one SV seta on segments
T2-T3. Seta L2 on segment T l and seta SD1 on segments T1-T3 are very attenuated.

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straight sensillae borne in darkly pigmented pits with densely microtrichiated walls
(tonosensillae: Rawlins, 1984). These setae are receptors of airborne sound in no-
todontid larvae (Tautz and Markl, 1978).

Abdomen: Figures 34-35. Four SV setae present on segment AblO, three SV setae
present on segments A2-A6, two SV setae present on segments Abl, Ab7, and Ab8,
and one SV seta present on segment Ab9. Seta LI posterior to spiracle on segments
Abl-Ab6 and Ab8; below spiracle on segment Ab7; spiracles subequal on segments
Abl-Ab7, enlarged on Ab8; seta SD1 on segment Ab9 normal in size and shape,
not a tonosensillum.

Thoracic legs: (Figs. 36-37). Coxa divided into anterior and posterior sclerites by
an anteroventral coxal suture extended as coxal condyle articulating with trochanter;
five primary coxal setae present, one (Cl) on anterior coxal sclerite, and four (C2-
C5) on posterior coxal sclerite, numbered in continuous sequence around coxa; one
small (or proprioceptor) seta on basal edge of anterior coxal sclerite near coxal suture
(MCI); one reduced, subprimary seta basal to seta Cl (DC1), and similar reduced
seta (DC2) basal to seta C5. Three proprioceptor setae in membranous gap between
trochanter and base of femur, numbered sequentially around segment from a rela-
tively anterodorsal position dorsal to coxal suture (MTR1), to an anteroventral po-
sition below coxal suture (MTR2), to a posteroventral position (MTR3); a pore
(MTRa) situated between seta MTR1 and point opposite coxal suture. Femur without
pores; with two setae, anterior (FE1) and posterior (FE2). Tibia with a single pos-
terodorsal pore (Tla) and six setae encircling apex, numbered as Gerasimov (1952)
in a continual sequence: anterodorsal (Til), anterolateral (T12), anteroventral (T13),
posteroventral (T14), posterolateral (T15), posterodorsal (T16). Tarsus with four setae,
numbered as the tibia: anterodorsal (TS1, a small seta), anteroventral (TS2), pos-
teroventral (TS3, just distad of TS2), and posterodorsal (TS4, larger than TS1); setae
TS2 and TS3 stout, slightly flattened and bladelike.

Discussion. Three features of the chaetotaxy of Aucula franclemonti deserve special
emphasis: the presence of two SV setae on segment Abl, two SV setae on segment
Ab7, and a normal, non-attenuated seta SD1 on segment Ab9. The number of SV
setae on the first abdominal segment has been an important character in the classi-
fication of Lepidoptera larvae, noctuoid and otherwise. The significance of three SV
setae on Abl was stressed by Gardner (1946, 1948) and by Crumb (1956) in rec-
ognizing several subfamilies of Noctuidae (all Catocalinae sensu lato, Herminiinae,
Bagisarinae, and Hypeninae) as well as several species of Plusiinae, Acontiinae sensu
lato (nearctic species of Lithacodia and Eublemma), and Amphipyrinae (species
variously placed by authors in Perigea, Platysenta, Condica, and Leuconycta ). All
agaristine larvae examined in this study had two S V setae on Ab 1 , the usual condition
for the “trifid” subfamilies.

The presence of two SV setae on segment Ab7 appears to be a diagnostic feature
of most Agaristinae, and it was utilized in a key to subfamilies for North American
species by Crumb (1956). I have examined larvae of species worldwide (listed above),
and all have two SV setae on Ab7 except those of Psychomorpha epimenis, a North
American species placed in the Agaristinae without comment by Crumb (1956). Only
a single SV seta was found on Ab7 in noctuid taxa of controversial affinities, including
those known to have additional SV setae on thoracic or other abdominal segments:
Litoprosopus futilis (Grote and Robinson), Diaphone eumela (Stoll), Xanthopastis

308

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

timais (Cramer), Diphthera Jestiva (Fabricius). The second SV seta on Ab7 is absent
in first instar larvae of Agaristinae, and its presence in subsequent instars appears to
be an excellent synapomorphy of most species in the subfamily.

Two SV setae have been observed on Ab7 of the very similar larvae of New World
Bagisara species ( B . rectifascia (Grote) and B. repanda (Fabricius)) and Old World
Xanthodes transversa Guenee. The former genus is placed by most authors in Acon-
tiinae, and the latter in Chloephorinae by Poole (1989), but both may best be treated
as a distinct taxon, Bagisarinae (or Bagisarini of the Acontiinae), as established by
Crumb (1956). The additional subventral seta on Ab7 in both subfamilies is always
smaller than the usual subventral seta (SV1), anterior to it, and never on the same
pinaculum.

In agaristines the additional seta is always distinctly dorsal to the level of SV 1 , in
a position spatially identical to that of setae SV2 on the first and second abdominal
segments, and is therefore labelled SV2. In most agaristines, SV2 on Ab7 is slightly
dorsal to SV1, but in T. chrysochlora SV2 is positioned so far dorsad that its pinac-
ulum is fused with that of seta L3.

In bagisarines, the additional subventral seta on Ab7 is far ventrad of SV1, and
may be almost as close to the ventral midline as seta V 1 in a position spatially similar
to that of SV3 on the second abdominal segment. This seta in bagisarines should be
labelled SV3, and is not considered homologous to SV2 in agaristines.

Aucula franclemonti and all other Agaristinae examined lack a tonosensillum on
segment Ab9. As on the thorax, the seta that is modified into a tonosensillum is
SD1. The presence of this tonosensillum in many “trifid” noctuids (Abrostolini,
Amphipyrinae, Cuculliinae, Glottulini, Hadeninae, Heliothinae, Noctuinae, Stiriini),
but not in other Noctuoidea, suggests that it may be synapomorphic for a major
lineage of Noctuidae. A comprehensive, worldwide survey of this setal modification
has not been undertaken, but its absence in Agaristinae does not support their in-
clusion in an extensive “trifid” lineage bearing the modified seta.

ACKNOWLEDGMENTS

Special appreciation is due the following for helping with this study: M. A. Klingler for his
detailed illustrations; G. L. Godfrey, C. W. Young and R. L. Davidson for review of the
manuscript; M. A. Daman and W. A. Zanol for assistance in specimen and manuscript prep-
aration; M. Smyers for exceptional success in caterpillar-hunting in 1983; I. Kitching and M.
Honey for facilitating study at The Natural History Museum, London; S. A. Thompson for
foodplant vouchering and obtaining an authoritative foodplant identification from M. O. Moore;
G. Onore for logistical help in Ecuador. Last, but not least, I thank J. G. Franclemont for so
generously sharing with me his truly insatiable passion for moths and their larvae.

LITERATURE CITED

Beck, H. 1960. Die larvalsystematik der Eulen (Noctuidae). Abhandl. Larvalsyst. Ins. (Aka-
demie-Verlag, Berlin) 4:1-406.

Common, I. F. B. 1990. Moths of Australia. E. J. Brill, New York.

Crumb, S. E. 1929. Tobacco cutworms. USDA Tech. Bull. 88:1-179.

Crumb, S. E. 1956. The larvae of the Phalaenidae. USDA Tech. Bull. 1 135:1-356.

1992

AUCULA FRANCLEMONTI

309

Fracker, S. B. 1915. The classification of lepidopterous larvae. Illinois Biol. Monogr. 2:1—
169.

Gardner, J. C. M. 1946. On larvae of the Noctuidae (Lepidoptera)— II. Trans. Roy. Ent. Soc.
Lond. 97:237-252.

Gardner, J. C. M. 1948. On larvae of the Noctuidae (Lepidoptera)— VI. Trans. Roy. Ent. Soc.
Lond. 9:291-318.

Gerasimov, A. M. 1952. Caterpillars, Part 1. Fauna SSSR: Insects— Lepidoptera 1(2): 1-338.
Zool. Inst. Acad. Nauk SSSR, New Series, No. 56. [In Russian]

Godfrey, G. L. 1972. A review and reclassification of larvae of the subfamily Hadeninae
(Lepidoptera: Noctuidae) of America north of Mexico. USDA Tech. Bull. 1450:1-265.

Godfrey, G. L. 1987. Noctuidae (Noctuoidea). Pages 549-578 in: F. W. Stehr (ed.), Immature
Insects. Kendall/Hunt Publishing Co., Dubuque, Iowa.

Grimes, L. R. and H. H. Neunzig. 1986a. Morphological survey of the maxillae in last stage
larvae of the suborder Ditrysia (Lepidoptera): palpi. Ann. Ent. Soc. Am. 79:491-509.

Grimes, L. R. and H. H. Neunzig. 1986b. Morphological survey of the maxillae in last-stage
larvae of the suborder Ditrysia (Lepidoptera): mesal lobes (laciniogaleae). Ann. Ent. Soc.
Am. 79:510-526.

Hasenfuss, I. 1963. Eine vergleichen-morphologische Analyse der regularen Borstenmuster
der Lepidopterenlarven. Studien zur Methodik der vergleichenden Morphologie der Bor-
stenmuster und zur phylogenetischen Deutung der Abwandlungen der regularen Bor-
stenmuster der Lepidopterenlarven. Z. Morph. Okol. Tiere 52:197-364.

Hinton, H. E. 1 946. On the homology and nomenclature of the setae of lepidopterous larvae,
with some notes on the phylogeny of the Lepidoptera. Trans. Roy. Ent. Soc. Lond. 97:
1-37.

Kiriakoff, S. G. 1977. Lepidoptera Noctuiformes, Agaristidae. III. (American Genera). Das
Tierreich 99:1-82.

Miller, J. S. 1991. Cladistics and classification of the Notodontidae (Lepidoptera: Noctuoidea)
based on larval and adult morphology. Bull. Am. Mus. Nat. Hist. 204:1-230.

Mosher, E. 1916. A classification of the Lepidoptera based on characters of the pupa. Bull.
Illinois State Lab. Nat. Hist. 12(2): 14-1 59, plates 19-27.

Nielsen, E. S. and N. P. Kristensen. 1 989. Primitive Ghost Moths. Morphology and taxonomy
of the Australian genus Fraus Walker (Lepidoptera: Hepialidae 5. lat.). Monographs on
Australian Lepidoptera 1:1-206.

Poole, R. W. 1989. Lepidopterorum Catalogus (New Series). Fascicle 118. Noctuidae, Part
1. E. J. Brill/Flora and Fauna Publications, New York, pp. 1-500.

Rawlins, J. E. 1984. Mycophagy in Lepidoptera. Pages 382-483 in: Q. Wheeler and M.
Blackwell (eds.), Fungus-Insect Relationships. Perspectives in Ecology and Evolution.
Columbia University Press, New York.

Ripley, L. B. 1923. The external morphology and postembryology of noctuid larvae. Illinois
Biol. Monogr. 8:1-169.

Schierbeek, A. 1917. On the Setal Pattern of Caterpillars and Pupae. E. J. Brill, Leiden.

Singh, G. P. and S. C. Goel. 1987. Body chaetotaxy of three noctuid caterpillars. Deutsch.
Ent. Zeit. 34:373-381.

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

Stehr, F. W. 1987. Immature Insects. Kendall/Hunt Publishing Co., Dubuque, Iowa.

Sugi, S. (ed.). 1987. Larvae of Larger Moths in Japan. Kodansha Co. Ltd., Tokyo. [In Japanese
with English summary]

Tautz, J. and H. Markl. 1978. Caterpillars detect flying wasps by hairs sensitive to airborne
vibration. Behavioral Ecol. Sociobiol. 4:101-1 10.

Todd, E. L. and R. W. Poole. 1981. A revision of the agaristid genus Aucula Walker (Lepi-
doptera: Noctuidae). J. Lep. Soc. 35:194-215.

310

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Watson, A. 1975. A reclassification of the Arctiidae and Ctenuchidae formerly placed in the
thyretid genus Automolis Hiibner (Lepidoptera) with notes on warning coloration and
sound. Bull. British Mus. Nat. Hist. (Ent.), Suppl. 4:1-104, 34 plates.

Received 31 July 1991; accepted 22 Nov 1991.

J. New York Entomol. Soc. 100(2):31 1—319, 1992

TWENTY YEARS OF FLUCTUATING PARAPATRY AND THE
QUESTION OF COMPETITIVE EXCLUSION IN THE
BUTTERFLIES PONTIA OCCIDENTALIS AND
P. PROTODICE (LEPIDOPTERA: PIERIDAE)

Arthur M. Shapiro

Department of Zoology and Center for Population Biology,

University of California, Davis, California 95616

Abstract.— The sibling species Pontia protodice and P. occidentalis occur on altitudinal gra-
dient across the Sierra Nevada in California, with a mid-elevation belt on the west slope in
which neither is a permanent resident. A similar pattern occurs in a variety of unrelated butterfly
genera. Studies of the occurrence and breeding of the two Pontia across the Sierra for 20 years
fail to demonstrate any association (positive or negative) between them in the “no man’s land”
on the west slope, thus arguing against competitive exclusion as the mechanism maintaining
the separation. Occurrence of P. protodice at 1,500 m and 2,100 m is also independent, implying
that its presence in the montane Sierra is not simply an epiphenomenon of outflow from a
single source in the lowlands. 1 99 1 was an unusual year in which the entire region was colonized,
seemingly in a wave emanating from the east. Sporadic competition between the species is not
ruled out by these findings, but is unlikely to be important.

Claims that “Competition from X limited the distribution of Y” used to be com-
mon in the ecological literature, but the days when they could be flung about with
great abandon are over (Begon et ah, 1990). That does not, however, remove the
inherent seductiveness of competition-based explanations for community structure.
In western North America a repeating pattern exists of altitudinally parapatric sibling
species pairs (rarely triplets) of butterflies for which competitive-exclusion expla-
nations appear attractive (Shapiro et ah, 1981). In the Sierra Nevada this pattern
includes Everes comyntas (Godart) and E. amyntula (Bdv.) and L. helloides (Bdv.)
and L. nivalis (Bdv.)(Lycaenidae), Phyciodes campestris (Behr) and P. montana
(Behr)(Nymphalidae), Anthocharis sara Lucas and A. Stella (W. H. Edwards) and
Pontia protodice (Bdv. & LeC.) and P. occidentalis (Reakirt) (Pieridae), Thorybes
pylades (Scudder), T. diversus Bell, and T. mexicana (H.-S.) ssp., and Polites sabuleti
(Bdv.) and P. tecumseh (Grinnell) (Hesperiidae), and others. In each case a widespread
low-elevation species (the first named) is replaced in the higher Sierra by a closely-
related entity of more or less ambiguous (species or subspecies) status, with a band
at mid-elevation (1,300-1,700 m) in which neither occurs (or, in the Thorybes case,
a third species occurs). Shapiro et ah (1981) were the first to point out the “natural
experiment” in northern California, where the Sierran endemics Phyciodes montana
and Anthocharis Stella do not occur, and P. campestris and A. sara move upslope
and fill more or less of the montane niches occupied by their congeners in the Sierra.
This situation suggests either competitive exclusion in the Sierra or its obverse,
competitive release, in the northern mountains.

Since 1972 detailed phenological data on all butterfly species have been taken along
a transect parallel to Interstate Highway 80, from sea level across the Sierra Nevada

312

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

via a series of sampling stations over the crest to mid-elevation on the east slope.
Each station is sampled at roughly two-week intervals in good weather whenever
there is no significant snow cover; all butterfly species observed are recorded. This
program is designed to permit an assessment of the spatial and temporal stability
and patterning of the faunas. The data document the fluctuating parapatry across the
west slope “no man’s land” as well as fluctuating sympatry on the east slope in several
species pairs. This paper considers the transect data on Pontia protodice and P.
occidentalis and their relevance to the competitive exclusion hypothesis.

THE ORGANISMS

Pontia protodice and P. occidentalis are sufficiently similar morphologically to be
confounded in virtually all museum and many private collections, as well as in many
published works. Although Reakirt and W. H. Edwards were clear in their concepts
of the species, most subsequent Lepidopterists were not. The nadir of confusion was
reached by Abbott et al. (1960), who confounded all the North American Pontia,
but Hovanitz (1962) mapped the distributions of P. protodice and P. occidentalis
reasonably accurately, and Chang (1963) provided diagnostic morphological char-
acters for them. Following several years of field and laboratory studies of their ecology,
physiology and variation, Shapiro (1976) reviewed the status of all Nearctic names
in the group and concluded that these two were amply distinct at the species level.
He also concluded that the entity nelsoni (W. H. Edwards) from subarctic Alaska
was conspecific with occidentalis and that both were probably conspecific with the
Palearctic P. callidice (Hbn.) complex, which extends from the Alps and Pyrenees
eastward to northeast Siberia as a series of disjunct, mostly high montane-alpine
populations. According to Shapiro ( loc . cit.): “{Pontia) occidentalis occurs upslope
and northward of P. protodice in western North America. The two species have been
found sympatric at various locations from 1000-2 500m in the Rocky Mountains
and Sierra Nevada, e.g., Donner Pass, California, where P. occidentalis is a permanent
resident and P. protodice a breeding immigrant. ... In areas of sympatry occasional
interspecific matings may occur, but only 2 of 339 specimens collected at Donner
Pass in 1973 were phenotypically ambiguous.” Shapiro and Geiger (1986), using
California populations including sympatric ones along the 1-80 transect, demonstrat-
ed electrophoretically that they are distinct species and found no evidence for hy-
bridization at Sierra Valley, where they are in massive contact. However, hybrid-
ization appears fairly frequent in Colorado (J. Kingsolver, D. Wiemasz, pers. comm.).

Both species occur only in non-forested environments, including tundra, grassland,
high desert and steppe; P. protodice extends to low (hot) desert. In areas of sympatry,
the two species often swarm in dense, mixed flights, breeding on weedy mustards in
grain fields and by roadsides. The life cycles of both are very similar, though diagnostic
differences exist in the larva and pupa (Shapiro, 1976). Both species are multivoltine
(except P. occidentalis at the highest elevations) and feed on a variety of Cruciferae,
both native and introduced. Both are highly vagile, colonizing species, and P. occi-
dentalis has a “hilltopping” epigamic strategy (Shapiro 1975, 1978).

THE DATA

Tables 1, 2 and 4 present the occurrence of both species at the Sierran stations
along the 1-80 transect: Lang Crossing, Nevada Co., ±1,500 m (LC); Donner Pass,

Table 1. Occurrence and breeding of Pontia protodice at Lang Crossing (LC), Donner Pass (DP), and Castle Peak (CP), 1972-1991.

1992

TWENTY YEARS OF PARAPATRY

313

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314

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Table 2. Occurrence and breeding of Pontia Occident alis at Lang Crossing (LC), 1972-1991.
This species has been present and breeding in all years at DP and CP.

Year

LC (1,500 m)

1972

—

1973

—

1974

—

1975

19, viii. 1 3

1976

—

1977

19, iv.l7a

1978

2<5, 19, viii. 17; 1(5, ix.14; 19, x.6; BRED

1979

—

1980

29, vii. 14; 16, viii.4b; 106, 39, viii. 26; 16, ix.10; BRED

1981

—

1982

—

1983

19, ix. 12

1984

—

1985

—

1986

—

1987

16, 19, vii. 20; 16, ix.7; BRED

1988

19, vii. 22

1989

19, vi.22; 19, vii. 17; 19, viii. 5; 16, viii. 16; BRED

1990

—

1991

—

a Implies colonization and overwintering undetected in 1976.
b Apparent hybrid.

Nevada and Placer Cos., ±2,100 m (DP); Castle Peak, Nevada Co., ±2,700 m (CP);
and Sierra Valley, Sierra Co., ± 1,500 m (SV). Lang Crossing is a west-slope locality
with a mosaic of mixed mesic forest, montane chaparral and Canyon Live Oak xeric
woodland on metasedimentary balds, and wet and dry meadows. Donner Pass con-
tains mostly upper montane coniferous forest on a mixture of granite and andesite
substrates, but is penetrated by a large wet and dry meadow complex and disturbed
roadsides and railway rights-of-way which extend completely through the pass, from
one slope to the other; these are lined with Crucifers. Castle Peak (Shapiro, 1978)
has subalpine coniferous forest and dry alpine fell-fields on andesite. Sierra Valley
is a very large wet and dry meadow complex on Pleistocene lake alluvium, with some
alfalfa cultivation along the margins. It is hemmed in by coniferous forest to the west
(on the east face of the Sierra) and by a low range of volcanic hills to the east,
historically covered with open pine forest but now occupied by sagebrush steppe and
juniper woodland; beyond these hills lies the Great Basin high desert.

LC and DP have been sampled for 20 years (the sampling stations were moved
several km from their 1972 locations to permanent ones in 1973). CP has been
sampled for 20 years, but only 4-6 times/year due to its short flight season. SV was
sampled sporadically before 1 980 and systematically thereafter, but early-spring data
(including the very early flight periods of post-diapause Pontia) were obtained only
since 1985.

Host plants recorded at the transect stations are presented in Table 5.

1992

TWENTY YEARS OF PARAPATRY

315

Table 3. Dates of first records of Pontia protodice at Sierra Valley (SV), where the species
is apparently always present in summer. In 1988 and 1990 the first (overwintered) brood,
“ vernalis was recorded in April, and the second date corresponds to other years as the beginning
of the summer flight.

Year

First date

1985

vi.20

1986

vii. 1 1

1987

vii. 1

1988

iv. 14; vi.19

1989

vi.3

1990

iv. 12; vii. 7

1991

vi. 13

The data are divided into “occurrence” and “breeding” years. Unavoidably, this
classification is somewhat arbitrary; deciding whether or not breeding has occurred
is sometimes not easy. Multiple captures on the same day at the same site are not
necessarily indicative of local origin (see discussion of dispersal, below). Interpre-
tation of the data involves: consideration of timing and previous frequency, and
intervals of sampling; numbers, condition, and sex-ratio of the animals observed;
and appropriateness of phenotype relative to season, both locally and in potential
source areas (both species are seasonally polyphenic). By these criteria many multiple
records have been interpreted as evidence of immigration rather than breeding from
a previously undetected colonist. The reality of undetected colonists, however, is
underscored by the discovery on 18 July 1986 of a large larva of P. protodice at DP
in a year when no adults of that species were seen there. For all years in which
breeding is claimed in the tables, the numbers and sexes of adults seen at the first
observation are noted. Criteria for recognizing breeding include: direct observation
of oviposition and/or early stages; ongoing (essentially continuous) presence of adults
over several consecutive samples; presence of newly-emerged or teneral individuals;
and appearance of two or more consecutive emergences at intervals appropriate for
successive generations in situ.

Table 4. Status of both species at all four sites, as fractions of years present or breeding
(1986 was not treated as a breeding year for P. protodice at DP). For criteria used to assess
breeding, see text.

LC

DP

CP

sv

Occurrence

P. protodice

15/20

16/20

3/20

10/10

P. occidentalis

8/20

20/20

20/20

10/10

Breeding

P. protodice

5/20

7/20

0/20

10/10

P. occidentalis

4/20

20/20

20/20

10/10

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Table 5. Hosts (all Cruciferae) recorded for Pontia protodice (P) and P. occidentalis (O) at
LC, DP and SV, 1972-1991. Additional hosts occur for O at CP.

LC

DP

sv

Lepidium virginicum var. pubescens (Greene) Thell.

p,o

p,o

p,o

Lepidium perfoliatum L.

p,o

Sisymbrium altissimum L.

p,o

p,o

Descurainia sophia (L.) Webb

p,o

p,o

Descurainia pinnata (Walt.) Brit.

p,o

Cradaria pubescens (Mey.) Roll.

p,o

p,o

Streptanthus tortuosus Kell.

o

o

RESULTS

Pontia occidentalis is a permanent resident at DP, CP and SV. This is shown by
the predictable occurrence of the vernal, post-diapause brood “ calyce (W. H. Ed-
wards)” (Shapiro, 1976) shortly after snowmelt, though it flies so early it is easily
missed. This is not the case at LC, and it very rarely overwinters there.

Pontia protodice is not a permanent resident at LC, DP or CP. Its status at SV is
uncertain. It has been common or abundant there in summer, with 2-3 generations,
every year for nearly 20 years, but early spring data are only available for seven years
and in only two of these was the vernal, post-diapause brood “ vernalis (W.H. Ed-
wards)” (Shapiro, 1976) observed (Table 3). In the Nevada desert and on the floor
of the Sacramento Valley (Shapiro, 1979 and unpublished), P. protodice overwinters
only locally and sporadically. Failure to detect it in early spring in Sierra Valley may
not establish its status in a broader, regional context.

Upslope movement by P. protodice on a seasonal basis is strongly supported by
circumstantial evidence. In 1991, for example, no protodice were seen at SV through
1 June. On 13 June, however, they were abundant and uniformly worn and of the
second-brood phenotype that was flying at that time around Reno. Females oviposited
on roadside Crucifers. Given the climatic and vegetational differences between Reno

Table 6. Comparisons of observed and (in parentheses) expected frequencies of P. protodice
(P) and P. occidentalis (O) alone and in combination at LC, 1972-1991. Neither distribution
differs statistically from mutual independence (unadjusted x2 values; x2 adjusted for continuity
also NS).

p + p -

Occurrence

o +

7(6)

1(2)

o -

- 8(9)

X2 = 1.055, NS

4(3)

Breeding

O +

2(1)

2(3)

o -

3(4)

X2 = 1.67, NS

13 (12)

1992

TWENTY YEARS OF PARAPATRY

317

Table 7. Comparisons of observed and (in parentheses) expected frequencies of occurrence
and breeding of P. protodice at LC vs. DP. (These are unadjusted x2', adjusted x2 are NS for
both comparisons, x2adJbreed,ng = 118*)

LC +

LC -

Occurrence

DP +

12 (12.75)

5 (4.25)

DP -

3 (2.25)
x2 = 1.54, NS

0 (0.75)

Breeding

DP +

3 (1.50)

3 (4.50)

DP -

2 (3.50)

X2 = 2.96, .05 < P < .10

12(10.50)

and SV, this apparent synchronization is unlikely to be coincidental. On 21 June a
male of this phenotype, in poor condition, was taken at DP. On 18 June both sexes
were common between Gardnerville, NV and Woodfords, Alpine Co., CA and along
the Kingsbury Grade highway to South Lake Tahoe, where they were flying upslope
along the roadside. This sort of movement seems to occur every year, albeit usually
less conspicuously, and could account for the abundance and reliable occurrence of
P. protodice at SV each year without overwintering.

What can 20 years of occurrence and breeding data along our transect tell us?
Given that neither species persists at LC, that site is particularly important as a
potential indicator of competitive exclusion: is the occurrence of the two species
either positively or negatively correlated? A negative association could imply either
competitive exclusion or reciprocal conditions for colonization (without regard to
competition — for example, wet years might favor one species, dry years the other).
A positive association would argue against competitive exclusion, suggesting that the
same conditions favor colonization by both. There is a single remarkable record of
a long-range dispersal by P. occidentalis, resulting in its capture in prime P. protodice
habitat near sea level (Shapiro, 1977)— suggesting that the two species have similar
habitat preferences as expressed in active habitat selection.

Table 6 shows that both occurrence and breeding data do not differ significantly
from the values expected if the two species were completely independent.

Also of interest is the extent to which occurrence and breeding of the two species
are linked among LC, DP and CP. If occurrence at these sites is an epiphenomenon
of dispersal from a uniform source (either east or west of the Sierra) we would expect
a strong association, while colonization from both sides would produce much weaker
association unless population densities in both potential source regions were tightly
coupled. Table 7 does not indicate strong association among sites. Mere occurrence
is indistinguishable from independence between LC and DP, while the hint of as-
sociation for breeding is entirely due to 1991 (and disappears if that year is not
considered). 1 99 1 was in fact a very unusual year for P. protodice. As Table 1 indicates,
it did eventually show up at LC and breed there. But it continued spreading westward
into the lower foothills, where it merged with an already established population in

318

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

the Sacramento Valley; 1991 was the first year since 1977 in which P. protodice
spread very widely (it reached the Oregon border in Modoc and Siskiyou Counties)
and the only year in the history of the transect that it occurred at all ten stations. P.
protodice is so rare at CP (Table 1) that no analysis is feasible, or necessary, beyond
noting that 1988 and 1991 were breeding years at DP, while 1986 was marked by
only the single larva mentioned above.

These data thus give no support to competitive exclusion as the cause of either
the “no man’s land” on the Sierran west slope or the altitudinal stratification of
Pontia overall. It must be stressed that the sort of comparison done here is not
powerful enough to detect competition itself; it can only be suggestive. Nor do these
results rule out the possibility of local competition arising as a consequence of the
spatial pattern of egg dispersion on hosts (Shapiro, 1975), with no larger-scale impact
on distribution.

As Begon et al. ( 1 990) point out, it may not be possible to exclude prior competition
and character displacement as factors leading to contemporary patterns of replace-
ment or apparent exclusion. Unless that hypothesis is testable, however, it is not
very interesting. Altitudinal parapatry is a common pattern in a variety of organisms.
In the tropical Andes much more complex patterns can be found in butterflies than
occur in the Sierra Nevada (Adams, 1973). Because these patterns could be involved
in the process of speciation, it is important to understand how they are maintained;
so far there is no persuasive evidence that competition is involved— in Pontia or
otherwise.

ACKNOWLEDGMENTS

Field work was supported by California Agricultural Experiment Station Project CA-D*-
AZO-3994-H, “Climatic Range Limitation of Phytophagous Lepidopterans.” I thank Bill Salt,
Tom Schoener, Cathy Toft and Ken Watt for discussion. And of course I thank Jack Francle-
mont, who taught me some 25 years ago that sound taxonomy is a prerequisite to sound ecology,
and there is no substitute for thoroughness in whatever one does. I have been separating the
protodice from the occidentalis ever since!

LITERATURE CITED

Abbott, W. P., L. S. Dillon and R. R. Shrode. 1960. Geographic variation in Pieris protodice
Boisduval and LeConte. Wasmann J. Biol. 18:103-127.

Adams, M. J. 1973. Ecological zonation and the butterflies of the Sierra Nevada de Santa
Marta, Colombia. J. Nat. Hist. 7:699-718.

Begon, M., J. L. Harper and C. R. Townsend. 1990. Ecology: Individuals, Populations and
Communities. Blackwell Scientific Publications, Boston. 945 pp.

Chang, V. C. S. 1963. Quantitative analysis of certain wing and genitalia characters of Pieris
in western North America. J. Res. Lepid. 2:97-125.

Hovanitz, W. 1962. The distribution of the species of the genus Pieris in North America. J.
Res. Lepid. 1:73-84.

Shapiro, A. M. 1975. Ecological and behavioral aspects of coexistence in six Crucifer-feeding
Pierid butterflies in the central Sierra Nevada. Amer. Midi. Nat. 93:424-433.

Shapiro, A. M. 1976. The biological status of Nearctic taxa in the Pieris protodice-occidentalis
group (Pieridae). J. Lepid. Soc. 30:289-300.

Shapiro, A. M. 1977. Apparent long-distance dispersal by Pieris occidentalis (Pieridae). J.
Lepid. Soc. 31:202-203.

1992

TWENTY YEARS OF PARAPATRY

319

Shapiro, A. M. 1 978. The alpine butterflies of Castle Peak, Nevada County, California. Great
Basin Nat. 37:443-452.

Shapiro, A. M. 1979. Weather and the lability of breeding populations of the checkered white,
Pieris protodice Bdv. & LeC. (Pieridae). J. Res. Lepid. 17:1-23.

Shapiro, A. M. and H. J. Geiger. 1986. Electrophoretic confirmation of the species status of
Pontia protodice and P. occidentalis (Pieridae). J. Res. Lepid. 25:39-47.

Shapiro, A. M., C. A. Palm and K. L. Wcislo. 1981. The ecology and biogeography of the
butterflies of the Trinity Alps and Mount Eddy, northern California. J. Res. Lepid. 18:
69-152.

Received 3 July 1991; accepted 22 November 1991.

J. New York Entomol. Soc. 100(2):320-324, 1992

HYDROPSYCHE FRANCLEMONTI, A NEW SPECIES OF
THE SCALARIS GROUP FROM EASTERN NORTH AMERICA
(TRICHOPTERA: HYDROPSYCHIDAE)

Oliver S. Flint, Jr.

Department of Entomology, Smithsonian Institution, Washington, D.C. 20560

Abstract. — The new hydropsychid caddisfly, Hydropsyche franclemonti, is described, illus-
trated and its distribution plotted (Connecticut, New York, North and South Carolina, Virginia
and Quebec). It is differentiated from its closest relatives in the scalaris group, H. rossi and H.
mississippiensis, and their distributional patterns are discussed.

The genus Hydropsyche, as it has been classically recognized, is found on most
continents of the world except for South America. However, recent works have
established separate genera (or subgenera) for many of the more distinctive groups
(Li and Tian, 1989; Neboiss, 1986; Ross and Unzicker, 1977). This has led to a great
deal of controversy and confusion in the literature (Schefter, Wiggins and Unzicker,
1986; Schuster, 1984) with the outcome still undecided pending a world-wide, full-
scale, revisionary study of all related species groups including their immature stages.

In America north of Mexico the species of Hydropsyche were placed into four
species groups by Ross (1944), a procedure followed by Nimmo (1987) who recog-
nized three groups, with one group subdivided into six subgroups. Ross christened
one of his groups as the scalaris group (“species group one” of Nimmo), and placed
13 species from eastern North America therein (another 5 eastern and 5 western
species in his 1 944 checklist would also be placed in the group). There are no European
species clearly belonging to the group, although the species related to H. pellucidula
are similar but nevertheless different on the basis of the shape of the tenth tergum
and apex of the phallus. Surprisingly, I am unable to find any illustrations of species
from Japan and China that would appear to be of species close to this group. Thus,
at this moment, the scalaris group appears to be restricted to North America with
one or two species in northern Mexico, a half dozen species in the western United
States and Canada, and another several dozen in the eastern United States. The
greatest number of species appears to be found in the Mississippi Valley, the Atlantic
Coastal Plain and Piedmont Regions. The number of species declines rapidly north
of Mason-Dixon Line, with only nine found north of the Canadian border (Nimmo,
1987).

The identification of the species in this group has been difficult, and misidentifi-
cations were (and still are) commonplace until Ross (1944) presented detailed illus-
trations of most of the eastern species. Even in 1979 in our study (Flint, Voshell and
Parker, 1979) of the scalaris group in Virginia it was necessary to borrow types of
most species to verify identifications which then uncovered a major misidentification
by Ross in 1944. More recent studies of problematic material have produced yet
another species which was either not present or misidentified in older material.

I take great pleasure in dedicating this new species to Dr. J. G. Franclemont who

1992

HYDROPSYCHE FRANCLEMONTI

321

5

Figs. 1-5. Hydropsyche franclemonti, male genitalia. 1. Lateral. 2. Clasper, postero ventral.
3. Ninth and tenth terga, dorsal. 4. Apex of phallus, ventral. 5. Same, lateral.

encouraged my studies on the Trichoptera at Cornell. His extensive collecting over
much of the range of this species in eastern North America— North Carolina, Virginia,
New York— while it never turned up this species, did provide many valuable records
and range extensions in the Trichoptera.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Hydropsyche franclemonti, new species
Figs. 1-5

Diagnosis. This species is a member of the Hydropsyche scalaris group (sensu Ross,
1 944). It belongs to that subgroup of species whose apical clasper segment is blunt
in posteroventral aspect, and whose mesal cavity of the phallus is widely open. In
this subgroup are placed Hydropsyche rossi Flint, Voshell & Parker and H. missis-
sippiensis Flint. The tenth tergum of H. rossi in lateral aspect differs in having the
dorsal margin nearly straight with the apicodorsal angle slightly prolonged and es-
pecially in the apex of the phallus. The inner margin of the lateral flange of the mesal
cavity of the phallus in H. rossi is, in ventral aspect, very narrow with that portion
of the lateral flange beyond the mesal cavity also narrow and of uniform width
throughout. The lateral aspect of the tip of the phallus and the apex of the clasper
in posteroventral aspect are almost identical in H. rossi and H. franclemonti. In H.
mississippiensis the tenth tergum and ventral aspect of the tip of the phallus are more
similar to H. franclemonti in appearance. However, the apical segment of the clasper
of H. mississippiensis in posteroventral aspect is bluntly rounded, and the tip of the
phallus in lateral aspect has a high mesal dome with the lateral flange close to the
midline.

Description. Length of forewing, 9.5-10.5 mm. Color brown; forewing mottled,
spotted with various shades of brown, as typical of most species of the genus. Eyes
of male very large; in frontal aspect each eye equal to the interocular distance. Male
genitalia: Ninth segment with anterior margin slightly rounded; with dorsal crest
pronounced, strongly compressed, dorsal margin almost semicircular. Tenth tergum
in lateral aspect with dorsal margin declivous, apical margin almost vertical with
very small apicodorsal bump; in dorsal aspect apex obliquely truncate, with small
apicomesal excision. Clasper with basal segment long, slender, slightly sinuous; apical
segment in lateral aspect with tip pointed, sharply curved dorsad, in posteroventral
aspect with apex obliquely truncate. Phallus tubular, phallobase not constricted before
lateral flanges; apex in lateral aspect with mesal dome slightly elevated, lateral flange
long, with ventral margin ascendant, tip rounded; in ventral aspect with inner margin
of lateral flange broad basally, tapering rapidly apicad, with mesal cavity thus wide
open apicad, lateral flange rounded laterally, much widened apicad.

Type material. Holotype, male: Virginia, Culpepper Co., Hazel River, off Rt. 707,
3 July 1981, B. C. Kondratieff (NMNH). Paratypes: Same data, except 14 July 1979,
1 <3 (NMNH). Connecticut, Litchfield Co., Housatonic Meadows State Park, 21-24
May 1983, Rodrigo Andrade, 4 <3, 2 9 (NMNH, YU). New York, Delaware Co.,
Beaverkill River near Horton, 13 June 1987, P. W. Schefter and R. MacCulloch, 1
<3, 2 9 (ROM). N.C. [North Carolina], Collection Morrison, 1 <3 (NMNH). South
Carolina, Oconee Co., Chattooga River at Burrell’s Ford near Route 107, 26-27 May
1981, Ent. 412/612 class, 1 <3 (CUC). Canada, Prov. Quebec, Station de Biologie de
l’Universite de Montreal, St. Hippolyte, 20 June 1973, P. P. Harper, 1 <3, 4 9 (UMDB);
Lac Monroe, Parc Provincial du Mont Tremblant, 18 July 1959, J. R. Mougeau &
R. Ouellet, 1 <3, 6 9 (UMDB).

Distribution. The distributions of the three related species (//. franclemonti, H.
mississippiensis and //. rossi), although apparently overlaping to a great part, do

Fig. 6. Known localities for Hydropsyche franclemonti (stars); ? placed at Black Mountain,
North Carolina, a known site for much of the North Carolina, Morrison, material.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

appear to differ in detail. Hydropsyche rossi and H. mississippiensis are species of
the Coastal Plan and lower Piedmont regions. The known range of H. mississippiensis
extends from Louisiana along the coast north to Virginia, that of H. rossi is the same
along the eastern coast but extends into Texas along the Gulf and up the Mississippi
Valley into Illinois. The range of H. franclemonti (Fig. 6), however, appears to follow
the eastern margin of the Appalachian Mountains from South Carolina north into
Quebec.

ACKNOWLEDGMENTS

I am indebted to those who supplied the material of this species: W. G. Downs, Yale University
(YU, Connecticut examples), P. W. MacCulioch, Royal Ontario Museum (ROM, New York
examples), S. W. Hamilton, Clemson University (CUC, South Carolina example), P. P. Harper,
Universite de Montreal (UMDB, Quebec examples) and C. R. Parker and B. C. Kondratieff
(Virginia examples).

LITERATURE CITED

Flint, O. S., Jr., J. R. Voshell, Jr. and C. R. Parker. 1979. The Hydropsyche scalaris group in
Virginia, with the description of two new species (Trichoptera: Hydropsychidae). Proc.
Biol. Soc. Wash. 92:837-862.

Li, Youwen and Lixin Tian. 1 989. Notes on a new subgenus and a new species of Hydropsyche.
J. Nanjing Agr. Univ. 12(4):44-45.

Neboiss, A. 1986. Atlas of Trichoptera of the SW Pacific— Australian Region. Series Ento-
moiogica (Dr. W. Junk Publishers) 37, 286 pages.

Nimmo, A. P. 1987. The adult Arctopsychidae and Hydropsychidae (Trichoptera) of Canada
and adjacent United States. Quaest. Ent. 23:1-189.

Ross, H. H. 1944. The Caddis Flies, or Trichoptera, of Illinois. Bull. 111. Nat. Hist. Surv.
23(1): 1—326.

Ross, H. H. and J. D. Unzicker. 1977. The relationships of the genera of American Hydro-
psychinae as indicated by phallic structures (Trichoptera, Hydropsychidae). J. Georgia
Entomol. Soc. 12:298-312.

Schefter, P. W., G. B. Wiggins and J. D. Unzicker. 1986. A proposal for assignment of
Ceratopsyche as a subgenus of Hydropsyche, with new synonyms and a new species
(Trichoptera: Hydropsychidae). J. N. Am. Benthol. Soc. 5:67-84.

Schuster, G. A. 1 984. Hydropsyche! — Symphitopsyche! — Ceratopsyche!: a taxonomic enigma.
Proc. 4th Int. Symp. Trichop. (Series Entomologica 30):339-345.

Received 21 December 1990; accepted 13 March 1991.

J. New York Entomol. Soc. 100(2):325-380, 1992

THE TRIBE LICININI (COLEOPTERA: CARABIDAE): A
REVIEW OF THE GENUS-GROUPS AND OF THE
SPECIES OF SELECTED GENERA

George E. Ball

Department of Entomology, University of Alberta,

Edmonton, Alberta T6G 2E3, CANADA

Abstract. — Evidence for monophyly of the Tribe Licinini is found in synapotypic character
states of mouthparts (mandibles, primarily reduction of occlusal ridges and grooves) and the
head capsule of larvae, with a distinct ventral apotome. Illustrations of mouthparts of adults
of representative licinines show four basic types, which distinguish four subtribes: Dicaelina,
Dicrochilina (new subtribe), Lestignathina (new subtribe) and Licinina (including the South
American genus Eutogeneius). Structure of the mandibles and of the fore coxal cavities indicate
relationships as follows: Dicaelina + ((Dicrochilina + Lestignathina) + Licinina). A review of
the genus Diplocheila Brulle reveals for the subgenus Diplocheila new synonymy: Pterostichus
( Platysma ) karikali Jedlicka, 1969 = Diplocheila distinguenda LaFerte, 1851. A review of the
geographical history of subgenus Isorembus Jeannel indicates that the striatopunctata group
was divided in two by opening of the Atlantic Basin, isolating one monophyletic segment in
the Nearctic Region, and one in the Palaearctic Region, the latter withdrawing subsequently
from Western Europe, and surviving in Western China. Review of recently collected material
of the subgenus Liodicaelus (gen. Dicaelus Bonelli) reveals a new species —D. franclemonti (type
locality— MEXICO, state of Nuevo Leon, Cerro Potosi, a mountain northwest of Galeana). Re-
examination of the type material of D. laevipennis dicaeloides Ball suggests that this taxon is
consubspecific with D. 1. laevipennis LeConte. The names are synonymized accordingly. Recon-
struction of the geographical history of Liodicaelus suggests that this lineage is of Late Miocene
origin (ca. 12 million years before present), and that differentiation has been the result of
vicariance and dispersal, with the oldest barrier between the Sierra Madre Oriental and the
Sierra Madre Occidental. Study of the mouthparts of Badister reflexus LeConte reveals that, in
the West Indies, the mandibles are dimorphic, with either the right or left mandible deeply
notched dorsally. This feature of asymmetry was used to diagnose subgenera: left mandible
notched, Baudia Ragusa; right mandible notched, Badister ( sensu stricto) Clairville. Because it
seems clear, now, that the genetic basis for mandibular difference is simple, the groups based
on this feature alone should be reassessed.

DEDICATION

Jack Franclemont was a superb mentor, guiding one along the way by example
rather than by precept. It was my good fortune to have been his first graduate student,
though it was rumored that on more than one occasion he declared that never again
would he have another coleopterist as a student— one of the few promises that he
did not keep.

Our association at Cornell was not confined to laboratory and office. I learned
under his tutelage that field trips could be civilized and even civilizing experiences,
rather than akin to military assault operations. And, on many Sundays, my wife Kay,
our two sons and I were invited to his home for dinner— a culinary experience that
we valued. But we valued even more Jack’s thoughtfulness in extending the invita-
tions.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Yol. 100(2)

Kay and I continue to enjoy Jack’s company when staying with him in his Snyder
Heights home, during our pilgrimages to Ithaca and Cornell. It has occurred to me
from time to time that one could spend profitably a sabbatical leave, sitting in his
living room, listening to music from his tapes and records, dining at his table, and
reading the rich supply of literature that comes to hand so readily.

Nearly 38 years ago, I completed a thesis about classification of licinine carabid
beetles. I think that Jack eventually became mildly interested in these fine creatures,
though initially he viewed with amused disdain my extolling of their virtues.

On this grand and joyous occasion, as we celebrate Jack’s 80 years of life, I am
pleased to have the opportunity to dedicate this review to him, with the hope that
it will bring back old and hopefully pleasant memories, and that he will find therein
some justification for the time and effort that he devoted to making my first review
of the licinines a presentable scientific document.

INTRODUCTION

Having completed a major treatment of a taxon, systematists tend to seek other
taxa, rather than to review again the group that has already been treated extensively—
even though a good taxonomic work poses nearly as many questions as it answers.
In part, I suppose, after an intensive study of a particular group, one loses a certain
amount of interest in it, and in part, at the time of doing the study, one becomes
aware of the possibilities at hand in other taxa that are readily accessible to analysis.

After publishing a revision of the North American Licinini (Ball, 1959), I proceeded
to investigate other taxa, though two short studies clarified some points that were of
special interest to me (El Moursy and Ball, 1959; Ball, 1966b). The idea for the
present publication came with the need to offer something suitable for the Francle-
mont Festschrift. It began with some observations of the West Indian species of the
licinine genus Badister. With interest in licinines rekindled, I thought about other
matters related to Badister. Then, because of my interest in Middle American Ca-
rabidae, the Mexican species of Dicaelus ( Liodicaelus ) came to mind. Certain inter-
esting observations about licinines that I had made during the past 30 years seemed
worth including. Previous statements that I had published that proved incorrect
required correction. And the mouthparts, especially the mandibles, that seemed of
great potential importance in licinine classification and that had not been studied in
detail, commended themselves for examination. Collectively, these items constitute
a highly eclectic review of licinine Carabidae, their relationships, and geographical
distribution.

MATERIAL AND METHODS

Material

This study is based on detailed examination of about 200 adult licinines, including
representatives of the four subtribes recognized and the genera known from North
America. Most of the material studied is in the Strickland Museum, Department of
Entomology, University of Alberta (UASM). Other collections from which material
was borrowed are listed below, along with names of the curators, and a four-letter
coden used in the text for identification of the lending institution.

1992

THE TRIBE LICININI

327

AMNH Department of Entomology, American Museum of Natural History, Cen-
tral Park West at 79th Street, New York, New York, U.S.A. 10024 (L.
H. Herman);

CASC Department of Entomology, California Academy of Sciences, Golden Gate
Park, San Francisco, California, U.S.A. 94118 (D. H. Kavanaugh);

CMNC Department of Zoology, Canadian Museum of Nature, P.O. Box 3443,
Station D, Ottawa, Ontario KIP 6P4, Canada (R. S. Anderson);

CNCI Canadian National Collection of Insects, Biological Resources Division,
Centre for Land and Biological Resources Research, K. W. Neatby Build-
ing, CEF, Ottawa, Ontario K1A 0C6, Canada (Y. Bousquet);

FSCA Florida State Collection of Arthropods, Division of Plant Industry, Florida
Department of Agriculture, Gainesville, Florida, U.S.A. 32601 (R. E.
Woodruff);

LSUC Louisiana State University Insect Collection, Department of Entomology,
Louisiana State University, Baton Rouge, Louisiana, U.S.A. 70803 (V.
L. Moseley);

MCZC Department of Entomology, Museum of Comparative Zoology, Harvard
University, Cambridge, Massachusetts, U.S.A. 02138 (D. G. Furth);

SEMC Snow Entomological Museum, The University of Kansas, Lawrence, Kan-

sas, U.S.A. 66045-2119 (R. Brooks);

UCBC Division of Entomology and Parasitology, University of California, Berke-
ley, California, U.S.A. 94720 (J. Chemsak);

UCDC The Bohart Museum of Entomology, University of California, Davis, Cal-
ifornia, U.S.A. 95616 (R. O. Schuster);

USNM Department of Entomology, United States National Museum of Natural
History, Smithsonian Institution, Washington, D.C., U.S.A. 20560 (T.
L. Erwin);

ZMUC Department of Entomology, Zoological Museum, University of Copen-
hagen, Universitetsparken, DK-2100, Kobenhavn O. (O. Martin); and

ZSMM Zoologische Staatssammlung; Miinchhausenstrasse 21, D-8000, Miinchen
60, Federal Republic of Germany (G. Scherer).

Methods

Comparative morphology, phylogenetic relationships and classification. — Studies
reported are at three hierarchical levels: subtribal, involving groups of genera; sub-
generic, and intrasubgeneric; and intraspecific. At the subtribal level, because my
principal goals were to assay and describe structural features used previously in
classification and to achieve thereby an improved understanding of licinine rela-
tionships, I used an exemplar method, i.e., one or a few taxa that were readily available
to me, to analyze the characters of a grouping recognized previously. To this study,
I added as well published studies of other character systems, including chromosome
number, defensive secretions, and characteristics of larvae.

Being interested in phylogenetic relationships as a basis for classification, I at-
tempted to establish transformation series, and to position the licinine taxa accord-
ingly. To polarize the transformation series, I used as outgroup the pterostichine
genus Loxandrus LeConte, because it seemed to be in most respects less derived

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than the licinines, and yet to exhibit some features of the licinine ground plan, such
as form of the male fore tarsomeres and having a pair of pits in the ventral surface
of the mentum.

For the subgenus Isorembus Jeannel, I reviewed a flawed previous effort to establish
the phylogenetic relationships of the species groups by using the outgroup method,
with the other two subgenera of the genus Diplocheila Brulle as outgroups.

For the subgenus Liodicaelus Casey, I had the opportunity to undertake a more
detailed study, having at hand representatives of all known taxa. However, the data
available were not sufficient to permit a very convincing evolutionary treatment, and
so 1 confined my activities to improving means of identifying the taxa and to doc-
umenting more fully their geographical ranges. Using essentially the commonality
principle in the absence of a clearly specified outgroup, I recognized one species as
the probable primitive adelphotaxon of the other four, and postulated that the latter
group comprised a monophyletic assemblage.

For the genus Badister, I postulated infraspecific dimorphism in mandibular asym-
metry of B. reflexus LeConte, a character that was regarded by most previous authors
as of generic or subgeneric value. To make such a postulate required a further postulate
about conspecificity. Recognition of the two mandibular morphs as members of the
same species was based on their overall detailed similarity in features (principally
male genitalia) whose variation signals specific differences throughout the genus
Badister.

Biogeographical analysis. — For the subgenus Isorembus Jeannel and Liodicaelus
Casey, I commented about their distribution patterns in an evolutionary context. I
compared the distribution patterns of these taxa to similar patterns of other taxa that
had been analyzed more rigorously, and reasoned by analogy that such similarities
reflected similar histories of dispersal and vicariance.

Measurements. — These were used to indicate size and proportions for specimens
of Liodicaelus. The following measurements were made, using an ocular micrometer
in a Wild S5 Stereobinocular Microscope, at a magnification of 25 x :

length of head— linear distance from base of the left mandible to the posterior margin
of the left eye;

width of head— maximum linear transverse distance across eyes, from outer margin
to outer margin;

length of pronotum— linear distance from anterior (apical) to posterior (basal) margin,
measured along the mid-line;

width of pronotum at apex— linear transverse distance between antero-lateral angles,
measured at right angle to the long axis;

maximum width of pronotum — maximum linear transverse distance across prono-
tum, measured at right angle to long axis;
length of elytra— linear distance from basal ridge to elytral apex;
width of elytra— maximum transverse distance across elytra, measured at right angle
to long axis.

These measurements were added or combined in ratios, and designated as follows:

SBL— Standardized Body Length— sum of length of head, pronotum, and elytra;
EW— width of elytra, and also maximum body width;

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PW/HW— width of pronotum/width of head;

PL/PWA— pronotum: length/width at apex;

EL/EW— elytra: length/width

For SBL, PW/HW, PL/PWA and EL/EW, means were calculated and results pre-
sented in Tables 3-6. Also, a scattergram was prepared (Fig. 1 5), based on PL/PWA
and EL/EW for discrimination among the species of Liodicaelus.

Dissections.— These were made using standard techniques. Genitalia were pre-
served in glycerine, in microvials, pinned beneath the specimens from which these
sclerites had been removed. Larger parts, or those that had been gold-coated for
study with the Scanning Electron Microscope, were glued to cards, and pinned beneath
the specimens from which they had been removed.

Illustrations.— A Cambridge S-250 Scanning Electron Microscope was used to
examine and photograph gold-coated sclerites. Line drawings were prepared by using
a camera lucida attached to a Wild S5 Stereobinocular Microscope.

Terms for structural features

Most of the words used to designate details of structures are found in entomology
textbooks or are used by coleopterists, generally. Other words, required to designate
particular structures or parts thereof, are not in general use. I provide information
about such words here.

Body parts.— The term “segment” is restricted to those body parts that reflect
embryonic somites. Thus, somite-like portions of the abdomen are referred to as
segments. Abdominal segments are designated by a Roman numeral that corresponds
to the respective somites. The first complete sternum is III, and the last one that is
completely sclerotized and normally exposed is VII.

Portions of limbs are designated by the suffix “-mere,” the prefix depending upon
the limb in question— antenno, palpo, tarso, etc.

Labium.— The word “ligula” is used for glossae + paraglossae. In turn, the fused,
sclerotized glossae characteristic of carabids are termed the glossal sclerite.

Elytra. — Longitudinal grooves on the dorsal surface, or remnants thereof (“striae”
of most authors) are designated as intemeurs (Erwin, 1974:3-5).

Ovipositor.— The terminal parts are referred to as stylomeres. The thick marginal
setae on stylomere 2 are ensiform setae.

Terms for phylogenetic aspects

These are standard, except for “adelphotaxon.” This word was introduced by Ax
(1987:36) as a substitute for sister taxon. The advantage of using adelphotaxon is
that, being of classical origin, it will be the same in all European languages and thus
does not require translation.

RELATIONSHIPS OF THE LICININI

Jeannel (1941) included in the Callistomorphi the families Callistidae (including
the oodines), Licinidae, and Panagaeidae. Kryzhanovskij (1983:88-89), using a more
traditional system of ranking, included the tribes Licinini, Callistini, and Oodini in
the supertribe Callistitae, placing the Panagaeini in the supertribe Panagaeitae, along

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with the Peleciini. Erwin (1985:468) separated also the panagaeines from the Callis-
titae, including in the Panagaeitae as well the Afrotropical Bascanini and the Peleciini
( sensu latiore). Considering the shared derived features of callistites ( sensu Kryzha-
novskij and Erwin) and at least panagaeines, exhibited by adults (including para-
median mental pits, and defensive secretions [see below]) and by larvae, it seems
best to include the two complexes in a single supertribe. Straneo and Ball (1989:85)
placed the peleciines as incertae sedis, near the base of the subfamily Harpalinae
{sensu Erwin, 1985:468).

Within the callistite assemblage, the tribe Licinini probably occupies a basal po-
sition—at least such is suggested by the distinctive pattern of volatile organic com-
pounds comprising the defensive secretions of the pygidial glands. A basal position
for the Licinini is indicated also by features of the fore coxal cavities— probably
plesiotypically uniperforate, whereas oodines, callistines and panagaeines exhibit the
probably apotypic feature of biperforate coxal cavities. I postulate the following
system of relationships: Licinini + (Oodini + (Callistini + Panagaeini)). See below
for details.

GENERIC CLASSIFICATION

Important recent references to Licinini are: Darlington (1968:14-19), treating the
genera and species of New Guinea; Lindroth (1968:939-944 and 1969:945-967 and
1121), treating the Canadian and Alaskan taxa; Kryzhanovskij (1983:274-276), treat-
ing the fauna of the USSR; Freude (1976:254-260), treating the fauna of Middle
Europe; and Moore et al. (1987:263-269), cataloguing the Australian taxa.

Freude (1976), evidently following Jeannel (1942), placed Licinus and Badister in
the subfamilies Licinitae and Badisteritae, respectively, thus separating these two
genera as widely as either is separated from other carabid subfamilies, such as Ptero-
stichitae and Harpalitae. Such ranking does not coincide with the affinities of these
genera, as explained by Ball (1959), and amplified below.

Ball (1959:7) arranged the licinine genera in four groups which he regarded as the
equivalent of subtribes. Two were named: Dicaelina (Group 1) and Licinina (Group
2), leaving Groups 3 and 4 unnamed. They are named here: Lestignathina (name
based on that of the first described genus included in this group— Lest i gnat hus Erichson
[1842:132]), new subtribe (Group 3); and Dicrochilina, new subtribe (Group 4).

Two genera, the northeastern Australian Mecynognathus MacLeay and Chilean
Eutogeneius Sober (1849:253), were excluded from the classification because Ball
had not seen specimens. Jeannel (1942:991) placed the enigmatic Mecynognathus in
the Licinini because of the strikingly modified mouthparts of the adults. Darlington
(1961:18) included Mecynognathus in the Tribe Pterostichini, as did Moore (1965:
22), based on overall similarity and chorological affinity to the clearly pterostichine
genus Paranurus Tschitscherine.

Although I have not studied material of Eutogeneius fuscus Sober (1849:255) in
great detail, I have seen the holotype in the Oberthiir-Chaudoir Collection (Mus.
Nat. Hist. Nat., Paris), and two additional specimens in the Canadian National
Collection of Insects, Ottawa. A male exhibits adhesive vestiture of articulo-setae
on the front tarsus (Stork, 1980:287), and the mandibles are short and broad, with
broad apices. On the basis of these observations, Eutogeneius is assigned to the
subtribe Licinina.

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Ball (1959:6-8) used features of the adult mandibles as the primary basis for
classifying the licinine genera, but he provided only a superficial account of these
structures. A more detailed description of the licinine mandibles is presented below,
based on Acorn and Ball (1991). Study of the mouthparts was extended to include
the labrum, maxillae, and labium. The adhesive vestiture of the anterior tarsi of
males was examined in some detail. Also, a few specimens were examined for the
condition of the fore coxal cavities. As noted in the Methods section above, the
pterostichine genus Loxandrus was used as an out-group.

Based on published information, additional characters for classification were sought
in the defensive secretions of adults, in structural features of the larvae, and in
chromosome number.

Mandibles

As explained by Acorn and Ball (1991) carabid mandibles are trigonal in form,
each tapered from a broad base to a narrow, more or less pointed apex (Fig. 1 A-B).
Laterally broad and with a triangular depression in the outer face (the scrobe), the
mandible is sloped toward the occlusal surface (Fig. 1C-D), or the edge that opposes
the other mandible, and is involved in biting or gripping the food items. In dorsal
aspect, two imprecisely defined areas are recognized: a terebra, extending from the
apical incisor tooth (Fig. 1A, i) to a transverse plane, extended through the terebral
tooth (tt). Posterad the terebra is the broader basal area. A series of ridges and teeth
are on the occlusal margin, as follows, anteriorly to posteriorly, and dorsally to
ventrally (Fig. 1A-E): incisor tooth; terebral ridge (tr), or if two, superior terebral
(str) and inferior terebral (itr); terebral tooth; retinaculum (r), in most groups with
teeth (anterior, art, and posterior, prt), and a ridge (rr). The end of the retinaculum
is marked in most taxa by a deep groove, the anterior occlusal groove (aog) posterior
to which is the molar area, divided or not by a posterior occlusal groove into a more
anterior premolar ridge and/or tooth, and a posterior molar ridge and tooth. These
distinctions are not required here, and the molar area is designated simply m. Ven-
trally, a groove extends diagonally or more or less parallel to the occlusal margin —
the ventral groove (Figs. 1E-F, vg). Arising from the ventral groove is a brush of
microtrichia (vmt).

Mandibles of Loxandrus LeConte (Fig. 1 A-F). — The terebra occupies about % of
the mandible, as is true of most carabid mandibles (Acorn and Ball, 1991). On the
left mandible (Fig. 1 A, C, E), the superior (str) and inferior (itr) terebral ridges meet
at the terebral tooth (tt). The tooth and ridges comprise the occlusal portion of the
terebra. Posterad the terebral tooth, the occlusal surface is occupied by the retinac-
ulum, or retinacular ridge (rr) which extends ventrad the terebral ridge. The anterior
occlusal groove (aog) at the posterior margin of the retinaculum, is marked distinctly.
Posterad the groove is the molar area (m), which in Loxandrus is ridged (Fig. 1C),
and extends anteriorly as a ridge (Fig. IF) beneath the retinaculum. The ventral
groove (Fig. IE, vg) is moderately long and its brush is comprised of rather short
microtrichia (vmt). The right mandible (Fig. IB, D, F) is similar to the left except
that the retinacular ridge (rr) is more prominent and terminates anteriorly in a
pronounced anterior retinacular tooth (art), referred to by Bell (1960:100) as the
canine tooth.

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Mandibles of representative species of Ch/aenius are described and illustrated by
Bell (1960:163, figs. 122-131). Spence (1982:571, fig. 5A) illustrated mandibles of
an oodine. Both groups are characterized by long terebrae, a complete set of teeth
and ridges, and occlusal grooves.

By contrast and in general, mandibles of licinine adults exhibit more extensive
ventral grooves (Fig. 1K-L, 2E-F, K-L, and 3E-F, K-L), and less complex occlusal
margins that lack an occlusal groove separating the molar area from the retinaculum
(Figs. 1G-L, 2A-L, 3A-L). The occlusal margin posterad the terebral tooth is named
here basal ridge (br), because it is unclear (except in D. daldorfi, Fig. 1G-L) how
much of this margin is retinacular and how much is molar.

Mandibles of subtribe Dicaelina. — These are illustrated by specimens of D. daldorfi
Crotch (Fig. 1G-L) and D. polita Dejean. See also the mandibles of Dicaelus ( Lio -
dicaelus) (Figs. 13A-L, 14A-F).

The terebral ridge is nearly straight, with the incisor quite broad (Figs. 1G-H, 2A-
B). In occlusal aspect (Figs. 1 1— J, 2C-D), the terebral surfaces are quite broad and
sloped diagonally ventro-laterally. In contrast to Loxandrus, dicaeline mandibles are
more slender basally. The ventral groove of Diplocheila mandibles bears brushes of
very long microtrichia (Figs. 1 K-L, 2E-F), but the microtrichia are shorter in Dicaelus
(Figs. 13E-F, K-L, and 14E-F). The dicaeline terebra is about half the length of the
mandible.

The terebrae of D. daldorfi (Fig. 1G-H) are about as broad as those of D. polita
(Fig. 2A-B), and the occlusal surface on the right mandible is smooth (Fig. 1 J) rather
than diagonally ridged, as in D. polita (Fig. 2D). The left mandible of D. daldorfi
exhibits a small supraterebral tooth (Fig. 1G, stt), a unique feature of this species.
(Unfortunately, the dorsal surface of the specimen illustrated is worn somewhat, and
irregular.) The retinaculum is small (Fig. 1G, I). The right mandible of D. daldorfi
(Fig. 1H, J) lacks a clearly marked retinaculum.

The left terebral tooth of D. polita (Fig. 2A, E) is more prominent, and the left
mandible has a small retinacular tooth (Fig. 2 A, art) and a rather indistinct retinacular
ridge (Fig. 2E).

The left terebral tooth of D. polita (Fig. 2A, E) is more prominent; the retinacular
tooth is small (Fig. 2A, prt), and the retinacular ridge is rather indistinct. The right
mandible (Fig. 2B, D) is without a clearly marked retinaculum.

Mandibles of subtribe Dicrochilina (Fig. 2G-L). — Terebrae are short, not more
than one-third the length of the mandibles (Fig. 2G-H), and the basal part is relatively
slender (Fig. 2I-J). The terebral teeth (tt) are small, the right one isolated from the
superior terebral ridge (str) by a notch. The left mandible exhibits two terebral ridges
(Fig. 2G, str and itr). The retinacula are absent. The ventral grooves bear long and
densely packed microtrichia (Fig. 2K-L).

Mandibles of subtribe Lestignathina (Fig. 3 A-E). — Markedly flattened dorso-ven-
trally, especially the terebrae (Fig. 3C-D), dorsally these mandibles exhibit broad
terebrae, very slender and sharp incisors, and very prominent and sharp terebral
teeth (Fig. 3 A-B, i and tt). The terebral area is less than half the length of the mandible.
Ventral retinacular or molar ridges are not indicated. The microtrichia of the ventral
grooves are short (Fig. 3E-F, vmt).

Mandibles of subtribe Licinina (Fig. 3G-L).— These sclerites are very broad, es-
pecially basally (Fig. 3G-H, K-L), and thick (Fig. 3I-J). The terebrae are narrow in

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the horizontal plane, comparatively short, with occlusal margins nearly straight, and
terebral teeth thick and prominent, and incisors blunt (Fig. 3I-J). The basal areas of
both mandibles are markedly thickened, especially that of the right mandible, which
has also a dorsal boss (Fig. 3I-J, b). The thickened terebral and basal areas of the
right mandible are separated from one another by a low area, referred to as a notch
(Fig. 3J, n). The left mandible does not exhibit such a notch. Such asymmetry between
mandibles is a common feature of the Licinina, some taxa with the right mandible
notched, others with the left, and still others with either mandible notched (see below,
under Badister reflexus LeConte). The ventral grooves have short microtrichia (Fig.
3K-L, vmt).

The mandibles of the Licinina are without retinacular and molar ridges.

Mandibles of Badister are illustrated in Figures 1 9 A-H and 20 A-F. The terebral
ridges of both mandibles are short, and are deflected ventrally, causing the mandibular
apices to appear very broad in the anterior aspect (Fig. 20A-B).

Labrum (Figs. 4A-I, and 5A-D)

Licinines exhibit variously modified labra, most with a more or less pronounced
medial notch. For comparison, illustrations of a more generalized labrum are pro-
vided {Loxandrus pactinullus Allen, Fig. 4A-B).

Labrum of Loxandrus (Fig. 4A-B).— This sclerite is about rectangular, with the
anterior margin slightly concave (Fig. 4A), or straight. Dorsally, six long setae form
a transverse row near the anterior margin. Ventrally, the labrum bears an epipharynx
comprised in carabids of trichoid sensilla (Fig. 4B, ets) flanked each side by a group
of microtrichia (Fig. 4F-G, emt). Liebherr (199 la: 10-1 1, and Figs. 2, 8, 14, 20)
showed a similar pattern for platynines, and that number of sensilla is correlated
positively with body size.

Labrum of subtribe Dicaelina (Fig. 4C-G).— The anterior margin is more or less
deeply notched, and in outline the labrum varies from transverse (Fig. 4E) to about
quadrate (Fig. 4C). Dorsal setae vary in number from four (Fig. 4C, E) to six. Ven-
trally, the trichoid sensilla of the epipharynx are long and submarginal, the central
part of the labrum being absent. The patches of microtrichia are more (Fig. 4F) or
less (Fig. 4D) extensive.

The labra of subgenus Isorembus and of genus Dicaelus are much less modified,
with the anterior margin less deeply and broadly notched, and trichoid sensilla of
the epipharynx shorter.

Labrum of subtribe Dicrochilina (Fig. 5A-B). — In outline, the labrum is longer
than wide, with anterior margin broadly concave. Dorsally are six marginal setae.
Ventrally, the epipharyngeal trichoid sensilla are of average length and number. The
microtrichia are numerous, and form a central “Y” figure, with the stem prominent
(Fig. 5B, etr).

Labrum of subtribe Lestignathina (Fig. 4H-I). — For the Licinini, the labrum is
markedly transverse and short. The central portion is present, and in fact is projected
as a small tooth. The dorsal marginal setae number four. Ventrally (Fig. 41), the
trichoid sensilla are few, and the microtrichia are grouped in two small basal patches,
one each side of the mid-line.

Although typical of Siagonyx, a labrum of this type is atypical for the Lestignathina

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as a whole, with adults of the other genera exhibiting a normal licinine labrum,
quadrate in outline, with the central area absent.

Labrum of subtribe Licinina (Fig. 5C-D).— The form varies appreciably from
asymmetric with an irregular anterior margin to symmetric, with a deep central notch
(Fig. 20A-B). Dorsally, six setae comprise the marginal row (Fig. 5C). Ventrally, the
epipharyngeal trichoid sensilla are few, and microtrichial patches are absent (Fig.
5D).

Maxillae (Fig. 6A-II)

In form and sclerites present, licinine maxillae are about average for adult carabids.
Palpomeres vary in proportions. Vestiture of the lacinia and development of the
lacinial tooth seem to offer the most useful features for classification.

Maxillae of subtribe Dicaelina (Fig. 6A-B).— The lacinial tooth (It) is curved slight-
ly. In dorsal aspect, the occlusal margin of the lacinia has a dense array of slender
setae which extends on to the dorsal surface near the apex (Fig. 6A). Ventrally, the
occlusal margin has several rows of stout setae which extend on to the ventral surface
basally (Fig. 6B). Palpomere 3 (Fig. 6A, mp-3) is subequal in length to palpomere 2
and is distinctly longer than palpomere 4.

Maxillae of subtribe Dicrochilina (Fig. 6C-D). — These are similar to the maxillae
of the Dicaelina (see above), except that the lacinial tooth is curved more sharply in
relation to the long axis of the lacinia.

Maxillae of subtribe Lestignathina (Fig. 6E-F).— The lacinia exhibits a sinuate
lateral margin, is widened apically, and the apical tooth is sharply bent, in fact slightly
recurved at apex (Fig. 6E, It). In dorsal aspect, the slender setae are relatively few
and quite short. Ventrally, the stout setae are rather few also (Fig. 6F). (Sparse setation
of the lacinia is exhibited also by oodines of the genera Anatrichis LeConte and
Oodinus Motschulsky ([Spence, 1982:572, fig. 7A, B]). Palpomere 3 (Fig. 6E, mp-3)
is much shorter than either 2 or 4.

Maxillae of subtribe Licinina (Fig. 6G-H).— The lacinial tooth (Fig. 6G, It) is very
prominent and is bent at a right angle to the long axis of the lacinia. Slender setae
are in several not particularly dense rows, and some of them are curved apically. A
few setae extend on to the dorsal surface preapically. Ventrally, the stout setae are
prominent and in several rows along the occlusal margin. The basal area has few
setae (Fig. 6H).

Labium (Figs. 7A-E and 8A-D)

The licinine labium is characterized by absence of a mental tooth, and by a bisetose
glossal sclerite. Other features vary from subtribe to subtribe, as noted below. The
bisetose submentum is taxonomically featureless.

Labium of subtribe Dicaelina (Fig. 7 A-C). — Separated from the submentum by a
complete suture, the mentum is transverse, with broad lateral lobes (1 1), and epilobes
(el) with apices terminated at the apices of the lateral lobes. Ventrally, the mentum
has a moderately deep median sinus, and bears a pair of paramedial setae, and near
the base a pair of pits (Fig. 7A, mp) with small pores in the surface (Fig. 7C). Allen
and Ball (1980:507, fig. 25B) illustrated similar structures for taxa of Loxandrus.
Mental pits are common in Chlaenius (Bell, 1960:100, and 163, figs. 132-135), in

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Panagaeini and in the Oodini (Spence, 1982:572, fig. 8A). Bell (1960:100) stated that
licinines were without these pits. Their function is unknown, but it seems more likely
that they are sense organs rather than . . simply the outward manifestations of
inwardly-directed apodemes” (Allen and Ball, 1980:488). The paraglossae (pg) of the
prementum are narrow lobes extended beyond the apex of the glossal sclerite (gl).
Dorsally, the paraglossae are covered with an extensive vestiture of microtrichia (Fig.
7B), as is true also for the oodine genus Anatrichis (Spence, 1982:572, fig. 9A).
Palpomere 2 is bisetose.

Labium of subtribe Dicrochilina (Fig. 7D-E). — In general, this complex of sclerites
is much like that of the Dicaelina, differing as follows. Lateral lobes (Fig. 7D, 11) are
longer, and the apices of the epilobes (el) extend clearly beyond the apices of the
lateral lobes. Palpomere 2 is quadrisetose.

Labium of subtribe Lestignathina (Fig. 8A-B).— The mentum is asetose and lacks
pits (Fig. 8A). The medial sinus is shallow, so that the medial part is quite broad.
The apices of the epilobes (el) are coextensive with the apices of the lateral lobes (11).
The paraglossae (pg) are small, and are not extended beyond the apex of the glossal
sclerite (gl), which is convex and is projected anteriorly. Dorsally, the surface of the
ligula (paraglossae + glossae) is covered with isodiametric microsculpture, the sculp-
ticells with surfaces convex (Fig. 8B). Each paraglossa has a row of small microtrichia.
Palpomere 2 is bisetose.

Mentum and submentum are fused, i.e., the mental suture is absent or incomplete.
Horn (1881:143) noted that the mental suture is absent from the specimen of Les-
tignathus that he examined. Sloane (1898:488) pointed out that this suture is absent
also from the labia of Lacordairia, Siagonyx, Hormacrus, Platylytron, and Micro-
feronia. Further, he countered Horn’s argument that Lestignathus is a platynine rather
than a licinine.

Labium of subtribe Licinina (Fig. 8C-D).— The mentum is bisetose (punctures
evident near posterior margin of the deep medial sinus, but setae broken), and the
ventral surface is without pits (Fig. 8C). Lateral lobes (11) are average, with apices of
epilobes (el) coextensive with apices of lateral lobes. The anterior margin of the
glossal sclerite (gl) is slightly convex and slightly projected. For Licinus, the paraglos-
sae (pg) are short and broad and not extended beyond the apex of the glossal sclerite.
Dorsally, each paraglossa has a covering of long setae apically (Fig. 8D, pts) and a
vestiture of microtrichia basally. In contrast, for Badister, the paraglossae are sub-
stantially longer than the glossal sclerite, and the dorsal surfaces of the paraglossae
have a covering of apically broad, flat microtrichia, and marginally, hair-like micro-
trichia. For both of these genera, palpomere 2 is bisetose.

Fore coxal cavities

According to Jeannel (1941:78-82), one of the features diagnostic of the callisto-
morph assemblage is biperforate fore coxal cavities. This is correct for the Callistini,
Oodini, and Panagaeini. However, condition of the fore coxal cavities varies within
the Licinini: Dicaelina and Licinina have uniperforate cavities (i.e., a sclerotized
bridge dividing each cavity is lacking), but Dicrochilina and Lestignathina have
biperforate cavities.

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Fig. 1 A-L. SEM photographs of mandibles of Loxandrus pactinullus Allen and Diplochei'la
daldorfi Crotch. A-F, L. pactinullus : A, C and E, left mandible— dorsal, occlusal and ventral
aspects, respectively; B, D, and F, right mandible— dorsal, occlusal and ventral aspects, re-
spectively. G-L, D. daldorfi Crotch: G, I and K, left mandible— dorsal, occlusal and ventral
aspects, respectively; H, J and L, right mandible— dorsal, occlusal and ventral aspects, respec-
tively. Legend: aog, anterior occlusal groove; art, anterior retinacular tooth; bb, basal brush;
br, basal ridge; i, incisor; itr, inferior terebral ridge; m, molar tooth; prt, posterior retinacular
tooth; r, retinaculum; rr. retinacular ridge; str, supraterebral ridge; stt, superior terebral tooth;
tt, terebral tooth; vg, ventral groove; vmt, ventral microtrichia. Scale bars: A-F = 200 jum; G-
I = 400 pm.

Male fore tarsi and adhesive vestitisre (Fig. 9A-I)

A diagnostic feature of the supertribe to which the Licinini belong (Callistomorphi
Jeannel, 1941; Kryzhanovskij, 1983; or Patellimanes of 19th Century authors) is size
of male fore tarsomeres and their adhesive vestiture. The group is characterized by

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Table 1 . Defensive compounds secreted by the pygidial glands of adult Licinini, and released
by spraying (data from Moore, 1979:195, table 1).

Compounds

Subtribe

Formic acid

Hydrocarbons

Aliphatic ketones

Dicaelina1

X

Dicrochilina

X

X

X

Lestignathina2

X

X

Licinina3

X

X

1 Dicaelus, only.

2 Siagonyx, only.

3 Licinus and Badister, only.

having at least some of the tarsomeres widened asymmetrically, and with a dense
vestiture of adhesive setae, designated articulo-setae by Stork (1980:287). Within the
Licinini, size of male fore tarsomeres and extent of vestiture vary appreciably.

Male fore tarsomeres of subtribe Dicaelina (Fig. 9 A-C).— Tarsomeres 1-3 slightly
expanded on medial side, with typical callistomorph adhesive vestiture ventrally
(Fig. 9B-C).

Male fore tarsomeres of subtribe Dicrochilina (Fig. 9 D-F).— Tarsomeres 1-3 ex-
panded diagonally medially (Fig. 9D), rather like loxandrine tarsomeres (Allen and
Ball, 1980:51 1, fig. 43A-B). Ventral adhesive vestiture of articulo-setae (Fig. 9E-F),
arranged in four diagonal parallel rows on tarsomeres 1 and 2, only two rows on
tarsomere 3.

Male fore tarsomeres of subtribe Lestignathina. — In form and vestiture, these are
like the tarsomeres of the Dicaelina.

Male fore tarsomeres of subtribe Licinina (Fig. 9G-I).— Tarsomeres 1 and 2 (Fig.
9G) expanded markedly, ventral adhesive vestiture of articulo-setae (Fig. 9H-I) on
tarsomeres 1 and 2 (or on 1-3), densely packed.

Defensive secretions of the Licinini

Moore (1979) compared the defensive secretions of the pygidial glands of a broad
taxonomic representation of carabid genera. He reported that callistomorphs produce
a wide range of volatile organic compounds: hydrocarbons, aliphatic ketones, satu-
rated esters, formic acid, phenols, aromatic aldehydes, and quinones. From an evo-
lutionary perspective, production of a mixture of hydrocarbons and formic acid
probably is plesiotypic for callistomorphs, this feature being widespread in the sub-
family Harpalinae, which includes the Callistitae. Within the latter group, however,
only the licinines exhibit this feature: oodines secrete unsaturated acids, only; cal-
listines, hydrocarbons, phenols, and quinone; and panagaeines, hydrocarbons, ali-
phatic ketones and phenols. Presumably, absence from the defensive repertoire of
hydrocarbons and/or formic acid is the result of loss, and the production of other
volatile organic compounds is a gain, both being apotypic. Thus, licinines are rela-
tively primitive compared to the other tribes with respect to production of defensive
secretions.

The pattern within the Licinini is shown in Table 1. Members of all subtribes

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Fig. 2 A-L. SEM photographs of mandibles of Diplocheila polita Dejean and Dicrochile goryi
Boisduval. A-F, Diplocheila polita : A, C and E, left mandible— dorsal, occlusal and ventral
aspects, respectively; B, D and F, right mandible— dorsal, occlusal and ventral aspects, respec-
tively. G-L, Dicrochile goryi : G, I and K, left mandible— dorsal, occlusal and ventral aspects,
respectively; H, J and L, right mandible— dorsal, occlusal and ventral aspects, respectively.
Legend: br, basal ridge; i, incisor; itr, inferior terebral ridge; prt, posterior retinacular tooth; rr,
retinacular ridge; str, superior terebral ridge; tr, terebral ridge; tt, terebral tooth; vg, ventral
groove; and vmt, ventral microtrichia. Scale bars = 200 yum.

produce formic acid, and all but the Dicaelina produce hydrocarbons. Only the
Dicrochilina produce aliphatic ketones. Absence of hydrocarbons and production of
aliphatic ketones are regarded as apotypic features.

Characteristics of larvae

Within the Carabidae, licinine larvae are distinguishable by three features: head
dorsally with frontal sutures extended posteriorly to the occipital suture; cervical

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Fig. 3A-L. SEM photographs of mandibles of Siagonyx species? and Licinus punctatulus
Fabricius. A-F, Siagonyx sp?: A, C and E, left mandible— dorsal, occlusal and ventral aspects,
respectively; B, D and F, right mandible— dorsal, occlusal and ventral aspects, respectively. G-
L, Licinus punctatulus : G, I and K, left mandible— dorsal, occlusal and ventral aspects, re-
spectively; H, J and L, right mandible— dorsal, occlusal and ventral aspects, respectively. Leg-
end: b, dorsal boss; bb, basal brush; br, basal ridge; bt, basal tooth; i, incisor; n, notch in dorsal
surface; tr, terebral ridge; tt, terebral tooth; vg, ventral groove; vmt, ventral microtrichia. Scale
bars = 200 pm.

suture forked, ventral apotome gula-like (Jeannel, 1942:989; Moore, 1966:160); an-
tennae elongate, at least 1.5 x as long as mandibles (van Emden, 1942:44-45). Most
known taxa have the occlusal margin of the terebra and posterior margin of the
retinaculum denticulate or serrate (van Emden, 1942:44-45).

Although the tribe is diagnosed clearly by structural features of larvae, and although
the few genera for which larval features are known (Dicaelus\ Diplocheilcr, Dicrochile;
Siagonyx ; Dilonchus\ Licinus\ and Badister) are readily distinguished from one an-

Fig. 4A-I. SEM photographs of the labrum of: Loxandrus pactinullus Allen (A-B); Diplo-
cheila daldorfi Crotch (C-D); D. polita Dejean (E-G); and Siagonyx species? (H-I). A, C, E and
H, dorsal aspects; B, D, F, G, and I, ventral aspects. Legend: emt, epipharyngeal microtrichia;
ets, epipharyngeal trichoid sensilla. Scale bars = 200 pm.

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Fig. 5A-D. SEM photographs of the labrum of: Dicrochile goryi Boisduval (A-B); and
Licinus punctatulus Fabricius (C-D). A and C, dorsal aspects; B and D, ventral aspects. Legend:
emt, epipharyngeal microtrichia; ets, epipharyngeal trichoid sensilla. Scale bars = 200 gm.

other, some groupings provided by these character states do not coincide with the
subtribal limits based on adult features. For example, Jeannel (1942:990) grouped
Licinus with Dicaelus in the Licinitae, and placed Badister by itself, in the Badisteritae.
The smooth mandibles characteristic of Diplocheila larvae isolate this genus from
all of the others, the latter exhibiting mandibles with serrate-edged terebrae and
retinacula. In contrast, Moore (1966:160) noted that the larvae of Lestignathus and
Dicrochile differed markedly from one another, and that such difference was in agree-
ment with placing them in different groups of genera.

Chromosome numbers of Licinini

Serrano (1986:224) postulated 37 as a basic diploid chromosome number for
Licinini and for “modem” groups of Carabidae generally (i.e., Harpalinae sensu
Erwin, 1985). Numbers are as follows: for subtribe Dicaelina, represented by Dip-
locheila (three species), 37-42; and for subtribe Licinina, represented by Badister
anomalus Perris, 45, and by Licinus (two species), 26 and 38 (Serrano and Yadav,

Fig. 6A-H. SEM photographs of the left maxilla of: Diplocheila daldorfi Crotch (A-B); Di-
crochile goryi Boisduval (C-D); Siagonyx species (E-F); and Licinus punctatulus Fabricius (G-
H). A, C, E and G, dorsal aspects; B, D, F and H, ventral aspects. Legend: 1, lacinia; It, lacinial
tooth; and mp-3, maxillary palpomere 3. Scale bars = 400 n m.

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Fig. 7A-E. SEM photographs of the labium (mentum and prementum) of: Diplocheila polita
Dejean (A-C); and Dicrochile goryi Boisduval (D-E). A, C and D, ventral aspects; B and E,
dorsal aspects; C, enlargement of mental pit, showing pores. Legend: el, epilobe; gl, glossal
sclerite; 11, lateral lobe; mp, mental pit; pg, paraglossa; and pmt, paraglossal microtrichia. Scale
bars: A-B, D-E = 200 jim; C = 10 gm.

1984:351, table 1; Serrano et al., 1986:134). These numbers indicate marked overlap
at the subtribal and generic levels, and thus, they are of little use in suprageneric
classification, except to confirm further the harpaloid affinities of the Licinini. Within
genera, at the species level, chromosome number likely is useful. Serrano (1986:325)
notes that, although similar in form and other structural features of adults, Licinus
aequatus Serville (2N = 26) and L. punctatulus Fabricius (2N = 38) differ markedly

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Fig. 8 A-D. SEM photographs of the labium (mentum and prementum) of: Siagonyx species?
(A-B); and Licinus punctatulus Fabricius (C-D). A and C, ventral aspects; B and D, dorsal
aspects. Legend: el, epilobe; gl, glossal sclerite; 11, lateral lobe; pg, paraglossa; and pts, paraglossal
trichoid sensilla. Scale bars = 200 jum.

in chromosome number, indicating that karyotypic change has been more rapid than
morphological change.

Food of Licinini

Lindroth ( 1 986:403) reported that the larvae and probably also the adults of Licinus
depressus Paykull ate snails. Harde (1984:106) reported that members of Licinus
hoffmannseggi Panzer and unspecified species of Badister were molluscivores. Ball
(1959:53) and Lindroth (1 969:948) cited records of Dicaelus purpuratus Bonelli eating
snails. I observed specimens of Diplocheila oregona Hatch in captivity eating snails.
To obtain the soft tissue, the beetles bit through the mollusc shells.

Mandibles of the lestignathines, with their long, slender incisors and terebral teeth,
appear rather like those of cychrines (cf. Ball, 1966a:723, fig. 2A-B), which are known
to prey on snails. However, rather than biting through the shell, a cychrine attacks
the soft tissue by inserting its narrow head into the operculum. Perhaps lestignathines
behave in the same manner.

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Based on these few observations, I postulate that molluscivory is a general feature
of licinines, and that such behavior is ancestral in this lineage.

Monophyly of the tribe Licinini

Efforts have not been made previously to establish that the tribe Licinini is mono-
phyletic. Ball (1959), for example, accepted implicitly the judgment of previous
workers that the Licinini is a natural group, based primarily on modifications of the
mouthparts, and principally the labrum and clypeus, with anterior margins of each
more or less markedly excavated. As noted above, these features vary markedly
within the Licinini. Other features of adults are likely not apotypic at the tribal level
(number of supraorbital setae; adhesive vestiture of the fore tarsi; form of male
genitalia) or are shared with other callistites (condition of the fore coxal cavities;
mental pits). Larval features, such as long cerci (or urogomphi) and long antennae
are shared variously with other callistites.

A few features of adults, reasonably interpreted as apotypic, characterize the Li-
cinini and help to establish monophyly of the group: reduction of the length of the
mandibular terebra relative to the basal area, absence of occlusal grooves, and an
extensive ventral groove; and the labial mentum without a tooth and with the median
sinus more extensive than usual (except in some lestignathines). Larval features that
seem likely to be apotypic for the Licinini are: dorsal surface of the head with frontal
sutures extended posteriorly to the occipital suture; and ventrally, with the cervical
suture branched, isolating a median apotome. Additionally, I infer that molluscivory
is a ground-plan feature, and is probably the driving selective force involved in
modifications of the mandibles.

Relationships of subtribes of Licinini

Within the Licinini, details of the mouthparts, especially the mandibles, seem to
establish the monophyly of the Dicrochilina, Lestignathina, and Licinina. Addition-
ally, within the Licinini, only the Dicrochilina have aliphatic ketones in their defen-
sive secretions.

Evidence for monophyly of the Dicaelina, however, is less convincing. The larva
of Diplocheila, for example, has a smooth, rather than serrate, terebral margin. This
latter feature is characteristic of Dicaelus and the other groups of Licinini. The
Dicaelina exhibit the most primitive mandibles (terebra relatively long, and various
taxa with indications of remains of the retinaculum). Adults of Dicaelus lack hydro-
carbons from their defensive secretions, producing formic acid only. Defensive se-
cretions of Diplocheila have not been studied. If it is found that members of this
genus also lack hydrocarbons, then this feature will serve as diagnostic for the Di-
caelina.

At present then, the monophyly of the subtribe Dicaelina has not been established:
either Diplocheila or Dicaelus may be more closely related to the derived subtribes
than to one another. For example, the terebral margin and retinaculum of the man-
dibles of larvae of Dicaelus are serrate or denticulate, like those of known members
of the other subtribes. In contrast, the larval mandibles of Diplocheila are smooth.
If, as seems likely, the serrate mandibles are derived, then this feature could indicate
that Dicaelus is related more closely to the other subtribes than to Diplocheila.

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Fig. 9A-I. SEM photographs of left fore tarsus of males of: Diplocheila polita Dejean (A-
C); Dicrochile goryi Boisduval (D-F); and Licinus punctatulus Fabricius (G-I). A, D and G,
dorsal aspects; B, C, E, F, H and I, ventral aspects. Scale bars: A-E, G, H = 200 /urn; F, I =
10 /urn.

However, until such relationship can be established firmly, it seems best to include
these two genera in a single dibasic subtribe rather than recognizing two monobasic
groups.

The markedly shortened terebra characteristic of the Dicrochilina, Lestignathina
and Licinina is accepted as evidence for the monophyly of these three groups, which

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collectively, I refer to as the derived assemblage. Within this assemblage, monophyly
of the Dicrochilina and Lestignathina is postulated, on the basis of the biperforate
coxal cavities exhibited by the members of this complex.

I believe, then, that the Dicaelina, if monophyletic, is the adelphotaxon of the
derived assemblage of subtribes. Within the latter, the Dicrochilina and Lestignathina
comprise a lineage that is the adelphotaxon of the Licinina. In summary, the subtribes
of Licinini are related as follows: Dicaelina + ((Dicrochilina + Lestignathina) +
Licinina).

KEY TO SUBTRIBES, BASED ON FEATURES OF THE
MOUTHPARTS OF ADULTS

1. Mandibular terebra longer, about Vi length of mandible (Fig. 2A); terebral tooth

small (Fig. 2A-B) or absent. Maxilla with lacinial tooth less prominent (Fig. 6A),
occlusal margin densely setose; palpomere 3 longer than palpomere 4 (Fig. 6A-B,
mp-3). Labium (Fig. 7A) with complete suture between submentum and mentum;
mentum ventrally bisetose, paramedian pits (mp) near base of mentum, and pal-
pomere 2 bisetose or quadrisetose Subtribe Dicaelina

1'. Mandibular terebra smaller, about Vi length of mandible (Figs. 2G, 3A, G); terebral
tooth various. Maxilla with lacinial tooth and proportions of maxillary palpomere
3 various. Labium with suture between mentum and submentum complete or not;
mentum bisetose or asetose, with or without paramedian pits, and labial palpomere
2 bisetose or quadrisetose 2

2. (1') Mandible with terebral tooth small (Fig. 2G-H, tt). Maxilla (Fig. 6C) with lacinial

tooth (It) moderately prominent, occlusal margin densely setose; maxillary palpo-
mere 3 (mp-3) longer than palpomere 4. Labium (Fig. 7D) with complete suture
between submentum and mentum; mentum bisetose, and with paramedian pits
basally (mp); labial palpomere 2 quadrisetose Subtribe Dicrochilina

2'. Mandible with terebral tooth prominent, either sharply pointed (Fig. 3A-B), or
broad and blunt (Fig. 3G-H). Maxilla with lacinial tooth prominent, occlusal margin
less densely setose (Fig. 6E-H), and maxillary palpomere 3 various. Labium suture
between submentum and mentum complete or reduced; mentum bisetose or asetose,
paramedian pits absent, and labial palpomere 2 bisetose 3

3. (2') Mandibles about symmetrical, with incisor and terebral tooth long, slender, and

pointed sharply (Fig. 3A-B). Maxilla with occlusal margin of lacinia sparsely setose
(Fig. 6E-F); and maxillary palpomere 3 (mp-3) much shorter than 4. Labium with
suture between submentum and mentum reduced or absent; mentum asetose, and
medial sinus shallow (Fig. 8A) Subtribe Lestignathina

3'. Mandibles markedly asymmetrical, either right or left with large basal boss (b) and
deep notch (n) dorsally (Figs. 3G-H, 1 7B-C, 1 8D-E); terebral tooth broad and blunt
(tt). Maxilla with occlusal margin of lacinia moderately setose (Fig. 6G-H); and
maxillary palpomere 3 (mp-3) subequal in length to 4. Labium with suture between
mentum and submentum complete; mentum bisetose, and medial sinus deep (Fig.

8C) Subtribe Licinina

Genus Diplocheila Brulle

Ball (1959:1 1) arranged the species of Diplocheila in three subgenera: one mono-
basic, the Oriental Neorembus Ball; and two polybasic— the Oriental-eastern Pa-

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laearctic Diplocheila ( sensu stricto ) and the widespread Isorembus Jeannel. Comments
about the latter subgenera are as follows.

Subgenus Diplocheila {sensu stricto)

The species of this subgenus are difficult to distinguish from one another, especially
using only external features of adults. More generally, these beetles resemble speci-
mens of Pterostichus Bonelli. Probably such resemblance accounts for the following
synonymy.

Pterostichus ( Platysma ) karikali Jedlicka

Type material of P. karikali Jedlicka, 1969 includes a holotype and 62 paratypes,
in the ZSMM. The holotype is labelled: “S. Indien 1964/ Pondicherry State/ Karikal
XI/ P. S. Nathan leg”; “Holotypus” [pink paper]; “Pterostichus/ karikali/ sp. n
[handwritten]/ det. Ing. Jedlicka” [pink paper]. Paratypes are from Karikal, also, and
labels are similar to those associated with the holotype. All specimens in the type
series are typical of Diplocheila {sensu stricto) distinguenda in the diagnostic features
of that species (Ball, 1959:3 1, and fig. 59). Accordingly, I propose the new synonymy:
Pterostichus karikali Jedlicka, 1969 = Diplocheila distinguenda Laferte, 1851.

Subgenus Isorembus Jeannel

Ball (1959:48-49) organized the species of Isorembus in three groups: zeelandica
group, with four species confined to the Oriental Region and eastern part of the
Palaearctic Region; the aegyptiaca group, with three species that range collectively
throughout the African continent and Madagascar, and on the southern edge of the
Palaearctic Region eastward to Pakistan; and the striatopunctata group, confined to
the Nearctic Region. Subsequently (Ball, 1966b:25), the southern Chinese species
Diplocheila minima Jedlicka (1931:103) was added to this species group. A formal
reconstructed phylogeny is offered for the species groups, incorporating D. minima
into the system that was proposed previously (Ball, 1959:91-94, and diagram 1).
Comments about geographical history of the striatopunctata group are offered, below.

Reconstructed phylogeny. — See Figures 1 0 and 1 1 . The basis for designation of
character states as plesiotypic or apotypic is outgroup comparison, using as outgroups
the genus Dicaelus and the hypothesized common ancestor of the subgenera Diplo-
cheila {sensu stricto) and Neor embus. Ball (1959:91, lineage 14) lists ancestral states
for Isorembus (Fig. 1 1 , lineage V, here).

Figure 1 0 shows that few characters were available for reconstructing the phylogeny.
It shows also that most of the changes were losses or reductions.

Evidence for group membership of D. minima was based originally on overall
similarity (Ball, 1966b), although in fact two synapotypic features support this as-
signment, as is shown in Figure 10. In the 1966 paper, I stated mistakenly that the
aegyptiaca group was characterized in part by a beaded (or margined) intercoxal
process of the prostemum, a feature of the striatopunctata group and of D. minima.
As indicated in Figure 10, the intercoxal process of members of the aegyptiaca group
is unmargined. Absence of ensiform setae from the ovipositor stylomeres of females
of the Nearctic species indicates that this geographically circumscribed assemblage
is monophyletic.

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Geographical distribution and history

Ball (1959:96-98 and 1966b:22-25) discussed the geographical history of Isorem-
bus against a geological background of continental stability, with the only possible
terrestrial connection between Eurasia and North America being Beringia, in the
north. More recently, it has become clear (and generally accepted by geologists) that
continents have shifted in relation to one another, and that the disappearance of
major epicontinental seaways has changed radically the configuration of land, and
of sea basins (Noonan, 1988:40-43, and references cited therein).

Allen (1983), Noonan (1988) and Liebherr (199 la: 137) have discussed the distri-
bution patterns of various insect taxa (carabids and others) in the light of this geo-
dynamic theory. It seems appropriate to extend such analysis to Isorembus, through
correlation of the reconstructed phylogeny with past geological events (Liebherr,
1986:161).

The distribution pattern of Isorembus was illustrated by Ball (1966b:23, fig. 7), and
it is tabulated in Figure 1 1 . In more detail, the aegyptiaca group is marginal in the
Oriental Region, being represented there by a single species in northern India, and
D. minima, of the striatopunctata group is known only from Szechuan, in southern
China— well to the west of the known range of the zeelandica group, which extends
from the tropics of the Oriental Region to the cool temperate parts of the eastern
Palaearctic Region. The ranges of the zeelandica and aegyptiaca groups are each more
or less continuous.

In contrast, the range of the striatopunctata group is broadly interrupted, with a
monophyletic assemblage of eight species in the Nearctic Region and a single species
known only from Szechuan. Climatic tolerance of the Nearctic assemblage of the
striatopunctata group extends from boreal conditions (two species) to the tropics (one
species), with six species in the general area of cool to warm temperate conditions.
To explain this pattern of geographical and ecological distribution, the following
sequence of events is postulated.

1. Origin of Isorembus in Laurasia, probably in early Tertiary time, this stock
becoming widespread, and occupying Euramerica, to the west of Turgai Straits,
and southern Asiamerica in the east (Fig. 11, lineage V; see also Noonan, 1988:
42, fig. 3).

2. The eastern fragment of ancestral Isorembus differentiated into the ancestral
stock of the zeelandica group (Fig. 1 1 , lineage W), while the western frag-
ment produced the ancestral stock of the aegyptiaca-striatopunctata groups
(lineage X).

3. Separation of lineage X by an unknown vicariance event into a more southern
stock (lineage Y), the ancestor of the aegyptiaca group, that eventually dispersed
into the Afrotropical Region, and a more northern stock (lineage Z), the ancestor
of the striatopunctata group.

4. Separation of the North American tectonic plate from Laurasia, with the con-
sequent formation of the Atlantic Ocean basin, leading to division of lineage
Z into the ancestor of the D. minima clade, and the ancestor of the extant
Nearctic species of the striatopunctata group.

5. The Nearctic clade of the striatopunctata group, differentiated, and extending
its overall climatic tolerance, entered the boreal forest, as the climate cooled in

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later Tertiary time. Also, by overseas dispersal, one species extended its range
to Cuba, in the West Indies.

6. The Palaearctic clade, meanwhile, extended its range across much of the tem-
perate parts of the Palaearctic Region, but eventually became extinct over most
of this area.

The large gap in the range of the striatopunctata group deserves additional com-
ment. Although Diplocheila minima is isolated from the Nearctic clade of the stria-
topunctata group by thousands of kilometres, the extent of the terrestrial component
of this separation may be comparatively recent. Based on fossil evidence, Coope
(1979:264) noted that:

“In response to climatic oscillations of the Quaternary, species of Coleoptera have

altered their geographic ranges on an enormous scale so that where a species is

today may give us little evidence of its geographical history.”

Two examples are offered in support of this statement, as follows. Coope (1973)
reported that the aphodiine scarabaeid species Aphodius holdereri Reitter, known
now from Tibet, only, was represented in Britain as recently as 26,000 years before
present. Hammond et al. (1979) reported that the staphylinid species Anotylus gib-
bulus (Eppelsheim) known today only from eastern China and the Caspian Sea region,
was present in Britain and eastern Canada during the Pleistocene, fossils having been
collected in deposits ranging in age from ca. 39,000 to 300,000 years before present.
Both of these species have undergone comparatively recent and dramatic range changes.
It would seem possible, therefore, that the Diplocheila striatopunctata group was
represented in Europe until the climatic changes of the Pleistocene epoch, with
extensive subsequent extinction. Only D. minima survives, whose range may have
extended far to the west of its present location, in southwestern China.

The main point of this discussion is that the distribution pattern of the striato-
punctata group can be fitted very readily to the vicariance models proposed by Allen
(1983) and Noonan (1988). Tempting as it is to propose a Jurassic origin for the
striatopunctata group (cf. Allen, 1983:626, fig. 65), with postulated vicariance events
paralleling so closely the reconstructed phylogeny of the striatopunctata group, it
seems better to think in terms of a more recent origin and differentiation, and thus
to postulate use of the warm temperate land connections across the North Atlantic
Ocean in early Tertiary time (cf. Matthews, 1979:37, fig. 23). A Beringian route,
postulated by Ball (1959, 1966b), seems less likely, in view of the present distribution
pattern of the Nearctic clade, with its species concentration in the east and in warm
temperate areas.

New World species

Although Lindroth (1968:939) referred to Balks (1959) revision of Licinini as
having been done “with utmost accuracy,” in fact I confused the names of two species
of Nearctic Diplocheila : D. modesta Casey, 1920 is conspecific with D. assimilis
LeConte, 1844, and the names therefore are synonymous; and D. assimilis Ball, 1959
(not LeConte) is conspecific with D. impressicollis Dejean, 1831. Lindroth (1968:
943-944) made the necessary corrections in the valid names for these species.

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Table 2. Distribution pattern of the Diplocheila striatopunctata group in the Nearctic Region.

Geographical zones

Mid-west

East coastal to 100th West of 100th West of No. geog.

Species

states

meridian

to Rockies

Rockies

zones

obtusa LeC.

X

X

X

X

4

striatopunctata LeC.

X

X

X

X

4

assimilis LeC.

X

X

2

impressicollis Dej.

X

X

2

major LeC.

X

X

2

oregona Hatch

X

X

2

undulata Carr

X

X

2

nupera Casey

X

1

Total species

6

6

4

3

Diplocheila nupera Casey

Locality record.— This rarely collected Floridian endemic, is known only from the
southern half of that state (Ball, 1959:73). An additional record is:

Monroe County, Big Pine Key, Long Beach, June 21,1 990; three males, six females,
collected by R. S. Anderson, from Sesuvium litter (CMNC).

The specimens were collected just above high tide line, by sifting debris consisting
of parts of dead Sesuvium plants and material washed on land by the sea. Like most
other members of this genus, adults of D. minima probably are hygrophilous, and
on Big Pine Key, probably they live near the fresh water marshes that occur there
(R. S. Anderson, pers. comm.). Thus, I believe that the sea beach record is accidental
for this species.

Diplocheila undulata Carr

Locality record.— This rarely collected species is known from a few localities in
Illinois, Manitoba, and Alberta. The species was described in 1920 on the basis of
specimens labelled “Edmonton, Alberta” (Ball, 1959:8). Lindroth (1969:946) noted
“Ecology not recorded for this species.” Because its putative adelphotaxon, D. obtusa
LeConte is unusual for Isorembus, living in mesic to dry situations, I assumed that
D. undulata lived in similar habitats.

However, five specimens were collected in May and June, 1985, in pitfall traps
placed at the edge of a permanent marl pond, and in adjacent sedge marsh, at Wagner
Bog, which is located ca. 1 3 km west of Edmonton’s city limits.

Technically, Wagner Bog is a mixed mire, with calcareous mud pools, and sup-
porting a rich flora, including brown mosses and some Sphagnum (Vitt, 1982:17-
22). Thus, Diplocheila undulata is hygrophilous, occupying a distinctive habitat. One
of 60 species of carabids taken at Wagner Bog, D. undulata shares this area with
many northern groups of ground beetles, such as B/ethisa, Elaphrus, Loricera, Pa-
trobus, Agonum (subgenus Europhilus), and Chlaenius.

352

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

CHAR.

No.

CHARACTER

CHARACTER STATES AND TAXA

zeeland. aegypt.

group group

striatopunctata group
D. minima Nearclic spp.

9 stylomere 2:
setae number

<D

Prosternum:

apex

Tarsomere 5:
ventral setae

Left mandible:
basal occ. area

Head: s-o setae
number

El: interneur 1 ,
base

El: discal seta

LEGEND

Character State

o

plesiotypic

apotypic, gain

apotypic, loss

apotypic, independent loss

Fig. 10. Characters and phylogenetic designation and distribution of character states among
the species groups of subgenus Isorembus Jeannel.

Genus Dicaelus Bonelli

Ball (1959:102-103) arranged the 14 species of this genus recognized at that time
into three subgenera: Liodicaelus Casey; Paradicaelus Ball; and Dicaelus ( sensu stric-
to). Casual examination of additional material of Paradicaelus and subgenus Dicaelus
that I have collected or received for identification has not revealed new taxa or any
marked flaws in the classification that I proposed. Lindroth (1969:948) stated that
he “. . . had nothing to add on the taxonomy of the genus,” and confined his treatment
to means of identification and characterization of the species of Dicaelus occurring
in Canada.

Work in Mexico during the past three decades produced new locality records and

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<

3EOGRAPHK

CAL DISTRIBUTION

Oriental

illlil

1111

E. Palaearctic

M’lvIvI'IvIvIvM

S. Palaearctic

X'MvIv!

Afrotropical

Nearctic

N. Neotropical

GEOLOGICAL

TIME

/ / J '/ &/

/ / §/ §/q-

Jvsy

//<? y *

Holocene
Eo-/ Oligocene

► 3,’ 4, 7a

,»7B-

'

► 5,' 6

■CD

'1

(

v)

Fig. 1 1 . Reconstructed phylogeny and geographical distribution of the species groups of
subgenus Isor embus Jeannel.

354

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

a new species of Liodicaelus, as noted below. Also, the new material has led to
modification of some previous decisions about ranking and recognition of taxa.

Because of the taxonomic importance of mandibles and diagnostic value of body
proportions, these features are reviewed separately from the following descriptive
treatment of taxa. A revised key to the species of Liodicaelus is provided, to accom-
modate the new species and taxonomic changes proposed.

Subgenus Liodicaelus Casey

Figure 1 2 illustrates the habitus of a specimen of Liodicaelus. The most important
feature for recognition is the nearly smooth elytra, resulting from absence of intemeurs
(or their reduction to rows of punctures), and flattening of the intervals. The subgenus
is circumscribed geographically, being represented: only in the Great Plains States
(and southern Alberta) west of the 100th meridian; in the southwest in New Mexico
and Arizona; and in the mountains of Mexico southward to the Trans- Volcanic Sierra
(Figs. 1 7 and 1 8). Habitats occupied are grassland and dry to mesic forest, ranging
in altitude from near sea level to more than 2,000 m.

Structural features: mandibles

The mandibles of Liodicaelus are average for Dicaelina: retinacular and molar
ridges, occlusal grooves, and basal brush absent; ventral grooves each with short
microtrichia (Fig. 13F, vmt). The left mandible lacks a terebral tooth, or has a small
one; the terebral tooth of the right mandible is prominent.

Mandibles of D. abbreviatus Bates (Fig. 13A-F). — Left mandible (Fig. 13 A, C, E)
with terebral tooth (Fig. 13 A) blunt, hardly projected; basal ridge narrow, not wid-
ened. Right mandible (Fig. 13B, D, F) average for Liodicaelus.

Mandibles of D. laevipennis LeConte (Fig. 13G-L). — Left mandible (Fig. 13G, I,
K) with terebral ridge (tr) and basal ridge (br) nearly continuous, terebral tooth (tt)
very small, hardly projected; basal ridge (br) slightly convex, evenly curved, in ventral
aspect hardly extended beyond tips of ventral microtrichia (Fig. 1 3K). Right mandible
(Fig. 13H, J, L) with terebral tooth prominent.

Mandibles of D.franclemonti, new species (Fig. 14A-F). — Left mandible (Fig. 14A,
C, E) with terebral ridge (tr) and basal ridge (br) in one continuous curve, terebral
tooth not evident; basal ridge projected well beyond tips of ventral microtrichia (Fig.
14F, vmt), and thickened as blunt basal tooth (bt). Right mandible (Fig. 14B, D, F)
average for Liodicaelus : terebral tooth (tt) blunt and somewhat worn.

