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JANUARY & FEBRUARY, 1998

US ISSN 0013-872X
NO. 1

ENTOMOLOGICAL NEWS

Dispersal & range expansion of introduced sand wasp,

Oxybelus bipunctatus (Hymenoptera: Sphecidae), in

northeastern North America Frank E. Kurczewski 1

Relationship of cell depth & soil moisture in Oxybelus

bipunctatus (Hymenoptera: Sphecidae)

F.E. Kurczewski, D.L. Wochadlo 7
Adventive lady beetles (Coleoptera: Coccinellidae)

in eastern Nova Scotia, Canada D.B. McCorquodale 15

Camelobaetidius variabilis (Ephemeroptera: Baetidae), a

new species from Texas, Oklahoma and Mexico N.A. Wiersema 21
Newly reported and little known mayflies (Ephemerop-
tera) of Texas N.A. Wiersema 27
Redescription of Nerthra praecipua (Heteroptera:

Gelastocoridae) from Chile J.T. Polhemus, T. Cekalovic K. 33

A new species of Symphitoneuria (Trichoptera: Lepto-

ceridae) from Sabah, Malaysia T. Andersen, J. Huisman 37

A new species of Tomocerus (5.5.) (Collembola:

Tomocerinae) from China Y-T. Ma, K.A. Christiansen 47

Tomocerus spinulus, (Collembola: Entomobryidae), a

new species of Chinese springtail J- X. Chen, K.A. Christiansen 51
Species distinction in abdominal pigmentation patterns

between females of Drosophila melanogaster and D.

simiila us (Diptera: Drosophilidae), from a

Spanish population K. Th. Eisses, M. Santos 56

Synonymic notes on some of Thomson's New World

Dolichopodidae (Diptera) Daniel J. Bickel 61

Dry weight of fresh and preserved spiders (Ara-

neida: Labidognatha) R.L. Edwards, W.L. Gabriel 66

Observations of interactive behavior in Parandra

glabra (Coleoptera: Cerambycidae) Steven W. Lingafelter 75

SCIENTIFIC NOTES:
Euhrychiopsis lecontei (Coleoptera: Curculionidae), a new

state report for Indiana R.D. Waltz, G.M. White, R.W. Scriballo 6
Recent interception of live khapra beetle, Trogoderma

granarium (Coleoptera: Dermestidae), at Port of

Baltimore, MD M.L. Zimmerman, J.A. Barron 20

Unusual occurrence of caddisflies (Trichoptera:

Phryganeidae) in a Penn. population of purple

pitcher plant, Sarracenia purpurea

R. Hamilton IV, R.L. Petersen, R.M. Duffield 36

CURTIS W. SABROSKY 60

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Vol. 109, No. 1, January & February, 1998

DISPERSAL AND RANGE EXPANSION OF AN

INTRODUCED SAND WASP, OXYBELUS

BIPUNCTATUS (HYMENOPTERA: SPHECIDAE),

IN NORTHEASTERN NORTH AMERICA 1

Frank E. Kurczewski^

ABSTRACT: Range expansion of an introduced sphecid, Oxybelus bipunctatus, is traced through
the northeastern United States and southeastern Canada. This species apparently is displacing O.
uniglumis quadrinotatus, a species with similar behavior and ecology, in abundance as it moves
northward through the Great Lakes Region, St. Lawrence River Valley, and Maritime Provinces
to become the most common Oxybelus in small inland sandy areas. Its southward dispersal
probably is limited by climatic, ecological and edaphic factors such as high summer and warm
winter temperatures, thermal soil temperature regime, and absence of appropriate sandy soils.

Oxybelus bipunctatus Olivier is a small digger wasp that usually excavates
1 - and 2-celled nests in sandy soil, impales adult flies on its sting, removes the
temporary nest closure with the fly still impaled, and stocks its cells mostly
with several male Brachycera and/or Cyclorrapha (Peckham et al. 1973;
Krombein 1979; Kurczewski 1996). This wide ranging Holarctic species evi-
dently was accidentally introduced into the northeastern United States from
Europe before 1935 (Pate 1943; Krombein 1979), perhaps in ship ballast or
molding sand deposited dockside. Based upon insect museum specimens, the
earliest dates of its occurrence in North America are 1935 from Essex County,
New Jersey and Hampden County, Massachusetts, and 1936 from Suffolk
County, New York (Table 1). By the 1940's O. bipunctatus was firmly estab-
lished in the Middle Atlantic Region and New England (Pate 1945, pers. obs.).
Since that time the range of this species has been expanding primarily north-
ward, most recently into western upper Michigan, central Wisconsin, and Nova
Scotia.

Within 10 years of its introduction into the U. S., the range of 0. bipunctatus
extended to Washington, D. C., a distance of over 300 km from its first North
American collection locality (Fig. 1). In only 20 years' time, this species dis-
persed to North Bay, Ontario and central lower Michigan, a distance of 660-
900 km. In less than 50 years, O. bipunctatus moved as far west as western
upper Michigan and central Wisconsin, a distance of 1,250 km. On a smaller
scale, this species dispersed from Ottawa to North Bay, Ontario, probably across
sandy sections of the Ottawa-Mattawa River Valley,a distance of about 300 km,

1 Received March 21, 1997. Accepted June 29, 1997.

2 Environmental and Forest Biology, State University of New York College of Environmental
Science and Forestry, Syracuse, New York 13210-2778.

ENT. NEWS 109(1 ): 1-6, January & February, 1998

ENTOMOLOGICAL NEWS

in nine years (1947-1956). Oxybelus bipunctatus moved from Clinton County
to Ogemaw County, Michigan, a distance of about 1 70 km, in five years ( 1 954-
1959). Using these durations and distances as gages of dispersal, the rate of
movement of O. bipunctatus amounted to approximately 30-45 km per year
assuming the species was not transported to these regions by some means other
than its own flight. This is a remarkably rapid rate of dispersal for a species
limited to short flights by its small size and stocky build, although the possibil-
ity of wind carriage should not be overlooked.

In only a half century, this species dispersed through New England to the
Maritime Provinces (New Brunswick, Nova Scotia) and St. Lawrence River
Valley (southern Quebec), across the Great Lakes Region (upstate New York,
southern Ontario, northwestern Pennsylvania, northern Ohio, upper and lower
Michigan, central Wisconsin), and down the Atlantic Coastal Plain to Mary-
land, District of Columbia, and northern Virginia (Table 1 , Fig. 1 ). The lack of
specimens of O. bipunctatus from southern Ohio and southern Virginia indicate
that its southward dispersal may be climatically and ecologically limited by
factors such as high summer and warm winter temperatures and/or thermal soil
temperature regime (USDA 1984), or it may simply reflect inappropriate habi-
tat (soils) for this species in these areas.

Despite its adventive status, O. bipunctatus is one of the most common
sphecids in small inland sandy areas in the northeast (Kurczewski and Harris
1968; Kurczewski and Acciavatti 1990). O'Brien (1996 pers. comm.) found it
to be the most numerous species of Oxybelus in Malaise traps at the Huron
Mountain Club, Marquette County, Michigan beginning in the early 1980's (Fig.
1). McCorquodale (1997 pers. comm.) indicated that this species is more com-
mon than any other species of Oxybelus in sandy areas on Cape Breton Island,
Nova Scotia. Its abundance in these regions probably can be attributed to cold
hardiness, long flight season, relatively short generation time, wide prey selec-
tion (Peckham et al. 1973; Krombein 1979; Kurczewski 1996), scarcity of natural
enemies (Spofford and Kurczewski 1990), and ubiquity in nesting habitat.
Oxybelus bipunctatus nests opportunistically in such varied places as flower
pots, children's sand boxes, temporary highway sand piles, and the sand bases
of swimming pools. Because of its small size and capability to nest shallowly
in soil, it is an excellent candidate for transport by human means from one place
to another including transoceanic portage.

Although quantitative data are unavailable, O. bipunctatus apparently is
supplanting O. uniglumis quadrinotatus in abundance in sandy and gravelly
areas as it disperses northward through the Great Lakes Region, St. Lawrence
River Valley, and Maritime Provinces. The displacement in upstate New York
began rather suddenly during the 1950's (Evans 1960 pers. comm.). Oxybelus
bipunctatus appeared in the Ithaca, New York area as early as 1937 (Pate 1943),
possibly transported there by human means, and by the 1960's it was flourish-

Vol. 109, No. 1, January & February, 1998

Fig. 1. Regional dispersal of Oxybelus bipunctatus by year of collection. Arrows indicate prob-
able dispersal pathways.

ENTOMOLOGICAL NEWS

ing in small inland sandy areas of upstate New York (Kurczewski and Harris
1968; Peckham et al. 1973; Kurczewski and Acciavatti 1990). Oxybelus
bipunctatus is now the most abundant member of the genus in northeastern
United States and southeastern Canada (pers. obs.)- Some nesting aggregations

Table 1. Pre-1955 dates and localities of collection of Oxybelus bipunctatus arranged in chrono-
logical order.

DATE

LOCALITY

SOURCE

July 4, 1935

Bloomfield, Essex County, NJ

AMNH

July 29, 1935

Holland, Hampden County, MA

MCZ

June?, 1936

Half Hollows, Suffolk County, NY

Pate 1943

July 11, 1937

Reading, Middlesex County, MA

MCZ

August 2, 1937

Durham, Middlesex County, CT

MCZ

August 2, 1937

Ithaca, Tompkins County, NY

Pate 1943

August 31, 1939

Keamy, Hudson County, NJ

AMNH

July 2, 1944

Princeton, Mercer County, NJ

USNM

May 4, 1945

Stinson Lake, Grafton County, NH

July 23-25, 1945

Washington, DC

USNM

July 24, 1945

Washington, DC

Pate 1945

June 23, 1946

Princeton, Mercer County, NJ

USNM

[F12], 1947

Princeton, Mercer County, NJ

USNM

June 11 -July 15, 1947

Washington, DC

USNM, MSU

June 22-29, 1947

Arlington, Fairfax County, VA

USNM

June 27, 1947

Silver Springs, Montgomery County, MD

USNM, UCD

July 3, 1947

Ottawa, Municipality of Ottawa-Carleton, ONT

CNC

June 15, 1949

East Hartford, Hartford County, CT

July 14, 1951

Parke Reserve, Kamouraska County, QUE

CNC

July 22, 1952

Rensselaerville, Albany County, NY

USNM

June 7- August 20, 1953

Ithaca, Tompkins County, NY

MCZ, USNM

July 26, 1953

Guelph, Wellington County, ONT

June 25, 1954

Guelph, Wellington County, ONT

JulyS, 1954

Clinton County, MI

MSU

July 9, 1954

Orangeville, Dufferin County, ONT

Septembers, 1954

Ithaca, Tompkins County, NY

MCZ

Abbreviations for insect museums are as follows: AMNH, The American Museum of Natural
History; CNC, Canadian National Collection; CU, Cornell University; MCZ, Museum of Com-
parative Zoology, Harvard University; MSU, Michigan State University; UCD, University of
California-Davis;UG, University of Guelph; and, USNM, United States National Museum.

Vol. 109, No. 1, January & February, 1998

of this species number several hundred individuals (Peckham et al. 1973;
Kurczewski and Acciavatti 1990).

Why is O. bipunctatus displacing O. uniglumis quadrinotatus in the north-
eastern United States and southeastern Canada? Although the two species
seemingly have identical nesting behaviors and similar ecological requirements
(Peckham et al. 1973), O. uniglumis quadrinotatus is almost twice the size of
O. bipunctatus. Because of this size difference the two species might be ex-
pected to prey upon different families, genera and species of flies. However,
they capture some of the same groups of prey with O. uniglumis quadrinotatus
taking a slightly greater proportion of Cyclorrapha and fewer Brachycera than
O. bipunctatus. Within the suborder Cyclorrapha, there is some predation on
the same species of flies (Peckham et al. 1973; Krombein 1979). Oxybelus
uniglumis quadrinotatus never captures Nematocera while O. bipunctatus oc-
casionally preys upon mosquitos, midges, and blackflies (Evans 1963;
Kurczewski 1996). A broader spectrum of prey, smaller size and inconspicu-
ousness, potentially fewer predators, and a lower rate of cleptoparasitism
(Spofford and Kurczewski 1990) are some reasons why O. bipunctatus is more
successful and, therefore, more numerous than O. uniglumis quadrinotatus in
many sandy areas of the northeastern U. S. and southeastern Canada.

ACKNOWLEDGMENTS

The following persons provided specimens, collection data, and/or abundance information
for O. bipunctatus: Bert Finnamore, Alberta Provincial Museum; Rick Hoebeke, Cornell Univer-
sity; Michael Kelly, Museum of Comparative Zoology, Harvard University; Lynn Kimsey, Uni-
versity of California-Davis; Steve Krauth, University of Wisconsin-Madison; Lubomir Masner,
Canadian National Collection; David McCorquodale, Guelph University, Nova Scotia Natural
History Museum, and University College of Cape Breton; Ron McGinley and Arnold Menke,
United States National Museum; Mark O'Brien, Museum of Zoology, The University of Michi-
gan; Mark Snethurst, The American Museum of Natural History; Andrey Sharkov, The Ohio
State University; Jeff Skevington, Pinery Provincial Park; Don Snitgen, Northern Michigan Uni-
versity; and John Wilterding, Michigan State University. I am grateful to David McCorquodale
and two anonymous reviewers for critically reviewing the manuscript. Julie Cushine-Rigg final-
ized Figure 1 .

LITERATURE CITED

Evans, H. E. 1963. Wasp Farm. Natural History Press, Garden City, N. Y. 178pp.
Krombein, K. V. 1979. Superfamily Sphecoidea. In: K. V. Krombein, P. D. Hurd, Jr., D. R.

Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America North of Mexico, vol 2,

pp. 1573-1740. Smithson. Inst. Press, Washington, D. C.
Kurczewski, F. E. 1996. Variation in nesting behavior of Oxybelus bipunctatus and O.

emarginatus, with notes on other species (Hymenoptera: Sphecidae). Mem. Entomol. Soc.

Wash. Contrib. Hym. , pp. 82-88.
Kurczewski, F. E. and R. E. Acciavatti. 1990. Late summer-fall solitary wasp fauna of central

New York (Hymenoptera: Tiphiidae, Pompilidae, Sphecidae). Great Lakes Entomol. 23:

57-64.

ENTOMOLOGICAL NEWS

Kurczewski, F. E. and B. J. Harris. 1968. The relative abundance of two digger wasps, Oxybelus
bipunctatus and Tachysphex terminatus, and their associates, in a sand pit in central New
York. J. N. Y. Entomol. Soc. 76: 81-83.

Pate,V. S. L. 1943. On some Holarctic sphecoid wasps (Aculeata, Hymenoptera). Bull. Brook-
lyn Entomol. Soc. 38: 14-16.

Pate, V. S. L. 1945. On two species of Oxybelus at Washington, D. C. (Hymenoptera, Sphecidae).
Bull. Brooklyn Entomol. Soc. 40: 165.

Peckham, D. J., F. E. Kurczewski, and D. B. Peckham. 1973. Nesting behavior of Nearctic
species of Oxybelus (Hymenoptera: Sphecidae). Ann. Entomol. Soc. Amer. 66: 647-661.

Spofford, M. G. and F. E. Kurczewski. 1990. Comparative larvipositional behaviours and
cleptoparasitic frequencies of Nearctic species of Miltogrammini (Diptera: Sarcophagidae).
J. Nat. Hist. 24: 731-755.

United States Department of Agriculture. 1984. General soil map of the northeastern United
States.

SCIENTIFIC NOTE

EUHRYCHIOPSIS LECONTEI

(COLEOPTERA: CURCULIONIDAE):

A NEW STATE REPORT FOR INDIANA 1

R.D. Waltz 2 , G.M. White 3 , R.W. Scribailo 4

The range of the native North American aquatic weevil species Euhrychiopsis lecontei (Dietz)
is poorly documented (Sheldon and O'Bryan 1996) although it has been reported from coast to
coast in various northern and central states including Iowa, Michigan, Wisconsin, Alberta, Brit-
ish Columbia, and Saskatchewan (O'Brien and Wibmer 1982), Washington (Creed and Sheldon
1994), Minnesota (Newman and Maher 1995), and Connecticut, Illinois, Massachusetts, New
York and Vermont (Sheldon and O'Bryan 1996). Because this species is recognized as a poten-
tially effective endemic biological control agent of watermilfoils (Haloragaceae: Myriophyllum
spp.) (Creed and Sheldon 1993; Sheldon and O'Bryan 1996), records of its distribution are of
special interest to those managing lakes and to those who have an interest in the documentation
of insect faunae within regional or politically delineated contexts.

Euhrychiopsis lecontei has not been reported previously from Indiana (above citations;
Blatchley and Leng 1916; Downie and Amett 1996). Herein, we report the first known record of
this species in Indiana. Voucher specimens were taken from: In: LaPorte Co., Saugany Lake,
May 16, 1997, G.M. White, collected in association with Eurasian watermilfoil (Myriophyllum
spicatum L.). Voucher specimens have been deposited at Purdue University Entomological Re-
search Collections, West Lafayette, Indiana.

Adults were collected and numerous eggs were observed on the stem apices of Eurasian

(continued on page 14)

1 Received June 7, 1997. Accepted July 4, 1997.

2 IDNR, Division of Entomology and Plant Pathology, 402 West Washington, Room W-290,
Indianapolis, IN 46204.

3 IDNR, Division of Soil Conservation, 402 West Washington, Room W-265, Indianapolis, IN
46204.

4 Biological Sciences and Chemistry Section, Purdue University - North Central, 1401 South
Hwy US 421, Westville, IN 46391.

Vol. 109, No. 1, January & February, 1998

RELATIONSHIP OF CELL DEPTH AND SOIL

MOISTURE IN OXYBELUS BIPUNCTATUS

(HYMENOPTERA: SPHECIDAE) 1

Frank E. Kurczewski, Donna L. Wochadlo^

ABSTRACT: Nests of Oxybelus bipunctatus were excavated and cell depth measured in control
and experimentally watered plots in a recently bulldozed sandy field in central New York in June
1987. There was a trend for cell depth to be inversely related to soil moisture under natural condi-
tions and in artificially watered replicates.

Soil moisture content is extremely critical for the survival of the immature
stages of ground inhabiting solitary wasps. Too little moisture in the soil results
in dessication of the paralyzed prey in the cells and the eventual death of the
wasp egg or larva. Too much soil moisture increases the likelihood of mold
development in the cell and this, too, can destroy the cell contents and wasp egg
or larva. A delicate balance in the amount of soil moisture is necessary for the
immature stages of the wasps to develop properly. Many physical and biotic
factors influence soil moisture content and regulate cell environment. Cell depth
and amount of precipitation are two such factors that can be readily measured
without elaborate or expensive equipment. The present study is an attempt to
investigate the relationship between the amount of rainfall, depth of soil mois-
ture line, and cell depth in a ground-nesting sphecid, Oxybelus bipunctatus
Olivier.

Oxybelus bipunctatus, a species adventive from Europe (Pate 1943, 1945;
Krombein 1979; Kurczewski 1998), is one of the most common digger wasps
in the northeastern U. S. (Kurczewski and Harris 1968; Kurczewski and
Acciavatti 1990). Ubiquity in nesting habitat, a long and continuous flight pe-
riod involving two or three generations per year, and high nest density make
this species an excellent subject for studying nesting behavior in a solitary wasp.
Peckham et al. (1973) investigated the behavior of O. bipunctatus over three
years. Although they carefully measured nest dimensions, these authors did
not examine the relationship between cell depth and soil moisture content.
Kurczewski ( 1 996) noted the difference in cell depth of O. bipunctatus between
localities in central New York and northwestern Pennsylvania but he, too, did
not account for soil moisture level. In a 15 weeks-long investigation of O.
bipunctatus at a site in upstate New York, the same author (in prep.) observed
weekly fluctuation in cell depth that seemed to be linked to soil moisture level

1 Received March 20, 1997. Accepted June 27, 1997.

2 Environmental and Forest Biology, State University of New York College of Environmental
Science and Forestry, Syracuse, New York 13210-2778.

ENT. NEWS 109(1): 7-14, January & February, 1998

ENTOMOLOGICAL NEWS

as influenced by the amount of rainfall. The present study is an extension of
this finding. Using measurements of rainfall, soil moisture line, and cell depth,
we tried to corroborate Kurczewski's (in prep.) results. By artificially adding
water to designated sand plots, we attempted to demonstrate a relationship be-
tween soil moisture content and cell depth.

MATERIALS AND METHODS

A recently bulldozed sandy field in northwestern Madison County, New
York, 5.6 km NW of Chittenango, a site used previously by Spofford et al.
(1986) for a study on the cleptoparasites of another sphecid, Tachysphex
terminatus (Smith), was selected as the study area. Natural conditions were
monitored during 5-15 June 1987. This involved collecting and measuring the
amount of rainfall from the previous day(s) and/or night(s) in mm, taking air
(shade), sand surface and cell depth temperature in degrees C, and noting any
change or disturbance to the area. Morning activity of individual wasps was
correlated with air and cell depth temperature.

Females were observed digging burrows in the morning on 5, 6, 10, 1 1 , and
14 June 1987, and their nest entrances were marked with color-coded tooth-
picks after completion of the temporary nest closures. (The wasps themselves
were too small to color-code).

The cells were excavated the following morning or afternoon after the wasps
completed their final closures, measurements were taken, and new nests were
found and marked for the next day, weather permitting.

The relationship between cell depth and soil moisture line, a readily visible
band of dark moist sand underlying a layer of light colored dry sand (Kurczewski
in prep.), was examined for 35 of the cells. Depths of soil moisure line and cell
to its bottom were measured with a metric ruler and recorded in mm. Depths of
53 other cells were also measured and recorded. Each excavation was then
levelled flush with the surrounding sand and lightly packed down to encourage
renesting.

Two areas of bare sand of equal size were selected and each area divided
into control and experimental plots (Fig. 1). The selection of these areas was
based upon local topography, lack of vegetation, and density of nesting wasps.
One area was divided in half by a north-south line into 240 X 280 cm plots and
labelled NC (North Control) and NE (North Experimental). The other area was
divided in half by an east-west line into 480 X 140 cm plots and labelled SC
(South Control) and SE (South Experimental). The direction of division into
control and experimental plots coincided with slight slopes in the respective
areas and was an attempt to eliminate any variables.

On June 16, we artificially watered the experimental plots to achieve a soil
moisture level sufficiently higher than that in the control plots yet not inhibit

Vol. 109, No. 1, January & February, 1998

wasp excavation or create soil surface run-off. Using a 1 .5 gal sprinkling can,
we evenly applied 10 gallons of water to each experimental area in rows 20 cm
wide beginning at 0920 hrs (EOT). Prior to watering, five 50 ml plastic gradu-
ated cylinders were set into the sand in each experimental plot, four of them
each 30 cm in from a corner and one directly in the center of the plot (Fig. 1 ).

240cm

12 9 8 7 6

13 10

15 1
5 14

9 6' 27 n a*.

, ] 3 5 Q\

6 2 8 ^>\

SE o

o
o

480cm

Fig. 1 . Design of control (NC, SC) and experimental (NE, SE) plots in Oxybelus bipunctatus study
area. Circles represent graduated cylinders set into sand to collect water. Numbers in NE and NC
are locations of nest entrances sequentially marked on 24 June 1987.

ENTOMOLOGICAL NEWS

The relative uniformity of our watering technique was reflected in the amount
of water collected by the graduated cylinders: (NE), 4.0, 4.5, 6.0 (central), 5.0,
5.0 ml; and, (SE), 5.0, 4.0, 4.0 (central), 4.5, 5.0 ml.

On 24 June, we applied 20 gal of water to the NE plot utilizing the same
technique, except the application was made between 2000-2100 hrs. We felt
that an application at dusk would substantially decrease water evaporation from
the sand surface and, by doubling the amount of water put into the soil, the
wasps might dig significantly shallower nests in this plot. After the application,
each of the five plastic graduated cylinders placed in the plot contained 7.5-8.0
ml of water. We intended to similarly water the SE plot the same evening, but
it had been severely disturbed the previous night by a truck carrying a bulldozer
and was abandoned for further use.

RESULTS

Temperature and rainfall. Only five of the 1 1 days (see above) set aside
to study O. bipunctatus under natural conditions were suitable for wasp nest-
ing. During these days, air temperatures averaged 22.8 C (18-25 C, n=5) and
soil temperatures at cell depth averaged 26.7 C (23-31 C, n=5) at 1100 hrs
(EOT). The six other days set aside for study under natural conditions were
non-conducive for wasp nesting because of cool temperatures, cloud cover, and
rain. It rained sporadically in three and heavily during two of the days (Fig. 2).

50
45
40
35

E 30

I 25
S. 20

Rainfall

D Mean cell depth

10
Day (June 1987)

50
49
48

47 ?
46 ~

45 "S

Fig. 2. Amount of rainfall and mean cell depth in Oxybelus bipunctatus study area, 5- 1 4 June
Amount of rainfall is a measurement from previous day(s) and/or night(s).

44
43
42
41
40

1987.

Vol. 109, No. 1, January & February, 1998

The average air and mean cell depth temperatures on these mornings at 1100
hrs (EOT) were only 17.2 C (15-18 C, n=6) and 21.1 C (18-22 C, n=6),
respectively.

Soil moisture line. The soil moisture line was indistinguishable from the
overlying sand on days when it rained heavily. On non-rainy days, this line was
clearly visible and cells were invariably excavated below it in moist sand. There
was an imperfect positive correlation between soil moisture line and cell depth
(Fig. 3). In some nests, the soil moisture line to cell depth value was more
disparate than in other nests (Fig. 3).

Cell depth. Females excavated fewer and deeper nests in drier soil. They
dug clumped and shallower nests in moister soil. Nests excavated in neither
dry nor wet sand were intermediate in depth and more evenly distributed than
nests in wet or dry sand (Figs. 1, 2). Mean cell depth and amount of rainfall
from the previous day(s) and/or night(s) were inversely related (Fig. 2). In
early to mid-June, slight increases or decreases in mean cell depth followed
corresponding decreases or increases, respectively, in the amount of rainfall
(Fig. 2). For example, shallowness in mean cell depth on 10, 11 and 14 June
was probably linked to a substantial amount of rainfall and increased soil mois-
ture content on 10-11 June. Nests excavated during 5-14 June under natural
conditions had, on average, shallower cells (x=44.45.38 mm; n=88; Fig. 2),
albeit not significantly so, than control plot nests dug on 1 6-24 June (x=47.36.29
mm; n=67; Table 1). This difference probably was due to the different amounts

Table 1 . Cell depth of Oxybelus bipunctatus, control and experimental plots, 16-24 June 1987.

Site Information

Cell Depth (mm)

Date

Plot

Range

Mean

6/16/87

38-59

46.6+5.99

6/16/87

38-58

47.8+6.23

6/17/87

38-56

45.8+5.54

6/17/87

26-45

37.8+8.16

6/17/87

35-55

48.1+5.79

6/17/87

37-65

45.2+8.22

6/24/87

34-63

48.2+7.88

6/24/87

39-61

47.6+6.77

ENTOMOLOGICAL NEWS

(uiui)

10
CO

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:X-

O V/

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(LULU) au;~| ajnjsio|/yj |;os

O
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CNJ

LO ^

- LO

Si ~

cfl 4>

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00 "

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;*
?

* y

*- C

S Si
c ^

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11

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Fig. 3. Di
surface, X

Vol. 109, No. 1 , January & February, 1998 13

of rainfall during these times of the month (5-14 June, 54.5 mm; 16-24 June,
45.7 mm).

Mean cell depth for 16 June NC and SC nests was similar (t=0.0793, df=18,
cc>0.20) (Table 1). On 17 June, wasps excavated significantly deeper cells in
SC than in SE (t=3.0415, df=25; oxO.Ol) (Table 1). There was no significant
difference in mean cell depth between two samples each comparing NC and
NE nests (June 17, t=l .0572, df=24, a>0.20; June 24, t=0.0397, df=25, a>0.20),
although the wasps did excavate slightly shallower cells in both NE plots (Table
1). All nests (n=12) in NE excavated on 24 June were clustered downslope in
the SW corner in moist sand (Fig. 1).

DISCUSSION

Few studies on fossorial sphecid wasps considered the influence of relative
soil moisture on cell depth. Moisture gradient influenced cell depth throughout
the nesting season in Microbembex nigrifrons (Provancher). Deeper cells were
dug in drier and shallower cells in wetter sand (Alcock and Ryan 1973). Soil
moisture level was related to cell depth in Ammophila harti (Fernald). Cells
excavated in drier sand remained deeper than those dug in wetter sand regard-
less of generation of wasp (Hager and Kurczewski 1986). Cell depth and, less
so, soil moisture line closely followed an increase or decrease in relative soil
moisture, as governed by the amount of rainfall, throughout the nesting season
in Oxybelus bipunctatus at one locality in central New York (Kurczewski in
prep.).

In the present study, both soil moisture line and cell depth were inversely
related to the amount of rainfall. There was, however, an imperfect positive
correlation between soil moisture line and cell depth with some values being
more disparate than others, even in perfectly flat sand. Regardless of possible
flaws in the experimental design, shallower cells were usually excavated in
moister and deeper cells in drier sand. However, only one control versus experi-
mental plot sample was conclusive. In this comparison, SE cells in artificially
watered sand were excavated at significantly shallower depths than SC cells.

ACKNOWLEDGMENTS

We thank M. G. Spofford for assisting with field work on the last day of the study. We are
grateful to D. J. Peckham for reading the manuscript critically. Use of a college vehicle was
made possible through an NYS-UUP Faculty Development Award to the senior author.

LITERATURE CITED

Alcock, J. and A. F. Ryan. 1973. The behavior of Microbembex nigrifrons (Hymenoptera:

Sphecidae). Pan-Pac. Entomol. 49: 144-148.
Hager, B. J. and F. E. Kurczewski. 1986. Nesting behavior of Ammophila Hani (Fernald)

(Hymenoptera: Sphecidae). Amer. Midi. Nat. 116: 7-24.
Krombein, K. V. 1979. Superfamily Sphecoidea. In: K. V. Krombein, P. D. Hurd, Jr., D. R. Smith,

and B. D. Burks, eds. Catalog of Hymenoptera in America North of Mexico, vol. 2, pp. 1 573-

1740. Smithson. Inst. Press, Washington, D. C.

14 ENTOMOLOGICAL NEWS

Kurczewski, F. E. 1 996. Variation in nesting behavior of Oxybelus bipunctatus and O. emarginatus,

with notes on other species (Hymenoptera: Sphecidae). Mem. Entomol. Soc. Wash. Contrib.

Hymen, pp. 82-88.
Kurczewski, F. E. 1998. Dispersal and range expansion of an introduced sand wasp, Oxybelus

bipunctatus, in northeastern North America (Hymenoptera: Sphecidae). Entomol. News 109:

1-6.
Kurczewski, F. E. and R. E. Acciavatti. 1990. Late summer- fall solitary wasp fauna of central

New York (Hymenoptera: TiphiidaePompilidae, Sphecidae).Great Lakes Entomol. 23: 57-64.
Kurczewski, F. E. and B. J. Harris. 1968. The relative abundance of two digger wasps, Oxybelus

bipunctatus and Tachysphex terminatus, and their associates, in a sand pit in central New York.

J. N. Y. Entomol. Soc. 76: 81-83.
Pate, V. S. L. 1 943. On some Holarctic sphecoid wasps (Aculeata, Hymenoptera). Bull. Brooklyn

Entomol. Soc. 38: 14-16.
Pate, V. S. L. 1945. On two species of Oxybelus at Washington, D. C. (Hymenoptera, Sphecidae).

Bull. Brooklyn Entomol. Soc. 40: 165.
Peckham, D. J., F. E. Kurczewski, and D. B. Peckham. 1973. Nesting behavior of Nearctic

species of Oxybelus (Hymenoptera: Sphecidae). Ann. Entomol. Soc. Amer. 66: 647-661.
SpofTord, M. G., F. E. Kurczewski, and D. J. Peckham. 1986. Cleptoparasitism of Tachysphex

terminatus (Hymenoptera: Sphecidae) by three species of Miltogrammini (Diptera: Sarcopha-

gidae). Ann. Entomol. Soc. Amer. 79: 350-358.

(continued from page 6)

watermilfoil. Based on the observation of adult activity in May and the presence of numerous
eggs, we believe E. lecontei is established in Indiana and we anticipate that the biology of E.
lecontei in Indiana will be very similar to that reported by Sheldon and O'Bryan (1996).

ACKNOWLEDGMENTS

We thank C.W. O'Brien and Peter Kovarik, Florida A&M, for providing comparative mate-
rial to confirm our identification of the species, and thank David Barnes, Mitchell Alix, and Bob
Jankowski, Purdue University North Central, for assisting in making collections of this species.

LITERATURE CITED

Blatchley, W.S. and C.W. Leng. 1916. Rhynchophora or Weevils of North Eastern America.
The Nature Publ. Co., Indianapolis, IN. 682 pp., illus.

Creed, R.P., Jr. and S.P. Sheldon. 1993. The effect of feeding by a North American weevil,
v Euhrychiopsis lecontei, on Eurasian watermilfoil (Myriophyllum spicatum). Aquat. Bot. 45:
245-256.

Creed, R.P., Jr. and S.P. Sheldon. 1994. Aquatic weevils (Coleoptera: Curculionidae) associ-
ated with northern watermilfoil (Myriophyllum sibiricum) in Alberta Canada. Entomol. News
105:98-102.

Downie, N.M. and R.H. Arnett. 1996. The Beetles of Northeastern North America, Volume II.
Sandhill Crane Press, Gainesville. 1721 pp.

Newman, R.M. and L.M. Maher. 1995. New records and distribution of aquatic insect herbi-
vores of watermilfoils (Haloragaceae: Myriophyllum spp.) in Minnesota. Entomol. News 106:
6-12.

O'Brien, C.W. and G.J. Wibmer. 1982. Annotated checklist of the weevils (Curculionidae
sensu lato) of North America, Central America, and the West Indies. Mem. Am. Entomol.
Inst. #34.

Sheldon, S.P. and L.M. O'Bryan. 1996. Life history of the weevil Euhrychiopsis lecontei, a
potential biological control agent of Eurasian watermilfoil. Entomol. News 107: 16-22.

Vol. 1 09, No. 1 , January & February, 1 998 15

ADVENTIVE LADY BEETLES

(COLEOPTERA: COCCINELLIDAE)

IN EASTERN NOVA SCOTIA, CANADA 1

D. B. McCorquodale 2

ABSTRACT: Specimens in three small insect collections document the arrival of four species of
adventive lady beetles in Nova Scotia. All four have experienced dramatic range expansions in
eastern North America in the past 25 years. Coccinella septempunctata arrived by 1984 and by
the 1990's was the most common coccinellid in disturbed habitats on Cape Breton Island, NS.
The more recent arrivals in Nova Scotia of Propylea quaiuordecimpunctata (1990), Harmonia
axyridis (1995) and Hippodamia variegata (1996) are reported.

Several species of lady beetles have become established in North America
after either accidental or intentional introductions in the past 25 years (Gordon
1985; Gordon and Vandenberg 1991). Populations of some of these intro-
duced species have burgeoned to the extent they have the potential to change
dramatically populations of both native and other introduced lady beetles (e.g.
Tedders and Schaefer 1994; Wheeler and Hoebeke 1995). Coccinellids de-
serve attention so that we can document which introduced species become
established, the speed of range expansion and major changes in populations.

Recently, Hoebeke and Wheeler (1996) recorded the range expansion of
adventive lady beetles in the Maritime provinces of Canada through the sum-
mer of 1995. Here I provide updated information from eastern Nova Scotia on
three of the four species they reported, Propylea qualuordecimpunciala,
Hippodamia variegata and Harmonia axyridis, and information on a species
established earlier, Coccinella septempunctata.

METHODS

All specimens of Coccinellidae in the small insect collections at the Uni-
versity College of Cape Breton, Sydney, Nova Scotia (UCCB), St. Francis
Xavier University, Antigonish, Nova Scotia (STFXU), and the Nova Scotia
Department of Natural Resources, Belmont, Nova Scotia (NSDNR) were ex-
amined. Identifications were based on Gordon ( 1 985), Gordon and Vandenberg
(1991), and comparison with specimens in the University of Guelph Collec-
tion, Guelph, Ontario. Representative specimens will be deposited in the Ca-
nadian National Collection, Ottawa, Ontario and/or the Nova Scotia Museum
of Natural History, Halifax, Nova Scotia.

1 Received June 6, 1997. Accepted July 19, 1997.

2 Department of Behavioural and Life Sciences, University College of Cape Breton, Sydney,
Nova Scotia B1P6L2.

ENT. NEWS 109(1): 15-20, January & February, 1998

16 ENTOMOLOGICAL NEWS

RESULTS

Coccinella septempunctata L.: Since 1973 this species has spread over
most of the continental United States and southern Canada (Gordon and
Vandenberg 1991). Schaefer et al. (1987) reported it in Nova Scotia in 1985.
Collections from mainland Nova Scotia in 1984 and Cape Breton Island in
1985 demonstrate that the species was widespread in the province by 1985.

Additional Locality Records:

Cumberland County, Chignecto, 5 June 1984, E. Georgeson, NSDNR; Cape Breton County,
Sydney, 26 August 1985, S. Tynski, UCCB; Cape Breton County, Sydney, 16 May 1986, S.
Tynski, UCCB

Since 1990, students at the University College of Cape Breton have col-
lected 171 specimens in the subfamily Coccinellinae. Collections are largely
from disturbed or anthropogenic habitats in the Sydney-Glace Bay area of
Cape Breton Island. About half of these specimens are Coccinella
septempunctata (86 or 50%), while native species are the next four most fre-
quently collected: Adalia bipunctata (24, 1 4%), C. trifasciata (19,11%), Anatis
mail (10, 6%) and Psyllobora vigintiduopunctata (9, 5%).

Propylea quatuordecimpunctata (L): This species was established in Que-
bec by 1968, had spread to Maine by 1988 and since then to much of the
northeastern United States (Day et al. 1994). There is a strong probability that
an independent establishment occurred in lower New York State in the mid-
late 1980's (Yanega 1996). Hoebeke and Wheeler (1996) provide the first
report for Nova Scotia and record collections from Cape Breton Island in 1995.
Collections in the summer of 1996 from Cape Breton and Antigonish Counties
indicate that it is now established in eastern Nova Scotia.

Additional Locality Records:

Cape Breton County, Georges River, 8 June 1996, DB McCorquodale, UCCB; Cape Breton
County, Sydney, UCCB Campus, 4 September 1996, S. Berrigan, UCCB; Cape Breton County,
Sydney, 5 September 1996, T. Rideout, UCCB; Antigonish County, Pomquet, 05 June 1996, R.
Lauff, STFXU; Antigonish County, Pomquet, 22 July 1996, R. Lauff, STFXU; Antigonish
County, Pomquet, 21 August 1996, R. Lauff, STFXU; Antigonish County, Antigonish, 15 July
1996, R. Lauff, STFXU

Propylea quatuordecimpunctata was established in Nova Scotia for sev-
eral years prior to the first collections reported by Hoebeke and Wheeler ( 1 996).
The Nova Scotia Department of Natural Resources collection has 35 speci-
mens collected between 1990 and 1995, mostly from Colchester County, but
also from Pictou and Halifax Counties. The earliest collections are from Debert,
Colchester County in 1990. Nine more were collected in Debert in 1991 . The
first collections from Pictou County were in 1994 and in Halifax County in
1995.

Vol. 1 09, No. 1 , January & February, 1 998 17

Additional Locality Records:

Colchester County, Debert, 13 June 1990, E. Georgeson, NSDNR; Colchester County, Debert,
16 June 1990, T.D. Smith, NSDNR; Colchester County, Debert, 19 June 1990, E. Georgeson,
NSDNR; Pictou County, Lyons Brook, 4 July 1994, E. Georgeson, NSDNR; Halifax County,
Peggy's Cove, 15 August 1995, J. Ogden, NSDNR

Harmonia axyridis (Pallas): Many attempts to introduce this species in North
America, including Nova Scotia, occurred between 1916 and 1983 (Gordon
1985; Hoebeke and Wheeler 1996). The first established population was
recorded in Louisiana in 1988 (Chapin and Brou 1991). Since then there has
been an explosive range expansion and population explosion in northeastern
North America (see Kidd et al. 1 995), with the first records for Canada in 1 994
(Coderre et al. 1995; Marshall 1995). This range expansion has continued into
the Maritimes with Hoebeke and Wheeler (1996) reporting it from three sites,
including two in Nova Scotia (Lunenburg and Digby Counties). They did not
record it from Cape Breton Island, where it has since been collected during the
summers of 1995 and 1996.

Additional Locality Records:

Cape Breton County, Sydney, UCCB Campus, 19 September 1995, K.J. Aucoin, UCCB; Cape
Breton County, Sydney, 15 July 1996, A.A. Risk, UCCB

Hippodamia variegata (Goeze): This species first became established at
Montreal, Quebec before 1984 (Gordon 1987). By 1992 it was widely
distributed from northern New Jersey through New York, Connecticut and
Vermont to Quebec (Wheeler 1993). The continuing range expansion to
southern New Brunswick in 1993 and to Prince Edward Island and Colchester
County, Nova Scotia in 1 995 was documented by Hoebeke and Wheeler ( 1 996).
One year later, the spread has continued another few hundred kilometres to
Antigonish County and Cape Breton County.

Specimens:

Antigonish County, Antigonish, 10 May 1996, B. Jones, STFXU;
Cape Breton County, Sydney, September 1996, J. Meagher, UCCB

The coccinellid fauna of Nova Scotia has changed dramatically in the past
15 years with the addition of four adventive species. The addition of these
species is similar to what has occurred over much of northeastern North
America (Gordon and Vandenberg 1991).

It is interesting to compare the rate of range expansion for the four spe-
cies, from their initial establishment in North America until they reached Cape
Breton Island (Table 1). Cape Breton is at the northeastern limit of possible
range expansion on continental North America. Further expansion requires a
major water crossing to Newfoundland. I have collated the information based
on the first established population in North America, as reported in Gordon

18 ENTOMOLOGICAL NEWS

Table 1 : Average range expansion per year for four species of adventive coccinellids now estab-
lished on Cape Breton Island, Nova Scotia. First established populations are from Gordon and
Vandenberg (1991). The distance to Sydney, NS was measured from "The International Atlas"
(McNally 1979).

Species First established First Approximate Average range

population Sydney, NS Distance expansion per

record (km) year (km)

Coccinella Bergen County, NJ 1985 1300 108

septempunctata

Propylea Quebec City, PQ 1995 850 31

quatuordecimpunctata 1 968

Harmonia axyridis Abita Springs, LA 1995 3100 442

1988

Hippodamia Montreal, PQ 1996 1050 80

variegata <1984

and Vandenberg (1991) and the first record for Sydney, Cape Breton Island.
Linear distance was based on a straight line measurement in "The Interna-
tional Atlas" (McNally 1979).

Rapid range expansion, with a minimum of 30 km/year and maximum of
more than 400km/year, is clear for all four species. The rates of range expan-
sion give a good picture of how quickly these species have colonized North
America. It is possible these overestimate 'real' rates of range expansion. There
have been many releases and attempts to introduce these species in many loca-
tions in eastern North America (e.g. Chapin and Brou 1991; Wheeler 1993)
and some releases or accidental introductions (e.g. Yanega 1996) closer to Nova
Scotia than the first established population, may have been successful. How-
ever, the rates are still useful because they emphasize the explosive potential
for range expansion of introduced species in North America.

Beyond impressive rates of range expansion, the newly established spe-
cies will have an effect on the existing assemblage of coccinellids. Within five
years of establishment on Cape Breton Island, Coccinella septempunctata be-
came the most common species in anthropogenic habitats, as documented by
the collections of UCCB students. Now, there are three more adventive spe-
cies in Cape Breton. It remains to be seen whether one or more of these will
displace C. septempunctata as the common coccinellid of disturbed habitats.
There is also the potential for these adventive species to reduce populations of

Vol. 1 09, No. 1 , January & February, 1 998 19

native species. Wheeler and Hoebeke (1995) have documented dramatic de-
clines in the populations of the native C. novemnotata in eastern North America.
Attention should be focused on the influence of these new adventives on the
common native species such asAdalia bipunctata and C. trifasciata in the next
few years.

This note documents useful additions to our knowledge on range expan-
sion and establishment of adventive coccinellids based on specimens in small
regional insect collections. This type of collection should play a primary role
in documenting population changes of these species in the next 15 years.

ACKNOWLEDGMENTS

I thank Eric Georgeson and Jeff Ogden of the Nova Scotia Department of Natural Resources
Insect Collection, Belmont, Nova Scotia, Randy Lauffof St. Francis Xavier University, Antigonish,
Nova Scotia, Barry Wright and Andrew Hebda of the Nova Scotia Museum of Natural History,
Halifax, Nova Scotia, and Steve Marshall, University of Guelph, Guelph, Ontario for allowing
me to examine specimens under their care and for discussions about coccinellids. Chris Thomson,
Steve Marshall and Randy Lauff kindly reviewed the manuscript. I also thank the students of
Biology 375 (Entomology) for providing many useful specimens. Financial support was pro-
vided by Natural Sciences and Engineering Research Council and UCCB.

