Title: Florida Entomologist
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00098813/00105
 Material Information
Title: Florida Entomologist
Physical Description: Serial
Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 1981
Copyright Date: 1917
Subject: Florida Entomological Society
Entomology -- Periodicals
Insects -- Florida
Insects -- Florida -- Periodicals
Insects -- Periodicals
General Note: Eigenfactor: Florida Entomologist: http://www.bioone.org/doi/full/10.1653/024.092.0401
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Bibliographic ID: UF00098813
Volume ID: VID00105
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: Open Access

Full Text

(ISSN 0015-4040)


(An International Journal for the Americas)

Volume 64, No.2 June, 1981

Published by the Florida Entomological Society
LUCAS, J., AND L. A. STANGE-Key and Descriptions to the Myrmeleon
Larvae of Florida (Neuroptera: Myrmeleontidae) -...-........ 207
CLARK, W. E.-The Weevil Genus Hamaba Casey (Coleoptera: Cur-
culionidae) ...----....... .. ..... .. .................... .. 217
AVERILL, A. L., AND R. J. PROKOPY-Oviposition Deterring Fruit Mark-
ing Pheromone in Rhagoletis -... -...... --..-------.-.. ....- ....... 222
BAER, R. G., AND M. KOSZTARAB-Two New Species of Gall-like Coccoids
on Oaks From the Nearctic Region .- ...-.......-.. ................ 226
PORTER, C. C.-Certonotus Kriechbaumer (Hymenoptera: Ichneum-
onidae) An Australian Genus Newly Recorded in South America 235
SNIDER, R.-Pseudachorutes (Pseudachorutes) quadrisetosus, A New
Species of Collembola from Florida (Hypogastruridae: Neanu-
rinae) .......----- ..... ........................ ... ..... 244
SLATER, J. A., AND M. HASSEY-The Distribution and Systematics of
Ozophora atropicta Barber, With the Description of a New Spe-
cies from the Neotropics .----.... -- ..-. ........ .................... 246
PORTER, C. C.-New Records for Monobia Saussure (Hymenoptera:
Eumenidae) in South Texas and Northeast Mexico ..-. -.....--. 260
AIKEN, R. B.-The Relationship Between Body Weight and Homosexual
Mounting in Palmacorixa nana Walley (Heteroptera: Corixidae) 267
HEPPNER, J. B.-Two New Dichrorampha (Lepidoptera: Torticidae)
From Florida ....----. --- ----- -- --- --.... -........ ........... 271
MUMA, M. H.-Adult Activity of Ground-Surface Spiders in Arid-
Grassland and Pinyon-Juniper Associations in Southwestern
New Mexico ...--.---------................... ......... 276
BROOKHART, J. 0., AND M. H. MUMA-The pallipes Species-Group of
Eremobates Banks (Solpugida: Arachnida) in the United States 283
HEPPNER, J. B.-Revision of the New Genus Diploschizia (Lepidoptera:
Glyphipterigidae) for North America --------............... .... ....... 309
FRANK, J. H.-A Review of Teratology in Staphylinidae With Descrip-
tion of a Teratological Specimen of Tachinus axillaris Erichson
(Coleoptera: Staphylinidae) From Florida -.-- --...... ---..............._ 337
Continued on Back Cover

Published by The Florida Entomological Society


President ....-........----------------..... ......-..---....-.....--------------- E. C. Beck
Vice-President .-................-.....-..-....--.... ------ ..................... W. L. Peters
Secretary ..-............--.......-...--------------.. F. W. Mead
Treasurer ................................ -.. --- D. P. Wojcik

SR. W. Flowers
N. C. Leppla
Other Members of Executive Committee R. H. Maltby
C. A. Musgrave Sutherland
J. L. Taylor


Editor ..........................---.. ---...------ C. A. Musgrave Sutherland
Associate Editors .................... -------------------- J. E. Lloyd
J. R. McLaughlin
C. W. McCoy
A. R. Soponis
H. V. Weems, Jr.
Business Manager ..............--.................----- -------------- D. P. Wojcik

FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September,
and December. Subscription price to non-members is $15.00 per year in
advance, $5.00 per copy. Membership in the Florida Entomological Society,
including subscription to Florida Entomologist, is $10 per year for regular
membership and $2 per year for students. Inquiries regarding membership
and subscriptions should be addressed to the Business Manager, P. 0. Box
12425, University Station, Gainesville, FL 32604. Florida Entomologist is
entered as second class matter at the Post Office in DeLeon Springs and
Gainesville, FL.
Authors should consult "Instructions to Authors" on the inside cover of
all recent issues while preparing manuscripts or notes. When submitting a
paper or note to the Editor, please send the original manuscript, original
figures and tables, and 3 copies of the entire paper. Include an abstract and
title in Spanish, if possible. Upon receipt, manuscripts and notes are ac-
knowledged by the Editor and assigned to an appropriate Associate Editor
who will make every effort to recruit peer reviewers not employed by the
same agency or institution as the authors (s). Reviews from individuals
working out-of-state or in nearby countries (e.g. Canada, Mexico, and
others) will be obtained where possible.
Manuscripts and other editorial matter should be sent to the Editor,
C. A. Musgrave Sutherland, Rt. 3, Box 115H, Las Cruces, NM 88001.
Business matters for other Society officers can be sent to that individual at
the University Station address above.

This issue mailed June 30, 1981

Lucas & Stange: Florida Myrmeleon Larvae



Descriptions of the 3rd instar larva of 5 of the 6 species of Myrmeleon
found in Florida are given. Only M. immaculatus De Geer has been described
previously. Myrmeleon insertus Hagen is a new record for the state. A key to
the species is provided with the diagnostic characters (chaetotaxy of the
mandible; pigmentation pattern of the ventral surface of the head) illus-

Clave y descripciones de las larvas de Myrmeleon de Florida. Se describe
el tercer estadio de las larvas de cinco de las seis species floridanas. Son M.
carolinus Banks, M. crudelis Walker, M. immaculatus De Geer, M. insertus
Hagen, y M. mobilis Hagen. La larva de M. heriocles Hagen es desconocida.
Se presentan dibujos y una clave para las species basada sobre las setas de
la mandibula y la pigmentacion de la superficie ventral de la capsula cefalica.

Five species of the ant-lion genus Myrmeleon have been recorded from
Florida. These are immaculatus De Geer, mobilis Hagen, crudelis Walker,
heriocles Banks, and carolinus (Banks 1928, 1943). In addition, we add to the
list a sixth species, insertus Hagen, known only from the Florida Keys and
the West Indies.
The larvae construct pit-fall traps and can move only backward, be-
havioral traits which distinguish them from all other ant-lion larvae in
Florida. Structurally, the short labial palps (shorter than basal width of
mandible) and the long 3-toothed mandibles with long setae between the
teeth (Fig. 1) serve to distinguish them from other genera in Florida
(Stange 1970). Only the larva of immaculatus has been described (Hagen
1873, Wheeler 1930). In this paper we provide descriptions and drawings to
distinguish 5 of the 6 known species (heriocles is unknown in the larval
stage). Myrmeleon carolinus, crudelis, mobilis, and insertus have been reared
several times from various localities in Florida, but immaculatus has been
found only as an adult; the larval description is based on material from other
All specimens examined are preserved in the Florida State Collection of
Arthropods, Gainesville.

'A joint contribution of the University of Florida, Department of Zoology, Gainesville, and
the Florida Dept. of Agr. and Consumer Services, Division of Plant Industry. Contribution No.
489, Bureau of Entomology, DPI.
2University of Florida, Department of Zoology, Gainesville, FL 32611 USA.
3Florida Department of Agriculture and Consumer Services, Division of Plant Industry,
Gainesville, FL 32602 USA.


208 Florida Entomologist 64(2) June, 1981

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Lucas & Stange: Florida Myrmeleon Larvae

I Inner Jaw

Distal Palpomere
S/ with Sensory Area

Fig. 1. Ventral view of head capsule of Myrmeleon (modified from
Principi 1943).

From the 1st to 3rd instar the ratio of head width to mandible length
generally increases for all species, and the ratio of head width to head length
generally decreases. Thus, earlier instars have relatively longer mandibles
and stouter heads than later instars.
The following descriptions pertain to the 3rd instar, the last before pupa-

Specimens were examined from 36 locations representing 21 different
counties in Florida. Larval specimens of M. immaculatus were studied from
Missouri and Ohio. We reared specimens of carolinus (89 reared), crudelis
133 reared), mobilis (66 reared), and insertus (8 reared). To characterize


Florida Entomologist 64 (2)

differences between species in size and shape of the head, we measured head
width, head length, and mandible length (Table 1). Head width was taken
near the mandibles where the width was maximum. Head length was meas-
ured from the point mid-way between the mandible bases to the median
posterior point of the head. Mandible length was measured from the tip of
the right mandible to its inner base. We also describe station patterns from
the ventral side of the mandibles that are species specific. These setae, which
we call inner jaw setae (Fig. 1), are found on the inner side of the mandi-
bles near the teeth. These setae originate from the ventral side of the mandi-
ble. All ant-lions have setae originating from the lateral inner border. All
counts are given for the right mandible. The 3 teeth on the inside of the jaw
are numbered 1-3, starting from the tooth nearest the base of the mandible
(as in Doflein 1916).
We studied 166 larvae of carolinus, 168 of crudelis, 106 of mobilis, and 19
of inserts. Nine specimens of immaculatus from Ohio and Missouri were
studied to provide tentative identification, although Florida specimens are
needed to confirm the pigment pattern.

Five species of ant-lions were examined: M. immaculatus, mobilis,
crudelis, carolinus, and insertus. The 6th species, heriocles, which is known
only from a few adult specimens, was not available as a larva and is not in-
cluded in this study.
The character that most easily distinguishes the species is the pigmenta-
tion on the ventral side of the head. Although one basic pattern is found in
all species, each exhibits a unique variation. The pattern consists of a
V-shaped mark below the labium with 2 spots, 1 on either side of the head,
below the "V" (Fig. 2-5). The pattern for each species is described below
with the species descriptions.
Banks (1928, 1943) separated immaculatus and mobilis from crudelis and
carolinus adults on the basis of several morphological characters, including
size. Insertus is approximately the same size as crudelis and thus the 5 spe-
cies can be broken into large (mobilis and immaculatus) and small (insertus,
carolinus, and crudelis) ant-lions.
In general, the color patterns on the ventral side of the head are retained
through all 3 instars. Occasionally, the head of the 1st instar carolinus is
entirely black below and black with a median tan stripe dorsally. The number
of inner jaw setae increases as the ant-lion grows from 1st instar through
the 3rd instar, and therefore this character is useful in distinguishing only
the 3rd instar (Table 1).

Myrmeleon carolinus Banks 1943
(Fig. 2)'

DESCRIPTION: Head oval-shaped, more elongate than other species. Head
width (1.40 mm) approximately equal to that of crudelis, smaller than im-
maculatus and mobilis. Mandibles shorter and broader than those of other
species. Inner jaw setae always extend past the 1st tooth and usually past
the 2nd (Table 1). Setae on head, especially lateral and ventral, shorter than
all other species.


June, 1981

Lucas & Stange: Florida Myrmeleon Larvae 211

PIGMENTATION: Head and mandibles brown. Ventral and dorsal side of body
with rows of black pigment on gray background. Ventral side of head: "V"
broad with rounded tips, arms separated. Spots large and circular, generally
well defined. Lighter pigmentation from "V" to spots forms "M" shape (Fig.
2). Dorsal side of head: Almost entirely dark brown. Median tan stripe in
posterior portion and 2 "V"-shaped tan marks behind each eye.
DISTRIBUTION: Found throughout North and Central Florida. We have rec-
ords only as far south as Highlands County. Banks (1943) reported carolinus
from Florida, Georgia, and North Carolina. M. carolinus is generally found



Fig. 2. Ventral view of head capsule of Myrmeleon carolinus Banks.

Florida Entomologist 64(2)

in open, sandy areas, although it is occasionally collected in sheltered areas
with crudelis.
SPECIMENS EXAMINED: FLORIDA: Alachua Co., Gainesville (12) III, V, X,
various collectors; Bay Co., St. Andrews State Park (10) 8-111-1980, B. Miller
& L. Stange; Dixie Co., 4 miles north Old Town (7) 1-V-1980, J. Lucas &
L. Stange; Highlands Co., Archbold Biological Station (4) 13-III-1980,
B. Miller & L. Stange, 5 miles south Sebring (15) 12-III-1980, B. Miller &
L. Stange; Franklin Co., Carrabelle (6) 10-IV-1978, L. Stange; Okaloosa Co.,
2 miles north Holt (3) 7-III-1980, B. Miller & L. Stange; St. Johns Co., Fort
Matanzas (2) 7-VII-1978, L. Stange, Ponte Vedra Beach (11) 7-VII-1978,
L. Stange.

Myrmeleon crudelis Walker 1852
(Fig. 3)

DESCRIPTION: Head somewhat rectangular with lateral indentations, similar
in shape to insertus. Head capsule width (1.38 mm) approximately same size
as carolinus, and smaller than mobilis and immaculatus. Mandible length to
head width ratio as well as head width to head length ratio almost identical

Fig. 3-4. Ventral view of head capsules; 3. Myrmeleon crudelis Walker,
4. M. insertus Hagen.


June, 1981

Lucas & Stange: Florida Myrmeleon Larvae

to insertus. Average of 8 inner jaw setae from mandible base to 1st tooth.
Generally none past 1st tooth. Head relatively long.
PIGMENTATION: Head and mandibles light brown to tan. Dorsal and ventral
body with rows of black spots on pink to light brown background. Ventral
side of head: "V" arms bar-shaped, not attached at base. Spots elongate and
angled toward "V". Pigment patterns very distinct (Fig. 3). Dorsal side of
head: Posterior portion dark brown with 4 large parallel bars. Anterior
light brown with 2 dark triangular spots behind antennae, entire area be-
tween antennae dark.
DISTRIBUTION: Found throughout Florida from the Panhandle to Dade and
Collier counties. Not found in the Florida Keys. Banks (1928) reported
crudelis from Florida, Georgia, New Jersey, North Carolina, Texas, and
Virginia. This species is generally found in sheltered areas in a variety of
soil types, although it is occasionally collected in open areas where carolinus
is abundant, often collected with mobilis.
SPECIMENS EXAMINED: FLORIDA. Alachua Co., Gainesville (29) year
round; Collier Co., Marco Island (20) 9-IV-1952, R. E. Woodruff; Dade Co.,
Matheson Hammock (8) 14-II-1978, L. Stange, Miami (2) 4-IV-1980,
L. Stange; Dixie Co., 4 miles north Old Town (9) 1-1-1980, J. Lucas &
L. Stange; Highlands Co., Archbold Biological Station (1) 13-III-1980,
B. Miller & L. Stange, 5 miles south Lake Placid (4) 21-III-1978, L. Stange,
Highlands Hammock (14) 4-III-1978, L. Stange; Hernando Co., Bayport (2)
22-IV-1978, L. Stange; Franklin Co., Alligator Point (5) 9-IV-1978,
L. Stange; Indian River Co., Sebastian (2) 25-1-1962, H. Weems; Lee Co.,
Sanibel Island (3) 20-IV-1978, L. Stange; Levy Co., Seahorse Key (5) 10-
III-1980, J. Lucas & L. Stange; Martin Co., Jonathan Dickinson State Park
(4) 15-II-1978, L. Stange; Pasco Co., Bayonet Pass (9) 22-IV-1978,
L. Stange; Pinellas Co., Tierra Verde (1) 5-VII-1978, L. Stange; St. Johns
Co., Fort Matanzas (3) 7-VII-1978, L. Stange.

Myrmeleon insertus Hagen 1861
(Fig. 4)

DESCRIPTION: Head somewhat rectangular with lateral indentations, very
similar in shape to that of crudelis. Head capsule width (1.31 mm) smallest
of the 5 species but similar to crudelis and carolinus. Mandible length to
head width ratio and head width to head length ratio almost like crudelis.
Average of 3 inner jaw setae near the mandible base, none past the 1st
tooth (Table 1). Head setae very long compared to other species.
PIGMENTATION: Head and mandibles light brown to tan. Ventral and dorsal
body with rows of black spots on pink to light brown background. Ventral
side of head: "V" short and blunt, arms not attached but each arm often
connected to elongated and bar-shaped spots. Together they form an "H"
shape (Fig. 4). Dorsal side of head: Posterior portion dark brown with 4
large parallel bars. Anterior light brown with 4 dark round spots, forming
a "Y" shape between antennae, with base of "Y" pointing anteriorly.
DISTRIBUTION: In Florida, insertus is known only from the Florida Keys.
Specimens were collected in sheltered sandy soils. It is also recorded from
Cuba (Hagen 1861, Alayo 1968), Haiti (Smith 1931), and the Dominican
Republic (Hagen 1861).
SPECIMENS EXAMINED: FLORIDA. Monroe Co., Big Pine Key (4) 24-III-


Florida Entomologist 64 (2)

1978, L. Stange, Ramrod Key (4) 25-III-1978, L. Stange, Bahia Honda Key
(11) 22-XI-1980, L. Stange.
Myrmeleon mobilis Hagen 1888
(Fig. 5)
DESCRIPTION: Head generally square with slight lateral indentations. Head
stouter than carolinus, crudelis, and inserts. Head width (1.77 mm) larger
than carolinus, crudelis, and insertus, and somewhat smaller than im-
maculatus. In relationship to head width, mandibles are the longest of the 5
species. Inner jaw setae entirely absent (Table 1). Setae on head longer than
those of carolinus but proportionately shorter than crudelis and insertus.
PIGMENTATION: Head and mandibles light brown to tan. Dorsal and ventral
body with rows of dark pigment on light brown background. Ventral side of
body: "V" broad at base, arms often connected, tips pointed. Spots generally
diffuse (Fig. 5). Dorsal side of head: Lighter than crudelis, carolinus, and
insertus. Four widely spaced blotches located posteriorly with 4 spots anterior
to blotches. Posterior 2 spots triangular as in crudelis, anterior most spots
are "T"-shaped.
DISTRIBUTION: We have records from the Panhandle south to Collier County.
Not found in the Florida Keys. Banks (1928) reported mobilis from Alabama

Fig. 5-6. Ventral view of head capsules; 5. Myrmeleon mobilis Hagen, 6.
M. immaculatus De Geer.


June, 1981

Lucas & Stange: Florida Myrmeleon Larvae

and Georgia. Myrmeleon mobilis is generally found in finer soils in sheltered
areas, often collected with crudelis.
SPECIMENS EXAMINED: FLORIDA. Alachua Co., Archer (1) 1-VIII-1967,
P. Choate, Gainesville (15) 1-V-1980, J. Lucas & L. Stange; Highlands Co.,
Archbold Biological Station (5) 23-III-1978, L. Stange, Highlands Hammock
(3) 4-III-1978, L. Stange, 5 miles south of Lake Placid (1) 21-III-1978,
L. Stange; Jackson Co., Florida Caverns (10) 14-IV-1978, L. Stange;
Okaloosa Co., 4 miles north Holt (3) 17-IV-1978, L. Stange.

Myrmeleon immaculatus De Geer 1773
(Fig. 6)
Hagen 1873: 291.
Wheeler 1930: 99, Fig. 13.
DESCRIPTION: Head square, more so than in mobilis, ratio of width to length
highest of the 5 species. Head width (2.05 mm) largest of all species. Mandi-
bles intermediate in length, not so long (in proportion to head) as mobilis.
Average of 3 inner jaw setae near the base of the mandibles (Table 1).
Setae on head short but not as short as carolinus.
PIGMENTATION: Head and mandibles light brown to tan. Ventral and dorsal
sides of body with rows of black pigment on pink to light brown background.
Ventral side of head: Less pigmentation than other species. "V" very short,
forms 2 almost round bars. Spots elongate, pointing toward "V" (Fig. 6).
Dorsal side of head: Three prominent black spots, 1 between the antennae
and 2 side by side between them. Two other lighter pigmented areas behind
the black spots.
DISTRIBUTION: Found only in the northernmost parts of Florida. Myrmeleon
immaculatus has one of the most restricted distributions of ant-lions in
Florida, but it is widely distributed in North America, found throughout the
eastern U.S. and southeastern Canada (Banks 1928).
SPECIMENS EXAMINED: OHIO. Hocking Co. (2) 1-V-1954, T. V. Wallace;
Ottawa Co. Rockyridge (1) 30-IX-1972, J. Balciunas; MISSOURI. Boone
Co., Columbia (3) 11-VIII-1967, S. L. Poe; Marion Co. (3) 24-VII-1972,
C. Childers.

Key to 3rd Instar Larvae of Myrmeleon of Florida
1. Mandible without inner jaw setae (Fig. 5) ; head width greater
than 1.6 mm ------------------ ---.---- ....... ........ mobilis Hagen
1'. Mandible with inner jaw setae (Fig. 2-4, 6); head width less
than 1.6 mm (except immaculatus) ...------ .. ................. .. 2
2. Mandible with inner jaw setae extending to or beyond 2nd tooth
(Fig. 2); head much darker than grayish body ....-. corolinus Banks
2(1'). Mandible with inner jaw setae extending no further than 1st
tooth; head coloration similar to that of light tan body .---......--------- 3
3. Pigmentation on ventral side of head with anterior set of bars
tilting toward center usually forming a "V" (Fig. 3) ; mandible
with inner jaw setae (usually more than 6) extending almost
to 1st tooth ... -.-........ ...------ ........... .. crudelis W alker
3(2'). Pigmentation on ventral side of head forming "H" mark
(insertus) or much reduced (immaculatus); mandible with
inner jaw setae (often less than 6) not extending to 1st tooth ........-- 4


Florida Entomologist 64(2)

4. Pigmentation on ventral side of head forming "H" mark (Fig.
4); head width less than 1.6 mm .-- ------ ----- inserts Hagen
4(3'). Pigmentation on ventral side of head much reduced (Fig. 6);
head width more than 1.6 mm immaculatus De Geer


The authors are indebted to Ms. Tina Pellicane for the preparation of
Figures 2-6.

ALAYO, P. 1968. Los Neur6pteros de Cuba. Poeyana (B) No. 2: 5-127.
BANKS, N. 1928. Revision of the nearctic Myrmeleonidae. Bull. Mus. Comp.
Zool. 68(1927): 3-84.
1943. New Neuroptera and Trichoptera from the United States.
Psyche 50: 74-81.
DOFLEIN, F. 1916. Der Ameisenlawe. Eine Biologische, Tierpsychologische
und Reflexbiologische Untersuchung. Jena, Gustav Fischer, p. 1-138,
Tab. 1-x.
HAGEN, H. 1861. Synopsis of the Neuroptera of North America with a list
of the South American species. Smithsonian Misc. Coll., p. 1-347.
1873. Die Larven von Myrmeleon. Stettin ent. Ztg. 34: 249-95.
1888. Stray notes on the Myrmeleonidae, Part. 5. Canadian Ent.
20: 204-11.
MOODY, H. L. 1873. Notes on the habits of the ant lion. Canadian Ent. 5:
PRINCIPI, M. 1943. Contributi allo studio dei Neurotteri italiania. II.
Myrmeleon inconspicuus Ramb. ed Euroleon nostras Fourcroy. Boll.
Ist. Ent. Univ. Bologna 14: 131-92.
REDTENBACHER, J. 1884. tbersicht der Myrmeleoniden-larven. Denk. Ak.
Wien 48: 335-68, pl. i-vii.
SMITH, R. 1931. The Neuroptera of Haiti, West Indies. Ann. Ent. Soc.
America 24: 798-823.
STANCE, L. A. 1970. Revision of the ant-lion tribe Brachynemurini of North
America. Univ. California Publ. Ent. 55: 1-192.
WALKER, F. 1853. List of the Neuropterous insects in the collection of the
British Museum. Part II (Sialidae-Nemopterides). London, p. 193-476.
WHEELER, W. M. 1930. Demons of the Dust. W. W. Horton & Co., New York,
378 p.

June, 1981

Clark: Hamaba Casey


Department of Zoology-Entomology,
Alabama Agricultural Experiment Station,
Auburn University, Auburn, AL 36849 USA

The genus Hamaba Casey contains 2 species, H. bahamensis Casey in the
Bahamas, southern Florida, and the island of Cozumel, Mexico, and H.
dispersa Casey in the Bahamas. Hamaba minima (Blatchley) from Florida is
newly placed in synonymy under H. bahamensis. The genus and species are
redescribed and illustrated, and characters by which the species can be dis-
tinguished are presented.

El genero Hamaba Casey contiene 2 species, H. bahamensis en las Ba-
hamas, el sur de Florida, y la Isla de Cozumel (Mexico) y H. disperse Casey
en las Bahamas. Hamaba minima (Blatchley) de Florida es nuevamente
puesto en sinonomia debajo H. bahamensis. Do nuevo se describe y se pre-
sentan dibujos del g6nero y de las species y se presentan los caracteres que
las distinguen.

The genus Hamaba was erected by Casey (1910) for 2 species from the
Bahamas, H. bahamensis Casey and H. disperse Casey. Champion (1910)
made passing mention of the taxon, but except for entries in catalogs and
checklists (Klima 1934, Blackwelder 1947), it did not receive further atten-
tion until Sleeper (1957) transferred to it the Floridian Paragoges minimus
Blatchley (1916). The genus was included in a key to the curculionid genera
of North America by Kissinger (1964) who placed it in the subfamily
Anthonominae, tribe Endaeini. It was subsequently transferred to the sub-
family Tychiinae, tribe Lignyodini, by Clark et al. (1977).
Renewed interest in Hamaba has come about because of ongoing re-
visionary studies of the lignyodine genera, Chionanthobius Pierce (Clark and
Anderson in Press), Lignyodes Dejean (Clark 1980a), Neotylopterus
Hustache (Clark 1980), Plocetes LeConte (see Clark et al. 1977), and Rosella
Whitehead (Clark 1980b). The species of Hamaba are smaller than other
known lignyodines (both are less than 2 mm in length) and have 6 rather
than 7 funicular articles. Discussion of relationships of Hamaba to the other
lignyodine genera is deferred until further progress has been made on the
revisionary studies.
Specimens examined in this study, including type-specimens of all
named taxa, were obtained by loan from the American Museum of Natural
History (AMNH), New York, NY; the Museum of Comparative Zoology
(MCZ), Cambridge, MA; Purdue University (PURC), West Lafayette, IN;
and the National Museum of Natural History (USNM), Washington, DC.