Comparisons.— These mandibles form a transformation series, from D. abbre-
viatus, with the best developed terebra, including tooth, to D. franclemonti, with
terebra reduced and basal area enlarged. However, this sequence cannot be the
complete basis for a reconstructed phylogeny, as indicated below (see notes about
phylogenetic relationships, for D. franclemonti).

Structural features: measurements and body proportions

Tables 3-6 provide quantitative data about variation in size (length) and in pro-
portions. For Liodicaelus, most such differences are not useful for distinguishing all

1992

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Table 3. Variation in Standardized Body Length (mm) among males and females of species
and subspecies of subgenus Liodicaelus Casey.

Males

Females

Species and population sample

N

Range

Mean

N

Range

Mean

D. franclemonti, n. sp.

Alazanas, Coahuila

1

14.9

Cerro Potosi, N.L.

1

16.2

1

16.8

D. sujfusus Casey

1959 data

6

13.4-14.8

14.3

6

12.1-15.5

14.4

Tres Rios (Son.), Chih.

5

11.1-12.2

11.7

3

12.0-12.4

12.3

vie. Col. Garcia, Chih.

7

12.3-13.2

12.8

6

13.1-13.3

13.2

vie. Madera, Chihuahua

12.7-13.6

13.1

vie. Tomochic, Chih.

7

12.9-14.3

13.5

5

12.8-13.9

13.3

D. abbreviatus Bates

1959 data

1

15.2

3

15.3-15.6

15.4

W Huejuquillo el Alto Jal.

1

14.9

W El Vergel, Durango

1

14.5

D. 1. laevipennis LeConte

1959 data (/. laev.)

16

16.5-20.0

18.8

9

17.2-19.0

18.4

1959 data (/. dicael.)

3

17.0-17.6

17.3

11

17.1-19.1

18.0

Pinos Altos Mtns., N.M.

14

13.4-15.6

14.7

8

14.1-15.7

14.9

Col. Zaragosa, Chih.
vie. Madera, Chihuahua

1

18.1

1

17.2

D. 1. flohri Bates

1959 data

2

18.7-20.5

19.6

1

19.2

Atlacomulco, Mexico

3

16.2-16.8

16.5

El Temascal, Durango

3

15.3-17.9

16.3

10

16.2-18.7

17.7

D. chermocki Ball

1959 data

Huachuca Mtns., Arizona

2

17.2-18.3

17.8

7

17.8-20.0

18.8

individuals of any one species from all other species, but they show trends, which
may be employed in analysis of relationships.

Figure 1 5 is a scattergram on which are plotted values for the ratios PL/PWA and
EL/EW. Because I do not have the original data summarized in the 1959 paper (tables
46, 47, 49), I have plotted only the mean values for these samples.

The position on the graph for each species is explained in simple terms, as follows.
Points for Dicaelus suffusus occupy a central area, indicating pronota not especially
narrowed apically, and elytra that are relatively narrow.

The points for both D. chermocki and D. franclemonti are low and more or less
central on the graph, indicating broad pronota apically (to accommodate an enlarged
head— cf. Fig. 12), and elytra of average proportions.

The points for D. abbreviatus occupy the left side of the graph, indicating short,
broad elytra and pronotal apices that vary from broad to relatively narrow.

356 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Table 4. Variation in values for the ratio PW/HW among males and females of species and
subspecies of subgenus Liodicaelus Casey.

Species and population sample

Males

Females

N

Range

Mean

N

Range

Mean

D. franclemonti, n. sp.

Alazanas, Coahuila

1

1.67

Cerro Potosi, N.L.

1

1.64

1

1.63

D. suffusus Casey

1959 data

6

1.75-2.00

1.87

6

1.73-1.84

1.78

Tres Rios (Son.), Chih.

5

1.72-1.90

1.79

3

1.66-1.77

1.70

vie. Col. Garcia, Chih.

7

1.70-1.87

1.77

6

1.80-1.88

1.84

vie. Madera, Chihuahua

3

1.84-1.88

1.85

vie. Tomochic, Chih.

7

1.78-1.91

1.84

5

1.86-1.88

1.87

D. abbreviatus Bates

1959 data

1

2.10

3

1.97-2.06

2.01

W Huejuquillo el Alto Jal.

1

1.94

W El Vergel, Durango

1

2.06

D. 1. laevipennis LeConte

1959 data (/. laev.)

16

1.99-2.20

2.10

9

1.94-2.20

2.09

1959 data (/. dicael.)

3

2.00-2.09

2.04

1 1

1.92-2.10

2.02

Pinos Altos Mtns., N.M.

14

1.86-2.09

1.97

8

1.88-2.03

1.97

Col. Zaragosa, Chih.

1

1.90

vie. Madera, Chihuahua

1

2.07

D. 1. flohri Bates

1959 data

2

2.06

1

2.12

Atlacomulco, Mexico

3

1.95-2.00

1.98

El Temascal, Durango

3

2.03-2.17

2.11

10

1.98-2.20

2.08

D. chermocki Ball

1959 data

7

1.53-1.71

1.62

Huachuca Mtns., Arizona

2

1.56-1.68

1.62

The points for D. laevipennis are relatively high on the graph, indicating average
elytral proportions but narrowed pronotal apices.

Overall, something of a radial pattern is indicated, with the center represented by
D. suffusus, and outliers in different directions being: D. chermocki and D. francle-
monti\ D. abbreviatus; and D. laevipennis. No doubt, this pattern will be of use in
phylogenetic analysis, although I doubt that the similarity exhibited by D. chermocki
and D. franclemonti indicate close relationship of these taxa.

KEY TO SPECIES AND SUBSPECIES OF SUBGENUS LIOD1CAELUS
CASEY (MODIFIED FROM BALL, 1959:169)

1. Elytral interval 7 not carinate. Elytra relatively short and broad, EL/EW 1.30-1.36

D. abbreviatus Bates

1992 THE TRIBE LICININI 357

Table 5. Variation in values for the ratio PL/PWA among males and females of species and
subspecies of subgenus Liodicaelus Casey.

Species and population sample

Males

Females

N

Range

Mean

N

Range

Mean

D. franclemonti, n. sp.

Alazanas, Coahuila

1

1.00

Cerro Potosi, N.L.

1

0.98

1

0.95

D. suffusus Casey

1959 data

6

1.19-1.23

1.20

6

1.10-1.21

1.16

Tres Rios (Son.), Chih.

5

1.12-1.20

1.16

3

1.09-1.15

1.13

vie. Col. Garcia, Chih.

7

1.08-1.18

1.14

6

1.11-1.17

1.14

vie. Madera, Chihuahua

3

1.08-1.14

1.11

vie. Tomochic, Chih.

7

1.10-1.18

1.15

5

1.08-1.14

1.11

D. abbreviatus Bates

1959 data

1

1.20

3

1.1 1-1.22

1.17

W Huejuquillo el Alto Jal.

1

1.07

W El Vergel, Durango

1

1.24

D. 1. laevipennis LeConte

1959 data (/. laev.)

16

1.15-1.34

1.23

9

1.18-1.26

1.21

1959 data (/. dicael.)

3

1.25-1.28

1.26

11

1.15-1.22

1.18

Pinos Altos Mtns., N.M.

14

1.18-1.29

1.24

8

1.15-1.30

1.21

Col. Zaragosa, Chih.

1

1.15

vie. Madera, Chihuahua

1

1.25

D. 1. flohri Bates

1959 data

2

1.24-1.30

1.27

1

1.24

Atlacomulco, Mexico

3

1.16-1.28

1.23

El Temascal, Durango

3

1.14-1.21

1.17

10

1.13-1.21

1.17

D. chermocki Ball

1959 data

7

0.98-1.05

1.00

Huachuca Mtns., Arizona

2

1.02-1.08

1.05

T. Elytral interval 7 carinate near humerus (cf. Fig. 1 2). Elytra of most specimens longer

and more slender, EL/EW 1.34-1.52 2

2(T). Head relatively large, PW/HW 1.53-1.71, and apical margin of pronotum relatively

broad, PL/PWA 0.95-1.08 (cf. Fig. 12) 3

2'. Head smaller, PW/HW 1 .66-2. 1 7 and apical margin of pronotum relatively narrow,

PL/PWA 1.08-1.34 4

3(2). Elytral intemeurs each a row of clearly impressed punctures. Range— mountains of

southwestern Arizona D. chermocki Ball

3'. Elytral intemeurs impunctate, each a very shallow indistinct stria, or elytra smooth,
without indication of intemeurs (Fig. 12). Range— mountains of Sierra Madre Ori-
ental, Mexico D. franclemonti, new species

4(2'). Size smaller, Standardized Body Length less than 15 mm (most specimens 12-14

mm). Pronotum anteriorly relatively broader (PL/PWA 1.23 or less) D. suffusus Casey
4'. Size larger, SBL 13 mm or more (most specimens 15-18 mm). Pronotum anteriorly

relatively narrower (most specimens with PL/PWA more than 1.20; cf. Fig. 15) ... 5

358 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Table 6. Variation in values for the ratio EL/EW among males and females of species and
subspecies of subgenus Liodicaelus Casey.

Species and population sample

Males

Females

N

Range

Mean

N

Range

Mean

D. franclemonti, n. sp.

Alazanas, Coahuila

1

1.44

Cerro Potosi, N.L.

1

1.46

1

1.47

D. sujfusus Casey

1959 data

6

1.46-1.52

1.50

6

1.44-1.55

1.48

Tres Rios (Son.), Chih.

5

1.41-1.50

1.45

3

1.41

1.41

vie. Col. Garcia, Chih.

7

1.40-1.50

1.46

6

1.36-1.46

1.40

vie. Madera, Chihuahua

3

1.38-1.43

1.40

vie. Tomochic, Chih.

7

1.36-1.42

5

1.33-1.41

1.38

D. abbreviatus Bates

1959 data

1

1.33

3

1.30-1.36

1.32

W Huejuquillo el Alto Jal.

1

1.29

W El Vergel, Durango

1

1.34

D. 1. laevipennis LeConte

1959 data (/. laev.)

16

1.42-1.55

1.48

9

1.40-1.50

1.44

1959 data (/. dicael.)

3

1.45-1.48

1.46

11

1.40-1.54

1.44

Pinos Altos Mtns., N.M.

14

1.34-1.46

1.39

8

1.34-1.44

1.39

Col. Zaragosa, Chih.

1

1.39

vie. Madera, Chihuahua

1

1.42

D. 1. flohri Bates

1959 data

2

1.50-1.52

1.51

1

1.46

Atlacomulco, Mexico

3

1.40-1.43

1.42

El Temascal, Durango

3

1.35-1.37

1.36

10

1.31-1.40

1.35

D. chermocki Ball

1959 data

7

1.45-1.52

1.48

Huachuca Mtns., Arizona

2

1.34-1.41

1.38

5(4'). Elytra with each intemeur indicated by row of distinct punctures. Range— Chihua-
hua and Sonora, north to southern Alberta D. 1. laevipennis LeConte

5'. Elytra with intemeurs 1-3 each indicated by row of distinct punctures; intemeurs
4-8 either not evident, or punctures of intemeurs 4-6 small, indistinct. Range—
southern Durango, in the Sierra Madre Occidental, south to the state of Mexico, in
the Trans- Volcanic Sierra .....< D. laevipennis flohri Bates

THE SPECIES OF LIODICAELUS’. NOTES,

DESCRIPTIONS, AND DISTRIBUTION RECORDS

Dicaelus (Liodicaelus) franclemonti, new species

Type material. — HOLOTYPE male, labelled: “MEX. Nuevo Leon/ Cerro de Po-
tosi/ 12.4 mi. n.w./ Galeana, 72007 e. slope/ X. 17.65”; “George E. Ball/ D. R.
Whitehead/ collectors” (USNM). ALLOTYPE female, labelled same as holotype

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359

Fig. 12. Photograph of allotype of Dicaelus ( Liodicaelus ) franclemonti, new species, illus-
trating habitus. SBL = 16.8 mm.

(USNM). One additional PARATYPE, labelled: “MEX. Coahuila/ 21 km. e. San/
Antonio de las Alazanas 2550 m/ oak-madrono/ fir forest/ July 25, 1975”; “MEX
EXP. 1975/ G. E. Ball &/ H. E. Frania/ collectors” (UASM).

Type locality.— Mexico, state of Nuevo Leon, the mountain Cerro Potosi, north-
west of Galeana.

Derivation of specific epithet.— Based on the surname of John G. Franclemont;
this species is named in his honor, to commemorate on his 80th birthday his ded-
ication to entomology and to the education of systematic entomologists.

Recognition.— Adults of this species share the following unique combination of
character states: integument black, elytra virtually smooth, with traces only of smooth,
impunctate intemeurs (Fig. 1 2); head and apical margin of pronotum relatively broad.
Males have a dorso-preapical projection on the median lobe, and the internal sac
has four spines, only.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Fig. 13A-L. SEM photographs of the mandibles of Dicaelus abbreviatus Bates and of D.
laevipennis LeConte. A-F, D. abbreviatus : A, C and E, left mandible— dorsal, occlusal and
ventral aspects, respectively; B, D and F, right mandible— dorsal, occlusal and ventral aspects,
respectively. G-L, D. laevipennis : G, I and K, left mandible— dorsal, occlusal and ventral aspects,
respectively; H, J and L, right mandible— dorsal, occlusal and ventral aspects, respectively.
Legend: br, basal ridge; i, incisor; tr, terebral ridge; tt, terebral tooth; vg, ventral groove; and
vmt, ventral microtrichia. Scale bars = 400 yum.

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Fig. 14A-F. SEM photographs of the mandibles of Dicaelus franclemonti, new species: A,
C and E, left mandible— dorsal, occlusal and ventral aspects, respectively; B, D and F, right
mandible— dorsal, occlusal and ventral aspects, respectively. Legend: br, basal ridge; bt, basal
tooth; i, incisor; tr, terebral ridge; tt, terebral tooth; vg, ventral groove; and vmt, ventral
microtrichia. Scale bars = 400 /urn.

362

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

1.30

1.30

1.30

1.30

EL/EW

1.30

Fig. 1 5. Scattergram for ratios PL/PWA and EL/EW for the species of subgenus Liodicaelus
Casey.

In body proportions, adults of D.franclemonti are most like those of D. chermocki.
Smooth elytra, with hardly distinguishable impunctate elytral intemeurs, distinguish
D. franclemonti. Reduced elytral intemeurs are characteristic of D. laevipennis flohri
and D. abbreviatus, but adults of these species have smaller heads and prothoraces
with narrower anterior margins.

Description. — With the character states of subgenus Liodicaelus. Habitus as in
Figure 12. Data about Standardized Body Length, elytral width, and ratios PW/HW,
PL/PWA, and EL/EW are presented in Tables 3-6. Integument black generally, with
palpomeres dark rufo-piceous. Microsculpture of body generally isodiametric, sculp-
ticells small and on dorsal surface slightly convex. Dorsal surface with soft silky
luster. Head and pronotum with surface features as for other species of Liodicaelus
(cf. Ball, 1959:176). Mandibles as in Fig. 14A-F. Elytra each with interval 7 toward
humerus carinate, each carina ca. 0.25 length of elytron, and broader basally than
in the other carinate species of Liodicaelus ; otherwise, dorsal surface virtually smooth,
intemeurs indicated by indistinct lines. Sternum VII with apical setae 4-5 in males,
and 2-4 in females.

Male genitalia. As in Figure 1 6 A, B. Median lobe in left lateral aspect with basal
part bent obliquely quite markedly; shaft with ventral surface straight, apex acute
(Fig. 16 A); in ventral aspect (Fig. 16B), apical portion straight, not curved to right;
apex obtusely and evenly rounded, not notched shallowly. Internal sac with four
large spines, only. Parameres average (cf. Ball, 1959, fig. 1 12c, d).

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1mm

Fig. 1 6A-D. Line drawings of male genitalia of species of Liodicaelus Casey. A-B, median
lobe of D. franclemonti, new species: A, median lobe, left lateral aspects; B, median lobe, apical
portion, ventral aspect, showing armature of internal sac. C-D, median lobe of D. chermocki
Ball: C, median lobe, left lateral aspect; and D, median lobe, apical portion, ventral aspect,
showing armature of internal sac. Legend: dr, dorsal ridge; is, spine of internal sac; and ss,
sclerotized strip.

Habitat. — On Cerro Potosi, the holotype and allotype of D. franclemonti were
collected at an altitude of 2,200 m, in a damp stream bed, under stones, with a
covering of oak and pine litter. The paratype male, collected near San Antonio de
las Alazanas at 2,550 m, was under a log, in a dry conifer-oak forest. Therefore, I
infer that D. franclemonti is an inhabitant of the mesic to dry montane forests that
are characteristic of the upper slopes of the northern sierras, in Mexico.

Geographical distribution (Fig. 1 7).— This species is known from the northern part
of the Mexican Sierra Madre Oriental, only.

Chorological affinities.— D. franclemonti is isolated geographically from all other
known species of Dicaelus. The other species of Liodicaelus occur to the north, in
Texas (Fig. 18), or westward, in the Sierra Madre Occidental, or southward, in the
Trans- Volcanic Sierra.

Phylogenetic relationships. — Based on features of the male genitalia, D. francle-
monti is postulated to be the most primitive extant member of Liodicaelus, and to
be the adelphotaxon of the other species of this subgenus. In turn, these species
comprise a monophyletic assemblage, based on absence of the dorsal preapical ridge
of the male median lobe, which is postulated to be a loss. This latter species assem-
blage is not analyzed further now, though the sequence of species used in the text

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and summarized here indicates how I believe they are related, based on details of
mandibular structure and body form: D. sujfusus-abbreviatus-laevipennis-chermocki.

Dicaelus sujfusus Casey

This species was known previously from “the Sierra Madre Mountains of Chi-
huahua, Mexico” (type area) and from the Chiricahua Mountains of Arizona (Ball,
1959:177-178). Additional material documents more fully the range of D. sujfusus.

Type locality. — Here restricted to the vicinity of Madera, Chihuahua, Mexico.

Material examined.— Forty four specimens, from the following localities (Fig. 17).

U.S.A. ARIZONA. Cochise County. Female, Chiricahua Mts.: X.4.1937 Bryant
Lot 65 (CASC). Male, Pinery Canyon, VII.9.1956; H. & A. Howden (CNCI). 2
females, Rustlers Park, 2,500 m, VII. 14. 1952, H. B. Leech, J. W. Green (CASC). 3
females, Rustlers Park, 2,560 m, VII.26-27.1979; A. Smetana (CNCI).

MEXICO. CHIHUAHUA. 4 males, 4.8 km S Colonia Garcia, pine-oak-madrono
forest, 2,130 m, VIII.4&28. 1980; G. E. Ball, D. Shpeley, & W. Fraga (UASM). Male,
4 females, Colonia Garcia, meadow & adj. pine-oak forest, 2,130 m, VIII. 5&28. 1980;
Ball, Shpeley & Fraga (UASM). 24 km E Creel, meadow, 2,200 m, VIII. 24. 1980;
Ball, Shpeley & Fraga (UASM). Male, 2 females, 1.3 km NE Ejido Zaragosa, pine-
oak forest, 1,850 m, VII.22.1979; J. S. Ashe, Ball & Shpeley (UASM). Male, ca. 14.8
km. E Ejido Zaragosa, pine-oak-aspen forest, 2,120 m, VIII.3.1980; Ball, Shpeley &
Fraga (UASM). 2 females, 8.8 km W Madera, 2,270 m, VIII. 30. 1974; Ball & H. E.
Frania (UASM). One female, 10.5 km W Madera, 2,320 m, pine-oak forest, 2,320
m, VIII. 29-30. 1974 (UASM). 3 males, 28.3 km E Mesa de Tres Rios (Sonora), pine-
oak forest, 2,160 m, VII.7.1983; Frania, R. Jaagumagi & Shpeley (UASM). 7 males,
five females, 25.4 km N Tomochic, pine-oak forest, 2,250 m, VIII.25.1980; Ball,
Shpeley & Fraga (UASM).

SONORA. Male, female. Sierra Huachinera, 32-34 km NE Nacori Chico, pine-
oak forest, 1,950 m, VIII.6-7. 1982; G. E. & K. E. Ball & S. McCleve (UASM).
Female, Sierra Huachinera, 56 km NE Nacori Chico, pine-oak forest, 2,330 m,
VIII. 8. 1982; Ball, Ball & McCleve (UASM). Male, female, 18.5 km E Mesa de Tres
Rios, pine-oak forest, 2,440 m, VII.7.1983; Frania, Jaagumagi & Shpeley (UASM).

Dicaelus abbreviatus Bates

Described originally as a variety of D. laevipennis LeConte (Bates, 1891:239), from
Refugio, Durango, this taxon is ranked as a distinct species on the basis of consistency
of its diagnostic features (including mandibles, Fig. 1 3 A-F) over its fairly extensive
geographical range, and though it has not been collected in the exact same places as
any other taxa of Liodicaelus, its range is partially overlapped by the range of D.
laevipennis Jlohri Bates (Fig. 1 7). Thus, D. abbreviatus maintains its distinctiveness
in parapatry, if not sympatry.

Material examined. — Four specimens from the following localities (Fig. 17).

MEXICO. DURANGO. Female, 68 km W Durango, 2,438 m, VI. 1 8. 1 97 1 (CNCI).
Male, 4.8 km E El Salto, VI.21.1964 (CNCI). Male, 72.9 km W Huajuquillo el Alto
(Jalisco), rd. to Jesus Maria (Nayarit), 2,540 m, VII. 28. 1 985, H. E. Frania, D. Shpeley
(UASM).

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SINALOA. Male, 199.4 km W El Vergel (Chihuahua), 2,400 m, pine-oak-madrono
litter, VII. 16. 1983 (UASM).

Dicaelus laevipennis LeConte

Ball (1959:169-175) recognized four geographical races of this species: D. /. lae-
vipennis?; D. 1. dicaeloides Ball; D. 1. abbreviatus, and D. 1. flohri Bates. See above for
taxonomic treatment of D. abbreviatus. The subspecies D. 1. dicaeloides (type locality
Cloudcroft, Otero Co., New Mexico [Ball, 1959:172]) was based on primarily the
mandibular features of the type material. Re-examination of part of the type series
satisfies me that the differences between the mandibles of this named form and
nominotypical D. laevipennis is the result of wear, and thus without taxonomic
significance. Accordingly, these two forms are regarded as consubspecific, and the
name D. 1. dicaeloides is a junior subjective synonym of D. 1. laevipennis (new syn-
onymy).

The gap in geographical range between D. 1. laevipennis and D. 1. flohri (Figs. 1 7
and 1 8) is substantial, but I think it represents a collecting artifact, only.

Material examined.— Forty five specimens of D. 1. laevipennis and 13 specimens
of D. 1. flohri, from the following localities.

D. 1. laevipennis LeConte (Fig. 1 8)

U.S.A. ARIZONA. Apache County. Male, St. Michael’s, VI.8.1971; D. Brzoska
(SEMC). Cochise Co. Male, Hwy. 666, 2.6 km SE Interstate Hwy 10, VII.29.1985;
D. Brzoska (SEMC). Coconino Co. Female, Hwy. 64, S Grand Canyon Nat. Park,
VIII. 3 1.1 967; P. S. Bartholemew (CASC). Mojave Co. Male, Peach Springs,
VIII. 22. 1967; S. McCleve (SMCC). Navajo Co. Male, Show Low, IX.5.1967, Bar-
tholemew (CASC). County not determined. Female, White Mts; D. K. Duncan (CASC).

COLORADO. Fremont Co. Female, Phantom Canyon. IX. 13. 1964; B. H. Banta
(CASC). Kiowa Co. Female, Arlington, VIII. 16. 1947; H. G. Rodeck (CASC). Routt
Co. Male, Steamboat Springs, 2,073 m, VI. 15. 1945; Bryant Lot (CASC). Teller Co.
Male, Manitou, VIII.21.1958; A. C. Cole (CNCI).

MONTANA. Gallatin Co. Male, Bozeman, V. 18. 1926 (CNCI).

NEW MEXICO. Bernalillo Co. Female, Albuquerque, IX. 19. 1965; P. S. Bartho-
lemew (CASC). Grant Co. Male, Hwy. 61,51.8 km E Beaverhead, 2,469 m, IX. 13.1 967;
Ball, T. L. Erwin & R. E. Leech (UASM). 5 males, 4 females, 16.1 km NE Pinos
Altos, IX. 12. 1967; Ball, Erwin, & Leech (UASM). Male, 2 females, Hwy. 25, 46.8
km NE Pinos Altos, 1,920 m, IX. 13. 1967; Ball, Erwin & Leech (UASM). Hidalgo
Co. Female, 3.2 km NE Rodeo, VII.30.1963; A. Rathke (CNCI). Otero Co. Female,
Lincoln Nat. Forest, 3.2 km NE Cloudcroft, 2,530 m, VII. 13. 1969; A. Smetana
(CNCI). Rio Arriba Co. Male, Capulin Mountain, IV.21.195 1; Bryant Lot 5 (CASC).
Socorro Co. Male, Magdalena Mountains, 32.2 km S Magdalena, 2,316 m,
VIII. 19. 1969; G. E. & K. E. Ball (UASM).

SOUTH DAKOTA. Fall River Co. Female, Hot Springs, IX. 19. 1906; E. Van Sant
(CASC).

MEXICO. CHIHUAHUA. Female, 4. 5-6. 5 km N Ejido Zaragosa, dry pine-oak
forest, under logs, 2,000 m, VII. 23. 1979; J. S. Ashe, G. E. Ball & D. Shpeley (UASM).

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Male, Hwy. 16, 21 km SE Madera, 2,100 m, grassland, IX. 1.1 974; Frania & Ball
(UASM).

D. 1. flohri Bates (Fig. 1 7)

MEXICO. DURANGO. 2 males, 1 1 females, Reserva de la Biosfera La Michilia,
El Temascal meadow, Potrillo La Laguna, 5 km S Trampa Las Casas, VIII. 19. 1980;
Ball, Shpeley & Fraga (UASM).

MEXICO (state of)- 3 males, 6.4 km N Atlacomulco, 2,652 m, VIII. 15. 1974; C.
W. & L. O’Brien & B. Marshall (UASM).

ZACATECAS. Male, 1 .9 km SW Villa de los ...[?], 2,350 m, VIII. 1 966; D. Thomp-
son (UASM).

Dicaelus chermocki Ball

The original description of this species was based on seven females, males being
unknown at that time (Ball, 1959:177). Subsequently, males were collected, and the
genitalia (Fig. 16C-D) were found to be identical to those of D. laevipennis and the
other taxa recognized in 1959.

Dicaelus chermocki is known only from the Huachuca and Chiricahua Mountains
of southeastern Arizona (Fig. 1 7). The few additional specimens collected since 1959
confirm the distinctness of the species, and extend its altitudinal range downward
from 2,286 m in the Huachuca Mountains to 1,402 m, at Sierra Vista, which is
located at the base of that mountain range.

Material examined.— Three specimens from the following localities (Fig. 17).

U.S.A. ARIZONA. Cochise County. Huachuca Mts.: male, Carr Cn., fir-pine forest,
2,150 m, VII. 23. 1976; Ball, J. M. Campbell & P. M. Hammond (UASM); male,
Ramsey Cn., III.22. 1956, F. G. Werner, G. D. Butler (UASM). Female, Sierra Vista,
VII.29. 1963, R. F. Stemitsky (CNCI).

Notes about geographical history of Liodicaelus

Ball (1959:185) noted the general association of Liodicaelus with the Madro-Ter-
tiary flora, but did not attempt a biogeographic analysis because the species were
known from few localities, and thus their ranges were understood inadequately. The
additional locality data presented above improves knowledge of the species ranges,
and the new species provides a new perspective. Furthermore, Liebherr ( 199 la: 1 37—
150, 1991b) provided a valuable general vicariance-based model and geological time-
frame for interpretation of distribution patterns of montane taxa in Mexico and
southwestern United States.

Briefly stated, Liebherr’s analysis explains the geographical history of vicariously
distributed taxa in terms of a sequence of range barriers operating in sequence:

1. isolation of the Trans-Volcanic Sierra (M) from the northern Sierra Madre
Oriental (A') + northern Sierra Madre Occidental (A + B) + southeastern
Arizona (A") + the rest of North America (N.A.);

2. isolation of A' from (A + B) + A" + N.A.;

3. isolation of (A + B) from A" + N.A.;

4. isolation of A" from N.A.

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Table 7. Distribution pattern of the taxa of subgenus Liodicaelus Casey, in terms of vicar-

iance areas of Liebherr (1991b).

Areas1

Taxon

M

A'

A + B

A"

N.A.

franclemonti, n. sp.

X2

suffusus Casey

X3

X

abbreviatus Bates

X

/. laevipennis LeC.

X3

X

X

/. flohri Bates

X

X4

chermocki Ball

X

1 Areas: M, Trans-Volcanic Sierra; A', northern Sierra Madre Oriental; A + B, northern Sierra
Madre Occidental; A", southwestern U.S.A. (see text for details); and N.A., North America,
north, west and east of A".

2 X, a species known from a single area, only.

3 In the northern part of A + B, only.

4 In the southernmost part, only.

The time span inferred for this sequence of events is ca. 1 2 million years, beginning
in Late Miocene.

The species of Liodicaelus are placed, in Table 7, in the context of Liebherr’s
analysis. Four points emerge: first, each area has a taxon that is endemic or nearly
endemic to it; second, although D. abbreviatus and D. suffusus are in the Sierra Madre
Occidental, their ranges seem not to overlap; third, the northern Sierra Madre Oriental
has in it a taxon that is postulated to be the most primitive member of the subgenus,
and to be the adelphotaxon of the remaining taxa; and fourth, ignoring the gap that
separates D. 1. laevipennis from D. 1. flohri, the total range of D. laevipennis bridges
all of the vicariance areas, except for the Sierra Madre Oriental. Points one and two,
regarding endemism, indicate that differentiation in Liodicaelus has been influenced
by the same sets of barriers and conditions that have influenced differentiation of
many other taxa in northern Mexico. To this extent, the data for Liodicaelus fit the
general geographical pattern described by Liebherr. However, concerning the phy-
logenetic pattern, the fit of the taxa of Liodicaelus is inexact (point three). One would
expect the most primitive adelphotaxon, postulated to be D. franclemonti, to be in
the Trans-Volcanic Sierra and perhaps the southern part of the Sierra Madre Occi-
dental—i.e., in the area occupied by D. 1. flohri. This element of discordance suggests
that the sequence of vicariance events for Liodicaelus was different from the sequence
indicated by Liebherr’s interpretation of the data that he used.

Because of the extensive range overlap of D. laevipennis with the ranges of the
other species, the distribution data suggest also that the barriers in the west, between
the Trans-Volcanic Sierra, the northern Sierra Madre Occidental and the Rocky
Mountain outliers in Arizona are more readily penetrated than the barrier between
the Sierra Madre Oriental and the Sierra Madre Occidental. Indeed, in spite of the
postulated Pleistocene connections between the eastern and western montane biotas,
there is evidence to the contrary— for example the Carabus forreri (S. M. Occidental-

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Fig. 17. Geographical distribution of the taxa of subgenus Liodicaelus Casey (except D. 1.
laevipennis ) in Mexico and southwestern United States.

southern Arizona)— C. hendrichsi (S. M. Oriental) species pair, absence of Scaphi-
notus from the Sierra Madre Oriental, and primitive species of subgenus Progaleritina
Jeannel and of Pasimachus Bonelli isolated in the eastern Mexican forests from their
western counterparts.

Nonetheless, I am disinclined to believe that a single general sequential pattern of
taxon/area relationships exists, just as I am inclined to believe that barriers between
adjacent mountain systems are permeable to semi-permeable as frequently as they
are impermeable to taxa on either side of them. Probably different taxa, even with
flightless adults and though they occupy similar habitats, differ in their tolerances to
environmental conditions. Thus, what is an impermeable barrier to one taxon is
crossed with difficulty by some others, and with ease by still others. Given a geo-
graphical situation such as one encounters in northern Mexico and southwestern
United States, one can expect to find a general concordance of taxon/area patterns,
but not one that is precise.

Nonetheless, I am intrigued by and appreciate the elegance of Liebherr’s analysis
that is based on principles of cladistic biogeography. As well, I concede that a cladistic
analysis of the taxa of Liodicaelus may, after all, support the pattern that he has
postulated.

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Fig. 18. Geographical distribution of Dicaelus l. laevipennis LeConte.

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Genus Badister Clairville

Specimens of Badister are rare in collections, primarily because they are small,
rather inconspicuous, and most of them live in habitats so wet that they are avoided
by most collectors. Nonetheless, I have seen many specimens during the past years,
and identifying them has given sufficient opportunity to test the delimitation of taxa
recognized in my treatment of the genus (Ball, 1959:191-241). Further, Lindroth
(1969:955-967) confirmed implicitly my species-level treatment, though he differed
with some of my interpretations of supraspecific relationships and classification. Such
differences require comment, and some new information has become available that
seems worthy of reporting, as detailed below. Also, I take this opportunity to offer
a more detailed treatment of microsculpture for several species.

In classifying the North American species of Badister (Ball, 1959:190-191), I
recognized three subgenera: Badister ( sensu stricto), Trimorphus Stephens and Baudia
Ragusa. These taxa were characterized primarily by details of mandibular structure
and tarsal setation: right mandible with dorsal surface notched, and tarsomere 5 with
a row of setae on each ventro-lateral margin ( Badister ); right mandible with dorsal
surface notched, tarsomere 5 with ventro-lateral margins asetose ( Trimorphus ); and
left mandible notched, tarsomere 5 with ventro-lateral margins asetose {Baudia). For
classification of the species of subgenus Badister, I used the ranks of complex and
group, as follows: bipustulatus complex— bipustulatus group (5 species), and neopul-
chellus group (5 species); pulchellus complex— pulchellus group (including B. pul-
chellus LeConte, only), and elegans group (5 species).

Lindroth (1969:955-967), who preferred not to use a formal subgeneric rank,
recognized four species groups: notatus group, including only B. notatus Haldeman,
that I included in subgenus Badister, the pulchellus group, for the other species of
subgenus Badister: ; the transversus group, for the single Nearctic species {B. trans-
versus ) that I included in subgenus Trimorphus ; and the reflexus group for most of
the species of Baudia. He did not recognize formally any ranks between group and
subgenus.

Lindroth’s assignment of B. notatus to a group equal in rank to a subgenus, was
based on the several striking features that are unique in the genus, such as reduction
of microsculpture, reduction of wings and loss of ability to fly, and development of
convex elytral intervals. However, B. notatus exhibits the derived feature of large
spines in the internal sac, which is characteristic of the elegans group. Thus, Lindroth
classified B. notatus on the basis of phenetic distinctness, whereas I classified it on
the basis of what is interpreted to be a shared derived feature; in other words, I used
phylogenetic principles.

Lindroth (1969:962) accepted my judgement that B. transversus was isolated from
the other subgenera of Badister, but he did not agree that it was related to the type
species of Trimorphus {B. sodalis Duftschmid), although both species are similar in
mandibular notching, and both have tarsomere 5 asetose. Noting the similarity
between members of Baudia and of B. transversus in reduced tarsal setation and
transverse pronotal microsculpture, Lindroth (1969:956) implied that the latter spe-
cies was a Baudia derivative, whose mandibles exhibited reverse asymmetry (i.e., the
right mandible notched, rather than the left). He concluded the argument by suggesting
that the difference in which one of the mandibles of a pair was notched was the result

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Table 8. Notching of mandibles among West Indian population samples of Badister reflexus
LeConte.

Island(s)

Left

Notched mandible

Right

Total

Bahamas

4

0

4

Cuba

4

2

6

Jamaica

2

0

2

Cayman Islands

1

10

11

Hispaniola

Haiti

1

4

5

Dom. Republic

3

3

6

Total

15

19

34

of a very simple genetic difference. If this is so, mandibular notching cannot be
regarded as a reliable character for grouping the species of Badister. This matter is
addressed below, following description of the clypeus, labrum and mandibles of
Badister.

Labrum, clypeus and mandibles of Badister (Figs. 19A-H and 20A-F)

The mouthparts of Badister adults are modified strikingly, particularly the labrum,
clypeus and mandibles. The clypeus (Fig. 20A-B, cl) is narrow, sharply bent in a
dorso-ventral plane, with two prominent lateral setae. The labrum (1) is deeply,
angularly notched, with six marginal dorsal setae.

Structure of the mandibles (Figs. 19A-H, 20C-F).— This pair of sclerites is typical
of the Licinina in being short and broad, lacking molar teeth, retinaculum and occlusal
grooves. Strikingly asymmetrical, one of the pair has a deep dorsal notch (Figs. 1 9 A,
B, E, and 20E-F), and ventrally (Fig. 19G-H) the ventral groove of the notched
mandible is markedly curved.

For the subgenus Baudia Ragusa, notching of the left mandible is normal (Fig.
1 9 A-B, E-G, n) the notch being located between an anterior much enlarged terebral
tooth (tt) and posterior basal boss (b). The occlusal margin in dorso-lateral aspect
(Fig. 19B-G) is markedly bent. The right mandible (Fig. 19C, D, F, H) is without
the deep dorsal notch, and without the basal boss. For the subgenus Badister ( sensu
stricto ), the right mandible is notched (as in Fig. 20E-F), with enlarged terebral tooth
and basal boss, and the left mandible is without notch or projections.

Most known species of Badister are constant in which mandible is notched— either
right or left, but not both. The West Indian populations of Badister reflexus LeConte
are exceptional, in that either mandible of a pair is notched, as illustrated in Figures
19 and 20. This species has been placed in the subgenus Baudia Ragusa (Ball, 1959:
221), a group characterized by having the left mandible notched.

Implications for classification.— The data for Badister reflexus seem to support
Lindroth’s opinion that the notching pattern of the mandibles is determined by a
simple genetic change. I doubt that such a character by itself is likely to distinguish
monophyletic lineages. Thus, it seems best either to abandon the subgenera as they

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Fig. 19A-H. SEM photographs of the mandibles of Badister reflexus LeConte: A, B, E, and
G, left mandible, lateral, dorsal, occlusal and ventral aspects, respectively; C, D, F and H,
dorsal, lateral, occlusal, and ventral aspects, respectively. Legend: b, basal boss; br, basal ridge;
i, incisor; n, notch in dorsal surface; tt, terebral tooth; and vg, ventral groove. Scale bars =
100 ^m.

are diagnosed at present, or to acknowledge that each is likely to be a polyphyletic
assemblage and to use it because of tradition or convenience.

Microsculpture of Badister (Fig. 21 A- H)

Elytral microsculpture in the bipustulatus complex.— Most species of Badister with
which I am familiar either lack elytral microsculpture ( B . notatus Haldeman) or have
markedly transverse and closely packed meshes ( B . neopulchellus Lindroth, Fig. 2 1 A-
B), with very narrow sculpticells (s), and an iridescent luster— a condition designated
as “grated” by Allen and Ball (1980:486). An exception is Badister obtusus LeConte
(Figs. 21C-D), with the mesh pattern slightly transverse, sculpticells wider, and
surface rather dull— lacking iridescence. There is an ecological correlate to this dif-
ference: B. neopulchellus is markedly hygrophilous, whereas B. notatus and B. obtusus
occupy the forest floor in damp to dry areas, but not in association with standing
water.

It is accepted generally that isodiametric microsculpture (Fig. 2 1 E-F) is plesiotypic,
and any other conditions are more or less apotypic, in a transformation series, with
the markedly transverse grated condition marking the far end. I had assumed that
the series was essentially irreversible. If this were so, then B. obtusus should occupy
a relatively primitive position. However, with Badister obtusus being part of a derived
clade in a lineage nearly all members of which exhibit grated microsculpture (Ball,
1 959:233, Diagram 3), it seems unlikely that this species can be regarded as ancestral
to the lineage as a whole. Accordingly, I conclude that the non-grated microsculpture
exhibited by B. obtusus is derived from the grated condition, and that this derivation

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Fig. 20 A-F. SEM photographs of anterior part of head and mandibles of Badister grandiceps
Casey and B. reflexus LeConte. A-B, B. grandiceps : A, head, labrum and mandibles, dorsal
aspect; B, same, frontal aspect. C-F, mandibles of B. reflexus : C-D, left mandible, lateral and
dorsal aspects, respectively; and E-F, right mandible— dorsal and lateral aspects, respectively.
Legend: b, basal boss; cl, clypeus; i, incisor; 1, labrum; n, notch; sop, supraorbital setigerous
puncture; tt, terebral tooth. Scale bars = 100 gm.

represents an evolutionary reversal. In turn, this reversal is associated with life in
an environment different from the environment occupied by most species of Badister.

Microsculpture of the head in the reflexus group. — Most species of Badister exhibit
isodiametric sculpture on the dorsal surface of the head capsule. Adults of B. ocularis
exhibit such sculpture (Fig. 21 E-F; the left supraorbital setigerous puncture [sop] is
shown, for orientation). A related species, B. grandiceps Casey, exhibits sculpture on
the vertex that is slightly transverse (Fig. 21G-H). Although the difference is not
much, and is not easily perceived even with relatively high power of a dissecting
microscope, it is sufficient to cause structural color, so that the vertex of a typical B.
grandiceps adult has a rather indistinct band of purple coloring on the vertex.

Geographical distribution of Badister

Ball (1959:1 90) reported Badister to be represented on every continent except South
America. Recently, I saw a specimen with the character states of Baudia from Tu-
cuman, Argentina, collected in January, 1979 by R. Goldbach (CNCI). Also, I par-
ticipated in collecting a series of specimens with the characteristics of subgenus
Badister in Amazonian Peru. These species will be studied and described, not nec-
essarily by me, at some future date.

Fig. 2 1 A-H. SEM photographs of microsculpture on various sclerites of members of Badister
Clairville: A-B, interval 3, left elytron, of B. neopulchellus Lindroth; C-D, same of B. obtusus
LeConte; E-F, vertex of head of B. ocularis Casey; G-H, same of B. grandiceps Casey. Legend:
1-2, intemeur 2; 1-3, intemeur 3; s, sculpticell; sop, supraorbital setigerous puncture. Scale bars:
A-D, F, H = 20 ^m; E, G = 100 ^m.

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Notes and new distribution records

Provided here for selected species are observations about variation, and about
geographical distribution, including range extensions. The species sequence is the
same as in Ball (1959:189-228).

Badister neopulchellus Lindroth

This is a wide-ranging species in North America, from the boreal forest in the
north southward to Georgia in the east, and Texas, in the west. A record not previously
published, extending the range of B. neopulchellus southward in the west, is based
on a single female: U.S.A. CALIFORNIA San Bernardino County. Mountain Pass,
IV. 1 1.1942; O. Bryant (CASC).

Badister obtusus LeConte

Basically northern and wide-ranging in the boreal forest, B. obtusus is distributed
southward only at higher altitudes in the Rocky Mountains and outlying ranges. New
records are the following.

CANADA. NORTHWEST TERRITORIES. 2 females, Fort Wrigley, IX.27. 1929;
O. Bryant (CASC).

U.S.A. NEW MEXICO. San Miguel County. Sangre de Cristo Mountains, 3.2 km
W Gascon, Rio de Gascon, 2,438 m, VIII.8.1963; G. E. & K. E. Ball (UASM).

TEXAS. Culberson County. Male, Guadalupe Mts., Nat. Park, Bocil, 2,407 m,
IX. 13. 1988; R. S. Anderson (UASM).

Badister elegans LeConte

This species was known previously from the Great Plains states, southward to
Brownsville, Texas. The following records establish its presence in northern Mexico.

COAHUILA. Female, 31.2 km E San Pedro, V.31.1981; J. Doyen (UCBC).

TAMAULIPAS. Male, 3 females, 15.9 km W La Pesca, at U-V light, X.28.1965;
G. E. Ball & D. R. Whitehead (UASM).

Badister vandykei Ball

The description of this species was based on a single male, collected in Baja
California. The following record establishes occurrence of this species on the adjacent
mainland of Mexico.

SINALOA. Male, female, 41.8 km N Perico, VIII. 13. 1960; P. H. Amaud, E. S.
Ross & D. C. Rentz (CASC).

Badister flavipes mexicanus Van Dyke

The description of this subspecies was based on a single female collected at Oaxaca
City, in southern Mexico. Because the other subspecies were northern (Ball, 1959:
21 1-213), ranging as far south as Texas on the mainland and the Bahamas in the
West Indies (Darlington, 1953), it was not unexpected that B. f. mexicanus would
occur in much of the intervening area, and thus would be wide-ranging in Mexico.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

The following records confirm this expectation. Mainly eastern in distribution, the
altitudinal range of B. f mexicanus extends from near sea level to about 1,500 m.
Specimens were collected in sites ranging from mesic to wet (i.e., palm savanna).
During daylight hours, specimens were found under cover (principally leaf litter).
Some were collected at U-V light at night, indicating nocturnal flight activity.

CHIAPAS. 4 females, Aguatenango, pasture-lake (under cow droppings, near wa-
ter), VI.26. 1972; G. E. Ball & P. E. Meyer (UASM). Male, Palenque ruins, U-V light,
91.4 m., V. 22. 1972; G. E. Ball, K E. Ball & P. E. Meyer (UASM).

MORELOS. 4 females, Santa Rosa, 5.1 km N Zacatepec, 975 m, 11.31.1966; M.
W. McFadden & D. R. Whitehead (UASM). 4 males, female, 8.7 km E Cuernavaca,
pedregal, 1,420 m, U-V light, V.29-30. 1966; Ball & Whitehead (UASM), Male,
Tejalpa, VI.23.1963; Whitehead (UASM).

NUEVO LEON. Female, Santa Rosa Canyon, 23.8 km W Linares, 732 m, U-V
light, VII. 6-7. 1966; Ball & Whitehead (UASM).

OAXACA. Female, Oaxaca (city), XI. 1968; G. Pollard (UASM). Female, Pueblo
Nuevo, VIII. 1-1 2. 1986; H. & A. Howden (UASM).

SAN LUIS POTOSI. 2 females, El Salto Falls, U-V light, VII.7.1966; R. E. Wood-
ruff (FSCA).

PUEBLA. Male, Puente Ahuehueyo, ca. 10 km W jet. Hwys 1 15 & 190, 1,280 m,
VIII. 10. 1965; Ball & Whitehead (UASM).

TABASCO. Male, female, S. Ciudad Pemex, 53 m, U-V light, V.22.1972; Ball,
Ball & Meyer (UASM).

TAMAULIPAS. Male, 2 females, Sierra de Tamaulipas, Hwy. 180, 76.6 km N
Aldama, ca. 1 50 m, palm savanna, XII. 8. 1978; Ball & Ball (UASM). 2 males, Gomez
Farias, Naciemiento del Rio Frio, VI.29.1969; S. & J. Peck (UASM).

Badister reflexus LeConte

Geographical variation.— As noted above, either the left or right mandible of adults
of this species is notched. On the mainland (Yucatan Peninsula to southern Ontario)
and in the Bahamas archipelago in the West Indies, only the left mandible is notched.
In the Greater Antilles and Cayman Islands, either the left or right mandible is
notched, as indicated in Table 8. A notched right mandible is predominant in His-
paniola and the Cayman Islands, while a notched left mandible is predominant in
Cuba and Jamaica. However, the sample sizes are too small to be confident that the
suggested pattern reflects reality.

Geographical distribution and localities.— This species was known to be wide-
ranging in eastern United States and adjacent parts of southern Canada, from Ontario
to “Texas” (no specific locality), and eastward to the Bahamas and Greater Antilles.
The following list documents occurrence of B. reflexus in Texas, the West Indies and
Mexico.

U.S.A. TEXAS. Cameron County. Female, Brownsville, X.42; E. S. Ross (CASC).

MEXICO. QUINTANA ROO. 4 males, 5 females, 5 km W Chetumal, Hwy. 186,
mercury vapor & U-V light, V.20.1987; E. G & T. J. Riley (LSUC).

WEST INDIES. BAHAMAS. 2 males, Rum Cay, nr. Port Nelson, III. 16. 1953;
Van Voast (AMNH). Male, female, Cat Island, Arthur’s Town, VII.29.1935; W. J.
Clench (MCZC).

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CAYMAN ISLANDS. 5 males, 5 females, Grand Cayman, South Sound, light trap
B, IV.17-VIII.26.1938; C. B. Lewis & G. H. Thompson (MCZC).

CUBA. Female, Baragua, XI. 12.1925; G. F. Stahl (MCZC). 2 males, 2 females,
Cienfuegos, Soledad, IV-X.1926, 1936; P. J. Darlington, Jr. (MCZC). Female, Imias
Camaguey, 8-1966, F. de Zayas (MCZC).

HISPANIOLA. Dominican Republic. 3 males, 2 females, Sanchez, VIII. 1938; P.
J. Darlington, Jr. (MCZC). Male, 2 females, Puerto Plata, VIII.29-IX.2.1938; P. J.
Darlington, Jr. (MCZC). Haiti. 3 females, swamps N. Dessalines, IX. 1 1.1934; P. J.
Darlington, Jr. (MCZC).

JAMAICA. Male, Clarendon Parish, Portland Ridge, nr. Jackson Bay Cave, 12
m, V.4.1973; D. & M. Davis (USNM). Female, Good Hope, VIII. 1 1. 1966; H. F.
Howden (CNCI).

Badister parviceps Ball

The original description of this species (Ball, 1959:225-226) was based on material
from the central and northeastern states, and from eastern Ontario. Lindroth (1969:
967) reported additional localities in Manitoba and eastern British Columbia, and
suggested that B. parviceps probably is transcontinental in distribution. This species
is less hygrophilous than the other members of the reflexus group, occurring in leaf
litter in mesic, shaded sites that are not in direct contact with standing water. Below
are noted localities from farther south, extending the known range of this species to
eastern Texas.

U.S.A. TEXAS. Brazos County. Female, Koppe’s Bridge, ca. 8 km SW college
Station, berlese, III. 10. 1988; R. S. Anderson (UASM). Male, same as above, but with
note: riparian ravine, leaf litter, berlese (UASM). Houston County. Male, female, Big
Slough Wild Area, FR 517 & Four C’s hiking trail, bottomland, hardwood, litter,
berlese, V.9.1988; Anderson (UASM). Tyler County. Male, female, 4.5 km W, 3 km
N, Spurger Beech Wood Trail, litter, berlese, IV.24. 1988; Anderson (UASM). Walker
County. Female, Huntsville State Park, pine-hardwood forest, litter, berlese,
IV. 14. 1988; Anderson (UASM).

Badister submarinus Motschulsky

I doubted that this species, known only from two females ostensively collected at
Sacramento, California, was a North American resident, imagining instead that the
specimens had been collected elsewhere and had been mislabelled. Lacking a male
for study, I included this species in subgenus Baudia, incertae sedis, because the
classification of the species of this group is based on male characters (Ball, 1959:
228). About eight years ago, I found in material for identification sent from the
Zoological Museum at Copenhagen (ZMUC), a third specimen (also a female) of B.
submarinus, labelled: “Oregon, U.S.A.”; ‘'‘Upper Klamath Lake at Algoma, Klamath
Co. Oregon”; “4100 ft. 11.X.1949 E. Suenson.” This record confirms that B. sub-
marinus is a western North American resident. However, it must remain incertae
sedis until a male is located and its genitalia studied.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

ACKNOWLEDGMENTS

I am grateful to J. K. Liebherr for undertaking to organize the Festschrift in honor of J. G.
Franclemont, and for inviting me to prepare a contribution to it.

A major part of this paper is the set of plates of photographs taken with the Scanning Electron
Microscope. They, and the structures illustrated, were prepared in a cooperative operation by
G. D. Braybrook, D. Shpeley, and J. S. Scott, who are technologists in my Department. Mr.
Scott prepared also the habitus photograph of Dicaelus franclemonti, and final versions of the
maps and other figures.

G. Scherer, of the Zoologische Staatssammlung, Miinchen, made me welcome during a visit
to his institution, and arranged for the loan of the type series of Pterostichus karikali. Y.
Bousquet, of what is now the Division of Biological Resources, and was formerly the Biosys-
tematics Research Centre, Ottawa, made it possible for me to examine the licinines in his care
during several visits to his institution. N. E. Stork, of The Natural History Museum, London,
responded with alacrity to my frantic request for last-minute assistance. David H. Kavanaugh,
of the California Academy of Sciences, San Francisco, arranged for the loan of an extensive
array of licinines that I had not seen previously. I appreciate very much the special assistance
of these colleagues, and as well, the cooperation of the other curators and collections managers
whose names appear in the list of institutions from the collections of which study material was
received.

As I learned so well from my association with Jack Franclemont, much of the pleasure of
systematic entomology is obtained through life in the field, where one hunts out and assembles
the specimens and knowledge that make possible all of the subsequent descriptions and analyses.
The pleasure that I enjoyed in finding the Liodicaelus material in the mountains of northern
Mexico and southwestern United States was enhanced by association with the late D. R.
Whitehead, and D. Shpeley, H. E. Frania, S. McCleve and W. Fraga. Not only did I enjoy their
company, but I appreciated as well the specimens that they found. My wife’s name appears on
many of the specimen labels, and I am pleased to acknowledge her contributions, also.

REFERENCES

Acorn, J. H. and G. E. Ball. 1991. The mandibles of some adult ground beetles: structure,
function and the evolution of herbivory. Can. J. Zool. 69:638-650.

Allen, R. T. 1 983. Distribution patterns among arthropods of the North Temperate Deciduous
Forest biota. Ann. Missouri Bot. Gard. 70:616-628.

Allen, R. T. and G. E. Ball. 1980 (1979). Synopsis of Mexican taxa of the Loxandrus series
(Coleoptera: Carabidae: Pterostichini). Trans. Am. Entomol. Soc. 105:481-576.

Ax, P. 1987. The Phylogenetic System: The Systematization of Organisms on the Basis of
Their Phylogenesis. John Wiley and Sons, Chichester.

Ball, G. E. 1959. A taxonomic study of the North American Licinini with notes on the Old
World species of the genus Diplocheila Brulle (Coleoptera). Mem. Am. Entomol. Soc.
No. 16, 258 pp., 15 plates.

Ball, G. E. 1 966a. The taxonomy of the subgenus Scaphinotus Dejean with particular reference
to the subspecies of Scaphinotus petersi Roeschke (Coleoptera: Carabidae: Cychrini).
Trans. Am. Entomol. Soc. 92:687-722.

Ball, G. E. 1966b. The Chinese species Diplocheila minima Jedlicka: a redescription and
observations on its relationships (Col.: Carabidae). Entomol. News 77:19-25.

Bates, H. W. 1891. Additions to the carabideous fauna of Mexico, with remarks on some of
the species previously recorded. Trans. Roy. Entomol. Soc. London 1 89 1 :223-278, plates
13, 14.

Bell, R. T. 1960. A revision of the genus Chlaenius Bonelli (Coleoptera: Carabidae) in North
America. Misc. Publ. Entomol. Soc. Am. 1:97-171.

1992

THE TRIBE LICININI

379

Coope, G. R. 1973. Tibetan species of dung beetle from Late Pleistocene deposits in England.
Nature 245:335-336.

Coope, G. R. 1979. Cenozoic fossil Coleoptera: evolution, biogeography and ecology. Ann.
Rev. Entomol. 10:247-267.

Darlington, P. J., Jr. 1961. Australian carabid beetles V. Transition of wet forest faunas from
New Guinea to Tasmania. Psyche 68:1-24.

Darlington, P. J., Jr. 1968. The carabid beetles of New Guinea. Part III. Harpalinae (Contin-
ued): Perigonini to Pseudomorphini. Bull. Mus. Comp. Zool. 137:1-253.

El Moursy, A. A. and G. E. Ball. 1959. A study of the diagnostic characters of two North
American species of the genus Diplocheila Brulle, with notes on their ecology. Coleopts.
Bull. 13:47-57.

Emden, F. I., van. 1942. A key to the genera of larval Carabidae. Trans. Am. Entomol. Soc.
92:1-99.

Erichson, W. F. 1 842. Beitrag zur Fauna von Vandiemensland, mit besonderer Riicksicht auf
die geographische Verbreitung der Insecten. Archiv fur Naturgeschichte 8:83-287.

Erwin, T. L. 1 974. Studies of the subtribe Tachyina (Coleoptera: Carabidae: Bembidiini), Part
II: a revision of the New World-Australian genus Pericompsus LeConte. Smithsonian
Contr. Zool. No. 162:iv + 96 pp.

Erwin, T. L. 1985. The taxon pulse: a general pattern of lineage radiation and extinction
among carabid beetles. Pages 437-493 in: G. E. Ball (ed.), Taxonomy, Phylogeny and
Zoogeography of Beetles and Ants: A Volume Dedicated to the Memory of Philip Jackson
Darlington, Jr. (1904-1983). Dr W. Junk Publishers, Dordrecht.

Freude, H. 1976. 1. Familie: Carabidae (Laufkafer). Pages 7-302 in: H. Freude, K. W. Harde
and G. A. Lohse (eds.), Die Kafer Mitteleuropas. Band 2, Adephaga 1. Goecke and Evers,
Krefeld.

Hammond, P., A. Morgan and A. V. Morgan. 1979. On the gibbula Group of Anotylus and
fossil occurrences of Anotylus gibbulus (Staphylinidae). Syst. Entomol. 4:215-221.

Harde, K. W. 1984. A Field Guide in Colour to Beetles. Octopus Books, London.

Horn, G. H. 1881. On the genera of Carabidae with special reference to the fauna of Boreal
America. Trans. Am. Entomol. Soc. 9:91-196, plates 1-10.

Jeannel, R. 1941. Coleopteres Carabiques, premiere partie, pp. 1-571, figs. 1-213. Faune de
France, 39. Paul Lechevalier et fils, Paris.

Jeannel, R. 1942. Ibid., 2me partie, pp. 573-1173, figs. 214-368. Ibid., 40. Ibid.

Jedlicka, A. 1931. Neue Caribiden aus Sud-China. Acta Soc. Entomol. Cechoslov. 28:102-
108.

Jedlicka, A. 1969. Neue Carabiden aus der Zoologischen Sammlung des Beyerischen Staats-
Museum (Coleoptera, Carabidae). Opusc. Zool. Miinchen 108:1-7.

Kryzhanovskij, O. L. 1983. Fauna of the USSR. Coleoptera. Vol. 1, No. 2. New Series, No.
128. Beetles of the Suborder Adephaga: Families Rhysodidae, Trachypachidae, Family
Carabidae [introductory part]. Science Press, Leningrad Division, pp. 1-341. [English
translation, by Multilingual Services Division, Translation Bureau, Agriculture Canada].

Laferte-Senectere, Marquis F. Thibault de. 1851. Revision de la Tribus Patellimanes. Ann.
Soc. Entomol. France (2me series) 9:209-294.

Liebherr, J. K. 1986. Cladistic analysis of North American Platynini and revision of the
Agonum extensicolle Group (Coleoptera: Carabidae). University of California Publi-
cations in Entomology 106:X + 198 pp.

Liebherr, J. K. 1991a. Phylogeny and revision of the Anchomenus clade: the genera Tetra-
leucus, Anchomenus, Sericoda, and Elliptoleus. Bull. Am. Mus. Nat. Hist. 202:1-163.

Liebherr, J. K. 1991b. A general area cladogram for Montane Mexico based on distributions
in the Platynine genera Elliptoleus and Calathus (Coleoptera: Carabidae). Proc. Entomol.
Soc. Wash. 93:390-406.

380

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Lindroth, C. H. 1968. The ground-beetles (Carabidae excl. Cicindelinae) of Canada and
Alaska. Part 5. Opusc. Entomol. Suppl. 33, pp. 649-944, figs. 329-480.

Lindroth, C. H. 1969. The ground-beetles (Carabidae excl. Cicindelinae) of Canada and
Alaska. Part 6. Opusc. Entomol. Suppl. 34, pp. 945-1192, figs. 481-557.

Lindroth, C. H. 1986. The Carabidae (Coleoptera) of Fennoscandia and Denmark. Fauna
Entomol. Scand. 15:233-497.

Matthews, J. V., Jr. 1979. Tertiary and Quaternary environments: historical background for
an analysis of the Canadian insect fauna. Pages 31-86 in: H. V. Danks (ed.), Canada
and Its Insect Fauna. Mem. Entomol. Soc. Canada, No. 108, 573 pp.

Moore, B. P. 1965. Studies on Australian Carabidae. 4— the Pterostichinae. Trans. Roy.
Entomol. Soc. London 117:1-32.

Moore, B. P. 1966. Australian larval Carabidae of the subfamilies Harpalinae, Licininae,
Odacanthinae and Pentagonicinae. Proc. Linn. Soc. New South Wales 90:157-163.

Moore, B. P. 1979. [Chapter] 2.24. Chemical defense and its bearing on phylogeny. Pages
193-203 in: T. L. Erwin, G. E. Ball, D. R. Whitehead and A. L. Halpem (eds.), Carabid
Beetles: Their Evolution, Natural History and Classification. Dr W. Junk b.v., Publishers,
The Hague.

Moore, B. P., T. A. Weir and J. E. Pyle. 1987. Carabidae. Pages 23-320 in: B. W. Wolton
(ed.), Zoological Catalogue of Australia, Volume 4, Coleoptera: Archostemata, Myx-
ophaga and Adephaga. Australian Government Publishing Service, Canberra.

Noonan, G. R. 1988. Faunal relationships between eastern North America and Europe, as
shown by insects. Pages 39-54 in: J. A. Downes and D. H. Kavanaugh (eds.). Origins
of the North American Insect Fauna. Mem. Entomol. Soc. Canada No. 144, 168 pp.

Serrano, J. 1986. A karyotypical approach to carabid evolution. Pages 221-234 in: P. J. den
Boer, M. L. Luff, D. Mossakowski and F. Weber (eds.), Carabid Beetles: Their Adap-
tations and Dynamics. Gustav Fischer, Stuttgart.

Serrano, J., J. Galian and A. Ortiz. 1986. A chromosome study of thirteen species of Spanish
Carabidae (Coleoptera, Adephaga). Fol. Biol. (Krakow) 34:319-328.

Serrano, J. and S. Yadav. 1984. Chromosome number and sex determination mechanism in
adephagan Coleoptera. Coleopts. Bull. 38:335-357.

Sloane, T. G. 1898. On Carabidae from West Australia sent by Mr. A. M. Lea (with descrip-
tions of new genera and species, synoptic tables, etc.). Proc. Linn. Soc. New South Wales
23:444-520.

Solier, A. J. J. 1849. Orden III. Coleopteros. Pages 105-380, and 414-511 in: C. Gay (ed.),
Historia Fisica y Politica de Chile Segun Documentos Adquiridos en Esta Republica
.... Volume 4 Published privately, Paris and Museo de historia natural de Santiago,
Santiago.

Spence, J. R. 1982. Taxonomic status, relationships, and biogeography of Anatrichis LeConte
and Oodinus Motschulsky (Carabidae: Oodini). Coleopts. Bull. 36:567-580.

Stork, N. E. 1980. A scanning electron microscope study of tarsal adhesive setae in the
Coleoptera. Zool. J. Linn. Soc. 68:173-306.

Straneo, S. L. and G. E. Ball. 1989. Synopsis of the genera and subgenera of the Tribe Peleciini,
and revision of the Neotropical and Oriental species. Insecta Mundi 3:73-179.

Vitt, D. H. 1 982. Peatland formation and vegetation of Wagner Bog. The Edmonton Naturalist
10:17-22. (Special issue: the Wagner Bog.)

Received 2 December 1991; accepted 3 December 1991.

J. New York Entomol. Soc. 100(2):38 1-398, 1992

TAXONOMY AND DISTRIBUTION OF
THE ATHETINE GENUS LYPOGLOSSA FENYES
(COLEOPTERA: ST APH YLINID AE: ALEOCHARINAE) IN
NORTH AMERICA, WITH DESCRIPTION OF A

NEW SPECIES

E. Richard Hoebeke

Department of Entomology, Comstock Hall, Cornell University,

Ithaca, New York 14853-0999

Abstract. — The athetine genus Lypoglossa Fenyes is reviewed and redescribed. Two North
American species are treated, of which one is described as new: franclemonti from the boreal
region of northern North America. The following new specific synonymy is proposed: fenyesi
Bemhauer 1907 ( =angularis Maklin, 1853). Lectotypes are designated for Lypoglossa fenyesi
(Bernhauer) and the Palearctic species L. lateralis (Mannerheim).

Adult mouthpart structures are illustrated for the first time for the genus; species distributional
data are recorded and mapped; all bionomic data are presented; habitus, male aedeagus, female
spermatheca, and other diagnostic characters are illustrated with the aid of line drawings and
scanning electron photomicrographs; and a key to species is provided.

The genus Lypoglossa was proposed by Fenyes in 1 9 1 8 to contain the single species
Dasyglossa fenyesi Bemhauer (1907:402) described from the Pacific Northwest. Dur-
ing the next half century, no important contributions were provided on the systematics
of this little known genus of athetine Aleocharinae. In 1978, C. H. Seevers, in his
tribal and generic revision of North American aleocharine rove beetles, gave little
information on this taxon and its characters, other than to include it in a key to
genera (p. 46). Our knowledge of the genus was further amplified in 1985, when
Lohse and Smetana, while studying numerous types of species of oxypodine and
athetine staphylinids described by Mannerheim and Maklin from northwestern North
America, noted that Myrmedonia angularis Maklin (1853:1 8 1) belonged to the genus
Lypoglossa ; in their paper they formalized this new combination, thus adding a second
species of this genus to our faunal list. In their discussion, Lohse and Smetana gave
a brief diagnosis to differentiate L. angularis and L. fenyesi, based on characters of
the antenna, color and shape of the pronotum, and male and female genitalia. They
also synonymized the Palearctic genus Megacrotona Scheerpeltz (1968:159), includ-
ing the single species M. lateralis (Mannerheim), with Lypoglossa.

The present paper resulted in part from my request to examine unidentified spec-
imens of Staphylinidae in the collection of the University of Vermont, in particular
the aleocharine rove beetles. To my surprise, included in this material were specimens
of a large, distinctive athetine from the high elevations of southern Vermont which
at first I was unable to name but later tentatively identified as belonging to the genus
Lypoglossa. Other borrowed material, from collections listed in Acknowledgments,
included specimens from high elevations of northern New York and New Hampshire,
a bog in Maine in the East, and numerous localities in western North America. In

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

this paper, I provide a detailed redescription and diagnosis of the genus Lypoglossa
with illustrations of adult habitus, mouthparts, and other structural characters, new
synonymy, lectotype designations, description of a new species, and a key to aid in
the identification of the North American species.

Genus Lypoglossa Fenyes

Lypoglossa Fenyes, 1918:23; 1920:239; Blackwelder, 1952:228; Moore and Legner,
1 975:448, Seevers, 1 978:46, 1 34. Type species: Dasyglossa Bernhauer, 1907:
402, by monotypy.

Megacrotona Scheerpeltz, 1 968: 1 59 (as new subgenus of Atheta ); Lohse and Smetana,
1 985:294 (as junior synonym of Lypoglossa ). Type species: Oxypoda lateralis Man-
nerheim, 1830:484, by monotypy.

Diagnosis. Members of Lypoglossa share a striking overall resemblance to Devia
prospera Erichson (Oxypodini), and to some large specimens of Oxypoda (Oxypodini),
but differ principally by the 4,5,5 tarsal formula (5,5,5 in Devia prospera and Oxypoda
spp.) and the unique punctation and pubescence of the dorsal abdominal segments
(Fig. 5) in which terga III-V are densely, finely punctured, with short, silky, dense,
appressed microsetae, whereas terga VI-VIII are very sparsely punctured, with sparse
microsetae (uniform punctation and microsetae on dorsal abdominal segments in
Devia prospera and Oxypoda spp.). Lypoglossa can be further characterized by the
following combination of characters: relatively large (3. 3-4. 5 mm), fusiform in shape;
pronotum exceptionally broad, 0.69-0.79 x wider than long; pronotal pubescence
along midline directed cephalad, remaining microsetae swirling laterad from midline
(Pattern E of Seevers, 1978) (Fig. 5); hypomera incompletely visible from lateral
view; mesocoxae narrowly separated; mesosternal process long, slender, acute at apex
(Fig. 8); mesosternal process: isthmus: metastemal process ratio, 12:5:1; tarsal for-
mula 4,5,5.

Redescription. Moderate to large sized, length of North American specimens 3.3-
4.5 mm. Body shape elongate, fusiform (Fig. 5). Body color testaceous to reddish-
brown, except head and abdomen dark reddish-brown to piceous. Body microsculp-
ture imbricate, dense and prominent on head, pronotum, and elytra (Fig. 9); that on
abdominal segments transversely striate, consisting of obsolete, interconnected, wavy
microlines; cuticular surface dull to slightly glossy; integument moderately to densely
pubescent, microsetae appressed; macrosetae present, conspicuous on abdomen;
punctures moderately dense throughout, very fine to slightly asperate, especially on
basal three abdominal segments.

Head more or less transverse, slightly wider than long (L/W ratio = 0.84), broadest
across temples just posterior to eyes, basal angles broadly rounded, neck absent. Eye
size small, 0.40 x length of head. Tempora moderate, subequal to length of eye,
broadly rounded. Dorsal pubescence prominent, directed more or less medially on
sides, and anteriorly along midline. Infraorbital ridge conspicuous posteriorly, absent
anteriorly. Antenna moderately elongate, about as long as, or slightly longer than
head and pronotum combined, slightly incrassate towards apex; articles 1-4 more
or less elongate, article 3 longer than 2, articles 5-10 becoming more gradually
transverse and increasing in width to apical articles; article 1 1 conical apically, about
as long as articles 9+10.

3 4

Figs. \-A. Lypoglossa franclemonti, adult mouthpart structures. 1, Labrum, dorsal aspect;
2, Maxilla, dorsal aspect; 3, Left mandible, dorsal aspect; 4, Labium, ventral aspect. (Scale line
= 1.0 mm)

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Labrum transverse (Fig. 1). Right and left mandibles with similar dentition (Fig.
3), without internal teeth, apices acute, entire; molar region slightly developed, with
short row of spinose teeth on inner edge; membranous prostheca finely serrated
medially. Maxilla (Fig. 2) with galea slightly longer than lacinia; galea densely pu-
bescent in apical 0.4; lacinia with comb of single row of 6-7 large recurved teeth
apically, and dense patch of recurved setae dorso-medially; maxillary palps 4-seg-
mented. Labium as in Figure 4; ligula moderately short, deeply cleft nearly to base
into 2 bluntly rounded, divergent lobes; 2 medial setae, broadly separated at base;
prementum with numerous pseudopores medially plus posterior to medial setae,
several pseudopores, 2 large real pores, and 1 spinose pore laterally on each side;
labial palps 3 -segmented, chaetotaxy of articles as in Figure 4; mentum and chaetotaxy
as in Figure 4.

Pronotum broadly transverse, about 0.69-0.79 x wider than long; anterolateral
margins obtusely rounded; sides broadly, evenly arcuate; posterior margin ranging
from broadly arcuate to distinctly bisinuate; pubescence with microsetae directed
cephalad in a narrow median strip, with remaining microsetae directed laterad from
median row in swirling pattern as in Figure 5 (Pattern E, sensu Seevers, 1978, or
Type I, sensu Hoeg, 1945).

Elytra relatively short, slightly longer than pronotum, parallel-sided; outer apical
angles strongly sinuate; pubescence dense, microsetae arranged in slightly sinuate
pattern, directed more or less posteriorly, as in Figure 5. Mesostemum with narrow
mesostemal process (Fig. 8), elongate, apically pointed, extending to slightly beyond
middle of coxal cavities; metastemal process short, broadly rounded; isthmus rela-
tively long; coxae narrowly separated, acetabulae finely margined posteriorly. Me-
sostemal process: isthmus: metastemal process ratio 12:5: 1. Legs slender, long; tarsal
formula 4,5,5; hind tarsus elongate, tarsomeres 1-4 decreasing in length.

Abdomen broad at base, elongate, tapered from basal segments to apex. Terga III—
VI slightly, transversely impressed basally; basal impressions impunctate, smooth.
Terga III-V dull, densely punctured, with short, silky, appressed microsetae. Terga
VI-VIII less dull, very sparsely punctured and pubescent.

Secondary sexual characteristics. Male: Dorsum of head, and broad to narrow
median area of pronotum moderately impressed (as in Fig. 6). Female: Dorsum of
head evenly convex; pronotum evenly convex with slight narrow impression along
median line in some specimens (as in Fig. 5). [The head and pronota of specimens
of the type series of the Palearctic species L. lateralis (Mannerheim), including both
sexes, are evenly convex dorsally without any apparent median impression.]

Distribution. At the present time, the known Nearctic species have a transconti-
nental, northern distribution, with relict populations in the mountains or high ele-
vations of more southern areas. The only known Palearctic species is also widely
distributed in the high elevations of northern and middle Europe, and Siberia (Scheer-
peltz, 1968; Benick and Lohse, 1974).

Bionomics. Limited distributional and ecological evidence would suggest that
members of Lypoglossa are probably restricted to the boreal forest zone of North
America (and Eurasia), a coniferous forest with a floor cover of mosses or of low
herbs and shrubs, interspersed with extensive lakes (Rowe, 1972), which extends in
a continuous belt from Newfoundland and the Labrador coast in the east, west to
the Rocky Mountains and then northwards to the mouth of the Mackenzie River

Figs. 5-9. Lypoglossa spp.: 5-6 & 8-9, L. franclemonti. 5, Dorsal habitus; 6, Pronotum,
dorsal aspect, male; 8, Meso- and metastemum; 9, Elytral microsculpture. 7, L. angularis,
Pronotum, dorsal aspect, female.

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and the central areas of Alaska (with relictual areas in the mountains of Arizona,
New Mexico, and Colorado, and high elevations of eastern New York through New
England) (Scudder, 1979). The principal tree species of the boreal forest include
White and Black Spruce, various pine and hr species, and White Birch, Trembling
Aspen and Balsam Poplar (Scudder, 1979).

Remarks. Lypoglossa, as now known, consists of 2 widely distributed boreal species
in North America and 1 species in the Palearctic region.

Having amassed a considerable number of specimens of both identified and un-
identified North American material from various collections, I assumed these spec-
imens, segregated into 2 distinct, but similar morphospecies, would be readily as-
signable to the two available North American names. However, this was not the
case. After careful examination of the types of Lypoglossa angularis (Maklin) and L.
fenyesi (Bemhauer), I discovered that they are conspecihc, thus necessitating the new
synonymy stated herein, and the description of a new species.

Most specimens of Lypoglossa in collections, if determined, are identified as L.
fenyesi, thought originally to be the only included species of the genus. The new
species, franclemonti, described herein, is often included in identified series of L.
fenyesi. Specimens of both species, however, can be separated readily by the shape
of the pronotum, and by key differences in the male genitalia.

KEY TO NORTH AMERICAN LYPOGLOSSA

1. Pronotum less transverse, broadly oval in shape (L/W ratio = 0.75-0.79), posterior
margin usually evenly arcuate, with posterior angles broadly rounded (as in Figs. 5-

6) , pronotal color testaceous to light reddish-brown, usually strongly contrasting with
darker color of head and elytra; median lobe of aedeagus and apical lobe of paramerite

as in Figs. 10-12 L. franclemonti, n. sp.

1'. Pronotum more transverse, rectangular in shape (L/W ratio = 0.69-0.71), posterior
margin usually distinctly bisinuate, with posterior angles strongly obtuse (as in Fig.

7) , pronotal color dark reddish-brown, usually concolorous with elytra, or only slightly
contrasting; median lobe of aedeagus and apical lobe of paramerite as in Figs. 15-17
L. angularis (Maklin)

Lypoglossa franclemonti, new species
Figs. 1-6, 8-14; Map 1

Lypoglossa fenyesi: sensu Lohse and Smetana, 1985:294 (not Bemhauer).

Diagnosis. Lypoglossa franclemonti is very similar externally to L. angularis, but
can be readily separated by the more oval, less transversely-shaped pronotum with
broadly arcuate posterior margin and evenly rounded posterior angles (Figs. 5-6),
by the reddish-yellow pronotum which is usually contrasting with the head and elytra,
and by the characteristic shape of the median lobe of the aedeagus (Figs. 10-1 1) and
the apical lobe of the paramerite (Fig. 1 2).

Description. Length 3. 3-4. 5 mm. Body color testaceous to yellowish brown, except
head, areas of elytra adjacent to scutellum and elytral suture, sometimes outer pos-
terior angles of elytra, and abdominal segments darker, reddish-brown to piceous;
in most specimens terga VI-VI1I with bluish-black iridescent sheen.

Head obsoletely imbricate with moderately dense, irregular, wide, transverse sculp-

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387

ticells, surface slightly glossy; microsetae moderately short, somewhat dense, ap-
pressed; punctures fine to very slightly asperate. Antenna elongate and slender, with
articles 1-3 elongate, article 1 slightly incrassate, article 2 more slender, slightly
shorter than article 3; article 4 distinctly longer than wide; articles 5-10 decreasing
in length and becoming more quadrate and transverse; article 1 1 subequal to articles
9 + 10.

Pronotum less transverse, tending to be somewhat oval-quadrate (L/W ratio =
0.75-0.79); posterior margin broadly arcuate, or very obsoletely bisinuate in some
specimens; posterior angles broadly rounded. Integument markedly imbricate, mod-
erately glossy; pubescence with microsetae moderately short, dense, appressed, di-
rected cephalad in narrow median strip, with remaining microsetae swirling laterad
from median strip; punctures moderately fine, very slightly asperate.

Elytra slightly narrower than maximum width of pronotum; microsculpture densely
imbricate (Fig. 9); pubescence with microsetae short, dense, appressed, directed pos-
teriorly in a slightly sinuate pattern; punctures dense, uniform, slightly asperate.

Abdominal terga III-V with moderately dense pubescence; microsetae very fine,
short, appressed; punctures fine, dense. Tergite VI with sparse, fine punctures; terga
VII-VIII with punctures even more sparse than that of tergite VI; microsetae of terga
VI-VIII moderately short, silky, fine, appressed; abdominal integument dull, with
fine, transverse microsculpture, appearing (in most specimens) iridescent (especially
terga VI-VII).

Male: Median lobe of aedeagus as in Figures 10-11. Paramere and apical lobe of
paramerite as in Figure 12.

Female: Spermatheca as in Figures 13-14.

Secondary sexual characteristics. Male: Dorsum of head, and broad area along
median line of pronotum impressed (Fig. 6). Female: Dorsum of head and pronotum
evenly convex (in most specimens pronotum may be narrowly impressed along
median line) (Fig. 5).

Types. Holotype, male, with labels: Mt. Whiteface, N.Y. [Essex Co.], 2,000-4,000
ft, 22 Aug. 1 9 1 6/HOLOTYPE Lypoglossa franclemonti <3, design. 1991, E. R. Hoe-
beke/HOLOTYPE Cornell U. No. 6510. Allotype, female, with labels: Artist’s Brook,
Essex Co., N.Y., June 1 1, 1933, H. Dietrich/ALLOTYPE Lypoglossa franclemonti
$, design. 1991, E. R. Hoebeke/ALLOTYPE Cornell U. No. 6510. Both holotype
and allotype are deposited in the Cornell University Insect Collection, Ithaca, N.Y.

Paratypes, 1 82, are deposited in the CNCI, MCZC, FMNH, CUIC, SEMC, UVCC,
and DENH.

CANADA: Alberta: 8 mi W Calgary, on 1A, 18-20-VI-1980 (CUIC, 2). George
Lake, ca. 53°57'N, 114°06'W, 1-VI, 12-VI, 16-VI, 2- VIII, 10-VIII-1980, JS Ashe
(SEMC, 16); George Lake, 53°57'N, 114°06'W, l-X-1966, 10-IX-1966, 25-IX-1966,
RE Leech (CNCI, 5). Lusk Creek, Kananaskis F.E.S., 14-VII-1971, JM & BA Camp-
bell (CNCI, 4). Manitoba: Riding Mtn. N.P., Moon Lake, 17-IX-1979, A Smetana
(CNCI, 6); nr Deep Lk., 7-IX-1979, A Smetana (CNCI, 4); Whirlpool Rv. at Hwy
19, 6-IX-1979, A Smetana (CNCI, 1). Northwest Territories: Hwy 3, 5 mi SE Ft.
Simpson, 2 1-VI- 1972, A Smetana (CNCI, 1). Nova Scotia: Cape Breton H.N.P., 410
m, Benjies Lake area, 18-IX-1984, JM Campbell & A Davies (CNCI, 1); 440 m, nr
Sammy’s Barren, 24. IX. 1984, JM Campbell & A Davies (CNCI, 1). Ontario: Lk.
Superior Prov. Pk., Garanyua, 9-VI-1973, JM Campbell & R Parry (CNCI, 10);

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Figs. 10-14. Lypoglossa franclemonti. 10, Median lobe of aedeagus, lateral aspect; 1 1, Me-
dian lobe of aedeagus, dorsal aspect; 12, Paramere; 13, Spermatheca (Bolton, Vermont); 14,
Spermatheca (Mt. Marcy, New York). (Scale line = 0.25 mm)

Frater, 9-1 3-VI- 1973, JM Campbell & R Parry (CNCI, 5); Old Woman Bay, 13-VI-
1973, Campbell & Parry (CNCI, 3); Agawa Bay, 1 3-VI- 1973, Campbell & Parry
(CNCI, 6); Baldhead Lk., 8-VI-1973, JM Campbell & R Parry (CNCI, 1). 46 mi N
Hurkett, Black Sturgeon Lk., 28-29-VI-1973, R Parry & JM Campbell (CNCI, 2).
49 mi N Savant Lake, 23-VI-1973, Campbell & Parry (CNCI, 1). Moosenoe, 30-VI-
1973, 2-VII-1973, Parry & Campbell (CNCI, 7). 21 mi & 41 mi N Pickle Lake, 19-
VI- 1973, Campbell & Parry (CNCI, 2). 36 mi S Pickle Lake, 22-VI-1973, Campbell
& Parry (CNCI, 5). Quebec: Mistassini Post, 1 3-VI- 1956, JR Lansway (CNCI, 1).
Parc des Laurentides, Mare-du-Sault, 2,700 ft, 1 5-1 7-VIII- 1 970, JM & BA Campbell
(CNCI, 2). Gaspesie Parc, Lac St. Anne, 12-VII-1972, 1,500 ft, JM & BA Campbell
(CNCI, 1); Lac du Diable, 12-VII-1972, 1,500 ft, JM & BA Campbell (CNCI, 2);

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Lac Cascapedia, 14-VII-1972, 1,700 ft, JM & BA Campbell (CNCI, 1); Mont Albert,
9-VII-1972, 1,000 ft, JM & BA Campbell (CNCI, 4).

UNITED STATES: Maine: Washington Co., Carrying Place Cove, Lubec, 23-VII/

24- VIII-1989, H. Frambs (CUIC, 22). New Hampshire: Coos Co., Crawford Notch,

25- IX-1908 (MCZC, 3); Bretton Woods (FMNH, 1); 1 mi NE East Inlet Dam, 27-
IX/17-X-1986, 26-IX-1986, DS Chandler (DENH, 2); Norton Pool, 2 mi E East
Inlet Dam, 7-IX-1984, DS Chandler (DENH, 2); White Mts., GlenBoulder Trail,
2,500 ft, 17-VIII-1976, JM & BA Campbell (CNCI, 1); Glen Ellis Falls, 1,900 ft, 15-
VIII- 1976, JM & BA Campbell (CNCI, 1). New York: Essex Co., Mt. Whiteface, 21-
X-1934, CR Crosby (CUIC, 3); Mt. Whiteface, 25-VIII-1921 (CUIC, 1); Top Mt.
Marcy, 22-X-1936, H Dietrich (CUIC, 1); Top of Mt. Whiteface, 22-X-1936, H
Dietrich (CUIC, 2); Essex Co. (no locality), 10-VII-1918 (CUIC, 4); Top Mt. Me-

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Intyre, 1 -VII- 1923, CR Crosby (CUIC, 1); Artist’s Brook, 1 1 -VI- 1933, H Dietrich
(CUIC, 1). Vermont: Chittenden Co., Camel’s Hump, Bolton, 3,800 ft, 7-VII-1972,
RT Bell (U VCC, 1 1 ); same locality, 9- 1 9-VIII- 1 972 (UVCC, 4); same locality, 1 -VIII-
1972, J Spence (UVCC, 6); same locality, VIII-IX-1972 (UVCC, 6); same locality,
24-VII-1972, 3790 ft (UVCC, 3); same locality, 18-VII-1973, 3,400 ft (UVCC, 2);
same locality, 18-VII-1973, 3,800 ft (UVCC, 4); same locality, 17-VII-1972, 3,800
ft (UVCC, 6); same locality, 23-VIII-1975, Langworthy (UVCC, 1); Mt. Mansfield,
4,000 ft, 1 l-X-1969, HP Wimmer (UVCC, 1).

Etymology ;. This species is named in honor of Dr. John G. Franclemont, emeritus
professor, Cornell University, who not only has made significant contributions to
our knowledge of North American Lepidoptera, but also has been responsible for
the training of many practicing North American insect systematists. In recognition
of his eightieth birthday, it is with great pleasure that I name this new species after
“Jack,” my academic advisor as a graduate student, and now, a dear friend and
colleague.

Geographic distribution. The known range of L. franclemonti includes localities in
Alberta, Northwest Territories, Manitoba, western Ontario, Quebec, and Nova Sco-
tia, with relict populations in the high elevations of eastern New York, Vermont,
and New Hampshire, and a bog in Maine (Map 1).

Bionomics. Little is known about the habits of this species. It appears to prefer
rather moist, rich, organic habitats. Most western specimens studied were taken from
sifted litter in birch, birch-liedum, and black spruce bogs; from sifted leaf litter; and
from sifted moss and lichens on rotten logs. A few additional eastern specimens bear
the following habitat data: “pit traps” in spruce forest at 3,400-3,800 ft, “flight
intercept trap,” and “sift conifer logs.” Most specimens examined were taken in June,
July, and August, but also in September and October.

Remarks. Externally the types of L. angularis and L. fenyesi are identical, and
detailed study of the genitalia of males from the syntype series of both supports the
synonymy proposed below. The taxon “ L. fenyesi” referred to in Lohse and Smetana
(1985, under “Discussion,” p. 294) is actually L. franclemonti. Apparently, Lohse
and Smetana studied only the type of L. angularis; had they also examined the type
series of L. fenyesi they would have discovered, as I did, that the types of both are
conspecific and thus would have known that the other “widespread” species of
Lypoglossa occurring in North America actually represented an undescribed species.
The male and female genitalia figured in Lohse and Smetana (1985) as that of L.
fenyesi (cf. Figs. 24-25, p. 295) is really that of L. franclemonti.

Lypoglossa angularis (Maklin)

Figs. 7, 15-18; Map 2

Myrmedonia angularis Maklin, 1853:181.

Lectotype male: UNITED STATES: Alaska: “Woskres.” (=Woskersensk, Kenai
peninsula) (UZMH). Specimen with labels: Coll. Maklin; Myrmedonia angularis
Makl. Woskres.; Dasyglossa prospera Er. det. Bernhauer; Mus. Zool. H. fors Spec,
typ. No. 2270 Myrmedonia angularis Makl.; Myremedonia angularis Maklin
Lectotypus Lohse design. 1983. Type specimen studied. Genitalia dissected and
mounted.

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ATHETINE GENUS LYPOGLOSSA

391

Dasyglossa fenyesi Bemhauer, 1907:402; Leng, 1920:126. New Synonymy.

Lectotype male: CANADA: British Columbia (FMNH). Specimen with labels:
Em[era]ld Lake, BC, 0.6.6.26, DrAFenyes; 330.; fenyesi Brh. Typus.; Chicago NHMus,
M. Bemhauer Collection; LECTOTYPE Dasyglossa fenyesi Bernhauer, design. 1991,
E. R. Hoebeke. One paralectotype, male, with labels: Banff, Alta, 0.6.6.24, DrAFenyes;
313.; 103. (in red ink); fenyesi Brh. Cotypus.; Chicago NHMus, M. Bernhauer Col-
lection; PARALECTOTYPE Dasyglossa fenyesi Bemhauer, design. 199 1 , E. R. Hoe-
beke (FMNH). Type material studied and lectotype here designated. Genitalia of
lectotype dissected and mounted.

Lypoglossa fenyesi: Fenyes, 1920:239, t. 5, f. 2; Bernhauer and Scheerpeltz, 1926:

597; Hatch, 1957:145; Moore and Legner, 1975:448; Seevers, 1978:271.
Lypoglossa angularis: Lohse and Smetana, 1985:281, 293.

Diagnosis. Lypoglossa angularis is practically indistinguishable externally from L.
franclemonti. Adults, however, differ from those of the latter species by having a
more transverse pronotum with rather pronounced, obtuse posterior angles (Fig. 7).
Moreover, the pronotal color is usually darker in L. angularis, and less contrasting
with the head and elytra. The shape of the median lobe of the aedeagus (Figs. 1 5-
1 6), and the apical lobe of the paramerite (Fig. 1 7) are the best diagnostic characters
for identification of this species.

Redescription. Length 3. 7-4. 2 mm. Body color generally dark, reddish-brown,
except mouthparts, antennae, legs, and oblique, indistinct band across each elytron
often paler. Head and abdomen generally darker, piceous. Abdominal terga VI-VII
usually with bluish-black iridescent sheen.

Head punctation, setation, and microsculpture similar to that in L. franclemonti.
Antennae slightly shorter, more robust than in L. franclemonti', articles 1-3 elongate;
article 4 slightly longer than or subequal to its width; articles 5-7 short, quadrate;
articles 8-10 tending to be more transverse; article 11 conical, pointed apically,
subequal to articles 9+10.

Pronotum strongly transverse (L/W ratio = 0.69-0.7 1), posterior margin distinctly
bisinuate, posterior angles broadly obtuse (Fig. 7). Integument distinctly imbricate,
moderately glossy; pubescence with microsetae as in L. franclemonti.

Elytra and abdomen as in L. franclemonti.

Male: Median lobe of aedeagus as in Figures 15-16. Paramere and apical lobe of
paramerite as in Figure 17.

Female: Spermatheca as in Figure 18.

Secondary sexual characteristics. Male: Dorsum of head, and narrow median strip
of pronotum impressed. Female: Dorsum of head and pronotum usually evenly
convex (sometimes narrow, median line finely impressed).

Geographic distribution. Lypoglossa angularis is widely distributed in northwestern
North America (Alberta, British Columbia, Yukon Territory, Manitoba, and Alaska),
with relict populations on the higher mountains in the Gaspe, eastern Quebec, and
northern New Hampshire, and in the high elevations of northern California, Oregon,
Washington, Colorado, New Mexico, Arizona, Wyoming, and Montana (Map 2).

Material examined. (1,087 specimens): CANADA: Alberta: Banff, 6-VI-1924 (CASC,
6); Banff, Dr A Fenyes (FMNH, 1). Banff N.P., Boom Lk., 6,000 ft, 16-VII-1971,

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17

Figs. 15-18. Lypoglossa angularis. 15, Median lobe ofaedeagus, lateral aspect; 16, Median
lobe of aedeagus, dorsal aspect; 17, Paramere; 18, Spermatheca (Glacier, British Columbia).
(Scale line = 0.25 mm)

JM & BA Campbell (CNCI, 1 1); Bow Lk., 16-VI-1968, 6,500 ft, Campbell & Smetana
(CNCI, 4); Agnes Lk., 7,000 ft, 17-VII-1971, JM Campbell (CNCI, 2); Consolation
Lk., 64-7,000 ft, 15-VI-68, Campbell & Smetana (CNCI, 26); Cirrus Mt., 5,500 ft,
16-VI-1968, Campbell & Smetana (CNCI, 1); Smith Lk., 15-VII-1971, JM & BA
Campbell (CNCI, 1); Mt. Temple Ski Lodge, 6,600 ft, 12-VIII-1971, JM Campbell
(CNCI, 1). Jasper N.P., Mt. Edith, Cavell, 6,000 ft, 22-VIII-1971, JM Campbell
(CNCI, 1). Kananaskis F.E.S., Lusk Creek, 14-VII-1971, JM & BA Campbell (CNCI,
31); 2.5 mi SE Kananaskis, F.E.S., 1 5- VIII- 1 970, EE Lindquist (CNCI, 2); 10 mi
SW Kananaskis F.E.S., Marmot Creek Basin, 6,500 ft, 1 3-VII- 1971, JM & BA Camp-
bell (CNCI, 1 9). Waterton Lks. N.P., Crypt Lake, 6,500 ft, 22-VI- 1 980, JM Campbell

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(CNCI, 1); Crypt Lk. Tr., 4,900 ft, 23-VI-1980, JM Campbell (CNCI, 1); Lower
Bertha Falls, 4,800 ft, 10-VI-1980, JM Campbell (CNCI, 3); Bertha Creek, 49-5,800
ft, 10-VI-1980, JM Campbell (CNCI, 5); Summit-Carthew Lks. Tr., 7,000 ft, 17-VI-
1980, JM Campbell (CNCI, 3). Lower Waterton Lk., 4,500 ft, 26-VI-1980, JM
Campbell (CNCI, 1); Cameron Cr., 5,200 ft, 24-VI-1980, JM Campbell (CNCI, 7);
Cameron Lk., 8-9-VI-1980, 9-14- VI-1980, 17-19-VI-1980, 5,450-5,500 ft, JM
Campbell (CNCI, 19); Cameron Lake, 5,500 ft, 31-VII-1980, DE Bright (CNCI, 1);
Cameron Lake, 7.VI.1980, 8-1 1 -VI- 1980, 23-VI-1980, IM Smith (CNCI, 4); Cam-
eron Ck. picnic ground, 1 2-1 6-VI- 1980, 17-28-VI-1980, 5,350 ft, IM Smith (CNCI,
29); Rowe Lks. Tr., 5,300-6,500 ft, 7-1 5-VI- 1980, JM Campbell (CNCI, 48); Mother
Duck Tr. above Cameron Lk., 17-VI-1980, IM Smith (CNCI, 9); Mother Duck Tr.,
58-6,200 ft, 17-VI-1980, JM Campbell (CNCI, 1 1); Crandell Lk., 19-VI-1980, 5,000

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ft, JM Campbell (CNCI, 1 1); Crandell Lk. Tr., 13-VI-1980, IM Smith (CNCI, 1 1);
Little Prairie picn. area, 23-VI-1980, I Smith (CNCI, 2); Rowe Creek, 5,300-5,500
ft, 7-VI-1980, 2 l-VI-1980, 4-VIII-1976, JM Campbell (CNCI, 14); Rowe Cr. Tr.,
60-6,300 ft, 1 5-VI- 1 980, JM Campbell (CNCI, 4); Rowe Trail, 6,400 ft, 1 6-VI- 1 980,
JM Campbell (CNCI, 4). British Columbia: Em[era]!d Lake, 6-VI- 1927 (CASC, 2).
Glacier, Dr. A Fenyes (CASC, MCZC, FMNH, AMNH, 23). Manning Prov. Pk.,
Mt. Frosty, 6,100 ft, 30-VII-1975, JM & BA Campbell (CNCI, 1); Blackwell Pk.,
6,000-6,750 ft, VI-20- 1968, Campbell & Smetana (CNCI, 26); Blackwell Peak, 3,000
ft, V-3 1-1968, Campbell & Smetana (CNCI, 6). 8 mi W, 21 mi W Creston, 10-VI-
1968, 9-VI-1968, Campbell & Smetana (CNCI, 3). Slide Mt., 10 mi E Barkerville,

4.500 ft, 23-24-VI- 1 968, Campbell & Smetana (CNCI, 25). Kootenay N.P., Kimpton
Cr., 4,000 ft, 21 -VII- 1971, JM & BA Campbell (CNCI, 7); Sinclair Cr., 6,000 ft, 12-

VI- 1968, Campbell & Smetana (CNCI, 22). Yoho N.P., Lk. McArthur, 7,000 ft, 28-

VII- 1971, JM & BA Campbell (CNCI, 4); Valley of Hagen Peak, 6-VIII-1971, JM
& BA Campbell (CNCI, 1); Lk. O’Hara, 6,700 ft, 30-VII-1971, JM & BA Campbell
(CNCI, 6); Otterhead R., 5,500 ft, 3-7-VIII- 1 97 1 , JM Campbell (CNCI, 38); Amiskwi
R., 6,000 ft, 5- VIII- 1971, JM & BA Campbell (CNCI, 4). Mi 56 Haines Hwy., 3
Guardsmen Pass, 4-VII-1968, 3,200 ft, Campbell & Smetana (CNCI, 26); Haines
Hwy., km 143-144, 2,400 ft, 22-V-1978, Smetana & Becker (CNCI, 4). 5 mi E
Whistler’s Mt., 29-V-1968, Campbell & Smetana (CNCI, 28). Mt. Revelstoke N.P.,

6.500 ft, 1 8- VI II- 1971, JM & BA Campbell (CNCI, 1); Eva Lk., 6,500 ft, 25-VII-
1971, JM & BA Campbell (CNCI, 61); Mt. Revelstoke, 6,000 ft, 8-VII-1952, GP
Holland (CNCI, 2); Mt. Revelstoke, 6,300 ft, 18-VI-1968, Campbell & Smetana
(CNCI, 7). 7.6 km E Hwy 25, 6.2 km S Terrace, Copper Mt. Rd., 24-VIII-1983, JM
Campbell (CNCI, 1). 46 km W McBride, 26-VIII-1983, JM Campbell (CNCI, 1).
Two Sisters Mt., 15 mi E Barkerville, 4,500-6,500 ft, 23-24-VI- 1968, Campbell &
Smetana (CNCI, 16). Courtney, Forbidden Plateau, nr Courtney Lookout, 25-VII-
1979, IM Smith (CNCI, 4); Forbidden Plateau, McPhee Lake, 3,700 ft, 19-VII-1975,
JM & BA Campbell (CNCI, 1). Garibaldi Prov. Pk., Diamond Head Tr., 4,000-

4.500 ft, 26-VII-1973, A & Z & D Smetana (CNCI, 1); Opal Cone, 5,200 ft, 3-VIII-
1975, JM & BA Campbell (CNCI, 1). Atlin, 2,200 ft, 23-VI-1955, H Huckel (CNCI,
1). Summit Lake, Mi 392 Alaska Hwy., 4,200 ft, 23-VIII-1959, RE Leech (CNCI,
1). Queen Charlotte Isl., Moresby Isl., Mt. Moresby, 2,100 ft, 25-VII-1983, JM
Campbell (CNCI, 5). Manitoba: Riding Mt. N.P., 1 km SE Long Lake, 16-IX-1979,
A Smetana (CNCI, 1 ); Stathclair Trail, 4.5 km S Kinnis Crk., 1 2-IX- 1 979, A Smetana
(CNCI, 6). Quebec: Parc Gaspesie, Lac Cascapedia, 14-VII-1972, 1,700 ft, JM & BA
Campbell (CNCI, 1); Mont Albert, 8-VII-1972, 1,000 ft, JM & BA Campbell (CNCI,
3); Mont Albert, 10-1 l-VII-1972, 2,800-3,700 ft, JM Campbell (CNCI, 8); Mt.
Albert, 3,150 ft, 20-VI-1954, 2-VI-1954, GP Holland (CNCI, 9). Parc des Lauren-
tides, Lac Arthabaska, 2,800 ft, 21-22-VIII-1970, JM & BA Campbell (CNCI, 3);
Mare-du-Sault, 2,700 ft, 1 5-1 7-VIII- 1 970, JM & BA Campbell (CNCI, 4); Bois Verts,
2,800 ft, 18-VIII-1970, JM & BA Campbell (CNCI, 1). Mont Jacques, Cartier, 4,000
ft, 22-VII-1972, JM Campbell (CNCI, 4). Yukon Territory: Dempster Hwy., Mi 29.5,
2,800 ft, 24-VII-1978, JM Campbell & A Smetana (CNCI, 2); Mi 45, 3,500 ft, 14-
VII- 1968, Campbell & Smetana (CNCI, 4); Mi 42, N Klondike R., 3,300 ft, 18-VII-
1978, Smetana & Campbell (CNCI, 14); Mi 53, North Fork Pass, 4,200 ft, 24-VII-
1978, A Smetana & JM Campbell (CNCI, 1).

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ATHETINE GENUS LYPOGLOSSA

395

UNITED STATES: Alaska: 2.5 mi S Lituya Bay, Mount Blunt, 2,500 ft, VI/3-
IX- 1977, 4-VI- 1977,1 -VII- 1977, Dan Mann (AMNH, 70). 7. 5 mi NNW Dillingham,
Wood River Red Bluff, 59°8'30", 158°33'W, 18-VI-1980, L Herman (AMNH, 4).
Hagemeister Island, North shore at base of long sand spit, 24-VI-1980, 58°38'N,
161°04'30"W, L Herman (AMNH, 5). Kenai Pen., 3 mi SE Kenai, 7-VI-1978, Sme-
tana & Becker (CNCI, 1); Kalifonsky Bch., nr Kenai, 8-VI-1978, Smetana & Becker
(CNCI, 9); Clam Gulch, 6-VI-1978, Smetana & Becker (CNCI, 3); Hope, 12- VI-
1951, WJ Brown (CNCI, 1); Nimilchik, 24-VI-1951 (CNCI, 1); Kasilof R. at Hwy.
1, 12-VI-1978, Smetana & Becker (CNCI, 1); 8 mi SE Kasilof, 9-VI-1978, Smetana
& Becker (CNCI, 3); Anchor Riv. at Hwy, 450 ft, 4-5-VI-1978, Smetana & Becker
(CNCI, 34); Anchor R. Cpgd., 12 mi N Homer, 450 ft, 5-VI-1978, Smetana & Becker
(CNCI, 8). 1 2 mi N mi 78 Denali Hwy., Windy Cr., 1 3-VII- 1 978, Smetana & Camp-
bell (CNCI, 1). Denali St. Pk., Byers Crk. at Hwy. 1, 20-VI-1978, Smetana & Becker
(CNCI, 2); Byers Lk. Cpgd., 23-25-VI-1978, Smetana & Becker (CNCI, 49). Kenai
Mts., 9 mi N Seward, 1 4-VI- 1980, L Herman (AMNH, 7); 15 mi N Seward, 400 ft,
29-V-1978, Smetana & Becker (CNCI, 12); 16 mi N Seward, 500-600 ft, 26-V-1978,
Smetana & Becker (CNCI, 2); 23 mi N Seward, 600-900 ft, 28-V-1978, Smetana &
Becker (CNCI, 7); 22 mi N Seward, 600-800 ft, 27-V-1978, Smetana & Becker (CNCI,
30); Ptarmigan Ck. Cpgd., 500-600 ft, 26-V-1978, Smetana & Becker (CNCI, 9).
Kenai Mts. Crk., above Tern Lk. Cpgd., 850-900 ft, 13-VI-1978, Smetana & Becker
(CNCI, 52); same locality as preceding, except 850 ft, 16-VI- 1978 (CNCI, 3 ). Arizona:
Coconino Co., San Francisco Mtns., Mt. Agassiz, 3,200 m, 28-VII-1976, JM Camp-
bell (CNCI, 2); Snow Bowl, 2,800 m, 27-VII-1976, JM Campbell (CNCI, 1); San
Franc[isco] Mts., Mann (CASC, FMNH, 3). California: El Dorado Co., L. Tahoe,
Cascade Lk., 6,300 ft, 1 1 -VIII- 1969, A Smetana (CNCI, 1). Deer Park Inn, Dr. A
Fenyes (MCZC, 3). Colorado: Huerfano Co., 15 mi SSW LaVeta, Cucharas Creek,
9,400 ft, 1 6-VIII- 1982, L Herman (AMNH, 3). Grand Co., Rollins Pass, 1 1,000 ft,

5- VIII-1973, JM Campbell (CNCI, 1). Coal Bank Pass, 10,550 ft, 3 l-VII-1973, JM
Campbell (CNCI, 3). Montana: Glacier Co., Glacier N.P., Logan Pass, 6,600 ft, 11-
VII- 1971, JM & BA Campbell (CNCI, 7). New Hampshire: Coos Co., Mt. Washing-
ton, 3,800 ft, 1 6-VIII- 1 976, JM & BA Campbell (CNCI, 3); above Lake of the Clouds,

6- VII-19 14, CA Frost (CASC, 1); Mt. Washington] (CASC, 5). Randolph, 18-IX-
1909 (CNCI, 1). Starr Lake, 4,890 ft (CASC, 1). New Mexico : Lincoln Co., Sierra
Blanca, 1 1 ,500 ft, 1 8-VII- 1 969, A Smetana (CNCI, 5); Sierra Blanca Ski Area, 1 0,600
ft, 1 8-VII- 1969, A Smetana (CNCI, 2). Oregon: Deschutes Co., 12 mi SW Sisters,
FSR 1551, 4,400 ft, 23-VII-1979, JM & BA Campbell (CNCI, 1). Hood River Co.,
Mt. Hood, Cloud Cap Rd., Tilly Jane Crk., 4,100 ft, 30-VI-1974, A & D Smetana
(CNCI, 8). Klamath Co., Cold Spring Camp, 33 mi NW Klamath Falls, 5,800 ft, 22-
VI- 1 978, J Schuh, L & N Herman (AMNH, 3). Wasco Co., 19 mi WNW of Dufur,
Wampus Springs, 4,800 ft, 9-VII-1978, L & N Herman (AMNH, 1). Washington:
Clallam Co., Olympic N. P., 13 mi S Port Angeles, 4,000 ft, 1 l-VIII-1979, JM &
BA Campbell (CNCI, 1). Pierce Co., Mt. Ranier N.P., N Puyallup River, 3,700 ft,
10-VIII-1973, A & Z & D Smetana (CNCI, 2); Sunbeam Falls, 4,000 ft, 17-V-1968,
Campbell & Smetana (CNCI, 2). Whatcom Co., Mt. Baker, 4 mi N Silver Fir Camp,
4,000 ft, 1 6-VIII- 1975, JM & BA Campbell (CNCI, 7). Wyoming: Sheridan Co., 27
mi WSW Ranchester, Big Horn Mts., nr Prune Creek Campgd., South Tongue River,
7,700 ft, 25-26-VIII-1982, L Herman (AMNH, 17).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Bionomics. Very little is known about the habits of this species. Some Alberta and
British Columbia specimens were taken in sifted moss and liverworts, and from sifted
alder, Populus, willow, spruce, Douglas fir, and Lodgepole pine litter. Other specimens
bear the following habitat data: “human dung trap, mouse nests, gopher burrows,
pitfall edge meadow, sifting bear dung, river debris, under bark, sifting pile of squirrel
midden, dung pan traps,” and “horse dung.” Most specimens have been taken in
May, June, July, and August, and occasionally in September.

Remarks. Male and female genitalia of L. angularis are accurately figured in Lohse
and Smetana (1985, Figs. 19-21, p. 295). The above authors also mentioned the
presence of “ 1 specimen [of L. angularis ] from Newfoundland” in the Canadian
National Collection (CNCI). I did not examine any CNCI specimens of L. angularis
collected from Newfoundland, but the distributional record would not surprise me.

Lypoglossa lateralis (Mannerheim)

Oxypoda lateralis Mannerheim, 1830:70.

Megacrotona lateralis: Benick and Lohse, 1974:103.

Lypoglossa lateralis: Lohse & Smetana, 1985:294.

In the course of examining specimens for this taxonomic study of Lypoglossa, I
had the opportunity to examine type specimens (syntype series) of L. lateralis (Man-
nerheim), the only known Palearctic representative of this holarctic genus. As a result,
it seems appropriate and warranted to designate a lectotype here.

The original syntype series in the Mannerheim collection, Zoological Museum,
University of Helsinki, Finland, consists of 4 pinned specimens (see Silfverberg,
1988). I examined all specimens of the series. They are labeled as follows: Spec. No.
1: [red triangle] = Villnas, Finland (a manor some 30 km NW of Turku/ Abo)/“ Abo”/
“Coll. Mannh.”/“Spec. typ.”/“108”/[red square]/“Mus. Zool. H:fors Spec. typ. No.
211 Oxypoda lateralis Mann.” Spec. No. 2: [red triangle]/“Abo”/“Coll. Mannh.”/
“Spec. typ.”/“109”/[red square]/“Mus. Zool. H:fors Spec. typ. No. 212 Oxypoda
lateralis Mann.” Spec. No. 3: [red triangle]/“l 17.” Spec. No. 4: [red triangle] only.