LITERATURE CITED

Chapin, J.B. and V.A. Brou. 1991. Harmonia axyridis (Pallas), the third species of the genus to

be found in the United States (Coleoptera: Coccinellidae). Proc. Entomol. Soc. Wash. 93:

630-635.
Coderre, D., E. Lucas, and I. Gagne. 1 995. The occurrence of Harmonia axyridis (Coleoptera:

Coccinellidae) in Canada. Can. Entomol. 127: 609-61 1 .
Day, W.H., D.R. Prokrym, D.R. Ellis, and RJ. Chianese. 1994. The known distribution of the

predator Propylea quatuordecimpunctata (Coleoptera: Coccinellidae) in the United States,

and thoughts on the origin of this species and five other exotic lady beetles in eastern North

America. Entomol. News 105: 244-256.
Gordon, R.D. 1985. The Coccinellidae (Coleoptera) of America North of Mexico. J. N. Y.

Entomol. Soc. 93:1-912.
Gordon, R.D. 1987. The first North American records of Hippodamia variegata (Goeze). J. N.

Y. Entomol. Soc. 95: 307-309.
Gordon, R.D. and N. Vandenberg. 1991. Field guide to recently introduced species of

Coccinellidae (Coleoptera) in North America, with a revised key to North American genera

of Coccinellini. Proc. Entomol. Soc. Wash. 93: 845-864.
Hoebeke, E.R. and Wheeler, A.G. 1996. Adventive Lady Beetles (Coleoptera: Coccinellidae)

in the Canadian Maritime Provinces, with new Eastern U.S. Records of Harmonia

quadripunctata. Entomol. News 107: 281-290.
Kidd, K.A., C.A. Nalepa, E.R. Day, and M.G. Waldvogel. 1995. Distribution of Harmonia

axyridis (Pallas) (Coleoptera: Coccinellidae) in North Carolina and Virginia. Proc. Entomol.

Soc. Wash. 97: 729-73 1 .

Marshall, S.A. 1995. Ladybird, Fly Away Home. Seasons Spring 1995: 30-33.
McNally, A. 1979. The International Atlas. Rand McNally Co., Chicago. 534pp.
Schaefer, P.W., R.J. Dysart, and H.B. Specht. 1987. North American distribution ofCoccinella

septempunctata (Coleoptera: Coccinellidae) and its mass appearance in coastal Delaware.

Environ. Entomol. 16: 368-373.

20 ENTOMOLOGICAL NEWS

Tedders, W.L. and P.W. Schaefer. 1994. Release and establishment of Harmonia axyridis (Co-

leoptera: Coccinellidae) in the southeastern United States. Entomol. News 105: 228-243.
Wheeler, A.G. Jr. 1993. Establishment of Hippodamia variegata and new records of Propylea

quatuordecimpunctata in the eastern United States. Entomol. News 104: 102-1 10.
Wheeler, A.G. Jr. and E.R. Hoebeke. 1995. Coccinella novemnotata in northeastern North

America: historical occurrence and current status (Coleoptera: Coccinellidae). Proc. Entomol.

Soc. Wash. 97:701-716.
Yanega, D. 1996. Records of Propylea quatuordecimpunctata (Coleoptera: Coccinellidae) from

Long Island, New York: evidence for a naturalized population before 1991. Entomol. News

107: 36-38.

SCIENTIFIC NOTE

RECENT INTERCEPTION OF LIVE KHAPRA

BEETLE, TROGODERMA GRANARIUM

(COLEOPTERA: DERMESTIDAE), AT THE PORT OF

BALTIMORE, MARYLAND 1

Michael L. Zimmerman, Julia A. Harron-

The khapra beetle, Trogoderma granarium Everts, is a quarantined insect pest under United
States Department of Agriculture (USDA) regulations. Each day, USDA Animal and Plant Health
Inspection Service - Plant Protection Quarantine officers (USDA-APHIS-PPQ) examine a mul-
titude of commodities entering the United States from all over the world looking for numerous
foreign insect / plant / animal pests in order to ensure that none are present prior to entering the
United States. Inspections are made of both commercial and personal freight arriving daily at
ports of entry in vehicle, rail, airline, and ocean liner luggage/baggage. The khapra beetle is one
of the major concerns for USDA-APHIS-PPQ officers when examining these imported goods. It
is a serious pest of food items including stored cereal grains, dried fruits and other plant and
animal materials, and does not occur in North America. Khapra beetles can survive long periods
without food and make a habit of hiding in cracks and crevices in the warehouse making them
hard to detect (Anonymous, 1983).

In June 1996, a shipment of Basmati rice from Pakistan was off loaded at the Dundalk
Marine Terminal, Port of Baltimore, Maryland, U.S.A. Investigators from the U.S. Food and
Drug Administration (FDA) collected a surveillance sample to check for compliance under the
adulterated food section of the Federal Food, Drug, and Cosmetic Act. The shipment consisted
of 550 nylon weave bags, each containing eight - 1 1 pound cloth bags (sewn closed) of hulled,
white grain rice. Examination of the rice at the FDA District Laboratory revealed the presence of
live and dead maize weevil adults, Sitophilus zeamais Motschulsky (Curculionidae); sawtoothed
grain beetle adults, larvae, and pupae, Oryzaephilus surinamensis (Linnaeus) (Cucujidae); red
flour beetle adults and pupae, Tribolium castaneum (Herbst) (Tenebrionidae); and Trogoderma
sp. dermestid adults and larvae. The Trogoderma larvae were initially identified as T. granarium
Everts (khapra beetle) by the authors using Kingsolver's (1991) key to dermestid beetles. As a
professional courtesy, FDA immediately contacted the local USDA-APHIS-PPQ entomologist
at the Baltimore Port Office and the specimens were hand delivered for positive confirmation.

(Continued on page 46)

1 Received June 10, 1997. Accepted July 1, 1997.

2 U.S. Food and Drug Administration, 900 Madison Avenue, Baltimore, Maryland 2 1 20 1 , U.S.A.

Vol. 109, No. 1, January & February, 1998 21

CAMELOBAETIDIUS VARIABILIS

(EPHEMEROPTERA: BAETIDAE), A NEW SPECIES

FROM TEXAS, OKLAHOMA AND MEXICO 1

N. A. Wiersema^

ABSTRACT: Camelobaetidius variabilis, new species, is described from larvae and from male
and female adults. This species is common in the creeks and rivers of the hill country region of
Texas and is also known from Oklahoma, and Nuevo Leon and Tamaulipas, Mexico. Characters
are presented to separate the larvae of this species from C. similis and other related species.

Lugo-Ortiz and McCafferty (1995) presented a taxonomic review of the
North and Central American species of the genus Camelobaetidius Demoulin.
As a result of their study, many previously accepted species were found to be
variants of more widespread species and two new species were described,
namely C. kondratieffi Lugo-Ortiz & McCafferty and C. similis Lugo-Ortiz &
McCafferty.

In 1996 specimens were collected by the author from the Guadalupe River
in Texas that exhibited tarsal claw denticulation not typical of the only nomi-
nal species, C. mexicanus (Traver and Edmunds), reported in Texas (Allen and
Chao 1978, Henry 1986, McCafferty and Davis 1992, McCafferty and
Provonsha 1993, Lugo-Ortiz and McCafferty 1995). Upon further examina-
tion it was observed that these specimens had gills or osmobranchia on the
pro- and mesothoracic segments. The presence of coxal gills on the middle
coxae was previously unreported in this genus. Camelobaetidius similis de-
scribed from Guerrero, Mexico, as well as other species known from Peru, do
have thoracic gills on the prothorax. After examination of many specimens
contained in my collection and the collection at the Brackenridge Field Labo-
ratory of the University of Texas at Austin it became evident that this was an
undescribed species. W. P. McCafferty and C. R. Lugo-Ortiz (Purdue Univer-
sity, West Lafayette, Indiana) confirmed that this was a previously undescribed
species and that the larvae differed from C. similis by size and tergal pattern-
ing as well as other differences. The adults of C. similis are presently un-
known. This new species is described here from the larvae and the adult male
and female stage as Camelobaetidius variabilis, new species.

All specimens were collected by the author and reside in the author's per-
sonal collection (NAW) unless otherwise indicated. Institutions housing the

1 Received April 26, 1997. Accepted May 28, 1997.

2 4857 Briarbend Drive, Houston, TX 77035.

ENT. NEWS 109(1): 21-26, January & February, 1998

22 ENTOMOLOGICAL NEWS

other materials used in this study and acronyms are as follows: Brackenridge
Field Laboratory, University of Texas at Austin (BFL), Purdue Entomological
Research Collection (PERC), University of North Texas Entomology Museum
(UNTEM), and the United States National Museum of Natural History (USNM).
Life stage and material abbreviations are as follows: L-larvae, M-adult male,
F-adult female E- larval exuvia slide mounted in Euparal.

Camelobaetidius variabilis, NEW SPECIES

(Figs. 1-4)

Larva, (in alcohol) Body length: 5.6-7.0 mm; caudal filaments: 2.4-3.2 mm. Head: Colora-
tion pale with area along coronal and frontal sutures pale olive-brown. Vertex pale olive-brown.
Ocelli black. Turbinate portion of eyes in males reddish brown to orange-yellow, lateral lobes
black. Antennae pale; sensilla not apparent. Labrum (Fig. 1 ) sclerotized posterolaterally, with
one submedial and two to three submarginal simple setae on each side; intermediate setae absent.
Right mandible 3+1+3 denticles; tuft of simple setae between prostheca and molar. Left
mandible 3+1+3 denticles with short, broad process at base of molar. Maxillae robust; maxil-
lary palps extending to galealacinia; palpal segment 2 broad about 0.5-0.75 length of segment 3.
Labium robust; palpal segment one long; segment two shorter than one, with four to five simple
setae dorsally and broad, rounded distal projection; segment three rounded apically, with numer-
ous fine, simple setae; glossae with 10-11 simple setae medially, three to four dorsally and four
to five laterally; paraglossae with five to six simple setae dorsally and two rows of simple setae
distally. Thorax: Color pale to olive-brown with distinct pattern in live larvae (Fig. 4), faint in
perserved specimens. Sterna pale to pale yellowish. Osmobranchia present on for- and midcoxae.
Legs pale; femora rounded and often darkened distally with numerous fine, long, simple setae
dorsally and pale olive-brown area medially. Tibiae armored ventrally with short, robust, simple
setae; tarsi distinctly darken distally with row of short, simple setae ventrally with distal 2-3
about 1 .5-2 times the length of preceding setae. Tarsal claws with 5-9 denticles (rarely 9, see
discussion) similar to Figs. 13, 15 & 16 in Allen and Chao (1978). Abdomen: Color pale olive-
brown; Terga with olive-brown coloration faint on segments 4-5, 7 and posterior half of 10, with
prominent submedian oblique markings and often a round pale area medially on each segment.
Gills on segments 1 & 7 small and without tracheation; those on segments 2-6 usually with very
little or no tracheation. Paraprocts with seven to eight spines. Caudal filaments pale, often
darken distally with numerous, fine, simple setae. Median caudal filament subequal in length to
cerci.

Adult Male, (in alcohol) Body length excluding cerci: 5.6-6.2 mm; forewings: 5.6-6.0 mm;
hindwing: 1 .0 mm; cerci: 8.8-9.2 mm. Body brownish tan, darkest at head, thorax, and abdomi-
nal terga 7-10. Head: Turbinate portion of eyes orange-yellow, semi-elliptical, well separated
and slightly divergent. Antennae pale. Thorax: Pronotum darker laterally. Legs pale. Wing
veins and membrane hyaline. Hindwings as in Fig. 3. Abdomen: Terga distinctly darker than
sterna. Terga brownish tan with segments 2-6 paler than segments 1 & 7-10. Usually darker
laterally giving the appearance of a wide pale median line dorsally. Tracheation along pleura
usually prominently darkened, but sometimes only faintly darkened. Sterna pale. Genitalia pale
and as in Fig. 2. Cerci usually entirely pale, but sometimes with dark articulation in basal seg-
ments.

Adult Female, (in alcohol) Body length excluding cerci: 5.6-6.2 mm; forewings: 5.8-6.2
mm; hindwing: 1 .0 mm; cerci 7.0-7.5 mm. Body coloration as in male with terga distinctly
darker than sterna. Abdominal segments 2-6 not distinctly paler than segments 1 & 7-10, due to
presence of eggs. Hindwing as in Fig. 3. Legs and cerci light tan.

Vol. 109, No. 1, January & February, 1998

Diagnosis. The larvae of C. variabilis are most similar to C. similis but
can be separated from it by having a single thread-like gill on the midcoxae as
well as forecoxae, labral setation lacking intermediate setae (as in Fig. 1), and
tergal patterning (as in Fig. 4). In addition, C. variabilis also differs from C.
similis by different mandibular denticulation and a greater number of marginal
spines of the paraprocts, but these latter difference are of inconsistent diagnos-
tic value. The larvae of C. variabilis are also very similar to C. mexicanus and

Figs. 1 -3. Camelobaetidius variabilis, n. sp.: 1 . Labrum, dorsal view. 2. Male genitalia, ventral
view. 3. Left hindwing.

Fig. 4. Live male larva of Camelobaetidius variabilis, n. sp. from Devils River, Texas. Scale bar
equals 1 mm. Photograph C.R. Nelson.

24 ENTOMOLOGICAL NEWS

C. waltzi McCafferty, and in fact share similar labral shape and setation pat-
terns. However, C. variabilis is easily separable by the presence of the coxal
gills discussed above. In addition, C. variabilis is further separable from C.
waltzi by having a rounded second segment of the labial palps, whereas C.
waltzi has a medially pointed second segment of the labial palps. The larvae
of C. waltzi also tend to have short, broad maxillary palps but this feature is of
inconsistent diagnostic value. Camelobaetidius warreni is also similar to C.
variabilis with respect to size and tarsal claw denticulation, but C. warreni
lacks coxal gills and has labral setation with intermediate setae as in Fig. 6
Lugo-Ortiz and McCafferty (1995).

Using the most recent key (Lugo-Ortiz and McCafferty 1995), the larvae
of C. variabilis will key as C. similis. The following added couplet can be
used to separate the larvae of these two species at that point in the Lugo-Ortiz
and McCafferty key.

Couplet to separate larvae C. variabilis from C. similis

1 . Coxal gills present on forecoxae only; labral setation as in Fig. 5 in Lugo-Ortiz and

McCafferty (1995) similis

1 ' Coxal gills on fore and middle coxae; labral setation as in (Fig. 1 herein) variabilis

According to Traver and Edmunds (1968) only Dactylobaetis sp. B de-
scribed from Metlac (Vera Cruz?), Mexico and C. warreni (C. zenobia and C.
cepheus were synonymized with C. warreni in Lugo-Ortiz and McCafferty
1995), are the only species known in the adult stage in North and Central
America which possess a darker terga than sterna. Adult males of D. sp. B can
be separated from C. variabilis by having semi-opaque abdominal segments
2-6 with tracheae not darkened and large turbinate eyes that are contiguous
dorsally. I have found no reliable characters for the separation of adults of C.
warreni from C. variabilis.

Known distribution. USA: Oklahoma, Texas; MEXICO: Nuevo Leon, Tamaulipas.

Material examined. HOLOTYPE: Male larva, USA: TEXAS: Williamson Co., Georgetown,
San Gabriel River, riffles below San Gabriel Park, 20 Jan 1997 (USNM). ALLOTYPE: Female
larva, same data and deposition as holotype. PARATYPES: One male and one female larvae
same data as holotype (PERC). ADDITIONAL PARATYPES: Three male and five female
larvae, USA: TEXAS: Comal Co., Guadalupe River about 11 mi., below Canyon Dam, 09 Nov
1996, same deposition as holotype.

Other Material Examined. USA: TEXAS: Comal Co., Guadalupe River about 12.5 mi.,
below Canyon Dam, 16 Feb 1997 (8M, 3F, 4E); same location as before, but 21 Feb 1997 (7M,
2F). USA: TEXAS: COMAL CO., Guadalupe River about 1 Imi. below Canyon Dam, 17 Feb
1997 (L); same location as before, but 21 Mar 1997 (10 M, 3F). USA: TEXAS: Travis Co.,
Austin, Little Walnut Creek at Cameron Rd., 25 Mar 1997 (L); same location as before but 12
Apr 1997 (L). USA: TEXAS: WILLIAMSON CO., Tejas Camp, unnamed creek flowing into
San Gabriel River, 19 Dec 1996 (L). USA: TEXAS: BASTROPCO., McKinney Roughs, Colo-
rado River at Wilbarger Bend. 23 Nov 1996 (L). USA: TEXAS: VAL VERDE CO., Devils
River, Dolan Falls Preserve. 27-29 Oct 1996 (L). USA: TEXAS: Edwards Co., Nueces River,

Vol. 109, No. 1 , January & February, 1998 25

Barksdale, Hwy 55, 1 5 Mar 1993 C.R. Nelson, K.D. Alexander & S.M. Stringer (L; BFL). USA:
TEXAS: Val Verde Co., Dolan Creek upstream from Devils River, 19 Mar 1993. C.R. Nelson,
K.D. Alexander & S.M. Stringer (L; BFL). USA: TEXAS: Val Verde Co., Devils River at Dolan
Falls Preserve, above falls, 1 3 Nov 1993 C.R. Nelson, K.D. Alexander & S.M. Stringer (L; BFL).
USA: TEXAS: TRAVIS CO., Walnut Creek at 1-35, 06 June 1995 (L); same data but Bull Creek
near Spicewoods Spring Road, 12 May 1995 (L). USA: OKLAHOMA: Murray Co., Davis,
Honey Creek at Turner Falls Park, 14 Oct 1993, P.P. Wagner (2M, 3F, L; UNTEM). MEXICO:
NUEVOLEON: Pobillo River at St. Hwy 115, near Linares, 15 May 1995, D.E. Baumgardner&
B.C. Henry (L; UTEM). MEXICO: NUEVO LEON: Pilon River at unnamed road, across from
General Teran, off St. Hwy 35, 15 May 1995, D.E. Baumgardner & B.C. Henry (L; UTEM).
MEXICO: TAMAULIPAS: Branch of Chihue River at Hwy 101, ca. 12 mi S of Juamave, be-
tween kilo marker 91 & 92; 17 May 1995, D.E. Baumgardner & B.C. Henry (L; UTEM).

Etymology. The specific epithet is a Latin word meaning changeable. It
is a reference to the apparent changeability in tarsal claw denticulation in early
instars of this species and possibly the genus as a whole, see discussion.

DISCUSSION

Tarsal claw denticulation is variable in this species, but it most commonly
exhibits 5-6 denticles, with 7-8 being less common. Of the approximately 300
larvae and exuvia examined by the author, one exuvia had 9 denticles on the
tarsal claw of one of the metathoracic legs. Due to the variability in tarsal
claw denticulation this character will not be useful in separating this species
from the known species with 10 or less denticles on their tarsal claws unless
specimens are collected in series.

A life history study of C. variabilis has been conducted by Paul Wagner
(Virginia Tech), and he has indicated to me that in early instars of this species
there is a greater number of denticles per tarsal claw than is seen in the later
instars and that a reduction in the number of denticles occurs within the first
few instars. Similar observations in tarsal claw development have been made
in species within the Family Heptageniidae, (see Traver and Edmunds 1968,
pg. 638; Ide 1935).

In Allen and Chao (1978) C. mexicanus was reported in Texas for the first
time as Dactylobaetis mexicanus (see McCafferty and Waltz 1 990 for discus-
sion on nomenclature change). The authors also presented a brief description
of these specimens, in which they stated that the tarsal claw denticulation was
of the warreni type 5-8 denticles. The original description of C. mexicanus
found in Traver and Edmunds (1968) states that the tarsal claws of C. mexicanus
have 5-6 denticles, this feature was also found to be consistent by Lugo-Ortiz
and McCafferty (1995) in their review. This, along with the general tergal
patterning described by Allen and Chao ( 1 978), suggests to me that their speci-
mens were in actuality C. variabilis. This also calls into question many of the
previous reports attributed to C. mexicanus in Texas.

Camelobaetidius mexicanus was reported in Oklahoma for the first time
by Reisen (1975) from Honey Creek in Murray County. I have examined

26 ENTOMOLOGICAL NEWS

specimens from this location provided to me by P.P. Wagner (Virginia Tech)
that are clearly Camelobaetidius variabilis.

ACKNOWLEDGMENTS

I would like to thank C. R. Nelson (University of Texas at Austin) for his suggestions and
critique of the manuscript and for allowing me access to The Brackenridge Field Laboratory of
the University of Texas at Austin (BFL). I would also like to thank P. F. Wagner (Virginia Tech)
and D.E. Baumgardner (Aberdeen Proving Ground, Maryland) for loan of specimens. I would
finally like to express my appreciation to W. P. McCafferty and C. R. Lugo-Ortiz (Purdue Uni-
versity, West Lafayette, Indiana) for all their assistance and for their continual work towards
Ephemeroptera systematics which made this description possible.

LITERATURE CITED

Allen, R. K. and S. M. Chao. 1978. Mayflies of the Southwest: new species and records of

Dactylobaetis. Pan-Pac. Entomol. 54: 300-304.

Henry, B.C., Jr. 1986. Mayflies of the Concho River, Texas. Southwest. Nat. 31: 15-21.
Ide, P.P. 1935. Postembryonical development of Ephemeroptera (Mayflies). External charac-
ters only. Canad. J. Research. 12: 433-78.
Lugo-Ortiz, C.R. & W.P. McCafferty. 1995. The mayflies (Ephemeroptera) of Texas and their

biogeograpic affinities. In:Current Directions in Research on Ephemeroptera (L.Corkum and

J. Ciborowski, eds.), pp. 151-169. Canadian Scholars' Press Inc., Toronto, Canada. 478 pp.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1995. Taxonomy of the North and Central American

species of Camelobaetidius (Ephemeroptera: Baetidae). Entomol. News 106: 178-192.
McCafferty, W. P. & J. R. Davis. 1992. New and additional records of small minnow mayflies

(Ephemeroptera: Baetidae) from Texas. Entomol. News 103: 199-209.
McCafferty, W.P. & A.V. Provonsha. 1993. New species, subspecies, and stage descriptions of

Texas Baetidae (Ephemeroptera). Proc. Entomol. Soc. Wash. 95: 59-69.
McCafferty, W.P. and R.D. Waltz. 1990. Revisionary synopsis of the Baetidae

(Ephemeroptera) of North and Middle America. Trans. Am. Entomol. Soc. 116: 769-799.
Reisen, W.K. 1975. The ecology of Honey Creek, Oklahoma: spatial and temporal distributions

of the macroinvertebrates. Proc. Okla. Acad. Sci. 55: 25-3 1 .
Traver, J.R. and G.F. Edmunds, Jr. 1 968. A revision of Baetidae with spatulate-clawed nymphs

(Ephemeroptera). Pac. Insects 10: 629-677.

Vol. 1 09, No. 1 , January & February, 1 998 27

NEWLY REPORTED AND LITTLE KNOWN MAYFLIES
(EPHEMEROPTERA) OF TEXAS 1

N. A. Wiersema^

ABSTRACT: Ten species are newly reported for Texas along with new distributions for little
known mayfly species in Texas. Baetis armillatus, B. bimaculatus, B. dubius, Labiobaelis
frondalis, Procloeon rufostrigatum, P. viridoculare, Ephemerella dorothea, Leptophlebia cupida,
L. intermedia and Ephoron album are reported in Texas for the first time.

Lugo-Ortiz & McCafferty (1995) presented the results of a biogeographic
study of mayfly fauna and a checklist of 76 nominal species reported for Texas.
Recent revisionary work (Henry 1993), a status change (Lugo-Ortiz &
McCafferty 1994), new reports (Baumgardner et al. 1997) and a new species
description (Wiersema 1 998) raised this number to 94 nominal species reported
for Texas.

Recent collections in the Big Thicket and Hill Country areas of Texas and
examination of the author's personal collection revealed several species of
mayflies previously unreported for Texas with new distributions for little known
species. Collection records reported here raise the number of nominal species
reported for Texas to 104 and cast doubt on the validity of a previous report of
Ephemerella inermis Eaton.

All specimens were collected by the author and reside in the author's per-
sonal collection unless otherwise indicated. Institutions housing the other ma-
terials used in this study and acronyms are as follows: Purdue Entomological
Research Collection (PERC) and the Burian Ephemeroptera Collection (BEC).
New state records are marked with an asterisk (*). Life stage and material
abbreviations are as follows: L-larvae, M-adult male, F-adult female, rM-reared
adult male, rF-reared adult female.

BAETIDAE

Apobaetis indeprensus Day

New Collection Records. WILLIAMSON CO., Georgetown, San Gabriel River at San Gabriel

Park, 07Oct 1995(2M, IF).

Remarks. This species was first reported in Texas by McCafferty & Davis (1992). There was

some hesitation by those authors in reporting their larval specimens as A. indeprensus because

the larval stage of A. etowah (Traver) is unknown. The adult specimens I collected clearly are A.

indeprensus. This species is now known to occur in the Balconian and Texan biotic provinces in

Texas (Lugo-Ortiz and McCafferty 1995).

1 Received June 2, 1997, Accepted July 14, 1997.

2 4857 Briarbend Drive, Houston, TX 77035.

ENT. NEWS 1 09( 1 ): 27-32, January & February, 1 998

28 ENTOMOLOGICAL NEWS

*Baetis armillatus McCafferty & Waltz

Collection Records. MONTGOMERY CO., New Caney, Caney Creek at US 59, 04 Jan 1997
(L); 08 Mar 1997 (L); 10 Mar 1997 (L). MONTGOMERY CO., New Caney, Peach Creek at FM
1485, 16 June 1997(L,7M, IF, IrM, IrF). AUSTIN CO., San Bernard River at I- 10, near Sealy,
20 May 1997(L, IrM).

Remarks. This species appears to be widely distributed across the eastern half of the country,
and is also known from Ontario, Alberta, (Berner & Pescador 1988) and has recently been re-
ported in Colorado in the West (McCafferty et al. 1993).

*Baetis bimaculatus (Bemer)

Collection Records. MONTGOMERY CO., New Caney, Caney Creek at US 59, 08 Mar 1997

(L); 10 Mar 1997 (L). MONTGOMERY CO., New Caney, Peach Creek at FM 1485, 16 June

1997(L, 2rM, 4rF).

Remarks. This species was described in Bemer (1946) from adults and larvae collected in

Florida and is currently known also from Alabama, Georgia, Mississippi, and South Carolina

(Bemer & Pescador 1988). The larval specimens collected on June 16 1997 agree well with

Berner's description of this species with one exception, the first as well as the seventh pair of

gills are suffused with reddish brown pigmentation. The collection of this species in Texas

extends its known range westward.

*Baetis dubius (Walsh)

Collection Records. COMALCO., Guadalupe River about 1 1-12 miles below Canyon Dam, 16

Feb 1997 (L, 1M); same data but about 7-8 miles below Canyon Dam, 16 Sept 1996 (1M).

COMAL CO., Sattler, Guadalupe River about 4-5 miles below Canyon Dam, 08 Feb 1997 (L).

HAYS CO., Blanco River at Post Road, near Kyle, 14 Apr 1997 (1M); 16 Apr 1997 (3M); 19Apr

1997(L, IrM, IrF), L(PERC); 20 Apr 1997 (L, IF); 24 Apr 1997 (L, 1M), 1 M (PERC); 05 May

1997 (L, IrM, IrF). WILLIAMSON CO., Georgetown, San Gabriel River at San Gabriel Park,

28 Apr 1997 (2F); 19 May 1997 (L, IrM, IrF).

Remarks. This species is widely dispersed throughout the eastern half of North America (Burks

1953; McCafferty & Waltz 1990) and the collection of this species in Texas extends its known

range westward.

Labiobaetis dardanus (McDunnough)

New Collection Records. HAYS CO., Blanco River at Post Rd., near Kyle, 1 4 Apr 1 997 (L); 1 6

Apr 1997 (L); 19 Apr 1997 (L); 24 Apr 1997 (L, IrM, 2rF). TRAVIS CO., Austin, Colorado

River at Hwy 1 83, 22 Sept 1 996 ( 1 F).

Remarks. This species has been reported in the Austroriparian biotic province by Baumgardner

et al. (1997). The data reported here place this species in the Balconian province as well.

* Labiobaetis frondalis (Traver)

Collection Records. NEWTON CO., Nichols Creek Hwy 87, 30 Dec 1996 (L). MONTGOM-
ERY CO., New Caney, Peach Creek at FM 1485, 05 Jan 1997 (L); 16 June 1997 (L, 3rM, 4rF).
MONTGOMERY CO., New Caney, Caney Creek at US 59, 08 Mar 1997 (L); 10 Mar 1997 (L),
26 May 1997(L, IrM, IrF).

Remarks. This species is widely distributed across the eastern half of North America (Morihara
& McCafferty 1979).

Labiobaetis propinquus (Walsh)

New Collection Records. SAN JACINTO CO., Winters Bayou at Hwy 150, 10 mi. SW of Cold
Springs, 23 Dec 1996 (L). MONTGOMERY CO., New Caney, Caney Creek at US 59, 26 May
1997(L, 3rM, 2rF).

Vol. 1 09, No. 1 , January & February, 1 998 29

Remarks. This species was first reported in Texas by McCafferty & Davis (1992) from the San
Bernard River which borders Colorado and Austin County, near the eastern limits of the Texan
biotic province. Lugo-Ortiz & McCafferty (1995) listed this primarily eastern species as exist-
ing in the Texan and Austroriparian provinces, but gave no distribution in the Austroriparian
province. The data reported here place this species in the Austroriparian province.

Paracloeodes minutus (Daggy)

New Collection Records. TRAVIS CO., Austin, Colorado River at Hwy 183 bridge, 22 Sept

1996 (6F). TRAVIS CO., Austin, Bull Creek near Spicewood Springs Rd., 12 Oct 1996 (2F).
MONTGOMERY CO., New Caney, Peach Creek at FM 1485, 16 June 1997 (L, IrM, 3M, IF).
Remarks. Paracloeodes minutus was first reported in Texas by McCafferty & Davis ( 1 992) and
is currently known to occur in the Austroriparian, Kansan, and Texan biotic provinces (Lugo-
Ortiz & McCafferty 1995). The Travis County data listed here extends its known range in Texas
into the Balconian biotic province.

*Procloeon rufostrigatum (McDunnough)

Collection Records. MONTGOMERY CO., New Caney, Caney Creek at US 59, 26 May 1997

(IrF).

Remarks. Procloeon rufostrigatum is known from Manitoba, New Brunswick, Ontario and

Quebec in Canada and Illinois, Kentucky, Maryland, Michigan, Minnesota, Pennsylvania, and

Wisconsin in the United States (Lowen & Flannagan 1992). The discovery of P. rufostrigatum in

southeast Texas significantly extends its known range southward.

*Procloeon viridoculare (Berner)

Collection Records. HAYS CO., Blanco River at Post Rd., near Kyle, 1 4 Apr 1 997 (L); 1 9 Apr

1997 (18M); 20 Apr 1997 (49M, IF); 24 Apr 1997 (52M), 2M (PERC); 05 May 1997 (larva).
AUSTIN CO., San Bernard River at 1-10, near Sealy, 20 May 1997 (L, IrM). MONTGOMERY
CO., New Caney, Caney Creek at US 59, 26 May 1997 (L). MONTGOMERY CO., New Caney,
Peach Creek at FM 1485, 16 June 1997 (L, 3F, 2rM, IrF), IrM, IF and L (PERC).
Remarks. This species is currently known from Alabama, Florida, Georgia, Mississippi and
South Carolina (Berner & Pescador 1988, Unzicker and Carlson 1982: p. 3.85) and was recently
reported in Michigan and Pennsylvania (Waltz & Munro 1996). The discovery of this species in
the Balconian, Texan and Austroriparian provinces in Texas extends its known range westward.

BAETISCIDAE
Baetisca obesa (Say)

New Collection Records. MONTGOMERY CO., New Caney, Caney Creek at US 59, 27 Dec
1 996 (L); 04 Jan 1997(L).

Remarks. This species was first reported in Texas by Lewis (1986) as occurring in the Neches
River drainage system in the Austroriparian province. The data reported here place Baetisca
obesa in the San Jacinto River drainage system within the Austroriparian province as well.

EPHEMERELLIDAE
*Ephemerella dorothea Needham

Collection Records. MONTGOMERY CO., New Caney, Peach Creek FM 1485, 05 Jan 1997
(L); 08 Mar 1997 (L, 2rM, 2rF); 10 Jan 1997 (L, 4rM, 3rF), 2rM, IrF and L (PERC).
Remarks. The adult specimens I reared vary somewhat from published descriptions of this
species (McDunnough 1931, Traver 1935, Burks 1953, Allen & Edmunds 1965) in that male
genitalia are reminiscent of E. inermis Eaton, a western mountainous species. Larvae of the
eastern E. dorothea and western E. inermis are also very similar morphologically, however the
Texas larvae are clearly of the dorothea type, possessing coarse vermiform cuticular stippling
typical of E. dorothea but not E. inermis. Adult variation may be a result of genetic drift in a

30 ENTOMOLOGICAL NEWS

fringe population and there is little doubt the two species are closely related, if not sister species
(pers. comm. W. P. McCafferty). Baumgardner et al. (1997) reported E. inermis from Texas
based on one larva apparently collected in the Balconian province in which the date and collector
where unknown. The farthest east E. inermis is known to occur are the headwaters of the Pecos
and Rio Grande Rivers in New Mexico and its presence in Texas does not fit any biogeographic
patterns (pers. comm. W.P. McCafferty). In light of this information and the presence of E.
dorothea in East Texas, it is assumed that the E. inermis report is the result of a misidentification
of E. dorothea.

Eurylophella doris (Traver)

New Collection Records. SAN JACINTO CO., Winters Bayou at Hwy 150, 23 Dec 1996 (L).

MONTGOMERY CO., New Caney, Caney Creek at US 59, 27 Dec 1996 (L). NEWTON CO.,

Trout Creek at Hwy 87, 30 Dec 1996.(L). MONTGOMERY CO., New Caney, Peach Creek at

FM 1 485, 05 Jan 1 997 (L). NEWTON CO., Cow Creek at Hwy 87, 1 4 Mar 1 995, D. Hulsen & C.

Mcduffie (L).

Remarks. This species has been reported in Texas by Baumgardner et al. ( 1 997). Using the key

of Funk & Sweeny ( 1 994), specimens from Texas have morphological characteristics of E. doris

as opposed to the closely related E. temporalis (broader fore femora, and longer posteral-lateral

projections of the 2nd and 3rd abdominal segments).

HEPTAGENIIDAE
Stenonema exiguum Traver

New Collection Records. AUSTIN CO., San Bernard River at I- 10, near Sealy, 03 Feb 1997
(L). MONTGOMERY CO., New Caney, Caney Creek at US 59, 27 Dec 1996 (L); 04 Jan 97 (L);
05 Jan 1997 (L, 14rM, 12rF). MONTGOMERY CO., New Caney, Peach Creek FM 1485, 05 Jan
1997 (L); 10 Mar 1997 (L, 1M).

Remarks. This species has been known from the Austroriparian biotic province (Lugo-Ortiz &
McCafferty 1995, Bednarik & McCafferty 1979) and the collection record listed here from Aus-
tin County extends its known distribution into the Texan biotic province.

Stenonema terminatum (Walsh)

New Collection Records. BELL CO., Lampasas River at FM 1 1 23 1 4 Dec 1 996 (L); Same data

as previous but at Dice Grove Rd., 14 Dec 1996(L). COMAL CO., Guadalupe River about 11

mi. below Canyon Dam, 7 Dec 1996 (L). HAYS CO., Blanco River at Post Rd., near Kyle, 24

Apr 1997 (IF). MONTGOMERY CO., New Caney, Caney Creek at US 59, 27 Dec 1996 (L); 04

Jan 1997 (L); 05 Jan 1997 (L, lOrM, 8rF). MONTGOMERY CO., New Caney, Peach Creek at

FM 1485, 05 Jan 1997 (L); 10 Mar 1997 (4M, IrF).

Remarks. This species has been known from the Austroriparian and Balconian biotic provinces

(Lugo-Ortiz & McCafferty 1995). The collection of this highly variable and widespread species

(Bednarik & McCafferty 1979) in the Texan province (Bell County data) was expected.

ISONYCHIIDAE
Isonychia arida (Say)

New Collection Records. MONTGOMERY CO., New Caney, Caney Creek at US 59, 04 Jan
1997 (L); 08 Mar 1997 (L); 10 Mar 1997 (L). MONTGOMERY CO., New Caney, Peach Creek
at FM 1485, 05 Jan 1997 (L); 08 Mar 1997 (L, 2rM, 3rF); 10 Mar 1997 (L, 3rM, 2rF).
Remarks. This species is currently being reported in Texas by Baumgardner et al. (1997). The
data reported here represent a new county record.

LEPTOPHLEBIIDAE
*Leptophlebia cupida (Say)

Collection Records. NEWTON CO., Trout Creek at Hwy 87, 30 Dec 1996 (3rM, IrF), IrM and
IrF(BEC).

Vol. 109, No. 1 , January & February, 1998 31

Remarks. Leptophlebia cupida was found cohabiting with L. intermedia (Traver) in Trout Creek.
This genus is being revised by Dr. Steve Burian, Southern Connecticut State University, who
provided confirmations. Besides L. intermedia (see below), the only other species of this genus
known for Texas is the distinctive L. bradleyi (Needham).

^Leptophlebia intermedia (Traver)

Collection Records. NEWTON CO., Trout Creek at Hwy 87, 30 Dec 1996 (8rM, 4rF), 2rM and

IrF(BEC).

Remarks. This species is widely distributed across the eastern half of North America.

Paraleptophlebia volitans (McDunnough)

New Collection Records. MONTGOMERY CO., New Caney, Caney Creek at US 59, 04 Jan

1997 (L); 08 Mar 1997 (L, 2rM, IrF); 10 Mar 1997 (L). MONTGOMERY CO., New Caney,
Peach Creek at FM 1485, 10 Mar 1997 (L).

Remarks. This species has been known from Quebec (McDunnough 1924) to Florida and west
into Louisiana, and it was recently reported in Texas by Baumgardner et al. (1997). The data
reported here represent a new county record.

POLYMITARCYIDAE
*Ephoron album (Say)

Collection Records. MONTGOMERY CO., New Caney, Peach Creek at FM 1485, 16 June
1997(2F).

Remarks. Ephoron album is a common western and midwestem species (McCafferty 1 994) and
Texas represents the southern limit of its known range.

ACKNOWLEDGMENTS

I would like to express my appreciation to the following individuals: W.P. McCafferty (Purdue
University) for confirmation and discussion of the Ephemerella dorothea specimens as well as
further assistance. S.K. Burian (Southern Connecticut State University) for confirmation of the
Leptophlebia specimens. C.R. Nelson (University of Texas at Austin) for loan of specimens and
for all his assistance. D. Hulsen for the loan of specimens. B.C. Kondratieff (Colorado State
University) for some helpful pre-manuscript advice. D.E. Baumgardner and J.H. Kennedy (Uni-
versity of North Texas) for providing me a copy of their manuscript.

LITERATURE CITED

Allen, R.K. & G.F. Edmunds, Jr. 1965. A revision of the genus Ephemerella (Ephemeroptera:

Ephemerellidae) VIII. The subgenus Ephemerella in North America. Ento. Soc. Am. Misc.

Publ. 4: 243-282.
Baumgardner, D.E., J.H. Kennedy, & B.C. Henry. 1997. New and additional records of Texas

mayflies (Insecta: Ephemeroptera). Trans. Am. Entomol. Soc. (in press)
Bednarik, A.F. & W.P. McCafferty. 1979. Biosystematic revision of the genus Slenonema

(Ephemeroptera: Heptageniidae). Can. Bull, of Fish. & Aq. Sci. 201: 1-72.
Berner, L. 1946. New species of Florida mayflies (Ephemeroptera). Florida Entomol. 28: 60-

82.
Berner, L. & M.L. Pescador. 1988. The mayflies of Florida, revised edition. Univ. Press,

Gainesville, Florida. 415 pp.

Blair, W.F. 1950. The biotic provinces of Texas. Texas J. Sci. 2: 93-1 17
Burks, B.D. 1953. The mayflies, or Ephemeroptera, of Illinois. Bull. 111. Nat. Hist. Surv. 26: 1-

216

32 ENTOMOLOGICAL NEWS

Funk, D.H. & B.W. Sweeny. 1994. The larvae of eastern North American Eurylophella Tiensuu

(Ephemeroptera: Ephemerellidae). Trans. Amer. Entomol. Soc. 120: 209-286.
Henry, B.C., Jr. 1993. A revision of Neochoroterpes (Ephemeroptera: Leptophlebiidae) new

status. Trans. Amer. Entomol. Soc. 119:317-333.
Lowen & Flannagan. 1992. Nymphs and imagoes of four North American species ofProcloeon

Bengtsson with description of anew species (Ephemeroptera: Baetidae). Can. Entomol. 124:

97-108.
Lugo-Ortiz, C.R. & W.P. McCafferty. 1994. New records of Ephemeroptera from Mexico.

Entomol. News 105:17-26.
Lugo-Ortiz, C.R. & W.P. McCafferty. 1995. The mayflies (Ephemeroptera) of Texas and their

biogeographic affinities. In: Current Directions in Research on Ephemeroptera (L. Corkum

and J. Ciborowski, eds.), pp. 151-169. Canadian Scholars' Press Inc., Toronto, Canada. 478

pp.
Lugo-Ortiz, C.R. & W.P. McCafferty. 1996. New Central American and Mexican records of

Ephemeroptera species. Entomol. News 107: 303-310.
Lewis, S.P. 1986. Records of Baetisca obesa (Say) (Ephemeroptera: Baetiscidae) in Southeast

Texas and Southwest Louisiana. Southwest. Nat. 31: 396-398.
McCafferty, W.P. 1994. Distributional and classificatory supplement to the burrowing mayflies

(Ephemeroptera: Ephemeroidea) of the United States. Entomol. News 105: 1-13.
McCafferty, W.P. & J.R. Davis. 1992. New and additional records of small minnow mayflies

(Ephemeroptera: Baetidae) from Texas. Entomol. News 103: 199-209.

McCafferty, W.P., R.S. Durfee, and B.C. Kondratieff. 1993. Colorado mayflies (Ephe-
meroptera): An annotated inventory. 1993. Southwest. Nat. 38:252-274.
McCafferty, W.P. & R.D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera)

of North and Middle America. Trans. Am. Entomol. Soc. 116: 769-799.
McDunnough, J. 1924. New Canadian Ephemeridae with notes, II. Canad. Entomol. 56: 90-98,

113-122, 128-133.
Morihara, D.K. & W.P. McCafferty. 1979. The Baetis larvae of North America (Ephemeroptera:

Baetidae). Trans. Am. Entomol. Soc. 105: 139-221.
Traver, J.R. 1935. Part II. Systematics, pp. 239-739. In: Needham, J.G., J.R. Traver, and Y.C.

Hsu. eds., The Biology of Mayflies. Comstock Publ., Ithaca, New York.
Unzicker, J.D. & P. Carlson. 1982. Ephemeroptera, pp. 3.1-3.97. In A.R. Brigham, W.U. Brigham,

and A. Gnika, eds. Aquatic insects and Oligocheates of North and South Carolina. Midwest

Aquatic Enterprises, Mahomet, Illionois.
Waltz, R.D. & J.B. Munro. 1996. Procloeon viridocularis (Ephemeroptera: Baetidae) from

Michigan and Pennsylvania, new range extensions with comments on the species. Entomol.

News. 107: 166-168.
Wiersema, N.A. 1998. Camelobaetidius variabilis, new species (Ephemeroptera: Baetidae) from

Texas and Oklahoma. Entomol. News. 109: 21-26.

Vol. 109, No. 1 , January & February, 1 998 33

REDESCRIPTION OF NERTHRA PRAEC1PUA

(HETEROPTERA: GELASTOCORIDAE)

FROM CHILE 1

J. T. Polhemus 2 , T. Cekalovic K. 3

ABSTRACT: Nerthra praecipua, known previously from the holotype female, has been redis-
covered in Chile. The species is redescribed, supplementing the incomplete original description
of the damaged female holotype. This species belongs to the Nerthra alalicollis group, which
occurs in Australia and Chile.

Nerthra praecipua Todd was described by Todd (1957) from a unique fe-
male in the Reed Collection (acquired by C. J. Drake, now housed in the Na-
tional Museum of Natural History, Washington, D. C.) that had been badly
damaged by dermestids, rendering a complete description impossible. A single
additional female has been collected by the junior author on Chiloe Island,
Chile, providing the first definitive locality and permitting further description
of the species. It is held in the J. T. Polhemus Collection (JTPC).

Nerthra praecipua Todd

Nerthra praecipua Todd, 1957:151, Fig. 9 (Holotype, female, Chile, Reed Collection, now
in USNM); Todd, 1961:472 (checklist).

Supplemental description to that of Todd (1957).