Florida Entomologist 64 (2)

Genus Hamaba Casey
Hamaba Casey 1910: 131. Type-species, Hamaba bahamensis Casey, by orig-
inal designation. Champion 1910: 191. Klima 1934: 3. Blackwelder 1947:
842. Sleeper 1957: 39. Kissinger 1964: 54. Clark et al. 1977: 10.
DIAGNOSIS-Small Lignyodini, length 1.15-1.80 mm; antennae with 6
funicular articles; elytra with more-or-less distinct, posteromedian and sub-
apical, zigzag-shaped, transverse bands of dark scales on dark integument;
base of spiculum gastrale of 8 genitalia narrow (Fig. 8).
DESCRIPTION-Head: vertex and gena with long, narrow scales, venter
with broad, slightly imbricated scales; eyes nearly round, height ca. 1.2x
length. Rostrum (Fig. 1-4) : subequal to pronotum in length; scrobe not de-
scending, dorsal margin subcarinate, ending just before lower portion of eye,
ventral marking obsolete. Prothorax: in dorsal view, sides strongly rounded,
narrower at apex than at base, subapical constriction feebly developed;
dorsum with admixture of narrow, acuminate, aeneus scales, and broader,
longer, whitish scales, the latter abundant laterally and in median vitta;
aeneus to fuscoaeneus scales on dark integument form broad, more-or-less
distinct, lateromedian vittae; pleuron with oblong, whitish, nonimbricated
scales, and intermixed, narrower, pallid aeneus scales, and with small mid-
dorsal patch of fuscous scales. Elytra: humeri and apices of interspaces 4-6
not prominent; interspaces subequal in width, odd-numbered interspaces not
prominent; each interspace with uniform vestiture of small, narrow, re-
cumbent scales, and sparser, slightly-to-much-larger, slightly raised scales
which form diffuse median rows, the latter most abundant on odd-numbered
interspaces and on declivities; most scales pallid aeneus to whitish, but
darker aeneus to fuscoaeneus scales present over darker integument, at least
in more-or-less prominent, more or less distinctly zigzag-shaped, transverse,
posteromedian band, and broader, subapical band which extends posteriorly
to apices on sutural interspaces. Abdomen: sterna 4 and 5 fused, interseg-
mental line feebly discernable medially; sterna 1-2 flat, sterna 4-5 broadly
convex medially; scales on median portions of all sterna seta-like. Legs:
femora unarmed, integument uniformly ferruginous, with sparse, long, white,
seta-like scales; tibiae straight, not sinuate on ventral margins, pro- and
meso-tibiae each with small, acute, apical uncus; metatibial uncus minute,
curved, parallel to long axis of tibia.

Hamaba bahamensis Casey
(Fig. 1, 2, 5, 7, 8)
Hamaba bahamensis Casey 1910: 131. Lectotype (here designated), 8,
labelled "Egg Is. Bahamas/May 13, Wickham," "Casey/bequest/1925,"
"bahamensis-4/PARATYPE USNM/36740" (USNM); paralectotypes, 4
9 9, 1 labelled "Eleuthera Bahamas/July 9-15, Wickham," "Casey/be-
quest/1925," and "TYPE USNM 36740." "Hamaba/bahamensis,/Csy.,"
and 3 with the first 2 labels the same, the third label bahamensiss 2 (3
and 5)/PARATYPE USNM/36740," and lacking the "Hamaba/ba-
hamensis,/Csy." label (all USNM). Champion 1910: 191. Klima 1934: 3.
Blackwelder 1947: 842. Sleeper 1957: 39. Clark et al. 1977: 10.
Paragoges minimus Blatchley 1916: 247. Holotype, 8, labelled "TYPE."
"Ft. Myers Fla./W.S.B. Col./3-11, 1911," "Purdue/Blatchley/ Collection,"


June, 1981

Clark: Hamaba Casey

(PURC). Leng 1920: 321. Blatchley 1930: 40. Klima 1934: 46. NEW
Hamaba minima (Blatchley) : Sleeper 1957: 39. Kissinger 1964: 54.
DIAGNOSIS-Rostrum (Fig. 1, 2) in profile most strongly curved at and
just above antennal insertions, distal portion tapered; whitish, recumbent
scales on midbasal and humeral portions of elytra ovate, acuminate, integu-
ment not broadly exposed; median lobe of & genitalia (Fig. 5) slightly ex-
panded and rounded or slightly emarginate apically.


1 2 3 4

% 4.
~-~2;;;~=1~ ~5L


Fig. 1-9. Hamaba spp. 1-4) heads and rostra, lateral views: 1) H. ba-
hamensis, $, 2) H. bahamensis, 3) H. dispersa, $, 4) H. disperse, 2, 5-6
median lobe and associated structures of S genitalia: 5) H. bahamensis, 6)
H. disperse. 7, 9) spermathecae: 7) H. bahamensis, 9) H. disperse. 8)
spiculum gastrale of S genitalia, H. bahamensis.


220 Florida Entomologist 64 (2) June, 1981

MALE-Length: 1.15-1.50 (1.41) mm. Width: 0.67-0.76 (0.71) mm. Head:
eyes separated by ca. 0.40x eye length; hind margin of eye very slightly
raised. Rostrum (Fig. 1): short, 0.90-1.03 (0.97) x pronotum length; in
profile, dorsal margin arcuate over and just behind antennal insertions,
nearly straight basally and distally; in dorsal view, sides of proximal portion
subparallel, sides of distal portion very slightly widened to tip; proximal por-
tion 53-61 (57)% of total rostral length; lateral sulcus broad, shallow,
dorsolateral sulcus narrower, deeper, dorsal midline broad, smooth; proximal
portion with sparse, narrow, inconspicuous scales, broader scales at extreme
base; distal portion, in lateral view, tapered, smooth, except for dorsolateral
sulci which end just beyond antennal insertions, glabrous. Elytra: recumbent
scales on interspaces small, short, ovate, apically attenuate, slightly-raised
scales mostly attenuate; distinct, prominently zigzag-shaped transverse
posteromedian and subapical dark bands present, posteromedian band usually
widened to form median, more-or-less distinctly diamond-shaped macula, dark
patches also present on midbasal portions of elytra in some specimens.
FEMALE-Length: 1.17-1.59 (1.47) mm. Width: 0.63-0.83 (0.76) mm.
Rostrum (Fig. 2): more slender than that of male, especially distally, 0.93-
1.14 (1.00) x pronotum length; in profile, dorsal margin strongly curved
just posterior to antennal insertions, straight posteriorly, slightly concave
distally; proximal portion 47-53 (51) % of total rostral length; distal portion
acuminate, smooth, glabrous. Spermatheca: (Fig. 7).
DIscussIoN-Hamaba bahamensis is easily distinguished from its only
known congener, H. dispersa, by the characters listed in the diagnosis. The
species is presently known from the Bahamas (Egg, Eleuthera, Grand
Bahama, and Harbor Islands), from Florida (Ft. Myers and Key West), and
from the island of Cozumel off the east coast of the Yucatan Peninsula of
Sleeper (1957: 39) considered Blatchley's (1916: 247) Paragoges mini-
mus (which he transferred to Hamaba) and H. bahamensis to be different
species. The holotype of H. minima and other Florida Hamaba cannot be
distinguished from examined specimens of H. bahamensis from the Bahamas
on the basis of the characters Sleeper presented: base of elytra "rather
strongly," or "not to very feebly" bisinuate, and vestiture "rather hispid,
many scales semi-erect to erect," or "prostrate or nearly so." Florida and
Bahamas specimens differ slightly in several other features, however. Most
Florida specimens are slightly larger (length 1.48-1.68 mm) than specimens
from the Bahamas (length 1.15-1.59 mm), and have the distal portion of the
rostrum somewhat shorter (males 30-33%, females 39% of total rostral
length) and stouter than Bahamas specimens (males 39-47%, females
47-50% of total rostral length). Florida specimens also have smaller por-
tions of the elytra covered by dark integument and dark scales. In addition,
Florida males have the apex of the median lobe rounded, whereas Bahamas
males have the extreme apex emarginate' (Fig. 5). It seems inadvisable,
however, to accord to these 2 forms, distinguished by minor differences, the
status of full species.
13-V, 1 1 9 (USNM); Eleuthera Island, 9-15 VII, 4 3, 1 9 (MCZ), 4 3,
5 9 (USNM); Grand Bahama Island, Pine Ridge, 13-V-1953, 3 S (AMNH);
Harbor Island, 8-VII, 1 3, 1 9 (USNM). MEXICO: Cozumel Island, San

Clark: Hamaba Casey 221

Miguel, 2-IV-1960, 1 S (USNM). UNITED STATES: Florida, Ft. Myers,
11-III-1911 (PURC); Key West, 1 S (USNM), IV, 1 2 9 (MCZ), 1-IV-
1903, 2 S (USNM).

Hamaba disperse Casey
(Fig. 3, 4, 6, 9)
Hamaba disperse Casey 1910: 131-2. Holotype, 8, labelled "Eleuthera Ba-
hamas/July 9-15, Wickham," ," "Casey/bequest/1925," "TYPE
USNM/36741," "dispersa/Csy." (USNM). Leng 1920: 320. Klima 1934:
3. Blackwelder 1947: 842. Sleeper 1957: 39.
DIAGNOSIs-Rostrum (Fig. 3, 4), in profile, strongly, evenly arcuate from
base to tip, distal portion not tapered; whitish, recumbent scales on midbasal
and humeral portions of elytra narrow, sparse, leaving ferruginous integu-
ment broadly exposed; median lobe of male genitalia (Fig. 6) narrowed
MALE-Length: 1.63 mm. Width: 0.83 mm. Head: eyes separated by ca.
0.33 x eye length, hind margin of eye not raised. Rostrum (Fig. 3) : short,
1.09 x pronotum length; in profile, dorsal and ventral margins strongly,
evenly arcuate from base to tip; in dorsal view, sides subparallel from base
to tip; proximal portion 58% of total rostral length; lateral and dorsolateral
sulci obsolete, represented by shallow punctures, dorsal midline narrow,
whitish scales laterally, distal portion glabrous. Elytra: recumbent scales on
interspaces sparse, very narrow, acuminate; slightly raised scales attenuate
to bluntly rounded apically; prominent posteromedian and subapical, in-
distinctly zigzag-shaped, transverse bands present, as well as irregular,
dorsomedian, longitudinal vittae which extend from posteromedian band to
base on portions of interspaces 2-5, and less distinct transverse bands which
extend laterally from longitudinal vittae to costal margins just behind
FEMALE-Length: 1.59-1.78 (1.69) mm. Width: 0.85-0.91 (0.88) mm.
Rostrum (Fig. 4): slightly longer than that of 8, 1.27-1.34 (1.31) x
pronotum length; proximal portion 56-58 (57)% of total rostral length.
Spermatheca: (Fig. 9).
DIscUssIoN-Hamaba disperse is distinguished from H. bahamensis by
the characters listed in the diagnosis. The species is presently known only
from the Bahamas (Eleuthera Island and Rum Cay).
Island, 9-15-VII 1 8 (USNM); Rum Cay, near Port Nelson, 16-III-1953,
1 9 (AMNH).


Thanks are extended to the curators of the museums and institutions cited
above for arranging for loan of specimens studied, and to R. S. Berger, M. H.
Bass, and 2 anonymous reviewers for critique of the manuscript.

BLACKWELDER, R. E. 1947. Checklist of the coleopterous insects of Mexico,
Central America, the West Indies, and South America. Part 5. Bull.
United States Natl. Mus. 185: 756-925.

Florida Entomologist 64 (2)

BLATCHLEY, W. S. 1916. In: W. S. Blatchley and C. W. Leng, Rhynchophora
or weevils of north eastern America. The Nature Publishing Company,
Indianapolis. 628 p.
1930. Blatchleyana. The Nature Publishing Company, Indianapolis.
77 p.
CASEY, T. L. 1910. On some new species of Balanini, Tychiini and related
tribes. Canadian Ent. 42: 114-44.
CHAMPION, G. C. 1910. Biologia Centrali-Americana (Curculionidae:
Calandrinae; appendix to Curculioninae). 4(7) : 79-212.
CLARK, W. E. 1980. Revision of the weevil genus Neotylopterus Hustache
(Coleoptera: Curculionidae). Ann. Ent. Soc. America 73: 216-30.
1980a. Revision of the weevil genus Lignyodes Dejean (Coleoptera:
Curculionidae) in America north of Mexico. Trans. America Ent. Soc.
106: 273-326.
1980b. Additions to the neotropical weevil genus Rosella White-
head (Coleoptera: Curculionidae) : R. triophori (Gyllenhal). n. comb.,
transferred from Lignyodes Dejean, and R. arcuata, n. sp. Coleopt.
Bull. 34(3) : 299-304.
AND D. M. ANDERSON. In Press. The genus Chionanthobius Pierce
(Coleoptera: Curculionidae): descriptions of a new species on
Forestiera (Oleaceae), and of the larva and pupa of C. schwarzi
Pierce. Proc. Ent. Soc. Washington.
D. R. WHITEHEAD, AND R. E. WARNER. 1977. Classification of the
weevil subfamily Tychiinae, with a new genus and species, new com-
binations, and new synonymy in Lignyodini (Coleoptera: Cur-
culionidae). Coleopt. Bull. 31(1) : 1-18.
KISSINGER, D. G. 1964. Curculionidae of America north of Mexico. A key to
the genera. Taxon. Publ., South Lancaster, MA. 143 p.
KLIMA, A. 1934. Curculionidae: subfamily Tychiinae. In: S. Schenkling
(ed.), Coleopterorum Catalogus. ... 29 (pars 138). Junk, The Hague.
61 p.
LENG, C. W. 1920. Catalogue of the Coleoptera of America, north of Mexico.
Mount Vernon, N.Y. 470 p.
SLEEPER, E. L. 1957. Notes on the Curculionoidea . Ohio J. Sci. 57(1):


Department of Entomology
University of Massachusetts
Amherst, MA 01003 USA

Following egg laying in rose hips, females of Rhagoletis basiola (Osten
Sacken) were found to deposit on the fruit surface a marking pheromone
which deters repeated oviposition attempts by other females into that fruit.
Rhagoletis basiola did not recognize the marking pheromone of R. pomonella

'Diptera: Tephritidae.
2Paper No. 2350, Massachusetts Agricultural Experiment Station.


June, 1981

Averill & Prokopy: Rhagoletis Pheromone

(Walsh), which likewise infests rose hips, while R. pomonella exhibited
partial recognition of R. basiola's marking pheromone.

Se encontr6 que, despues de ovipositar en los frutos del rosal, las hembras
de Rhagoletis basiola (Osten Sacken) depositan un feromono sobre la super-
ficie del fruto que detiene otras hembras de ovipositar en el mismo fruto.
Rhagoletis basiola no reconoci6 el feromono marcador de R. pomonella
(Walsh), dicha especie igualmente ataca los frutos del rosal, mientras R.
pomonella mostr6 reconocimiento parcial del feromono marcador de R.

Several species of plant parasitic insects, including 12 species of
Tephritidae, have been found to deposit marking pheromones following
oviposition which deter repeated oviposition (Prokopy 1981). The ecological
significance of these pheromones is hypothesized to be mediation of uni-
formity in egg dispersion, and consequently insurance of ample food re-
sources for each developing larva (Prokopy 1981).
Here, we present evidence that, following egg laying into a host fruit
(rose hips), a Rhagoletis basiola (Osten Sacken) female drags its ovipositor
on the fruit surface and thereby deposits a trail of marking pheromone
which deters subsequent oviposition attempts in that fruit. In light of the
recent discovery that the apple maggot, Rhagoletis pomonella (Walsh), as
well as R. basiola infest Rosa rugosa Thunb. rose hips in New England
(Prokopy and Berlocher 1980), we also compared the response of R. basiola
and R. pomonella females to one another's marking pheromones.

All assay females originated from field collected pupae from various
localities in Massachusetts. Studies were conducted under controlled labora-
tory conditions of 22-260 C and 55-65% RH. Maintenance and preparation of
assay females prior to testing, as well as the experimental procedure, were
modeled after Prokopy et al. (1976), with the exception that wild rose hips
were used in assays. These hips were either ca. 12 mm diam hips (tenta-
tively identified as fruits of the pasture rose, Rosa rubinigosa L.) or 17-22
mm hips (tentatively identified as fruits of Rosa virginiana Mill.). Same-
sized fruits were used within a replicate. Test fruit were clean control hips,
hips with oviposition punctures only, or hips marked with oviposition punc-
tures plus ovipositor dragging. Owing to the tendency of a R. basiola fe-
male to drag its ovipositor ca. 20 sees longer following egg laying than a
R. pomonella female (mean = 66 vs. 46 sec), a correspondingly greater num-
ber of ovipositor dragging was collected for R. pomonella. Also, to equalize
the amount of trail substance on each assay fruit, small (12 mm) fruits
were marked with fewer dragging than large (17-22 mm) fruits. Thus,
each large hip was marked by 15-17 R. basiola females and each small hip
by 7-9 R. basiola females. In cross-recognition assays, large and small hips
were marked as above by R. basiola females, while large hips were marked
by 18-23 R. pomonella females and small hips by 12 R. pomonella females.


Florida, Entomologist 64(2)


Behavioral observations of 15 R. basiola females revealed that only a
single egg was laid per oviposition bout, a phenomenon previously reported
for this tephritid species by Balduf (1959). In 14 of 15 cases in which egg
laying occurred, ovipositor dragging on the fruit surface followed invariably.
In all 9 cases in which boring attempts were not followed by ovipositor
dragging, no egg had been deposited. The hardness of the fruit hypanthium
is the most probable explanation for this high frequency of unsuccessful
oviposition, a possibility also suggested by Balduf (1959).
The data in Table 1 provide clear evidence that R. basiola females de-
posit an oviposition deterring substance during ovipositor dragging following
egg laying. Only 20% of arriving females attempted to bore in fruit which
had oviposition punctures plus ovipositor dragging, while 91% attempted
boring into clean control fruits. There was no evidence of deterrence re-
sulting from the presence of eggs or boring punctures: 88% of arriving fe-
males bored into fruits where other females had been allowed to oviposit
but not to drag the ovipositor.
Table 2 shows results of the assay of R. basiola and pomonella female
responses to each other's oviposition deterring pheromones. Rhagoletis
basiola females were not deterred from boring into fruits marked with R.
pomonella pheromone as evidenced by the fact that the same proportion of
R. basiola females attempted boring into control fruits as into fruits heavily
marked by R. pomonella females. However, the data indicate that R.
pomonella females are substantially deterred from boring into fruits marked
by R. basiola. Response of R. pomonella to R. basiola marking pheromone has
been confirmed by electrophysiological assays (E. S. Bowdan, Zoology De-
partment, University of Massachusetts, unpublished data).
A similar situation was demonstrated by Prokopy et al. (1976) wherein


No. arrivals
No. 2 culminating
arrivals in boring % boring*
Treatment on fruit attempt attempts

Control 47 43 91(a)
Oviposition punctures
only** 25 22 88 (a)
Oviposition punctures
plus ovipositor
draggingst 41 8 20 (b)

*Values followed by different letters are significantly different at the 1% level by X2 analysis.
**Two sizes of rose hip were used: 12 mm diam hips with 7-9 oviposition punctures, or
17-22 mm hips with 15-17 punctures.
t12 mm and 17-22 mm hips were oviposited in and dragged on by 7-9, and 15-17 ?, respec-


June, 1981

Averill & Prokopy: Rhagoletis Pheromone

TABLE 2. PROPORTION OF Rhagoletis basiola AND R. pomonella FEMALES

No. arrivals
No. 9 culminating
arrivals in boring % boring*
Treatment on fruit attempts attempts

R. basiola
Control 42 39 93 (a)
by R. basiola 44 13 30 (b)
by R. pomonella 36 33 92 (a)
R. pomonella
Control 47 44 94 (a)
by R. basiola 56 25 45 (b)
by R. pomonella 44 11 25 (c)

*Values in the same column followed by different letters are significantly different at the
5% level by X2 analysis. Comparisons are only within species.
**R. basiola marked fruits were either: large (17-22 mm diam) hips dragged on by 13-16
R. basiola 9 or small (12 mm diam) hips dragged on by 7-9 R. basiola $.
+R. pomonella marked fruits were either: large (17-22 mm diam) hips dragged on by 18-23
R. pomonella 9 or small (12 mm diam) hips dragged on by 12 R. pomonella Y.

R. pomonella showed recognition of the oviposition marking pheromones of
R. mendax Curran and R. cornivora Bush, while the latter 2 species recog-
nized the pheromone of R. pomonella only partially or not at all. They addi-
tionally found that R. pomonella, R. fausta (Osten Sacken) and R. cingulata
(Lowe) do not recognize one another's pheromones.
This latter finding, as well as our finding of incomplete cross-recognition
of marking pheromones between R. pomonella and R. basiola, are of ecological
interest. Within the last hundred years, populations of R. pomonella, R.
fausta and R. cingulata have shifted from their native hosts onto the intro-
duced host Montmorency sour cherry, and populations of R. pomonella and
R. basiola have shifted from their native hosts onto the introduced host
Rosa rugosa (Prokopy and Belocher 1980). In both cherry and rose hip
hosts, simultaneous colonization by all of the respective Rhagoletis species
has been observed. In such situations, any advantage of intra-specific regula-
tion of larval spacing through oviposition deterring pheromone deposition
may be reduced by effects of unregulated inter-specific competition.


We thank Ms. P. J. Spatcher, Ms. C. M. Bardinelli and Mr. K. Spatcher


Florida Entomologist 64 (2)

for assistance. This project was supported by Massachusetts Agricultural
Experiment Station Project 380.

BALDUF, W. V. 1959. Obligatory and facultative insects in rose hips, their
recognition and bionomics. Illinois Biol. Monogr. No. 26.
PROKOPY, R. J. 1981. Epideictic pheromones that influence spacing patterns
of phytophagous insects. In: D. A. Nordland, R. L. Jones and W. J.
Lewis (ed.) Semiochemicals: their role in pest control. Wiley and Sons.
(In press).
-- AND S. H. BERLOCHER. 1980. Establishment of Rhagoletis pomonella
(Diptera: Tephritidae) on rose hips in southern New England. Ca-
nadian Ent. 112: 1319-20.
W. H. REISSIG AND V. MOERICKE. 1976. Marking pheromones de-
terring repeated oviposition in Rhagoletis flies. Ent. Exp. Appl. 20:

Delta States Research Center, USDA, SEA-AR
Southern Weed Science Laboratory, Biological Weed Control Research
P.O. Box 225, Stoneville, MS 38776
in cooperation with the
Department of Entomology
Mississippi State University
Mississippi State, MS 39762 USA

The first instars of Kermes iselini and K. prinus new species are described
and illustrated. Keys are provided to separate closely related Nearctic species
based on first instar morphology.

Se described y se presentan dibujos de los primeros instares de Kermes
iselini y K. prinus, nuevas species. Se presentan claves basadas en la
morfologia de los primeros instares para separar las species neArcticas
cercanas en relaci6n.

The Kermesidae or gall-like scale insects are found throughout the
Nearctic Region on oaks (Quercus) and chinquapins (Castanopsis) in the
family Fagaceae. There are 2 genera in the family Kermesidae: Kermes

'Research Associate, Department of Entomology, Mississippi State University, Mississippi
State, MS 39762.
Professor of Systematic Entomology, Department of Entomology, Virginia Polytechnic
Institute and State University, Blacksburg, VA 24061.


June, 1981

Baer & Kosztarab: New Coccoids on Oak

Boitard (1828) and Olliffiella Cockerell (1896). The genus Olliffiella consists
of 2 leaf-gall forming species found in the southwestern United States and
Mexico. Twenty-nine species have been included in the genus Kermes.
Ferris (1955) stated that, "The genus Kermes presents to the student an
almost impossible situation" when referring to the external morphology and
classification of the group. The difficulty stems from the fact that early de-
scriptions were based on the external appearance of mature, sclerotized adult
females. The characters (i.e. size, shape and color patterns) used in the de-
scriptions are subject to excessive intraspecific variations and are unreliable
for species determination.
Hamon et al. (1976) and Bullington (1978) determined that post-
reproductive females do not provide much information even after they are
slide mounted. However, the morphology of the 1st instars provides useful
characters for identification of Kermes species. Kuwana (1931) and Bala-
chowsky (1950) separated 10 species in the Palearctic Region based on this
stage. Baer (1980) described the 1st instar of Kermes kosztarabi Baer and
presented a key to separate the 1st instars of 4 species in the K. galliformis
Riley group. With the 2 species described here, 11 species or species com-
plexes comprising 3 species groups can now be distinguished based on 1st
instars in the Nearctic Region.
In concurrent studies, Bullington (1978) and Baer (1978) found that
certain Kermes species can only be separated at certain stages of develop-
ment. Kermes iselini and K. prinus new species can be separated morpholog-
ically in the 1st instar.
Each description is based on the holotype and 9 paratype 1st instars. The
holotype measurements are followed in parentheses by the average and the
range of the 9 paratype specimens. Measurements are given in microns. The
descriptions and illustrations include only half the dorsum and venter and
should be doubled for a complete view of each surface. A key to separate
closely related species follows each description. Sexual differentiation was
not apparent among the 1st instars.

Kermes iselini Baer and Kosztarab, NEW SPECIES
(Fig. 1 a-t)

TYPE MATERIAL STUDIED: On leaves, petioles and buds of Quercus
mohriana Buckl., Mohrs oak. Holotype and 74 paratype 1st instars, 15 mi.
N. of El Paso Gap, Otero Co., New Mexico, 15-VI-1976, coll. W. A. Iselin.
TYPE DEPOSITION: Holotype 1st instar with 4 paratypes on same slide and
20 other paratypes on 4 slides deposited in the United States National Mu-
seum of Natural History, Washington, D.C. Other paratype 1st instars de-
posited in the following collections: 20 specimens on 4 slides in the British
Museum (Natural History), London; 20 specimens on 4 slides in the Uni-
versity of California at Davis; and 10 specimens on 2 slides in the Virginia
Polytechnic Institute and State University, Blacksburg (VPI). Additional
dry material curated at VPI.
ETYMOLOGY: Kermes iselini was named for Mr. William A. Iselin, Agri-
cultural Biologist for the State of California, whose diligent collecting efforts
in southwestern United States resulted in the discovery of this species.