Specimen No. 1 bearing the label “Spec. typ. No. 211” is hereby designated as
lectotype (sex undetermined, undissected) of Oxypoda lateralis; the label LECTO-
TYPE Oxypoda lateralis Mann., design. 1991, E. R. Hoebeke, is attached. Specimen
No. 4, male, bearing only a small [red triangle] label, was dissected and remounted
on a small cardboard placard; the aedeagus is mounted in a drop of Hoyer’s medium
on the card. The label PARALECTOTYPE Oxypoda lateralis Mann., design. 1991,
E. R. Hoebeke, is attached to Specimen Nos. 2-4 of the syntype series.

ACKNOWLEDGMENTS

Specimens used in this study were borrowed from several institutional and university col-
lections, and I gratefully acknowledge the assistance of the curators responsible. The abbrevi-
ations listed below refer to material mentioned in the text.

American Museum of Natural History, New York (AMNH), L. H. Herman
California Academy of Sciences, San Francisco (CASC), D. H. Kavanaugh
Canadian National Collection, Ottawa (CNCI), J. M. Campbell
Cornell University Insect Collection, Ithaca, NY (CUIC)

1992

ATHETINE GENUS LYPOGLOSSA

397

Field Museum of Natural History, Chicago (FMNH), A. F. Newton, Jr.

Museum of Comparative Zoology, Cambridge, MA (MCZC), D. G. Furth

Snow Entomological Museum, University of Kansas, Lawrence, KS (SEMC), J. S. Ashe

University of Vermont, Burlington, VT (UVCC), R. T. Bell

University of New Hampshire, Durham, NH (DENH), D. S. Chandler

Zoological Museum, University of Helsinki, Finland (UZMH), H. Silfverberg

I also appreciate the assistance given by Herbert Frambs, a visiting German scholar (Uni-
versity of Bremen) studying at Cornell University, who donated staphylinids to the Cornell
collection that included specimens of Lypoglossa collected from bogs in Maine. The loan of
type specimens was made possible by H. Silfverberg (Zoological Museum, University of Hel-
sinki, Finland) and A. F. Newton, Jr. (Field Museum of Natural History, Chicago), and special
thanks are due to these individuals. I acknowledge my colleagues James K. Liebherr and Quentin
D. Wheeler for helpful suggestions and for editing the manuscript, and J. Howard Frank (Uni-
versity of Florida, Gainesville) and J. S. Ashe (Snow Entomological Museum, University of
Kansas, Lawrence) also provided valuable criticism of the manuscript.

LITERATURE CITED

Benick, G. and G. A. Lohse. 1974. Aleocharinae, Tribus 14 (Callicerini). Pages 72-220 in:
H. Freude, K. W. Harde and G. A. Lohse (eds.). Die Kafer Mitteleuropas, 5.

Bernhauer, M. 1907. Neue Aleocharini aus Nordamerika. Deut. Ent. Zeits. 1907:381-406.

Bemhauer, M. and O. Scheerpeltz. 1926. Coleopterorum catalogus. Pars 82, Staphylinidae 6:
499-988.

Blackwelder, R. 1952. The generic names of the beetle family Staphylinidae, with an essay
on genotypy. U.S. Nat. Mus. Bull., no. 200. 483 pp.

Fenyes, A. 1918. Genera Insectorum, Coleoptera, fam. Staphylinidae, subfam. Aleocharinae.
fasc. 1 73A: 1—1 10.

Fenyes, A. 1920. General Insectorum, Coleoptera, fam. Staphylinidae, subfam. Aleocharinae.
fasc. 1736:111-414.

Hatch, M. H. 1957. The beetles of the Pacific Northwest. Part II: Staphyliniformia. Univ.
Wash. Publ. Biol. 16:1-384.

Hoeg, N. 1 945. Beitrag zur Systematik der Aleocharinen. Uber die Behaarung des Thorax bei
der Gattung Atheta Thomson. Entomol. Medd. 4:264-286.

Leng, C. W. 1 920. Catalogue of the Coleoptera of America, North of Mexico. John D. Sherman,
Mount Vernon. 470 pp.

Lohse, G. A. and A. Smetana. 1985. Revision of the types of species of Oxypodini and
Athetini (sensu Seevers) described by Mannerheim and Maklin from North America
(Coleoptera: Staphylinidae). Coleopt. Bull. 39:281-300.

Maklin, F. 1853. In: Mannerheim, C. Dritter Nachtrag zur Kafer-Fauna der Nord-Ameri-
kanischen Laender des Russischen Reiches. Bull. Soc. Imp. Nat. Moscou 26:95-273.

Mannerheim, C. G. 1830. Precis d’un nouvel arrangement de la famille brachelytres de l’ordre
des insectes coleopteres. Mem. Acad. Sci. St. Petersbourg 1:415-501.

Moore, I. and E. F. Legner. 1975. A catalogue of the Staphylinidae of America north of Mexico
(Coleoptera). Div. Agric. Sci., Univ. Cal., Spec. Publ. 3015:1-514.

Rowe, J. S. 1972. Forest regions of Canada. Dept. Environ. Can. For. Ser., Publ. 1 300, Ottawa.
172 pp.

Seevers, C. H. 1978. A generic and tribal revision of the North American Aleocharinae
(Coleoptera: Staphylinidae). Fieldiana: Zoology 71:1-289.

Scheerpeltz, O. 1968. Coleoptera-Staphylinidae. Catalogus Faunae Austriae, 15(fa): 1-279.

Scudder, G. G. E. 1979. Present patterns in the fauna and flora of Canada. Pages 87-179

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

in: H. V. Danks (ed.), Canada and its Insect Fauna. Mem. Entomol. Soc. Can. 108.
573 pp.

Silfverberg, H. 1988. Lists of the insect types in the Zoological Museum, University of Helsinki.
8. Coleoptera: Staphylinidae. Acta Entomol. Fenn. 52:15-40.

Received 15 July 1991; accepted 22 November 1991.

J. New York Entomol. Soc. 100(2):399— 414, 1992

THE NEW WORLD BLEPHARIDA GROUP, WITH A KEY TO
GENERA AND DESCRIPTION OF A NEW SPECIES
(COLEOPTERA: CHRY SOMELID AE)

David G. Furth

Department of Entomology, Museum of Comparative Zoology,

Harvard University, Cambridge, Massachusetts 02138

Abstract.— Using external and internal adult morphology, clarification of status and deter-
mination keys are provided for Blepharida Chevrolat and the eight New World genera either
related to it or placed near it in recent catalogues. The morphological reasons (e.g., pattern) for
maintaining Blepharida and Notozona Chevrolat as separate genera are discussed. The Afro-
tropical genus Blepharidina Bechyne is lowered to subgenus status within the genus Blepharida
in light of morphological similarities and differences with New World Blepharida. Blepharida
haitiensis Blake is transferred to Acrocyum Jacoby. Morphological evidence (e.g., metatibial
emargination, eye shape, and metafemoral spring shape) for placing the monotypic Euplec-
troscelis Crotch close to Blepharida is given. Blepharida franclemonti is described as a new
species known to date only from the Yucatan Peninsula of Mexico.

In the catalogue-like checklist of Leaf Beetle genera of the world followed by most
workers (Seeno and Wilcox, 1982), the Alticinae (Chrysomelidae) begin with a group
of three genera, formerly Galerucinae (Decarthrocerini of Laboissiere, 1937). This
group is followed by three groups of genera that are confused in keys and collections.
The first of these groups consists of three Neotropical genera ( Elithia Chapuis, Pro-
calus Clark— originally in Galerucinae, and Crimissa Stal). The second group consists
of 14 genera from around the world, including 7 found in the New World (Nearctic
and/or Neotropical): Blepharonycha Fall; Blepharida Chevrolat; Chrysogramma Ja-
coby; Acrocyum Jacoby; Notozona Chevrolat; Pseudorthygia Csiki; and Phydanis
Horn. The third group contains only two Afrotropical genera ( Diamphidia Ger-
staecker and Polyclada Chevrolat) which probably should be combined with the
previous group. As stated by Seeno and Wilcox (1982), these groups (family-groups)
are quite artificial and should not be considered as accepted groupings or tribal
arrangements. Unfortunately the separation of these family groups in the Seeno and
Wilcox (1982) checklist and their indication of nominate genera for most of these
groups is very misleading and has caused some authors to refer to these family-groups
incorrectly as tribes. In the present paper the author will attempt to clarify some
morphological relationships among the New World genera of the above-mentioned
family-groups surrounding the genus Blepharida.

Morphological relationships of most of these New World Blepharida relatives are
apparent in Scherer’s 1983 key to the Neotropical Alticinae (a translation of Scherer,
1962); Pseudorthygia and Phydanis, considered by Scherer to be Nearctic, were not
included; however, these two genera are included in Arnett’s (1971) keys to North
American Alticinae. The genus Euplectroscelis Crotch is not included in Arnett (1971)
or Scherer (1983), even though in Wilcox (1975) it is placed next to the Blepharida-
related genera. The relationships of some of these genera have been questioned by

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

several workers. Bechyne (1968) considered the genera of the second family-group
from Seeno and Wilcox (1982) to be the Blepharidini. He disagreed with Scherer’s
(1961) interpretation of the genus Blepharida as including the New World Blepharida
( sensu stricto) together with the Old World Calotheca Heyden and Blepharidella
Weise. Bechyne (1968) argued that chaetotaxy and other morphological details of
the clypeus and the tarsal claws were good reasons for leaving these all as separate
genera. Based on the presence of a transverse series of setiferous punctures along the
anterior border of the clypeus and the absence of a transverse depression in the
clypeus (forming a carina above), Bechyne (1968) claimed that only three species ( B .
rhois Forster, B. irrorata Chevrolat, and B. haitiensis Blake) formed the true Bleph-
arida and that all other New World Blepharida were congeneric with Notozona.

Bechyne (1968) then created a new genus ( Blepharidina ) to encompass all non-
New World Blepharida, i.e., Afrotropical and Madagascan, possessing bifid tarsal
claws, closed procoxal cavities, prostemum with a non-emarginate apex, and a pro-
epimeron separated at its lateral border by a deep groove from the edge of the
pronotum. Bechyne (1968) considered Blepharonycha to be a valid genus because,
although it shares the character of bifid tarsal claws with Blepharida and Notozona,
it differs by having open procoxal cavities. He also considered Acrocyum to be
congeneric with Chrysogramma which he wrote had appendiculate tarsal claws but
variable procoxal cavities.

New World species of Blepharida and Notozona are very difficult to separate using
the morphological characters used in traditional dichotomous keys. Baly (1865) sep-
arated these two genera using several characters which were not true for all species,
especially the subapical tooth on the dorsal edge of the metatibiae ( Blepharida ) and
the more swollen metafemora with a ventral tooth of Notozona. Chapuis (1875)
separated these genera using the maxillary palps; narrow and filiform in Blepharida
versus robust and claviform in Notozona. However, none of these or other single
morphological characters work for separation of all species in these two genera, as
pointed out by Jacoby (1885). Jacoby (1885, 1891) stated that because he could not
find any structural differences between these two genera, Notozona might be included
with Blepharida, but he still kept them as separate genera based on consistent color
pattern differences. Scherer (1983) acknowledges Bechyne’s (1968) treatment of
Blepharida, but offers his opinion that the true Blepharida includes one Nearctic
species [B. rhois] and many of the Afrotropical species. In his 1983 keys, Scherer
stated that it is too early for him to form an opinion about the Neotropical species,
but that many belong to Notozona. Although Scherer mentions several places in his
1983 keys the potential for confusing Blepharida and Notozona, he nevertheless does
separate and list them based on the deep transverse impression in the anterior part
of the frons. Most of Scherer’s (1983) characters for separating Blepharida and No-
tozona from other genera seem to be based on description of Afrotropical Blepharida.

METHODS

Specimens including types were borrowed from a variety of institutions (see Ac-
knowledgments). The metafemoral spring (a generic-level character) was studied for
all the genera concerned using the methods described in Furth (1989) (see also Furth,
1985, 1988). In addition, the female spermatheca and male aedeagus were studied

1992

NEW WORLD BLEPHARIDA GROUP

401

2

Figs. 1-3. 1. Blepharida mexicana metatibia (Length = L = 2.89 mm). 2. Notozona his-

trionica metafemur (L = 3.55 mm) and metatibia (L = 3.37 mm). 3. Elithia sp. metatibia (L
— 3.96 mm).

in more than one species of each genus whenever possible. These genitalic structures
were extracted after prolonged heating (not boiling) of entire specimens in water. All
dissections were made with Wild M 5 stereomicroscope, including a calibrated reticle
and camera lucida attachment. All photographs were made with a Wild M 400
Photomakroskop using Kodak TMAX 400 film.

Types of all the described species of New World Blepharida ( sensu lato ) and of
Notozona have been studied by the author. Examples of three of the five described
species each of Elithia and Crimissa and the two valid species (I. Askevold, personal
communication) of Procalus were also examined. In addition the following number
of species (of the known described species) were examined for this study: Blepharo-
nycha (1 of 1); Chrysogramma (4 of 5); Acrocyum (2 of 4); Euplectroscelis (1 of 1);
Pseudorthygia (1 of 2); and Phydanis (1 of 2).

KEY TO THE GENERA OF THE NEW WORLD BLEPHARIDA-GROUP

1 . Apex of metatibia with dorsal edges emarginate or flattened (Figs. 1 , 2); eyes oriented
dorso-mesally, elliptical-shaped, large (dorso-ventral length equal to or greater than
the dorsal interocular distance) (Fig. 4); metafemoral spring without recurve flange
from ventral lobe, with an acute basal angle of ventral lobe, and with a long dorsal
lobe extended arm (Fig. 8) [ Phydanis has the first but not the second and third of the

three characters listed above] 2

F. Apex of meta tibia not emarginate or flattened, but with continuous, morphologically
unchanged dorsal edges (Fig. 3); eyes round, not oriented dorsally towards each other,
small (diameter usually considerably less than interocular distance) (Figs. 5, 6, 7);
metafemoral spring with different morphology than Figure 8, most (except Acrocyum
and Pseudorthygia ) with basal angle of ventral lobe obtusely angled or with recurve
flange (Figs. 9, 10, 11, 12, 13, and 16— metafemoral spring morpho-groups #1 and

#3, respectively, see Furth, 1989) 5

2. Tarsal claws simple Euplectroscelis

2'. Tarsal claws appendiculate Phydanis

2". Tarsal claws bifid 3

3. Procoxal cavities open B/epharonycha

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Figs. 4-7. 4. Blepharida franclemonti face and eyes (interocular distance dorsally = 1.04

mm). 5. Procalus mutans face and eyes (interocular distance =1.18 mm). 6. Crimissa cruralis
face and eyes (interocular distance = 1.78 mm). 7. Chrysogramma pictipennis face and eyes
(interocular distance = 0.85 mm).

(Blepharida flavocostata keys to here, but has 6 thin, longitudinal stripes (costae) on
non-striate elytra)

3'. Procoxal cavities closed (even very narrowly) 4

4. Elytral pattern mottled (irregular/variable, sometimes asymmetrical; one species, en-
demic to Baja California [Mexico] has all black elytra) (Figs. 23, 24, 25), occasionally
with vague irregularly bordered, transverse bands or series of small/tiny light-colored
spots; metatibial apex emarginated with a preapical tooth or protuberance on outer
dorsal edge (Fig. 1); elytral punctures usually not striate (arranged in rows), but with
some exceptions (e.g., B. rhois ); clypeus (lower frons) often flat or only slightly im-
pressed transversely; metafemur without ventral subapical teeth; metatibia not strongly

curved Blepharida

4'. Elytral pattern with distinct (large/wide) transverse bands or spots, variable in size,
but never mottled (Figs. 26, 27, 28); metatibial apex with dorsum flattened, but not
emarginate and without preapical protuberance (Fig. 2); elytral punctures striate, some-
times appearing as double puncture rows, punctures fine and close together; clypeus
(lower frons) usually with a deep transverse impression; metafemur often with a sub-
apical, ventral tooth and with metatibia strongly curved, especially in males (Fig. 2)
Notozona

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12

Figs. 8-12. 8. Blepharida mexicana metafemoral spring (L = 1.23 mm). 9. Elithia sp.

metafemoral spring (L = 1.41 mm). 10. Procalus mutans metafemoral spring (L = 0.81 mm).
1 1. Crimissa cruralis metafemoral spring (L = 1.15 mm). 12. Pseudorthygia nigritarsis meta-
femoral spring (L = 0.41 mm).

5. Tarsal claws bifid 6

5'. Tarsal claws simple 7

5". Tarsal claws appendiculate 8

6. Procoxal cavities open; metafemoral spring with basal angle of ventral lobe somewhat
obtuse (ca. 100°) (Fig. 10); body smaller (size 6-10 mm) and elongate oval in shape
Procalus

6'. Procoxal cavities closed; metafemoral spring with basal angle of ventral lobe obtuse
(more than 120°) (Fig. 9); body larger (9-15 mm) and more spherical in shape . . . Elithia

7. Procoxal cavities closed; metafemoral spring with basal angle of ventral lobe obtuse

(Fig. 11); body larger (9-15 mm); elytral punctation confused; distributed in South
America north to Panama Crimissa

7'. Procoxal cavities open; metafemoral spring with basal angle of ventral lobe acute (Fig.

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1 2); body smaller (ca. 4 mm); elytral punctation striate; distributed in Mexico

Pseudorthygia

8. Metatibial apex with dorsal edges flattened (Fig. 2); body smaller (3-4 mm); elytral

punctation striate; elytral color solid blue/black (no pattern) Phydanis

8'. Metatibial apex with dorsal edges continuous and morphologically unchanged (Fig.

3); body larger (4-8 mm); elytral punctation confused; elytra with distinct pattern of
large dark spots or transverse bands on a light background 9

9. Procoxal cavities open (almost closed, but not); metafemoral spring with recurve flange

from ventral lobe (morpho-group #3, see Furth, 1989) (Fig. 16); spermatheca with
distinctly separate pump and receptacle and with spermathecal ductus simple and
uncoiled (Fig. 17) Chrysogramma

9'. Procoxal cavities completely closed; metafemoral spring (Fig. 13) without recurve
flange from ventral lobe (morpho-group # 1 ); spermatheca without distinctly separate
pump and receptacle and some species with a coiled spermathecal ductus (Fig. 14, 15)
Acrocyum

RESULTS AND DISCUSSION

Admittedly the separation of Blepharida from Notozona is difficult. The combi-
nation of characters given in the key above should provide adequate means for
separation of these two genera. Generally the reliance on color or pattern as a primary
key character is not advisable; however, at present it appears to be the best possibility
for these taxa. A variety of characters has been suggested for separation of Blepharida
and Notozona by previous workers, but most have not proved 100% applicable.
Therefore, it seems most practical to include several characters that each pertain to
a majority of the species in each genus, the combination of which will yield almost
complete reliability of determination. Some previous workers have also used a few
other less reliable characters such as: antennal segment 2 distinctly shorter than
segments 3 or 4 in Blepharida, whereas segments 2-4 subequal but shorter than the
others in Notozona ; and maxillary palpus narrow and filiform in Blepharida, but
more robust/swollen and claviform apically in Notozona. These characters are of
some use for separation of these two genera, but are quite variable in certain species.
Further knowledge of the biology (e.g., Furth, 1982), foodplants, larvae, and other
biological information may provide additional characteristics for the separation of
Blepharida and Notozona. Preliminary information indicates that certain species in
these genera feed on related foodplants (Furth, unpubl data); however, foodplant
preference testing, leaf biochemical differences, and even foodplant ecology may be
useful for indicating relationships.(Furth and Young, 1 988). Also, the status of several
species of Blepharida and Notozona will be changed in a species-level revisionary
study of these two genera (Furth, unpubl. data). Therefore, the author prefers at this
point to consider Blepharida and Notozona as valid separate genera in much the
same way as Jacoby (1885, 1891) and Scherer (1983).

Bechyne (1968) states that the true Blepharida consisted of only three species based
on a transverse series of setiferous punctures on the anterior clypeus and the absence
of a transverse clypeal groove (with accompanying carina above). However, the author
finds these characters present only in B. rhois, not in B. irrorata or B. haitiensis. In
fact, the status of the last of these species is quite different, as discussed below.
Bechyne (1968) lumped all other New World Blepharida ( sensu lato) into Notozona ;
the present author disagrees and retains them as Blepharida.

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405

17

Figs. 13-17. 13. Acrocyum haitiensis metafemoral spring (L = 0.56 mm). 14. Acrocyum

haitiensis spermatheca (L = 0.59 mm). 1 5. Acrocyum maculicollis spermatheca (L = 0.59 mm).
16. Chrysogramma pictipennis metafemoral spring (L = 0.85 mm). 17. Chrysogramma picti-
pennis spermatheca (L = 0.70 mm).

In the present study the aedeagus, spermatheca, and metafemoral spring of several
species of both Blepharida and Notozona are studied to look for generic level char-
acteristics. The aedeagi of these two genera are quite variable at the species level and
show no generic level characters. The spermathecae of Blepharida and Notozona are
quite similar in the shape of the main body of the spermatheca— receptacle and pump
(Figs. 19, 18, respectively), but they do show some species differences in the ductus
shape. The metafemoral spring of Blepharida and Notozona is essentially identical
belonging to metafemoral spring morpho-group #1 (see Furth, 1988, 1989)— a long
extension (extended arm) of the dorsal lobe, the ventral lobe without a recurved
flange and with an acute basal angle (pointed basally) (Fig. 8). These three internal
characters (aedeagus, spermatheca, and metafemoral spring) were also examined for

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all the genera included in this study to reveal generic level characters. The results of
this are partially reflected in the above key and the associated figures. As with Bleph-
arida and Notozona, the aedeagus reveals little obvious definitive, generic level char-
acters; however, a more comprehensive survey of species in these genera is needed
in order to be certain of this conclusion. The spermatheca and metafemoral spring
are much more revealing as generic characters, as demonstrated in the above key.

As indicated in the Introduction, there has been some confusion and disagreement
as to the proper inclusions for the genus Blepharida. Scherer (1961) basically followed
the catalogue arrangement of Heikertinger and Csiki (1940) which was continued by
Seeno and Wilcox (1982). These authors considered Blepharida to be composed of
three subgenera: Blepharida {sensu stricto), from the New World, possessing bifid
tarsal claws; Calotheca, from East and South Africa and the Celebes, with simple
tarsal claws; and Blepharidella, from East and South Africa, with appendiculate tarsal
claws. Bechyne (1968) raised these subgenera to generic status, divided New World
Blepharida into Blepharida { sensu stricto) and Notozona, and created a new genus,
Blepharidina, for the Afrotropical species with bifid tarsal claws. Seeno and Wilcox
(1982) treat Blepharidina as a valid genus. In the present study it has become apparent
that there is a group of external morphological differences between the Afrotropical
and New World Blepharida that possess bifid tarsal claws, other than those mentioned
by Bechyne (1968). The author considers Bechyne’s Blepharidina to be only a sub-
genus of Blepharida from the Afrotropical and Madagascan Regions, with bifid tarsal
claws and possessing the following external morphological characters: proepimeron
laterally separated from pronotum by a deep grove or suture; frons with deeply
impressed, sinuate grooves extending from lower to upper frons between antennae;
pronotum with antero-lateral quadrants with a short row of sublateral, longitudinal
punctures, usually connected to a transverse (horizontal) row of coarse punctures
reaching the lateral pronotal border, thus forming a square or circle in the antero-
lateral comers of the pronotum; elytra always striate; clypeus without transverse
groove or carina; anterior border of clypeus without transverse series of setiferous
punctures; elytral epipleuron with some relatively large, dark spots/pattern of color.
Preliminary examination also indicates some characteristic differences of sperma-
thecal morphology (e.g., relatively longer pump). These characters, or at least in
combination, are not present in the New World subgenus Blepharida {sensu stricto).

Another change in status is Blepharida haitiensis Blake, considered to be a true
Blepharida, especially by Bechyne (1968). The following characters place B. haitiensis
as a species of Acrocyum: form of the metafemoral spring (Fig. 13); female sper-
mathecal morphology (Fig. 14); male aedeagus short and stout in form; absence of
any flattening or emargination of the dorsum of the metatibial apex; appendiculate
tarsal claws; small round eyes; elytral pattern primarily composed of dark spots; and
elytral punctation confused, very fine, sparse. Even though Bechyne (1968, unpubl.
data) considered Acrocyum congeneric with Chrysogramma, evidence (i.e., metafem-
oral spring and female spermatheca) provided in the key above proves that they are
truly separate genera.

The genus Euplectroscelis Crotch is a monotypic genus endemic to Baja California
(Mexico). The six species described by Baly as Euplectroscelis and listed in Heikertin-
ger and Csiki (1940) all belong to Heikertingerella Csiki. E. Riley and A. Gilbert
(personal communication) have observed Euplectroscelis in nature. They report that

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NEW WORLD BLEPHARIDA GROUP

407

Figs. 18-19. 18. Notozona histrionica spermatheca (L = 1.22 mm). 19. Blepharida francle-

monti spermatheca (L = 1 .04 mm).

this and a Blepharida species feed on Bursera (Burseraceae) and that their larvae are
similar morphologically and behaviorally to those of Blepharida. As indicated in the
above key, the present study has revealed that Euplectroscelis adults are morpho-
logically very similar to Blepharida, and that the placement of Euplectroscelis in
catalogues or checklists should be just after Blepharida rather than far away near
Chaetocnema Stephens as it is in Seeno and Wilcox (1982).

The author finds it curious that the arrangement of scientific names in catalogues
or checklists (e.g., Heikertinger and Csiki, 1940, Seeno and Wilcox, 1982) is sup-
posedly based on some type of similarities, presumably morphological; however,
these similarities or the rationale of such catalogue arrangements is not explained or
referred to and such “accepted” arrangements are often passed down through several
generations. Admittedly, these traditional “catalogue phytogenies” are often con-
venient for curating collections and some doubtlessly reflect valid relationships;
however, they must be viewed with great caution and restraint when inferring true
phylogeny or broad-spectrum relationships. Therefore, until more is known about
the true phylogeny of the Blepharida- group genera worldwide, for the convenience
of such catalogue arrangements, the author proposes that (based on the present
morphological study) the following order of New World genera be used in catalogues
and checklists:

Blepharida Chevrolat, 1837
( Blepharida Chevrolat sensu stricto)

(Blepharidina Bechyne, 1968)

(Calotheca Hey den, 1887)

{Blepharidella Weise, 1 909)

Notozona Chevrolat, 1837
Euplectroscelis Crotch, 1873
B/epharonycha Fall, 1927
Acrocyum Jacoby, 1885
Ch rysogra mma J acoby ,1885

Genera from the Afrotropical, Madagascan, and Pacific Ocean regions should be

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Figs. 20-22. 20. Blepharida franclemonti aedeagus. a) ventral view b) lateral view (L = 3.40

mm). 21. Blepharida suturalis aedeagus. a) ventral view b) lateral view (L = 3.22 mm). 22.
Blepharida mexicana aedeagus. a) ventral view b) lateral view (L = 3.52 mm).

listed geographically after these New World genera, primarily for convenience, until
their relationships to the other genera can be studied.

Elithia Chapuis, Procalus Clark, and Crimissa Stal should probably be left together
as in the Seeno and Wilcox, 1982 checklist. Their relationship to the Blepharida-
group genera is not apparent from the present study of external and internal mor-
phology.

Pseudorthygia Csiki and Phydanis Horn are not related to the Blepharida- group
and should not be included with them. Currently it is not clear where they should
be placed in catalogues.

Blepharida franclemonti, new species
Figs. 4, 19, 20, 23

Antennae, color all yellow; average segment (1-11) lengths in millimeters 0.67,
0.23, 0.56, 0.59, 0.74, 0.56, 0.56, 0.52, 0.48, 0.44, 0.56.

Head, color yellow/light brown; clypeus smooth and impunctate; upper clypeus
(lower frons) with deep transverse furrow, creating a prominent carina below (slightly
wider in center); upper frons between antennal sockets with faint longitudinal carina,
not raised, flattened with a few fine punctures, and only faintly delimited laterally
by impressions; frontal bossae round, but poorly delimited, not raised and bordered
only mesally and dorso-laterally by small depressions; a few very fine punctures along
mesal eye margins; vertex with finely shagreened surface and very fine, sparse, con-
fused punctures; eyes elliptical converging dorso-mesally (Fig. 4); average dorso-
ventral eye length =1.11 mm, average interocular distance (at dorso-mesal margins)
= 1.04 mm.

Pronotum. color yellow/light brown; rectangular; lateral margins evenly, gradually
rounded, slightly narrower at antero-lateral angles; antero-lateral angles obtuse, pro-
truding laterally somewhat; punctation very fine, with a few sparse, coarser punc-
tures antero-lateral to midline; surface smooth with slight evidence of shagrination;

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409

23a

24a

23b 24b

Figs. 23-24. 23. Blepharida franclemonti elytral pattern a) dorsal view b) lateral view (Length

of elytra = Le = 6.80 mm). 24. Blepharida suturalis elytral pattern a) dorsal view b) lateral
view (Le = 7.04 mm).

laterally often with small, gently depressed areas; average width at middle = 3.52
mm; average length at middle = 1.92 mm.

Elytra, striate with deep, medium-sized punctures; interstriae surface finely sha-
greened; background color chestnut/mahogany brown, with yellow markings; yellow
pattern (Fig. 20a, b) often slightly assymetrical (fine details only); yellow pattern
consisting mostly of yellow spots, sometimes contiguous, always on interstriae and
distributed longitudinally, not forming transverse bands; third interstriae (between
full puncture rows number 2 and 3 from suture, not including incomplete basal
scutellar row) often with few or no spots; occasionally also small yellow spots near
suture in middle or posteriorly; base of elytra with continuous, transverse yellow
band (posterior borders irregular) surrounding the brown scutellum and continuous
as a thin yellow longitudinal stripe along the base of the suture for approximately
one quarter of elytral length; humeral callus prominent and surrounded by yellow
(interstriae 6-9 with fused yellow spots surrounding brown humeral spot); fused
yellow spots forming partial longitudinal stripes on interstriae 6 and 1 0 (lateral-most
stria); spots on interstriae 6-10 often laterally forming small transverse bands of up
to three interstriae wide; apically interstriae tapered and narrow, yellow spots often
fused to form partial apical yellow band joining four or more interstriae; male elytral

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

25b 26b

Figs. 25-26. 25. Blepharida mexicana elytral pattern a) dorsal view b) lateral view (Le =

7.52 mm). 26. Notozona bifasciata elytral pattern a) dorsal view b) lateral view (Le = 6.24
mm).

length = 6.72-6.96 mm, width (at middle) = 5.04-5.20 mm; female elytral length =
7.04-7.68 mm, width = 5.28-5.60 mm; general body form similar to Chrysomelinae.

Venter, color yellow/light brown; pubescent throughout; procoxal cavities narrowly
closed; prostemal process flat, apically expanded; mesostemal process with posterior
margin emarginate; mesostemum vertically oriented; male apical stemite sublaterally
cleft forming median, U-shaped lobe; apical tergite U-shaped; all stemites finely,
densely punctate.

Legs, male first fore and midtarsal segments greatly expanded; metafemora elon-
gate-oval in shape; metatibial dorsal edges, apically strongly emarginate, each edge
with preapical protuberance; tarsal claws strongly bifid.

Aedeagus. in ventral view (Fig. 20a) apex evenly rounded with very rugose margin,
subapical ventral surface with triangular raised area, tapered basally (also evident in
lateral view); in lateral view (Fig. 20b) apex almost at right angle (dorsum/venter),
extreme apex with small step-like angle, dorsal apical projection broadly hook-like;
in dorsal view surface flat, dorsal projection with centrally tapering, subparallel,
sclerotized ribs expanding apically into a short, umbel-shaped, sclerotized structure.

Spermatheca. Figure 19; slightly twisted ductus, visible only from ductus (right)
side.

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NEW WORLD BLEPHARIDA GROUP

411

27b 28b

Figs. 27-28. 27. Notozona histrionica elytral pattern a) dorsal view b) lateral view (Le =

7.20 mm). 28. Notozona nicaraguensis elytral pattern a) dorsal view b) lateral view (Le = 8.56
mm).

Differences from related species. B. franclemonti is closest to B. suturalis Jacoby,
1885 (page 385, plate 22, fig. 12) and quite similar to B. mexicana Jacoby, 1885
(page 386, plate 22, fig. 16) differing from these primarily in elytral pattern and
aedeagus morphology. B. suturalis differs from the new species by having the following
characteristics: Elytra (Fig. 24a, b): more elytral yellow pattern, especially often
forming transverse bands at apex and just in front of the middle; the yellow pattern
usually as longitudinal stripes, not spots, especially along the apical two thirds of the
second interstriae; first (sutural) interstriae without any yellow markings on apical
two thirds, thus forming an apical brown sutural stripe; subbasal large brown areas;
humeral and posthumeral brown spots; apex almost always with fused yellow, sub-
marginal stripes. Aedeagus: Ventrally (Fig. 21a) deeply emarginate apex with less
rugose margin; more prominently raised area subapically, diverging before tapering
basally. Laterally (Fig. 21b) apical part (dorsum to venter) obtusely angled; extreme
apex without step-like angle; dorsal apical projection broadly hook-like in shape;
subapical raised area prominent. Dorsally surface deeply concave; dorsal projection
with subparallel, more broadly tapered, median, sclerotized ribs, expanding apically
into broad, umbel-shaped, more sclerotized apex than in franclemonti (more bullet-
shaped apex). Distribution: Guatemala; El Salvador; Nicaragua; and Costa Rica.

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B. mexicana differs from B. franclemonti as follows: Elytra: pattern (Fig. 25a, b),
especially by having mostly yellow pattern with only a humeral brown spot and a
few brown longitudinal, partial stripes, most notably on interstriae 3 and 5; most of
sutural and subsutural interstriae yellow; apex entirely yellow. Pronotum: more quad-
rate (relatively longer than wide) than in the other two species. Aedeagus: ventrally
(Fig. 22a) has a less rugose, more sinuate apical margin; laterally (Fig. 22b) no
subapical raised area; dorsal projection (only slightly hook-shaped) pointing straight
out apically; dorsally flat surface, dorsal projection with narrow, subparallel ribs,
diverging apically, expanding laterally into lightly sclerotized, subumbel-shape (al-
most T-shape) with lateral angles acutely pointed. Distribution: Mexico (Oaxaca and
Vera Cruz States).

Etymology. This new species is named for Prof. Emer. John G. Franclemont
(Cornell University) who offered me valuable guidance and encouragement during
my first few instars as a coleopterist, including during my first research on the genus
B/epharida.

Material examined. HOLOTYPE: male, (MCZ type number 33406), Mexico, label
data: “Mex: Yucatan, Chichen Itza 18-19 V 1987 D. A. Rider, E. G. & T. J. Riley.”
Deposited in the Department of Entomology, Museum of Comparative Zoology
(MCZ), Harvard University as a donation from Edward G. Riley (EGR). ALLO-
TYPE: female (MCZ type number 33406), “Mexico, Yuc. Merida VII 29-30 1964
Paul J. Spangler.” Deposited in the Department of Entomology, Museum of Com-
parative Zoology (MCZ), Harvard University as a donation from the National Mu-
seum of Natural History, Washington, D.C. — (USNM). PARATYPES: male (EGR),
female (J. E. Wappes), “Mex., Quin. Roo 18-24 km N. San Felipe Carr. pte. 5/27/
-6/1/84, J. E. Wappes”; 3 males (1 EGR, 2 Robert Turnbow-RT), female (RT),
“Mex, Quintana Roo, 18 km N. Carrillo Puerto, at lite. 31 May 1984 R. Turnbow”;
3 females (USNM), same data as Allotype; female (American Museum of Natural
History, New York— AMNH), “Piste. Yucatan Mex. VI 8-10 59 P. & C Vaurie”;
female (AMNH), “Mex., Yucatan Piste June 3-5, 59 P. & C. Vaurie”; female (The
Natural History Museum, London), “55 24,” “Jac,” “Baly Coll”; female (MCZ),
“Mex”; female (MCZ), “Yucatan.”

ACKNOWLEDGMENTS

The author thanks the following people and their institutions for the loan of specimens in
genera treated in this study: B. S.. Bechyne, L. J. Joly, V. Savini (Universidad Central de
Venezuela, Maracay); N. Berti (Museum National D’Histoire Naturelle, Paris); J. A. Chemsak
(University of California, Berkeley); A. J. Gilbert (California Department of Food and Agri-
culture, Fresno); G. Haghebaert (Institut Royal des Sciences Naturelles de Belgique, Brussels);
L. H. Herman (American Museum of Natural History, New York); J. Jelinek (Narodni Museum
V Praze, Prague); D. H. Kavanaugh and N. D. Penny (California Academy of Sciences, San
Francisco); J. K. Liebherr and E. R. Hoebeke (Cornell University, Ithaca); E. G. Riley (E. G.
Riley Collection, and Texas A&M University, College Station); S. Santiago (Universidad Na-
tional Autonoma de Mexico, Mexico City); T. N. Seeno (California Department of Food and
Agriculture, Sacramento); S. L. Shute (The Natural History Museum, London); A. Solis and
R. W. Flowers (Instituto Nacional de Biodiversidad, San Jose); R. Turnbow (Ft. Rucker, AL);
J. E. Wappes (Chadds Ford, PA); R. E. White (National Museum of Natural History, Wash-
ington, D.C.). The author is also grateful to C. N. Duckett, E. G. Riley, and G. Scherer for

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NEW WORLD BLEPHARIDA GROUP

413

information and discussion about the genera treated here. Support for travel to the Bechyne
collection in Venezuela was from an Ernst Mayr Grant, Harvard University.

LITERATURE CITED

Arnett, R. H. 1971. The Beetles of the United States. The American Entomological Institute,
Ann Arbor.

Baly, J. F. 1865. Diagnosis of a new genus and some undescribed species of African Phy-
tophaga. Entomol. Mon. Mag. 14:204-206.

Bechyne, J. 1968. Contribution a la faune du Congo (Brazzaville). Mission A. Villiers et A.
Descarpentries. LXXXI. Coleopteres Alticidae. Bull. Inst. Franc. Afrique Noire, Ser A.
30(4): 1687—1728.

Chapuis, F. 1 875. In: Lacordaire, Histoire naturelle des insectes. Genera des coleopteres. Vol.
1 1, Familie des Phytophages, Paris.

Chevrolat, L. A. 1837. In: Dejean, Catalogue des Coleopteres de la collection de M. le comte
Dejean. Troisieme edition, revue, corigee, et augmentee, livr 5, pp. 385-503. Paris.
Crotch, G. R. 1873. Materials for the study of the Phytophaga of the United States. Proc.
Acad. Natur. Sci. Phila. 1873:19-83.

Fall, H. C. 1927. Expedition of the California Academy of Sciences to the Gulf of California.

The Chrysomelidae (Coleoptera). Proc. Calif. Acad. Sci. (4) 16:381-395.

Furth, D. G. 1982. The metafemoral spring of flea beetles (Chrysomelidae: Alticinae). Spix-
iana, Suppl. 7:1 1-27.

Furth, D. G. 1985. Relationships ofPalearctic and Nearctic genera of Alticinae. Entomography
31:375-392.

Furth, D. G. 1988. The jumping apparatus of flea beetles (Alticine)— the metafemoral spring.
In: P. Jolivet, E. Petitpierre, and T. Hsiao (eds.), Biology of Chrysomelidae. Ch. 1 7:285—
297.

Furth, D. G. 1989. Metafemoral spring studies of some Neotropical genera of Alticinae.
Entomography 6:497-510.

Furth, D. G. and D. A. Young. 1988. Relationships of herbivore feeding and plant flavonoids
(Coleoptera: Chrysomelidae and Anacardiaceae: Rhus). Oecologia 74:496-500.
Heikertinger, F. and C. Csiki. 1940. Coleopterorum catalogus, Chrysomelidae: Halticinae,
pars 169:335-635.

Heyden, L. von. 1887. Kleine coleopterische Mittheilungen. Wiener Entomol. Zeit. 6(3):98.
Horn, G. H. 1889. A synopsis of the Halticini of Boreal America. Trans. Amer. Entomol.
Soc. 16:163-320.

Jacoby, M. 1885. Biologia Centrali-Americana, Insecta, Coleoptera, Galerucidae 6(1):337-
408.

Jacoby, M. 1891. Biologia Centrali-Americana, Insecta, Coleoptera, Supplement to Phytophga
6( 1 ):233— 3 1 2.

Laboissiere, V. 1937. Nouveaux Galerucinae palearctiques. Bull. Soc. Ent. France 42:27-31.
Scherer, G. 1961. Bestimmungschlussel der Alticinen-Genera Afrikas. Entomol. Arbeiten Frey
12:251-288.

Scherer, G. 1962. Bestimmungschlussel der neotropischen Alticinen-Genera. Entomol. Ar-
beiten Frey. 13:497-607.

Scherer, G. 1983. Diagnostic key for the Neotropical alticine genera (Coleoptera: Chryso-
melidae: Alticinae). Entomol. Arbeiten Frey. 31/32:1-89.

Seeno, T. N. and J. A. Wilcox. 1982. Leaf Beetle genera (Coleoptera: Chrysomelidae). En-
tomography 1:1-221.

Weise, J. 1909. Chrysomelidae und Coccinellidae. In: Sjostedt, Ergebnisse der schwedischen
Expedition nach Kilimandjaro, vol. 7(1 2): 153-266.

414 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(2)

Wilcox, J. A. 1975. Checklist of the Chrysomelidae of Canada, United States, Mexico, Central
America, and the West Indies. Family No. 104. N. Amer. Beetle Fauna Proj. pp. 1-166.

Received 1 1 July 1991; accepted 9 October 1991.

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Journal of the

New York Entomological Society

VOLUME 100 APRIL 1992 NO. 2

CONTENTS

Contributions from former students
in honor of the 80th Birthday of
Professor John G. Franclemont

A remembrance of John G. Franclemont Ronald W. Hodges

Neotropical Microlepidoptera XXV, a new arboreal species of Acrolophus from
Peru (Lepidoptera: Acrolophidae) Donald R. Davis and Charles L. Hogue

Two new species of Mompha from California (Lepidoptera: Momphidae)

Ronald W. Hodges

Six new species of Catastega (Lepidoptera: Tortricidae: Eucosmini) from Mexico
and southwestern United States Richard L. Brown

Pupal morphology and the subfamily classification of the Notodontidae (Lepi-
doptera: Noctuoidea) James S. Miller

Franclemont la interrogans (Walker), a new genus for an old species (Lepidoptera:
Noctuidae: Amphipyrinae) Douglas C. Ferguson

An annotated key and illustrations to the species of the genus Cucullia (Lepidop-
tera: Noctuidae: Cuculliinae) from Mexico and Central America

Robert W. Poole

A revision of the genus Hypsoropha Hiibner (Lepidoptera: Noctuidae: Ophider-
inae) Tim L. McCabe

Life history and systematics of the West Andean moth Aucula franclemonti with
description of a new species from Eduador (Lepidoptera: Noctuidae: Agaristi-
nae) John E. Rawlins

Twenty years of fluctuating parapatry and the question of competitive exclusion
in the butterflies Pontia occidentalis and P. protodice (Lepidoptera: Pieridae)

Arthur M. Shapiro

Hydropsyche franclemonti, a new'species of the scalaris group from eastern North
America (Trichoptera: Hydropsychidae) Oliver S. Flint, Jr.

The Tribe Licinini (Coleoptera: Carabidae): a review of the genus-groups and of
the species of selected genera George E. Ball

Taxonomy and distribution of the athetine genus Lypoglossa Fenyes (Coleoptera:
Staphylinidae: Aleocharinae) in North America, with description of a new spe-
cies E. Richard Hoebeke

The New World Blepharida group, with a key to genera and description of a new
species (Coleoptera: Chrysomelidae) David G. Furth

185-190

191-202

203-208

209-227

228-256

257-266

267-272

273-285

286-310

311-319

320-324

325-380

381-398

399-414

JULY 1992

No. 3

L

61

867

NT ol. 100

Journal

of the

New York

Entomological Society

(ISSN 0028-7199)

Devoted to Entomology in General

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY

Editor: James K. Liebherr, Department of Entomology, Comstock Hall,

Cornell University. Ithaca, New York 14853
Assistant Editor: E. Richard Hoebeke, Department of Entomology, Com-

stock Hall, Cornell University, Ithaca, New York 14853
Book Review Editor: James S. Miller, Department of Entomology, Amer-

ican Museum of Natural History, Central Park West at 79th Street,
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Publications Committee: Randall T. Schuh, American Museum of Natural

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of Agriculture, Harrisburg.

The New York Entomological Society
Incorporating The Brooklyn Entomological Society

President: Durland Fish, Medical Entomology Laboratory, New York

Medical College, Valhalla, New York 10595

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of Natural History, New York, New York 10024

Trustees: Class ofl 990— Dennis J, Joslyn, Department of Biology, Rutgers

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York. Class of 1991 — Sule Fischl, New York, New York; Theodore
Weisse, Northport, New York.

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J. New York Entomol. Soc. 1 00(3):4 1 5—4 1 7, 1992

NEW SYNONYMS OF FRA NKLINIELLA BONDARI AND
A REVIEW OF THE SYNONYMS OF F. CEPHALICA
(THYSANOPTERA: THRIPIDAE)

Sueo Nakahara

Systematic Entomology Laboratory, PSI, Agricultural Research Service, USD A,
10300 Baltimore Avenue, Beltsville, Maryland 20705-2350

Abstract.— Frankliniella alba Moulton and F. bratleyi Watson are junior synonyms of F.
bondari Hood (New Synonymy). The seven species and varieties of Frankliniella treated as
junior synonyms of F. cephalica (Crawford) in the “Catalogue of the Thysanoptera of the World”
(1974), are not synonyms. Frankliniella bispinosa (Morgan), F. bruneri Watson, F. echinodora
Moulton, F. melanommata Williams and F. reticulata (Crawford) are treated as valid species.
Euthrips tritici var. projectus Watson and Frankliniella cephalica var. masoni Watson are junior
synonyms of F. bispinosa (New Synonymy).

Frankliniella bondari Hood (1 942:622) and F. bratleyi Watson (1942: 1 7) described
from tuberose, Polianthes tuberosa L. (Agavaceae), are synonyms based on my ex-
amination of the types. The types of F. bondari were collected in Bahia, Brazil, and
consist of only macropterous females. The types of F. bratleyi were collected in
Gainesville, Florida, on damaged bulbs and consist of macropterous and brachyp-
terous females. Hood’s paper was published in January, 1942 and Watson’s paper
in June, 1942. Thus, F. bondari has priority and F. bratleyi is treated here as the
junior synonym (New Synonymy).

Frankliniella alba Moulton (1948:1 13) was described from two females (macrop-
terous and brachypterous) found on gladiolus from Mexico. The macropterous ho-
lotype examined in this study is similar to F. bondari except for its longer posteroangu-
lar setae on the pronotum. The lengths of these setae vary in material of F. bondari
examined, and I conclude that F. alba is another junior synonym of F. bondari (New
Synonymy). The holotype of F. alba is deposited in the Thysanoptera collection of
the U.S. National Museum of Natural History, not in the California Academy of
Sciences as indicated by Jacot-Guillarmod (1974:762).

Frankliniella bondari has pale yellow body, wings and setae, and the head is slightly
produced anterior of the eyes. In this country, it resembles F. unicolor Morgan.
Frankliniella bondari has the basal one-third to one-half of antennal segment VI
yellow and the interocellar setae are positioned between the anterior and posterior
ocelli, whereas F. unicolor has completely brown antennal segment VI and the in-
terocellar setae are positioned laterad of the anterior ocellus.

The known distribution of F. bondari is Brazil, Mexico, and the United States
(Florida, Georgia and Hawaii). I have also examined a quarantine interception from
Israel and another from Cuba on Polianthes sp.

The preferred host is Polianthes tuberosa and some of the other recorded hosts
probably are incidental. According to “Hortus Third,” the species of Polianthes
originated in Mexico. The thrips apparently also originated in Mexico and was in-
troduced into other countries on the rhizomes of tuberose, which is grown com-

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

mercially and in gardens for the waxy-white, fragrant flowers. This species was re-
cently discovered in the state of Hawaii on the islands of Molokai and Oahu where
it damages tuberose.

Jacot-Guillarmod (1974:767) treated the following seven taxa as junior synonyms
of Frankliniella cephalica (Crawford) (1910:153) in his “Catalogue of the Thysa-
noptera of the World.”

Euthrips cephalicus var. reticulata Crawford, 1910:153.

Euthrips tritici var. bispinosus Morgan, 1913:10.

Frankliniella melanommatus Williams, 1913:213.

Euthrips tritici var. projectus Watson, 1915:51.

Frankliniella cephalica var. masoni Watson, 1919:4.

Frankliniella cephalica var. bruneri Watson, 1926:54.

Frankliniella cephalica var. echinodora Moulton, 1948:107.

In this review of the synonyms, I conclude that none of these taxa are junior synonyms
of F. cephalica.

The lectotype of E. cephalicus var. reticulata, which is deposited in the Canadian
National Collection, was examined. On the type slide are also seven yellow specimens,
of which five are F. borinquen Hood and two are F. cephalica. In my opinion,
reticulata is a distinct species in Frankliniella (New Status). The body is uniform
brown as described by Crawford (1910:153). It also differs morphologically from F.
cephalica in the shape of antennal segment II and enlargement of the pedicel of
antennal segment III, and by having on abdominal tergite VIII, a complete postero-
marginal comb which is medially incomplete in F. cephalica.

Sakimura (1981:484, 1986:354) treated F. bruneri and F. melanommata as good
species.

Although a formal redesignation has not been published, F. bispinosa in recent
years has been given species status in identifications, reports and publications (i.e.,
Beshear, 1979:210) for the following reasons: the pedicel of antennal segment III of
F. bispinosa is larger and differently shaped than that of F. cephalica, and in Florida
and Bermuda, F. cephalica is normally found in the flowers of Bidens sp. and F.
bispinosa in various flowers. Frankliniella bispinosa is recorded from Florida, Geor-
gia, South Carolina, Bahama Islands and Bermuda.

In my opinion, F. cephalica var. masoni and E. tritici var. projectus are junior
synonyms of F. bispinosa (New Synonymy).

The holotype and allotype of F. cephalica var. echinodora differ from F. cephalica
in the shapes of the enlargement of the pedicel and basal part of antennal segment
III. For these reasons, I treat the two taxa as different species (New Status). Frank-
liniella echinodora is known only from Puerto Rico. Jacot-Guillarmod (1974:770)
stated that the types were deposited in the California Academy of Sciences. The
paratypes are deposited in that museum but the holotype and allotype are deposited
in the Thysanoptera collection of the U.S. National Museum of Natural History.

ACKNOWLEDGMENTS

I thank H. A. Denmark, Florida Dept, of Agriculture and Consumer Services, Gainesville,
R. J. Beshear, University of Georgia, Griffin, T. J. Henry, Systematic Entomology Laboratory,
Washington D.C. and R. L. Smiley, same laboratory, Beltsville, Maryland, for their reviews of

1992

FRANKLINIELLA SYNONYMS

417

the manuscript and useful comments. I also thank R. Foottit, Biosystematic Research Center,
Agriculture Canada, Ottawa, for the loan of F. cephalica and F. reticulata types.

LITERATURE CITED

Beshear, R. J. 1979. Additional records of thrips in Georgia (Thysanoptera: Terebrantia). J.
Georgia Entomol. Soc. 4:209-21 1.

Crawford, D. L. 1910. Thysanoptera of Mexico and the South. II. Pomona College J. Entomol.
2:153-170.

Hood, J. D. 1942. A century of new American Thysanoptera. Ill Rev. Entomol., Rio de
Janeiro (1941) 12:547-678.

Jacot-Guillarmod, C. F. 1974. Catalogue of the Thysanoptera of the world, part 3. Ann. Cape
Province Mus. (Nat. Hist.) 7:517-976.

Morgan, J. 1913. New genera and species of Thysanoptera with notes on distribution and
food plants. Proc. U.S. Nat. Mus. 46:1-55.

Moulton, D. 1948. The genus Frankliniella Kamy, with keys for the determination of species
(Thysanoptera). Rev. Entomol., Rio de Janeiro 19:55-1 14.

Sakimura, K. 1981. A review of Frankliniella bruneri Watson and description of F. kelliae,
n. sp. (Thysanoptera: Thripidae). Florida Entomol. 64:483 — 49 1 .

Sakimura, K. 1986. Thrips in and around the coconut plantations in Jamaica, with a few
taxonomical notes (Thysanoptera). Florida Entomol. 69:348-363.

Watson, J. R. 1915. New Thysanoptera from Florida. Entomol. News 26:49-52.

Watson, J. R. 1919. Additions to the Thysanoptera of Florida. V. Florida Buggist 3:2-7.
Watson, J. R. 1926. Two new Thysanoptera from Cuba. Florida Entomol. 9:53-55.

Watson, J. R. 1942. A new Frankliniella from Florida (Thysanoptera). Florida Entomol. 25:
17-18.

Williams, C. B. 1913. On two new species of Thysanoptera from the West Indies. J. Econ.
Biol. 8:209-215.

Received 14 May 1991; accepted 9 December 1991.

J. New York Entomol. Soc. 100(3):4 18-423, 1992

THE FIRST RECORDS OF THE FAMILIES OCHTERIDAE AND
HEBRIDAE (HETEROPTERA) FROM THE GRANITIC
SEYCHELLES, WITH DESCRIPTIONS OF
TWO NEW SPECIES

Dan A. Polhemus

Applied Research Group, Bishop Museum, P.O. Box 19000-A,

Honolulu, Hawaii 96817

Abstract. — The families Ochteridae and Hebridae are recorded for the first time from the
granitic Seychelles, based on collections from mountain streams on the island of Mahe. Two
new species, Ochterus seychellensis and Hebrus seychellensis, are described, with dorsal habitus
and male genitalic figures.

The aquatic Heteroptera fauna of the granitic Seychelles is of interest from a
biogeographical standpoint, since it is hypothesized that these oceanic granitic islands
represent small continental fragments stranded in the wake of drifting India (Davies,
1968). Up to the present time the fauna has appeared to be relatively similar to that
occurring on the high islands of the Mascarenes, consisting primarily of vagile, wide-
spread species of common occurrence throughout the western Indian Ocean basin.
Several recent discoveries, however, have revealed that the granitic Seychelles do in
fact harbor a significantly richer fauna than the Mascarenes, including the unusual
mesoveliid genus Phrynovelia (Andersen, in press) and two species of the genus
Rhagovelia (J. Polhemus, 1990). To these discoveries may now be added two new
species belonging to the families Ochteridae and Hebridae, which were previously
unknown from any of the high Indian Ocean islands between Madagascar and Ceylon.
The relationships of these new species have yet to be resolved, since the genera
Ochterus and Hebrus to which they belong are both large and in need of revision in
the Eastern Hemisphere, but their presence indicates that the granitic Seychelles may
indeed harbor the remnants of an old continental aquatic Heteroptera fauna that was
fragmented by vicariance events during the breakup of the Gondwana supercontinent.

The collections on which the new taxa in this paper are based were made by the
author during his participation in the Smithsonian Institution’s Aldabra Program
and were undertaken in cooperation with the Seychelles Islands Foundation. The
holotypes are deposited in the collections of the National Museum of Natural History,
Washington, D.C. (USNM); paratypes are also deposited in the J. T. Polhemus
collection, Englewood, Colorado (JTPC). All measurements are given in millimeters.
The CL numbers following locality data refer to codes used by the author to reference
ecological notes.

Ochterus seychellensis, new species

Diagnosis. This species may be separated from any other Ochterus presently de-
scribed from India, Ceylon, Africa or Madagascar by the following combination of
characters: the rostrum exceeds the hind coxae; the clavus and corium are smooth.

1992

HETEROPTERA FROM SEYCHELLES

419

without deep punctations; the hemelytra lack gold scale-like setae; and the embolium
is only weakly punctate. The absence of punctations on the distal part of the inner
corium is a character state not seen in any other Old World species.

Description. Male: general form broadly ovate, body length 4.31 mm; maximum
width 2.42 mm (Fig. 1). General coloration dark velvety green, with scattered pru-
inose lavender markings as follows: narrow linear patches at base of ocular flanges
and along anterior pronotal margin adjoining vertex; four small irregular patches
arranged transversely between anterior and posterior lobes of pronotum; single small
triangular patch medially along anterior margin of scutellum; broad irregular patches
basally on each half of clavus and corium; two large roughly circular patches on each
embolium; 5 to 6 irregular patches of varying size on each half of central corium;
irregular network of partially interconnected patches on wing membrane. Head shin-
ing black; anterior margin of clypeus, anterolateral and posterior margins of pro-
notum, anterolateral margin of embolium, rostrum, legs and abdominal venter dark
yellow; antennal segments I and II pale yellow, segments III and IV pale brown;
thoracic venter dull pruinose black; hind tibiae spotted with black at bases of stout
erect spines.

Head glabrous, length (along midline as measured from directly above)/width
(across eyes) = 0.39/1 .34, angled downward at 45° when viewed laterally; vertex with
numerous striae; eyes large, protruding, projecting posteriorly behind vertex; anten-
nae with segments I and II short, globose, segments III and IV slender, filiform, lengths
of segments I-IV = 0.11, 0.11, 0.28, 0.31; rostrum long, length = 2.58, exceeding
hind coxae and extending onto base of abdominal venter.

Pronotum length (midline)/width = 1.06/2.21, bearing numerous small glabrous
punctations, lacking evident setae; anterior collar prominent, flat; anterolateral mar-
gins explanate, well demarcated from disk; calli weakly tumescent; posterior margin
multisinuate, posteriorly concave centrally above base of scutellum. Scutellum tri-
angular, length/width = 0.73/1.34, weakly tumescent, bearing numerous small gla-
brous punctations centrally plus a double row of slightly larger, more closely spaced
punctations along lateral margins, lacking evident setae. Hemelytra long, attaining
tip of abdomen, corium, clavus and embolium well defined, surfaces smooth, sparsely
set with very tiny gold setae, lacking evident punctations except for a few very tiny
punctations on embolium; anterolateral embolar margin explanate; length of clavus
along outside margin = 1.79; membrane venation evident, defining 7 closed cells.

Ventral surface of thorax smooth and pruinose, lacking setae, bearing scattered
punctations on pro-, meso-, and metaepistema; abdominal venter covered with very
short fine appressed gold setae, this setal covering interrupted by ovate glabrous
patches surrounding spiracles laterally on paratergites. Leg segments covered with
short, fine, recumbent gold setae; anterior and middle tibiae bearing numerous short
bristly gold setae, these setae becoming more numerous distally; fore and middle
tibiae with a few longer semi-erect spinelike setae on posterior margins; hind tibia
with double longitudinal row of large erect gold spines; all tibiae with transverse rows
of spines at distal ends; claws gold, gently curving, arolia long, exceeding length of
claws when viewed laterally.

Genital segment well retracted into abdomen; subgenital plate with V-shaped in-
dentation medially on posterior margin. Pygophore quadrate, apex transverse, lateral
lobes well removed from apex (Fig. 2). Right paramere with head of moderate height,

420

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Figs. 1-3. Ochterus seychellensis, n. sp. 1. Male, dorsal habitus (legs omitted). Locations of
pruinose lavender patches defined by stippling. 2. Male pygophore, caudal tip. 3. Male genital
complex.

Figs. 4-7. Hebrus seychellensis, n. sp. 4. Male, dorsal habitus (legs omitted). Locations of
white markings on hemelytra defined by stippling. 5. Male right paramere, top view. 6. Male
right paramere, lateral view. 7. Male genital capsule, left lateral view.

1992

HETEROPTERA FROM SEYCHELLES

421

evenly domed and convex; appendices moderately long, apices indented; shaft stout,
broadened distally (Fig. 3).

Female: Similar to male in general structure and coloration but slightly smaller,
body length 4.06, width 2.35.

Discussion. O. seychellensis n. sp. is immediately separable from the widespread
O. marginatus occurring on India and Ceylon, which possesses numerous punctations
on the clavus and corium, and alternating brown and yellow spots on the margins
of the hemelytra, and from the somewhat similar O. minor from Africa, which also
has punctate hemelytra and dark yellow spots along the wing margins, a yellowish
area centrally on the posterior margin of the pronotum, and a shorter rostrum that
does not exceed the hind coxae. O. seychellensis n. sp. is also separable from the
south and east African O. cajfer, which is much larger, has punctate hemelytra and
a short rostrum which does not exceed the hind coxae, and lacks the bluish pruinosity
at the base of the clavus.

The general habitus and smooth hemelytra of O. seychellensis n. sp. are reminiscent
of a group of Neotropical ochterids which also occur on vertical rock faces, but these
resemblances appear to be due to convergence rather than monophyly, since the New
World species have a very different male genitalic plan.

Etymology. The name “seychellensis” refers to the Seychelles islands.

Habitat data. The type series was taken from a wet vertical rock wall along the
Foret Noire road near the Congo Rouge trailhead; this is close to the highest point
that can be reached by road on the island of Mahe. The rock face was formed by the
road cut, with disturbed secondary rain forest immediately above it. A shallow flow
of water trickled down across the face, collecting in a narrow concrete channel at its
base. The face itself was irregular and covered with alternating patches of moss and
open wet rock, and the ochterids moved across it with great agility, being quick to
fly if pursued or otherwise disturbed. To slow their movements a small amount of
pyrethrin insecticide was sprayed on the rock face; this application resulted in the
appearance of many previously undetected ochterids which had apparently been
secreted amid the moss or in cracks in the rock. This treatment also resulted in the
appearance of an undescribed species of Phrynovelia (Andersen, in press) which
appears to share the rock face habitat with O. seychellensis. Another single specimen
of O. seychellensis was taken along the upper Riviere Grande Bois on the opposite
side of the road from the rock face, and several other individuals were seen along
this stream. All were on shaded mossy rock walls and overhangs near the water, and
were once again very fast and difficult to catch.

On the basis of these observations O. seychellensis seems to be confined to vertical
wet rock faces, a preference quite distinct from that of many other Old World ochterids
which more typically occupy sandy horizontal substrates ajacent to streams and
ponds. It seems likely that this species is probably more widespread on Mahe than
the present records indicate, but that its obscure habitat and difficulty of capture
have precluded its collection.

Holotype. Male: SEYCHELLES, Mahe, seeping rock faces along Foret Noire road
near Congo Rouge trailhead, 390 m, 3 April 1989, CL 8047, D. A. Polhemus (USNM).

Paratypes. SEYCHELLES, Mahe: 15 66, 22 22, same data as holotype (USNM,
JTPC); 1 <5, headwaters of Riviere Grande Bois, along Foret Noire road near Congo
Rouge trailhead, 420 m, water temp. 22.5°C., 3 March 1989, CL 8016, D. A. Pol-

422

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

hemus (USNM); 1 6, Riviere Grand Bois, 6.2.74, F/Sey/4., F. Starmlihlner (JTPC).

Additional specimens not paratypes. SEYCHELLES, Mahe: 18 immatures (stored
in alcohol), same data as holotype (USNM).

Hebrus seychellensis, new species

Diagnosis. This species may be separated from other Hebrus presently known in
Africa, Madagascar and India by the following characters: the large keeled triangular
male proctiger thickly set with erect curving setae (Fig. 7); the distinctive male
paramere shape with a spur on the inner face (Figs. 5, 6); the bowed hind femur; the
unmodified hind tibia; the shape of the posterior margin of the bucculae; the setose
areas distally on the anterior and middle tibiae; the tumescent abdominal ventrite
VI in females; and the pattern of white markings on the hemelytra (Fig. 4).

Description. Macropterous male: elongate, body length/width (across humeri) =
2.02/0.77. Ground color dark brown; eyes reddish brown; rostrum dark yellow; lateral
portions of anterior lobe of pronotum dark brown; hemelytra dark blackish brown
with white patches basally on clavus to either side of scutellum, membrane fumate,
with 3 elongate white streaks posteriorly (Fig. 4); legs dark yellow brown, becoming
more infuscated on distal portions of femora and tibiae; venter of thorax medium
brown, venter of abdomen black.

Head long, declivant anteriorly, length (as measured from directly above)/width
(across eyes) = 0.40/0.43, width across antennal tubercles = 0.32; vertex lacking a
median sulcus. Eyes moderately large, exserted, globose, coarsely faceted; ocellar pit
located mesad of posterior portion of eye; ocelli of moderate size, evident, eye width/
interocular space = 0.09/0.24. Antennae long, slender, lengths of segments I-IV =
0.29, 0.20, 0.45, 0.27; segments I and II stoutest, of similar diameter, bearing short
pale recumbent setae, segments III and IV more slender, bearing longer erect pale
setae whose lengths exceed the segment diameters. Rostrum long, length = 0.88,
extending past hind coxae and onto base of abdomen. Bucculae long, prominent,
bearing 5 foveae, with 2 large foveae along dorsal margin adjoining gula and 3 smaller
foveae evenly spaced around the posterior buccular margin.

Pronotum long, length/width (across humeri) = 0.50/0.79, weakly constricted me-
dially, bearing median longitudinal sulcus. Anterior lobe with 2 (1 + 1) deep pits to
either side of longitudinal midline, anterior margin with prominent collar. Posterior
lobe tumid, raised above anterior lobe, lacking significant pits or depressions pos-
teriorly, humeri prominent. Scutellum length/width = 0.22/0.41; mesoscutellum short,
raised, length 0.05; metanotum moderately long, length/width = 0.16/0.29, with
weak longitudinal median carina, posterior margin raised to form a horizontal lip.

Hemelytra complete, attaining tip of abdomen; costal margin and corium thick-
ened, set with numerous fine recumbent gold setae.

Legs moderately stout, straight, all segments covered with short semi-recumbent
gold setae, fore and middle tibiae with thicker patches of setae distally on ventral
faces; proportions of leg segments as follows: fore femur/tibia/tarsal 1 /tarsal 2 =
1.10, 1.06, 0.80, 0.31; middle femur/tibia/tarsal 1/tarsal 2 = 1.06, 1.02, 0.08, 0.27;
hind femur/tibia/tarsal 1/tarsal 2 = 1.33, 1.57, 0.08, 0.47.

Venter of thorax weakly pruinose, sparsely set with short gold setae; abdominal
venter shining, more thickly covered with short recumbent gold setae, central portion

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HETEROPTERA FROM SEYCHELLES

423

of ventrite VI glabrous and broadly but distinctly tumescent, this tumescence most
evident when viewed laterally.

Genital segment of moderate size, mostly retracted into abdomen; proctiger tri-
angular, bearing an anteromedial keel-like structure thickly set with erect curving
setae (Fig. 7); parameres symmetrical, hooked at tips with spur on inner face, shape
as in Figures 5, 6.

Macropterous female: similar to macropterous male in general structure and col-
oration but slightly larger. Body length 2.15 mm, width (across humeri) 0.86 mm.

Discussion. The genus Hebrus is in serious need of revision in the Old World, and
the number of species in Africa, Madagascar and tropical Asia is so large that in-
dividual comparisons with H. seychellensis n. sp. would be impractical. The char-
acters given in the diagnosis and the geographical provinence should, however, be
sufficient to allow easy recognition of this species.

Etymology. The name “seychellensis” refers to the Seychelles islands.

Habitat data. The type series was taken along the Riviere Grande Saint Louis
above the Le Niol water intake, a clear rocky stream coming down out of a disturbed
primary rain forest catchment. The insects were very uncommon, and were found
only on the dark undersides of large midstream rocks which projected out over the
flowing water.

Holotype. Macropterous male: SEYCHELLES, Mahe, Riviere Grande St. Louis
above Le Niol water intake, 330 m, water temp. 22.5°C., 2 April 1989, CL 8046, D.
A. Polhemus (USNM).

Paratypes. 1 macropterous 6, 1 macropterous 9, same data as holotype (USNM,
JTPC).

ACKNOWLEDGMENTS

I wish to thank Brian Kensley of the Smithsonian Institution for allowing me the opportunity
to participate in the Aldabra Project, and the people of Seychelles for their kindness during my
stay in the country. Early drafts of this paper were read by Thomas J. Henry of the Systematic
Entomology Laboratory, USDA % National Museum of Natural History, and John T. Polhemus
of Englewood, Colorado, who also provided useful advice regarding comparisons with other
species in the Indian Ocean region.

LITERATURE CITED

Davies, D. 1968. When did the Seychelles leave India? Nature 220:1225-1226.

Polhemus, J. T. 1990. Miscellaneous studies on the genus Rhagovelia Mayr (Heteroptera:
Veliidae) in Southeast Asia and the Seychelles Islands, with keys and descriptions of new
species. Raffles Bull. Zool. 38:65-75.

Received 14 May 1991; accepted 21 October 1991.

J. New YorkEntomol. Soc. 100(3):424-479, 1992

A REVIEW OF THE SPECIES OF ANTHOCORIDAE
(HEMIPTERA: HETEROPTERA) FOUND ON
PINUS CONTORTA

J. D. Lattin and N. L. Stanton

Systematic Entomology Laboratory, Department of Entomology,

Oregon State University, Corvallis, Oregon 97331-2907; and
Department of Zoology and Physiology, University of Wyoming,

Laramie, Wyoming 82071

Abstract. — The Anthocoridae fauna found on the widespread western North American pine,
Pinus contorta, is reviewed. Ten genera and 18 species are reported. Three of these species
( Anthocoris spp. and Orius tristicolor ) are considered casual visitors to the tree. Biological,
ecological, and taxonomic information is presented for each species and the pertinent literature
reviewed.

Pinus contorta Dough, ex Loud, with four subspecies, is the most widespread species
of pine in North America (Critchfield, 1957, 1980, 1985; Critchfield and Little, 1966;
Wheeler, Guries, and O’Malley, 1983; McCune, 1988). Commonly known as lodge-
pole pine throughout much of its range, the northern coastal subspecies is referred
to as shore pine and the unusual dwarf subspecies is called pygmy pine. Forrest
(1980a, b) recognized a number of chemical regions for Pinus contorta, based upon
studies on the variation found in foliar monoterpenes. The subspecies occur in north-
ern Baja California, on the north coast of California, north through Oregon, Wash-
ington and British Columbia, in the Sierra Nevada and Cascade Mountains north to
the Yukon Territory, and south through the Rocky Mountains and adjacent areas to
southern Colorado (Critchfield, 1957). It grows from sea level in Oregon to 12,000
feet (3,900 m) in the southern Sierra Nevada mountains (Critchfield, 1980).

We have collected Hemiptera: Heteroptera from this tree species regularly at six
sites in Oregon and Wyoming. Other collections have been made at various localities
in different parts of the range of P. contorta, and to a lesser extent, on some of the
other species of conifers occurring with P. contorta. The family Miridae is particularly
well represented on this conifer (approximately 57 species known to date; unpublished
data), and so it seems are the Anthocoridae. The diverse habits of the various species
of anthocorids and their particular habitat requirements, often remarkably specific,
combined with the enormous ecological and geographical range of P. contorta result
in a diverse fauna of 10 genera and 18 species (24 genera and 90 species occur in
America north of Mexico [Henry, 1988]).

The objective of this paper was to coalesce information on the systematics, habitats,
plant hosts, prey species, and geographic distribution for those 18 species of North
American Anthocoridae found on Pinus contorta. The primary source of information
is based upon our regular sampling of three sites in Oregon on P. contorta contorta,
P. c. latifolia, and P. c. murrayana, and three sites on P. c. latifolia in Wyoming.
Additional records come from specimens examined in several collections and from

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ANTHOCORIDAE ON PINUS CONTORT A

425

the literature. We have summarized the available literature on each species treated,
including taxonomic, biological, and ecological information. Appropriate literature
records are presented in the condensed bibliographic style of taxonomic papers for
brevity.

SYSTEMATICS

The Anthocoridae comprise a small family of Hemiptera: Heteroptera with about
80 genera and 450 species world-wide (Pericart, 1972). Henry (1988) reported 23
genera and 89 species from America north of Mexico. An additional genus and
species, Brachysteles parvicornis (Costa), has been reported recently from the East
Coast where it was taken under the bark of pine trees and beaten from Picea and
Pinus branches (Asquith and Lattin, 1 990; Lattin and Asquith, 1991). Approximately
one-third of the species north of Mexico occur on conifers. Others are found on
broad-leafed plants or in leaf litter.

Pericart (1972) provided an extensive treatment of the Anthocoridae of western
Europe and permitted the comparison of elements of our fauna with those of the
European continent. Herring (1976) published a key to the genera of Anthocoridae
north of Mexico. Anderson (1962b) provided the most detailed summary of the
Anthocoridae of the Pacific Northwest. Kelton (1978) is the only modem, compre-
hensive treatment of the family for North America. Earlier, he published several
generic revisions of the North American species (Kelton, 1963, 1966a, 1967, 1976b)
as Herring (1966) did for Orius. Much additional biosystematic work remains to be
done on the Anthocoridae.

The higher classification of the Heteroptera is undergoing substantial change, with
increased emphasis on cladistic analysis (see Schuh, 1986, for an important review).
The suborder Cimicomorpha, which contains the Anthocoridae, is being reviewed
by several investigators. Major changes in classification of the Anthocoridae have
been suggested by Schuh (1986; pers. comm., April, 1991), based on evidence given
by Ford (M.S. thesis). Schuh and Stys (1991) provided a detailed account of some
of these changes. We have taken a rather conservative approach in this paper with
the recognition that future phylogenetic studies are certain to modify the current
definition of the family. Such information will aid in interpreting biological and
ecological data.

METHODS

We collected ten genera and 18 species on P. contorta. The genera and species
reported here are arranged alphabetically for ease of reference. Table 1 shows the
same taxa arranged by systematic scheme (Carayon, 1972a; Pericart, 1972; Henry,
1988).

Collection sites: We have collected regularly at three sites in Oregon and three sites
in Wyoming from April to October in 1986. Rather than repeat these localities in
full, we will describe them here and refer to them together with the dates of collection.
Collections made at other times and places are described in full. Branches of conifers
were firmly beaten with an axe handle and the dislodged anthocorids were collected
from the beating sheet held under the branch.

Oregon: Site 5: Lincoln Co., 1.6 km S Newport, South Beach State Park, R1 1W,

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Table 1 . Species of Anthocoridae collected from Pinus contorta subspp. arranged by current
classification. * Indicates incidental occurrence.

Anthocoridae

Anthocorinae Reuter, 1884
Anthocorini Reuter, 1884
Acompocoris lepidus (Van Duzee)

*Anthocoris antevolens White
*Anthocoris musculus (Say)

Elatophi/us dimidiatus (Van Duzee)

Elatophilus pullus Kelton and Anderson
Elatophilus sp. A
Melanocoris longirostris Kelton
Melanocoris nigricomis Van Duzee
Tetraphleps canadensis Provancher
Tetraphleps latipennis Van Duzee
Tetraphleps pilosipes Kelton and Anderson
Tetraphleps uniformis Parshley
Oriini Carayon, 1955
*Orius tristicolor (White)

Lyctocorinae Reuter, 1884
Dufouriellini Van Duzee, 1916
Cardiastethus borealis Kelton
Lyctocorini Reuter, 1884

Lyctocoris tuberosus Kelton and Anderson
Scolopini Carayon, 1954

Scoloposcelis flavicornis Reuter
Xylocorini Carayon, 1972
Xylocoris californicus (Reuter)

Xylocoris cursitans (Fallen)

T1 IS, SW'/4 Sec. 20. ex Pinus contorta contorta Dough ex Loud. Site 7: Crook Co.,
40 km E Prineville, 1,476 m elev., R19E, T13S, SW*/4 Sec. 2. ex Pinus contorta
latifolia Engelm. Site 8: Deschutes Co., Three Creeks Meadow, 2,069 m, 26 km S
Sisters, R9E, T17S, SW'/t Sec. 13 ex Pinus contorta murrayana (Grev. & Balf.)
Engelm.

Wyoming: Site 2: Albany Co., Medicine Bow Nat. For., Happy Jack Road, 19 km
E Laramie, 2,500 m, R72W, T1 5N Sec. 25. ex. Pinus contorta latifolia. Site 3: Albany
Co., Snowy Range Mountains, North Fork Camp ground. Sand Lake Road, 2,800
m, 60 km W. Laramie, R78W, T16N, Sec. 17, ex Pinus contorta latifolia. Site 4:
Carbon Co., Snowy Range Mountains, French Creek Road, 2,750 m, 90 km W.
Laramie, R81W, T15N, Sec. 1 & 17. ex. Pinus contorta latifolia.

Collections examined: All specimens are housed in the Systematic Entomology
Laboratory (OSU-SEL), Department of Entomology, Oregon State University, unless
otherwise indicated. Additional material was examined from the Canadian National
Collection, Ottawa, Canada (CNC), Oregon Department of Agriculture (ODA), Sa-
lem, and from the U.S. Forest Service, Pacific Northwest Research Station, LaGrande,
Oregon, collection (USFS-LaGrande). Supportive material was examined from the

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ANTHOCORIDAE ON PINUS CONTORT A

427

California Academy of Sciences, San Francisco, the University of California, Riv-
erside, and the United States National Museum, Washington, D.C.

The section that follows presents information about each species of Anthocoridae
found on or reported from Pinus contorta. A bibliography is given for each species
to provide access to the biological and systematic literature. Diagnostic characters
are included to assist in recognition. Information on habitat and host plant associ-
ation, prey, life history, and distribution is reviewed and summarized followed by
documentation of specimens collected and examined during this study.

BIONOMICS
Acompocoris Reuter

Acompocoris Reuter, 1875:63; Harris and Shull, 1944:207; Pericart, 1972:146; Her-
ring, 1976:146; Kelton, 1978:32; Henry, 1988:13.

Diagnosis: Normally macropterous (membrane of female may be slightly reduced);
shiny, usually pubescent; rostrum almost reaching or exceeding hind coxae; hind
coxae proximate; ostiolar canal broad, slightly curved in anterior direction, apex
extended anteriorly as fine carina (Pericart, 1972; Kelton, 1978). North American
species with head, pronotum, and scutellum, dark brown to black in contrast to pale
brown hemelytra.

Acompocoris Reuter contains six species in the Holarctic Region (Pericart, 1972).
Species of the genus are found only on conifers where they appear to feed on aphids,
especially members of the genus Cinara Curtis. Two occur in North America, A.
lepidus (Van Duzee) from California and the introduced European species, A. pyg-
maeus (Fallen). Host plant genera for North American Acompocoris include Pinus
(most common), Picea, Larix, and Abies (Pericart, 1972; Kelton, 1978). Only A.
lepidus occurs on P. c. murrayana. Acompocoris pygmaeus was reported from P.
sylvestris in Nova Scotia and New Brunswick (Kelton 1977a) and P. strobus (Kelton,
1978).

Acompocoris lepidus (Van Duzee)

(Figure 1)

Tetraphleps lepidus Van Duzee, 1921:142.

Acompocoris lepidus: Kelton and Anderson, 1 962: 1 307; Mitchell, 1 962:53; Anderson,
1962b: 1 332; Kelton, 1977a:245; Kelton, 1978:32; Henry, 1988:13.

Acompocoris sp. Harris and Shull, 1944:207.

Diagnosis: Macropterous species about 3.5 mm long; shiny with long pubescence;
apex of rostrum attaining or surpassing hind coxae. Color dark brown to black,
pronotum entirely black, hemelytra brown, shiny; antennal segment 2 dark, pale at
middle. Resembles species of Tetraphleps but apex of ostiolar canal not elevated
above metapleuron, continued forward as fine carina (apex of canal elevated above
metapleuron in Tetraphleps ).

Habitat: Little specific information is available on the preferred habitat of this
species, which was originally described from specimens collected on Pinus contorta
murrayana in California (Van Duzee, 1921); and later reported from P. ponderosa

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

and Picea engelmanni in British Columbia (Kelton, 1978). We report it here from
P. c. murrayana in the central Oregon Cascade Mountains and on P. c. latifolis in
British Columbia, the Yukon Territory, and in Wyoming.

Prey: Mitchell (1962) considered A. lepidus an effective predator of the balsam
woolly aphid.

Phenology and life history: Type specimens were collected in late July in southern
California. Adults were collected in July in Wyoming, August in British Columbia,

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ANTHOCORIDAE ON PINUS CONTORT A

429

Oregon, Wyoming, and Yukon Territory, and September in Colorado, Oregon, and
Wyoming. No information is available on the life history of this species.

Distribution: Acompocoris lepidus was described from Huntington Lake, 2,277 m,
Fresno Co., California (as a species of Tetraphleps ). Harris and Shull (1944) reported
an unidentified species of Acompocoris from McCall, Idaho (presumably A. lepidus).
Mitchell (1962) reported A. lepidus from the central Cascade Mountains of Oregon.
Anderson (1962b) reported the species from several localities in British Columbia
(Princeton on P. ponderosa and Cathedral Lakes on Picea engelmanni). Kelton (1977a)
compared A. lepidus with the newly collected A. pygmaeus (Fallen) from Europe, a
species found on Pinus sylvestris in Nova Scotia and New Brunswick. We report A.
lepidus here from the Yukon Territory, Colorado, Oregon, and Wyoming.

Specimens examined: CANADA: British Columbia: Summit Lake, 16 August
1982, P. c. latifolia, 1 male (CNC); Yukon Territory, Rancheria, 1 1 August 1982, P.
c. latifolia, 5 males, 1 female (CNC). UNITED STATES: Colorado: Gilpin Co., 3
km S Guanella Pass, 3,015 m, 4 September 1991, P. c. latifolia, 17 females; as before
but P. aristata Engelm., 7 females; Larimer Co., 1 km SW Milner Pass, 3,298 m,
R76W T5N, Rocky Mt. N. P., 3 September 1991, P. c. latifolia, 3 females. Oregon:
Deschutes Co., Three Crks. Mdw., 26 km S Sisters, 2,000 m, 19 September 1979,
P. c. murrayana, 1 female; Site 8, 6 August 1986, P. c. murrayana (trees 3, 19), 1
female, 1 male; Site 8, 26 August 1986, P. c. murrayana (general collecting) 1 female;
Site 8, 20 August 1986, P. c. murrayana (tree 3), 1 female; Site 8, 3 September 1986,
Picea engelmanni (10 trees pooled) 1 female. Wyoming: Albany Co., Site 2.1, 9 July
1986, Pinus flexilis (10 trees pooled), 3 males, 1 female; Site 2.1, 11 July 1986, P.
c. latifolia (tree 15), 1 male; Site 2.2, 25 July 1986, P. c. latifolia (tree 18), 1 female;
Site 2.1, 25 July 1986, P. flexilis (10 trees pooled), 1 1 females; Site 2.1, 5 September
1986, P. flexilis { 10 trees pooled), 1 female; Site 3.1, 11 July 1986, P. c. latifolia (tree
16), 2 males; Site 3.1, 30 July 1986, P. c. latifolia (trees 2, 3, 6, 15, 16, 18, 20), 31
females; Site 3.1, 27 August 1986, P. c. latifolia (trees 5, 13, 16), 4 females; Carbon
Co., Site 4.2, 17 July 1986, P. c. latifolia, 1 female. Park Co., Yellowstone Nat. Pk.,
16.2 km N Norris, 2,277 m, 16 September 1980. P. c. latifolia, 3 females; Park Co.,
Yellowstone Nat. Pk., 1.6 km N Indian Crk. Cmpgd., 19.5 km N Norris, 2,270 m,
16 September 1980, P. c. latifolia, 2 females; Yellowstone Nat. Pk., 8.1 km S Tower
Falls, 2,262 m, 17 September 1980, P. c. latifolia, 9 females.

Anthocoris Fallen

Anthocoris Fallen, 1814:9; Reuter, 1884:66; Lethierry and Severin, 1896:243; Van
Duzee, 1917a:292; Collyer, 1953:86; Hill, 1957: 17 1; Sands, 1957:296; Southwood
and Leston, 1959:173; Anderson, 1962a:67; 1962b: 1327; Anderson and Kelton,
1963:439; Cobben and Amoud, 1969:5; Carayon, 1972a:345; Pericart, 1972:1 10;
Herring, 1976:146; Evans, 1976a:85; 1976b:157; 1976c:163; 1976d:283; Kelton,
1978:34; Kerzhner, 1988:772; Henry, 1988:14.

Diagnosis: Length 2.4 to 4.6 mm, usually macropterous (except A. dimorphicus
Anderson and Kelton); pruinose or shiny; dorsal pubescence; usually black or brown
and white. Ocelli distinct; rostrum short, apex reaching or slightly surpassing front
coxae. Pronotum with distinct collar and calli; lateral margin carinate. Hemelytra

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smooth, pruinose or shiny. Hind coxae proximate. Ostiolar canal curved anteriorly,
apical portion attached to metapleura, continued in anterior direction as a carina.

Approximately 30 species of Anthocoris are known, and most occur in the Northern
Hemisphere (Lethierry and Severin, 1896; Pericart, 1972). Twelve species are known
from North America (Henry, 1988). No recent world monograph for the genus exists.
The best current treatment is that of Pericart (1972) who treats 14 species in the
western Palearctic Region. No revision is available for North America, but Hill (1957)
produced a key to the 1 1 species known to him. The best references to North American
species are those of Anderson (1962b) and Kelton (1978).

Species of Anthocoris occur most commonly on broad-leafed plants, particularly
on trees. Only two species in North America have been reported from conifers. They
are searching predators, moving over the surface of the branches and leaves (Collyer,
1953; Sands, 1957; Southwood and Leston, 1959; Anderson, 1962a, b; Cobben and
Amoud, 1969; Pericart, 1972; Evans, 1976a, b, c, d; Kelton, 1978). Kelton (1978)
listed host plants and summarized some of the literature dealing with five species
considered important predators. A considerable literature on this genus documents
the activities of the different species in agroecosystems (see e.g., Evans, 1976a, b,
c, d).

Anthocoris antevolens White
(Figure 2)

Anthocoris antevolens White, 1879:146; Reuter, 1884:77; Lethierry and Severin,

1896:243; Van Duzee, 1917a:293; b:262; Hill, 1957:172; Anderson, 1962b: 1327;

Anderson and Kelton, 1963:439; Kelton, 1978:37; Evans, 1983:45; Henry,

1988:14.

Diagnosis: Length 3. 6-4. 4 mm; macropterous, shiny; pubescence long and erect.
Pronotum black, shiny, basal half usually pale brown. Hemelytra with clavus black
except outer margin pale; basal third of corium and embolium pale, apical two-thirds
of corium, embolium and all of cuneus black; membrane pale with middle and apical
third fuscous.

Habitat: Anthocoris antevolens is an active, surface predator like A. musculus and
other species of Anthocoris. Evans (1983) reported this species on P. c. latifolia in
northwestern British Columbia. This species is a common predator in orchards
(Anderson, 1962b; Wilde and Watson, 1963; McMullen and Jong, 1967; Kelton,
1978). According to Van Duzee (1917a) and Anderson (1962b), willow is the main
host plant. Kelton (1978) added a number of host plants, mostly deciduous trees,
including fruit crops where it feeds largely on Homoptera.

Prey: Anderson (1962b) reported aphids and mites as prey of A. antevolens. He
documented this species biting humans, especially in orchards. Anderson and Kelton
(1963) reported predation on several species of Pemphigus aphids on poplar and
cottonwood and on the gall-forming aphid Mordwilkoja vagabunda (Walsh). Kelton
(1978) reviewed some of the literature on the feeding habits of this species and noted
that it feeds on psyllids (including the pear psylla) and mites. The reference to this
species on Pinus contorta latifolia by Evans (1983) did not include prey records.

Phenology and life history: Anderson (1958, 1962b) detailed the life history of A.

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ANTHOCORIDAE ON PINUS CONTORT A

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Fig. 2. Anthocoris antevolens White.

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antevolens. Individuals, usually fertilized females, overwinter under bark scales and
other parts of trees. The ratio of females to males was 8:1 in western Oregon. There
appeared to be three generations per year in the Pacific Northwest, but four gener-
ations per year are possible in the laboratory. The egg stage lasts about 5 days. The
duration of nymphal instars varies from 2.7 in the third- to 7.9 for fifth-instar females.
Adults are long lived, especially the females. Under laboratory conditions, a mated
female lived for 50 days. Naturally, the overwintering females live considerably
longer.

Distribution: Anthocoris antevolens is widely distributed across Canada and north-
ern United States, especially in the western part where it extends south along the
coast into California, its type locality. Within the range of P. contorta, it has been
recorded from Alaska, Alberta, British Columbia, California, Colorado, Idaho, Mon-
tana, Nevada, Oregon, and Wyoming. Much of its range overlaps that of A. musculus,
with the former more prevalent in the west and the latter in the east.

Specimens examined: Many specimens from throughout the range of the species.
The record of this species from P. contorta latifolia is that of Evans (1983) from
British Columbia.

Anthocoris musculus (Say)

Reduvius musculus Say, 1832:32; Reuter, 1884:164.

Anthocoris borealis Dallas, 1852:588; Drake, 1921:202; 1922:66; Blatchley, 1926:

635.

Anthocoris musculus: Uhler, 1876:321; Lethierry and Severin, 1896:244; Van Duzee,

1917a:293; Strickland, 1953:199; MacPhee and Sanford, 1954:129; Reid, 1957:

111; Hill, 1957:172; Anderson, 1962b:1329; Anderson and Kelton, 1963:439;

Kelton, 1978:40; Henry, 1988:14.

Diagnosis: Length 3.3-4. 1 mm; macropterous, shiny; pubescence short and scat-
tered, inconspicuous and appressed. Pronotum black, shiny; basal half pale colored,
especially on postero-lateral angles. Hemelytra with clavus black, pale along outer
margin; basal half of corium and embolium pale, apical half of corium, embolium
and all of cuneus, black; membrane pale with dark pattern in middle and all of apical
third.

Habitat: Kelton (1978) reported this species to be an active predator on a variety
of surface-dwelling arthropods. Drake (1921, 1922) reported this species (as A. bo-
realis) as an occasional visitor on conifers but stated it was more commonly found
on deciduous trees, particularly on the branches and leaves of willow. He reported
taking a few specimens under bark scales and in the burrows of bark beetles, including
a few specimens from burrows of bark beetles on Picea. In England, Hamilton (1978)
reported the occurrence of A. nemorum (L.), a similar species, under bark in the
winter in England, often occurring with Xylocoris. Drake (1922) expanded his ob-
servations on A. musculus (as A. borealis), again noting its more common occurrence
on willow where it feeds on lace bugs (Tingidae). Reid (1957) also reported A. mus-
culus from the galleries of Ips in Pinus contorta latifolia. Kelton (1978) reported the
species to be an active predator on a variety of surface-dwelling arthropods and added
many additional host plants.

Anderson (1962b) reported it from alder and willow in British Columbia as do

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two closely related species, A. dimorphicus Anderson and Kelton and A. antevolens
(Anderson and Kelton, 1963; Kelton, 1978). Kelton (1978) added many other genera
of deciduous plants as habitats and referred to its importance as a predator in or-
chards, especially apple orchards in eastern Canada. It appears that A. musculus is
only an occasional visitor to conifers, although its occurrence in bark beetle galleries
seems somewhat different from its usual surface-feeding habit.

Prey: Drake (1922) reported A. musculus feeding on several species of Corythucha
(Hemiptera: Tingidae) on willow. Reid (1957) reported predation on eggs of Ips (/.
pini Say and I. perroti Swain) on P. contort a latifolia in Alberta. It has been observed
feeding on red mites (Kelton, 1978), the eyespotted bud moth in apple orchards in
Nova Scotia (MacPhee and Sanford, 1954), and aphids (Strickland, 1953).

Phenology and life history: Little is known about this species except that it is a
predator in orchards, particularly apple orchards, in eastern Canada (MacPhee and
Sanford, 1954).

Distribution: Originally described from north-central United States, Anthocoris
musculus is widespread from the Maritime Provinces of Canada west to British
Columbia and Alaska and south to Oregon, Colorado, Kansas, Missouri, New York,
and North Carolina. Part of its range overlaps that of A. antevolens.

Specimens examined: We have examined many specimens from throughout its
range. The record for Pinus contorta is that of Reid (1957) from Alberta where it
occurred in the galleries of Ips pini and I. perroti Swain. There are two specimens
from Washington, Whatcom Co., Heather Meadows, R9E T39N Sec 19, 7 September
1979, Abies amabilis.

Cardiastethus Fieber

Cardiastethus Fieber, 1860:266; Van Duzee, 19 17a:295; Herring, 1966:150; Pericart,

1972:249; Kelton, 1977a:246; Kelton, 1978:57; Henry, 1988:22.

Diagnosis: Macropterous, densely pubescent, long hairs on pronotum and hemel-
ytra; antennal segments 3 and 4 hardly thinner than segment 2; pronotum punctate,
lateral margin carinate; collar narrow, disc of pronotum with distinct transverse
groove; ostiolar canal evenly curved forward and prolonged into carina that reaches
anterior margin of metastemum. Male with single clasper, female with poorly de-
veloped ovipositor (Pericart, 1972; Herring, 1976; Kelton, 1978).

Cardiastethus is a cosmopolitan genus containing about 45 species (Pericart, 1972).
Six species occur in the United States and Canada; all but one of these are found in
the southern United States (Henry, 1988). A single species, C. borealis Kelton, is
northern in distribution and may be the only species occurring on a conifer in North
America. One European species, C. fasciiventris (Garbiglietti), occurs on conifers
{Abies and Picea ), but also is found on broad-leaf trees where it feeds on Psocoptera
(Pericart, 1972). According to Southwood and Leston (1959), this latter species is
mainly subcorticular and overwinters as an adult. There is a single generation per
year. Perris (cited in Pericart, 1972) reported nymphs from male flowers of pine in
France. It will eat aphids in captivity. Another European species feeds upon scale
insects.

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Cardiastethus borealis Kelton
(Figure 3)

Cardiastethus borealis Kelton, 1977a:246; Kelton, 1978:57; Henry, 1988:22.

Diagnosis: Small (2.25-2.70 mm), reddish-brown species, i.e., pronotum and hem-
elytra densely pubescent. Pronotum, scutellum and cuneus dark, most of wing pale.
Habitus similar to species of Orius, but larger, brown rather than black, and more
pubescent. Ostiolar canal distinctive, long and slender, evenly curved forward, ter-
minal carina reaching anterior margin of metastemum. Single, sickle-shaped male
clasper.

Habitat: Little information is available on the microhabitat of C. borealis. It has
been collected on Pinus ponderosa in British Columbia, P. banksiana in Manitoba,
and P. sylvestris in Nova Scotia (Kelton, 1977a, 1978). We record it here from P.
contorta latifolia in central Oregon (Ochoco Mountains). Pinus ponderosa occurs at
this locality in some abundance. The 1965 Oregon specimen, from northwest of
Lakeview, lacked host information but probably occurred on P. ponderosa (J.D.L.).

Prey: No prey information is available for C. borealis. The single European species
of Cardiastethus [C. fasciiventris (Garbiglietti)] occurring on conifers (. Abies and Picea ),
is reported to feed on Psocoptera (Pericart, 1972). A nonconifer inhabiting species
in Europe is reported to feed on scale insects (Pericart, 1972).

Phenology and life history: Southwood and Leston (1959) reported that the British
species of Cardiastethus overwinters as an adult. Perris (cited in Pericart, 1972) stated
that the nymphs of C. fasciiventris occurred on the male strobili of pine in France.
The early appearance of the adults on the strobili suggests they overwinter as adults
or as eggs (e.g., most Miridae). The original type series (Kelton, 1977a) contained
only a single male collected in early June in British Columbia. The other six speci-
mens, all females, were collected in June (British Columbia and Manitoba) and July
(Nova Scotia). The two specimens recorded here from central Oregon, a male and a
female, were collected in July and August, respectively. Much remains to be learned
about the habits of this species. Limited evidence suggests most species of Cardiasteth-
us occur on deciduous trees: only two are found on conifers, one in Europe and the
other in North America. C. borealis appears to occur on different hosts in different
parts of its range.

Distribution: The type locality of C. borealis is Aspen Grove, British Columbia
(Kelton, 1977a). Included in the type series were specimens from Seddon’s Comer
and Telford, Manitoba, and Mt. Uniacke, Nova Scotia. To these records we add two
localities from central Oregon. A comprehensive systematic review of this genus is
needed.

Specimens examined: Oregon: Crook Co., Ochoco Mts., Site 7, 9 July 1986, Pinus
c. latifolia (tree 13), 1 male; Klamath Co: L. Elder Ck., 48.7 km W Lakeview, 18
August 1965, 1 female.

Elatophilus Reuter 1884

Elatophilus Reuter, 1884:616; Blatchley, 1928:86; Sands, 1957:303; Southwood and
Leston, 1959:173; Pericart, 1967:52; Biliotti and Riom, 1967:1 103; Cobben and
Amoud, 1969:8; Carayon, 1972a:345; Pericart, 1972:97; Herring, 1976:146; Kel-

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ANTHOCORIDAE ON PINUS CONTORT A

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ton, 1976b:631; Kelton, 1977b: 10 17; Kelton, 1978:19; Henry, 1988:15; Mendel,
Carmi, and Podoler, 1991:502.

Xenotracheliella Drake and Harris, 1926:38 (Synonymized by Kelton and Anderson,
1962: 1306).

Diagnosis: Length 2.5-4 mm, flattened, usually dark with some pale markings,
pubescence short. Head elongate, eyes prominent (especially in male). Antenna with

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segment two linear or enlarged. Apex of rostrum extending to or beyond anterior
coxae. Pronotum with poorly developed collar, dorsal surface roughened. Hemelytra
dull, dark, usually marked with pale, macropterous or brachypterous (males and
females), membrane with four veins. Ostiolar canal appressed to metapleuron, curved
forward, apex rounded, hind coxae widely separated. Male clasper short, broad and
curved.

Elatophilus Reuter contains 18 species distributed throughout the Holarctic Re-
gion, which are divided about equally between the Old and New World (Hiura, 1966;
Pericart, 1972; Kelton, 1976b, 1977b, 1978; Mendel, Carmi, and Podoler, 1991).
Seven of the eight described Nearctic species occur in America north of Mexico
(Henry, 1988). Species of Elatophilus occur only on conifers and almost exclusively
on the genus Pinus (Blatchley, 1928; Pericart, 1972; Kelton, 1976b; Mendel, Carmi,
and Podoler, 1991). Elatophilus appears to be the conifer inhabiting analogue of the
genus Temnostethus Fieber, a genus found on deciduous trees in the Palearctic
(LeQuesne, 1955).

Where known, species of the genus Elatophilus appear to have a special relationship
with species of the scale genus Matsucoccus Cockerell (Homoptera: Magarodidiae)
(Morrison, 1939; McKenzie, 1942a, b; Kosztarab and Kozar, 1988). Biliotti and
Riom (1967) reported that E. nigricornis (Zett.) preyed on Matsucoccus feytaudi Due.,
a pest of pine in France. These two authors cited unpublished observations of J.
Carayon reporting the anthocorid from male strobili of Pinus sylvestris in the spring
in France. Sands (1957) reported the same bug feeding only on aphids occurring on
P. sylvestris in Britain. Cobben and Amoud (1969) recorded E. nigricornis from The
Netherlands and called attention to the discrepancy in the observed feeding reported
by Sands with that of Biliotti and Riom. Cobben and Amoud also pointed out the
likelihood of associations between other species of Elatophilus and scales of the genus
Matsucoccus. Pericart (1967) and Woodroffe (1969) described morphological differ-
ences among populations of E. nigricornis in Scotland and southern England. It is
possible that such differences may extend to prey preference as well.

Mendel, Carmi, and Podoler (1991) summarized information on the 18 known
species of Elatophilus and pointed out the presumed relations with species of the
scale genus Matsucoccus and species of the conifer genus Pinus. Some corrections
are needed on the information they provided for the eight North American species;
these will be published elsewhere except for the species treated in our paper. They
report confirmed observations of scale predation for five Palearctic and three Nearctic
species, however, only E. inimicus (Drake and Harris) has been documented in North
America as feeding on several species of Matsucoccus (Lussier, 1965).

In North America, Bean and Godwin (1955), Hartzell (1957) and Doane (1965)
associated E. inimica with the red pine scale, M. resinosae Bean & Godwin, in New
England. Lussier (1965) wrote a thesis on E. inimica and its role as a predator of the
scale. He discussed the life history of this anthocorid in detail, and found a close
correlation between the presence of the prey scale and the occurrence of the bug on
the tree. Although Lussier found the anthocorid associated chiefly with M. resinosae
on P. resinosa, he did cite one occurrence of the bug on P. rigida in association with
M. gallicolus and on P. sylvestris in association with aphids and mites without the
apparent presence of specimens of Matsucoccus. Drooz (1985) recorded the associ-
ation of E. inimica with both species of Matsucoccus and summarized the life history

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of both scale species. The anthocorid was first reported feeding on M. resinosae in
plantations only outside the natural range of Pinus resinosa. Anderson, Ford, Kegg,
and Risley (1976), among others, have suggested that M. resinosae may be an intro-
duced species in North America since the scale is very abundant on plantations of
the pine outside the tree’s natural range.

Direct observations of predation upon species of Matsucoccus by species of Ela-
tophilus have not been made in western North America. Several species of Matsu-
coccus occur on western species of pine, including Pinus contorta. Fumiss and Carolin
(1977) described host plants, seasonal history, and distributional information of the
western species of Matsucoccus. Ray and Williams (1984) provided a key to North
American species and gave additional host data. Anderson (1958) reported the as-
sociation of E. pullus Kelton and Anderson (as Elatophilus sp. b) with the aphids
Essigella fusca Gillette and Palmer and Cinara sp., and with the pine needle scale,
Chionapis ( Phenacaspis ) pinifoliae (Fitch), on P. ponderosa in British Columbia. He
reared E. pullus on the aphid Myzocallis coryli (Goeze) in the laboratory (1958,
1962b). We have encountered species of Essigella and Cinara on trees from which
specimens of Elatophilus have been collected. A number of other potential predators
of these aphids were taken at the same time, including Miridae and other species of
Anthocoridae.

Elatophilus dimidiatus (Van Duzee)

(Figure 4)

Anthocoris dimidiatus V an Duzee, 1921:139; Hill, 1957:173.

Elatophilus dimidiatus: Kelton, 1976b:632; Mendel, Carmi, and Podoler, 1991:503.
Elatophilus (Elatophilius) dimidiatus: Henry, 1988:15.

Diagnosis: Length 3.0 mm. Males macropterous, females macropterous or bra-
chypterous; ocelli present in all forms and sexes. Head with apex of rostrum extending
to middle of mesosternum. Basal half of hemelytra white or pale, including most of
the clavus, corium, and embolium, apical half brown to black, wing membrane pale
on basal half, dark brown to black on apical half; pubescence of wing moderately
dense and long.

Habitat: Conventional beating of branches and small trees usually produced few
specimens. Very rigorous beating with the edge of a straight axe handle produced
more. Branches and trunks with scaly bark seemed most productive. The only re-
corded host for E. dimidiata is Pinus contorta murrayana (Grev. and Balf.) Engelm,
the Sierra lodgepole pine (Little, 1979). Forrest (1980b) provided information about
the monterpenes of P. contorta oleoresin and defined some geographical areas based
upon these chemical characteristics, including one he calls the Sierra Nevada Division
that encompasses many of the collection sites of E. dimidiatus. All specimens that
we have collected were beaten from the branches of P. c. murrayana.

Prey: Although Mendel, Carmi, and Podoler (1991) cite Matsucoccus spp. as the
prey of E. dimidiatus no specific host prey have been observed or identified thus far.
Pinus contorta hosts at least one species of Matsucoccus, its presumed prey, as well
as the aphids belonging to Cinara and Essigella.

Phenology and life history: This species appears to overwinter as a nymph or an
adult. The males seem to disappear much earlier than the females (see Pericart,

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Fig. 4. Elatophilus dimidiatus (Van Duzee).

1967). Males and females were collected above Big Bear Lake, California, at an
elevation of 2,215 m on June 8, 1989; a male was collected at an elevation of 2,954
m, west of Big Pine, California on June 6, 1 989; brachypterous females were collected
on June 23, 1981 at Childs Meadow, California (1,538 m) (Fig. 4); and the type series

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(only females) from Cayton, California was collected on July 15, 1918. Near Sisters,
Oregon; only females were collected on July 6, 1989.

Distribution: Elatophilus dimidiatus was described by Van Duzee (1921) (as a
species of Anthocoris) from the Felch Ranch, near Cayton, Shasta Co., California
based on three females. The type is mature and macropterous, the second specimen
is teneral and macropterous, and the third, described by Van Duzee as a nymph, is
actually a brachypterous female. As indicated in the section on wing polymorphism,
species of this genus have always been regarded as macropterous in contrast to its
deciduous tree analogue, Temnostethus. This brachypterous specimen of E. dimidia-
tus was subsequently included as a paratype of Temnostethus fastigiatus Drake and
Harris (1926). In their paper (Drake and Harris, 1926), they erroneously cited the
collection date of the specimen as 1913 rather than 1918. The taxonomic problems
resulting from a mixed-type series will be dealt with in a subsequent publication.

As currently recognized, E. dimidiatus is known to occur above Big Bear Lake
(San Bernardino Co., California); west of Big Pine (Mono Co., California); near Child’s
Meadow (Tehama Co., California); the type locality, Cayton (Shasta Co., California);
and northward along the east side of the Cascade Mountains of Oregon at least as
far as Sisters (Deschutes Co.). Brachypterous specimens, all females, are thus far
known only from the San Bernardino, Tehama, and Shasta Co., California, localities.

Specimens examined: California: Mono Co., Cienega Mirth, 19.5 km W Big Pine,
6 June 1989, Pinus c. murrayana, 1 macropterous male; San Bemadino Co., above
Big Bear Lake, end FS2N1 1, Champion tree site, R1W T2N NW lA Sec 34; 2,215
m, 8 June 1989, P. c. murrayana, 4 macropterous males, 12 brachypterous females;
Tehama Co., Brokenshire Picnic area, 3.2 km W Child’s Meadow, 1,754 m, 23 June
1981, P. c. murrayana, 3 brachypterous females. Oregon: Deschutes Co., Jet Hwy
20 and Metolius road, 1 ,077 m, 6 July 1 989, P. c. murrayana, 6 macropterous females;
9. 1 km NW Sisters, Hwy 20, 1 ,0 1 5 m, 26 August 1 989, P. ponderosa, 2 macropterous
females; as above, 14 September 1989, P. ponderosa, 1 macropterous female; as
above, 26 September 1989, P. ponderosa, 2 macropterous females; 4.9 km S Sisters,
1,066 m, 14 September 1989, P. ponderosa, 1 macropterous female; as above, 24
September 1989, P. ponderosa, 1 brachypterous female; 14.6 km NW Sisters, Jet
Hwy 20 and FS 2,060, 1,031 m, 6 July 1989, P. c. murrayana, 6 macropterous
females; as above, 16 August 1989, P. ponderosa, 1 macropterous female; as above,
29 August 1989, P. ponderosa, 1 macropterous male, 1 macropterous female; as
above, 14 September 1989, P. ponderosa, 2 macropterous, 1 brachypterous female;
as above, 26 September 1989, P. ponderosa, 1 macropterous female.

Elatophilus pullus Kelton & Anderson
(Figure 5)

Elatophilus pullus Kelton & Anderson, 1962:1306; Anderson, 1962b: 1332; Kelton,

1978:22; Mendel, Carmi, and Podoler, 1991:503.

Elatophilus ( Elatophilius ) pullus: Henry, 1988:16.

Diagnosis: Length 2.6 to 3.0 mm. Males and females macropterous. Head with tip
of rostrum reaching middle of mesostemum. Hemelytra dark, dull; bases of corium
and embolium and middle of clavus often pale; membrane fumate, transparent;
pubescence silvery, sparse, short and semi-erect.

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Fig. 5. Elatophilus pullus Kelton and Anderson.

Habitat: Elatophilus pullus occurs on Pinus ponderosa from British Columbia south
to Oregon (Evans, 1983). The type series was collected from P. ponderosa. The
exception is the John’s Prairie, Washington site where we collected it from P. contorta.
This locale, northwest of Olympia, is on the west side of the Cascade Mountains at
the southern end of the Puget Sound. Pinus ponderosa occurs very rarely on the west

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side of the mountains in Washington (Critchfield and Little, 1966). According to
Forrest (1980b), the populations of P. contorta contorta in the Puget Sound area are
discrete chemically and not particularly similar to the populations to the north or to
the south. The trees looked more like the subspecies P. c. murrayana or P. c. latifolia
than the coastal subspecies. No Elatophilus have yet been taken farther south from
the coastal subspecies, P. c. contorta. Kelton (1978) lists Pinus flexilis James and
Picea glauca (Moench) Voss as additional hosts.