Female: Length 8.10 mm; width of pronotum 5.50 mm; width of abdomen 6.02 mm.
General color stramineous, with scattered dark markings on veins of hemelytra, head, me-
dian lobe of pronotum, scutellum; depressions on head, median lobe of pronotum, fuscous to
piceous; anterior 2/3 of lateral margins of abdominal tergites III-VI dark; scutellum with lateral
tumescence fuscous, anterolateral angles piceous. Thoracic venter mostly stramineous, mesos-
ternum medially mostly piceous except median anterior margin and medial protuberance; ab-
dominal venter mostly dark, with scattered yellowish markings, laterally and distally lighter.
Legs stramineous, with fuscous markings; fore femora broadly infuscated, with fascia at distal 21
3; middle and hind femora with fascia at basal 1/3, irregular annulus at distal 2/3, distal extreme,
fuscous; tibia dark distally.

Structural characters: Head excavate anteriorly, without apical tubercle but with small api-
cal protuberance, with superapical and lateral tubercles (Fig. 4), set with clavate setae, ocelli
absent. Pronotum with median lobe raised, sculptured, set with scattered short clavate setae.
Abdominal tergites sinuate, extending laterally beyond hemelytra (see Todd, 1957, fig. 9), with
groups of moderate-length clavate setae on posterolateral angles. Hemelytra without membrane,
partially fused, with thin medial line of demarcation between hemelytra, set with short clavate
setae along lateral margins; network of veins more extensive than depicted by Todd (loc. cit.).

1 Received March 15, 1997. Accepted May 15, 1997.

2 Colorado Entomological Museum, 31 15 S. York St., Englewood, CO. 801 10.

3 Casilla 764, Concepcion, CHILE.

ENT. NEWS 109(1): 33-36, January & February, 1998

ENTOMOLOGICAL NEWS

Fore femur with edge of anterior dilation forming about a 50 angle with posterior edge;
anterior trochanter with two black-tipped denticles (Fig. 5).

Abdominal stemites almost symmetrical (Fig. 2); ovipositor lobes slightly asymmetrical,
located posterodorsally (Figs. 1 , 3).

Figures 1 - 5. Nerthra praecipua Todd, female, (scale bar - 1.0 mm). 1 - 3. Abdominal
terniinalia. 1. dorsal view. 2. ventral view. 3. lateral view. 4. Head, dorsal view. 5. Anterior
trochanter, with black-tipped denticles anteriorly.

Material examined. 1 female, Chile, Chiloe Island, Puente la Caldera, 41 40' S, 74 02' W,
15 Feb. 1996, T. Cekalovic K. (JTPC).

Comparative notes. TV. praecipua does not resemble any other South
American species, and is the only American member of the N. alaticollis group.
It is the only American species with sternite VII prolonged posteriorly and
lacking emargination, with the female genital structures located dorsally. In
Todd's (1960) key to Australian species of Nerthra, N. praecipua drops at
couplet 3 because it lacks ocelli; thus, in this regard, it is unique in the alaticollis
group. Todd (loc. cit.) stated that species of the alaticollis group have an api-
cal tubercle, but in N. adspersa (Stal) and TV. stall (Montandon) it is no more
than a small apical protuberance, similar to that of N. praecipua.

Habitat. The habitat in which the single specimen was collected is sur-
rounded on all sides by forest, situated in hills of moderate altitude. The veg-
etation is native and semihumid. The trunks and limbs of trees in various
states of decay are covered with mosses, lichens, and adhering ferns of the
genus Hymenophyllum. The vegetative community is a typical damp
Nothofagus forest.

The specimen was separated from a sample of humus (TC-427) by means

Vol. 1 09, No. 1 , January & February, 1 998 35

of a Berlese funnel, in company with various arachnids and insects including
Peloridiidae (Coleorrhyncha: Peloridora sp.).

Zoogeography. N. praecipua provides another link between the hemi-
pteran faunas of Austral South America and Australia. Concerning N. praecipua,
Todd (1957) stated that "The projecting median portion of the last abdominal
sternite and general appearance would seem to indicate that this species is
most closely related to the species of the alaticollis group found in Australia."
This is an interesting observation (supported by independent investigations by
JTP) because Chile and adjacent austral Argentina are Gondwanan in origin,
and several diverse groups found there are phylogenetically linked to New
Zealand, Australia, and New Guinea (e.g. mayflies, Edmunds, 1 975; Aradidae,
Isoderminae, Monteith, 1 982:649; Peloridiidae, Evans, 1 98 1 ; Nothofagus, Ash,
1982:355).

The projecting abdominal sternite VII of females and dorsal location of the
female genital structures is a synapomorphy for the clade comprising the
Nerthra alaticollis, laticollis, elongata, and rugosa groups established by Todd
(1955, 1960). The laticollis group occurs in Australia and Melanesia (mainly
Solomon Islands and New Guinea), the elongata and alaticollis groups occur
on Australia, but the latter also on Chile. The rugosa group has only a few
species that occur on various islands and continental land masses, and appar-
ently all are halophilous and are rarely if ever found far from the seashore.

The associated Peloridiidae, a hemipteroid family of almost exclusively
flightless bugs (only one species with rare macropters), is of interest because
of its association with, and zoogeographical parallels to, flightless Nerthra
species. The primary ecological difference between them is that peloridiids
are primarily cool-adapted moss inhabitants not yet found on New Guinea,
whereas the flightless Nerthra are mostly warm-adapted litter inhabitants that
are absent from New Zealand. Evans (1981:395) speculated that peloridiids
will be found in the Nothofagus forests of New Guinea.

ACKNOWLEDGMENTS

We thank D. A. Polhemus and R. T. Schuh for their constructive reviews. Support for this
research was provided in part by grant BSR-9020442 from the National Science Foundation,
Washington, D. C. This research was carried out as an adjunct faculty member of Colorado State
University. We thank these organizations for their continued support of research into the system-
atics and zoogeography of aquatic Heteroptera.

LITERATURE CITED

Ash, J. 1982. The Nothofagus Blume (Fagaceae) of New Guinea, pp. 355-380 In: Gressitt, J. L.

ed., Biogeography and Ecology of New Guinea, Junk, The Hague.
Edmunds, G. F., Jr. 1975. Phylogenetic biogeography of mayflies. Ann. Missouri Bot. Gard.

62:251-263.

36 ENTOMOLOGICAL NEWS

Evans, J. W. 1981. A review of the present knowledge of the family Peloridiidae and new
genera and new species from New Zealand and New Caledonia (Hemiptera: Insecta). Rec.
Austr. Mus. 34:381-406.

Monteith, G. B. 1982. Biogeography of the New Guinea Aradidae (Heteroptera), pp. 645-657
In: Gressitt, J. L. ed., Biogeography and Ecology of New Guinea, Junk, The Hague.

Todd, E. L. 1955. A taxonomic revision of the family Gelastocoridae (Hemiptera). Univ. Kans.
Sci. Bull. 37:277-475.

Todd, E. L. 1957. Five new species of Gelastocoridae with comments on other species (Hemi-
ptera). Proc. Entomol. Soc. Wash. 59:145-162.

Todd, E. L. 1960. The Gelastocoridae of Australia (Hemiptera). Pac. Insects 2:171-194.

Todd, E. L. 1961. A checklist of the Gelastocoridae (Hemiptera). Proc. Haw. Entomol. Soc.
17:461-476.

SCIENTIFIC NOTE

AN UNUSUAL OCCURRENCE OF CADDISFLIES

(TRICHOPTERA: PHRYGANEIDAE) IN A

PENNSYLVANIA POPULATION OF THE PURPLE

PITCHER PLANT, SARRACENIA PURPUREA

R. Hamilton IV 2 , R. L. Petersen 2 , R. M. Duffield 2 ' 3

While studying the contents of one hundred and four leaves collected from a population of
purple pitcher plants in ChristnerBog, Somerset County, Pennsylvania, on March 13, 17, and 22,
1996, five caddisfly larvae were recovered. Three larvae were alive in their cases, one larva was
dead in a case, and one was dead and covered by a fungus identified as a species of Saprolegnia
(Saprolegniaceae) (Coker, 1923). The larvae were identified as belonging to the genus Oligostomis
(Phryganeidae). Adult caddisflies collected in the same site in May 1996, were identified as
Oligostomis ocelligea (Walker). This is the first report of Oligostomis larvae in pitcher plants.
The presence of this caddisfly is probably related to the flooding of the habitat during the late
winter and early spring.

Purple pitcher plants are carnivorous plants, usually found in nitrogen-deficient, Sphagnum-
dominated bogs or swamps. A wide variety of species of insects and other arthropods are asso-
ciated with the pitcher plant. These invertebrates can be classified as regular inhabitants, prey,
occasional associates, or plant herbivores (Rymal and Folkerts, 1982). Caddisfly larvae have
been reported in pitcher plants (Brower and Brower, 1971; Cresswell, 1991), yet it is not clear
what their presence signified.

The water level in wetlands can rise significantly after snow melt or rain, submerging the
pitcher plant leaves. This would permit aquatic organisms to move inside and/or on top of the
leaves. Pittman et al. (1996) proposed that aquatic predators may gain entrance to the pitchers
during periods of high water and flooding, and feed on the inhabitants. While it is possible
caddis larvae may have inadvertently entered the pitcher plant leaves due to flooding, we suggest
that the larvae recovered in this study entered the pitcher plant leaves in search of food. Oligostomis

(Continued on page 50)

1 Received April 28, 1997. Accepted July 28, 1997.

2 Department of Biology,Howard University, Washington, D.C. 20059.

3 To whom correspondence should be sent.

Vol. 109, No. 1, January & February, 1998 37

A NEW SPECIES OF SYMPHITONEURIA

(TRICHOPTERA: LEPTOCERIDAE)

FROM SABAH, MALAYSIA '

Trond Andersen^, Jolanda Huisman^

ABSTRACT: Symphitoneuria sabaensis n. sp. from Sabah (Borneo), East Malaysia is described
and figured as male and female imago. The species is the second Symphitoneuria to be described
from outside the Australian biogeographical region. A key to the adult males of Symphitoneuria
is provided.

The genus Symphitoneuria was described by Ulmer ( 1 906) for Notanatolica
exigua (McLachlan). Later, Mosely (1936) described the genus Loticana for
Notanatolica opposita (Walker), at that time only known from the female. Banks
(1939) suggested that Loticana was a synonym of Symphitoneuria, an opinion
formally confirmed by Mosely and Kimmins (1953). Symphitoneuria is char-
acterized by the male forewing venation in which the media and cubitus are
fused with the lower margin of the discoidal cell. In the female the wing vena-
tion is normal, and the hind wings are similar in both sexes. The male genitalia,
however, closely resemble those found in the genus Triplectides Kolenati.

In his study on the phylogeny and classification of the longhorned caddisfly
family Leptoceridae, Morse (1981) included Symphitoneuria in the tribe
Triplectidini of the subfamily Triplectidinae. The phylogeny of Triplectidinae
was outlined by Morse and Holzenthal (1987), but they did not resolve the
phylogeny of Triplectides sensu lato, and were inclined to regard the various
monophyletic lineages identified as subgenera only. By suggesting the alterna-
tive, that Symphitoneuria, Lectrides Mosely in Mosely and Kimmins,
Symphitoneurina Schmid, and Triplectidina Mosely, presently considered sepa-
rate genera, apparently evolved from within Triplectides, the latter genus would
be rendered paraphyletic. As implied by Morse (1989) further studies on the
taxonomy, morphology and zoogeography of the Triplectides complex are es-
sential to resolve this problem.

Including the new species described here, the genus Symphitoneuria now
comprises seven species. With the exception of S. dammermanni Ulmer, all
previously described species appear to be restricted to the Australian biogeo-
graphic region (Morse, 1989). Symphitoneuria dammermanni was described
by Ulmer ( 1 95 1 ) from the Sumba Islands, Indonesia, just west of Weber's line.
Ulmer (1906) recorded females of S. opposita from Sulawesi (Celebes). Later,

1 Received June 19, 1997. Accepted July 28, 1997.

2 University of Bergen, Museum of Zoology, Museplass 3, N-5007 Bergen, Norway.

3 University of Minnesota, Department of Entomology, 219 Hodson Hall, 1980 Folwell Avenue,
St.Paul, Minnesota 55108-6125 USA.

ENT. NEWS 109(1): 37-46, January & February, 1998

38 ENTOMOLOGICAL NEWS

Martynov ( 1 93 1 ) reported a male of this species (as Notanatolica opposita),
also from Sulawesi, but did not describe it, even though the male of 5. opposita
was not known at that time. Banks (1913) further recorded 5. opposita (as
Notanatolica opposita) from the Philippines. Neboiss (1977) listed 5. opposita
from Sulawesi, but Morse (1989) suggested that these records might be
misidentifications of S. dammermanni or the specimens might belong to some
yet undescribed species. With the distribution of 5. opposita in doubt, the de-
scription of S. sabaensis, n. sp., from Sabah (Borneo) confidently extends the
known range of the genus further into the Oriental biogeographic region.

The material treated here was collected by the junior author during several
trips to Sabah, Sarawak and Brunei between 1986 and 1990. Habitats collected
ranged from lowland to montane (50 to 3300 m a.s.l.) primary forest types.

MATERIAL AND METHODS

The material was collected predominately with light traps and the speci-
mens were preserved in 70% alcohol. Methods used in preparing, examining,
and illustrating genitalia are those commonly used in the study of Trichoptera.
The terminology is adopted from Morse and Neboiss (1982). Measurements
are reported as total lengths, antennae length is given as the longest measured.
Measurements are given as ranges, followed by the mean when more than three
measurements were taken, and (in parentheses) by the number of specimens
measured.

The holotype and paratypes of Symphitoneuria sabaensis n. sp. are depos-
ited in the Nationaal Natuurhistorisch Museum (formerly Rijksmuseum van
Natuurlijke Historic), Leiden, The Netherlands (RMNH); paratypes are also
deposited in the University of Minnesota Insect Collection, St. Paul, Minne-
sota, USA (UMSP), the National Museum of Natural History, Smithsonian In-
stitution, Washington, USA(USNM), and in the Museum of Zoology, University
of Bergen, Norway (ZMBN).

Symphitoneuria sabaensis, Andersen and Huisman, NEW SPECIES

Figs. 1-16

Adult male (n=10, if not otherwise stated). Forewing length 8.8-10.9, 10.0 mm; hindwing
length 6.0-7.8, 7.2 mm. Eye 0.43-0.56, 0.5 1 mm wide. Antennae at least 35.2 mm long including
0.45-0.52, 0.48 mm long antennal scape. Maxillary palp segment lengths (in mm): 0.42-0.52,
0.46; 0.61-0.72, 0.67; 0.71-0.84, 0.76; 0.37-0.47, 0.42; 0.72-0.85, 0.81. Color (in alcohol) yel-
lowish brown.

Wings (Figs. 1 -2). Forewing with crossvein sc-r broadened, and stigma present; apical forks
I, II, and V present, i.e. forks of Sj+2, $3+4, and Cui+2; f r ^ I reaching basally beyond S; discoi-
dal cell short and broad, slightly shorter than its stem, and nearly as broad as fork II; thyridial cell
vestigial, veins M and Cu seemingly forming single, thick, composite longitudinal vein with 83+4
for apical five-sevenths of discoidal cell, with five veins, 83, 84 + MA, MP, Cuj, and Cu2, arising

Vol. 109, No. 1, January & February, 1998

SC-R

Cu1

Cu2

Figures 1-4. Symphitoneuria sabaensis n. sp. wings. 1. Male forewing. 2. Male hindwing. 3.
Female forewing. 4. Female hindwing.

Abbreviations: Cu - cubitus, dc - discoidal cell, MA - anterior media, MP = posterior media, R
- radius, S - sector, SC - subcosta, tc - thyridial cell; I, II, V - primary apical cells, or "forks",
I, II, and V.

40 ENTOMOLOGICAL NEWS

from it; Cu2 recurved about 100 basally, and arising from composite vein slightly basally to
where 83+4 joins the composite vein; nygma faint, rectangular, present in fork II. Hindwing
shorter and broader than fore wing; apical forks II and V present; fork V originating about one-
half of distance between origin of discoidal cell (fork of S) and fork of M; nygma faint, rounded,
present in fork II.

Genitalia (Figs. 5-9). Segment IX narrow, subrectangular. Superior appendage about three-
fifths as long as tergum X, narrow, setose, in dorsal view broader with rounded apex; with small
lobe mesally at base bearing one strong seta. Tergum X about as long as inferior appendage; in
lateral view with distinct step-like elevation middorsally and narrow apical portion projecting
caudad; in dorsal view rounded, deeply cleft apically, with broadly rounded projections laterally,
single to partly double row of 6-9, 8 (5) setae dorsally on each side; and 6-10, 9 (5) sensilla
apically; with transverse reinforcing band ventrally at base of cleft and lateral, diagonal reinforc-
ing ridge internally towards base. Inferior appendage with broad base, dorsally slightly convex;
in ventral view with posteromesal corner rounded, forming nearly straight angle (90); subbasally
bearing few, long setae ventrolaterally; apicodorsal lobe rounded apically, setose, slightly shorter
than harpago, with accessory projection mesally bearing 4-5, 5 (5) strong, spine-like setae, long-
est 0.064-0.076, 0.069 (5) mm long; in ventral view harpago slender, slightly sinuous, with apex
hooked mesad; basoventral lobe curved caudad, setose, about one-third as long as inferior ap-
pendage. Phallus subcylindrical with slightly sinuous sides; ejaculatory duct narrow, conspicu-
ous; phallotremal sclerite about half as long as tubular portion of phallus.

Adult female (n-10, if not otherwise stated). Forewing length 8.9-9.9, 9.4 mm; hindwing
length 6.5-7.7, 7.2 mm. Eye 0.45-0.50, 0.47 mm wide. Antennae at least 27.9 mm long including
0.40-0.48, 0.45 mm long antennal scape. Maxillary palp segment lengths (in mm): 0.43-0.52,
0.47; 0.60-0.72, 0.68; 0.71-0.84, 0.79; 0.40-0.45, 0.42; 0.76-0.92, 0.85. Color (in alcohol) as in
male.

Wings (Figs. 3-4). Forewing with crossvein sc-r broadened and stigma present; apical forks
I, II, and V present; fork I sessile; discoidal cell long, subequal to thyridial cell in length; nygma
faint, rectangular, present in fork II. Hindwing shorter and broader than forewing; apical forks II
and V present; fork V long, originating about one-third of distance between origin of discoidal
cell (fork of S) and fork of M; nygma faint, rounded, present in fork II.

Genitalia (Figs. 10-14). Abdominal segment VIII with sternum narrow, triangular, setose; in
ventral view with posterior margin broadly rounded. Segment IX with tergum broad; pleuron
with distinct, rounded corner laterally. Dorsal setose lobe narrow, triangular, attached along ven-
tral surface; in dorsal view broadly rounded to subtriangular; with short, triangular semimembra-
nous sensilla-bearing process, bearing one or occasionally two setae. Lamella with longitudinal
striae on ventral surface; with double to triple row of slightly curved setae along posterolateral
margin; apparently flexible both in posterior and in ventral direction, when in dorsal, (open)
position narrowly triangular in lateral view (Fig. 10), in dorsal view subrectangular and pointing
posterolaterad (Fig. 1 3); when turned more ventrad, (closed position), subtriangular with broadly
rounded apex in lateral view (Fig. 11), in dorsal view rounded, pointing posteriad (Fig. 14); in
some specimens apex turned further ventrad, (fully closed position), overlapping, forming roof-
shaped hood. Gonopod plate apparently somewhat flexible, with straight or convex ventral mar-
gin in lateral view; in ventral view with posterior margin subrectangular, with small, rounded,
median excavation. Spermathecal sclerite oval, broadly rounded anteriorly; in lateral view slightly
curved, subrectangular; in all spent females twisted or positioned askew.

Egg mass (Figs. 15-16). Consisting of strings of eggs apparently glued together with trans-
parent secretion, with scattered long setae, probably originating from the wing fringe, encrusted
in the surface. Outline oval or slightly ovoid, with one side rounded, other side flatter with dis-
tinct groove where next layer of eggs begin. Egg mass 2.19-2.62 (2) mm long, 1.87-1.89 (2) mm
wide, and 1.34-1.38 (2) mm thick.

Larva and pupa. Unknown.

Vol. 109, No. 1, January & February, 1998

sup app

-ap do

Iffl

ph sc

-- ej du

sup app

Figures 5-9. Symphitoneuria sabaensis n. sp. male genitalia. 5. Lateral. 6. Ventral. 7. Dorsal. 8.
Phallus, ventral. 9. Phallus, lateral.

Abbreviations: ac pr - accessory projection of apicodorsal lobe of inferior appendage, ap do -
apicodorsal lobe of inferior appendage, bv - basoventral lobe of inferior appendage, ej du -
ejaculatory duct, har- harpago, inf app- inferior appendage, IX - segment IX, ph sc - phallotremal
sclerite, sup app - superior appendage, X - tergum X.

ENTOMOLOGICAL NEWS

, do se

_-- s-b p

- viii

lam

lam --__

do se

IX-

go pi

sp sc

Figures 10-14. Symphitoneuria sabaensis n. sp. female genitalia. 10. Lateral, with lamella in
open position. 1 1 . Lateral, with lamella in closed position. 12. Ventral. 13. Dorsal, with lamella
in open position. 14. Dorsal, with lamella in closed position.

Abbreviations: do se = dorsal setose lobe, go pi - gonopod plate, IX - segment IX, lam - lamella,
s-b p - sensilla-bearing process, sp sc - spermathecal sclerite, VIII - sternum VIII.

Vol. 109, No. 1, January & February, 1998 43

Figures 15-16. Symphitoneuria sabaensis n. sp. egg mass. 15. Ventral (flat side). 16. Lateral
(blunt end).

Type material. HOLOTYPE: Cf, Malaysia, Sabah, Long Pa Sia area E of kampong, 25. xi-
8.xii.l987, 1000 m a.s.l.. Malaise trap 4, C. van Achterberg (RMNH). PARATYPES: 1 Cf , Long
Pa Sia, riverbank near airstrip, 0425'N 11543'E, 1000 m a.s.l., 21 .x. 1986, J. Huisman (UMSP);

5 9 , Long Pa Sia, airstrip, 0424'N 11543'E, 1000m a.s.l., 1 6.iv. 1987, at light, J. Huisman &J.
van Tol (RMNH 3 9 , NMNH 9 , ZMBN 9 ); 1 9 , Long Pa Sia, 04"25'N 1 1543'E. 1000 m
a.s.l., 1 . xii. 1 987, J. Huisman & C. van Achterberg (UMSP); 4 9 , Long Pa Sia, airstrip, 0424'N
11543'E, 1000m a.s.l., 7. xii. 1987, at light, J. Huisman (RMNH); 1 Cf, Sapulut, kampong, road-
side, 4.V.1987, 0442'N 1 1629'E, 290 m a.s.l., at light, J. Huisman (ZMBN); 5 Cf , 4 9 , 7 km S
Nabawan, near old airstrip, pond, 1 4.xi. 1 987, 0502'N 1 1 627'E, 400 m a.s.l., at light, J. Huisman

6 R. de Jong (RMNH); 1 Cf, 2 9, 1 1 km S Nabawan, pond on roadside, 15.xi.1987, 04"57'N
11627'E, 400m a.s.l., at light, J. Huisman &R.de Jong (UMSP); 1 Cf, 1 9, 12.5 km S Nabawan,
on road to Kg. Pamuntariah, 1st bridge, 16.xi.1987, 0457'N 11627'E, 400 m a.s.l., at light, J.
Huisman & R. de Jong (RMNH Cf , ZMBN 9 ); 3 Cf , Kundassang kampong, Sg. Liwagu, on
bridge, 23.viii.l 986, 0600'N 1 1634'E, at light, J. Huisman (UMSP 2 Cf, ZMBN Cf); 1 Cf , 1 9,
Kundassang kampong, Sg. Liwagu, on bridge, 23.xi.1986, 0600'N 11634'E, 1185 m a.s.l., J.
Huisman (NMNH Cf , UMSP 9 ).

Etymology: named after Sabah, the northeastern part of the Island of Borneo,
using the Latin suffix -ensis, denoting place, locality or country of origin.

KEY TO MALES OF SYMPHITONEURIA

1 Apicodorsal lobe of inferior appendage with setose, accessory projection mesally 2

1 ' Apicodorsal lobe of inferior appendage lacking setose, accessory

projection mesally 6

2 Tergum X about as long as inferior appendage; fork I in forewing sessile 3

2' Tergum X distinctly longer than inferior appendage, in dorsal view triangular,

pointed, with deeply cleft apex; fork I in forewing with short stem (Mosely and

Kimmins, 1953: 260, figs. 180-181) 5. exigua (McLachlan)

3 Forewing with discoidal cell short and broad, with crossvein r-s reaching
discoidal cell in distal one-fifth; fork V in hindwing long, originating distinctly

closer to base than fork of M 4

44 ENTOMOLOGICAL NEWS

3' Forewing with discoidal cell long and narrow, with crossvein r-s reaching

discoidal cell about two-thirds from base; fork V in hindwing short, originating
about the same distance from base as fork of M (Mosely and Kimmins, 1953: 263, fig.
183) 5. opposita (Walker)

4 Tergum X in dorsal view with apex rounded to bluntly triangular 5

4' Tergum X in dorsal view with apex subtruncate (Morse, 1989: 207, fig. 2)

5. dammermanni Ulmer

5 Tergum X in dorsal view gradually widened basally, with apex bluntly triangular;
accessory mesal projection of apicodorsal lobe of inferior appendage with more than 6
spine-like setae (Neboiss, 1987: 140, fig. 40) S. ampla Korboot

5' Tergum X in dorsal view with broadly rounded and projecting sides and rounded
apex; accessory mesal projection of apicodorsal lobe of inferior appendage with
4-5 strong spine-like setae (Figs. 5-7) S. sabaensis n. sp.

6 Inferior appendage with harpago fused almost entirely to apicodorsal
lobe, basoventral lobe not forked (Mosely and Kimmins, 1953: 268,

fig. 187) 5. wheeleri Banks

6' Inferior appendage with harpago free, basoventral lobe forked (Neboiss,

1986: 222, figs. 38-40) 5. licmetica Neboiss

DISCUSSION

The male wing venation of 5. sabaensis most closely resembles that of S.
ampla Korboot and 5. dammermanni, especially in the comparatively short and
broad forewing discoidal cell and the long hindwing fork V. However, it differs
from both in having Cu2 arising from composite vein slightly basally to where
83+4 joins the vein. The setose mesal accessory projection on the apicodorsal
lobe of the inferior appendage of the male genitalia, groups 5. sabaensis with 5.
ampla, S. dammermanni, S. exigua, and S. opposita. It can be distinguished,
however, from all known species of Symphitoneuria by the rounded apex and
broadly rounded and projecting sides of tergum X.

The female of S. sabaensis can be separated from other described females
except 5. exigua in having the posterior margin of the gonopod plate subrec-
tangular, with a small, rounded, median excavation. However, S. exigua ap-
pears to have the posterior margin of sternite VIII with a shallow, broadly
V-shaped, median excision, while the posterior margin in S. sabaensis is evenly
rounded.

Biology. Little seems to be known about the ecology and habitat requirements
of most of the Symphitoneuria species. St Clair (1994) described the larvae of
S. exigua and stated that it is usually found in small, sluggish, often turbid
lowland streams and swamps. According to lilies (1969) 5. ampla appeared to
be the most abundant Trichoptera species in Lake Pinde and Lake Aunde at
about 3,600 m a.s.l. on Papua New Guinea. The present specimens were taken
in localities ranging from 290 to 1 1 85 m a.s.l., mostly along fast flowing rivers,
but also close to stagnant blackwater pools.

Vol. 109, No. 1 , January & February, 1998 45

In some genera of Triplectidini the females carry egg masses at the tip of
their abdomen (see e.g. St Clair, 1993). Korboot (1963) studied the life history
of S. exigua, figuring and describing the egg mass as spherical and covered in
a layer of mucilage. The egg mass was dropped by the female from a height of
3 to 4 inches into the water. Although none of the females of S. sabaensis n. sp.
had their egg masses still attached when studied, the females apparently carry
the egg mass between sternum VIII and the gonopod plate. Sternum VIII ap-
pears to be movable and a cavity is formed in the pleural region of segment
VIII. Setae probably originating from the wing fringe and imbedded in the egg
mass were seen inside the cavity in several of the females. The apparent flex-
ibility of the lamella, which allow for movement in both posterior and ventral
directions, might also be an adaptation for manipulation of the egg mass.

ACKNOWLEDGMENTS

The senior author wants to express his gratitude to Ralph Holzenthal and the University of
Minnesota for making it possible for him to spend his sabbatical at the Minnesota Insect Collec-
tion, and to the University of Bergen for financial support.

The junior author received financial support for the field work in Sabah from Uyttenboogaart-
Eliasen Foundation, MelchiorTreub Foundation, and the National Museum of Natural History in
Leiden. In Sabah, the staff of Kinabalu National Park was very supportive particularly Anthea
Lamb-Phillips and Fui-Lian Inger-Tan; Jan van Tol, Cees van Achterberg and Rienk de Jong
participated in the fieldwork.

We both want to thank Ralph Holzenthal and John C. Morse for all support and useful com-
ments on the manuscript.

LITERATURE CITED

Banks, N. 191 3. On a collection of neuropteroid insects from the Philippine Islands. Proc. Entomol.

Soc. Wash. 15: 170-180.
Banks, N. 1939. New genera and species of neuropteroid insects. Bull. Mus. Comp. Zool. Harv.

Univ. 85: 439-504.
lilies, J. 1969. Trichoptera from the high mountain Lakes Pinde and Aunde, New Guinea. Pac.

Insects 11: 487-493.
Korboot, K. 1963. Biological studies of some Caddis Flies (Trichoptera) from south east

Queensland. Univ. Queensland Pap., Ent., Brisbane 1: 241-274.
Martynov, A.B. 1931. Resultats scientifiques du voyage aux Indes orientates neerlandaises de

LL. AA. RR. le prince et la princesse Leopold de Belgique. Trichoptera. Mem. Mus. Hist.

Nat. Belg., Brussels (h.s.) 4 (2): 13-15.
Morse, J.C. 1981. A phylogeny and classification of family-group taxa of Leptoceridae

(Trichoptera). Pp. 257-264. In: G.P. Moretti, [ed.], Proceedings of the Third International

Symposium on Trichoptera. W. Junk Publ., The Hague, Ser. Entomologica 20: 472 pp.
Morse, J.C. 1989. A redescription of Symphitoneuria dammermanni ; Ulmer, 1951 (Trichoptera:

Leptoceridae). Entomol. Mitt. Zool. Mus. Hambg 9: 205-212.
Morse, J.C. and Holzenthal, R.W. 1987. Higher classification of Triplectidinae (Trichoptera:

Leptoceridae). Pp. 139-144. In: M. Bournaud and H. Tachet [eds.] Proceedings of the Fifth

International Symposium on Trichoptera. W. Junk Publ., The Hague, Ser. Entomologica 39:

397 pp.

46 ENTOMOLOGICAL NEWS

Morse, J.C. and Neboiss, A. 1982. Triplectides of Australia (Insecta: Trichoptera: Leptoceridae).

Mem. Nat. Mus. Vic. 43: 61-98.
Mosely, M.E. 1 936. A revision of the Triplectidinae, a subfamily of the Leptoceridae (Trichoptera).

Trans R. Entomol. Soc. London 85: 91-130.
Mosely, M.E. and Kimmins, D.E. 1953. The Trichoptera (caddisflies) of Australia and New

Zealand. Br. Mus. (Nat. Hist.), Lond. 550 pp.
Neboiss, A. 1977. A taxonomic and zoogeographic study of Tasmanian caddis-flies (Insecta:

Trichoptera). Mem. Nat. Mus. Vic. 38: 1-208.

Neboiss, A. 1986. Taxonomic changes in caddis-fly species from the south' west Pacific- Austra-
lian Region with descriptions of new species (Insecta: Trichoptera). Mem. Mus. Vic. 47: 213-

223.
Neboiss, A. 1987. Identity of species of Trichoptera described by K. Korboot 1964-65 (Insecta).

Mem. Mus. Vic. 48: 131-140.
St Clair, R.M. 1993. Life histories of six species of Leptoceridae (Insecta: Trichoptera) in Victoria.

Aust. J. Mar. Freshwater Res. 44: 363-379.
St Clair, R.M. 1994. Some larval Leptoceridae (Trichoptera) from South-eastern Australia. Rec.

Australian Mus. 46: 171-226.
Ulmer, G. 1906. Neuer Beitrag zur Kenntnis Auser-Europaeischer Trichopteren. Notes Leyden

Mus. 28: 1-128.
Ulmer, G. 1951. Kocherfliegen (Trichoptera) von den Sunda-Inseln. Teil I. Arch. Hydrobiol.,

Suppl. 19: 1-528.

(continued from page 20)

The Basmati rice was immediately detained by FDA after live insects were found and the
presence of khapra beetle suspected. Once the larvae found in the rice were positively confirmed
as khapra beetle by USDA, the rice shipment was fumigated with methyl bromide under USDA
supervision. A reconditioning plan was proposed by the importer to salvage the rice, but it was
subsequently refused. After no further response by the consignee, the shipment was issued a
notice of refusal and was ordered to be exported or destroyed under U.S. Custom's supervision.

A personal communication with J. F. Cavey, Entomologist, USDA-APHIS-PPQ in Riverdale,
MD, provided additional records of khapra beetle interceptions into the United States. He stated
that within the last five years, 35 interceptions of khapra beetle by USDA-APHIS-PPQ officers
were recorded at ports of entry into the United States, with the majority of these interceptions
occurring in Texas, Georgia, and California. Ours is the only record of the khapra beetle found
entering Baltimore, MD during this time.

ACKNOWLEDGMENTS

We thank Susan Broda-Hydorn, Identification Specialist, USDA-APHIS-PPQ, Baltimore,
MD for her confirmation of our identifications and Joseph F. Cavey, Entomologist, USDA-APHIS-
PPQ, Riverdale, MD, for his contribution of unpublished data on khapra beetle interception records
by the USDA into the U.S. since October, 1992.

LITERATURE CITED

Anonymous. May 1983. Khapra Beetle. APHIS Facts, Animal and Plant Health Inspection

Service, US. Dep. Agric., Wash., DC.
Kingsolver, J. M. 1991. Dermestid beetles (Dermestidae, Coleoptera) [pp. 115-135]. In: Insect

and Mite Pests in Food: An Illustrated Key, Volume 1 . J. R. Gorham (ed.), USDA Handbook

No. 655, US. Gov't. Print. Off., Wash., DC. vii + 767 pp.

Vol. 109, No. 1, January & February, 1998 47

A NEW SPECIES OF TOMOCERUS (S.S)
(COLLEMBOLA: TOMOCERINAE) FROM CHINA 1

Y-T. Ma 2 - K. A. Christiansen 3

ABSTRACT: A new Chinese species, Tomocerus cheni, from China is described. It is similar to
the Japanese T. cuspidatus, Borner 1 909, but differs in body color, unguiculus and other features.

Nearly 50 species have been described in the genus Tomocerus (s.l.)\ how-
ever, only 12 species were described or reported from China: Tomocerus
(Tomocerina) minutus Tullberg 1 876 from Shanxi and 1 1 species in the genus
Tomocerus (s.s.): 4 from Tibet (monticolus, obsculus, parvus and zayensis
Huang and Yin 1981), 2 from Yunnan (varius Folsom 1899 andfolsomi Denis
1929), caputiviolaceus Lee 1975 and cuspidatus Borner 1909 from Taiwan,
kinoshitai Yosii 1954 from Hunan, ocreatus Denis 1948 from Zhejiang, and
sibiricus Reuter 1891 from Hebei. A new species of the subgenus Tomocerus
(s.s.) cheni, from Anhui Province, is described here.

Tomocerus (S.S.) cheni, NEW SPECIES

Color: Background color pale yellow with purplish blue pigment. Head dark with irregular pale
spots; eye patches dark. Antennal segments I, IV and basal part of III pigmented, II and most
distal part of III pale. Thoracic segments and anterior margin of abdominal segment I irregularly
pigmented. Abd. V & VI sometimes with a few scattered pigment patches. Base of coxae with
dark pigment. Tibiotarsus with scattered pigment (Fig. 1 ).

Head: Antennae short, respectively 0.5-0.8 and 2.7-3.1 times as long as body and Cephalic di-
agonal; ratios of Ant. I-IV - 1.0/1.4-2.0/4.5-7.3/1.2-2. Eyes 6+6, A & B largest, E & F smallest
(Fig. 2). Labral setae 4/5,5,4, marginally with 4 recurving spinules. Head capsule anteriorly with
2,4 large setae, posteriorly with 43-54 small setae (Fig. 3).

Body: Thoracic macrochaetae and bothriotricha as shown in Fig. 4. Trochanteral organ not clearly
seen. Unguis rather slender; a pair of well developed pseudonychia 1/3-1/2 as long as inner edge
of unguis; inner teeth 5-7, 5-7 & 5-6 respectively on legs I-III. Unguiculus lanceolate with 1
outer tooth and 1 -2 inner teeth. Tenent hair well developed, as long as inner edge of unguis, apex
spatulate (Fig. 5). Tibiotarsus with numerous pointed smooth setae in different sizes; ventral side
with 3-4(5), 6(4,8), 6-8 large blunt spinelike setae respectively on legs I-III (Fig. 6).

Abdominal macrochaetae and bothriotricha on segments I-V as shown in Fig. 4. Tenaculum
unsealed with 4+4 teeth, corpus with 8-12 smooth setae (Fig. 7). Ventral tube scaled, posterior
face with numerous smooth setae in different sizes, anterior face not clearly seen, lateral flap
with about 70 smooth setae in different sizes (Fig. 8). Ratios of manubrium/dens/mucro - 2.S-4/
5-5.6/1. Manubrium laterally with a row of large ciliate setae on each side, these setae more *
strongly tapered near tip; dorsally with 2 longitudinal bands (setaceous stripes, Yosii 1967) of
numerous weakly ciliate to striate, acuminate setae in different sizes, about 20 of them very

1 Department of Biology, Nanjing University, Nanjing 210093 P.R. China

2 Grinnell College Grinnell Iowa 50112 U.S.A.

ENT. NEWS 109(1 ): 47-50, January & February, 1998

ENTOMOLOGICAL NEWS

Tomocerus cheni, All figures of type specimens. Fig. 1. Habitus; 2. Eyes of left side; 3. Cephalic
chaetotaxy; 4. Chaetotaxy of body; 5. Hind foot complex; 6. Large setae of hind tibiotarsus; 7.
Tenaculum; 8. Lateral flap of ventral tube; 9. Left half of distal part of manubrium (dorsal view);
10. Dental spines; 1 1. Mucro; 12. Setulae at base of macrochaeta and bothriothrix.

Vol. 109, No. 1, January & February, 1998 49

large; no scales present between setaceous bands (Fig. 9). Dental spines as 4<3)-5/3(2)-5,I,2( 1 ,4), I;
heavy chestnut brown and each with 2(3) secondary teeth (spinules) near base. 1-2 (rarely 3)
small, finely ciliate, spiny setae present interior to basal dental spines (Fig. 10). Mucro elongate
with numerous ciliate setae; outer dorsal lamella with 5-7 intermittent teeth; outer basal tooth
with a corner toothlet; apical and anteapical teeth subequal (Fig. 1 1 ). Upper anal flap of Abd. VI
with 7 large, striate primary setae arranged in an irregular transverse row.

Scales brownish, hyaline and heavily striated. Each trunk macrochaeta surrounded by 3-6
setulae, bothriotricha without setulae at base (Fig. 12).

Size: Maximum length 3.6 mm.

Type materials Holotype: 9. China: Anhui Province, Yellow Mt., VII- 16- 1990, leaf litter in
deciduous forest and in moss, collection number 8223.

Paratypes: 799. same data as holotype, collection numbers 8223 & 8213

Other locality: 1 9- Anhui: Jinzhai County: Tiantangzhai Park, collection number 8306. All
specimens will be deposited in the Department of Biology, Nanjing University.

Etymology: This species is named after Prof. Jian-xiu Chen in the Department
of Biology, Nanjing University, whose help was essential.

DIAGNOSIS

The large manubrial dorsal setae ("principal setae" of Yosii 1967) are acumi-
nate in T. cheni sp. nov. rather than blunt. This species is very similar to T.
cuspidatus Borner 1909; however, it differs from the latter as shown below:

cheni

cuspidatus*

cuspidatus**

Maximum body length
Scales on tenaculum
Blunt "principal" setae
on manubrium

3.6
absent
absent

6.0
present
2+2,1

6.5

Spinules on dental spine

2(3)

3-6

3-5

* sensu Yosii 1967
** sensu Lee 1975
? unknown

ACKNOWLEDGMENTS

We would like to express our thanks to Mr. Liu Ren-hua who inked all the final drawings
for the present paper. Thanks are also given to Prof. Byung-Hoon Lee and Prof. Jian-xiu Chen
for their great help to this work. This work was published with a grant from Grinnell College.

50 ENTOMOLOGICAL NEWS

LITERATURE CITED

Denis, J. R. 1929. Notes sur les Collemboles recoltes dans ses voyages par le Prof. F. Silvestri.

I. Seconde note sur les Collemboles d'Extreme-Orient. Bull. Lab. Zool. Portici 22: 305-320.
Denis, J. R. 1948. Collemboles d'Indochine recoltes de M. C. Dawydoff. Notes d'Entomologie

Chinoise, Musee Heude, Vol XII, Fasc. 17: 183-259.

Huang, Fu-Sheng, 1981. Insects of Xizang, Vol. 1, pp. 41-46, China Science Press,
Huang, Fu-Sheng, 1 995. Three new recorded species of Tomocerus Nicolet in China (Collembola:

Tomoceridae). Sinozoologia 12: 192-193.
Lee, B. H., 1975. Etude de la Faune coreenne des Insectes Collemboles. VI. Sur la Famille des

Tomoceridae, edaphiques, avec la Description de quatre nouvelles especes et d'une nouvelle

sous-espece. Bull. Mus. Nat. Histoire Nat. 3e sen, no 317, juillet-aout, Zool. 224: 946-961.
Stach, J., 1964. Materials to the Knowledge of Chinese Collembolan Fauna. Acta Zoologica

CracoviensialX(l): 1-26.
Yosii. R., 1967. Studies on the Collembolan Family Tomoceridae, with Special Reference to

Japanese Forms. Contributions from Biol. Lab. Kyoto Univ. No. 20: 1-54.

(continued from page 36)

is primarily a predator, then secondarily an herbivore and detritivore (Wiggins, 1984). The internal
environment of the pitcher plant leaves offers live larvae of other species, plant materials, and
numerous decomposing remains. Analysis of the larval gut contents revealed numerous pieces of
insect cuticle that may have been consumed while the larvae were inside the pitchers.

ACKNOWLEDGEMENTS

We would like to thank the U.S.D. A.-Forest Service (Eastern Region), especially Keith Jensen
and Martin MacKenzie for partial support during the course of this project. We also thank Dr.
Oliver S. Flint, Jr. (Department of Entomology, Smithsonian Institution, Washington, D.C.) for
helpful suggestions. R. Hamilton is supported in part by a fellowship from a National Science
Foundation Graduate Traineeship in Plant Biology (GER 93549 1 6). Voucher specimens are held in
the Department of Biology, Howard University, Washington, D.C. (RMD). Ecological report #5.

LITERATURE CITED

Brower, J. H. and Brower, A. E. 1 97 1 . Notes on the biology and distribution of moths associated

with the pitcher plant in Maine. Entomol. Soc. Ont. Proc. 101:79-83.
Coker, N. C. 1923. The Saprolegniaceae with notes on other water molds. Univ. N. Carolina

Press. Chapel Hill, N.C., 201 pp.
Cresswell, J. E. 1 99 1 . Capture rates and composition of insect prey of the pitcher plant Sarrace-

nia purpurea. Am. Midi. Nat. 125:1-9.
Pitt man, J. L., T. S. Turner, L. Frederick, R. L. Petersen, M. E. Poston, M. Mackenzie, and R.

M. Duffield. 1 996. Occurrence of alderfly larvae (Megaloptera) in a West Virginia population

of the American pitcher plant, Sarracenia purpurea L. (Sarraceniaceae). Ent. News 107:137-

140.
Rymal, D. E. and G. W. Folkerts. 1982. Insects associated with pitcher plants (Sarracenia:

Sarraceniaceae), and their relationship to pitcher plant conservation: a review. J.Alabama Acad.

Sci. 53:131-151.
Wiggins, G. B. 1984. Trichoptera, In: An Introduction to the Aquatic Insects of North America

(2 nd ed.) Merrill, R. W. and Cummins, K. W. (eds). Kendall/Hunt, Dubuque, Iowa, pp. 27 1 -3 1 1 .

Vol. 109, No. 1, January & February, 1998 51

TOMOCERUS (s.s.) SPINULUS

(COLLEMBOLA: ENTOMOBRYIDAE),

A NEW SPECIES OF CHINESE SPRINGTAIL 1

J-X. Chen 2 , K. A. Christiansen 3

ABSTRACT: A new Chinese species, Tomocerus spinulus, from Anhui Province China is de-
scribed here. It is closest to the Pakistan species T. asoka Yosii, 1965 and the Japanese species T.
viridis Yosii, 1967.