Florida Entomologist 64 (2)

Fig. la-t. Kermes iselini Baer and Kosztarab, n. sp. first instar; a, dorso-
ventral view of body. Dorsum: b,c, marginal setae; d,e, submedial setae f,
simple pore; g, derm; h, intersegmental membrane; i, anal lobes. Venter: j,
antenna; k, trilocular pore; 1, derm; m, enlargement of tibia, tarsus and claw
of metathoracic leg; n, sensory pore; o, quinquelocular pore laterad of
spiracle; p, bilocular pore; q, submedial trilocular pore; r-t, body setae.

BODY (Fig. la). Oblong, widest at mesothorax, tapering posteriorly,


June, 1981

Baer & Kosztarab: New Coccoids on Oak

antennae, legs and anal lobes well developed, with long apical setae. Length
423 (483,408-544), width 181 (191,166-227).
DORSUM: Marginal setae (Fig. Ib). Conical, 33-35, 16 (16,12-23) long, 8
(8,5-9) wide. Each abdominal segment with 2 pr.
Marginal setae on anal lobe (Fig. Ic). Slightly longer and thicker than
other marginal setae, 18 (18,16-24) long, 9 (8,8-9) wide.
Submedial setae on head and thorax (Fig. Id). Conical, 4 pr. on head and
thorax, similar to marginal setae but different from submedial setae on
abdomen, 14 (14,12-16) long, 9 (7,6-9) wide.
Submedial setae on abdomen (Fig. le). Thin, elongate, 7 pr., 4 (5,4-6)
long, ca. 1 wide.
Simple pores (Fig. If). Composed of a membranous duct opening into a
sclerotized ring, arranged in 1 longitudinal row along submargin of body,
several irregularly spaced pores in submedial area, 13-15 pores.
Derm (Fig. Ig). Composed of overlapping plate-like areas.
Intersegmental membrane (Fig. Ih). Apparent, due to type of derm.
Anal lobes (Fig. la,i). Sclerotized. Conical seta on inner margin of each
lobe, 16 (21,16-23) long, another at apex, 19 (19,16-21) long; apical seta
106 (124,91-196) long.
VENTER: Antennae (Fig. lj). Six segmented. Total length 84 (90,80-102).
Scape 12 (13,12-14) long, 26 (26,23-33) wide. Segments II to VI: 16 (17,
16-21), 16 (18,16-19), 9 (11,9-12), 12 (11,9-14), 19 (21,16-26) long, respec-
tively. One trilocular pore at each scape base (Fig. 1k). Segment I with 3
slender setae, II with 2 slender setae and 1 sensory pore, III with 1 slender
seta, IV with 1 fleshy seta, V with 4 slender setae and 1 fleshy seta and VI
with 4-6 slender setae and 3 fleshy setae.
Clypeolabral shield. Sclerotized, length 74 (74,74-95), width 58 (59,51-70).
Stylet loop extending to the 1st abdominal segment.
Labium. Sclerotized, triangular, composed of 3 segments. Basal segment
small, with 2 pr. of setae, the middle segment with 1 pr. of setae and apical
segment with 4 pr. of setae. Length 65 (73,58-86), width 46 (49,44-53).
Derm (Fig. 11). Membranous.
Legs (Fig. 1m). Numerous, slender elongate setae on each segment. Four
sensory pores on each trochanter (Fig. In). Sensory pore on proximal lateral
margin of each tarsus. Tarsal digitules long, enlarged at apices; claw dig-
itules longer than claw, enlarged at apices; claw notched apically.

Segments Lengths
and Claw Prothoracic Mesothoracic Metathoracic

Coxa 19 (21,19-23) 19 (20,16-23) 19 (20,16-23)
Trochanter 19 (25,19-30) 23 (26,23-30) 23 (27,23-35)
Femur 44 (49,44-53) 44 (47,37-53) 46 (50,44-58)
Tibia 33 (30,23-35) 33 (33,26-35) 30 (33,26-37)
Tarsus 49 (42,35-49) 39 (45,39-46) 42 (48,42-53)
Claw 9 (14,9-19) 9 (15,9-21) 14 (15,9-21)

Entire leg 173 (182,160-200) 167 (189,162-202) 174 (193,167-218)


Florida Entomologist 64 (2)

Spiracles and associated pores (Fig. lo). Each anterior and posterior
spiracle with 1 quinquelocular pore; diam. ca. 4. Anterior spiracle 14 (15,
12-19) long, 5 (5,5-7) wide, atrium ca. 2 wide. Posterior spiracle 13 (13,
12-17) long, 5 (5,5-6) wide, atrium ca. 2 wide.
Bilocular pores (Fig. Ip). Located near bases of submarginal, setae; 7-10
along each margin.
Submedial pores (Fig. lq). Four pr. in 1 submedial longitudinal row on
abdomen, 4 pr. on head and thorax, mostly triloculars but occasionally sev-
eral quinqueloculars in each row, diam. ca. 4.
Body setae (Fig. 1 r-t). In 3 longitudinal rows on abdomen and partly
on thorax: medial 6 (7,5-11) long, ca. 1 wide (Fig. Ir); submedial 7 (7,5-9)
long, ca. 1 wide (Fig. Is); submarginal 6 (7,5-9) long, 2 (1,1-2) wide (Fig.
It). One long elongate seta near base of each coxa and 3 setae submedial
to antennal base. One pair of setae similar morphologically to dorsal mar-
ginal setae on anterior margin of head.
Anal lobes and anal ring (Fig. la,i). Lobes sclerotized, with 1 submedial
seta on each lobe, 10 (10,9-12) long. Anal ring oval, 14 (12,9-19) long, 16 (18,
16-21) wide, with 16-22 translucent cells, 6 anal ring setae 16 (19,14-23)
long; 1 pr. of thick setae slightly anterior to anal ring 7 (9,7-14) long.
ADDITIONAL MATERIAL STUDIED: On Quercus grisea Liebm., gray oak, Tex
Canyon, about 18 mi. N. from Rt. 80, Arizona, 20-VI-1976, colls. M. and
M. Kosztarab, VPI; On Q. rugosa N6e, near Portal, 1.5 mi. up on S. Fork Rd.,
Arizona, 19-VI-1976, colls. M. Kosztarab, D. Hanna, and I. Storks, VPI;
Coronado National Forest, New Mexico, 20-VI-1976, colls. M. and M. Kosz-
tarab, VPI.
AFFINITIES: Based on 1st instar morphology, Kermes iselini n. sp. is
closely allied with K. pubescens Bogue (1898). Kermes pubescens is desig-
nated to represent a group containing the 2 species. Both species have conical
dorsal marginal setae (with 2 pr. on each abdominal segment) and tri-
locular submedial pores on the venter. They can be separated with the fol-
lowing key:
Key to the First Instars of Kermes pubescens Group
1, Dorsal submedial setae conical on abdomen; 2 or 3 quinquelocular
pores laterad of each anterior spiracle ..._- Kermes pubescens Bogue
1'. Dorsal submedial setae elongate on abdomen; 1 quinquelocular
pore laterad of each anterior spiracle ........- -------....-......-.
-...-- ------------------------- Kermes iselini Baer and Kosztarab, n. sp.
CHARACTERISTIC INJURY: Old sclerotized females of this southwestern
species were found attached to the leaf petiole and midvein, causing distor-
tion. Damage was similar to that caused by K. pubescens found in central
and eastern United States (McConnell and Davidson 1959).

Kermes prinus Baer and Kosztarab, NEW SPECIES
(Fig. 2 a-v)

TYPE MATERIAL STUDIED: On twigs near bud regions of Quercus prinus
L., chestnut oak. Holotype and 63 paratype 1st instars, at overlook, near
tennis courts of Pipestem Lodge, Pipestem State Park, West Virginia, 27-VI-
1978, colls. M., M. and E. Kosztarab and D. and M. Suhayda.
TYPE DEPOSITION: Holotype and 3 paratypes on same slide and 12 other


June, 1981

Baer & Kosztarab: New Coccoids on Oak

Fig. 2a-v. Kermes prinus Baer and Kosztarab, n. sp. first instar; a, dorso-
ventral view of body. Dorsum: b,c, marginal setae; d,e, submedial setae; f,
simple pore; g, derm; h, intersegmental membrane; i, anal lobes. Venter; j,
antenna; k, trilocular pore; 1, derm; m, enlargement of tibia, tarsus and claw
of metathoracic leg; n, sensory pore; o, quinquelocular pore laterad of
spiracle; p, bilocular pore; q, submedial quinquelocular pore; r-v, body setae.

paratypes on 3 slides deposited in the United States National Museum of
Natural History, Washington, D.C. Other paratype 1st instars deposited in
the following collections: 16 specimens on 5 slides in the British Museum


Florida Entomologist 64 (2)

(Natural History), London; 16 specimens on 4 slides in the University of
California at Davis; and 12 specimens on 4 slides in the Virginia Polytechnic
Institute and State University, Blacksburg.
ETYMOLOGY: Kermes prinus was named after Quercus prinus, which was
its first and only recorded host.

BODY (Fig. 2a). Oblong, widest at mesothorax, tapering posteriorly, an-
tennae, legs and anal lobes well developed, with long apical setae. Length 550
(539,513-563), width 250 (244,225-263).
DORSUM: Marginal setae (Fig. 2b). Elongate, thick, 21-24, 26 (25,23-33)
long, 4 (4,3-4) wide. Each abdominal segment with 1 pr.
Marginal setae on anal lobe (Fig. 2c). Longer and thicker than other
marginal setae, 37 (29,28-39) long, 5 (4,4-5) wide.
Submedial setae on head and thorax (Fig. 2d). Elongate, thick similar to
abdominal submedial setae, 4 pr., 25 (24,15-26) long, 3 (3,3-4) wide.
Submedial setae on abdomen (Fig. 2e). Elongate, thick, 7 pr., 22 (24,
17-26) long, 2 (3,3-4) wide.
Simple pores (Fig. 2f). Composed of a membranous duct opening into a
sclerotized ring, arranged in 2 longitudinal rows, 17-23 pores.
Derm (Fig. 2g). Simple, membranous.
Intersegmental membrane (Fig. 2h). Apparent, due to type of derm.
Anal lobes (Fig. 2a,i.). Sclerotized. One thick seta on inner margin of
each lobe, 32 (28,27-33) long; another at apex, 36 (35,32-37) long; apical
seta 166 (142,125-168) long.
VENTER: Antennae (Fig. 2j). Six-segmented. Total length 101 (97,88
106). Scape 13 (13,12-15) long, 37 (28,27-37) wide. Segments II to VI: 15
(16,15-19), 25 (25,23-25), 9 (9,9-12), 13 (13,12-14), 26 (27,24-28) long, re-
spectively. One trilocular pore at each scape base (Fig. 2k). Segment I with
3 slender setae, II with 2 slender setae and one sensory pore, III with 1
slender seta, IV with 1 fleshly seta, V with 4 slender setae and 1 fleshy seta
and VI with 4-5 slender setae and 3 fleshy setae.
Clypeolabral shield. Sclerotized, length 101 (98,97-110), width 71 (70,
68-72). Stylet loop extending to 1st or 2nd abdominal segments.
Labium. Sclerotized, triangular, composed of 3 distinct segments. Basal
segment with 2 pr. of setae, the middle segment with 1 pr. of setae and the
apical segment with 4 pr. of setae. Length 81 (79-78-83), width 58 (60,
Derm (Fig. 21). Membranous.
Legs (Fig. 2m). Numerous, slender elongate setae on each segment. Four
sensory pores on each trochanter (Fig. 2n). Sensory pore on proximal lateral
margin of each tarsus. Tarsal digitules long, enlarged at apices; claw dig-
itules longer than claw, enlarged at apices; claw notched apically.
Spiracles and associated pores (Fig. 20). Each anterior spiracle with 3
quinquelocular pores, rarely 1 of these a trilocular pore, each posterior
spiracle with 1 quinquelocular pore; diam. ca. 4. Anterior spiracle 15 (14,
13-16) long, 5 (5,4-6) wide, atrium ca. 2 wide. Posterior spiracle 14 (13,
13-17) long, 5 (5,5-8) wide, atrium ca. 2 wide.
Bilocular pores (Fig. 2p). Located near bases of submarginal setae. As-
sociated with anterior setae; 4-7 along each margin.


June, 1981

Baer & Kosztarab: New Coccoids on Oak

Segments Lengths
and Claw Prothoracic Mesothoracic Metathoracic

Coxa 26 (25,24-30) 26 (23,23-27) 26 (23,22-27)
Trochanter 26 (25,24-29) 30 (26,24-30) 31 (27,26-32)
Femur 70 (68,64-70) 68 (64,58-70) 70 (67,64-71)
Tibia 37 (35,30-37) 35 (33,32-39) 39 (36,34-40)
Tarsus 52 (49,47-53) 54 (54,53-58) 57 (57,56-59)
Claw 19 (19,19-20) 23 (20,19-23) 23 (20,19-23)

Entire leg 230 (219,210-237) 236 (225,215-245) 243 (239,231-253)

Submedial pores (Fig. 2q). Five pr. in 1 submedial longitudinal row on
abdomen, 4 pr. on head and thorax, usually quinquelocular but sometimes
several triloculars in each row, diam. ca. 4.
Body setae (Fig. 2r-v). In 4 longitudinal rows on abdomen and partly on
thorax: medial 15 (14,9-19) long, ca. 1 wide (Fig. 2r) ; submedial 8 (8,7-10)
long, ca. 1 wide (Fig. 2s); submarginal 9 (8,7-10) long, ca. 1 wide (Fig. 2t);
marginal 15 (18,12-24) long, ca. 2 wide (Fig. 2u). Posteriormost seta in
marginal row thicker than rest in row 20 (20,18-26) long (Fig. 2v). One
very elongate seta near base of each coxa and 3 setae submedial of an-
tennal base. One pair of setae similar morphologically to dorsal marginal
setae on anterior margin of head.
Anal lobes and anal ring (Fig. 2a,i). Lobes sclerotized, with 1 submedial
seta on each lobe 21 (17,13-23) long. Anal ring oval 12 (13,9-14) long, 20
(21,15-22) wide, with 14-20 translucent cells, 6 anal ring setae 19 (18,16-22)
long; 1 pr. of thick setae slightly anterior to anal ring 19 (17,16-23) long.
AFFINITIES: Kermes prinus n. sp. is closely allied to K. concinnulus
Cockerell (in Bogue 1898), K. cockerelli Ehrhorn (1898), K. shastensis
Ehrhorn (1911) and K. rimarum Ferris (1955). Kermes concinnulus, the
oldest described species, is designated to represent a group which includes
all the above listed species. All species in this group have elongate or parallel-
sided dorsal marginal setae and quinquelocular submedial pores on the venter.
They can be separated with the following key:
Key to the First Instars of Kermes concinnulus Group
1. All or some dorsal submedial setae on head, pro- and meso-
thorax thickened and similar to dorsal marginal setae -....--..2.....__ 2
1'. Dorsal submedial setae on head, pro- and mesothorax slender,
small and different from dorsal marginal setae -- ......- .. ......--_ 4
2(1). Parallel-sided and blunt dorsal,marginal setae; 1 7-locular
pore laterad of each anterior spiracle -.-_. Kermes shastensis Ehrhorn
2'. Elongate and pointed dorsal marginal setae; quinquelocular
pore(s) laterad of each anterior spiracle --. ................. 3
3(2). Dorsal submedial setae only on head, pro- and mesothorax
similar to dorsal marginal setae; 1 quinquelocular pore laterad
of each anterior spiracle .................. __ Kermes cockerelli Ehrhorn
3'. Dorsal submedial setae along entire body, similar to dorsal


Florida Entomologist 64 (2)

marginal setae; 3 quinquelocular pores laterad of each anterior
spiracle ----.--- Kermes prinus Baer and Kosztarab, n. sp.
4(1). Four to 7 bilocular pores along each ventral body margin;
terminal antennal segment 21-25 long Kermes concinnulus Cockerell
4'. Eight to 10 bilocular pores along each ventral body margin;
terminal antennal segment 26-30 long -......__ Kermes rimarum Ferris
Information on the first, second and third instars and the adult females
will be presented in the forthcoming revision of the Kermesidae in the
Nearctic Region to be published as a Research Division Bulletin of the
Virginia Polytechnic Institute and State University, Blacksburg.

The authors express appreciation to Ms. M. Rhoades for assistance with
slide mounting of specimens and editing and to Ms. S. B. Walker and Ms.
D. R. Tabb, Southern Weed Science Laboratory for typing this manuscript.

BAER, R. G. 1978. A morphological and systematic study of the first and
second instars of the Kermesidae in the Nearctic Region. (Homoptera:
Coccoidea). Ph.D. Dissertation, Va. Polytech. Inst. & State Univ.,
Blacksburg, 192 p.
1980. A new species of gall-like coccid from southeastern United
States (Homoptera: Coccoidea: Kermesidae). J. Georgia Ent. Soc.
15: 20-5.
BALACHOWSKY, A. 1950. Les Kermes (Hom. Coccoidea) des chines en Europe
et dans le bassin M6diterraneen. Proc. Internat. Cong. Ent. (Stock-
holm) 8: 739-54.
BOGUE, E. E. 1898. Two new species of Kermes from Kansas. Canadian Ent.
30: 172.
BOITARD, P. 1828. Les gallinsectes. In Manuel d'Entomologie ou Histoire
Naturelle des Insectes. 2: 169-72.
BULLINGTON, S. W. 1978. A revision of the tribe Kermesini (Homoptera:
Kermesidae) in North America. M.S. Thesis, Virginia Polytech. Inst.
and State Univ., Blacksburg, 212 p.
COCKERELL, T. D. A. 1896. A gall-making coccid in America. Science 4: 299-
EHRHORN, E. M. 1898. New Coccidae from California. Ent. News. 9: 185-6.
1911. New Coccidae with notes on other species. Canadian Ent. 43:
FERRIS, G. F. 1955. Atlas of the scale insects of North America, The fam-
ilies Aclerdidae, Asterolecaniidae, Conchaspididae, Dactylopiidae, and
Lacciferidae. Vol. 7, Stanford Univ. Press. 233 p.
HAMON, A. B., P. L. LAMBDIN, AND M. KOSZTARAB. 1976. Life history and
morphology of Kermes kingi in Virginia. (Homoptera: Coccoidea:
Kermesidae). Virginia Polytech. Inst. and State Univ. Res. Div. Bull.
No. 111: 1-31.
KUWANA, I. 1931. The genus Kermes of Japan. Min. Agr. & For., Dept. Agr.
(Tokyo, Japan), Sci. Bull. No. 2: 15-9.
MCCONNELL, H. S., AND J. A. DAVIDSON. 1959. Observations on the life his-
tory and morphology of Kermes pubescens Bogue (Homoptera: Coc-
coidea: Dactylopiidae). Ann. Ent. Soc. America 52: 463-8.


June, 1981

Porter: New Certonotus


Biology Department, Fordham University
Bronx, NY 10458 USA

Certonotus is cited for the 1st time from temperate South America. The
genus formerly was believed to be endemic to the Australian zoogeographical
region and to some nearby islands. Certonotus invictus n. sp. and C. um-
brarum n. sp. inhabit moist, cool Nothofagus woods of south Chile and ad-
jacent Argentina. These species differ in many characters but may be dis-
tinguished most easily because C. invictus has slender, largely fulvous legs,
while the legs of C. umbrarum are black with white markings and much more

Certonotus Kriechbaumer (Hymenoptera: Ichneumonidae), un g6nero
Australiano por primera vez citado de America del sur. Certonotus, cuyas
species estin concentradas en la Regi6n Australiana y que parecia estar
restringida a dicha zona biogeogrifica, es reportado ahora de America del
Sur. Certonotus invictus n. sp. y C. umbrarum n. sp. habitan los himedos y
frescos bosques de Nothofagus del sur de Chile y parties adyacentes de la
Argentina. Muchas caracteristicas separan estas species, las que pueden
distinguirse mis facilmente, porque en C. invictus las patas son relativamente
tenues y de color mayormente anaranjado, mientras que C. umbrarum tiene
las patas mis robustas y negras con disefios blancos.

Certonotus, a genus of large labenine ichneumonids evidently parasitic
on wood-boring coleopterous larvae, so far has been known only from Aus-
tralia, Tasmania, New Guinea, New Zealand, and some associated islands.
However, in 1964 the Chilean entomologist, Sr. Luis Pefia G., generously
gave me a single Certonotus captured in south Chile and obviously different
from any of the Australian species. Since then, I have collected in South
America every year, have returned several times to Chile, and have studied
most of the important collections of Latin American Ichneumonidae, all
without finding another Certonotus. Finally, in 1980, I received 2 more
specimens obtained by Ing. Sergio Schajovskoi at Pucari in Neuqu6n Prov-
ince of Argentina. One of these individuals is conspecific with Pefia's Chilean
Certonotus, but the other represents a trenchantly distinct species.
I describe the new South American Certonotus, offer a revised definition
of the genus, and discuss the biogeographic implications of its presence in
southern South America.

1Contribution No. 493, Bureau of Entomology, Division of Plant Industry, Florida Depart-
ment of Agriculture and Consumer Services, Gainesville, FL 32602 USA.
'Research Associate, Florida State Collection of Arthropods, Florida Department of Agri-
culture and Consumer Services, Gainesville.


Florida Entomologist 64 (2)

Certonotus belongs to the Tribe Labenini of the Subfamily Labiinae (as
defined by Townes 1969: 194-6). Within these categories, it may be recog-
nized by the following combination of characters: occipital carina broadly
interrupted or absent dorsally; upper edge of pronotum with a deep, more or
less V-shaped emargination; mesoscutum anteriorly rounded-off and without
a transverse ridge, covered throughout by sharp transverse wrinkles; fore
tibia apico-externally either with a broad and only moderately projecting
triangular tooth or without a tooth; nervellus intercepted near middle; 8th
tergite of female produced apicolaterally into a broad subligulate or sub-
triangular lobe; ovipositor sheath 1.1 as long as fore wing.
Certonotus much resembles the Neotropic genus Apechoneura Kriech-
baumer but in Apechoneura the upper edge of the pronotum is much more
shallowly notched, the front face of the mesoscutum is vertical and bears a
salient transverse ridge, the areolet averages larger, the discoidella often
arises from the mediella, and the ovipositor may be longer.
The Neotropic, Nearctic, and Australian Labena Cresson seems intimately
related to both the foregoing genera and may represent the ancestral stock
from which each was derived. Labena differs from Certonotus in its dorsally
complete occipital carina, roundedly emarginate upper pronotal edge, lack of
transverse mesoscutal wrinkling, presence on the fore tibia of a markedly
outstanding thorn-like, apico-external spine, and apico-laterally unmodified
female 8th tergite. Except for the upper pronotal contour, all these features
likewise distinguish Labena from Apechoneura.
In addition to the "traditional" diagnostic characters discussed above,
many Labena show other noteworthy differences from Certonotus as repre-
sented in South America. For example, I have in hand specimens of L.
grallator (Say), the type species, as well as representatives of undescribed
species from Panama, Ecuador, and Bolivia. These differ collectively from
Certonotus invictus Porter and C. umbrarum Porter by the following fea-
tures: eye strongly emarginate just above antennal socket (slightly emargi-
nate in Certonotus); parapsidal furrows and notauli absent or obsolete
(traceable in Certonotus) ; upper end of prepectal carina slopes anteriad and
terminates at or near front margin of mesopleuron (prepectal carina of
Certonotus is nearly vertical and ends far from front margin of meso-
pleuron); pleural carina absent between base of propodeum and spiracle
(strong in Certonotus) ; 1st gastric sternite ends at or near level of spiracles
(well beyond spiracles in Certonotus); female mid tibia medially inflated
and twisted, basally narrowed (unmodified in Certonotus); female hind
coxa without a strong, triangular projection below toward base; metacarpel-
lan hamuli mostly separated by more to much more than the length of each
hamulus (uniformly close-set in Certonotus); 2nd recurrent approximately
vertical on lower 1/2, outcurved on upper 1/2 (in Certonotus deeply and
triangularly angled proximad with vertex of angle near mid-height of vein).
Certonotus, Labena, and Apechoneura have scores of species each and
include numerous undescribed forms. As comprehensive revisions of these
conspicuous but rare genera become possible, it will undoubtedly be necessary
to redefine them as well as to split off some new taxa.


June, 1981

Porter: New Certonotus

Key to South American Species of Certonotus
1. Hind femur fulvous, 5.7-6.0 as long as deep; hind tibia and tarsus
fulvous with black or dusky markings; hypostomal carina mod-
erately raised; scutellum at apical 0.6 with a strong, arcuate
carina that partly delimits a semicircular area on the sclerite's
apical face; mesopleural suture coarsely foveolate; area-basalis
complete; 3rd pleural area 1.0 as high as long; 1st gastric tergite
3.5-5.0 as long as wide at apex, with dorsal, dorso-lateral, and
ventro-lateral carinae more or less well defined; 2nd recurrent at
apical 1/3 of areolet ...-----------------------.- 1. Certonotus invictus n. sp.
1'. Hind femur black, 3.8 as long as deep; hind tibia and tarsus black
with white markings; hypostomal carina very strongly raised be-
tween juncture with occipital carina and base of mandible; scutel-
lum with a transverse carina at apical 0.6; mesopleural suture
scarcely foveolate; area-basalis open behind; 3rd pleural area 2.0
as high as long; 1st gastric tergite 2.7 as long as wide at apex,
with numerous longitudinal wrinkles between base and spiracle
but without well defined dorso, dorso-lateral, or ventrolateral
carinae; 2nd recurrent at basal 1/4 of areolet -----------
-- .............-------------------------------- 2. Certonotus um brarum n. sp.