Prey: Although the aphids Essigella and Cinara are taken regularly with E. pullus
and other predatory Heteroptera, the relatively small numbers of the anthocorids
collected suggest that the prey may be scale insects of the genus Matsucoccus. Fumiss
and Carolin (1977) provided host tree information on western species of Matsucoccus
that differed from the scale species cited by Mendel, Carmi, and Podoler (1991) as
possible prey for E. pullus.

Phenology and life history: According to Anderson (1962b), E. pullus overwinters
as a fifth-instar nymph in the region around Summerland, British Columbia. Second
and third instar nymphs were taken in mid-November and only fifth-instar nymphs
were taken the following March. Males appeared first at the beginning of April,
followed by the females about two weeks later. The males died within a month. Eggs
were deposited during late April and hatched in early May. The occurrence of early
instars in September suggested the possibility of a second generation. The series we
collected from John’s Prairie near Olympia, Washington, in early July from P. con-
torta, were all females. This bug has not yet been collected from the coastal form of
P. contorta further south where we have collected extensively. In the vicinity of
Sisters, Oregon, at an elevation of 1,046 m, we collected females of E. pullus on P.
ponder osa on May 3, June 7, July 6, and August 2, 1989; fifth-instar nymphs from
June 7 to August 16 (Fig. 5), and third-instar nymphs from July 6 to August 2, 1989.
The early occurrence of females at this locale, and the absence of males suggests that
fertilized females overwinter. A rather protracted generation may occur through the
summer, with the males and females appearing in late August.

Distribution: Elatophilus pullus was described by Kelton and Anderson ( 1 962) from
specimens collected in British Columbia and Oregon. The type came from Westbank,
British Columbia. Anderson (1962b) reported the species from Westbank and Sum-
merland, British Columbia, and Adel, Chandler State Park, and Klamath Falls,
Oregon, all from P. ponderosa. We have seen additional specimens from John’s
Prairie, near Shelton, Washington, and in the vicinity of Sisters, Oregon. With the
exception of the John’s Prairie record, all localities are on the east side of the Cascade
Mountains from southern British Columbia, south to at least Klamath Falls, Oregon,
near the Oregon-Califomia border. At Sisters, Oregon, it occurs at the same locality
as E. dimidiata, although on different host trees.

Specimens examined: CANADA: British Columbia: Summerland, 1 April 1958,
P. ponderosa, 1 male; Summerland, 5 November 1957, lab reared ex P. ponderosa,
1 male; 16 km W Westbank, 26 June 1957, P. ponderosa, 1 female; Wilson’s Landing,
14.6 km N Westbank, 6 May 1958, P. ponderosa, 2 females; as above, but 12 June
1956, 1 female. UNITED STATES: Oregon: Deschutes Co., 14.6 km NW Sisters,
Jet Hwy 20 and FS 2060, 1,031 m, 7 June 1989, P. ponderosa, 1 female; as above,
20 July 1989, P. ponderosa, 1-IV instar nymph; as above, 16 August 1989, P. pon-
derosa, 1-V instar nymph; 4.9 km S Sisters, 1,066 m, 3 May 1989, P. ponderosa, 2
females; as above, 11 May 1989, P. ponderosa, 1 female; as above, 24 May 1989,

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P. ponderosa, 1 female; as above, 7 June 1989, P. ponderosa, 1-IV instar nymph; as
above, 2 August 1989 ,P. ponderosa, 1 -II, 1 -III instar nymphs; as above, 14 September
1989, P. ponderosa, 1 female; 9.1 km NW Sisters, Hwy 20, 1,015 m, 7 June 1989,
P. ponderosa, 2-IV, 2-V instar nymphs; as above, 21 June 1989, 5-V instar nymphs;
as above, 6 July 1989, 3 females; as above, 20 July 1989, 1 -III, 1-IV instar nymphs;
as above 2 August 1989, 1 female; as above, 16 August 1989, 3-V instar nymphs;
as above, 26 September 1989, 2 females. Washington: Mason Co., John’s Prairie, nr
Shelton, 2 July 1988, P. contorta, 4 females. Wyoming: Park Co., Yellowstone Nat.
Pk., 1.6 km N Indian Crk. Cmpgd., 19.5 km N Norris, 2,269 m, 16 September 1980,
P. contorta latifolia, 2 females.

Elatophilus sp. A.

An undescribed species of Elatophilus from Pinus contorta is referred to here as
species A; it will be formally described when the generic revision of Elatophilus, now
in progress, is completed. Species A may be separated from the other species of
Elatophilus occurring on P. contorta by the enlarged second antennal segment. All
specimens thus far collected have been macropterous.

Habitat: One specimen was taken on Picea engelmanni on Washington Pass, Wash-
ington; the other specimens have come from Pinus contorta murrayana in Oregon
and northern California. No specific prey information is available other than the
usual aphids beaten from the same hosts. Scale insects have not yet been collected
from P. contorta, although at least one species of Matsucoccus is known to occur
on it.

The specimens we collected were all females. The collection dates (from north to
south) were Washington Pass, Washington, late September; southern Oregon Cas-
cades, late June and mid-July; northern California, mid-July and mid- August. No
nymphs have been collected that can be referred to this species with certainty.

Distribution: The species occurs in the Cascade Mountains from northern Wash-
ington (Washington Pass) south into northern California (near McCloud) and in the
Sierra Nevada of California (near Calpine, Sierra Co.). Thus, it occurs in several
different regions of P. contorta murrayana, as defined chemically by Forrest (1980b).

Specimens examined: California: Sierra Co., Plumas Nat. For., Hwy. 89, 5 km W
Calpine, 1,631 m, 15 July 1987, P. c. murrayana, 1 female; Siskiyou Co., 24.4 km
NNE McCloud, Swamp Creek Road and Hwy. 19, 14 August 1988, P. c. murrayana,
3 females. Oregon: Deschutes Co., 14.6 km NW Sisters, Jet 20 and FS 2060, 1,031
m, 6 July 1989, P. c. murrayana, 1 female; as above but 29 August 1989, 1 male;
14 September 1989, 1 female; 4.9 km S Sisters, 1,066 m, 14 September 1989, P.
ponderosa, 1 male, 2 females; Harney Co., 16 km W Denio, 23 September 1959, 1
female; Klamath Co., 1.6 km W Crescent, 1,385 m, 17 July 1979, P. c. murrayana,
1 female; Klamath Co., nr Klamath Falls (above Geary Ranch), 26 June 1959, 1
female. Washington: Okanogan Co., Washington Pass, 1,615 m, 20 September 1979,
Picea engelmanni, 1 female.

Lyctocoris Hahn

Lyctocoris Hahn, 1836:19; Reuter, 1884:6; Van Duzee, 19 17a:228; Sands, 1957:307;

Kelton and Anderson, 1962:1302; Anderson, 1962b: 1325; Kline and Rudinsky,

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1964:8; Kelton, 1966b:320; Kelton, 1967:807; Chu, 1969:1; Carayon, 1972a:337;

Pericart, 1972:200; Herring, 1976:149; Kelton, 1978:57; Dixon and Payne, 1979:

12, 13; Berisford, 1980:250; Goyer, Lenhard, Nebeker, and Schmitt, 1985:4;

Kerzhner, 1988:770; Henry, 1988:23.

Diagnosis: Macropterous; length 3.30-4.90 mm; slightly flattened, shiny, pale to
dark brown, pronotum and hemelytra punctate. Antennal segments three and four
slender, setose; tip of rostrum reaching or exceeding middle coxae. Anterolateral
margins of pronotum expanded, collar present and narrow, calli distinct. Hemelytra
punctate and pubescent, four veins in membrane. Ostiolar canal slender, angulate,
apex pointing forward. Left and right claspers present; ovipositor well-developed.

About 25 species of Lyctocoris have been described (Chu, 1969; Pericart, 1972);
eight occur in the United States and Canada. The genus is distributed over much of
the world. At least one species, L. campestris (Fabricius), is almost cosmopolitan,
occurring very commonly in stored grain facilities. The extensive movement of grain
throughout the world may be responsible for its wide distribution. This species also
occurs in bird and mammal nests, hay stacks and thatch (Sands, 1957; Hicks, 1959).
While largely predaceous on other arthropods in these habitats, it has been reported
biting a person in New Zealand (Woodward, 1951; Anderson, 1962b). Anderson
(1962b) reported it from a hen house, on apple trees, and in grain storage facilities.
He recorded its occurrence under bark scales of maple and oak trees (fifth-instar
nymphs and adults) during the winter in southern England. Moore (1972) reported
L. campestris as a predator of the southern pine beetle. Kinzer (1976) reported this
species from Abies concolor in New Mexico. This identification needs to be confirmed.

Dixon and Payne (1979) and Goyer, Lenhard, Nebeker, and Schmitt (1985) re-
ported L. elongatus Reuter under pine bark in association with southern pine beetles,
chiefly Dendroctonus frontalis Zimmerman but including three species of Ips as well.
The species of Ips arrived slightly later to the trees than did Dendroctonus frontalis,
and Lyctocoris elongatus (Reuter) showed a closer correlation with the Ips spp. than
with D. frontalis. Kline and Rudinsky (1964) reported a Lyctocoris sp. as a possible
predator of the Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins.

Kelton (1967) published a synopsis of the genus Lyctocoris for North America,
providing information on the nine known species. He discussed the habits of some
of the species treated. Most species are found under the bark of a wide variety of
trees. Thus far, only a single species of Lyctocoris has been collected from P. contorta.

Lyctocoris tuberosus Kelton and Anderson
(Figure 6)

Lyctocoris elongatus: Downes, 1927:1 1.

Lyctocoris tuberosus Kelton and Anderson, 1962:1303; Anderson, 1962b: 1326; Kel-
ton, 1967:812; Kelton, 1978:62; Evans, 1983:45.

Lyctocoris (Lyctocoris) tuberosus: Henry, 1988:24.

Diagnosis: Length 3.6 to 3.9 mm. Head and pronotum reddish brown, scutellum
dark brown, hemelytra yellow brown tinged with reddish brown; membrane clear,
outer vein well developed, other three faint; legs pale yellow brown. Tip of rostrum
reaching hind coxae. Male with front and middle tibiae with a row of short, brown

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Fig. 6. Lyctocoris tuber osus Kelton and Anderson.

pegs on inner surface, hind tibiae with several long spines; abdominal segment seven
with small tubercle on each side; left clasper flattened, apex obliquely truncate, right
clasper smaller. Female with median portion of abdominal sternum six extended in
posterior direction, inner margin of abdominal sternum seven gently curved, posterior
margin almost straight, postero-medial angle rounded. Ovipositor well developed.

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Habitat: Most records provide only host plants, but Kelton (1967) stated that the
species occurs under the bark of P. ponderosa, P. contorta, and Picea engelmanni in
association with bark beetles. Kline and Rudinsky (1964) reported Lyctocoris sp.
from under the bark of Pseudostuga menziesii, also in association with bark beetles.

The holotype was collected from P. ponderosa at Aspen Grove, British Columbia.
Other host plants listed in the type series included P. contorta in British Columbia,
Picea engelmanni in British Columbia, P. ponderosa in British Columbia and Col-
orado, and Pinus monticola in British Columbia. Anderson (1962b) added P. pon-
derosa in northeast Washington, Pseudotsuga menziesii (Mirbel) Franco in Oregon
and Pinus contorta var. latifolia Engelmann in Oregon (note: this should be P. c.
murrayana ). Evans (1983) cited P. ponderosa as a host in southeastern British Co-
lumbia.

Prey: Kelton (1967) listed a species of the bark beetle genus Ips occurring on several
species of Pinus and on Picea engelmanni as potential prey. Kline and Rudinsky
(1964) cited a species of Lyctocoris as a possible predator of the Douglas-fir bark
beetle, Dendroctonus pseudostuga, in the Pacific Northwest. Evans (1983) cited L .
tuberosus as a “small active predator.” Mosier, Thatcher and Pickard (1 97 1) reported
L. elongatus (Reuter) as an associate of D. frontalis and Ips spp. Goyer, Lenhard,
Nebeker and Schmitt (1985) reported L. elongatus (Reuter) as a predator of southern
pine bark beetles.

Phenology and life history: Very little is known about this species of Lyctocoris.
Adults have been collected from April 28 through August 31 in southeastern British
Columbia with both sexes being taken in late August. Specimens from southwestern
Colorado were collected on April 23 (Kelton and Anderson, 1962). How any species
overwinters is not yet known. No information is yet available on the immature stages.

Distribution: The type series contained specimens from Aspen Grove, Femie,
Loma, Midday Valley, Merritt, Kane Valley and Trinity Valley, British Columbia,
and from Pine River, La Plata Co., Colorado (Kelton and Anderson, 1 962). Anderson
(1962b) added Northport, Stevens Co., Washington, and Keno, Elsie and Pringle
Falls, Oregon. He also stated the 1927 record of Downes of L. elongatus (Reuter)
from Midday Valley, British Columbia, actually referred to a specimen of L. tubero-
sus. Kelton (1967) added South Dakota, with no specific locality. This is a widespread
species, but with few known localities within that range.

Specimens examined: CANADA: British Columbia: Aspen Grove, 15 June 1933,
P. ponderosa, #17480, lot 23, 1 female; Stanley, 26 June 1932, Abies lasiocarpa,
#17424, lot 6, 1 female. UNITED STATES: Oregon: Jefferson Co., 3 km S Camp
Sherman, 20 October 1962, under P. ponderosa bark, 1 female.

Melanocoris Champion

Melanocoris Champion, 1900:329; Van Duzee, 191 7a:8 1 9; Kelton and Anderson,

1962:1307; Herring, 1976:144; Kelton, 1977a:246; Kelton, 1978:23; Henry,

1988:16.

Diagnosis: Macropterous; shiny; pubescence on hemelytra pale, flattened, scale-
like or elongate and simple, may be serially arranged. Head short, tip of rostrum
extending beyond anterior coxae; pronotum with collar indistinct, lateral margins
broadly expanded; ostiolar canal short, straight, apex elevated; hind coxae widely
separated, metastemum with apex straight or evenly rounded.

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The genus Melanocoris contains four described species, all found in North America.
The type of the genus is M. obovatus Champion, described from a female collected
at Totonicapam, Guatemala (Los Altos region), between 8,500 and 10,500 feet, likely
from a pine tree (Champion, 1900). The other three species occur in western Canada
and United States (Drake and Harris, 1926; Kelton and Anderson, 1962; Kelton,
1978; Henry, 1988). Species of Melanocoris occur only on conifers, chiefly species
of Pinus but occasionally on Picea and Abies (Champion, 1900; Van Duzee, 1921;
Drake and Harris, 1926; Kelton and Anderson, 1962; Anderson, 1962b; Kelton,
1977a, 1978).

Melanocoris longirostris Kelton

Melanocoris longirostris Kelton, 1977a:246; Kelton, 1978:25; Henry, 1988:16.

Diagnosis: Length 2.60-3.00 mm, head and pronotum black, shiny; hemelytra
brown, punctate, shiny, pubescence short and simple. Tip of rostrum extending
beyond hind coxae.

Habitat: Kelton (1977a) reported M. longirostris from Pinus monticola in British
Columbia; P. ponderosa and P. strombiformis in Arizona; and Pinus flexilis, Picea
glauca, and Abies lasiocarpa in Colorado. We have collected it on P. c. latifolia and
P. flexilis in Colorado; P. c. murrayana and P. monticola in Oregon; and P. c. latifolia
and P. flexilis in Wyoming.

Prey: No specific prey information is available for this species. It was collected
with woolly aphids on P. monticola in Corvallis, Oregon.

Phenology and life history: Adults have been collected in June, July, and August
throughout the known range of the species. No specific life history information is
available although both nymphs and adults were collected on July 25, and adults
only on July 27 at site 2 in Wyoming.

Distribution: The species was described from a series of specimens collected at
New Denver, British Columbia, and other localities in British Columbia, Colorado,
New Mexico, Arizona and Utah. We add localities from eastern and western Oregon,
Nevada, and southeastern Wyoming. The various host plants are listed under habitat.

Specimens examined: California: San Bernardino Co., above W end Big Bear Lake,
R1W T2W NW'/t Sec 34, 2,215 m, 8 June 1989, champion P. c. murrayana, 3
females. Colorado: Larimer Co., Rocky Mt. Nat. Pk., 8.1 km W Estes Park, R13W
T5N Sec 19, 2,677 m, 29 September 1990, P. c. latifolia, 1 female; Las Animas Co.,
4 km S Cucharas Pass, 2,892 m, 5 September 1991, P. aristata, 1 female; Pitkin Co.,
10 km S Aspen, 2,646 m, 1 1 September 1991, P. c. latifolia, 10 females, 1 male.
Nevada: Washoe Co., Galena Cr., Mt. Rose, 14 June 1964, 1 female. Oregon: Benton
Co., Corvallis, OSU campus, 1 July 1988 (teneral), P. monticola, 1 male, 1 female;
Klamath Co., base of Gearhart Mt., 7 June 1959, P. c. murrayana, 1 female; Lane
Co., H. J. Andrews Exp. For., 18 km NE Blue River, 18 August 1988, P. monticola,
2 males, 5 females, 2 V-instar nymphs. Wyoming: Albany Co., Sites 2.1 and 2.2, 25
July 86, P. c. latifolia adults and nymphs, 1 female; Site 2.2, 27 July 86, P. c. latifolia
adults; Site 2.1, 9 July 86, P. flexilis (10 trees pooled), 1 female; Site 2.2, 25 July 86,
P. c. latifolia (T-20), 1 female.

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Melanocoris nigricornis Van Duzee
(Figure 7)

Melanocoris obovatus: Van Duzee, 1917a:819, b:262.

Melanocoris nigricornis Van Duzee, 1921:143; Kelton and Anderson, 1962:1307;

Anderson, 1962b: 1332; Kinzer, Ridgill and Watts, 1972:10; Kelton, 1977a:246;

Kelton, 1978:23; Henry, 1988:16.

Tetraphleps novitus Drake and Harris, 1926:41 (Colo.); Kelton and Anderson, 1962:

1307.

Diagnosis: Moderate length (2.66-3.50 mm), head, and pronotum black, base and
margins of pronotum often brown; hemelytra dark brown, finely sculptured, shiny;
pubescence on hemelytra scalelike and serially arranged. Tip of rostrum reaching
anterior coxae.

Habitat: The original series was collected from pines in the southern part of the
Sierra Nevada of California. One specimen was taken from Jeffrey pine (Van Duzee,
1921). Anderson (1962b) reported M. nigricornis from Pinus ponderosa and P. c.
latifolia in British Columbia and P. ponderosa in Oregon. Kinzer (1976) reported M.
nigricornis from P. ponderosa in New Mexico. Kelton (1978) reported the species
from Pinus sylvestris, P. contorta, P. ponderosa, and Picea engelmanni in British
Columbia. We add P. c. murrayana in Oregon, California, and Mexico, P. c. latifolia
in Wyoming, and P. radiata in California.

Prey: Melanocoris nigricornis was collected in association with the aphids Cinara
sp. and Essigella fusca G. & P., as well as the pine needle scale, Phenacaspis pinifoliae
(Anderson, 1962b). Kelton (1978) speculated that the species probably feeds upon
aphids, scales and other small arthropods. We collected aphids belonging to both
Cinara and Essigella from the Oregon and Wyoming sites during 1986.

Phenology and life history: Adults have been collected from April (California and
Mexico) to September (Oregon and Wyoming) in various parts of the range (Van
Duzee, 1921; Drake and Harris, 1926; Anderson, 1962b; Kelton, 1978). According
to Anderson (1962b), M. nigricornis overwinters as an adult in southern British
Columbia and adults and nymphs were common from July to October. Reproduction
continued into the fall with last instar nymphs collected in the latter part of October.
Males and females were collected in early winter but only the females survived until
spring. Under laboratory conditions, development from egg to adult took 24 days,
and two consecutive generations were reared in the laboratory in only 67 days. Thus
two generations are possible under field conditions.

Distribution: Melanocoris nigricornis was originally described by Van Duzee (1921)
from three female specimens from California. One specimen was taken at Summit,
Placer Co., at 2,154 m; another on pine trees on Mt. Tahquitz above Keen Camp,
Riverside Co., at 2,154 m; the last was taken on Jeffrey pine at Monmouth, Mono
Co. Previously, Van Duzee (1 9 1 7a) reported this species (under the name Melanocoris
obovatus Champion) from Summit, in Placer Co., California locality. Tetraphleps
novitus was described by Drake and Harris (1926) from specimens collected in Col-
orado on pine at Stonewall and Trinidad, Las Animas Co., 2,770 m and Estes Park,
Larimer County [note: Pinus ponderosa scopulorum is the common pine at these
locales]. This species was synonymized with Melanocoris nigricornis by Kelton and

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Fig. 7. Melanocoris nigricornis Van Duzee.

Anderson (1962). Anderson (1962b) reported M. nigricornis from several localities
in the Okanagan Valley of British Columbia: Oliver to Westbank, Princeton, and
from Adel, Oregon. Kelton (1978) reported it from several localities in southeastern
British Columbia. We report here additional localities in Oregon and California and
add Washington, Wyoming, and Mexico.

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ANTHOCORIDAE ON PINUS CONTORT A

449

Specimens examined: CANADA: British Columbia: 19.5 km W Princeton, 1,169
m, 31 July 1957, P. contorta, 1 male. MEXICO: Baja California Norte, 90 km E San
Telmo, 18 April 1979, P. contorta, 2 males, 4 females. UNITED STATES: California:
Fresno Co., Pioneer Basin, 27 October 1959, 1 female (UCR). Mono Co., June Lake,
Hwy. 158, 16 June 1989, P. c. murrayana. 3 females; Monterey Co., Pt. Lobos St.
Res., E Hwy 1, Huckleberry Ridge Oak area, 8 April 1979, P. radiata, 1 female;
Sierra Co., Tahoe Nat. For., off FS Rd. 07, 27.7 km W Hwy. 89, 2,092 m, 16 July
1987, P. c. murrayana, 1 female. Colorado: Larimer Co., Estes Park YMCA Camp,
6 km SW Estes Park, 2,465 m, R13W T4N Sec 4, 26 September 1990, P. ponderosa
scopularum, 1 male, 3 females; Las Animas Co., 4 km S Cucharas Pass, 2,892 m, 5
September 1991, P. ponderosa scopulorum, 1 female. Oregon: Deschutes Co., Three
Creeks Mdw., 26 km S Sisters, 2,065 m, 1 September 1977, P. c. murrayana, 3
males; Three Creeks Lake, 28 km S Sisters, 2,100 m, 14 June 1990, P. c. murrayana,
8 females; Black Butte Ranch, 13 km NW Sisters, 1,015 m, 24 March 1990, P.
sylvestris, 3 males, 3 females; Linn Co., Big Lake, 1,431 m, 1 October 1979, P. c.
murrayana, 4 females. Washington: Kittatas Co., 8 km W, 5 km S of Cle Elum, 6
June 1970, 1 female. Wyoming: Park Co., Yellowstone Nat. Pk., 8 km S Tower Falls,
2,262 m, 17 September 1980, P. c. latifolia, 3 females; Teton Co., Grand Teton Nat.
Pk., 2,123 m, 18 September 1980, P. c. latifolia, 3 males.

Orius Wolff

Orius Wolff, 1811:5; Carayon, 1953b: 1206; Wagner, 1952:22; Carayon and Steffan,

1959:53; Kelton, 1963:631; Herring, 1966:1093; Carayon, 1972a:346; Pericart,

1972:160; Herring, 1976:147; Kelton, 1978:46; Henry, 1988:18.

Triphleps Fieber, 1860:266; Van Duzee, 1917a:293.

Diagnosis: Small, shiny, ovate, macropterous. Head short, ocelli distinct; tip of
rostrum reaching anterior coxae; antennal segments II, III, and IV of equal thickness.
Pronotum with punctures; short collar, not well differentiated; calli distinct, smooth.
Hemelytra shiny, finely punctate, three veins in membrane. Anterior tibiae with fine,
stout spinules on inner surface. Ostiolar canal curving forward, evaporative area
broad and shiny.

More than 70 species of Orius are recognized and almost half of these occur in the
Northern Hemisphere (Pericart, 1972). This is a large, complex genus with much
additional work needed to achieve an understanding of the taxon. Wagner (1952)
monographed the European species and recognized four subgenera. Kelton (1963)
published a synopsis of the North American species, and Herring (1966) mono-
graphed the species of the Western Hemisphere. Carayon and Steffan (1959) reviewed
the feeding habits of the genus. The North American fauna still requires additional
work, for many of the described taxa are known from very few individuals. Two
native species, Orius insidiosus (Say) and O. tristicolor( White), are abundant through-
out their respective ranges in North America, the former chiefly eastern and southern,
and the latter northern and western (Kelton, 1963; Herring, 1966). Both have been
deliberately introduced into Hawaii as biological control agents for two different
Lepidoptera (Weber, 1953; Davis and Krauss, 1 963). A selected bibliography of these
two species was published by Ryerson and Stone (1979). A third species, Orius
minutus (L.), represents an introduction into North America (Lattin, Asquith, and
Booth, 1989).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Orius tristicolor (White)

(Figure 8)

Triphleps tristicolor White, 1879:145; Van Duzee, 1914:15; Van Duzee, 1917a:293;

b:262.

Orius insidiosus var. tristicolor: Blatchley, 1926:637.

Orius tristicolor: Harris and Shull, 1944:207; Anderson, 1962b: 1330; Kelton, 1963:

634; Herring, 1966:1103; Kelton, 1978:50; Henry, 1988:19.

Diagnosis: Length 1.80-2.3 mm. Head black, shiny; tip of rostrum extending to
anterior coxae. Pronotum black, shiny, punctate and rugose; calli smooth, raised.
Scutellum black, shiny. Hemelytron with anterior half of corium, parts ofclavus pale
and shiny; balance of wing black; pubescence short, scattered; membrane fuscous,
with two visible veins.

Habitat: Species of Orius occur on the surface of forbs, shrubs and trees including
conifers. They are mobile predators that patrol the plant surfaces, feeding on small
arthropods and their eggs (Collyer, 1953). Most species seem to be generalist pred-
ators, taking prey scaled to their size. Orius tristicolor has been found on a wide
variety of plants and associated with many prey species (Anderson, 1962b; Ryerson
and Stone, 1979). This species is often encountered in flower heads as well as on
other parts of the plant.

Along with O. insidiosis, this is a ubiquitous species of Anthocoridae. Not primarily
a conifer-inhabiting species, we have seen specimens from Pinus ponderosa (Oregon)
and from P. c. murrayana (Baja California, Norte, Mexico). They Oregon locality
(near Sisters) had a scattered shrub understory of Chrysothamnus and Purshia.

Prey: Orius tristicolor feeds especially on Homoptera (Anderson, 1962b; Wilde and
Watson, 1963; McMullen and Jong, 1967). However, Barber (1936) found that the
early nymphal instars of O. insidiosus, a closely related species, were largely phy-
tophagous and in fact could complete development on plant food alone. Dicke and
Jarvis (1962) indicated that pollen is an important food source for O. insidiosus.
Carayon and Steffen (1959) reported similar feeding habits for other species of Orius.

Phenology and life history: According to Anderson ( 1 962b), this species overwinters
as an adult (mostly females) and has been collected during winter from ground litter
and has been beaten from deciduous and coniferous trees. He stated that three
generations per year were likely in the Pacific Northwest but a fourth was possible.
Developmental time from egg to adult was approximately 3 weeks. Eggs are inserted
into leaves (including the veins) and hatch in about 7 days. Five instars occur with
the duration of each stage about 4 days.

Distribution: The species ranges widely over Canada, northern and western United
States, south through Mexico into Central and South America and the Bahamas
(Kelton, 1963; Herring, 1966). In the eastern part of Canada and United States, it
is largely replaced by O. insidiosis, which ranges through the southern and central
states into Mexico and Central and South America and several of the West Indies
(Kelton, 1963; Herring, 1966).

Specimens examined: Many specimens from throughout the range of the species.
MEXICO: Baja California Norte, 90 km E San Telmo, 18 April 1979, P. contorta.
UNITED STATES: Oregon: Deschutes Co., 4.9 km S Sisters, 26 September 1989,
Pinus ponderosa, 2 females; Linn Co., Big Lake, 1,431 m, 1 October 1979, P. c.
murrayana, 1 female.

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451

Fig. 8. Orius tristicolor (White).

Scoloposcelis Fieber

Scoloposcelis Fieber, 1864:62; Van Duzee, 1917a:296; Carayon, 1953a:65; Carayon,
1972a:341; Pericart, 1972:279; Kelton, 1976a:196; Herring, 1976:149; Kerzhner,
1988:771; Henry, 1988:25.

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Diagnosis: Body flattened, sides subparallel, smooth, shiny. Macropterous, corium,
and clavus impunctate. Legs stout, anterior and posterior femora enlarged, anterior
femur with short, stout teeth on inner surface. Long setae at end of abdomen. Male
clasper short, stout, slightly curved, parallel-sided except apex.

Twelve species of Scoloposcelis are known, nine in the Holarctic Region (Pericart,
1972). Two species, S. flavicornis Reuter and S. basilicus Drake and Harris, are
known from America north of Mexico (Henry, 1988). The former species occurs
throughout the United States and Canada from Mississippi and Texas to the Yukon
Territory, whereas the latter is known only from Arizona and New Mexico. It is
likely that additional species will be recognized. Species of Scoloposcelis occur under
the bark of conifers, chiefly species of the genus Pinus where they feed mainly on
Scolytidae and other subcorticular insects (Drake, 1921, 1922; Carayon, 1953a; Mo-
sier, Thatcher, and Pickard, 1971; Pericart, 1972; Moore, 1972; Goyer, Lenhard,
Nebeker, and Schmitt, 1985). Carayon (1953a) provided a detailed discussion of the
habits and habitats of S. obscurella (Zett.). The range of this species extends from
Europe to the far-eastern U.S.S.R. (Kerzhner, 1988). Carayon stated that the species
occurs not so much under bark, but between the layers of bark where its exceptionally
flat shape permits it to enter crevices most insects could not. Besides scolytids,
Carayon reported that S. obscurella fed upon the larvae of Tribolium, Gnathocerus,
Anobium and Oryzaephilus in captivity.

Carayon (1953a) described the life cycle of S. obscurella in France. Eggs deposited
in tree bark hatch in about 1 2 days. The resulting nymphs are reddish, and gradually
darken as they become older. The nymphs are distinctive in appearance: very flat,
parallel-sided, with the anterior and posterior femora greatly enlarged. The anterior
pair have a row of short teeth on the inner surface. Several very long setae extend
from the end of the abdomen. There are five nymphal instars and a single, protracted
generation per year.

Scoloposcelis flavicornis Reuter
(Figure 9)

Scoloposcelis flavicornis Reuter, 1871:561; Van Duzee, 1917a:297; Drake and Harris,
1926:44; Harris and Shull, 1944:207; Anderson, 1 962b: 1 332; Kelton, 1976a:196;
Kelton, 1978:52; Berisford, 1980:250; Dixon and Payne, 1979:13, 25; Ohmart,
1981:5; Ohmart and Voight, 1981:677, 680; Henry, 1988:25.

Scoloposcelis mississippensis Drake and Harris, 1926:45 (Miss.); Mosier, Thatcher
and Pickard, 1971:73; Moore, 1972:63; Kelton, 1976a: 196; Dixon and Payne,
1979:13, 25; Berisford, 1980:250; Goyer, Lenhard, Nebeker, and Schmitt,
1985:4.

Scoloposcelis occidentalis Drake and Harris, 1926:45 (Calif.); Kelton, 1976a: 196.
Anthocoris sp.: Drake, 1921:201; Drake, 1922:67; pi. II, fig. b.

Diagnosis: Body elongate, parallel-sided and very flat; dorsum shiny, glabrous;
anterior and posterior femora enlarged, anterior femur with short teeth on inner
surface. Several long setae at end of the abdomen. Head and pronotum dark, shiny;
macropterous, hemelytra shiny, glabrous; clavus and corium largely pale, dark along
claval suture, embolium and most of cuneus, with membrane clear.

Some morphological differences in some of the specimens examined suggest the
likelihood of at least one other species, perhaps a northern and southern species.

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453

Fig. 9. Scoloposcelis flavicornis Reuter.

Only a thorough examination of available specimens will clarify this situation. Be-
cause both S. obscurella Zett. and S. pulchella Zett. have been reported from the far-
eastern Palearctic Region by Kerzhner (1988), specimens from the far northwestern
portion of North America should be examined with special care.

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Habitat: Scoloposcelis flavicornis occurs under the bark of trees where it is often
found in the burrows of bark and ambrosia beetles (Drake, 1921, 1922; Drake and
Harris, 1926; Dahlsten, 1970; Mosier, Thatcher, and Pickard, 1971; Moore, 1972;
Kelton, 1976a; Dixon and Payne, 1979; Ohmart and Voight, 1981; Ohmart, 1981;
Goyer, Lenhard, Nebeker, and Schmitt, 1985). We collected adults of S. flavicornis
in the Cascade Mountains of Oregon in July beneath bark flakes of a recently dead
branch of P. c. murrayana (needles brown, attached) together with three species of
Scolytidae. Fifth-instar nymphs were collected from P. resinosa at Harvard Forest
(near Petersham), Massachusetts, in mid-October. These specimens were beaten from
dead branches on the ground that had been cut earlier in the year (needles brown,
attached). Only branches with scaly bark produced specimens. By mid-November,
the specimens were still in the nymphal state. The Oregon record of S. flavicornis is
based on specimens from a subalpine meadow at 2,000 m on the east side of the
Cascade Mountains on P. c. murrayana in mid-July. This is in contrast to the Wash-
ington specimens taken at Bremerton, Washington, just west of Seattle at sea level
from shore pine (P. c. contorta).

This species has been collected from Pinus contorta contorta, P. c. latifolia, P. c.
murrayana (this paper), P. banksiana (Kelton, 1978), P. palustris (Drake and Harris,
1926), P. ponderosa (Dahlsten, 1970; Kelton, 1978), P. radiata (Ohmart and Voight,
1981; Ohmart, 1981), P. resinosa (Kelton, 1978), Picea glauca (Kelton, 1978), and
Picea sp. (Drake, 1921, 1922).

Prey: The chief prey of S. flavicornis appears to be the eggs and larvae (and perhaps
callow adults) of Scolytidae. Drake (1921, 1922) reported it (as Anthocoris sp.) from
the burrows of 1 1 species of bark and ambrosia beetles in spruce logs in New York
and reported feeding on both types of beetles. He provided an excellent drawing of
the fifth-instar nymph, showing the characteristically enlarged front and hind femora.
Drake and Harris (1926) reported nymphs and adults (as S. mississippensis) in the
burrows of Ips spp. in long-leaf pine. Dahlsten (1970) reported likely predation on
Dendroctonus brevicomis LeConte in California. Mosier, Thatcher, and Pickard (1971)
reported S. flavicornis as a predator of the eggs and larvae of Dendroctonus frontalis
Zimmerman and Ips sp., as did Moore (1972). Ohmart (1981) reported it from the
brood gallery of Ips paraconfusus Lanier, while Dixon and Payne (1979) and Goyer,
Lenhart, Nebeker, and Schmitt (1985) recorded it as an associate of D. frontalis.
According to the latter, another anthocorid, Lyctocoris elongatus Reuter, occurred
with S. flavicornis.

Phenology and life history: Winter is normally spent as a mature nymph (at least
in northern locales). Adults appear in the spring and usually disappear in early fall.
Carayon (1953a) provided a detailed account of S. obscurella, collected in the moun-
tains of central France, and the general phenology was the same. The original de-
scription of S. mississippensis (Drake and Harris, 1926) mentioned males, females,
and fourth and fifth-instar nymphs being collected together at Port Gibson, Missis-
sippi, on Pinus palustris in late July. Mosier, Thatcher and Pickard (1971) reported
the occurrence of S. flavicornis (as S. mississippensis ) in January and March through
November. Dixon and Payne (1979) showed a close correlation between the arrival
of Scoloposcelis and the bark beetles, especially Dendroctonus frontalis, attacking
pines in southern United States. The beetles are multivoltine and it appears that the
anthocorid is too, thus explaining the occurrence of the bug throughout the year.

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Ohmart (1981) stated that S. flavicornis was collected throughout the year on Pinus
radiata in coastal California, but was most abundant from November to March. He
did not indicate life stages collected.

Distribution: Scoloposcelis flavicornis was described from Texas by Reuter (1871).
This widespread anthocorid has since been recorded from Alberta, British Columbia,
California, Florida, Idaho, Indiana, Manitoba, Mississippi, New Brunswick, Nova
Scotia, Ontario, Pennsylvania, Quebec, Saskatchewan, Yukon Territory, Guatemala,
and Mexico (Henry, 1988). To these records we add Oregon, Massachusetts and
Washington. Two species described by Drake and Harris (1926), S. mississippensis
from Mississippi and S. occidentalis from Ventura Co., California, were synonymized
with S. flavicornis by Kelton (1976a).

Specimens examined: Massachusetts: Worchester Co., Harvard Forest, nr Peter-
sham, 28 September 1989, under bark of dead branches of Pinus resinosa, five fifth
instar nymphs. Montana: Ravalli Co., Sula, 26 June 1929, Pinus contorta latifolia,
Hopk. US/ 19509, 7 males, 2 females (USFS, LaGrande, Ore.); Sula, 8 July 1929,
Pinus contorta latifolia, Hopk. US/ 19509, 6 males, 1 female (USFS, LaGrande, Ore.);
Sula, 27 July 1929, Pinus contorta latifolia, Hopk. US/ 19509, 7 males, 2 females
(USFS, LaGrande, Ore.) Oregon: Deschutes Co., Three Creeks Meadow, 2,069 m,
26 km S Sisters, 1 1 July 1988, under bark of dead branch of P. c. murrayana together
with three species of Scolytidae, 2 males, 6 females. Washington: Kitsap Co., Brem-
erton, 1 7 February 1975, emerged indoors, 19 February from gallery of Pityophthorus
confertus, in P. c. contorta, 1 female.

Tetraphleps Fieber 1860

Tetraphleps Fieber, 1860:262; Reuter, 1884:56, 85; Lethierry and Severin, 1896:245;
Reuter, 1908:86; Parshley, 1920:83; Van Duzee, 1921:143; Drake, 1921:203; 1922:
67; Drake and Harris, 1 926:4 1 ; Kelton and Anderson, 1 962: 1 307; Anderson, 1 962b:
1331;Ghauri, 1964:675; Kelton, 1966a: 199; Mitchell and Wright, 1967:143; Harris,
Holms, and Dawson, 1968:5-6; Kelton, 1968:1072; Carayon, 19 72a: 34 5; Pericart,
1972:154; Herring, 1976:146; Kelton, 1978:26; Evans, 1983:45; Kerzhner, 1988:
771; Henry, 1988:17.

Diagnosis: Macropterous, shiny, pubescent. Tip of rostrum reaching or exceeding
front coxae. Pronotum with distinct collar, lateral margins carinate, calli smooth,
surface punctate. Hemelytra with punctures, shiny, pubescent, membrane with four
veins. Hind coxae close together. Ostiolar canal straight, apex rounded, raised above
surface of metapleuron.

The genus Tetraphleps contains approximately 12 species (Pericart, 1972). The
species occur in the Northern Hemisphere, with about equal numbers in the Old and
New World. Five species are known to occur in North America (Kelton, 1966a, 1978;
Henry, 1988). Keys to the North American species are found in the two papers by
Kelton (1966a, 1978), together with information on distribution and host associations.
Species of Tetraphleps are found only on conifers, especially of the genus Pinus, and
to a lesser extent on Larix, Picea, Abies, and Pseudotsuga (Reuter, 1908; Pericart,
1972; Kelton, 1978). Kinzer, Ridgill, and Watts (1972) and Kinzer (1976) recorded
Tetraphleps sp. from Pinus ponderosa cones in New Mexico. Evans (1983) reported
Tetraphleps sp. from lodgepole pine in the northern interior of British Columbia.

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Species of Tetraphleps are active, surface feeding predators, often abundant on
their host plants. Their main food seems to be aphids. Pericart (1972) mentioned
several aphids of the genus Cinara as primary prey of T. biscuspis (H.-S.) in Europe.
Cinara spp. were commonly collected from P. c. murrayana and P. c. latifolia at a
number of localities in the present study. Specimens of the aphid genus Essigella
also were taken.

Mitchell and Wright (1967) reported the introduction of Tetraphleps spp. from
Pakistan and India in 1964. Specimens from Pakistan were released at Randle,
Washington, and Portland, Oregon (Hoyt Arboretum), and those from India at Wind
River, Washington (Wind River Arboretum), and into a laboratory culture. It is not
known if they have become established (Dr. R. G. Mitchell, pers. comm., 12 March
1990). These anthocorids were released as potential predators of the balsam woolly
adelgid, Adelges piceae (Ratzeburg). These same species, ultimately identified as
Tetraphleps abdulghani Ghauri from Pakistan, and T. raoi Ghauri from India (Gha-
uri, 1 964; Henry, 1 988), were introduced onto Vancouver Island (Thetis Park), British
Columbia in 1965 (Harris, Holms, and Dawson, 1968). According to Dr. L. M.
Humble, Forest Insect and Disease Survey, Forestry Canada, Victoria, British Co-
lumbia, these species apparently have not become established (pers. comm., 2 March
1990). According to Kelton (1978), they were also introduced into New Brunswick
and Nova Scotia, and as of 1971, had not been recovered. Two native species, T.
latipennis Van Duzee and T. sp., were collected at the Thetis Park release site in
1964 and 1965.

A shortage of specimens from a number of localities makes it difficult to assess
the degree of species variation. Females predominate in most collections, mainly
because of the timing of collections; and thus the variation in male genitalia, especially
the claspers, is largely unknown. A thorough revision of Tetraphleps, with exami-
nation of all type material, almost certainly will result in the recognition of additional
species.

Tetraphleps canadensis Provancher

Tetraphleps canadensis Provancher, 1886:90; Parshley, 1920:85; Van Duzee, 1921:
142; Drake and Harris, 1928:50; Anderson, 1958:50; Kelton and Anderson, 1962:
1307; Anderson, 1962b: 1331; Kelton, 1966a: 199; Kelton, 1968: 1072; Kelton, 1978:
27; Henry, 1988:17.

Lyctocoris canadensis : Van Duzee, 1912:320.

Tetraphleps americana Parshley, 1920:84; Drake and Harris, 1928:50.

Tetraphleps osborni Drake, 1922:67; Kelton and Anderson, 1962:1307.
Acompocoris ( Tetraphleps ) osborni: Drake and Harris, 1926:43.

Tetraphleps edacis Drake and Harris, 1926:43; Kelton and Anderson, 1962:1307.

Diagnosis: Length 3.1 to 3.9 mm. Head, pronotum and scutcllum black, shiny,
pubescence dense and rather long. Hemelytra pale brown, shiny, finely punctate, with
rather long, dense pubescence. Apex of rostrum reaching middle of mesostemum.
Hind tibiae with short setae. Ostiolar canal with posterior margin concave, apex
broadly rounded.

Habitat: Drake (1921) reared a specimen (as Tetraphleps n. sp.) from the burrows
of a weevil from Bebb’s willow ( Salix bebbiana Sarg.) in New York, but he thought

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that pine was its usual host. Drake (1922) mentioned it (as Tetraphleps osborni) was
a predator of leaf-feeding insects on Pinus strobus in New York. Drake and Harris
(1926) reported Larix laricina as a host (as Tetraphleps edacis) in New York. Kelton
(1966a, 1978) listed Abies balsamea L., Pinus banksiana, Picea glauca, P. mariana,
and Larix laricina as host plants.

Prey: Drake (1921, 1922) cited the beetle, Cryptorhynchus lapathi Linnaeus, as
prey of Tetraphleps osborni Drake (=77 canadensis Provancher), but thought that
herbivorous insects on conifers, in particular Pinus strobus, provided most of its
food. Kelton (1966, 1978) mentioned its association with the balsam woolly adelgid
in eastern Canada.

Phenology and life history: The type of Tetraphleps americana Parshley (=77 can-
adensis Prov.), a female, was collected in Maine in early August and the paratype,
also a female, was collected at Ottawa, Canada, in early September. Adult Tetraphleps
osborni Drake (=77 canadensis Prov.) were taken in New York during July, August,
and September (Drake, 1922). Drake (1922) stated that nymphs and adults were
found during the summer, but the larger nymphs and adults (males and females)
were found only during the latter part of the summer. He reported only a single
generation per year. How this species overwinters is not known. According to Pericart
(1972), adults of T. bicuspis (H.-S.) overwinter under the bark of their coniferous
hosts. Parshley (1920) recorded no host data when he described T. americana.

Distribution: Originally described from Cape Rouge, Quebec, Tetraphleps cana-
densis Provancher is now known to occur across southern Canada from Newfound-
land west to British Columbia (Kelton, 1 978). Kelton (1966a) also cited Alaska, Maine,
Michigan, and Wisconsin. Tetraphleps americana Parshley (=77 canadensis Pro-
vancher) was described from Maine; T. osborni Drake (=77 canadensis) from New
York (Barber Point, Wanakena, and The Plains); and T. edacis Drake and Harris
(=77 canadensis ), from The Plains, Wanakena, New York. We report it here from
coastal Oregon.

Specimens examined: We have examined several specimens of what appears to be
this species along the Oregon coast. Oregon: Lincoln Co., Driftwood Beach Wayside,
8.5 km N Waldport, 8 August 1985, P. c. contorta, sample #91,1 female.

Tetraphleps latipennis Van Duzee
(Figure 10)

Tetraphleps latipennis Van Duzee, 1921:140; Downes, 1927:11; Harris and Shull,

1944:206; Anderson, 1958:50 (in part); Kelton and Anderson, 1962:1307; An-
derson, 1962b: 1331; Kelton, 1966a:201; Harris, Holms, and Dawson, 1968:15;

Fumiss and Carolin, 1977:81; Kelton, 1978:29; Henry, 1988:17.

Tetraphleps profugus Drake and Harris, 1926:42; Kelton and Anderson, 1962:1307;

Kelton, 1966a:201; Henry, 1988:17.

Diagnosis: Length 3.50 to 4.35 mm, dorsum shiny; pubescence distinct, short and
appressed. Tip of rostrum reaching anterior coxae. Pronotum with calli black, balance
red brown; scutellum black; hemelytra red brown, shiny, membrane brown, narrowly
pale around veins.

Habitat: This is a surface-feeding species, primarily a predator of small arthropods.
It forages for aphids on the branches and needles of various conifers, most commonly

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Fig. 10. Tetraphleps latipennis \ an Duzee.

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on pines. The type series was collected on white-bark pine ( P . albicaulis) (Van Duzee,
1921); Abies, Pseudotsuga, Picea, Pinus, and Larix (Anderson, 1962b); Pseudotsuga
menziesii (Mirb.); Pinus ponderosa Laws., P. c. latifolia Engelm., Abies grandis (Dough),
Picea sp., Larix sp., and on Alnus sp. (Kelton, 1966a); Abies grandis (Harris, Holms,
and Dawson, 1968); Abies, Pseudotsuga, Picea, Pinus, and Larix (Fumiss and Car-
olin, 1977); and Abies balsamea, A. amabilis, A. lasiocarpa, Larix occidentalis, Picea
engelmanni, P. glauca, Pinus monticola, P. albicaulis (Kelton, 1978). We add P. c.
murrayana from Oregon.

Prey: Kelton (1966a) reported T. latipennis feeding on the balsam woolly adelgid
in eastern Canada. Fumiss and Carolin (1977) stated that this species is the most
common anthocorid on conifers and reported it feeding on Adelges cooleyi. Kelton
(1978) added “. . . other aphids in British Columbia.”

Phenology and life history: Anderson (1962b) thought there were two generations
per year and that the adults overwintered beginning in late August or September;
the spring generation adults matured in late June or July. The only immature stages
collected were fifth-instar nymphs in southeastern British Columbia in June and July
together with males and females. The earliest adults in British Columbia were col-
lected in late May and the latest, September 12. The type series, all females, were
collected in California on July 28 (Van Duzee, 1921).

Distribution: The type series was taken on the top of Mt. Eddy (2,800 m), Siskiyou
Co., California (Van Duzee, 1921). Also included in the type series were specimens
from Huntington Lake (2,308 m), Fresno Co., and near Walker, Siskiyou Co., Cal-
ifornia as well as a specimen from Nanaimo, Vancouver Island, Canada. Downes
(1927) reported it from Victoria, British Columbia; Harris and Shull (1944) from
Moscow Mountain, Idaho; Anderson (1958) listed localities in British Columbia,
Idaho, Washington; Anderson (1962b) recorded it from several other British Colum-
bia localities, McCall, Idaho and Corvallis, Oregon. Kelton (1966a) added many
localities from Alberta, British Columbia, California, New Brunswick, Newfound-
land, Oregon and Wyoming; and (1978) provided a distribution map of the species
in Canada. A considerable gap existed between the western and eastern localities and
the map showed several localities in Saskatchewan and one in Manitoba besides the
provinces already recorded.

Specimens examined: CANADA: British Columbia; Westbank, 12 June 1956,
Douglas-fir, 1 female. UNITED STATES: California; Shasta Co., 14 km E Shasta
City, 8 August 1956, Abies shastensis, 1 male, 2 females; Siskiyou Co., Medicine Lk.
Rd., 1,846 m, 26 June 1979, Abies concolor, 4 females. Idaho: Benewah Co,, Emida,
Charlie Crk. Rd., 8 July 1979, Abies sp., Abies concolor, 4 females; Latah Co.,
Moscow, 25 July 1920, 1 female. Oregon: Benton Co., Mary’s Peak, Saddle, 17
August 1978, Douglas-fir, 1 male, 1 female; Mary’s Peak, summit, 4 October 1977,
Abies procera, 2 females; Clackamas Co., Molalla, 1,108 m, 26 September 1977,
Abies procera, 3 females; Deschutes Co., Three Creeks Lake, 26 km S Sisters, 2,154
m, 14 June 1990, Pinus contorta murrayana, 4 females; Black Butte Ranch, 13 km
NW Sisters, 1,015 m, 23 March 1990, under bark of Pinus sylvestris, 1 female; Hood
River Co., Mt. Hood, nr Silcox Hut, 2,154 m, 24 August 1980, on snow bank, 1
female; 5 km N Barlow Pass Summit, R9E T3S Sec 15 NW~, 1,372 m, 5 September
1979, Abies amabilis, 2 females; Jackson Co., Tubb Springs, 10 km W Pinehurst,
1,323 m, 19 May 1960, 15 females; Klamath Co., 8 km E Kirk, 1,385 m, 21 May

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

1958, P. c. murrayana, 1 female; Lake of the Woods, Rainbow Camp, 1,508 m, 23
May 1958, Pinus c. murrayana, 3 females; same but Abies concolor, 25 females;
same, but Pinus monticola, 1 8 females; Lane Co., 3 km W Willamette Summit, 1 ,43 1
m, 17 July 1979, Tsuga mertensiana, 1 male, 1 female; Lane-Linn Co., H. J. Andrews
Exp. For. R6E T15S Sec 7, SW~ 1,323 m, 31 July 1979, Abies amabilis, 1 male, 1
female; same but McRae Crk. crossing, FS rd 1506-359, 1,237 m, 309 August 1978,
Abies procera, 1 female; same, 25 August 1980/860825-11 Psme-1 ( Pseudotsuga
menziesii), 1 female; Marion Co., 5 km W Breitenbush Lk., 5 June 1966, Abies
lasiocarpa, 2 females; Multnomah Co., Portland, Black light trap, 13 August 1963,

1 male, (OSDA): Union Co., 6 km ESE Tollgate, 1,538 m, 26 June 1960, 1 female.
Washington: Chelan Co., Pump Chance Crk., 4 km E Swauk Pass, 1,415 m, 10
August 1978, 1 male, 4 females; King Co., Summit, Stevens Pass, R13E T26N Sec
14, 22 August 1979, Abies amabilis, 5 males, 3 females; Okanagan Co., Washington
Pass Mdw., 1,662 m, 1 1 August 1978, Pinus albicaulis, 1 male; Whatcom Co., Mt.
Baker Nat. For., R9E T39N Sec 17, 7 September 1979, Tsuga mertensiana, 1 male,
1 female; Yakima Co., 5 km E White Pass Summit, 1,292 m, R12E T14N Sec 33
SW~, 22 August 1979, Abies procera, 5 females.

Tetraphleps pilosipes Kelton and Anderson

Tetraphleps pilosipes Kelton and Anderson, 1962:1307; Anderson, 1962b: 1331; Kel-
ton, 1966a:200; Kelton, 1978:28; Henry, 1988:17.

Diagnosis: Length 3.8 to 4.2 mm; head, pronotum, and scutellum black and shiny;
hemelytra brown, shiny, and punctate; pubescence long and dense on pronotum,
scutellum, and hemelytra. Hind tibia with long, distinct bristles. Ostiolar canal straight,
slender, with narrowly rounded apex.

Habitat: Tetraphleps pilosipes is commonly beaten from the branches of its host
trees. This species was originally described from specimens collected on Abies lasio-
carpa in British Columbia and on A. procera in Oregon, and specimens without host
data from Manitoba and the Yukon Territory (Kelton and Anderson, 1 962; Anderson
1 962b). Kelton ( 1 966a) added Larix laricina, Picea glauca and P. mariana\ he reported
Pinus albicaulis and P. contorta in his 1978 paper.

Prey: The only positive prey information was provided by Kelton (1966a, 1978),
who reported its association with the balsam woolly adelgid. Aphids of the genus
Cinara are commonly found on Abies procera at the Oregon locality of Mary’s Peak.

Phenology and life history: This species has been collected from early May to mid-
October on Mary’s Peak (1,231 m) near Corvallis, Oregon, and early May (British
Columbia) to early September at Churchhill, Manitoba. Both males and females were
collected in late August at Blackwall, Manning Park, British Columbia.

Distribution: Described from Blackwall, Manning Park, British Columbia, the type
series also contained specimens from other localities in British Columbia, Manitoba,
the Yukon Territory, and Oregon (Kelton and Anderson, 1962). Records for Alaska
and Newfoundland were added by Kelton (1966a) and repeated in his 1978 publi-
cation. We add records of a disjunct population in southeastern Wyoming. The
occurrence of T. pilosipes in Newfoundland, well removed from the main area of
occurrence, is perhaps paralleled by T. latipennis Van Duzee.

Specimens examined: Specimens were examined from Oregon (mainly from Abies

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spp.), Washington, and British Columbia. We include a selection of these specimens.
The record of collection from P. contorta given by Kelton (1978) did not give a
specific locality but the distribution map shows several localities in British Columbia
and the Yukon Territory that fall within the range of P. contorta. Specimens from
Site 3.1 in Wyoming from Pinus contorta latifolia are reported. Colorado: Grand
Co., 9 km S Fraser on Hwy 40, 17 August 1980, Abies lasiocarpa, 1 female. Oregon:
Benton Co., Mary’s Peak, nr Summit prkg. lot, 1,123 m, R7W T12S Sec 21 SW~,
2 October 1979, Abies procera, 2 males, 2 females; same, summit mdw., 4 October
1977, Abies procera, 4 males, 2 females; same, summit, 23 August 1988, Abies procera,

1 male, 2 females, 3 nymphs; Grass Mountain, 8 km NW Alsea, R8W T13S Sec 21
SW~, 1,092 m, 17 September 1979, Abies procera, 6 females; same, 985 m, 14
August 1980, Abies procera, 1 nymph; Clackamas Co., Molalla, 1,108 m, 26 Sep-
tember 1977 , Abies procera, 4 males, 31 females; Mt. Hood, 1.5 km below Timberline
Lodge, 9 September 1976, 1 male. Washington: Okanagan Co., Washington Pass
Mdw., 1,662 m, 11 August 1978, 2 males, 1 female; Yakima Co., 5 km E White
Pass Summit, R12E T14N Sec 33 SW~, 1,292 m, 22 August 1979, Abies procera,

2 females. Wyoming: Albany Co., Site 3.1, 30 July 1986, P. c. latifolia (trees 3, 15),
1 male, 1 female.

Tetraphleps uniformis Parshley
Tetraphleps n. sp.: Slosson, 1906:326.

Tetraphleps uniformis Parshley, 1920:85, Van Duzee, 1921:143; Kelton and Ander-
son, 1962:1307; Kelton, 1966a:202; Kelton, 1978:31; Henry, 1988:17.
Tetraphleps furvus Van Duzee, 1921:141; Anderson, 1958:52; Kelton and Anderson,
1962:1307; Anderson, 1962b: 1331; Kelton, 1966a:202; Kelton, 1978:31; Henry,
1988:17.

Tetraphleps concolor Drake, 1922:68. Nomen nudum.

Diagnosis: Length 3.5 to 4.2 mm; shiny, uniformly light brown with head slightly
darker and hemelytra slightly paler; pubescence dense, long. Rostrum just reaching
beyond anterior coxae. Lateral margins of pronotum narrowly carinate. Membrane
long, pale brown, with pale area at base end along middle of each vein. Ostiolar
canal with posterior margin shallowly concave, apex broadly rounded.

Habitat: Little is known about the habits of this species. The original description
by Parshley (1920) did not include host information, but he speculated that it would
occur on conifers. Kelton (1966a) reported it from white and black spruce ( Picea
glauca and P. mariana), balsam fir (Abies balsamea) and on Pinus contorta latifolia
and P. sylvestris. We add P. c. murrayana and Picea engelmanni from Oregon.

Prey: Kelton (1966a) stated that T. uniformis is associated with the balsam woolly
adelgid. He repeated this prey record for eastern Canada and added that it likely
feeds on other aphids on other hosts (1978). This anthocorid was very abundant on
several P. c. murrayana heavily infested with a species of Cinara near Sisters, Oregon.
Melancoris nigricornis, another anthocorid, was taken in lesser numbers from the
same trees.

Phenology and life history: Although locality records of this species have been
published (Parshley, 1920; Kelton, 1966a), no collecting dates were included. The
specimens we have examined bore dates ranging from May to September.

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Distribution: Parshley (1920) described this species from a female collected on Mt.
Washington, New Hampshire. Kelton (1966a) added many localities from Newfound-
land west to British Columbia and the Yukon Territory in Canada and added Maine,
New York, and Colorado in the United States. He also gave a map showing trans-
continental distribution in Canada.

We saw some distinct differences in specimens from the Rocky Mountains of
Wyoming and Colorado, especially in the shape and length of the pronotum, and
length of the membrane. The color pattern of the typical specimen, rather uniformly
brown, differs considerably from those of Kelton (1966) (head, pronotum and scu-
tellum black). We have examined specimens from Colorado and Wyoming that fit
the description of T. furvus very well. Careful study of specimens from throughout
the range is likely to disclose more than a single species.

Specimens examined: Colorado: Grand Co., 8.6 km S Fraser on Hwy 40, 1 7 August
1980, P. contorta latifolia, 1 female; Gilpin Co., 3 km S Guanella Pass 3,015 m, 4
September 1991, P. c. latifolia, 1 female, 1 male. Oregon: Deschutes Co., Site 8, 20
August 1986, Picea engelmanni (10 pooled trees), 1 female; Site 10, 10 June 1986,
Abies lasiocarpa, 1 female; Three Creeks Lake, 26 km S Sisters, 2,069 m, 14 June
1990, P. c. murrayana, 8 females; 7 October 1989, P. c. murrayana, 2 females. Wy-
oming: Albany Co., Site 3.1, 11 July 1986, P. c. latifolia (tree 19), 1 female; Site 3.1,
30 July 1986, P. c. latifolia (trees 3, 13, 15, 16), 5 males, 2 females; Carbon Co., Site
4.1, 17 July 1986, P. c. latifolia (tree 2), 1 male; Site 4.2, 1 1 July 1986, P. c. latifolia
(tree 19), 2 females; Site 4.2, 12 July 1986, P. c. latifolia (tree 10), I-V instar nymph,
Site 4.2, 17 July 1986, P. c. latifolia (trees 6, 11), 2 females.

Xylocoris Dufour

Xylocoris Dufour, 1831:423; Van Duzee, 1917a:290; Chu, 1969:47; Carayon, 1972a:

337; 1972b:579; Pericart, 1972:210; Herring, 1976:70; Kelton, 1976a: 193; Kerzh-

ner and Elov, 1976:364; Kelton, 1977b: 10 17; Kelton, 1978:63; Kerzhner, 1988:

771; Henry, 1988:26.

Diagnosis: Small, smooth, shiny species; both sexes may be macropterous or bra-
chypterous. Antennal segments one and two of normal thickness, segments three and
four slender, with long setae. Pronotum shiny, collar narrow, lateral margins carinate.
Hemelytra punctate or smooth, with or without pubescence; membrane, when pres-
ent, with four veins. Metacoxae proximate; ostiolar canal well developed, curved or
angled. Males with anterior and middle tibiae with pads. Left clasper only; ovipositor
well developed.

The genus Xylocoris contains approximately 30 described species. Although widely
distributed in the world, most of the species are found in the northern temperate
regions (Chu, 1969; Carayon, 1972a, b; Pericart, 1972). The type of the genus is
Xylocoris rufipennis Dufour, a junior synonym of Lygaeus cursitans Fallen. Chu
(1969) presented a list of world species. The genus contains four subgenera (Carayon,
1 972b; Pericart, 1972). Representatives of three of the four subgenera occur in North
America and twelve species are known in America north of Mexico (Henry, 1988).
Three of these, including X. cursitans, are considered introductions into the Nearctic
(Kelton, 1978; Henry, 1988).

Species of Xylocoris occur in quite different habitats, ranging from leaf litter, stored-

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grain facilities, to subcorticular environments, sometimes entering the galleries of
bark beetles (Tragardh, 1914; Sands, 1957; Southwood and Leston, 1959; Anderson,
1962b; Pericart, 1972; Kelton, 1978). Aside from the several species known to occur
in stored grain, little is known about the habits of the North American Xylocoris.
Chu (1969) provided a detailed study of X. galactinus (Fieber) in Japan. Papers by
Arbogast (1979) and Arbogast, Flaherty, and Press (1983) provided similar informa-
tion on X. flavipes (Reuter) and X. sordidus (Reuter), respectively, in North America.

The North American fauna needs careful study to clarify the status of almost all
of the species, including those considered to be introductions. Because of the low
vagility of brachypterous forms, for example, it is difficult to imagine their rapid
dispersal in natural forests. At present, no key exists for the North American species.
Kelton (1976a, 1978) provided descriptions of several species and a key to the four
species found in Canada and Alaska. Chu (1969), Carayon (1972b), Pericart (1972),
Kerzhner and Elov (1976), and Kerzhner (1988) provide valuable information on
the Palearctic fauna.

Xylocoris californicus (Reuter)

Piezostethus californicus Reuter, 1884:600; Lethierry and Severin, 1896:239; Van

Duzee, 1917a:291.

Xylocoris californicus: V an Duzee, 1916:34; 1917a:291; 1921:138; Harris and Shull,

1944:207; Anderson, 1962b:1326; Chu, 1969:18.

Xylocoris (Xylocoris) californicus: Henry, 1988:27.

Diagnosis: Macropterous. Length 3 mm. Dark brown to black, smooth, shiny.
Apex of rostrum attaining middle coxae. Pronotum polished, calli indistinct, trans-
verse impression absent. Hemelytron largely pale with inner margin of clavus and
commissural margin piceus, also apex of embolium and all of cuneus except inner
angle, piceus. Ostiolar canal curved, apex obtuse, ending some distance from anterior
margin of metapleuron; area anterior to canal, broad and striate.

Xylocoris californicus is similar to X. umbrinus Van Duzee. It may be separated
from the latter by the paler portions of the hemelytra ( X . umbrinus tends to be largely
brown to dark brown), and the shorter ostiolar canal with more of the evaporative
area anterior to the canal than in X. umbrinus, where the canal is longer and ap-
proaches the anterior edge of the metapleuron. Van Duzee (1921) compared the two
species.

Habitat: Little is known about the habitat of this species. The female reported here
was collected from the branches of P. c. murrayana near Bartle, California. Since
this species appears close to X umbrinus Van Duzee, it is possible that some of the
records for X. californicus actually refer to X. umbrinus. Van Duzee (1921) reported
taking X. californicus at Cayton and Bryson, California, the same localities as the
type series of X. umbrinus. He mentioned a female of X. umbrinus taken at Cayton
under the bark of an old fir log. The only host plant information available is that of
Pinus contorta murrayana from the specimen taken near Bartle, Siskiyou Co., Cal-
ifornia.

Prey: No information is available regarding the specific prey of this species.

Phenology and life history: Harris and Shull (1944) reported collections from July

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14 to September 19 in Idaho. The female we report here was collected in late June
in northern California. No information is available about its life history.

Distribution: Xylocoris californicus was originally described from Mariposa, Mar-
iposa Co., California (as Piezostethus californicus) (Reuter, 1884). Van Duzee re-
ported it from Utah, New Mexico and California (1917a) and from Cayton, Shasta
Co., and Bryson, Monterey Co., California (1921). Harris and Shull (1944) reported
the species from Hagerman and Twin Falls, Idaho. Anderson (1962b) added Merrill
and Klamath Falls, Oregon.

Specimens examined: California: Siskiyou Co., 2 km E Bartle, Hwy. 89, 1,246 m,
21 June 1981, P. c. murrayana, 1 macropterous female.

Xylocoris cursitans (Fallen)

(Figure 1 1 )

Lygaeus cursitans Fallen, 1807:74.

Piezostethus cursitans: Reuter, 1871:411; 1884:43; Van Duzee, 1914:14.

Xylocoris cursitans: Van Duzee, 1916:34; 191 7a:29 1 ; 1921:137; Harris and Shull,

1944:207; Sands, 1957:305; Southwood and Leston, 1959:185; Anderson, 1962b:

1326; Carayon, 1972b:582, 593; Pericart, 1972:228; Kelton, 1978:68; Henry,

1988:27.

Xylocoris vicarius: Anderson, 1962b: 1326.

Diagnosis: Length 1 .8-2.5 mm, width 0.70-1 mm, dark red brown to black, shiny,
smooth. Tip of rostrum reaching middle of mesosternum. Pronotum black, shiny,
calli smooth, basal portion transversely rugose. Scutellum with basal portion smooth,
apical portion rugose, a slightly elevated median carina. Hemelytra (macropterous
form) smooth, shiny, with moderately long, semi-erect and dense golden pubescence;
corium and clavus adjacent to corium pale, balance of wing dark brown to black;
membrane pruinose, only outer vein distinct. Hemelytra of brachypterous form
reduced, extending to posterior margin of abdominal segment three; membrane great-
ly reduced. Ostiolar canal strongly elbowed. Male clasper slender and sinuate.

Habitat: The principal habitat is beneath the bark of dead trees, both deciduous
and coniferous (Tragardh, 1914; Sands, 1957; Southwood and Leston, 1959; An-
derson 1962b; Pericart, 1972; and Kelton, 1978). Rarely it has been found in grana-
ries, leaf litter, and on plants (only macropterous forms in the latter) (Sands, 1957;
Pericart, 1972). It seems most abundant under bark of trees in the early stages of
decay (Southwood and Leston, 1959). Such genera as Quercus, Fagus, Fraxinus and
Populus have been cited as examples of deciduous trees, while Pinus has been reported
as the coniferous example (Sands, 1957; Anderson, 1 962b; Pericart, 1 972). We report
its occurrence here under the bark of P. c. contorta.

Prey: Anderson (1962b) cited its association with dipterous larvae (likely Myce-
tophilidae) under bark. He was able to rear it on aphids in the laboratory. Morkrzecki
(1923) reported this species feeding on Ips typographus Linnaeus. Pericart (1972)
suggested a variety of prey, such as the larvae of many insects and other subcorticular
arthropods including larval Scolytidae, other beetles, thrips, and Collembola.

Phenology and life history: Anderson (1962b) reported that the number of gener-
ations per year had not been established although most life stages were present
throughout most of the year. Both nymphs and adults overwinter under the bark of

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trees, although true diapause appears to be lacking (Pericart, 1972). Pericart (1972)
suggested at least two or three generations occur per year. The eggs are deposited on
the inner surface of the bark or rotten wood (Anderson, 1962b). The same author
reported that the eggs hatched in 8 to 10 days. The nymphs develop slowly, even
under laboratory conditions, requiring from 58 to 62 days to reach the adult stage.
He reported that adults were long lived, one living for 80 days. This long adult life
could account for the overlap between generations.

Distribution: Xylocoris cursitans is a Palearctic species, originally described from
Sweden. It is considered to be an introduction into North America, first recorded by
Van Duzee (1916). According to Henry (1988), it has been reported from Alberta,
British Columbia, Connecticut, Idaho, Indiana, Michigan, New Jersey, Nova Scotia,
New York, Ontario, Oregon, and Quebec. To these records, we add Washington.
Within the range of P. contorta, the species is known from southern British Columbia
and Alberta, south into Idaho, western Washington, through western Oregon (An-
derson, 1962b). Some of these localities are well removed from areas where intro-
ductions might be considered reasonable, especially since this species does not appear
to be particularly vagile.

Specimens examined: Washington: Kitsap Co., Bremerton, 2 November 1974,
under bark of Pinus contorta killed in 1972; abandoned by Scolytidae, 1 macropterous
female, 1 brachypterous female.

DISCUSSION

HOST PLANT ASSOCIATION

Although the Anthocoridae are primarily predaceous, some show remarkable fi-
delity to particular plants at various taxonomic levels (i.e., conifers or a particular
genus of plants). Of the 28 genera of Anthocoridae known from Canada and the
United States, 8 have some or all of their species occurring on conifers and two
(. Anthocoris and Orius ) are occasionally found on conifers, although they live chiefly
on broad-leafed plants. Five of the eight genera ( Acompocoris , Elatophilus, Melan-
ocoris, Tetraphleps, and Scoloposcelis ) occur only on conifers. Cardiastethus, Lyc-
tocoris, and Xylocoris occur on non-coniferous trees as well.

All of the species reported in this paper were taken on conifers (Table 2), although
two of the genera {Anthocoris and Orius) should be regarded more as casual visitors
from broad-leafed plants. They are, however, broadly foraging predators and do
become involved with insects on Pinus contorta (Reid, 1957). Species of several of
the genera {Acompocoris, Elatophilus, Melanocoris, Cardiastethus, Scoloposcelis, and
Xylocoris ) occur chiefly on species of Pinus. Ten species have been taken on several
species of Picea, six species on several species of Abies, four species on species of
Larix, and two species on Pseudotsuga. The diversity of coniferous hosts in western
North America certainly contributes to the diversity of the Anthocoridae found on
them. All of these hosts are in the Pinaceae; none has been taken from plants of the
family Cupressaceae. Specimens of Dufouriclla ater (Dufour) were taken from Ju-
niperus occidcntalis near Sisters, Oregon, but this species was not taken from Pinus
contorta murrayana or P. pondcrosa occurring in the immediate vicinity. Several of
the taxa, notably Melanocoris longirostris, all five species of Tetraphleps, and Scolopo-
scelis flavicornis, show wide host plant associations. How many of these represent

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Table 2. Occurrence of Anthocoridae on different subspecies of Pinus contorta.

Pi nus

contorta Pinus contorta Pinus contorta

contorta murrayana latifolia

Acompocoris lepidus
Anthocoris antevolens
Anthocoris musculus
Elatophilus dimidiatus
Elatophilus pullus
Elatophilus sp. A
Melanocoris longirostris
Melanocoris nigricornis
Tetraphleps canadensis
Tetraphleps latipennis
Tetraphleps pilosipes
Tetraphleps uniformis
Orius tristicolor
Cardiastethus borealis
Lyctocoris tuberosus
Scoloposcelis flavicornis
Xylocoris californicus
Xylocoris cursitans

OR

BC

CA, OR

WA

CA, OR
CA, OR
CA, OR, MEX

OR

OR

OR

OR, MEX

OR

WA

OR

CA

WA

BC, YT, COLO, WY

ALB

WY

COLO, WY
BC, WY

BC

BC, WY
COLO, WY

OR

BC

MT

hosts upon which the insects breed rather than “sitting” records remains to be
determined. The capability to identify immature individuals to species would be of
great value. It does appear that greater host plant breadth occurs in the northern
regions (i.e., Canada and northern United States) but that may be an artifact of
collecting or the dominance of conifers over broad-leafed trees in the same area.
Again, close attention to the occurrence of immature stages should clarify this matter.

FEEDING HABITS

The Anthocoridae are generally regarded as predaceous, although there is ample
evidence that some plant feeding occurs in some species (e.g., Dicke and Jarvis, 1962;
Chu, 1969; Bachelor and Baranowski, 1975). Members of this family are part of the
guild of predators found on a variety of plants (Carayon, 1961; Anderson, 1962b;
Pericart, 1972; Evans, 1976a, b, c, d), but detailed biological studies have been
confined largely to species occurring in agroecosystems (e.g., Collyer, 1953; Sands,
1957; Carayon, 1961; Anderson, 1962a, b; Chu, 1969; Evans, 1976a, b, c, d; Ryerson
and Stone, 1979; Arbogast, 1979, 1984). Some predaceous species have been intro-
duced as biological control agents (e.g., Davis and Krauss, 1 963; Mitchell and Wright,
1967; Harris, Holms, and Dawson, 1968).

Less information is available on the genera and species found on conifers, and
much of it is contained in taxonomic papers or those dealing with collections of
insects from different localities or from different hosts. Some have well-defined feed-
ing habits that suggest prey specificity, a phenomenon also noticed in some other
insect predators (Thompson, 1951).

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Generalist Feeders

The genus Anthocoris contains many species that feed on a variety of prey, usually
scaled to their size (Carayon, 1961; Evans, 1976a). They forage over the surface of
trees, and, to a lesser extent, shrubs for aphids, scales, young lepidopterous larvae,
thrips, mites, and other small arthropods. Orius contains several very small species
that occur on many different plants, including a variety of forbs and grasses. Both
genera are normally found on deciduous plants but Anthocoris antevolans and A.
musculus, and Orius tristicolor were taken on Pinus contorta (Evans, 1983; Reid,
1957; this study). The genus Tetraphleps is a coniferous analogue of Anthocoris, but
it seems more specific in its feeding habits— chiefly Homoptera, and especially aphids
(Pericart, 1972).

Lyctocoris beneficus (Hiura), a species similar to L. campestris, occurs in Japan
and Korea (Chu, 1969). Chu (1969) provided a detailed study of this species and
Xylocoris galactinus (Fieber). While both species were primarily predaceous, they
were able to survive for short periods feeding on moldy grain. Reproduction did not
occur when restricted to such food.

Specialist Feeders

Aphids: Anthocorids that feed on aphids are surface dwellers. This group contains
the generalist feeders {Anthocoris and Orius)-, but it also contains three genera {Acom-
pocoris, Melanocoris, and Tetraphleps) that appear to feed chiefly upon aphids. Ac-
cording to Pericart (1972), Acompocoris pygmaeus feeds on Cinara pinicola Kalten-
bach and Schizo/achnus pi net i Fabricius in Europe. Acompocoris lepidus occurs on
Pinus contorta murrayana in North America, together with the aphids Cinara spp.
and Essigella spp. Foottit and Mackauer (1990) provide information on populations
of Cinara nigra (Wilson), a widely distributed species found on Pinus contorta.

Melanocoris nigricornis has been taken in association with the aphids Cinara sp.,
Essigella fusca and the pine needle scale, Phenacaspis pinifoliae (Anderson, 1962b).
Both genera of aphids were taken in our collections.

The balsam woolly adelgid, Adelges piceae, is a major pest of the true firs {Abies
spp.) in boreal North America. A native of Europe, it was first recorded from North
America in 1908 from Maine (Drooz, 1985). Today, it occurs in the eastern and far
western parts of Canada and the United States (Fumiss and Carolin, 1977; Foottit
and Mackauer, 1983). Acompocoris lepidus is reported to feed on this insect in the
central Cascade Mountains of Oregon (Mitchell, 1962). Several native species of
Tetraphleps have been reported occurring with the aphid, including T. canadensis
(Kelton, 1966a, 1978), T. latipennis (Kelton, 1966a, 1978; Harris, Holms, and Daw-
son, 1968; Fumiss and Carolin, 1977), and T. uniformis (Kelton, 1966a, 1978).

Bark beetles: A number of genera and species of Anthocoridae are found associated
with bark beetles (Coleoptera: Scolytidae). These bugs are generally found in the
galleries where they feed upon the eggs and larvae of the beetles.

Although not usually regarded as a conifer inhabitant, Anthocoris musculus (Say)
has been recorded as an egg predator of Ips bark beetles on lodgepole pine by Reid
(1957). Kline and Rudinsky ( 1 964), Furniss and Carolin (1977), and Dahlsten (1982)
reported Lyctocoris sp. or spp. from bark beetle galleries. Fronk (1947) reported L.
elongatus (Reuter) as a predator of the southern pine beetle, Dendroctonus frontalis

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Zimmerman, as did Dixon and Payne (1979), Mosier, Thatcher, and Pickard (1971),
Berisford (1980), and Goyer, Lenhart, Nebeker, and Schmitt (1985). Moore (1972)
and Berisford (1980) also recorded L. campestris (F.) feeding on the same beetle.
Kelton (1967) reported L. tuberosus Kelton and Anderson as an associate of Ips spp.
found on several hosts, including P. contorta. He (Kelton, 1967) also reported L.
mexicanus Kelton in association with the bark beetles Ips and Dendroctonus spp.,
the weevils Cossonus spp., and the buprestids Chrysobothris spp.

Carayon (1953a) provided a detailed study of Scoloposcelis obscurella (Zett.) in
France, where the bug feeds upon the larvae of several species of Scolytidae. In North
America, Drake (1922) reported Scoloposcelis flavicornis Reuter (as Anthocoris sp.)
from the galleries of a number of bark and ambrosia beetles in New York. Drake
and Harris (1926) described several species of Scoloposcelis, including S. mississip-
pensis, taken from the burrows of bark beetles on long-leaf pine. Mosier, Thatcher,
and Pickard (1971) reported this species as an associate of the southern pine beetle,
Dendroctonus frontalis Zimmermann, as did Moore (1972), Dixon and Payne (1979),
Berisford (1980) and Goyer, Lenhart, Nebeker, and Schmitt (1985). Dahlsten (1970)
recorded a species of Scoloposcelis (likely S. flavicornis) from Pinus ponderosa bark
disks infested with Dendroctonus brevicomis LeConte. Ohmart (1981) reported S.
flavicornis from the foliage of Pinus radiata D. Don, and from the brood galleries of
Ips paraconfusus Lanier. Dahlsten (1982) found Scoloposcelis spp. common in adult
and egg galleries of bark beetles. He pointed out the problems predators face in coping
with the clumped distribution of their prey and that they may locate them by chance
encounters; however, Borden (1982), citing Dixon and Payne (1979), reported a
species of Scoloposcelis that was attracted to the aggregation pheromones of Scoly-
tidae.

Xylocoris cursitans (Fallen) is common under bark. According to Chu (1969),
Tragardh (1914) was the first to report it as a predator of bark beetle larvae. He also
cited Morkrzecki’s (1923) record of X. cursitans feeding on Ips typographus Linnaeus
in Europe.

Bark-lice: Cardiastethus fasciiventris (Garbiglietti), a European species, is reported
to feed on Psocoptera (Pericart, 1972), but no specific prey data have been given for
C. borealis Kelton, our only North American species found on conifers. Cardiastethus
borealis has been found on Pinus ponderosa, P. banksiana, and P. sylvestris (Kelton,
1978) and report it here on P. contorta latifolia in central Oregon. Psocoptera are
usually abundant on most of these species of pines, and thus, potentially available
as prey.

Mites: In Europe, Brachysteles parvicornis (Costa), reported recently from North
America by Asquith and Lattin (1990), feeds on members of the mite suborder
Oribatida in Europe (Carayon, cited in Pericart, 1972). Specimens were collected in
Maine and Massachusetts, on Picea glauca (Moench) Voss and Pinus rigida Miller,
in association with the cosmopolitan oribatid Humerobates rostrolamellatus Grand-
jean (Family Ceratozetidae) (Lattin and Asquith, 1991).

Needle miners: Struble (1967) reported occasional predation on the lodgepole nee-
dle miner, Coleotechnites milleri (Busck) (Lepidoptera: Gelechiidae) on Pinus contorta
murrayana by an unidentified species of Anthocoridae in Yosemite National Park,
California.

Scale insects: Anderson (1958) associated Elatophilus pullus (as Elatophilus sp. b.)

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and Melanocoris nigricornis (Anderson, 1962b) with the pine needle scale, Chionapis
(Phenacaspis) pinifoliae. Species of the genus Elatophilus feed primarily on various
Coccidoidea, chiefly species of the scale genus Matsucoccus (Homoptera: Magarodi-
dae) (Biliotti and Riom, 1967; Cobben and Amaud, 1969; Drooz, 1985; Mendel,
Carmi, and Podoler, 1991). Lussier (1965) provided a detailed study of Elatophilus
inimica preying on red pine scale, Matsucoccus resinosae. Although there is at least
one species of Matsucoccus and several species of Elatophilus on lodgepole pine, no
direct feeding observations have yet been made.

Subcorticular insects: The flattened shape of Anthocoridae makes them well adapt-
ed for preying on insects under bark (Hamilton, 1978). Some species of Elatophilus
are known to feed on scales in the genus Matsucoccus (Lussier, 1965; Mendel, Carmi
and Podoler, 1991). Lyctocoris campestris (F.) has been taken under bark (Anderson,
1962b), as have other species of Lyctocoris (e.g., Kelton, 1967; Goyer, Lenhart,
Nebeker, and Schmitt, 1985). Scoloposcelis jlavicornis Reuter is a subcorticular spe-
cies that also enters bark beetle burrows (e.g., Drake, 1922; Mosier, Thatcher, and
Pickart, 1971; Goyer, Lenhart, Nebeker, and Schmitt, 1985). Xylocoris cursitans is
commonly collected under the bark of dead trees (e.g., Tragardh, 1914; Anderson,
1962b; Pericart, 1972; Kelton, 1978). One from the type series of Xylocoris umbrinus
was collected under the bark of an old fir log (Van Duzee, 1921). Many other species
of Xylocoris are found in leaf litter (e.g., Anderson, 1962b; Carayon, 1972b; Pericart,
1972; Kerzhner and Elov, 1976; Kelton, 1976a).

INTRODUCTIONS

At least twelve species of Anthocoridae have been introduced into North America
either accidentally [(e.g., Acompocoris pygmaeus (Fallen), Brachysteles parvicornis
(Costa), and Orius minutus (Linneaus)] or deliberately [e.g., Anthocoris nemoralis
(Fabricius)]; none has yet been taken on Pinus contorta (Anderson, 1 962b; McMullen
and Jong, 1967; Kelton, 1978; Henry, 1988; Lattin, Asquith, and Booth, 1989;
Asquith and Lattin, 1990; Lattin and Asquith, 1991). At least four species are conifer
associates (e.g., Acompocoris pygmaeus, two species of Tetraphleps and Brachysteles
parvicornis), while the others are found on broad-leafed hosts, in litter, or in stored-
grain facilities. Tetraphleps abdulghanii Ghauri and T. raoi Ghauri have been intro-
duced into North America from Pakistan and India, respectively, as predators of the
balsam woolly adelgid, a pest of trtie firs (Ghauri, 1964; Mitchell and Wright, 1967;
Harris, Holms and Dawson, 1968), but apparently they have not become established
(L. M. Humble and R. G. Mitchell, pers. comm., March, 1990). Xylocoris cursitans
(Fallen), found on P. contorta, may be naturally Holarctic, rather than an introduction;
it occupies essentially the same habitat in both hemispheres— the galleries of beetles,
usually Scolytidae, beneath the bark of both coniferous and deciduous trees. Xylocoris
galactinus (Fieber), found chiefly in organic litter and stored grain, sometimes under
bark, may be an introduction rather than a naturally Holarctic species.

Two species, A. confusus Reuter and A nemoralis (Fabricius), have been introduced
accidentally from Europe. The latter species subsequently was introduced into south-
ern British Columbia from Switzerland in 1 963 as a potential biological control agent
for the pear psylla (McMullen and Jong, 1967; McMullen, 1971; Fields and Beirne,
1973). This species seems to have adapted to orchards in the Pacific Northwest.

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Much less is known about A. confusus, but its range has been expanding (Kelton,
1978; Scudder, 1986).

WING POLYMORPHISM

In the Anthocoridae wing reduction is usually correlated with cryptic habits; under
and within cracks and crevices of bark; and, to a lesser extent, within the litter layer.
Wing reduction, either by brachyptery or microptery, occurs in a number of genera
of Anthocoridae (Pericart, 1972; Kelton, 1978), including several collected during
this study (. Anthocoris , Elatophilus, and Xylocoris). Brachyptery is more common
than microptery (Pericart, 1972). While wing modification may occur in both sexes,
it is most common in females. Psocoptera show a similar trend (Thornton, 1985).

Hamilton (1978) considered the subcorticular habitat to have been of considerable
evolutionary importance in the insects. He included examples of several genera of
Anthocoridae ( Xylocoris and Anthocoris) in his discussions; representatives of both
of these genera have been collected from P. contorta.

The reduction of wing size and development, accompanied by the loss of flight, is
often associated with some degree of habitat stability (Usinger, 1950; Southwood,
1961, 1962; Hamilton and May, 1977; Slater, 1977; Hamilton, 1978; Denno and
Grissell, 1979; Greenwood, 1988; Rolf, 1990). Insects with reduced wings, or lacking
wings entirely, are found frequently on large bodies of water, including the oceans
(e.g., Halobates ), in the ground or in the litter layer, as ectoparasites, and under bark.
In contrast, insects with high flight activity are normally associated with temporary
habitats (Southwood, 1960).

Wing reduction in the Heteroptera sometimes results in fewer nymphal instars
during development. Stys and Davidova-Vilimova (1989) reviewed unusual numbers
of instars in the Heteroptera and, citing Pericart (1972) for Temnostethus gracilis
Horvath, suggested that Anthocoridae displaying wing reduction may have a reduced
number of developmental stages.

We collected three brachypterous females of Elatophilus dimidiatus (Van Duzee)
from the branches of P. contorta murrayana outside of Lassen National Park in
Tehama Co., California. Another series of specimens was collected above Big Bear
Lake, San Bernardino Co., California, on the same host; the four males were macrop-
terous and the 12 females were brachypterous. We also collected a brachypterous
male and brachypterous females of an undescribed species of Elatophilus from Pinus
ponderosa near Sisters, Oregon. The genus Elatophilus had been regarded as always
macropterous (Pericart, 1972; Kelton, 1976b, 1978), but see our discussion under
E. dimidiatus. A brachypterous female of Xylocoris cursitans (Fallen) was taken under
the bark of P. c. contorta killed by Scolytidae at Bremerton, Washington; a macrop-
terous female was collected at the same time. Kelton (1978) found X. cursitans in
Canada under the bark of dead trees, both coniferous and deciduous. Both sexes
occur in the macropterous and brachypterous state in Europe and North America.
Two additional species, X. galactinus (Fieber) and X. umbrinus (Van Duzee), are
known to occur under the bark of trees in North America, but neither has yet been
collected from P. contorta. Xylocoris galactinus is only known in the macropterous
state, while both sexes of X. umbrinus occur in the macropterous and brachypterous
state.

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It is interesting that brachypterous males and females have ocelli. Many taxa in
the Heteroptera lose their ocelli when the wings are reduced (e.g., Lygaeidae). Ac-
cording to Pericart (1972), brachypterous females in the family Microphysidae lack
ocelli, or else they are vestigial, and the strongly brachypterous Cimicidae lack ocelli
in both sexes. However, Pericart (1972) reported that the brachypterous Xylocoris
thomsoni (Reuter) was the only brachypterous anthocorid lacking ocelli known from
the western Palearctic Region.

RESEARCH NEEDS

This is an interesting family of insects about which we know relatively little. Most
species are predators of small insects and other arthropods, including such pests as
the balsam woolly adelgid, some scale insects and bark beetles. The various genera
and species often show specific host plant and habitat preference, the details of which
are largely unknown. Biological and ecological studies of most species would be useful.
Our knowledge of the distribution of nearly all species is fragmentary at best with
much additional collecting needed. Generic revisions are needed to allow accurate
identification of the various taxa, including consideration of species variation
throughout their range. Sands (1957), Anderson (1958), and Pericart (1972) have
shown that eggs and nymphs can be recognized, often to species, thus encouraging
further work with these stages. Finally, a well-illustrated and updated key to the
genera of the Anthocoridae of North America would be of value.

ACKNOWLEDGMENTS

We thank A. Asquith, G. M. Cooper, J. A. DiGiulio, A. R. Moldenke (all Oregon State
University), R. T. Schuh (American Museum of Natural History), L. M. Shults (University of
Wyoming), M. D. Schwartz (Biosystematics Research Centre, Ottawa), and G. M. Stonedahl
(Commonwealth Institute of Entomology, London) for assistance in collecting specimens; N.
H. Anderson (Oregon State University) for sharing his knowledge of Anthocoridae; J. Carayon
(Natural Museum of Natural History, Paris), J. Pericart, and L. A. Kelton, Ottawa, for their
significant contributions to our knowledge of Anthocoridae; P. H. Amaud (California Academy
of Science), R. Beckwith and T. Torgersen (U. S. Forest Service), R. G. Foottit (Biosystematic
Research Centre, Ottawa), S. Frommer (University of California, Riverside), T. J. Henry
(U.S.D.A., Systematic Entomology Laboratory), and R. L. Westcott (Oregon State Department
of Agriculture), for the loan of specimens and literature under their care; B. Hall for the fine
habitus drawings; D. Watkins, C. Betts, and J. Eggers, for assistance in preparation of the
manuscript, A. Asquith and P. Oman for the review of the manuscript, and to two anonymous
reviewers for their careful work and thoughtful comments on the manuscript. Support to J.D.L.
from NSF grant BSR85 14325 is acknowledged.

LITERATURE CITED

Anderson, J. F., R. P. Ford, J. D. Kegg and J. H. Risley. 1976. A report to the Eastern Plant
Board. The red pine scale in North America. Connecticut Agr. Exp. Sta. Bull. No. 765.

6 pp.

Anderson, N. H. 1958. The distribution and biology of some Anthocoridae (Hemiptera:
Heteroptera) in the Pacific Northwest. M.S. thesis, Oregon State Univ. 90 pp.
Anderson, N. H. 1962a. Bionomics of six species of Anthocoris (Heteroptera: Anthocoridae)
in England. Trans. Royal Entomol. Soc. London 1 14:67-95.

1992

ANTHOCORIDAE ON PINUS CONTORT A

473

Anderson, N. H. 1962b. Anthocoridae of the Pacific Northwest with notes on distributions,
life histories, and habits (Heteroptera). Can. Entomol. 94:1325-1334.

Anderson, N. H. and L. A. Kelton. 1963. A new species of Anthocoris from Canada, with
distribution records for three other species (Heteroptera: Anthocoridae). Can. Entomol.
95:439-442.

Arbogast, R. T. 1 979. The biology and impact of the predatory bug Xylocoris flavipes (Reuter).

Proc. Second Int. Working Conf. Stored-Products Entomol.: 91-105.

Arbogast, R. T. 1984. Demography of the predaceous bug Dufouriellus ater (Hemiptera:
Anthocoridae). Environ. Entomol. 13:990-994.

Arbogast, R. T., B. R. Flaherty and J. W. Press. 1983. Demography of the predaceous bug
Xylocoris sordidus (Reuter). Am. Midi. Nat. 109:398-405.

Asquith, A. and J. D. Lattin. 1990. Brachysteles parvicornis (Costa), an Anthocoridae new to
North America (Hemiptera: Heteroptera). J. New York Entomol. Soc. 98:352-356.
Bachelor, J. S. and R. M. Baranowski. 1975. Paratriphleps laeviusculus, a phytophagous
anthocorid new to the United States (Hemiptera: Anthocoridae). Florida Entomol. 58:
157-163.

Barber, G. W. 1936. Orius insidiosus (Say), an important natural enemy of the com earworm.
U.S. Dept. Agr., Tech. Bull. 504. 24 pp.

Bean, J. L. and P. A. Godwin. 1955. Description and bionomics of a new pine scale, Mat-
sucoccus resinosae. For. Sci. 1:164-176.

Berisford, C. W. 1980. Natural enemies and associated organisms. Pages 31-52, 250-257 in:
R. C. Thatcher, J. L. Searcy, J. E. Coster, and G. D. Hertel (eds.), The Southern Pine
Beetle. U.S. Dept. Agr., For. Ser., Tech. Bull. 1631. 266 pp.

Biliotti, E. and J. Riom. 1 967. Faune corticole du pin maritime, Elatophilus nigricornis (Hem.

Anthocoridae). Ann. Soc. Entomol. France (N.S.) 3:1 103-1 108.

Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America, with Especial
Reference to the Faunas of Indiana and Florida. Nature Pub. Co., Indianapolis.

1116 pp.

Blatchley, W. S. 1928. Two new anthocorids and a new microphysid from Florida (Heter-
optera). Entomol. News 29:85-88.

Borden, J. H. 1982. Aggregation pheromones. Pages 74-139 in: J. B. MittonandK. B. Sturgeon
(eds.), Bark Beetles in North American Conifers. Univ. Texas Press, Austin. 527 pp.
Carayon, J. 1953a. Observations sur Scoloposcelis obscurella (Zett.) (Hemipt. Anthocoridae).
Rev. Sci. Nat. d’ Auvergne 19:65-73.

Carayon, J. 1953b. Existence d’un double orifice genital et d’un tissu conducteur des sper-
matozoides chez les Anthocorinae (Hemipt. Anthocoridae). Comp. rend, seances de
l’Acad. des Sci. 236:1206-1208.

Carayon, J. 1961. Quelques remarques sur les Hemipteres-Heteropteres: leur importance
comme insects auxiliaries et les possibitites de leur utilisation dan la lutte biologique.
Entomophaga 6:133-141.

Carayon, J. 1972a. Caracteres systematiques et classification des Anthocoridae (Hemipt.).
Ann. Soc. Entomol. France (N.S.) 8:309-349.

Carayon, J. 1 972b. Le Genre Xylocoris : subdivision et especes nouvelles (Hem. Anthocoridae).
Ann. Soc. Entomol. France (N.S.) 8:579-606.

Carayon, J. and J. R. Steffan. 1959. Observations sur le regime alimentaire des Orius et
particulierement d 'Orius pallidicornis( Heteroptera: Anthocoridae) (Reuter). Cah. Natur.,
Bull, des Nat. Parisiens (N.S.) 15:53-63.

Champion, G. C. 1897-1901. Insecta: Rhynchota (Hemiptera-Heteroptera). Vol. II, 416 pp.

(1897:1-32; 1898:33-192; 1899:193-304; 1900:305-344; 1901:i-xvi + 345-416).

Chu, Y. I. 1969. On the bionomics of Lyctocoris beneficus (Hiura) and Xylocoris galactinus
(Fieber). (Anthocoridae, Heteroptera). J. Fac. Agr., Kyushu Univ. 15:1-136.

474

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Cobben, R. H. 1978. Evolutionary Trends in Heteroptera. Part II. Mouthpart-Structures and
Feeding Strategies. Mededel. Landbouwhogeschool Wageningen 78-5. J. Veenman and
Zonen B. V., Wageningen, Nederland. 407 pp.

Cobben, R. H. and B. Arnoud. 1 969. Anthocoridae van Viseum, Buxus en Pinus in Nederland
(Heteroptera). Pub. van bet Natuurhist. Genootschap in Limburg 19:5-16.

Collyer, E. 1953. Biology of some predatory insects and mites associated with the fruit tree
red spider mite ( Metatetranychus ulmi (Koch)) in south-eastern England. II. Some im-
portant predators of the mite. J. Hort. Sci. 28:85-97.

Critchfield, W. B. 1957. Geographical variation in Pinus contorta. Harvard University, Cam-
bridge. Maria Moors Cabot Found., Pub. No. 3. 1 18 pp.

Critchfield, W. B. 1980. The genetics of lodgepole pine. U.S. Dept. Agr., For. Ser., Res. Pap.
WO-37:l-37.

Critchfield, W. B. 1985. The late Quaternary history of lodgepole and jack pines. Can. J. For.
Res. 15:749-772.

Critchfield, W. B. and E. L. Little, Jr. 1 966. Geographic distribution of the pines of the world.
U.S. Dept. Agr., For. Ser., Misc. Pub. 991. 97 pp.

Dahlsten, D. L. 1970. Parasites, predators, and associated organisms reared from western
pine beetle infested bark samples. Pages 75-79 in: R. W. Stark and D. L. Dahlsten (eds.),
Studies on the Population Dynamics of the Western Pine Beetle, Dendroctonus brevi-
comis (Coleoptera: Scolytidae). Univ. California, Div. Agr. Sci. 174 pp.

Dahlsten, D. L. 1982. Relationships between bark beetles and their natural enemies. Pages
140-182 in: J. B. Mitton and K. B. Sturgeon (eds.), Bark Beetles in North American
Conifers. Univ. Texas Press, Austin. 527 pp.

Dallas, W. S. 1851-1852. List of the specimens of hemipterous insects in the collection of
the British Museum. Taylor and Francis Inc. London 1851, 1:1-368; 1852, 2:369-592.

Davis, C. J. and N. L. H. Krauss. 1963. Recent introductions for biological control in Hawaii,
8. Proc. Hawaiian Entomol. Soc. 18:248.

Denno, R. F. and E. E. Grissell. 1979. The adaptiveness of wing-dimorphism in the salt
marsh-inhabiting plant hopper, Prokelisia rnarginata (Homoptera: Delphacidae). Ecol-
ogy 60:221-236.

Dicke, F. F. and J. L. Jarvis. 1962. The habits and seasonal abundance of Orius insidiosus
(Say) (Hemiptera-Heteroptera: Anthocoridae) on com. J. Kansas Entomol. Soc. 35:339-
344.

Dixon, W. N. and T. L. Payne. 1979. Sequence of arrival and spatial distribution of ento-
mophagous and associate insects on southern pine beetle-infested trees. Texas Agr. Exp.
Sta. MP-1432. 27 pp.

Doane, C. C. 1965. The red pine scale. Connecticut Agr. Exp. Sta., Circ. 207. (revised). 7 pp.

Downes, W. 1927. A preliminary list of the Heteroptera and Homoptera of British Columbia.
Proc. Entomol. Soc. British Columbia. No. 23:1-22.

Drake, C. J. 1921. A new ambrosia beetle from the Adirondacks; notes on the work of
Xyloterinus politus Say. Ohio J. Sci. 21:201-204.

Drake, C. J. 1922. Heteroptera in the vicinity of Cranberry Lake. Tech. Pub. No. 16, New
York State Coll. For., Syracuse LTniv. 22:54-86.

Drake, C. J. and H. M. Harris. 1926. Notes on American Anthocoridae with descriptions of
new forms. Proc. Biol. Soc. Washington 39:33-36.

Drake, C. J. and H. M. Harris. 1928. Tetraphleps canadensis Provancher, a true Tetraphleps
(Hemip.). Can. Entomol. 60:50.

Drooz, A. T. (Ed.) 1985. Insects of eastern forests. U.S. Dept. Agr., For. Ser., Misc. Pub. No.
1426. 608 pp.

Dufour, L. 1831. Description et figure du Xylocoris rufipennis, Hemiptere nouveau. Ann. Sci.
Natur., Paris 22:423-426.

1992

ANTHOCORIDAE ON PINUS CONTORT A

475

Evans, D. 1983. Annotated checklist of insects associated with native pines in British Co-
lumbia. Environ. Canada, Can. For. Ser., Inf. Rep. BC-X-244. 1 15 pp.

Evans, H. F. 1976a. The role of predator-prey size ratio in determining the efficiency of
capture by Anthocoris nemorum and the escape reactions of its prey, Acyrthosiphon pisum.
Ecol. Entomol. 1:85-90.

Evans, H. F. 1976b. The effect of prey density and host plant characteristics on oviposition
and fertility in Anthocoris confusus (Reuter). Ecol. Entomol. 1:157-161.

Evans, H. F. 1976c. The searching behavior of Anthocoris confusus (Reuter) in relation to
prey density and plant surface topography. Ecol. Entomol. 1:163-169.

Evans, H. F. 1976d. Mutual interference between predatory anthocorids. Ecol. Entomol. 1:
283-286.

Fallen, C. F. 1807. Monographia Cimicum Sueciae. C. G. Proft, Hafniae. 123 pp.

Fallen, C. F. 1814. Specimen Novam Hemiptera Disponendi Methodum Exhibens. Lunde,
Berlin-qualanus. 26 pp.

Fieber, F. X. 1860. 1860-61. Die europaischen Hemipteren. Halbfliigler (Rhynchota Het-
eroptera). Wien. 1860, 1-112; 1861, 113-444.

Fieber, F. X. 1864. Erorterungen zur Nomenclatur der Rhynchoten (Hemiptera) Livland’s.
Wein. Entomol. Monatschrift 7:53-62 (1863).

Fields, G. J. and B. P. Beime. 1973. Ecology of anthocorid (Hemiptera: Anthocoridae) pred-
ators of the pear psylla (Homoptera: Psyllidae) in the Okanagan Valley, British Columbia.
J. Entomol. Soc. British Columbia 70 (1973): 18-1 9.

Foottit, R. G. and M. Mackauer. 1 983. Subspecies of the balsam woolly aphid, Adelges piceae
(Homoptera: Adelgidae) in North America. Ann. Entomol. Soc. Am. 76:299-304.

Foottit, R. G. and M. Mackauer. 1990. Morphometric variation within and between popu-
lations of the pine aphid, Cinara nigra (Wilson) (Homoptera: Aphidoidea: Lachnidae),
in western North America. Can. J. Zool. 68:1410-1419.

Forrest, G. I. 1980a. Variation in monoterpene composition of the shoot cortical oleoresin
within and between trees of Pinus contorta. Biochem. Syst. Ecol. 8:337-341.

Forrest, G. I. 1980b. Geographical variation in the monoterpenes of Pinus contorta oleoresin.
Biochem. Syst. Ecol. 8:343-359.

Fronk, W. D. 1947. The southern pine beetle, its life history. Tech. Bull. Virginia Agr. Exp.
Sta., Tech. Bull. No. 18. 12 pp.

Fumiss, R. L. and V. M. Carolin. 1977. Western forest insects. U.S. Dept. Agr., For. Ser.,
Misc. Pub. No. 1339. 654 pp.

Ghauri, M. S. K. 1964. Notes on the Hemiptera from Pakistan and adjoining areas. Ann.
Mag. Nat. Hist. 7 (13th Series): 673-688.

Goyer, R. A., G. J. Lenhard, T. E. Nebeker and J. J. Schmitt. 1985. Distinguishing immatures
of insect associates of southern pine bark beetles. U.S. Dept. Agr., For. Ser., Coop. St.
Res. Ser., Agr. Handb. 641. 19 pp.

Greenwood, S. R. 1 988. Habitat stability and wing length in two species of arboreal Psocoptera.
Oikos 52:235-238.

Hahn, C. W. 1 832-1836. Die Wanzenartigen Insecten. C. H. Zeh’schen Buchhandlung, Num-
burg 3:17-34. (1836).

Hamilton, W. D. 1978. Evolution and diversity under bark. Pages 154-175 in: L. A. Mound
and N. Waloff (eds.), Diversity of Insect Faunas. Symp. Royal Entomol. Soc. London.
No. 9. Blackwell Sci. Pub., Oxford. 204 pp.

Hamilton, W. D. and R. M. May. 1977. Dispersal in stable habitats. Nature 269:575-581.

Harris, H. M. and W. E. Shull. 1944. A preliminary list of Hemiptera of Idaho. Iowa State
Coll. J. Sci. 18:199-208.

Harris, J. W. E., J. C. V. Holms and A. F. Dawson. 1968. Balsam woolly aphid predator
studies. British Columbia, 1959-1967. Canada Dept. For. Rural Devel., For. Res. Lab.,
Victoria, British Columbia, Inf. Rep. BC-X-23. 18 pp.

476

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Hartzell, A. 1957. Red pine scale with special reference to its host plants and cold hardiness.
Contr. Boyce Thompson Inst. 18:421-428.

Henry, T. J. 1988. Anthocoridae. Pages 12-28 in: T. J. Henry and R. C. Froeschner (eds.),
Catalog of the Heteroptera, or True Bugs of Canada and the Continental United States.
E. J. Brill, Leiden. 958 pp.

Herring, J. L. 1966. The genus Orius of the Western Hemisphere (Hemiptera: Anthocoridae).
Ann. Entomol. Soc. Am. 59:1093-1 109.

Herring, J. L. 1976. Keys to genera of Anthocoridae of America north of Mexico, with
description of a new genus (Hemiptera: Heteroptera). Florida Entomol. 59:143-150.

Hicks, E. A. 1959. Checklist and Bibliography on the Occurrence of Insects in Birds’ Nests.
Iowa State Coll. Press, Ames, Iowa. 681 pp.

Hill, A. R. 1957. A key to the North American members of the genus Anthocoris Fallen
(Hemiptera: Anthocoridae). Pan-Pac. Entomol. 33:171-174.

Hiura, I. 1966. Contribution to the knowledge of Anthocoridae from Japan and its adjacent
territories (Hemiptera-Heteroptera). Bull. Osaka Mus. Nat. Hist., No. 19:29-37.

Kelton, L. A. 1963. Synopsis of the genus Orius Wolff in America north of Mexico (Heter-
optera: Anthocoridae). Can. Entomol. 95:631-636.

Kelton, L. A. 1 966a. Synopsis of the genus Tetraphleps Fieber in North America (Hemiptera:
Anthocoridae). Can. Entomol. 98:199-204.

Kelton, L. A. 1966b. A new species of Lyctocoris Hahn from Mexico (Heteroptera: Antho-
coridae). Can. Entomol. 98:320-321.

Kelton, L. A. 1967. Synopsis of the genus Lyctocoris in North America and description of a
new species from Quebec (Heteroptera: Anthocoridae). Can. Entomol. 99:807-814.

Kelton, L. A. 1968. On the Heteroptera in the Provancher collection (Hemiptera). Natur.
Canadien 95:1065-1080.

Kelton, L. A. 1976a. Three new species of Xylocoris from North America, and a note on the
status of species in the genus Scoloposcelis Fieber (Heteroptera: Anthocoridae). Can.
Entomol. 108:193-198.

Kelton, L. A. 1976b. The genus Elatophilus Reuter in North America with descriptions of
two new species (Heteroptera: Anthocoridae). Can. Entomol. 108:631-634.

Kelton, L. A. 1977a. New species of Cardiastethus Fieber and Melanocoris Champion, and
new records of European Acompocoris Reuter and Temnostethus Fieber in Canada (Het-
eroptera: Anthocoridae). Can. Entomol. 109:243-248.

Kelton, L. A. 1977b. A new species of Elatophilus Reuter from Ontario, and new synonymy
for Piezostethus flaccidus Van Duzee (Heteroptera: Anthocoridae). Can. Entomol. 109:
1017-1018.

Kelton, L. A. 1978. The insects and arachnids of Canada. Part 4. The Anthocoridae of Canada
and Alaska. Heteroptera: Anthocoridae. Canada Dept. Agr., Pub. 1639. 101 pp.

Kelton, L. A. and N. H. Anderson. 1962. New Anthocoridae from North America, with notes
on the status of some genera and species (Heteroptera). Can. Entomol. 94:1302-1309.

Kerzhner, I. M. 1988. Family Anthocoridae. Pages 768-778 in: P. H. Ler (ed.), Keys to the
Identification of Insects of the Soviet Far East. Vol. 2: Homoptera and Heteroptera.
Nauka. Leningrad. 972 pp.

Kerzhner, I. M. and E. S. Elov. 1976. Bugs of the genus Xylocoris Duf., subgenus Proxylocoris
Carayon (Heteroptera, Anthocoridae) in the USSR and neighboring countries. Entomol.
Rev. 55:364-368.

Kinzer, H. G. 1976. A synoptic list of insects associated with seeds and cones of New Mexico
conifers. New Mexico State Univ. Agr. Exp. Sta., Res. Rep. 311:6 pp.

Kinzer, H. G., B. J. Ridgill and J. G. Watts. 1972. Seed and cone insects of ponderosa pine.
New Mexico Agr. Exp. Sta., Bull. No. 594. 36 pp.