Nearly 50 species have been described in the genus Tomocerus s.l.; how-
ever, only 12 species were described or reported from China: Tomocerus
(Tomocerina) minutus Tullberg 1 876 from Shanxi and 1 1 species in the genus
Tomocerus (s.s.; 4 from Tibet (monticolus, obsculus, parvus, and zayensis
Huang and Yin 1981), 2 from Yunnan (varius Folsom 1899 andfolsomi Denis
1929), caputiviolaceus Lee 1975, cuspidatus Borner 1909 from Taiwan,
kinoshitai Yosii 1954 from Hunan, ocreatus Denis 1948 from Zhejiang, and
sibiricus Reuter 1891 from Hebei. A species of the subgenus: Tomocerus (s.s.)
spinulus, is described here.

Tomocerus (s.s.) spinulus, NEW SPECIES

Color: background pale yellow with purplish blue pigment. Head with pale, scattered pigment
on anterior margin between antennae. Eye patches dark. Lateral sides of Ant. I & II pale purplish
blue; Ant. Ill gradually darker from base to apex. Ant. IV totally dark. Pattern as shown in Fig. 1 .
Distal half of tibiotarsus with pale, scattered pigment.

Head: Eyes 6+6, A & B larger, others subequal. Antennae up to 1 .3 mm long; ratio of antenna/
cephalic diagonal: 2.9/1 to 3.2/1; average segment ratios: 1-4= 1/1.2-1.6/6.67-7.67/1-1.4. Head
capsule with 2,4 anterior setae and about 40 small setae in a transverse row at posterior margin
(Fig. 2). Labral setae 4,5,5,4 with 4 marginal recurving spinules (Fig. 3).
Body: Thorax with dorsal chaetotaxy and bothriotricha as in Fig. 4. Unguis rather slender; a pair
of well developed pseudonychia, about 0.25-0.35 as long as inner edge of unguis; inner teeth 5-
6 on all 3 pairs of legs. Unguiculus lanceolate with 1 inner tooth. Tenent hair well developed and
spatulate, about as long as inner edge of unguis (Fig. 5). Tibiotarsus with numerous pointed
smooth setae in different sizes, and legs I-III respectively with 3-4, 4-6, 6-8 apically tapered
spine-like setae on ventral side (Fig. 6). Trochanteral organ reduced with 1/1 smooth setae (Fig.

7).

Abdominal segments 1-5 with dorsal chaetotaxy and bothriotricha as in Fig. 4. Tenaculum
unsealed, corpus with 4-7(9)* smooth setae (Fig. 8). Ventral tube scaled, anteriorly with 1 8 smooth

1 Received March 15, 1997. Accepted May 15, 1997.

2 Department of Biology, Nanjing University, Nanjing 210093, P. R. China

3 Department of Biology, Grinnell College, Grinnell, I A 501 12, U. S. A.
* Numbers in parentheses represent unusual conditions.

ENT. NEWS 1 09( 1 ): 5 1 -55, January & February, 1 998

ENTOMOLOGICAL NEWS

Tomocerus spinulus. All figures of type specimens. 1 . Habitus; 2. Head, 3. Labrum, 4. Chaetotaxy
of body, 5. Hind claw, 6. Hind tibiotarsus, 7. Trochanteral organ, 8. Tenaculum, 9. Anterior
face of ventral tube, 10. Lateral flap of ventral tube.

Vol. 109, No. 1, January & February, 1998

16 a-

Tomocerus spinulus. All figures of type specimens. 11. Distal part of manubrium (dorsal view),
12. Dental spines, 13. Single dental spine, 14. Mucro, 15. Upper anal valve of Abd. VI,
16. Microsetae at base of macrochaeta, 17. Bothriothrix.

54 ENTOMOLOGICAL NEWS

setae (holotype) on each side (Fig. 9), posteriorly with numerous smooth setae in different sizes;
lateral flap with about 60 smooth setae in different sizes (Fig. 10). Ratios of manubrium/dens/
mucro = 2. 3-3.0/3. 3-4.0/ 1 .0. Manubrium with a longitudinal band of lateral setae on each side
and 2 longitudial bands of setae on dorsal side; lateral setae large, weakly ciliated, gradually
tapering but more strongly tapering and pointed distally; each dorsal band of setae consists of
numerous setae in different sizes, those in outer row and at distal part very large; all setae very
weakly ciliate to striate and pointed; no blunt "principal setae" present; scales present between
longitudinal bands of setae (Fig. 11). Dental spines dark chestnut brown, formula 4(3)/4(3)-5,I,
each with many tiny teeth (spinules) (Fig. 1 2 & 13). Denies without inner differentiated swollen
scales or outer spine-like setae. Mucro elongate, with numerous ciliate setae; outer dorsal lamella
bearing 3-6 intermediate smaller teeth; outer basal tooth with a corner toothlet; apical and
anteapical teeth subequal (Fig. 1 4). Dorsal anal valve of Abd. VI with 7 large setae arranged in an
irregular transverse row (Fig. 15). Body scales brownish, hyaline and heavily striate. Each body
macrochaeta surrounded by 0(1-2) setulae (Fig. 16). Bothriotricha not surrounded by setulae
(Fig. 17).

Size: Maximum body length 2.7 mm.

Etymology: The name of this species is derived from the Latin spinula = spines.
It refers to the numerous spinules on dental spines.

Type materials. Holotype: male, China: Anhui Province, Yellow Mt., VII- 16- 1990, collection
numbers 8220 & 8221. Jian Xiu Chen coll. Deposited in the Department of Biology, Nanjing
University.

Paratypes: 10 females & 4 males, same data as Holotype.

DIAGNOSIS

This species bears some similarity to the widespread T. ocreatus but can
easily be distinguished by the much finer denticulations on the dental spines,
the smaller number of tenacular setae and the unguis shape. It also resembles
the Korean species jesonicus Yosii 1967 and spinistriatus Lee 1975 as well as
the Tibetan species zayuensis Huang and Li 198 1 but may be readily separated
as shown below:

Character spinulus jesonicus spinistriatus zayuensis

Dental spines 4(3)/4(3)-5,I 5-6/5-6,1,1,1 5-6,1/6-7,1 3,3/3-4,11

Tenaculum setae 4-7(9) 15 1 ?

Unguiculus tooth + +

Vol. 109, No. 1, January & February, 1998 55

Tomocerus spinulus* is much more similar to the Pakistan species T. asoka
Yosii & Ashraf 1965 & Japanese species T. viridis Yosii 1967. It shares fea-
tures such as the structure of claw and the number of dental spines; however, it
differs from them in the body color and the features listed in the following
table.

SPECIES

Character spinulus asoka viridis

Spinules on dental spines tiny but numerous absent absent except on

distalmost spine

Thick spine-like setae 3-4, 4-6, 6-8 ? 5, 5, 5

on tibiotarsi

Tenacular setae 4-7(9) 15 2

Maximum body

length (in mm.) 2.7 5.0 2.0

* Found in leaf litter in a deciduous forest.

ACKNOWLEDGMENTS

We would like to thank Mr. Ma Yi-tong in the Department of Biology of Nanjing University,
whose help was essential for this work. Publication of this work was made possible by a grant
from Grinnell College.

LITERATURE CITED

Denis, J. R. 1929. Notes sur les Collemboles recoltes dans ses voyages par le Prof. F. Silvestri.

I. Seconde note sur les Collemboles d'Extreme-Orient. Bull. Lab. Zool. Portici 22: 305-320.
Denis, J. R. 1948. Collemboles d'Indochine recoltes de M. C. Dawydoff. Notes d'Entomologie

Chinoise, Musee Heude, Vol XII, Fasc. 17: 183-259.
Huang, F-S and H-F Yi. 1 98 1 . Insects of Xizang, Vol. 1 , pp. 41-46, China Science Press, Beijing.

No. 13031.1657. In Chinese.
Huang, Fusheng, 1 995. Three new recorded species of Tomocerus Nicolet in China (Collembola:

Tomoceridae). Sinozoologia 12: 192-193.
Lee, B.H. 1975 .Etude de la faune coreene des Insectes Collemboles VI. Sur la famille des

Tomoceridae, edaphiques avec la description de quatre nouvelles especes et d'une nouvelle

sous-espece. Bull Mus. natn. Histoire Natur. Paris 317: 945-961.
Yosii, R. & Ashraf, M. 1965. On Some Collembola of West Pakistan. IV. Pakistan J. Sci. Res.

17: 153-160.
Yosii, R. 1967. Studies on the Collembolan Family Tomoceridae, with Special Reference to

Japanese Forms. Contrib. Biol. Lab. Kyoto Univ. No. 20, 1-54.
Zhao, Lijun et al., 1 997. Tentative checklist of Collembolan species from China (Insecta). Publ.

Itako Hydrobiol. Stn., 9: 15-40.

56 ENTOMOLOGICAL NEWS

SPECIES DISTINCTION IN ABDOMINAL
PIGMENTATION PATTERNS BETWEEN FEMALES OF

DROSOPH1LA MELANOGASTER AND
D. SIMULANS, FROM A SPANISH POPULATION 1

Karel Th. Kisses^, Mauro Santos^

ABSTRACT : The sibling species Drosophila melanogaster and D. simulans coexist in natural
conditions. Whereas males are easily recognizable by their genital arches, females were consid-
ered to be indistinguishable but for their eye sizes. In many papers separate female counts were
omitted because of this difficult characteristic. However, the abdominal pigmentation pattern
was found to be different between the two species in a Spanish population. The discrimination of
the females based on pigmentation differences was checked by electrophoresis and found to be
very reliable.

Since the discovery of Sturtevant (1919) that Drosophila melanogaster has
a closely resembling sibling species D. simulans, both species are known to be
cosmopolitan and coexistent (Lachaise et al., 1988). In some population screens
the authors make no effort to distinguish the females of the two species, and
only mention their grand total (Tantawy & Soliman, 1967; references in Lachaise
et al., 1988). Most often, research starts with isofemale lines and checking their
progeny in which the males of the two species are distinguishable due to differ-
ent genital arches (Burla, 1951;Coyne, 1983; Sturtevant, 19 19). Based on mea-
surements of eye sizes of D. melanogaster and D. simulans, it is possible to
make a distinction between the females (Burla, 1951; Gallo, 1973; McNamee
& Dytham, 1993) but it is a painstaking job when large numbers of flies have to
be examined. Okada (1956) described a way of discrimination based on differ-
ences in egg guides, but this character also necessitates much practice to distin-
guish the two species. A high number (up to 45 %) of misqualifications of D.
melanogaster have been reported, based on different eye size definitions
(McNamee & Dytham, 1993 and references therein).

We used flies captured in traps in Carboneras (Almeria, Spain) to see if a
way of morphological distinction by abdominal pigmentation differences (Gallo,
1 973) might be applicable in our population of D. melanogaster and D. simulans.
Eye size was used as the character to separate the species, but we also checked
the pigmentation of the sixth tergite to see whether a useful correlation existed.

1 Received November 5, 1996. Accepted July 23, 1997.

2 Universitat Autonoma de Barcelona, Departament de Genetica i de Microbiologia, 08 193 Bellaterra
(Barcelona), Spain.

3 Present address and address for correspondence: Utrecht University, Department of Plant Ecol-
ogy and Evolutionary Biology, Padualaan 8, 3584 CH Utrecht, The Netherlands, fax: 31-30-
2513655, e-mail:kteisses@dds.nl

ENT. NEWS 109(1): 56-60, January & February, 1998

Vol. 109, No. 1 , January & February, 1998 57

Females emerging from Opuntia ficus-indica fruits were primarily discrimi-
nated by the pigmentation pattern of the sixth tergite. All flies deemed D.
melanogaster were subjected to electrophoresis for other reasons (Hisses and
Santos, 1997).

MATERIALS AND METHODS

Flies were captured with mashed banana traps during five days in the
Carboneras area (Almeria, Spain; 3700'N; 153'W) and locations nearby (Eisses
and Santos, 1997). Opuntiaficus-indica fruits (prickly pears) were put in trays
in a semi-abandoned O. ficus-indica plantation and left for almost seven days
in the field. After recollection, the fruits were placed in glass jars, and emerging
flies were aspirated. Captured and emerged flies were checked for D.
melanogaster morphology and frozen at -29 C until electrophoresis.

ADH is a diagnostic enzyme between D. melanogaster and D. simulans
because of clearly distinctive bands in gel electrophoresis (Eisses, Van Dijk &
VanDelden, 1979).

RESULTS

The apparent D. melanogaster females trapped in the O. ficus-indica plan-
tation near Carboneras were separated from D. simulans by eye size only, whereas
flies from the other locations were separated at the species and sex level by eye
size and genital arches. After electrophoresis the number of misqualifications
of D. melanogaster was calculated (Table I A). Almost 21 % of the female flies
turned out to be D. simulans (Table I A 1). For females and males together a
general misqualification of 1 1 % was obtained (Table I A 2).

Table I. Number of flies initially separated as D. melanogaster and percentage of misqualifications
based on electrophoresis of flies trapped in banana baits in a semi abandoned O. ficus-indica
plantation (A 1) and other locations close to Carboneras (A 2), and of flies emerging from O.
ficus-indica fruits collected at the plantation (B).

Method of distinction No. initially separated % actually

as D. melanogaster determined as D. simulans

'A 1 Eye size 226 females 20.8

2 Eye size / Genital arch 437 females / males 1 1 .0

B Pigmentation of 6th

tergite and eye size 1 078 females 1 .68 0.26*

Genital arch 1 092 males 0.64 0.034
*Empirical Standard Deviation

ENTOMOLOGICAL NEWS

Flies emerging from O. ficus-indica fruits were separated primarily by the
morphological distinction of the pigmentation pattern of the sixth tergite (Fig.
1) and in cases of doubt the eye size was examined as well. Approximately
equal numbers of female and male flies were checked by each of us. After
checking the flies with electrophoresis, the average percentage of misquali-
fications of the females was calculated to be 1 .68 % +.0.26. This is in the same
order as misqualifying male flies (Table I B).

The most important difference between D. melanogaster and D. simulans
females is the black pigmentation of the sixth tergite, which runs to the ventral
margin in D. melanogaster, whereas the pigmentation border line in D. simulans
makes an angle with the tergite margin. It forms a continuous line with the
pigmentation border line in the seventh tergite (Fig. 1 a). In contrast with an
apparently monomorphic D. simulans, we observed large variation in abdomi-
nal pigmentation patterns in this natural population of D. melanogaster and
also in some laboratory strains (Fig. 1 b - i).

Figure 1 . Pigmentation patterns of the 6th and 7th tergite of D. simulans (a) and D. melanogaster
(b - i). Within D. melanogaster variation was present in wild type populations and in homozy-
gous or isogenic laboratory strains Groningen SSN (b) and Groningen FFF (d). None of the D.
melanogaster strains was monomorphic.

Vol. 109, No. 1, January & February, 1998 59

We have demonstrated that distinction between females of a natural popu-
lation of D. melanogaster and D. simulans in Spain can be made in an easy and
reliable way. As similar observations have been made on females from a Brazil-
ian population (Gallo, 1973) and from a midwestern U.S. population (Thomp-
son, Hisey & Woodruff, 1979) it might be generalized to more populations of
D. simulans. It seems worthwhile to excavate information about other D.
simulans populations with respect to female abdominal pigmentation of the
sixth tergite to establish whether or not D. simulans is world wide monomor-
phic for this character in contrast to D. melanogaster (Robertson, Briscoe &
Louw, 1977; David, Capy & Gauthier, 1990). Robertson, Briscoe and Louw
(1977) described the focus fap (female abdomen pattern) to be residing on the
extreme tip of the 3L chromosome, with some effects from the fourth chromo-
some. This might be the reason why the D. melanogaster Groningen-FFF strain,
used as a reference in electrophoresis, showed a pigmentation pattern in the
sixth tergite like D. simulans (Fig. 1 d). This strain is partly homozygous for the
second chromosome and the third chromosome.

ACKNOWLEDGMENTS

Montse Peiro is thanked for helping with electrophoreses of the flies and Albert Kamping
(University of Groningen) for providing the Groningen reference strains of Drosophila
melanogaster. Prof. Wim Scharloo and Dr. Agusti Galiana are thanked for their comments on an
earlier draft. This work was funded by Contract No. CHRX-CT92-0041 from the Commission of
the European Communities, and grant PB-0843 from the DGICYT (Spain) to A. Eontdevila.

LITERATURE CITED

Burla, H. 1 95 1 . Systematik, Verbreitung und Oekologie der Drosophila-Arten der Schweiz. Rev.
SuisseZool. 58, 23-175.

Coyne, J.A. 1983. Genetic basis of difference in genital morphology among three sibling species
of Drosophila. Evolution 37, 1101-1117.

David, J.R., Capy, P., & Gauthier, J.-P. 1990. Abdominal pigmentation and growth tempera-
ture in Drosophila melanogaster: Similarities and differences in the norms of reaction of
successive segments. J. Evol. Biol. 3, 429-445.

Kisses, K.Th., & Santos, M. 1 997. Short distance differences in ADH and alphaGPDH allozyme
polymorphisms and linkage disequilibrium in D. melanogaster in Spanish desert popula-
tions. Dros. Inf. Serv. 80: in press.

Kisses, K.Th., van Dijk, H. & van Delden, W. 1979. Genetic differentiation within the
Melanogaster species group of the genus Drosophila (Sophophora). Evolution 33,1 063- 1 068.

Gallo, A J. 1 973. Morphological distinction between female Drosophila melanogaster and fe-
male D. simulans. Ciencia & Cultura 25, 341-345.

Lachaise, D., Cariou, M.-L., David, J.R., Lemeunier, F., Tsacas, L. & Ashburner, M. 1988.
Historical biogeography of the Drosophila melanogaster species group. Evol. Biol. 22,
159-225.

ENTOMOLOGICAL NEWS

McNamee, S. & Dytham, C. 1993. Morphometric discrimination of the sibling species Droso-
phila melanogaster (Meigen) and D. simulans (Sturtevant) (Diptera: Drosophilidae). Syst.
Entomol. 18,231-236.

Okada, T. 1956. Systematic study of Drosophilidae and allied families of Japan. Gihodo Co.,
Tokyo, Japan.

Robertson, A., Briscoe, D.A. & Louw, J.H. 1977. Variation in abdomen in Drosophila
melanogaster females. Genetica 47, 73-76.

Sturtevant, A.H. 1919. A new species closely resembling Drosophila melanogaster. Psyche 26,
153-156.

Tantawy, A.O. & Soliman, M.H. 1967. Studies on natural populations of Drosophila. VI. Com-
petition between Drosophila melanogaster and Drosophila simulans. Evolution 21, 34-40.

Thompson Jr., J.N., Hisey, B.N. & Woodruff, R.C. 1979. Morphological differences between
the females of Drosophila melanogaster and Drosophila simulans. Southwest Nat. 24,
204-205.

CURTIS W. SABROSKY

The American Entomological Society deeply regrets the recent passing of Dr. Curtis W. Sabrosky,
a friend and Honorary Member of the Society. Although a long time member of the Entomologi-
cal Society of Washington, DC, in recent years, following his move to Medford Leas, Medford,
NJ, Curtis regularly attended meetings of the American Entomological Society.

Because Curtis was honored in 1982 in a "festschrift" (Vol. 10) edition of the Memoirs of
the Entomological Society of Washington, only a brief notice is now planned by the Wash-
ington Society, together with publication of a complete bibliography of his entomological
contributions.

-H.P.B

Vol. 1 09, No. 1 , January & February, 1 998 61

SYNONYMIC NOTES ON SOME OF THOMSON'S
NEW WORLD DOLICHOPODIDAE (DIPTERA) 1

Daniel J. Bickel '

ABSTRACT: Types of four New World Dolichopodidae described by C.G. Thomson ( 1 869) from
material gathered during the voyage of the Swedish frigate Eugenie were examined. Three new
synonyms are established. Dolichopus lamellicornis is a senior synonym of Hygroceleuthus afflictus.
Of three overlooked species described by Thomson from Puna, Ecuador, Chrysotus ochropus is
redescribed and regarded as a senior synonym of C. flavipalpus, Psilopus zonatulus is a junior
synonym of Condylostylus longicornis, and Psilopus pleuralis, known only from the female type,
is referred to the Condylostylus caudatus group.

While completing a faunal study of the Galapagos Dolichopodidae (Bickel
& Sinclair, in press), I had the opportunity to examine some New World
dolichopodid types described by C.G. Thomson, 1869 from material gathered
during the voyage of the Swedish frigate Eugenie (see Persson, 1 97 1 for an au-
thoritative account of the collecting localities). Some of these species had been
overlooked in revisions of New World dolichopodid genera, and/or had been
omitted from major catalogues. Three of the species proved to be synonyms.
These matters are discussed below.

Dolichopus lamellicornis Thomson.

Dolichopus lamellicornis Thomson, 1 869: 511.

Hygroceleuthus afflictus Osten Sacken 1877: 313. NEW SYNONYM.

Dolichopus lamellicornis was overlooked in the principal revision of North
American Dolichopus (Van Duzee, et al., 1921) and subsequently was listed
under "Unplaced Species of Dolichopodidae" in the Catalog of the Diptera of
America North of Mexico (Foote, et al., 1965).

The lectotype of Dolichopus lamellicornis, here designated, is a female,
bearing the labels: "California/ Kinb./Type". The body length of the lectotype
is 6.0 mm, not 7.0 mm as stated in Thomson's description.

I used the key to female Nearctic Dolichopus (Van Duzee & Curran, 1934)
and the specimen was confidently identified as the widespread western North
American Dolichopus afflictus (Osten Sacken), an identification substantiated
by the description in Van Duzee, et al., 1921. Although the lectotype of

1 Received April 14, 1997. Accepted May 3, 1997.

2 Entomology Section, Australian Museum, 6 College Street, Sydney, NSW 2000 AUSTRALIA,
e-mail: danb@amsg.austmus.oz.au

ENT. NEWS 109(1): 61-65, January & February, 1998

62 ENTOMOLOGICAL NEWS

Dolichopus lamellicornis bears a locality label with only the word "Califor-
nia", the ship's records note that the only Californian port of call was San
Francisco, from July 3 to August 11, 1852, and the specimen probably was
collected in vicinity of that city. The type locality of D. afflictus is San Rafael,
Marin County, adjoining San Francisco Bay.

Osten Sacken 1877, in his description of Hygroceleuthus afflictus (Hygro-
celeuthus is a synonym of Dolichopus) thought that on the basis of Thomson's
description, D. lamellicornis was possibly the female of the species. He rea-
soned that if Thomson's female specimen had an indentation on hind wing mar-
gin similar to that of male D. afflictus, then the two species possibly were
conspecific, especially since both sexes of the related species D. crenatus had
indented hind wing margins. Since Thomson's description did not note any
wing indentation, Osten Sacken therefore considered them to be different spe-
cies. Therefore, it is noted here that the wing of the Dolichopus lamellicornis
female lectotype does have a marginal indentation at the apex of vein CuA, as
has been described for female D. afflictus.

The following three species were collected at Puna, an island in the Gulf of
Guayaquil, Ecuador, and are not listed in the "Catalogue of the Diptera of the
Americas south of the United States" (Robinson, 1970), nor in any other re-
cent Diptera catalog. Becker (1922a: 220) thought that the locality "Puna"
given in Thomson's descriptions was Puma (sic.), India, and therefore he listed
one of the species, Psilopus zonatulus (as Condylostylus zonatulus) in his mono-
graph on the Indo-australian fauna. None of the three species appears in his
treatment of the New World Dolichopodidae (Becker, 1922b).

Chrysotus ochropus Thomson.

Chrysotus ochropus Thomson, 1 869: 505.

Chrysotus flavipalpus Van Duzee, 1930: 76. NEW SYNONYM.

The lectotype, here designated, is a male in fair condition bearing the la-
bels "Puna/ Kinb./Type". A diagnosis based on the holotype is given below,
and the species is placed in synonymy with Chrysotus flavipalpus Van Duzee,
described from San Bartolome, Lima, Peru. In the following diagnosis, the
relative lengths of the podomeres are representative ratios, not measurements,
and for each leg are given in the following formula and punctuation: trochanter
+ femur; tibia; tarsomere 1/2/3/4/5; the abbreviations (MSSC) means "male
secondary sexual character(s)," the non-genitalic characters found only on the
male body.

Diagnosis: Male: length: 2.0 mm; wing: 1.7 x 0.6 mm.

Head: vertex and frons metallic blue-green with some grey pruinosity;
pairs of strong diverging ocellars, strong verticals, and short postverticals
present; eyes almost joined across face-clypeus, but separated by narrow band

Vol. 109, No. 1, January & February, 1998 63

of grey pruinose cuticle; palp yellow; proboscis brown; scape and most of
pedicel yellow; distalmost pedicel and first flagellomere brown; first
flagellomere subtriangular with apical arista; ventral postcranium with pale
postorbitals.

Thorax: mostly obliterated by pin, but metallic green with bronze reflec-
tions; setae black; lateral scutellar setae about 1/4 the length of medians.

Legs: coxa I yellow; coxae II and III brown; remainder of legs mostly
yellow; coxae I and II with pale anterior setae; coxa III with strong brownish
lateral seta; leg I: 2.7; 2.3; 1 .0/ 0.6/ 0.3/ 0.2/ 0.3; femur I in distal half with row
of 7 projecting brown pv setae (MSSC); tibia I with crestlike row 12-15 pale
ventral hairs which are longer than normal vestiture, and which continue as
row of short pale hairs on tarsus I (MSSC); leg II: 3.2; 2.8; 1.2/ 0.5/ 0.4/ 0.3/
0.3; femur II with group of 4-5 brown subapical pv setae (MSSC?); tibia II
with very strong ad and much weaker pd near 1/4, and with weak offset ad and
pd setae near 1/2, and with strong ventroapical seta; leg III: 3.5; 3.7; l.O/ 0.8/
0.4/ 0.3/ 0.3; femur III with 3-4 long brownish av and pv setae from 1/2 to 3/4
(MSSC), and in apical quarter with very strong av seta followed distally by
weaker seta; tibia III with ad and pd setal pairs at 1/5 and 1/2, and with some
short dorsal setae.

Wing: hyaline; 1*4+5 and M parallel to apex; CuAx ratio: 0.3; lower calypter
yellow with fan of brownish setae; halter yellow.

Abdomen: metallic blue-green with bronze reflections, and covered with
short brownish vestiture; hypopygium with dark brown capsule and yellow
cercus.

Remarks: Chrysotus ochropus is now known from coastal Ecuador and
Peru. It is closely related to C. brevicornis Van Duzee, found in the Galapagos,
Central America and Mexico. Both species have a similar overall description
and coloration, and have similar leg setation: pv setae on femur I (MSSC),
tibia I with row of pale ventral hairs (MSSC), similar tibia II setation, femur
III with long brownish pv setae from 1/2 to 3/4 (MSSC). They belong in the
picticornis group (see Bickel & Sinclair, in press, for further discussion).

Condylostylus longicornis (Fabricius)

Musca longicornis Fabricius, 1775: 783.

Psilopus zonatulus Thomson, 1 869: 509. NEW SYNONYM

Condylostylus zonatulus (Thomson) [Becker, 1922a: 220].

The lectotype of Psilopus zonatulus, here designated, is a female bearing
the labels "Puna/ Kinb./Type". It is conspecific with Condylostylus longicornis
(Fabricius), agreeing in every detail with this common and widespread New
World species. The earlier synonyms and distribution of this species are dis-
cussed in Bickel (1994: 108).

64 ENTOMOLOGICAL NEWS

It should be noted that Psilopus zonatulus Thomson is also a junior hom-
onym of the Palearctic Psilopus zonatulus Zetterstedt, 1843, a valid species in
the genus Sciapus (see Meuffels & Grootaert, 1990).

Condylostylus pleuralis (Thomson).

Psilopus pleuralis Thomson, 1869: 510.

Psilopus pleuralis is known only from the female lectotype, here desig-
nated, which bears the labels the labels "Puna/ Kinb./ Type". It was referred to
Condylostylus in Bickel (1994: 107).

This species clearly belongs in the widespread New World group of spe-
cies centered around Condylostylus caudatus (Wiedemann) (see Becker, 1922b
and Van Duzee, 1927). However, females in this group cannot be accurately
identified. Previously, C. caudatus itse\f was thought to be widespread through-
out both North and South America, but both Van Duzee (1927) and Robinson
( 1 970, 1 975) regard most of the tropical records as being that of C. graenicheri
(Van Duzee) or some other related species. Without further study, it is prema-
ture to make a decision, but C. pleuralis is possibly a senior synonym of C.
graenicheri (Van Duzee) or C. barbatus (Aldrich), or a junior synonym of two
other South American caudatus group species, C. anceps (Wiedemann) and
C. smaragdulus (Wiedemann). Unfortunately, both Wiedemann species are
described from females and remain unidentifiable. Until the entire New World
caudatus group is revised, there will be uncertainty regarding names. The large
number of old species based on unidentifiable females is regrettable, espe-
cially since they have the potential to become senior synonyms of subsequent
well-described species.

ACKNOWLEDGMENTS

I thank Bert Viklund of the Naturhistorika Riksmuseet, Stockholm, for kindly making the
Thomson material available to me. F.C. Thompson, United States National Museum, Washing-
ton, D.C., and C.E. Dyte, University of Reading, England, refereed the manuscript and provided
valuable comments.

LITERATURE CITED

Becker, T. 1922a. Dipterologische Studien, Dolichopodidae der indo-australischen Region. Capita

Zoologica 1(4): 1-247.
Becker, T. 1922b. Dipterologische Studien. Dolichopodidae. B. Nearktische und neotropische

Region. Abhandlungen der Zoologische-Botanische Gesellschaft in Wien (1921) 13(1): 1-

394.
Bickel, D.J. 1994. The Australian Sciapodinae (Diptera: Dolichopodidae), with a review of the

Australasian and Oriental faunas, and a world conspectus of the subfamily. Records of the

Australian Museum , Supplement 21: 1-394.

Vol. 109, No. 1, January & February, 1998 65

Bickel, D.J. & B.J. Sinclair, (in press). The Dolichopodidae (Diptera) of the Galapagos Islands,

with notes on the New World fauna. Entomologica Scandinavica.
Fabric ins, J.C. 1775. Systema entomologiae. 832pp. Flensburgi et Lipsiae.
Foote, R.H., J.R. Coulson, & H. Robinson 1965. Family Dolichopodidae, p. 482-530, In: A.

Stone et al., eds. A Catalog of the Diptera of America North of Mexico. United States

Department of Agriculture, Agricultural Handbook 276, 1696 pp.
Meuffels, H.J.G. & P. Grootaert. 1 990. The identity of Sciapus contristans (Wiedemann, 1817)

(Diptera: Dolichopodidae), and a revision of the species group of its relatives. Bulletin de

1'Institut Royal des Sciences Naturelles de Belgique 60: 161-178.
Osten Sacken, C.R. 1877. Western Diptera: descriptions of new genera and species of Diptera

from the region west of the Mississippi and especially from California. Bulletin of the United

States Geological and Geographical Survey of the Territories. 3: 189-354.
Persson, P.I. 1971. "Eugenics resa". Localities, dates and labels of the insects collected during

the voyage around the world by the Swedish frigate "Eugenie" in the years 1851-1853.

Entomologisk Tidskrift 92: 1 64- 1 72.
Robinson, H. 1970. Family Dolichopodidae. In: N. Papavero, ed. A catalogue of the Diptera of

the Americas south of the United States. 40, 92 pp. Museu de Zoologia, Universidade Sao

Paulo, Brasil.
Robinson, H. 1975. Bredin-Archbold-Smithsonian biological survey of Dominica. The family

Dolichopodidae with some related Antillean and Panamanian species. Smithsonian Contri-
butions to Zoology 185: 1-141.
Thomson, C.G. 1 869. Diptera, pp. 443-61 4, In: Kongliga svenska fregatten Eugenics resa omkring

jorden under befal af C. A. Virgin, aren 1851-1853. 2 (Zoologi), I (Insekter), 617 pp.,

Stockholm.
Van Duzee, M.C. 1927 .Three new species ofPsilopus from North America, and notes on caudatus

Wied. (Dipt.: Dolichopodidae). Entomological News 38: 72-76.
Van Duzee, M.C. 1930. Dolichopodidae, Part 5(1), Pp. 1-92, pis. 1-3. In: Diptera of Patagonia

and South Chile. British Museum (Natural History), London.
Van Duzee, M.C., F.R. Cole, & J.M. Aldrich. 1921 . The dipterous genus Dolichopus Latreille

in North America. Bulletin of the United States National Museum. 1 16, 304 pp.
Van Duzee, M.C. & C.H. Curran. 1934. Key to the females of Nearctic Dolichopus Latreille

(Diptera). American Museum Novitates 684, 17 pp.

66 ENTOMOLOGICAL NEWS

DRY WEIGHT OF FRESH AND

PRESERVED SPIDERS
(ARANEIDA: LABIDOGNATHA) 1

Robert L. Edwards 2 , Wendy L. Gabriel 3

ABSTRACT: Data on the dry weight for 1 9 taxa (suborder Labidognatha) of fresh and preserved
spiders are presented. The variation in weight at length for individual species is also provided.
With the notable exception of the genus Tetragnatha, Family Tetragnathidae, the Family Theridi-
idae, and most of the genera of the Family Thomisidae, the families examined are similar to one
another in their weight-length relationships and are not readily separable on that basis. Dry
weight can be approximated using either the weight of fresh specimens or preserved material.
The 'typical' spider, based on the material examined, increases in length faster relatively than it
increases in weight, and dry weight decreases relative to fresh weight as length increases.

With the exception of the papers by Clausen, 1983, that included data on
fresh and dry weight-length relationships for nine species (five families), and
Breymeyer, 1967, for three species of the family Lycosidae, there is very little
general information available on the dry weight of spiders. This report serves
to increase the information available on spider weight and explores the degree
of difference between taxa from the weight-length perspective.

MATERIAL AND METHODS

The bulk of preserved material was collected in 1989 and 1990 in the Frances
Crane Wildlife Management area, Hatchville, Falmouth Township, Barnstable
County (Cape Cod), Massachusetts, in connection with another study (Edwards
1993). Collection details are provided therein. All were preserved in 75%
denatured ethanol and all had their alcohol replaced at least once, typically
within 48 hours of collection. The total length was measured from the clypeus
to the distal end of the abdomen using an ocular micrometer for specimens
< 1 2 mm in total length and vernier calipers for those > 1 2 mm. The total length,
as described above, was measured to the nearest 0.1 mm, and the specimens
damp dried on absorbent paper before weighing. Obviously distorted speci-
mens were not used. In those cases where the pedicel had elongated, the sepa-
ration of the thorax from the abdomen was measured and the total measure-
ment corrected accordingly.

The fresh material for this study was collected in the months of June through
September, 1996, from the same area and habitats as the preserved material
with one exception. The collection of Leucauge regnyi Simon, Family Tetra-
gnathidae, was made in Puerto Rico in September, 1996. All collections were

1 Received April 9, 1997. Accepted July 7, 1997.

2 Research Associate, United States National Museum, Box 505, Woods Hole, Ma. 02543

3 Wendy L. Gabriel, Northeast Fisheries Center, 166 Water St., Woods Hole, Ma. 02543

ENT. NEWS 109(1): 66-74, January & February, 1998

Vol. 109, No. 1, January & February, 1998

made in the afternoon, the spiders were immobilized in an ethyl acetate col-
lecting jar, identified and measured that day, refrigerated overnight at 3 C,
and weighed the following day on a Mettler A200 balance, accurate to 1 mg.
Following this, the material was oven-dried at a temperature of 40 C for seven
days. To check the efficacy of the drying regime, three samples of 50 or more
mixed species samples were dried for an additional seven days, with the great-
est additional loss of weight observed of less than 3% in all cases. Specimens
of all species used in this study have been deposited in the United States Na-
tional Museum.

RESULTS AND DISCUSSION

The families, genera, number of individuals weighed, and the range of
total lengths are provided in Table 1 . A total of 2,315 measurements of fresh,
dry fresh, and preserved specimens representing 78 genera and 17 families
were made (Table 1). Note in Fig. 1 that the various taxa are identified with
the first four letters of the taxon as listed in Table 1 .

DC
O

O
O

0.25

0.2-

o.i5-

t o.H

0.05-

FRESH

O DRY FRESH

-f DRY PRESERVED

n a

a a
a + a

a a

THOMTMAR LINY LEUC ARAN LYCO SALT PISA PHOL TETR

THEROXYO PHIL ANYPAMAUCLUB CORI AGEL GNAP

TAXA

Figure 1 . Condition factors for the taxa listed in Table 1 , using the equation k = (aL h ) I L 3 . The
taxa are identified using the first four letters of each taxon. There are values for each treatment
(fresh, dry fresh, dry preserved) with the exception of AMAU, TMAR, PHOL, and OXYO, for
which there was no dry preserved data available.

68 ENTOMOLOGICAL NEWS

The log transformed least squares equation, In weight = lna + b(ln length),
was used to estimate weight (mg) at length (mm). The statistical parameters
are provided in Table 2. The coefficient of determination (r 2 ) ranged from
0.743 (Oxyopidae, dry fresh) to 0.985 (Corinnidae, fresh). Averages, mini-
mum, and maximum values are listed for all parameters. The average values
for coefficient In a (intercept) varied considerably and were greatest
(-1.976) for fresh and least for dry preserved specimens (-3.580), while the
average values for exponent b (slope) varied relatively little, from 2.739 for
dry preserved to 2.800 for dry fresh specimens (Table 2).

The number of individuals and genera obtained for each family varied con-
siderably (Table 1). It is clear from the statistical parameters presented in
Table 2 that there was little difference in the weight-length relationships be-
tween taxa. To compare weight at length using different taxa, while taking
into consideration the differing length ranges over which the parameters were
estimated, we calculated the condition factor for each taxon, k = (aL b ) I L 3 (L
= mid-point of lengths (mm) in sample, a = Exponent In a) and the results
shown in Fig. 1 . Of the 1 9 taxa the genus Tetragnatha Latreille (Tetragnathidae),
the family Theridiidae, and the rotund crab-like members of the family
Thomisidae (genera Xysticus C. L. Koch, Ozyptila Simon, Misumena Latreille,
Misumenops F.O.P.-Cambridge, and Misumenoides F.O.P.-Cambridge) stand
apart from the rest. The genus Tmarus Simon, family Thomisidae, is rela-
tively slender (less crab-like and rotund), differing in this respect from the
other genera of the family Thomisidae listed above and is plotted separately
(TMAR). Similarly, the less elongate genera Leucauge White, and Pachygnatha
Sundevall, family Tetragnathidae, differ in body form from the elongate mem-
bers of the genus Tetragnatha and are also plotted separately (LEUC). The
uniqueness of Tetragnatha was noted by Greenstone, et. al. 1985.

Breymeyer (1967) in a study concerning the dry weight of preserved spi-
ders reported that alcohol dissolves and extracts some parts of spider bodies.
This appears to be the case in this study as well. It is worth noting that undried
specimens preserved in denatured alcohol weigh considerably more than fresh
material (Edwards 1996). Clausen (1983, p. 143-144) noted that "the ratio of
dry over wetweight increases with decreasing size of specimens", and sug-
gested that "With decreasing size, the exocuticle may make up a relatively
greater part of the animal's weight because of the relatively greater surface.
And, there may be a minimum thickness of the cuticle, which, in effect, will
give the same result." To test Clausen's (op. cit.) suggestion further, the entire
data set available was examined by treatment, i.e. fresh, dry fresh and dry
preserved. The statistical parameters for each treatment are provided in Table
3. The percent of fresh weight for dry fresh and dry preserved material is shown
in Fig. 2. The results are consistent, for the average spider, with the observa-
tions of Clausen (op. cit.), with the caveat that more than the cuticle is un-

Vol. 109, No. 1, January & February, 1998

y 0.35
LJJ

X
CO
LJJ
DC

LJJ

DC
LJJ
Q.

0.25-

0.2-

0.15

10 15

LENGTH (MM)

DRY FRESH

DRY PRESERVED

Figure 2. Percent of fresh weight represented by dry fresh weight and dry preserved weight,
based on average data for each treatment (Table 3).

doubtedly involved since drying does not reduce the specimens to the cu-
ticle only. Other tissues are involved which may also vary in the degree to
which they are present in different taxa.

The variability seen in individual species in the ratio of dry weight to fresh
weight was examined for 1 1 species collected in October and November, 1 996.
The individual species collections were each made within an hour in restricted
localities to reduce environmental variabilities as much as possible. The col-
lections were treated and analyzed as described earlier (see Table 4). On aver-
age the ratio of dry to fresh weight was 0.314 0.038, varying from 0.250 to
0.390. The average slope (b) of dry on fresh weight was slightly in excess of
1 with two notable exceptions, that of Pardosa lapidicina (b = 1 .224) and
Phidippus clarus (b = 1 .537). The average intercept (In a) values for these two
species were also well in excess of the average value, -2.015 and -3.161 re-
spectively. Both of these collections were of immature individuals that would
have matured the following year, although the Tmarus angulatus collection,
also of immature individuals due to mature the following year, did not have a
similar departure from the average values. The average r 2 for the weight-
length regressions of these species was 0.892 for fresh and 0.864 for dry fresh.

ENTOMOLOGICAL NEWS

For studies requiring precision, e.g. those of a single or a set of closely
related species, it would be best to use a sclerotized part of the body such as
the head capsule, to reduce the problems associated with measurement error
(cf. Jocque 1981). Whatever method is used, it is obvious that the weight of
individual spiders is highly variable.

In one survey (Edwards 1993) over 12,000 specimens were collected. Many
of these were archived against the future. It was encouraging to find out that
preserved material also served the purpose of realistically estimating dry weight.

Table 1 . Fresh and preserved spiders examined. Number of individuals = n, number of genera -
genera, lengths (mm) included in sample - range. Family Tetragnathidae is subdivided into the
genus Tetragnatha and a second category of rounder body forms, including the genera Leucauge
and Pachygnatha, listed as Leucauge. The family Thomisidae includes all genera sampled
except the genus Tmarus which is listed separately.

Fresh

Fresh, dry

Preserved, dry

Taxon

n genera range

Agelenidae

4.5

- 19.1

7.4

- 19.1

3.7-

16.5

Amaurobiidae

4.0

- 14.1

4.0

- 14.1

Anyphaenidae

2.7

- 7.6

3.2

- 7.8

3.6-

9.2

Araneidae

2.7

-21.2

2.7

-20.5

2.3-

14.8

Clubionidae

2.3

- 8.8

2.5

- 11.1

2.0-

9.0

Corinnidae

2.2

- 8.6

3.2

-11.2

2.3-

7.1

Gnaphosidae

2.8

- 10.1

3.4

- 9.4

3.2-

11.6

Linyphiidae

1.5

- 5.5

2.5

- 5.4

2.0-

6.5

Lycosidae

1.5

- 16.8

4.0

- 16.8

2.6-

13.5

Oxyopidae

4.2

- 7.3

4.2

- 7.5

Philodromidae

2.0

- 9.0

2.9

- 12.5

2.0-

6.6

Pisauridae

5.5

- 19.3

4.0

- 11.1

2.1 -

12.0

Pholcidae

2.3

- 8.5

2.3

- 10.8

Salticidae

2.3

- 10.1

3.4

- 10.8

2.2-

9.0

Tetragnathidae

Tetragnatha

2.5

- 11.0

3.0

- 10.2

2.4-

8.5

Leucauge

2.5

- 7.6

2.8

- 7.6

1.5-

7.1

Theridiidae

1.5

- 8.3

3.0

- 8.2

1.7-

6.3

Thomisidae

1.9

- 8.6

2.6

- 8.2

1.6-

9.1

Tmarus

4.3

- 6.4

3.0

- 8.2

Totals

928 78

808 69

579 68

Vol. 109, No. 1, January & February, 1998

Table 2. Statistical parameters for spider weight-length equations (In weight ug = In a + b (In
length mm), for fresh, dry fresh and dry preserved material. No. of individuals - n, standard error
- SE., coefficient of determination - r% exponent of In a = Exp In a.