1. Certonotus invictus Porter, NEW SPECIES
(Fig. 1, 3, 5, 6)

FEMALE: Color: antenna black with flagellum mostly pale yellow on seg-
ments beyond 21, except for some dusky on last 4 or 5 segments; palpi pale
brown; head black with yellow on most of clypeus, a dull yellow line on facial
orbit, a large brighter yellow spot on vertical orbit, and a broad brownish
yellow band on subdorsal 1/4 of hind orbit; mesosoma black with dull yellow
on basal 1/2 of tegula; gaster shining black, in places with some obscure
brown staining; fore and mid legs pale fulvous with a little dusky on base of
coxa, on apex of trochantellus and base of femur, and at least slightly on
last tarsomere, especially apicad; hind leg similar to fore leg but with bright
fulvous on coxa, trochanters, and femur, tibia more dully fulvous on basal
1/3 but otherwise mostly black except for obscure brown staining toward
apex, and tarsus rather dull fulvous with brownish apicad on last segment;
wings uniformly medium brown with a subtle yellowish sheen, stigma black
with ventral margin narrowly yellowish brown.
Length of fore wing: 18.8-22.3 mm. Hypostomal carina: between base of
mandible and occipital carina forming a moderately strong flange that be-
comes higher apicad and which at its juncture with occipital carina is 0.05
as high as length of 1st flagellomere. Clypeus: with disc mostly smooth,
weakly concave mesad and a little swollen on lateral 1/3, lateral margins
not reflexed. Face: silky-shining with well developed trans-biased wrinkling
on upper 1/2 and with punctures mostly small and sparse; 0.69-0.76 as high
as wide. Temple: in dorsal view moderately receding and slightly rounded
off. Mesoscutum: throughout with very long and outstanding, overlapping
but not really dense setae. Scutellum: at apical 0.6 on median 1/3-1/2 with
a strongly raised arcuate carina that extends ventrad and partially delimits
a semicircular area on the steeply declivous hind face of the sclerite.


Florida Entomologist 64(2)

Fig. 1-4. 1) Certonotus invictus, 9 Paratype. Lateral view of propodeum
and 1st gastric tergite; 2) Certonotus umbrarum, 9 Holotype. Lateral view
of propodeum and 1st gastric tergite; 3) C'ertonotus invictus, 9 Paratype.
Areolet; 4) Certonotus umbrarum, 9 Holotype. Areolet.
Prepectal carina: very broadly and sharply angled rearward on upper
mesosternum and lower mesopleuron, above which it curves forward almost
as strongly and finally bends back again to terminate on about upper 0.4-
0.3 of mesopleuron. Mesopleural suture: coarsely foveolate. Mesepimeron:
practically throughout with very long, overlapping setae. Metapleuron:


~c~--~--------- .
: `::

June, 1981

Porter: New Certonotus

lower division on disc with numerous, very widely spaced small punctures
that emit unusually long and overlapping setae. Submetapleural carina: a
high, percurrent, foveolate flange that becomes gradually a little higher
basad. Mesosternum: with median longitudinal sulcus strong and foveolate.
Propodeum: area-basalis strongly defined, approximately rectangular; areola
0.6-0.8 as wide at apex as long, varying from almost parallel-sided through-
out to considerably narrowed between costulae and base; 1st lateral area
with very long, outstanding and overlapping setae; 2nd lateral area with
some very long, outstanding setae; 2nd pleural area 0.70-0.75 as high as long,
3rd abscissa of basal trans-carina becoming obsolete dorsally; 3rd pleural
area 1.0 as high as long, profoundly concave, scarcely wrinkled. First gastric
tergite: 3.5-5.0 as long as wide at apex; dorsal carinae traceable about 2/3
the distance from base to spiracle; dorso-lateral carina strong from base to
spiracle; ventro-lateral carina sharp throughout. Second gastric tergite:
thyridium 1.0-2.0 as long as wide. Ovipositor: sheathed portion 1.1 as long
as fore wing. Fore femur: 5.0-5.4 as long as deep. Fore tibia: on apex above
with a stout, triangular tooth which is best seen in front view; apical spur
with an almost percurrent translucent ventral flange that averages 1/3 as
high as sclerotized portion of spur and is vertically truncate subapically.
Hind femur: 5.7-6.0 as long as deep. Wing venation: 2nd recurrent inserted
at apical 0.3 of areolet; ramellus absent; intercubitella 0.53-0.58 as long as
radiella; costella strong but translucent pale brown; 16-17 metacarpellan
MALE: Unknown.
VARIATION: The paratype shows only the following differences from the
holotype: no yellow on facial orbit, yellow on base of tegula dull and in-
conspicuous, length of fore wing 18.8 mm (versus 22.3 mm), face 0.69
(versus 0.76) as high as wide, prepectal carina reaching upper 0.4 (versus
0.3) of mesopleuron, areola 0.8 as wide at apex as long (versus 0.6) and
considerably narrowed based of costulae (versus almost parallel-sided
throughout), 1st gastric tergite 5.0 as long as wide at apex (versus 3.5),
and thyridium 2.0 as long as wide (versus 1.0).
TYPE MATERIAL: Holotype 9 : CHILE, Arauco Province, Pichinahuel, Cord.
Nahuelbuta, 31-XII-1958, G. Barria. Paratype 9: ARGENTINA, Neuqu6n
Province, Pucara nr. San Martin de los Andes, 640 m, X-1971, S. Schajov-
skoi. Holotype in Florida State Collection of Arthropods, Gainesville. Para-
type in collection of Fundaci6n Miguel Lillo, Universidad Nacional de Tucu-
man, Argentina:
RELATIONSHIP: Certonotus invictus seems but distantly related to the sym-
patric C. umbrarum. It differs from umbrarum by the key characters and
also in such features as its smooth (instead of trans-striate) clypeal disc,
unreflexed lateral margins of clypeus, sparsely punctate face with con-
siderable wrinkling dorsad (face densely punctate but scarcely wrinkled in
umbrarum), densely (instead of sparsely) Setose mesoscutum, biangulate (in-
stead of bisinuate) prepectal carina, moderately (instead of very strongly)
raised submetapleural carina, deep and foveolate mesosternal sulcus, and
absence of a ramellus.
FIELD NOTES: For a week in December 1969 I collected at the paratype
locality of C. invictus, on Lago Lacar between the Chilean border and San
Martin de los Andes, Argentina. The area bestrides an ecotone annectant


Florida Entomologist 64 (2)

between the Valdivian and Deciduous Forest Districts of the Subantarctic
Life Zone (as these communities are defined by Cabrera and Willink 1973:
97-101). It is dominated by Nothofagus forests. On the pre-Andean slopes
the deciduous N. pumilio (Poepp. & Endl.) is prominent. At low altitude
near Lago Lacar (where most collecting was done) there are dense, lofty
forests of the evergreen Nothofagus dombeyi (Mirb.) Bl., and of such
deciduous species as N. antarctica (Forest. f.) Oerst., N. obliqua (Mirb.)
Bl., and N. procera (Poepp. & Endl.) Oerst. Chusquea, a large bambusacean
cane, and the hygrophile myrtacean tree, Myrceugenella apiculata (DC.)
Kaus., also characterize this distinctive flora.
Pucara has a cool, wet, almost ceaselessly windy climate. The annual
temperature averages about 100 C, and yearly rainfall oscillates between
1000 and 2000 mm. There is essentially no summer, although occasional
maxima of 22-250 C may be registered between November and February,
but winters are mild, with average minima not falling much below freezing.
Ichneumonids abound in the humid woodlands of PucarA, where they fly
from September to May, with an extraordinarily pronounced maximum in
December. During my fieldwork at Pucara, I found no Certonotus but did
collect about 1500 specimens belonging to Calliephialtes, Eruga, Polysphincta,
Coccygomimus, Netelia, Clasis, Macrogrotea, Bilira, Surculus, Anacis, Doch-
midium, Dotocryptus, Trachyshyrus, Alophophion, Scallama, Thymebatis,
and a host of other genera.
SPECIFIC NAME: From the Latin adjective invictus, "invincible", used here in
reference to the imposing size of this splendid wasp and in honor of the great
Chilean nation.

2. Certonotus umbrarum Porter, NEW SPECIES
(Fig. 2, 4, 7, 8)

FEMALE: Color: antenna black with flagellomeres 20-30 white and last
flagellomere (31) white on basal 1/2 and brownish black apically; palpi
white with some dusky staining; head and mesosoma shining black; gastric
tergites shining black with white markings as follows: 1st tergite with an
ovoid area near apex dorso-laterally and with a narrow subapical band on
median 0.6 of dorsum; 2nd tergite with a broad, somewhat irregular band on
all but base and apex of lateral margin and with a narrow subapical band
on median 0.8 of dorsum; 3rd-5th tergites similar to 2nd but with both
lateral and dorso-subapical white bands successively a little narrower and
the subapical white bands tending to reach farther laterad; 6th tergite like
proceeding but with lateral and subapical white bands substantially nar-
rower and the subapical band several times briefly interrupted; 7th tergite
with a subapico-lateral white band that is interrupted on whole dorsum of
tergite; 8th tergite with a small white area medially on apex; legs with coxae
shining black; fore trochanter shining black with whitish narrowly on apex;
fore trochantellus narrowly black above and mostly white below; mid and
hind trochanter and trochantellus almost wholly shining black; fore femur
shining black with a small white blotch on apex dorso-posteriorly and with
a very broad, basally narrowed and finally effaced, dorsal and dorso-anterior
white band; all tibial spurs white; fore tibia white; fore tarsus white with
black briefly on apex of 1st and 2nd segments, a little more broadly on apex


June, 1981

Porter: New Certonotus


Fig. 5-8. 5) Certonotus invictus, 9 Paratype. Lateral view of hind leg,
showing color pattern and general proportions; 6) Certonotus invictus, 9
Paratype. Spur of fore tibia, showing translucent ventral lamella; 7)
Certonotus umbrarum, 9 Holotype. Spur of fore tibia, showing translucent
ventral lamella; 8) Certonotus umbrarum, 9 Holotype. Lateral view of hind
leg, showing color pattern and general proportions.

Florida Entomologist 64(2)

of 3rd segment, on almost all of 4th segment, which becomes dully whitish
below, and on apical 1/2 of 5th segment; mid femur shining black with a
large, basally much tapered, anterio-dorsal white area occupying most of
distal 0.6 except near apex; mid tibia white with black broadly below on
apical.0.7 and narrowly throughout on apex; mid tarsus white with black
markings much as those described for front tarsus except that those on
apices of segments 1 and 2 are broader and that on 3 occupies more than
apical 1/2 of segment; hind femur shining black; hind tibia black with white
on most of basal 0.3 above and 0.4 below; and hind tarsus about as described
for mid tarsus; wings hyaline with stigma black.
Length of fore wing: 12 mm. Hypostomal carina: between base of man-
dible and occipital carina unusually elevated and becoming a very high
flange apicad, at juncture with occipital carina 0.23 as high as length of 1st
flagellomere. Clypeus: disc largely trans-striate, gently concave almost
throughout, lateral margins conspicuously reflexed. Face: shining with
abundant, moderately small, sharp, uniformly distributed punctures that
mostly are separated by at least 1.0 their diameters; 1.1 as high as wide.
Temple: in dorsal view moderately rounded and a little receding. Meso-
scutum: with setae mostly short and widely spaced. Scutellum: at apical 0.6
on median 2/5 with a strong transverse carina that sets off the steeply
declivous apical face. Prepectal carina: broadly bisinuate but not angled on
upper mesosternum and lower mesopleuron. Mesopleural suture: smooth,
foveolate only in extreme lower hind corner. Mesepimeron: practically
glabrous, except for some long setae on upper 1/3. Metapleuron: lower
division on disc polished, the upper 1/2 with numerous moderately small
punctures that emit long and overlapping setae, the lower 1/2 almost im-
punctate and glabrous. Submetapleural carina: very strongly raised, bluntly
triangular, highest at about mid point, margined by gross foveae. Meso-
sternum: median sulcus fine, weakly impressed, scarcely foveolate. Pro-
podeum: area-basalis not defined, open posteriorly; combined area-basalis
and areola 1.5 as wide at apex as long and considerably narrowed basad of
costula; 1st lateral area with rather few, very long, well spaced setae; 2nd
lateral area practically glabrous; 2nd pleural area 1.3 as high as long; 3rd
abscissa of basal trans-carina strong throughout; 3rd pleural area 2.0 as
high as long, gently concave, strongly wrinkled except dorsad. First gastric
tergite: 2.7 as long as wide at apex; dorsal carinae obsolete; dorso-lateral
carina not distinguishable among the many irregularly longitudinal wrinkles
extending between base and spiracle; ventro-lateral carina obsolete basad of
spiracle and absent between spiracle and apex. Second gastric tergite:
thyridium large, 0.5 as long as wide. Ovipositor: sheathed portion 1.1 as long
as fore wing. Fore femur: 3.8 as long as deep. Fore tibia: with a broad,
vestigial, appressed tooth on dorsal margin apically; tibial spur more deli-
cate than in C. invictus, the translucent ventral flange about 1/4 as high as
sclerotized part and terminated by an oblique truncation so that the apex of
the spur appears asymmetrically bifid. Hind femur: 3.7 as long as deep.
Wing venation: 2nd recurrent inserted at basal 0.4 of areolet; ramellus well
,developed; intercubitella 0.80 as long as radiella; costella strongly sclerotized,
dark brown; 10 metacarpellan hamuli.
MALE: Unknown.
TYPE MATERIAL: Holotype 9: ARGENTINA, Neuqu6n Province, Pucari nr.


June, 1981

Porter: New Certonotus

San Martin de los Andes, 640 m, 30-1-1974, S. Schajovskoi. Holotype in col-
leciton of Fundaci6n Miguel Lillo, Universidad Nacional de TucumAn, Argen-
RELATIONSHIPS: This is a distinctive species with no close relatives among
either the South American or the Australian Certonotus.
FIELD NOTES: Certonotus umbrarum inhabits the same temperate Nothofagus
forests already discussed under my treatment of C. invictus.
SPECIFIC NAME: From the Latin noun umbra, "shadow" or "shade", used
here in the genitive plural and chosen with regard to this species' dank,
storm-swept habitat.

The South American species here assigned to Certonotus fit well in this
primarily Australian genus, as diagnosed by modern descriptive taxonomists
(Townes 1969: 196-8). Assuming that such phenotypic resemblance connotes
also close evolutionary relationship, Certonotus becomes another member of
the temperate-adapted biota that is shared in impressive numbers and
diversity between the Antarctic Beech Forests of austral South America and
those of south Australia (Kuschel 1960: 540-50). The genus perhaps origi-
nated in some part of the Southern Hemisphere. Then it could have dis-
persed between South America and Australia via Antarctica, which had a
benign climate during the Cretaceous and early Tertiary and which also at
that time seems to have offered direct land connections between the 2 south-
ern continents.
Interestingly, 2 genera which show affinity to Certonotus have similar
distributions. Labena, which belongs to the same tribe and is surely a close
relative, ranges through North, Middle, and South America (including south
Chile) as well as Australia. Labium, a more distantly associated genus in-
cluded by Townes in the same subfamily, has many Australian species but
also a single representative in southeast Brasil.
Their apparent extreme rarity constitutes another remarkable attribute
of the South American Certonotus. Sr. Luis Pefia has collected unceasingly
in Chile for more than 3 decades and Ing. Sergio Schajovskoi spent more
than 10 years at Pucara, where he was active with nets and Malaise Traps
and entertained a host of visiting entomologists. It seems incredible that such
protracted and assiduous labor should have yielded only 3 specimens of these


The fieldwork and museum study in South America, which allowed me to
complete this project, were done under a Faculty Fellowship from Fordham
University and under United States National Science Foundation Grants
GB-6925 and DEB-75-22426. I am indebted to Sr. Luis Pefia G. of Santiago,
Chile and to Ing. Sergio Schajovskoi of Pucara, Argentina for the donation
or loan of crucial specimens and for much additional help.

CABRERA, A. L., AND A. WILLINK. 1973. Biogeografia de America Latina.
Organizaci6n de los Estados Americanos, Serie de Biologia. Mono-
grafia No. 13: 1-117.


Florida Entomologist 64 (2)

KUSCHEL, G. 1960. Terrestrial zoology in southern Chile. Proc. Roy. Soc.
(London), B, 152: 540-50.
TOWNES, H. K. 1969. The genera of Ichneumonidae, Pt. 1, Mem. American
Ent. Inst. 11: 1-300.


Institute of Ecology
University of Georgia
Athens, GA 30602 USA

A new species, Pseudachorutes (Pseudachorutes) quadrisetosus Snider,
is described from Florida. It bears similarities to Pseudachorutes (Pseu-
dachorutes) curtus Christiansen and Bellinger (1980), but can easily be
separated on the number of tenacular teeth, dental setae and mucro shape.
The type locality is Lee County, Florida, collected from Nymphaea odorata

Una nueva especie, Pseudachorutes (Pseudachorutes) quadrosetosus
Snider se describe desde Florida. Tiene similaridades a P. curtus Christiansen
y Bellinger (1980), pero se puede distinguir de aquel facilmente por el
inmero de dientes tenaculares, setas dentales, y la forma del mucro. La
localidad del tipo es el Condado de Lee, Florida, coleccionado de Nymphaea

Recently I completed the identification of 28 species of Collembola col-
lected by Dr. E. S. Del Fosse from semi-aquatic and aquatic plants. His
sample sites were located in Broward, Collier, Glades and Lee Counties,
Florida. While sorting through a sample from Lee County, an unusual species
of Pseudachorutes was detected. Careful analysis showed it to be undescribed.
The purpose of this report is to present that new species.

Pseudachorutes (Pseudachorutes) quadrisetosus Snider, NEW SPECIES
COLOR DESCRIPTION: Dark blue throughout body and appendages; with very
narrow intersegmental lines, small pale dots scattered over body segments,
and pigment on tarsal complex and mucronal bases.
MORPHOLOGICAL DESCRIPTION: Eyes 8 + 8. Postantennal organ with 5 lobes
arranged in a circle (Fig. 1), approximately 2/3 diameter of nearest ocellus.
Antennal segments in ratio of 1:2.1:1.8:2 (Fig. 2) ; segment IV with trilobed
subapical bulb, 2 apical falciform (Fig. 3) and 4 blunt setae (Fig. 4); seg-
ments III and IV with long outstanding setae, longest setae occur on segment
III and are 3/4 as long as segment IV. Mouthparts form a pronounced

'Mailing Address: Department of Zoology, Michigan State University, East Lansing, MI
48824 USA.


June, 1981

Snider: New Pseudachorutes


buccal cone; mandible with 4 teeth, sometimes with minute subapical 5th
tooth (Fig. 5), maxilla crochet-like (Fig. 6). Tibiotarsus without differ-
entiated tenent hair; pretarsus with anterior and posterior setula; unguis
with distinct inner tooth, curving lanceolate; unguiculus absent (Fig. 7).
Ventral tube with 1 + 1 setae. Corpus of tenaculum without setae, rami
with 3 + 3 teeth (Fig. 8). Dens 2 times as long as mucro, with 4 dorsal
setae. Mucro tapering, apically reflexed, lamellae narrow, evident distally
(Fig. 9). Dorsal anal lobe with 5 dorsal setae and 1 short median seta (Fig.
10). Body clothed sparsely with curved setae, becoming longer on posterior,
longest setae 3/4 longer than inner margin of unguis. Length: 1.5 mm.
DIAGNOSIS: Pseudachorutes quadrisetosus keys out nearest to Pseudachorutes
algidensis Carpenter in Massoud (1967). It differs from that species in color,
length, mucronal shape, and claw characteristics. Pseudachorutes algidensis
is described from New Zealand and appears unrelated to P. quadrisetosus.
In Christiansen and Bellinger (1980) P. quadrisetosus keys out nearest to
Pseudachorutes curtus Christiansen and Bellinger. While the 2 species share
similar morphology with respect to postantennal organ shape, maxilla, urguis
and absence of tenent hairs, they are dissimilar in the following respects:

P. A. O. with 5 lobes
Mandible with inner teeth
similar in shape





P. A. O. with 6 lobes
Inner teeth dissimilar


"K J


\ *.

1 I .

/ I ^



t 10.

Fig. 1-10. Pseudachorutes (Pseudachorutes) quadrisetosus n. sp. 1) Post-
antennal organ and anterior ocelli; 2) antenna segments; 3) falciform seta
of ANT IV; 4) tip of ANT IV; 5) mandible; 6) maxilla; 7) meta unguis;
8) tenaculum; 9) mucro-dens; 10) anal papilla.

Florida Entomologist 64 (2)

Ventral tube with 1 + 1 setae With 5 + 5 or 6 + 6 setae
Tenaculum 3 + 3 teeth With 4 + 4 teeth
Dens with 4 setae With 6 setae
Mucro with narrow lamellae With large, broad lamellae
Long posterior setae acuminate Setae truncate or weakly clavate
In addition, on the 4th antennal segment, P. curtus has a ventral "file"
which is lacking in P. quadrisetosus.
TYPES: Holotype (male) and 1 paratype on slides; 1 additional partype in
alcohol. Holotype and paratypes are deposited in the Entomology Museum,
Michigan State University. All specimens were collected in Florida, Lee
County, junction of SR 82 and SR 82a, in a pond on Nymphaea odorata Ait.,
13-III-1978, E. S. Del Fosse, collector.

Special thanks are offered to Dr. D. A. Crossley, Jr. for laboratory facil-
ities and manuscript preparation, and also to the Department of Entomology,
University of Georgia at Athens.

CHRISTIANSEN, K. A., AND P. F. BELLINGER. 1980. The Collembola of North
America North of the Rio Grande. Part I, Families Poduridae and
Hypogastruridae. Grinnell College, Grinnell, Iowa. 386 p.
MASSOUD, Z. 1967. Monographie des Neanuridae. Biol. Amer. Austr. 3: 7-


Section of Systematic & Evolutionary Biology
University of Connecticut
Storrs, CT 06268 USA

The distribution and geographic variation of Ozophora atropicta Barber
is analyzed. A discussion is given of the likelihood of the present distribution
being due to dispersal, vicariance or a combination of the above. Ozophora
heydoni Barber and Ashlock is considered a junior synonym of atropicta.
Ozophora parapicta is described as a new species from Central and South
America. The 5th instar nymph of the latter is briefly described and com-
pared with that of Ozophora concava Distant.
Charts and graphs of intraspecific variation in atropicta are included as
is a dorsal view illustration of atropicta and figures of the male genitalia of
both species.

'This work was supported by a grant from the National Science Foundation.


June, 1981

Slater & Hassey: Ozophora atropicta

Se analizan la distribuci6n y la variaci6n geogrAfica de Ozophora
atropicta Barber. Se discute la posibilidad que la distribuci6n actual se debe
o a la dispersion o a la disyunci6n o a una combinaci6n de las dos factors.
Ozophora heydoni Barber Y Ashlock se consider un sin6nimo junior de O.
atropicta. Ozophora parapicta se describe como una especie nueva de America
Central y America del Sur. El 5 instar de la ninfa del anterior se describe
brevamente y se lo compare con la de 0. concava Distant.
Se incluyen cartas y graficos de la variaci6n intraspecifica en 0. atropicta,
un dibujo de la vista dorsal de 0. atropicta, y dibujos de las genitalias de los
machos de ambas species.

The status of Ozophora populations in the West Indies is a difficult tax-
onomic problem. The present paper deals with one aspect of this-the status
of the 2 nominal species atropicta and heydoni. This taxonomic situation is
complicated by a nomenclatoral problem involving the type series. In addi-
tion a new related species from Central and South America is described.
In the following discussion, we synonymize heydoni with atropicta and
discuss in detail the distribution and variation of the latter.
Ozophora atropicta was described by Barber (1939) from Puerto Rico,
Hispaniola, and Andros Island in the Bahamas. It was later reported by
Ramos (1946) from Mona I., by Barber (1954) from Cuba, by Scudder
(1958) front the Caymans and by Barber and Ashlock (1960) from many
islands in the Bahamas and from the Turks and Caicos (the Slater 1964
Catalogue is in error regarding the distribution given in this latter paper).
Ozophora heydoni was originally described by Barber and Ashlock (1960)
from New Providence Island, the Abaco Cays, Great Abaco I., and Andros
Island (all in the Bahamas). It apparently has not been reported subse-
quently in the literature.
Unfortunately the type series of atropicta is mixed. The holotype from
Puerto Rico and 3 of the paratypes from the Dominican Republic (San
Lorenzo (1), Sanchez (2)) are conspecific with the holotype (and all of the
rest of the type series) of heydoni. The remaining paratypes of atropicta
represent at least one quite distinct species. Thus, heydoni Barber and Ash-
lock becomes a junior synonym of atropicta Barber. What we discuss in the
present paper as atropicta is, therefore, in the old sense of heydoni.
The status of the remaining material that has previously been called
atropicta is beyond the scope of this paper but we will comment on it briefly.
Material from the Upper Bahamas (Great Abaco) as well as from Florida
represents an undescribed species which is being treated in a separate paper
by R. M. Baranowski and the senior author on the Ozophora of Florida. Ma-
terial from the Turks and Caicos, Fortune I. and Mayaguana is, as noted by
Barber and Ashlock (1960), very distinctly colored and may prove to rep-
resent another undescribed species. Specimens from the Greater Antilles all
appear to represent still an additional species. We have not examined ma-
terial of this complex from the Caymans so cannot comment on Scudder's
record other than to note that pallidofemora Scudder and fuscofemora
Scudder described from the Cayman's are quite distinct from any of the
Ozophora atropicta (=heydoni) as now restricted has a much more ex-
tensive range than has previously been suspected. It not only occurs in the

248 Florida Entomologist 64 (2) June, 1981

2 A


Fig. 1. Ozophora atropicta Barber-Dorsal view.
.,and thus a single species appears to be involved (Fig. 2, 4).-

.. ..