Kline, L. N. and J. A. Rudinsky. 1964. Predators and parasites of the Douglas-fir beetles:

1992

ANTHOCORIDAE ON PINUS CONTORT A

All

description and identification of the immature stages. Oregon State Univ., Agr. Exp.
Sta., Tech. Bull. 79. 52 pp.

Kosztarab, M. and F. Kozar. 1988. Scale Insects of Central Europe. Dr. W. Junk, Dordrecht.
456 pp.

Lattin, J. B. and A. Asquith. 1991. Contemporary records of Brachysteles parvicornis (Costa)
in the United States (Hemiptera: Heteroptera: Anthocoridae). J. New York Entomol.
Soc. 99:240-241.

Lattin, J. D., A. Asquith and S. Booth. 1989. Orius minutus (Linneaus) in North America
(Hemiptera: Heteroptera: Anthocoridae). J. New York Entomol. Soc. 97:409-416.

LeQuesne, W. J. 1955. An examination of the genus Temnostethus Fieb. (Hem., Anthocoridae)
in Britain and an addition to the British list. Entomol. Mon. Mag. 91 (1095):259-262.

Lethierry, L. F. and G. Severin. 1896. Catalogue general des Hemipteres 3:1-275. R. Fried-
lander et fils, Bruxelles et Berlin.

Little, E. L., Jr. 1979. Checklist of United States trees (native and naturalized). U.S. Dept.
Agr., For. Ser., Agr. Handb. No. 541. 375 pp.

Lussier, S. J. 1965. A study of Elatophilus inimica D. & H. (Hemiptera: Anthocoridae) and
its role in the natural control of the red-pine scale Matsucoccus resinosae B. & G.
(Homoptera: Margarodidae). M.S. thesis, Univ. Massachusetts, Amherst. 42 pp.

MacPhee, A. W. and K. H. Sanford. 1954. The influence of spray programs on the fauna of
apple orchards in Nova Scotia. VII: effects on some beneficial arthropods. Can. Entomol.
86:128-135.

McCune, B. 1988. Ecological diversity of North American pines. Amer. J. Bot. 75:353-368.

McKenzie, H. L. 1942a. New species of pine-infesting Margarodidae from California and
southwestern United States (Homoptera: Coccoidea: Margarodidae). Microentomology
7:1-18.

McKenzie, H. L. 1 942b. Season history of the margarodid scale, Matsucoccus bisetosus Mor-
rison, occurring on ponderosa and Jeffrey pines in California (Homoptera: Coccoidea:
Margarodidae). Microentomology 7:19-24.

McMullen, R. D. 1971. Psylla pyricola Foerster, pear pyslla (Hemiptera: Psyllidae). Pages 33-
38 in: Biological Control Programs against Insects and Weeds in Canada. Tech. Comm.
Commonwealth Inst. Biol. Contr. Vol. 4.

McMullen, R. D. and C. Jong. 1967. New records and discussions of predators of the pear
psylla, P. pyricola Foerster, in British Columbia. Proc. Entomol. Soc. British Columbia
64:35-40.

Mendel, Z., E. Carmi and H. Podoler. 1991. Relations between the genera Matsucoccus
(Homoptera: Margarodidae) and Elatophilus (Hemiptera: Anthocoridae) and their sig-
nificance. Ann. Entomol. Soc. Am. 84:502-507.

Mitchell, R. G. 1962. Balsam woolly aphid predators native to Oregon and Washington.
Oregon State Univ., Agr. Exp. Sta., Tech. Bull. 62: 63 pp.

Mitchell, R. G. and K. H. Wright. 1967. Foreign predator introductions for control of the
balsam woolly aphid in the Pacific Northwest. J. Econ. Entomol. 60:140-147.

Moore, G. E. 1 972. Southern pine beetle mortality in North Carolina caused by parasites and
predators. Environ. Entomol. 1:58-65.

Morkrzecki, Z. 1923. Sprawozdanie z. dzialalnosci Zakladow ochrony Lasu i Entomologji w
Skiemiewicach. Ecole Superior Agriculture a Varsovie L (1922-23):32.

Morrison, H. 1939. Descriptions of new species of Matsucoccus. (Homoptera: Coccidae).
Proc. Entomol. Soc. Washington 41:1-20.

Mosier, J. C., R. C. Thatcher and L. S. Pickard. 1971. Relative abundance of Southern Pine
Beetle associates in east Texas. Ann. Entomol. Soc. Am. 64:72-77.

Ohmart, C. P. 1981. An annotated list of insects associated with Pinus radiata D. Don in
California. Commonwealth Scientific and Industrial Research Organization, Canberra,
Australia. Div. For. Res., Div. Rep. No. 8. 50 pp.

478

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Ohmart, C. P. and W. G. Voight. 1981. Arthropod communities in the crowns of the natural
and planted stands of Pinus radiata (Monterey pine) in California. Can. Entomol. 1 13:
673-684.

Parshley, H. M. 1920. Hemiptera from Peaks Island, Maine, collected by Mr. G. A. Moore.
Can. Entomol. 52:80-87.

Pericart, J. 1967. Note taxonomique au sujet du genre Elatophilus Reuter. Description d’une
espece nouvelle et observations diverses. Bull. Soc. Entomol. France 72:52-60.

Pericart, J. 1972. Hemipteres Anthocoridae, Cimicidae et Microphysidae de l’ouest-Palearc-
tique. Faune de L’Europe et du Bassin Mediterraneen No. 7. Masson et Cie, Paris.
402 pp.

Provancher, L. 1885-1890. Petite faune entomologique de Canada et particulierement de la
Province de Quebec. Vol. 3, Cinquieme Ordre les Hemipteres. Natur. Canadien 1885:
1-64; 1886:65-112; 1887:113-184; 1888:185-204; 1889:205-282; 1890:283-354, pi.
1-5.

Ray, C. H., Jr. and M. L. Williams. 1984. Two new species of Matsucoccus (Homoptera:
Margarodidae) from Arizona and Mexico with a key to species in North America. Ann.
Entomol. Soc. Am. 77:765-769.

Reid, R. W. 1957. The bark beetle complex associated with lodgepole pine slash in Alberta.
Part III — notes on the biologies of several predators with special reference to Enoclerus
sphegeus Fab. (Coleoptera: Cleridae) and two species of mites. Can. Entomol. 89:1 1 1-
120.

Reuter, O. M. 1871. Acanthiidae americanae, descriptae. Ofversight af Kongliga Svenska
Vetenskaps-Akademiens Forhandlinger 28:557-568.

Reuter, O. M. 1 875. Genera Cimicidarum Europae. Bihang till Kongliga Svenska Vetenskaps-
Akademiens Handlingar 3:1-66.

Reuter, O. M. 1884. Monographia Anthocoridarum orbis terrestris. Acta Soc. Sci. Fennicae
14:555-758.

Reuter, O. M. 1908. Charakteristik und entwickelungsgeschicte der Hemipteren-Fauna (Het-
eroptera, Auchenorrhyncha und Psyllidae) der Palaerktischen coniferen. Acta Soc. Sci.
Fennicae 36:1-129.

Rolf, D. A. 1990. The evolution of flightlessness in insects. Ecol. Monogr. 60:389-421.
Ryerson, S. A. and J. D. Stone. 1979. A selected bibliography of two species of Onus: the
minute pirate bug, Onus tristicolor, and Orius insidiosus (Heteroptera: Anthocoridae).
Bull. Entomol. Soc. Amer. 25:131-135.

Sands, W. A. 1957. The immature stages of some British Anthocoridae (Hemiptera). Trans.
Royal Entomol. Soc. London 109:295-310.

Say, T. 183 1-32. Descriptions of New Species of Heteropterous Hemiptera of North America.
New Harmony, Indiana. 39 pp.

Schuh, R. T. 1986. The influence of cladistics on heteropteran classification. Ann. Rev.
Entomol. 31:67-93.

Schuh, R. T. and P. Stys. 1991. Phylogenetic analysis of Cimicomorphan family relationships
(Heteroptera). J. New York Entomol. Soc. 99:298-350.

Scudder, G. G. E. 1986. Additional Heteroptera new to British Columbia. J. Entomol. Soc.
British Columbia 83 ( 1 986):63— 65.

Slater, J. A. 1977. The incidence and evolutionary significance of wing polymorphism in
lygaeid bugs with particular reference to those of South Africa. Biotropica 9:217-229.
Slosson, A. T. 1906. Additional list of insects taken in alpine region of Mt. Washington.
Entomol. News 17:323-326.

Southwood, T. R. E. 1960. The flight activity of Heteroptera. Trans. Royal Entomol. Soc.
London 112:173-220.

Southwood, T. R. E. 1961. A hormonal theory of the mechanism of wing polymorphism in
Heteroptera. Proc. Royal Entomol. Soc. London (Series A) 36:63-66.

1992

ANTHOCORIDAE ON PINUS CONTORT A

479

Southwood, T. R. E. 1962. Migration of terrestrial arthropods in relation to habitat. Biol.
Rev. 37:171-214.

Southwood, T. R. E. and D. Leston. 1959. Land and Water Bugs of the British Isles. Wame
and Co., London. 436 pp.

Strickland, E. F. 1953. An annotated list of the Hemiptera (S.L.) of Alberta. Can. Entomol.
85:193-214.

Struble, G. R. 1967. Insect enemies in the natural control of the lodgepole needle miner. J.
Econ. Entomol. 60:225-228.

Stys, P. and J. Davidova-Vilimova. 1989. Unusual numbers of instars in Heteroptera: A
review. Acta Entomol. Bohemoslovaca 86:1-32.

Thompson, W. R. 1951. The specificity of host relations in predacious insects. Can. Entomol.
83:262-269.

Thornton, I. W. B. 1985. The geographical and ecological distribution of arboreal Psocoptera.
Ann. Rev. Entomol. 30:175-196.

Tragardh, I. 1914. Skogsentomogiska bidrag. 1-5. Entomol. Tidskrift 3-4:188-209.

Uhler, P. R. 1876. List of the Hemiptera of the region west of the Mississippi River, including
those collected during the Hayden explorations of 1873. Bull. U.S. Geol. Geog. Surv.
Terr. 1:267-361.

Usinger, R. L. 1950. The origin and distribution of apterous Aradidae. Proc. Eighth Int. Cong.
Entomol., Stockholm 1948:174-179.

Van Duzee, E. P. 1912. Synonymy of the Provancher collection of Hemiptera. Can. Entomol.
44:317-329.

Van Duzee, E. P. 1914. A preliminary list of the Hemiptera of San Diego County, California.
Trans. San Diego Soc. Nat. Hist. 1:1-57.

Van Duzee, E. P. 1916. Checklist of the Hemiptera (excepting the Aphididae, Coccidae and
Aleurodidae) of America, north of Mexico. New York Entomol. Soc., New York. XI +

111 pp.

Van Duzee, E. P. 1917a. Catalogue of the Hemiptera of America, north of Mexico excepting
the Aphididae, Coccidae and Aleurodidae. Univ. California Pub. Tech. Bull., Entomol.
No. 2: 902 pp.

Van Duzee, E. P. 1917b. Report upon a collection of Hemiptera made by Walter M. Giffard
in 1916 and 1917, chiefly in California. Proc. California Acad. Sci. (4th Series) 7:249-
318.

Van Duzee, E. P. 1921. Characters of eight new species of North American Anthocoridae or
flower bugs. Proc. California Acad. Sci. (4th Series) 1 1:137-144.

Wagner, E. 1952. Die europaischen Arten der Gattung Orius Wff. (Hemiptera Heteroptera
Anthocoridae). Not. Entomol. 32:22-59.

Weber, P. W. 1953. Recent liberations of beneficial insects in Hawaii, 2. Proc. Hawaiian
Entomol. Soc. 15:128.

Wheeler, N. C., R. P. Guries and D. M. O’Malley. 1983. Biosystematics of the genus Pinus,
subsection Contortae. Biochem. Syst. Ecol. 1 1:333-340.

White, F. B. 1879. Descriptions of new Anthocoridae. Entomol. Mon. Mag. 16:142-148.
Wilde, W. H. A. and T. K. Watson. 1963. Bionomics of the pear psylla, Psylla pyricola
Foerster, in the Okanagan Valley of British Columbia. Can. J. Zool. 41:953-961.

Wolff, J. F. 1800-1811. leones Cimicum Descriptionibus Illustrate. J. J. Palm, Erlangen.

208 pp.

Woodroffe, G. E. 1969. Notes on some Hemiptera-Heteroptera from Aviemore, Inverness-
shire. Entomol. Mon. Mag. 105:165-166.

Woodward, T. E. 1951. A case of persistent attacks on a human by Lyctocoris campestris (F.)
(Hem., Anthocoridae). Entomol. Mon. Mag. 87:44.

Received 3 July 1991; accepted 10 December 1991.

J. New York Entomol. Soc. 100(3):480-487, 1992

SEASONAL OCCURRENCE OF THE MIRIDAE
(HETEROPTERA) ASSOCIATED WITH OHIO BUCKEYE,
AESCULUS GLABRA WILLD., IN MISSOURI

R. L. Blinn

Department of Entomology, North Carolina State University,

Raleigh, North Carolina 27695-7613

Abstract. —Seasonal history of the mirids Lygocoris aesculi (Knight), Microphylellus modestus
Reuter, and Orthotylus aesculicola Blinn was followed on Aesculus glabra Willd. in central
Missouri during 1985-86. The activity period of these univoltine plant bugs was confined to
late March to early June, concurrent with the onset of flowering and leaf flush, with almost
complete overlap between the species. The adult male of L. aesculi and the fifth-instar nymphs
of all 3 species are described and illustrated.

The large plant bug genera Lygocoris Reuter and Orthotylus Fieber are better known
compared to many other mirid genera, with respect to their host plants and distri-
bution; much less is known about the genus Microphylellus Reuter. Members of these
genera are in general arboreal and feed on a variety of trees and shrubs. Ohio buckeye,
Aesculus glabra Willd., is interesting because it is a host of 1 species each of Lygocoris
and Orthotylus, both of which are known only from Missouri. The discovery in 1984
of breeding populations of these little-known plant bugs, Lygocoris aesculi (Knight),
and Orthotylus aesculicola Blinn, and the more widely known Microphylellus mo-
destus Reuter, was the motivation for this study.

Herein I summarize the distribution, seasonal occurrence, and food habits for the
3 mirids found breeding on Ohio buckeye in Missouri. Descriptions of the adult male
of L. aesculi (Knight) and of the fifth instars of the 3 species are also given.

STUDY SITE AND METHODS

The study site in Columbia (Boone Co.), Missouri, was a rocky, southern-exposed
slope in a wooded area dominated by white oak, Quercus alba L., and shagbark
hickory, Carya ovata (Mill.) K. Koch, with an understory of mostly Ohio buckeye,
A. glabra Willd., and serviceberry, Amelanchier arborea (Michx. f.) Fern. The buck-
eyes were a mixture of mature and young trees ranging from 2-10 meters high.

Sampling began at, or slightly before, bud break to determine earliest eclosion (9
April in 1985; 31 March in 1986) and continued every 3 to 7 days until adults were
no longer collected (early June). Mirids were sampled visually by inspecting the foliage
and flowers of Ohio buckeye trees for ca. 30 minutes, and any specimens observed
were captured. If an adequate sample was not obtained by visual inspection, the
branches were shaken into a standard 15 -inch beating net to obtain additional spec-
imens. The numbers were generally small, with a typical sample consisting of 1 0 or
fewer specimens of each species. Immatures were placed in alcohol and sorted to
species and instar in the laboratory; preserved adults and immatures were deposited

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MIRIDAE ON OHIO BUCKEYE

481

as voucher specimens in the North Carolina State University Insect Collection,
Raleigh. All measurements are in millimeters.

RESULTS AND DISCUSSION
Lygocoris aesculi (Knight)

Known only from Missouri, this mirine was originally described by Knight (1953)
as Neolygus aesculi from three females collected on “buckeye leaves” in Kansas City
(Jackson Co.). Blinn and Yonke (1985) collected this species in Columbia (Boone
Co.) on Ohio buckeye.

Adult male (Fig. la). Length 4.83-5.25 (N = 5), width 2.04-2.30. Head: width
0.98-1.03, vertex 0.35-0.38, yellowish, frons with transverse red lines, juga and lora
chiefly red, clypeus with basal half yellowish with red streaks, distal half fuscous.
Rostrum, length 1.60-1.64, reaching to middle of metacoxae, pale, apex fuscous.
Antennae: segment I, length 0.53-0.58, fuscous, dorsal aspect (especially distally)
yellowish brown in some specimens; II, 1.65-1.83, fuscous; III, 0.93-1.05, fuscous;
IV, 0.50-0.65, fuscous. Pronotum: length 0.95-1.05, width at base 1.73-1.88; uni-
formly yellowish brown. Scutellum pale yellowish to yellowish brown, transversely
striate, mesoscutum reddish. Hemelytra yellowish brown to brown. Embolium and
cuneus translucent. Membrane and veins clear to slightly dusky, variously marked
with brownish blotches near anal vein, within areoles near their apices and near
areoles. Ventral surface pale to yellowish. Fuscous stripe extending from venter of
propleura to and including genital segment. Legs pale, apical half of hind femora
reddish to fuscous; tibial spines yellowish; third tarsal segment fuscous. Genitalia:
Figure lb-d.

Remarks. Males of L. aesculi run to viburni (Knight) in Knight’s (1941) key, but
may be distinguished by the black antennal segment II and male genitalia. The left
clasper in lateral view is evenly rounded at the angle between the sensory lobe and
shaft ( sensu Schwartz, 1984), whereas in viburni the angle is sinuate. In Kelton’s
(1971) key, L. aesculi (Knight) runs to knighti Kelton, but may be distinguished by
the more uniform yellowish brown color, lack of dark rays behind the calli (present
in knighti ), and by the structure of the male genitalia. The left clasper of aesculi lacks
the dorsal prongs present in knighti.

Fifth instar (Fig. 2). Length 3.40-3.96 (N = 5), width 1.60-1.82. Elongate oval,
uniformly green, legs and antennae green to yellowish green, tarsal segment II fuscous
apically, apex of labial segment IV fuscous. Rather densely clothed with pale, re-
cumbent setae. Head: width 0.85-0.93, vertex 0.45-0.49. Rostrum, length 1.25-1.33,
reaching base of mesocoxae. Antennae: segment I, length 0.33-0.36; II, 1.07-1.1 1;
III, 0.82-0.87; IV, 0.45-0.56. Pronotum: length 0.45-0.49, width at base 1.18-1.29.
Wing pads reaching abdominal segment IV; dorsal scent gland opening pale, indis-
tinct. Parempodia fleshy, divergent apically.

Microphylellus modestus Reuter

This mirid is widely distributed in eastern North America, extending from Quebec
south to North Carolina and west to Saskatchewan and Texas (Henry and Wheeler,
1988). In addition to Ohio buckeye, M. modestus has been recorded from Acer sp.,

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Figs. 1-4. Fig. la-d. Lygocoris aesculi adult, la. Adult male habitus, lb. Left paramere,
lateral view. lc. Left paramere, dorsal view. Id. Right paramere, ventral view. Fig. 2. Lygocoris
aesculi, fifth-instar nymph. Fig. 3. Microphylellus modestus, fifth-instar nymph. Fig. 4. Ortho-
tylus aesculicola, fifth-instar nymph.

Catalpa sp., Fraxinus pennsylvanica Marsh., Quercus velutina Lam. (Blinn and Yonke,
1985), Q. macrocarpa Michx. (Kelton, 1980), Carya sp., Corylus americana Walt.,
Crataegus mollis (T. & G.) Scheele, Q. alba L., Ulmus sp. (Knight, 1941), and Q.
stellata Wangenh. (Wheeler et al., 1983).

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MIRIDAE ON OHIO BUCKEYE

483

Although adults have been collected from a large number of plants and are believed
to breed on Ulmus sp., there is no published evidence that any of these plants, other
than Ohio buckeye, serve as true breeding hosts. Knight (1941) observed this species
on elm leaves curled by aphids, where it fed to some extent on honeydew; less often,
he observed it feeding on the eggs of the elm leaf beetle, Pyrrhalta luteola (Muller).

Fifth instar { Fig. 3). Length 2.15-2.64 (N - 10), width 0.87-1.18. Elongate oval,
uniformily green, setigerous spot near apex of antennal segment I, basal annulus on
antennal segments II and III, setigerous spot located anteriorly near apex of pro- and
mesofemora, metafemora with 2 setigerous spots near apex, 1 anteriorly and a smaller
1 posteriorly, and spots at base of spines on tibiae fuscous. Densely clothed with
pale recumbent setae. Head: width 0.55-0.62; vertex 0.31-0.37. Rostrum, length
0.80-0.96, extending to between metacoxae. Antennae: segment I, length 0.16-0.19;
II, 0.49-0.55; III, 0.40-0.45; IV, 0.32-0.36. Pronotum: length 0.35-0.39, width at
base 0.64-0.75. Wing pads reaching to or slightly beyond abdominal segment IV,
dorsal scent gland opening slitlike. Parempodia to setiform, parallel.

Orthotylus aesculicola Blinn

This orthotyline was originally described from material collected in Columbia
(Boone Co.), Missouri, on Ohio buckeye by Blinn (1987), who illustrated the adult
and male genitalia.

Fifth instar { Fig. 4). Length 3.09-3.81 (N = 8), width 0.96-1.16. Elongate slender,
uniformly green to yellowish green, head tinged with orange near eyes. Sparsely
clothed with pale, recumbent setae. Head: width 0.73-0.76; vertex 0.35-0.44. Ros-
trum, length 0.98-1.11, extending to apex of mesocoxae. Antennae: segment I, length
0.33-0.38; II, 1.02-1.09; III, 0.69-0.75; IV, 0.49-0.56. Pronotum: length 0.38-0.45,
width at base 0.75-0.85. Wing pads reaching abdominal segment IV, dorsal scent
gland opening slitlike, anterior area tinged with orange (faded in preserved specimens).
Parempodia fleshy, convergent apically.

Seasonal History

Sampling results are given in Figures 5 and 6. Figure 5 gives the percentage of all
stages of each species relative to the total number of Miridae collected per sampling
date. Figure 6 shows the immatures and adults of each species separately as a per-
centage of the total immature, and adult catch, respectively, per sampling date.

The life cycles of these 3 species are similar and can be summarized as follows. All
three species were present from late March or early April to the end of May, with
almost complete overlap among the species (Fig. 5). Overwintering eggs begin to
hatch from late March to early April, coinciding with leaf flush and the onset of
flowering. All instars of these species are active feeding stages, and development takes
place during a time when the host plant is producing new growth and flowers. Early
instars (I— III) of both L. aesculi and O. aesculicola were observed feeding on the
expanding leaflets and flowers where, with their uniform green color, they resembled
the color of the young foliage and flowers. Later instars (IV-V) and adults of L.
aesculi and O. aesculicola were observed feeding on both the flowers and leaves at
about equal frequencies, with the former species preferring the flowers and the latter
species the underside of leaves. Feeding by M. modestus was not observed.

484

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

0)

CQ

*u

o

L_

o

LU

O

DC

LU

CL

SAMPLING DATES (1985 & 1986)

Fig. 5. Seasonal occurrence of the three species of Miridae on Aesculus glabra during 1985
and 1986. Adults and immatures combined for each plant bug species. Solid line = Lygocoris
aesculr, dotted line = Microphylellus modestus; dashed line = Orthotylus aesculicola.

\

Developmental time from first instar to the adult molt takes ca. 3.5 to 4.5 weeks,
with each of the 5 nymphal instars taking from 4 to 7 days. Adults can be found in
the field for 3 to 4 weeks following eclosion from late April through May where they
mate and the females deposit the overwintering eggs. By late May or early June the
active portion of the life cycle of these mirids has ended, with the eggs overwintering
to complete the life cycle. There was no evidence of more than a single generation
per year for each species, which is consistent with what has been reported for con-
geners (Knight, 1917; Pinto, 1982; Southwood and Leston, 1959; Wheeler, 1982).

M. modestus was the first species to appear in the sampling. First instars were
present from 3 1 March to 4 April. Instars II and III were first collected on 7 April,
and 9 April, respectively and were present until 21 April and 25 April, respectively.
Fourth instars were collected only from 21 April to 23 April. Fifth instars were
present from 23 April to 3 May. Adults first appeared in the samples on 3 May and
were present until 30 May.

For L. aesculi the earliest specimen observed was a first instar collected on 2 April

Fig. 6. Seasonal occurrence of the three species of Miridae on Aesculus glabra during 1985
and 1986. Dotted lines indicate immatures, solid lines adults.

PERCENT OF TOTAL IMMATURE/ADULT Miridae

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MIRIDAE ON OHIO BUCKEYE

485

SAMPLING DATES (1985 & 1986)

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

1986. First instars were present from 2 April through 9 April. Instars II to IV were
present from 9 April through 1 6 April, 1 1 April, and 30 April, respectively. Fifth
instars were present from 1 1 April through 23 April. Adults began to appear during
the last week of April and were present only for 3-4 weeks; the last male was collected
on 1 7 May, the last female on 2 1 May. In the sampling, adults of this univoltine
mirid were no longer present after 21 May in 1985 and 20 May in 1986.

For O. aesculicola instars I to III were first collected on 7 April and were present
through 1 6 April for the first instar and 2 1 April for both instars II and III. Fourth
instars were present in the samples from 16 April through 30 April. Fifth instars
were present from 23 April through 6 May. Adults first appeared during the last week
of April and were present for ca. 4 weeks. In the sampling, adults of this univoltine
mirid were no longer present after 30 May in 1985 and 20 May in 1986.

No sign of damage was observed in the field as a result of feeding by these 3 plant
bug species, perhaps due to their low population densities. In the laboratory, both
nymphs and adults failed to produce any visible damage when confined to unblem-
ished leaves of Ohio buckeye.

ACKNOWLEDGMENTS

I would like to thank J. R. Baker, L. L. Deitz, and H. H. Neunzig (Department of Entomology,
North Carolina State University) for their comments on the manuscript. Funds for this project
were in part provided by the Missouri Agricultural Experiment Station, Columbia, MO 6521 1
and the North Carolina Agricultural Research Service, Raleigh, NC 27659-7601.

LITERATURE CITED

Blinn, R. L. 1987. Orthotylus aesculicola : a new plant bug from Missouri (Heteroptera: Miri-
dae: Orthotylinae). J. New York Entomol. Soc. 95:57-61.

Blinn, R. L. and T. R. Yonke. 1985. An annotated list of the Miridae of Missouri (Hemiptera:
Heteroptera). Trans. Missouri Acad. Sci. 19:73-98.

Henry, T. J. and A. G. Wheeler, Jr. 1988. Family Miridae Hahn, 1 833 (= Capsidae Burmeister,
1 835) the plant bugs. Pages 251-507 in: T. J. Henry and R. C. Froeschner (eds.), Catalog
of the Heteroptera or True Bugs, of Canada and the Continental United States. E. J.
Brill, Leiden and New York. 958 pp.

Kelton, L. A. 1971. Review of Lygocoris species found in Canada and Alaska (Heteroptera:
Miridae). Mem. Entomol. Soc. Canada 83:1-87.

Kelton, L. A. 1 980. The plant bugs of the prairie provinces of Canada. Heteroptera: Miridae.

The Insects and Arachnids of Canada. Part 8. Agriculture Canada. Ottawa. 408 pp.
Knight, H. H. 1917. A revision of the genus Lygus as it occurs in America north of Mexico,
with biological data on the species from New York. Cornell Univ. Agric. Exp. Stn. Bull.
391:555-645.

Knight, H. H. 1941. The plant bugs, or Miridae of Illinois. Bull. Illinois State Nat. Hist. Surv.
22:1-234.

Knight, H. H. 1953. New species of Miridae from Missouri (Hemiptera). Iowa State Coll. J.
Sci. 27:509-518.

Pinto, J. D. 1982. The phenology of the plant bugs (Hemiptera: Miridae) associated with
Ceanothus crassifolius in a chaparral community of southern California. Proc. Entomol.
Soc. Wash. 84:102-1 10.

Schwartz, M. D. 1984. A revision of the black grass bug genus Irbisia Reuter (Heteroptera:
Miridae). J. New York Entomol. Soc. 92:193-306.

1992

MIRIDAE ON OHIO BUCKEYE

487

Southwood, T. R. E. and D. Leston. 1959. Land and Water Bugs of the British Isles. Frederick
Wame, London & New York. 436 pp.

Wheeler, A. G., Jr. 1982. Coccobaphes sanquinarius and Lygocoris vitticollis (Hemiptera:
Miridae): seasonal history and description of fifth-instar, with notes on other mirids
associated with maple. Proc. Entomol. Soc. Wash. 84:177-183.

Wheeler, A. G., Jr., T. J. Henry and T. L. Mason, Jr. 1983. An annotated list of the Miridae
of West Virginia (Hemiptera-Heteroptera). Trans. Am. Entomol. Soc. 109:127-159.

Received 16 July 1991; accepted 5 September 1991.

J. New York Entomol. Soc. 100(3):488-490, 1992

PRIAPISMUS PINIy A NEW SPECIES OF
DISCOCEPHALINI FROM HONDURAS ON PINE
(HEMIPTERA: PENTATOMIDAE: DISCOCEPH ALIN AE)

L. H. Rolston

Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University Agricultural Center, Louisiana State University,

Baton Rouge, Louisiana 70803

Abstract. — A new species of pentatomid, Priapismus pini, is described from Honduras. Both
nymphs and adults were collected on Pinus maximinoi H. E. Moore.

The genus Priapismus Distant, 1889, ranges from Ecuador to Honduras and cur-
rently contains five species (Rolston, 1984). None of the species is commonly col-
lected, and there has been no host record for any of them until now. Several nymphal
instars as well as adults of the species described here were collected under the bark
of Pinus maximinoi H. E. Moore in Honduras. Although the trees were stressed and
covered with sooty mold, the bug’s role in this situation is unclear.

Priapismus pini, new species

Flavescent ground color of dorsum largely obscured on head, pronotum and scu-
tellum by black punctation and suffusion from punctures; ground color predominant
on coria. Rufous flecks often present in some pale areas of dorsum, especially along
R + M vein of each corium and on midline of pronotum. Venter predominantly black
with small, pale markings, especially on lateral borders of abdomen.

Head densely punctate including tylus or at least postclypeal part (Fig. 1). Vertex
sometimes bearing small, flavescent, irregularly shaped macule partially surrounding
cluster of black punctures. Each antenna black excepting superior surface of basal
segment, joints between segments, and basal or subbasal band of fifth segment pale.
First two segments of rostrum flavescent, third somewhat infuscated basally, fourth
darkening gradually to black apically; apex reaching to or onto fourth visible ab-
dominal sternite.

Pronotum consistently marked with pale, mesial vitta of very irregular outline;
other pale markings on pronotal disk consisting of scattered, usually small macules.

Scutellum consistently marked with pale callus adjacent to fovea in each basal
angle; fovea black, punctate. Scutellar apex with three inconspicuous, pale macules,
one mesial and one on each side just cephalad of mesial macule; these lateral macules
may meet, forming transverse bar, or all three macules become confluent, forming
Y. Additional small, pale macules usually present on scutellar disk, particularly on
midline.

Each corium bearing large, black macule at distal end of R + M vein; punctation
moderately strong, irregular, black, tending to form blotches on endocorium; mem-
brane lightly fumose, its four vestigial veins darker. Connexiva moderately exposed,
alternated flavescent and black with black bordering both sides of transverse sutures.

1992

PRIAPISMUS PINI

489

Figs. 1-9. Priapismus pini. 1. Head. 2. Genital plates. 3. Spermathecal bulb and pump. 4.
Genital cup. 5. Pygophore, caudal view. 6. Paramere, lateral view. 7. Same, rotated ca. 90°
toward viewer. 8. Phallotheca, ventral view. 9. Same, right lateral view. Symbols: bpl, basal
plate; brp, basal projection; c, conjunctiva; d, diverticulum; pa, paramere; pe, penisfilum; pf,
proximal flange; pr, proctiger; sb, spermathecal bulb; t, tubercle.

Venter predominately fuscous; head except bucculae sometimes extensively fla-
vescent; posterolateral border of each metapleuron and middle of lateral margins of
each abdominal stemite flavescent; a few small macules usually scattered elsewhere
on venter. Legs flavescent, heavily marked with fuscous spots and macules. Abdom-
inal venter impunctate mesially; lateral punctation becoming denser and stronger as
far as spiracles, then weaker. Spiracles and surrounding area fuscous.

Measurements (mm, both sexes). Total length 6. 8-7. 5. Width of head 1.75-1.90,
length 1.20-1.35, length to ocelli 0.95-1.05; interocular width 1 .00- 1.15; width across
ocelli 0.80-0.90. Segments 1-5 of antennae: 0.30-0.35; 0.30-0.40; 0.55-0.70; 0.65-
0.75; 1.00-1.10. Segments 1-4 of rostrum: 0.75-0.95; 1.00-1.15; 0.90-1.00; 1.10-
1.15. Pronotal width 3. 5-3.9, mesial length 1.6- 1.8. Scutellar width at base 2.4-2. 6,
width at distal end of frena 1.6- 1.8, length 3. 1-3.5.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Genitalia. 9. Posterior margin of basal plates sigmoid from caudoventral view,
border curving abruptly dorsad from disk; mesial margin of each plate raised in
narrow ridge, adjacent half of disk slightly impressed (Fig. 2). Spermathecal duct
simple; sclerotized duct of spermathecal pump constricted distad of proximal flange
(Fig. 3); spermathecal bulb globose, bearing three short diverticula.

6. Proctiger largely covering genital cup, rather thickly clothed distally with mod-
erately long hairs, widely separated by membrane from anterior margin of genital
cup (Fig. 4). Only flattened, distal ends of parameres visible in cup. Pygophore
noticeably impressed adjacent to anterolateral rim of genital cup; posterior pygopho-
ral margin sinuously transverse from dorsal view, widely and shallowly emarginate
from caudal view (Fig. 5). Each paramere with large, compressed, basal projection;
apex flattened and bent dorsolaterad; small, setose tubercle located on shaft near
apical bend (Fig. 6); shaft essentially triangular with many small hairs on dorsal
surface (Fig. 7). Phallotheca without appendages; penisfilum sigmoid, asymmetrically
flanged where conjunctiva joins (Figs. 8, 9); conjunctiva lacking diverticula.

Types. Holotype, male labeled “Honduras: Cortes St. Cusuco Nat. Park[.] 26 I
1 99 1 [.] On Pinus maximinoi[.] Coll: R.S. Cameron.” Deposited in National Museum
of Natural History, Washington, D.C. Paratypes: 4 males, 5 females with same
labeling as holotype. Punctuation in brackets is not on labels.

Comments. This species is most like P. costaricensis Rolston, 1984, a species
described from females. Measurements of these two species are quite similar, but
their color and punctation are dissimilar. P. pini is the only known species of the
genus that is predominantly dark in color, and the denser punctation, particularly
on the head, separates it from P. costaricensis.

ACKNOWLEDGMENTS

I am indebted to R. S. Cameron, of the Texas Forest Service, for making the specimens of
this species available to me, and to J. B. Chapin, J. A. Moore and D. A. Rider for reviewing
the manuscript.

Approved for publication by the director of the Louisiana Agricultural Experiment Station
as manuscript 91-17-5202.

LITERATURE CITED

Rolston, L. H. 1984. A revision of the genus Priapismus Distant (Hemiptera: Pentatomidae).
J. Kans. Entomol. Soc. 57(1): 1 19-126.

Received 17 April 1991; accepted 3 July 1991.

J. New York Entomol. Soc. 100(3):49 1-497, 1992

LIFE HISTORY AND DESCRIPTIONS OF THE IMMATURE
STAGES OF THE PLANTHOPPER PROKELISIA CROCEA
(VAN DUZEE) (HOMOPTERA: DELPHACIDAE)

Monte W. Holder and Stephen W. Wilson

Department of Biology, Central Missouri State University,
Warrensburg, Missouri 64093

Abstract.— The life history of Prokelisia crocea (Van Duzee) was studied in Missouri from
May to October 1988 and the immature stages described. P. crocea feeds and reproduces on
prairie cordgrass, Spartina pectinata Link, is trivoltine and overwinters as nymphs. Nymphal
instars differed in body size, development of wingpads, number of metatibial spines, and shape
and dentition of metatibial spurs.

Extensive ecological studies of two of the five species of Prokelisia, P. marginata
(Van Duzee) and P. dolus (Wilson), have focused on laboratory rearing, nutrition,
parasites, predators, and wing morphology (literature summarized in Denno et ah,
1985, 1986). The host plants of these salt marsh species are Spartina alterni flora
Loisel and S.foliosa Trin. No information is available on the biology of planthoppers
occurring on the inland species of Spartina, S. pectinata Link.

Prokelisia crocea (Van Duzee) is monophagous on prairie cordgrass (S. pectinata)
and has been recorded from Maine south to Connecticut, and west to North Dakota
and New Mexico (Metcalf, 1943; Wilson, 1982). Wilson (1982) and Heady and
Wilson (1990) provided detailed descriptions of male and female adults of P. crocea.

The present study summarizes the life history of P. crocea on prairie cordgrass in
Missouri, and provides information on laboratory rearing and descriptions and il-
lustrations of immature stages with a key to nymphal instars.

MATERIALS AND METHODS

Field study. The field study was conducted on plots of Spartina pectinata along
roadsides and native tallgrass prairies in Saline, Pettis, and Johnson Counties, Mis-
souri. Weekly sweep and D-vac suction samples (Holder and Wilson, in prep.) were
taken from 24 May to 1 0 October 1988 and 26 April to 1 1 May 1 989. Some specimens
were placed in vials and returned to the laboratory for rearing. Most were preserved
in 70% isopropyl alcohol for phenological analysis, measurements and descriptions.
Information recorded from these samples included number of individuals captured
and number of each nymphal instar. Feeding and oviposition sites were determined,
in both field and laboratory studies, by observing individuals on the host plants.
Attempts were made to collect overwintering nymphs by gathering thatch from host
plant sites in five 20-gallon plastic garbage bags in February, March, November and
December, 1989, and placing it in a large modified Berlese funnel; these samples
produced no planthoppers. A further attempt at collecting specimens with a modified
leaf vacuum (Wilson et al., in prep.) was made on 24 November 1990.

Laboratory study. Plants were collected from roadside areas and potted in 1 2 cm

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

diam pots. An environmental chamber was unavailable thus plants were maintained
in a laboratory under fluorescent lights on a 1 2-hr light-dark cycle; temperature ranged
from 18 to 32°C. Adults were collected and placed on potted plants (Calvert and
Wilson, 1986), nymphs were collected and placed on individual plants but survival
was poor. Immatures were also placed in 2.5 cm diam culture tubes containing a
fresh S. pectinata leaf and plugged with cotton (Calvert et al., 1987). Insufficient data
were collected to determine stadia of nymphal instars due to low survival of nymphs.

The 5th instar is described in detail but only major differences are described for
4th through 1 st instars. Measurements are given in mm as mean ± SD. Length was
measured from apex of vertex to apex of abdomen, width across the widest part of
the body, and thoracic length along the midline from the anterior margin of the
pronotum to the posterior margin of the metanotum. Eggs were obtained by excising
them with a fine needle from the potted host plants.

RESULTS AND DISCUSSION

Field study. P. crocea is trivoltine in Missouri and overwinters as fifth instar nymphs
which were collected on 24 November and 26 April (Figs. 1, 2). Adults were found
from 26 April to 9 June and 23 June to 20 September. First instar nymphs were first
collected on 9 June and found sporadically until 2 October. Second through fifth
instar nymphs showed similar seasonal distributions.

Nymphs and adults were observed feeding along the midrib on the axial surface
of leaves near the base of the leaf and on the stem. Two percent of adult P. crocea
(N = 100) were found to be brachypterous.

Laboratory study. Numerous attempts to rear field collected specimens on potted
S. pectinata in the laboratory were only partially successful. Because an incubator
was unavailable, plants were kept in a laboratory where low humidity apparently
reduced survival of nymphs during hatching and ecdysis.

Females deposit eggs singly in slits made with the ovipositor along the midrib of
either the axial or adaxial surface of a leaf. After eggs have been inserted into the
plant tissue they are covered with wax from the 9th abdominal tergite.

Descriptions. Fifth instar { Figs. 3-6, 16). Length 2.7 ± 0.21; thoracic length 0.9 ±
0.09; width 0.9 ± 0.13. N = 13.

Body straw color with white middorsal line extending from vertex almost to end
of the abdomen. Form elongate, subcylindrical, slightly flattened dorsoventrally,
widest across mesothoracic wingpads.

Vertex subtriangular, narrowing anteriorly; posterior margin concave, lateral mar-
gins concave in anterior V2. Frons yellowish with white longitudinal median line;
border with clypeus slightly convex; lateral margins convex and carinate (outer ca-
l'! nae) and paralleled by second pair of carinae (inner carinae) continuous with lateral
margins of vertex; area between inner carinae forming trough; area between inner
and outer carinae with 9 pits on each side (8 visible in frontal view); 4 pits between
each outer carina and eye. Clypeus yellow, narrowing distally, consisting of subconical
basal postclypeus and cylindrical distal anteclypeus. Beak 3-segmented, cylindrical,
segment 1 hidden by anteclypeus, segment 2 ca. 1.3 x length of segment 3, segment
3 with black apex. Antennae 3-segmented; scape short, cylindrical; pedicel subcy-
lindrical, 2 x length of scape, with ca. 8- 1 0 pit-like sensoria; flagellum bulbous basally.

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PR OKELISIA CROCEA

493

MM JUN JUL AUG SEP OCT MAY JUN JUL AUG SEP OCT

Figs. 1-2. Seasonal occurrence of P. crocea in Missouri. 1. Number of individuals of each
stage is expressed as percentage of total individuals of all stages collected per collection date.
2. Number of individuals of each stage is expressed as percentage of all individuals of that stage
collected throughout the study (NA = 236, N5 = 98, N4 = 127, N3 = 130, N2 = 89, N, = 25).

with elongate bristle-like extension distally, bulbous base ca. 0.2 x length of pedicel.
Eyes red.

Thoracic nota divided by middorsal line into three pairs of plates. Pronotal plates
subtriangular; anterior margin convex following posterior border of eye; posterior

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Figs. 3-6. P. crocea fifth instar. 3. Habitus. 4. Frontal view of head. 5. Apical part of venter
of male abdomen. 6. Apical part of venter of female abdomen. Vertical bar = 1.0 mm (3, 4);
= 0.5 mm (5, 6).

border sinuate; each plate with row of 7 pits originating at anterior margin curving
distally from middorsal line to lateral margin (lateralmost pits often not visible in
dorsal view). Mesonotum with median length 1 .5 x that of pronotum; subrectangular;
elongate lobate wingpads extending to tips of metanotal wingpads; each plate with
posterolaterally directed carina originating on anterior margin in median lA and
terminating on posterior margin in lateral V2; area between carinae elevated and
triangular; 2 pits near carina and 1 pit on lateral VL Metanotum with median length
ca. 1.25 x that of mesonotum; subrectangular, lobate wingpads extending to middle
of 4th tergite; each plate with weak longitudinal carina and 2 pits near middle of
plate. Pro- and mesocoxae elongated and directed posteromedially; metacoxae fused
to sternum. Metatrochanter short and subcylindrical. Pro- and mesofemora and tibiae
with 2 ventral and 1 dorsal longitudinal rows of setae. Metatibia with 2 spines on
lateral aspect of shaft, an apical transverse row of 5 black-tipped spines on plantar
surface and a subtriangular flattened movable spur with 13-14 teeth on posterior
margin. Pro- and mesotarsi with 2 tarsomeres, tarsomere 1 wedge-shaped; tarsomere

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PR OKELISIA CROCEA

495

Figs. 7-11. P. crocea immature stages. 7. Egg. 8. First instar. 9. Second instar. 10. Third
instar. 1 1. Fourth instar. Vertical bar = 0.5 mm (7); = 1.0 mm (8-1 1).

2 subconical, ca. 4 x length of tarsomere 1 , with pair of apical claws and median
membranous pulvillus. Metatarsi with 3 tarsomeres; tarsomere 1 with apical trans-
verse row of 6 black tipped spines on plantar surface; tarsomere 2 cylindrical, ca.
4 x length of tarsomere 1 , with apical transverse row of 4 black tipped spines on
plantar surface; tarsomere 3 subconical, ca. 2 x length of tarsomere 2 with pair of
apical claws and median pulvillus.

Abdomen 9-segmented; slightly flattened dorsoventrally; widest across 5th abdom-
inal segment. Tergite 1 small, subtriangular; 2 subrectangular, ca. 0.5 x width of 3;
tergites 4-8 each with 2 lateral pits on either side (lateralmost pits not always visible
in dorsal view). Segment 9 surrounding anus, with 3 pits; female with 1 pair of acute
processes extending from juncture of stemites 8 and 9; males lacking processes.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Figs. 12-16. P. crocea apices of metathoracic legs. 12. First instar. 13. Second instar. 14.
Third instar. 15. Fourth instar. 16. Fifth instar. Vertical bars = 0.1 mm.

Fourth instar { Figs. 11, 15). Length 2.1 ± 0.26; thoracic length 0.7 ± 0.06; width
0.7 ± 0.09. N = 11.

Antennal pedicel with 4-6 sensoria; basal portion of antennal flagellum 0.3 x length
of pedicel.

Mesonotal wingpads shorter, each covering ca. % of metanotal wingpad laterally.
Metanotal median length 1.125 x that of mesonotum; carinae weaker; wingpad ex-
tending to tergite 2. Metatibial spur slightly smaller, with 8-10 teeth on margin.
Metatarsi with 2 tarsomeres; tarsomere 1 with apical transverse row of 6 black-tipped
spines on plantar surface; tarsomere 2 subconical with 3 black-tipped spines in middle
of tarsomere on plantar surface.

Third instar (Figs. 10, 14). Length 1.8 ± 0.14; thoracic length 0.6 ± 0.09; width
0.6 ± 0.06. N = 11.

Basal flagellum of antenna ca. 0.5 x length of pedicel.

Mesonotal wingpads shorter, each covering lh of metanotal wingpad laterally.
Metanotal wingpad extending to juncture of tergites 1 and 2. Metatibial spur smaller;
with 5 marginal teeth. Metatarsomere 1 with apical transverse row of 5 black-tipped
spines on plantar surface.

Second instar (Figs. 9, 13). Length 1.4 ± 0.15; thoracic length 0.4 ± 0.08; width
0.4 ± 0.07. N = 13.

Mesonotal plates each with 1 pit; wing pads undeveloped. Metanotal wingpads
undeveloped. Metatibia with apical row of 4 black-tipped spines on plantar surface;
spur small with 3 marginal teeth, ca. 3 x length of longest metatibial spine; meta-
tarsomere 1 with 4 apical black-tipped spines.

First instar (Figs. 8, 12). Length 1.1 ±0.15; thoracic length 0.3 ± 0.04; width 0.2
± 0.05. N = 15.

Bulbous base of antennal flagellum subequal in length to that of pedicel.

Metatibia lacking spines on shaft, with apical row of 3 black-tipped spines on
plantar surface; metatibial spur smaller, ca. 1.5 x length of longest metatibial spine,
lacking marginal teeth.

Egg (Fig. 7). Length 0.7 ± 0.05; width 0.2 ± 0.02. N = 8.

Eggs laid singly; white; cylindrical, slightly flattened; curved in anterior lA; chorion
translucent, smooth.

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PROKELISIA CROCEA

497

KEY TO P. CROCEA NYMPHAL INSTARS

1. Metatibial spur with more than 7 marginal teeth; mesonotal wingpads overlapping

more than half length of metanotal wingpads (Figs. 3, 1 1, 15, 16) 2

- Metatibial spur with fewer than 7 marginal teeth; mesonotal wingpads overlap less

than half length of metanotal wing pads (Figs. 8-10, 12-14) 3

2. Metatarsi with 3 tarsomeres; metatibial spur with more than 1 1 marginal teeth (Fig.

16) 5th Instar

- Metatarsi with 2 tarsomeres; metatibial spur with fewer than 1 1 marginal teeth (Fig.

15) 4th Instar

3. Metatibia with transverse row of 5 apical spines, spur with 5 marginal teeth (Fig. 14)

3rd Instar

- Metatibia with fewer than 5 apical spines, spur with fewer than 5 marginal teeth (Figs.

12, 13) 4

4. Metatibia with 2 lateral spines on shaft and 4 apical spines, spur with 3 marginal teeth

(Fig. 13) 2nd Instar

- Metatibia without lateral spines on shaft and with 3 apical spines, spur without marginal

teeth (Fig. 12) 1st Instar

ACKNOWLEDGMENTS

We thank P. D. Calvert, Department of Biology, Central Missouri State University, War-
rensburg, Missouri, 64093, for assistance in sample collection and R. F. Denno, Department
of Entomology, University of Maryland, College Park, Maryland, 20742, for discussion.

LITERATURE CITED

Calvert, P. D., J. H. Tsai and S. W. Wilson. 1987. De/phacodes nigrifacies (Homoptera:
Delphacidae): field biology, laboratory rearing and descriptions of immature stages. Fla.
Entomol. 70:129-134.

Calvert, P. D. and S. W. Wilson. 1986. Life history and descriptions of the immature stages
of the planthopper Stenocranus lautus (Homoptera: Delphacidae). J. New York Entomol.
Soc. 94:118-125.

Denno, R. F., L. W. Douglass and D. Jacobs. 1985. Crowding and host plant nutrition:
environmental determinants of wing-form in Prokelisia marginata. Ecology 66:1588-
1596.

Denno, R. F., L. W. Douglass and D. Jacobs. 1 986. Effects of crowding and host plant nutrition
on a wing-dimorphic planthopper. Ecology 67:1 16-123.

Heady, S. E. and S. W. Wilson. 1990. The planthopper genus Prokelisia (Homoptera: Del-
phacidae): morphology of female genitalia and copulatory behavior. J. Kansas Entomol.
Soc. 63:267-278.

Metcalf, Z. P. 1943. General catalogue of the Hemiptera. Fasc. IV. Fulgoroidea, Part 3.
Araeopidae (Delphacidae), 556 pp.

Wilson, S. W. 1982. The planthopper genus Prokelisia in the United States (Homoptera:
Fulgoroidea: Delphacidae). J. Kansas Entomol. Soc. 55:532-546.

Wilson, S. W., J. L. Smith and A. H. Purcell III. In prep. An inexpensive gas-powered vacuum
collector for insect sampling.

Received July 23, 1991; accepted October 7, 1991.

J. New York Entomol. Soc. 100(3):498-502, 1992

A UNIQUE AMYCLE NYMPH (HOMOPTERA: FULGORIDAE)
THAT MIMICS JUMPING SPIDERS
(ARANEAE: SALTICIDAE)

Gregory Zolnerowich

Department of Entomology, Texas A&M University,

College Station, Texas 77843

Abstract. — An undescribed Amycle fulgorid planthopper nymph that mimics jumping spiders
when viewed from behind is recorded from Guerrero, Mexico. The nymph has 4 smooth,
polished dark areas on its metathorax and hind wing pads that resemble the anterior eyes of
jumping spiders. The middle and hind legs, which project posteriorly and have flattened femora
and tibiae, were moved in an up and down manner similar to those of salticid legs and pedipalpi.
It is hypothesized that such mimicry affords protection from predation by salticids only, not
predators in general.

Many spiders mimic insects, especially ants (Reiskind, 1970, 1977; Edmunds,
1978; Foelix, 1982; Jackson, 1982), but the converse has only been observed in the
fly family Tephritidae. Monteith (1972) commented on the similarity in appearance
and behavior between the apple maggot Rhagoletis pomonella (Walsh), the cherry
fruit fly R. cinqulata (Loew) (Tephritidae), and the sympatric jumping spider Para-
phidippus marginatus (Walck.) (Salticidae). Eisner’s (1985) report that the tephritid
Zonosemata vittigera (Coquillett) mimicked jumping spiders stimulated a flurry of
experiments dealing with the protection afforded by spider mimicry (Mather and
Roitberg, 1987; Greene et al., 1987; Whitman et ah, 1988).

It has been suggested, but never demonstrated, that other insects also may mimic
spiders. O’Brien (1967) reported the behavior and appearance of male Caliscelis
bone/lii (Latreille) (Homoptera: Issidae) strongly resembled that of jumping spiders.
Santiago-Blay and Maldonado-Capriles (1988) suggested Emesa tenerrima (Dohm)
(Hemiptera: Reduviidae) may mimic the spider Modisimus signatus (Banks) (Phol-
cidae) in Puerto Rico.

I report here a nymph of Amycle sp. (Homoptera: Fulgoridae) which closely re-
sembles a salticid spider when viewed from behind. The single specimen, a fourth
instar nymph, was collected by sweep-netting in a dryland forest 4.5 miles northwest
of El Ocotito, Guerrero, Mexico, on July 7, 1987, by R. A. Wharton. Although seven
entomologists were collecting in the area, only the single specimen was caught.

The mimic has four smooth, polished dark areas on the metathorax which imitate
the anterior eye row of a salticid, even to the point of reflecting highlights (Figs. 1-
3). Furthering the illusion are the middle and hind legs with flattened femora and
tibiae. These project posteriorly and simulate the anterior legs and pedipalpi of a
jumping spider. The specimen, which is 7.2 mm long and 3.9 mm wide, is approx-
imately the same size as the largest salticid collected from the same locality (Fig. 4).
This fulgorid nymph was observed moving its middle and hind legs up and down
in a manner similar to that of a salticid courtship or aggression display. The fulgorid
was killed and preserved in 70 percent ethanol, critical-point dried, and mounted

1992

UNIQUE AMYCLE NYMPH

499

Fig. 1. Posterior view of Amycle sp. spider mimic.

Fig. 2. Posterior view of metathorax and abdomen of Amycle sp. showing smooth areas
that imitate salticid eyes.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Fig. 3. Posterior dorsal view of Amycle sp.

for 35 mm photography. The mimic has been deposited as Voucher Specimen No.
554 in the Texas A&M University Insect Collection. Five earlier instar nymphs of
Amycle sp. were collected screen-sweeping the area. They lack the flattened femora
and tibiae and false eyes of the spider mimic and are not believed to be conspecific
with the mimic.

Jumping spiders were the most common cursorial spider collected by screen-
sweeping the habitat. All spiders collected were approximately the same size or
smaller than the mimic. Salticids present at the collection site were: Habronattus
zebranus F.O.P.C., Habronattus sp., Metaphidippus sp., Nycerella delecta (Peckhams),
Peckhamia sp., and Thiodina sp. Other cursorial spiders collected from the same
habitat were: Strotarchus sp. (Clubionidae), Oxyopes sp. (Oxyopidae), Apollophanes
sp. nr. punctipes F.O.P.C. (Philodromidae), Misumenoides sp., Misumenops sp. nr.
dubius (Keyserling), and Xysticus facetus F.O.P.C. (Thomisidae).

As only the single specimen was collected and observed for a short period, it is
unknown whether the mimic is modelled after a specific salticid species or jumping

1992

UNIQUE AMYCLE NYMPH

501

Fig. 4. Amycle sp. and the salticid Nycerella delecta (Peckhams) collected from the same
locality.

spiders in general. Also unknown is whether such mimicry affords protection from
a wide range of predators or only from salticids.

The most likely case is that the mimicry is directed solely toward jumping spiders.
The behavior of salticids is often stereotyped and mediated by their highly developed
visual system. Once an object is visually perceived, a jumping spider will advance
in a predatory, agonistic, or courtship mode. If the fulgorid is perceived as another
salticid the risk of predation is lessened, since jumping spiders often signal each other
during interspecific and intraspecific encounters (Crane, 1949; Forster, 1982) and
actual cannibalism or combat does not appear to be common (Crane, 1 949).

Previous work supports the hypothesis that this mimicry is designed to deter only
jumping spiders. Greene et al. (1987) found salticid mimicry by the tephritid Z.
vittigera did not deter predation by nonsalticid spiders, mantids, assassin bugs, and
whiptail lizards. Whitman et al. (1988) reported salticid mimicry by the same fly
protected it from a number of sympatric salticid species. Jumping spiders perceived
the flies as other jumping spiders and engaged in agonistic or courtship displays.

Using Vane- Wright’s (1 976) terminology, this case of a fulgorid mimicking salticids
would be Class VI antergic defensive mimicry (Batesian). Here the model and op-
erator (predator) would be the same. Since many species of jumping spiders and
fulgorids occur in similar habitats, this type of mimicry may be more common than
reported.

The only other apparent case of mimicry in fulgorids occurs in the lantemflies

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

( Fulgoria spp.). Hogue (1984) speculated they avoid predation by mimicking several
genera of arboreal lizards.

ACKNOWLEDGMENTS

I am grateful to L. B. O’Brien for identifying the fulgorids and G. B. Edwards and M. L.
Jimenez for identifying spiders. S. W. Wilson provided literature on homopteran nymphs. J.
M. Heraty, L. B. O’Brien, R. A. Wharton, J. B. Woolley, and an anonymous reviewer provided
useful comments. Special thanks goes to M. J. Rose for his advice and the use of his photographic
equipment. This paper is Technical Article No. 26073 from the Texas Agricultural Experiment
Station.

LITERATURE CITED

Crane, J. 1949. Comparative biology of salticid spiders at Rancho Grande, Venezuela. Part
IV. An analysis of display. Zoologica 34:159-214.

Edmunds, M. 1978. On the association between Myrmarachne spp. (Salticidae) and ants.
Bull. Br. Arachnol. Soc. 4:149-160.

Eisner, T. 1985. A fly that mimics jumping spiders. Psyche 92:103-104.

Foelix, R. F. 1982. Biology of Spiders. Harvard University Press, Cambridge, Mass.

Forster, L. 1982. Visual communication in jumping spiders (Salticidae). Pages 161-212 in:
P. Witt and J. Rovner (eds.), Spider Communication: Mechanisms and Ecological Sig-
nificance. Princeton University Press, Princeton, N.J.

Greene, E., L. J. Orsak and D. W. Whitman. 1987. A tephritid fly mimics the territorial
displays of its jumping spider predators. Science 236:310-312.

Hogue, C. L. 1984. Observations on the plant hosts and possible mimicry models of “Lantern
Bugs” ( Fulgora spp.) (Homoptera: Fulgoridae). Rev. Biol. Trop. 32:145-150.

Jackson, R. R. 1982. The biology of ant-like jumping spiders: intraspecific interactions of
Myrmarachne lupata (Araneae; Salticidae). Zool. J. Linn. Soc. 76:293-319.

Mather, M. H. and B. D. Roitberg. 1987. A sheep in wolfs clothing: tephritid flies mimic
spider predators. Science 236:308-310.

Monteith, L. G. 1972. Status of predators of the adult apple maggot, Rhagoletis pomonella
(Diptera: Tephritidae), in Ontario. Can. Entomol. 104:257-262.

O’Brien, L. B. 1967. Caliscelis bonellii (Latreille), a European genus of Issidae new to the
United States (Homoptera: Fulgoroidea). Pan-Pacific Entomologist 43:130-133.

Reiskind, J. 1970. Multiple mimetic forms in an ant-mimicking clubionid spider. Science
169:587-588.

Reiskind, J. 1977. Ant-mimicry ip Panamanian clubionid and salticid spiders (Araneae:
Clubionidae, Salticidae). Biotropica 9:1-8.

Santiago-Blay, J. A. and J. Maldonado-Capriles. 1988. Observations on the true bugs Emesa
tenerrima, a possible spider mimic, and Ghilianella borincana (Hemiptera: Reduviidae:
Emesinae) from Puerto Rico. Proc. Entomol. Soc. Wash. 90:369-372.

Vane-Wright, R. I. 1976. A unified classification of mimetic resemblances. Biol. J. Linn. Soc.
8:25-56.

Whitman, D. W., L. Orsak and E. Greene. 1988. Spider mimicry in fruit flies (Diptera:
Tephritidae): further experiments on the deterrence of jumping spiders (Araneae: Sal-
ticidae) by Zonosemata vittigera (Coquillet). Ann. Entomol. Soc. Am. 81:532-536.

Received 1 February 1991; accepted 21 October 1991.

J. New York Entomol. Soc. 100(3):503-509, 1992

BEMBIDION FEMORATUM STURM AND AMARA COMMUNIS
(PANZER) (COLEOPTERA: CARABIDAE) NEW TO

NORTH AMERICA

Yves Bousquet

Centre for Land and Biological Resources Research, Agriculture Canada,

Ottawa, Ontario K1A 0C6, Canada

Abstract. —The Palaearctic species Bembidion femoratum Sturm and Amara communis (Pan-
zer) are reported for the first time from North America. The first species is known from New
Brunswick and Nova Scotia since 1967 and the second from New Brunswick since 1988. Both
species were probably introduced through nursery stock. A short description and comments
are provided for each species.

From a practical point of view, introduced arthropods are important to mankind.
Some are pests, others beneficial, and several could have an impact on local fauna
by displacing native species or reducing their reproductive success. Early detection
of exotic species has been advocated as a national priority in the U.S.A. (Hoebeke
and Wheeler, 1983) and a computerized data base on immigrant arthropods has been
recently developed (Knutson et al., 1990).

Recent collecting in The Maritimes has yielded two European carabids, Bembidion
femoratum Sturm, 1 825 and Amara communis (Panzer, 1797), previously not known
to occur in North America. The purpose of this paper is to report the occurrence of
these two species on this continent. A short description and comments are provided
for each species to assist in their recognition.

Bembidion femoratum Sturm, 1825

Description. Body black, upper surface with faint metallic lustre; elytra with large
humeral and apical rufous spots on each side, spots not linked laterally; apical spot
usually extending to apex and suture; first antennomere and base of following 2 or
3 antennomeres pale, rufo-testaceous; penultimate maxillary and labial palpomeres
and usually also preceding maxillary palpomere infuscated; femora infuscated, tibiae
clearly paler than femora. Pronotum with sides less rounded than in B. tetracolum\
basal punctures shallow, often confluent into longitudinal wrinkles; microsculpture
weakly impressed laterally, absent on disc. Elytra parallel-sided; strial punctures
smaller than in B. tetracolum; punctures of seventh stria distinctly smaller than those
of sixth stria; microsculpture rather strong, markedly transverse, not forming distinct
meshes. Aedeagus as illustrated (Fig. 1).

Length of body: 4. 3-5.0 mm.

Distribution. This species occurs over most of Europe, from Scandinavia south to
Spain and Italy (including Sicily), in Asia Minor, and in Siberia east to the Lena
River (Lindroth, 1945); it was also reported from Mongolia (Poppius, 1907). In
North America, this Bembidion is known from Cape Breton Island, continental Nova
Scotia, and along the southern coast of New Brunswick. The first specimen collected

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Fig. 1. Median lobe (left lateral view) of Bembidion femoratum (Sydney, Nova Scotia).

in the Nearctic Region was found in Lawrencetown near Halifax, Nova Scotia, on
July 19-20, 1967. The specimen was identified by C. H. Lindroth in 1971.

New Brunswick. St. John Co.: St. John’s, 7. VII. 1988 (Larochelle & Lariviere col-
lection) 4.

Nova Scotia. Halifax Co.: Lawrencetown, 19-20.VII.1967 (Canadian National
Collection) 1. Cape Breton Co.: Sydney, 12.VI.1983 (Canadian National Collec-
tion) 14.

Habitat. According to Lindroth (1985), the species occurs in Europe mainly near
water, notably on river banks, on clay or clay-mixed sand, silt or gravel, often among
sparse vegetation. It is also found, though less frequently, in gravel and clay pits, at
roadsides and in cultivated fields, far from water. All North American specimens,
except the one from Lawrencetown, were collected under rocks and debris on mod-
erately moist, clayey soil in abandoned fields. At Sydney, it was found with several
other European introduced carabids, including B. tetracolum Say.

Wing Condition. The wings are fully developed in this species and flight obser-
vations have been recorded (Lindroth, 1945).

Discussion. This Bembidion belongs to the tetracolum group as defined by Lindroth
(1963:330-33 1). It differs from all other North American species of the group, except
B. petrosum attuense Lindroth, 1963 and B. poppii Netolitzky, 1914, which occur in
the Nearctic Region only in Alaska, by having the penultimate palpomeres, as well
as the second and third antennomeres, infuscated at least apically, and no microsculp-
ture on the disc of the pronotum.

To include this species in Lindroth’s key (1963) to the Canadian and Alaskan
species of Bembidion, the following changes should be made:

151. Pronotum with sides straight or very slightly sinuate toward base . . B. nevadense var.

- Pronotum with sides distinctly sinuate toward base 152

152. Seventh elytral stria with punctures as strong as those of sixth stria on basal half.

[Alaska] 152'

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IMMIGRANT CARABIDAE

505

- Seventh elytral stria with punctures smaller than those of sixth stria on basal half.

[Maritime Provinces] B. femoratum

1 52'. Elytral spots poorly defined. Elytral microsculpture consisting of meshes on average

less than twice as wide as long. Body length 4. 9-5. 9 mm B. petrosum attuense

- Elytral spots well defined. Elytral microsculpture consisting of meshes more than

twice as wide as long. Body length 3. 8^1. 4 mm B. poppii

Bembidion femoratum is morphologically very similar to the European B. andreae
(Fabricius) (cf. Lindroth, 1985) and has been considered as its subspecies by several
authors.

Amara communis (Panzer, 1797)

Description. Body black, upper surface brassy or occasionally greenish or bluish;
first two antennomeres and base of third antennomere pale, yellowish, third anten-
nomere infuscated on apical half; palpomeres, except basal ones, piceous to black;
femora and tarsi rufopiceous to black, tibiae paler than femora. Head narrow with
flat eyes. Pronotum with sides more or less rounded on basal half; anterior angles
markedly, angularly produced; basal punctation variable; basal foveae superficial,
more or less distinct; posterior lateral seta on each side widely separated from lateral
bead. Elytron with striae deepened apically; intervals more convex toward apex;
setigerous punctures along eighth stria interrupted near middle; seventh stria with
three subapical setae; parascutellar seta absent; microsculpture more or less isodia-
metric to slightly transverse, rather weakly impressed in male, well-impressed in
female. Male hind tibia with setal brush on distal half of medial surface. Female with
only two apical setae on last visible sternum. Aedeagus with apical lamella sym-
metrical, tapered; right paramere with marked hook.

Body length: 6.0-7. 5 mm.

Distribution. This species occurs over most of Europe, from Scandinavia south to
Spain and Italy, Asia Minor, the Caucasus, and across Siberia to the Kamchatka
Peninsula (Lindroth, 1945). In North America, A. communis is known from two
localities in southern New Brunswick.

New Brunswick. St. John Co.: West Quaco, 7.VII.1988 (Larochelle & Lariviere
collection) 2. Albert Co.: Waterside, 8. VII. 1 988 (Larochelle & Lariviere collection) 6.

Habitat. According to Lindroth (1986), this eurytopic species is found on almost
every kind of moderately dry soil in meadows, fields and woods, often under moss
and dry leaves. The adults feed on plant seeds. The species is a spring breeder.

Wing Condition. The wings are fully developed and functional.

Discussion. Amara communis can be distinguished from all other North American
species of Amara occurring in the northeast by the combination of having the first
two antennomeres pale with the third one infuscated at apical half, the femora
infuscated, no parascutellar seta, and the seventh elytral stria with three subapical
setae.

To include this species in Lindroth’s key (1966) to the Canadian and Alaskan
species of Amara, the following changes should be made to couplets 51 and 54:

51. Shoulder tooth obsolete. Seventh elytral stria with 2 subapical setae 52

- Shoulder tooth often small but distinct. Seventh elytral stria with 3 subapical setae 53

54. Inner basal foveae of pronotum deep, linear and virtually parallel to median line . .

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

A. aenea

- Inner basal foveae of pronotum shallow, if linear then somewhat oblique 54'

54'. Elytral striae deepened toward apex, intervals more convex apically. Third anten-

nomere infuscated in apical half A. communis

- Elytral striae shallow throughout, intervals quite flat. Third antennomere entirely pale

or almost so 55

This species is closely related to the European A. convexior Stephens. The latter
differs by being more parallel-sided, by the more or less continuous setigerous punc-
tures in the eighth stria and by the somewhat constricted apical lamella of the aedeagus
(Lindroth, 1986).

DISCUSSION

With the addition of the two species herein reported, the fauna of northeastern
North America now comprises 45 exotic carabid taxa (Table 1). In contrast, 21
introduced species are known from western North America, 15 being common to
both sides of the continent (Spence, 1990).

Among the exotic carabids established in eastern North America, all but two were
accidentally introduced. Calosoma sycophanta (Linne) and Carabus auratus Linne
were intentionally introduced from Europe to Massachusetts in 1905-1910 for the
control of the gypsy moth, Lymantria dispar (Linne) (Smith, 1959; Weseloh, 1986).
Several other European species of Carabus and Calosoma were deliberately intro-
duced into eastern North America but only the two above-mentioned species are
established.

Bembidion femoratum and Amara communis are known in North America only
from The Maritimes which probably represent the point of introduction. To be more
specific, St. John, New Brunswick, could well be the port of entry for both species.
Twenty-three species, about 50% of the exotic carabids found in northeastern North
America, were first spotted on this continent in Newfoundland or The Maritimes.
There is no doubt that ports in the Atlantic Provinces of Canada have been and are
still the primary points of introduction of exotic species in northeastern North Amer-
ica. Other prime introduction areas in the northeast include ports along the Saint
Lawrence, particularly Montreal (cf. Blemus discus, Clivina fossor ), the Great Lakes
region (cf. Trechus quadristriatus,' Bembidion obtusum), and along the United States
coast (cf. Asaphidion flavipes, Clivina collaris, Harpalus puncticeps, Harpalus rubri-
pes).

Brown (1940) and Lindroth (1957) claimed that many introduced species in North
America crossed the Atlantic with bulk rock and soil regularly taken aboard sailing
vessels as ballast. As a rule, ballast was dumped ashore on the North American coast.
Because this practice was abandoned after World War I (Lindroth, 1957) and the
introductions of B. femoratum and A. communis are likely of recent origin, the
presence of these two carabids in North America cannot be explained by this mode
of introduction. So how did these species get here? Spence and Spence (1988), Ka-
vanaugh and Erwin (1985), and Kelton (1983) considered it likely that European
carabids and mirids were accidentally introduced to North America through nursery
stock. I have no reason not to believe that the two species herein reported also could
have been introduced with nursery stock.

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IMMIGRANT CARABIDAE

507

Table 1. List of exotic carabid species likely established in northeastern North America.

Year of

Introduced species

Area of first record

first record

Abax parallelepipedus (Piller & Mitterpacher)

Cape Breton, NS

1965

Acupalpus meridianus (Linne)

Quebec area, PQ

1969

Agonum muelleri (Herbst)

Newfoundland

<1840

Amara aenea (DeGeer)

?

<1828

Amara apricaria (Paykull)

?

<1875

Amara aulica (Panzer)

Cape Breton, NS

1929

Amara bifrons (Gyllenhal)

Cape Breton, NS

1929

Amara communis (Panzer)

New Brunswick

1988

Amara eyrinota (Panzer)1

Newfoundland

1971

Amara familiaris (Duftschmid)

Long Island, NY

1915

Amara fulva (Muller)

Newfoundland

1905

Asaphidion flavipes (Linne)

Long Island, NY

1930

Bembidion bruxellense Wesmael

Newfoundland

1907

Bembidion femoratum Sturm

Nova Scotia

1967

Bembidion lampros (Herbst)

Newfoundland

1949

Bembidion obtusum Audinet-Serville

Great Lakes area

1956

Bembidion properans (Stephens)

Nova Scotia

1947

Bembidion stephensii Crotch

Ottawa, ON

1891

Bembidion tetracolum Say

?

<1823

Blemus discus (Fabricius)

Montreal area, PQ

1933

Broscus cephalotes (Linne)

Cape Breton, NS/

1987

Prince Edward Is.

Calosoma sycophanta (Linne)

Massachusetts

1905-10

Carabus auratus Linne

Massachusetts

1908

Carabus g. granulatus Linne

Montreal, PQ

1952

Carabus g. hibernicus Lindroth

New Brunswick

1890

Carabus nemoralis Muller

New Brunswick

1870

Clivina collaris (Herbst)

Massachusetts

<1838

Clivina fossor (Linne)

Montreal area, PQ

1915

Harpalus affinis (Schrank)

?

<1798

Harpalus puncticeps (Stephens)

Long Island, NY

1954

Harpalus rubripes (Duftschmid)

New Hampshire

1981

Harpalus rufibarbis (Fabricius)

Montreal area, PQ

1953

Harpalus rufipes (DeGeer)

Prince Edward Is.

1937

Laemostenus terricola (Herbst)

Nova Scotia

<1894

Leistus ferrugineus (Linne)

Newfoundland

1977

Notiophilus biguttatus (Fabricius)

Newfoundland

1923

Notiophilus palustris (Duftschmid)

Nova Scotia

1967

Paranchus albipes (Fabricius)

Newfoundland

<1840

Perigona nigriceps (Dejean)

?

<1853

Porotachys bisulcatus (Nicolai)

? Massachusetts

<1900

Pterostichus melanarius (Illiger)

Nova Scotia

1926

Pterostichus strenuus (Panzer)

Newfoundland

1937

Stomis pumicatus (Panzer)2

Cape Breton, NS

1984

Trechus quadristriatus (Schrank)

Great Lakes area

1965

Trechus rubens (Fabricius)

Nova Scotia

<1875

1 The name eurynota, used by most authors for this taxon, is an incorrect subsequent spelling.

2 The first North American record of this species from Hemmingford in Quebec was based
on a mislabelled specimen (cf. Bousquet, 1987:124).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

To establish the source area of the two species herein reported is impossible.
Lindroth (1957) argued that most European carabids introduced before World War
I originated from England, particularly the southwestern region, because most of the
transatlantic ballast traffic originated from that country. However, for B. femoratum
and A. communis the best guess seems to be Holland. Both species are common in
that country (Turin et ah, 1977) and Holland has been by far the main source of
nursery stock imported from Europe to Canada for at least the past 20 years (Duguay
and Anderson, 1986).

ACKNOWLEDGMENTS

I thank A. Larochelle and M.-C. Lariviere for making the material in their collection available
to me, and A. Smetana and S. Laplante for critically reviewing the manuscript.

LITERATURE CITED

Bousquet, Y. 1987. The carabid fauna of Canada and Alaska: range extensions, additions and
descriptions of two new species of Dyschirius (Coleoptera: Carabidae). Coleopts. Bull.
41:111-135.

Brown, W. J. 1 940. Notes on the American distribution of some species of Coleoptera common
to the European and North American continents. Can. Entomol. 72:65-78.

Duguay, C. and R. W. Anderson. 1986. The Canadian cut flower industry. Agriculture Canada,
Working Paper 12/86.

Hoebeke, E. R. and A. G. Wheeler, Jr. 1983. Exotic insects reported new to northeastern
United States and eastern Canada since 1970. J. New York Entomol. Soc. 91:193-222.

Kavanaugh, D. H. and T. L. Erwin. 1985. Trechus obtusus Erichson (Coleoptera: Carabidae),
a European ground beetle, on the Pacific Coast of North America: its distribution,
introduction, and spread. Pan-Pac. Entomol. 61:170-179.

Kelton, L. A. 1983. Four European species of Psallus Fieber found in Canada (Heteroptera:
Miridae). Can. Entomol. 115:325-328.

Knutson, L., R. I. Sailer, W. L. Murphy, R. W. Carlson and J. R. Dogger. 1990. Computerized
data base on immigrant arthropods. Ann. Entomol. Soc. Am. 83:1-8.

Lindroth, C. H. 1945. Die Fennoskandischen Carabidae: Eine Tiergeographische Studie. I.
Spezieller Teil. Kungl. Vetensk. Vitterh. Samh. Handlingar, Ser. B, 4:1-709.

Lindroth, C. H. 1 957. The Faunal Connections between Europe and North America. Almqvist
& Wiksell, Stockholm.

Lindroth, C. H. 1963. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and
Alaska. Part 3. Opusc. Entomol. Suppl. 24:201-408.

Lindroth, C. H. 1966. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and
Alaska. Part 4. Opusc. Entomol. Suppl. 29:409-648.

Lindroth, C. H. 1985. The Carabidae (Coleoptera) of Fennoscandia and Denmark. Fauna
Entomologica Scandinavica, vol. 15, part 1. E. J. Brill/Scandinavian Science Press Ltd.,
Leiden.

Lindroth, C. H. 1986. The Carabidae (Coleoptera) of Fennoscandia and Denmark. Fauna
Entomologica Scandinavica, vol. 15, part 2. E. J. Brill/Scandinavian Science Press Ltd,
Leiden.

Panzer, G. W. F. 1797. Faunae Insectorum Germanicae Initia; oder Deutschlands Insecten.
Part 40. Numberg.

Poppius, B. 1907. Beitrage zur Kenntnis der Coleopteren-Fauna von West-Sibirien und der
NW Mongolei. Ofvers. Finska Vet. Soc. Forh. 49 (No. 17): 1-24.

1992

IMMIGRANT CARABIDAE

509

Smith, M. E. 1959. Carabus auratus L. and other carabid beetles introduced as gypsy moth
predators (Coleoptera, Carabidae). Proc. Entomol. Soc. Wash. 61:7-10.

Spence, J. R. 1990. Success of European carabid species in western Canada: preadaptation
for synanthropy? Pages 129-141, in: N. E. Stork (ed.), The Role of Ground Beetles in
Ecological and Environmental Studies. Intercept Ltd., Andover.

Spence, J. R. and D. H. Spence. 1988. Of ground-beetles and men: introduced species and
the synanthropic fauna of western Canada. Mem. Entomol. Soc. Can. 144:151-168.
Sturm, J. 1825. Deutschlands Fauna in Abbildungen nach der Natur mit Beschreibungen. V.

Abtheilung. Die Insecten. Vol. 6. Verfasser, Niimberg.

Turin, H., J. Haeck and R. Hengeveld. 1977. Atlas of the Carabid Beetles of The Netherlands.
North-Holland, Amsterdam.

Weseloh, R. M. 1986. Biological control of gypsy moths: help from a beetle. Front. Plant Sci.
39:2-3.

Received 3 July 1991; accepted 8 October 1991.

J. New York Entomol. Soc. 100(3):5 10-521, 1992

DESCRIPTIONS OF NEW OR POORLY KNOWN SPECIES
OF GASTROSTICTA CASEY, 1918 AND
PARAFERONINA BALL, 1965 (COLEOPTERA:
CARABIDAE: PTEROSTICHUS BONELLI, 1810)

Yves Bousquet

Centre for Land and Biological Resources Research, Agriculture Canada,

Ottawa, Ontario K1A 0C6, Canada

Abstract. — Three new species of Pterostichini from North America are described: Pterostichus
enodis (type locality: Texas), and P. mutoides (type locality: Georgia) which belong to the
subgenus Gastrosticta Casey, 1918, and P. spathifer (type locality: Isabella Creek, Clearwater
Co., Idaho) which is a member of the subgenus Pseudoferonina Ball, 1965. Three species of
Gastrosticta are redescribed: P. obesulus LeConte, 1873; P. punctiventris Chaudoir, 1878; and
P. ophryoderus Chaudoir, 1878. Pterostichus amnicola (Casey, 1918) is listed as a new junior
synonym of P. punctiventris (Chaudoir, 1878). A key to all known species of Gastrosticta is
included.

The purpose of this paper is to provide names for three undescribed species of
Pterostichus Bonelli belonging to the subgenera Gastrosticta Casey and Pseudoferoni-
na Ball and redescribe three species of Gastrosticta previously known only from the
original descriptions. The new species are described herein so that the names could
be used in a forthcoming publication dealing with the supraspecific classification of
the North American Pterostichini. As members of Gastrosticta and Pseudoferonina
are rare in collections, I hope this publication will stimulate anyone interested in
North American Carabidae to collect and study these beetles.

Acronyms used in this publication are as follow: CAS— California Academy of
Sciences, San Francisco, California; CMNH— Carnegie Museum of Natural History,
Pittsburgh, Pennsylvania; CNC— Canadian National Collection, Ottawa, Ontario;
MCZ— Museum of Comparative Zoology, Cambridge, Massachusetts; MNHP— Mu-
seum National d’Histoire Naturelle, Paris; U ASM— University of Alberta Strickland
Museum, Edmonton, Alberta; UICM — Department of Plant, Soil & Entomological
Sciences, University of Idaho, Moscow, Idaho; USNM — United States National Mu-
seum (Natural History), Washington, DC.

Subgenus Gastrosticta Casey, 1918

Gastrosticta Casey, 1918:323. Type species: Feronia ventralis Say, 1823, by original
designation.

Gastrosticta is one of the most poorly known North American subgenera of Ptero-
stichus. It includes ten species: P. ventralis (Say, 1823); P. putus Casey, 1913; P.
sayanus Csiki, 1930 [replacement name for P. obscurus (Say, 1834)]; P. subacutus
(Casey, 1 9 1 8); P. tumesccns LeConte, 1 863; P. obesulus LeConte, 1 873; P. punctiven-
tris (Chaudoir, 1878); P. ophryoderus (Chaudoir, 1878); and the two species described
in this paper. The group is restricted to the temperate regions of eastern United States.

1992

GASTROSTICTA AND PARAFERONINA

511

According to Ball (1960) members of this subgenus live in dry, open, coniferous
woodlands; all species are brachypterous.

Adults of Gastrosticta are distinguished from those of other North American Ptero-
stichini by the laterally punctate sterna in combination with the absence of scutellar
stria, lack of seta on the hind trochanter, glabrous last tarsomere, and poorly devel-
oped metatarsal carina. All these character states are apotypic within the Pterostichini;
likely Gastrosticta is a monophyletic taxon.

Three species of this group, P. obesulus, P. punctiventris, and P. ophryoderus, are
known only from the original descriptions which are difficult to interpret. These
species are redescribed herein along with two new species; a key to the species of the
subgenus is also provided.

Pterostichus obesulus LeConte, 1873
Pterostichus obesulus LeConte, 1873:314. Type locality: Georgia.

Diagnosis. Unique among the species of Gastrosticta by the characteristic shape
of the pronotum (Fig. 2) with the lateral depressions explanate basally.

Description. Habitus (Fig. 1). Coloration. Body rufopiceous to piceous dorsally;
antennae, palpi, legs, lateral margins of pronotum and sometimes first elytral intervals
testaceous to dark ferrugineous; elytra iridescent. Microsculpture. Frons with poorly
impressed, isodiametric meshes. Pronotum with very transverse meshes. Elytra with
linear microsculpture (i.e., meshes linear, narrow, parallel, with few anastomoses).
Head. Frontal impressions shallow, more or less parallel, impunctate. Pronotum (Fig.
2). Widest around middle; sides rounded on posterior half; posterior angles rounded;
lateral bead thickened basally; lateral depression explanate basally; base not or faintly
and partly marginated laterally; laterobasal impression single, short, shallow, punc-
tate; area between laterobasal impression and lateral margin depressed. Elytra. Mod-
erately long (index length elytra/length pronotum = 2.06-2.28); sixth and seventh
striae not impressed, at most suggested; striae punctate, except on apical fourth and
sometimes also on basal fourth; intervals convex; second discal setigerous puncture
on or near second stria. Legs. Apical spur of fore tibia markedly dilated laterally.
Ventral parts. Proepistemum without punctures. Metepistemum punctate. Sternum
VII with punctures medially, punctation as dense as on lateral areas. Aedeagus.
Median lobe with short apex, apical lamella rounded (Fig. 8); right paramere elongate,
markedly curved apically (Fig. 8).

Length of body: 8. 1-9.5 mm.

Type material. LeConte’s collection in MCZ includes a single specimen under the
name of P. obesulus. The specimen, a female, bears the following labels: “Ga/ Horn
Coll H554/ P. obesulus LeC/ Lectotype MCZ 32326/ Designated 1975 R.T. Allen/
UAIC 00015001.” LeConte’s original description was based on a single specimen
since he wrote (1873:314) “I have seen only the specimen kindly communicated by
Dr. Horn.” The specimen therefore is the holotype (by monotypy).

Distribution. The species is known from Georgia, Florida, and Louisiana. In ad-
dition to the holotype, I have seen five specimens of this species from the following
localities. Florida: “Fla”, Liebeck (MCZ) 1; Alachua Co., Gainesville, 29.1.1972, D.
R. Whitehead (UASM) 1; idem, 9-1 7. IV. 1986, G. Gibson (CNC) 1; boundary of

512

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Fig. 1. Pterostichus obesulus LeConte, female

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GASTROSTICTA AND PARAFERONINA

513

5

6

7

Figs. 2-7. Pronotum. 2. Pterostichus obesulus-, 3. P. ophryoderus (lectotype); 4. P. punctiven-
tris (lectotype); 5. P. mutoides (paratype); 6. P. ventralis\ 7. P. putus (lectotype).

Levi-Alachua Co., 14-1 6. IV. 1986, G. Gibson (CNC) 1. Louisiana: “La”, Liebeck
(MCZ) 1.

Comments. LeConte (1873:307) originally included P. obesulus in his section D of
the genus Pterostichus along with P. fallax (Dejean, 1828) and P. sculptus LeConte,
1852. Lindroth (1966:535) did not study any specimens ofP. obesulus and, following
LeConte, included the species in Abacidus LeConte, a group that contains both P.

Figs. 8-14. Median lobe (left lateral view), right paramere, and apical lamella. 8. Pterostichus
obesulus (Gainesville, Florida); 9. P. ophryoderus (5 mi S. Toomsuba, Mississippi); 10. P.
sayanus (Charleston, Illinois); 11. P. enodis (holotype); 12. P. mutoides (holotype); 13. P.
ventra/is (neotype); 14. Apical lamella of median lobe of P. putus (lectotype).

2.5 mm

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GASTROSTICTA AND PARAFERONINA

515

fallax and P. sculptus. The species, however, without doubt belongs to the subgenus
Gastrosticta.

Pterostichus ophryoderus (Chaudoir, 1878)

Feronia ophryodera Chaudoir, 1878:64. Type locality: Mississippi.

Diagnosis. Distinguished from the other Gastrosticta by the characteristic shape
of the pronotum (Fig. 3) with the wide lateral depressions.

Description. Coloration. Body ferrugineous with antennae, palpi, legs, sides of
pronotum and elytra fulvous. Microsculpture. Frons without distinct microsculpture.
Pronotum and elytra with linear microsculpture. Head. Frontal impressions shallow,
more or less parallel, impunctate. Pronotum (Fig. 3). Widest in front of middle; sides
rounded on posterior half, more constricted than in P. obesulus\ posterior angles
rounded; lateral bead regular; lateral depression rather wide, not explanate basally;
base laterally marginate; laterobasal impression shallow, barely punctate; area be-
tween laterobasal impression and margin convex. Elytra. Proportionally long (index
length elytra/length pronotum = 2.54); sixth and seventh striae not impressed, at
most suggested; striae punctate, except on apical fourth and sometimes also on basal
fourth; intervals slightly convex; second discal setigerous puncture situated on or
near third stria. Legs. Apical spur of fore tibia feebly dilated laterally. Ventral parts.
Proepistemum and metepistemum smooth. Sternum VII with few punctures re-
stricted to anterolateral areas. Aedeagus. Median lobe with moderately long apex,
apical lamella rounded (Fig. 9); right paramere elongate, slightly curved apically
(Fig. 9).

Length of body: 9.7-10.2 mm.

Type material. Chaudoir’s collection in MHNP contains a single specimen, a fe-
male, under this name labelled: “ “Mississippi” Salle (handwritten)/Holotype/ Fero-
nia ophryodera Chaud. Holotype (handwritten)/ Museum Paris 1952 Coll R Ober-
thur.” It is obvious that these labels are not the original ones. The specimen, however,
agrees with Chaudoir’s description and I have no doubt that it is part of the type
series. Chaudoir did not mention the number of specimens in the original series and,
since he did not properly label his type material (Ball and Erwin, 1983), the specimen
can not be regarded as the holotype. The specimen is herein selected as lectotype
and the label “Lectotype Feronia ophryodera Chaudoir Des. by Y. Bousquet 1989”
has been attached to it.

Distribution. In addition to the lectotype, I have seen one male and one female of
this species from the following localities. Mississippi: Lauderdale Co., 5 mi S Toom-
suba, U.S. 1 1, 14.VI.1965, R. Freitag & D. Larson (CNC) 1; Rankin Co., Flowood,
28.11.1988, D. & M. Hildebrandt (Hildebrandt Coll.) 1 [cypress swamp on banks of
Pearl River].

Pterostichus punctiventris (Chaudoir, 1878)

Feronia punctiventris Chaudoir, 1878:66. Type locality: Texas.

Gastrosticta amnicola Casey, 1918:372 (New Synonymy). Type locality: St. Louis,
Missouri.

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Diagnosis. Distinguished by the combination of having the sides of the pronotum
obliquely convergent in posterior half, the elytral microsculpture transverse, and the
second discal seta in or close to second stria.

Description. Coloration. Body ferrugineous with antennae and palpi fulvous; tibiae
darker than femora. Microsculpture. Frons with poorly impressed, isodiametric
meshes. Pronotum and elytra with transverse meshes. Head. Eyes less convex than
in P. obesulus. Frontal impressions shallow, more or less parallel, impunctate. Pro-
notum (Fig. 4). Widest in front of middle; sides obliquely convergent on posterior
half; posterior angles obtuse; lateral bead regular; lateral depression not explanate
basally; base marginated laterally; laterobasal impression slightly longer, deeper and
more punctate than in P. obesulus; area between laterobasal impression and margin
convex. Elytra. Proportionally long (index length elytra/length pronotum = 2.42-
2.48); sixth and seventh striae feebly impressed on anterior half, more so on posterior
half; striae punctate, except on apical fourth and sometimes also on basal fourth;
intervals slightly convex; second discal setigerous puncture situated on or near second
stria. Legs. Apical spur of fore tibia feebly dilated laterally. Ventral parts. Proepister-
num impunctate or with few punctures anteriorly. Metepisternum punctate. Sternum
VII smooth medially or with few punctures, punctation if present much sparser than
on lateral areas. Aedeagus. Apical lamella rounded apically much like that of P.
ventralis ( cf Fig. 1 3); right paramere more or less elongate, slightly curved apically
(as in Fig. 12).

Length of body: 9.7-10.2 mm.

Type material. Chaudoir’s collection in MHNP contains a single specimen under
this name, a female labelled: “ “Texas” Schmelb. (handwritten)/ Holotype/ Feronia
punctiventris Chaud. Holotype (handwritten)/ Museum Paris 1 952 Coll R Oberthur.”
As for the preceding species, it is obvious that these labels are not the original ones.
The specimen, however, agrees with Chaudoir’s description and I have no doubt
that it is part of the type series. It is selected herein as lectotype and the label
“Lectotype Feronia punctiventris Chaudoir Des. by Y. Bousquet 1989” has been
attached to it.

Casey’s collection in USNM contains two specimens under the name Gastrosticta
amnicola. One of them, a male labelled “MO./ Gastrosticta amnicola Csy (hand-
written)/ Type USNM 47116/ Casey bequest 1925/ UAIC 00044070/ Lectotype
Gastrosticta aminicola [sic!] by (handwritten) R. T. Allen,” was selected lectotype
by Allen (1977).

Distribution. In addition to the type material, I have seen 2 specimens of this
species in the CNC. Arkansas. Stone Co., Blanchard Springs St. Park, 18.V.1973,
Kavanaugh & Goulet. Missouri. St. Louis Co., Ranken, 4.X.36, E. P. Meiners.

Comments. I have found no external character states to distinguish the type spec-
imens of P. amnicola (Casey) from the lectotype of P. punctiventris (Chaudoir).
Therefore, Gastrosticta amnicola Casey, 1918 is herein considered as a junior syn-
onym of Feronia punctiventris Chaudoir, 1878.

Pterostichus enodis, new species

Diagnosis. Distinguished from other species of Gastrosticta by the rather well
impressed sixth and seventh elytral striae.

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517

Description. Coloration. Body black with antennae, palpi, and legs rufopiceous.
Microsculpture. Frons without distinct microsculpture. Pronotum and elytra with
linear microsculpture. Head. Frontal impressions deeper than on preceding species,
more or less parallel, punctate. Pronotum. Widest at middle; sides rounded on pos-
terior half; posterior angles rounded; lateral depression not explanate basally; later-
obasal impression short, shallow, faintly punctate; area between laterobasal impres-
sion and lateral margin convex. Elytra. Moderately long (index length elytra/length
pronotum = 2.18); sixth and seventh striae impressed, as much as, or nearly as, fifth
stria; striae punctate in basal half; intervals convex; second discal setigerous puncture
on or near second stria. Legs. Apical spur of fore tibia markedly dilated laterally.
Ventral parts. Proepistemum punctate on anterior half; metepistemum impunctate.
Aedeagus. Median lobe with short apex, apical lamella rather rounded (Fig. 1 1 ); right
paramere markedly curved apically (Fig. 1 1 ).

Length of body: 10.8 mm.

Type material. Holotype. Texas: “Tex/ Henry Ulke Beetle Coll. CMNH Acc. No.
1645/ Holotype Pterostichus enodis Bousquet.” The specimen is deposited in the
CMNH.

The specimen, a male, has lost the last 2 left antennomeres, the last 2 left labial
palpomeres, the right maxillary palpus, the tibia and tarsus of the right front, left
middle and right hind legs, and the tarsus of the right middle and left hind legs.

Distribution. The species is known only from the holotype collected in Texas.

Etymology. The specific name is derived from the Latin adjective enodis, e (without
knots, smooth). It refers to the fact that the integument of the species is smooth and
polished, the microsculpture on the pronotum and elytra being of the linear type.

Comments. The shape of the median lobe and particularly the markedly curved
apical half of the right paramere suggest that this species and P. obesulus are probably
sister species.

Pterostichus mutoides, new species

Diagnosis. Separated from the other Gastrosticta by the characteristic shapes of
the pronotum (Fig. 5) and of the median lobe (Fig. 12).

Description. Coloration. Body ferrugineous with antennae, palpi, epipleura, tibiae,
and tarsi flavous. Microsculpture. Frons without distinct microsculpture. Pronotum
with very transverse meshes, meshes poorly impressed and distinct only by spots.
Elytra with transverse meshes. Head. Frontal impressions shallow, more or less
parallel, impunctate. Pronotum (Fig. 5). Widest in front of middle; sides straightly
convergent on posterior half; posterior angles markedly obtuse; lateral bead regular;
lateral depression not explanate basally; laterobasal impression short, shallow, punc-
tate; area between laterobasal impression and lateral margin convex. Elytra. Pro-
portionally long (index length elytra/length pronotum = 2.40); sixth and seventh
striae feebly impressed; striae impunctate; intervals slightly convex; second discal
setigerous puncture on or near second stria. Legs. Apical spur of fore tibia feebly
dilated laterally. Ventral parts. Proepistemum impunctate. Metepistemum with few
punctures. Sternum VII with punctation restricted to anterolateral areas. Aedeagus.
Median lobe with moderately long apex, apical lamella with minute jag (Fig. 12);
right paramere more or less elongate, slightly curved apically (Fig. 1 2).

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Length of body: 9.6-10.0 mm.

Type material. Holotype. Georgia: “Ga./ Holotype Pterostichus mutoides Bous-
quet.” The specimen lacks the tarsus of the right middle leg.

One paratype labelled “Ga/ Paratype Pterostichus mutoides Bousquet.” The ho-
lotype is deposited in the CMNH, the paratype in the CNC.

Distribution. The species is known only from the type material collected in Georgia.

Etymology. The specific name is derived from the specific name mutus and the
suffix oides (like, resembling); it refers to the fact that the adults superficially look
like those of P. mutus, a species of the subgenus Bothriopterus Chaudoir.

KEY TO SPECIES OF GASTROSTICTA

The key is preliminary and is given to facilitate the interpretation of the species
described above.

1 . Frontal impressions deep, grooved, distinctly divergent posteriorly. Laterobasal im-
pressions of pronotum deep, convergent posteriorly 2

- Frontal impressions superficial, not grooved, more or less parallel in most species;
if moderately deep and slightly divergent posteriorly, then laterobasal impressions

of pronotum superficial and more or less parallel 3

2(1). Sides of pronotum rounded on posterior half; posterior angles minutely denticulate

P. subacutus (Casey)

- Sides of pronotum straightly convergent on posterior half; posterior angles obtuse
P. tumescens LeConte

3(1). Second discal puncture of elytron in or close to third stria 4

- Second discal puncture of elytron in or close to second stria 5

4(3). Elytral striae smooth. Microsculpture on elytra slightly transverse. Apical lamella of

median lobe rather long, more or less triangular apically (Fig. 10) .... P. say anus Csiki

- Elytral striae punctate, punctures rather small and sparse but distinct. Microsculpture
on elytra linear. Apical lamella of median lobe rather short, rounded apically (Fig.

9) P. ophryoderus (Chaudoir)

5(3). Sixth and seventh striae clearly impressed in basal half, as much, or nearly so, as

fifth stria P. enodis, new species

- Sixth and especially seventh striae indistinct, or much less impressed in basal half

than fifth stria 6

6(5). Lateral depression of pronotum explanate basally (Fig. 2); base of pronotum not or
faintly and partly marginate laterad of laterobasal impression. Right paramere mark-
edly curved in apical half (Fig. 8) P. obesulus LeConte

Lateral depression of pronotum regular, not explanate basally (Figs. 4-7); base en-
tirely and distinctly marginate laterad of laterobasal impression. Right paramere at
most slightly curved in apical half (Figs. 12-13) 7

7(6). Elytra without microsculpture, except on apical fourth. Last visible stemite with

punctures medially, punctation as dense as that on lateral areas 8

- Elytra with microsculpture all over. Last visible sternite with very few punctures

medially or without punctures, punctation if present much sparser than that on
lateral areas 9

8(7). Pronotum markedly contricted basally (Fig. 6); greatest width at or near level of
anterolateral seta. Apical lamella of median lobe with rounded apex (Fig. 13) ....
P. ventralis (Say)

- Pronotum less constricted basally (Fig. 7); greatest width behind level of anterolateral
seta. Apical lamella of median lobe slightly distorted to right (Fig. 14) . . P. putus Casey

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519

9(7). Pronotum with sides regularly rounded in posterior half (Fig. 5); basal margin ex-
tending medially beyond level of laterobasal impression P. mutoides, new species

- Pronotum with sides rather obliquely convergent in posterior half (Fig. 4); basal

margin not extending medially beyond level of laterobasal impression

P. punctiventris (Chaudoir)

Subgenus Pseudoferonina Ball, 1965

Pseudoferonina Ball, 1965:107. Type species: Pterostichus lanei Van Dyke, 1925, by
original designation.

Members of the subgenus Pseudoferonina are known only from the states of Idaho,
Oregon, and Washington. Six species have been described: P. vexatus Bousquet, 1985;
P. shulli (Hatch, 1949); P. lanei Van Dyke, 1925; P. humidulus (Van Dyke, 1943);
P. smetanai Bousquet, 1985; and P. campbelli Bousquet, 1985. A key to these species
was published recently (Bousquet, 1985); a new species from Idaho is described
herein.

Pterostichus spathifer, new species

Diagnosis. Adults of this species are externally similar to those of P. shulli and can
be differentiated only by the examination of the median lobe of the aedeagus (see
Figs. 15-16). In P. spathifer the median swelling on the left side of the median lobe
is less rounded, the submedian sinuosity is slightly more anterad, and the apex is
wide, spoonlike, and slightly distorted to the right.

Description. Coloration. Dorsal surface of body black; antennae and legs rufous to
rufopiceous; palpi usually rufous. Microsculpture. Frons with isodiametric meshes.
Pronotum and elytra with moderately transverse meshes. Pronotum. Sides oblique
to sinuate in posterior half; anterior angles markedly produced; posterior angles right
to obtuse; outer laterobasal impressions slightly to distinctly impressed; inner latero-
basal impressions punctate. Legs. Mesotibia of male slightly curved apically. Ventral
parts. Sternum VII of male with shallow, medial depression more or less delimited
anteriorly by arcuate carina. Aedeagus (Fig. 1 5). Left side of apical portion of median
lobe (in left lateral view) sinuate submedially; apex of median lobe (in left lateral
view) wide, spoonlike; apical portion of median lobe (in ventral view) with lightly
sclerotized diagonal band.

Length of body: 10.0-10.8 mm.

Type material. Holotype (male). Idaho: “Idaho, Clearwater Co. Isabella Creek,
1,700', S31, T41N, R7E, nr N.Fk. Clearwater River, 17.IX.1984, P. J. Johnson, J.
B. Johnson.” The specimen is deposited in the CAS.

Paratypes. Idaho: same data as holotype (8 males, 9 females), or date 24.VI.1985
(8 males, 8 females). The specimens are deposited in the CNC (10 specimens), USNM
(5), CAS (7), UICM (6) and P.J. Johnson’s collection (5).

Distribution. The species is known only from the type-locality in Idaho.

Habitat. All specimens were collected under alluvial cobbles partly imbedded in
sand and partly covered with moss in a seasonally mostly dry side channel of Isabella
Creek (P. Johnson, pers. comm.).

Etymology. The specific name is derived from the Latin noun spatha, -ae, f (spoon)

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Figs. 1 5-16. Median lobe (left lateral view), right paramere, and apical half of median lobe
(ventral view). 15. P. spathifer (holotype); 16. P. shulli (holotype).

and the verb ferre (to bear). It refers to the spoonlike apex of the median lobe in left
lateral view.

Comments. To include the new species in Bousquet’s key (1985), the following
changes should be made.

3. Median lobe of aedeagus in left lateral aspect strongly sinuate behind middle 3'

- Median lobe of aedeagus in left lateral aspect not sinuate behind middle

» P. lanei Van Dyke

3'. Median lobe of aedeagus with apex wide, spoonlike, and slightly distorted to right in

left lateral aspect (Fig. 15) P. spathifer, new species

Median lobe of aedeagus with apex narrower, not spoonlike, and rather straight in left
lateral aspect (Fig. 16) P. shulli (Hatch)

ACKNOWLEDGMENTS

I thank R. L. Davidson, Carnegie Museum, Pittsburgh; P. J. Johnson, University of Idaho,
Moscow; S. Shaw, Museum of Comparative Zoology, Cambridge; and P. J. Spangler, United
States National Museum of Natural History, Washington D.C. for making the material under
their care available for study. The manuscript has been critically reviewed by A. Smetana and
H. Goulet. The habitus and inking of the drawings were done by G. Sato of the Centre for Land
and Biological Resources Research.

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GASTROSTICTA AND PARAFERONINA

521

LITERATURE CITED

Allen, R. T. 1977. Designation of holotypes and lectotypes for species described by Thomas
L. Casey in the subtribe Pterostichi (Coleoptera: Carabidae: Pterostichini). Coleopts.
Bull. 31:283-286.

Ball, G. E. 1960. Carabidae (Latreille, 1810). Pages 55-181, in: R. H. Arnett Jr. (ed.), The
Beetles of the United States. Catholic University of America Press, Washington, D.C.

Ball, G. E. 1 965. Two new subgenera of Pterostichus Bonelli from western United States, with
notes on characteristics and relationships of the subgenera Paraferonia Casey and Feroni-
na Casey (Coleoptera: Carabidae). Coleopts. Bull. 19:104-1 12.

Ball, G. E. and T. L. Erwin. 1983. The Baron Maximilien de Chaudoir: inheritance, associates,
travels, work, and legacy. Coleopts. Bull. 36[1982]:47 5—50 1 .

Bousquet, Y. 1985. The subgenus Pseudoferonina Ball (Coleoptera: Carabidae: Pterostichus ):
description of three new species with a key to all known species. Pan-Pac. Entomol. 6 1 :
253-260.

Casey, T. L. 1913. Memoirs on the Coleoptera. Vol. IV. The New Era Printing Company,
Lancaster (PA).

Casey, T. L. 1918. Memoirs on the Coleoptera. Vol. VIII. The New Era Printing Company,
Lancaster (PA).

Chaudoir, M. de. 1878. Descriptions de genres nouveaux et d’especes inedites de la famille
des Carabiques. Bull. Soc. Imp. Nat. Moscou 53:1-80.

Csiki, E. 1930. Carabidae: Harpalinae IV (Pars 112). Pages 529-737, in: W. Junk & S.
Schenkling (eds.), Coleopterorum Catalogus. W. Jung, Berlin.

Dejean, P. F. M. A. 1828. Species general des Coleopteres, de la collection de M. le Comte
Dejean. Vol. 3. Mequignon-Marvis, Paris.

Hatch, M. H. 1949. Studies on the Coleoptera of the Pacific Northwest III: Carabidae: Har-
palinae. Bull. Brooklyn Entomol. Society 44:80-88.

LeConte, J. L. 1852. Synopsis of the species of Pterostichus Bon. and allied genera inhabiting
temperate North America. J. Acad. Nat. Sci. Philadelphia (Ser. 2) 2:225-256.

LeConte, J. L. 1 863. New species of North American Coleoptera. Prepared for the Smithsonian
Institution. Part 1. Smiths. Misc. Coll. No. 167:1-86.

LeConte, J. L. 1873. The Pterostichi of the United States. Proc. Acad. Nat. Sci. Philadelphia
[25]:302-320.

Lindroth, C. H. 1966. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and
Alaska. Part 4. Opusc. Entomol. Suppl. 29:409-648.

Say, T. L. 1823. Descriptions of insects of the families of Carabici and Hydrocanthari of
Latreille, inhabiting North America. Trans. Amer. Philos. Soc. (New Ser.) 2 [ 1 825]: 1—
109.

Say, T. L. 1834. Descriptions of new North American insects and observations on some
already described. Trans. Amer. Philos. Soc. (New Ser.) 4:409-470.

Van Dyke, E. C. 1925. New species of Carabidae in the subfamily Harpalinae, chiefly from
western North America. Pan-Pac. Entomol. 2:65-76.

Van Dyke, E. C. 1943. New species and subspecies of North American Carabidae. Pan-Pac.
Entomol. 19:17-30.

Received 3 July 1991; accepted 4 February 1992.

J. New York Entomol. Soc. 100(3):522-526, 1992

A HIGH ANDEAN NEW SPECIES OF TERRA
(LEPIDOPTERA, LYCAENIDAE)

Kurt Johnson

Department of Entomology, American Museum of Natural History,

Central Park West at 79th Street, New York, New York 10024

Abstract. — Terra altilineata, new species, is described from a small spring-fed oasis isolated
in high montane arid terrain (3,500-3,650 m), Jujuy Province, Argentina. Several other un-
described butterflies are endemic to this site.

I recently revised the Neotropical hairstreak butterfly genera Nesiostrymon Clench
and Terra Johnson and Matusik (Theclinae, Eumaeini) (Johnson, 1991a). Both genera
occur throughout the mainland neotropics and in the Greater Antilles and, as revised,
included five and six respective species. Neither genus contained a high Andean
representative, though Nesiostrymon australivaga Johnson was described from low
latitude scrub-steppe in Mendoza Province, Argentina. Subsequently, a new species
of Terra was collected in isolated high montane habitat, Jujuy Province, Argentina,
at altitudes over 2,000 meters higher than any previously known congener. This
distinctive new species is described below, following on the generic diagnosis of
Johnson and Matusik (1988) and Johnson (1991a).

Terra altilineata, new species
Figs. 1A, 2B

Diagnosis. Hindwing under surface with thin lineal black medial band (congeners
exhibit lunulate yellow, or wide lineal red or yellow-brown, medial bands); male
forewing upper surface with narrowly elliptic black androconial “brand” ( sensu Eliot,
1973) contrasting lighter brown apical wing color (lowland Argentina congener T.
cana (Hayward) with black wing apices in both sexes, obscuring dark ovate andro-
conial brand in males). Valvae of male genitalia with distinctive lateral shoulders.

Description. Male. Upper Surface of Wings: ground color of both wings dusted
light silvery blue; forewing with brown apices contrasted by black elliptic brand at
distal end of discal cell; hindwing with thin white line along margin, adjacent hairlike
tails at termini of veins CuAl and CuA2. Under Surface of Wings: ground color of
both wings concolorous gray; forewing with dull gray lineal band across postmedial
area; hindwing with thin, undulate, black band across medial area; limbal area with
small black marginal spot in cell CuAl. Length of forewing: 1 1.0-12.5 mm. Female.
Unknown. Male Genitalia. Figure 2B (congeners, Johnson, 1991a, figs. 59, 69-80,
and as noted below). Terminal tergites typical of genus but brush organs uniquely
absent. Genitalia with vinculum typical of genus but saccus stouter, terminus flat-
tened; aedeagus elongate as in congeners but with heavily sclerotized, basally prong-
like, terminus; valvae shouldered with distinctively elliptic sclerotized elements (Fig.
2, B5y), each bordered inwardly by slight ventral ridge extending caudally beneath
the terminal microtrichia otherwise characteristic of the genus (Fig. 2, A lx; B3x).