Fresh specimens

Taxon

aSE

& SE

r 2

Exp In a

Agelenidae
Amaurobiidae

-1.657 0.325
-2.303 0.1 50

2.553 0.1 13
2.999 + 0.077

0.888
0.984

0.191
0.100

Anyphaenidae
Araneidae

28
90

-1.697 0.186
-1.726 0.374

2.514 + 0.116
2.746 0.066

0.948
0.952

0.183
0.178

Clubionidae

- .928 + 0.212

2.636 0.1 13

0.951

0.145

Corinnidae

-2.002 0.1 11

2.595 0.074

0.985

0.135

Gnaphosidae
Linyphiidae
Lycosidae
Oxyopidae
Pisauridae

82
60
92
23
16

-2.492 0.237
-2.766 0.260
-1.746 + 0.277
-1.7060.147
-2.963 0.207

2.930 + 0.098
2.647 + 0.108
2.695 0.080
2.571 +0.209
3.272 + 0.177

0.918
0.919
0.926
0.878
0.961

0.083
0.171
0.174
0.182
0.052

Philodromidae

-1.707 0.233

2.740 0.1 35

0.947

0.181

Pholcidae

-2.453 0.1 54

2.905 0.076

0.984

0.086

Salticidae

-2.403 0.248

3.027 0.071

0.957

0.090

Tetragnathidae
Tetragnatha
Leucauge
Theridiidae

58
49

-2.268 0.240
- .966 + 0.237
- .456 0.222

2.431 0.113
2.853 0.1 27
2.839 0.071

0.892
0.915
0.958

0.103
0.140
0.233

Thomisidae

- .447 + 0.199

2.945 0.086

0.959

0.229

Tmarus

- .860 0.108

2.743 + 0.177

0.903

0.156

Average
Minimum

- .976 0.199
-2.963 0.1 08

2.771 0.1 10
2.431 0.066

0.938
0.878

0.148
0.052

Maximum

-1.447 0.374

3.272 + 0.209

0.985

0.233

Dry fresh specimens
Agelenidae
Amaurobiidae

29
26

-4.504 0.326
-4.045 + 0.221

3.184 + 0.308
3.1980.119

0.798
0.968

0.011
0.018

Anyphaenidae
Araneidae

31
84

-2.652 0.23 1
-2.401 +0.368

2.406 0.2 14
2.61 5 0.077

0.813
0.934

0.070
0.091

Clubionidae

-3.722 0.265

2.999 0.1 39

0.951

0.024

Corinnidae

-3.896 0.1 65

3.054 + 0.175

0.975

0.020

Gnaphosidae
Linyphiidae
Lycosidae
Oxyopidae
Pisauridae

43
43
83
42
16

-3.584 0.329
-2.761 0.260
-3.253 0.271
-3.473 0.269
-3.107 + 0.178

2.845 0.1 98
2.530 0.209
2.804 0.093
2.905 + 0.270
2.743 + 0.182

0.835
0.830
0.917
0.743
0.942

0.028
0.060
0.039
0.03 1
0.045

Philodromidae

-2.643 + 0.388

2.617 0.196

0.860

0.071

Pholcidae

-3.460 + 0.462

3.354 0.181

0.953

0.014

Salticidae

-3.330 0.289

2.904 0.1 20

0.875

0.036

Tetragnathidae
Tetragnatha
Leucauge

42
52

-2.350 0.285
-3.253 0.272

1.9140.158
2.920 0.1 63

0.785
0.866

0.095
0.039

ENTOMOLOGICAL NEWS

Taxon n

a + SE

bSE

Exp In a

Theridiidae 55

-3.436 0.3 17

3.22910.177

0.863

0.032

Thomisidae 52

-2.41410.329

2.741 10.147

0.874

0.089

Tmarus 29

-3.043 0.203

2.790 + 0.195

0.884

0.048

Average
Minimum

-3.22810.278
-4.50410.165

2.829 + 0.175
1.91410.077

0.871
0.743

0.046
0.011

Maximum

-2.35010.388

3.229 + 0.308

0.975

0.095

Dry preserved specimens
Agelenidae 52
Anyphaenidae 33
Araneidae 56

-5.38010.375
-3.284 1 0.267
-3.607 1 0.484

3.38610.145
2.482 1 0.224
3.00410.138

0.916
0.798
0.898

0.005
0.037
0.027

Clubionidae 19

-3.35610.253

2.45810.146

0.943

0.035

Corinnidae 19

-2.946 1 0.222

2.46310.122

0.960

0.053

Gnaphosidae 34
Linyphiidae 56
Lycosidae 53
Pisauridae 25

-4.380 1 0.297
-3.308+0.288
-3.54210.294
-3.64310.307

3.011 +0.193
2.64610.120
2.772 1 0.088
2.74410.101

0.883
0.900
0.951
0.970

0.053
0.037
0.029
0.026

Philodromidae 39

-2.801 10.363

2.474 + 0.213

0.784

0.061

Salticidae 49

-4.13910.280

3.10910.124

0.931

0.016

Tetragnathidae
Tetragnatha 3 1
Leucauge 32
Theridiidae 40

-3.59010.371
-3.58910.378
-2.952 1 0.280

2.182 + 0.181
2.79610.183
2.553 + 0.133

0.833
0.886
0.906

0.028
0.028
0.052

Thomisidae 41

-3.18410.210

3.001 10.087

0.968

0.041

Average
Minimum

-3.58010.311
-5.38010.210

2.73910.147
2.18210.087

0.902
0.784

0.032
0.005

Maximum

-2.801 +0.484

3.386 1 0.224

0.970

0.061

Table 3. Statistical parameters for spider weight-length equations for
treatment. Equation and table headings as in Table 2.

all available

material by

Treatment n

In a SE

b SE

Exp. In a

Fresh 928

-1.87410.447

2.73310.031

0.891

0.153

Dry fresh 808

-2.857 1 0.436

2.637 1 0.039

0.847

0.057

Dry preserved 579

-3.27910.537

2.581 10.046

0.829

0.038

Vol. 109, No. 1, January & February, 1998

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y 3

- OL \

fl
e

fi 1 ^

60_flJ "S

<L>
&o

'C "S.

i/5

2 .s 1

Agelenopsis

Callobius b

Pardosa lap

Oxyopes sal

Pholcus pha

Phidippus a

Leucauge re

Tetragnatha

Achaearane

Tmarus ang

r5 ">

| e J2

1 II

> I 1

Agelenidae.

Amaurobiidae.

Araneidae.

Lycosidae.

Oxyopidae.

Pholcidae.

Salticidae.

Tetragnathidae.

Theridiidae.

Thomisidae.
Means

74 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

The assistance of Eric and Annabel Edwards in the field was greatly appreciated. The com-
ments and suggestions of Jess Zimmermann, University of Puerto Rico, Los Piedras, and Stan
Sady, Massachusetts General Hospital, Boston, who kindly critiqued the initial manuscript were
most helpful. We thank the two anonymous reviewers for their comments and suggestions. Nancy
McHugh, Northeast Fisheries Center, Woods Hole, kindly made laboratory facilities available.

LITERATURE CITED

Breymeyer, A. 1967. Correlations between dry weight of spiders and their length and fresh weight.

Bull. acad. Pol. Sci. CI-II, XV(5):263-265.

Clausen, I.H.S. 1983. Weight-length relations of eight species of spiders (Araneae) from Den-
mark. Ent. Meddr 50: 139-144. Copenhagen, Denmark 1983.
Edgar, W. D. 1971. Seasonal weight changes, age structure, natality and mortality in the wolf

spider Pardosa lugubris in Central Scotland. Oikos 22:84-92.
Edwards, R. L. 1993. Can the species richness of spiders be determined? Psyche. 100:(3-4)185-

208.
Edwards, R. L. 1996. Estimating live spider weight using preserved specimens. J. Arachnol.

24:161-166.
Greenstone, M. H., C. E. Morgan and A. Hultsch. 1985. Ballooning methodology: equations

for estimating masses of sticky-trapped spiders. J. Arachnol. 13:225-230.
Jocque, R. 1981 . Size and weight variations in spiders and their ecological significance. Biol.

Jb. Dodonaea. 155-165.

Vol. 109, No. 1 , January & February, 1998 75

OBSERVATIONS OF INTERACTIVE BEHAVIOR

IN PARANDRA GLABRA
(COLEOPTERA: CERAMBYCIDAE) 1

Steven W. Lingafelter^

ABSTRACT: The first observed behavioral interactions for a species of Parandrinae (Coleoptera:
Cerambycidae) are reported. Two sets of observations between males and females of Parandra
glabra were made under artificial conditions: 1) individual male/female interactions; 2) interac-
tions of multiple males in the presence of each female. Results of individual interactions indi-
cate consistent patterns of behavior including antennation, mandibulation, genitalic exsertion,
mounting, and dorsal-oblique mating posture. Males display aggression in first encounters with
females, but subsequent antennation leads to mounting and copulation attempts. When more
than one male is present with a female, these behavioral categories are directed toward other
males more frequently than to the female.

Intraspecific interactions in longhorn beetles can range from violent,
indiscretionary encounters to complex, deliberate engagements, but the litera-
ture contains few descriptions of these. Most observations are simply noted in
more expansive papers concerning other biological or systematic issues.
Michelsen (1966) focussed on interactive behavior when he provided the most
detailed and widest coverage for behavior (especially with regard to courtship
and copulation) in longhorned beetles. He provided data on species represent-
ing four different subfamilies of longhorns (Aseminae, Spondylinae, Lepturinae,
Cerambycinae). His behavioral observations were placed into 25 categories
and a wealth of additional anecdotal information was included. Other refer-
ences are restricted to fewer (usually one) species and often are much more
general: Webster (1904) made interesting observations on Oberea ulmicola
Chittenden (Lamiinae); Goldsmith (1987a, 1987b, 1989) examined mating
systems of three species, Trachyderes mandibularis Dupont, Perarthrus linsleyi
(Knull), and Stenaspis verticallis arizonicus Casey (all Cerambycinae); Hughes
( 1 98 1 ) examined mating behavior in Monochamus scutellatus (Say) (Lamiinae);
Piper (1977) discussed mating behavior in Hippopsis lemniscata (Fabricius)
(Lamiinae); Chemsak (1965) commented on habits of Oeme costata LeConte
(Cerambycinae); Chemsak and Linsley (1971) observed mating behavior in
Rosalia funebris Motschulsky (Cerambycinae); and Wang, et al. ( 1 990) looked
at the complex mating behavior of Paraglenea fortunei Saunders (Lamiinae).
This study on Parandra glabra (DeGeer) represents the first information on
behavior in Parandrinae.

1 Received May 8, 1997. Accepted July 30, 1997.

2 Systematic Entomology Laboratory, PSI, ARS, USDAc/o U.S. Natural History Museum, NHB-
168, Washington, D.C. 20560.

ENT. NEWS 109(1): 75-80 January & February, 1998

76 ENTOMOLOGICAL NEWS

MATERIALS AND METHODS

This study was undertaken at Monteverde Biological Reserve, Monteverde,
Costa Rica. Three males and three females ofParandra glabra (DeGeer) were
collected at lights, measured, and their date of collection noted. Measure-
ments were taken of the total body length (from the base of the mandibles to
the apex of the elytra; body width (distance between elytral humeri); length of
mandibles (from the base to farthest point along medial axis) for correlation
analyses. These data are listed in Table 1 .

Table 1 . Data for specimens of Parandra glabra (DeGeer) used in this study.

Specimen code

(sex/#)

Length

Width

Mandible
length

Date of Capture

29mm

9.5mm

5.0 mm

22/May/1993

31 mm

9.5 mm

3.0 mm

22/May/1993

30 mm

10.0mm

5.0 mm

23/May/1993

28mm

9.0mm

2.5mm

25/May/1993

32mm

10.5 mm

3.0 mm

28/May/1993

32mm

10.0mm

6.0 mm

30/May/!993

Specimens were marked on the base of the right elytron with an indelible black-
ink marker. The beetles were simply coded with the number of the collection,
"1-6". The beetles were kept separately in a small plastic container which was
equipped with plastic dividers creating twelve separate holding cells. Damp
wood was placed in each cell with the beetles.

After several days of observing the beetles, their highest activity appeared
to be at night, from 22:00 - 02:00. Formal observations were made during this
period under low light conditions. Although specimens were strongly phototaxic
to acute sources of light, low and diffuse lighting did not appear to alter their
behavior.

Two sets of investigations were made. The first involved nine pair
comparisons of behavior between each male and female. The last focussed on
interactive behavior when each female was placed with all three males. Each
combination was observed for 20 minutes.

RESULTS

Observations: one on one male/female interactions

Noted behaviors included antennation, mandibulation, genital ic exsertion,
genitalic contact with substrate, mounting, and copulation. The number of
incidents (or merely presence/absence) of these behavioral activities for each
20 minute observation period is indicated in Table 2.

Vol. 109, No. 1, January & February, 1998 77

Antennation and mandibulation

Males and females each opened their mandibles widely after initial contact. In
most cases, the male relaxed and closed his mandibles after antennation of the
female. In two instances, males bit the females upon initial contact but this
was never repeated after further antennation.

Genitalic exsertion and substrate contact

In most cases, after prolonged antennation of the terminal abdominal segments
of the female, the males exserted their genitalia partially or distended the region
between the last two ventrites. This occurred an average of 2.9 times per male
for each 20 minute observation period. In most cases the males would rub
their genitalia on the substrate (the wood, container bottom, or container side)
and in several cases on the female (the prothorax, mandibles, or elytra). It was
distinctly obvious that muscular control of the genitalia, directing it downward,
was being affected, and in most cases, the males would reverse walking direction
several times during this behavior, rubbing the genitalia back and forth.

Mounting

Mounting of the female occurred an average of 2.2 times for each 20 minute
observation period. In two pair combinations it did not occur. Both of these
instances involved female #2, the only female not observed to copulate.
Mounting usually occurred after contact with the female and antennation by
the male, usually around her terminal abdominal segments. Usually in this
scenario the females continued walking, somewhat oblivious to the male
antennation. The male would then follow the female, maintaining contact and
would mount her. In many cases, mounting would occur from the side as the
female became "trapped" in a corner or against the piece of wood. This method
was similar to that observed commonly in R.funebris (Cerambycinae) (Chemsak
and Linsley, 1971). In some cases, after head-to-head contact between the
pair, the male climbed on top of the female opposite her orientation, and in
some cases, exserted his genitalia so it contacted her mandibles and prothorax.
Often, a mounting involved probing by the male genitalia in search of the
female's. The ratio of mountings to copulations was 4: 1 .

Copulation

Copulation is here defined as a visible, sustained contact between male and
female genitalia. Transfer of sperm to the female could not be determined and
thus was not a criterion for this category. Copulations were observed in four
of the nine pair combinations. In one pair combination (male #1 ; female #5),
copulation occurred twice during the twenty minute observation period. Copula-
tions ranged in duration from 50 to 240 seconds, averaging 101 seconds. I
interrupted a copulation in one instance as I was attempting to document it
with a photograph. In every instance, despite preliminary mounting orientation,
copulations occurred with the male atop the female in a slightly oblique position
(ca. 30 from female central axis). The males extended their terminal abdominal

78 ENTOMOLOGICAL NEWS

segments outward and probed with their genitalia for the female genital opening.
The males did not forcibly extrude the female ovipositor as has been witnessed
in other longhorn species (Michelsen, 1966). In many other longhorn species,
Michelsen (1966) observed the male to actually leave the female's dorsum
during copulation and in many cases face the opposite direction. During four
of the five copulations, the female showed motility. In the one copulation
where the female remained stationary (the second copulation of male #1 and
female #5), pulsation of the male occurred about 45 times (about once every
5.5 seconds). This pulsation behavior was also noted in R.funebris by Chemsak
and Linsley (1971). They witnessed this to last from 30 to 60 seconds and
occur about once every four seconds. During this time the female remained
motionless during the four-minute copulation. Interestingly, four of the five
copulations involved female #5; female #2 had no copulations; and female #4
had only one.

Observations: all males in presence of individual females

For each of the three females, all three males together were placed with her
and their actions were observed for about 20 minutes. Copulation was never
observed in any of the three sets of observations; genitalic exsertion and rubbing
on other males was observed in the presence of each female; same-sex mounting
was observed in each combination; and a few instances of aggression between
males was observed.

Female #2

All three males contacted and mounted one another. Antennation of another
male's sternites by each male was observed. Two instances of aggression were
observed (male #6 biting male #1 ; male #6 biting male #3). Genitalic exsertion
among the males was common during their juxtaposition, and they often rubbed
their genitalia on one another. Mounting of the female occurred only once (by
male #6); otherwise attention by males was directed toward other males.

Female #4

The same behaviors as above were noted with the addition of the following:
male #6 was seen rubbing his genitalia on the container. For a few minutes, all
the males were in contact with the female and began mounting one another
with their genitalia exserted.

Female #5

Little attention was given to her as the males continued to mount and congregate

with one another.

DISCUSSION

Because of the artificial conditions of this study and the small sample size
of individuals, interpretation and generalization of the observations is made

Vol. 109, No. 1, January & February, 1998 79

with caution. No correlations between size and exhibited behavior were evident
from this study. Antennation seems to play an important role in the identification
of females by males. Although observations of antennation were focussed on
and biased toward males, it is evident that antennae are used to a greater extent
by males than females. Obvious female antennation subsequent to a male
encounter was rarely observed. Based on the rapidity in which males determined
females by antennal contact (and then relaxed their mandibles and aggressive
behavior), the antennae may have a great sensitivity to some chemical exudate
of the female. Once the male established the presence of the female, subsequent
contacts were the stimulus for his genital ic exsertion. In many of these instances,
the male would rub his genitalia on the substrate and/or female. Based on
observations of this behavior in both sets of experiments, it is possible that the
male may be secreting some chemical, perhaps as a stimulus to cause the female
to be receptive for copulation or as a chemical indicator of his presence to
other males.

Further study, both in natural conditions and with individual, laboratory
reared specimens, is essential to understand the observations and interpretations
presented herein. The phenomena of sperm precedence, male and female
pheromones, body and mandible size correlations to mating success, and
average number of mountings for each copulation or number of copulations
required for successful sperm transfer, are interesting questions that may be
addressed upon further study of the Parandrinae.

Table 2. Summary of behaviors expressed by Parandra glabra (DeGeer) during the individual
male/female pair combinations. A - antennation; MA - mandibulation; GE - genitalic exser-
tion; GS - rubbing of genitalia on substrate; MO - mounting; C - copulation. For each category,
numbers indicate number of observations of behavior, except for antennation which occurred
numerous times in all pairs and is simply indicated with a checkmark. Duration of the five
copulations is measured in seconds.

Pair code
(sex/# X sex/#)

M1XF2

M1XF4

1 (50s)

M1XF5

2 (65s & 240s)

M3XF2

M3XF4

M3XF5

1 (60s)

M6XF2

M6XF4

M6XF5

1 (90s)

80 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

I thank the Department of Entomology (University of Kansas) for providing funds for field-
work in Costa Rica. M. Greenfield provided logistical support for this work. I thank T. A. Erwin,
D. G. Furth, R. S. Hanley, A. S. Konstantinov, A. L. Norrbom, and M. A. Solis for helpful sug-
gestions on this manuscript.

LITERATURE CITED

Chemsak, J. A. 1965. A new subspecies of Oeme costata with observations on the habits of

larvae and adults. Journ. Kans. Entomol. Soc. 38 (4):35 1-355.
Chemsak, J. A. and E. G. Linsley . 1 97 1 . Some aspects of adult assembly and sexual behavior of

Rosalia funebris Motschulsky under artificial conditions. Pan-Pac. Entomol. 47(2): 1 49- 1 54.
Goldsmith, S. K. 1987a. The mating system and alternative reproductive behaviors ofDendrobias

mandibularis (Coleoptera: Cerambycidae). Behav. Ecol. Sociobiol. 20: 1 1 1-1 15.
Goldsmith, S. K. 1987b. Resource distribution and its effect on the mating system of a longhorned

beetle, Perarthrus linsleyi (Coleoptera: Cerambycidae). Oecologia 73:317-320.
Goldsmith, S. K. 1 989. Feeding ecology and the mating system of Stenaspis verticalis arizonicus

Casey (Coleoptera: Cerambycidae). Journ. Kans. Entomol. Soc. 62(4): 528-533.
Hughes, A. L. 1981. Differential male mating success in the white spotted sawyer Monochamus

scutellatus (Coleoptera: Cerambycidae). Ann. Entomol. Soc. Am. 74: 180-184.
Michelsen, A. 1966. The sexual behavior of longhorned beetles. Entomol. Medd. 34:329-355.
Piper, G. L. 1977. Biology and habits of Hippopsis lemniscata (Coleoptera: Cerambycidae).

Coleop's. Bull. 31(3):273-278.
Wang, Q., Zeng, W. Y., and J. S. Li. 1 990. Reproductive behavior of Paraglenea fortunei

(Coleoptera: Cerambycidae). Ann. Entomol. Soc. Am. 83(4): 860-866.
Webster, F. M. 1904. Studies of the life history, habits, and taxonomic relations of a new

species ofOberea (Oberea ulmicola Chittenden). Bull. Illinois State Lab. Nat. Hist. 7 (1): 1-

14+2 pis.

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MARCH & APRIL, 1998

NO. 2

NTOMOLOGICAL NEWS

J\J ]" istruma memorialis (Hymenoptera: Formicidae),
^ new species of ant from Kentucky Cumberland

Plateau

Mark Deyrup 81

A new species of Aphaenogaster (Hymenoptera: Formi-
cidae) from upland habitats in Florida M. Deyrup, L. Davis 88

Laboratory rearing of Mesovelia cryptophila

(Heteroptera: Mesoveliidae) S.J. Taylor, J.E. McPherson 95

Two new species of microcaddisflies (Trichoptera: Hydrop-

tilidae) from Kentucky R.E. Houp, K.H. Houp, S.C. Harris 99

Seasonal flight periodicities of six microcaddisflies
(Trichoptera, Hydroptilidae, Glossosomatidae)
in the Brazos River, Texas, with notes on
larval biology D.C. Houghton, K.W. Stewart 103

A new species of Pseudocentroptiloides (Ephemeroptera:
Baetidae), with revisions to previously unnamed
baetid species from Texas NA. Wiersema, W.P. McCafferty 110

The adult of Moribaetis macaferti (Ephemeroptera:

Baetidae) W. P. McCafferty, C.R. Lugo-Ortiz 117

First report and new species of genus Cloeodes
(Ephemeroptera: Baetidae) from Australia

C.R. Lugo-Ortiz, W.P. McCafferty 122

Diel emergence patterns of Tricorythodes stygiatus
(Ephemeroptera: Leptohyphidae) on Little Lehigh
Creek near Allentown, PA F. Gough, B.L. Haase 129

First fossil Dorylinae with notes on fossil Ecitoninae

(Hymenoptera: Formicidae) Mark B. DuBois 136

Coccinellidae (Coleoptera) in apple orchards of
eastern West Virginia, and impact of invasion
by Harmonia axyridis M.W. Brown, S.S. Miller 143

SOCIETY MEETING OF OCTOBER 22, 1997

BOOKS RECEIVED AND BRIEFLY NOTED

NOTE FROM EDITOR RE PRIORITY OF NEW
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152
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152

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Vol. 109, No. 2, March & April, 1998 81

SMITHISTRUMA MEMORIALIS (HYMENOPTERA:

FORMICIDAE), A NEW SPECIES OF ANT FROM THE

KENTUCKY CUMBERLAND PLATEAU 1

Mark Deyrup^

ABSTRACT: A new species of dacetine ant, Smithistruma memorialis, is described. It was found
on top of a ridge on the Cumberland Plateau in Kentucky. This species is distinguished from
other North American species of Smithistruma by its short, erect, unmodified hairs on the oc-
ciput, body and legs. It appears to be a member of the pulchella group of Smithistruma. It is
named in honor of the late William L. Brown, Jr., who greatly advanced our knowledge of dacetine
ants.

The genus Smithistruma was described in 1948 by a pioneer of ant system-
atics, the late William L. Brown, Jr., and his revision of the Nearctic species
(1953) was so competent that it still serves well, almost 45 years after its pub-
lication. The species described below is dedicated to Dr. Brown, in gratitude
for his original work in bringing order to our knowledge of dacetine ants, and
making the systematics and biology of these ants accessible to other natural-
ists. This was only one of Dr. Brown's many contributions to myrmecology,
biogeography, and evolutionary biology.

Smithistruma includes about 104 described species (Bolton 1995). There
are probably additional species to be discovered, as these ants are small (work-
ers about 2 mm long), slow-moving, and usually hidden in leaf litter, rotten
wood, or soil. Even in the eastern United States, where the cryptic ant fauna is
relatively well known, there are a number of species that are rare in collec-
tions, and it is still possible to find undescribed species.

Smithistruma memorialis Deyrup, NEW SPECIES

Diagosis. Distinguished from all other Nearctic Smithistruma by the presence
of abundant, erect, unmodified hairs on the occiput, body and legs, and the
lack of reclinate widened hairs in these areas (Fig. 1 ). The clypeus is similar to
that of S. missouriensis (M. R. Smith) in shape and arrangement of enlarged
hairs.

Description. Holotype worker. Measurements in mm: Total length: 1 .98; head length: 0.5 1 ; maxi-
mum head width: 0.40; length of alitrunk: 0.46.

Features described below as in Fig. 1 . Head with preocular laminae not continuing the out-
line of the occipital lobes, so outline of head in frontal view not cuneiform. Mandible in lateral
view not tapering, but abruptly decurved at tip; mandibular diastema conspicuous at full closure;

1 Received June 25, 1997. Accepted August 1, 1997.

2 Archbold Biological Station, P. O, Box 2057, Lake Placid, FL 33862

ENT. NEWS 1 09(2) 8 1 -87, March & April, 1 998

ENTOMOLOGICAL NEWS

mandibles with four principle teeth in the subapical series, the second largest, the first next larg-
est, the fourth next largest, the third smallest. Clypeus with the apex rounded and thickened,
central area of clypeus minutely reticulate, completely bare, slightly raised and diamond shaped;
each side of clypeus with three small spoon-shaped hairs on the edge of the basal external corner,
two large, spoon-shaped, anteriorly directed hairs on the anterior margin, at points about one-
fifth and one half the distance to the apex, a large, spoon-shaped, posteriorly directed submedian
hair, and two small, submarginal, submedian spoon-shaped hairs. Antennal scapes with enlarged,
erect, untapered hairs on the inner margin as follows: one subbasal, directed slightly toward the
apex of the scape, a series of four or five in descending size, more or less evenly spaced, directed
slightly toward the base of the scape. Frontal area with small, sparse, inconspicuous, medially
directed, reclinate spoon-shaped hairs. Occiput with sparse, erect, straight, blunt hairs, no espe-
cially elongate or widened hairs on the lateral margins of the occiput. Upper part of head finely
reticulate as in other members of the pulchella group, with inconspicuous, sparse, longitudinal
rugae.

Pronotum finely reticulate dorsally, sparsely and evenly covered with erect hairs, no elon-
gate or otherwise distinctive hairs in the humeral area, smooth and shining laterally, convex in
profile. Remainder of alitrunk with dorsum finely reticulate, with a few erect hairs, lateral areas
smooth and shining. Propodeal teeth short, broad, infradental carinae narrow, evenly concave.
Legs, including tarsi, with sparse, suberect hairs, without reclinate enlarged or spatulate hairs, no
outstanding elongate curved or crimped hairs on the apical third of the mid or hind tibiae or
basitarsi.

Petiole finely reticulate dorsally, with sparse suberect hairs, infrapetiolar lamina narrow,
without a basal lobe or extension, spongiform process small, in lateral view not extending to
lower margin of petiole. Postpetiole with dorsal suberect hairs, lower spongiform process small,
not extending down past upper half of basal face of first stemite of gaster. Caster with sparse,
short erect hairs on dorsal surface, long dorsal hairs absent.

Figure 1 . Smithistruma memorialis, new species

Vol. 109, No. 2, March & April, 1998 83

Paratype female. Measurements in mm: Total length: 2.45, head length: 0.68, maximum head
width: 0.47, length of alitrunk: 0.57.

Usual queen modifications present: ocelli present, compound eyes large, alitrunk modified
for flight. Otherwise, queen resembles worker, including abundant short, erect hairs on dorsum
of thorax, gaster and legs. This vestiture distinguishes this specimen from queens of other dacetine
species.

Paratypes. Paratype material is 61 workers and one queen.

Collecting data for type material. All type material shares the same data: KENTUCKY: Laurel
Co., Daniel Boone National Forest, Bald Rock picnic area; 23 March 1997; collected by Stephen
and Mark Deyrup. Habitat open, grassy, with low herbs, scattered large pines. Nest with holotype
worker, allotype queen, and 53 workers found (by Stephen) in a small chamber in clay soil a few
centimeters below the surface near the base of a large pine, near the restrooms; 8 workers from a
small soil sample about three meters away. Other ants with nearby nests were: Aphaenogaster
tennesseensis (Mayr) (dealate queen), Brachymyrmex depilis Emery, Leptothorax pergandei
Emery, Mornomorium minimum (Buckley), Paratrechinafaisonensis Forel, Ponera pennsylvanica
Buckley, Prenolepis imparts (Say), Smithistruma pulchella (Emery) (dealate queen), Solenopsis
carolinensis (Forel).

Deposition of type material. Holotype, allotype, 12 paratypes: Museum of Comparative Zool-
ogy, Harvard University, Cambridge, Mass.; 6 paratypes: National Museum of Natural History,
Smithsonian Institution, Washington, D.C.; 6 paratypes; The Natural History Museum, London;
6 paratypes: Los Angeles County Museum of Natural History; 6 paratypes: Florida State Collec-
tion of Arthropods, Gainesville, Fla.; 3 paratypes, collection of Lloyd Davis, Gainesville, Fla.; 3
paratypes: Collection of Mark DuBois, Washington, III.; 3 paratypes, collection of William
MacKay, El Paso, Tex.; remaining specimens in the collection of the Archbold Biological Sta-
tion, Lake Placid, Fla.

Etymology. The specific epithet, translated "of remembrance," dedicates this
species to Bill Brown in place of the more usual patronym. For some years Bill
Brown had held an antipathy toward patronyms. His main objection, as far as
I can tell, was that patronyms often honor people who have only the most
trivial association with the species bearing their name, and in the worst cases
could be assigned to stoke the egos of sponsors or patrons. I do not think these
objections would apply in the case of an honoree who had made great and
lasting contributions to our knowledge of a genus, especially when the name is
applied posthumously. Nevertheless, I am respecting his feelings by avoiding
a direct patronym.

DISCUSSION

Myrmecologists who deal with dacetines in general and with Smithistruma
in particular place a well-justified faith in the taxonomic value of the elabora-
tions of the clypeus and mandibles as species-specific character states. In the
absence of any plausible theories explaining the remarkable diversity of these
features, it is easy to develop an illogical feeling that their biological function
is also associated with species recognition, like the modified palps and facial

ENTOMOLOGICAL NEWS

hairs that distinguish certain male dolichopodid flies in their courtship antics.
In S. memorialis we see a species that might never have been recognized on
the basis of its clypeal structure, which is similar to that of the variable species
S. missouriensis (Fig. 3). This is a useful, if somewhat worrisome reminder
that in Smithistruma the shape of the clypeus and the pattern of its pilosity
could remain constant in a group of related species.

Smithistruma memorialis clearly belongs in Brown's pulchella group, along
with missouriensis, reflexa, and cloydi. Smithistruma memorialis (Fig. 1),
reflexa (Fig. 2), and missouriensis have mandibles that in lateral view are broad
and abruptly decurved in their apical third, while the mandibles of pulchella
(Fig. 4) and cloydi (Fig. 5) are slender and tapering. Smithistruma pulchella

Figure 2. Smithistruma reflexa (Wesson and Wesson)

Figure 3. Smithistruma missouriensis (Smith)

Vol. 109, No. 2, March & April, 1998

Figure 4. Smithistruma pulchella (Emery)

Figure 5. Smithistruma cloydi Pfitzer

86 ENTOMOLOGICAL NEWS

and cloydi differ markedly in the shape of the clypeus and in the number and
arrangement of clypeal hairs (Figs. 4, 5). Smithistruma reflexa and missouriensis
are distinguished by differences in the orientation of clypeal hairs (Figs. 2, 3),
a character that is somewhat variable, and reflexa could be a junior synonym
of missouriensis (Brown 1953); the small series of missouriensis that I have
studied appears virtually identical to reflexa in lateral view. In Brown's 1953
key, memorialis keys to missouriensis (second half of couplet 23), but is easily
distinguished by its bristling hairs. Now that it is clear that species recognition
in this complex may depend on more than clypeal hairs, other structural char-
acters should be examined more closely, and we should be cautious in synony-
mizing species.

The described native species of Smithistruma in the southeastern U. S.
now number 24, and there are at least two additional undescribed species
(Deyrup and Cover, MS in preparation), bringing the known species from the
region to 26. Southeastern North America is therefore a major center of diver-
sity for the genus, and Brown (1953) suggested that this fauna is most closely
related to that of Asia, rather than to the Neotropical fauna, as had been sug-
gested earlier. Ward (1988) described three species from relict warm mesic
areas in the southwestern U.S. This pattern closely matches the distribution of
relict concentrations of warm temperate arcto-tertiary flora (Raven and Axelrod
1978). The genus Smithistruma, therefore, appears to be our only clear ex-
ample among the ants of a diverse assemblage left over from the gloriously
speciose warm temperate forests of the Miocene. Most of the known species
of North American Smithistruma are quite widely distributed through the mixed
deciduous forests of the Middle Atlantic states, south through north Florida,
and west into the more mesic woodlands of eastern Texas. If, however, the
genus Smithistruma in North America mirrors the distribution of arcto-tertiary
flora, there are probably some species confined to isolated habitat types in the
southern Appalachians. The species described here, found on top of a ridge on
the Cumberland Plateau, could represent such a species.

It seems appropriate to place memorialis in Brown's genus Smithistruma,
even though there are some indications that this genus may disappear in a
small implosion of dacetine genera. Unfortunately, Bill Brown will not be
around to offer his comments. In a general way, he felt that some changes were
necessary: in a recent (2 February, 1997) letter he stated, "...there is no doubt
that generic slaughter is overdue." However, he also wrote, in the same letter,
"...I hate to see all the names go down, and I'll be watching." As most contem-
porary myrmecologists know, Bill Brown thought that defining a genus on a
strictly phyletic basis could lead to a foolish nomenclature. Paraphyletiphobia
was not one of his afflictions. In his address at the 1987 meeting of the Ento-
mological Society of America, he publicly expressed the opinion that super-
specific names, such as the names of genera, are inevitably derived by an exer-

Vol. 109, No. 2, March & April, 1998 87

cise of judgement, and these judgements should be openly informed by ecol-
ogy and convenience, as well as by phylogenetics. He strongly objected to
examples of supposedly objective phyletic nomenclature that were really de-
rived from the secret manipulation of an arcane analysis. I wish I could re-
member all his much more humorous private comments at this convention.
There was one particularly funny comparision of some cladistic taxonomy to
astrology, in which all the character states and accomplishments of life are
subsidiary to, and mystically influenced by, the exact moment of separation
from the mother.

Within the narrow confines of a paper describing one species of dacetine
ant, I have tried to touch on the extreme importance of Bill Brown to our
understanding of the systematics and biogeography of this group. I would not
want to leave the impression that the study of dacetines will falter; for some
years the tremendously talented and energetic Barry Bolton of The Natural
History Museum, London, has been carrying on the work of large-scale revi-
sions of dacetines. What we have lost is the presence of the man, fantastically
knowledgeable, honest, critical, humorous, creative and intuitive, who could
inspire both nervousness and relief when he said, "I'll be watching."

ACKNOWLEDGMENTS

I am proud to acknowledge our son Stephen as the collector of the large nest series of
Smithistruma memorialis I also thank the custodians of the Daniel Boone National Forest for
watching over the beautiful and diverse habitats of the area, where I am sure that many more
undescribed species of interesting insects will be found.

LITERATURE CITED

Bolton, B. 1995. A new general catalog of the ants of the World. Harvard Univ. Press, Cam-
bridge, Mass. 504 pp.

Brown, W. L., Jr. 1948. A preliminary generic revision of the higher dacetini. Trans. Amer.
Entomol. Soc. 74: 101-129.

Brown, W. L., Jr. 1953. Revisionary studies in the ant tribe Dacetini. Amer. Midi. Natur. 50: 1 -
137.

Raven, P. H., and D. I. Axelrod. 1978. Origin and relationships of the California flora. Univ.
Cal. Publ. Bot. 72: 1-134.

Ward, P. S. 1988. Mesic elements in the western Nearctic ant fauna: taxonomic and biological
notes on Amblyopone, Proceratium, and Smithistruma ). Kans. Entomol. Soc. 61: 102-124.

88 ENTOMOLOGICAL NEWS

A NEW SPECIES OF APHAENOGASTER

(HYMENOPTERA: FORMICIDAE) FROM UPLAND

HABITATS IN FLORIDA

Mark Deyrup^, Lloyd Davis^

ABSTRACT: Aphaenogaster umphreyi, n. sp., is described from sandy uplands of peninsular
Florida. It appears to be closely related to A.fulva Roger. The new species, which may be entirely
subterranean in habits, is characterized by unusually small eyes, coarse sculpture, short propodeal
spines, and unusually small hind tibial spurs.

The genus Aphaenogaster tends to accumulate taxonomic problems. A
number of species show conspicuous variation between populations, within
populations, and even within colonies, while camouflaged in this tangle of
intraspecific variation are cryptic species that can only be detected by the most
sophisticated methodology, such as that used by Umphrey (1996) for the A.
rudis group. One species that emerged some time ago from the taxonomic
thicket \sA.fulva Roger, which, once divested of the set of "varieties" that are
now recognized as the rudis group, seemed to be a single, easily recognized
species (Creighton, 1950). It now appears that there is a second species that
shares most of the features previously ascribed solely to fulva. Fortunately,
since this species is rare, or at least difficult to find, it is unlikely to have been
the basis of many (if any) published records of fulva.

Aphaenogaster umphreyi, Deyrup and Davis, NEW SPECIES

Figure 1

Description: Holotype worker measurements (mm): head length (anterior edge of clypeus to
occiput) 1.15; head width (above eyes): 0.95; malar space ( = distance from lower edge of eye to
mandible in lateral view): 0.38; length of eye: 0.15; distance from propodeal spiracle to tip of
propodeal spine: 0.31.

In frontal view, head with convex vertex; coarse reticulate rugae covering dorsum and sides
of head, including occiput; venter of head with prominent carinae diverging from midline. Man-
dible, antenna, and clypeal area resembling those of fulva (cf. Figs. 1 and 2).

Mesosoma with strongly raised rugae on the pronotum, mesonotum, and propodeum, these
rugae zigzagging, not smoothly undulating, except less elevated and more undulating on pronotal
disc. Propodeal spine short, compared to that of fulva (Fig. 2), strongly upturned, approaching a
right angle with long axis of propodeum. Legs generally similar to fulva, including transverse
ridges on front coxae, except hind and middle tibial spurs reduced, shorter than width of basitarsus
on respective legs.

1 Received June 6, 1997. Accepted August 30, 1997.

2 Mark Deyrup, Archbold Biological Station, P. O. Box 2057, Lake Placid, FL 33862.

3 Lloyd Davis, 3920 NW 36th Place, Gainesville, FL 32606.

ENT. NEWS 109(2) 88-94, March & April, 1998

Vol. 109, No. 2, March & April, 1998

Figure 1 . Aphaenogaster umphreyi, new species, worker.

1mm

Figure 2. Aphaenogaster fulva, worker, Florida specimen.

90 ENTOMOLOGICAL NEWS

Petiole and gaster similar lofulva (cf. Figs. 1 and 2), except petiole with more conspicuous
rugae.

Color reddish brown, legs and gaster yellowish brown.

Diagnosis. Similar to A.fulva, but differs in having much smaller eyes, shorter
propodeal spines, coarser and more extensive sculpture on the head and
mesosoma, more convex vertex in frontal view, and reduced hind tibial spurs
(compare Figs. 1 and 2).

Type material. Holotype worker: FLORIDA, Putnam Co., 3 miles east of Melrose, 20 Aug.
1995 (Lloyd R. Davis), Ordway Preserve, sandhill habitat, nest in ground, at base of small oak;
deposited in Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts.

Paratypes: FLORIDA: 22 workers from nest series of holotype; same site and collector as
holotype: 1 worker: 1 Oct. 1995; 1 worker: 24 Feb. 1995; 1 worker: 27 Aug. 1995; 1 worker: 6
Sept. 1996; 2 workers: 14 Sept. 1996; 2 workers: 3 Mar. 1995. One worker: Highlands Co.,
Archbold Biological Station, 16 May 1988 (M. Deyrup), sifted from sand, Florida scrub habitat;
1 worker: Highlands Co. Sebring, 11 Mar. 1987 (M. Deyrup), Red Water Lake, Florida scrub
habitat; 2 workers: Highlands Co., Sebring 17 Sep. 1990 (M. Deyrup), Flamingo Villas develop-
ment, Florida scrub habitat; 10 workers (callows, nest series): Marion Co., 16 Oct. 1990 (M.
Deyrup), Ocala Waterway development, Florida scrub habitat; 3 workers: Alachua Co., 5.5 miles
west of Gainesville (L. Davis), in soil beside rotten pine log, open oak woodland, 22 Mar. 1992;
6 workers: Highlands Co., Placid Lakes Development (M. Deyrup), 1 Jan. 1997, Florida scrub
habitat, collector's yard, root mat below Quercus inopina.

Deposition of paratypes: 4: Museum of Comparative Zoology, Harvard University; 5: Na-
tional Museum of Natural History, Smithsonian Institution, Washington, D.C.; 5: Florida State
Collection of Arthropods, Gainesville; 5: The Natural History Museum, London; 4: Los Angeles
County Museum of Natural History; 3: collection of Gary Umphrey, London, Ontario; 2 paratypes:
collection of Mark Dubois, Washington, Illinois; 2: collection of Kye Hedlin, Raleigh, North
Carolina; 2: collection of William MacKay, El Paso, Texas; 5: collection of Lloyd Davis,
Gainesville, Florida; 17: Arthropod Collection, Archbold Biological Station.

Etymology. This species is named in honor of Dr. Gary Umphrey, in recognition
of his long labors working to elucidate the taxonomy and phylogeny of the
intractable A. rudis group.

DISCUSSION

Although we have known of specimens of an aberrant Aphaenogaster for
almost nine years, we were wary about assigning them to a new species be-
cause of the notorious intraspecific variation within the genus, and the result-
ing history of synonymy. Our hypothesis was that there might be a southern
isolate offulva that differed in various ways from northern forms, and it was
not until we had a good series of the new species from within the range of
fulva in north Florida (Fig. 3) that this hypothesis became untenable. We had
also hoped to find many more colonies, and associated sexuals, though these
aims still elude us. Meanwhile, we gathered specimens offulva from much of
its range, so that variation within that species is now clearer to us.

Vol. 109, No. 2, March & April, 1998

The evidence that we use to establish the species-level distinctness of
umphreyi is as follows: 1 . A. umphreyi is sympatric with/w/va, at least in Putnam
and Marion Counties, and probably farther north as well, so umphreyi is not
likely to be a geographic subspecies of fulva. 2. There is no overlap in the
following structural character states used to distinguish the new species: rela-
tively small eyes (Fig. 4); relatively small propodeal spines; heavy zigzag cari-

Aphaenogaster fulva

Aphaenogasler umphreyi

Figure 3. Collection sites for Aphaenogaster umphreyi and A. fulva in Florida.

ENTOMOLOGICAL NEWS

nae on the sides of the pronotum and on the propodeum; reduced spurs on the
middle and hind tibiae. 3. The occurrence ofumphreyi in xeric habitats, while
fulva (at least in the southeast) is in mesic, often wet sites. 4. The kinds of
morphological differences between the two species go far beyond the kinds of
intraspecific variation that seem to be directly influenced by environmental
conditions in different habitats (e.g.: in xeric habitats Pheidole dentata Mayr
seems paler, Odontomachus brunneus (Patton) paler and smaller).

The evidence available suggests that umphreyi and fulva are a closely re-
lated species pair. They share exclusively the following character states: coarse
sculpture on the head and mesosoma; upward-pointing propodeal spines;
strongly elevated, notched anterior edge of the mesonotum. This combination
of features brings umphreyi out to couplet 18 in Creighton's key (1950) to
Aphaenogaster, but the short propodeal spines produce an impasse.

Color is not a very reliable character in fulva; it is generally dark brown,
but we have seen reddish specimens, particularly from its western range. It

26 XvIylvXvXvXvXvXvXvXvIvXvXvXv

CD
C
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.25
.24
23
.22
.21
.20
.19
.18
.17
16
15
14
.13
.12
.11

f3 f2

f12 f7

'5 'vX:X;

te . : : : x : x :

111

U 7

us ::::::

u8 u6

u 1 2 u11u9 u 1

u3
u10

.27 28 29 .30 31 .32 .33 .34 35 .36 37 .38 39 40 .41

Malar Space (=distance from lower margin of eye to mandible)

Figure 4. Malar space and eye length in Aphaenogaster umphreyi (u), representing 10 colonies,
and A. fulva (f), representing 12 Florida colonies.

Vol. 1 09, No. 2, March & April, 1 998 93

would still be useful to check reddish brown individuals assigned to fulva in
collections to see whether they have smaller eyes and the other characteristics
of umphreyi. We have provisionally identified as umphreyi a pair of pale speci-
mens with small eyes collected in Decatur County, Georgia, but have not in-
cluded them among the paratypes because they are small specimens with much
less conspicuous rugae than normal in umphreyi', these specimens are in the
Museum of Comparative Zoology, Harvard University.

The reduced hind and middle tibial spurs of umphreyi are quite variable. In
some specimens they are absent, in others they are present, but short. The hind
tibial spurs, when present, are shorter than the middle tibial spurs. The hind
tibial spurs in Aphaenogaster as a whole are remarkably expressive, compared
with most ant genera. In A.floridana Smith they are broad and abruptly acumi-
nate. In A. pallida (Nylander), which also has very small eyes, they are ex-
tremely reduced. In A.flemingi Smith, the hind tibial spurs are somewhat re-
duced, shorter than the middle tibial spurs. In A. tennesseensis (Mayr), they
are much reduced, thick, and sometimes curved. In A. sardoa Mayr, and A.
senilis Mayr, the basal third is noticeably swollen. In A. campana Emery, they
are bristle-like. In A. cockerelli Andre, they have apparently been lost, and
replaced by enlarged lateral bristles. Aphaenogaster fulva and several other
species have evenly tapering spurs. Nobody knows, of course, what ecological
and evolutionary factors are affecting tibial spur morphology, but a study of
the habits of umphreyi and other species that have unusual spurs might help us
understand the function of tibial spurs in ants.