West Indies and Bahamas but in Central sh brown and Souwith thAmericark areas ofwell. (Asthe

hmcolor and relaticove proportions. Despite this the genitalia, which usually are

hemelytra covering much less of the total wing surface than do the pale

Slater & Hassey: Ozophora atropicta

\ 4 -' 5 \
Fig. 2-5. 2) Ozophora atropicta paramere. 3) 0. parapicta n. sp. para-
mere. 4) 0. atropicta sperm reservoir. 5) 0. parapicta n. sp. sperm reservoir.

areas. Specimens from Cuba are also small and often as pale as are those
from the Bahamas, but frequently the posterior pronotal lobe is suffused with
dark coloration and most specimens have more extensive dark hemelytral
markings. Somewhat surprisingly specimens from Hispaniola are appreciably
larger and darker than other specimens from the West Indies (Fig. 6).
Specimens from Puerto Rico resemble those from Hispaniola more closely
in color than they do material examined from Cuba.
Mainland specimens are generally large (Fig. 6) and dark, but there is a
great deal of individual variation. Nevertheless, the difference in appearance
is appreciable and if one compares a large dark Trinidad or Mexican speci-
men with a small pale Bahaman or Cuban one it is difficult to believe that
they represent the same species. Mainland populations also have proportion-
ately longer antennae (Fig. 7, 8).
Bahaman and Cuban specimens have a noticeably longer labium than do


Florida Entomologist 64 (2)







I I I I I I I Ii I I I l I I I I
4.5 5.0 5.5 6.0 6.5
Fig. 6. Comparison of total length of males of various populations of
Ozophora atropicta.1
mainland specimens. In the former the labium exceeds the posterior margin
of the metacoxae and may reach well into abdominal sternum 2. In mainland
populations the labium extends at most only between the metacoxae. Inter-
estingly, Hispaniolan specimens have a relatively shorter labium and thus,
in this respect, are similar to the mainland, rather than other West Indies
The number of fore femoral spines is variable. Some specimens have 3
spines on a fore femur and others have 4. The condition is not always the
same on the 2 femora of the same specimen. While numbers of individuals
available from most areas is small and it is possible that 3-spined and 4-
spined conditions are equally common (see Hispaniola and Trinidad Columns,
Fig. 9) available data suggests otherwise., Specimens from the West Indies
tend to show a significantly higher proportion of individuals that have 3
spines' than do those from the mainland (Fig. 9).
All of the data plus the series of measurements (Table 1, 2) indicates
that West Indies population are, as one might expect, more similar to one
another in most respects than any of them are to a mainland population.

1In Fig 6, 7, 8 "N" numbers are given in Fig. 10 and diagrams show (a) mean (vertical
central line), (b) range (horizontal line), (c) one standard deviation (horizontal box).


June, 1981

Slater & Hassey: Ozophora atropicta







2.5 3.0 3.5 4.0 4.5 5.0
Fig. 7. Comparison of ratio of length of second antennal segment to inter-
ocular space in females of various populations of Ozophora atropicta.

What is unexpected is that the Hispaniolan population is more similar,
in size, color, antennal, and labial length to the mainland populations than it
is to Cuban. Today situations such as this must be considered from alternate
viewpoints. Are the population differences found in atropicta likely to be the
result of a dispersal phenomenon, a vicariant phenomenon, or a combination
of both? We believe the species is a good example of how information can be
used to arrive at a "most probable" hypothesis where a "3 taxon" statement
is not available.
For the present situation to have resulted entirely from dispersal across
a water barrier probably 4 (or 5) dispersal events would have had to have
taken place. Three of these would be minor dispersals (Cuba to the Bahamas;
Hispaniola to Puerto Rico, or the reverse) and 2 major (Mainland to Cuba;
Mainland to Hispaniola). It can, of course, be argued with some credibility
that only a single major dispersal event need have occurred, resulting in the
establishment of an ancestral population on Hispaniola which subsequently
reached Cuba and differentiated there. Given the nature of the winds of great
force and speed that periodically sweep over the islands it can be argued that
dispersal of small insects is a sweepstakes situation with nearly as much
chance for a population to be established on Hispaniola as on Cuba even
though the latter is nearer to a potential mainland source area.

Florida Entomologist 64 (2)







2.5 3.0 3.5 4.0 4.5
Fig. 8. Comparison of ratio of length of third antennal segment to inter-
ocular space in females of various populations of Ozophora atropicta.
With a dispersal theory there seems to be no way to differentiate between
2 alternate possibilities. (1) An original dispersal event that established
a population on Cuba which then differentiated and subsequently dispersed
through the Bahamas. Subsequent to the differentiation on Cuba a 2nd dis-
persal event occurred which established (from the mainland) a population
on Hispaniola which then differentiated slightly and dispersed to Puerto Rico.
This dispersal hypothesis would attribute the greater similarity of His-
paniolan specimens to those on the mainland to its relatively recent arrival
there as compared to the population on Cuba. (2) An alternative dispersal
hypothesis would be that an original dispersal event established a population
on Hispaniola. After this population had, differentiated, a dispersal event
occurred from Hispaniola that established a population on Cuba that further
differentiated and dispersed to the Bahamas. This latter hypothesis would
consider the Hispaniolan population as ancestral on the islands because it is
most similar to those on the mainland. Its plausibility would, of course, be
based upon the progressive restriction of the gene pool as successive prop-
agules dispersed and established new populations each of which possessed
only a portion of the original genetic variability.


June, 1981

Slater & Hassey: Ozophora atropicta


N= 3 19 10 8 74 6 7 10 10 49 72

Fig. 9. Proportion
phora atropicta.

of population showing 3 forefemoral spines in Ozo-

The principal argument against either dispersal option is where atropicta
is not found. It seems difficult to believe that had atropicta been able to at
least reach Hispaniola or Puerto Rico and subsequently disperse to the other
islands, that it should be completely absent from Jamaica and the Lesser
Antilles. Yet this seems to be the case. The absence of atropicta from Jamaica
is particularly important. The island is varied ecologically and has a rich
Ozophora fauna, of which several species also occur on Cuba and Hispaniola.
Furthermore, this island has been collected far more intensively for Ozophora
species than have any of the islands upon which atropicta occurs. Although
we recognize the danger of unequivocally stating that an insect such as this
is absent from a large island such as Jamaica we find it hard to believe that
if it does occur there (especially given its proclivity for coming to lights),
that not a single example would be known. This is true especially since the

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Florida Entomologist 64 (2)


























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Slater & Hassey: Ozophora atropicta

senior author and several colleagues have concentrated upon collecting speci-
mens of Ozophora on several trips to Jamaica and have maintained a light
trap there for an extended period. Its absence from the Lesser Antilles is
less well established but here again the senior author and colleagues have
collected on many of the islands where species of Ozophora are abundant
without taking a single specimen, nor has one appeared in any collection
examined. If atropicta is a species with high dispersal potential its absence
from all of these islands seems extremely difficult to explain.
The vicariance hypothesis takes into account the position of Jamaica
south of the Cayman trench, and thus the likelihood that the island has never
had a land connection with Cuba and Hispaniola. Also, Jamaica appears to
have been completely submerged in the Miocene, whereas western Cuba and
probably parts of Hispaniola were above water. This suggests the plausibility
of an hypothesis that ancestral atropicta populations were present in the
Greater Antilles at the time the latter were part of the Mainland and were
isolated on Hispaniola. Differentiation took place subsequent to the formation
of a water barrier between Hispaniola and Cuba. This vicariance hypothesis
seems the most reasonable and parsimonious hypothesis to us. We do believe
that insects disperse across water barriers and suggest that atropicta has
reached the Bahamas in just that way.
We have discussed the alternate possibilities above in some detail so that
it may serve as a basis for future comparisons with the distribution of other
West Indian species of Ozophora and to emphasize the possibility that the
hitherto dominant idea that the West Indian insect fauna has been the result
of dispersal may only in part be true.
Ozophora atropicta (Fig. 1) may be readily recognized by the pale yellow
or white calloused margins on the anterior pronotal lobe and anterior collar
that contrast strongly with the dark red brown or chocolate brown of the
dorsal surface of the anterior lobe and by the presence (always) of a pair of
conspicuous white or pale yellow diagonal streaks on the scutellum that
contrast strongly with the darker coloration. In some specimens the dark
posterior pronotal rays so characteristic of many species of Ozophora are
completely developed. The head is broad and relatively short with large eyes.
The 1st and 2nd antennal segments are usually entirely pale yellow, the third
segment chocolate brown distally (sometimes as much as the entire distal
half), the 4th segment has the usual conspicuous white annulus. The para-
meres and sperm reservoir are distinctive (Fig. 2, 4).
The only species that atropicta is likely to be confused with is parapicta
n.sp. described below from the mainland of South and Central America.
Despite the widespread distribution and apparent abundance in some
areas the only information concerning the biology of atropicta appears to be
P. D. Ashlock's collection of a single adult under a species of Ficus on Barro
Colorado I. in a population of adults and nymphs of 0. parapicta and
O.concava. Most of the specimens listed below were taken at light.
MATERIAL EXAMINED: BAHAMAS: Eleuthera I.-7, 14 9 Current Cut 26-XI-
1964 ((on ship at light) (D. Dean); 5 9 same Powell Pt. 24-XI-1964. 1 9
Arthurs Town Cat I. VII-VIII-1935 (W. J. Clench). CUBRAS 10 S, 1 9
Varadero (Prov. Matanzas) 1-5 m. 11-IV-1966 (F. Gregor). 1 9 Habana
Marianao 15 m. 20-VII-1966 (F. Gregor). 1 $ Vinales (Prov. Pinar del Rio)
200 m. 26-VII-1966 (F. Gregor). 2 8, 1 9 Los Jazmines (Prov. Pinar del


Florida Entomologist 64(2)

Rio 200 m. 16-17-VI-1966 (F. Gregor). 1 9 same 26-VII-1966. 1 9 Habana-
Almar-Cojimar (Prov. Habana 2-10 m., 26-31-VII-1966 (Jar. Prokop). 1 9
same 1-5-VII-1966. 1 9 same 10-24-VIII-1966. 1 9 San Bias. Sta. Clara 21-
VII-1932 (B. B. Leavitt). 1 9 San Vincente 6-10-VII-1956 (C. & P. Vaurie).
HAITI: 1 9 Ennery nr. 1000 ft 11-IX-1934 (Darlington). DOMMINICAN RE-
PUBLIC: 71 8, 64 9 2 km. N. Cabo Rojo (Pedernales Prov.) 18-19-VI-1976
(R. E. Woodruff). 3 8, 1 9 Altagracia Nisibon 8-VI-1976 (R. E. Woodruff).
3 $, 1 9 La Cumbres (Prov. Santiago) 15-VI-1976 (R. E. Woodruff). 1 9
Santo Domingo 12-VII-1967 (J. C. Schaffner). 1 9 Jarabacoa (Prov. La
Vega) 8-VIII-1967 (J. C. Schaffner), MEXICO: 1 9 1 mi. S. Tamazunchale
7-VII-1966 (P. M. & P. K. Wagner). 1 S, 1 9 ca. 25 mi. N.C. Monte
Nacimiento del Rio Frio Tam. 31-VII-1970 (C. W. O'Brien). 1 8 Coyame,
Lake Catemaco (Vera Cruz) 2-VII-1962 (D. H. Janzen). 1 $, 1 9 Rio
Quezalapan, 2 mi. E. Lago Catemace (Vera Cruz) 21-VI-5-VII-1964 (J. R.
Meyer). 1 9 Lake Catemaco (Vera Cruz) 2-VII-1971 (Clark, Schaffner, Hart,
Murray). 1 $, 1 9, Temascal (Oaxaca) 1-VII-1971 (Clark, Murray, Hart,
Schaffner). HONDURAS: 3 8, 7 9 San Alejo U.F. Co. (Dept. Atlantida) 4-VI-
1964 (R. E. Woodruff). 1 9 Lk. Yojda Agua Azul U.F. Co. (Dept. Cortes)
31-V-1964 (F. S. Blanton, A. B. Broce, R. E. Woodruff). COSTA RICA: 1 $
Turrialba 20-VI-1974 (C. W. & L. O'Brien, Marshall). PANAMA: Canal Zone:
Barro Colorado I. 4 S, 3 9 IV-1940 (Jas Zetek); 2 S same V-24-40; 1 3,
1 9 same IV-1941; 1 9 same VII-1941; 1 8 same IV-1940; 1 9 same 18-28-
IV-1964 (W. D. & S. S. Duckworth); 2 $, 1 9 same 10-17-V-1964; 1 $ same
20-23-V-1964; 1 9 same 1-9-V-1964; 1 $, 1 9 same 24-28-V-1964; 1 9 same
28-IV-1962 (H. Ruckes); 2 9 same 21-22-IV-1962; 1 9 same 25-IV-1962;
3 $ same 5-XI-1973 (H. Wolda) ; 1 3, 1 9 same 25-IV-1972 (D. Engleman);
1 9 same 18-V-1973 (Under Ficus) (P. D. Ashlock). Coco Solo Hospital
(Canal Zone) 1 9 14-V-1975 (D. Engleman); 1 S same 31-VIII-1972; 1 9
same 30-VIII-1972; 1 9 same 7-XI-1972; 1 9 same 24-1-1974; 1 $ same 14-
V-1973 (P. D. Ashlock); 1 S same 16-V-1973; 1 9 same 15-V-1973; 1 $
same 19-V-1973; 2 $ same 7-V-1973 (D. Engleman); 2 S same 19-XII-1972;
1 S same 8-V-1973; 1 $ same 21-V-1976; 4 9 same 19-XII-1972; 1 & same
14-V-1975; 1 $ same 24-1-1974; 1 $ same 17-IX-1972; 1 9 same 11-VII-1973;
1 8 same 30-VIII-1972. 1 & Fort Sherman (CZ) 24-VII-1974 (D. Engle-
man). 1 S Mojina SWp. (CZ) 16-IV-1952 (F. S. Blanton). 1 $ H. Kohle
(CZ) 3-X-1951 (F. S. Blanton). 1 8 Canal Zone 769 AAA IX-24-1952 (F. S.
Blanton). 1 8 Canal Zone 17-IV-1952 (F. S. Blanton). 1 8 Las Cumbres
09006'N 72032'W 9-V-1974 (H. Wolda); 1 $ same 16-V-1974; 2 S same 18-
V-1974; 1 9 same 19-IV-1974. 1 $, 1 9 Rio LasLajas nr. Coronado Bch.
17-IX-1952 (F. S. Blanton); 1 $, 1 9 same 19-IX-1952. 1 8, 5 9 Rio Hato
(Prov. Panama) 9-XI-1952 (F. S. Blanton); 1 $ same Code 8-XI-1952 (F. S.
Blanton); 2 $ same 9-XI-1952. 2 $ Rio Lajas nr. Coronado Bch. 19-IX-1952
(F. S. Blanton). 1 9 Rovira (Prov. Chiriqui) 5-VII-1964, 2500 ft.
(A. Bruce). 2 8 Fort Amador II-1964 (Ch. Keenan). 1 $ nr. LaCanarca 8-
VIII-1952 (F. S. Blanton). 1 8 Trinidad River 2-V-1911 (August Busck).
VENEZUELA: 1 9 Barinus-Reserve Forestal Ticoporo 280 m. 26-29-II-1968
(F. Fernandez & E. Osuna). TRINIDAD: 1 $, 1 9 St. Augustine 14-VI-1973
(Baranowski, O'Rourke, Picchi, Slater). 49 8, 70 9 Simla, Arima-Blanchis-
seuse Rd. 600 ft. 20-VII-1975 (J. Price) ; 1 $ same 12-VII-1975; 1 3, 4 9
same 14-VII-1975; 1 $ same 11-VII-1975; 1 9 same 1-VIII-1975. 1 9 Simla,


June, 1981

Slater & Hassey: Ozophora atropicta

Arima Valley 4-II-1965 (J. A. Slater & N. T. Davis). 1 & Arima Valley; 1 9
same 800-1200 ft. 27-II-1966 (J. G. Rosen). BRAZIL: 1 3, 19 Corupa (Hans
Humbolt) S. Cat. XI-1944. 1 9 Espiritu Santo; Conceicao da Barra X-1972
(M. Alvarenga). 1 $ Piracicaba (S.P.) 2-XX-1965 (C. A. Triplehorn). 1 9
Guanabara, Repressa Rio Grand VII-VIII-1972 (M. Alvarenga). In Amer-
ican Museum Natural History, National Museum of Natural History (Wash.,
D.C.), U. Venezuela, U. California, Florida State Coll. of Arthropods, Museu
Brno, Texas A & M Univer., G. G. E. Scudder, P. D. Ashlock, D. Engleman,
R. M. Baranowski, and J. A. Slater collections.

Ozophora parapicta Slater and Hassey, NEW SPECIES
(Fig. 3, 5)
Similar in form and color to 0. atropicta but more slender. Head and
anterior pronotal lobe red brown, lateral pronotal margins and anterior
collar a strongly contrasting pale yellow. Posterior pronotal lobe yellow with
weakly developed darker rays including meson. Scutellum dark red brown,
apex white; smooth calloused divergent elevated areas slightly lighter red
brown than ground color but never a strongly contrasting white or pale
yellow. Hemelytra chiefly testaceous with conventionally darker corial areas.
Pale spot at inner angle of corium large and elliptical. Membrane fumose
with pale veins and apex. Pleural and ventral surfaces reddish brown, paler
on abdomen. Legs uniformly pale yellow. Antennal segments I, II and prox-
imal one-third to one-fourth of III testaceous; distal portion of segment III
chocolate brown; segment IV with a broad basal white annulus, distal half
dark chocolate brown. Dorsal surface pruinose but lacking conspicuous up-
standing hairs.
Head relatively short and broad, non-declivent, head length 0.902, width
0.98, interocular space 0.40. Pronotum with complete but shallow transverse
impression, calli surface with numerous punctures, posterior pronotal margin
concave, pronotal length 1.10, width 1.58. Scutellum slightly excavated near
center, scutellar length 1.02, width 0.90. Hemelytra with lateral corial mar-
gins conventionally concave, claval commissure length 0.75, apex clavus-apex
corium distance 1.30, apex corium-apex membrane distance 0.95. Metathoracic
scent gland auricle straight, rather elongate not curving posteriorly. Fore-
femora slender with 4 or 5 small sharp ventral spines. Labium exceeding
mesocoxae, labial segments I 0.88, II 0.85, III 0.60, IV 0.38 long respectively.
Antennae elongate, conventionally terete, antennal segments I 0.65, II 1.92,
III 1.58, IV 1.58 long respectively. Total length 5.90.
Paramere with shaft broad tapering to an acute blade, inner projection
elongate tooth-like, rather strongly hooked; inner "tooth" extremely broad
and short, flange-like (Fig. 4). Sperm reservoir with cup nearly circular,
wings short, thick and broad (Fig. 5). Vesica with 5 coils.
HOLOTYPE: & PANAMA: Barro Colorado Isl. 18-28.IV.64 (W. D. and S. S.
Duckworth). In National Museum of Natural History no. 73799.
PARATYPES: PANAMA: 1 & same as holotype 28-30.IV.1964; 1 9 same 24-28.
V.1964; 1 9 Maje Station 909N 78047W V.17.1975 (light) (Engleman,
Ramirez); CANAL ZONE: 1 9 Coco Solo Hosp. 16.V.1973 (at light) (P. D.
Ashlock), 1 9 same 8.V.1973 (D. Engleman); 1 9 Barro Colo. Is. IV.1940

'All measurements are in millimeters.

Florida Entomologist 64 (2)

(Jas Zetek) No. 4647 Lot no. 40-8104, 1? 1 $, 3 9 same IV.1941 (at light)
(Jas Zetek) No. 4776 Lot no. 41-7231, 1 & same 7.VIII.1967 (L. & C. W.
O'Brien), 2 9 same 7.VIII.1967 (at light) (C. W. & L. O'Brien), 1 3, 1 9
same 21.IV.1962 (H. Ruckes) 7 8, 1 9 same 18-V-1973 (Under Ficus)
(P. D. Ashlock), 1 9 same 20.VII.1962. GUATEMALA: 1 9 Chiquimula XI.
1930 (J. J. White) J. C. Lutz collection 1961, 1 9 Guatemala City (el. 5000
ft.) 11.1932 (C. N. Ainslie). VENEZUELA: 1 9 Monagas Jusepin 4.X.1965
(F. Fernandez, C. J. Rosales) Venezuela Ind. Zoo. Agricola. Fac. Agronomia
Univ. Central. TRINIDAD: 6 8, 3 9 Simla Arima-Blanchisseuse Rd. 20.VII.
1975 (el. 600 ft.) (Blacklight trap) (J. Price), 1 9 St. Augustine 14.VI.1973
(light trap) (R. Baranowski, F. O'Rourke, V. Picchi, J. Slater). 1 9 Curepe
14-VII-1975 (light trap) (F. D. Bennett). In P. D. Ashlock, J. A. Slater,
American Museum of Natural History, National Museum of Natural History,
R. M. Baranowski and Venezuela University Central collections.
This species is similar in general appearance to 0. atropicta. The chief
distinguishing features externally are the lack in parapicta of diagonal white
markings on the scutellum, a more extensively darkened (distal two-thirds)
3rd antennal segment and the dull pruinose dorsal body surface. In atropicta
the dorsal surface of the body is shining to subshining. This pruinosity dif-
ference between parapicta and atropicta is subtle over most of the body un-
less both species are available for direct comparison. The presence of
pruinosity in parapicta can be seen most easily on the anterior pronotal lobe
before, between, and immediately behind the calli. There are striking differ-
ences in the genitalia between the 2 species. The parameres, sperm reservoir
(see Fig. 2, 3, 4, 5) and number of coils of the vesica are quite different in
the 2 species. (Atropicta has 10 vesical coils in addition to the paramere and
reservoir differentiation). The degree of dark coloration on the 3rd antennal
segment is somewhat variable and while usually there is a considerably
greater portion darkened in parapicta than in atropicta the character is not
definitive. Insofar as we have been able to determine the differences in the
scutellar markings are definitive and provide a ready means of separating
the 2 species.
BIOLOGY: P. D. Ashlock (U. Kansas) took a series of adults and nymphs in
seed litter below a species of Ficus on Barro Colorado I. in the Canal Zone
(Panama). These specimens were taken together with a series of nymphs
and adults of Ozophora concava Distant and a single adult of 0. atropicta.
FIFTH INSTAR NYMPH: Nymphs of this species and those of Ozophora con-
cava were taken together on Barro Colorado Island (Canal Zone) Panama by
P. D. Ashlock on 18-V-1973. Both species show a variegated striped dark and
pale pattern similar to that described for Ozophora hirsuta by Slater and
Baranowski (1979). The 2 species are easily separable. Parapicta has a
glabrous dorsal surface, the dark longitudinal head stripes on either side of
the midline are continuous with the dark tylus and do not expand laterally
before the epicranial arms, the dark thoracic stripes laterad of the midline
are separated completely by a pale stripe from the still more lateral dark
areas, the lateral margin of the explanate margin is pale, the apical portion
of the mesothoracic wing pad, while dark, has a narrow pale vitta (some-
times reduced to a small spot), the dark sclerites around the dorsal abdom-
inal scent glands are short and subquadrate and the labium extends to or
slightly beyond the posterior margin of the metacoxae.


June, 1981

Slater & Hassey: Ozophora atropicta

Ozophora concava agrees with hirsuta and differs from parapicta in
having numerous conspicuous hairs on the dorsal surface, the dark longi-
tudinal head vittae not reaching the dark coloration of the tylus but ex-
panding laterally along the epicranial arms, having the dark coloration of
the pronotum confluent between the inner and outer striping so that the pale
markings appear as separated spots and streaks rather than as distinct
complete stripes, in having completely dark distal portions of the meso-
thoracic wing pads, noticeably wider, more slender and elliptical sclerites
about the dorsal abdominal scent gland openings and with the labium usually
only attaining or extending slightly between the metacoxae.


We should like to extend our appreciation to the following for allowing us
to examine material from their respective institutions or personal collections.
Dr. P. D. Ashlock (U. Kansas), Dr. R. M. Baranowski (U. Florida, Home-
stead), Dr. Dodge Engleman (Coco Solo Hospital C. Z.), Drs. F. Fernandez-
Yepez & Eduardo Osuna (Universidad Central de Venezuela), Dr. R. C.
Froeschner (National Museum of Natural History, Washington, D.C.), Dr.
Jerry Powell (U. California), Dr. J. C. Schaffner (Texas A & M University),
Dr. R. T. Schuh (American Museum of Natural History), Dr. G. G. E.
Scudder (U. British Columbia), Dr. J. Stehlik (Museum Brno), and Drs.
H. V. Weems, Jr. & R. E. Woodruff (Florida State Collection of Arthropods).
We are indebted to Mr. Stephen Thurston for the dorsal view illustration;
to Ms. Mary Jane Spring and Ms. Molly Hubbard (U. Connecticut) for
preparation of the charts and to Mrs. Elizabeth Slater (U. Connecticut) for
assistance in the preparation of the manuscript.

BARBER, H. G. 1939. Scientific survey of Porto Rico and the Virgin Islands:
Insects of Porto Rico and the Virgin Islands: Hemiptera-Heteroptera
(excepting the Miridae and Corixidae). Sci. Surv. Porto Rico 14(3):
263-441. New York Acad. Sci. Vol. 14(3): 226-441.
1954. The family Lygaeidae (Hemiptera-Heteroptera) of the island
of Cuba and the Isle of Pines-Part II. Mem. Soc. Cubana Hist. Nat.
22: 335-53.
AND P. D. ASHLOCK. 1960. The Lygaeidae of the Van Voast-Amer-
ican Museum of Natural History expedition to the Bahama Islands
1953. Proc. Ent. Soc. Washington 62 (2) : 117-24.
RAMOS, J. A. 1946. The Insects of Mona Island (West Indies). J. Agric.
Univ. Puerto Rico, Rio Piedras 30(1) : 1-74.
SLATER, J. A., AND R. M. BARANOWSKI 1979. New Species of Ozophora from
the Lesser Antilles with notes on the Biology and Immature Stages
(Hemiptera: Lygaeidae). Florida Ent. 62(3): 244-59.


Florida Entomologist 64 (2)


Biology Department, Fordham University
Bronx, NY 10458 USA

Monobia is a Tropical American genus, which ranges also into the south-
western and northeastern United States. The Neotropic M. biangulata
Saussure, the Nearctic M. quadridens (Linnaeus), and the Sonoran M.
texana (Cresson) represent Monobia in the Lower Rio Grande Valley of
Texas and in adjacent Mexico. Populations of these species inhabiting the
study area show some differences in phaenology, habitat preference, and
flower selection. Monobia biangulata is recorded for the 1st time from the
United States and M. quadridens is cited as new for M6xico.