1992

HIGH ANDEAN TERRA

523

Fig. 1. Adults of Terra altilineata (A): holotype, upper surface right, under surface left; and
T. cana (B): same aspects, male (Tucuman Prov, Dept. Yerba Buena, Rte. 338, km post 15,
Cumbres San Javier, 700 m, 8 February 1991), AMNH.

Types. Holotype male (forewing 11.0 mm) (Fig. 1A), Quebrada de las Cruces,
Dept. Tilcara, Jujuy Province, Argentina, 3,500 m, 1,130 hr, February 2, 1991, in
boulder-strewn, dry wash some 1 00 m south of small spring-fed oasis (see Remarks);
deposited American Museum of Natural History (AMNH). Paratypes. Same data as
holotype, two males deposited Instituto Miguel Lillo, Tucuman, Argentina.

Distribution. Currently known only from type locality (see Remarks).

Remarks. Habitat. — T. altilineata is currently known from a boulder-strewn dry
wash approximately 100 m south of a long, narrow, spring- fed oasis within “Quebrada
de las Cruces” (found on most Jujuy Province topographic maps about 5 km W of
Huacalera village [2,700 m, see site description 53A, Johnson et al., 1990]). On the
sampling date, extent of oasis vegetation was approximately 2 km by 30 m along
spring-fed rivulets extending from 3,500-3,650 m altitude, km 8-10, of the “Abra
de las Cruces” footpath (known to most local residents and originating from the road
leading south from the east end of Huacalera’s Rio Grande River bridge). The larval
foodplant of T. altilineata probably occurs in the oasis; however, the type specimens
were collected in the adjacent un vegetated wash. Other butterflies were observed
some 1 00-200 m from the oasis, including various Lycaenidae (Theclinae, Polyom-
matinae) and Hesperiidae “mud-puddling” at rivulets up to 100 m north of the dry
arroyo bed. In certain years, small spring- and snow-fed lakes or marshes occur about
5 km north and upland from the oasis (the “Lagunas de las Cruces” of most maps,
4,250 m). Thus, more extensive vegetation apparently occurs at certain times along
the quebrada footpath upland of 3,500 m. In 1991 the “lagunas” were dry; aside
from occasional upland xerophiles of the Pieridae and Satyridae, few butterflies were
observed there. Robert C. Eisele (Jujuy, Argentina, pers. comm.) reports the 1991
sampling at Quebrada de las Cruces as the second known visit there by lepidopterists.
To date, three other undescribed species of butterflies, two Lycaenidae and one
Satyridae (all of uncertain generic assignment) are known only from this isolated
locality. Johnson (1992) placed one new thecline from the oasis in a new genus
( Shapiroana matusikorum). It is the southernmost member of a group occurring at
very high altitudes north to the Sierra Nevada de Santa Marta of Colombia.

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Fig. 2. Male genitalia of Terra altilineata, T. cana and T. andevaga (showing aspects illus-
trated and described in generic revision, Johnson, 1991a). A. T. cana, 1, genitalia (ventral view,
x, terminal microtrichia); 2, valvae enlarged (ventral view). B. T. altilineata, 3, genitalia (ventral
view, x, terminal microtrichia); 4, valvae, lateral view; 5, valvae enlarged (ventral view, y,
distally shouldered element); 6, aedeagys terminus, lateral view; 7, terminal tergite, dorsal view.
C. T. andevaga, valvae enlarged (ventral view). Stipples, areas of generally transparent sclerotiza-
tion.

Congeners and phylogenetic position. —Regional congener Terra cana (Fig. IB) is
restricted to dense wet deciduous forest (700-1,250 m) in Tucuman and Jujuy prov-
inces (Hayward, 1949, 1973; Johnson, 1991a). Contrasting T. altilineata on the
hindwing under surface, T. cana has disjunct lunulate yellow-orange bands, an ad-
jacent orange-bordered discal slash or line in cells RS and/or CuAl, and lunulate
orange marginal spot in cell CuAl. Also, the forewing upper surface of T. cana
exhibits wide blackish apices which obscure the usual distinction of the sexes by the
males’ dark forewing brand (Hayward, 1949, 1973; Johnson, 1991a, figs. 45, 47).
The forewing brand in T. altilineata appears thinly elliptic and black against much

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HIGH ANDEAN TERRA

525

lighter brown wing apices, a condition typical of many other eumaeines (like taxa of
species rich Strymon Hiibner [Johnson et al., 1991]) but less like members of Terra,
Nesiostrymon or their six immediate outgroup genera (Johnson, 1991b) with their
mostly diffusive brands. Male genitalia in T. cana have brush organs attached along
the vincular dorsum and, contrasting T. altilineata, exhibit an elongate saccus and
terminally blunted aedeagus. Valvae of T. cana (Fig. 2A) are elliptic and of generally
even contour, lacking any prominently shouldered elements. Valvae of T. altilineata
only slightly resemble those of T. andevaga Johnson (TL submacrothermic rain forest,
Pinchincha Dept., Ecuador, 1,325 m) which exhibit a robust, rather square, base
contrasting short caudal extensions (Fig. 2C; Johnson, 1991a, fig. 74).

Considering cladograms of Terra and Nesiostrymon (Johnson, 1991b, fig. 1, 2,
tables 1-4), the phylogenetic position of T. altilineata is obscured by the prepon-
derance of female structural characters used in constructing the Terra ingroup clado-
gram. Geographic distributions of Terra species are greatly disjunct and interspecific
differences in male genitalia (particularly valvae) do not reflect the intrageneric group-
ings suggested by wing stripe coloration and female genitalia. If, however, one assumes
T. altilineata is part of the clade of Terra species restricted to South America (Johnson,
1991b, fig. 2, clade F), the following observations are consistent with character po-
larities recently attributed to the two genera.

For T. altilineata : (1) thin black under surface bands are a retained primitive
character (consistent with the condition in Nesiostrymon and the reduced bands of
the plesiotypic Antillean endemic Terra Hispaniola Johnson and Matusik); (2) widely
shouldered valvae are most likely autapomorphic (non-homologous with distally
lobate, ventrally multiplanar, conditions characterizing valvae of Nesiostrymon since,
as typical of all Terra species, the valval ventrum in T. altilineata is of even contour
[Fig. 2, B4]).

Etymology. The name combines the Latin roots for high (altus) and line (linea),
referring to the high montane habitat and lineal under surface bands typifying the
species.

ACKNOWLEDGMENTS

Robert C. and Barbara Eisele hosted the eleven member 1991 AMNH Argentine collecting
expedition and chose Quebrada de las Cruces as one of the collecting sites. David and Kathy
Matusik, Nick and Jake Pritzker, David and Karl Kroenlein, David and Seth Grae, and Steve
Grossman also accompanied the author in the field. F. H. Rindge (AMNH) administered the
AMNH Theclid Research Fund in support of this work.

LITERATURE CITED

Eliot, J. N. 1973. The higher classification of the Lycaenidae (Lepidoptera) a tentative ar-
rangement. Bull. Brit. Mus. Nat. Hist. 28:373-505.

Hayward, K. J. 1949. Nuevas especies del Lycaenidae de la Argentina. (Lep. Rhop.). Acta
Zool., Lilloana, Tucuman 8:567-581.

Hayward, K. J. 1973. Catalogo de los Ropaloceros Argentinos. Opera Lilloana 23:1-318.
Johnson, K. 1991a. Cladistics and the biogeography of two trans-Caribbean hairstreak but-
terfly genera: Nesiostrymon and Terra (Lepidoptera, Lycaenidae). Am. Mus. Novitates
3011:1-43.

Johnson, K. 1991b. Neotropical hairstreak butterflies: genera of the “Calycopis/Calystryma

526

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Grade” of Eumaeini (Lepidoptera, Lycaenidae, Theclinae). Rept. Mus. Nat. Hist. Univ.
Wisconsin (Stevens Point) 21:1-127.

Johnson, K. 1992. Genera and species of the Neotropical “elfin”-like hairstreak butterflies
(Lepidoptera, Lycaenidae, Theclinae). Rept. Mus. Nat. Hist., Univ. Wisconsin (Stevens
Point) 22:[vol. 1] 1-135; [vol. 2] 135-270.

Johnson, K., R. Eisele and B. MacPherson. 1990. The “hairstreak butterflies” (Lycaenidae,
Theclinae) of northwestern Argentina. II. Strymon, sensu stricto. Bull. Allyn Mus. 130:
1-77.

Johnson, K. and D. Matusik. 1988. Five new species and one new subspecies of butterflies
from the Sierra de Baoruco of Hispaniola. Ann. Carnegie Mus. 57:221-254.

Received 16 July 1991; accepted 22 August 1991.

J. New York Entomol. Soc. 100(3):527-532, 1992

OVIPOSITIONAL BEHAVIOR AND LARVAL DEVELOPMENT
OF APHIDENCYRTUS APHIDIVORUS
(HYMENOPTERA: ENCYRTIDAE),

AN APHID HYPERPARASITOID

Michael J. Kanuck1 and Daniel J. Sullivan2

department of Biology, Bronx Community College of CUNY,
University Avenue and West 181 Street, Bronx, New York 10453;

2 Department of Biological Sciences, Fordham University,

Bronx, New York 10458

Abstract.— In most hyperparasitoids, the female wasp attacks the primary parasitoid host
either while the parasitized aphid is still alive or after the mummy is formed. Aphidencyrtus
aphidivorus (Mayr) had been known to have a “dual” ovipositional behavior. Whether the adult
female wasp attacked the primary parasitoid host through a live parasitized aphid or a mum-
mified aphid, her egg was always deposited inside the host larva and not on the surface. The
egg hatched inside the host larva, whereupon the hyperparsitoid larva first fed endophagously,
and then ectophagously until the primary host larva was consumed. In “choice” experiments,
82% of the female A. aphidivorus wasps preferred to attack the primary parasitoid larva through
the mummy, while only 1 8% chose the live parasitized aphids for oviposition.

Aphid hyperparasitoids or secondary parasitoids can be divided into two categories
based on their combined adult ovipositional and larval feeding behavior (Sullivan,
1986, 1987, 1988): (1) endophagous species in the families Charipidae ( Alloxysta ,
Phaenoglyphis, Lytoxysta ) and Eulophidae ( Tetrastichus ) in which the female wasp
deposits her egg inside the primary parasitoid larva while it is still developing inside
the live aphid (before the aphid is mummified); the egg does not hatch until the
mummy is formed, at which time the hyperparasitic larva feeds internally on the
primary parasitoid larval host, and (2) ectophagous species in the families Pteromali-
dae {Asaphes, Pachyneuron, Coruna ) and Megaspilidae (Dendrocerus). In these hy-
perparasitoids, the female wasp deposits her egg on the surface of the primary par-
asitoid larva after the aphid is killed and mummified. The hyperparasitic larva then
feeds externally on the primary parasitoid larval host while both are still within the
mummy.

Earlier studies on the biology of Aphidencyrtus aphidivorus (Mayr) (Encyrtidae) by
Silvestri (1909), Griswold (1929), and Maple (1947) have documented “dual” ovi-
positional behavior in this species. Matteson (1977) reported that female Aphiden-
cyrtus aphidivorus attack and oviposit into both live, parasitized aphids and mum-
mified aphids. Details of this dual ovipositional behavior as well as the endo- and
ectophagous development of the larva are described. In addition, the present labo-
ratory study includes results of “choice” experiments by female A. aphidivorus be-
tween a primary parasitoid larval host in a live aphid or in a mummy, which might
indicate preferential ovipositional behavior.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

MATERIALS AND METHODS

Host plant and aphid. — The broad bean, Vicia faba Linnaeus (Windsor variety),
was used to rear the pea aphid, Acyrthosiphon pisum (Harris) (Homoptera: Aphid-
idae), according to the method described by Bennett and Sullivan (1978). Plants and
aphids were kept in a Percival bioclimatic chamber. The daytime regime had a
photoperiod of 16 hr, a temperature of 21.1 ± 0.6°C and 75 ± 5% RH. At night the
photoperiod was 8 hr of darkness, at 15.5 ± 0.6°C and 85 ± 5% RH. This bioclimatic
chamber and the same environmental conditions were also used for separate, caged
rearings of the primary parasitoids, hyperparasitoids, and for the “choice” experi-
ments.

Primary parasitoid and hyperparasitoid. —Aphidius smithi Sharma and Subba Rao
(Hymenoptera: Aphidiidae) was the primary parasitoid. Our colony originated from
the Division of Biological Control Gill Tract (Albany), of the University of California
at Berkeley. The secondary parasitoid or hyperparasitoid was Aphidencyrtus aphi-
divorus (Mayr) (Hymenoptera: Encyrtidae), which was reared from pea aphid mum-
mies collected on alfalfa in Lafayette (Sussex County), New Jersey. In preparation
for the photographic studies on egg deposition and larval development, we dissected
a minimum of 30 live parasitized aphids and 30 mummified aphids.

“Choice” experiments. — After mating, 3 female A. smithi were introduced into a
stinging-tube (glass cylinder 15 cm long and 3.3 cm in diameter and covered at both
ends with a fine mesh organdy cloth, secured by rubber bands) containing a cut broad
bean plant stem and 10-15 pea aphids always in the 4th instar. The stinging-tube
was kept in the bioclimatic chamber for a 6 h ovipositional period. At the end of
that time, the primary parasitoids were removed, and the aphids placed on growing
bean sprouts and returned to the bioclimatic chamber. After 8 d, the aphid was killed
by the primary parasitoid larva which had been feeding internally, and a mummy
was formed, followed by pupation.

In the “choice” experiments, however, we started by first using one of the live,
but parasitized, pea aphids that had been attacked by A. smithi 7 d earlier in a
stinging-tube. At this stage of development, the parasitoid would be a 4th instar
larva. This parasitized aphid was paired with a dead or mummified aphid (containing
a 9-day old A. smithi prepupa or early pupa). Both were placed on a segment of
broad bean leaf or stem in a Petri dish. One mated female A. aphidivorus, that had
not been given the opportunity to oviposit previously, was put into the Petri dish
equidistant between the two potential hosts (live parasitized aphid and mummy) and
permitted to oviposit. The behavior of A. aphidivorus was observed continuously
during a period of 2 hr maximum. Once the ovipositor was withdrawn from either
one of the two potential hosts, the female A. aphidivorus was removed from the Petri
dish. This might be much less than the 2 hr oviposition period, for in no case was
the female permitted to oviposit into the 2nd host, nor remain beyond this maximum
time limit even if she had not oviposited at all.

Once the hyperparasitoid was removed from the Petri dish, the mummy was placed
in a gelatin capsule to await emergence of an adult wasp. The live aphid, however,
was permitted to continue feeding on the broad bean for 48 hr or until it was killed
and mummified by the A. smithi parasitoid larva inside. If a live aphid was not really
parasitized and killed, then this particular paired replicate was discarded from the

1992

APHIDENCYR TVS APHID1 VOR US

529

experiment. This newly formed mummy was also placed into a separate gelatin
capsule. If the primary parasitoid, A. smithi, emerged (approximately 4 d after mum-
my formation), then oviposition by A. aphidivorus would not have occurred, or at
least hyperparasitism would not have been successful. If on the other hand, the
hyperparasitoid, A. aphidivorus, emerged (about 21-23 d after the attack), then this
indicated effective hyperparasitism of whatever primary host was chosen: either in
a live aphid or in a mummy. A total of 1 7 5 paired replicates was used in these choice
experiments that resulted in 175 adult emergences of A aphidivorus.

RESULTS AND DISCUSSION

Oviposition into a live parasitized aphid.— After initially antennating the live par-
asitized aphid (usually for less than a minute), the female A. aphidivorus mounted
the dorsum, always facing caudally (Fig. 1). With her antennae pointed downward,
motionless and touching the aphid, the female inserted her ovipositor through the
integument of the aphid, pushing it into the hemocoel. With up and down thrusts,
she probed for the larva of the primary parasitoid, and later dissections showed that
the egg is laid internally in the A. smithi larva. After withdrawing her ovipositor, the
A. aphidivorus sometimes host fed on the fluid that may ooze from the puncture hole
of the aphid. The process of oviposition lasted 3-5 minutes.

Oviposition into a mummy. —This is similar to the behavior observed when at-
tacking a live parasitized aphid. There was the usual antennating by the female, and
she eventually climbed onto the dorsal surface of the mummified aphid. As in the
behavior described when attacking a live parasitized aphid, the antennae were pointed
downward and remained motionless, with the female facing caudally (Fig. 2). She
then drilled a hole through the mummy wall with her ovipositor (Fig. 3), moving
her body rapidly up and down. The ovipositor was pushed into the mummy, and as
later dissections have shown, the egg was again laid inside the larva of A. smithi.
This is unusual among hyperparasitoids that attack a primary parasitoid larva within
a mummy, because the egg is normally deposited on the surface of the host in the
mummy (Sullivan, 1986, 1987, 1988). After oviposition, the female withdraws her
ovipositor and sometimes host feeds at the drill hole. However, a “feeding tube” is
not formed as has been described for the host feeding behavior at the mummy by
Asaphes lucens (Keller and Sullivan, 1976). The duration of oviposition into the
mummy took 4-8 minutes, somewhat longer than when ovipositing into a live
parasitized aphid. This was probably because of the need to penetrate the hard
mummy wall.

Developmental stages of Aphidencyrtus aphidivorus.— As mentioned earlier, the
female always placed the egg inside the primary parasitoid larva, never on the surface,
regardless of whether the host was attacked in a live aphid or in an aphid mummy.
When a 9-day old A. smithi was attacked through the mummy, this host was a
prepupa or early pupa. In this case, A. aphidivorus required approximately 21 days
to develop from egg deposition to adult emergence. However, if A smithi was attacked
while the aphid was still alive, the host was a 4th instar larva and the hatching of
the egg was delayed 24-48 hr until the primary parasitoid larva killed the aphid and
formed the mummy. This time difference was probably due to the developmental
stage of the A. smithi host in relation to the aphid being alive or mummified, so that

530

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

1992

APHIDENCYR TUS APHID I VOR US

531

the adult emergence of A. aphidivorus occurred 22-23 days after oviposition when
a live parasitized aphid was attacked.

While still within the aphid mummy, the 1 st and 2nd larval instars of A aphidivorus
fed endophagously within the primary parasitoid host, but the 3rd instar larva ate
through the exoskeleton of the A. smithi larva and emerged from the host (Figs. 4-

5) . This occurs 9 d after egg deposition, and for the next 48 h, the A. aphidivorus
larva continued to feed ectophagously until the primary parasitoid host was con-
sumed.

When feeding ceased, the hyperparasitoid larva voided its meconium in the form
of 35-40 small, sticky, yellow-orange, spherical pellets and became a prepupa (Fig.

6) . These meconium pellets were easily distinguishable from the sausage-shaped,
black meconium of A. smithi. After 48 hr, the pupa (Fig. 7) is recognizable and lasts
7-9 d, until the adult cuts its way out of the mummy by making an uneven, saw-
edged exit hole (Fig. 8) in the aphid mummy, usually through the posterior dorsal
surface. Then the adult A. aphidivorus pushed its head through the hole and pulled
itself out of the mummy. This emergence occurs about 21-23 d after the original egg
deposition; males usually emerged before females.

Choice experiments. — 82% (N = 287) of the A. aphidivorus females oviposited into
mummies, while only 18% (N = 63) chose the live parasitized aphids for oviposition.

When describing host selection by A. aphidivorus, Matteson (1977) reported that
“Of the 60 hosts accepted for probing, 33 were mummies and 27 were live aphids.
A total of 42 eggs were laid, 22 in T. pallidus larvae inside live aphids. These figures
indicate no preference between hosts in live aphids and hosts in mummies for probing
or oviposition.” Trioxys pallidus (Haliday) is a primary parasite of the walnut aphid,
Chromaphis juglandicola (Kaltenbach). Our data, on the other hand, are based on a
different primary parasitoid and a different aphid species; and show a parametric
statistically significant preference (P = <0.01) by A. aphidivorus for hyperparasitism
of the primary parasitoid larval host in mummies (82%) over that in live aphids
(18%). Perhaps this preference is elicited by some factor or combination of factors,
such as visual and/or chemical stimuli, but this important aspect is beyond the scope
of the present study.

One might speculate about the evolution of this “dual” ovipositional behavior of
A. aphidivorus. Data from related species are needed to clarify whether this hyper-
parasitoid primitively exhibited ovipositional behavior whereby it attacked the pri-
mary parasitoid larva in a live aphid, or whether it primitively waited for the host
larva to kill the aphid, form a mummy, and only then oviposit into it. At present,
it is not clear which of these two types of behavior is “primitive” or “advanced.”

Figs. 1-8. 1. Aphidencyrtus aphidivorus ovipositing into live parasitized aphid (x 10); 2. A.

aphidivorus ovipositing into mummified aphid ( x 6.4); 3. A. aphidivorus with exserted ovipositor
(a) drilling into mummified aphid (x 13); 4-5. Nine-day old 3rd instar A. aphidivorus larva (a)
emerging from Aphidius smithi larva (b) and beginning to feed as an ectoparasitoid ( x 1 3, x 32);
6. Ventral view of 1 1-day old A. aphidivorus prepupa (a) voiding meconium at caudal end (b)
(x 13); 7. Ventral view of 16-day old A. aphidivorus pupa with meconium (x 19); 8. Saw-edged
exit hole of (a) A. aphidivorus located on posterodorsal side of mummified aphid (x 13).

532

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

Also unusual about this hyperparasitoid is that unlike other genera of secondary
parasitoids that attack the primary parasitoid through the mummy and deposit the
egg externally on the surface of the host, A. aphidivorus oviposits inside the primary
host within the mummy, thus behaving more like the endophagous hyperparasitoids
that attack live parasitized aphids.

LITERATURE CITED

Bennett, A. W. and D. J. Sullivan. 1978. Defensive behavior against tertiary parasitism by
the larva of Dendrocerus carpenteri, an aphid hyperparasitoid. J. New York Entomol.
Soc. 86:153-160.

Griswold, G. H. 1929. On the bionomics of a primary parasite and two hyperparasites of the
geranium aphid. Ann. Entomol. Soc. Am. 22:438-457.

Keller, L. J. and D. J. Sullivan. 1976. Oviposition behavior and host feeding of Asaphes
lucens, an aphid hyperparasitoid (Hymenoptera: Pteromalidae). J. New York Entomol.
Soc. 84:206-211.

Maple, J. D. 1947. The eggs and first instar larvae of Encyrtidae and their morphological
adaptations for respiration. Univ. Calif. Publ. Ent. 8:25-122.

Matteson, P. C. 1977. A study of aphid parasite complexes, with special reference to the
hyperparasite Aphidencyrtus aphidivorus (Mayr), (Hymenoptera: Encyrtidae) Ph.D. dis-
sertation, University of California, Berkeley, 1 12 pp.

Silvestri, F. 1909. Contribuzioni alia conoscenza biologica delgi imenotteri parassiti. Boll.

Lab. Zool. Gen. Agraria R. Scuola Superior Agr. Portici 3:54-71.

Sullivan, D. J. 1986. Aphid hyperparasites: taxonomy and ovipositional behavior. Pages 5 1 1-
518, in: I. Hodek (ed.), Ecology of Aphidophaga. Academica, Prague.

Sullivan, D. J. 1987. Insect hyperparasitism. Ann. Rev. Entomol. 32:49-70.

Sullivan, D. J. 1988. Aphid hyperparasites. Pages 189-203, in: P. Harrewijn and A. K. Minks
(eds.), Aphids, Their Biology, Natural Enemies and Control, Vol. 2B. Elsevier Science
Publ., Amsterdam.

Received 17 June 1991; accepted 9 December 1991.

NOTES AND COMMENTS

J. New York Entomol. Soc. 1 00(3):533— 534, 1992

NEW DISTRIBUTIONAL RECORDS FOR SEMIAQUATIC
HETEROPTERA, INCLUDING A MICROVELIA
NEW TO THE UNITED STATES (HETEROPTERA:
HYDROMETRIDAE, SALDIDAE, VELIIDAE)

During a recent collecting trip to Arizona I collected a single female of Microvelia
rufescens Champion along Bear Creek, about 6.5 miles west of the western edge of
Coronado National Monument (Cochise Co., CL 2543, 2-IV-1991). This is a north-
western extension of the range of this species and the first record for the United
States. The species occurs as far south as Guatemala and Costa Rica, and is widespread
in central Mexico but rare in northern Mexico. The previous northernmost record
was in northern Chihuahua (Smith, C. L. 1980, A taxonomic revision of the genus
Microvelia Westwood [Heteroptera: Veliidae] of America including Mexico, unpub-
lished dissertation, University of Georgia, Athens).

The following records extend the ranges of the listed species into states or provinces
of North America not recorded in the recent catalog of North American Heteroptera
(Henry, T. J. and R. C. Froeschner, Eds., 1988, Catalog of the Heteroptera, or True
Bugs of Canada and the Continental United States, E. J. Brill, Leiden & New York).
They are given so that they will be available before the next revision of that work.
Collection abbreviations are given in the acknowledgments.

Hydrometridae

Hydrometra martini Kirkaldy. Colorado, Yuma Co., Hale Ponds, 6 mi NE of Hale,
10-VII-1987, J. T. Polhemus (JTPC).

Saldidae

Micracanthia bergrothi (Jakovlev). Quebec, R. Port-Daniel (Bona.), #17, 20- VIII-
1975, L. LeSage (CNCI).

Pentacora sphacelata (Uhler). Quebec, Becancour (Nicolet), #797, 16-VI-1979, L.
LeSage (CNCI).

Saldula opacula (Zetterstedt). Ontario, Elmira, Salem Creek, #T4, 14-VIII-1977
and #T5, 7-VII-1977, L. LeSage (CNCI).

Saldula orbiculata (Uhler). Ontario, Elmira, Salem Creek, #T10, 20-VIII-1977, L.
LeSage (CNCI).

Teloleuca pellucens (Fabricius). Quebec, Becancour (Nicolet), #797, 16-VI-1979,
L. LeSage (CNCI).

Veliidae

Microvelia austrina Torre Bueno. Kentucky, Fayette Co., Lexington, S. Travers
(JTPC).

534

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(3)

— J. T. Polhemus, University of Colorado Museum, 3115 S. York, Englewood, Col-
orado 80110.

Acknowledgment. For the opportunity to study a small collection of Saldidae that
he collected, I am indebted to Laurent LeSage, Biosystematics Research Institute,
Ottawa (CNCI). Material held in the Polhemus collection is coded (JTPC).

Received 31 July 1991; accepted 9 December 1991.

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Journal of the

New York Entomological Society

VOLUME 100 JULY 1992 NO. 3

CONTENTS

New synonyms of Frankliniella bondan and a review of the synonyms of F.
cephalica (Thysanoptera: Thripidae) Sueo Nakahara

The first records of the families Ochteridae and Hebridae (Heteroptera) from the
granitic Seychelles, with descriptions of two new species Dan A. Polhemus

A review of the species of Anthocoridae (Hemiptera: Heteroptera) found on Pinus
contort a J. D. Lattin and N. L. Stanton

Seasonal occurrence of the Miridae (Heteroptera) associated with Ohio Buckeye,
Aesculus glabra Willd., in Missouri R. L. Blinn

Priapismus pini, a new species of Discocephalini from Honduras on pine (He-
miptera: Pentatomidae: Discocephalinae) L. H. Rolston

Life history and descriptions of the immature stages of the planthopper Prokelisia
crocea (Van Duzee) (Homoptera: Delphacidae) Monte W. Holder and

Stephen W. Wilson

A unique Amycle nymph (Homoptera: Fulgoridae) that mimics jumping spiders
(Araneae: Salticidae) Gregory Zolnerowich

Bembidion femoratum Sturm and Amara communis (Panzer) (Coleoptera:
Carabidae) new to North America Yves Bousquet

Descriptions of new or poorly known species of Gastrosticta Casey, 1918 and
Paraferonina Ball, 1965 (Coleoptera: Carabidae: Pterostichus Bonelli, 1810)

Yves Bousquet

A high Andean new species of Terra (Lepidoptera, Lycaenidae) Kurt Johnson

Ovipositional behavior and larval development of Aphidencyrtus aphidivorus (Hy-
menoptera: Encyrtidae), an aphid hyperparasitoid Michael J. Kanuck and

Daniel J. Sullivan

Notes and Comments

New distributional records for semiaquatic Heteroptera, including a Microvelia
new to the United States (Heteroptera: Hydrometridae, Saldidae, Veliidae)

J. T. Polhemus

415-417

418-423

424-479

480-487

488-490

491-497

498-502

503-509

510-521

522-526

527-532

533-534

iooa 5

0 OCTOBER 1992 No. 4

Journal

of the

New York

Entomological Society

(ISSN 0028-7199)

Devoted to Entomology in General

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY

Editor: James K. Liebherr, Department of Entomology, Comstock Hall,

Cornell University. Ithaca, New York 14853
Assistant Editor: E. Richard Hoebeke, Department of Entomology, Com-

stock Hall, Cornell University, Ithaca, New York 14853
Rook Review Editor: James S. Miller, Department of Entomology, Amer-

ican Museum of Natural History, Central Park West at 79th Street,
New York, New York 10024

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J. New York Entomol. Soc. 100(4):535-539, 1992

A FOSSIL EMPICORIS WOLFF
(REDUVIIDAE: HETEROPTERA) FROM MEXICAN AMBER
WITH REMARKS ON THE PHYLOGENETIC STATUS OF
THE FOSSIL GENUS ALUMEDA POPOV

Donald B. Thomas1

USDA-ARS, Apartado Postal 544, Tuxtla Gutierrez, Chiapas, Mexico;
present address ARS Subtropical Agriculture Research Laboratory,

2301 S. International Blvd., Weslaco, Texas 78596

Abstract.— A fossil emesine reduviid (Heteroptera) is described from the Oligo-Miocene
amber from the Simojovel formation of Chiapas, Mexico. The single specimen is described as
a new species, Empicoris electricus. The characteristics of the fossil genus Alumeda are found
to link the extant genera Empicoris and Ctydinna.

Fossil Emesine reduviids are known from both European and New World deposits.
A specimen assigned to the genus Ploiaria Scopoli is reported from rock at Aix,
France (Scudder, 1890). Bachofen-Echt (1949) illustrated an emesine nymph which
was not assigned to genus, from the Baltic amber, and Schlee (1980) published a
photograph of an emesine embedded in Dominican amber. Wygodzinsky (1966)
mentions having seen a specimen of Empicoris Wolff in the Chiapas amber but
offered no further elaboration. The most comprehensive reports on fossil emesines
are those of Popov ( 1 987a, b, 1989) who studied material from the Dominican amber.
These included one species of Malacopus Stal, three species of an extinct genus
Alumeda Popov, and a species of Empicoris in subfossil Dominican copal.

Wygodzinsky’s (1966) report of Empicoris in Mexican amber is now confirmed,
although his specimen was not available to me. The following description is based
on a specimen purchased in Tuxtla Gutierrez, Chiapas, with a stated provenance of
Simojovel, Chiapas, Mexico. The Simojovel formation, from which amber is mined,
straddles the Oligo-Miocene boundary (Frost and Langenheim, 1974) giving an age
of ca. 23 million years.

Empicoris electricus, new species

Description. Narrow, elongate, small; length 3.8 mm. Thorax appearing unusually
bilaterally compressed. Wings present, membranous with pattern of spots. Antennae
and legs with numerous dark annuli; protarsi two-segmented.

Flead constricted behind eyes; posterior lobe slightly larger than anterior lobe; eyes
and antennal tubercle large. Rostrum without spines or stiff setae; segment II ap-
pearing distinctly swollen. Antennal segmental ratios I-IV: 0.09:1.00:0.40:0.15.

Pronotum with complete lateral carinae as well as pair of mesial carinae on anterior
dorsum. Humerus with small, spinous tooth at dorsal terminus of each lateral carina.
Posterior dorsal border of pronotum without medial tubercle or projection. Scutellum
with non-spinous projection; mesonotum with small, median tubercle; metanotum
and first abdominal segment each with prominent erect spine (Fig. 2).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Figs. 1-2. Empicoris electricus, n. sp. 1. Hemelytron with venation: M+Cu = Medi-
al + Cubital vein, Sc = Subcostal vein, R = Radial vein, M = Medial vein, PCu + A, = Postcu-
bital + Anal vein one, Cu = Cubital vein, DsC = Discal Cell. 2. Head and thorax, lateral view
(legs, wings and antennae not shown): Mes. Car. = mesial carinae (=dorsal carinae or submedial
carinae), Lat. Car. = lateral carinae. Hum. Sp. = humeral spine, Scut. = scutellum, Ms
mesonotum, Mt = metanotum, Ab, = first abdominal tergite, Cx = coxae. All figures drawn
with Camera Lucida.

Hemelytron with pterostigma far from apex: distance from insertion of M vein on
pterostigma to tip of pterostigma about half distance from latter to apex of wing.
M + Cu vein fused to form single stem emitting from base of discal cell. Discal cell
without smaller closed cell at its base; basal angle truncate; separated from costal
margin by intervening membrane. R vein connected to margin of wing by two veinlets
(sc-r crossveins). Cu vein bent in apical portion oblique to wing axis. False or spurious
submarginal vein present along posterior and apical portion of wing (Fig. 1).

Profemur with posteroventral series of minute spinules, longer proximally; longest
ca. one-third diam. of femur. Tarsomere II ca. twice length of I.

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FOSSIL EMPICORIS

537

Holotype. Male. Mexico, Chiapas, Simojovel Area. Cat. No. DT-055. Embedded
in a cut and polished piece of fossil amber, flat, tear-shaped, measuring 26 x 1 3 x
6 mm in dimension. Deposited American Museum Natural History, New York.

Remarks. Empicoris is a cosmopolitan genus of about 50 species of which five are
reported to occur in Mexico (Maldonado-Capriles and Brailovsky, 1983). The fossil
species Empicoris electricus is unique in having humeral spines, although E. incre-
dibilis Wygodzinsky and E. copal Popov have flaps in the same position. Dorsal
pronotal carinae (termed submedial carinae by Wygodzinsky [1966]) occur in E.
barberi McAtee & Malloch and E. mirabundus Wygodzinsky. The carinae are more
pronounced in the fossil species but this may be because the specimen is bilaterally
compressed; possibly as a result of the heterogenous distortion common in amber
fossilization. The thorax in Empicoris is typically dorso-ventrally compressed. The
presence or absence of spines on the scutellum, mesonotum, metanotum and first
abdominal tergite are important in species recognition. Empicoris electricus shares
with E. barberi and E. mirabundus the absence of a scutellar spine commonly found
in other species but has a metanotal spine that is absent in the latter two species.
The new fossil species is also unusual in having the second rostral segment distinctly
swollen. In the Emesinae the second rostral is at most slightly swollen (Wygodzinsky,
1966). A swollen second rostral segment is characteristic of the related subfamily
Saicinae. However, in that subfamily the rostrum and venter of the head is bristled
with spinous setae and the acetabulae of the procoxae are not directly forward (Blinn,
1990). In these regards the specimen is a typical emesine, and again the apparent
condition may be an artifact of distortion in fossilization.

DISCUSSION

When Popov (1989) described the early Miocene fossil genus Alumeda he listed
three characteristics by which it differed from all other ploarioline genera. These were
(briefly): (1) the pterostigma shortened; (2) the base of the discal cell “shortly pointed”;
and (3) the discal cell in broad contact with the wing margin. In fact these characters
will not adequately separate Alumeda from all species of Empicoris including the
fossil species described above. Popov reiterated these characters in a key to those
emesine genera related to Alumeda and Empicoris, which have a single stem vein
(M + Cu) emitting from the base of the discal cell. The key was modified from Wy-
godzinsky’s (1966) key to genera of Ploiariolini to include only the five extant genera
with the single stem vein and the fossil genus Alumeda. The couplet [10(7)] leading
to Alumeda and separating Empicoris and Ctydinna states: “Distance from apex of
pterostigma to tip of forewing no less than twice as long as distance from pterostigmal
apex to insertion of M on same; base of discal cell shortly pointed, only a basal
quarter or fifth of anterior border of cell separated from wing margin and connected
to it by one cross vein.” Actually, in some common species of Empicoris, such as
E. vagabundus (L.) and E. orthoneuron McAtee & Malloch, the ratio in the length
of the distal pterostigma to the length to the wing tip, is twice as in Alumeda. Also,
in the fossil species E. electricus, which has venation very similar to that of E.
vagabundus, this ratio is 2 x . With regard to the character of the discal cell relative
to the wing margin; in most species of Empicoris the discal cell is separated from
the wing margin by an intervening membrane and connected by two bridging cross-

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

veins. However, E. orthoneuron is like Alumeda spp. in having the discal cell in
broad contact with the margin. Similarly, some Asian species, including the holarctic
species E. culiciformis, have the discal cell in broad contact with the wing margin
(Putshkov, 1989). Thus, most of the wing venation characters emphasized by Popov
will not separate Alumeda from Empicoris. The only reliable venation character
separating the genera is the presence of two sc-r crossveins in Empicoris and Ctydinna
(only one in Alumeda). The presence of the proximal crossvein forms the angle which
results in the discal cell base being truncate instead of pointed; Popov’s second
distinguishing characteristic for Alumeda.

Popov’s (1989) and Wygodzinsky’s ( 1 966) key character separating Empicoris from
Ctydinna will also separate Alumeda. Empicoris characteristically has lateral pronotal
carinae that are lacking in Alumeda and Ctydinna. Bergroth (1909) proposed the
separation of two genera Ploiariodes White and Ploiariola Reuter (synonyms of
Empicoris ) based on the presence or absence of the pronotal carinae. McAtee and
Malloch (1925) noted that the carinae varied from complete to reduced to obsolete
(particularly in unnamed specimens from the South Pacific) and therefore disregarded
the value of the character, stating that it was worth at most subgeneric distinction.
Subsequently, China ( 1 930) proposed the subgenus Dictynna for a species from Samoa
in which a salient feature was the lack of lateral pronotal carinae. In his monograph,
Wygodzinsky (1966) elevated Dictynna to full genus, changing the name to the ana-
gram Ctydinna Wygodzinsky; Dictynna being preoccupied in the Hymenoptera. Cty-
dinna is monotypic; its one species nitidicollis China, has wing venation similar to
that of a typical Empicoris. When China proposed Dictynna as a subgenus of Em-
picoris he emphasized the lack of an emargination on the posterior border of the
basal abdominal sternite. Wygodzinsky (1966) characterized the posterior border of
this sternite as “faintly emarginated,” and did not include this character in the generic
key. Thus, Wygodzinsky elevated the monotypic Dictynna to genus level while dis-
pensing with the character considered by China to be its defining character, and
emphasizing the character (lateral pronotal carinae) considered by McAtee and Mal-
loch (1925) to be worth at most subgeneric distinction. In Empicoris rubromaculatus
McAtee & Malloch, for example, the lateral carinae, though present, are greatly
reduced. -

Thus, Ctydinna and Empicoris are closely related and distinguished only by a
character whose discreteness and significance is questionable. The fossil genus Alu-
meda differs from Ctydinna in wing venation as described by Popov (1989), and
from Empicoris by the lack of lateral pronotal carinae. Whether these differences
deserve generic or only subgeneric distinction is clearly subjective. A reanalysis of
the character-states exhibited by the fossil genus Alumeda and the extant genera,
Empicoris and Ctydinna indicates a closer relationship among them than prior treat-
ments would suggest.

ACKNOWLEDGMENTS

I am grateful to Randall T. Schuh (American Museum Natural History) and Richard C.
Froeschner (United States National Museum) for access to Emesinae specimens in their care.
I am also grateful to David Grimaldi for reviewing the manuscript.

1992

FOSSIL EMPICORIS

539

LITERATURE CITED

Bachofen-Echt, A. 1949. Der Bernstein und seine Einschliisse. Vienna. 204 pp.

Bergroth, E. 1909. E. ploeariodes B. White und Ploeariola Reut. (Hemiptera-Heteroptera,
Reduviidae). Rev. Russe Entomol. 9:324.

Blinn, R. L. 1990. Pseudosaica panamaensis, a new genus and species of assassin bug from
Panama (Heteroptera: Reduviidae: Saicinae). J. New York Entomol. Soc. 98:347-351.

China, W. E. 1930. Insects of Samoa, Pt. II. Hemiptera, Fasc. 3. Heteroptera. British Museum
(Natural History), London. 8 1 pp.

Frost, S. H. and R. L. Langenheim. 1974. Cenozoic reef biofacies, Tertiary larger foraminifera
and scleractinian corals from Chiapas, Mexico. Northern Illinois Univ. Press, DeKalb.

Maldonado-Capriles, J. and H. Brailovsky. 1983. Estudio sobre los reduviidos Mexicanos,
Parte I: Emesiinae (Hemiptera: Reduviidae). Carib. J. Sci. 19:71-90. [In Spanish]

McAtee, W. L. and J. R. Malloch. 1925. Revision of the American bugs of the reduviid
subfamily Ploiariinae. Proc. U.S. Nat. Mus. 67:1-153.

Popov, Y. A. 1987a. A new species of the bug genus Empicoris Wolff from Dominican copal,
with the redescription of E. nudus McAtee & Malloch (Heteroptera: Reduviidae: Eme-
sinae). Stuttgarter Beitr. Naturk. 134:1-9.

Popov, Y. A. 1987b. Synopsis of the neotropical bug genus Malacopus Stal, with the descrip-
tion of a new fossil species from Dominican amber (Heteroptera: Reduviidae, Emesinae).
Stuttgarter Beitr. Naturk. 130:1-15.

Popov, Y. A. 1989. Alumeda n. g., a new bug genus erected for three fossil species from
Dominican amber (Heteroptera: Reduviidae, Emesinae). Stuttgarter Beitr. Naturk. 1 50:
1-14.

Putshkov, P. V. 1989. Far eastern species of the genus Empicoris (Heteroptera, Reduviidae).
Zool. Zh. 68:146-150. [In Russian]

Schlee, D. 1980. Bernstein-Raritaten. Staatlich Museum fur Naturkunde, Stuttgart. 88 pp.

Scudder, H. S. 1890. The tertiary insects of North America. Rept. U.S. Geol. Surv. Terr. 13:
1-662.

Wygodzinsky, P. W. 1966. A monograph of the Emesinae (Reduviidae, Hemiptera). Bull.
Am. Mus. Nat. Hist. 133:1-614.

Received 17 June 1991; accepted 9 December 1991.

J. New York Entomol. Soc. 100(4):540-573, 1992

DISCOVERY OF SEXUAL WING DIMORPHISM IN
STAPHYLINIDAE (COLEOPTERA): “OMALIUM” FLAVIDUM,

AND A DISCUSSION OF WING
DIMORPHISM IN INSECTS

Margaret K. Thayer

Department of Zoology, Field Museum of Natural History,
Chicago, Illinois 60605-2496

Abstract. — Wing reduction and wing dimorphism in both sexes of an insect species are
widespread phenomena. Sexual dimorphism in wing development is less common, and is
previously unrecorded in the large beetle family Staphylinidae. Omalium jlavidum Hamilton,
a little-known forest staphylinid widespread in northeastern North America, has unusual wing
dimorphism: flightless females and males, with minute vestigial wings and modifications often
associated with wing loss; and fully-winged males, with distinctly elongate antennae, elytra, and
legs. Occurrence of this and other patterns of sex-linked wing dimorphism in insects is reviewed
and discussed. Omalium jlavidum is redescribed and illustrated and a lectotype is designated
for the species.

The presence of wings and the usually-associated ability to fly are conspicuous and
fundamental features of most insects in the vast assemblage Fterygota. Nevertheless,
secondary loss of wings in some descendants of winged insects has long been known
(e.g., Wollaston, 1854, whose data were discussed by Darwin, 1859) and is quite
widespread. In addition to orders whose members are universally wingless, strong
wing reduction or loss has been recorded within all orders of insects except Odonata,
Ephemeroptera, and Megaloptera (Rolf, 1986a); although some Ephemeroptera have
the hind wings vestigial or absent, they have not lost the ability to fly. Varying degrees
of wing reduction have been recognized, which may be roughly categorized as: aptery,
complete lack of wings; microptery, presence of very small wing vestiges (venation
highly reduced or absent); and brachyptery, possession of distinctly shortened wings
(venation almost normal to noticeably reduced, but apical region of wing strongly
reduced). In fact, these are merely approximate points on a continuum of wing
development, and additional terms have been proposed by some workers.

Most species are uniform with regard to wing development, but many show vari-
ation to different degrees. The occurrence of some kind of wing dimorphism (or
polymorphism) within species is reasonably common in many orders of insects (Rolf,
1986a and references therein; CSIRO, 1970). There are, however, different classes
of dimorphism with respect to the distribution of morphs within the sexes, and some
of these classes appear to be far more common than others (see Discussion and Table
4). The type of wing dimorphism reported here in Omalium jlavidum Hamilton,
namely micropterous females and both micropterous and macropterous males, ap-
pears to be extremely rare, as it has seldom been reported.

1992

SEXUAL WING DIMORPHISM

541

MATERIALS, METHODS, AND TERMINOLOGY

I examined a total of 161 specimens of Omalium flavidum, originating from or
deposited in the following collections (acronyms as used in collection records. Ap-
pendix B):

AMNH American Museum of Natural History, New York
ANMT A. F. Newton, Jr. and M. K. Thayer collection, Chicago
CAS California Academy of Sciences, San Francisco
CM Carnegie Museum of Natural History, Pittsburgh
CNC Canadian National Collection, Ottawa

CU Cornell University, Ithaca, New York

FMNH Field Museum of Natural History, Chicago
GHN G. H. Nelson collection, Pomona, California
MCZ Museum of Comparative Zoology, Cambridge, Massachusetts
UNH University of New Hampshire, Durham, New Hampshire
USNM National Museum of Natural History (Smithsonian Institution), Wash-
ington

Specimens were examined dry (point- or card-mounted), in alcohol, and KOH-
cleared and slide-mounted. Drawings were made with a drawing tube on a Leitz
Dialux 20 differential-interference contrast microscope; photomicrographs were taken
on the Dialux with a Leitz Vario Orthomat 2 apparatus. Scanning electron microscopy
was done with an AMRAY 1810; the specimens used were cleaned, critical-point
dried, and gold-coated. Measurements, made with an ocular micrometer in a Leitz
stereomicroscope, are defined as follows (all viewed perpendicular to line of mea-
surement):

Antennal length: from constriction at base of scape to apex of last antennomere
Head length: along midline, clypeal apex to posterior ocellar margins, dorsal view
Head width: maximum width including eyes, dorsal view
Ocular length: maximum longitudinal distance, dorsal view

Interocular width: minimum distance between dorsal margins of eyes, dorsal view
Pronotal length: along midline, base to apex, dorsal view
Pronotal width: maximum width, dorsal view

Elytral length: longitudinal from front of humerus to elytral apex, dorsal view
Elytral width: maximum combined width of closed elytra, dorsal view
Hind tibial length: base to apex, not including apical spurs or setae
Hind tarsus, tarsomeres 1-4: base of tarsomere 1 to base of tarsomere 5
Hind tarsus, tarsomere 5: base to apex, not including claws

Metathoracic length: along midline, apex of mesosternal intercoxal process to apex
of metasternum, ventral view

Metathoracic width: maximum width (at posterior margin), ventral view

Calculated figures in Tables 1 and 2 are:

Calculated length = Head length + Pronotal length + Elytral length
Eyes, 2 x width = Head width — Interocular width

Dorsal “area” (both eyes) = (Head width — Interocular width) x Ocular length

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Statistical analyses (Mests for pairwise differences between means, Fmax tests for
homogeneity of variances) were done using the computer program MYSTAT, a
product of SYSTAT, Inc. Because of the disproportionately high number of winged
males in the material available, some problems with condition of specimens, and
receipt of some material very late in the project, not all specimens were measured.
The map was produced using QUIKMap (ver. 2.5), a product of Axys Systems, Inc.

I did not attempt an exhaustive literature search regarding wing reduction or
dimorphism/polymorphism in insects. Literature on Coleoptera was searched more
thoroughly than that on other orders, and the paper is mainly focused on Coleoptera.
In the Discussion, ordinal placement of non-Coleopteran families is indicated.

Some species include individuals with only slight reduction in wing length, which
are probably capable of flight; these individuals are here lumped with fully-winged
individuals as macropterous (=fully-winged), since they are functionally so. For
simplicity, forms incapable of flight are likewise generally lumped together. For
brevity, I will often refer to brachypterous or micropterous forms as — W and macrop-
terous forms as +W. Likewise, for simplicity of expression, I will use the term wing
dimorphism to encompass both dimorphism and polymorphism of wing develop-
ment, since aptery, microptery, and brachyptery are functionally the same with re-
spect to flight capability.

OMALIUM FLAVIDUM

Omalium flavidum Hamilton is a widespread (Fig. 1) but very poorly known eastern
North American species of staphylinid beetle. It belongs to the mainly temperate
subfamily Omaliinae, and more specifically to the tribe Omaliini. Its placement in
Omaliini is supported by three derived characters (Thayer, in preparation): the pres-
ence of a (vestigial) seta-edged groove (Fig. 22) on the third abdominal tergite (Ham-
mond, 1979, figs. 16, 17), a two-chambered sclerotized spermatheca in the female
(Fig. 30), and the extremely reduced anterior tentorial arms. Females and microp-
terous males also have tarsi characteristic of Omaliini, with the basal four tarsomeres
together shorter than the fifth (Hatch, 1957; Moore and Legner, 1979).

When Hamilton (1 896:347) described Omalium flavidum, the genus Omalium was
generally used in a much broader sense than now. There is no basis for placing the
species in Omalium as now recognized (e.g., Lohse, 1964; Moore and Legner, 1979;
Zanetti, 1987), since O. flavidum lacks the characteristic carinate mesosternum and
aedeagal structure of that genus (abparameral surface of median lobe truncate well
before apex; see Zanetti, 1987, figs. 45, 46, 47a-p). The head structure of O. flavidum
is suggestive of the Palearctic genera Carcinocephalus and Dialycera, but I prefer not
to change its generic placement or erect a new genus before completing a compre-
hensive study of the genera of Omaliini (Thayer, in preparation). Since the species
was originally so incompletely described, I provide a more thorough redescription
in Appendix A of the present paper.

Sexual and male dimorphism. Since this species was named and described very
briefly almost 100 years ago (Hamilton, 1896), its only mention in the literature has
been in catalogs. I have discovered that O. flavidum is a dramatically variable species,
consisting of macropterous males (Figs. 4, 16a) and extremely micropterous females
and males (Figs. 5, 16b). According to the conventions given above, O. flavidum (or

1992

SEXUAL WING DIMORPHISM

543

Fig. 1 . Northeastern U.S.A. and adjacent Canada, showing known distribution of Omalium
flavidum Hamilton. Scalloped and dotted line shows approximate maximum limit of late
Wisconsinan glaciation (after Morgan and Morgan, 1981).

the males collectively) can thus be called ±W. Micropterous ( — W) individuals,
including the syntype specimens, are all similar in appearance and readily distin-
guishable from the + W males (compare Figs. 4, 6, 8, 16a, 17, 23, with 5, 7, 9, 16b,
18, 24). Table 1 summarizes a number of features differentiating the morphs and
Table 2 quantifies these and some other mensural characters.

Both wing morphs of O. flavidum differ from macropterous species of Omaliini in
readily observable ways. The — W forms (S and 9) have several modifications (derived
features) often associated with wing loss: shortened elytra (common in — W Staph-
ylinidae); reduced eyes, elytral humeri, and metathorax (Fig. 2a); strengthened elytral
interlocking; loss of abdominal tergal wing-folding patches; and (9 only) loss of the
palisade fringe on tergite 7. The +W<3 differ from other macropterous Omaliini (and
— W O. flavidum) in having: (1) filiform (instead of clavate) antennae, composed of
slightly elongate antennomeres (Fig. 4); (2) elongate tarsi and tarsomeres (Figs. 2b,
c, 17); and (3) very long elytra (Fig. 4). The first two of these resemble probably-
primitive features for the subfamily, but appear to be secondarily derived within
Omaliini; the third occurs sporadically within Omaliinae (presumably having evolved
several times), but is unknown to me among other Omaliini.

Conspecificity of morphs. Despite these dramatic differences between +W and — W
morphs, there are several morphological characters that argue for the different wing

Metatarsomere 5 length (mm) Metatarsal length (mm) Metathoracic length (mm)

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morphs being very closely related, and indeed conspecific. A feature unique to O.
flavidum is the form of the empodial setae. Normally, Omaliinae (like most Staph-
ylinidae) have a pair of subequal setae on each tarsal empodium. In some taxa one
of the pair is lost, leaving the empodium unisetose (e.g., in the genus Empelus, related
to Omaliinae, Thayer, 1987; Newton and Thayer, 1992); in others, for instance
Corneolabiini (see Thayer, 1985), both are lost and the empodium is asetose. All
morphs of Omalium flavidum, however, have one empodial seta greatly reduced in
size and the other distinctly longer than usual (Fig. 19). I interpret this modification
as a unique autapomorphy linking the morphs, although not necessarily indicating
conspecificity. ( Carcinocephalus ( Scribaia ) blandus (Luze) has unequal empodial se-
tae, but the shorter one is much less reduced and the difference between the two is
less.)

Male genitalia of Omaliinae, including Omaliini, usually provide characters di-
agnostic at the species level (Steel, 1957, 1960, 1964; Lohse, 1964; Smetana, 1981,
1985; Thayer, 1985; Zanetti, 1987; Watanabe, 1990), as in most Staphylinidae (and
indeed most Coleoptera and many other Insecta). In several genera of Omaliini, there
are also male secondary sexual characters of the abdominal stemites, hind tibiae,
and/or hind coxae that differ between and serve to distinguish species (e.g., Steel,
1957; Zanetti, 1987; Watanabe, 1990; Thayer, in preparation). The aedeagi of the
male morphs of O. flavidum (Fig. 28) are indistinguishable from each other in size,
external morphology, and armature of the internal sac. In addition, the arrangement
of setae at the apices of abdominal stemites 7 and 8 (Fig. 25) and the sclerotization
pattern at the apex of stemite 8 (Fig. 25) are the same in both morphs and different
from any I have seen in other Omaliinae. In addition, males of O. flavidum have
dense long blunt sensilla on antennomeres 3-1 1; it is very unusual, if not unique
among at least Omaliini, to have these sensilla located more basally than antenno-
mere 6.

The pronotum of O. flavidum (females and both male morphs; Figs. 4, 5) is
distinctive in having a raised median triangular area containing an impression on
either side of the raised midline. This is unlike the pronotum of any other Omaliini
known to me.

At least some genera of Omaliini show interspecific variation in the structure of
the defensive gland associated with the anterior projection on abdominal stemite 8.
These structures appear to be identical in females and the two male morphs of O.
flavidum (Fig. 26). Similarly, the occurrence of apical membranous fringes (and their
form, if present) and the distribution of apical spinules and ventral sensory structures
(sensilla basiconica?) on the epipharynx varies among genera and, to some extent,
species of Omaliinae. Females and both male morphs of O. flavidum all have the
same epipharyngeal ornamentation (Fig. 14), which is distinguishable (albeit some-

Fig. 2. Omalium flavidum Hamilton, (a) Metathoracic length vs. Head length + Pronotal
length; (b) Metatarsal length vs. Metatibial length; (c) Length of metatarsomere 5 vs. combined
length of metatarsomeres 1-4. 0 +W<3, x — W<5, □ $. Arrows mark points representing spec-
imens whose measurements are not included in Tables 1 and 2 because all their measurements
could not be taken.

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flight

litter

logs

snow

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SEXUAL WING DIMORPHISM

547

Table 1. Differences among wing morphs of Omalium flavidum (qualitative characters and means of
some quantitative characters and ratios). See Table 2 for sample sizes, additional mensural characters,
and standard deviations of measurements. All measurements are in mm.

Feature

6 +W

6 -W

* (-W)

Color

Dark brown (almost
black) to reddish-brown

Reddish-brown; elytra
lighter, abdomen darker

Head reddish-brown,
rest yellowish- to
reddish-brown

Antennae

Long (1.56)

Short (1.18)

Short (1.07)

Antennomeres

Elongate (Fig. 4)

Shorter (Fig. 5)

Shorter (cf. Fig. 5)

Eye size

Larger (Fig. 4)

Smaller (Fig. 5)

Smaller (cf. Fig. 5)

•Ocular length

0.17

0.12

0.12

•2 x Width

0.15

0.11

0.10

•Dorsal "area"

0.03mm2

0.01mm2

0.01mm2

Elytral length

1.47

0.78

0.80

Elytral humeri

Normal, prominent
(Fig. 4)

Reduced (Fig. 5)

Reduced (cf. Fig. 5)

Elytral locking

Loose

Tightly interlocked with
each other and
scutellum (not fused)

Tightly interlocked with
each other and
scutellum (not fused)

Wings

Fully developed, ca.
4mm long (Fig. 16a)

Small stubs < 0.1mm
long (Fig. 16b)

Small stubs < 0.1mm
long (Fig. 16b)

Metathoracic length

Longer (Fig. 2a), 0.60

Shorter (Fig. 2a), 0.37

Shorter (Fig. 2a), 0.35

Hind tarsi

Longer (0.42), tarso-
mere 5 relatively short
(Figs. 17, 2b-c)

Shorter (0.31), tarso-
mere 5 relatively long
(Figs. 18, 2b-c)

Shorter (0.29), tarso-
mere 5 relatively long
(Figs. 2b-c)

•Tarsal length/

0.67

0.58

0.56

Hind tibial length

•Tarsomere 5

0.92

1.07

1.29

length/ Tarso meres
1-4 together

Basal abdominal
spiracles (1-3)

Larger (Figs. 4, 6, 21)

Smaller (Figs. 5, 7, 22)

Larger (cf. Figs. 4, 6,
21)

•Spiracles 1-3

Distinctly larger

Slightly larger

Distinctly larger

compared to 4

Abdominal wing-
folding patches

Present on tergites
4 and 5 (Fig. 23)

Absent (Fig. 24)

Absent

Tergite 7 palisade

Present (Fig. 10)

Present (Figs. 11-12)

Absent

fringe

Fig. 3. Oma/ium flavidum Hamilton, (a) Number of specimens of each morph seen, by
month, with number above each bar indicating number of samples included; one collection of
2 — W 9 in February not included; (b) Number of specimens of each morph seen, by generalized
microhabitat, with number above each bar indicating number of samples included; “flight”
includes intercept traps and “in flight.”

548 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Table 2. Mensural characters of wing morphs of Omalium flavidum. Mean ± standard deviation for
indicated sample size (N) of each morph. Boldfaced entry indicates mean of that feature for that
morph is significantly different (by t-test) from means of both other morphs at the P < 0.001 level or
at the higher (still P < 0.05) probability level indicated. All features with means differing
significantly between morphs had variances not differing significantly (P < 0.05) among morphs, so
the t-test is appropriate. Non-bold means with no P given are not significantly different from
unmarked means of the other morphs at the P ^ 0.05 level. Means of ratios are the means of
individual ratios, not ratios of mean measurements. All measurements in mm.

Measurement or ratio

6 +W

6 -W

? (-W)

(N)

8

9

7

Calculated length

2.38 ± 0.09

1.66 ± 0.15

1.74 ± 0.14

Antennal length

1.56 ± 0.10

1.18 ± 0.11

(P < 0.045 vs. 9)

1.07 ± 0.08

Head length

0.41 ± 0.02

0.39 ± 0.03

0.42 ± 0.04

Head width

0.53 ± 0.02

0.50 + 0.04

0.54 ± 0.05

Ocular length

0.17 ± 0.07

0.12 ± 0.01

0.12 ± 0.01

Interocular width

0.38 ± 0.02

0.39 + 0.04

0.44 ± 0.03

(P < 0.002 vs. +W 6 )
(P < 0.011 vs. -Wd )

Pronotal length

0.50 ± 0.02

0.50 ± 0.05

0.53 ± 0.04

Pronotal width

0.61 ± 0.02
(P < 0.019 vs. 9)

0.62 ± 0.06

0.67 ± 0.05

Elytral length

1.47 ± 0.06

0.78 ± 0.09

0.80 ± 0.07

Elytral width

0.97 ± 0.05

0.77 ± 0.08

0.83 ± 0.07

Hind tibial length

0.62 ± 0.03

0.54 ± 0.04

0.51 ± 0.03

Hind tarsus, length
tarso meres 1-4
combined

0.22 ± 0.02

0.15 ± 0.02

(P < 0.003 vs. 9)

0.13 ± 0.02

Hind tarsus, length
tarsomere 5

0.20 ± 0.02

0.16 ± 0,02

0.16 ± 0.01

Metathoracic length

0.60 ± 0.02

0.37 ± 0.05

0.35 ± 0.04

Metathoracic width

0.83 ± 0.05

(P < 0.002 vs. 9)

0.65 ± 0.05

0.70 ± 0.07

times subtly) from that of numerous other Omaliinae examined (Thayer, unpub-
lished). Thus, characters on several parts of the body support the idea that the three
forms discussed belong to a single dimorphic species.

Distribution and ecology ;. No additional collection records have been published
since O. flavidum was described; the species is, in fact, distributed far more widely
than just western Pennsylvania (type-locality “near St. Vincent”). As shown in Figure
1, O. flavidum occurs in the northeastern U.S. (District of Columbia and western
Pennsylvania to Maine) and southeastern Canada (extreme southeastern Ontario and
southern Quebec). Relatively little habitat information is available for most collec-

1992

SEXUAL WING DIMORPHISM

549

Figs. 4-7. Omalium flavidum Hamilton. 4. Habitus, dorsal view, right elytron and wing
removed, +W<3. 5. Same, — W<5. 6. Detail of pterothorax, dorsal view, + W<3. 7. Same, — W<3.
(Scale line: Figs. 4-5, 1 mm; Figs. 6-7, 200 /urn.)

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

tions of O. flavidum (Appendix B), but it appears to be a forest-associated species.
From explicit ecological data associated with some specimens and inference from
locality and potential-vegetation data (Kuchler, 1964) for others, it appears that O.
flavidum occurs in Appalachian oak, northern hardwood, and northern hardwood-
spruce-fir forests (classification according to Kuchler, 1964). Its distribution in a
densely (human-) populated and heavily-collected area makes it surprising that O.
flavidum has been so seldom collected. The reason, as with some other Omaliinae
(e.g., Glypholoma rotundulum, Thayer and Newton, 1979; Omalorphanus aenigma,
Campbell and Chandler, 1987), appears to be its seasonality, presumably reflecting
a preference for (or at least tolerance of) cool temperatures. Available records are
concentrated in October through December, with activity (as indicated by trap catch-
es) continuing through the winter (Fig. 3a). Only five collections of one individual
each have been made from April through September (Fig. 3a), the more usual en-
tomological collecting season in the area.

Of the 163 specimens examined (116 +W males, 23 — W males, 24 — W females),
some ecological or microhabitat data are available for 141; these are plotted in
generalized terms in Figure 3b. The relatively large number of +W males in flight
intercept traps (mostly from four samples at three different sites; see Appendix B)
indicates that they do fly, although the presence of two — W males in one sample
and one each in two others suggests that the traps also acted as pitfall traps to a
limited extent (see Chandler, 1987). Forest leaf and log litter are the only microhab-
itats from which O. flavidum has been collected, but the relatively low number of
individuals thus collected (compared to flight intercept traps) and the absence of +W
males in those collections raise the question of whether that is their real microhabitat.
More investigation of the litter fauna in late autumn and winter could show them
to be more abundant there than is now apparent; the species may occupy a more
specialized microhabitat, such as mammal or bird nests. The few specimens cleared
had no visible gut contents; the beetles are probably extra-orally digesting predators.

The available data show no obvious geographical pattern in the distribution of
+ W and — W males of O. flavidum. Both forms occur more or less throughout the
species’ range (Fig. 1) and both forms have been collected at four sites and three
additional pairs of close sites. Eight sites are represented in collections by only +W
males, 5 by only — W males, 7 by only females, 4 by only — W males and females,
2 by only males ( + W and — W), and 2 by all forms.

Are there really no winged females? One must consider whether present collections
accurately reflect the nature of the species or the lack of winged females is an artifact
of relatively small sample size. A reasonable starting point in assessing this question
is the null hypothesis that winged females of O. flavidum actually exist at the same
within-sex frequency as winged males. (This is the case for, e.g., the omaliine Gly-
pholoma rotundulunr, see Discussion.) Testing this null hypothesis using all available
data (listed in Appendix B) results in decisive rejection at the P = 0.05 level (Table
3, x2 and P columns, line 1). The large number of winged males from four flight
intercept trap samples (and the mostly-fulfilled expectation that only winged speci-
mens would be caught that way) may skew the result toward rejection of the null
hypothesis by overestimating the real frequency of winged compared to wingless
males. A more stringent test, therefore, can be made by reducing the influence of the
flight intercept samples. This can be done in several ways; the x2 values and associated

1992

SEXUAL WING DIMORPHISM

551

Table 3. Chi-square tests of within-sex wing-morph frequencies of O. flavidum (using Yates’ correction
term for small expected numbers in one or more classes; see Sokal and Rohlf, 1969). Right column:
maximum within-sex frequency of winged females (as a fraction of within-sex frequency of winged males)
that does not require rejection of the null hypothesis that winged females exist.

Null hypothesis:

+ W frequency in $ =
+ W frequency in 6

X2 P

Maximum
+ W9 frequency
as a fraction of
+ W<3 frequency

1 . # of specimens

115.08

«« 0.001

0.20574

2. tt of specimens except those from flight
intercept traps

11.15

<0.001

0.49793

3. tt of collections

23.41

«0.001

0.36381

4. tt of collections except flight intercept traps

9.34

<0.005

0.57527

5. tt of specimens (flight intercept trap

collections each counted as single specimen)

25.18

«0.001

0.32279

probabilities are given in Table 3 (x2 and P columns), lines 2-5, for several possible
treatments of the data. Even with these manipulations, the equal-frequency null
hypothesis is overwhelmingly rejected at the P = 0.05 level.

An alternative class of null hypotheses can also be proposed: that winged females
exist at a lower within-sex frequency than winged males. There are no obvious grounds
for hypothesizing a particular ratio of female frequency to male frequency, but one
can take an exploratory approach and calculate, under each method of counting the
data (Table 3, lines 1-5), the highest frequency of winged females compared to winged
males that could exist but be statistically indistinguishable (P > 0.05) from the
observed values. These threshold frequencies are given in the right-most column of
Table 3. Such statistical explorations show that the existence of winged females cannot
be ruled out completely; if such females do exist, however, they occur at a distinctly
lower frequency than winged males. Flight timing of conspecific males and females
is often different, so there would not necessarily be a concurrent peak abundance of
the hypothetical +W females. The series of samples from West Lebanon, Maine (see
Appendix B) further supports the non-existence of + W$, as trapping before and after
the +W3 peak produced a few +W<3, but no females.

DISCUSSION

Types and taxonomic distribution of dimorphism. Wing dimorphism (sensu lato)
including flightless and flying morphs is the central concern of this paper, rather than
monomorphic aptery, microptery, or brachyptery. Not all variably-winged taxa show
the same pattern of dimorphism, however. Of seven possible intraspecific patterns
of variation in wing development with respect to sex, two are relatively common;
the others seem to be much rarer and one is as yet unknown. Table 4 summarizes
the higher-taxon placements of wing-dimorphic species reported in the literature,
categorized by the wing condition in each sex. One of the two most common types

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Table 4. Distribution of wing-dimorphisms with respect to sex; each taxon shown
includes at least 1 species with the type of dimorphism indicated.

+Wd

+w$

MONOMORPHICALLY +W SPECIES:

Most members of most orders of pterygote insects

±W9

Homoptera: Aphididae [2,7]

Hemiptera: Miridae [6]

Hymenoptera: Bethylidae, Formicidae [2]

Lepidoptera: Arctiidae, Geometridae, Lymantriidae, Pyralidae [8]
Diptera: Sciaridae, Tipulidae [5]

-W9

Embioptera [32]

Phasmatodea: Phasmatidae [2]

Orthoptera: Tettigoniidae [2]

Blattodea: Blaberidae [2]

Man tod ea: Amorphoscelidae, Mantidae [2]

Psocoptera [2]

Homoptera: Coccoidea [2]

Hemiptera: Microphysidae, Miridae [6]

Strepsiptera [2]

Coleoptera: Cerambycidae, Dascillidae, Elateridae, Lucanidae, Phengodidae (as Cantharidae),
Scarabaeidae [31]

Corylophidae, Drilidae, Lagriidae, Lampyridae, Meloidae, Ptinidae [17]

Rhipiphoridae [2]

Hymenoptera: Bethylidae, Cleptidae, Diapriidae, Dryinidae, Mutillidae, Sclerogibbidae [2]
Lepidoptera: Arctiidae (s. 1.), Gelechiidae, Geometridae, Hepialidae, Lasiocampidae, Lymantriidae,
Megalopygidae, Momphidae, Noctuidae, Oecophoridae, Olethreutidae, Psychidae (and related
families), Pyralidae, Symmocidae, Syntomidae, Tortricidae [8]

Diptera: Cecidomyiidae, Chironomidae, Phoridae, Sciaridae, Tipulidae (some as Limoniidae) [5]

References: [1] Crowson (1981); [2] CSIRO (1970); [3] Kalmus (1945) (Agaonidae as Chalcididae); [4] Venturi
(1964); [5] Hackman (1964); [6] Southwood & Leston (1959); [7] Hille Ris Lambers (1966); [8] Hackman (1966);
[9] Darlington (1936); [10] Darlington (1943); [11] Lindroth (1979); [12] Den Boer et al. (1980); [13] Langor &
Larson (1984); [14] Ljebherr & Hajek (1986); [15] Desender (1989); [16] Shute (1980); [17] Jackson (1928);

[18] Pope (1977); [19] Hammond (1985); [20] Paulian (1988); [21] Dybas (1978); [22] Taylor (1978; 1981);

1992

SEXUAL WING DIMORPHISM

553

Table 4 (continued). "-W" includes brachypterous, micropterous, and apterous.
Numbers [x] indicate sources listed below; see Literature Cited for full references.

±Wd

-Wd

No examples found

Psocoptera [2]
Thysanoptera [3]
Coleoptera: Scolytidae [1]
Hymenoptera:

Agaonidae [2,3]
Torymidae [2]

Diptera: Sciaridae [4]
Tipulidae (as
Limoniidae) [5]

Orthoptera: Acrididae [2]

Psocoptera [3]

Gryllidae, Pyrgomorphidae [26]

Homoptera: Adelgidae,

Blattodea: Blaberidae [2]

Pemphigidae [2]

Isoptera [2]

Thysanoptera [3]

Zoraptera [2]

Homoptera: Aphididae [2]

Delphacidae [29]

Hemiptera: Aradidae, Berytidae, Cimicidae, Dipsocoridae, Hebridae,

Hydrometridae, Mesoveliidae, Miridae, Nabidae, Pyrrbocoridae, Reduviidae,
Rhopalidae, Saldidae, Tingidae, Veliidae [6]

Genridae [6,27]

Lygaeidae [6,28]

Thysanoptera [3]

Coleoptera: Carabidae [9-15]

Chrysomelidae [16]

Cicindelidae, Curculionidae, Endomychidae, Hydrophilidae, Nitidulidae [17]
Coccinellidae [18,19]

Dermestidae [9]

Dryopidae [20]

Ptiliidae [21,22]

Staphylinidae [19,23,24,thispaper]

Trogidae [25]

Lepidoptera: Elachistidae, Gelechiidae, Noctuidae, Tineidae, Tortricidae,
Yponomeutidae(as Hyponomeuthidae) [8]

Diptera: Chloropidae, Sphaeroceridae (as Borboridae), Tipulidae [5]

Sciaridae [30]

Diptera: Cecidomyiklae [5]

Embioptera [32]

MQNQMQRPHICALLY j

Homoptera: Aphididae [7]

-W SPECIES:

Coleoptera: Carabidae? [33]

Some members of most

Dermestidae [34]

1 Staphylinidae [this paper]

Diptera: Sciaridae [5,35]

Tipulidae (as Limoniidae) [5]

orders of neopterous insects

[23] Hammond (1979); [24] Campbell (1983); [25] Scholtz (1981); [26] Harrison (1980); [27] Vepsalainen (1978);
[28] Slater (1975; 1977); [29] Denno & GrisseU (1979); [30] Steffan (1973; 1975); [31] Van Dyke (1933); [32] Ross
(1970); [33] Liebherr (1989): Tanystoma diabolica may be ±W<3/-W9; it was described from only 5 specimens
(2 +Wd, 1 -W<J, 2 -W9); [34] Barber (1947; 1948); [35] Steffan (1966).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

is that in which both sexes show similar variation in wing development (± W<5/± W9),
although morph frequencies may differ between the sexes. Such non-sexual dimor-
phism has been recorded in some members of at least 49 families in ten orders (Table
4). Among these, Isoptera are a highly specialized case, with macropterous reproduc-
tives and apterous soldier and (in higher termites) worker castes occurring in both
sexes.

The other commonest pattern of intraspecific wing variation is +W<3/— W9, one
of the six types involving sexual dimorphism. This pattern occurs in all Strepsiptera
(CSIRO, 1970), most Embioptera (Ross, 1970), and in some members of at least 49
families in 1 1 other orders.

Other sex-related patterns of wing variation appear to be distinctly rarer than these
two, especially among Coleoptera. The pattern of + W<3/±W9 has been reported in
at least ten families in five orders (Table 4). Most species of Formicidae (Hymenop-
tera) can be regarded as a special case of this category, having two or more female
castes ( + W reproductives, — W workers and sometimes soldiers) and only +W males.
Aphididae (Homoptera) (Hille Ris Lambers, 1966) and Lymantriidae and Pyralidae
(both Lepidoptera) show seasonal (generation-to-generation) rather than simulta-
neous (within-generation) dimorphism (Hackman, 1966).

I found no reports of species that are ±W<3/+W9, and only a few (in four orders)
that are — W<5/±W9. Species with the pattern — W<5/+W9, in at least seven families
in five orders, include Agaonidae (Hymenoptera) (Kalmus, 1945, as Chalcididae;
CSIRO, 1970) and some of the Torymidae (Hymenoptera) that parasitize them
(CSIRO, 1970).

The pattern of primary interest in this paper is the also rare one of ±W<3/— W9.
This pattern has been reported previously in four orders (see Table 4), mostly in a
single species per family. Discovery of this pattern in Omalium flavidum Hamilton
provides the first case of this (and in fact any) pattern of sexual wing dimorphism in
Staphylinidae.

Associated morphological changes. Various morphological changes beyond lack of
wings are commonly associated with the evolution of flightlessness in insects. Co-
leoptera, with their front wings modified into elytra and related specializations of
thoracic structure, have diverged significantly from other insects in general body
organization. Many modifications accompanying wing loss in Coleoptera thus have
no close parallel in other orders, and the following discussion deals principally with
Coleoptera.

The most extreme modifications are generally found in monomorphically bra-
chypterous species, particularly those belonging to entirely flightless lineages. In the
latter cases flight capability presumably was lost in a temporally rather distant an-
cestor of any extant species, allowing more time for development of other modifi-
cations than would be expected for a wingless species having all fully- winged relatives.
At its fullest development (short of reduction to a larviform morph), what could be
called a “flightless beetle syndrome” includes: loss of wings and flight muscles (direct
and indirect); rounding of elytral humeri; tighter than normal locking, or even fusion
(ankylosis), of the elytra to each other; reduction in length and width of the meta-
thorax; reduction in elytral length (sometimes to tiny vestiges); loss of abdominal
wing-folding and wing-toiletry devices (Hammond, 1979, 1985); reduction in dorsal
sclerotization of the pterothorax and (except in brachelytrous forms, e.g., Staphylin-

1992

SEXUAL WING DIMORPHISM

555

idae and Meloidae: Meloe ) abdomen; reduction in size of spiracles of abdominal
segment 1 (Riischkamp, 1927; Jackson, 1956); loss of eyes; and sometimes (depending
at least partly on microhabitat) reduction in antennal length and/or body pigmen-
tation. Within a number of insect orders, ocelli are sometimes reduced or lost along
with wings (Kalmus, 1945). Very atypically for Coleoptera, most Omaliinae (Staph-
ylinidae) have ocelli, but no correlation of wing and ocellar loss has yet been found
in this group.

Jackson (1928) found no external differences between macropterous and brachyp-
terous individuals of Sitona hispidula (Curculionidae). The brachypterous morphs
of many other wing-dimorphic species of Coleoptera (also Lygaeidae [Hemiptera];
Slater, 1977) do, however, exhibit some features of the “flightless beetle syndrome”
despite normal development of the corresponding features in macropterous conspe-
cifics. A few examples from Coleoptera follow. Reduction of eyes in wingless morphs,
reported here in O. flavidum, has also been reported in Aglyptinus dimorphicus
(Leiodidae, Peck, 1978), Leistus americanus (Carabidae, Darlington, 1936), and spe-
cies of the genus Ptinellodes (Ptiliidae, Dybas, 1978). The last-named also show
reduction of elytra in the wingless morph (“vestigial morph” of Dybas, 1978), as do
Platystethus nitens (Staphylinidae, Hammond, 1985) and Omalium flavidum (re-
ported here). Reduction of the elytral humeri occurs in wingless morphs of Trox spp.
(Trogidae, Scholtz, 1981), Pseudomacronychus spp. (Dryopidae, Paulian, 1988), O.
flavidum, and rarely in dimorphic Carabidae (though commonly in monomorphically
— W species; Darlington, 1936). Reduction in metathoracic size has been found in
wingless morphs of Platystethus nitens (Staphylinidae, Hammond, 1985) and Tany-
stoma maculicolle (Carabidae, Liebherr and Hajek, 1986) as well as O. flavidum,
while reduction in dorsal meso- and metathoracic sclerotization occurs in Sitona
hispidula (without reduction in size; Curculionidae, Jackson, 1928) as well as O.
flavidum. Reduction in size of the spiracles of abdominal tergite 1 seems not to have
been reported previously in dimorphic beetle species, but occurs in — W males of O.
flavidum.

Discussion of the state of flight muscles is seldom part of normal taxonomic work,
but in special studies of the subject, reduction or loss of these muscles in wingless
and some winged individuals has been found in Sitona hispidula (Curculionidae,
Jackson, 1928), Bembidion lampros (Carabidae, Langor and Larson, 1984), and sev-
eral other species of Carabidae (Den Boer et al., 1980). In many cases (including
some monomorphically macropterous species discussed by Den Boer et al., 1980),
there is seasonal change in development of flight muscles in individuals. Undoubtedly
more such examples will be found when more species are investigated in this regard.
Fusion of elytra seems not to have been recorded in dimorphic Coleoptera, although
Lindroth (1949) mentioned two species of Carabidae in which some brachypterous
individuals show a slight amount of fusion.

Two features of the “flightless beetle syndrome” that are quite rare in wingless
morphs of dimorphic species are loss of abdominal wing-folding patches and loss of
“palisade” or “wing-toiletry” fringes (Hammond, 1979). Hammond (1985) surveyed
over 300 species of wing-dimorphic beetles in several families and found (in addition
to Ptinellodes spp., Dybas, 1978) only one, Rhyzobius litura (Coccinellidae), dimor-
phic in presence of wing-folding patches and only R. litura and some species of Othius
and Mycetoporus (both Staphylinidae) dimorphic in presence of the palisade fringe.

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Glypholoma rotundulum (Staphylinidae: Omaliinae) is dimorphic in the size, not
presence, of its wing-folding patches (Thayer, unpublished). Omalium flavidum, as
reported here, provides an additional example of dimorphism in each of these features
(see above and Appendix A), although the distributions of the reductions are not
concordant. This non-concordance runs counter to the sometimes-observed pattern
of morphological changes associated with wing loss apparently evolving in a certain
sequence (at least within a given lineage), yielding a nested pattern of losses (Jeannel,
1926; Hammond, 1979; Scholtz, 1981).

Mechanisms producing and maintaining wing dimorphism. Rolf (1986a, b) re-
viewed mechanisms of wing length determination; the number of species whose wing-
determination has been studied in detail is still extremely small. At least many cases
of wing dimorphism are what Clark (1976) called “genetically determined polymor-
phism” (Rolf [1986a, b] and Rolf and Fairbaim [1991] cited a total of 29 examples).
Sequential wing dimorphism (e.g., seasonal changes in many aphids: Hille Ris Lam-
bers, 1966; Lees, 1966; Clark, 1976; MacKay et al., 1983) and some cases of non-
seasonal dimorphism appear, on the other hand, to be largely what Clark termed
“environmentally cued polymorphism,” in which environmental factors interact with
the genotype to produce different morphs under different conditions. Such a mech-
anism was suggested by Dybas (1978) for some Ptiliidae and by Denno and Grissell
(1979) for a species of Delphacidae (Homoptera). There can also be genetic variation
in presence or thresholds of environmental response (Vepsalainen, 1978; Denno and
Grissell, 1979). Morph determination in individuals is probably hormonally medi-
ated, possibly by levels of juvenile hormone (Southwood, 1961; Wigglesworth, 1961;
Rolf, 1986a, b).

Even where genetic control of wing morphs occurs, there appears to be variation
in its nature. In the 29 examples cited by Roff and Fairbaim (1991) and Roff ( 1 986a,
b), single-locus systems are more common in holometabolous orders and polygenic
systems far more common in paurometabolous orders. Roff ( 1 986b) argued that even
if wing dimorphism arose in a species as a single-locus mutation, selection would
favor replacement of single-locus control by a system of polygenic control because
of the greater flexibility of polygenic systems in responding to environmental fluc-
tuations. Rolf ( 1 990a) and Rolf and Fairbaim (1991) gave evidence for the occurrence
of antagonistic pleiotropy in maintenance of wing dimorphism in Gryllus firmus
(Orthoptera) (and potentially in other species) in the form of a tradeoff between
migration ability and earlier age at reproduction. The contention in the former work,
however, that “expression of wing form . . . in . . . wing dimorphic insects in general,
is highly dependent on environmental conditions such as temperature and photo-
period . . .” appears to be based almost entirely on paurometabolous insects (e.g.,
Honek, 1976, 42 paurometabolous, 5 holometabolous species). Roff and Fairbaim
(1991), in contrast, said “. . . the available evidence suggests a genetic basis [for wing
dimorphism] in most species. . . .” The seeming difference between holometabolous
and paurometabolous insects in the type of genetic control of wing polymorphism
and the still-limited data bearing on the question leave the importance of environ-
mental conditions to morph determination in Holometabola uncertain.

I have found no explicit discussions of the genetic or non-genetic basis of sexually
dimorphic wing development; Rolf (1986a) listed two species of Aphididae (Ho-
moptera) as having polygenic wing morph determination, but did not specifically

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discuss sexual wing dimorphism. Work done on sexual dimorphism of wing devel-
opment in aphids (e.g., Lees, 1966; MacKay et al., 1983) suggests that determination
of morphs in female aphids is under complex environmental control. Determination
of wing morphs in male aphids has been studied far less, and it is not clear whether
it is produced by genetic or environmental factors (or both). In any case, the extreme
complexity of aphid life cycles leaves some question as to how applicable the aphid
results are to other insects.

Ecology and evolution of wing loss. On the basis of observed occurrences and
theoretical considerations, various workers have summarized or predicted conditions
under which brachyptery is favored, including: small-area stable (“permanent”) hab-
itats (Darlington, 1943; Vepsalainen, 1978), including those on mountains (Darling-
ton, 1943; Scholtz, 1981); isolated habitats (in combination with stability and areal
limitation; Lindroth, 1949; Vepsalainen, 1978); cold areas or seasons of activity
(Downes, 1965; Byers, 1969); caves (Jeannel, 1926; for different reasons, Barr, 1968);
and areas providing distinctly suboptimal conditions for a species (Jeannel, 1926,
but with little or no support from other workers). It is also well known that many
nidicolous or parasitic insects are wingless (e.g., Arixeniina and Hemimerina [Der-
maptera]; Phthiraptera [loss of wings probably preceded parasitism; Lyal, 1985];
Cimicidae and Polyctenidae [Hemiptera]; Staphylinidae: Amblyopinini and Leiod-
idae: Platypsyllinae [Coleoptera]; Siphonaptera; Nycteribiidae, Hippoboscidae, and
Streblidae [Diptera]). Hackman (1964) attributed most cases of wing loss in Diptera
(aside from halobionts and parasites) to the adults’ living in “concealed terricolous
habitats” (including galleries of social insects), where he suggested selection might
favor morphological changes improving the ability to run, even at the expense of the
ability to fly. The unusual groundplan of Coleoptera, with the flight wings protected
by elytra, facilitates the coexistence of good running and flying abilities and presum-
ably would reduce selective pressure against flight in microhabitats with restricted
passageways. Wing loss does appear to be less common in litter-dwelling Coleoptera
than Diptera, although soil-inhabiting beetles are more likely to be wingless (e.g.,
Coiffait, 1960).

Rolf ( 1 990b) judged available data to be consistent with the idea of habitat stability
favoring evolution of flightlessness, but not yet adequate to test the importance of
other potential factors such as temperature or physical constraints. Barbosa et ah
(1989) argued convincingly that habitat stability alone may not be a sufficient ex-
planation for evolution of brachyptery or wing dimorphism; the way in which a
species interacts with its habitat may also play a key role.

In contrast to the idea of strongly directional selection for brachyptery in stable
environments, Hamilton and May (1977) demonstrated that even in relatively un-
changing habitats an evolutionarily stable strategy (ess) is likely to include a significant
probability of migration of offspring from the parent’s “unitary site” (individual site
occupied by one indivdual in the model). Although these authors mentioned the
relevance of their models to insect species having flying males and flightless or flight-
dimorphic females, they did not discuss the key relationship between unitary site
size and capability of dispersal by means other than flight. How large is an insect’s
unitary site? In a species whose individuals can move beyond their natal unitary site
by walking, swimming, or larval ballooning (Barbosa et al., 1989), even complete
loss of flight capability would not preclude an ess involving significant probability

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of dispersal. This potential nonidentity of flight capability and dispersal/migration
ability, admittedly difficult to investigate, is often overlooked in discussions of wing
and flight loss.

Reasons for the origin, persistence, and spread of wing loss remain somewhat
uncertain and may well vary among species. Although Jeannel suggested “senility”
of lineages ( 1 926) or inheritance of changes caused directly by effects of cave envi-
ronments (1943), there seems little reason to doubt that random mutations are the
underlying cause of wing loss. Darlington (1936) suggested that wing atrophy arises
through spontaneous mutations (occurring persistently at a significant rate), and might
in some cases increase through simple accumulation of the mutations in the absence
of natural selection against wing loss. On the other hand, it seems likely that natural
selection could often be involved, whenever advantages of winglessness (or flight-
lessness) outweigh selective pressures favoring flight. Both flight and the synthesis of
muscle and other tissues needed for it are energetically expensive (Rolf and Fairbaim,
1991 and references therein). Wingless forms, by rechanneling this biosynthetic and
locomotory energy into earlier or greater total reproduction, might gain a substantial
fitness advantage over + W forms under circumstances where flight was not essential
(Darlington, 1936, 1943; Byers, 1969; Dybas, 1978; Rolf and Fairbairn, 1991; but
not Taylor, 1978). (Evidence for such an “energy conservation hypothesis” in bac-
terial systems is mixed [Zamenhof and Eichhorn, 1967; Dykhuizen, 1978].) In con-
trast, Regal’s (1977) “noise suppression theory” proposes that natural selection favors
loss of non-essential structures because such elimination removes informational noise
(thus potential errors) from the process of transcription of the genome.

Wing dimorphism has been predicted to occur when the habitat (or microhabitat)
of a species is mostly stable with occasional disruptions (Darlington, 1936; South-
wood, 1962; Hammond, 1985; Rolf, 1986a). When a species has different habitat
requirements at different times of year, the selective advantage of dispersing may
vary seasonally (Cohen, 1967); under such conditions, sequential wing dimorphism
may be favored in bi- or multivoltine species (Harrison, 1980). Some authors (e.g.,
Lindroth, 1949; Den Boer et al., 1980; Rolf and Fairbairn, 1991) have regarded wing
dimorphism or polymorphism as an intermediate state in the transition to brachyp-
tery, in essence predicting that dimorphic species should be found in the same sit-
uations as monomorphically brachypterous ones, but in habitat patches that have
been stable a shorter time than those with brachypterous species.

There has been relatively little discussion of possible reasons for sex-biased wing
dimorphism. Downes (1965) and Byers (1969) suggested that in the Arctic the severe
energetic demands of the climate have caused selection for a +W<3/— W9 pattern.
The short available reproductive season there makes it advantageous for females to
produce their eggs as soon as possible after emerging in the spring; the resulting extra
weight of mature eggs would make female flight energetically very expensive, perhaps
impossible. In a number of arctic species, therefore, females appear to have maxi-
mized their reproductive effort by abandoning flight ability and males have special-
ized in mate location and dispersal by flight, the latter permitting outcrossing. (Some
other arctic species have instead abandoned outcrossing or even mating.) Such ar-
guments may apply equally well to cold montane areas at lower latitudes.

Other circumstances associated with the pattern of +W<$/— W$ are: (1) in some

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559

Hymenoptera, adaptations for burrowing (or host-seeking behavior in other tight
places) by females and transport of therefore — W females by males (CSIRO, 1970)
and (2) in some eastern North American forest Lepidoptera, a complex of life-history
traits including polyphagy on dominant trees with common chemical defenses, uni-
voltinism, larval ballooning, and overwintering as eggs or larvae (Barbosa et al.,
1989). Clearly neither these nor the cold hypothesis provides a general explanation
for sexual wing dimorphism, but all three point out the complex connections between
such dimorphism and life-history traits.

Rolf and Fairbaim (1991) found reproductive advantages of brachyptery for fe-
males, but not males. This inequality may, in fact, be one factor behind the relative
commonness of + W<3/— W9 species. Evolution of such dimorphism would, of course,
depend on preexistence of a suitably modifiable genetic control system. Given that
sex determination is usually chromosomal, it seems possible that a locus on one of
the sex chromosomes might modify the effects of a somatic allele for brachyptery to
produce various kinds of sex-linked patterns of dimorphism.

How does Omalium flavidum fit in? Omalium flavidum is one of the few insect
species known to have ±W males and — W females, and is the first such species
recorded in the large and biologically diverse beetle family Staphylinidae.

Omalium flavidum is unusual among wing-dimorphic beetle species in exhibiting
dimorphism of two wing-associated abdominal structures (palisade fringe and tergal
wing-folding patches). Hammond (1979, 1985) found only a few cases of variability
in one or both of these structures in wing-dimorphic beetle species; in those cases,
the T W morph retained and the — W morph lacked the structure in question. Gly-
pholoma rotundulum, mentioned above, has reduced wing-folding patches in the
micropterous morph (see below). In O. flavidum, the loss of wing-folding patches is
fully correlated with wing loss, but the palisade fringe is lost only in females (see
Table 1). The tergite 3 groove usually present in winged (but not wingless) species
of Omaliini is vestigial in females and both male morphs.

The patterns of variation found in O. flavidum differ from the other cases of wing-
variation known so far in Omaliinae in being sexually dimorphic and apparently
geographically undifferentiated. Glypholoma rotundulum was described (Thayer and
Newton, 1979) from southeastern Australia on the basis of over 200 micropterous
(there called “brachypterous”) males and females from several sites, plus 2 truly
brachypterous individuals ( 1 6, 19) and 1 macropterous <3 from the two (very close)
northernmost sites. Based on the type series and additional material I have seen
(including macropterous males and females from the same and two additional lo-
calities; Thayer, unpublished), tests like those in Table 3 support the hypothesis that
fully-winged individuals occur at the same (low) frequency in both sexes of G. ro-
tundulum. The abdominal wing-folding patches are dimorphic in this species, being
larger in macropters and brachypters than in micropters. Brachypterous and macrop-
terous individuals of this species are still known only from the northernmost localities
for the species and the micropterous form, universal elsewhere, has not been collected
at those sites. Hammond (1985) mentioned wing dimorphism in four other species
of Omaliinae, none of which appears to be sexually wing-dimorphic. Three of these
( Olophrum fuscum (Gravenhorst), Eucnecosum (as Arpedium ) brachypterum (Gra-
venhorst), and Acidota cruentata Mannerheim), with macropters rare to very rare,

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

have the wing-folding patches monomorphic and smaller than those of related ma-
cro ptero us species. The fourth, Anthobium unicolor, has varying frequencies of ma-
cropters over its range; Hammond did not discuss its wing-associated structures.

There is also a slightly odd pattern of abdominal spiracle size in O. flavidum. All
morphs have the spiracles of segment 1 larger than those of segments 2 and 3, which
are in turn larger than those of segment 4; this seems to be the normal configuration
in at least Omaliini (Thayer, unpublished) and probably a much wider group. The
spiracles on segments 1-3 of macropterous males are larger than the corresponding
ones of — W males (Figs. 4, 6, 21 vs. 5, 7, 22). Winged species of Omaliini similarly
have larger spiracles 1-3 than do wingless species, and such a difference has also
been found between + W and — W species in Chrysomelidae (Riischkamp, 1927) and
Dytiscidae (Jackson, 1956). The ( — W) females of O. flavidum also have larger spir-
acles than — W<3, however. This could reflect higher oxygen needs of egg production
compared to sperm production, or perhaps greater locomotory activity among females
than — W males. I have not seen a sex-based difference in spiracle size in wingless
species of Omaliini, and neither Riischkamp (1927) nor Jackson (1956) mentioned
any differences between the sexes in this regard. Divergence in spiracle size may be
greatly underreported, however, since (like wing-muscle development) it is not a
character usually examined in taxonomic work.

As discussed earlier, wing dimorphism has been shown to be variously genetically
or environmentally controlled (or a combination) in different species. From the
limited data available, it appears that in Coleoptera (as in other Holometabola) it is
usually genetically determined (Roff, 1986a). In the absence of any rearing data or
evidence regarding seasonal change in morph frequencies, it is impossible to say how
wing dimorphism is determined in O. flavidum. If there are indeed no + W females,
the mechanism in this species is clearly not a simple single-locus, two-allele, auto-
somal genetic system with brachyptery dominant, like that found in several species
of Coleoptera (Carabidae, Curculionidae) and Diptera (Sphaeroceridae) cited by Roff
(1986a). Two possibilities are: (1) some form of direct genetic sex-linkage or (2) sex-
determined effects on hormonal mediation of wing development, with females re-
acting in one way and males in either of two ways.

Most previously reported ±W<3/— W9 species have male morphs that are at least
partly sympatric (Dermestidae: Thylodrias contractus, Barber, 1947; Sciaridae [Dip-
tera]: Pnyxia scabiei, Hackman, 1964; some Embioptera, CSIRO, 1970; but not
Tipulidae [Diptera]: Pedicia hannai, Hackman, 1964; Byers, 1969). Not all known
O. flavidum localities are represented by specimens of all morphs (or even both
sexes), but both male wing morphs are widely distributed over the range of the species;
no clinal variation in occurrence vs. latitude is evident (Fig. 1), in contrast to nu-
merous species mentioned by Roff (1990b). There is no other obvious pattern of
geographical restriction, and the two male wing morphs have been collected together
at four sites so far; O. flavidum is thus not unusual among ±W3/ — W9 species in the
geographical distribution of its male morphs.

The driving force favoring evolution of wing dimorphism in Omalium flavidum
(or brachyptery, if its dimorphism is a transitional stage) is not yet clear, perhaps in
part because of the limited ecological data available. As mentioned above, the species
inhabits the widespread eastern North American deciduous and deciduous-coniferous
forests. Before the widespread forest clearing that accompanied European settlement.

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561

this climax forest biome had a far more continuous distribution than it does now.
This would have provided large areas of relatively unbroken favorable habitat for
O. flavidum, presumably with scattered areas of disturbance caused by severe storms
or occasional treefalls. Several authors (Southwood, 1962; Harrison, 1980; Ham-
mond, 1985; Roff, 1 986a) have suggested that conditions such as these favor evolution
(in both sexes) of either wing-dimorphism or brachyptery, depending on the level of
habitat stability. Could bisexual dimorphism be a stage that Omalium flavidum has
already passed through, en route to brachyptery? Such a pathway to brachyptery can
be envisioned: under conditions allowing brachyptery, if the energy conservation
hypothesis is valid, stronger (hence faster) selection for wing loss in females than
males might occur because of the greater reproductive investment by females than
males. If this were the case, however, one would expect wing dimorphisms like that
of O. flavidum to be far more common.

On an ecological time scale, the eastern forests have constituted a relatively stable
habitat for a long time; their stability, however, does not extend very far back on a
geological scale. All but the southernmost records of O. flavidum (southern Penn-
sylvania, District of Columbia vicinity) are from areas that were ice-covered during
the Pleistocene glaciations (Fig. 1; Flint, 1971; Morgan and Morgan, 1981). Thus,
the eastern forests only came to occupy a large part of the present range of O. flavidum
during the last 8,000-10,000 years (Matthews, 1979), presumably moving in from
the south (and/or east, if the continental shelf formed a glacial refugium; Flint, 1971).
Obviously, stable forest did not appear immediately behind the disappearing glaciers,
so the time available for colonization by mature-forest inhabitants such as O. flavidum
would have been even less.

When did wing loss develop in O. flaviduml Three alternatives can be proposed
if the wing loss is assumed to be a result of habitat stability. First, if wing loss
developed independently in many newly established local populations as a result of
relatively stable habitats, one would expect to find a distribution pattern like that of
Calathus mollis (Carabidae) in Scandinavia (Lindroth, 1979), with brachypterous
forms predominating in the longest-occupied (southern) areas. No such pattern is
apparent in the data available for O. flavidum, so this appears not to be the expla-
nation. Second, if dimorphism arose only after O. flavidum had fully occupied more
or less its present range, the phenomenon would not necessarily occur throughout
the species’ range, and morph frequencies might be expected to vary more or less
randomly over space. Multiple population samples (taken throughout the year) would
be needed to test this hypothesis; the data now available for O. flavidum are far too
scanty for such purposes. Finally, the species might already have had its present form
of wing-dimorphism while in a glacial refugium and when the ice sheet began to
retreat, managed to disperse over a large area despite the flightlessness of females.
Although this seems absurd at first glance, over a period of 8,000 years it would
require movement on the order of 100 m per year, which might be possible. (Morgan
and Morgan, 1981, arrived at a similar conclusion for a flightless species of Carab-
idae.)

Alternatively, O. flavidum's seasonal pattern of occurrence, seeming to reflect a
preference for (adaptation to?) cool to cold conditions, and the (partly) sexual nature
of its dimorphism, suggest that wing-dimorphism in this species might be an example
of a cold-adaptation pattern such as discussed above (Downes, 1965; Byers, 1969).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Further knowledge of the distribution and seasonality of the species, study of larger
samples of multiple populations, investigation of the genetic basis of its wing di-
morphism, and more detailed information about its habits and microhabitat may
help to clarify the factors involved in the evolution of the unusual wing dimorphism
found in Omalium flavidum. Understanding this case might also help to elucidate
the reasons for the evolution of a similar pattern in other taxa and for the rarity of
this pattern.

ACKNOWLEDGMENTS

I thank the following for providing access to specimens in their care: L. H. Herman (AMNH);
D. H. Kavanaugh (CAS); J. E. Rawlins, G. Wallace, and C. W. Young (CM); J. M. Campbell
(CNC); L. L. Pechuman, J. K. Liebherr, and E. R. Hoebeke (CU); D. S. Chandler (UNH); G.
N. House (USNM); and G. H. Nelson (GHN). 1 owe special thanks to Don Chandler, probably
the only person in the world getting tired of collecting O. flavidum. B. Chemoff (FMNH) provided
valuable advice on the statistical treatments; he and A. F. Newton (FMNH), C. H. C. Lyal
(Natural History Museum, London) and four anonymous reviewers made comments that sig-
nificantly improved the manuscript. This work was supported by grant NSF BSR-8806625;
preliminary work was supported by an NSF Graduate Fellowship at Harvard University.

LITERATURE CITED

Barber, H. S. 1947. On the odd, or tissue-paper, beetle supposed to be Thylodrias contractus
(Coleoptera: Dermestidae). Ann. Entomol. Soc. Amer. 40:344-349.

Barber, H. S. 1948. Postscript on the “Odd Beetle.” Ann. Entomol. Soc. Amer. 41:478.
Barbosa, P., V. Krischik and D. Lance. 1989. Life-history traits of forest-inhabiting flightless
Lepidoptera. Amer. Midi. Nat. 122:262-274.

Barr, T. C. 1968. Cave ecology and the evolution of troglobites. Pages 35-102 in: T. Dob-
zhansky, M. Hecht and W. Steere (eds.). Evolutionary Biology. Vol. 2. Appleton-Century-
Crofts, New York, xi + 452 pp.

Byers, G. W. 1969. Evolution of wing reduction in crane flies (Diptera: Tipulidae). Evolution
23:346-354.

Campbell, J. M. 1983. A new species of Pycnoglypta Thomson (Coleoptera: Staphylinidae)
from eastern Canada. Can. Entomol. 1 15:361-370.

Campbell, J. M. and D. S. Chandler. 1987. Omalorphanus aenigma, an unusual new genus
and species of Omaliinae (Coleoptera: Staphylinidae) from Oregon. Can. Entomol. 1 19:
315-327.

Chandler, D. S. 1987. Species richness and abundance of Pselaphidae (Coleoptera) in old-
growth and 40-year-old forests in New Hampshire. Can. J. Zool. 65:608-615.

Clark, W. C. 1976. The environment and the genotype in polymorphism. Zool. J. Linn. Soc.
58:255-262.

Cohen, D. 1967. Optimization of seasonal migratory behavior. Amer. Nat. 101:5-17.
Coiffait, H. 1960. Les coleopteres du sol. Vie et Milieu Supplement No. 7:1-204.

Crowson, R. A. 1981. The Biology of the Coleoptera. Academic Press, London, 802 pp.
CSIRO Division of Entomology. 1970. The Insects of Australia. Melbourne University Press,
Carlton, xiii + 1029 pp.

Darlington, P. J., Jr. 1936. Variation and atrophy of flying wings of some carabid beetles
(Coleoptera). Ann. Entomol. Soc. Amer. 29:136-179.

Darlington, P. J., Jr. 1943. Carabidae of mountains and islands: data on the evolution of
isolated faunas, and on atrophy of wings. Ecol. Monogr. 13:37-61.

Darwin, C. 1859. The Origin of Species by Means of Natural Selection. J. Murray, London
(seen in 1979 reprint by Avenel Books, New York), xi + 460 + 16 pp.

1992

SEXUAL WING DIMORPHISM

563

Den Boer, P. J., T. H. P. van Huizen, W. Den Boer-Daanje, B. Aukema and C. F. M. Den
Bieman. 1980. Wing polymorphism and dimorphism in ground beetles as stages in an
evolutionary process (Coleoptera: Carabidae). Entomol. Gen. 6:107-134.

Denno, R. F. and E. E. Grissell. 1979. The adaptiveness of wing-dimorphism in the salt-
marsh inhabiting planthopper, Prokelisia marginata (Homoptera: Delphacidae). Ecology
60:221-236.

Desender, K. 1989. Heritability of wing development and body size in a carabid beetle,
Pogonus chalceus Marsham, and its evolutionary significance. Oecologia 78:513-520.

Downes, J. A. 1965. Adaptations of insects in the Arctic. Ann. Rev. Entomol. 10:257-274.

Dybas, H. S. 1978. Polymorphism in featherwing beetles, with a revision of the genus Pti-
nellodes (Coleoptera: Ptiliidae). Ann. Entomol. Soc. Amer. 71:695-714.

Dykhuizen, D. 1978. Selection for tryptophan auxotrophs of Escherichia coli in glucose-limited
chemostats as a test of the energy conservation hypothesis of evolution. Evolution 32:
125-150.

Flint, R. F. 1971. Glacial and Quaternary Geology. John Wiley and Sons, New York, ix +
892 pp.

Hackman, W. 1964. On reduction and loss of wings in Diptera. Not. Entomol. 44:73-93.

Hackman, W. 1966. On reduction and loss of wings in Lepidoptera. Not. Entomol. 46:1-16.

Hamilton, J. 1896. Catalogue of the Coleoptera of Southwestern Pennsylvania, with notes
and descriptions. Trans. Amer. Entomol. Soc. 22:317-381 (1895).

Hamilton, W. D. and R. M. May. 1977. Dispersal in stable habitats. Nature 269:578-581.

Hammond, P. M. 1979. Wing-folding mechanisms of beetles, with special reference to in-
vestigations of Adephagan phylogeny (Coleo.). Pages 1 13-180 in: T. L. Erwin, G. E. Ball
and D. R. Whitehead (eds.), Carabid Beetles: Their Evolution, Natural History, and
Classification. W. Junk, The Hague, x + 635 pp.

Hammond, P. M. 1985. Dimorphism of wings, wing-folding and wing-toiletry devices in the
ladybird, Rhyzobius litura (F.) (Coleoptera: Coccinellidae), with a discussion of inter-
population variation in this and other wing-dimorphic beetle species. Biol. J. Linn. Soc.
24:15-33.

Harrison, R. G. 1980. Dispersal polymorphism in insects. Ann. Rev. Ecol. Syst. 1 1:95-1 18.

Hatch, M. H. 1957. The beetles of the Pacific Northwest. Part II: Staphyliniformia. Univ.
Wash. Publ., Biol. 16:1-384 + 9 pp.

Hille Ris Lambers, D. 1966. Polymorphism in Aphididae. Ann. Rev. Entomol. 1 1:47-78.

Honek, A. 1976. Factors influencing the wing polymorphism in Pyrrhocoris apterus (Heter-
optera, Pyrrhocoridae). Zool. Jahrb., Abt. Syst., Okol., Geogr. Tiere 103:1-22.

Jackson, D. J. 1928. The inheritance of long and short wings in the weevil, Sitona hispidula,
with a discussion of wing reduction among beetles. Trans. Roy. Entomol. Soc. Edinburgh
55:665-735.

Jackson, D. J. 1956. Observations of flying and flightless water beetles. J. Linn. Soc. London,
Zool. 43:18-42.

Jeannel, R. 1926. Mongraphie des Trechinae. L’Abeille 32:221-550.

Jeannel, R. 1943. Les Fossiles Vivant des Cavemes. Librairie Gallimard, Paris, 321 pp.

Kalmus, H. 1 945. Correlations between flight and vision, and particularly between wings and
ocelli, in insects. Proc. Roy. Entomol. Soc. London (A)20:84-96.

Kuchler, A. W. 1964. Potential natural vegetation of the conterminous United States. Amer-
ican Geographical Society, Special Publication No. 36, v + 39 + 110 pp., map.

Langor, D. W. and D. J. Larson. 1984. Alary polymorphism and life history of a colonizing
ground beetle, Bembidion lampros Herbst. Coleopt. Bull. 37:365-377.

Lees, A. D. 1966. The control of polymorphism in aphids. Adv. Insect Physiol. 3:207-277.

Liebherr, J. K. 1989. Tanystoma diabolica new species (Coleoptera: Carabidae: Platynini)
from Baja California and its biogeographic significance. J. New York Entomol. Soc. 97:
173-186.

564

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Liebherr, J. K. and A. E. Hajek. 1986. Geographic variation in flight wing development and
body size of the Tule Beetle, Tanystoma maculicolle (Coleoptera: Carabidae). Pan-Pac.
Entomol. 62:13-22.

Lindroth, C. H. 1949. Die Fennoskandischen Carabidae. Eine tiergeographische Studie. III.
Allgemeiner Teil. Goteb. K. Vetensk. Vitterhets-Samh. Handl. 6, Ser. B, 4(3): 1-9 11.

Lindroth, C. H. 1979. The theory of glacial refugia. Pages 385-394 in: T. L. Erwin, G. E. Ball
and D. R. Whitehead (eds.), Carabid Beetles: Their Evolution, Natural History, and
Classification. W. Junk, The Hague, x + 635 pp.