Aphaenogaster umphreyi may be almost entirely subterranean in habits,
and if it does emerge, it may do so at night. All the specimens were collected
underground, either in sand under a thick layer of dead leaves and roots, or
under piles of litter and trash. The senior author has spent hundreds of hours
over the last 14 years prowling scrub habitat at the Archbold Biological Sta-
tion, without seeing a single specimen of umphreyi in the open. The small eyes
and pale color of umphreyi are consistent with a subterranean life.

There are no sexuals associated with workers of umphreyi, but we believe
that we may have collected an unassociated queen. Carroll (1975) states that
queens of fulva are easily distinguished by the "heavily rugose mesothoracic
episternite and sternite." The worker-associated queens we have seen from
Florida, South Carolina, Maryland, and Arkansas seem to agree with this de-
scription, the rugosity consisting of long, gently undulating longitudinal rugae
on a granulate background. We have one dealate queen from Archbold Bio-
logical Station (where fulva is unknown) that has zigzag rugae on the me-
sothoracic episternite and reduced spurs on the hind and middle tibiae. The
eyes and propodeal spines are not reduced. The specimen was collected in a
window trap in Florida scrub habitat in November, 1987.

Nothing is known about umphreyi other than it is a subterranean inhabitant

94 ENTOMOLOGICAL NEWS

of sandy uplands of the southeast, it is difficult to collect, and it appears to be
closely related tofulva. We have no long series showing intraspecific varia-
tion, no associated sexuals, no details of its geographic range, and no informa-
tion on diet or behavior. Until myrmecologists develop an effective method for
finding colonies, this will remain one of our most obscure species of
Aphaenogaster.

ACKNOWLEDGMENTS

We thank Gary Umphrey (University of Western Ontario) and Stefan Cover (Harvard Uni-
versity) for reviewing drafts of this paper. We thank Stefan Cover for arranging the loan of speci-
mens from the Museum of Comparative Zoology, Harvard University, and Walter Suter, Zachary
Prusak, and David Corey for contributing specimens of Aphaenogaste rfulva, and Fabrizio Rigato
for contributing specimens of several species of European Aphaenogaster. We thank our wives,
Nancy and Marie, for waiting patiently whenever we left them by the roadside and went off into
the woods for "a few minutes" of ant collecting.

LITERATURE CITED

Carroll, J. F. 1 975. Biology and ecology of ants of the genus Aphaenogaster in Florida. Unpub-
lished Ph.D. Thesis, Univ. Florida, Gainesville. 177 pp.

Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. Harvard Univ. 104:
585 pp.

Umphrey, G., J. 1996. Morphometric discrimination among sibling species in the fulva-ru dis-
texana complex of the ant genus Aphaenogaster (Hymenoptera: Formicidae). Can. J. Zool.
74: 528-559.

Vol. 109, No. 2, March & April, 1998 95

LABORATORY REARING OF MESOVELIA
CRYPTOPHILA (HETEROPTERA: MESOVELIIDAE) 1

Steven J. Taylor^, J. E. McPherson^

ABSTRACT: Mesovelia cryptophila was reared from egg to adult at 26.70.6 C under a 1 4L: 1 OD
photoperiod. The incubation period averaged 14.93 days; and the four nymphal stadia, 3.24,
2.51, 3.15, and 4.85 days, respectively.

Mesovelia cryptophila Hungerford occurs from New Jersey south to Florida
and west to Michigan, Iowa, Oklahoma, and Mississippi (Smith 1988); it also
has been reported from Minnesota (Bennett and Cook 1981) and Texas
(Polhemus 1997).

Little is known about the biology of this infrequently collected species. It
has been collected from a shaded pool in Mississippi (Wilson 1958); a cypress
swamp in South Carolina (Sanderson 1982); bog, lake, and impoundment habi-
tats in New Jersey (Chapman 1959); a small pond and shaded, stagnant back-
waters of a stream in Iowa (Harris 1943); and the margin of a bog lake in
Michigan, similar to the habitat of Mesovelia amoena Uhler (reported as
Mesovelia douglasensis Hungerford) (Hoffmann 1932, Hungerford 1924).

Hoffmann (1932) reported only apterous adults in Michigan, and Sanderson
(1982) stated that macropterous adults were unknown. Chapman (1959) re-
ported specimens (stages not given) collected in May and August through Oc-
tober in New Jersey; Sanderson (1982) collected specimens (stages not given)
in July in South Carolina; and Hungerford (1924) and Hoffmann (1932) col-
lected adults in June and July, and in July, respectively, in Michigan.

Hoffmann (1932) reared this species in the laboratory under unspecified
conditions with limited success (i.e., five individuals reached adult) and briefly
described the immature stages. He reported only four nymphal instars.

On 9 July 1991, one of us (SJT) discovered a small population of M.
cryptophila in Gallatin County, Illinois. Because so little is known about this
insect, and because Hoffmann's (1932) rearing data were limited and collected
under unspecified conditions, we decided to rear the insect in our laboratory
under controlled conditions.

1 Received June 23, 1997. Accepted August 23, 1997.

2 Center for Biodiversity, Illinois Natural History Survey, 607 E. Peabody Dr., Champaign, IL
61820.

3 Department of Zoology, Southern Illinois University at Carbondale, Carbondale, IL 62901-
6501 USA.

ENT. NEWS 109(2) 95-98, March & April. 1998

96 ENTOMOLOGICAL NEWS

MATERIALS AND METHODS

The Gallatin County site was a small, shaded bay of Crab Lake ( = Hulda
Lake) located 6 mi NE of Shawneetown. The water was covered with a light
(approximately 10 plants per m^) covering of duckweed (Lemanaceae). Thirty
apterous adults (15C?C?,1599) were collected with an aquatic D-net up to 2
m from shore, brought to the laboratory, and placed in glass creamers (1C?,
1 9 /container). The creamers (4.5 cm deep x 3.0 cm diam.) were filled with 1
cm of deionized water and 3 floating plastic disks (0.6 cm in diam.) were
added. The curved sides of the containers were sufficient to prevent the insects
from escaping. Two paper strips of cardstock (approximately 1.25 x 2.5 cm)
were angled against opposite sides of each container with the tops above the
water. The disks and strips served as oviposition sites and allowed individuals
to leave the water.

Containers were checked daily for eggs, all of which were laid beneath the
water surface. Plastic disks and paper strips, with attached eggs, were trans-
ferred to new containers. If eggs were deposited on the walls of the container,
adults were transferred to a new container. As eggs hatched, the newly emerged
first instars were transferred to new containers prepared similarly to those for
adults, but without paper strips. Nymphs of the same instar molting into the
subsequent instar on the same day were transferred to new containers if other
nymphs in the container had not molted. The water level was maintained just
above (0.1 - 0.3 cm) the eggs. Maximum numbers of individuals reared per
container were: 6 first instars, 4 second instars, 3 third instars, and 1 fourth
instar. Adults reared from these eggs were preserved in 70% ethanol.

Each adult was fed 1, and each nymph 1/2, frozen adult fruit fly (Droso-
phila melanogaster Meigen) per day. Flies were crushed or torn slightly for
nymphs to facilitate feeding and were replaced daily.

All individuals were maintained in incubators at 26.7 0.6 C under a
1 4L: 10D photoperiod. All containers were changed at least once per week but
more frequently when water became cloudy.

Data were analyzed with the SAS (SAS Institute 1988) TTEST procedure.
Level of significance was set at 0.05.

RESULTS AND DISCUSSION

Eggs (n = 96) were deposited singly on the paper strips, sides of the plastic
disks, and walls of the containers. The incubation period averaged 14.93 days
(Table 1). There were four nymphal instars, thus confirming Hoffmann's (1932)
results. The first through fourth stadia averaged 3.24, 2.51, 3.15, and 4.85
days, respectively. Duration of total developmental time from egg to adult
averaged 28.56 days. No sexual difference was detected for either the fourth

Vol. 109, No. 2, March & April, 1998

stadium (T = -0.9685, df = 45, p = 0.3304) or for total length of development
(T = -0.8387, df = 45, p = 0.4061). Of the 30 field-collected adults, five fe-
males and four males were still alive after one month.

Hoffmann (1932) collected adults in Michigan but reared their offspring in
Kansas. Females laid up to 75 eggs (mean = 55, n = 4), which were inserted
into plant tissue. He reported an egg laid on 27 July hatched on 1 4 March (229
days), and two eggs laid on 24 and 25 July hatched between 21 and 25 Septem-
ber (58-63 days). The large discrepency in incubation period between
Hoffmann's study (58-229 days) and ours (12-19 days) (Table 1) suggests his
eggs were in diapause. Hoffmann's (1932) five specimens reared from first
instar to adult averaged 1 7.6 days (range = 1 6-21 , no rearing temperature given),
approximately four days longer than ours.

Although the occurrence of only four instars is rare in Gerromorpha (Stys
and Davidova-Vilimova 1989), it is not unprecedented in the Mesoveliidae;
Mesoveliafurcata Mulsant and Rey, a European species, has only four nymphal
instars (Zimmerman 1984).

Table 1 . Durations (in days) of immature stages of laboratory-reared Mesovelia cryptophila.

Number
completing
Stage Sex stadium

Mean Std. Err.

Range

Egg a

14.93 + 0.16

12-19

First instar

3.24 0.07

2-5

Second instar

2.51 0.07

2-4

Third instar

3. 15 0.07

2-4

Fourth instar Males + Females' 3

4.85 0.11

3-7

Males

4.74 0.1 5

3-7

Females

4.95+0.15

3-6

Egg through
fourth instar Males + Females^

28.56 + 0.20

25-31

Males

28.41 0.26

25-31

Females

28.75 + 0.32

26-31

a 96 eggs were laid.

" One individual died during molting and could not be sexed.

98 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

We thank the following individuals for their critical reviews of the manuscript: R. A. Bran-
don, J. A. Beatty, B. M. Burr, Department of Zoology; and D. Ugent, Department of Plant Biol-
ogy, Southern Illinois University at Carbondale.

LITERATURE CITED

Bennett, D. V. and E. F. Cook. 1981. The semiaquatic Hemiptera of Minnesota (Hemiptera:

Heteroptera). Minn. Agric. Exp. Stn. Tech. Bull. 332:1-59.
Chapman, H. C. 1959. Distributional and ecological records for some aquatic and semi-aquatic

Heteroptera of New Jersey. Bull. Brooklyn Entomol. Soc. 54:8-12.
Harris, H. M. 1943. A note on the range of Mesovelia cryptophila Hungerford (Hemiptera:

Mesoveliidae). J. Kansas Entomol. Soc. 16:53-54.
Hoffmann, C. H. 1932. The biology of three North American species of Mesovelia (Hemiptera-

Mesoveliidae). Can. Entomol. 64:88-95, 113-120, 126-134.
Hungerford, H. B. 1924. A second new Mesovelia from the Douglas Lake, Michigan region

(Hemiptera-Mesoveliidae). Ann. Entomol. Soc. Am. 17:453-456.
Polhemus, J. T. 1997. New state and U. S. records and other distributional notes for Heteroptera

(Insecta). Entomol. News 108:305-310.
Sanderson, M. W. 1982. Aquatic and semiaquatic Hemiptera, pp. 6.1-6.94. In: A. R. Brigham,

W. U. Brigham, and A. Gnilka (eds.). Aquatic insects and oligochaetes of North and South

Carolina. Midwest Aquatic Enterprises, Mahomet, Illinois. 837 pp.
SAS Institute. 1988. SAS/STAT user's guide, version 6, 4th ed. Vol. 2. SAS Institute, Gary,

North Carolina. 479 pp.
Smith, C. L. 1988. Family Mesoveliidae Douglas and Scott, 1867, pp. 247-248. In: T. J. Henry

and R. C. Froeschner (eds.). Catalog of the Heteroptera, or true bugs, of Canada and the

continental United States. E. J. Brill, New York. 958 pp.
Stys, P. and J. Davidova-Vilimova. 1989. Unusual numbers of instars in Heteroptera: a review.

Acta Entomol. Bohemoslovaca 86: 1-32.
Wilson, C. A. 1 958. Aquatic and semiaquatic Hemiptera of Mississippi. Tulane Stud. Zool. 6: 1 16-

170.
Zimmermann, M. 1984. Population structure, life cycle and habitat of the pondweed bug

Mesovelia furcata (Hemiptera, Mesoveliidae). Revue Suisse Zool. 91: 1017-1035.

Vol. 109, No. 2, March & April, 1998 99

TWO NEW SPECIES OF MICROCADDISFLIES

(TRICHOPTERA: HYDROPTILIDAE)

FROM KENTUCKY 1

Ronald E. Houp 2 , Katherine H. Houp 3 , Steven C. Harris 4

ABSTRACT: Two new species of microcaddisflies, Hydroptila howelli and Hydroptila kuehnei
(Trichoptera: Hydroptilidae), from Kentucky are described and their affinities noted.

The caddisfly fauna of the southeastern United States has been fairly well
studied, but new species continue to be found. This paper describes two new
species in the genus Hydroptila from a single locality along Salt Lick Creek in
Kentucky, an unaltered Reference Reach stream, draining a portion of the
"Knobs", an area within the Interior Plateau ecoregion. Terminology used in
the descriptions follows that of Marshall (1979). Specimen length was mea-
sured from the tip of the head to the end of the wings and is given as a range.
Type material will be deposited at the National Museum of Natural History,
Smithsonian Institution (NMNH), the Illinois Natural History Survey (INHS),
the Branley A. Branson Museum of Zoology, Eastern Kentucky University
(BABMZ) and the collections of the authors.

Hydroptila howelli NEW SPECIES

(Fig. 1)

Description. Male. Length 1.9 - 2.1 mm. 27 antennal segments. Brown in alcohol. Venter of
abdominal segment VII with short apicomesal process. Segment VIII triangular in lateral view,
acute posteroventrally; in ventral view, deep rounded incision posteromesally, laterally terminat-
ing in several sclerotized teeth; nearly rectangular in dorsal aspect. Segment IX retracted within
segment VIII in ventral view; in dorsal view, anterior portion retracted within VIII and mesally
incised, posterior portion divided into pair of truncate lateral lobes, widely separated mesally.
Tenth tergum broadly triangular, laterally with elongate, sinuate sclerotized processes with acute
apices turned inward; in lateral view these thin processes sharply turned ventrad. Subgenital
plate a rounded shelf in ventral view, bearing pair of short setae posteromesally. Inferior append-
ages in lateral view thin and elongate, clublike at apex; in ventral view widely separated, nearly
parallel along mesal margin, outer margins sinuate, apices acute and strongly turned outward.
Phallus tubular, widening at base and narrowing at midlength, ejaculatory duct protruding apically
about 1/4 phallus length, thin paramere at midlength encircling shaft.

Female and larva. Unknown.

Type material. Holotype, male. Kentucky, LaRue-Marion County line, Salt Lick Creek on Salt
Lick Road, 17 May 1996, at blacklight, R. Houp and K. Houp (NMNH). Paratype, same locality
asholotype, 1 male (BABMZ).

1 Received May 21, 1997, Accepted August 20, 1997.

2 Kentucky Division of Water, 14 Reilly Road, Frankfort, KY 40601 .

3 Department of Biology, Midway College, Midway, KY 40347.

4 Department of Biology, Clarion University, Clarion, PA 16214.

ENT. NEWS 109(2) 99-102, March & April, 1998

100 ENTOMOLOGICAL NEWS

Etymology. Named for the late Dr. Henry H. Howell, in honor of his con-
tributions to aquatic ecology. A teacher, friend and mentor.

Diagnosis. This species is similar to several species in the H. waubesiana group,
including H. delineata Morton, H. tridentata Holzenthal and Kelly, H. englishi
Hamilton, and H. grandiosa Ross. From the latter two species, H. howelli is
separated by the lack of elongate, heavy spines from the sternum of abdominal
segment IX. The new species is separated from other species in the waubesiana
group by the ventral elongation of segment VIII, the elongate, sinuate lateral
straps of the tenth tergite, and the thin, widely separated inferior appendages
which are strongly hooked apically.

Hydroptila kuehnei NEW SPECIES

(Fig. 2)

Description. Male. Length 2.8 - 3.1 mm. 28 antennal segments. Brown in alcohol. Venter of
abdominal segment VII with short apicomesal process. Segment VIII annular; slightly incised along
posterior margin in ventral view; dorsally with pair of crescent-shaped sclerites posteriorly.
Segment IX elongate posterodorsally in ventral view; in dorsal view, emarginate posteriorly, elon-
gate processes laterally, deeply incised anteriorly. Segment X fused with IX, dorsum divided at
base into pair of thin arms, nearly parallel sided, distally with acute points apically and subapi-
cally; in lateral view, arms separated distally with apical points dorsad and ventrad. Subgenital
plate in ventral view thin, lateral margins curved inward, rounded apically bearing pair of mesa!
setae. Inferior appendages elongate and thin in lateral view, sharply curved downward at base,
apically with sclerotized ventral point; in ventral view short and slightly elbowed outward, heavy
peglike setae subapically, elongate seta on lateral margin subapically and basally. Phallus tubular,
widening at base, ejaculatory duct protruding distally, elongate paramere encircling shaft near
midlength.

Type material. Holotype, male. Kentucky, LaRue-Marion County line, Salt Lick Creek on Salt
Creek Road, 7 August 1996, at blacklight, R. Houp and K. Houp (NMNH). Paratypes, same data
as holotype, 5 males (NMNH, INHS, BABMZ, REH, SCH).

Etymology. Named for the late Dr. Robert A. Kuehne, and his contributions to
aquatic ecology. A friend, teacher and mentor.

Diagnosis. This species, another member of the H. waubesiana group, is most
similar to H. patriciae Harris and H. eramosa Harper. With these two species,
H. kuehnei shares the lateral elongation of abdominal segment IX and the
division of the tenth tergite into two lateral arms. These lateral arms are elongate
in the new species, but short in both H. patriciae and H. eramosa and terminate
in long apical extensions in H. patriciae, but short acute extensions in H. kuehnei
and H. eramosa. The inferior appendages are strongly elbowed in H. patriciae
and to the lesser degree in H. kuehnei, but they are nearly straight in H. eramosa.
As well, these appendages in both H. eramosa and the new species have a
strong peglike seta apically which is absent in H. patriciae. The subgenital
plate, in ventral view, in both H. patriciae and H. kuehnei is thin apically, but

Vol. 109, No. 2, March & April, 1998

101

broadly rounded in H. eramosa. This combination of characters serves to
distinguish the new species from other members of the H. waubesiana group.

^S-4-T -' jy^H
\ .'.i I )- xCs/M
,\ i^ /\' : /'$ I '

^K^K ' - ' /*^

Figure 1. Hydroptila howelli, n. sp. male genitalia. A. Lateral view; B. Dorsal view; C. Ventral
view; D. Phallus, ventral view.

102

ENTOMOLOGICAL NEWS

Figure 2. Hydroptila kuehnei, n. sp. male genitalia. A. Lateral view; B. Dorsal view; C. Ventral
view; D. Phallus, ventral view.

ACKNOWLEDGMENTS

The authors appreciate the interest and assistance of G. A. Schuster, Eastern Kentucky Uni-
versity.

LITERATURE CITED

Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bull. Brit. Mus.
(Nat. Hist.), Entomol. Ser. 39: 135-239.

Vol. 109, No. 2, March & April, 1998 103

SEASONAL FLIGHT PERIODICITIES OF SIX

MICROCADDISFLIES (TRICHOPTERA:

HYDROPTILIDAE, GLOSSOSOMATIDAE) IN

THE BRAZOS RIVER, TEXAS, WITH NOTES ON

LARVAL BIOLOGY AND SITE RECORDS 1

David C. Houghton 2 ' 3 , Kenneth W. Stewart 2

ABSTRACT: The seasonal flight periodicities of six small caddisflies were studied from April,
1995, to November, 1996, at a large riffle of the Brazos River in north-central Texas. Light trap
samples for adults suggested a synchronized univoltine cycle for Protoptila alexanderi, asyn-
chronous univoltine cycles for Hydroptila icona and Orthotrichia cristata, bivoltine cycles for
Ithytrichia clavata and Ochrotrichia tarsalis, and a multivoltine cycle for Hydroptila angusta;
the last species accounted for 82% of the total hydroptilid adult abundance. Larval collections of
P. alexanderi, H. angusta and /. clavata confirmed their presence in the benthic community.
Seven site records and one drainage record of Neotrichia vibrans were documented, increasing
the total number of caddisfly species found at this site by Moulton et al. (1993) to 30.

The caddisfly family Hydroptilidae, often referred to as the "micro-
caddisflies", contains the smallest members of the Trichoptera, with adults of
many of the 220 Nearctic species reaching only 2-3 mm in length (Wiggins
1996). Hydroptilidae is the most species rich trichopteran family, and often
ten or more congeners can occur together at a single site (Moulton and Stewart
1996). Despite this, very little is known concerning hydroptilid life histories
and behavior. Nielson (1948) studied the biology of five Danish species; eluci-
dating univoltine cycles for Orthotrichia tetensii Kolbe and Ithytrichia
lamellaris Eaton, and bivoltine cycles for Agraylea multipunctata Curtis,
Hydroptila femoralis Eaton and Oxeythira costalis Curtis. This work remains
the source for much of the knowledge of larval morphology, feeding, case-
building behavior and general biology of the family (Marshall 1979, Wiggins
1996).

Other ecological studies of hydroptilids have dealt with the drift patterns
of larvae, including those of Hydroptila rono Ross and H. icona Mosely, re-
spectively, in an Oregon and Texas stream (Anderson 1967, Cloud and Stewart
1974a) from which an interpretation of univoltine life cycles was made by
Wiggins (1996). Resh and Houp (1986) studied the biology of Dibusa angata
Ross and found it to be bivoltine in a Kentucky stream and dependent on the
red alga Lemanea australis Atkinson for food and case-building material.
Marshall (1979) reviewed the world genera of Hydroptilidae.

1 Received May 15, 1997. Accepted July 25, 1997.

2 Department of Biological Sciences, University of North Texas, P.O. Box 5218, Denton, TX
76203-0218.

3 Current address: Department of Entomology, University of Minnesota, St. Paul, MN 55108.

ENT. NEWS 109(2) 103-109, March & April, 1998

104 ENTOMOLOGICAL NEWS

Moulton et al. (1993) documented 23 species of Trichoptera from a riffle
of the Brazos River in Palo Pinto County, Texas as part of their investigation of
the Brazos River caddisfly fauna. This documentation included 10 hydroptilid
species and one member of the glossosomatid caddisfly subfamily Protoptilinae,
Culoptila cantha (Ross). Protoptiline glossosomatids are often field-sorted in
with hydroptilids due to their small size; no life history information has been
reported for any of the 18 North American species (Wiggins 1996). The pur-
pose of our study was to document the seasonal flight periodicities of micro-
caddisflies at this Brazos River site as suggestive of their respective voltinisms.

MATERIALS AND METHODS

Study site. The Brazos River arises on the Caprock Escarpment in eastern
New Mexico and flows southeasterly through Texas to the Gulf of Mexico.
Our study was conducted at a 200 m riffle located approximately 35 km down-
stream of the Morris Shepard Dam in Palo Pinto County; research on the biol-
ogy of many aquatic insects has been ongoing at this site for 25 years (Stewart
et al. 1973; Cloud and Stewart 1974a,b; Vaught and Stewart 1974; McClure
and Stewart 1976; Rhame and Stewart 1976; Houghton 1997). Recent descrip-
tions of this site can be found in Moulton et al. (1993) and Houghton (1997).

Adults. Adults were collected from April, 1995, to November, 1996, using
two 8-watt portable ultra-violet lights placed over two 22.5 X 35 cm white
porcelain pans filled with 70% ETOH. These samples were generally taken
monthly from October to March and twice monthly from April to September.
The pans were set about 1.0 m from the water's edge; one at the head of the
riffle, and the other 16.5 m downstream. Preliminary observations had indi-
cated predominately nocturnal flight periods for Brazos microcaddisflies; there-
fore lights were run for 20 minute intervals beginning at dusk, separated by 40
minutes between samples, until three sets of samples were taken or until a
sample yielded no adults of any species. A small number ofHydroptila angusta
Ross adults were aspirated from riparian rocks and vegetation prior to dusk
during February and March, 1996. Each sample taken was sub-sampled by the
following procedure: the specimens were placed in a 9.5 cm diameter round
petrie dish and mixed thoroughly; a 3 cm diameter ring was randomly placed
into the sample and all individuals within the ring area were counted; the num-
ber of males and females of each species found was extrapolated to estimate
the total number present in each sample; the samples were combined to esti-
mate the number present on each sampling date.

Larvae and Pupae. Larvae and pupae were collected off rocks with soft-
touch forceps on the same sampling dates as adults and either preserved in
70% ETOH or transported alive back to the laboratory in Styrofoam "six-pack"
coolers (Szczytko and Stewart 1979). Larvae and pupae were reared in wire

Vol. 109, No. 2, March & April, 1998 105

mesh baskets within a Frigid Units Living Stream with Brazos River simu-
lated flow, temperature, photoperiod, and natural algal food to estimate pupa-
tion period and associate the life stages using the metamorphotype method
(Milne 1938). Behavior of fifth instar larvae was observed by placing indi-
viduals in a 5 cm diameter petrie dish under a Wild M2E dissecting micro-
scope equipped with a Dyonics fiberoptic light source. Voucher specimens of
all species studied are deposited in the University of North Texas Entomologi-
cal Collection.

RESULTS AND DISCUSSION

Seasonal flight periodicities were determined for the hydroptilid species
Hydroptila icona, H. angusta, Ithytrichia clavata Eaton, Ochrotrichia tarsalis
(Hagen) and Orthotrichia cristata Morton, and for Protoptila alexanderi Ross
(Glossosomatidae: Protoptilinae) (Fig. 1); the life history of the protoptiline
glossosomatid Culoptila cantha, which was very common at this site, was stud-
ied by Houghton (1997) and will be published in a separate paper. Protoptila
alexanderi and Ochrotrichia tarsalis represented site records not reported by
Moulton et al. (1993), as did the following species which did not occur in
enough abundance to discern a flight periodicity pattern: Hydroptila waubesiana
Betten (n = 63), Neotrichia minutisimella (Chambers) (n = 1 ), N. vibrans Ross
(n = 2), Oxeythira aculea Ross (n = 3), and O. azteca (Mosely) (n = 2).
Neotrichia vibrans also represented a new drainage record.

Protoptila alexanderi Ross. Adults of this species were caught from early
July to late September, 1995, and early August to October, 1996 (Fig. 1 ). Peak
abundances occurred in late August, 1995, and early September, 1996, sug-
gesting a synchronized univoltine cycle. Males were slightly more abundant
than females; both sexes exhbited similar seasonal periodicity. We collected a
series of twelve fifth instar larvae in early July, 1995, on clean 10-20 cm diam-
eter stones in the upstream half of the riffle. Moulton et al. (1993) collected
adults from the Paluxy River and Ham Creek, Brazos River tributaries, in June
and October. Previously, the species had been reported in Texas only from the
San Antonio and San Marcos Rivers (Edwards 1973).

Hydroptila angusta Ross. Adults of this species were collected in all months
of the year (Fig. 1 ) and accounted for 82% of the total hydroptilid sample. As
many as four generations may have occurred during both years as suggested
by the peaks in abundance, although an exact determination is not possible
without extensive larval data. Males and females exhibited similar seasonal
periodicity. Larvae were reared to adults during April, July, and September,
1 995, and March, April, and June, 1 996. Moulton et al. ( 1 993) collected adults
throughout their Brazos Drainage study area during the months of March- April,

106 ENTOMOLOGICAL NEWS

May -June, and October. Cloud and Stewart ( 1 974a) did not report this species
in the drift of the Brazos River. We found substantial numbers of fifth instar
larvae in the filamentous algal growth (mostly Cladophora sp.) that was com-
mon on most of the rocks in the riffle during summer months. When placed on
small stones in a petrie dish, larvae were sedentary and held their cases at
acute angles to the faces of the stones.

Hydroptila icona Mosely. Adults of this species were collected from early
May to November during both years (Fig. 1). Peak abundances of both sexes
occurred in late June, 1995, and early July, 1996, suggesting a univoltine cycle;
this would corroborate the one-year cycle proposed by Cloud and Stewart
(1974a) and Wiggins (1996). However, voltinism is confounded by a second
peak in male abundance that occurred in late August, 1995, and early August,
1996. Moulton et al. (1993) collected adults throughout their study area during
the months of June and October-November. Cloud and Stewart (1974a) re-
ported H. icona as one of the two common hydroptilids at this site and noted a
nocturnal drift of both cased and caseless larvae. We did not find larvae of this
species in our benthic samples.

Ithytrichia clavata Eaton. Adults of this species were collected from early
April to November, 1995, and early April to October, 1996 (Fig. 1). Adult
abundance peaked during early May and early September, 1995, and late April
and late September, 1996, suggesting a synchronized bivoltine cycle. Males
were slightly more abundant during the early generation and females were
slightly more abundant in the later generation during both years. A series of 1 2
pupae collected in early April, 1995, emerged in the laboratory in early May,
1995; a series of 6 pupae collected in early April, 1996, emerged in late April,
1996. We also collected approximately 50 empty cases in early April, 1996,
and only one cased larva in early May, 1995. These pupae and empty cases
were found on rocks in quiet areas of the riffle. Moulton et al. (1993) collected
adults throughout most of their study area, including this site, during the months
of March- April, June and October. Despite the low numbers of larvae and pu-
pae that we found, Cloud and Stewart (1974a) reported that /. clavata was a
common hydroptilid at this site and that it drifted nocturnally. The one fifth
instar larva was found on a rock near the head of the riffle. When placed on a
small stone in a petrie dish under microscopic light and videotaped, this indi-
vidual exhibited strong negative phototaxicity and actively sought the under-
side of the stone. A torsion angle of 90 between the first two abdominal seg-
ments, causing the flat sides of the case to be held parallel to the ground, was
evident. This phenomenon was also noted for/, lamellaris (Nielson 1948).

Ochrotrichia tarsalis (Hagen). Adults of this species were collected from
early June to November, 1995, and early April to October, 1996. Adult abun-

Vol. 109, No. 2, March & April, 1998

107

200

100

10,000

,C 5,000

U o

2,000

^2 1 ,000

J> 800

3 400

~CS 400

."&? 200

600

300

-*- Males
Females

Protoptila alexanderi

Hydroptila angusta

Hydroptila icona

Ithytrichia clavata

Ochrotrichia tarsalis

Orthotrichia cristata

AMJ J ASONDJ FMAMJ J ASON

1995

1996

Month

Figure 1 . Estimated number of adults caught from a riffle on the Brazos River, Texas from April,
1995, to November, 1996, based on subsamples.

108 ENTOMOLOGICAL NEWS

dance peaked in late July and late September, 1995, and early July and Octo-
ber, 1996 (Fig. 1), suggesting a synchronized bivoltine cycle. Males and fe-
males had similar seasonal periodicity. Moulton et al. (1993) did not collect
this species at our site but found adults in much of their Brazos Drainage study
area during June and October. Cloud and Stewart (1974a) reported Ochrotrichia
larvae in low abundance in their drift samples but did not identify a species.
We did not find larvae of this species in our study riffle.

Orthotrichia cristata Morton. Adults of this species were collected from
late June to November, 1995, and early June to November, 1996 (Fig. 1). Both
sexes peaked in abundance in late September during both years, suggesting a
univoltine cycle. However, females were not collected until early August both
years, while males were present for 3-4 months before their peak abundance.
Moulton et al. (1993) collected adults at our site during June and July. Cloud
and Stewart (1974a) report Orthotrichia larvae in low abundance in their drift
samples but did not identify a species. We did not find larvae of this species in
our sampling area.

ACKNOWLEDGMENTS

The authors thank S.R. Moulton, II for confirming the identity of several species and the
following people for assisting us in the field and laboratory: J.C. Abbott, I.W. Agado, G.L. Akin,
R.G. Jones, C.J. Larson, J.C. Norwood, S.A. Peterson, J.P. Snow and N.L. Witt. We also thank
K.D. Alexander, S.W. Szczytko and two anonymous reviewers for commenting on earlier ver-
sions of our manuscript.

LITERATURE CITED

Anderson, N. H. 1967. Biology and downstream drift of some Oregon Trichoptera. Can. Entomol.

99: 507-521.
Cloud, T. J. and K. W. Stewart. 1974a. Seasonal fluctuations and periodicity in the drift of

caddisfly larvae (Trichoptera) in the Brazos River, Texas. Ann. Entomol. Soc. Am. 67: 805-

810.
Cloud, T. J. and K. W. Stewart. 1974b. The drift of mayflies (Ephmeroptera) in the Brazos

River, Texas. J. Kans. Entomol. Soc. 47: 379-396.
Edwards, S. W. 1973 Texas caddisflies. Texas J. Sci. 23: 491-516.
Houghton, D. C. 1 997. Descriptions, life history and case-building behavior of Culoptila cantha

(Ross) (Trichoptera: Glossosomatidae) in the Brazos River, Texas. M.S. thesis. Univ. North

Texas, Denton, TX. 62 pp.
Marshall, J. E. 1979. A re view of the genera of Hydroptilidae (Trichoptera). Bull. British. Mus.

Natur. Hist., Entomol. Sen 39: 135-239.
McClure, R. G. and K. W. Stewart. 1976. Life cycle and production of Chorterpes (Neo-

chorterpes) mexicanus Allen (Ephemeroptera: Leptophlebiidae). Ann. Entomol. Soc. Am.

69: 134-148.
Milne, M. J. 1938. The "metamorphotype" method in Trichoptera. J. N.Y. Entomol. Soc. 46:

435-437.
Moulton II, S. R. and K. W. Stewart. 1996. Caddisflies (Insecta: Trichoptera) of the Interior

Highlands of North America. Mem. Am. Entomol. Inst. 58: 1-313.

Vol. 109, No. 2, March & April, 1998 109

Moulton II, S. R., Petr, D., and K. W. Stewart. 1993. Caddisflies (Insecta: Trichoptera) of the

Brazos river drainage in north-central Texas. Southw. Natur. 38: 19-23.
Nielson, A. 1 948. Postembryonic development and biology of the Hydroptilidae. Kongelige Danske

Videnskabernes Selskab, Biolgiske Skrifter 5( 1 ). 203 pp.

Resh, V. H. and R. E. Houp. 1986. Life history of the caddisfly Dibusa angata and its associa-
tion with the red alga Lemanea australis. J. N. Am. Benthol. Soc. 5: 28-40.
Rhame, R. E. and K. W. Stewart. 1976. Life cycles of three Hydropsychidae (Trichoptera)

species in the Brazos River, Texas. Trans. Am. Entomol. Soc. 102: 65-99.
Szczytko, S. W. and K. W. Stewart. 1979. Stonefly drumming as a model classroom study of

aquatic insect behavior. In: Resh, V. H. and D. M. Rosenberg (eds.) Innovative teaching in

aquatic entomology. Can Spec. Publ. Fish. Aquat. Sci. 43: 31-37.
Stewart, K. W., G. P. Friday and R. E. Rhame. 1973. Food habits of hellgrammite larvae,

Corydalus cornutus (Megaloptera: Corydalidae), in the Brazos River, Texas. Ann. Entomol

Soc. Am. 66: 959-963.
Vaught, G. E. and K. W. Stewart. 1974. The life history and ecology of Neoperla clymene

(Newman) (Plecoptera: Perlidae). Ann. Entomol. Soc. Am. 67: 167-178.
Wiggins, G. B. 1996. Larvae of the North American caddisfly genera (Trichoptera). Univ. Toronto

Press. Toronto, Canada. 457 pp.

1 10 ENTOMOLOGICAL NEWS

A NEW SPECIES OF PSEUDOCENTROPT1LOIDES

(EPHEMEROPTERA: BAETIDAE), WITH REVISIONS

TO OTHER PREVIOUSLY UNNAMED BAETID

SPECIES FROM TEXAS 1 ' 2

N. A. Wiersema 3 , W. P. McCafferty 4

ABSTRACT: Pseudocentropriloides morihari, new species, is described from larvae and male
and female adults from Texas, USA. The species represents the second of the genus to be discov-
ered in North America, where it is distinguished from P. usa primarily by size, labral and maxil-
lary morphology, and ventral abdominal pattern in the larvae. Pseudocentroptiloides morihari
was previously reported as Centroptilum sp. 2. Some other unnamed but numbered species of
Baetidae reported from Texas are resolved to known nominal species ofAcerpenna, Baetis, and
Centroptilum, and the status of others is reviewed. Difficulty in separating adult males of
Pseudocentroptiloides and some Procloeon is noted.

The small minnow mayfly genus Pseudocentroptiloides Jacob (Ephe-
meroptera: Baetidae) was revised by Waltz and McCafferty (1989) and has
included P. ceylonica Glazaczow (Oriental), P. christineae Waltz and McCaf-
ferty (Oriental), P. shadini (Kazlauskas) (Palearctic; type species), and P. usa
Waltz and McCafferty (Nearctic). Larvae are characterized by a V-shaped emar-
gination on the labrum, broad maxillae, labial palp segments 3 that are greatly
expanded and truncated, and glossae that are relatively short and truncate.

In 1977, one of us (WPM) collected very small and then unidentifiable
larvae in Texas that were simply referred to as Centroptilum sp. 2, and subse-
quently reported as such by McCafferty and Davis (1992) and Lugo-Ortiz and
McCafferty (1995).Recently,the other of us (NAW) collected additional larvae
of this species in Texas and was able to rear adults. Close examination of the
material in light of recent revisions of North American Baetidae (see e.g.,
McCafferty and Waltz 1990, Edmunds and Waltz 1996) shows that it is clearly
Pseudocentroptiloides and that it represents a previously undescribed species.
Prior to this discovery, P. usa, from the Midwest, was the only species of Pseudo-
centroptiloides known from the Western Hemisphere, and it and P. shadini were
the only species known as adults. McCafferty and Waltz (1990) had predicted
that additional species of Pseudocentroptiloides eventually would be found in
North America. We are most honored to name this new species after Dennis
Morihara (Pukalani, Hawaii), who helped collect the original material, and who
contributed significant systematic work that catalyzed our modern understand-
ing of the diverse and challenging family Baetidae in North America.

1 Received August 18, 1997. Accepted September 10, 1997.

2 Purdue Agricultural Experiment Station Journal No. 15478.

3 4857 Briarbend, Houston, TX 77035.

4 Dept. Entomology, Purdue University, West Lafayette, IN 47907.

ENT. NEWS 109(2) 110-116, March & April, 1998

Vol. 109, No. 2, March & April, 1998

Because of the state of knowledge at the time, a number of baetids col-
lected from Texas could not be assuredly assigned to nominal species and thus
were reported as numbered species by McCafferty and Davis ( 1 992) and Lugo-
Ortiz and McCafferty (1995). New analyses of much of that material, often in
light of subsequent collections, have allowed us to update the identification
status of those species. We provide this new information under the heading
Additional Species Revisions following the new species description below.

Pseudocentroptiloides morihari, NEW SPECIES

(Figs. 1-8)

Centroptilum sp. 2 McCafferty and Davis, 1992:207.

Larva. Lengths: body 4.0-5.5 mm; caudal filaments 1.6-2.0 mm. Head: Coloration pale,
darkest around coronal and frontal sutures. Antennae broken but extending at least as far as fore
coxae; scapes, pedicels, and flagella with sparse, fine, simple setae. Labrum (Fig. 1) sclerotized
posterolaterally, slightly wider than long, and with broad triangular emargination apically; dorsal
subapical setae present across entire width of labrum, relatively short and not extending beyond,
or much beyond, distal margin of labrum. Mandibles (Figs. 2 and 3) with relatively broad bases.
Maxillae (Fig. 4) with crest of fine setae extending basally well below galealacinia crown, setae
longer or subequal to galealacinia denticles; digitate setae of galealacinia comb poorly devel-
oped (1-2 setae); palpi three segmented. Labium as in Fig. 5, with glossae nearly as wide as long
and two-thirds to three-fourths length of paraglossae. Thorax: Nota cream, without distinct pat-
terns. Hindwingpads present. Legs similar to Figs. 6 and 7 of Waltz and McCafferty (1989);
claws subequal in length to respective tarsi. Abdomen: Coloration pale; terga light brown, dark-
est at anterior margins of each tergum. Sterna unicolorous cream, except for wide transverse
brown band anteriorly on sternum 8 [mature individuals also with purplish transverse markings
of adults (see below) on sterna 7 and 8]. Paraprocts with five to six marginal spines. Median
caudal filament subequal in length to cerci.

Male adult. Lengths: body 4.6-5.8 mm; forewings 3.8-5.0 mm; hindwings 0.8-1.0 mm;
cerci 8.0-10.0 mm. Head: Turbinate portion of compound eyes yellow, slightly divergent anteri-
orly, oval, ca. 1.6 times longer than wide in dorsal view, and on low stalks. Thorax: Coloration
generally cream to light brown, sterna slightly paler than nota. Forewings (Fig. 6) with marginal
intercalaries relatively long, especially posterior to MA2 (longer than respective distal crossveins
connecting longitudinal veins between MA2 and ICuA)). Hindwings (Fig. 7) slightly broad-
ened, with short marginal veins following 2nd main longitudinal vein. Abdomen: Segments 2-6
translucent white; segments 7-10 slightly darker; small longitudinal dashes laterally on segments
1-9; prominent black, complex, internal maculation visible in dorsal view on segments 7 and 8.
Sterna 7 and 8 often with transverse, pale reddish purple line anteriorly. Genitalia as in Fig. 8.

Female adult. Lengths: body 5.6-5.8 mm; forewings 5.0 mm; hindwings 1 .0 mm; cerci 6.5-
7.0 mm. Body more robust than male, generally pale throughout except for prominent black
spiracular markings dorsolaterally on abdomen, black line in cervical area, and faint dark mark-
ings laterally on thorax.

Material examined. HOLOTYPE: male larva, TEXAS, Austin Co., San Bernard River at I-
10, west of Sealy, V-9-1977, W. P. McCafferty, A. V. Provonsha, D. Morihara, in the Purdue
Entomological Research Collection. PARATYPES: one male and one female larva, same data
and deposition as holotype. Additional material: One female larva and one reared male adult
(some parts mounted on slides), TEXAS, Austin Co., San Bernard River at I- 10 west of Sealy, V-
1 9- 1 997, N. A. Wiersema, same deposition as holotype. Three male and two female adults, TEXAS,
Williamson Co., Georgetown, San Gabriel River at San Gabriel Park, X-7- 1 996, N. A. Wiersema,

1 1 2 ENTOMOLOGICAL NEWS

same deposition of holotype, except one male in private collection of NAW. One male adult,
TEXAS, Comal Co., Guadalupe River about 1 1 miles below Canyon Dam, XII-7-1996, N. A.
Wiersema, in private collection of NAW.

Diagnosis. Larvae of P. usa are much larger and more robust than those of
P. morihari, with mature individuals nearly twice as large. Structurally, P.
morihari can be further easily distinguished from P. usa by the shape of the
distal emargination of the labrum (Fig. 1 ), dorsal labral subapical setae that
extend in a row along the entire width of the labrum (Fig. 1), the much more
extensive crest of setae on the galealaciniae (Fig. 4), and relatively broader
glossae and labial palps segment 3 (Fig. 5) [see comparative figures off. usa
in Waltz and McCafferty (1989): Figs. 1,4, and 5)]. In addition, whereas the
paraprocts of P. morihari have five or six marginal spines, re-examination of
the type material of P. usa indicated that they have six or seven. This latter
difference is of inconsistent value for diagnosis; however, significant color
pattern differences were also revealed from the type material of P. usa. Whereas
P. morihari larvae have a somewhat broad band of pigment anteriorly on ab-
dominal sternum 8, in P. usa such a band is prominent on both sterna 8 and 9
and weakly present on sternum 7. In addition, sublateral paired spots that are
present on all the abdominal sterna of P. usa, but particularly well developed
on sterna 7-9, are entirely absent in P. morihari.

In the larval key to world species of Pseudocentroptiloides provided by
Waltz and McCafferty (1989), P. morihari can most easily be incorporated
into it simply by adding another couplet at the end. First, however, the couplet
part 3' should be modified to read: "Maxillae [Figs. 3 1 (Keffermuller and Sowa
1984) and 4 herein] with crest of setae extending basally below crown of
galealacinia, subequal to or longer than denticles of galealacinia . . . 4." Cou-
plet 4 should then be added as follows:

4. Labrum (Fig. 1 herein) with relatively short dorsal subapical setae not extending beyond, or

much beyond, distal margin of labrum P. morihari.

4'. Labrum [Fig. la of Jacob and Glazaczow (1986)] with long dorsal subapical setae extending
well beyond distal margin of labrum P. shadini.