Monobia es un genero neotropical, cuya distribuci6n ademas se extiende
hasta el suroeste y el nordeste de los Estados Unidos. Monobia estA rep-
resentada en el Valle del Bajo Rio Grande de Tejas y en parties limitrofes de
M6xico por 3 species: M. biangulata Saussure, de distribuci6n neotropical
mesoamericana; M. quadridens (Linnaeus), de distribuci6n principalmente
neartica; y M. texana (Cresson), de afinidades sonorenses. Poblaciones de
dichas species, presents en el Area bajo studio, demuestran algunas dif-
erencias entire si en cuanto a fenologia, ambientes y micro-ambientes, y
preferencias florales. M. biangulata es citada por primera vez para los
Estados Unidos, dAndose asimismo las primeras citas para M. quadridens en

In 1973 I began to survey Lower Rio Grande Eumenidae and have pub-
lished 2 contributions on taxonomy and ecology of local species in the genus
Zethus (Porter 1975, 1978a). I now review Monobia, another mainly Neo-
tropic taxon, all of whose United States representatives become sympatric at
the southern tip of Texas.
Monobia are large eumenids which may be recognized by the following
combination of characters: labial palpus with 3 segments, tegula elongate
basally, prestigma as long as hind margin of stigma, prepectal carina well
developed, propodeum with more or less triangular or tooth-like cristae,
gaster sessile. The above diagnosis (modified from Parker 1966: 173) will
separate south Texas Monobia from other local Eumenidae. Some large sym-
patric Euodynerus closely resemble Monobia quadridens or M. texana in
color and habitus. In Euodynerus, the labial palpus always has 4 well de-
veloped segments and the maxillary palpus has 6 segments, whereas the
palpal formula of Monobia is 3:5.

1Contribution No. 496, Bureau of Entomology, Division of Plant Industry, Florida Depart-
ment of Agriculture and Consumer Services, Gainesville, FL 32602 USA.
2Research Associate, Florida State Collection of Arthropods, Florida Department of Agri-
culture and Consumer Services, Gainesville.


June, 1981

Porter: Records for Monobia

The biology of this genus is practically unstudied. The species nidify by
refurbishing diverse preformed cavities in various substrates (tunnels in
wood, burrows in the ground, hollow stems, abandoned mud-dauber nests,
etc.). Their nests consist of linear series of cells separated by mud parti-
tions. They store several prey in each cell. Judging from the records sum-
marized by Krombein (1979: 1498), each species may select prey from among
several different families of Lepidoptera.
The 26 described species and subspecies of Monobia are confined to the
New World. Most of them occur in tropical South and Middle America, but
some have reached such peripheral areas as central Argentina, the north
Chilean Coastal Desert, the West Indies, and the eastern United States.

Key to United States Monobia

1. Apical rim of gastric tergites 2 and 3 forming a broad, subtly
differentiated, impunctate, translucently brownish lamella;
lamella of 2nd tergite about 0.1 as long as entire tergite;
macro-punctures on lower 2/3 of mesopleural disc medium-sized,
mostly separated by more than their diameters and becoming
sparse ventrad; upper 1/3 of mesopleuron with a large yellow
blotch; at least 1st and 2nd gastric tergites with yellow sub-
apical margins; legs black with variably profuse yellow marks
that include blotches on mid and hind coxae M. biangulata Saussure
1'. Apical rims of tergites 2 and 3 unmodified or at most forming
a narrow, concolorous lamella that is scarcely 0.03 as long as
whole tergite; macro-punctures on lower 2/3 of mesopleural
disc uniformly large and mostly separated by no more or by
less than their diameters, not thinning out ventrad; meso-
pleuron without yellow blotch; pale markings of gaster usually
confined to 1st tergite; legs black or ferruginous, without con-
spicuous yellow marks .. ..... -- .------ .....---- -----. 2
2(1). Black with creamy white marks which include broad areas on
pronotum and on postpetiole; 2nd gastric sternite abruptly de-
clivous at base, angular in profile; propodeum with dorsal and
apical faces usually separated mesad by a sharp crest; 13th $
flagellomere slender and tapered in profile, gently decurved
apicad and with ventral margin palpably concave
.......-------------.....----------- ........ - M quadridens (Linnaeus)
2'. Ferruginous with some black staining and often with yellow
marks that may include faint marginal stripes on pronotum
and a moderately broad subapical band on postpetiole; 2nd
gastric sternite evenly convex at base, not angulate in profile;
dorsal and apical faces of propodeum at no point separated by
a sharp crest; 13th & flagellomere comparatively stout in pro-
file, weakly tapered and only slightly concave ventrally -
.................--------- ---------------- -----M -exana (Cresson)

1. Monobia biangulata Saussure
(Fig. 3)

Florida Entomologist 64 (2)

MATERIAL EXAMINED: 2 9 and 2 : MEXICO, San Luis Potosi State,
Ciudad Valles, Hotel Covadonga, 1 9, 2-VI-1977, C. Porter, A. Cerbone;
UNITED STATES, Texas, Hidalgo County, Bentsen Rio Grande Valley
State Park, 1 9, 2 3, 1- and 4-VIII-1980, C. Porter.
The Texas records given above are the 1st for M. biangulata in the United
FIELD NOTES: The Texas M. biangulata were netted in dense shade as
they flew around a small sapling of Celtis lindheimeri Engelm. They were
caught between 1300 and 1400 CST, when the air temperature had reached
350 C. The collecting locality was in deep woods near 1 of the few perma-
nent water holes in Bentsen Park. Unusually large Celtis dominated the
overstory along with tall Fraxinus and Ulmus, while a few small Taxodium
mucronatum Ten. grew close to the water, and most trees were draped with
Spanish Moss.
In M6xico this species was collected from subtropical palm and Taxodium
mucronatum gallery woods along the Rio Valles on the grounds of the Hotel
Covadonga, 20 km south of Ciudad Valles.
PHAENOLOGY: 1 9 in VI, 1 9 and 2 $ in VIII.

2. Monobia quadridens (Linnaeus)
(Fig. 1, 5)
MATERIAL EXAMINED: 2 9 and 1 : MEXICO, Nuevo Le6n, Cola de
Caballo ca. El Cercado, 1 9, 1-VI-1974, C. Porter, C. Calmbacher; UNITED
STATES, Texas, Hidalgo County, McAllen Botanical Gardens at McAllen,
1 9,1 $, 7-IX-1978, C. Porter.
The literature offers no previous records of M. quadridens from M6xico.
FIELD NOTES: Both Texas specimens were collected in partial sun from
flowers of the introduced Brasilian ornamental shrub, Schinus terebinthi-
folius Raddi. Nearby natural vegetation was semihumid woods dominated by
Celtis, Condalia, Xanthoxylum, Acacia, Baccharis, and Prosopis.
In M6xico I have found M. quadridens only in the humid Cola de Caballo
ravine with its dense forest of Acer, Carya, Juglans, Platanus, Pinus,
Quercus, and Taxodium.
PHAENOLOGY: 1 9 in VI, 1 9 and 1 & in IX.

3. Monobia texana (Cresson)
(Fig. 2, 4, 6)
Hidalgo County, Bentsen Rio Grande Valley State Park, 1 9, 20-V-1977,
1 3, 8-VII-1980, 1 9, 11-VII-1980, 1 9, 2 S, 15-VII-1980, 1 9, 9-VIII-1980,
1 9, 22-VIII-1979, 4 9, 7-IX-1976, 1 9, 26-XI-1979, C. Porter; McAllen
Botanical Gardens at McAllen, 1 $, 28-III-1975, 2 9, 16-V-1975, 1 $, 25-V-
1975, 1 3, 1-VI-1976, 1 3, 5-VI-1975, 1 '9, 22-VIII-1974, 1 8, 1-IX-1978,
1 S, 2-IX-1978, 1 9, 1 8, 4-IX-1975, C. Porter.
FIELD NOTES: M. texana flies beneath full to partial sun throughout open
subtropical "thorn scrub" or along the edges of denser woods. I have collected
it, usually at 1-3 m above ground level, on flowers of such trees and shrubs
as Baccharis glutinosa (Ruiz L6pez & Pav6n) Pers., B. neglect Britton,
Condalia obovata Hook., Salix interior Rowlee, and Schinus terebinthifolius


June, 1981

Porter: Records for Monobia

Fig. 1-5. 1) Monobia quadridens, 3, McAllen, Texas. Dorsal view of 1st
gastric tergite, showing color pattern; 2) Monobia texana, $, Bentsen
Rio Grande Valley State Park, Texas. Dorsal view of 1st gastric tergite,
showing color pattern; 3) Monobia biangulata, $, Bentsen Rio Grande Val-
ley State Park, Texas. Dorsal view of apex of 2nd and all of 3rd gastric
tergite, showing apical lamellae and color pattern; 4) Monobia texana, $,
Bentsen Rio Grande Valley State Park, Texas. Lateral view of flagellomeres
10-13; 5) Monobia quadridens, $, McAllen, Texas. Lateral view of flagel-
lomeres 10-13.
Raddi. It becomes active when the air temperature reaches 250 C and con-
tinues flying even at 36 or 370 C.
MONTHLY PHAENOLOGY: 1 8 in III, 2 9 and 2 in V, 3 a in VI, 2 9
and 3 & in VII, 3 9 in VIII, 5 9 and 3 & in IX, and 1 9 in XI.



Florida Entomologist 64(2)

Fig. 6. Habitat of Monobia texana photographed during January 1978
near the McAllen Botanical Gardens at McAllen, Texas. The large, dark-
leaved shrub in the middle of the picture is Condalia obovata. The spindly
shrubs at the far left are Baccharis neglecta. During late summer and fall,
when in flower, both plants attract many specimens of Monobia texana.

ANNUAL PHAENOLOGY: 3 9 in 1974, 1 9 and 5 8 in 1975, 5 9 in 1976,
1 9 in 1977, 2 & in 1978, 2 9 in 1979, 3 9 and 3 3 in 1980.

ZOOGEOGRAPHY: Monobia fulfills a paradigmatic post-Pleistocene Neo-
tropic distributional pattern. Bequaert (1940: 882) divided the genus into
26 species and subspecies. Eighteen occupy various ranges in South America
from Colombia or Venezuela to Brasil or Argentina (1 attains PanamA),
and 12 of these form a diversity nucleus situated between the Amazon Basin
and Paraguay. A secondary radiation in Middle America includes 3 species
which extend from M6xico into northern South America, plus 1 that is re-
stricted to M6xico and south Texas. The West Indies have 1 species each on
Cuba and Hispaniola. Southwest North America possesses 1 endemic species.
The eastern United States has a single apparently relict form. No Monobia
have reached (or now survive) at the Newv World's temperate antipodes in
sub-Atacaman Chile.
The data given above show that Monobia primarily inhabits tropical
evergreen and deciduous forests but that it also has entered deserts at sev-
eral extremes of its range. Two Monobia occupy the Argentine Chaco Thorn
Scrub and Subandean Desert, 2 occur in the Peruvian Coastal Desert, and 1
frequents the southwestern United States and northern M6xico. These wasps


June, 1981

Porter: Records for Monobia


also fly vigorously and are versatile in prey selection. Monobia thus should
be able to disperse effectively, and that is reflected in the ample ranges of
most species. Like Zethus (Porter 1978a), this eumenid genus also appears
to be limited mainly by cold and, perhaps for that reason, avoids temperate
Chile and has only 1 species in temperate North America.
The Lower Rio Grande Valley Monobia belong to 3 major zoogeographic
categories and in part show correspondingly divergent habitat preferences.
Monobia biangulata is a Middle American Neotropical sylvatic species that
ranges from south Texas throughout M6xico. I have collected it only
beneath the shade of lush subtropical forests and always near water. Among
sympatric Eumenidae, Hypalastoroides mexicanus (Saussure) and Zethus
otomitus Saussure most closely parallel M. biangulata both in range and
habitat, but many Ichneumonidae and other hygrophile biota fit the same
pattern even more perfectly (Porter 1978a: 164). Monobia texana belongs
to the Sonoran distributional category and occurs from the Mexican states of
Veracruz, Sonora, and Baja California Sur north to Texas and Arizona. It
flies mostly under bright sun in open brushland or at the edges of denser
woods. As discussed in an earlier contribution (Porter 1978b: 174), numerous
south Texas biota (e.g., the eumenid, Dolichodynerus tanynotus Cameron)
show geographic and environmental distributions similar to those of M. tex-
ana. However, the Sonoran fauna is a heterogeneous complex of elements
derived from Holarctic, Nearctic, and several different Neotropic sources.
Monobia texana is Neotropic in origin, but more data will be needed on
relationships within the genus before it can be decided whether M. texana is
a dry-adapted offshoot of an old sylvan Middle American stock or whether
its nearest relatives are in South American deserts and it spread northward
during some late Tertiary xerothermic episode. Monobia quadridens ranges
from Florida to New Hampshire west to Wisconsin, Kansas, and Texas. It
also has apparently disjunct populations in New MBxico and northeast
Mexico. Monobia quadridens is a northeast North American Neotropic ele-
ment that differentiated from an ultimately South American ancestor while
isolated during Pleistocene glacial maxima somewhere in the southeastern
United States and which in the present interglacial has extended its range
both northeast and southwestward. Other zoogeographically comparable
North American Eumenidae are Pachodynerus erynnis (Lepeletier), Para-
nortonia symmorphus (Saussure), Pseudodynerus quadrisectus (Say), and
Zethus spinipes Say. Monobia quadridens in south Texas and adjacent
M6xico appears to occupy a spatial niche intermediate between the deep
woods preferred by M. biangulata and the open, sunny haunts of M. texana,
but more field observations will be needed to confirm this impression.
The study region thus has 3 taxonomically, ecologically, and distribution-
ally well differentiated Monobia. Monobia biangulata and M. quadridens are
more or less hygrophile eumenids which seem in the Lower Rio Grande
Valley to approach the limits of their environmental tolerance. Monobia
texana, with xeric proclivities, is common and finds in the Valley an almost
optimum habitat. These facts probably reflect not only the relative aridity
prevailing in south Texas during the current interglacial period, but also
the cultural desertification which man has inflicted on the region with his
intensive farming and flood control projects.
PHAENOLOGY: Monobia biangulata and M. quadridens have been col-

Florida Entomologist 64(2)

elected very sporadically in the study area and only during summer. More
records are needed to confirm their patterns of seasonal distribution.
Monobia texana flies from March to November and is conspicuous every
year. It reaches its maximum in September (8 of 25 specimens). A spring to
late fall activity period with an absolute or relative peak in September
characterizes numerous Lower Rio Grande Valley Hymenoptera. Among
these are the eumenids, Zethus aztecus (Porter 1978a: 165), the sphecids,
Bicyrtes capnoptera (Porter 1980: 284), Sphex dorsalis Lepeletier, S.
habenus Say, and S. texanus Cresson (Porter 1978b: 175), and the thermo-
philic ichneumonids Mesostenus longicaudis Cresson and Diapetimorpha
macula Cameron (Porter 1977: 82-3). September, with its tropical storms, is
the region's most consistently verdant month. The concurrent upsurge of
insect activity might also be caused by this climatic factor.
FLOWER PREFERENCE: Monobia texana visits flowering shrubs and trees
where it may feed at heights beyond reach of the standard insect net. I have
not seen it on herbaceous plants. Most other Lower Rio Grande Valley
eumenids (Hypalastoroides, Stenodynerus, Euodynerus, Pachodynerus,
Eumenes, and Zethus) seek nourishment not only in the shrub and tree
strata of their communities, but also from low herbs, such as Rivina,
Teucrium, Aster, Helianthus, Parthenium, and Pluchea.

Material covered in this study has been deposited in the Florida State
Collection of Arthropods (Division of Plant Industry, P. O. Box 1269, Gaines-
ville, FL 32602) and in the author's private collection (301 N. 39th Street,
McAllen, TX 78501).

Fieldwork in south Texas and M6xico was supported in 1980 by a Faculty
Fellowship from Fordham University, during 1973-75 and in 1979 by grants
from the Committee for Research and Exploration of the National Geo-
graphic Society, and from 1976-77 by United States National Science Founda-
tion Grant DEB-75-22426. The Texas Parks and Wildlife Department has
issued permits for collecting in Bentsen Park (current permit number 2-80).
Mr. Anthony F. Cerbone of Fordham University has assisted me during
some of the field work involved in this study.

BEQUAERT, J. 1940. Synopsis of Monobia de Saussure, an American genus
of solitary wasps. Rev. de Ent. 11: 822-42.
KROMBEIN, K. V. 1979. Family Eumenidae, p. 1472-1510 In Krombein, K. V.,
P. D. Hurd, D. R. Smith, and B. D. Burks, Catalog of Hymenoptera in
America north of Mexico, Vol. 2, Aculeata. Smithsonian Institution
Press, Washington, D. C.
PARKER, F. D. 1966. A revision of the North American species in the genus
Leptochilus. Ent. Soc. America, Misc. Pub. 5: 153-229.
PORTER, C. 1975. New records for Zethus from Texas. Florida Ent. 58:


June, 1981

Porter: Records for Monobia

1977. Ecology, zoogeography, and taxonomy of the Lower Rio
Grande Valley mesostenines. Psyche 84: 28-91.
1978a. Ecology and taxonomy of Lower Rio Grande Valley Zethus.
Florida Ent. 61: 159-67.
1978b. Ecological notes on Lower Rio Grande Valley Sphecini.
Florida Ent. 61: 169-78.
---. 1980. Bicyrtes Lepeletier (Hymenoptera: Sphecidae) in the Lower
Rio Grande Valley of Texas and in northeast Mexico. Florida Ent. 63:


Department of Zoology
Erindale College
University of Toronto
Mississauga, Ontario, Canada L5L 1C6

Homosexual mounting is common in insects. Half the mounting attempts
of Palmacorixa nana Walley males are homosexual and weights of mounting
and mounted animals show that mounters are seeking animals larger than
themselves. Since females are larger than males, this behaviour results in
some proportion of correct choice of mates.

La monta homosexual es comin entire los insects. La mitad de los intentos
de montar de los machos de Palmacorixa nana Walley son de naturaleza
homosexual y los pesos de los animals montados y de los que montan revelan
que los que montan buscan a los animals mas grandes que ellos mismos.
Como las hembras son mas grandes que los machos, este comportamiento di
por resultado en que el macho escoja correctamente una hembra en alguna
proporci6n de los casos.

Homosexual mounting has been reported for Diptera (Spieth 1952, Tauber
and Toschi 1965), Neuroptera (Henry 1979), Lepidoptera (Benz 1973,
Palaniswamy et al 1979) and Hemiptera (Constanz 1974). Most of these
encounters are briefer than heterosexual'encounters (Benz 1973, Henry 1979)
but can represent a substantial investment in time and effort.
Adults of Palmacorixa nana Walley are small water bugs that live in
dense aggregations in quiet permanent water throughout the summer months.
Since homosexual mounting accounts for more than half the mounting ac-

1Present address: Department of Entomology, University of Alberta, Edmonton, Alberta,
Canada, T6G 2E3.


Florida Entomologist 64(2)

tivity of males (Aiken 1980), I attempted to discover the physical basis of
such errors. Tauber and Toschi (1965) suggested that in species where homo-
sexual mounting is common, males and females may present similar stimuli
to sexually active males, but the authors did not speculate on what these
stimuli may be. Since the most obvious difference between male and female
P. nana is the larger size of the female, I investigated this factor as a po-
tential source of error in mounting.

Males were collected in Black Creek, Ontario. They were housed in 57
liter glass aquaria in an environmental room set at 200C with a 16:8 light:
dark cycle. All bugs were used within 48 hours.
The animals were observed in a 10 x 15 cm section of a glass aquarium
with a water depth of 15 cm. Thirty males were introduced into this aquarium
and allowed 30 min to acclimate. The animals were then observed con-
tinually for one hour. Six such observation sessions were completed.
Coupled males were captured in a siphon tube, transferred to separate
shell vials and 2 more males were added to the aquarium from a stock
culture. Each member of the pair was then removed from its respective shell
vial, air dried for one minute and weighed on a Mettler H20 analytical
balance. This procedure was repeated until weights for 30 pairs were ob-
tained. Weight was used as an index of size in order to avoid excessive
handling of animals that were needed for use in subsequent experiments.
Weights of 171 males and 42 females were also taken to determine the
distribution of weights in both sexes. The number of each sex represents the
sex ratio in the stream at the time of collecting.


Most attempts at mounting by P. nana males consist of one animal
swimming rapidly onto the dorsum of another, grasping the pronotum with
the forelimbs and attempting to intromit by moving the tip of the abdomen
under the venter of the inferior animal. In homosexual encounters, the
mounting male disengaged after 1-2 seconds of genital probing.
Figure 1 shows that males are lighter than females and that there is no
overlap in the weights of the sexes.
Weights of mounted and mounting animals (Fig. 2) show that in 25 of 30
encounters, the mounted animal was heavier than the mounting animal (p <
0.02 sign test, Siegel 1956). In those instances where the mounting animal
was smaller than the mounted animal, the former comprised 90.16% of the
weight of the latter. This weight relationship was consistent throughout the
entire range of male weights.

Male P. nana attempt to mate with any other conspecific individual,
principally attempting to mate with any bug larger than themselves. Since
all females are larger than males and are present in the population through-
out the season, this behaviour would result in some proportion of correct
choices. For some males, this is a more productive strategy than for others.


June, 1981

Aiken: Palmacorixa nana




25 3 35 4 45 5 55
Weight in milhgrams

Fig. 1. Weights (mg.) of 171 male and 42 female Palmacorixa nana.

The chance of error is relatively great for any low weight male with
essentially the entire population as a pool of potential mates. For heavy
males, the proportion of females in the pool of heavier animals is greater,
enhancing the chances for making a correct choice. Consequently, there
should be proportionally more light males in the population of homosexual
mounters than in the population of males as a whole. A test of the data con-
firmed this (t = 43.81, p < 0.005).
Two possible explanations come to mind for the presence of such ap-
parently wasteful behaviour. First, the incidence of homosexual mounting
might be viewed as an inconsequential accident. Because of the ephemeral
nature of each mounting attempt, little time and energy are expended. This
behaviour is then just a tolerable inefficiency in the system. Indeed, it seems
to be most prevalent in those insects exhibiting low male investment (usually
only sperm) in a particular mating. Secondly, it could serve as an indicator
of male density and/or sex ratio. The frequency with which a given male
(depending on size) mounts other males or is mounted could be an indicator
of excessively high male density and/or a sex ratio heavily skewed toward
males. Aiken (1980) has already shown that males actively avoid areas of
already high male density. In either case, the incidence of homosexual mount-
ing could serve as a behavioral indicator of the mate acquiring potential of
one area over another.


I thank G. K. Morris, P. D. Bell and G. E. Ball for critical comments on
the manuscript. This study was supported by Natural Sciences and Engi-
neering Research Council of Canada Operating Grant #4946 to G. K. Morris.


270 Florida Entomologist 64 (2) June, 1981


equal weight-.

0 *
35- 0 "


3.0- .

* *

2-5 3-0 3-5 4-0

Weight of mounted male

Fig. 2. Comparison of weights of mounted and mounting males in 30
homosexual encounters. Points falling below the dashed line indicate that the
mounted male is heavier.

AIKEN, R. B. 1980. The role of acoustic signalling in the aggregating and
mating behaviour of Palmacorixa nana Walley (Heteroptera: Corixi-
dae). Ph.D. thesis. University of Toronto. 119 p.
BENZ, G. 1973. Role of sex pheromone and its insignificance for heterosexual
and homosexual behaviour of larch bud moth. Experientia 29: 553-4.
CONSTANZ, G. 1973. The mating behaviour of a creeping water bug, Am-
brysus occidentalis (Hemiptera: Naucoridae). American Midi. Nat.
92: 230-9.
HENRY, C. 1979. Acoustical communication during courtship and mating in
the green lacewing, Chrysopa carnea (Neuroptera; Chrysopidae).
Ann. Ent. Soc. America 72: 68-79.
PALANISWAMY, P., W. D. SEABROOK, AND R. ROSs. 1979. Precopulatory be-
haviour of males and perception of potential male pheromone in spruce
budworm, Choristoneura fumiferana. Ann. Ent. Soc. America 72:
SIEGEL, S. 1956. Non-parametric statistics. McGraw-Hill. Toronto.
SPIETH, H. 1952. Mating behaviour within the genus Drosophila (Diptera).
SPIETH, H. 1952. Mating behaviour within the genus Drosophila (Diptera).

Aiken: Palmacorixa nana

Bull. Amer. Mus. Nat. Hist. 99: 401-74.
TAUBER, M., AND C. TOSCHI. 1965. Bionomics of Euleia fratria (Loew)
(Diptera: Tephritidae). I. Life history and mating behaviour. Ca-
nadian J. Zool. 43: 369-79.


Department of Entomology, Smithsonian Institution
Washington, D.C. 20560 USA

Dichrorampha sapodilla, new species, and Dichrorampha manilkara, new
species, are described from Florida. The former species feeds as a larva on
flowers of sapodilla (Manilkara zapota (Linnaeus) Van Royen) and the
latter species feeds on flowers of wild dilly (Manilkara bahamensis (Baker)
Lamarck & Meeuse).

Dichrorampha sapodilla, nueva species, y Dichrorampha manilkara,
nueva species, describir de Florida. Las species antecedente alimentarse de
sapodilla flores (Manilkara zapota (Linnaeus) Van Royen) como la larva y
las species iltimo alimentarse de "wild dilly" [dilla inculto] (Manilkara
bahamensis (Baker) Lamarck & Meeuse).

In anticipation of a future paper on the biologies of 2 new Dichrorampha
tortricids from subtropical Florida, the new species are described herein.
Both have been reared from host plants of the family Sapotaceae, one from
commercial sapodilla, Manilkara zapota (Linnaeus) Van Royen, and the
other from what is locally referred to as "wild dilly," Manilkara bahamensis
(Baker) Lamarck & Meeuse. Larvae of both species of tortricids utilize the
flowers of their respective hosts. Both species are currently known only from
southern Florida, mainly from the Florida keys.