Lohse, G. A. 1964. Fam. Staphylinidae I. (Micropeplinae bis Tachyporinae). H. Freude, K.
W. Harde and G. A. Lohse (eds.), Die Kafer Mitteleuropas. Vol. 4. Goecke and Evers,
Krefeld, 263 pp.

Lyal, C. H. C. 1985. Phylogeny and classification of the Psocodea, with particular reference
to the lice (Psocodea: Phthiraptera). Syst. Entomol. 10:145-165.

MacKay, P. A., D. J. Reeleder and R. J. Lamb. 1983. Sexual morph production by apterous
and alate viviparous Acyrthosiphon pisum (Harris) (Homoptera: Aphididae). Can. J. Zool.
61:952-957.

Matthews, J. V. 1979. Tertiary and Quaternary environments: historical background for an
analysis of the Canadian insect fauna. Pages 31-86 in: H. V. Danks (ed.), Canada and
Its Insect Fauna. Mem. Entomol. Soc. Can. No. 108, 573 pp.

Moore, 1. and E. F. Legner. 1979. An illustrated guide to the genera of the Staphylinidae of
America north of Mexico exclusive of the Aleocharinae (Coleoptera). Division of Ag-
ricultural Sciences, University of California Priced Publication 4093, 332 pp.

Morgan, A. V. and A. Morgan. 1981 (1980). Faunal assemblages and distributional shifts of
Coleoptera during the Late Pleistocene in Canada and the northern United States. Can.
Entomol. 112:1 105-1 128.

Newton, A. F., Jr. and M. K. Thayer. 1992. Current classification and family-group names
in Staphyliniformia (Coleoptera). Fieldiana: Zool., N. S., No. 67, 92 pp.

Paulian, R. 1988. Biologie des Coleopteres. Editions Lechevalier, Paris, xi + 719 pp.

Peck, S. B. 1978. New records and species of Leiodinae and Catopinae (Coleoptera: Leiodidae)
from Jamaica and Puerto Rico, with a discussion of wing dimorphism. Psyche 84:243-
254 (1977).

Pope, R. D. 1977. Brachyptery and wing-polymorphism among the Coccinellidae. Syst. En-
tomol. 2:59-66.

Regal, P. J. 1977. Evolutionary loss of useless features: is it molecular noise suppression?
Amer. Nat. 1 1 1:123-133.

Rolf, D. A. 1986a. The evolution of wing dimorphism in insects. Evolution 40:1009-1020.

Rolf, D. A. 1986b. Evolution of wing polymorphism and its impact on life cycle adaptation
in insects. Pages 204-221 in: F. Taylor and R. Karban (eds.), The Evolution of Insect
Life Cycles. Springer- Verlag, New York, x + 287 pp.

Rolf, D. A. 1 990a. Antagonistic pleiotropy and the evolution of wing dimorphism in the sand
cricket, Gryllus firmus. Heredity 65:169-177.

Rolf, D. A. 1990b. The evolution of flightlessness in insects. Ecol. Monogr. 60:389-421.

Rolf, D. A. and D. J. Fairbairn. 1991. Wing dimorphisms and the evolution of migratory
polymorphisms among the Insecta. Amer. Zool. 31:243-251.

Ross, E. S. 1970. Biosystematics of the Embioptera. Ann. Rev. Entomol. 15:157-172.

Riischkamp, F. 1927. Der Flugapparat der Kafer. Zoologica 28(3/4):vi + 1-88, pis. 1-8.

Scholtz, C. H. 1981. Aptery in Trox (Coleoptera: Trogidae): morphological changes and their
relationship to habitat. J. Entomol. Soc. So. Afr. 44:83-87.

Shute, S. L. 1980. Wing-polymorphism in British species of Longitarsus beetles (Chryso-
melidae: Alticinae). Syst. Entomol. 5:437-448.

Slater, J. A. 1975. On the biology and zoogeography of Australian Lygaeidae (Hemiptera:

1992

SEXUAL WING DIMORPHISM

565

Heteroptera) with special reference to the southwest fauna. J. Aust. Entomol. Soc. 14:
47-64.

Slater, J. A. 1977. The incidence and evolutionary significance of wing polymorphism in
lygaeid bugs with particular reference to those of South Africa. Biotropica 9:217-229.

Smetana, A. 1981. A new species of Phloeonomus Heer from the Kuril Islands (Coleoptera:
Staphylinidae). Entomol. Scand. 12:78-80.

Smetana, A. 1985. Systematic position and review of Deinopteroloma Jansson, 1946, with
descriptions of four new species (Coleoptera, Silphidae and Staphylinidae (Omaliinae)).
Syst. Entomol. 10:471-499.

Southwood, T. R. E. 1961. A hormonal theory of the mechanism of wing polymorphism.
Proc. Roy. Entomol. Soc. London (A)36:63-66.

Southwood, T. R. E. 1962. Migration of terrestrial arthropods in relation to habitat. Biol.
Rev. Camb. Philos. Soc. 37:171-214.

Southwood, T. R. E. and D. Leston. 1 959. Land and Water Bugs of the British Isles. Frederick
Wame, London, xi + 436 pp.

Steel, W. O. 1957. Notes on the Omaliinae. (8). The genus Acrolocha Thomson. Entomol.
Mon. Mag. 93:157-164.

Steel, W. O. 1960. Three new Omaliine genera from Asia and Australia previously confused
with Phloeonomus Thomson. Trans. Roy. Entomol. Soc. London 112:141-172.

Steel, W. O. 1964. Insects of Campbell Island. Coleoptera: Staphylinidae. Pac. Ins. Monogr.
7:340-375.

Steffan, W. A. 1966. A generic revision of the family Sciaridae (Diptera) of America north
of Mexico. Univ. California Publ., Entomol. 44:1-77.

Stelfan, W. A. 1973. Polymorphism in Plastosciara perniciosa. Science 182:1265-1266.

Steffan, W. A. 1975. Morphological and behavioral polymorphism in Plastosciara perniciosa
(Diptera: Sciaridae). Proc. Entomol. Soc. Wash. 77:1-14.

Taylor, V. A. 1978. A winged elite in a subcortical beetle as a model for a prototermite.
Nature 276:73-75.

Taylor, V. A. 1981. The adaptive and evolutionary significance of wing polymorphism and
parthenogenesis in Ptinella Motschulsky (Coleoptera: Ptiliidae). Ecol. Entomol. 6:89-98.

Thayer, M. K. 1985. Revision, phylogeny, and biogeography of the austral genus Metacor-
neolabium Steel (Coleoptera: Staphylinidae: Omaliinae). Pages 113-179 in: G. E. Ball
(ed.), Taxonomy, Phylogeny, and Zoogeography of Beetles and Ants. Series Entomologica
33. W. Junk, Dordrecht, xiv + 514 pp.

Thayer, M. K. 1987. Biology and phylogenetic relationships of Neophonus bruchi, an anom-
alous south Andean staphylinid (Coleoptera). Syst. Entomol. 12:389-404.

Thayer, M. K. and A. F. Newton, Jr. 1979. Revision of the south temperate genus Glypholoma
Jeannel, with four new species (Coleoptera: Staphylinidae: Omaliinae). Psyche 85:25-
63 (1978).

Van Dyke, E. C. 1933. Peculiarities of the coleopterous fauna of semi-arid southwestern North
America. Ve Congr. Int. Entomol., Paris, 1932, 2:471^477.

Venturi, F. 1964. Saggio sui Licoridi (Insecta, Diptera) brachitteri ed atteri europei, con
particolare reguardo alle forme sicula. Atti Accad. Gioenia Sci. Nat. Catania 16:87-1 14.

Vepsalainen, K. 1978. Wing dimorphism and diapause in Gerris : determination and adaptive
significance. Pages 218-253 in: H. Dingle (ed.), Evolution of Insect Migration and Dia-
pause. Springer- Verlag, Berlin, xvi + 284 pp.

Watanabe, Y. 1 990. A taxonomic study on the subfamily Omaliinae from Japan. Mem. Tokyo
Univ. Agric. 31:57-391.

Wigglesworth, V. B. 1961. Insect polymorphism— a tentative synthesis. Symp. Roy. Entomol.
Soc. London 1:103-113.

Wollaston, T. V. 1854. Insecta Maderensia, Being an Account of the Islands of the Madeiran
Group. Van Voorst, London, 634 pp.

566

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Zamenhof, S. and H. Eichhom. 1 967. Study of microbial evolution through loss of biosynthetic
functions: establishment of ‘defective’ mutants. Nature 216:456-458.

Zanetti, A. 1987. Fauna d’ltalia. Coleoptera: Staphylinidae: Omaliinae. Edizioni Calderini,
Bologna, xii + 472 pp.

APPENDIX A

Redescription of “Omalium” flavidum Hamilton
and Lectotype Designation

Redescription. Yellowish- or reddish-brown to dark brown, varying among morphs
(see Table 1). Body sparsely punctate and sparsely setose (Figs. 4, 5); dorsal surfaces
of head (except neck), pronotum, and elytra without microsculpture; abdomen with
reticulate microsculpture of varying distinctness.

Head (Figs. 8, 9) with pair of ocelli on vertex, often concolorous with surrounding
cuticle (thus difficult to see); head width 1.2-1. 3 x head length; distinct neck present;
frontal area elevated, delimited posteriorly by oblique ridges running from behind
antennal bases to just in front of ocelli; eyes well-developed (somewhat reduced in
— W<5 and 9), with seta-fringed ridge posterior to each eye (Figs. 8, 9); cuticle above
and behind eye with curved ridges roughly paralleling margin of eye; gular sutures
close but not confluent between eyes, diverging posteriorly. Labral apex arcuately
emarginate; epipharynx with sensilla and setae arranged as in Figure 14. Mandibles
apically acute, asymmetrical, each with well-developed articulated mola bearing ver-
tical ridges on adoral surface; ventral surface of each mandible near middle with
band of mostly apically bifid setae running from anterolateral to posteromedial; right
mandible with a single sharp preapical tooth, left with a blade-like preapical area.
Galea (Fig. 1 3) with densely setose apex; lacinia with medial comb of spine-like setae
and unarticulated apical spine (Fig. 13); maxillary palp with 4 articles, all but first
subequal in maximum width, ultimate about 2.5 x as long as penultimate (Figs. 8,
9). Labial palp of 3 articles subequal in width, the ultimate longest, penultimate
shortest. Antenna 1 1 -segmented, without distinct club, longer than head and pro-
notum together (length varying among morphs: see Tables 1, 2); with dense long
blunt sensilla on antennomeres 3-1 1 in 3, 5-11 only (and concentrated at apices) in
9; +W<3 with all antennomeres longer than wide, 9 and — W 6 with antennomeres 1-
3 and 1 1 longer than wide, 6-10 wider than long, and 4-5 quadrate or slightly wider
than long; antennomere 3 slightly wider in — W<3 than in 9.

Pronotum (Figs. 4, 5) slightly transverse, width about 1.25 x length, widest slightly
anterior to middle, narrowed slightly sinuately from there toward base; lateral margins
narrowly explanate; disc with raised roughly triangular median area enclosing lon-
gitudinal impressions on either side of midline.

Elytra of +W<3 loosely interlocked, with arcuate apices, length about 1.5 times
conjoint width and nearly 3 times as long as pronotum, covering part or all of
abdominal tergite 5, humeri prominent; of 9 and — W<5 very tightly interlocked (though
not fused), with truncate apices, about as long as their conjoint width and about 1.5
times as long as pronotum, only covering tergite 3, humeri reduced and rounded.
Mesosternum not carinate. Metathorax of + W3 (Fig. 6) normal in size (Fig. 2a, Table
1) and sclerotization, normal flight muscles present, metendosternite as in Figure

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567

Figs. 8-13. Omalium flavidum Hamilton. 8. Head, dorsal view, +W3. 9. Same, — W3. 10.
Median area of abdominal tergite 7 apex, +W<3. 11. Same, — W<$. 12. Detail of 11. 13. Apex
of left galea, +W<3, dorsal view. (Scale line: Figs. 8-1 1, 100 /tm; Figs. 12-13, 10 /mi.)

15a; metathorax of 9 and — W 6 (Fig. 7) reduced in length (Fig. 2a, Table 1), width,
and dorsal sclerotization, without flight muscles, metendostemite as in Figure 1 5b.
Wings normally developed ( + W<3, Fig. 16a) or minute (9, — W S, Fig. 16b).

Tarsi 5-segmented; empodium bisetose, one seta much longer than other (Fig. 19);

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Figs. 14-16. Omalium flavidum Hamilton. Scale lines = 0.1 mm, except in Fig. 16a, 1 mm.
14. Epipharynx, 9, ventral view. 15. Metendostemite, dorsal view, (a) + W <3; (b) — W<5. Right
wing, dorsal view, (a) +W3; (b) — W<3.

+ W 6 with hind tarsus (Figs. 2b, 1 7) about two-thirds as long as hind tibia, tarsomeres
1-4 together longer than 5 (Fig. 2c, Table 1), 9 and — W<3 with hind tarsus (Figs. 2b,
18) no more than 0.6 as long as hind tibia, tarsomeres 1-4 together shorter than 5
(Fig. 2c, Table 1).

Abdomen with spiracles on segments 1-8, located in tergite of segments 3-7, in
membrane adjacent to tergite of segments 1-2 and 8; spiracles 1-3 distinctly (9, + W<3)
or slightly ( — W<$) larger than following ones; one pair of paratergites on each of
segments 3-7; intersegmental membranes connecting segments 3-7 bearing brick-

Figs. 17-20. Omalium flavidum Hamilton, tarsi. 17. Hind tarsus, +W<3, dorsal view. 18.
Same, — WS, oblique ventral view. 19. Detail of 18, showing unequal empodial setae. 20. Front
tarsus, — W<5, dorsal view. (Scale line: Figs. 17-18, 20, 100 yum; Fig. 19, 25 yum.)

Figs. 21-24. Omalium flavidum Hamilton, abdominal tergites. 21. Tergite 3, with arrow
indicating vestigial row of setae near apex, +W 6. 22. Same, — W<5. 23. Tergites 4 and 5, +W8.
24. Same, — W<1 (Scale line: Figs. 21-22, 100 yum; Figs. 23-24, 200 yum.)

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■0? |

25

Figs. 25, 26. Omalium flavidum Hamilton, abdominal sternites. 25. Sternites 7 and 8, — W<3,
ventral view. 26. Anterior median projection of stemite 8 with associated gland tubules, +W<3,
ventral view. (Scale line: 0.1 mm.)

wall pattern of sclerites (Figs. 21-24); tergite 3 posteriorly with vestiges of transverse
seta-edged groove (Figs. 21, 22); sternite 3 with distinct hind coxal impressions
delimited by a ridge; wing-folding patches present on tergites 4-5 of +W<3 only (Fig.
24, cf. Fig. 23); tergite 7 with (±W<3, Figs. 10-12) or without (2) apical palisade fringe;
sternite 8 with anterior projection bearing ducts of defensive gland cells (Fig. 26).

Male. Front tarsus with tenent setae on tarsomeres 1-4 (Fig. 20). Sternites 7 and
8 with characteristic arrangement of short stout setae apically (Fig. 25); genital seg-
ment (Fig. 27) with tergite 9 continuous across dorsum. Parameres of aedeagus dorsal
when aedeagus is retracted within abdomen; aedeagus as in Fig. 28, with small th«n
basal piece, median lobe with median desclerotized band at base and on abparameral
side, and internal sac with complex armature of small spines and sclerotized plates.

Female. External genitalia (Fig. 29) lacking stemite 9; tergite 9 divided dorsally;
tergite 10, two pairs of gonocoxites, and one pair of styli present; lightly sclerotized
structure present internally, extending anteriorly from tergites 9 and 10 when genitalia
are extended, including a flat portion in median sagittal plane and small pale oval
“window” in horizontal plane on each side of it, near anterior end (Fig. 29, broken
lines). Spermatheca (Fig. 30) lightly sclerotized, two-chambered.

Lectotype designation. Hamilton (1896) stated that “several examples occurred
near St. Vincent.” His collection is at the Carnegie Museum, which houses two
apparent syntypes of O. flavidum. I here designate as lectotype the intact female,
bearing the labels: St. Vine. Penn./type/Carn. Mus. Ace. 349/[b!ank red label]/LEC-
TOTYPE Omalium flavidum Hamilton des. M. K. Thayer 1991. I designate the
male ( — W morph, abdomen missing) a paralectotype; its labels read: St. Vine. Penn./
Carn. Mus. Ace. 349/PARALECTOTYPE Omalium flavidum Hamilton des. M. K.
Thayer 1991.

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APPENDIX B

Material Examined (all 9 are — W)

Canada: ONTARIO: 2 3 ( — W), 39, Chaffeys Locks, birch-maple litter beside logs,
23.x. 1985 (A. Davies) (CNC); 13 (+W), Kemptville, intercept, 25.x-8.xi. 1 983 (L.
Dumouchel and J. Denis) (CNC); QUEBEC: 13 (-W), Johnville, TP-066, 4.V.1988
(C. Levesque) (CNC). United States: DISTRICT OF COLUMBIA: [no locality], 13
(-W) (CM); MAINE: Androscoggin Co.: \6 ( + W), Poland Spring, 27.xi.1909 (JHE)
(MCZ); York Co.: 5 <3 (+W), West Lebanon, 20-29.xi.90, FIT, disturbed forest (DW
Barry) (UNH); 3 6 ( + W), West Lebanon, FIT, disturbed forest, 30.xi-5.xii.90 (DW
Barry) (UNH); 203 ( + W), West Lebanon, FIT, disturbed forest, 1 l-17.xii.9Q (DW
Barry) (UNH); 163 ( + W), West Lebanon, FIT, disturbed forest, 18-26.xii.90 (DW
Barry) (UNH); 13 ( + W), West Lebanon, FIT, disturbed forest, 27.xii.90-7.i.91 (DW
Barry) (UNH); 23 ( + W), West Lebanon, FIT, disturbed forest, 6-12.ii.9 1 (DW Barry)
(UNH); 23 ( + W), West Lebanon, FIT, disturbed forest, 15— 24.iii.91 (DW Barry)
(UNH); 13 ( + W), West Lebanon, FIT, disturbed forest, 25— 31.iii.91 (DW Barry)
(UNH); MARYLAND: Prince Georges Co.: 13 ( + W), Takoma Pk., 3 mi E, flying,
1 7.xii. 1949 (G. H. Nelson) (GHN); MASSACHUSETTS: Hampshire Co.: 13 (-W),
Northampton, 14.xi. 19 17 (CNC); Middlesex Co.: 19, Framingham, sifting, 17.xi.1934
(C. A. Frost) (MCZ); 33 (2 +W, 1 -W), 19, Tyngsboro, 17.xi. 1901 (MCZ); 19,
Tyngsboro, 3.xi. 1915 (MCZ); 13 ( — W), Tyngsboro, sifting leaves, 7.xi. 1924 (MCZ);
NEW HAMPSHIRE: Carroll Co.: 33 ( — W), 19, Passaconaway Cpgd., spruce-fir-pine-
hdwd., 340 m, leaf and rain-washed litter, berl., 1 7-1 8.x. 1 973 (A. Newton) (ANMT);
19, Passaconaway Cpgd., spruce-fir-pine-hdwd., 340 m, leaf and rain-washed litter,
berk, 15.x. 1978 (A. Newton and M. Thayer) (ANMT); 13 ( + W), The Bowl, 2.5 mi
NW Wonalancet, 590 m, FIT, 20.x-7.xi. 1984 (D. S. Chandler) (UNH); 13 (-W),
The Bowl, 2.5 mi NW Wonalancet, 590 m, sift birch log, l.xi.1984 (D.S. Chandler)
(CNC); 19, The Bowl, 2.5 mi NW Wonalancet, 590 m, sift conifer logs, 17.x. 1985
(D. S. Chandler) (UNH); 13 (-W), The Bowl, 2.5 mi NW Wonalancet, 590 m, sift
birch logs, 3 1.x. 1985 (D. S. Chandler) (UNH); 30 3 (28 + W, 2 -W), The Bowl, 2.5
mi NW Wonalancet, 590 m, FIT, 1— 19.xi. 1985 (D. S. Chandler) (UNH); 33 (2 +W,
1 — W), The Bowl, 2.5 mi NW Wonalancet, 590 m, on snow, 19.xi.1985 (D. S.
Chandler) (UNH; CNC); Coos Co.: 13 ( — W), Mt. Washington toll rd, 0.3 mi below
Halfway Hse., spruce-fir-birch for., 1 100 m, litter, berk, 15.x. 1978 (A. Newton and
M. Thayer) (ANMT); Grafton Co.: 19, Franconia, 26.xii. 1960 (K. W. Cooper) (USNM);
19, Hubbard Brook Exp. For., Bear Brook, 460 m, sift rotten wood, 15.x. 1982 (D.
S. Chandler) (UNH); 33 ( — W), 39, Kancamagus Pass, spruce-fir-birch, 850 m, litter,
berk, 19.x. 1973 (A. Newton) (ANMT); 19, Kancamagus Pass, spruce-fir-birch, 850
m, litter, berk, 15.x. 1978 (A. Newton and M. Thayer) (ANMT); Strafford Co.: 53
( + W), Durham, 1 mi SW, FIT, 15.x-4.xi.1987 (D. S. Chandler) (UNH); 23 (1 +W,
1 -W), Durham, 1 mi SW, FIT, 18.xi-4.xii. 1987 (D. S. Chandler) (UNH); 153 (14
+W, 1 — W), Spruce Hole, 3 mi SW Durham, FIT, 5-24. xk 1 987 (D. S. Chandler)
(UNH; CNC); 63 (+W), Spruce Hole, 3 mi SW Durham, FIT, 4-15.xii.1987 (D. S.
Chandler) (UNH); NEW YORK: St. Lawrence Co.: 19, Wanakena [as Wananeka],
15.x. 1982 (Lee Herman) (AMNH); Tompkins Co.: 13 ( + W), Caroline, on snow,
12.iii. 1963 (M. A. Deyrup) (AMNH); 13 ( + W), Ithaca, 10.iv.1926 (Fletsher) (CU);
19, Jacksonville, N, maple + forest, dead maple buttress, FMHD #82-49, 24. v. 1982
(W. S. Suter) (FMNH); PENNSYLVANIA: 19, [state only] (CM); Philadelphia Co.:

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16 (— W), Manayunk [as Manyunk], 5.x (J. W. Green colln.) (CAS); Schuylkill Co.:
1<5 (+W), Pottsville, forest, trap #203, 1-1 l.iv. 1967 (S. Peck) (FMNH); Westmore-
land Co.: 19, Jeannette, x (H. G. Klages) (CM); 19, Jeannette, xii (H. G. Klages) (CM);
19, Jeannette, “VIII. 16” (H. G. Klages) (CM); 1<3 (+W), Powdermill Nature Res., nr
Rector, cabins area on snow, 25.ii.1958 (CM; CM Ace. 18464); 1<5 ( — W) (Paralec-
totype), 19 (Lectotype), St. Vincent [as St. Vine.] (CM).

Received 4 November 1991; accepted 1 1 February 1992.

Figs. 27-30. Omalium flavidum Hamilton. Scale lines = 0.1 mm. 27. Male genital segment
(segments 9-10), +W. (a) dorsal view; (b) ventral view. 28. Aedeagus. (a) — W<3, parameral view
(dorsal in repose); (b) +W<3, lateral view. 29. Female external genitalia, anterior broken lines
indicating lightly sclerotized internal sclerite. (a) dorsal view; (b) ventral view. 30. Spermatheca
(spermathecal gland attached to subapical duct not shown).

J. New York Entomol. Soc. 100(4): 5 74-5 80, 1992

TWO NEW SPECIES OF DIPLOT AXIS FROM MEXICO
(COLEOPTERA: MELOLONTHIDAE)

Leonardo Delgado1 and Fabricio Capistran2

^Institute) de Ecologia, A.C., Apdo. Postal 63,

91000 Xalapa, Veracruz, Mexico, and
2Parque de la Flora y Fauna Silvestre Tropical,

Apdo. Postal 57, Catemaco, Veracruz, Mexico

Abstract.— Two new Mexican species of Diplotaxis from Veracruz state are described. Mod-
ifications to Vaurie’s keys to the genus are included to allow their identification.

The American genus Diplotaxis includes 221 species, most of which (about 77%)
are distributed in Mexico (Vaurie, 1958, 1960, 1962; Delgado-Castillo, 1990). How-
ever, only 18 species have been cited from Veracruz state: six are localized in the
lowland tropical rain forest with four of these six extending their distribution to
Central America; five others occur principally in areas above 1,500 m; two others
are distributed almost exclusively in the cloud forest (one of which, D. veracruzana
Vaurie, is a Veracruz endemic); and five others with various habitat preferences ( D .
aenea Blanchard, D. angustula Moser, D. atramentaria Bates, D. coriacea Bates and
D. puberea Bates) seem to be very scarce in Veracruz and need to be confirmed.

In this paper we describe two new species from Veracruz state, and incorporate
them into Vaurie’s keys.

Diplotaxis xalapensis, new species
(Figs. 1-2)

Type material. Holotype 6, labeled: “MEXICO, Veracruz, Xalapa, Rancho Gua-
dalupe, Km 2.5 Antigua carr. a Coatepec. Bosque mesofilo-cafetal. Alt. 1,360 m, Luz.
13-11-1991. L. Delgado, col.” Allotype 9 labeled as holotype, except: 6-7-III- 1992,
F. Capistran, J.R. Hernandez y L. Delgado, cols. Paratypes 54 66, 29 99. One male
labeled as holotype. 32 66, 21 99 labeled as allotype. 17 66, 8 99 labeled as holotype
except: 8-III-1991 (16), 14-01-1991, L. Arellano, col. (466); 8-IV-1991 (16); 10-III-
92, P. Rojas, col. (16); 20-111-92, F. Capistran y E. Santos, Cols. (1066, 799). 4 66
labeled: “MEXICO, Veracruz, Xalapa, Briones, Alt. 1,360 m. 28-III- 1 99 1 . M.A.
Moron, col.” One female labeled Mexico: Veracruz, 2 km al NE de Calcahualco.
AH. 1,650 m Bosque mesofilo 4-V-92 a la luz, Capistran y Delgado, cols.

Holotype and four paratypes deposited at the Institute de Biologia-Universidad
Nacional Autonoma de Mexico (Mexico City), paratypes deposited at the following
collections: British Museum Natural History (London), Museum National d’Histoire
Naturelle (Paris), Zoologisches Museum Humboldt Universitat (Berlin), Canadian
National Collection (Ontario), United States National Museum (Washington), Amer-
ican Museum of Natural History (New York), Museu de Zoologia da Universidade
de Sao Paulo, Museu Nacional da Universidade Federal do Rio de Janeiro, Universi-
dad Nacional de Venezuela (Maracay), Museo Argentine de Ciencias Naturales (Bue-
nos Aires), Laboratorio Natural Las Joyas-Universidad de Guadalajara (Jalisco),

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NEW D I PLOT AX IS

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Figs. 1-4. New species of Diplotaxis. 1-2. D. xalapensis 1. Parameres, frontal
view. 2. Parameres, lateral view. 3-4. D. squamisetis. 3. Parameres, frontal view. 4.
Parameres, lateral view. Scale = 1 mm.

Museo de Historia Natural de la Ciudad de Mexico, Instituto de Ecologia (Veracruz),
M. Bunalski (Poznan), H. & A. Howden (Ontario), B. D. Gill (Ontario), B. C. Ratcliffe
(Nebraska), W. B. Warner (Arizona), S. McCleve (Arizona), P. K. Lago (Mississippi),
A. A. De Ascencao (Merida), A. Martinez (Salta), G. Halffter (Veracruz), M. A.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Moron (Veracruz), J. Blackaller (Mexico City), J. R. Hernandez (Veracruz), F. Cap-
istran (Veracruz) and L. Delgado (Mexico City).

Description. Holotype male. Length 7.2 mm; elytral maximum width 3.0 mm.
Body elongate and flattened; front of head, disc of pronotum, abdomen and pygidium
metallic green; elytra opaque green, sericeous; clypeus, marginal regions of pronotum
and scutellum brilliant reddish; venter and legs yellowish-red. Clypeus trapezoidal,
as long as the front, with obtuse front angles, broadly emarginate anteriorly and sides
indented in front of eyes; surface deeply concave, except for a tumid small central
area; some short setae present only behind the margins and with scarce, minute setae
on disc; fronto-clypeal suture well marked and sinuate at middle; clypeal surface
densely rugose. Front with anterior half abruptly descending to clypeus, with scarce
medium-sized punctures and concave at central region, posterior half flattened and
with rugose punctation; transverse eye diameter almost 0.25 x as wide as head.
Antenna 10-jointed. Labrum slightly concave with dense punctures, level with and
0.75 x the length of the reflexed underside of clypeus; mandibles slender; mentum
slightly convex, with anterior declivity marked by transverse sharp, setiferous ridge;
last article of maxillary palpus not impressed dorsally.

Pronotum hexagonal; fore angles acute, lateral angles obtuse and situated nearly
at the middle, hind angles obtuse; pronotal surface with longitudinal mesial furrow
and four irregular depressions each side of midline as follows; one on anterolateral
corner, one lateral, one on posterolateral corner and one central, last two larger than
the others; pronotal surface irregularly elevated with large punctures, some of them
confluent, the punctation between them becoming finely reticulate to marginal regions;
pronotum laterally and basally with cariniform, narrow margin, with the adjacent
region depressed, basal margin crenate. Scutellum with scarce medium-sized punc-
tures.

Elytra longer than wide (1:0.66), with irregular rugose swellings, many of them
branching off from convex costae; punctures smaller than those on pronotum, irreg-
ularly placed and with scarce, minute setae overall (only visible at high magnification);
striae not evident, except the two outer ones; marginal setae scarce and very short,
present only on basal quarter. Abdomen laterally rounded; propygidium not grooved
above pygidium; abdominal sternites 1-5 subequal in length, with long setae more
abundant laterally; pygidium wider than long and with coarse and deep punctures,
some confluent, and with setae more or less abundant, those on apical half longer.

Protibiae tridentate, basal tooth scarcely marked and situated in the distal half;
claws slightly curved, cleft subapically, tooth entire and shorter than apex; tarsi longer
than respective tibiae; basal mesotarsomere almost as long as second one; metacoxal
plates margined laterally and truncate; metafemora straight and slender; metatibial
spurs slender, long and acute; basal metatarsomere shorter than the second one and
of the same length as the longest spur.

Genitalia with basal piece a little longer than parameres, which are jointed on
inner margin at basal third and abruptly bent at the apex (Figs. 1-2).

Allotype female. Length 7.5 mm; elytral maximum width 3.2 mm. Differs from
holotype in the following respects: elytra with metallic reflections; clypeus with only
the marginal regions yellowish-red, a little shorter than the front, scarcely reflexed,
the anterior margin truncate and the front angles rounded, half posterior of the front
a little convex and with punctures not confluent; transverse eye diameter 0.20 x as

1992

NEW DI PLOT AX IS

577

wide as head; labrum as long as the underside of clypeus; last article of maxillary
palpus shorter; pronotum with fore angles almost right, with the surface almost evenly
elevated, with punctures not confluent and the reticulated punctation reduced; fifth
abdominal sternite shorter than each stemites 1-4; pygidium with coarser and deeper
punctures almost crater-like; femora wider and robust; protibia wider and with the
basal tooth clearly marked, meso and metatibiae more flared out at apex; metatibial
spurs wider; protarsi shorter; basal metatarsomere shorter than the longest spur;
claws shorter, tooth of the same length as the apex, obliquely truncate and more
separated from the apex.

Variation. Males, length; 5. 8-7. 7 mm, elytral maximum width; 2. 6-3. 5 mm; fe-
males, length 6. 1-7.7 mm, elytral maximum width 2. 7-3. 6 mm. In the males the
color of elytra is generally sericeous green; and in the females metallic-green; the
brilliant reddish color on the marginal regions of clypeus and pronotum varies a little
in extent. Setae on clypeus, scutellum and elytra varies from more or less abundant
to nearly absent; pygidial punctures vary in density and depth; length of the basal
metatarsomere varies from a little longer to shorter than the longest metatibial spur.
The shape of clypeus, the transverse eye diameter, the length of labrum, the last
article of palpus maxillary, the pronotal fore angles and the legs are considered
dimorphic characters.

Type locality. Km 2.5 Antigua carr. a Coatepec, Rancho Guadalupe, Xalapa, Ve-
racruz, Mexico.

Remarks. D. xalapensis presents most of characters given by Vaurie (1960:191)
to the “Trapezifera” group as: clypeus hairy (only behind front edge), eye large,
antennae 10-jointed, mandibles small, elytra about three times longer than the pro-
notum, abdomen not ridged laterally, propygidium without groove, metacoxal plates
margined laterally and not angulate, and basal tooth of protibiae in the distal half.
However D. xalapensis shows characters not shared with other species of this group,
such as the unevenly elevated dorsal surface, the hexagonal shape of pronotum, and
the elytral punctation and microsculpture with a metallic or sericeous cast, which
give it an unusual and distinctive appearance, and the shape of the claws.

A species that shows the same elytral punctation and microsculpture is D. aurata
Bates, which differs from D. xalapensis by the labrum longer than underside of
clypeus, the pronotum arcuate near middle, and the elytra with long, erect setae. We
key out D. xalapensis in the “trapezifera” group for convenience, however, this species
seems not to have close affinities to this group. Rather it shows possible relationships
with D. aurata, a species only cited from Guatemala.

D. xalapensis is an early flying species, and this phenology could explain why it
has not been collected before, since the area has been visited by many entomologists
over a long time.

Etymology. This species is dedicated to the City of Xalapa de Enriquez, where it
was collected, and one of the classical localities of the faunal work “Biologia Centrali-
Americana.”

The key to the “trapezifera” group proposed by Vaurie (1960:193), is modified as
follows to incorporate D. xalapensis :

27 Pronotum with sides strongly arcuate behind middle and hind angles rounded off
without angulation; side margins of clypeus almost parallel to front; Guerrero, Puebla

incisa

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

27' Pronotum with sides scarcely arcuate or angulate, and hind angles distinctly angulate;
side margins ofclypeus converging obliquely to front; Durango, (Hidalgo?), Veracruz

27a

27a Color wholly light red; clypeus with dentiform front angles; elytral striae clearly

distinguished; Durango, (Hidalgo?) saltensis

27a' Color of the elytra and the most of the pronotum and front green, marginal regions
of clypeus reddish; clypeus with obtuse or rounded front angles; striae of the elytral
disc not distinguished; Veracruz xalapensis

Olplotaxis squamisetis, new species
(Figs. 3-4)

Type material. Holotype 6, labeled: “MEXICO: Veracruz, Los Tuxtlas, Sierra de
Santa Marta, Ocotal Chico. 17-1 8- VII- 1982. Alt. 700 m, colecta nocturna. H. Perez,
col.” Allotype 9, labeled: “MEXICO: Veracruz, Catemaco, Pipiapan, Parque de la
Flora y Fauna Silvestre Tropical. 16-IV-91. Alt. 600 m, selva mediana perennifolia/
comiendo hojas de Guarea glabra (Meliaceae). O. Aquino y F. Capistran, cols.”
Paratypes 6<3<3, 19. 2 66 labeled as holotype; 1 9 labeled as allotype, except: “13-15-
IV- 1 99 1 . Luz ultravioleta. G. Cabrera y F. Capistran, cols”; 4 66 labeled: “MEXICO:
Veracruz, Estacion Biologia “Los Tuxtlas,” 250', V-20-1983/ C.W. & L. O’Brien &
G.B. Marshall.”

Holotype and allotype deposited at the Institute de Biologia, Universidad Nacional
Autonoma de Mexico (Mexico City). Paratypes deposited at the following collections:
W. B. Warner (Arizona), S. McCleve (Arizona), M. A. Moron (Veracruz), F. Capistran
(Veracruz) and L. Delgado (Mexico City).

Description. Holotype male. Length 9.3 mm, elytral maximum width 4.6 mm.
Body ovate, robust, reddish-brown, densely clothed with yellowish scale-like setae.
Clypeus rectangular, its length nearly equal to 0.33 x the cephalic length, slightly and
broadly emarginate in front and with the fore angles rounded; punctation large and
dense with erect scale-like setae. Fronto-clypeal suture without setae, except at center;
frons with punctation similar to that of clypeus but with longer and semierect setae;
transverse eye diameter equal to 0.25 x the cephalic width. Labrum twice longer than
underside of clypeus and with dense, fine punctures; mandibles robust; mentum
almost flat, with lateral depressions posteriorly and with anterior declivity little
marked. Antennae 10-jointed, club reddish; last article of the maxillary palpi not
impressed.

Pronotum evenly rounded laterally, wider at middle, fore and hind angles obtuse;
pronotal surface with dense setiferous punctures, setae erect directed to midline,
surface between the punctures finely punctate; anterior margin with long, erect setae;
posterior margin with short, decumbent setae. Scutellum densely punctate and with
decumbent setae overall.

Elytra longer than wide (1:0.70); punctation similar to that of pronotum; striae
little evident; elytral surface with setae shorter than the length of scutellum, most of
them decumbent; some erect setae on the costae and sutural interval. Abdomen
ridged laterally; propygidium not grooved; abdominal stemites of equal length, clothed
with decumbent setae and with a transverse row of erect setae; pygidium a little
wider than long, densely punctate-setiferous, setae long semierect and short decum-
bent.

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NEW D I PLOT AX IS

579

Protibiae tridentate, teeth placed in the distal half, basal tooth slightly marked;
protarsi shorter than protibiae; mesotarsi longer than mesotibiae; first and second
mesotarsomere of same size; basal metatarsomere of same length as the longest
metatibial spur; claws abruptly bent and subapically cleft, tooth wider than the apex
but of the same length and with distal half broadly separated from the apex. Genitalia
with basal piece a little longer than parameres, which are jointed behind the middle
and widened on apical third (Figs. 3-4).

Allotype female. Length 9.1 mm; elytral maximum width 4.9 mm. Differs from
holotype in the following respects: pygidium a little longer and the basal metatar-
somere shorter than longest metatibial spur.

Variation. Female, length 8.6 mm; elytral maximum width 4.5 mm. Males, length
6. 8-8.4 mm; elytral maximum width 3.4-4. 3 mm. The color of the elytra varies from
reddish-brown to reddish, the density of the setae varies a little, possibly by abrasion,
and the apex of clypeus is almost truncate in the smaller specimens.

Type locality. Ocotal Chico, Sierra de Santa Marta, Los Tuxtlas, Mexico.

Remarks. Diplotaxis squamisetis does not fit into any presently defined groups of
the genus. The presence of dense vestiture over all of the body, formed by decumbent,
semierect and erect yellowish scale-like hairs is exclusive to this species, and very
different from all other setiferous species of Diplotaxis, as the species of the “puberu-
la,” “pilipennis” and “pilifera” groups. In the first two groups the vestiture is formed
by erect and fine setae and in the last one by white, short, scale-like setae apressed
to the surface. Other species with thick scaly vestiture are D. clypeata Bates and D.
mus Fall, the former differing principally from D. squamisetis by the smaller eyes,
dark antennal club, lack of pubescence on elytral striae, shape of clypeus and different
vestiture (similar to that of the “pilifera” group); the second one differing by the
white, scaly vestiture apressed to surface, laterally rounded abdomen, straight-sided
pronotum, and rounded clypeus.

The food plant, Guarea glabra Vahl (Meliaceae) is a deciduous tree, with height
of 3-15 m, ranged from Mexico to Colombia, Venezuela and Ecuador. In the region
of “Los Tuxtlas,” Veracruz, Mexico, it is a dominant species within the forest (Ibarra-
Manriquez, 1985).

Etymology. The specific epithet refers to the conspicuous scale-like setae over all
of the body.

The key to species and species-groups of Diplotaxis in Vaurie (1960:188) should
be modified as follows to incorporate D. squamisetis.

87(86) Clypeus rounded from side to side as a semicircle, its front margin not reflexed;

Arizona; Chihuahua mus

87' Clypeus trapezoid, quadrate or rectangular in shape, angulate at sides, front margin

reflexed; central, southern and eastern Mexico 87a

87a Dorsal vestiture formed by yellow scale-like setae, most of which are semierect;
clypeus rectangular with the front angles rounded, its frontal margin slightly re-
flexed; labrum twice longer than the reflexed under side of clypeus; Veracruz . . .

squamisetis

87a' Dorsal vestiture formed by white, depressed scales; clypeus trapezoid or quadrate
with the front angles obtuse or straight, its front margin strongly reflexed; labrum
of the same length or shorter than under side of clypeus; central and southern
Mexico 88

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

ACKNOWLEDGMENTS

The authors offer thanks to S. McCleve for his valuable comments on the manuscript, to W.
B. Warner, S. Santiago and J. Blackaller for the loan of specimens, to M. A. Moron for the
donation of specimens collected by him, to O. Aquino for the determination of the host plant
and his aid during collecting trips, to G. Cabrera, L. Arellano, P. Rojas, E. Santos and J. R.
Hernandez for assistance in the collects, to R. Novelo for the revision of our English version.

This work represents contribution No. 13 of the “Studies of the Coleoptera associated with
the rhizosphere of the grass crops in Mexico” (P228CCOX891679) project supported by CON-
ACYT, Mexico.

LITERATURE CITED

Delgado-Castillo, L. 1 990. Dos nuevas especies mexicanas de Diplotaxis del grupo “Puberea”
(Coleoptera: Melolonthidae; Melolonthinae). Folia Entomol. Mex. 78:61-70.
Ibarra-Manriquez, G. 1985. Estudios preliminares sobre la flora lenosa de la Estacion de
Biologia Tropical Los Tuxtlas, Veracruz, Mexico. Tesis. Fac. Ciencias, UNAM, Mexico,
264 pp.

Vaurie, P. 1958. A revision of the genus Diplotaxis (Coleoptera, Scarabaeidae, Melolonthinae).

Part 1. Bull. Am. Mus. Nat. Hist. 1 15:267-396.

Vaurie, P. 1 960. A revision of the genus Diplotaxis (Coleoptera, Scarabaeidae, Melolonthinae).

Part 2. Bull. Am. Mus. Nat. Hist. 1 20: 1 6 1 — 434.

Vaurie, P. 1 962. New synonymy for Diplotaxis ebenina from Martinique and new distribution
records for Diplotaxis (Coleoptera: Scarabaeidae: Melolonthinae). Coleopts Bull. 16:
97-98.

Received 12 September 1991; accepted 30 April 1992.

J. New York Entomol. Soc. 100(4):58 1—593, 1992

STUDIES OF NEOTROPICAL CADDISFLIES, XLVII;
KUMANSKIELLA , A NEW GENUS OF
MICROCADDISFLIES FROM CUBA AND PUERTO RICO

Steven C. Harris1 and Oliver S. Flint, Jr.2

'Department of Biology, The University of Alabama,

Tuscaloosa, Alabama 35487, and

2Department of Entomology, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560

Abstract.— A new genus of Neotrichiini, Kumanskiella, and new species K. karenae, from
the Greater Antilles are described and illustrated. Mayatrichia aliena is transferred into this
new genus. Keys to the genera of Neotrichiini are provided and a proposed phylogeny of the
tribe presented. Kumanskiella appears most closely related to Neotrichia with Taxatrichia the
sister group to Mayatrichia, Kumanskiella and Neotrichia.

In 1987, Kumanski described Mayatrichia aliena from a single male collected from
Sierra de Trinidad, Cuba. He placed the species in the genus Mayatrichia primarily
on the basis of the four tibial spurs on the hindleg, but mentioned that in most
characters of the male genitalia the species was similar to Neotrichia. Kumanski
concluded that the species might be better placed in a new generic taxon, but with
only a single specimen he left the taxonomic position unresolved.

In 1989, a series of microcaddisflies similar to M. aliena were collected from a
malaise trap over a small stream in Puerto Rico by E. C. Masteller. A comparison
of this material with the type of M. aliena revealed the specimens to represent a new,
but closely related species. Larvae of this species were collected by Karen Buzby and
were found, as with the adults, to possess characteristics intermediate between the
genera Mayatrichia and Neotrichia. With both species fitting poorly within the generic
limits of either Mayatrichia or Neotrichia, we herein erect the new genus Kuman-
skiella.

Morphological terminology used in the species descriptions follows that of Mar-
shall, 1979. Length is measured from the top of the head to the tip of the forewings
and is given as a range with more than one specimen. Type material will be deposited
in the National Museum of Natural History (Smithsonian Institution), Center for
Energy and Environmental Research (Puerto Rico), and the University of Alabama.
The type of Mayatrichia aliena is deposited in the National Natural History Museum,
Sofia, Bulgaria.

Kumanskiella, new genus

Type species: Kumanskiella karenae, new species

Adult. — Head light brown in color, yellow between posterior warts; 3 ocelli; ten-
torium with anterior arms complete; antennae light brown, 1 8-segmented, about xh
length of forewing. Thorax light brown; pronotum narrow with pair of mesal and

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

lateral warts; mesoscutellum without transverse suture; metascutellum pentagonal.
Forewings narrow, brown with no discernible pattern in alcohol, small lobe on pos-
terobasal margin; hindwings brown, attenuate, patch of long setae on anterobasal
margin. Legs yellow, tibial spur formula 0, 2, 4, outer spurs barely Vi length of inner
spurs. Body circular in cross-section. Abdomen yellow; segment VI with short api-
comesal projection.

Male genitalia. —Abdominal segment IX elongate posteroventrally in lateral as-
pect, bearing spine or sclerotized projection. Bracteoles present as lightly sclerotized
lobes or acute projections from lateral margins of segment IX. Segment X membra-
nous and indistinct dorsally. Inferior appendages well developed and elongate.
Subgenital plate prominent, narrowing to distal beak in lateral aspect, in ventral view
rounded basolaterally with mesal protuberance distally. Phallus with long tubular
basal portion, sinuate asymmetrial median area bearing spiral process, apical portion
bulbous beyond spiral process then tapering to narrow apex.

Female genitalia. — Segment VIII with pair of elongate apodemes; thin tongue-like
process arising anteroventrally, tapering and projecting posteriorly. Segment IX with
pair of short apodemes. Segment X short, rounded apically bearing pair of papillae
subapically. Bursa copulatrix simple, narrow and elongate anteriorly and posteriorly,
median portion oblong with thin cuneiform mesal sclerite.

Larva. — Head unicolorous brown, narrowing anteriorly in dorsal view, coronal
and frontoclypeal sutures distinct, elongate seta posterior to eye, labrum well devel-
oped and covering mandibles. Thoracic nota unicolorous brown with medial ecdysial
line; short, stout setae along anterior and anterolateral margins, few scattered short
setae on dorsal surface; pleural sclerites well developed on meso- and metathorax,
reduced on prothorax; prothoracic legs short with widened coxa and femur, meso-
and metathoracic legs slender and elongate. Abdomen nearly cylindrical with distinct
intersegmental grooves, lateral fringe of short setae, elongate seta dorsally and ven-
trally; abdominal segments IX and X with dorsal sclerites each bearing numerous
elongate setae; anal prolegs elongate, projecting from body, anal claw short and stout.
Larval case constructed of sand grains, slightly compressed dorsoventrally, gently
tapering posteriorly, anterior and posterior openings semicircular.

Etymology. —Named in recognition ofKrassimir P. Kumanski, who first collected
the genus in Cuba.

Kumanskiella karenae, new species
Figs. 1-4

Male. — Length 1.5-1. 9 mm. Brown in alcohol, no discernible pattern on wings.
Antennae 1 8-segmented. Sternum of abdominal segment VI with short apicomesal
process. Segment VIII square. Segment IX in lateral view narrowing posteriorly to
a mesal, setose lobe, which bears a large distally-projecting spine; short rounded
bracteoles arising mesally from inner surface; in ventral view deeply incised ante-
riorly, posteriorly broadly incised, each lateral arm bearing large spine; in dorsal
aspect, broadly incised posteriorly and anteriorly, pair of oblong bracteoles originating
near lateral margins and projecting distally. Segment X narrow in lateral view, mem-
branous near apex; in dorsal aspect short and broadly rounded distally. Inferior
appendages thin in lateral view, narrowing to rounded apex; in ventral view narrowly

1992

K UMA NSKIELLA , NEW GENUS

583

Fig. 1 . Kumanskiella karenae, new species. Adult, male: A. Head and thorax, dorsal view.
B. Prothoracic leg. C. Mesothoracic leg. D. Metathoracic leg. E. Forewing. F. Hindwing.

separated basally and tapering distally from broad base, slightly diverging at mid-
length. Subgenital plate narrow, generally rectangular in lateral view; in ventral and
dorsal aspects wide basally, with setae bearing knobs on inner margin, rounded
laterally, narrowing to rounded apex, bearing pair of setae. Phallus with long, tubular
basal portion, sinuate asymmetrical median area bearing spiral process, apical portion
bulbous beyond spiral process then tapering to narrow apex.

Female. — Length 1. 5-2.0 mm. Brown in alcohol. Antennae 18-segmented. Ab-
dominal segment VI with short apicomesal process on venter. Segment VII truncate.
Segment VIII elongate and narrow, fringe of setae on posterior margin, elongate

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Fig. 2. Kumanskiella karenae, new species. Male genitalia: A. Lateral view. B. Dorsal view.
C. Ventral view. D. Phallus, dorsal view.

tongue-like process arising anteroventrally, narrowing distally, pair of thin apodemes
extending from lateral margins at juncture with segment VII and extending just into
segment VI. Segment IX short, pair of median apodemes originating anterolaterally
and extending anteriorly through segment VI. Segment X narrow and tapering to
rounded apex, bearing pair of papillae subapically. Bursa copulatrix simple, narrow
and elongate anteriorly and posteriorly, anterior extension about half length of pos-
terior, median portion oblong with a thin mesal sclerite which is deeply incised
anteriorly, smaller anterior sclerite with posterior notch.

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KUMANSKIELLA, NEW GENUS

585

Fig. 3. Kumanskiella karenae, new species. Female genitalia: A. Terminal abdominal seg-
ments, ventral view. B. Bursa copulatrix, ventral view.

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Fig. 4. Kumanskiella karenae, new species. Larva: A. Lateral view. B. Head and thorax,
dorsal view. C. Pro-, meso-, and metathoracic legs, lateral view. D. Abdominal segments IX
and X, dorsal view. E. Case, dorsal view, insert of posterior opening.

Larva. — Length 0.8- 1 .2 mm ( 1 0 larvae measured). Head unicolorous brown, slight-
ly darker at posterior margin, narrowing anteriorly in dorsal view, coronal and fron-
toclypeal sutures distinct, the latter less so in anterior portion, numerous setae along
frontoclypeal suture, elongate seta posterior to eye; antennae short, 2-segmented with
second segment about Vi length of first; labrum well developed and elongate, covering
mandibles. Thoracic nota heavily sclerotized, unicolorous brown, slightly darker at
posterior margins, each notum with median eedysial line; pronotum enlarged, about
as wide as long, narrowing posterolaterally, setae along anterior and anterolateral

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KUMANSKIELLA, NEW GENUS

587

margins, seta short and stout, few scattered short setae near ecdysial line; meso- and
metanota similar in shape, wider than long, short setae along anterior and antero-
lateral margins and scattered over the dorsal surface; pleural sclerites well developed
on meso- and metathorax, reduced to a small triangle on prothorax; prothoracic legs
short with widened coxa and femur; meso- and metathoracic legs slender and elon-
gate. Abdomen nearly cylindrical, with intersegmental grooves well defined, lateral
fringe of short setae, elongate seta dorsally and ventrally, segments IX and X with
dorsal sclerites each bearing numerous elongate setae; anal prolegs elongate, projecting
from body, lateral sclerite of proleg elongate, bearing several setae posteriorly, anal
claw short and stout. Larval case constructed of sand grains, slightly compressed
dorsoventrally, and gently tapering posteriorly, anterior and posterior openings semi-
circular. Length of case 0.9- 1.6 mm (10 cases measured).

Etymology. — Named in recognition of Karen Buzby who has faithfully removed
insects from the malaise trap and who collected and first recognized the larvae in
her field samples.

Type material. — Holotype <5. Puerto Rico, El Verde Field Station, Quebrada Prieta,
malaise trap, 370 m, 6-10 February 1990, O.S. Flint, Jr.

Paratypes. Puerto Rico, same data as holotype, 136, 329; same locality, but all
following from emergence trap; 22-27 February 1989, E.C. Masteller, 16, 39; March-
May 1989, K. Buzby and E.C. Masteller, 16, 19; 20 February 1990, 19; 19 June 1990,
19; 25 June 1990, 16; 4 July 1990, 19; 7 July 1990, 19; 25 July 1990, 29; 8 August

1990, 16; 13 October 1990, 19; 10 December 1990, 26; 11 February 1991, 19; 13
February 1991, 16; 20 February 1991, 16, 19; 22 February 1991, 26, 29; 4 March

1991, 19; 6 March 1991, 19; 11 March 1991, 16, 29; 19 March 1991, 16, 29; 23 March
1991, 19; 27 March 1991, 16; 30 March 1991, 16; 5 April 1991, 16; 8 April 1991,
29; 17 April 1991, 26; 20 May 1991, 26; 19 June 1991, 16; 9 July 1991, 19; 13 July
1991, 16; 17 July 1991, 26; 29 July 1991, 16, 19. Puerto Rico, El Verde Field Station,
Quebrada Toronja, bottom sample, 1 1 August 1990, K. Buzby, 2 larvae; 31 August
1990, 4 larvae; 1 1 October 1990, 1 larvae; 22 October 1990, 2 larvae; 12 December
1990, 4 larvae; 16 January 1991, 2 larvae.

Habitat. — Quebrada Prieta is a second order, high gradient mountain stream,
averaging 20% slope, near El Verde in the Luquillo Experimental Forest of north-
eastern Puerto Rico. The stream originates at approximately 600 meters above sea
level (masl) and flows into the Quebrada Sonadora at 310 masl; the emergence trap
was placed at 390 masl. The predominant substrates are bedrock, large boulders and
cobble. The larvae were taken in the Quebrada Toronja, a very similar stream that
is adjacent to the Prieta. These streams are all tributaries of the Rio Espiritu Santo.
They flow through tabonuco forest, the predominant type on the Luquillo Mountains
below 600 masl (preceding data supplied by K. Buzby, pers. comm.).

Kumanskiella aliena (Kumanski), New Combination

Fig. 5

Mayatrichia aliena Kumanski, 1987:24.

Male. — Length 1.7 mm. Brown in alcohol. Antennae broken, but with at least 18
segments. Sternum of abdominal segment VI with short apicomesal process. Segment
VIII square. Segment IX in lateral view narrowing posteriorly to form setose lobe,

588

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Fig. 5. Kumanskie/la a/iena, new combination. Male genitalia: A. Lateral view. B. Dorsal
view. C. Ventral view. D. Phallus, dorsal view.

postero ventral margin with elongate, acute process; dorsally with pair of long, finger-
like processes mesally, each bearing long seta apically; incised mesally in ventral
view, each lateral arm with heavy sclerotized spine directed inward; bracteoles scler-
otized and originating on inner posterolateral margins, in lateral view falciform, in
ventral view elongate and narrow, slightly emarginate apically. Segment X mem-
branous and indistinct dorsally, ventrolateral margins developed as an obliquely
curved sclerite, in dorsal aspect visible as acute lateral projections, directed poste-
riorly. Inferior appendages in lateral view rectangular with elongate dorsal extension

1992

KUMANSKIELLA, NEW GENUS

589

distally; in ventral view narrowly separated along straight inner margins, lateral
digitiform extension distally. Subgenital plate in lateral view with prominent distal
beak projecting somewhat ventrad; in dorsal view narrow with sinuate lateral margins,
distally with mesal protuberance bearing pair of subapical setae. Phallus with long,
tubular basal portion, sinuate asymmetrical median area bearing spiral process, apical
portion slightly bulbous beyond spiral process then tapering to thin, slightly flared
apex.

Female. — Unknown.

Larva. — Unknown.

Type locality. — Cuba: Province Las Villas, Sierra de Trinidad, small torrent on
the road Trinidad-Topes de Collantes, 150 m, 3 December 1982.

SYSTEMATIC RELATIONSHIPS

The Neotrichiini as presently defined (Marshall, 1 979) are restricted in distribution
to the New World with species occurring throughout North, Central, and South
America, including the West Indies. The tribe is now comprised of four genera,
Taraxitrichia, with a single species from Venezuela (Flint and Harris, 1991), Ku-
manskiella, with two species from the Greater Antilles, Mayatrichia with five species
from North, Central and northern South America (Harris and Holzental, 1990) and
Neotrichia with over 80 species and a widespread distribution over the Americas.
These genera are readily distinguished as adults and larvae, as far as they are known
(the immature stages of Taraxitrichia are as yet unknown) (Table 1).

KEY TO ADULT MALE NEOTRICHIINI

1. Mesothoracic leg with three tibial spurs; without ocelli Taraxitrichia

— Mesothoracic leg with two tibial spurs; with ocelli 2

2(1). Metathoracic leg with three tibial spurs Neotrichia

— Metathoracic leg with four tibial spurs 3

3(2). Bracteoles elongate (Harris and Holzenthal, 1990, fig. 1 A); phallus tubular and lack-
ing spiral process (Harris and Holzenthal, 1990, fig. ID) Mayatrichia

— Bracteoles not clearly evident (Fig. 2A); phallus with spiral process and divided into

basal and distal portions (Fig. 2D) Kumanskiella

TENTATIVE KEY TO LARVAE1

1. Thoracic nota with elongate hairs along the margins and scattered on the surface

(Wiggins, 1977, fig. 7.8F) Neotrichia

— Thoracic nota with short, stout setae along the margins and scattered on the surface

(Fig. 4A, B; Wiggins, 1977, fig. 7.7F) 2

2(1). Larval case constructed of sand grains (Fig. 4E); abdominal segments well defined

(Fig. 4A) Kumanskiella

— Larval case constructed of silk, walls reinforced with ridges (Wiggins, 1977, fig. 7.7B;

Ross 1944, fig. 558); abdominal segments poorly defined (Wiggins, 1977, fig. 7.1 A)
Mayatrichia

1 The key is tentative since only a few species of Neotrichiini have associated larval stages
and the larva of Taraxitrichia is unknown. Since Taraxitrichia appears to be closely related to
Mayatrichia, we would expect the larvae to be similar in overall appearance.

590

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

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1992

K UMANSKIELLA , NEW GENUS

591

Fig. 6. Proposed phylogeny of the genera of Neotrichiini.

Although Marshall (1979) summarized much of the current knowledge of the
Hydroptilidae, she only suggested phylogenetic placement of the tribes, giving neither
characters nor producing a cladistic analysis. In her Chart II, Marshall places the
Ochrotrichiini as the basal sister group to a clade inclusive of the Neotrichiini,
Hydroptilini, Orthotrichiini, and several unplaced genera. The Ochrotrichiini are
characterized by having a tibial spur formula of 1, 3, 4 or 0, 3, 4, simple phallus
without spiral process, 3 ocelli, and a transverse suture on the mesoscutellum.

The Ochrotrichiini are here considered the out-group to the Neotrichiini in order

592

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

to root the proposed phylogeny of Figure 6. Monophyly of the Neotrichiini is sup-
ported by the following synapomorphies:

1 . Bracteoles originating from posterior margin of abdominal segment IX;

2. Lack of a transverse suture on the mesoscutellum;

3. Larvae constructing cylindrical cases;

4. Anal prolegs of larvae, elongate, cylindrical and projecting prominently.

Taraxitrichia appears to be the sister group to the other genera in the tribe as it
retains the plesiomorphic condition of the third tibial spur on the mesothoracic leg
and the simple phallus which lacks a spiral process. However, the absence of ocelli
(5) in this genus is autapomorphic.

Mayatrichia, Kumanskiella and Neotrichia share the apomorphic condition of the
mesothoracic leg having two tibial spurs (6). Mayatrichia is distinguished on the
basis of the larvae having poorly defined abdominal segments (7). Mayatrichia retains
the simple phallic structure which is plesiomorphic. With Taraxitrichia, Mayatrichia
shares the character of the females having a bursa copulatrix with a pair of anterior
lobes (Harris and Holzenthal, 1990, fig. 2B). As well, both these genera have several
species with an elongate sternal process on the abdomen of the males. If we consider
these to be plesiomorphic characters possessed by the hypothetical ancestor of the
tribe (HTU), then a shortening of the sternal process, a character reversal, seen in
both Kumanskiella and Neotrichia, as well as a loss of the anterior lobes of the bursa
copulatrix seen in these two genera is apomorphic.

Monophyly for Kumanskiella and Neotrichia is inferred by three synapomorphies:

8. Complex phallic structure, usually with distinct distal and basal sections and
spiral process;

9. Larvae with cylindrical sand cases;

10. Bursa copulatrix with narrow, cuneiform vaginal sclerite.

Both genera also have in common, larvae with well defined abdominal segments
which is probably plesiomorphic since larvae of Ochrotrichiini also have this char-
acter. However, another possibility is that the HTU had larvae with poorly defined
abdominal segments, with a reversal of the character state in Kumanskiella and
Neotrichia. The interpretation of this larval character will be clearer with the discovery
of the larvae of Taraxitrichia.

Kumanskiella is distinguished by the modified bracteoles (11) which are difficult
to discern, and the asymmetrical median portion of the phallus (12). Neotrichia is
distinctive in that the metathoracic leg has only three tibial spurs (13) and the larvae
have elongate setae on the thoracic nota (14).

ACKNOWLEDGMENTS

Dr. Krassimir Kumanski, Director, National Natural History Museum, Sofia, Bulgaria, kindly
lent the type of Mayatrichia aliena for comparison as well as several Neotrichia from Cuba and
was very helpful in our diagnosis of the new genus. Dr. E. C. Masteller, Behrend College,
Pennsylvania State University, Erie, Pennsylvania, first installed the emergence traps on Que-
brada Prieta and has continued to oversee this operation and has sent all the caddisflies to OSF
for identification. Karen Buzby, College of Environmental Science and Forestry, State University
of New York, Syracuse, New York, has emptied these traps several times a week over the last

1992

KUMANSKIELLA, NEW GENUS

593

two years, and recognized the larvae of Kumanskiella in the bottom samples taken for her thesis
work. The research of Masteller and Buzby on the Quebrada Prieta was supported by grant
BSR-88 1 1 902 from the National Science Foundation to the Center for Energy and Environment
Research (University of Puerto Rico) and the Institute of Tropical Forestry (Southern Forest
Experiment Station) as part of the Long-Term, Ecological Research Program in the Luquillo
Experimental Forest. R. L. Mayden, University of Alabama, was very helpful in the phyogenetic
analysis and construction of Figure 6.

LITERATURE CITED

Flint, O. S., Jr. and S. C. Harris. 1991. Studies of Neotropical caddisflies, XLII: Taraxitrichia
amazonensis, a new genus and species of microcaddisfly from Venezuela (Trichoptera:
Hydroptilidae). Proc. 7th Intemat. Symp. Trich. (in press).

Harris, S. C. and R. W. Holzenthal. 1990. Hydroptilidae (Trichoptera) of Costa Rica: the
genus Mayatrichia Mosely. J. New York Entomol. Soc. 98:453 — 460.

Kumanski, K. P. 1987. On caddisflies (Trichoptera) of Cuba. Acta Zool. Bulgarica 34:3-35.
Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bull. Brit.
Mus. (Nat. Hist.), Entomol. Ser. 39:135-239.

Ross, H. H. 1944. The caddis flies, or Trichoptera, of Illinois. Illinois Nat. Hist. Surv. Bull.
23:1-326.

Wiggins, G. B. 1977. Larvae of the North American Caddisfly Genera (Trichoptera). Uni-
versity of Toronto Press, Toronto. 401 pp.

Received 24 September 1991; accepted 1 1 February 1992.

J. New York Entomol. Soc. 100(4):594-614, 1992

FIRST INSTAR DESCRIPTION OF KORSCHELTELLUS
GRACILIS (GROTE) AND STHENOPIS AURATUS
(GROTE) (LEPIDOPTERA: HEPIALIDAE) WITH A
CONSIDERATION OF CLADISTIC RELATIONSHIPS

BETWEEN SETAE

Jonathan G. Leonard, John R. Grehan, and Bruce L. Parker

Entomology Research Laboratory, University of Vermont,

655-B Spear Street, South Burlington, Vermont 05403

Abstract. — The external morphology and chaetotaxy of first instar Korscheltellus gracilis (Grote)
and Sthenopis auratus (Grote) are described. The larvae of K. gracilis are smaller and have
fewer microtrichia than S. auratus. The only difference in chaetotaxy involves the absence of
pore Va from the head of S. auratus. The setal nomenclatures of Hasenfuss (1969) and Wagner
(1987) are evaluated and used for description. The classifications of Hinton (1946) and Hasen-
fuss (1969) are compared for the homology of cranial setal groups. In a comparison of the P
and V setae we suggest that homologies are treated as apomorphies by Hasenfuss, while Hinton’s
setal homologies represent plesiomorphies. Hasenfuss’s ( 1 969) nomenclature renders the cranial
chaetotaxy of Hepialidae identical to the Ditrysia. We suggest that the prothoracic association
of SD1, SD2 and D2 may represent an exoporian apomorphy. Cladistic criteria provide a useful
technique for resolving homology problems and choosing between alternative systems of chae-
totaxy.

Korscheltellus gracilis (Grote) is a common eastern North American insect of upper
elevation boreal forest soils (Wagner, 1988; Tobi et al., 1989; Wagner et al., 1989;
Leonard et al., 1991; Wagner et al., 1991; Grehan et al., 1992; Tobi et ah, 1992).
Larvae feed on mosses, ferns and tree roots, including roots of red spruce ( Picea
rubens Sargent) and balsam fir (Abies balsamea (L.) Miller), which are species of
major concern for environmental scientists interested in boreal forest dynamics and
tree decline (e.g., Reiners and Lang, 1979; Siccama et ah, 1 982; Hornbeck and Smith,
1985; Main, 1987; Klein and Perkins, 1988; Vogelmann et ah, 1988; Bonan and
Shugart, 1989; Pitelka and Raynal, 1989; Silver et ah, 1991; Smith, 1991).

Larvae of Sthenopis auratus (Grote) feed on roots, stems and leaf bases of ostrich
fern (Matteuccia struthiopteris (L.) Todaro) (McCabe and Wagner, 1989) and wood
fern (Dryopteris campyloptera Clarkson) (JGL unpubh data). They are infrequently
recorded from the spruce-fir zone (1,000 m elev.) on Camels Hump Mt., Huntington,
VT, but are common at 200 m in the University of Vermont Experimental Research
Forest, Jericho, VT. At our Camels Hump field site one S. auratus and 1,200 K.
gracilis larvae were recovered from 1 10 soil pits (Leonard et ah, 1991).

The post-first instar chaetotaxy of K. gracilis was described by Wagner et ah ( 1 989)
using Stehr’s (1987) modification of Hinton (1946). The post-first instar of S. auratus
was described and the chaetotaxy discussed, but not illustrated, by McCabe and
Wagner (1989). Hepialid larvae have been described by a number of authors (e.g.,
Dyar, 1895; Packard, 1895; Fracker, 1915; Forbes, 1923; Gerasimov, 1937, 1952;
Hinton, 1946; Hasenfuss, 1963, 1969; Aitkenhead and Baker, 1964; Hardy, 1973;

1992

HEPIALID LARVAE

595

Table 1 . Hepialidae head chaetotaxy according to Hinton ( 1 946), Hasaenfuss ( 1 963, 1 969),
Wagner (1987), and Wagner et al. (1989) (modified from Stehr, 1987). Setal names in paren-
theses, mislabeling in square brackets.

Hinton

Hasenfuss

Wagner

Al (Anterior)

A3

Al

A2

A2

A2

A3

P2

A3

Ol (Ocellar)

Al

SI (Stemmatal)

02

Ol

S2

SOI (Subocellar)

SOI

SSI (Sub-stemmatal)

S02

S02

SS2

S03

SO 3

SS3

FI (Frontal)

FI

FI

Fa (pore)

Fa

Fa

Cl (Clypeal)

Cl

C2

C2

C2

Cl

LI (Lateral)

LI

LI

La (pore)

La

La

PI (Posterodorsal)

PI

PI

P2

VI (Vertexal)

P2

V2

V2

MD2 [MD1]

V3

V3

MD3

Va (pore)

Va

MDa [MD2]

G1 (Genal)

G1

MG1 (Micro-genal)

G2

03

MG2

Grehan, 1981; Grehan et al., 1983; Yasuda and Abe, 1986; Wagner, 1987; McCabe
and Wagner, 1989; Nielsen and Kristensen, 1989; Wagner et al., 1989; Common,
1990; Boudinot, 1991). Nomenclatural systems for hepialid chaetotaxy and cranial
pores proposed as modifications to Hinton (1946) include Hasenfuss (1969), Nielsen
and Kristensen (1989), Yasuda and Abe (1986), Stehr (1987), and Wagner (1987).
With the exception of Hasenfuss ( 1 963, 1 969), the alternatives are presented without
extensive comparative corroboration.

The chaetotaxy of first instars is different from second and later instars and may
be more conservative. First instars, therefore may contribute significant information
about phylogenetic relationships within the Exoporia (Nielsen, 1989). However, out
of an estimated 500 species in the Hepialidae (Kristensen, 1978), descriptions of first
instar Hepialidae are limited to about 10 species (Dyar, 1895; Fracker, 1915; Hinton,
1946; Aitkenhead and Baker, 1964; Grehan, 1981; Grehan et al., 1983).

This description of K. gracilis and S. auratus first instars will refer to Hasenfuss’s
(1969) setal nomenclature for the head, Wagner’s (1987) system for the prothorax
(TI) (with the addition of the Hasenfuss [1963] notation for the XD pores which were
not illustrated in Wagner [1987]), and Hinton (1946) for the meso- and metathorax
(Til, Till) and abdomen (A1-A10). We will discuss these nomenclatural systems in
relation to our observations on the first instars and comparisons with Hepialidae
described in the literature. Tables 1 and 2 compare the hepialid nomenclatural sys-
tems currently in use.

596 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Table 2. Prothorax chaetotaxy used for Hepialidae according to Hinton (1946), Nielsen and
Kristensen (1989), Wagner (1987), and Wagner et al. (1989).

Hinton

Nielsen and
Kristensen

Wagner

D1 (Dorsal)

D1

D1

D2

D2

SD2 (Sub-dorsal)

XD1 (Extra-dorsal)

XD1

XD1

XD2

XD2

XD2

XDb (pore)

XDa

—

XDa

XDb

—

XDc

XDc

—

SD1

SD1

SD1

SD2

SD2

D2

LI (Lateral)

LI

LI

L2

L2

L2

L3

L3

L3

SV 1 (Sub- ventral)

SV1

SV1

SV2

SV2

SV2

MV2 (Mid-ventral)

MV2

MV2

MV3

MV3

MV3

V 1 (Ventral)

VI

VI

We have confined our description to body setae and pores with the exclusion of
the larval thoracic leg and abdominal proleg setae. Comparative chaetotaxy of tho-
racic and abdominal legs has received little attention but several alternative systems
have been proposed (e.g., Gerasimov, 1952; Birket-Smith, 1984; Nielsen and Kris-
tensen, 1989).

MATERIALS AND METHODS

Korscheltellus gracilis eggs were collected on 25 July 1990 from a female that was
in copula on a balsam fir branch apex 1 m above ground at 1,000 m elevation on
Camels Hump Mountain, Huntington, Vermont. The mating pair and fir branch
were enclosed in a plastic bag after the evening flight (approximately 9 pm EST) and
eggs recovered the following morning. Sthenopis auratus eggs were collected on 14
July 1990 from a female found in copula on the frond tip of ostrich fern at the
University of Vermont Experimental Research Forest in Jericho, VT. Eggs were
incubated on moist filter paper inside petri dishes in a saturated atmosphere at 18°C.

First instars were preserved in 70% EtOH. The head capsules of 8 S. auratus and
24 K. gracilis were measured to the nearest 0.001 mm. Means and 95% confidence
intervals for head capsule measurements were computed using univariate descriptive
procedures in SAS (1985). Sub-samples of larvae were randomly chosen for exam-
ination by scanning electron (SEM) and light microscopes. Light microscope slides
were prepared as described in Ailkenhead and Baker ( 1 964). In addition, some larvae
were cleared with KOH and temporarily slide-mounted in glycerine. Descriptions
presented refer to both species unless otherwise indicated.

1992

HEPIALID LARVAE

597

RESULTS

Eggs of K. gracilis hatched in 26 days and those of S. auratus hatched in 30 days.
Eggs of S. auratus were larger than those of K. gracilis.

Head. The head capsule is straw-colored and large relative to the body size, es-
pecially for K. gracilis. The head capsule widths of S. auratus (mean = 0.546 mm,
standard error ± 0.002 mm, N = 8) were significantly larger than that of K. gracilis
(0.332 mm ± 0.002 mm, N - 24). Head setae of S. auratus were shorter relative to
the head size than on K. gracilis and the shape of the head capsule, adfrontal suture,
and mandible was different between species (Figs. 1-6).

The antenna appears 3-segmented, but may be 2-segmented as described for Fraus
by Nielsen and Kristensen (1989). Antenna is shorter and thicker than in the post-
first instar (cf. McCabe and Wagner, 1989; Wagner et al., 1989). Each antenna bears
three conical sensilla and three trichoid sensilla with the lateral trichoid sensillum
1.5 times the length of the antenna. The anterior-most trichoid sensillum arises from
the base of the most anterior conical sensillum (Figs. 7-8).

The labrum is deeply cleft with 5 pairs of setae. The mandible has 5 teeth. The
middle three teeth are serrate on ventrad edge. The mandible has 2 setae; one small
seta located on the dorsal tooth, the second 1.5 times the mandible length and inserted
into the mandible base (Figs. 3, 6).

The maxillae of both species are as described for post-first instar K. gracilis (Wagner
et al., 1 989), except the palpi are thinner and more elongate (Figs. 9-13). Each maxilla
bears a stout trichoid sensillum on the dististipes, a 3-segmented maxillary palpus,
and a mesal lobe (lobarium) with 1 trichoid sensillum on the base and 7 sensilla on
the apex. Unlike later instars, the basal trichoid sensillum on the mesal lobe is
subequal in length to the lobe. The apical sensilla of the mesal lobe consists of 2
styloconic sensilla, 3 small trichoid sensilla, and 2 larger trichoid sensilla. The basal
segment of the maxillary palpus bears a pore on the lateral side. The apical end of
the palpus bears larger, more prominent sensilla than in later instars including at
least 4 trichoid sensilla and 1 squamiform sensillum (Fig. 11-13). (The maxillary
palpus of K. gracilis is incorrectly labeled the labial palpus in Wagner et al. [1989:
721, Fig. 11].)

The hypopharynx and labium for both K. gracilis and S. auratus are nearly as
described for post-first instars by Wagner et al. (1989). The labial palpus is 2-seg-
mented, but the number is uncertain for the later instars (Wagner et al., 1989). The
distal segment is short and conical, bearing a trichoid seta subequal in length to the
palpus (Fig. 12). The spinneret of both species is approximately equal to the length
of the maxillolabial-hypopharyngeal complex and is oriented ventro-caudad (Figs.
1, 4, 14-15). The hypostomal plates are not fused at the midline for both K. gracilis
and S. auratus first instars.

In larvae of K. gracilis the pore Va (=MD2 of Wagner et al., 1989) is positioned
between the two vertex (V) setae (N = 6) as in other Hepialidae (Figs. 1-2), but
missing in S. auratus (N = 7) (Figs. 4-5, 16, 17). The pore MDa was misidentified
as microseta MD2 in Wagner et al. (1989), resulting in the anterior microseta MD2
being misidentified as MD1. Setae Al and SI were also mislabelled (Wagner et al.,
1989:721, fig. 8) as A3 and S3 respectively.

First instars of both K. gracilis and S. auratus show all of the setal characters
present in later instars except for the lack of subprimary setae (see Tables 1 and 2

Figs. 1-3. Korscheltellus gracilis first instar head chaetotaxy. 1 . Dorso-frontal view. 2. Lateral
view. 3. Mandible.

1992

HEPIALID LARVAE

599

1 00 jt/m

Figs. 4-6. Sthenopis auratus first instar head chaetotaxy. 4. Dorso-frontal view. 5. Lateral
view. 6. Mandible.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Figs. 7-8. Scanning electron micrographs. 7. Antenna of Korscheltellus gracilis, 1095 x. 8.
Antenna of Sthenopis auratus, 535 x .

for nomenclature). First instars lack: L2 on TII, Till and A9; SV2 on TII and Till,
and A7-A9; L3 on all segments; SV3 on A1-A9.

Thorax. Both species have microtrichia on all thoracic and abdominal segments
but S. auratus is more densely covered. Conical microtrichia were present in the TI
spiracular opening of -S’, auratus (Fig. 1 8) but absent in K. gracilis. The number and
relative positioning of setae is the same as previously reported for other hepialid first
instars. TI lateral setae (LI, L2) are on a laterad sclerotized plate just ventrad of the
prothoracic shield (Figs. 19, 21). Setae SD1, SD2, and D2 are on a thinly sclerotized
latero-caudad extension of the prothoracic shield. In comparison to D2, setae SD1
and SD2 are thicker, longer and inserted into larger sockets (unlike post-first instars).
SD setae are not subtended by dense microtrichia as in post-first instars. D2 is closer
to SD setae than to XD setae as in post-first instars. MV setae are of approximately
equal length.

On TII and Till of both species, D1 pinacula are fused across the midline. Setae
D2, SD1 and SD2 are on a single non-melanized pinaculum in K. gracilis on both
Til and Till (Fig. 19) unlike post-first instars where on Till D2 is on a separate
melanized pinaculum (Wagner et al., 1989). In S. auratus D2 is on a separate mela-
nized pinaculum on Till (Fig. 21), as in post-first instars (McCabe and Wagner,
1989). MD1 is the same length as MSD setae in K. gracilis first instars which differs
from post-first instars where MD1 is longer. MD1 in the first instar of S. auratus is
slightly longer than the MSD setae. The thoracic leg claw basal tooth ends near one-

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Figs. 9-15. Scanning electron micrographs. 9. Maxillolabial-hypopharyngeal complex of
Korscheltellus gracilis, 175 x. 10. Maxillolabial-hypopharyngeal complex of Sthenopis auratus,
155x. ll. Maxilla of Sthenopis auratus, 225 x. 12. Maxillolabial detail of Sthenopis auratus,
225 x. Arrows point to labial palps. 13. Maxilla of Korscheltellus gracilis, 465 x. 14. Head of
Korscheltellus gracilis in lateral view, 310x. 15. Head of Sthenopis auratus in lateral view,

1 25 x .

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Figs. 16-18. Scanning electron micrographs. 16. Vertex setae of Sthenopis auratus, 145x.
17. Detail of box from Figure 16 showing V setae. 18. Detail of spiracle of Sthenopis auratus
showing conical microtrichia, 640 x .

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19

20

Figs. 19 20. Chaetotaxy of Korscheltellus gracilis first instar larvae; spiracle represented by
shaded ellipse. 1 9. Thorax and first two abdominal segments; CX, coxa. 20. Abdominal segments
3-10; non-melanized pinacula bounded by solid lines; P, pore; PRL, proleg.

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third of the claw length (Figs. 23-24), but in second to final instars the tooth ends
near midlength (Wagner et al., 1989: fig. 12).

Abdomen. The number and relative positioning of abdominal setae is the same as
for other first instar hepialids. MV3 is a proprioceptor microseta on Al, A8, and
A9, but a tactile macroseta on A2-A7 (Figs. 20, 22). In S. auratus a darkened
melanized pinaculum encloses SD1 and SD2 while the L setae are each enclosed in
a separate melanized pinaculum as in post-first instars. Crochets in both species are
in biserial ellipses with a prominent inner row (Figs. 25, 26). Segment A 10 has 1 1
setae and two pores: anal shield with 3 setae and a pore; remaining pore and 8 setae
below shield.

DISCUSSION

Chaetotaxy is important in the systematics of the Lepidoptera because it is used
to distinguish taxa and to define monophyletic groups. At present there is a lack of
explicit analytical criteria for identifying chaetotactic homologies that confer natural
setal groups. Hinton’s (1946) major review emphasized comparisons among the
Ditrysia and some Monotrysia including the Hepialidae. Earlier systems were already
in existence (e.g., Fracker, 1915) and Hinton’s ( 1 946) nomenclatural system modified
earlier proposals by Heinrich (1916) and Gerasimov (1935) (Hinton, 1946:9, Table
1). Hinton’s (1946) nomenclature has received wide application, but several authors
have identified problems with respect to the Hepialidae and offered modifications
(Hasenfuss, 1963, 1969; Wagner, 1 987; Nielsen and Kristensen, 1989). The different
alternatives and their justifications are discussed as follows:

Head. Hinton’s ( 1 946) nomenclature (Fig. 27) was critically examined by Hasenfuss
(1969), who was concerned with Hinton’s designation of a unique genal micro seta
(G2) and apparent absence of micro seta VI that renders the Hepialidae anomalous
to the Ditrysia where G2 is absent and VI is present. Hinton’s ( 1 946) contrast between
Hepialidae and Ditrysia is based upon differences in size and position of setae (the
G and V setal groups being defined by their small size as microsetae). Hinton (1946:
6) was unable to decide which of the G setae were represented in the Ditrysia and
VI was presumed absent from Hepialidae because the two micro V setae present
were necessarily identified as V2 and V3 by the interposition of the pore Va (Fig.
27).

Hasenfuss (1969) argued that setal length was a consequence of head capsule
orientation and degree of retraction of the head against the thorax. He also pointed
out that in leaf mining groups the longer tactile setae are present only on the leading
edge and sides of the head capsule. He suggested that setal length alone does not
qualify as a defining character (i.e., the equivalent of apomorphy) for inclusion of
setae within particular setal groups. He noted that there are normally three vertex
setae in the Ditrysia (VI, V2 and V3) with the pore Va almost always between V2
and V3 (Va is sometimes offset to one side), while the genal region supports only a
single seta G1 (Fig. 28). While the homology of Ditrysian setae was stable, there were
cases where the boundary between setal groups (“Musterelementgruppe”) became
blurred. Hasenfuss (1969) presented the Yponomeuta as an example with V2 and V3
present as microsetae while VI was very long and having the appearance of a mac-
roseta. The Hepialidae show the same total number of head setae (although fewer

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22

Figs. 21-22. Chaetotaxy of Sthenopis auratus first instar larvae; spiracle represented by
shaded ellipse. 21. Thorax and first two abdominal segments. 22. Abdominal segments 3-10;
melanized pinacula bounded by solid lines; P, pore; PRL, proleg.

pores) as the Ditrysia, but only two microsetae occur on both the vertex (normally
three in the Ditrysia), and the genal region (normally one in the Ditrysia). In Hinton’s
system, all the long tactile setae of Hepialidae are regarded as homologous with the
Ditrysia and VI therefore is assumed missing and a new seta, G2, considered present.

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Figs. 23-26. Scanning electron micrographs. 23. Prothoracic (TI) claw of Korscheltellus
gracilis, 1,3 15 x. 24. Prothoracic (TI) claw of Sthenopis auratus, 920 x. 25. Abdominal proleg
of Sthenopis auratus, 820 x, arrow on outer crochet. 26. Abdominal proleg of Korscheltellus
gracilis, 1 ,455 x , arrow on outer crochet.

Since Yponomeuta exhibits Ditrysian chaetotaxy except for the presence of only two
micro-vertex setae, the macro seta anterior to V2 is necessarily VI. This arrangement
of micro and macro setae is identical to the Hepialidae and, therefore, the hepialid
macro seta P2 of Hinton (1946) is regarded as VI (compare Figs. 27 and 28). Hasen-
fuss (1969) suggests that the resulting rearrangement supports the view that Hinton’s
micro seta G2 is actually homologous with 03, with the result that the chaetotaxy
of the Hepialidae is identical to that of the Ditrysia.

Stehr (1987) produced a system adopted by Wagner (1987) (Fig. 29) modifying
Hinton’s (1946) homology and terminology of the head. The major concern was that
the micro-vertex setae (V series) and genal seta (G series) were proprioceptors ho-
mologous with the rest of the body and should be renamed in accordance with
Hinton’s system as microdorsal (MD) and microgenal setae (MG) (although to be
consistent with Hinton’s nomenclature, the genal setae should have been designated
by MV). J. E. Rawlins (pers. comm.) suggests that while there may be some indications
for homonomy of genal proprioceptors with the MV group on the thorax and ab-
domen, priority of usage favors retention of G and MV respectively as symbols for

27. Hinton (1946)

S02

29. Wagner et al. (1989)

Figs. 27-29. Comparison of head chaetotaxy systems. Head of Korscheltellus gracilis re-
drawn from Wagner et al. (1989). 27. Hinton (1946) system. 28. Hasenfuss (1969) system. 29.
Wagner et al. (1989) system; SS and MG setae marked with * were not labeled in Wagner et
al. (1989), Stehr (1987) system is used for these setae.

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those groups. In Stehr (1987:290-291, 300) the designation of Hinton’s Cl and C2
was accidentally reversed due to labeling error (F. W. Stehr, pers. comm.).

Nielsen and Kristensen ( 1 989) adopted the nomenclature of Hasenfuss ( 1 969) with
the exception of the genal setae where they retain Hinton’s (1946) G2 instead of 03
as used by Hasenfuss (1969). Kristensen (1984), in a discussion of the ground plan
of the lepidopteran larval head, stated the retention of G2 instead of 03 “seems most
natural from a phylogenetic point of view,’’ because the genal position of the seta is
ancestral in the Lepidoptera (N. P. Kristensen, pers. comm.). We have retained 03
in this paper pending future reevaluation of cranial chaetotaxy.

The contrast between the classifications of Hinton (1946) and Hasenfuss (1969)
can be represented cladistically for the P and V setae. In a cladistic representation,
setae are terminals and homologies are nodes. The use of branching diagrams has
no necessary implication for the ultimate origins of these morphological features
although one might attempt to explore the implications with respect to ontogeny or
phylogeny (Nelson, 1989).

The classification of seta P2 is inferred by Hinton ( 1 946) to share closer relationship
to the macroseta PI than to microsetae V2/V3 (Fig. 30). The principles by which
homologies are recognized for different setal groups were not explicitly stated by
Hinton or Hasenfuss, although Hasenfuss (1963) supported a cladistic approach to
phylogenetic classification in general. Hasenfuss (1969) suggested that the relative
distinction between “long” and “short” setae did not provide an adequate defining
character for setal groupings in this context. The fact that P2 is long was not sufficient
to place it only with PI because the “long” character would also place it within
several other setal groups. Hasenfuss argued that the position of pore Va provides a
defining character identifying a sister group relationship between V3 and V2 (i.e.,
the setae share a closer relationship to each other than with any other setae). To this
group is added VI (formerly P2) by virtue of its position adjacent to and in line with
V2-V3 as also found in the Ditrysia (Fig. 31). With this reformulation of setal
relationships, the setae of Hepialidae are not incongruent with respect to the Ditrysia
and we support the nomenclature of Hasenfuss for the head capsule because the
modifications are cladistically explicit and justifiable with reference to the Ditrysia.
The status of G2 also requires further clarification. The retention of G2 by Nielsen
and Kristensen (1989) while accepting VI of Hasenfuss requires the elimination of
03. We acknowledge that the present setal classifications are provisional pending
future comprehensive evaluation of Lepidoptera larvae.

Thorax. Nielsen and Kristensen (1989) adopted Hardy’s (1973) thoracic nomen-
clature, except for the MD and MSD microsetae where they used Hinton’s system.
Hardy (1973) applied a modified Hinton system but the XD pores on the prothorax
(TI) were not labeled (Hardy, 1973:122, fig. 1). Hasenfuss (1963) reversed Hinton’s
subscripts for the XD pores so they were in linear order (a, b, c) and this order was
adopted by Nielsen and Kristensen ( 1 989). It is unclear why Hinton ( 1 946) originally
labeled these pores in the order b, a, c. He may have ordered the pores by their
proximity to XD1. The pore labeled XDa is closest to the XD1 (Hinton, 1946:19,
fig. 23). We use Hasenfuss’s (1963) subscripts for the XD pores.

Wagner (1987) adopted Stehr’s (1987) system of nomenclature (which is a mod-
ification of Hinton, 1946) for the prothorax (TI), but reversed the notation of D2
and SD2. The identity of Hinton’s D2 as SD2 presents a further example of prob-

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HEPIALID LARVAE

609

30

Hinton (1946)

PI P2 V2 V3

31

Hausenfuss (1969)

Pi VI (P2) V2 V3

Figs. 30-31. Cladistic representation of the vertex setae using (30) Hinton (1946) and (31)
Hasenfuss (1969). Hinton’s classification treats setal size as an informative character for PI
and P2. This classification is altered by Hasenfuss to include Hinton’s “P2” as part of the
vertex group with the informative character being its proximity to V2/V3.

lematical setal relationships. Nielsen and Kristensen (1989) acknowledged Wagner’s
(1987) notation, but retained Hinton’s usage pending further explanation of the
rationale. Wagner (1987) noted that the designated SD setae are thin in comparison
to D2 (SD2 of Hinton, 1 946). Based on first instars, the two pronotal setae most
similar in length and socket size are what Wagner (1987) calls SD1 and SD2 (D2 of
Hinton) (Figs. 19, 21). However, unlike post-first instars, setae SD1 and SD2 are
thicker, longer and insert into larger sockets than D2.

The SD setae (sensu Wagner, 1987) are subtended by dense microtrichia in later

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instars of K. gracilis and S. auratus, as in some other Hepialidae (e.g., Fig. 32) (see
also Martyn, 1960; Elder, 1978; Grehan, 1981; Nielsen and Kristensen, 1989). In
still other hepialid species the seta D2 (sensu Wagner, 1987) shares a microtrichical
bed with both SD setae, but never with only one of them. The SD association with
microtrichia may represent an exoporian trait as it is also present in the hepialoid
sister group Mnesarchaeidae (Fig. 33) but not outside the Exoporia. Nielsen and
Kristensen (1989:94) suggest that the presence of all three setae within a single
“melanized depression” (=microtrichial bed of this paper) is an apomorphy for the
Hepialidae sensu lato. First instar K. gracilis and S. auratus have no microtrichial
bed associated with the SD setae or D2 (also the case for other first instar Hepialidae),
but they share a thinly melanized section of the shield. The setal size and presence
of microtrichia surrounding the SD setae vary, but the setae consistently share ho-
mologies that exclude D2 in all species reported and we therefore support Wagner’s
(1987) revision of nomenclature.

Abdomen. Hinton’s (1946) system of abdominal nomenclature has not received
modification in the literature, but requires critical examination comparable to that
applied to the head capsule by Hasenfuss (1969). In particular are problems of serial
homology or homonomy of the thoracic and abdominal L setae (D. L. Wagner, pers.
comm.). Mutuura (1980) has suggested abdominal musculature as one source of
characters for comparing pinacula/setal homologies between abdominal segments.

PHYLOGENY AND HEPIALID CHAETOTAXY

Chaetotaxy has historically provided useful phylogenetic characters, but the alter-
native nomenclatural choices currently available for hepialids represent a significant
problem for recognizing setal homologies. Setal and pore characters currently rec-
ognized by all workers as “characteristic” for larvae of Hepialidae are based on
Gerasimov (1937, 1952) and Hinton (1946) and include the following; The frons
pore (Fa) is laterad and slightly ventral to FI seta. The Adfrontal pore (Afa), anterior
pore (Aa), posterior pores (Pa, Pb), ocellar (stemmatal) pores (Oa, Ob) and genal
pore (Ga) are absent. Setae A3, A2, and P2 (of Hasenfuss, 1969) are almost in line
running between the stemmata and VI, with P2 caudad of PI. Two microsetae are
present in the genal region: 03 (of Hasenfuss, 1969) and Gl. Stemmata (ocelli) are
arranged in two vertical columns. The prothoracic shield includes LI, L2 and some-
times L3 in post-first instars. The prothorax has two subventral setae (SV1 and SV2)
and two mid ventral setae (MV2 and MV3). Prothoracic setae SD1 and SD2 (of
Wagner, 1987) are diagonally positioned and thin in comparison to D2 in post-first
instars. Seta MV3 is elongate on TI in post-first instar larvae. The mesothorax (TII)
has MD1, MSD1 and MSD2 on the same pinacula anterad of SD1 and SD2. Seta
MD1 is often elongate in comparison with the MSD setae. There is only one sub-
ventral seta (SV1) on TII and Till; there are two (SV1 and SV2) on the prothorax.
On the abdomen, setae SD1 and SD2 are on the same pinacula with SD2 shorter
and thinner than SD1. Setae LI and L2 are near the spiracle with LI being farthest
caudad and L2 being closer and ventrad of the spiracle (L2 is incorrectly described
as being caudad of LI and spiracle, and LI ventrocaudad to spiracle in Wagner
[1987]). On post-first instars, L3 is anterad and ventrad of the spiracle on segments
A 1-A8. On segments A2-A7, MV3 is elongate and grouped with the SV setae. Wagner
(1987) identified additional hepialid larval characters: Head: Seta 03 (of Hasenfuss,

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Figs. 32-33. Scanning electron micrographs. 32. SD and D2 setae of Trioxycanus sp. showing
the microtrichial bed shared by the SD setae, 65 x . 33. SD and D2 setae of Mnesarchaea sp.
showing the microtrichial bed shared by the SD setae, 80 x .

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1969) is approximately half-way between G1 and S03. Prothorax: SD1 and SD2
are diagonally positioned in a darkened pinacula. In some species SD pinacula en-
compassing D2 in a darkened pigmented area with dense microtrichia.

Nielsen and Kristensen (1989) commented on several characters that appear to
represent exoporian or hepialid apomorphies, but are also represented in other Lep-
idoptera. They suggest the presence of MXD1 to be a glossatan groundplan character,
and its absence an Exoporian groundplan trait (but not a synapomorphy since this
proprioceptor is found in some but not all Eriocraniidae). Wagner (1987) identified
the presence of two microgenal setae on the head as unique characteristics of Hepiali-
dae, but this does not apply using the classification of Hasenfuss. Adult Hepialidae
provide only one recognized synapomorphy (absence of tibial spurs) and this is not
a universal character for the Hepialidae sensu lato (Viette, 1 949; Wagner and Tindale,
1988; Nielsen and Kristensen, 1989:1 13). The cladistic status of many chaetotactic
characters recognized for the Hepialidae require clarification. A cladistic classification
of setae, as proposed in this paper for the V series, provides a measure of confidence
for comparative chaetotaxy and may help determine which setal characters qualify
as hepialid synapomorphies. Complementary to this kind of evaluation is the inclu-
sion of a greater range of larval records, including first instars which are extremely
easy to obtain from fertilized eggs.

ACKNOWLEDGMENTS

Thanks to Kristi Kieth, Jennifer Neat, Lila Porter, Diane Riggleman, Dan Tolces, Dan Van
der Vliet, and Laura Wells for help with field collections. Greg Hendricks was invaluable for
help and suggestions with the scanning electron microscope. Eva Doane (Entomology Research
Laboratory) kindly translated sections of Hasenfuss (1963, 1969) and George Gibbs provided
the photo of the mesarchid sensory pit. Gary Nelson provided very helpful discussion on cladistic
methodology and its application to setal characters. John Dugdale, Niels Kristensen, John
Rawlins, Judy Rosovsky (Entomology Research Laboratory), Frederick Stehr, David Wagner
and James Liebherr (editor) contributed important suggestions for improving the manuscript.

LITERATURE CITED

Aitkenhead, P. and C. R. B. Baker. 1964. The larvae of the British Hepialidae. Entomologist
97:25-38.

Birket-Smith, S. J. R. 1984. Prolegs, legs, and wings of insects. Entomonograph 5:1-128.
Bonan, G. B. and H. H. Shugart. 1989. Environmental factors and ecological processes in
boreal forests. Annu. Rev. Ecol. Syst. 20:1-28.

Boudinot, J. 1991. Biologie d’Aenetus cohici Viette (Lepidoptera: Hepialidae). In: J. Chazeau
and S. Tillier (eds.), Zoologia Neocaledonica, Vol. 2. Mem. Mus. natn. Hist, nat., (A),
149:167-175. Paris.

Common, I. F. B. 1990. Moths of Australia. E. J. Brill, New York.

Dyar, H. G. 1895. The classification of the Lepidoptera on larval characters. Am. Nat. 29:
1066-1072.

Elder, R. J. 1978. Notes on the biology and descriptions of the life stages of Oncopera parva
Tindale (Lepidoptera: Hepialidae). J. Aust. Entomol. Soc. 17:5-1 1.

Forbes, W. T. M. 1923. The Lepidoptera of New York and neighboring states. Cornell Univ.
Agr. Exp. Sta. Mem. 68:1-729.

Fracker, S. B. 1915. The classification of lepidopterous larvae. 111. Biol. Monogr. 2:1-169.
Gerasimov, A. M. 1935. Zur Frage der Homodyamie der Borsten von Schmetterlingsraupen.
Zool. Anz. 1 12:117-194.

1992

HEPIALID LARVAE

613

Gerasimov, A. M. 1937. Hepialidenraupen (Lepid.). Ent. Zeit. 51:81-84.

Gerasimov, A. M. 1952. Lepidoptera, Part 1. Caterpillars. Zool. Inst. Acad. Nauk USSR.
New Ser. No. 56, Fauna USSR. Insects-Lepidoptera. 2:1-338.

Grehan, J. R. 1981. Morphological changes in the three-phase development of Aenetus vi-
rescens larvae (Lepidoptera: Hepialidae). New Zealand J. Zool. 8:505-514.

Grehan, J. R., A. Moeed and M. J. Meads. 1983. Observations on Trioxycanus enysii (Butler)
( sensu Meyrick, 1890) (Lepidoptera: Hepialidae) on Kapiti Island, New Zealand, with a
description of larval chaetotaxy. New Zealand Entomol. 7:408-413.

Grehan, J. R., B. L. Parker, D. L. Wagner, J. Rosovsky and J. Aleong. 1992. Root damage
by the conifer swift moth: a mortality factor in montane red spruce regeneration. For.
Sci. (in press).

Hain, F. P. 1987. Interactions of insects, trees, and air pollutants. Tree Phys. 3:93-102.

Hardy, R. J. 1973. Larval chaetotaxy of thorax and abdomen of Fraus simulans Walker
(Lepidoptera: Hepialidae). J. Aust. Entomol. Soc. 12:121-123.

Hasenfuss, I. 1963. Eine vergliechend-morphologische Analyse der regularen Borstenmuster
der Lepidopterenlarven. Studien zur Methodik der vergleichenden Morphologie der Bor-
stenmuster und zur phylogenetischen Deutung der Abwandlungen der regularen Bor-
stenmuster der Lepidopterenlarven. Zeit. Morph. Okol. Tiere 52:197-364.

Hasenfuss, I. 1 969. Zur Homologie der Borstenmusterelemente der Larven-Kopfkapsel einiger
monotrysischer Lepidoptera. Beitr. Ent. 19:289-301.

Heinrich, C. 1916. On the taxonomic value of some larval characters in the Lepidoptera.
Proc. Entomol. Soc. Wash. 18:154-164.

Hinton, H. E. 1946. On the homology and nomenclature of setate of lepidopterous larvae,
with some notes on the phylogeny of the Lepidoptera. Trans. Roy Entomol. Soc. Lond.
97:1-37.

Hombeck, J. W. and R. B. Smith. 1985. Documentation of red spruce growth decline. Can.

J. For. Res. 15:1 199-1201.

Klein, R. and T. D. Perkins. 1988. Primary and secondary causes and consequences of
contemporary forest decline. Bot. Rev. 54:2-24.

Kristensen, N. P. 1978. A new family of Hepialoidea from South America, with remarks on
the phylogeny of the suborder Exoporia (Lepidoptera). Entomol. Germ. 4:272-294.

Kristensen, N. P. 1984. The larval head of Agathiphaga (Lepidoptera, Agathiphagidae) and
the lepidopteran ground plan. Syst. Entomol. 9:63-81.

Leonard, J. G., D. R. Tobi and B. L. Parker. 1991. Spatial and temporal distribution of
Korscheltellus gracilis larvae (Lepidoptera: Hepialidae) in the Green Mountains, Ver-
mont. Environ. Entomol. 20:371-376.

Martyn, E. J. 1960. The ecology of Oncopera intricata Walker and Oncopera rufobrunnea
Tindale (Lepidoptera, Hepialidae) with notes on the classification of the genus Oncopera
in relation to larval taxonomy. Tasmanian Depart, of Agric. Res. Bull. 3.

McCabe, T. L. and D. L. Wagner. 1 989. The biology of Sthenopis auratus (Grote) (Lepidoptera:
Hepialidae). J. New York Entomol. Soc. 97:1-10.

Mutuura, A. 1980. Morphological relations of sclerotized and pigmented areas of lepidop-
terous larvae to muscle attachments, with applications to larval taxonomy. Can. Entomol.
1 12:697-724.

Nelson, G. 1989. Cladistics and evolutionary models. Cladistics 5:275-289.

Nielsen, E. S. 1989. Phylogeny of major Lepidopteran groups, pp. 28 1-294. In: B. Femholm,

K. Bremer and H. Jornvall (eds.), The Hierarchy of Life. Elsevier Science Publishers,
Amsterdam.

Nielsen, E. S. and N. P. Kristensen. 1 989. Primitive Ghost Moths, morphology and taxonomy
of the Australian genus Fraus Walker (Lepidoptera: Hepialidae 5. lat.). Monographs on
Australian Lepidoptera, Vol. 1. CSIRO.

Packard, A. S. 1895. On the larvae of the Hepialidae. J. New York Entomol. Soc. 3:69-73.

614

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Pitelka, L. F. and D. J. Raynal. 1989. Forest decline and acidic deposition. Ecology 70:2-10.

Reiners, W. A. and G. E. Lang. 1979. Vegetational patterns and processes in the balsam fir
zone, White Mountains, New Hampshire. Ecology 60:403-417.

SAS Institute. 1985. SAS User’s Guide: Basics, Version 5 Ed. SAS Institute, Cary, N.C.

Siccama, T. G., M. Bliss and H. W. Vogelmann. 1982. Decline of Red Spruce in the Green
Mountains of Vermont. Bull. Torrey Bot. Club 109:162-168.

Silver, W. L., T. G. Siccama, C. Johnson and A. H. Johnson. 1991. Changes in red spruce
populations in montane forests of the Appalachians 1982-1987. Am. Mid. Nat. 125:
340-347.

Smith, W. H. 1991. Air pollution and forest damage. Chem. Engin. News 69:30-43.

Stehr, F. W. 1987. Order Lepidoptera. Pages 288-305 in: F. Stehr (ed.), Immature Insects.
Kendall-Hunl, Dubuque, Iowa.

Tobi, D. R., W. E. Wallner and B. L. Parker. 1989. The conifer swift moth, Hepialus gracilis,
and spruce-fir decline. Pages 361-363 in: Proceedings of the US-FGR Symposium: Effects
of Atmospheric Pollutants on the Spruce-Fir Forests of the Eastern United States and
Federal Republic of Germany. USDA Forest Service, Northeastern For. Exp. Sta. Gen.
Tech. Rep. Broomall, PA.

Tobi, D. R., J. G. Leonard, B. L. Parker and W. E. Wallner. 1992. Survey methods, distri-
bution, and seasonality of Korscheltellus gracilis (Lepidoptera: Hepialidae) in the Green
Mountains, Vermont. Environ. Entomol. 21:(in press).

Viette, P. 1949. Contribution a l’etude des Hepialidae (12c note). Genres et synonymie.
Lambillionea 1949:101-104.

Vogelmann, H. W., T. D. Perkins, G. J. Badger and R. M. Klein. 1988. A 21 -year record of
forest decline on Camels Hump, VT USA. Eur. J. For. Pathol. 18:240-249.

Wagner, D. L. 1987. Hepialidae. Pages 347-349 in: F. Stehr (ed.), Immature Insects. Kendall-
Hunt, Dubuque, Iowa.

Wagner, D. L. 1988. Taxonomic status of Korscheltellus Bomer in North America (Lepidop-
tera: Hepialidae). J. New York Entomol. Soc. 96:345-354.

Wagner, D. L. and N. B. Tindale. 1988. An appraisal of Gazoryctra Hubner (Hepialidae) and
description of a new species from Arizona and New Mexico. J. Lepidopterists’ Soc. 43:
204-212.

Wagner, D. L., D. R. Tobi, B. L. Parker, W. E. Wallner and J. G. Leonard. 1989. Immature
stages and natural enemies of Korscheltellus gracilis (Lepidoptera: Hepialidae). Ann.
Entomol. Soc. Am. 82:717-724.

Wagner, D. L., D. R. Tobi, W. E. Wallner and B. L. Parker. 1991. Korscheltellus gracilis
(Grote): a pest of red spruce and balsam fir roots (Lepidoptera: Hepialidae). Can. Entomol.
123:255-263.

Yasuda, T. and K. Abe. 1986. Endoclita hosei Tindale (Lepidoptera: Hepialidae) attacking
Eucalyptus in Sabah, with descriptions of the immature and imaginal stages. Appl.
Entomol. Zool. Tokyo 21:41 7 — 423.

Received 4 September 1991; accepted 30 April 1992.

J. New York Entomol. Soc. 1 00(4):6 1 5—620, 1992

A NEW SPECIES OF ORCYA FROM THE MARONI
RIVER BASIN OF FRENCH GUIANA
(LEPIDOPTERA: LYCAENIDAE)

Kurt Johnson

Department of Entomology, American Museum of Natural History,

Central Park West at 79th Street, New York, New York 10024

Abstract. — First report of pan-Neotropical Orcya from the Guyana Shield is Orcya snyderi,
a distinctive new species described from recently disturbed riparian rain forest margin, upper
Maroni River, French Guiana. Sister genus Noreena has a regional endemic of similar
distribution.

I recently revised the Neotropical “ Thecla orcynia Group” (Draudt, 1919) placing
twelve pan-Neotropical species in the new genus Orcya (Johnson, 1990) and related
this genus to a number of sister- and outgroups by numerical cladistic analysis
(Johnson, 1989a, 1990). No member of Orcya was known from the Guyana Shield
region, though a species endemic to this area occurs in a sister genus ( Noreena
guianivaga Johnson) (Johnson, 1989a, b).

Calvin Snyder (volunteer field worker, American Museum of Natural History
[AMNH]) recently returned from extensive field work along the Maroni River in
French Guiana and his collections included a female of a distinctive new species of
Orcya. Subsequently, a male and five females from French Guiana were located at
the Museum National d’Histoire Naturelle (Paris) (MNHN). This new species is
described below, following on the generic diagnosis of Orcya (Johnson, 1990). For
brevity, the abbreviations DFW, DHW (dorsal fore- and hindwings) and VFW, VHW
(ventral fore- and hindwings) are used.

Orcya snyderi, new species
Figs. 1, 2

DIAGNOSIS. Male dorsal iridescence limited on FW to baso-medial cells CuAl-
CuA2, HW to centro-medial patch; female DFW, DHW non-iridescent dark gray
(congeners brightly iridescent in both sexes). Unique pattern elements: (1) DHW
black marginal band basad of greatly elongate tails (terminus vein CuA2), (2) VFW,
VHW each with (i) prominent black submarginal band, (ii) black medial band (costa
to cell M3) basad of, and broadly paralleling, usual white wing band of genus. Male
genitalia with diminutive, elliptically shouldered valvae and thin, elongate, vincular
structures; female genitalia with anterior and posterior elements of ductus bursae
both undulate.

DESCRIPTION. Male. Upper surface of wings: DFW lacking androconial brand
as typical of genus, FW with dark azure blue iridescence confined to baso-medial
areas of cells CuAl-CuA2 and to a centro-medial HW patch; tails on known specimen
broken (short tail apparent only at terminus vein CuA2, see Female below), anal
lobe with bold orange spot. Under surface of wings: ground dark gray, FW with faint

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Fig. 1 . Adults of Orcya snyderi, new species. A. holotype female (upper surface, left; under
surface, right). B. allotype male (same view).

black postmedial and submarginal bands (known specimen worn); HW with basal
area charcoal gray with slight white slash in discal cell, medial area with meandering
white band, broadly bordered basally with black, angled in wide “W”-shape before
the anal margin; submarginal area with jagged white band bordered distally with
black extending from costa to anal margin and surrounding submarginal orange spots
in cell CuAl and at base of anal lobe. Length of forewing: 1 1.5 mm (allotype).