The adults of P. morihari and P. usa are remarkably similar. Structurally,
the only apparently significant differences found involve the wings. In P.
morihari, the marginal intercalaries of the forewing are relatively long, espe-
cially in the posterior half of the wing. Whereas the intercalaries are longer
than the respective crossveins connecting longitudinal veins between MA2 and
ICuA[ in P. morihari (Fig. 6), they are shorter by the same comparison in P.
usa [Fig. 8 of Waltz and McCafferty (1989)]. Also, the hindwings of P. morihari
(Fig. 7) are slightly broader and have an additional short vein posteriorly. The
hindwings of P. usa are very narrow and have only two long longitudinal veins
[Fig. 9 of Waltz and McCafferty (1989)]. The abdominal segments 1-6 of the
male adults are light in color in both species, as is common in many baetid

Vol. 109, No. 2, March & April, 1998

Figs. 1-8. Pseudocentroptiloides morihari, n. sp., 1-5. Larva. 1. Labrum (right: dorsal, left: ven-
tral). 2. Left mandible. 3. Right mandible. 4. Left maxilla. 5. Labium (half, dorsal view). 6-8.
Male adult. 6. Forewing. 7. Hindwing (enlarged relative to forewing). 8. Genitalia (ventral).

1 1 4 ENTOMOLOGICAL NEWS

males, but the thorax and abdominal segments 7-10 are slightly more darkly
contrasting in P. usa than in P . morihari. One other possible difference be-
tween the males of these species is that in all the material known thus far, the
anterior line on abdominal sterna 7 and 8 is broken in the middle in P. usa, thus
appearing as a pair of lines rather than a single line as in P. morihari. Unfortu-
nately, we cannot be sure if any of the adult differences we have reviewed
above will be consistent between the species when larger populations are known.
All such characteristics have been known to be variable in other adults of cer-
tain baetids.

Although the female adult of P. usa was not described by Waltz and
McCafferty (1989), our examination of the female adult paratype of P. usa
indicated that the females of P. morihari and P. usa are very similar, except for
the venational differences noted above. In addition, based on the few compara-
tive specimens available, the branching spiracular maculations appear more
developed in P. usa, and there is a long, horizontal, unbroken, thin, dark line
connecting abdominal segments 8 and 9 laterally in P. usa that is not present as
such in P. morihari.

Remarks. The San Bernard River where larvae of P. morihari have been
collected has a shifting sand substrate. Larvae were kicked into a screen or
dredged with a D-net from sand and vegetation in relatively shallow water.
Adults have been collected at lights at night (as subimagos) from the Guadalupe
and San Gabriel Rivers. All of the localities are in the Hill Country of central
Texas. Lugo-Ortiz and McCafferty (1995) showed that this area had the high-
est mayfly diversity in Texas. They also indicated that this area was strongly
influenced by eastern North American faunal elements.

Caution should be taken when sorting Texas adults with single marginal
intercalaries in the forewings, including P. morihari. We have recently reared
Procloeon viridoculare (Berner) from Texas that as adults have genitalia with
a medial protrusion on the forceps that will make them key out to Pseudo-
centroptiloides when using the generic key by Edmunds and Waltz (1996).
Because there is no doubt that they are P. viridoculare, and because we have
seen the slight development of such a protrusion on other Procloeon in the
South and Midwest, there is a potential problem in separating Procloeon from
Pseudocentroptiloides when only adults are in hand.

ADDITIONAL SPECIES REVISIONS

Acentrella sp. 1 Lugo-Ortiz and McCafferty [= Baetis virilis (McDun-
nough)]. The original report of Acentrella sp. 1 by Lugo-Ortiz and McCafferty
(1995) was based on the male larval stage of Baetis virilis. This species lacks
hindwings, and because of leg setation was provisionally placed in Acentrella
at the time. This also represents the first report of B. virilis from Texas.

Acentrella sp. 2 Lugo-Ortiz and McCafferty [= Baetis virilis (McDun-

Vol. 109, No. 2, March & April, 1998 115

nough)]. The fact that the female larvae originally referred to a&Acentrella sp.
2 by Lugo-Ortiz and McCafferty (1995) also is referable to B. virilis reflects
the fact that the larvae of B. virilis are sexually dimorphic (Ide 1937). Differ-
ences in the male and female adults of B. virilis were first noted by McDunnough
(1924).

Acerpenna sp. 1 McCafferty and Davis [= Acerpenna pygmaea (Hagen)].
McCafferty and Davis (1992) stated that this species was similar to A. harti
(McDunnough). We now interpret A. sp. 1 as an adult variant of the relatively
widespread species A. pygmaea.

Baetis sp. 1 McCafferty and Davis [= Baetis punctiventris (McDunnough)].
These larvae represent a pale color variant of B. punctiventris that lacks the
ventral spotting usually typical of the species. It is the most common two-
tailed Baetis in the Hill country of Texas, where we have additionally seen it
from Medina River, Bandera County; Guadalupe River, Comal County; Blanco
River, Hays County; Block Creek, Kendall County; and Gorman Creek, San
Saba County.

Camelobaetidius sp. 1 McCafferty and Davis. This species was originally
reported from larvae from the Rio Grande River along the Texas/Mexico bor-
der by McCafferty and Davis (1992), and more recently larvae were reported
from Colorado by McCafferty et al. (1993). Formal description of a new spe-
cies has been postponed until more mature specimens are available.

Centroptilum sp. 1 McCafferty and Davis [= Centroptilum triangulifer
(McDunnough)]. McCafferty and Davis (1992) initially noted the similarity of
C. sp. 1 to C. triangulifer. Observations of variation in this species since that
time indicate that the Texas material falls within the range of variation, and C.
triangulifer should be added to the list of Texas mayflies.

Centroptilum sp. 3 Lugo-Ortiz and McCafferty. The single larva that the
report of Lugo-Ortiz and McCafferty (1995) was based on was sent to another
co-worker for an opinion as to its identification and has since been lost. Thus,
we presume that it will never be resolved. Procloeon viridoculare (Berner),
which was recently reported from Texas by Wiersema (1998), has been taken
from the same locality as C. sp. 3, on the South Llano River in Kimble County,
but we cannot be sure of any association.

Procloeon sp. 1 McCafferty and Davis. Due primarily to the fact that this
report was based on female adults, the material still cannot be placed to any
nominal species.

Procloeon sp. 2 McCafferty and Davis [= Centroptilum album McDun-
nough]. Centroptilum album was first reported by Moore (1950) from Texas
(San Marcos River); however, that report generally has not been referenced in
subsequent inventories of mayflies in Texas. We still cannot verify the accu-
racy of Moore's report, but must assume it was correct, based on the now
confirmed presence of C. album in Texas. Procloeon sp. 2 was based on larvae
that fall within the expected range variation of C. album, as had been sug-

1 1 6 ENTOMOLOGICAL NEWS

gested by McCafferty and Davis (1992). This species has been placed both in
Centroptilum and Procloeon recently; however, it is more consistent with the
concept of Centroptilum reflected in the key to North American mayfly genera
by Edmunds and Waltz ( 1 996).

Procloeon sp. 3 McCafferty and Davis. This new but unnamed species
was based on highly distinctive larvae from the J. R. Davis private collection
taken from Pinto Creek, Kinney County Texas (see McCafferty and Davis 1992).
While we were in the process of writing up the formal new description and
new name for the McCafferty and Davis species for publication, we learned
that other workers had also decided to describe this species independently.
Rather than cause an unnecessary synonym in the literature, we are deferring
to those others, although to our knowledge a formal description is not in press
at this time, nor can we be sure when or if it will be published in the near
future.

ACKNOWLEDGMENTS

We thank C. R. Lugo-Ortiz and R. P. Randolph (West Lafayette, Indiana) for discussing and
reviewing this paper.

LITERATURE CITED

Edmunds, G. F M Jr. and R. D. Waltz. 1996. Ephemeroptera. Pp. 126-163 In: R. W. Merritt and
K. W. Cummins (eds.), An introduction to the aquatic insects of North America, 3rd edition.
Kendall/Hunt, Dubuque.

Ide, F. P. 1937. Descriptions of eastern North American species of baetine mayflies with particu-
lar reference to the nymphal stage. Can. Entomol. 69: 219-213, 235-243.

Jacob, U. and A. Glazaczow. 1986. Pseudocentroptiloides, new baetid genus of Palearctic and
Oriental distribution (Ephemeroptera). Aq. Insects 8: 197-206.

Keffermuller, M. and R. Sowa. 1984. Survey of central European species of the genera
Centroptilum Eaton and Pseudocentroptilum Bogoescu (Ephemeroptera: Baetidae). Polsk.
Pismo Entomol. 54: 309-340.

Lugo-Ortiz, C. R. and W. P. McCafferty. 1995. The mayflies (Ephemeroptera) of Texas and
their biogeographic affinities. Pp. 151-169 In: L. Corkum and J. Ciborowski (eds.), Current
directions in research on Ephemeroptera. Can. Schol. Press, Toronto.

McCafferty, W. P. and J. R. Davis. 1992. New and additional records of small minnow mayflies
(Ephemeroptera: Baetidae) from Texas. Entomol. News 103: 199-209.

McCafferty, W. P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera)
of North America. Trans. Am. Entomol. Soc. 1 16: 769-799.

McCafferty, W. P., R. S. Durfee, and B. C. Kondratieff. 1993. Colorado mayflies (Ephe-
meroptera): an annotated inventory. Southwest. Nat. 38: 252-274.

McDunnough, J. 1924. New Canadian Ephemeridae with notes, II. Can. Entomol. 56: 90-98,
113-122, 128-133.

Moore, L. F., Jr. 1950. Distribution of mayfly nymphs in streams of Dallas County, Texas. Field
Lab. 18: 103-112.

Waltz, R. D. and W. P. McCafferty. 1989. New species, redescriptions, and cladistics of the
genus Pseudocentroptiloides (Ephemeroptera: Baetidae). J. New York Entomol. Soc. 97: 151-
158.

Wiersema, N. A. 1998. Newly reported and little known mayflies (Ephemeroptera) from Texas.
Entomol. News, 109: 27-32.

Vol. 109, No. 2, March & April, 1998 117

THE ADULT OF MORIBAETIS MACAFERTI
(EPHEMEROPTERA: BAETIDAE) 1

W. P. McCafferty, C. R. Lugo-Ortiz 2

ABSTRACT: Alate stages of the common and distinctive Central American and southern Mexi-
can species Moribaetis macaferti are described for the first time based on males of both the adult
and subimago stages. The species is distinct as an adult in having forewings with extensive stain-
ing, an abdomen with distinctive patterning, and genital forceps with a smooth medial margin.
The species' abdominal color pattern is also apparent in the subimago. A key to the male adults
of Moribaetis species is provided. Moribaetis macaferti is reported from Panama for the first
time.

The distinctive mayfly genus Moribaetis was described by Waltz and
McCafferty (1985). It was later restricted by Lugo-Ortiz and McCafferty
(1996a) and shown to be a monophyletic, relatively ancestral lineage within
the New World Baetodes complex of genera, which also includes the genera
Baetodes Needham and Murphy, Mayobaetis Waltz and McCafferty, and
Prebaetodes Lugo-Ortiz and McCafferty. Moribaetis is Neotropical in origin
(McCafferty et al. 1992), and the entire Baetodes complex is evidently South
American in origin (McCafferty 1998), although several species of Baetodes
are found in the southwestern Nearctic region.

Moribaetis essentially is a Central American genus. It currently consists of
M. macaferti Waltz, M. maculipennis (Flowers), and M. salvini (Eaton). Men-
tion of the genus as occurring in South America (Lugo-Ortiz and McCafferty
1996a) was inadvertent, although an unspecified report of the genus by Rojas
de Hernandez et al. (1995) could actually be applicable to either Moribaetis or
Mayobaetis because they were considered subgenera of Moribaetis at the time.
The latter may be more likely because Mayobaetis is known from Ecuador and
Peru as well as Central America.

Moribaetis macaferti has recently been reported from southern Mexico in
addition to Costa Rica, Honduras, and Guatemala (Lugo-Ortiz and McCafferty
1994, 1996b). This is a relatively large, striking species of Baetidae that has
been collected by WPM from streams and waterfalls, including splash zones,
in Costa Rica. A key to the larvae of all three species of Moribaetis was pro-
vided by Waltz and McCafferty (1985). Although M. maculipennis and M.
salvini are known as adults (Eaton 1885, Flowers 1979), the adult of M.
macaferti has remained unknown up to this time.

Boris Kondratieff (Fort Collins, Colorado) has kindly given us larvae and
an adult of M. macaferti that he recently collected from Puebla, Mexico in

1 Received August 2, 1997. Accepted September 2, 1997.

2 Department of Entomology, Purdue University, West Lafayette, IN 47907, USA.

ENT. NEWS 109(2) 1 17-121, March & April, 1998

1 1 8 ENTOMOLOGICAL NEWS

1994. In addition, Wills Flowers (Tallahassee, Florida) has kindly loaned us
larvae and a reared subimago of M. macaferti that represent a new country
record (see material examined, below). We are thereby able to provide a first
description of the adult M. macaferti and the first species key to male adults of
Moribaetis.

Moribaetis macaferti Waltz

Male adult. Dimensions in mm: body length, 9.7; forewing length, 7.6; forewing width, 3.5;
hindwing length, 1.2; caudal filaments length, 20.0. Head (Figs. 1 and 2): Coloration generally
light yellowish brown. Turbinate portion of compound eyes smoke yellow, nearly round with
anterior-posterior length only slightly longer than width, nearly contiguous, and mounted on
short stalks with encircling band of suffused black. Scape and pedicel of antennae distally marked
with brown. Thorax (Figs. 1 and 3): Pronotum (Fig. 1 ) cream medially and light brown laterally.
Antecosta of mesotergum (Fig. 3) suffused dark brown. Meso- and metanota cream with medium
and light brown elongated markings as illustrated in Figure 3, including conspicuous narrow-
elongate U-shaped submedian medium brown mark on scutum. Forefemora light to medium
brown; foretibiae medium brown, with length 1 .32x that of femora; foretarsal segment 1 medium
brown; foretarsal segments 2-5 cream basally and light brown distally. Mid- and hindlegs cream
with femora and tarsi light brown distally; tibiae subequal in length to femora. Claws dissimilar,
each consisting of one hooked-acute and one straight-ovoid member. Forewings with distinct
markings as illustrated in Figure 4; yellowish brown stains generally associated with crossvenation
but more extensive in basal area of membrane with large free spots in basal costal and subcostal
area and extensions from crossveins near MA2 and M?2- Hindwings as in Figures 5 and 6;
membrane devoid of staining; costal process acute and asymmetrical (straight edged posteri-
orly); first elongate vein unforked; second elongate vein forked in distal half, with forks encom-
passing free marginal intercalaries; third elongate vein short, terminating in anal margin. Abdo-
men (Fig. 7): Terga light basally, becoming more suffused with yellowish light brown shading
posteriorly (terga 6-10 markedly becoming progressively more shaded); tergum 1 with narrow
lateral medium brown markings; terga 2-8 with additional triangular medium brown markings
extending length of terga and filling posterolateral corners of terga; tergum 9 with lateral mark-
ings more extensive anteriorly and not well developed posteriorly, with additional pair of small
oblique dashes sublaterally at posterior margin; tergum 10 pale and unmarked. Sterna 1-6 whit-
ish, almost translucent; sterna 7-9 pale yellow, almost translucent; sternum 9 pale yellow. Geni-
talia (Fig. 8) with short, medially smooth basal segment, elongate second segment, and small,
rounded terminal segment weakly demarcated from second segment. Caudal filaments, for most
of length, distinctly banded in alternating pattern of one white and three darkened segments.
Male subimago. Generally similar to adult. Turbinate portion of compound eyes appearing slightly
more oval-elongate. Color pattern not well developed on body and not yet evident in forewings
as described above. Submedian elongate marking on thoracic scutum not formed into U-shape,
but unconnected posteriorly. Subcostal process of hindwings similar to adults but appearing mi-
nutely hooked, probably as an artifact of the subimaginal pellicle; only one major intercalary
evident between forks of second elongate vein. Triangular posterolateral abdominal tergal mark-
ings evident but not highly contrasting as in adult. Genitalia and caudal filaments missing.

Material examined. MEXICO, PueblaSta, Pahuatlan, Hwy 106, VIII- 17- 1994, B. C. Kondratieff,
male adult and associated larvae (left fore- and hindwings slide mounted). PANAMA, Bocas del
Toro Prov., trib. of Rio Guabo at pipeline rd., V-28-1985, R. W. Flowers (male subimago and
associated larvae).

Vol. 109, No. 2, March & April, 1998

119

Figs. 1-8. Moribaetis macaferti, male adult. 1. Head and pronotum. 2. Head (lateral view). 3.
Meso- and metanota. 4. Left forewing. 5. Left hindwing (to scale with forewing). 6. Left hindwing
(enlarged). 7. Abdomen (lateral view). 8. Male genitalia (ventral view).

DISCUSSION

The adult male described above was not reared; however, we have no doubt
that it is M. macaferti. It was taken where only M. macaferti larvae were taken
(B. C. Kondratieff, pers. comm.), is specifically distinct from other adults in
the genus, and essentially agrees with the subimago from Panama that was
indeed reared from larvae of M. macaferti (R. W. Flowers, pers. comm.).

The forewings of M. macaferti have more extensive staining than those of
M.maculipennis [see Flowers (1979): Fig. 1] and M. salvini [see Eaton (1885):
Fig. 29a]. The hindwings are most similar to those of M. maculipennis in vena-
tion but differ in not having any staining [see Flowers (1979): Figs. 2 and 3]. It
is possible that the number of marginal intercalaries between the forks of the
second elongate vein in the hindwings is variable within M. macaferti, based
on a comparison of the hindwings of the subimago from Panama and the adult
from Mexico, and thus we do not recommend attempting to identify adults of
species of Moribaetis on that basis. The abdominal terga of M. macaferti (Fig.

120 ENTOMOLOGICAL NEWS

7) possess distinct posterolateral markings on segments 2-8, and additionally
lack the distinctive large brown submedian markings of M. maculipennis [see
Flowers (1979): Fig. 5] and the pitch-black posterior marginal markings of the
terga described by Eaton (1885). There are, however, very slight and difficult
to detect indications of a pair of pale spots on most abdominal terga in M.
macaferti. The terminal segment of the genital forceps of M. macaferti (Fig. 8)
is most similar to that of M. maculipennis [see Flowers (1979): Fig. 4] in that
it is weakly demarcated from segment 2. However, M. macaferti lacks the
roughened medial margin of the basal segment of the forceps that is present in
both M. maculipennis and M. salvini [see also Kimmins (1934): Fig. 13], in
addition to the well-defined rounded and setose medioapical protrusion of the
basal forceps segment seen in M. maculipennis [Flowers (1979): Fig. 4].

Key To Male Adults ofMoribaetis Species

[key to larvae in Waltz and McCafferty (1985)]

1. Forewings with extensive staining throughout membrane (Fig. 4); abdominal terga 2-8 with
triangular posterolateral markings (Fig. 7); forceps with medial margin of basal segment smooth

medially (Fig. 8) M. macaferti

I 1 . Forewings [Flowers (1979): Fig. 1; Waltz and McCafferty (1985): Fig. 43] and abdominal
terga not as above; forceps [Kimmins (1934): Fig. 13; Flowers (1979): Fig. 4] with medial mar-
gin of basal segment roughened 2

2. Cerci banded with two to three dark segments alternating with two light segments [Waltz and
McCafferty (1985): Fig. 33] M. maculipennis

2'. Cerci banded with one slightly darkened segment alternating with three light segments [Waltz
and McCafferty (1985): Fig. 42] M. salvini

ACKNOWLEDGMENTS

We thank R. W. Flowers (Florida A & M Univ.) and B. C. Kondratieff (Colorado State Univ.)
for collecting and providing the specimens that have made this paper possible. The paper has
been assigned Purdue Agricultural Research Program Journal No. 15467.

LITERATURE CITED

Eaton, A. E. 1883-88. A revisional monograph of recent Ephemeridae or mayflies. Trans. Linn.

Soc. Lond., 2nd Ser. Zool. No. 3, 352 pp.
Flowers, R. W. 1979. A new species of Baetis from Panama (Ephemeroptera: Baetidae). Pan-

Pac. Entomol. 55: 187-191.
Kimmins, D. E. 1934. Notes on the Ephemeroptera of the Godman and Salvin collection, with

descriptions of two new species. Ann. Mag. Nat. Hist., Ser. 10. 14: 338-353.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1994. New records of Ephemeroptera from Mexico.

Entomol. News 105: 17-26.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1996a. Phylogeny and classification of the Baetodes

complex (Ephemeroptera:Baetidae), with description of a new genus. J. N. Am. Benthol.

Soc. 15:367-380.

Vol. 1 09, No. 2, March & April, 1 998 121

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996b. New Central American and Mexican records
of Ephemeroptera species. Entomol. News 107: 303-310.

McCafferty, W. P. 1998. Ephemeroptera and the great American interchange. J. N. Am. Benthol.
Soc., in press.

McCafferty, W. P., R. W. Flowers and R. D. Waltz. 1992. The biogeography of Mesoamerican
mayflies, pp. 173-193 In: S. P. Darwin and A. L. Weldon(eds.). Biogeography of Mesoamerica:
proceedings of a symposium. Tulane Stud. Zool. Bot, Suppl. Publ. No. 1.

Rojas de Hernandez, A. M., M. del Zuniga de Cardoso, M. H. Burbano T. and C. Serrato H.
1 995. Altitudinal distribution of Ephemeroptera in the Farallones de Cali National Park, Co-
lombia, pp 121-129 In: L. D. Corkum and J. J. H. Ciborowski (eds.). Current directions in
research on Ephemeroptera. Can. Scholars' Press, Toronto.

Waltz, R. D. and W. P. McCafferty. 1985. Moribaetis: a new genus of Neotropical Baetidae
(Ephemeroptera). Proc. Entomol. Soc. Wash. 87: 239-251.

BOOKS RECEIVED AND BRIEFLY NOTED

METHODS IN ECOLOGICAL & AGRICULTURAL ENTOMOLOGY. D.R.
Dent & M.P. Walton, eds. 1997. CAB International. 387 pp.

Citing the need for an up-to-date and comprehensive textbook on experimental and analyti-
cal methods in both ecological and agricultural entomology, this book seeks to integrate new
technologies and approaches to research with traditional methods, to provide a balanced view of
the subject for final year undergraduates and postgraduates, and to be a useful source of refer-
ence for research workers. Twenty three authors contributed chapters on thirteen subjects rang-
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populations and insect migration, host plant selection, insect pollination, insecticide efficacy and
efficacy of natural enemies, and modelling.

THE EVOLUTION OF MATING SYSTEMS IN INSECTS AND
ARACHNIDS. J.C. Choe & B.J. Crespi, eds. 1997. Cambridge University Press.
387 pp. $44.95 pbk, $100 hdbk.

The complex mixtures of conflict and cooperation that typify sex and sociality make these
among the most endlessly fascinating and difficult topics in ecology and evolution. The twenty
one chapters in this volume, contributed by thirty six authors, explore the intricacies of sexual
and social competition and the variety of ways in which males and females pursue, persuade,
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insect mating systems by Thomhill and Alcock (1983) and complement the recent synthesis of
the study of sexual selection by Andersson (1994).

PROGRESS AND PROSPECTS IN EVOLUTIONARY BIOLOGY: THE
DROSOPHILA MODEL. J.R. Powell. 1997. Oxford University Press. 562
pp. Cloth $70.00.

The author states the primary impetus for this book is the conviction that if a major advance
in understanding evolution is to be achieved, it is most likely to come from integration of levels
of knowledge centered on a single closely related set of organisms. Drosophila has the greatest
chance of being this organism. Thus, this book is about empirical studies of Drosophila. and is
directed primarily toward graduate students but also to evolutionary biologists and colleagues
studying development, molecular biology, and neurobiology of Drosophila.

1 22 ENTOMOLOGICAL NEWS

FIRST REPORT AND NEW SPECIES OF THE GENUS

CLOEODES (EPHEMEROPTERA: BAETIDAE)

FROM AUSTRALIA 1 ' 2

C. R. Lugo-Ortiz, W. P. McCafferty 3

ABSTRACT: Cloeodesfustipalpus, new species, and C. illiesi, new species, are described from
larvae from eastern Australia. The two species represent the first report of Cloeodes from the
continent. Cloeodes fustipalpus is distinguished by the irregular labral setation, clublike labial
palps segment 3, and abdominal color pattern. Cloeodes illiesi is distinguished by the bifid right
prostheca with a medially setose branch, reduced maxillary palps, medially bulbous labial palps
segment 3, abdominal color pattern, and narrow-elongate gills. Numerous morphological char-
acteristics indicate that C. fustipalpus and C. illiesi are most closely related to the Afrotropical
C. inzingae and the Oriental C. longisetosus and C. soldani. Three biogeographic scenarios are
discussed that would explain the world distribution of Cloeodes.

Traver (1938) erected the genus Cloeodes (Ephemeroptera: Baetidae) for
the Caribbean species C. maculipes Traver and C. consignatus Traver. The
genus is distinct among small minnow mayflies because its larvae have eden-
tate tarsal claws (Fig. 6; Waltz and McCafferty 1987b: Fig. 8), a conspicuous
subproximal arc of long, fine, simple setae on the tibiae (Fig. 6; Waltz and
McCafferty 1987b: Fig. 7), and setal tufts on sterna 2-6 (Waltz and McCafferty
1987a: Fig. 5; Waltz and McCafferty 1987b: Figs. 9, 44). Adults of Cloeodes
are distinguished by having segment 2 of the male genital forceps basally bul-
bous and with abundant minute, fine, simple setae (Waltz and McCafferty
1987b:Fig. 34).

Cloeodes has been reported from the Afrotropics, Neotropics, Orient, and
southwestern Nearctic (Traver 1938, Waltz and McCafferty 1987ab, 1994,
Kluge 1991, Flowers 1991, Lugo-Ortiz and McCafferty 1993, 1994, 1995,
McCafferty and Lugo-Ortiz 1995, McCafferty et al. 1997). Herein we report
Cloeodes for the first time from Australia. The report is based on two new
species described from larvae collected from Queensland and New South Wales.
The specimens studied are housed in the Purdue Entomological Research Col-
lection, West Lafayette, Indiana.

The discovery of Cloeodes in Australia is of considerable biogeographic
interest because it is the first genus of Baetidae known to occur throughout the
Southern Hemisphere. Cloeodes may have been widespread in Gondwanaland
during the Jurassic approximately 1 80 million years ago (mya). Southern Hemi-
sphere landmasses drifting to their present positions would have carried an-

1 Received July 28, 1997. Accepted August 27, 1997.

2 Purdue Agricultural Research Program Journal No. 15460.

3 Department of Entomology, Purdue University, West Lafayette, IN 47907.

ENT. NEWS 109(2) 122-128 , March & April, 1998

Vol. 109, No. 2, March & April, 1998 123

cestral species of Cloeodes with them. The South East Asian distribution could
be explained by the northward drifting and abutment of the Indian subconti-
nent approximately 45 mya. The southern Nearctic distribution of the genus
has been explained by northward dispersal during and after the formation of
the Isthmus of Panama approximately 6-5 mya, as reviewed by McCafferty
(1998). Alternatively, Cloeodes may have originated somewhat later on the
African-Indian-South American landmass (essentially West Gondwanaland)
during the Early Cretaceous approximately 1 40 mya. This explanation is simi-
lar to the first, except that dispersal of the genus from Asia into Australia dur-
ing the Middle Miocene approximately 15 mya is assumed. A third historical
explanation would have Cloeodes originating in Africa-South America during
the Middle Cretaceous approximately 1 10-100 mya. At that time, however, the
genus might not have been present on the Indian subcontinent, and the pres-
ence of Cloeodes in the Orient, Australia, and North America would all be
attributed to dispersal events beginning in the Eastern Hemisphere when Af-
rica and Eurasia reunited during the middle Miocene approximately 17 mya.
Interchange between Africa and Asia in this period is consistent with some
other animal groups (e.g., see Cox and Moore 1985).

We cannot at this time be sure which of these three biogeographic explana-
tions is the most likely because only a small number of species of Cloeodes are
known and cladistic analysis is not possible. Considerable insular evolution of
the genus has occurred in South America since the isolation of that continent,
as evidenced by the fact that Western Hemisphere Cloeodes are a closely re-
lated, distinctive grouping (Waltz and McCafferty 1987b). The fact that all
species in the Eastern Hemisphere are closely related to each other (see spe-
cies discussions below) would further suggest that African, Asian, and Austra-
lian lineages have dispersed relatively recently, not having been isolated from
each other to the extent in which Western and Eastern Hemisphere lineages of
the genus have.

Cloeodes fustipalpus Lugo-Ortiz and McCafferty, NEW SPECIES

Larva. Body length: 5.1-6.0 mm. Caudal filaments length: 2.3-2.5 mm. Head: Coloration
light yellow-brown, with faint vermiform markings on vertex. Antennae approximately 1 .5x length
of head capsule. Labrum (Fig. 1 ) with submedial pair of long, fine, simple setae and submarginal
row of six to eight fine, simple setae of various lengths. Hypopharynx similar to Figure 1 1 . [Left
and right mandibles (Figs. 2, 3) with outer incisors worn in material examined.] Left mandible
(Fig. 2) with inner incisor with three denticles; prostheca robust, apically denticulate; minute
denticles present between prostheca and mola. Right mandible (Fig. 3) with inner incisor with
four denticles; prostheca somewhat slender, apically acute; minute denticles present between
prostheca and mola. Maxillae (Fig. 4) with three long, fine, simple setae near medial hump;
maxillary palps reaching galealaciniae; palp segment 1 approximately 0.50x length of segment
2. Labium (Fig. 5) with glossae and paraglossae equal in length; palp segment 1 approximately
O.SOx length of segments 2 and 3 combined; segment 2 approximately 1 .20x length of segment 3;

1 24 ENTOMOLOGICAL NEWS

segment 3 bulbous, clublike (medially broader than apical width of segment 2). Thorax: Colora-
tion pale yellow-brown, with complex markings. Hindwingpads absent. Legs (Fig. 6) cream;
femora with dorsal row of five to eight long, robust, simple setae, last two almost contiguous and
longer than others; tibiae with dorsal row of long, fine, simple setae; tarsi with dorsal row of
long, fine, simple setae. Abdomen (Fig. 7): Coloration pale brown and cream; segment 1 cream;
segment 2 pale brown, with submedial and sublateral pairs of large, oblong, cream markings;
segment 3, 5, and 6 pale brown, with submedial pair of circular cream markings; segment 4
anteriorly and posteriorly pale brown, medially cream; segment 7 anteriorly pale brown, posteri-
orly cream; segment 8 cream; segment 9 pale brown, with semicircular anteromedial cream mark-
ing; segment 10 pale brown. Sterna cream. Gills (Fig. 8) subtriangular, well tracheated, with
smooth margin. Paraprocts (Fig. 9) with 18-20 sharp marginal spines, increasing in size apically;
abundant scale bases scattered over surface. Caudal filaments whitish; medial caudal filament
approximately O.SOx length of cerci.

Adult. Unknown.

Material examined. Holotype: Larva, AUSTRALIA, New South Wales, Chandler R., 26
mi E of Armidale, no date, G. F. Edmunds. Paratypes: Larva, same data as holotype [mouthparts,
left foreleg, gill 4, and paraproct mounted on slide (medium: Euparal)]; two larvae, AUSTRA-
LIA, New South Wales, Bellinger R., at Bellinger, 11-23-1966, G. F. Edmunds. Additional mate-
rial: Three exuviae, AUSTRALIA, New South Wales, Serpentine R., New England National Park,
11-19-1966, G. F. Edmunds; three larvae, AUSTRALIA, New South Wales, Bellinger R., at
Bellinger, 11-23-1966, G. F. Edmunds.

Etymology. The specific epithet is a combination of the Latin words fustis
(club) and palpus (palps). It is in reference to the clublike labial palps.

Discussion. Cloeodes fustipalpus is distinguished from other members of
the genus by the irregular setation of the labrum (Fig. 1), clublike segment 3 of
the labial palps (Fig. 5), and abdominal color pattern (Fig. 7). The abdominal
color pattern varies somewhat among specimens, but the most consistent is the
one shown in Figure 7.

Cloeodes fustipalpus appears to be related to the Oriental species C.
longisetosus (Braasch and Soldan) and C. soldani (Muller-Liebenau), the
Afrotropical species C. inzingae (Crass), and C. illiesi, new species, from Aus-
tralia (see below). The larvae of all these species have a bulbous labial palp
segment 3 (Figs. 5, 15; Crass 1947: Fig. 9a; Braasch and Soldan 1980: Fig. 12;
Muller-Liebenau 1983: Fig. 3b), well-developed rows of long, fine, simple
setae on the tibiae and tarsi (Fig. 6; Braasch and Soldan 1980: Figs. 2, 3; Waltz
and McCafferty 1994: Fig. 2), and lack hindwingpads.

Cloeodes illiesi Lugo-Ortiz and McCafferty, NEW SPECIES

Larva. Body length: 3.9 mm. Caudal filaments length: unknown. Head: Coloration yellow-
brown to medium brown, with faint vermiform markings on vertex. Antennae approximately
1.5x length of head capsule. Labrum (Fig. 10) with submedial pair of long, simple setae and
submarginal row of three to four long, fine, simple setae. Hypopharynx as in Figure 11. Left
mandible (Fig. 12) with six denticles; prostheca robust, apically denticulate; minute denticles
between prostheca and mola absent. Right mandible (Fig. 13) with outer incisor with four den-
ticles; inner incisor with three denticles; prostheca slender, bifid, one branch medially with minute,
fine, simple setae; minute denticles between prostheca and mola absent. Maxillae (Fig. 14) with

Vol. 109, No. 2, March & April, 1998

125

Figs. 1-9. Cloeodesfustipalpus, new species, larva. 1. Labrum. 2. Left mandible. 3. Right man-
dible. 4. Right maxilla. 5. Labium (left-ventral; right-dorsal). 6. Left foreleg. 7. Abdomen (dor-
sal). 8. Gill 4. 9. Paraproct.

126

ENTOMOLOGICAL NEWS

three long, fine, simple setae near medial hump; maxillary palps not reaching galealaciniae; palp
segment 1 approximately 0.60x length of segment 2. Labium (Fig. 1 5) with glossae and paraglossae
equal in length; palp segment 1 as long as segments 2 and 3 combined; segment 2 approximately
0.74x length of segment 3; segment 3 bulbous, apically flattened. Thorax: Coloration pale to
medium yellow-brown, with complex markings. Hindwingpads absent. Legs (similar to Fig. 6)
cream; femora with dorsal row of five to seven long, robust, simple setae, last two almost con-
tiguous; tibiae with dorsal row of long, fine, simple setae; tarsi with dorsal row of long, fine,
simple setae. Abdomen (Fig. 16): Coloration pale brown and yellow-brown; segment 1 yellow-

Figs. 10-18. Cloeodes illiesi, new species, larva. 10. Labrum. II. Hypopharynx. 12. Left man-
dible. 13. Right mandible. 14. Right maxilla. 15. Labium (left-ventral; right-dorsal). 16. Abdo-
men (dorsal). 17. Gill 6. 18. Paraproct.

Vol. 1 09, No. 2, March & April, 1 998 1 27

brown; segment 2 pale brown, with submedial anterior pair of large yellow-brown oblong mark-
ings and sublateral oblong yellow-brown markings; segments 3-6 pale brown, with submedial
anterior pair of small subtriangular yellow-brown markings; segment 7 pale brown; segment 8
yellow-brown; segment 9 pale brown, with faint medial streak; segment 10 pale brown. Sterna
cream to yellow-brown. Gills (Fig. 1 7) narrow-elongate, poorly tracheated, with smooth margin.
Paraprocts (Fig. 18) with 9-10 sharp marginal spines; scale bases scattered over surface. Caudal
filaments whitish.

Adult. Unknown.

Material examined. Holotype: Larva, AUSTRALIA, Queensland Province, nr. Cairns,
Cascade Falls, sea level, 20C, X- 13- 1966, J. lilies [mouthparts, left foreleg, and paraproct mounted
on slide (medium: Euparal)].

Etymology. This species is named after the late renowned entomologist-
limnologist Joachim lilies, who collected it.

Discussion. Cloeodes illiesi is distinguished from other members of the
genus by the bifid right prostheca with a setose branch (Fig. 1 3), reduced max-
illary palps (Fig. 14), medially bulbous labial palps segment 3 (Fig. 15), ab-
dominal color pattern (Fig. 16), and narrow-elongate gills (Fig. 17). Its pos-
sible relationships to other species of Cloeodes are discussed above under C.
fustipalpus.

ACKNOWLEDGMENTS

We thank G. F. Edmunds, Jr. (Salt Lake City, Utah) for the donation of the material used in
this study.

LITERATURE CITED

Braasch, D. and T. Soldan. 1 980. Centroptella n. gen., eine neue Gattung der Eintagsfliegen aus

China (Baetidae, Ephemeroptera). Reichenbachia 18: 123-127.
Cox, C. B. and P. D. Moore. 1985. Biogeography: an ecological and evolutionary approach, 4th

edition. Blackwell Scientific Pub., London.

Crass, R. S. 1947. The may-flies of Natal and the Eastern Cape. Ann. Natal Mus. 11: 37-1 10.
Flowers, R. W. 1991. Diversity of stream-living insects in northwestern Panama. J. N. Am.

Benthol. Soc. 10: 322-334.
Kluge, N. 1991. Cuban mayflies of the family Baetidae (Ephemeroptera) 1. Genera Callibaetis,

Cloeodes, and Paradoeodes. Zool. Zh. 12: 128-136. [in Russian]
Lugo-Ortiz, C. R. and W. P. McCafferty. 1993. Genera of Baetidae (Ephemeroptera) from

Central America. Entomol. News 104: 193-197.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1994. New records of Ephemeroptera from Mexico.

Entomol. News 105: 17-26.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1995. Annotated inventory of the mayflies

(Ephemeroptera) of Arizona. Entomol. News 106: 131-140.
McCafferty, W. P. 1998. Ephemeroptera and the great American interchange.]. N. Am. Benthol.

Soc., in press.

128 ENTOMOLOGICAL NEWS

McCafferty, W. P. and C. R. Lugo-Ortiz. 1995. Cloeodes hydation, n. sp. (Ephemeroptera:

Baetidae): an extraordinary, drought tolerant mayfly from Brazil. Entomol. News 106: 29-

35.
McCafferty, W. P., C. R. Lugo-Ortiz, and G. Z. Jacobi. 1997. The mayfly fauna of New

Mexico. Gr. Basin Natural., 57: 283-314.
Miiller-Liebenau, I. 1983. Three new species of the genus Centroptella Braasch and Soldan,

1 980, from Sri Lanka (Insecta: Ephemeroptera). Arch. Hydrobiol. 97: 486-500.
Traver, J. R. 1938. Mayflies of Puerto Rico. J. Agric. Univ. P. R. 22: 5-42.
Waltz, R. D. and W. P. McCafferty. 1987a. Generic revision of Cloeodes and description of two

new genera (Ephemeroptera: Baetidae). Proc. Entomol. Soc. Wash. 89: 177-184.
Waltz, R. D. and W. P. McCafferty. 1987b. Revision of the genus Cloeodes Traver

(Ephemeroptera: Baetidae). Ann. Entomol. Soc. Am. 80: 191-207.
Waltz, R. D. and W. P. McCafferty. 1994. Cloeodes (Ephemeroptera: Baetidae) in Africa. Aq.

Insects 16: 165-169.

BOOKS RECEIVED AND BRIEFLY NOTED

THE BIONOMICS OF GRASSHOPPERS, KATYDIDS AND THEIR KIN.
S.K. Gangwere & M.C. & M. Muralirangan, eds. 1997. CAB International.
529 pp.

This volume consists of twenty chapters authored by an international group of twenty eight
scientists and is written from a broad, comparative biological, behavioral, and evolutionary ap-
proach best expressed by the term bionomics. It focuses on history and recent developments in
grasshopper and plague locust biology as well as the biology of katydids, crickets, and other
Orthoptera, an insect group of exceptional economic and biological interest.

INSECT ECOLOGY, 3rd ed. P.W. Price. 1997. J. Wiley & Sons. 874 pp.

Moving from the dynamics of plant-insect interactions, predation, parasites and hosts, as
well as mutualistic relationships, including pollination ecology, this book examines the themes
central to understanding the role of insects in our environment. It describes the various levels of
insect interaction, from trophic relationships, populations, and communities, while unfolding the
infinite variety of insect species and their visible legacy in the fossil record. This new edition
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Vol. 109, No. 2, March & April, 1998 129

DIEL EMERGENCE PATTERNS OF

TRICORYTHODES STYGIATUS
(EPHEMEROPTERA: LEPTOHYPHIDAE)

ON THE LITTLE LEHIGH CREEK
NEAR ALLENTOWN, PENNSYLVANIA 1

Fran Cough, Bruce L. Haase^

ABSTRACT: Tricorythodes stygiatus subimagos and adults show a diel periodicity in drift rates.
Most of their activity is centered around the hours of sunrise and sunset. There is a differential
emergence of male and female subimagos, males emerge after sunset and females emerge after
sunrise. Light periodicity is the primary influence in the timing of T. stygiatus emergence, molt-
ing, and oviposition. This species is univoltine at this study site.

Tricorythodes stygiatus McDunnough (Ephemeroptera: Leptohyphidae) is
a common species in the limestone streams of southeastern Pennsylvania . The
species emerges throughout the summer. Tricorythodes species have been de-
scribed as univoltine (Newell and Minshall 1978), bivoltine (Hall 1975), or
multivoltine (Newell and Minshall 1978) depending on geographic location
and water temperatures.

Tricorythodes species are atypical of most mayflies in that the subimago
emerges from the larval shuck underwater and swims/floats/crawls to the sur-
face, with males emerging after dark and females emerging early the next
morning (Hall 1975; Edmunds and McCafferty 1988).

In Hall's (1975) study of T. allectus Needham, the surface drifting of male
subimagos took place from sunset until the early hours of the morning, while
female subimagos showed a peak in surface drift rate around sunrise. The
emergence of male subimagos of T. allectus was correlated with low light inten-
sity in the evening, while female subimago emergence is correlated with in-
creasing light intensity in the morning (Hall 1975). However, in a study by
Newell and Minshall (1978) T. minutus always emerged at the water surface,
usually in the afternoon. T. minutus seems to be little influenced by photope-
riod.

T. allectus male subimagos molt to adults before dawn, so the length of the
male subimago stage is 5-7 hours. The female's subimago stage is very short.
Some female subimagos molt immediately after emergence, others may wait
up to 2 hours (Hall 1975). The male alate life of T. allectus lasts about 9-10
hours, beginning after dusk until the sun is well above the horizon. The female's
alate stage spans a few short hours after dawn (Hall, Berner and Cook 1975).

1 Received June 9, 1997. Accepted September 20, 1997.

2 Department of Biological Sciences, East Stroudsburg University, East Stroudsburg, Pa. 1 830 1 .

ENT. NEWS 109(2) 129-135 , March & April, 1998

130 ENTOMOLOGICAL NEWS

In T. minutus, the subimago stage usually lasts less than 30 minutes and the
alate forms live less than 6 hours (Newell and Minshall 1978).

The purpose of this study is to determine if there is a differential emer-
gence of male and female subimagos of T. stygiatus in the Little Lehigh Creek
in Allentown, PA., and to determine if this differential emergence can be cor-
related with water temperature or time of day.

METHODS

This study took place on the Little Lehigh Creek, a 4th order stream in
Salisbury Township, Lehigh County, Pennsylvania. The study site was at the
end of a riffle in the main channel of the stream. It is a limestone spring creek
with a typical riffle-pool complex, with a substrate composed mainly of marl
and silt.

Sampling was done using a Munro style drift net with a 250 micron mesh
size and a net opening measuring 39cm x 39cm and a length of approximately
one meter. A removable (250 cm) catch basket, composed of 3 inch PVC pipe
and mesh screen was attached to the end of the drift net. The frame, made of
0.5 inch PVC pipe, was slid over steel reinforcing rods that were hammered
into the stream bottom. When in place, the Munro drift net extended from the
stream bottom through the water's surface (anywhere from 5 cm to 15 cm
above the surface depending on water levels).

The net was placed in the selected riffle of the study area for 24 hours at a
time. Samples were removed and the net replaced every hour during the study
period. At this time, the stream temperature was also taken and recorded. These
24 hour study periods were conducted three times, about one month apart, on
July 1 3, August 1 1 , and September 8 of 1995.

The samples were rinsed from the net into a pan, concentrated with an
aquarium net and preserved in 70% ethanol. Using a 20x Swift dissecting mi-
croscope, these insects were counted, aged (as subimagos or adults) and the
subimagos sexed. To determine the life stage of the alate T. stygiatus, the meth-
ods described by Edmunds and McCafferty (1988) were used. Subimagos had
translucent wings with cilia along the hind edge. Adults had transparent wings
without cilia on the hind edge of the wing. Subimagos and adults were sexed
as males if claspers were present at the end of the abdomen, and as females if
claspers were absent. Voucher specimens are deposited at the Department of
Biological Sciences, East Stroudsburg University, East Stroudsburg, Pa.

RESULTS

The largest numbers of alate T. stygiatus were caught in the July 1995
sample with 891 organisms collected, including 73 subimagos and 818 adults.
During the July 13th sample period, the water temperatures ranged from 16
C. in the early morning hours to 21 C. in the late afternoon. Sunrise was at

Vol. 109, No. 2, March & April, 1998

131

0542 and sunset was 2033 so the day length was 14 hours 51 minutes. The
peak drift time for male subimagos occurred between 2200 and 2400; for
female subimagos this peak occurred between the hours of 0600 and 0900
(Table 1 , Figure 1 ). Adults showed a peak drift rate between the hours of 0800
and 1 100 (Table 1, Figure 2).