Dichrorampha sapodilla Heppner, NEW SPECIES
Fig. 1, 2, 5-8
Hemimene sp.-Kimball 1965: 263; Bacheler and Baranowski 1975: 158.
SIZE: Forewing length 3.8-4.5 mm.
HEAD: Gray mottled with tan; frons tah; labial palpus cream white; an-
tenna gray and tan mottled.
THORAX: Gray fuscous; venter tan-white; legs tan. Forewing: gray with
dark fuscous bar from basal 1/3 of anal margin to midwing, angled toward
apex; costa with 6-7 major dark fuscous strigulae continued diagonally
toward tornus, surrounded by yellow, with white borders immediately along
costal margin; apical 4 fuscous strigulate continued to tornus after sharp

Florida Entomologist 64(2)



wr -*
ArF .-

3 4

Fig. 1-4. Adults of Dichrorampha species: 1, D. sapodilla Heppner, new
species, paratype 8; 2, same, paratype 9 ; 3, D. manilkara Heppner, new
species, paratype ; 4, same, paratype 9 [all from Florida].
bend at radius, surrounded by yellow except at tornus where a large dark
fuscous mark has yellow only on distal side of mark; 5 small black subdistal
(speculum) spots in yellow field; fringe fuscous; ventral side lustrous bronze
fuscous. Hindwing: fuscous; fringe pale fuscous; ventral side lustrous
bronze fuscous.
ABDOMEN: Fuscous; venter tan-white. Male genitalia: tegumen stout, arched
to acute point; gnathos a narrow band across membrane at center; vinculum
reduced; anellus a small semi-circular plate; valva elongate with setaceous
dorsally directed cucullus, rounded along ventral margin; a narrow
neck prior to broad base of valva; aedeagus small, wedge-shaped, with
numerous (ca. 40) cornuti. Female genitalia: ovipositor with setaceous
papilla anales; apophyses with anterior pair somewhat longer than posterior
pair; ostium a cup-shaped sclerotized area posterior to 7th sternite, with a
pointed fold extended over ostium; ductus bursae sclerotized to near a
sclerotized area at bursa, with ductus seminalis emerging from near bursa;
bursa copulatrix ovate, rugose interior walls; one large thorn-like signum.
TYPES: Holotype 8 : Homestead, Dade Co., Florida, 4-IV-1974 (larva), R. M.
Baranowski, emerged 18-IV-1974 ex "Achras zapota" (USNM type no.
76751). Paratypes: (6 $, 6 9) Florida.-Homestead, Dade Co., 4-IV-1974
(larvae), R. M. Baranowski, emerged ex "Achras zapota" 18-IV-1974 (2 3,
3 9) (FSCA); Miami, Dade Co., 18-V-1967, D. H. Habeck, on sapodilla


June, 1981


Heppner: New Dichrorampha

flowers (A160), (1 8) (FSCA); Subtropical Res. Sta., Homestead, Dade
Co., 3-VI-1974, J. B. Heppner, emerged ex "Achras zapota" 13-VI-1974 (2 3,
2 ) (JBH); Dear Res., Big Pine Key, Monroe Co., 19-VI-1973, J. A.
Powell, (1 ) (UCB); Homestead, Dade Co., 4-XI-1958, D. O. Wolfenbarger,
(1 Y) (CPK). Paratypes to BMNH and USNM.
ADDITIONAL RECORDS: Florida.-Homestead, Dade Co., 31-III-1959, ex "Calo-
carpus sapota flowers" 2-IV-1963 (CPK); Siesta Key, Sarasota Co., 11-V-
1957, (CPK). [Kimball (1965) records].
HOST: Manilkara zapota (Linnaeus) Van Royen (Sapotaceae) flowers [plant
is sometimes referred to as Manilkara zapotilla (Jacq.) Gilly and in other




" t' ^
K:,. It.




t __ ,

&k: -4.
'1. .%

r 'flo \

Fig. 5-8. Dichrorampha sapodilla Heppner, new species (Florida): 5,
paratype & genitalia; 6, same aedeagus (enlarged); 7, paratype 9 genitalia;
8, same, ductus bursae.



~... ~h~L~ij'

Florida Entomologist 64(2)

REMARKS: This and the following new species are superficially rather similar
but the genitalia are markedly different. The maculation of D. sapodilla is
distinguished by the dark forewing mark near the wing base and the lack of
the silvery tornal line of the hindwing which is present in males of
Dichrorampha manilkara, new species. The genitalia will distinguish the
species from other Dichrorampha.
Bacheler and Baranowski (1975) noted that D. sapodilla (as "Hemimene
sp.") competes with the anthocorid Paratriphleps laeviusculus Bacheler and
Baranowski on flowers of sapodilla.

Dichrorampha manilkara Heppner, NEW SPECIES
Fig. 3-4, 9-12

SIZE: Forewing length 3.5-4.5 mm.
HEAD: Gray with tan frons; labial palpus tan-white, apical segment gray;
antenna fuscous.
THORAX: Fuscous; patagia tan; venter tan-white; legs tan with fuscous on
distal sides. Forewing: gray with 7 dark fuscous strigulae bordered by
orange (white on costal margin) from costal margin angled toward tornus;
anal margin dark fuscous; 5 dark fuscous strigulae from basal 1/3 of anal
margin angled toward apex and bordered by orange; distal 1/3 of wing
orange with extensions of dark strigulae intruding and large silvery line
from tornus straight to radius, then sharply angled away from apex to 4th
strigula along costal margin; apical margin dark fuscous; fringe dark
fuscous; ventral side lustrous dark fuscous. Hindwing: uniform dark fuscous
becoming darker only along anal margin in both sexes; males with a silver
iridescent line along tornal margin; ventral side as forewing.
ABDOMEN: Fuscous; venter tan-white. Male genitalia: tegumen with a dorsal
point; vinculum reduced; gnathos a broad band; anellus a large semi-circular
plate; valva elongate and curved upwards, with a large setaceous cucullus
and a mesally directed stoutly setaceous lobe midway along ventral margin;
valval neck almost absent; ventral valval margin with several stout setae;
aedeagus small, curved, with numerous (ca. 40) small spine-like cornuti.
Female genitalia: ovipositor setaceous on papilla anales; apophyses subequal,
of average stoutness; ostium a small sclerotized entrance behind an asym-
metrical circular sternal plate; ductus bursae partially sclerotized near
bursa, with ductus seminalis diverging from near bursa; bursa copulatrix
ovate, rugose, with one large thorn-like signum.
TYPES: Holotype : Middle Torch Key, Monroe Co., Florida, 12-VI-1974
(larval), J. B. Heppner, emerged ex "Manilkara emarginata" (rearing 74F4)
6-VII-1974 (USNM type No. 76750). Paratypes: (21 8, 33 9) Florida.-
Middle Torch Key, 12-VI-1974 (larvae), J. B. Heppner, emerged ex "Manil-
kara emarginata" (rearing 74F4) 24-VI, (1 S, 1 a), 25-VI (1 8), 27-VI
(1 9),29-VI (1 3 9), 30-VI (1 S,3 9), 2-VII (1 1 9), 6-VI (1 $),
10-VII (2 $,3 9), 11-VII (1 8),12-VII (3 9),20-VII (1 $) (JBH); Key
Largo, Monroe Co., 27-II-1967, S. Kemp (1 9) (CPK), 19-III-1968 (1 $),
15-VII-1967 (8 o, 11 9), 7-XI-1965 (1 9) (CPK); 2 mi. NE. Tavernier,
Key Largo, Monroe Co., 17-VI-1974, J. B. Heppner, (2 9) (JBH); 2 mi.
NE. Tavernier, Key Largo, Monroe Co., 20-VI-1973, J. B. Heppner & J. A.
Powell, (2 8-UCB; 3 9-JBH). Paratypes to BMNH, FSCA, USNM.


June, 1981

Heppner: New Dichrorampha

J I-k

9 10

Fig. 9-12. Dichrorampha manilkara Heppner, new species (Florida) : 9,
paratype S genitalia; 10, same, aedeagus (enlarged); 11, paratype 9 gen-
italia; 12, same, sternal plate.

HosT: Manilkara bahamensis (Baker) Lamarck & Meeuse (Sapotaceae)
flowers [called "wild dilly" locally].
REMARKS: The remarks for the previous species will serve to distinguish the
2 species. Dichrorampha manilkara superficially also resembles a more
northern species, Dichrorampha leopardana (Busek), but is smaller and has
less orange on the forewings in addition to very different genitalia. Both the
male and female genitalia are very unusual for the genus Dichrorampha.
This species appears to be common locally wherever wild dilly bushes are

The following collections, with their respective abbreviations, were con-
sulted: Charles P. Kimball Collection, West Barnstable, Massachusetts


Florida Entomologist 64 (2)

(CPK); Florida State Collection of Arthropods, Gainesville, Florida
(FSCA); John B. Heppner Collection, Washington, D.C. (JBH); University
of California, Berkeley, California (UCB); and the National Museum of
Natural History, Smithsonian Institution, Washington, D.C. (USNM). In
addition to paratype deposition with the above collections, paratypes will be
deposited with the British Museum (Natural History), London (BMNH).
R. L. Brown and J. F. G. Clarke (Smithsonian Institution) kindly provided
helpful criticisms. I wish also to thank the University of Florida (Depart-
ment of Entomology, Institute of Food and Agricultural Sciences) for sup-
port of my research efforts while at the University of Florida, which made
possible several field trips to southern Florida as well as study in Gainesville.

BACHELER, J. S., AND R. M. BARANOWSKI. 1975. Paratriphleps laeviusculus,
a phytophagous anthocorid new to the United States (Hemiptera:
Anthocoridae). Florida Ent., 58: 157-63.
KIMBALL, C. P. 1965. The Lepidoptera of Florida. An annotated checklist.
Arthrop. Florida Neighbor. Land Areas, 1: 1-363; 26 pl.


Silver City, NM 88061 USA

Maturity and adult activity data were accumulated for 21 species of
ground-surface spiders from 2 locations in the pinyon-juniper life zone and
2 in the arid-grasslands of southwestern New Mexico. Web-building Psilo-
chorus species were found to be the most dominant ground-surface adult
spiders in both associations. The 2 species concerned were not strongly
seasonal in maturity and adult activity indicating that they either matured
erratically or the adults were long lived. Except for Herpyllus propinquus
Keyserling, other less dominant or prevalent species were found to vary
seasonally in periods of maturity and adult activity. Some were also re-
stricted to 1 or the other of the 2 plant associations. A few were restricted to
1 plot.

Se acumularon datos sobre la madurez y la actividad adulta de 21 species
de arafias de la superficie del suelo en 2 localidades en la zona de pifion -
junipero y en 2 localidades en los terrenos pastorales aridos del suroeste de

'Contribution No. 458. Bureau of Entomology, Division of Plant Industry, Florida Depart-
ment of Agriculture and Consumer Services, Gainesville, FL 32602 USA.
2Contribution of the Office of Research, Western New Mexico University, Silver City, New
Mexico 88061 USA.
"Entomologist Emeritus, IFAS, University of Florida, Gainesville. Research Associate,
Bureau of Entomology, Division of Plant Industry, Florida Department of Agriculture and
Consumer Services, Gainesville, and Western New Mexico University, Silver City.


June, 1981

Muma: Ground-Surface Spiders 277

Nuevo Mexico. Se encontr6 que las species de Psilochorus que construyen
telas son las arafias adults mas dominantes en la superficie del suelo en
ambas asociaciones. Las 2 species estudiadas no mostraron relaciones
temporales fuertes con respect a la madurez y a la actividad adulta, in-
dicando o que maduraron erraticamente o que los adults vivian largo tiempo.
Exceptuando Herpyllus propinquus Keyserling, otras species menos domi-
nante o menos prevaleciente se variaron en periods de madurez y de
actividad adulta segfin la temporada. Algunas species fueron restringidas
a una o otra de las 2 asociaciones de plants. Unas pocas fueron restringidas
a una sola parcela.

The adult maturity and activity data presented and analyzed here indi-
cate the seasonal succession of adults of 21 dominant and prevalent spider
species in 2 different plant associations in southwestern New Mexico. Com-
bined with previously published data, they suggest that Barnes' (1953) 2
contentions: 1) that species attaining peak population in the spring would
not be rare in the same community in the fall, and 2) that apparent seasonal
successions were the result of a workers lack of identification of immatures,
are both open to question.
Within the past 50 years, studies on populations of North American
spiders have increased in number dramatically. Although many are little
more than species lists, as noted by Turnbull (1973), at least 20 studies have
involved efforts to quantify populations and species for analyses of ecological
importance. Eight of these papers have included tables and/or graphs indi-
cating the annual periods of maturity and adult spider activity: Elliott
(1930), Gibson (1947), Muma and Muma (1949), Chew (1961), Whitcomb
et al. (1963b), Coppel and Smythe (1963), Peck (1966), and Muma (1973).
Comparative population studies such as those of Lowrie (1948), Whitcomb
et al. (1963a), and Muma (1975a) have not included adult activity data.
Further, 2 important papers, those of Lowrie (1942) and Barnes (1953),
also have not included information on adult activity. Barnes discussed the
omission of such data from his study in his section on seasonal succession.
His contentions were that if a species attained a peak population in the
spring, it would not be rare in the same community in the fall, and that
apparent seasonal successions were the result of a worker's identification of
adults and lack of identification of immatures. Close examination of the data
of Chew (1961), Coppel and Smythe (1963), Peck (1966), Muma (1973,
1980 and here) indicate that neither of Barnes' contentions are necessarily
true; a species may be abundant during 1 season and rare or absent during
another, both as adults and as immatures. Some of this variation in abund-
ance is unquestionably caused by the often stated "sampling inadequacy."
On the other hand it is unwise to overlook the fact that population variations
are most often the result of critical changes in temperature, moisture, food
supply, and natural enemies, as stated by Muma (1973). Such factors can
and do limit the periodicity, size, and temporal span of adult activity. Many
earlier workers obviously did not overlook this problem, since they published
maturity and activity data.
Other aspects of the populations producing the maturity and adult ac-
tivity data presented here were recorded in another publication (Muma 1980).

Florida Entomologist 64 (2)

The study areas or plots were located near Silver City and in the Burro
Mountains in pinyon-juniper associations, and near Hurley and near Lords-
burg, New Mexico, in arid-grassland associations. They were the same as
those utilized by Muma (1974) for solpugid studies. Critical data on plot
location, elevation, soil, and plants are given in that paper.
Can traps, those described by Muma (1970) and evaluated by Muma
(1975b), containing 250 cc. of a killing-preserving medium (a 1:1 mixture
of 70% isopropyl alcohol and commercial ethylene glycol), were the only
collecting devices utilized in this study. Ten traps were set in each study
area; 5 in a north-south transect at about 10 m intervals and 5 in an east-
west transect at similar intervals. Traps were visited every 2 weeks from 1
April 1972 to 1 December 1973; specimens were screened from the medium,
the medium was reconstituted with a 3:1 mixture of isopropyl alcohol and
ethylene glycol, and the traps were reset.
In order to facilitate sampling and interpretation of data, the year was
arbitrarily divided into 4 series of 3 months each, approximating but not
coinciding with the seasons. March, April, and May represent spring; June,
July, and August, summer; September, October, and November, fall; De-
cember, January, and February, winter.
Although sampling was continuous for 20 months, only the data from 1
April through 30 November of each year are included in order to present
uniform, comparative, quantitative data for both 1972 and 1973. Specimens
collected from 1 December 1972 through 31 March 1973 are mentioned only
in the discussion.
Spiders were sorted, counted, and identified in the laboratory. Although
some spider genera and species are identifiable as immatures, many are not,
so immatures were identified only to family and are not cited or discussed
here (see Muma 1980). Species of certain genera, such as Meioneta of the
Linyphiidae and Micaria of the Gnaphosidae, were not identified owing to the
lack of adequate generic reviews and are also not included in this paper. Dr.
Willis J. Gertsch of Portal, Arizona, identified or confirmed the identification
of most of the species recognized here.
Luczak's (1960) terms, dominant and influent, are recognized respectively
for those species that were strikingly abundant, and those that were reg-
ularly collected. Luczak's accessory species, those that were uncommon or
rare, are discussed but not included here (see Muma 1980).
Since Muma (1975b) indicated that 30 traps per study area were re-
quired for precision limits of certain spider families and species within 30%
of the mean (x) number of specimens collected per trap, only those species
with populations differing by 50% or more of their means are considered
significantly different here.

Quantitative data on species maturity and adult activity of 5 dominant
spider species and 16 influents are presented in Table 1 for the pinyon-
juniper plant associations and Table 2 for the arid-grasslands.

June, 1981

Muma: Ground-Surface Spiders

(M TI tr r4 r4 i r4 i r 0 r4 r4 0S 0 c

+1 +1 +1 +1 +1 +1 + 1 +1 +1 +1 +i +1

Lr2 0 CO ] N N N C] C' (=

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280 Florida Entomologist 64(2) June, 1981

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Muma: Ground-Surface Spiders

The data presented in the tables are relatively easy to interpret. However,
from the standpoint of adult activity, several factors inherent in the loca-
tion of the plots and the periods of sampling introduce certain biases that
are not apparent. These are discussed in the following paragraphs.
The web-building Psilochorus imitatus Gertsch and Mulaik, which appears
to be the most abundant ground-surface spider in the area, was dominant in
both pinyon-juniper plots and the Hurley arid-grassland plot but was re-
placed by another dominant, Psilochorus utahensis Chamberlin, on the Lords-
burg arid-grassland plot. Adults of P. imitatus were either more abundant
and/or more active during the summer months. Psilochorus utahensis adults
were more uniformly abundant and active than P. imitatus throughout the
sampled period which was biased to the summer and fall, since only 2 months,
April and May, were sampled in the spring. Both of these species either
mature erratically, from spring through the fall, and/or adults live for long
periods of time, perhaps 2 or more years. In the laboratory, adult females
have lived for 6 months on ants. Further, as indicated by Muma (1975b),
populations of these spiders may be lower than the data indicate, since they
use the cans as web-building sites.
Zelotes tuobus Chamberlin, the dominant gnaphosid in the pinyon-juniper
associations, also occurred as an influent in the arid-grasslands. The spring-
summer peak of adults and/or adult activity on the pinyon-juniper plots did
not occur on the arid-grassland plots. A slight fall peak on the latter plots
indicated that the species matured earlier in the arid grasslands.
Schizocosa new species (near avida Walckenaer) was a dominant lycosid
on both pinyon-juniper plots with a summer peak of adult activity but was
absent from the arid-grasslands. On the other hand, Schizocosa mimula
Gertsch was an influent with a summer peak of adult activity only on the
Silver City pinyon-juniper plot. It occurred as an accessory species on the
Burro Mountain pinyon-juniper plot and the Hurley arid-grassland plot but
was absent from the Lordsburg arid-grassland.
Euryopis spinigerus (0. P. Cambridge) was an influent theridiid with a
summer peak of adult activity only on the Burro Mountain pinyon-juniper
plot. It was absent from all other plots.
The pinyon-juniper thomisid influent, Xysticus cunctator (Thorell), was
much more abundant on the Silver City plot than on the Burro Mountain
plot. Further the spring peak of adults and adult activity would have been
much more pronounced if March records could have been included with those
of April and May. The latter statement is true also for the arid-grassland
influent, Xysticus lassanus Chamberlin, which was an influent only on the
Lordsburg plot. Additionally, X. cunctator was a rare accessory species in
the arid-grassland, but X. lassanus was a relatively common accessory species
in the pinyon-juniper association.
Two pinyon-juniper influents, Drassyllus mormon Chamberlin and
Haplodrassus chamberlini Platnick and Shadab, also would have had higher
spring peaks of adult activity if March records could have been included.
However, the adult activity peaks of the pinyon-juniper influent, Herpyllus
propinquus Keyserling, and the arid-grasslands dominant, Drassyllus mumai
Gertsch and Reichert, would not have been affected by the inclusion of March


Florida Entomologist 64 (2)

records. There is no explanation for the fact that adults of H. propinquus
were relatively common for 9 months during the year.
One other aspect of the data is worthy of mention, since it seemed to vary
from year to year and plot to plot. That involved relative abundance of adult
dominants and influents. In the pinyon-juniper associations, the 3 dominants
were obviously and significantly more abundant than any of the 12 recorded
influents, even though each varied considerably from year to year and to
some extent, from plot to plot. However, the same was not true of the ob-
vious dominants and influents in the arid-grasslands. Although the 3
recorded dominants were more than twice as abundant as any of the 6 cited
influents during both years, the 2 less common dominants were not signif-
icantly more numerous than the more common influents during any year,
according to the statistic used. In fact, only 1 species, P. imitatus, was con-
sistently dominant, and only 1, Zelotes new species, consistently influent
during the test period. All other species were dominant during 1 year and
influent the other.
With the exceptions of reference specimens, unique, and primary types,
the specimens and species recorded here have been or will be deposited in the
Florida State Collection of Arthropods at Gainesville, FL 32602.

BARNES, R. D. 1953. The ecological distribution of spiders in non-forest
maritime communities at Beaufort, North Carolina. Ecol. Mono. 23:
CHEW, R. M. 1961. Ecology of the spiders of a desert community. New York
Ent. Soc. 69: 5-41.
COPPEL, H. C., AND R. B. SMYTHE. 1963. Occurrence of spiders on eastern
white pine trees infested with the introduced pine sawfly, Diprion
similis (Htg.). Univ. Wisconsin Forestry Res. Notes (102) : 1-7.
ELLIOTT, F. R. 1930. An ecological study of the spiders of the beech-maple
forest. Ohio J. Sci. 30 (1) : 1-22.
GIBSON, W. W. 1947. An ecological study of the spiders of a river-terrace
forest in western Tennessee. Ohio J. Sci. 47: 38-44.
LOWRIE, D. C. 1942. The ecology of the spiders of the xeric dunelands in the
Chicago area. Bull. Chicago Acad. Sci. 6(9) : 161-89.
1948. The ecological succession of spiders of the Chicago area dunes.
Ecology 29(3) : 334-51.
LUCZAK, J. 1960. Rozmieszczenie pietrowe pajakow w lesie. Ekol. Polska B.
6: 39-50.
MUMA, M. H. 1970. An improved can trap. Notes Arachnol. S. W. 1: 16-8.
1973. Comparison of ground surface spiders in four central Florida
ecosystems. Florida Ent. 56(3) : 173-96.
1974. Solpugid populations in southwestern New Mexico. Florida
Ent. 57(4) : 385-92.
1975a. Two vernal ground-surface arachnid populations in Tularosa
Basin, New Mexico. S. W. Nat. 20(1) : 55-67.
1975b. Long term can trapping for population analyses of ground-
surface, arid-land arachnids. Florida Ent. 58(4): 257-70.
1980. Comparison of ground-surface spider populations in pinyon-
juniper and arid-grassland associations in southwestern New Mexico.
Florida Ent. 63(1) : 211-22.
AND KATHARINE E. MUMA. 1949. Studies on a population of prairie
spiders. Ecology 30(4) : 485-503.


June, 1981

Muma: Ground-Surface Spiders

PECK, W. B. 1966. The population composition of a spider community in
west central Missouri. American Mid. Nat. 76(1) : 151-68.
TURNBULL, A. L. 1973. Ecology of the true spiders (Araneamorphae). Ann.
Rev. Ent. 18: 305-48.
WHITCOMB, W. H., H. EXLINE, AND M. HITE. 1963a. Comparison of spider
populations of ground stratum in an Arkansas pasture and adjacent
cultivated field. Proc. Arkansas Acad. Sci. 17: 1-6.
AND R. C. HUNTER. 1963b. Spiders of the Arkansas cotton
field. Ann. Ent. Soc. America 56(5) : 653-60.



The pallipes species-group of Eremobates Banks is reviewed following an
investigation of the stability of diagnostic characters utilized in distinguish-
ing species of the group and the discovery of new reliable specific characters.
Three new species are described: E. docolora from western Colorado, Utah,
Montana, and Wyoming; E. dentilis from southeastern Arizona; and E.
woodruffi from the Big Bend region of Texas. Eremobates arizonica (Roewer)
is resurrected from synonymy with E. pallipes (Say), and Eremothera
barber Muma from southern Texas is reassigned to this group of the genus

Se present una revision del grupo de species cerca de pallipes del
g6nero Eremobates Banks siguiendo la investigaci6n de la estabilidad de
caracteres diagn6sticos utilizados en distinguir las species del grupo y el
descubrimiento de nuevos caracteres especificos y consistentes. Se described
3 nuevas species: E. docolora del oeste de Colorado, Utah, Montana y Wy-
oming; E. dentilis del sureste de Arizona; y E. woodru.fi de la region de Big
Bend de Tejas. Se resucita E. arizonica (Roewer) de sinonimia con E.
pallipes (Say), y Eremothera barberi Muma del sur de Tejas se reasigna a
este grupo del genero Eremobates.

This review of the pallipes species-group of Eremobates Banks (1900) is
the result of an investigation of the diagnostic characters utilized in dis-
tinguishing members of this group of the genus.
The pallipes group was established by Muma (1951) for those species of

'Partially supported by NSF Grant No. 43566.
2Contribution No. 419, Bureau of Entomology, Division of Plant Industry, Florida Dept. of
Agriculture and Consumer Services, Gainesville, FL 32602 USA.
3Western New Mexico University, Office of Research.
4Science Teacher, Cherry Creek High School, Englewood, CO 80119 USA.
'Entomologist Emeritus, IFAS, Univ. of Florida, Gainesville, FL 32611 USA, Research
Associate, Bureau of Entomology, Div. of Plant Industry, Florida Dept. of Agriculture and
Consumer Services, Gainesville, FL 32602, and Western New Mexico University, Silver City,
NM 88061 USA.


Z84 Florida Entomologist 64(2) June, 1981

the genus Eremobates Banks with the following common characters; males
with a constriction but not distinct notch at the base of the fixed finger from
a dorsal view, a mesoventral groove dilated basally from a mesal view, and
the first post-stigmatic abdominal sternite with or without ctenidia. Fe-
males with roughly triangular genital opercula that have mesal margins
adjacent for the anterior third to half of their length but slightly to mod-
erately separated posteriorly. Both sexes with fondal teeth graded in size
When the group was erected by Muma, 6 species were recognized; E.
pallipes (Say), E. durangonus Roewer, E. californicus (Simon), E. dilatatus
(Putnam), E. putnami (Banks), and E. suspects Muma. E. pallipes was
designated the typical species of the group. Characters utilized in distin-
guishing these species included: for males, the presence or absence of a
palpal scopula, the number of papillae in the palpal scopula, width of the
fondal notch, presence or absence of abdominal ctenidia, number of ab-
dominal ctenidia, and color of the malleoli; for females, the form and size
of the caudal notch of the genital opercula, and color of the malleoli. Other
characters such as general coloration patterns, curvature of the male fixed
cheliceral finger, and the presence of denticules on the lower margin of the
male fixed finger were cited but not stressed.
After examination of solpugid types in European museums, Muma (1970)
maintained the same number of group species, distinguished by the above
characters but placed more stress on palpal coloration and curvature of the
male fixed cheliceral finger. Muma (1970) renamed as E. simoni Muma, the
species he had called californicus in 1951, after determining that the speci-
men in the type vial of the latter species was an unidentifiable immature.
Brookhart (1972) reported that previously identified Colorado specimens
showed intergrades among E. pallipes, E. durangonus, and E. suspects on
the basis of established diagnostic characters. He dropped durangonus from
the Colorado fauna on the bases of palpal coloration and curvature of the
male fixed cheliceral finger. After discussing character intergradation and
seasonal reproductive isolation of Colorado specimens of pallipes and
suspects, he questioned the validity of suspects.
In 1973 additional variations among Colorado specimens of the species-
group were noted, and a peculiar overlapping of the specific ranges of speci-
mens identified as pallipes and durangonus was discovered in southwestern
New Mexico. At that time, the 2 authors conferred on the problem and sub-
mitted a request to National Science Foundation for a grant to collect addi-
tional specimens, study known diagnostic characters, search for new diag-
nostic characters, and statistically test the numerical findings. The present
paper is the result of these studies.