Female. Upper surface of wings: ground dull gray, FW margins and apices black,
HW with black marginal line; HW margin with short black tail, terminus vein CuAl,
elongate white-tipped black tail, terminus vein CuA2, anal lobe at anal margin with
bold orange spot. Under surface of wings: ground gray, FW with (i) white postmedial
line, bordered basally by black, extending in straight line from costa to cell M3 then
basally angled to inner margin; (ii) white submarginal line, bordered basally with
black, extending in an arc from costa to cell M3 (see Remarks); HW with (i) medial
band distally white, basally black, arched widely from costa to cell M3, thereafter
jagged in wide “W”-shape before anal margin; (ii) finely jagged, basally white-suffused,
black submarginal band extending from costa to anal margin and surrounding sub-
marginal orange orbicular markings in cell CuAl and at base of anal lobe. No white
discal slash that typifies other species of genus. Length of forewing: 1 1.5 mm (ho-
lotype); 1 1.5- 12. 5(x 1 1.9) (five paratypes).

Male genitalia. Figure 2B. Generally typical of genus (Johnson, 1990: figs. 6, 8)
but with ventral vincular parts elongate, slim; valval ventrum parabolic in bilobed
area (2B,a,l), terminating in relatively short tapered caudal extensions (2B,a,2). Ae-
deagus elongate, exceeding rest of genitalia length by about two-fifths; caecum com-
prising about one-fourth aedeagal length.

Female genitalia. Figure 2A. Generally typical of genus (Johnson, 1990: figs. 7, 9)
but distinctive as follows: anterior and posterior elements of ductus bursae undulate,
length of former about one-half that of latter, posterior element with produced ventral
lip; cervix bursae with hood more prominent than in congeners; corpus bursae with
paired signa more robust and terminally pronged than in any congener.

TYPES. Holotype female (Fig. 1A), FRENCH GUIANA, Maripasoula; environs

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617

Fig. 2. Genitalia of Orcya snyderi, new species. A. holotype female: a, ductus bursae, ventral
view; b, lamella! keel, lateral view; c, corpus bursae signum, lateral view; d, cervix bursae [1]
with adjoining ductus seminalis [2] and anterior terminus of ductus bursae [3], all in lateral
view. B. allotype male: a, genitalia with aedeagus removed (valval bilobed area [1], caudal
extension [2]), ventral view; b, valvae, lateral view; c, aedeagus, lateral view; d, aedeagus
terminus, ventral view.

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of Maripasoula, 125 m, 31 March 1991, leg. C. Snyder, deposited AMNH. Allotype
male (Fig. IB) labelled “Guyane Francaise, Rives [=“banks of’] de Maroni, E. Le
Moult 1909,” deposited MNHN. Paratypes. MNHN: five females labelled “Guyane
Francaise, Rives de Maroni, E. Le Moult 1909” (2) and “Guyane Francaise, Maroni,
E. Le Moult 1909” (3).

DISTRIBUTION. Currently known from the Maroni River basin in French Gui-
ana/Surinam (see Remarks).

REMARKS. Habitat. Only the holotype has precise field data. Snyder (pers. comm.)
reports that the specimen was collected in the morning in moist secondary riverbank
forest and scrub, in an area of high annual rainfall (up to 3,000 mm) broken by a
2Vi month dry season. Time of collection coincided with the so-called “March sum-
mer” of the region, a period of scant rainfall at the midpoint of the November to
July rainy season. Before disturbance the habitat supported lowland rain forest typical
of the Guyana Shield on both sides of the river (the east bank now being in Surinam).

Orcya snyderi and historical names in “Theda.” Because wing markings of the O.
snyderi female might be construed as resembling the short original description (“OD”)
of Thecla imma Prittwitz (1865), explanation is needed concerning why I do not
associate this name with Orcya. Thecla imma was described from a single female,
now lost, from Corcovado, Rio de Janeiro, Brazil (Bridges, 1988) and the name has
been variously applied by curators and catalogers. Draudt (1919) placed T. imma in
his Thecla “strephon- Group,” a diverse assemblage of species showing a wide “W”-
shaped bend in the VHW band and reduced DFW structural color (Draudt, 1919,
pi. 151 d— g). Draudt (1919:781) also noted similarities between the OD of T. imma
and members of his ” orcynia-G roup,” some of which I included in Orcya (Johnson,
1990). Early lycaenid specialists W. P. Comstock and E. I. Huntington, curating at
the AMNH, attached the name T. imma to a variety of Central and South American
specimens (some of the latter noted as “new species nr. imma”). I included mor-
phological data from these specimens in data matrices for numerical cladistic analysis
(Johnson, 1989a, 1990). The final rooted tree (Johnson, 1989a, fig. 8; 1990, fig. 1)
included some South American specimens as “undescribed Outgroup X” (Johnson,
1989a, =“Group 3” Johnson, 1990). The other specimens, including the Central
American ones, were rejected from the Orcya ingroup and placed with specimens of
Thecla orios Godman & Salvin (1879-1901 [1887]) based on characters of its ho-
lotype, a Guatemalan male (Johnson, 1990 “Group 1,” an outgroup at the base of
the cladogram). I thus consider “ Thecla imma ” objectively ambiguous, its OD being
applicable to any number of specimens of divergent morphology. It is particularly
noteworthy that an arbitrary application of Thecla imma, based on its OD and a
regional view of its type locality, would likely include specimens of either “Outgroup
X” or Contrafacia Johnson (1989a). As can be seen from figure 7A-C in Johnson
(1989a), the morphology of these latter groups differs greatly from that of O. snyderi
and its congeners discussed below. I have proposed elsewhere (Johnson, 1991a) that
certain widely used historical names, ambiguous in a revisionary context, may even-
tually require formal suppression under the ICZN Code.

Intrageneric relations. Distinctive characters in both the male and female of O.
snyderi are intriguing. Unique wing characters include: (1) reduced and absent DFW,
DHW surface structural color in males and females, respectively; (2) bold under
surface submarginal bands on the fore- and hindwing; (3) small size (congeners

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NEW ORCYA

619

generally 13.5-17.0 mm [15.0-17.0 mm in the common and widespread taxa, 12.0-
14.0 mm in the few austral species]). Because of these characters, O. snyderi is not
readily associated with the genus from a superficial view.

Features appearing somewhat like other congeners require careful consideration.
The distinctive VH W pattern, for instance, cannot be considered informative. Austral
species O. larseni (Lathy) and O. obliqua Johnson also have VHW medial bands
departing from the thin, extremely jagged pattern typifying most of the genus but
neither species’ band resembles the robust band of O. snyderi. Similarly, southeastern
Brazil xerophile O. cat hah na (Draudt) shows reduction of the VHW discal slash but
it is never completely absent in fresh specimens as is the case in O. snyderi.

Truly compelling similarities involve other wing characters and some structural
features emphasized by Johnson (1990) in the generic analysis. Most prominent are
several wing and genitalic characters shared by O. snyderi, O. hewitsoni Johnson
(humid montane forest, northern Ecuador south to central Peru) and O. anthracea
(Hewitson) (humid montane forest, southeastern Brazil). These three taxa show bold
hindwing medial bands marked near the anal margin with an emphatic “W”-shaped
element and bright orange orbs. In the female genitalia the ductus bursae is undulate
and the respective anterior and posterior elements of similar size. Typifying the
tendency toward autapomorphy in male genitalia of Orcya (Johnson, 1990, figs. 6,
8), details in O. snyderi (ventral valval and vincular shape, elongate aedeagus) do
not argue for or against any particular intrageneric relation.

Biogeography. Affinity of O. snyderi to O. hewitsoni and O. anthracea mirrors
biogeographic relations in several other groups of butterflies where the Amazon basin
segregates sister species into respective Guyana Shield, western Andean, and south-
east Brazilian distributions. This pattern occurs in sister genus Noreena (Johnson,
1989a) and numerous other Theclinae (Johnson, 1991a, b). Regarding an Andean/
Maroni River basin sister species relationship, perhaps the most oft-cited example
is Heraclides maroni (Moreau) and H. rhodostictus (Butler and Druce) (Papilionidae),
where the superficial resemblance is so striking that first reported specimens of the
former were thought to be mislabelled examples of the latter (D’Abrera, 1981; John-
son and Rozycki, 1986; Johnson and Matusik, 1989).

Etymology. Patronym for Calvin Snyder who collected the holotype.

ACKNOWLEDGMENTS

I thank Calvin Snyder for bringing the holotype to my attention and Dr. Jacques Pierre
(MNHN) for locating specimens at the MNHN.

LITERATURE CITED

Bridges, C. A. 1988. Catalogue of Lycaenidae & Riodinidae (Lepidoptera: Rhopalocera).

Published by the author, Urbana, Illinois, vii + 811 pp.

D’Abrera, B. 1981. Butterflies of the Neotropical realm. Part 1. Papilionidae and Pieridae.

Landsdowne Editions, East Melbourne, 172 pp.

Draudt, M. 1919. Thecla. Pages 794-81 1 in: A. Seitz (ed.), Macrolepidoptera of the World.

Alfred Kemen Verlag, Stuttgart, Vol. 5 (“Vol. II”), pp. 593-1 139, Vol. 5 plates (“Vol.

II Plates”), 194 pis.

Godman, F. and O. Salvin. 1879-1901 [1887], Biologia Centrali-Americana. Zoology: Insecta,

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Lepidoptera. 3 volumes. Vol. 2 (Pieridae, Papilionidae, Lycaenidae, Hesperidae) 1 887—
1901. Taylor & Francis, London, 782 pp.

Johnson, K. 1989a. A revisionary study of the Neotropical hairstreak butterfly genus Noreena
and its new sister genus Contrafacia (Lepidoptera: Lycaenidae). J. New York Ent. Soc.
97(1): 1 1-46.

Johnson, K. 1989b. The hairstreak butterfly genus Noreena (Lycaenidae, Theclinae) and a
new species from western Peru. Insecta Mundi 3(3):247-248.

Johnson, K. 1990. The new hairstreak butterfly genus Orcya, a revision of the Neotropical
“ Thecla ” orcynia assemblage (Lepidoptera: Lycaenidae). J. New York Ent. Soc. 98(1):
50-87.

Johnson, K. 1991a. Neotropical hairstreak butterflies: genera of the “Calycopis/Calystryma
Grade” of Eumaeini (Lepidoptera, Lycaenidae, Theclinae). Reports, Mus. Nat. Hist.,
Univ. Wisconsin (Stevens Point) 21:1-127.

Johnson, K. 1991b. Cladistics and the biogeography of two trans-Caribbean hairstreak but-
terfly genera: Nesiostrymon and Terra (Lepidoptera, Lycaenidae). Amer. Mus. Novitates
301 1:1-43.

Johnson, K. and D. Matusik. 1986. Additional notes on type specimens of little-known
Papilionidae. Papilio International 6(2):45 1-457.

Johnson, K. and R. Rozycki. 1 986. A new species of the anchisiades group of Heraclides from
Venezuela (Lepidoptera: Papilionidae). J. New York Ent. Soc. 94(3):383-393.

Prittwitz, O. F. W. L. von. 1865. Beitrag zur Fauna des Corcovado. Stett. Ent. Zeit. 26:123-
143, 307-325.

Received 5 September 1991; accepted 25 February 1992.

J. New York Entomol. Soc. 1 00(4):62 1 —633, 1992

NOVEL TAX A OF TRICHOGRAMMA FROM THE
NEW WORLD TROPICS AND AUSTRALIA
(HYMENOPTERA: TRICHOGRAMMATIDAE)

John D. Pinto

Department of Entomology, University of California,

Riverside, California 92521

Abstract. —Three new species of Trichogramma from the New World tropics and one new
species from Australia are described. These taxa differ from all congeners by the unjoined
funicular and club segments of the male antenna, and the two-segmented maxillary palp. The
New World species also are distinguished by the unique structure of the aedeagus. All four new
species are retained tentatively within the nominate subgenus of Trichogramma. Their rela-
tionship to other species of the genus is discussed.

The study of recent Trichogrammatidae collections from the Neotropics and Aus-
tralia has revealed four new species of Trichogramma which extend considerably the
known anatomical variation in this cosmopolitan genus of insect egg parasitoids.
These species share certain derived features that define Trichogramma, but are be-
lieved to represent primitive lineages that diverged before the attainment of fusion
of the antennal flagellomeres in males. Pending an evaluation of phylogenetic rela-
tionships in the genus these taxa are tentatively placed in two informal species groups
within the nominate subgenus, the Lachesis and Primaevum groups.

In this paper the new species groups and included species are described. These
taxa are compared to other elements of Trichogramma, and their phylogenetic po-
sition and taxonomic treatment are discussed. Also, terms are utilized for the male
genitalia that conform to those adopted for other genera of Trichogrammatidae.

TERMINOLOGY

The male genitalia comprise the most important character complex in Tricho-
gramma. Species identifications were virtually impossible prior to Nagarkatti and
Nagaraja (1968) in which this rich source of variation was first considered. Because
of questions of homology, however, these authors introduced a unique terminology
for certain genitalic structures in Trichogramma and Trichogrammatoidea (Nagar-
katti and Nagaraja, 1968, 1971, 1977; Nagaraja, 1978). This was adopted by other
authors (e.g., Ertle and Davis, 1975; Pinto and Oatman, 1985; Pinto et al., 1978,
1989; Schulten and Feijen, 1978, 1982; Vincent and Goodpasture, 1986; Voegele
and Pintureau, 1982) and became standard in studies of both genera. In contrast,
Viggiani (1971), and Viggiani and Laudonia (1989) utilized a terminology for all
Trichogrammatidae which stems from generalized works on Hymenoptera genitalia
(Snodgrass, 1941, 1957; Domenichini, 1953). This is largely employed here, and it
is suggested that other investigators working with Trichogramma and Trichogram-
matoidea also adopt these more conventional terms. Maintaining two distinct ter-
minologies within the family is awkward and potentially confusing.

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The descriptions of male genitalia in the new species treatments, for the most part,
are consistent with Viggiani and Laudonia (1989). The structures, acronyms (also
see Figs. 10, 11, 13) and, where appropriate, counterparts (in brackets) used by
Nagarkatti and Nagaraja (1968, 1971) are as follows:

Genital capsule (GC)— the entire genitalia except the aedeagus; phallobase (PB) [=
gonobase, GB]; dorsal aperture of the phallobase (DA); dorsal lamina (DLA) [=dorsal
expansion of gonobase, DEG & DEG IT; parameres (PM) [=gonoforceps, GF]; vol-
sellar digiti (VS) [=chelate structures, CS]; intervolsellar process (IVP) [=median
ventral projection, MVP]; intervolsellar bridge (IB)— surface between the base of the
volsellae, includign the IVP; ventral ridge (VR) [=chitinized ridge, CR]; ventral
processes (VP) — paired papillae or tubercle-like structures lateral to the ventral ridge,
at or near base of IVP [^lateral tubercles (Nagaraja, 1978), and paired ventral pro-
tuberances (Nagaraja and Nagarkatti, 1 969)]. GC, IB, VR and VP were not considered
by Viggiani and Laudonia (1989).

Acronyms terminating with L, D or W signal length, distance and width mea-
surements, respectively. Included are HTL (hind tibial length), FWW (maximum
forewing width), FWL (forewing length, taken from tegula to wing apex), GW (max-
imum width of genital capsule), GL (maximum length of genital capsule), AD (apical
distance of genital capsule: from base of volsellae and IVP to apex of PM), BD (basal
distance of genital capsule: GL minus AD, or distance from base of volsellae and
IVP to base of genital capsule), AL (length of aedeagus, including apodemes), OL
(maximum length of ovipositor).

Acronyms for two types of sensilla on the antenna are BCPS (=basiconic peg
sensilla), and PS (=linear, placoid sensilla). Sensilla formulae in descriptions refer to
these sensilla on each of the funicular and club segments. Funicular and club segments
are referred to as FI -2, and Cl -3, respectively.

Antennal, forewing and genitalic features referred to in the descriptions are illus-
trated below. Hindwing and scutellar features are illustrated in Pinto et al. (1978).

DEFINING FEATURES OF TR ICHOGRA MM A

Trichogramma is easily distinguished from other Trichogrammatidae. The fore-
wing is relatively broad with distinct vein tracks including an RSt; the stigmal vein
is well developed but not abruptly delimited from the marginal vein — instead it
gradually curves into the latter, resulting in the characteristic “sigmoid” venation
(Fig. 7). The female antenna consists of two funicular segments and a single club
segment. The male genitalia have a dorsal lamina and the aedeagus is not fused to
the genital capsule.

Trichogramma is believed to be closest to Trichogrammatoidea (Nagarkatti and
Nagaraja, 1 977; Nagaraja, 1978). They share the “sigmoid” venation in the forewings,
the antennal structure of females is identical, and they have similar male genitalia.
Although similar, Trichogrammatoidea lacks the RS, vein track in the forewing as
well as the dorsal lamina of the genital capsule. The latter feature, so far as known,
is unique in the family (see Viggiani 1971, 1984) and is the derived feature defining
T richogramma.

Until now, Trichogramma was also distinguished from Trichogrammatoidea and
other genera by its male antenna. In Trichogrammatoidea the two funicular and three
club segments are distinct. In virtually all species of Trichogramma, these five seg-

Figs. 1-7. Male antennae, maxillary palp, and forewing venation in the Lachesis and Pri-
maevum groups of Trichogramma. 1. Trichogramma lachesis, antenna (left). 2. Trichogramma
clotho, funicle. 3. Trichogramma atropos, funicle. 4. Trichogramma primaevum, antenna (left).
5. Trichogramma lachesis, maxillary palp. 6. Trichogramma clotho, maxillary palp. 7. Trich-
ogramma lachesis, forewing venation (arrow points to submarginal seta).

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ments, as well as the second anellus, are consolidated into a single, elongate segment.
The only previous exceptions known were species in the subgenus Trichogrammanza
Carver which has the two funicular (but not the club) segments unjoined (Carver,
1978; Oatman and Pinto, 1987).

The four new species described below necessitate a modest redefinition of Trich-
ogramma. In these species, forewing venation and vein tracks, and male genital
structure (with the dorsal lamina) are typical of the genus. However, unlike all con-
geners, the two funicular and three club segments are unjoined as in Trichogram-
matoidea (Figs. 1, 4). In addition, the maxillary palp is two-segmented (Figs. 5, 6).
In all congeners, as well as in Trichogrammatoidea, the palp is one-segmented. It
should be noted that although the number of flagellomeres in these new species agrees
with Trichogrammatoidea, antennal structure (e.g., shape, sensilla type and number)
does not. Instead, in all respects, their antennae more closely resemble other tricho-
grammatine genera, particularly Australufens and Trichogrammatomyia. For ex-
ample, the club is compact, suboval in shape, and C3 is subconical and pointed
apically. In Trichogrammatoidea the club is more elongate, the segments are more
loosely joined, and C3 is rounded apically (Doutt and Viggiani, 1968). The phylo-
genetic implications of these new species and the justification for placing them in
species groups in the nominate subgenus is discussed following the descriptions.

LACHESIS GROUP

As in other Trichogramma except as follows; Male antenna with two funicular and
three club segments; C3 subconical, pointed apically; funicular segments lacking linear
PS. Aedeagus (Fig. 12) apparently bilobed with phallotreme more extensive than in
congeners, extending a distance from apex along ventral surface; aedeagus longer
than genital capsule. Maxillary palp two-segmented (Figs. 5, 6). Forewing with a seta
near posterior border of marginal vein (Fig. 7). Hindwing without a posterior vein
track.

Notes. The Lachesis Group contains three species from Central America and north-
ern South America. The female is indistinguishable from congeneric females except
for the structure of the maxillary palp, and wing setation.

Etymology / of specific names. After the three Fates of Greek mythology.

Trichogramma lachesis, new species
Figs. 1, 5, 7, 10-12

Description based on two males and one female, slide-mounted in Canada balsam.
Unless referring to the unique female, quantitative data are means with an n of 2 or
3, as appropriate; range is reported if specimens differ considerably.

Diagnosis. Trichogramma lachesis is distinguished from other Lachesis Group
species by the very broad ventral ridge (VR) and the absence of an intervolsellar
process (I VP) (Fig. 1 1).

Description. Color light yellow brown; 0.47 mm in length; HTL = 0.12-0.13 mm
(males), 0.14 mm (female). Maxillary palp with segment II short, obsolescent (Fig.
5). Forewing broad, 0.20 mm; FWW/FWL = 0.57; longest fringe setae 0.16 FWW;
relatively few setae between major vein tracks, 8-13 between 4th and 5th tracks.
Hindwing without anterior or posterior tracks, but two small, widely spaced setae

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625

Figs. 8-9. Thoracic microsculpturing in the Lachesis Group of Trichogramma. 8. Tricho-
grarnma atropos. 9. Trichogramma clotho.

slightly posterior to basal half of middle track may represent posterior track setae.
Mesoscutum and scutellum smooth, microsculpturing indistinct; scutellum with an-
terior pair of setae elongate, stout, ca. 0.6 length of posterior pair.

Male. Antenna (Fig. 1) with funicle 0.70 length of club, subequal to length of scape;
FI subquadrate, apical socket acentrically positioned; F2 1.6 as long as wide, 1.3 as
long as FI, asymmetrically tapered from middle to relatively narrow apex, only
slightly narrower than F 1 at basal half; length of club segments 12:14:11, respectively;
C2 slightly wider than Cl; C3 narrower, subconical; longest setae on flagellum 2.3-
2.7 width of FI; BCPS large, bulbous, formula 2-2-2- 1-1; PS extending considerably
beyond apex of segments, conspicuously so on C3, formula 0-0- 1-1-2.

Genital capsule (Figs. 10, 1 1) 0.37 as wide as long, tapering moderately from widest
point to apex; PM evenly curved, convergent to apex, appendiculate at apex; AD
short, only 0.16 GL; DA large, its length 0.72 GL; DLA not lobed or notched at
base, extremely broad its entire length, width of posterior extension subequal to that
across parameres, broadly arcuate at apex, extending to apical % of AD; VS abruptly
produced ventromedially at apical half, very narrowly separated from one another,
extending slightly beyond apex of DLA, occupying ca. 0.7 AD; I VP absent; IB narrow;
VR conspicuous, extremely broad, elongate, occupying ca. Vi BD; VP not visible.
Aedeagus length (Fig. 12) 1.10 GL, 0.90 HTL; apodemes 0.51 AL; aedeagus widest
at apodemes.

Female. Antenna with FI and F2 subquadrate, about as broad as long, each with
1 BCPS on dorsal surface; club length subequal to length of scape, with two BCPS
on dorsal surface (at basal and apical lA, respectively) and five PS. Ovipositor mod-
erately long, OL/HTL = 1.12.

Types. Male holotype. COSTA RICA. Guanacaste: Santa Rosa National Park,
“Hacienda 3-0”; v-24/vi-14-1986. Allotype female, same data as holotype. Paratype

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Figs. 10-14. Male genitalia in the Lachesis Group of Trichogramma. 10. Trichogramma
lachesis, genital capsule (dorsal) (VS partially visible in dorsal view but not shown). 1 1. Trich-
ogramma lachesis, genital capsule (ventral). 12. Trichogramma lachesis, aedeagus (ventral).

1992

NOVEL TRICHOGRAMMA

627

male, same data as holotype except “Hacienda 2-C”; i-3 l/ii-2 1-1987. The types are
deposited as follows: holotype and allotype— United States National Museum; para-
type— University of California, Riverside.

Santa Rosa is the former name of Guanacaste National Park. According to infor-
mation received from Ian Gauld and Paul Hanson (in litt.) the types were collected
in two different Malaise traps (one in shade, the other in a clearing) placed on the
plateau (250-300 m) immediately west of park headquarters in young scrubby wood-
land.

Notes. The female allotype was collected with the holotype. Their association is
based on correspondence of all comparable characters including color, structure of
the maxillary palp, scutellar setal length, thoracic sculpturing, and hindwing setation.

Trichogramma clotho, new species
Figs. 2, 6, 9, 13

Description based on the unique male holotype (slide-mounted in balsam).

Diagnosis. Trichogramma clotho is distinguished from other Lachesis Group spe-
cies by the distinctive rugulose microsculpturing on the thorax (Fig. 9), and the more
elongate and apically tapered second funicular segment (Fig. 2).

Description. Color dark brown; 0.55 mm in length; HTL = 0.14 mm. Forewing
broad, 0.23 mm in width; FWW/FWL = 0.57; longest fringe setae 0.15 maximum
wing width; setation on disk similar to lachesis, 5 setae between 4th and 5th tracks,
RS, consisting of only 2 setae. Maxillary palp (Fig. 6) with segment II longer, more
distinct than in lachesis. Hindwing without anterior and posterior tracks. Mesoscutum
and scutellum with distinct, rugulose microsculpturing (Fig. 9); scutellum with an-
terior pair of setae elongate, stout, ca. 0.6 length of posterior pair (type has a super-
numerary posterior seta on left side).

Antenna with funicle 0.70 length of club and subequal to length of scape; FI
subrectangular, 0.8 as long as wide, apical socket acentrically positioned; F2 elongate,
considerably narrower and longer than FI (Fig. 2), strongly asymmetrically tapered
from basal Vs to apex, 2.3 as long as wide, 1.9 as long as FI; length of club segments
14:14:18, respectively; C2 slightly wider than Cl, C3 apparently considerably nar-
rower and linear (partially collapsed in type); longest setae on flagellum 3.0 width of
FI; BCPS shape and formula as in lachesis ; PS extending considerably beyond apex
of segments, formula 0-0-0- 1-2.

Genital capsule (Fig. 13) 0.45 as wide as long; AD short, 0.15 GL; PM straight,
narrow, convergent, apparently not appendiculate; DA large, its length 0.69 GL; DLA
as in lachesis, extending 0.4 length of AD; VS slightly less strongly produced ven-
tromedially than in lachesis, occupying ca. 0.8 AD; I VP present, moderately long,
robust, subtriangular, extending 0.3 AD; IB wider, slightly greater than basal width
of IVP; VR narrow, occupying 0.3 BD; VP short, slightly protuberant, at base of

showing bilobed structure and elongate phallotreme at apex. 13. Trichogramma clotho, genital
capsule (ventral). 14. Trichogramma atropos, genital capsule (dorsal) (VS partially visible in
dorsal view but not shown). DA = dorsal aperature of phallobase; DLA = dorsal lamina; IVP
= intervolsellar process; PM = parameres; VR = ventral ridge; VS = volsellar digiti.

628

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

IVP. Aedeagus length 1.21 GL; AL/HTL = 0.79; apodemes occupying 0.49 AL,
aedeagus not wider at apodemes.

Female. Unknown.

Type. Holotype male. COSTA RICA. Puntarenas : Golfo Dulce, 13 km S Rincon,
10 m elev.; ii/iii- 1989; Malaise trap; P. Hanson coll. Deposited in the United States
National Museum.

Trichogramma atropos, new species
Figs. 3, 8, 14

Description based on the unique male holotype, slide-mounted in Canada balsam.

Diagnosis. Trichogramma atropos is distinguished from other Lachesis Group
species by the subquadrate second funicular segment (Fig. 3), the presence of an
incomplete anterior vein track in the hindwing, and the triangular posterior extension
of the dorsal lamina (DLA) (Fig. 14).

Description. Color yellow brown; 0.48 mm in length; HTL = 0.15 mm. Maxillary
palp II as in lachesis. Forewing broad, 0.20 mm in width, FWW/FWL = 0.54; longest
fringe setae 0. 19 wing width; disk moderately densely setate, with 7 linearly arranged
setae between 4th and 5th tracks. Hindwing with anterior track of 3 setae; posterior
track absent. Mesoscutum and scutellum smooth, microsculpturing indistinct; scu-
tellum with anterior pair of setae elongate, stout, ca. 0.7 length of posterior pair
(Fig. 8).

Antenna with funicle 0.70 length of club, subequal to length of scape; FI subquad-
rate; F2 (Fig. 3) 1.2 as long as wide, 1.2 as long as FI, asymmetrically tapered only
at apex of segment; length of club segments 12:12: 1 1, respectively; C2 slightly wider
than Cl; C3 considerably narrower, subconical; flagellar setae relatively short and
stout, abruptly tapered at apex, longest setae 1.8 width of FI; BCPS large, bulbous,
formula 2-2-2-2-1; PS formula 0-0-0- 1-2.

Genital capsule (Fig. 14) 0.43 as wide as long, gradually tapering from widest aspect
to apex; PM relatively straight, evenly convergent to apex, apparently not appendicu-
late; AD 0.27 GL; DA relatively narrow, its length 0.58 GL; DLA not obviously
notched or lobed at base, its posterior extension broadly subtriangular, tapering evenly
from base to bluntly rounded apex, extending to apex of VS; VS asymmetrically
narrowed apically, produced ventromedially, occupying 0.70 AD; IVP present but
inconspicuous, short, linear, occupying only 0. 1 AD; IB narrow; VR difficult to discern
in type, apparently narrow and relatively short; VP not protuberant, positioned
considerably basal to IVP. Aedeagus length 1.11 GL, 0.70 HTL, apodemes 0.41 AL;
aedeagus not wider at apodemes.

Female. Unknown.

Type. Holotype male. VENEZUELA. Merida: La Montana Station, 2,435 m elev.;
iv-3/ 1 3- 1 988; Malaise trap; A. T. Finnamore & C. E. Baxfield, collrs. Type deposited
in United States National Museum.

PRIMAEVUM GROUP

As in the Lachesis Group except as follows: FI of male antenna with a linear PS.
Aedeagus not bilobed, phallotreme restricted to apex. Hindwing with an incomplete
but distinct posterior vein track.

1992

NOVEL TR1CHOGRAMMA

629

Figs. 15-16. Male genitalia in the Primaevum Group of Trichogramma. 1 5. Trichogramma
primaevum, genital capsule (dorsal). 16. Trichogramma primaevum, aedeagus (ventral).

Notes. This group includes a single species from Queensland, Australia. The female
is unknown. Male antennal segmentation, maxillary palp structure, and marginal
vein setation are as in the Lachesis Group. Aedeagus structure, however, is as in all
other Trichogramma.

Trichogramma primaevum, new species
Figs. 4, 15, 16

Description based on the unique male holotype, slide mounted in Canada balsam.

Diagnosis. Trichogramma primaevum is distinguished from members of the Lach-
esis Group by the simple aedeagus and its shorter phallotreme (Fig. 1 6), and by the
presence of a placoid sensillum (PS) on the second funicular segment of males. The
segmented male club separates the species from all other congeners.

Description. Color yellow brown, legs and abdomen lighter than rest of body; 0.60
mm in length; HTL = 0.15 mm. Maxillary palp with a short second segment. Forewing
broad, 0.26 mm wide; FWW/FWL = 0.60; stigmal vein relatively short, robust, with
a slightly constricted neck; fringe unusually short, longest setae only 0.06 FWW; 18
setae between 4th and 5th tracks. Hindwing with posterior track of 5 setae extending
0.45 distance from hamuli to apex; anterior track complete to apex. Mesoscutum
and scutellum relatively smooth, microsculpturing indistinct; scutellum with pos-
terior pair of setae short (0.0 1 6 mm long), only ca. half the length of those in congeners;
anterior setae slightly shorter.

Antenna (Fig. 4) with funicle 0.58 length of club, 0.73 length of scape; scape
somewhat broader than in congeners; FI subquadrate, apical socket not acentrically
positioned; F2 1 .4 as long as wide, 1 .4 as long as FI , asymmetrically tapered at apex;

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Table 1 . Characters distinguishing species of the Lachesis and Primaevum groups of Trich-
ogramma.

Species

Characters1 2

lachesis

clotho

atropos

primaevum

1.

0

1

0

0

2.

0

0

0

1

3.

0

0

1

2

4.

1

2

0

0

5.

0

1

0

0

6.

0

0

0

1

7.

1

1

1

0

8.

1

1

1

0

9.

1

0

0

0

10.

0

0

1

2

11.

0

2

1

1

12.

1

0

0

0

13.

1

1

1

0

1 Characters: 1 . Dorsum of thorax smooth, with indistinct microsculpturing (0) (Fig. 8); with
distinct, rugulose microsculpturing (1) (Fig. 9). 2. Hindwing with posterior vein track absent

(0) , present (1). 3. Hindwing with anterior vein track absent (0), incomplete (1), complete to
apex (2). 4. F2 of male antenna slightly (0) (Fig. 3), moderately (1) (Fig. 1), strongly (2) (Fig.
2), tapered to apex. 5. Length of F2 of male antenna less than 1.5 (0) (Fig. 1), or almost 2.0 (1)
(Fig. 2), that of FI. 6. F2 of male antenna without (0) (Fig. 1), or with (1), a PS (Fig. 4). 7. AL
shorter (0) (Figs. 15, 16), or longer (1) (Figs. 11, 12), than GC. 8. Aedeagus not bilobed at apex,
phallotreme restricted to apex (0) (Fig. 16); aedeagus bilobed apically, phallotreme more ex-
tensive, extending from apex onto ventral surface (1) (Fig. 12). 9. PM appendiculate apically

(1) (Fig. 10), or not (0) (Fig. 13). 10. Posterior extension of DLA broad, ligulate (0) (Fig. 10);
broad, triangular (1) (Fig. 14); extremely narrow, filiform (2) (Fig. 15). 11. IVP absent (0) (Fig.
1 1), minute (1) (Fig. 15), well developed (2) (Fig. 13). 12. VR narrow (0) (Fig. 13), very broad
(1) (Fig. 11). 13. VS relatively straight, unmodified (0) (Fig. 15); contorted, produced ventro-
medially (1) (Figs. 11, 13).

2 Character codes are ordered phenetically.

maximum length of club segments subequal; Cl, C2 subequal in width, C3 consid-
erably narrower, subconical; flagellar setae relatively short and stout, abruptly tapered
at apex, longest setae 1.67 width of FI; BCPS relatively small, subglobose, difficult
to discern in type, formula apparently 1 -2-2-1- 1; PS formula 0-1 -1-1-2.

Hind tibia atypical for genus, broadly notched dorsoapically.

Genital capsule (Fig. 1 5) 0.34 as wide as long, strongly, evenly tapered from widest
point to apex and to base; PM narrow, slightly curved at apex only, not appendiculate;
AD 0.17 GL; DA smaller than in Lachesis Group, its length 0.45 GL; DLA unique;
base of structure consists of a broadly and lightly sclerotized area intermediate to a
basal and an apical membranous region, basal membranous area subtriangular and
well defined, apical area poorly differentiated; posterior extension of DLA uniformly
very narrow, subliliform, curving ventrally, extending from apex of subtriangular
membranous area to near base of volsellae; DLA not notched or lobed at base; VS
straight elongate, narrowly separated, occupying ca. 0.90 AD; IVP minute, subtrian-
gular; IB narrow, no broader than basal width of IVP; VR narrow, elongate, occupying
0.43 BD; VP not protuberant, extremely basal in position, near midpoint of VR.
Aedeagus length (Fig. 16) 0.93 GL, 0.82 HTL; apodemes 0.47 AL.

1992

NOVEL TRICHOGRAMMA

631

Female. Unknown.

Type. Holotype male. AUSTRALIA. Queensland: Westwood, 9.6 km NE, on Hwy.
66 at Valentine Creek; iv- 12- 1988; screen sweeping; G. Gordh & J. D. Pinto, collrs.
Type deposited in the Queensland Museum, Brisbane, Australia.

Etymology. The specific name primaevum is a Latin word for young or early
(primeval).

Notes. T. primaevum is similar to species of the Lachesis Group only in antennal
segmentation, the two-segmented maxillary palp and the presence of a submarginal
seta on the marginal vein. Hindwind setation, aedeagus and genital capsule structure,
scutellar setal length, and the PS formula on the antenna all differ.

The PS on F2 of the type’s left antenna is oriented longitudinally as is normal in
Trichogrammatidae; its counterpart on the right antenna is oriented transversely.

DIAGNOSES AND SPECIES RELATIONSHIPS

The four new species share characters absent in all other Trichogramma. The male
antenna has both the funicular and club segments unjoined, the maxillary palp is
two-segmented, and the marginal vein has a seta distinctly behind its anterior border.
Species differences are summarized in Table 1. Trichogramma primaevum is clearly
differentiated from the Lachesis Group. Its relationship to this group and other
components of the genus cannot yet be resolved. The phenetically similar species of
the Lachesis Group are tied by aedeagal structure and hindwing setation. Species
relationships within the group also are questionable. The more highly modified FI
in lachesis and clotho suggsts that they are more closely related than either is to
atropos. The occurrence of an IVP in clotho and atropos, and not in lachesis is in
contradiction, but only if the absence of the IVP is plesiomorphic. The distinctive
thoracic microsculpturing in T. clotho separates it from related species and all other
congeners (cf. Figs. 8, 9).

DISCUSSION

Trichogramma currently is divided into two subgenera, the nominate subgenus
and Trichogrammanza. Trichogrammanza was described by Carver (1978) based on
male antennal segmentation (two distinct funicular segments, single club segment).
It includes three Australian species and was recently reviewed by Oatman and Pinto
( 1 987). The nominate subgenus, world-wide in distribution and with over 1 60 named
species, is divided into a variable number of species groups depending on authority
(Nagarkatti and Nagaraja, 1977; Voegele and Pintureau, 1982; Viggiani and Lau-
donia, 1989).

The differences between the species of the Lachesis and Primaevum groups, and
congeners are of a greater magnitude than between the nominate subgenus and Tricho-
grammanza, and would also justify subgeneric, perhaps even generic, treatment on
phenetic grounds. Thus, the placement of these new taxa within the nominate sub-
genus is somewhat discordant to the current classification.

This conservative approach is proposed until a better understanding of phylogenetic
relationships in the genus is gained. Phylogenetic questions concern the relationship
of Trichogrammanza to the nominate subgenus, and the relationship of the Lachesis
and Primaevum groups to one another and to congeners.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

With regard to Trichogrammanza, male antennal segmentation prompted Nagar-
katti and Nagaraja (1977) to consider the taxon intermediate to Trichogrammatoidea
and Trichogramma (Trichogramma). However there is reason to suggest that, rather
than representing the sister taxon of the nominate subgenus, Trichogrammanza is
of more recent origin, and is actually derived from an ancestor with complete antennal
ankylosis in males. The presence of rare males with typical Trichogramma (Trich-
ogramma) antennae (completely fused flagellar segments) in an isofemale-derived
laboratory culture of T. ( Trichogrammanza ) funiculatum Carver (unpubl. data), sug-
gests that the unjoined funicular segments may not be plesiomorphic, but a reversal
instead. Also, the male genitalia of the three species of Trichogrammanza are not
distinctive in any way but instead are similar to those of the Palearctic T. (Tricho-
gramma) principium Sugonjaev and Sorokina, and relatives (Oatman and Pinto,
1 987). The possibility that the latter assemblage represents the sister taxon of Tricho-
grammanza warrants investigation. Of course, subgeneric treatment for Trichogram-
manza could not be supported on cladistic grounds if such a relationship to this or
any other component of the nominate subgenus is shown.

In my opinion the Lachesis and Primaevum groups do represent primitive lineages,
basal to all other Trichogramma. This is suggested by male antennal structure which
is remarkably similar to that in other genera of Trichogrammatini, namely Austral-
ufens and Trichogrammatomyia (cf. figs, i and 4 with figs. 3D & 57B in Doutt and
Viggiani, 1968), and by the two-segmented maxillary palp. Because these similarities
presumably are plesiomorphic they do not argue for the Lachesis and Primaevum
groups representing a single lineage. The bilobed aedeagus and its elongate apicov-
entral phallotreme in the Lachesis Group are unique and perhaps justify isolation at
the subgeneric level. The former trait probably is derived. The ventral extension of
the phallotreme, however, was considered primitive in Hymenoptera by Snodgrass
(1941, 1957). A detailed comparison of aedeagal structure in Trichogramma is re-
quired before a more definitive estimate of polarity of these characters can be given.
Unfortunately, the paucity of Lachesis Group specimens prevents this. In any case,
formal treatment of the Lachesis Group would still leave the position of T. primaevum
unsettled. Although placing both groups in the nominate subgenus is not entirely
satisfactory, it seems preferable to erecting one or two inadequately supported sub-
genera.

ACKNOWLEDGMENTS

Specimens were mounted on slides by Rob Velten; Linda Bobbitt assisted with Figs. 1-7,
10-12, and 14-16; Robert Orth prepared Fig. 13. Gary Platner helped with Figs. 8 and 9, and
the preparation of plates.

This study was supported by Grant BSR-9006177 from the National Science Foundation.

LITERATURE CITED

Carver, M. 1978. A new subgenus and species of Trichogramma Westwood (Hymenoptera:
Chalcidoidea) from Australia. J. Aust. Entomol. Soc. 17:109-112.

Domenichini, G. 1953. Studio sulla morfologia delfaddome degli Hymenoptera Chalcidoidea.
Estr. Ann. Fac. Agr. Milano 2:147-261.

Doutt, R. L. and G. Viggiani. 1968. The classification of the Trichogrammatidae (Hyme-
noptera: Chalcidoidea). Proc. Calif. Acad. Sci. (4th ser.) 35:477-586.

1992

NOVEL TRICHOGRAMMA

633

Ertle, L. R. and C. P. Davis. 1975. Trichogramrna nubilale new species (Hymenoptera:
Trichogrammatidae), an egg parasite of Ostrinia nubilalis (Hiibner). Ann. Entomol. Soc.
Am. 68:525-528.

Nagaraja, H. 1978. Studies on Trichogrammatoidea (Hymenoptera: Trichogrammatidae).
Oriental Insects 12:489-530.

Nagaraja, H. and S. Nagarkatti. 1969. Three new species of Trichogramrna [Hymenoptera:
Trichogrammatidae] from India. Entomophaga 14:393^100.

Nagarkatti, S. and H. Nagaraja. 1968. Biosystematic studies on Trichogramrna species: I.
Experimental hybridization between Trichogramrna australicurn Girault, T. evanescens
Westwood and T. minutum Riley. Tech. Bull. CIBC 10:81-96.

Nagarkatti, S. and H. Nagaraja. 1971. Redescriptions of some known species of Trichogramrna
(Hym., Trichogrammatidae), showing the importance of the male genitalia as a diagnostic
character. Bull. Entomol. Res. 61:13-31.

Nagarkatti, S. and H. Nagaraja. 1977. Biosystematics of Trichogramrna and Trichogram-
matoidea species. Annu. Rev. Entomol. 22:157-176.

Oatman, E. R. and J. D. Pinto. 1987. A taxonomic review of Trichogramrna ( Trichogram -
manza ) Carver (Hymenoptera: Trichogrammatidae), with descriptions of two new spe-
cies from Australia. J. Aust. Entomol. Soc. 26:193-201.

Pinto, J. D. and E. R. Oatman. 1985. Additions to Nearctic Trichogramrna (Hymenoptera:
Trichogrammatidae). Proc. Entomol. Soc. Wash. 87:176-186.

Pinto, J. D., G. R. Platner and E. R. Oatman. 1978. Clarification of the identity of several
common species of North American Trichogramrna (Hymenoptera: Trichogrammati-
dae). Ann. Entomol. Soc. Am. 71:169-180.

Pinto, J. D., R. K. Velten, G. R. Platner and E. R. Oatman. 1989. Phenotypic plasticity and
taxonomic characters in Trichogramrna (Hymenoptera: Trichogrammatidae). Ann. En-
tomol. Soc. Am. 82:414-425.

Schulten, G. G. M. and H. R. Feijen. 1 978. Two new species of Trichogramrna (Hymenoptera;
Trichogrammatidae) from Malawi; egg parasitoids of Diops is macrophthalma Dalman
(Diptera: Diopsidae). Entomol. Berich. 38:25-29.

Schulten, G. G. M. and H. R. Feijen. 1982. A new species of Trichogramrna (Hymenoptera:
Trichogrammatidae) from Malawi, parasitizing eggs of Chilo dijfusilineus (de Joannis).
Entomol. Berich. 42:142-144.

Snodgrass, R. E. 1941. The male genitalia of Hymenoptera. Smithson. Misc. Coll. 99(14): 1-
86, 33 plates.

Snodgrass, R. E. 1957. A revised interpretation of the external reproductive organs of male
insects. Smithson. Misc. Coll. 135(6): 1-60.

Viggiani, G. 1971. Ricerche sugli Hymenoptera Chalcidoidea. XXVIII. Studio morfologico
comparative dell’armatura genitale estema maschile dei Trichogrammatidae. Boll. Lab.
Entomol. Agric. Portici 29:181-222.

Viggiani, G. 1 984. Further contribution to the knowledge of the male genitalia in the Tricho-
grammatidae (Hym. Chalcidoidea). Boll. Lab. Entomol. Agric. Portici 41:173-182.

Viggiani, G. and S. Laudonia. 1989. Le specie italiane di Trichogramrna Westwood (Hy-
menoptera: Trichogrammatidae), con un commento sullo stato della tassonomia del
genere. Boll. Lab. Entomol. Agric. Portici 46:107-124.

Vincent, D. L. and C. Goodpasture. 1 986. Three new species of Trichogramrna (Hymenoptera:
Trichogrammatidae) from North America. Proc. Entomol. Soc. Wash. 88:491-501.

Voegele, J. and B. Pintureau. 1982. Caracterisation morphologique des groupes et especes
due genre Trichogramrna Westwood. Pages 45-75 in: Les Trichogrammes, Les Colloques
de PINRA, No. 9, Antibes, France.

Received 24 September 1991; accepted 22 April 1992.

NOTES AND COMMENTS

J. New York Entomol. Soc. 100(4):634-637, 1992

OBSERVATIONS ON THE BIZARRE JELLY
MASS HABIT OF LARVAL GERANOMYIA
(DIPTERA: TIPULIDAE: LIMONIINAE)

An unusual specimen of a larval Geranomyia tipulid was found at the Finca La
Selva Biological Station in Heredia, Costa Rica, on 8 February, 1989. It was found
while examining leaves among trees in the old coffee finca on the station property.
Suspended beneath one leaf, at about 6 feet from the ground, was a globular mass
about 2.5 cm in diameter, made of a clear jelly-like substance and harboring a verdent
green larva (Fig. 1 ). The larva was obviously a mature instar nematoceran, with an
evident head capsule and thickened thoracic region. Posterior spiracles barely pen-
etrated the surface of the jelly mass, no doubt allowing the larva to respire. When
disturbed, the larva rolled around in the jelly mass, even retracting the posterior
spiracles. With sunlight coming through the leaf the green larva became almost
invisible. It is unknown as to what the larva may have been feeding upon, how it
found its way to such an inaccessible spot, and whether or not the jelly is a feature
of the last (fourth) instar. The specimen, with leaf, was placed in a moist petri dish
and 4 days later it emerged as a female Geranomyia, with intricately patterned wings
typical of the genus. Being such an unusual habit for any fly, this prompted us to
describe the habit and morphological modifications that the larva might have for
this peculiar existence. Other references to the jelly mass habit of larval Geranomyia
are reviewed, and possible origins of the habit are given.

The larval exuvium (with head capsule) was originally pointed on the same pin as
the adult specimen, but it was then softened in KOH and slide mounted for compound
microscopy. Specimens are in the AMNH. Dorsal and ventral views of the head
capsule are shown in Figures 2 and 3, respectively. The rest of the larva was not
examined, since the exuvium was not well preserved. The larva has a typical limoniine
head capsule (Hennig, 1950), with broad lateral plates which are fused to the fronto-
clypeus dorsally forming a blackened, strongly sclerotized area along much of its
length (ventrally the lateral plates are connected only anteriorly to form the mentum).
The antennae are 2-segmented, inserted above the base of the mandibles (the typical,
terminal papilla may have been lost during molting). The mandible is of the cutting-
chewing type, strongly sclerotized, and with one pointed tooth at the apex and five
smaller teeth along the inner margin. The maxilla possesses a sclerotized cardo and
stipes (the lacinia not being well defined). Eight visible pointed teeth occur ventrally
on the hypopharynx (the number dorsally was not visible). The mentum possesses
1 1 rather blunt teeth, the median one largest and slightly elevated. Features of the
head capsule do not differ significantly with other limoniines (cf. Geranomyia cana-
densis [Alexander and Malloch, 1920], see Cramer, 1969), indicating that the diet
may not be so unusual as to require distinct morphological specialization. No evi-
dence of folivory was evident. Such a habit is very rare in tipulids, and occurs, for
example, in Limonia ( Dicranomyia ) kauaiensis, which mines leaves of Crytandra in

1992

NOTES AND COMMENTS

635

Fig. 1. Photograph of Geranomyia sp. fourth instar/prepupal larva in situ in jelly mass
under leaf. Arrow points to posterior spiracle opening.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

2

3

Figs. 2, 3. Head capsule from larval exuvium of Costa Rican Geranomyia female. 2. Dorsal
view. 3. Ventral view.

Hawaii (Swezey, 1915). The Geranomyia specimen might have been grazing upon
an algal film on the leaves.

The bizarre jelly mass habit of tipulids has actually been reported elsewhere, by
Hingston (1932) for a specimen in British Guiana. So far, this habit is known only
in Geranomyia. Hingston reported (p. 342): . . in the forest there was a crane-fly,

Geranomyia, which suspended its pupa in a globule of jelly wrapped around the tip
of Turn palm leaf. The globule was a firm gelatinous substance, rather smaller than
a hazel-nut and clear as water. Indeed, its appearance was that of a large drop of
water about to fall from the end of the palm leaf. . . . The pupa was clearly visible
inside it. It rested in a nearly vertical position, its head upward in the broad part of
the globule, its tail below and close to the leaf. Its colour was grass green with five
dark bands that half-crossed its body.” The Costa Rican specimen was not found in
a pendulous jelly mass the way Hingston reported (the larva in fact was parallel to
the leaf surface), nor were there obvious dark bands across the body. Hingston’s
specimen, with others, were later described as Limonia ( Geranomyia ) gelatifex (Ed-
wards, 1934). Alexander and Alexander (1970) indicate G. gelatifex to be a junior
synonym of Limonia (G.) recondita (Alexander), 1921. They report the distribution
of recondita to be from Guyana and Brazil to Ecuador, Peru, and Panama. The
species identity of the Costa Rican female could not be determined with certainty
since tipulid species are generally defined on the basis of male genitalia.

It is evident that the jelly mass is produced directly from the larva. There is no
evidence that leaves exude the substance, nor even that the larva sequesters the

1992

NOTES AND COMMENTS

637

substance from the plant. In fact, a similar jelly substance is produced in much
smaller quanitities by Limonia triocellata. This is a broadly mycophagous species
breeding in various macrofungi in the eastern United States. As the mature larvae
become pharate pupae, they form mucilaginous tunnels in the fungal context or
substrate below, in which they pupate. Production of the jelly substance may, in fact,
be widespread in limoniines and just produced in overabundance by Geranomyia.
Geranomyia pupating on such exposed surfaces as leaves would probably be highly
susceptible to hymenopterous parasitoids, and the jelly mass may afford at least some
concealment or physical barrier .—David Grimaldi, Department of Entomology,
American Museum of Natural History, Central Park West at 79th Street, New York,
New York 10024-5192; and Chen Young, Section of Invertebrate Zoology, Carnegie
Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213.

ACKNOWLEDGMENTS

We wish to thank Jon Gelhaus (Academy of Natural Sciences, Philadelphia) for citations and
copies of the Geranomyia literature, Doug Futuyma for bringing the specimen to our attention,
and especially an anonymous reviewer for such helpful comments on the manuscript.

LITERATURE CITED

Alexander, C. P. and M. M. Alexander. 1970. Family Tipulidae, Fasc. 4. In: N. Papavero
(ed.), A Catalogue of the Diptera of the Americas South of the United States. Univ. Sao
Paulo, 259 pp.

Alexander, C. P. and J. R. Malloch. 1920. Notes on the life-history of a cranefly of the genus
Geranomyia Haliday (Tipulidae, Diptera). Trans. 111. St. Acad. Sci. 13:310-319.
Cramer, E. 1969. Die Tipuliden des Naturschutzparkes Hohr Vogelsberg. Deut. ent. Zeitr.
15:133-232.

Edwards, F. W. 1934. Oxford University Expedition to British Guiana, 1929. Diptera Nema-
tocera. Ann. Mag. Nat. Hist. 14:632-635.

Hennig, W. 1950. Die Larvenformen der Dipteren. Akademie-Verlag, Berlin, 457 pp.
Hingston, R. W. G. 1932. A Naturalist in the Guiana Forest. E. Arnold & Co., London.
Swezey, O. H. 1915. A leaf-mining cranefly in Hawaii. Proc. Haw. Ent. Soc. 3:87-89.

Received 4 June 1990; accepted 1 1 May 1992.

BOOK REVIEW

J. New York Entomol. Soc. 100(4):638-64Q, 1992

Women in the Field, America’s Pioneering Women Naturalists. — M. M. Bonta. 1991.

Texas A&M Press, College Station, TX. $13.95.

Marcia Myers Bonta’s Women in the Field is a must read for men and women
naturalists, botanists, ornithologists, ecologists, entomologists and historians. It is
written in a style that is enjoyable for light bedtime reading; yet provides rigorous
historical information rendering it a valuable reference book. Chapters are dedicated
to 25 women who, in the author’s estimation, made the most important contributions
to their field, and about whom adequate information could be obtained. They are
grouped in 6 sections: I. The pioneers, II. The naturalists, III. The botanists, IV. The
entomologists, V. The ornithologists, and VI. The ecologists. Each section contains
a preface that highlights the accomplishments of several other women who either
had less impact or whose lives were less well documented. The book is illustrated
with 30 photographs of the women in their element or their artwork. Reference
material is not included in the text, but a selected bibliography is provided at the
end for readers who wish to obtain source material used by the author. The book
also includes an index. Each chapter stands alone, such that readers can selectively
learn about the careers of specific women. However, the author often points out
connections between these women, since many of them were acquainted with each
other.

It is remarkable how many women naturalists emerged “in the days when women
were perceived as fragile beings fit only to keep house and raise children.” Here I
describe some of my favorite chapters to give readers the flavor of Bonta’s writing
style. Martha Maxwell was a mid 19th century intrepid Colorado huntress and taxi-
dermist who is pictured with a gaudy display of stuffed deer, elk and bighorn sheep
at the Philadelphia Centennial. She often went camping alone in the mountains and
also had a passion for wildflowers. Mary Treat, a late 19th century naturalist of the
New Jersey Pine Barrens, spent many productive hours in her backyard surrounded
with an arbor vitae hedge 15 feet high and 150 feet in circumference, which became
known as her “insect menagerie” or “arachnidian menagerie.” Annie Montague
Alexander, a late 1 9th century quintessential naturalist and daring explorer, preferred
to sleep outside in the desert “where half the universe shown down upon us, not a
tree to break the wonderful arch of the Milky Way reaching from horizon to horizon.”
Alice Eastwood, best known for her early 20th century botanical contribution to the
western US flora, was completely self-taught, raised mostly by relatives and foster
parents after her mother died when Alice was 6. At age 91, she was made honorary
president of the International Botanical Congress in Sweden where she sat in the
chair of Linneaus. Ann Haven Morgan was trained as an aquatic ecologist and mayfly
expert by J. G. Needham at Cornell University in the early 20th century. She was
once described as “a gray-haired lady knee-deep in muck and water ferociously
pursuing nasty little herbivora and carnivora with a net” never wanting to “come
out of some particularly oozy mudhole.” I personally aspire to be described similarly
when my hair becomes more gray than brown.

1992

BOOK REVIEW

639

The last chapter describes the inspiring career of Rachel Carson, who was labeled
by some critics as a “mere birdwatcher,” due to the fact that she was much too
qualified a scientist to treat as a “hysterical woman” for publishing Silent Spring,
one of the most influential and controversial conservation works of all time. Unlike
many of her predecessors who had enjoyed long and healthy lives, Rachel Carson
tragically died of cancer at the age of 56, “mourned throughout the world by those
who love the earth.”

The lives of three entomologists are described in detail. Annie Trumbull Slosson,
known in the late 1 9th century as “the old bug woman” was often observed on the
slopes (or summit) of Mt. Washington in New Hampshire with her “butterfly net.”
She also spent time collecting insects in Florida and in the Delaware Water Gap.
She discovered many new insect species including Lepidoptera and Hymenoptera.
After having a new moth species named after her, she remarked, “Without a thought
of irreverence I felt like expressing my willingness to depart in peace now that I had
seen this greatest of all earthly honors descend upon my humble head. An insect
named for unworthy me! And the name so sonorous and delightful in sound. In-
guromorpha slossonil Like the proverbial Mesopotamia of the eloquent preacher it
stirred one’s soul and I think I murmured it even in my sleep.” When that particular
species was superseded, she lamented that it sunk “into the dread valley of synon-
ymy.”

Edith Patch, best known for her work on biology of aphids and also for her children’s
books on natural history, was one of the few women naturalists who was classically
trained, graduating as an English major from the University of Minnesota in 1901,
and eventually receiving a M.S. degree from the University of Maine and a Ph.D.
from Cornell University under John Henry Comstock. She relentlessly pursued the
study of entomology despite early warnings that entomology was not a suitable
profession for a woman. She was given an early opportunity by Dr. Charles D. Woods,
director of the Maine Agricultural Experimental Station who came under some crit-
icism for hiring a woman. “When one of the leading Bureau of Entomology men
remarked to Dr. Woods, ‘I hear you have appointed a woman as entomologist. Why
on earth did you do that? A woman can’t catch grasshoppers,’ he received the drawled
reply, ‘It will take a lively grasshopper to escape Miss Patch.’ ”

The chapter on Anna Botsford Comstock was touching and masterfully written.
Working in the shadow of her husband, at the turn of the century she was appointed,
and then unappointed as an assistant professor of nature study because the Board of
Trustees at Cornell University objected to giving a woman professorial status. John
Henry Comstock was highly supportive collaborating with Anna on numerous proj-
ects for which she not only did the artwork, but also some of the writing. In addition
she amassed an impressive list of scholarly achievements of her own. The most
famous was her Handbook of Nature Study, which has affected the lives of many
children. My personal favorite is The Ways of the Six-footed, a charming account of
her own observations of insects. After many years of teaching and scholarship, she
was eventually appointed as an assistant, then a full professor the same year that
women finally received the vote in this country. She and her husband, childless, but
not by choice, “adopted” a long line of children and graduate students for whom
they provided not only emotional but also financial support. Tragically, John Henry
Comstock suffered a stroke, after which Anna continued to work while caring for

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

him. Four years later she died of cancer two weeks after teaching her last class in her
home in Ithaca. Liberty Hyde Bailey described her as an unfailing humanitarian who
left “a fragrant memory of high achievement, noble service, unselfish cooperation,
constructive counsel, inspired teaching, loving kindness and unforgettable compan-
ionship.”

In the preface the author points out many attributes that these pioneer women
naturalists had in common. One of them was modesty, as so aptly illustrated in the
quote from Annie Trumbull Slosson. None liked typically “women things” like
housework or clothes, although most of them did enjoy cooking for their friends.
Some had been favorite daughters of enlightened fathers who appreciated nature
themselves, and others had independent mothers as role models. Of the 25 women
portrayed in this book, most of them never married or else started their careers after
their husbands died. Six of them had supportive husbands while two were actively
discouraged by their husbands. Only two ever had children. Most were self-taught
and did not have professional positions. Some even turned over their materials to
men professionals for publication in books and journals.

The author’s hope was that “the lives and thoughts of my women naturalists will
not only convince others that women and nature were and still are compatible, but
also that the women’s contributions to the study of nature were important— in fact
amazing, given the prejudices they had to contend with.” I found the book delightful,
captivating and inspiring. I will encourage both my young son and daughter to read
it some day to gain an appreciation of the relatively supportive environment within
which their ecologist mother and hopefully they will we able to grow, in contrast to
the harsh social climate that acted as a major hurdle during the 19th and much of
the 20th centuries for women who were interested in pursuing the study of nature.—
Barbara L. Peckarsky, Department of Entomology, Cornell University, Ithaca, New
York 14850-0999.

J. New York Entomol. Soc. 1 00(4):64 1—65 1 , 1992

OBITUARY

In Memoriam
La Verne L. Pechuman
1913-1992

Dr. LaVerne L[eRoy]. Pechuman, or Verne to nearly everyone who knew him.
Professor Emeritus of the Department of Entomology, Cornell University, and an
international authority on the taxonomy of the blood-sucking flies of the family
Tabanidae, died Monday, March 30, 1 992, at Strong Memorial Hospital, Rochester,
New York, after a short battle with cancer. He was 78 years old, and had continued
to be active in his research until about three weeks before his death.

He was born October 18, 1913, in Lockport, New York, the son of Henry J. and
Anna Brege Pechuman. He attended the Lockport public schools, and graduated from
Lockport High School in 1930. After one year of post-graduate work, he matriculated
at Cornell University, and graduated in 1935 with a bachelor’s degree in Entomology.
In July 1935, he was appointed an assistant in Entomology for the Dutch elm disease
investigation being conducted jointly by the Departments of Entomology and Plant
Pathology. He was admitted to the Graduate School at Cornell in September 1935.
During the initial course of work on the Dutch elm disease, it was thought advisable
to conduct a comprehensive survey of the various insects found associated with elm,
and with special reference to their potential to transmit the causative organism of
Dutch elm disease. Verne carried out these early biological investigations, first begun
in 1934 by Drs. Philip A. Readio and Henry Dietrich, under the aforementioned
professors’ direction and later that of Dr. D. L. Collins. Pechuman’s work culminated
in a thesis entitled “Preliminary account of the insects found in the bark and wood
of the American elm ( Ulmus americana L.),” and in February 1937 he successfully
completed requirements for a master’s degree in Entomology. Verne’s doctorate work
was a continuation of his research on the Dutch elm disease, again under the able
direction of Dr. Readio. A thesis entitled “The insects found in the bark and wood
of the American elm ( Ulmus americana L.)” was submitted to the Faculty of the
Graduate School of Cornell for the degree of Doctor of Philosophy in June 1939.

Upon the completion of Verne’s doctorate, he gained employment with the Ortho

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Division of the Chevron Chemical Company (formerly California Chemical Co.).
Beginning as a field representative (1939-45) in the company, he rose through the
ranks, first from a Branch Manager (1945-47), to a District Manager (1947-61), and
finally to a senior research scientist (1961-62). After nearly 25 years of dedicated
service to Chevron Chemical Co., he returned to academia and Cornell University,
joining the entomology faculty in 1962 as an associate professor and curator of the
world famous insect collection. He was named full professor in 1972. Soon after his
retirement in 1982, he was awarded the title of professor emeritus.

Among his academic peers and colleagues, Verne Pechuman will be best remem-
bered as a world authority on the systematics, biology, and distribution of horse flies
and deer flies (Tabanidae). During his 20 years as curator of the Cornell University
Insect Collection, he is credited with amassing a collection of well in excess of 40,000
tabanid specimens, the largest and most geographically diverse assemblage of world
Tabanidae in a North American university collection.

In the early 1970s, while collaborating with Dr. Mathias J. Kemen, Jr., a Cornell
veterinarian, on the annoying biting and blood-sucking habits of horse and deer flies,
and, in an attempt to answer the plight of New York dairymen and horse owners,
Verne designed a mechanical trap (a prototype of which he used previously to make
general collections of tabanids) to capture and significantly reduce the number of
flies in a given area. It is a known fact that dairy cattle attacked by these biting flies
often show a significant drop in milk production, and in horses these flies can be
vectors of a debilitating viral disease known as equine infectious anemia. His broad-
based knowledge of tabanid biology and especially of some of the behavior traits of
these flies was the basis for the efficacy of this mechanical trap. Tabanid flies, which
are attracted to dark, moving objects, are lured to the trap by a swinging, shiny, black
ball, and a carbon dioxide supply (dry ice), which fools the flies into thinking there
are live animals about (anthropomorphically speaking!). Once inside the trap, the
flies move upward towards light and are captured and killed by an insecticide in the
frap head. This trap design is still in general use today.

Although his primary taxonomic and biological work focused principally on the
horse and deer flies, other interests of his included following the emergence of the
various broods of the periodical cicada (the 17-year locust) in the eastern United
States, a fascination with the effect of Pleistocene glaciation on several groups of
insects, and yearnings in the history and prehistory of Upstate New York, local native
American history, archaeology, and botany.

Verne loved the out-of-doors, and on many occasions he seized the opportunity
to go collecting, usually to his favorite bogs around central New York to find the
pesky horse and deer flies. He also had an admiration for plants, of which he knew
many, if not most, of the local native and cultivated species. He had an inordinate
fondness for trilliums, however— so much so that he authored several articles in the
early 1960s. In Science on the March, a Buffalo Museum of Science publication,
April and June 1962 issues, Verne reported on a multiplicity of forms of an uncertain
status growing on the Tonawanda Indian Reservation in Genesee and Erie Counties.
While living in western New York, he built up a significant collection of most of the
North American species of Trillium, including some natural hybrids and a number
of specimens of aberrations of some species. He also exchanged Trillium specimens
with botanists in Japan, which are still known to be growing in his native Lockport.

1992

L. L. PECHUMAN MEMORIAM

643

Drs. Mathias J. Kemen, Jr. (left) and LaVeme L. Pechuman examine a mechanical trap,
designed by Pechuman, that captures horse and deer flies.

In Spring of 1963 he donated specimens of his extensive collection, which included
five Japanese species, to the Cornell Plantations.

Perhaps one of Verne’s rarest tributes was his welcome as a guest at Tonawanda
and Tuscarora Indian ceremonies. As well he was an adopted member of the Ton-
awanda Band (Hawk Clan) of the Seneca Indians. Through the years, Verne was
always deeply concerned about the welfare of native Americans. On his many travels
about New York State, he was known occasionally to detour to an Indian reservation
to speak to and share times with his “adopted” brethren. In 1961, as a director and
editor of The Niagara County Historical Society, Verne wrote an introduction to one
of the early writings of David Cusick, a Tuscarora Indian: “Ancient History of the
Six Nations,” which was reprinted by the Society. In Verne’s closing statement in

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

this introduction, he wrote “It is hoped that by reprinting this work of a Tuscarora
Indian, it will be more readily available not only to students of anthropology but to
anyone interested in a peculiar bit of Americana.”

His numerous professional affiliations included the American Association for the
Advancement of Science (AAAS), Entomological Society of America, Entomological
Society of Ontario, American Entomological Society, New York Entomological So-
ciety (of which he was a long-time sustaining member), American Mosquito Control
Association, Sociedade Brasileira de Entomologia, Buffalo Society of Natural Sci-
ences, New York State Archaeological Association, and the Society for Pennsylvania
Archaeology. As an active advocate for habitat preservation locally and globally, he
belonged to The Nature Conservancy, Nature Sanctuary Society of Western New
York, Bergen Swamp Preservation Society, and the Wilderness Society. He held
advisory positions in many of these associations. Among his honors he was a member
of Sigma Xi, appointed a fellow of AAAS in 1964 and a fellow of the Rochester
Museum of Arts and Sciences in 1965, and listed in the American Men of Science.
In 1960 he was made an honorary member of the Rochester Academy of Science,
and also in that same year received a citation as “distinguished scientist” from the
University of Buffalo.

Verne was author and coauthor of nearly 100 scientific papers and monographs
(see “publications ...” below). He described 23 species (and subspecies) of horse
and deer flies from around the globe as new to science (see “new taxa . . .” below).
Reflecting the respect that other entomologists had for his work, Verne had 26 species
named after him as of 1986, mostly flies, but also species of other insect orders and
a protozoan (see “taxa named . . .” below).

He is survived by a sister and brother-in-law, Dorothy and Benjamin Neal of San
Jose, California; daughters and sons-in-law, Patricia and William Ferris of Bergen,
New York, and Jean and James McIntyre of Waterville, Maine; grandchildren Wil-
liam and Michael Ferris; and one niece and several cousins. His wife of 52 years,
Berta, preceeded him in death on December 3, 1991. Following a memorial service
held in Ithaca, New York, he was buried in the family plot (Cold Springs Cemetery)
in Lockport, New York, on April 4, 1992.

Professor Pechuman has left an enduring mark on the systematics of the Tabanidae
to which he did much to fashion. As well he leaves behind a legacy of scholarship
and dedicated service to Cornell University that includes not only his own publi-
cations but, also, his large collection of tabanid flies. The lives of his long-time
colleagues and students are much richer today for having known and worked with
him. We will remember him with fondness and great respect, and for his generosity
of time, dash of good humor, and as an enthusiastic conversationalist and delightful
companion. He will be dearly missed by his colleagues in the profession and by his
many other friends.

new taxa described by l. l. pechuman

1. Silvius philipi n. sp. 1938. USA: 10 mi. SE Lebanon, Oregon.

2. Si/vius quadrivittatus (Say) var. texanus n. var. 1938. USA: College Station, Texas.

3. Chrysops mutata n. sp. 1939. MEXICO: Juan Manuel (near El Salto), Durango.

4. Chrysops berta n. sp. 1940. BRAZIL: Nova Teutonia, Santa Catharina.

5. Chrysops striatula n. sp. 1943. CHINA: Suifu, Szechuen.

1992

L. L. PECHUMAN MEMORIAM

645

Professor L. L. Pechuman, curator of the Cornell University Insect Collection from 1 962—
1982, examines one of the more than 12,000 drawers which house approximately 4-5 million
insect specimens from around the world.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

6. Chrysops aenea n. sp. 1940. CHINA: Yellow Dragon Gorge near Songpan, Szechuen.

7. Glutops rossi n. sp. 1945. CANADA: Port Haney, British Columbia.

8. Catach/orops d’almeidai n. sp. 1946. BRAZIL: Tingua, Rio de Janeiro.

9. Chrysops atlantica n. sp. 1949. USA: Rehoboth Beach, Delaware.

10. Chrysops flavida celata n. subsp. 1949. USA: Medford Lakes, New Jersey.

1 1. Chrysops patricia n. sp. 1953. PARAGUAY: Cerro Pelado.

12. Chrysops argentina n. sp. 1953. ARGENTINA: Tucuman.

13. Tabanus dietrichi n. sp. 1956. USA: Phelps Botanical Area, White Mountains, Arizona.

14. Si/vius pollinosus jeanae n. subsp. 1960. USA: Naval Air Station, Corpus Christi, Texas.

15. Hybomitra frosti n. sp. 1960. CANADA: Orrville, Ontario.

16. Tabanus wilsoni n. sp. 1962. USA: Arkansas River, Arkansas Co., Arkansas.

17. Pelecorhynchus penai n. sp. 1967. CHILE: Pichinahuel, Arauco.

18. Chrysops calvus n. sp. (Pechuman & Teskey). 1967. USA: Allegany State Park, New York.

19. Asaphomyia floridensis n. sp. 1974. USA: Archbold Biological Station, Lake Placid, High-
lands County, Florida.

20. Chrysops dixianus n. sp. 1974. USA: Wedge Plantation, McClellanville, South Carolina.

21. Chrysops stonei n. sp. 1977. JAPAN: Shirikishinai, Kamedagun, Hokkaido.

22. Atylotus woodin. sp. 1981. CANADA: S. of Lot 31, Cone. Gore, Puslinch Twp., Wellington
Co., Ontario.

23. Apatolestes philipi n. sp. 1986. USA: Dougout Wells, Big Bend National Park, Texas.

TAX A NAMED IN HONOR OF L. L. PECHUMAN

Protozoa-Eugregarinida

1. Cometoides pechumani Anderson & Magnarelli. 1978. J. Invert. Path. 31:324-328 (orig.
descrip, p. 324). USA: Housatonic River, Milford, Connectict.

Insecta

Odonata-Gomphidae

2. Epigomphus pechumani Belle. 1970. Studies on the fauna of Suriname and other Guyanas
1 1:1-158 (orig. descrip, p. 19). COLOMBIA: (no other data given).

Plecoptera-Capniidae

4. Allocapnia pechumani Ross & Ricker. 1964. Trans. 111. State Acad. Sci. 57:88-93 (orig.
descrip, p. 88). USA: Starkville, Herkimer County, New York, Otsquago Creek.

Homoptera-Aphididae

4. Macrosiphum pechumani MacGillivray. 1966. Can. J. Zool. 44: 1085-1088 (orig. descrip,
p. 1085). USA: Lockport, New York, ex Smilacina racemosa.

Diptera-Ceratopogonidae

5. Culicoides pechumani Cochrane. 1974. Florida Entomol. 57:125-135 (orig. descrip, p.
133). USA: Ringwood Game Preserve, Tompkins Co., New York.

6. Forcipomyia ( Euprojoannisia ) pechumani Bystrak & Wirth. 1978. U.S.D.A., Tech. Bull,
no. 1591:51 pp. (orig. descrip, p. 34). USA: Cranberry Lake, St. Lawrence Co., New York.

1992

L. L. PECHUMAN MEMORIAM

647

Diptera-Tabanidae

7. Chrysops pechumani Philip. 1941. Proc. Entomol. Soc. Wash. 43:113-130 (orig. descrip,
p. 128). USA: Niles, Inyo Co., California.

8. Amphichlorops pechumani Barretto. 1 948. Revista de Entomol. 19:401-417 (orig. descrip.
p. 407). BRAZIL: “Faz. Serra, Itatiaia, Est. do Rio de Janeiro.”

9. Cydistomyia pechumani Philip. 1959. Fieldiana:Zool. 33:543-625 (orig. descrip, p. 557).
PHILIPPINES: Negros Orientale, Mount Canlaon.

10. Tabanus imitans var. pechumani Philip. 1960. Florida Entomol. 43: 1 7 1-1 74 (orig. descrip,
p. 171). USA: “Gunntown,” Levy Co., Florida.

1 1. Stenotabanus ( Aegialomyia ) pechumani Philip. 1966. Ann. Entomol. Soc. Amer. 59:51 9—
527 (orig. descrip, p. 522). MEXICO: 10 mi S. Telolutla, Vera Cruz.

12. Dasybasis pechumani Coscaron & Philip. 1967. Segundas jornadas entomoepidemiologicas
argentinas 1 ( 1 965):95— 1 03 (orig. descrip, p. 97). CHILE: Tarapaca, Arica.

13. Diachlorus pechumani Fairchild. 1972. Florida Entomol. 55:219-229 (orig. descrip, p.
223). PERU: Quince Mil, Cuzco.

14. Haematopota pechumani Stone & Philip. 1974. U.S.D.A., Tech. Bull. no. 1489:240 pp.
(orig. descrip, p. 159). LAOS: Vientiane Province, Ban Van Eue.

15. Promycteromyia pechumani Coscaron & Philip. 1979. Proc. Calif. Acad. Sci. 41:427-
452 (orig. descrip, p. 436). CHILE: Aconcagua, Valle de Piuquenes.

16. Hybomitra pechumani Teskey & Thomas. 1979. Can. Entomol. 1 1 1:343-350 (orig. de-
scrip. p. 346). CANADA: Alfred, Ontario.

17. Esenbeckia pechumani Wilkerson & Fairchild. 1983. J. Nat. Hist. 17:519-567 (orig.
descrip, p. 549). PERU: Cuzco, Quincemil.

Diptera-Otitidae

18. Euxesta pechumani Curran. 1938. Amer. Mus. Nov. no. 975:7 pp. (orig. descrip, p. 4).
USA: Bronx, New York City, New York.

Diptera-Sciomyzidae

19. Dictya pechumani Valley. 1977. Search Agric., N.Y. State Agric. Exp. Stn. (Ithaca), no.
7:44 pp. (orig. descrip, p. 4). USA: Ipswich, Massachusetts.

Diptera-Chloropidae

20. Elachiptera pechumani Sabrosky. 1948. J. Wash. Acad. Sci. 38:365-382 (orig. descrip,
p. 377). USA: 6 mi S of Middleport, New York, reared ex larvae from base of wild iris
( Iris versicolor).

Diptera-Agromyzidae

21. Liriomyza pechumani Spencer. 1986. U.S.D.A., Agric. Handb. no. 638:478 pp. (orig.
descrip, p. 291). USA: Copeland, Collier Co., Florida.

Diptera-Tachinidae

22. Vibrissotheresia pechumani Reinhard. 1943. Bull. Brooklyn Entomol. Soc. 38:78-90 (orig.
descrip, p. 87). USA: Brewster, New York.

648

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

Hymenoptera-Braconidae

23. Opius pechumani Fischer. 1970. Anz. ost. Akad. Wiss. Mathematische-Naturwisser-
schaftliche Klasse 107:12-32 (orig. descrip, p. 22). USA: near Portal, Arizona.

Hymenoptera-Ichneumonidae

24. Ceratophygadeuon pechumani Townes. 1983. Mem. Amer. Entomol. Inst. 35:281 pp.
(orig. descrip, p. 39). USA: Ludlowville, New York.

Hymenoptera-Sphecidae

25. Psammaletes pechumani Pate. 1936. Trans. Amer. Entomol. Soc. 62:49-56 (orig. descrip,
p. 53). USA: Fairmount and Kearney Avenues, The Bronx, New York, New York. [Now
a synonym of Psammaletes mexicanus (Cameron)].

26. Tachysphex tarsatus subsp. pechumani Krombein. 1938. Ann. Entomol. Soc. Amer. 31:
467^170 (orig. descrip, p. 468). USA: Manahawkin, New Jersey. [Now raised to specific
rank.]

PUBLICATIONS OF L. L. PECHUMAN

1. 1937. An annotated list of insects found in the bark and wood of Ulmus americanus L.
in New York State. Bull. Brooklyn Entomol. Soc. 32:8-21.

2. 1937. Notes on some Neotropical species of the genus Chrysops (Dipt. Taban.). Rev. de
Entomologia 7:134-141.

3. 1937. Lariophagus distinguendus Foerst. parasitic on Sitodrepa panicea L. in New York.
J. Econ. Entomol. 30:563.

4. 1938. A synopsis of the New World species of Vermileo (Diptera-Rhagionidae). Bull.
Brooklyn Entomol. Soc. 33:84-89.

5. 1938. The identity of Tabanus moerens Fabr. Bull. Brooklyn Entomol. Soc. 33:136-137.

6. 1938. A preliminary study of the biology of Scolytus sulcatus LeC. J. Econ. Entomol. 31:
537-543.

7. 1938. Two new Nearctic Silvius (Diptera). Can. Entomol. 70:165-171.

8. 1938. Additions to the New York state list of Tabanidae. J. N. Y. Entomol. Soc. 46:457-
460.

9. 1 939. Notes on some Mexican species of Chrysops (Diptera-Tabanidae) and the description
of a new species. Bull. Brooklyn Entomol. Soc. 34:240-244.

1 0. Tyler, L. J., K. G. Parker and . 1939. The relation of Saperda trident at a to infection

of American elm by Ceratostomella ulmi. Phytopathology 29:547-549.

11. 1 940. Notes on the feeding and breeding habits of Saperda tridentata Oliv. Bull. Brooklyn
Entomol. Soc. 35:113-116.

12. 1940. A new Chrysops from Brazil (Dipt. -Tabanidae). Rev. de Entomologia 1 1:886-888.

13. 1942. Lista provisional de los Tabanidos de Venezuela. Bol. de Entomol. Venezolana 1:
51-58.

14. 1943. Two new Chrysops from China (Diptera-Tabanidae). Proc. Entomol. Soc. Wash.
45:42-44.

15. 1943. Epitrimerus pyri Nal. in New York. J. Econ. Entomol. 36:1 16.

16. 1943. Scolytus sulcatus LeC. in western New York. J. Econ. Entomol. 36:1 19.

17. 1943. Notes on the biology of Chyliza notata (Diptera, Psilidae). Bull. Brooklyn Entomol.
Soc. 38:97.

18. 1945. A new species of Glutops (Diptera, Coenomyiidae). Can. Entomol. 77:134-135.

1992

L. L. PECHUMAN MEMORIAM

649

19. 1946. A new Catachlorops from Brazil (Diptera, Tabanidae). Livro de Homenagem a R.
F. d’Almeida, no. 32:283-284.

20. 1949. Somes notes on Tabanidae (Diptera) and the description of two new Chrysops. Can.
Entomol. 81:77-84.

21. 1952 (Ed.). Selections from “A Narrative Along Three-Fourths of a Century” by Anson
Augustus Boyce. Occ. Contribr. Niagara Co. Hist. Soc., no. 8:28 pp.

22. 1953. A new Neotropical Chrysops (Diptera, Tabanidae). Psyche 60:1 15-1 18.

23. 1953. A new Chrysops from Argentina (Diptera, Tabanidae). Acta Zool. Lilloana 13:381—
385.

24. 1954. Seasonal distribution of some Tennessee Tabaninae (Diptera, Tabanidae). Bull.
Brooklyn Entomol. Soc. 49:128-130.

25. 1955. Some undescribed or little known males of Tabanidae (Diptera). Bull. Brooklyn
Entomol. Soc. 50:57-59.

26. 1956. An unusual new Tabanus from Arizona. Pan-Pacif. Entomol. 32:39 — 42.

27. 1956. An Orleans County bog. Sanctuary News 16:6-8.

28. 1957. Descriptions of Tabanidae previously known from one sex only (Diptera). Bull.
Brooklyn Entomol. Soc. 52:29-34.

29. 1957. A Tabanus not previously known from the United States (Diptera, Tabanidae).
Entomol. News 68:1 18.

30. 1957. Some Tabanidae (Diptera) not previously recorded from Canada. Rept. Entomol.
Soc. Ont. 87:27-28.

31. 1957. The Tabanidae of New York. Proc. Roch. Acad. Sci. 10:121-179.

32. 1958. Niagara County and its towns. Occ. Contribr. Niagara Co. Hist. Soc., no. 13:38 pp.

33. 1958. Some comments on Tabanus longiusculus Hine (Diptera: Tabanidae). Bull. Brooklyn
Entomol. Soc. 53:74-75.

34. 1958. Another radiocarbon date for New York State: the Oakheld site, Genesee County.
Bull. N. Y. State Archaeol. Assoc., no. 14:2-4.

35. 1958. A brief history of Trinity Lutheran Church, Lockport, New York. 10 pp.

36. Frost, S. W. and . 1958. The Tabanidae of Pennsylvania. Trans. Amer. Entomol.

Soc. 84:169-215.

37. 1960. Some new and little-known North American Tabanidae (Diptera). Can. Entomol.
92:793-799.

38. 1961 (Ed.). “Ancient history of the Six Nations” by David Cusick. Occ. Contribr. Niagara
Co. Hist. Soc., no. 15:42 pp.

39. and W. W. Wirth. 1961. A new record of Ceratopogonidae (Diptera) feeding on

frogs. J. Parasitol. 47:600.

40. , H. J. Teskey and D. M. Davies. 1961. The Tabanidae of Ontario. Proc. Entomol.

Soc. Ont. 91:77-121.

41. 1962. Trilliums of western New York. Science on the March 42:68-71.

42. 1962. Trillium variations in western New York. Science on the March 42:96-102.

43. 1 962. A new Nearctic Tabanus of the fulvulus group (Diptera: Tabanidae). Bull. Brooklyn
Entomol. Soc. 57:66-70.

44. 1 963. Another record of Tabanidae as prey of Crabro monticola (Packard) (Hymenoptera:
Sphecidae). J. N. Y. Entomol. Soc. 71:218-219.

45. 1 964. A synopsis of Merycomyia (Diptera: Tabanidae). Proc. Entomol. Soc. Ont. 94:62-67.

46. 1964. A list of the Tabanidae (Diptera) of Quebec. Can. Entomol. 96:1495-1496.

47. 1965. A second specimen of Neochrysops globosus Walton (Diptera, Tabanidae). Entomol.
News 76:89-90.

48. 1967. A new Pelecorhynchus from Chile (Diptera: Tabanoidea: Pelecorhynchidae). Can.
Entomol. 99:554-555.

49. 1967. Symphoromyia biting in New York State (Diptera, Rhagionidae). Mosquito News
27:183-184.

650

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 100(4)

50. 1967. Observations on the behavior of the bee Anthidium manicatum (L.). J. N. Y.
Entomol. Soc. 75:68-73.

51. and H. J. Teskey. 1967. A new eastern Nearctic Chrysops (Diptera: Tabanidae).

Can. Entomol. 99:645-650.

52. 1967. Lectotype designation for Chrysops shermani Hine (Diptera, Tabanidae). Entomol.
News 78:260.

53. 1968. Diptera. TIEG Newsletter 4:10-13.

54. 1968. The periodical cicada. Brood VII (Homoptera: Cicadidae, Magicicada). Trans.
Amer. Entomol. Soc. 94:137-153.

55. and A. Stone. 1968. A new synonymy in Hybomitra (Diptera: Tabanidae). Proc.

Entomol. Soc. Wash. 70:302.

56. 1969. Recent gifts to the Cornell University insect collection. Entomol. News 80:43.

57. 1969. A record of Rachiceridae from New York (Diptera). Entomol. News 80:78.

58. and J. J. S. Burton. 1969. Seasonal distribution of Tabanidae (Diptera) at Texas

Hollow, New York in 1968. Mosquito News 29:216-220.

59. 1969. A synopsis of Holcopsis (Diptera: Tabanidae). Can. Entomol. 101:1 107-1 111.

60. Thompson, P. H. and . 1970. Sampling populations of Tabanus quinquevittatus

about horses in New Jersey, with notes on the identity and ecology. J. Econ. Entomol. 63:
151-155.

61. 1971. Shapiro collection at Cornell. J. Lepidopt. Soc. 25:82-83.

62. 1971. A table of the insects that are found in Canada. Bull. Entomol. Soc. Can. 3:69.

63. 1972. The horse flies and deer flies of New York (Diptera, Tabanidae). Search Agric.,
N.Y. State Agric. Exp. Stn. (Ithaca), no. 2:72 pp.

64. 1973. Horse flies and deer flies of Virginia (Diptera: Tabanidae). Va. Polytech. Inst, and
State Univ. Res. Div. Bull. 81:1-92.

65. Ezell, W. B., D. C. Sheppard, T. R. Adkins, Jr. and . 1974. An annotated list of

the Tabanidae (Diptera) of South Carolina. J. Georgia Entomol. Soc. 9:91-99.

66. 1974. Chloromyia formosa established in the United States (Diptera: Stratiomyiidae).
Entomol. News 85:54-55.

67. 1974. A second specimen of Holcopsis fenestr at a (Diptera: Tabanidae). Can. Entomol.
106:559.

68. 1974. N. Y. vs. W. T. Entomol. News 85:62-63.

69. 1974. Two new Tabanidae from southeastern United States (Diptera). J. N. Y. Entomol.
Soc. 82:183-188.

70. Allen, W. A. and . 1975. New distribution records of thirty-eight species of Virginia

tabanids (Diptera: Tabanidae). U.S.D.A., Coop. Econ. Insect Rept. 25:305-309.

7 1 . Magnarelli, L. A. and . 1975. Ovarian studies of Tabanus quinquevittatus (Diptera:

Tabanidae). J. Med. Entomol. 1 1:687-690.

72. Krinsky, W. L. and . 1975. Trypanosomes in horse flies and deer flies in central

New York State. J. Parasitol. 61:12-16.

73. 1975. University collections. Bull. Entomol. Soc. Amer. 21:91-93.

74. 1975. James Chester Bradley (obituary). Ann. Entomol. Soc. Amer. 68:923.

75. Allen, W. A. and . 1976. New geographical and seasonal distribution records for

fifty-three species and subspecies of Virginia tabanids (Diptera: Tabanidae). U.S.D.A.,
Coop. Plant Pest Rept. 1:853-858.

76. 1977. Chrysops stonei, a new tabanid from Japan (Diptera: Tabanidae). Proc. Entomol.
Soc. Wash. 79:107-1 10.

77. Allen, W. A. and . 1977. New geographical and seasonal distribution records for

forty-four species and subspecies of tabanids from Virginia, Tennessee, and West Virginia.
U.S.D.A., Coop. Plant Pest Rept. 2:887-891.

78. Burnett, A. M., C. V. Coveil, Jr. and . 1978. The horse flies and deer flies of

Kentucky: new state records and a revised annotated checklist (Diptera: Tabanidae). En-
tomol. News 89:197-200.

1992

L. L. PECHUMAN MEMORIAM

651

79. Hacker, J. D., L. Butler and . 1979. New geographical distribution records for

forty-four species and subspecies of tabanids in West Virginia (DipteraiTabanidae). U.S.D.A.,
Coop. Plant Pest Rept. 4:220-224.

80. Hacker, J. D., L. Butler and . 1979. New geographical distribution records for

twenty-eight species and subspecies of tabanids in West Virginia (Diptera: Tabanidae).
Entomol. News 90:200-202.

81. Drees, B. M., L. Butler and . 1980. Horse flies and deer flies of West Virginia: an

illustrated key (Diptera: Tabanidae). West Virginia Univ. Agric. and For. Exp. Sta. Bull.
674:1-67.

82. 1981. Notes on Atylotus and description of a new species from eastern North America
(Diptera: Tabanidae). Entomol. News 92:1-6.

83. Burnett, A. M., C. V. Coveil, Jr. and . 1981. Records of Kentucky Tabanidae

(Diptera) including species new to the state fauna. Entomol. News 92:56-58.

84. and H. J. Teskey. 1981. Tabanidae. In Manual of Nearctic Diptera, Vol. 1. Can.

Dept. Agric. Res. Brch. Monograph no. 277:463—478.

85. and A. G. Wheeler, Jr. 1981. Northern range extension of Acanalonia conica

(Homoptera: Acanaloniidae). Entomol. News 92:98-100.

86. 1981. The horse flies and deer flies of New York (Diptera, Tabanidae). Second edition.
Search Agric., N.Y. State Agric. Exp. Stn. (Ithaca), no. 18:68 pp.

87. Hoebeke, E. R. and . 1982. New North American records for the Palearctic soldier

fly, Chloromyia formosa (Scopoli) (Diptera: Stratiomyidae). Entomol. News 93:177-179.

88. and E. R. Hoebeke. 1983. European Rhagionidae in eastern North America:

records of newly discovered species (Diptera). Entomol. News 94:86-88.

89. , D. W. Webb and H. J. Teskey. 1983. The Diptera, or true flies of Illinois. 1.

Tabanidae. Bull. 111. Nat. Hist. Surv. 33:1-121.

90. Mullens, B. A., and R. A. Rutz. 1983. A second finding of Entomophthora ta-

banivora and observations on Atylotus thoracicus (Diptera: Tabanidae). J. Invert. Pathol.
42:270-272.

91. White, D. J., K. Green and . 1983. New distribution records of nine species of

Tabanidae from the northern Adirondack Mountain region of New York State, USA. J.
Med. Entomol. 20:689.

92. 1985. The periodical cicada— Brood VII revisited (Homoptera: Cicadidae). Entomol. News
96:59-60.

93. 1985. Apatolestes philipi, a new horse fly from Texas (Diptera: Tabanidae). Miya 3:361—
365.

94. Wilkerson, R. C., J. F. Butler and . 1985. Swarming, hovering, and mating behavior

of male horse flies and deer flies (Diptera: Tabanidae). Miya 3:515-546.

95. Burger, J. F. and . 1 986. A review of the genus Haematopota (Diptera: Tabanidae)

in North America. J. Med. Entomol. 23:345-352.

96. Wright, R. E., R. K. Whittle and . 1986. Range extensions, new state records and

annotated checklist for Oklahoma Tabanidae (Diptera). J. Kans. Entomol. Soc. 59:235-
245.

97. Burger, J. F. and . 1989. The identity of Corizoneura caucasica Krober and con-

sequent change of name for some Palearctic species of Silvius (Diptera, Tabanidae). En-
tomofauna 10:445-452.

98. Burger, J. F., L. A. Martinez, and L. V. Bermudez. 1990. A revision of the horse

fly genus Agkistrocerus Philip (Diptera: Tabanidae). Pan-Pacif. Entomol. 66:181-194.

— E. Richard Hoebeke, Department of Entomology, Comstock Hall, Cornell Univer-
sity, Ithaca, New York 14853-0999.

1992

HONORARY MEMBERS

Dr. F. S. Chew
Dr. James Forbes
Dr. John G. Franclemont
Ms. Su Zan Nouguchi
Dr. James A. Slater

SUSTAINING MEMBERS

Dr. Richard S. Beal, Jr.

Dr. R. C. Froeschner
Dr. Dennis J. Joslyn
Mr. Ralph F. Kirchner
Mr. Raymond A. Mendez
Mr. Richard G. Robbins
Mr. Enrico Sismondo

LIFE MEMBERS

Dr. Annette Aiello
Prof. Clive F. Bailey

Dr. David G. Casdorph
Dr. Howard E. Evans
Mr. Irving Granek
Prof. Dipl. Ing. Ernst Heiss
Dr. Donald F. J. Hilton
Prof. Hussein S. Hussein
Mr. Mark Indenbaum
Dr. Gary G. Kennen
Dr. Brobson Lutz, M.D.

Dr. Gordon A. Marsh

Dr. John C. Morse

Mr. M. S. Moulds

Mr. Kikumaro Okano

Dr. John A. Shetterley

Dr. Kathryn M. Sommerman

Mr. Hubert J. Thelen

Dr. F. Christian Thompson

Mr. George F. Townes

Mr. Theodore H. Weisse

REVIEWERS

The outgoing editors acknowledge those who provided timely, critical reviews of
our 1992 manuscripts. Our thanks go to: J. S. Ashe, G. E. Ball, R. L. Brown, C.
Darling, R. L. Davidson, G. C. Eickwort, J. H. Frank, J. K. Gelhaus, G. L. Godfrey,
P. A. Goloboff, R. D. Gordon, D. A. Grimaldi, S. W. Hamilton, T. J. Henry, R. W.
Hodges, R. W. Holzenthal, D. J. Larson, J. D. Lattin, T. L. McCabe, S. McCleve,
J. E. McPherson, J. S. Miller, L. A. Mound, L. B. O’Brien, J. Pakaluk, S. B. Peck,
G. O. Poinar, Jr., R. W. Poole, J. E. Rawlins, R. K. Robbins, K. G. Ross, M. E.
Schauff, R. T. Schuh, A. M. Shapiro, O. Shields, J. Shuey, F. W. Stehr, K. R. Valley,
A. Venables, D. L. Wagner, A. G. Wheeler, Jr., S. W. Wilson, and J. B. Woolley.

CHANGE IN EDITOR

James M. Carpenter, of the Department of Entomology, American Museum of
Natural History, will assume the editorship of the Journal effective 1 November
1992. As of that date, all manuscripts should be submitted to him, using the con-
ventions of the Instructions to Authors on the inside back cover.

STATEMENT OF OWNERSHIP,

MANAGEMENT, AND CIRCULATION

1. Title of Publication: Journal of the New York Entomological Society (ISSN

0028-7199).

2. Date of Filing: November 1992.

3. Frequency of Issue: Quarterly.

4. Complete Mailing Address of Known Office of Publication: % American Mu-
seum of Natural History, Central Park West at 79th Street, New York, NY 10024-
5192.

5. Complete Mailing Address of the Headquarters or General Business Offices of
the Publishers: New York Entomological Society, % American Museum of Natural
History, Central Park West at 79th Street, New York, NY 10024-5192.

6. Full Names and Complete Mailing Addresses of Publishers, Editors, and Man-
aging Editor: Publisher: New York Entomological Society, % American Museum of
Natural History, Central Park West at 79th Street, New York, NY 10024-5192.

Editor and Managing Editor: James K. Liebherr. Assistant Editor: E. Richard Hoe-
beke. Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY
14853-0999.

7. Owner: New York Entomological Society (non-profit), % Department of En-
tomology, American Museum of Natural History, Central Park West at 79th Street,
New York, NY 10024-5192.

8. Known Bondholders, Mortgages, and Other Security Holders Owning or Hold-
ing 1% or More of Total Amount of Bonds, Mortgages, or Other Securities: None.

9* Purpose: The purpose, function, and non-profit status of this organization and
exempt status for federal income tax purposes have not changed during the preceding
12 months.

19. Extent and Nature of Circulation:

Avg. no. copies Actual no. copies

each issue of single issue

during preceding published nearest to
12 mo. filing date

A. Total no. copies (net press run)

B. Paid circulation

700

700

1. Sales through dealers and carriers,

street vendors, and counter sales

—

—

2. Mail subscription

552

531

C. Total paid circulation

D. Free distribution by mail, carrier, or

552

531

other means; samples, complimenta-
ry, and other free copies

19

18

E. Total distribution

F. Copies not distributed

571

549

1. Office use, left over, unaccounted.

spoiled after printing

129

151

G. Total

700

700

Journal of the

New York Entomological Society

published by

The New York Entomological Society

Contents Volume 100, 1992, Numbers 1-4

Number 1

Key and diagnoses for the genera of Ochlerini (Hemiptera: Pentatomidae: Dis-
cocephalinae) L. H. Rolston 1-41

Revision of the genus Thyanta Stal, 1862 (Heteroptera: Pentatomidae) II. North
America, Central America, and the West Indies

D. A. Rider and Joan B. Chapin 42-98

Revision of Arocera Spinola, with the description of two new species (Heteroptera:

Pentatomidae) D. A. Rider 99-136

Description of the male of Oxyporus balli Campbell, with notes on distribution
of Oxyporus in Mexico (Coleoptera: Staphylinidae)

Jose Luis Navarrete- Heredia 137-141

Review of the species of Telephanus Erichson from the Malagasy region, with
description of a new species (Coleoptera: Silvanidae) M. C. Thomas 142-154

Hydroptilidae (Trichoptera) of Costa Rica: The genus Oxyethira Eaton

Ralph W. Holzenthal and Steven C. Harris 155-177

On the spider subfamily Zavattaricinae (Araneae, Gnaphosidae)

Norman I. Plat nick 178-179

Notes and Comments

The identity and synonymy of Hydroessa fusca Germar (Veliidae: Heteroptera)

John T. Polhemus 180-181

Dispersal of the ergot fungus Claviceps purpurea by the lauxaniid fly Minettia
lupulina Kathleen M. Lemon 182-184

A new homonym in the Mezirinae (Hemiptera: Aradidae)

Nicholas A. Kormilev 1 84

Number 2

Contributions from former students
in honor of the 80th Birthday of
Professor John G. Franclemont

A remembrance of John G. Franclemont Ronald W. Hodges 185-190

Neotropical Microlepidoptera XXV, a new arboreal species of Acrolophus from
Peru (Lepidoptera: Acrolophidae) Donald R. Davis and Charles L. Hogue 191-202

Two new species of Mompha from California (Lepidoptera: Momphidae)

Ronald W. Hodges

Six new species of Catastega (Lepidoptera: Tortricidae: Eucosmini) from Mexico
and southwestern United States Richard L. Brown

Pupal morphology and the subfamily classification of the Notodontidae (Lepi-
doptera: Noctuoidea) James S. Miller

Franc/emontia interrogans (Walker), a new genus for an old species (Lepidoptera:
Noctuidae: Amphipyrinae) Douglas C. Ferguson

An annotated key and illustrations to the species of the genus Cucullia (Lepidop-
tera: Noctuidae: Cuculliinae) from Mexico and Central America

Robert W. Poole

A revision of the genus Hypsoropha Hiibner (Lepidoptera: Noctuidae: Ophider-
inae) Tim L. McCabe

Life history and systematics of the West Andean moth Aucula franclemonti with
description of a new species from Eduador (Lepidoptera: Noctuidae: Agaristi-
nae) John E. Rawlins

Twenty years of fluctuating parapatry and the question of competitive exclusion
in the butterflies Pontia occidentalis and P. protodice (Lepidoptera: Pieridae)

Arthur M. Shapiro

Hydropsyche franclemonti, a new species of the sea laris group from eastern North
America (Trichoptera: Hydropsychidae) Oliver S. Flint, Jr.

The Tribe Licinini (Coleoptera: Carabidae): a review of the genus-groups and of
the species of selected genera George E. Ball

Taxonomy and distribution of the athetine genus Lypoglossa Fenyes (Coleoptera:
Staphylinidae: Aleocharinae) in North America, with description of a new spe-
cies E. Richard Hoebeke

The New World Blepharida group, with a key to genera and description of a new
species (Coleoptera: Chrysomelidae) David G. Furth

Number 3

New synonyms of Frankliniella bondari and a review of the synonyms of F.
cephalica (Thysanoptera: Thripidae) Sueo Nakahara

The first records of the families Ochteridae and Hebridae (Heteroptera) from the
granitic Seychelles, with descriptions of two new species Dan A. Polhemus

A review of the species of Anthocoridae (Hemiptera: Heteroptera) found on Pinus
contorta J. D. Lattin and N. L. Stanton

Seasonal occurrence of the Miridae (Heteroptera) associated with Ohio Buckeye,
Aesculus glabra Willd., in Missouri R. L. Blinn

Priapismus pini, a new species of Discocephalini from Honduras on pine (He-
miptera: Penlalomidae: Discocephalinae) L. H. Rolston

Life history and descriptions of the immature stages of the planthopper Prokelisia
crocea (Van Duzee) (Homoplera: Delphacidae) Monte W. Holder and

Stephen W. Wilson

203-208

209-227

228-256

257-266

267-272

273-285

286-310

311-319

320-324

325-380

381-398

399-414

415-417

418-423

424-479

480-487

488-490

491-497

A unique Amycle nymph (Homoptera: Fulgoridae) that mimics jumping spiders
(Araneae: Salticidae) Gregory Zolnerowich 498-502

Bembidion femoratum Sturm and Amara communis (Panzer) (Coleoptera:

Carabidae) new to North America Yves Bousquet 503-509

Descriptions of new or poorly known species of Gastrosticta Casey, 1918 and
Paraferonina Ball, 1965 (Coleoptera: Carabidae: Pterostichus Bonelli, 1810)

Yves Bousquet 5 1 0-52 1

A high Andean new species of Terra (Lepidoptera, Lycaenidae) Kurt Johnson 522-526

Ovipositional behavior and larval development of Aphidencyrtus aphidivorus (Hy-
menoptera: Encyrtidae), an aphid hyperparasitoid Michael J. Kanuck and

Daniel J. Sullivan 527-532

Notes and Comments

New distributional records for semiaquatic Heteroptera, including a Microvelia
new to the United States (Heteroptera: Hydrometridae, Saldidae, Veliidae)

J. T. Polhemus 533-534

Number 4

A fossil Empicoris Wolff (Reduviidae: Heteroptera) from Mexican amber with
remarks on the phylogenetic status of the fossil genus Alumeda Popov

Donald B. Thomas 535-539

Discovery of sexual wing dimorphism in Staphylinidae (Coleoptera): “Omalium”
flavidum, and a discussion of wing dimorphism in insects

Margaret K. Thayer 540-573

Two new species of Diplotaxis from Mexico (Coleoptera: Melolonthidae)

Leonardo Delgado and Fabricio Capistran 574-580

Studies of Neotropical caddisflies, XLVII; Kumanskiel/a, a new genus of micro-
caddisflies from Cuba and Puerto Rico

Steven C. Harris and Oliver S. Flint, Jr. 581-593

First instar description of Korscheltellus gracilis (Grote) and Sthenopis auratus
(Grote) (Lepidoptera: Hepialidae) with a consideration of cladistic relationships
between setae Jonathan G. Leonard, John R. Grehan, and Bruce L. Parker 594-614

A new species of Orcya from the Maroni River Basin of French Guiana (Lepi-
doptera: Lycaenidae) Kurt Johnson 615-620

Novel taxa of Trichogramma from the New World tropics and Australia (Hy-
menoptera: Trichogrammatidae) John D. Pinto 621-633

Notes and Comments

Observations on the bizarre jelly mass habit of larval Geranomyia (Diptera:

Tipulidae: Limoniinae) David Grimaldi and Chen Young 634-637

Book Review

Women in the field, America’s pioneering women naturalists

Barbara L. Peckarsky 638-640

Obituary

In Memoriam LaVeme L. Pechuman (1913-1992) E. Richard Hoebeke 641-651

Honorary, Life, and Sustaining Members 652

Reviewers for 1992 652

Change in Editor 653

Statement of Ownership, Management, and Circulation 654

INSTRUCTIONS TO AUTHORS

The Journal of the New York Entomological Society publishes original research resulting
from the study of insects and related taxa. Research that contributes information on taxonomy,
classification, phylogeny, biogeography, behavior, natural history, or related fields will be con-
sidered for publication. The costs of publishing the Journal are paid by subscriptions, mem-
bership dues, page charges, and the proceeds from an endowment established with bequests
from the late C. P. Alexander and Patricia Vaurie.

Manuscripts should be submitted in triplicate to: Dr. James K. Liebherr, Editor, Journal of
the New York Entomological Society, Department of Entomology, Comstock Flail, Cornell
University, Ithaca, New York 14853-0999. Contributions must be written in English, double-
spaced (including references, tables, captions, etc.) and prepared in the format of a recent issue
of the Journal. Manuscripts of any length will be considered for publication. Longer papers will
be printed as articles, shorter ones as “scientific notes.” Book reviews will be solicited by the
Book Review Editor.

Longer manuscripts intended for submission as articles should be accompanied by a brief
abstract. Footnotes should be avoided. Tables should be prepared as separate pages; they should
be kept to a minimum because of the high cost of typesetting, but may be submitted as
photographically reproducible material (see below). The list of references is headed “Literature
Cited” and should follow the format indicated in the CBE Style Manual or as found in a recent
issue of the Journal. Manuscripts should be printed in roman, bold roman, or underlined roman
or bold roman to conform to journal typesetting style. Manuscripts should not contain any
italic fonts.

Illustrations (originals whenever possible) should be submitted flat, never rolled or folded,
with the manuscript. Size of illustrations should be kept manageable and suitable for shipment
through the US mail without being damaged. Proportions of illustrations should conform to
single column format, actual page size 1 1.5 x 17.5 cm. Please assemble illustrations in a com-
pact fashion in order to use journal space effectively. Mount line drawings and halftones on
separate plates. Figures should be numbered consecutively. Figure captions should be double-
spaced, grouped together and placed at the end of the manuscript. If tables are submitted for
photographic reproduction, they should be neat and clean and conform to journal format
proportions.

Authors will receive page proofs, with the original manuscripts and illustrations, directly
from the printer. Corrected proofs should be returned promptly to the Editor to avoid publi-
cation delays. Revisions in proof should be kept to the absolute minimum. Alterations in proof
will be charged to the author at the rate of $3.25 per revision line.

Society members will be charged a fee of $23.00 per printed page and $8.50 per plate of
figures. Non-members will be charged $65.00 per printed page and $15.00 per plate of figures.
Member authors who are students, or who do not have institutional affiliation may petition to
the Society for waiver of page charges for no more than eight pages on a once a year basis.
Because of limited funds, all such requests will be handled on a first-come first-serve basis.

Authors will receive a reprint order blank with the proofs. Reprints are ordered directly from
the printer with no benefit accruing to the Society.

Journal of the

New York Entomological Society

VOLUME 100 & OCTOBER 1992 NO. 4

CONTENTS

A fossil Empicoris Wolff (Reduviidae: Heteroptera) from Mexican amber with
remarks on the phylogenetic status of the fossil genus A/umeda Popov

Donald B. Thomas

Discovery of sexual wing dimorphism in Staphylinidae (Coleoptera): “ Omalium ”
flavidum, and a discussion of wing dimorphism in insects

Margaret K. Thayer

Two new species of Diplot axis from Mexico (Coleoptera: Melolonthidae)

Leonardo Delgado and Fabricio Capistran

Studies of Neotropical caddisflies, XLVII; Kumanskiella, a new genus of micro-
caddisflies from Cuba and Puerto Rico

Steven C. Harris and Oliver S. Flint , Jr.

First instar description of Korscheltellus gracilis (Grote) and Sthenopis auratus
(Grote) (Lepidoptera: Hepialidae) with a consideration of cladistic relationships
between setae Jonathan G. Leonard , John R. Grehan, and Bruce L. Parker

A new species of Orcya from the Maroni River Basin of French Guiana (Lepi-
doptera: Lycaenidae) Kurt Johnson

Novel taxa of Trichogramma from the New World tropics and Australia (Hy-
menoptera: Trichogrammatidae) John D. Pinto

Notes and Comments

Observations on the bizarre jelly mass habit of larval Geranomyia (Diptera:
Tipulidae: Limoniinae) David Grimaldi and Chen Young

Book Review

Women in the held, America’s pioneering women naturalists

Barbara L. Peckarsky

Obituary

In Memoriam LaVerne L. Pechuman (1913-1992) E. Richard Hoebeke

Honorary, Life, and Sustaining Members

Reviewers for 1992

Change in Editor

Statement of Ownership, Management, and Circulation

535-539

540-573

574-580

581-593

594-614

615-620

621-633

634-637

638-640

641-651

652

652

653

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