Study specimens were obtained by recalling previously identified material
from major museums and collectors, by utilizing ecological and biological
study specimens collected by the authors, and by conducting collecting trips
into western Texas, southern Colorado, New Mexico, and eastern and south-
ern Arizona. Not all of the study specimens were included in the statistical


_1 _

Brookhart & Muma: Eremobates pallipes Group

Preliminary diagnoses were made using only material that had been col-
lected in the same geographic area at approximately the same time. Collec-
tions of this type were available from the San Simon valley near Portal,
Arizona; White River, Arizona; Craig, Colorado; Denver, Colorado; Peyton
Road near Colorado Springs, Colorado; southeastern Colorado from Pueblo
to Lamar; Wet Mountains in Colorado; and from the Deming, Lordsburg,
Silver City area in New Mexico (Fig. 1). Miscellaneous specimens from the
same geographic regions also were used, if they fitted an a posteriori assess-
ment of species suitability. This was done for the most part for areas in
central and southern Texas and to augment meager collections from Craig,
Colorado and White River, Arizona.
Quantitative data consisted of counts of abdominal ctenidia and palpal
papillae; measurements of male cheliceral fondal notches; of female oper-
cula; of the lengths of palpi, leg I, and leg IV; of the length and width of
propeltidia; and of the length and width of chelicerae. These latter measure-
ments were analyzed both alone and as ratios, i.e., cheliceral L/W, propel-
tidial L/W, and A/CP. The latter was obtained by summing the length of

S u ran- woodruffs


Fig. 1. Study areas and presently known ranges of 9 species of the pal-
lipes-species group of Eremobates Banks.


Florida Entomologist 64 (2)

the palpus, leg I and leg IV, and dividing by the length of the propeltidium
plus the length of a chelicera. The number was expressed as a ratio, i.e.
3.56:1 and 4.25:1.
The A/CP was established to demonstrate the relationship of appendages
(A) to body length (CP). As considered here, body length is the combined
lengths of the chelicerae and propeltidium excluding the abdomen which is
subject to size variability caused by the presence or absence of food, eggs, or
Analysis of variance (Sokal and Rohlf 1973) was used to determine those
computations that were statistically useful in this study. We considered dif-
ferences at or above the 95% level of confidence to be significant. In order to
check validity of samples collected from year to year or from month to
month, 2 sample groups, Portal, Arizona, 1963-64, and Deming, etc. New
Mexico, 1973-74 were tested for variance between years and between months
in terms of palpal papillae number and A/CP ratio.
Qualitative data consisted of notation of coloration of palpi, malleoli,
propeltidia, and legs; dentition of male and female chelicerae; location and
shape of the mesoventral groove and basal flange of the mesoventral groove
of the male fixed finger; and the shape, notch, and sclerotization of the
female opercula.

The A/CP ratio proved to be a reliable species indicator for both males
and females (Table 1).
Grouped males from Portal, Arizona and from Deming, New Mexico,
analyzed for variance of the A/CP ratio and the number of palpal papillae,
exhibited no significant difference either between years or between months,
indicating that the specimens could be considered members of the same popu-
lation. We then assumed this to be true of populations from other geographic
Males from different collection areas exhibited the following similarities
or differences at or above the 95% level of confidence for the same characters.
Those from Colorado were similar for all measurements and counts except
those from Laramie Basin, Craig, Colorado, which differed in both A/CP
ratio and number of abdominal ctenidia and those from the Peyton Road
area, which differed in number of palpal papillae from all other Colorado
specimens. All of the Colorado males differed from those from New Mexico
and Arizona. Males from Deming, Lordsburg, and Silver City, New Mexico
were all similar, but differed from those from the San Simon valley and
White River, Arizona. Males from the San Simon valley differed from those
from White River. Texas males from the 2 collection areas differed from
each other and from all other males in the A/CP. Comparative data from
both male and female A/CPs for the 10 major collection areas are cited in
Table 1.
Peyton Road, Colorado males complicated the analysis of palpal papillae
variation among populations. If the Peyton Road populations were included
in the total population, there was no significant difference between Colorado
and New Mexico populations for this character. However, if this population
was excluded, then a significant difference in both A/CP ratio and number
of papillae existed between the 2 populations.


June, 1981

Brookhart & Muma: Eremobates pallipes Group


Males Females
No. No.
Sample area spms. Mean SD spms. Mean SD

Laramie Basin 6 5.80 0.17 3 5.50 0.20
Denver 23 5.47 0.20 15 4.82 0.38
Peyton Road 21 5.45 0.49 10 4.75 0.22
Wet Mountains 10 5.66 0.51 14 4.61 0.18
Southern 22 5.74 0.30 16 5.01 0.25
New Mexico
Deming, Lordsburg,
Silver City Area 32 6.18 0.34 36 5.26 0.29
San Simon Valley 20 5.35 0.25 10 4.55 0.32
White River Area 6 6.00 0.26 2 5.11 0.15
Central 4 5.60 0.38 3 4.78 0.66
Southern 5 6.20 0.40 3 5.10 0.06

*Ratio of the combined length of the palpus, leg 1, and leg 4 to the combined lengths of
the chelicerae and propeltidium.

Careful examination of the mesoventral groove and basal flange of the
mesoventral groove of the male fixed finger revealed subtle but consistent
differences among populations examined. These differences were helpful in
separating and identifying specimens where aberrant patterns of dentition
or palpal papillae existed. The size of the mesal tooth on the movable finger
of the male was also useful when combined with other characters.
Likewise we were able to recognize a consistent pattern in the female
opercula that we had overlooked in previous studies. The opercula of each
species are shown in the species figures. This morphology was consistent for
all localities except for a population in northern Colorado near Denver where
nearly one-half of all females displayed female opercula similar to those of
E. arizonica (Roewer), although they were E. pallipes in all other respects.
These observations combined with statistical data and numerical counts
not requiring statistical treatment have resulted in the recognition of the
following diagnosed and illustrated group-species.


MALES: Fixed cheliceral finger straight or slightly curved upward with a
constriction but no distinct notch or spur at the base. Mesoventral groove of
fixed finger dilated basally into a cup-like structure. Basal flange of meso-

Florida Entomologist 64 (2)

ventral groove of fixed finger distinct but variable in length, width, and
pitch. Movable cheliceral finger with a large principal tooth, an anterior
tooth, 2 small intermediate teeth, and a mesal tooth. Palpi with or without a
scopula of papillae. First post-stigmatic abdominal sternite with or without
ctenidia. A/CP 5.45-6.20.
FEMALES: Fixed cheliceral finger with principal and medial teeth large,
an anterior tooth half as large as principal tooth, 2 intermediate teeth be-
tween principal and medial teeth, and 1 between medial and anterior teeth.
Movable cheliceral finger with a large principal tooth, an anterior tooth, 2
intermediate teeth, and a mesal tooth. Palpi usually without a scopula and
first post-stigmatic abdominal sternite without distinct ctenidia. Genital
opercula roughly triangular and adjacent anteriorly but slightly to mod-
erately separated posteriorly. A/CP 4.55-5.50.
Both sexes have the fondal teeth graded in size I, III, II, IV.
Diagnostic characters utilized in distinguishing species are presented in
Table 2.

Key to the pallipes species-group of Eremobates Banks

1. Pale species: propeltidia pale, at most, dusky marginally; legs
pale to slightly dusky; palpi pale, at most slightly dusky on
tarsi and apical ends of metatarsi. Males with cheliceral fondal
notch slightly longer than wide. Female opercula with median
notch obscure .----- ... ..-..-....--- ....--------........ .. .. ......------------------- .-... 2
1'. Dusky or dark species: propeltidia dusky or dark with a pale
median stripe; legs dusky with 1 or more segments decidedly
darker than metatarsi and/or tarsi; palpi dusky throughout
and/or dark on 2 or more segments. Males with fondal notch
wider than long or much longer than wide. Female opercula
with median notch small to large but distinct -....-... ..-----....- 5
2(1). Males with 2 ctenidia on first post-stigmatic abdominal sternite.
Females with mesal margins of the opercula sclerotized for
70% or more of their length. A western Colorado, Utah,
Montana, and Wyoming species -.--..-......---------- docolora new species
2'. Males without ctenidia. Females with mesal margins of the
opercula sclerotized for no more than 60% of their length .------....---- 3
3(2). Male mesoventral groove of fixed cheliceral finger distinctly
narrowed above first fondal teeth. Female opercular mesal mar-
gins sclerotized for 50-60% of their length and curved pos-
teriorly. An eastern Colorado, Nebraska, Kansas, Oklahoma,
N.W. Texas, eastern New Mexico, and eastern Wyoming spe-
cies .---.............------ ........ ....... .... pallipes (Say)
3'. Male mesoventrol groove broad throughout its length. Female
opercular mesal margins sclerotized for 40-50% of their length
and truncate posteriorly. New Mexico and Arizona species ---...--._4 4
4(3). Male fixed cheliceral finger with a distinct ventral tooth. Fe-
males unknown. A S.E. Arizona species -......- dentilis new species
4'. Male fixed cheliceral finger without teeth. Females diagnosed in
couplet 3. A western New Mexico and eastern Arizona species
.........---------------.--.- ----- arizonica (Roewer)


June, 1981

Brookhart & Muma: Eremobates pallipes Group


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Florida Entomologist 64 (2)

5(1). Dusky species: propeltidia lightly to distinctly dusky with pale
median stripe often indistinct; legs more distinctly dusky at
union of femora and tibia; palpi dusky throughout and slightly
to distinctly darker on metatarsi and tarsi. Males with chelic-
eral fondal notch slightly wider than long; fixed cheliceral
finger straight or lightly curved mesally; no abdominal cten-
idia. Females with median notch delineated by mesally curved
posterior margins ......-.....- ..........---..- ..............-- 6
5'. Dark species; propeltidia dark for all or most of length with
pale median stripe distinct; legs distinctly dusky or dark on 3
or more segments; palpi distinctly dusky or dark from apical
ends of femora through tarsi. Males with fondal notch vari-
able; tip of fixed cheliceral finger lightly curved ectally; 2 ab-
dominal ctenidia. Females with median notch wide but not well
delineated ---..~.-- -.. -- ....-- ---- ---------.... ............. ..-----------------. .. 9
6(5). Males with fixed cheliceral finger distinctly curved upward.
Females with median notch of the opercula occupying more
than 1/2 of the opercular length. A northcentral and south-
central Texas species -........-.~---....- . simoni Muma
6'. Males with fixed cheliceral finger straight or indistinctly curved
upward. Females with notch occupying less than /2 of the oper-
cular length --....... .. .. ....-...-.--.. -------------. ------------------- 7
7(6). Males with mesoventral groove of fixed finger narrowed for
most of its length. Females with median opercular notch small,
only 1 of the opercular width ---......- -. ...........-- 8
7'. Males with msoventral groove wide for most of its length. Fe-
males with median opercular notch moderate in size, about 1/3
the opercular width; an east-central Arizona species .
..-------- ------------------------- -------suspectus M um a
8(7). Males diagnosed in couplet 5, 6, 7. Females with median oper-
cular notch variable in form but not invading vulva. A south-
eastern Arizona and southwestern New Mexico species -
.- ---- ---.- ..-------------------- ..... durangonus Roewer
8'. Males unknown. Females with the mesal margins of the median
opercular notch apparently invading vulva. Distribution un-
known ...... ...........--- ........-------------------------- dilatatus (Putnam)
9(5). Males with fixed cheliceral finger bearing 3-5 tiny denticules.
Females diagnosed in couplet 5. A southern Texas species ...-
..---------....----...---... ------.. -......------------------ barber (M um a)
9'. Male fixed finger without denticules. Females unknown. Posi-
tively known only from type locality in Big Bend National
Park, Texas ...-----. ........-- ---.. -- ---.--... woodruffi, new species

Eremobates pallipes (Say)
(Fig. 2-14)

Galeodes pallipes Say, 1823, footnote on p. 3 ( 9).
Galeodes subulata Say, 1823, footnote on p. 3 ( ).
Eremobates pallipes, Fichter, 1940: 335, Fig. 1-4; Muma, 1951: 72, Fig. 106-
114; Muma, 1970: 22 ( and 9).


June, 1981

Brookhart & Muma: Eremobates pallipes Group

Fig. 2, 3. Eremobates pallipes (Say), $, scanning electron micrographs.
2. Fixed cheliceral finger showing mesoventral groove and apical plumose
seta, mesal view 60X. 3. Fixed cheliceral finger showing basal cup and mesal
flange of mesoventral groove, mesal view 120X.

Florida Entomologist 64(2)

TYPES: Say's specimens cannot be located in North American (Muma
1951) or European (Muma 1970) type depositories and are presumed lost.
We hereby designate the following: male neotype from Highway 205c, Byers,
Colorado, 17-VII-1973 by J. 0. Brookhart; female allotype from Castle Rock,
Colorado, 27-VI-1973 by J. O. Brookhart. Both are deposited in the American
Museum of Natural History, New York, N. Y.
DIAGNOSIS: Males and females pale yellow to light amber; palpi pale
except slightly dusky tarsal tips; other appendages pale (Fig. 7, 9-13).
Malleoli white. Males with mesoventral groove narrow, flared on the ventral
side, and a deep basal cup (Fig. 2, 3, 5, 6). Basal flange elongate forming a
shallow ventral hollow the entire length of finger. Mesal tooth medium. A/CP
5.56 -0.38, fondal notch (Fig. 4) L/W 1.3/1. Palpal papillae 55.6-40.5 -13.8-
18.1. Ctenidia absent (Fig. 8). Female genital opercula triangular with the
anterior margins slightly separated, somewhat bowed posteriorly, and the
posterior edge curved (Fig. 14). A/CP 4.84 0.37.
REMARKS: Species statistics were compiled from 4 populations ranging
from Denver, Colorado southward to Lamar, Colorado. There is more in-
consistency in this species than in any other member of the group. Occasion-
ally a male will have a ctenidia and a female will have a scopula. The spe-
cies may be highly variable or presently undergoing speciation. The popu-
lation from Peyton Road near Colorado Springs was variant in many re-
spects and may well represent a still unrecognized sibling species.
Muma's records (1951) probably are valid for all of Colorado, Kansas,
Oklahoma, Nebraska, North and South Dakota, and northern New Mexico
east of the Rocky Mountains. His records from Idaho and Utah probably
are E. docolora new species. The single record from Kirkwood, Missouri may
be an error in locality label.
Since this study has demonstrated that species of the pallipes group are
geographically isolated, the synonyms of pallipes suggested by Muma (1951,
1970) must be considered to be largely invalid. At the present time only
Galeodes subulata Say can be validly cited as a synonym of pallipes. On the
bases of type examinations and original descriptions the following are Mex-
ican species: Gluvia cinerascens L. L. Koch (1842), Gluvia formicarius
C. L. Koch (1842), Datames lentiginosus Kraepelin (1899, 1901), and
Eremostata dinamita Roewer (1934). Eremostata arizonica Roewer (1934),
and Eremostata californica Roewer (1934) are recognized here as separate
distinct species, but E. californica may well be the female of Eremothera
sculpturata Muma (see Muma 1962), so is not considered here.
STUDY SPECIMENS: Colorado: Arapahoe Co., 27 8, 14 9; El Paso Co.,
32 S, 17 9. Wet Mtns., 12 $, 14 9; Pueblo Co., 10 5 9; Otero Co., 9 3,
9 9; Powers Co., 2 8, 2 9. Boulder, 3 $, 1 9. Nebraska: Alliance, 1 ;
Dawson, 1 9; Chadron, 1 9 ; McCook, 1 9. Montana: Culbertson, 1 9. South
Dakota: Rapid City, 1 $. Texas: Palo Duro Canyon, 1 9; Colorado City,
Mitchell Co., 1 $ ; Salt Flats, 1 3. Wyoming: Cheyenne, 1 ; Douglas, 1 8.

Eremobates docolora Brookhart and Muma, NEW SPECIES
(Fig. 15-25)

TYPES: Male holotype from 1 mi. N. W. Craig, Colorado, Moffet Co., 26-
VI-1971 by J. O. Brookhart; female allotype from 8 mi. S. W. Encampment,


June, 1981

Brookhart & Muma: Eremobates pallipes Group


y /,ij

Fig. 4-14. Eremobates pallipes (Say). 4-13, $. 4. Right chelicera, ectal
view. 5. Left cheliceral fixed finger, mesal view. 6. Left cheliceral fixed finger,
dorsal view. 7. Propeltidium, dorsal view. 8. First post-stigmatic abdominal
sternite. 9. Palpus, anterolateral view. 10. Leg 1, anterolateral view. 11. Leg
2, anterolateral view. 12. Leg 3, anterolateral view. 13. Leg 4, anterolateral
view. 14. Female genital opercula, ventral view.


Florida Entomologist 64 (2)




Fig. 15-25. Eremobates docolora, new species. 15-24, 8. 15. Right chelicera,
ectal view. 16. Right cheliceral fixed finger, mesal view. 17. Right cheliceral
fixed finger, dorsal view. 18. Propeltidium, dorsal view. 19. First post-stig-
matic abdominal sternite. 20. Palpus, anterolateral view. 21. Leg 1, antero-
lateral view. 22. Leg 2, anterolateral view. 23. Leg 3, anterolateral view. 24.
Leg 4, anterolateral view. 25. Female genital opereula, ventral view.


June, 1981

Brookhart & Muma: Eremobates pallipes Group

Wyoming, Carbon Co., 5-VIII-1967 by F. P. & M. Rindge. These localities
may represent the easternmost extension of its range. Types are deposited in
the American Museum of Natural History, New York, N. Y.
DIAGNOSIS: Males and females pale yellow to ivory in coloration with
palpi and legs pale except for slightly dusky palpal tarsus and metatarsus
(Fig. 18, 20-24). Malleoli white. Male fixed finger as in E. pallipes with a
narrow mesoventral groove. The shallow ventral hollow formed by the basal
flange is traceable the entire length of the finger, (Fig. 16, 17), and is more
rounded than in pallipes. Mesal tooth small. A/CP 5.78 0.17. Fondal notch
(Fig. 15) L/W 1.2/1, palpal papillae 49-7.4; two short flat ctenidia (Fig.
19). Female genital opercula with straight, only slightly separated, interior
margins and an almost straight posterior edge (Fig. 25). A/CP 5.5 0.30.
REMARKS: This species is closely related to E. pallipes and E. arizonica
(Roewer). It is identified by pale coloration, 2 ctenidia on the male, and
shape of the female genital opercula. The specific name is an anagram of
the word Colorado. Canadian records of E. pallipes (Muma 1951) may well
be this species.
STUDY SPECIMENS: Colorado: Craig, 5 3. Montana: Baker, 1 9; Bose-
man, 1 9; Columbus, 1 9. Wyoming: Encampment, 1 9; Laramie, 1 3.

Eremobates arizonica (Roewer)
(Fig. 26-36)

Eremostata arizonica, Roewer, 1934: 572, Fig. 324w, 324e (9).
Eremobates pallipes (Say), Muma, 1951: 73. Muma, 1970: 34, Fig. 27 (mis-
TYPES: Female type (here designated holotype) of Eremostata arizonica
from Arizona, Roewer No. 8481, is deposited in Zoologisches Staatinstitut ad
Museum, Hamburg, West Germany. Male allotype from under cow dung,
Hurley, New Mexico, 17-VIII-1974, Martin H. Muma. The allotype is de-
posited, along with a typical female, in the American Museum of Natural
History, New York, N.Y.
DIAGNOSIS: Males and females yellow to dusky in coloration with lightly
dusky legs and palpi and a darker tip on the palpal tarsus (Fig. 29, 30-35);
ventral hollow shallow, rotated mesially and joining the mesoventral groove
just posterior to the tip. Mesal tooth medium. A/CP 6.8- 0.34. Fondal notch
(Fig. 26). L/W 1.2/1. Palpal papillae 48.1-19.9. No ctenidia (Fig. 30).
Female genital operculae with mesal margin straight and only slightly
separated, and with the posterior edge truncate, (Fig. 36). A/CP 5.26-0.29.
REMARKS: This species is closely related to E. pallipes.
STUDY SPECIMENS: Arizona: Holbrook, 2 3, 1 9. New Mexico: Hurley,
14 3, 12 9 ; Lordsburg, 5 3, 11 9; Jemez Springs, 2 3, 1 9 ; Elephant Butte
St. Park, 2 2 9; Newcombe, 1 3 ; Pleasanton, 1 3.

Eremobates dentilis Brookhart and Muma, NEW SPECIES
(Fig. 95-104)
TYPE: Male holotype yellow with anterior and posterior edges of pro-
peltidium dusky purple; palpal tarsus tinged purple; legs yellow (Fig. 98,
100-104) ; malleoli white. Mesoventral groove wide and deep, without a widely
flared ventral edge, (Fig. 96, 97). Ventral hollow shallow and rotated


Florida Entomologist 64(2)

June, 1981


-7 ^ 2eO





Fig. 26-36. Eremobates arizonica (Roewer). 26-35, 8. 26. Right chelicera,
ectal view. 27. Right cheliceral fixed finger, mesal view. 28. Right cheliceral
fixed finger, dorsal view. 29. Propeltidium, dorsal view. 30. First post-stig-
matic abdominal sternite. 31. Palpus, anterolateral view. 32. Leg 1, antero-
lateral view. 33. Leg 2, anterolateral view. 34. Leg 3, anterolateral view. 35.
Leg 4, anterolateral view. 36. Female genital opercula, ventral view.


Brookhart & Muma: Eremobates pallipes Group

mesially. Fixed finger slightly recurved with a conspicuous denticule located
above the principal tooth (Fig. 85). A smaller denticule is visible on the
fixed finger in the fond (Fig. 97). One large and 2 minute denticules are
located on the lower margin of the fond. Movable finger typical of the group.
Mesal tooth large and distinct. A/CP 5.33. Fondal notch L/W, 1/1.2; no
abdominal ctenidia, (Fig. 99). Palpal papillae 50-43.
REMARKS: The species is known only from the type and what may be an
immature female in the type vial. This species keys out with, and seems to
be more closely related to the pallipes lineage. It appears to be most closely
related to E. arizonica.

Eremobates durangonus Roewer
(Fig. 37-47)

Eremobates durangonus Roewer, 1934: 557, Fig. 323a, 324b ( 9).
Eremobates durangonus, Muma, 1951: 78, Fig. 119-23; Muma 1970: 22 (S
and 9).
TYPES: The 2 female types from Durango, Mexico cannot be located and
must be presumed to be lost (Muma 1970). However, we have no specimens
from Durango, so will refrain from designating a neotype until such are
DIAGNOSIS: Males and females yellow to dark amber in coloration with
propeltidia dark anteriorly, dusky laterally and light medially; palpi dark
on tarsus and dusky on the apical half to two thirds of metatarsus; legs pale
to lightly dusky (Fig. 40, 42-46). Malleoli white. Male mesoventral groove
deep without flared ventral edge as in E. pallipes (Fig. 38, 39); ventral hol-
low rotated mesially. Mesal tooth small to indistinct. A/CP 5.35 0.25. Fondal
notch (Fig. 37), L/W 1/1.1. Palpal papillae 22.5 9.1. Ctenidia absent (Fig.
41). Female genital opercula with separated sclerotized interior margins and
a small distinct notch or caudal indentation at the posterior edge (Fig. 47).
Posterior edges otherwise nearly straight (Fig. 47). A/CP 4.55 0.32.
REMARKS: This is a sonoran species possibly ranging from the type
locality in Durango, Mexico to its northern limits in southeast Arizona and
southwest New Mexico. It is closely related to E. suspects. We expect to
find other closely related species in Mexico.
It is probable that Muma's (1951) records of this species from California
and Texas are erroneous. The female from the White Mountains is probably
E. suspects. Muma's (1951) opercular variations (Fig. 121, 123) have not
been located.
STUDY SPECIMENS: Arizona: Portal, 23 $, 24 9 ; Rustler's Park, Chiri-
cahua Mtns., 1 ; Chiricahua Nat. Mon., 1 $; Sierra Vista, 2 $. New Mex-
ico: Rodeo, 16 9; Lordsburg, 1 9.

Eremobates suspects Muma
(Fig. 48-58)

Eremobates suspects Muma, 1951: 19: Muma, 1970: 25 ( S and 9).
TYPES: Male holotype and female allotype from White Mountains, 10
miles northeast of White River, Arizona, 8-11-VII-1940, Gertsch and Hook,
in the American Museum of Natural History, New York, N. Y.

Florida Entomologist 64(2)




k1' /
' "''I

Fig. 37-47. Eremobates durangonus Roewer. 37-46, &. 37. Right chelicera
ectal view. 38. Right cheliceral fixed finger, mesal view. 39. Right cheliceral
fixed finger, dorsal view. 40. Propeltidium, dorsal view. 41. First post-stig-
matic abdominal sternite. 42. Palpus, anterolateral view. 43. Leg 1, antero-
lateral view. 44. Leg 2, anterolateral view. 45. Leg 3, anterolateral view. 46.
Leg 4, anterolateral view. 47. Female genital opercula, ventral view.



June, 1981

Brookhart & Muma: Eremobates pallipes Group



0 OZ

r 53

a~i-l, \


Fig. 48-58. Eremobates suspects Muma. 48-57, 8. 48. Right chelicera,
ectal view. 49. Right cheliceral fixed finger, mesal view. 50. Right cheliceral
fixed finger, dorsal view. 51. Propeltidium, dorsal view. 52. First post-stig-
matic abdominal sternite. 53. Palpus, anterolateral view. 54. Leg 1, antero-
lateral view. 55. Leg 2, anterolateral view. 56. Leg 3, anterolateral view. 57.
Leg 4, anterolateral view. 58. Female genital opercula, ventral view.

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