Fifty- four alate T. stygiatus were collected in the August drift samples,
including 8 subimagos and 46 adults. During the August 1 1th sample period,
the water temperatures ranged from 15 C. in the early morning hours to 19
C. in the late afternoon. Sunrise was at 0608 and sunset was 2006, so the day
length was 13 hours and 58 minutes. The peak drift time for male subimagos
occurred between 2100 and 2300, for female subimagos this peak occurred
between 0900 and 1000 (Table 1, Figure 1). Adults showed a peak drift rate
between the hours of 0900 and 1000 (Table 1 , Figure 2).

Table 1 . Hourly drift rates of Tricorythodes stygiatus subimagos and adults taken over a 24 hour
period in the Little Lehigh Creek.

Time

0100
0200
0300
0400
0500
0600
0700
0800
0900
1000
1100
1200
1300
1400
1500
1600
1700
1800
1900
2000
2100
2200
2300
2400
Total

Subimago Male Subimago Female Adults

7/13/95 8/11/95 9/08/95 7/13/95 8/11/95 9/08/95 7/13/95 8/11/95 9/08/95

1
1

8
4
11
28

1
1

349

335

818

132

ENTOMOLOGICAL NEWS

Fifty-five alate T. stygiatus were collected in the September drift samples,
including 4 subimagos and 51 adults. During the September 8th sample pe-
riod, the water temperatures ranged between 15 C. and 16 C. Sunrise was at
0636 and sunset was 1921, so the day length was 12 hours 45 minutes. The
peaks in drift time for male subimagos were at 1800 and 2100, for female
subimagos these peaks were at 0700 to 0900 (Table 1 , Figure 1 ). Adults showed
a peak in drift rate from 0700 to 1 100 (Table 1 , Figure 2).

Number of
Subimagos

7/13/95
8/11/95 Date
9/08/95

2400

Figure 1A. Hourly drift rate of Tricorythodes stygiatus male subimagos in the Little Lehigh
Creek.

Number of
Subimagos

7/13/95
8/11/95 Date
9/08/95

2400

Figure IB. Hourly drift rate of Tricorythodes stygiatus female subimagos in the Little Lehigh
Creek.

Vol. 109, No. 2, March & April, 1998

133

400

350

300

250

Number of
Adulti

200

150

100

1 Q B O D D

IB a n

a fa) a o e

0600

1200
Time

1800

2400

Figure 2. Hourly drift rate of Tricorythodes stygiatus adults in the Little Lehigh Creek.

DISCUSSION

Alate forms of T. stygiatus adults on the Little Lehigh show a diel period-
icity in their drift rates. Water temperature did not appear to trigger the differ-
ential emergence of T. stygiatus in this study. The temperature of the stream
did not fluctuate much (+ 2 C.) on a 24 hour basis, or on a seasonal basis ( 1 5-
21 C.) throughout the study period. Instead, the peak for subimago and adult
drift revolves around the hours of sunrise and sunset. This can be seen in the
data as the season progresses and the day length shortens. It is apparent that
light periodicity is the primary influence in the timing of the T. stygiatus alate
life activities of emergence, molting, mating and oviposition. This agrees with
the findings of Hall (1975) and Friesen, Flannagan and Lautersweiler (1980)
(T. allectus).

There was differential emergence of male and female subimagos of T.
stygiatus in the Little Lehigh. This is similar to what Hall (1975), Friesen,
Flannagan and Lautersweiler ( 1 980) and Edmunds and McCafferty ( 1 988) have
found in other parts of North America for other species of Tricorythodes. In

134 ENTOMOLOGICAL NEWS

found in other parts of North America for other species of Tricorythodes. In
this study 83% (30 of 36) of the male subimagos emerged in the first 3 hours
after sunset, and 93% (43 of 49) of the female subimagos emerged in the first
3 hours after sunrise. About 98% (894 of 915) of the spent adults were col-
lected from 1 to 5 hours after sunrise, so mating and ovipostion had occurred
prior to this event. Mating and oviposition occur for about 3-4 hours after
sunrise based on the first/last appearance of spent wing adults in the drift
samples. This is a somewhat longer period than was observed by Hall, Berner
and Cook (1975) in T. allectus. These findings were similar to R. J. Hall's
( 1 975) study of T. allectus except that male subimago emergence did not occur
steadily through the night, while the peak emergence of female subimagos
occurred after sunrise.

The short life span of Tricorythodes species has been well documented
(Hall 1975; Hall, Berner and Cook 1975; Edmunds and McCafferty 1988). On
the Little Lehigh, alate males live 8-14 hours and alate females live 2-5 hours
on average, based on the time from first/last appearance of subimagos and the
first/last appearance of adults.

It appears that the T. stygiatus on the Little Lehigh Creek are univoltine
based on this study. This would agree with Newell and Minshall's (1978) study
on T. minutus. The drift rate of alate T. stygiatus was many times greater in
July than in August or September. In this study, 89% (891 of 996) of the total
drift is accounted for in the July sample alone.

In a number of the T. stygiatus larvae, subimagos and adults, a distinct red
coloration on the thorax was noted. In the 0600 hour sample on July 13, 1995,
70% of the female subimagos had a red thorax. The cause of this red colora-
tion is another area for further investigation. A number of causes have been
hypothesized from 1) an element of water chemistry (Gary Borger personal
communication 1995), 2) parasitic mites (Greg Hoover personal communica-
tion 1996, Brittain 1982), to 3) a type of polymorphism (Peters & Peters 1977).

ACKNOWLEDGMENTS

We would like to acknowledge the following people who helped make this project a success:
for much needed assistance from Rod Rohrbach, Rich Heiserman, Steve Hoovler, Joe Kohler,
Gary Pyle and other members of the Little Lehigh Fly Fishers and Little Lehigh Trout Unlimited;
to Andy Shiels of the PA Fish and Boat Commission and Don Marushak of the Allentown Parks
Dept., to Jim Munro for his drift nets; to Mark Gerber for his word processing skills; and the
primary author gives special thanks to his wife, Maura, for her endless patience and understand-
ing. The identification of this species as T. stygiatus was verified by Don Baylor of Aquatic
Resource Consulting of Saylorsburg, PA and by Greg Hoover, Penn State University Department
of Entomology. The authors would like to thank R.D. Waltz and W.P. McCafferty for their sug-
gestions in reviewing this article.

Vol. 109, No. 2, March & April, 1998 135

LITERATURE CITED

Brit lain, J. E. 1 982. Biology of mayflies. Ann. Rev. Entomol. 27: 1 1 9- 1 47.

Edmunds, G. F and W. P. McCafferty. 1988. The mayfly subimago. Ann. Rev. Entomol. 33:
509-529.

Friesen, M. K., J. F. Flannagan and P. M. Laufersweiler. 1 980. Diel emergence patterns of
some mayflies (Ephemeroptera) of the Roseau River (Manitoba, Canada). Advances in
Ephemeropteran Biology . Plenum Publishing Co. p. 287-296.

Hall, R. J. 1975. Life history, drift and production rate of the stream mayfly, Tricorythodes
atratus McDunnough in the headwaters of the Mississippi River. Ph.D. Thesis, Univ. of Min-
nesota. 288 pp. (T. allectus as T. atratus)

Hall, R. J., L. Berner and E. F. Cook. 1975. Observations on the biology of Tricorythodes
atratus McDunnough. Proc. Entomol .Soc. Wash. 77 (1): 34-49.

Hall, R. J., T. F. Waters and E. F. Cook E. F. 1980. The role of drift dispersal in production
ecology of a stream mayfly. Ecology 61 (1): 37-43.

Newell, R. L., and G. W. Minshall. 1978. Life history of a multivoltine mayfly, Tricorythodes
minutus: an example of the effect of temperature on the life cycle. Ann. Entomol. Soc. Am.
71: 876-881.

Peters, W. L., and J. G. Peters. 1 977. Adult life and emergence ofDolania americana in north-
western Florida. Int. Revue ges Hydrobiol. 62(3): 409-438.

136 ENTOMOLOGICAL NEWS

THE FIRST FOSSIL DORYLINAE WITH NOTES ON

FOSSIL ECITONINAE
(HYMENOPTERA: FORMICIDAE) 1

Mark B. DuBois 2 > 3 ' 4

ABSTRACT: Two army ant species, Dorylus molestus and Neivamyrmex iridescens, are reported
from copal of undetermined age (possibly Upper Pliocene and Pleistocene respectively). The
presence of such fossils is remarkable given the disposition of many such species to foraging
underground. Until more precise dates for this material are available, historical and biogeographic
implications of these fossils remain uncertain.

Ants are represented in the fossil record from Cretaceous through Pleis-
tocene (Carpenter 1992; Bolton 1994). The currently recognized classification
includes 16 extant subfamilies and four extinct subfamilies (Bolton 1994). A
number of species are known from impressions in Tertiary shale (Carpenter
1930) and are mostly represented by reproductives. Many ants have been pre-
served in amber (Wheeler 1914; Mayr 1868; Wilson 1985) and are mostly
represented by workers. Foraging workers were trapped in resins which be-
came amber, while reproductives flying over bodies of water were drowned
and buried in silt (which later became shale). Some species, such as Stenamma
berendti (Mayr), are known only from reproductives preserved in amber (Mayr
1868). Most publications have concentrated on pre-Pleistocene fossils; a few
have dealt with sub-fossils (Francoeur and Elias 1985; DuBois 1993). Limited
work has been done on specimens preserved in copal, which is usually Pleis-
tocene or Pliocene in age (Poinar 1992, Larsson 1978).

Most fossil worker ants belong to the extant subfamilies Formicinae,
Dolichoderinae, Myrmicinae, and Ponerinae. Though some wing fragments
have been tentatively identified as belonging to army ants, until recently, no
fossil worker army ants were known. Wilson (1985) described Neivamyrmex
ectopus from workers preserved in amber in the Dominican Republic. These
fossils are thought to be late Oligocene or early Miocene in age. The rarity of
fossil worker army ants is probably due to their behavior because many spe-
cies principally forage underground.

Recently, I had the opportunity to study fossil ants of both Ecitoninae and
Dorylinae preserved in copal. Although copal is considered much younger in

1 Received Dec. 27 1996. Accepted January 31 1997.

2 1 16 Burton St., Washington, Illinois 61571-2509, U.S.A.

3 Research Affiliate, Center for Biodiversity, Illinois Natural History Survey, 607 East Peabody
Drive, Champaign, Illinois 61820, U. S. A.

4 Send reprint requests to Washington, Illinois address.

ENT. NEWS 109(2) 136-142, March & April, 1998

Vol. 109, No. 2, March & April, 1998 137

age (Recent, Pleistocene or Pliocene), these specimens provide a link (geo-
logically and biogeographically) with older material. Both species are briefly
discussed below.

There appears to be a vast number of ant specimens preserved in copal
from various deposits. The species discussed above represent a minuscule por-
tion of available material. I hope this paper will stimulate further study of such
fossils and subfossils. The precise age of the material discussed below is un-
known (this is true for much copal). Although ages ranging from Recent through
Pliocene have been proposed for various copal deposits, the stratigraphy of
sites containing this material (and associations with micro-fossils) must be
established to determine proper age.

Dorylus molestus (Gerstacker)

This species can be distinguished from related Dorylus species through the
following combination of characters: head with convex sides; posterior angles
of head pointed; ventral lobe of petiole well developed (Raignier and Van Boven
1955). Specimens appear identical to modern forms (as described in Raignier
and Van Boven 1955). Bill Gotwald examined selected specimens and con-
cluded they were this species ("... the common driver ant of East Africa, espe-
cially Kenya and Tanzania.") (Gotwald, pers. comm.). Wheeler (1922: 740)
gave the distribution of this species in Eastern Africa (including Mombassa,
Mt. Kenia (2400 - 2800 m), Bura Mountains, Freretown, and Naivasha, Rift
Valley).

Two hundred and seventy one specimens contained in 12 pieces of copal
were examined. This material was collected along the Tanzania - Kenya bor-
der (Tanzania side, approximately 100 km south-west of Mombassa, Kenya).
Individual pieces of copal were covered in caliche and found in a layer 1 1 m
below the soil surface (Alan Graffham, pers. comm.). Specimen disposition is
as follows. Lloyd Davis, Jr. provided two pieces of copal containing Dorylus.
These have been returned to him. All remaining pieces discussed are from my
personal collection and were obtained from Allen Graffham and other sources.
Of these, one piece containing Dorylus has been donated to the British Mu-
seum of Natural History and another piece containing Dorylus has been do-
nated to the Museum of Comparative Zoology.

Schluter and von Gnielinski (1987) indicated this copal was derived al-
most entirely from resin of Hymenaea verrucosa (Leguminosae: Caesalpina-
ceae). Deposition sites were listed as in the forest soil (primary), in the man-
grove fringed estuary (secondary), and along the beach (tertiary).

These deposits are part of the Mikindani beds (Stockley 1928). Stockley
(1928) assigned an age of Upper Pliocene. Poinar (1992) indicated this Tanza-
nian copal may be Pliocene in age (1.6-5 Ma). Schluter and von Gnielinski

1 38 ENTOMOLOGICAL NEWS

(1987: 11) presented an age of Pleistocene, but cautioned that ". . . strati-
graphically well defined associations of copal with index- or guide-fossils do
not exist." von Gnielinski (pers. comm.) reiterated the uncertainty with the
dating of copal from that region (Pleistocene or Pliocene could be argued).
Additionally, Ken Anderson who is currently studying amber and copal depos-
its from around the world (pers. comm.) indicated these were "resins of unde-
termined geological age, but probably not of great antiquity."

Schluter and von Gnielinski ( 1 987: 1 8) further stated: "Generally, it can be
assumed that the fauna of the East African copal does not show significant
differences from the fauna living today, i.e., all the trapped specimens repre-
sent recent species..." Although a number of insects have been reported from
these deposits (including nine species of Hymenoptera), no ants have previ-
ously been reported (Schluter and von Gnielinski 1987).

Figures 4 through 8 illustrate typical features of these specimens. Figures
1 1 and 12 show the general appearance of these specimens within the copal.
Other arthropods were preserved with the ants. Their state of preservation,
coupled with problems in identification of many East African insects, did not
permit precise identifications. Specimens include other ants (several male
Ponerinae, two male Myrmicinae, three worker Dolichoderinae, one minor
worker Pheidole sp., one worker Crematogaster sp.), four Chalcidoidea, one
alate termite, and a number of flies, beetles, leafhoppers, and spiders. After
examining the fossils in cross section, it appears a foraging column of driver
ants became trapped on the sticky resin and were then covered with another
resin flow.

Neivamyrmex iridescens Borgmeier

This species can be distinguished from related Neivamyrmex species
(Borgmeier 1955 - Group VII) through the following combination of charac-
ters: postpetiole about five-sixths as long as high; head with violet reflections
(difficult to see in fossil material); postpetiole about as long as wide, higher
than long with posterior surface of node more steeply sloping than anterior
surface; head smooth and distinctly shining; apex of scape closer to upper
margin of head than eye level (Watkins 1976). Julian Watkins II identified the
Neivamyrmex in one of these samples. He indicated the fossil is most similar
to specimens of Neivamyrmex iridescens from Cundinamarca, Colombia in
his collection. The only significant difference is a slightly longer than usual
petiole. (Julian Watkins II, pers. comm.). Specimens key to N. iridescens using
Watkins (1976) and fit the description provided by Borgmeier (1955: 540 -
542). This species is presently known from scattered localities: Panama,
Guianas, Surinam, Bolivia, Colombia (Borgmeier 1955; Watkins 1976).

Four specimens contained in two pieces of copal were examined from Co-
lombia. Disposition of specimens: these Neivamyrmex remain in my collec-

Vol. 109, No. 2, March & April, 1998

139

tion. They originally came from Colombia via Mr. Allen Graffham.

The specific locality for this material is unknown, but Schlee (1984) indi-
cated this may be from "Pena Blanca." This material bears locality informa-
tion of Santander Department, Colombia. Allen Graffham (pers. comm.) indi-
cated it came from a roadcut and provided several photographs. He indicated
that he doubts any deposition has occurred in this area for a very long time as
it is quite mountainous with very steep slopes. I have examined fossil ants in
copal from both Santandar and Boyaca departments. Most material was col-
lected at a depth of 2 - 3 meters in Santandar and up to 10 meters in Boyaca.
Poinar (1992) indicated all known Colombian material is of Pleistocene age.
Ken Anderson (pers. comm.) also considered these were "resins of undeter-
mined geological age, but probably not of great antiquity."

This copal is presumed to be derived from a species of Hymenaea, but
details regarding this locality, deposition, and origin of this material are sketchy
at best. It is not known to this author whether any species of Hymenaea live in

Figures 1 - 8. Scale varying. Legs, pilosity, and sculpture omitted. Figures 1 - 3. Neivamyrmex
iridescens. Figure 1 . Worker, lateral view. Figure 2. Head, profile of occipital vertex. Remainder
of head obscured by position within copal. Figure 3. Petiole and postpetiole, dorsal view. Figures
4 - 8. Dorylus molestus. Figure 4. Minor worker, lateral view. Figure 5. Major worker, head, full
face view. Figure 6. Major worker, lateral view. Figure 7. Worker, petiole and postpetiole, dorsal
view. Figure 8. Major worker, mandibles and anterior clypeal margin.

140

ENTOMOLOGICAL NEWS

Figures 9-12. Photographs of fossilized army ants and driver ants. Scale varying. Figures 9-10.
Neivamyrmex iridescens. Figure 9. Damaged specimen (damaged previously by polishing). Fig-
ure 10. Worker, lateral view. Figures 11-12. Dorylus molestus. Figure 1 1 . Numerous workers in
copal. Figure 12. Major worker, lateral view.

Vol. 109, No. 2, March & April, 1998 141

the area today.

Figures 1 - 3 show typical features of these specimens. Figures 9 and 10
depict the general appearance of these specimens within the copal. Other
arthropods were preserved with the ants. Their state of preservation coupled
with problems in identification of South American insects did not permit pre-
cise identifications. Specimens include two other ants (one minor worker
Pheidole sp., one worker A nochetus sp.), one Chalcidoidea, one fly, one worker
termite, one set of termite wings, and one spider. Since only four ants were
trapped, it is possible that they were on the periphery of a foraging column
which encountered the resin.

ACKNOWLEDGMENTS

This paper is dedicated to the memory of the late Frank M. Carpenter (Museum of Compara-
tive Zoology, Harvard Univ. advisor and friend for the past 2 decades). His encouragement was
appreciated.

My continuing thanks is extended to Jeri and Benjamin DuBois for their support and under-
standing. They make it all possible. Lloyd Davis (Gainesville, Florida) provided some specimens
of Dorylus. Julian Watkins II (Baylor University, Waco, Texas) identified some specimens of
Neivamyrmex iridescent. Bill Gotwald (Utica College of Syracuse University, Utica, New York)
identified some specimens of Dorylus molestus. Merrill Foster (Bradley University, Peoria, Illi-
nois) put me in contact with Allen Graffham (Ardmore, Oklahoma) who provided numerous
comments regarding localities and possible ages for the ant fossils discussed herein. The major-
ity of fossil ants discussed herein also came from Allen Graffham. F. von Gnielinski (Geological
Survey of Queensland, Brisbane, Queensland) provided insights into the age and conditions of
deposition of the African copal. Thanks also to Ken Anderson (Argonne National Laboratory,
Argonne, Illinois) who reviewed his current analyses of assigning ages to copal and amber world-
wide and discussed his view of the age of these resins with me.

This paper has been reviewed by Wallace LaBerge and Don Webb (Illinois Natural History
Survey, Champaign, Illinois), Julian Watkins II, Stefan Cover (Museum of Comparative Zool-
ogy, Harvard University), and three anonymous reviewers. Where possible, I have tried to incor-
porate their comments. I accept responsibility for any remaining errors or omissions.

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Cambridge, MA. 222 pp.
Borgmeier, T. 1955. Die Wanderameisen der Neotropischen Region (Hym. Formicidae). Studia

Entomol. 3: 1 -716.
Carpenter, F. M. 1930. The fossil ants of North America. Bull. Mus. Comp. Zool., Harvard

Univ. 70: 1 - 66.
Carpenter, F. M. 1992. Part R, Arthropoda 4, Volumes 3 and 4, Superclass Hexapoda. In Kaesler,

R. J., E. Brosius, J. Keim, J. Priesner (eds.). Treatise on Invertebrate Paleontology, xxii + 655

p. 1489 figs. Univ. Kansas Press, Lawrence, KS. [Formicidae, pp. 490 - 495].
DuBois, M. B. 1993. What's in a name? A clarification of Stenamma westnoodi, S. debile, and 5.

lippulum (Hymenoptera: Formicidae: Myrmicinae). Sociobiology 21(3): 299 - 334.
Francoeur, A. and S. A. Elias. 1985. Dolichoderus taschenbergi Mayr (Hymenoptera: Formicidae)

from an early Holocene fossil insect assemblage in the Colorado Front Range. Psyche 92

(2-3): 303 - 307.

142 ENTOMOLOGICAL NEWS

Larsson, S. G. 1978. Baltic amber - a palaeobiological study. Entomonograph Vol. 1 . Klampen-
borg, Denmark. 192 pp.

Mayr, G. 1868. Die Ameisen des baltischen Bemsteins. Beitr. Naturk. Preussens 1:1- 102.

Poinar, G. O., Jr. 1992. Life in Amber. Stanford Univ. Press, Stanford, Calif, xiii + 350 pp., 8 pi.

Raignier, A. and J. Van Boven. 1 955. Etude taxonomique, biologique et biometrique des Dorylus
du sous-genre Anomma (Hymenoptera Formicidae). Ann. Musee Royal Congo Beige n.s. 4
(Sci. Zool.) 2: 1 - 359.

Schlee, D. 1984. Notizen ubereinige Bemsteine und Kopule aus aller Weit. Stuttg. Beitr. Natkde.
18:29-38.

Schluter, T. and F. von Gnielinski. 1987. The East African Copal Its geologic, stratigraphic,
palaeontologic significance and comparison with fossil resins of similar age. Nat. Mus. Tan-
zania Occ. Pap. 8: 1 - 34.

Stockley, G. M. 1928. Report on the geology of the Zanzibar Protectorate. 126 pp. Zanzibar.

Watkins, J. F., II. 1976. The identification and distribution of New World army ants (Dorylinae:
Formicidae). Markham Press Fund Baylor Univ. Press, Waco, TX, x + 102 pp.

Wheeler, W. M. 1914 (1915). The ants of the Baltic amber. Schriften der Physikalisch
Okonomischen Gesellschaft zu Konigsberg 55: 1-142.

Wheeler, W. M. 1922. Ants of the American Museum Congo Expedition, a contribution to the
myrmecology of Africa, I: On the distribution of the ants of the Ethiopian and Malagasy
regions; II: The ants collected by the American Museum Congo Expedition; VII: Keys to the
genera and subgenera of ants; VIII: A synonymic list of the ants of the Ethiopian region; IX:
A synonymic list of the ants of the Malagasy region. Bull. Amer. Mus. Nat. Hist. 45( 1 ): 13-
27, 39 - 269, 631 - 710, 71 1 - 1004 1005 - 1055.

Wilson, E. O. 1985. Ants of the Dominican amber (Hymenoptera: Formicidae). 2. The first
fossil army ants. Psyche 92 (1): 11 - 16.

Vol. 109, No. 2, March & April, 1998 143

COCCINELLIDAE (COLEOPTERA)

IN APPLE ORCHARDS OF EASTERN

WEST VIRGINIA AND THE IMPACT

OF INVASION BY HARMONIA AXYRIDIS 1

M. W. Brown, S. S. Miller 2

ABSTRACT: Twenty-five species of Coccinellidae were found on apple in eastern West Virginia
from 1983 to 1996. From 1989 through 1994, the exotic coccinellid, Coccinella septempunctata
(first collected in 1983), has dominated the fauna of the tribe Coccinellini. Another adventive
species, Harmonia axyridis (first collected in 1994), became the dominant species in the tribe
Coccinellini in 1995, and continues to dominate the coccinelline guild on apple. Harmonia axyridis
has displaced C. septempunctata and is providing better biological control of Aphis spiraecola
on apple in eastern West Virginia.

Coccinellids (Coleoptera: Coccinellidae) are an important group of aphid
predators in many ecosystems. Because of their ability to contribute to the
control of pests, many coccinellids have been selected for introduction in bio-
logical control programs. As of 1985, there had been 179 introductions of
coccinellids into the U. S., with 26 species having become established; how-
ever, 8 were a result of accidental introductions (Gordon 1985). Two recent
coccinellid immigrants have made an obvious impact in the mid-Atlantic states.
In 1973, Coccinella septempunctata L. was first found in New Jersey (Angalet
et al. 1979), and it gradually spread throughout the Northeast (Schaefer et al.
1987). In 1988, H. axyridis was first found in Louisiana (Chapin and Brou
1991), and has since spread into the mid- Atlantic region (Day et al. 1994).
Although both species had been released intentionally, their establishment ap-
pears to be a result of accidental introductions (Day et al. 1994). Both species
are now abundant throughout much of the northeastern United States and east-
ern Canada (Hoebeke and Wheeler 1996) and have even become subjects of
public concern because of occasional massive migrations of C. septempunctata
(Schaefer et al. 1987) and by large numbers of H. axyridis overwintering in
houses (Knodel and Hoebeke 1996).

There have been numerous studies cataloguing the coccinellids in apple
orchards in Europe and North America. In Europe, more than 23 species of
coccinellids have been collected on apple, but usually 3 to 14 species in any
one study (Hodek and Honek 1996). In a study spanning 4 years in 3 regions
of Hungary, 23 coccinellid species were found on apple (Marko et al. 1 995). In
Europe, Adalia bipunctata (L.) and C. septempunctata dominate in apple

1 Received July 7, 1997. Accepted September 6, 1997.

2 USDA, ARS, Appalachian Fruit Research Station, 45 Wiltshire Road, Keameysville, WV 25430.

ENT. NEWS 109(2) 136-142, March & April, 1998

144 ENTOMOLOGICAL NEWS

orchards. The number of coccinellid species recorded from North American
orchards has been similar to that reported in Europe: 7 species in Ontario (Smith
1957), 17 in another study in Ontario (Hagley 1974), 15 in Quebec (LeRoux
1960), 8 in Washington state (Carroll and Hoyt 1984), and 6 in Pennsylvania
(Horsburgh and Asquith 1 968). Putman ( 1 964) found 1 species of coccinellids
in peach orchards in Ontario. In Washington, Coccinella transversoguttata
Faldermann was the most abundant (Carroll and Hoyt 1984) but in the east, the
dominant species varied temporally and spatially. In this paper, we character-
ize the coccinellid fauna of apple orchards in eastern West Virginia. Data are
taken from several studies covering a 14-year period, 1983 to 1996. These
years span the time of the invasion of//, axyridis into the region and are used
to evaluate the effect of H. axyridis on other species in the tribe Coccinellini,
and on Aphis spiraecola Patch (Homoptera: Aphididae) populations.

MATERIALS AND METHODS

Data from several studies were used to characterize the Coccinellidae in
apple orchards in eastern West Virginia. Except for the years 1993 to 1995,
both conventionally managed and unsprayed orchards were sampled in each
study. Data from 1983 to 1984 were taken from Brown and Adler (1989), and
Brown et al. (1988). These studies covered the region from Virginia to New
York, but only data from West Virginia are used in the present paper. Sampling
consisted of visual examination of 7 branches from 6 trees in each of 4 or-
chards, repeated 6 times per year. The purpose of the study was to examine the
phytophagous community structure on apple, but occurrence, not abundance,
of natural enemies was also recorded. Data from 1 984 to 1 988 were from Brown
and Welker (1992) using whole-tree visual sampling from 5 to 10 trees from
each of 3 orchards, 4 to 6 times per year. Data from 1989 to 1990 were from
Brown (1993), sampling in 2 orchards, 2 branches from each of 10 trees in
1989 and 5 trees in 1990, 6 samples per year. Data from 1991 were taken from
limb-jarring samples from 3 branches of 6 apple trees each hour over a 24-
hour period 4 times during the summer. Limb-jarring samples from 2 peach
trees per hour and one sour cherry tree every other hour provide data for
Coccinellidae on those tree species. A second study in 1991 added data for
Coccinellidae on peach trees (Brown and Puterka 1997) and used observations
on 6 branches from each of 6 trees in 3 orchards at 5 times during the summer.

Coccinellid data from 1992 to 1996 were based largely on two experimen-
tal orchards planted in 1992: one managed with conventional methods, and the
second managed with reduced insecticides and diverse ground cover plantings
underneath the trees. In 1992, sampling involved two, 15-tree transects per
orchard 6 times per year (Brown and Lightner 1997). Every terminal on each
sample tree was examined for aphids and predators. Monthly samples from 10
randomly selected trees per orchard, 10 shoots per tree, were collected from

Vol. 109, No. 2, March & April, 1998 145

1993 to 1995 (Brown etal. 1997). In 1996 the same two orchards were sampled
more intensively to estimate populations of A. spiraecola and aphid predators.
Ten terminals on 20 randomly selected trees were sampled weekly in May and
June, then biweekly into September whenever aphids were present. In 1992
and 1996, three other orchards were also sampled with the same methodology
as described above for those years.

Identification of adult Scymninae, Sticholotidinae, and Psylloborini were
made by R. D. Gordon, USDA, ARS, Systematic Entomology Laboratory,
Washington, DC. Adult Coccinellinae (except Psylloborini) and Chilocori-
nae were identified using Dillon and Dillon (1961).

Data on populations of A. spiraecola, the predominant aphid on apple trees
in West Virginia (Pfeiffer et al. 1989), resulted from pest monitoring in three
conventionally managed apple orchards. Ten terminals on two trees from each
orchard were sampled and the number of leaves per terminal infested with
aphids was recorded. Sampling was conducted every week from early May to
September from 1992 to 1996.

Assessment of the impact of//, axyridis on coccinellids was limited to the
tribe Coccinellini. Any effect on the endemic fauna would be expected to be
greatest on members of the same tribe because of their generally similar biolo-
gies and niche requirements.

RESULTS AND DISCUSSION

Twenty-five species of coccinellids in 4 subfamilies and 9 tribes were found
on apple in eastern West Virginia from 1983 to 1996 (Table 1). Fourteen spe-
cies of coccinellids were found on peach, including Scymnus caudalis LeConte,
which was not found on apple, and 7 on sour cherry in limited sampling on
these host trees (Table 1). In addition to those species listed, 4 others, all be-
longing to the Coccinellini, were caught in sticky or blacklight traps located in
apple orchards but were not seen foraging on fruit trees: C. transversoguttata,
Hippodamia convergens Guerin, H. tredecimpunctata tibialis (Say), and
Mulsantina picta (Randall). The 25 species listed in Table 1 represent a larger
and more comprehensive list of the coccinellid fauna on apple than other stud-
ies because it covers a 14-year period, whereas all previous studies, except for
Marko et al. (1995), covered only 1 to 3 years.

For the 8 years in which numerical data are available (1989 to 1996), the
two exotic species, C. septempunctata and H. axyridis, dominated the
coccinelline fauna (Table 2). Although 7 species of Coccinellini were recorded
on apple during these years, over 70% of the individuals collected belonged to
the two exotic species. Coccinella septempunctata was first collected from
apple in West Virginia in 1983. By 1985, C. septempunctata was the most
frequently encountered coccinelline in orchards, and it was the only coccinelline

146

ENTOMOLOGICAL NEWS

Table 1. Coccinellidae, by subfamily and tribe (after Kovar 1996), found in
apple orchards of eastern West Virginia, 1983-1996.

Scymninae

Stethorini

Stethorus punctum (LeConte) 1

Scymnillini (Zilini)
Zilus horni Gordon

Scymnini

Diomus terminates (Say) 1 ,2
Scymnus fraternus LeConte
S. circumspectus Horn 2
5. iowensis Casey 1 ,2
S. rubricaudus Casey
Scymnus sp. 1

Brachiacanthini

Brachiacantha ursina (F.)

Hyperaspidini

Hyperaspis proba (Say) 1,2
Chilocorinae

Chilocorini

Chilocorus stigma (Say) 1,2

Coccinellinae

Coccinellini

Adalia bipunctata (L.) '
Anatis labiculata (Say)
Anatis mali (Say) 1
Coccinella novemnotata Herbst
C. septempunctata L. 1 ,2
C. transversoguttata Faldermann
Coleomegilla maculata lengi
Timberlake 1
Cycloneda munda (Say) 1
C. sanguinea (L.)
Harmonia axyridis (Pallas)
Hippodamia parenthesis (Say)
Olla v-nigrum (Mulsant) 1

Psylloborini

Psyllobora vigintimaculata (Say) 2
Sticholotidinae

Microweiseini

Microweisea misella (LeConte) 1

Also found on peach trees; Scymnus caudalis LeConte (Scymninae: Scymnini) was found only
on peach trees.

Also found on sour cherry trees.

Table 2. Percentage composition of the coccinelline fauna on apple in eastern West Virginia,
1989-1996.

Species

1989-90

1991

1992

1993

1994

1995

1996

C. septempunctata

72.7

86.4

99.5

93.2

79.4

9.1

10.2

A. bipunctata

9.1

2.3

1.7

C. munda

9.1

4.5

0.5

A. labiculata

9.1

C. maculata lengi

2.3

5.1

2.9

27.3

1.1

O. v-nigrum

4.5

H. axyridis

17.7

63.6

88.7

Number observed

192

265

Vol. 109, No. 2, March & April, 1998

147

seen in 1987 and 1988 during sampling of the same orchards (Kozar et al.
1994). Coccinella novemnotata Herbst, which Wheeler and Hoebeke (1995)
suggest may be negatively correlated with C. septempunctata, has not been
seen in West Virginia apple orchards since 1985. In a survey prior to the arrival
of C. septempunctata in nearby south-central Pennsylvania (Horsburgh and
Asquith 1968), the only Coccinellini found were A. bipunctata, C. munda, and
H. convergens.

Harmonia axyridis, first collected in 1994, has replaced C. septempunctata
as the dominant coccinelline in apple orchards (Fig. 1). In 1995, C.
septempunctata was reduced to the third most abundant species of Coccinellini
behind H. axyridis and Coleomegilla maculata lengi Timberlake (Table 2). Of

1 00%

C. septumpunctata _J Harmonia axyridis LJ All others

Fig. 1. Percentage composition of the coccinelline fauna in apple orchards of eastern '.Vest Vir-
ginia from 1990 to 1996, showing the displacement of C. septempunctata by H. axyridis.

148 ENTOMOLOGICAL NEWS

the less abundant coccinellids, A. bipunctata, Cycloneda munda (Say), Anatis
labiculata (Say), and Olla v-nigrum (Mulsant) have not been seen in apple
orchards in the three years since H. axyridis appeared. Only C. maculata lengi
has seemed to become more abundant on apple since the arrival of//, axyridis.
Harmonia axyridis has shown its ability to rapidly dominate in other ecosys-
tems in other regions where it has invaded (Tedders and Schaefer 1994, Day et
al. 1994).

Comparing the two years in which intensive sampling of aphid predators
was conducted, 1992 and 1996, gives insight into the interaction between C.
septempunctata and H. axyridis (Table 3). In 1992, all but one coccinelline

Table 3. Number of Coccinellini collected in two intensive surveys of aphid predators in eastern
West Virginia apple orchards.

1992

1996

Species

Adults

Immatures

Adults

Immatures

C. septempunctata

C. munda

H. axyridis

187

C. maculata lengi

Unidentified

was C. septempunctata, with about equal numbers of adults and immatures.
By 1996, two years after H. axyridis was first detected, no C. septempunctata
immatures were seen on apple trees. The 12 unidentified immatures were
either egg masses or first instar larvae that we could not identify to species.
The reduction in total numbers of C. septempunctata from 1992 to 1996 sug-
gests that //. axyridis not only replaced C. septempunctata as the dominant
species but also largely displaced it from apple, especially in its use of apple as
a larval habitat. It has been shown in Japan that H. axyridis larvae prey on C.
septempunctata brucki Mulsant, but predation in the reverse direction does not
occur (Hironori and Katsuhiro 1997).

The displacement of C. septempunctata by H. axyridis has had a positive
impact on biological control of A. spiraecola in West Virginia apple orchards
(Fig. 2.). Aphid populations were much lower from 1994 to 1996 than in pre-
vious years; peak aphid populations were lower and the duration of aphid in-
festations was shorter, particularly in 1996 (Fig. 2). The aphid data were from
orchards sprayed with insecticides, explaining the sharp declines in popula-
tion abundance in most years. Harmonia axyridis was first found in apple or-
chards in 1994, but it did not dominate until 1995, when aphid populations
appeared to be suppressed. Only in 1995 and 1996, the two years in which //.

Vol. 109, No. 2, March & April, 1998

149

120

140

160

1991

180
Day
1992

200

220

240

1993

I
I

240

1996

Fig. 2. Population estimates of A. spiraecola by day of the year in three conventionally managed
orchards in eastern West Virginia; A, 1991 to 1993, prior to arrival of H. axyridis; B, 1994-1996,
after arrival of//, axyridis.

150 ENTOMOLOGICAL NEWS

axyridis dominated the coccinelline fauna, were insecticide sprays not applied
to control aphids. Data on aphids were obtained only from a few trees in three
orchards in only one location and, therefore, cannot be considered conclusive.
The indication, however, is that the arrival of H. axyridis has improved the
biological control of aphids on apple.

Conclusions: Exotic species of Coccinellini have greatly affected the
coccinelline fauna of West Virginia apple orchards. First, C. septempunctata
dominated the fauna, but it was in turn displaced by the newly arrived H. axyridis
in 1995. The overall effect of H. axyridis on native Coccinellini has not been
evaluated, but since the arrival of C. septempunctata, native coccinellines have
represented only a very minor component of the fauna on apple. Aphid bio-
logical control has been enhanced since the arrival of//, axyridis, so this latest
addition to the coccinelline fauna has been beneficial in the apple agroeco-
system.

ACKNOWLEDGMENTS

We thank R. D. Gordon (USDA ARS, Systematic Entomology Laboratory, Beltsville, MD)
for identifications of the Scymninae, Sticholotidinae, and Psylloborini; L. Claire Stuart, Cynthia
R. L. Adler, Donald C. Weber, Jeffrey J. Schmitt, and V. Larry Crim for their data collection; and
William H. Day, Henry W. Hogmire, Jeffrey J. Schmitt, and Natalia J. Vandenberg for their com-
ments on an earlier draft of this paper.

LITERATURE CITED

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States: recolonizations and notes on its ecology. Environ. Entomol. 8: 896-901.

Brown, M. W. 1993. Resilience of the natural arthropod community on apple to external distur-
bance. Ecol. Entomol. 18: 169-183.

Brown, M. W. and C. R. L. Adler. 1989. Community structure of phytophagous arthropods on
apple. Environ. Entomol. 18: 600-607.

Brown, M. W., C. R. L. Adler, and R. W. Weires. 1988. Insects associated with apple in the
mid-Atlantic states. New York's Food and Life Sci. Bull. No. 124, 1-31.

Brown, M. \V., D. M. Glenn, and T. van der Zwet. 1997. Impact of ground cover plants on pest
management in West Virginia, USA, apple orchards. Hortic. Sci. (Prague) 24:39-44.

Brown, M. W. and G. W. Lightner. 1997. Recommendations on minimum experimental plot
size and succession of aphidophaga in West Virginia, USA, apple orchards. Entomophaga.
42: 259-269.

Brown, M. W. and G. J. Puterka. 1997. Orchard management effects on the arthropod commu-
nity on peach with comparisons to apple. J. Entomol. Sci. 32: 165-182.

Brown, M. W. and W. V. Welker. 1992. Development of the phytophagous arthropod commu-
nity on apple as affected by orchard management. Environ. Entomol. 21: 485-492.

Carroll, D. P. and S. C. Hoyt. 1984. Natural enemies and their effects on apple aphid, Aphis
pomi DeGeer (Homoptera: Aphididae), colonies on young apple trees in central Washington.
Environ. Entomol. 13: 469-481.

Chapin, J. B. and V. A. Brou. 1 99 1 . Harmonia axyridis (Pallas), the third species of the genus to
be found in the U.S. (Coleoptera: Coccinellidae). Proc. Entomol. Soc. Wash. 93: 630-635.

Vol. 109, No. 2, March & April, 1998 151

Day, W. H., D. R. Prokrym, D. R. Ellis, and R. J. Chianese. 1994. The known distribution of
the predator Propylea quatuordecimpunctata (Coleoptera: Coccinellidae) in the United States,
and thoughts on the origin of this species and five other exotic lady beetles in eastern North
America. Entomol. News 105: 244-256.

Dillon, E. S. and L. S. Dillon. 1961. A manual of common beetles of eastern North America.
Row, Peterson & Company. Evanston, 111.

Gordon, R. D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. J. New York
Entomol. Soc. 93: 1-912.

Hagley, E. A. C. 1974. The arthropod fauna in unsprayed apple orchards in Ontario. II. Some
predacious species. Proc. Entomol. Soc. Ont. 105: 28-40.

Hironori, Y. and S. Katsuhiro. 1997. Cannabalism and interspecific predation in two ladybirds
in relation to prey abundance in the field. Entomophaga. 42: 153-163.

Hoebeke, E. R. and A. G. Wheeler. 1 996. Adventive lady beetles (Coleoptera: Coccinellidae) in
the Canadian maritime provinces, with new eastern U.S. records. Entomol. News 107: 281-
290.

Hodek, I. and A. Honek. 1996. Ecology of Coccinellidae. Kluwer Academic Publishers. Bos-
ton.

Horsburgh, R. L. and D. Asquith. 1968. Initial survey of arthropod predators of the European
red mite in south-central Pennsylvania. J. Econ. Entomol. 61: 1753-1754.

Knodel, J. J. and E. R. Hoebeke. 1996. Multicolored Asian lady beetle, Harmonia axyridis
(Pallas) Coleoptera: Coccinellidae. Cornell Coop. Ext., Misc. Pest Management Fact Sheet,
page 101.00.

Kovar, I. 1996. Phylogeny. In: Hodek, I. and A. Honek. Ecology of Coccinellidae. Kluwer Aca-
demic Publishers. Boston, pp: 19-31.

Kozar, F., M. W. Brown and G. Lightner. 1994. Spatial distribution of homopteran pests and
beneficial insects in an orchard and its connection with ecological plant protection. J. Appl.
Entomol. 117:519-529.

LeRoux, E. J. 1960. Effects of "modified" and "commercial" spray programs on the fauna of
apple orchards in Quebec. Ann. Soc. Entomol. Quebec 6: 87-121.

Mark6, V., O. Merkl, A. Podlussany, K. Vig, Cs. Kutasi, and S. Bogya. 1995. Species compo-
sition of Coleoptera assemblages in the canopies of Hungarian apple and pear orchards. Acta
Phytopathol. Entomol. Hung. 30: 221-245.

Pfeiffer, D. G., M. W. Brown, and M. W. Yarn. 1989. Incidence of spirea aphid (Homoptera:
Aphididae) in apple orchards of Virginia, West Virginia, and Maryland. J. Entomol. Sci. 24:
145-149.

Putman, W. L. 1964. Occurrence and food of some coccinellids (Coleoptera) in Ontario peach
orchards. Can. Entomol. 89: 1 149-1 155.

Schaefer, P. W., R. J. Dysart, and H. B. Specht. 1 987. North American distribution of Coccinella
septempunctata (Coleoptera: Coccinellidae) and its mass appearance in coastal Delaware.
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Smith, B. C. 1957. Notes on relative abundance and variation in elytral patterns of some com-
mon coccinellids in the Belleville district (Col.: Coccinellidae). Rep. Entomol. Soc. Ont. 88:
59-60.

Tedders, W. L. and P. W. Schaefer. 1994. Release and establishment of Harmonia axyridis
(Coleoptera: Coccinellidae) in the southeastern United States. Entomol. News 105: 228-243.

Wheeler, A. G., Jr. and E. R. Hoebeke. 1995. Coccinella novemnotata in northeastern North
America: Historical occurrence and current status (Coleoptera: Coccinellidae). Proc. Entomol.
Soc. Wash. 97: 701-716.

152 ENTOMOLOGICAL NEWS

SOCIETY MEETING OF OCTOBER 22, 1997

Dr. Susan P. Whitney

University of Delaware, Cooperative Extension

SUBTERRANEAN TERMITE COMMUNITY ECOLOGY

Dr. Whitney began by explaining that her research is an effort to answer basic questions
about termite ecology, sparked by the need to find new controls. Chlordane, the old standby, has
been unavailable since 1 988. Subterranean termites are the number two household insect in terms
of damage caused.

After providing a brief account of the natural history of termites, Dr. Whitney reported on
studies of three species of subterranean termites in two sites in Delaware. Reticulitermesflavipes,
the eastern subterranean termite, is found in wooded areas in northern Delaware. R. virginicus
and R. hageni are found in dry habitats in southern Delaware. During the summers (May-Sep-
tember) of 95 and 96, field sites were monitored for termite activity with pine stakes buried
approximately 20 cm in the ground. Stakes that showed feeding were replaced with "bucket
traps" plastic buckets with the bottom cut off. Into each bucket was placed a wood "sandwich"
six pieces of pine arranged to allow gaps for workers to build mud tubes. Termites were
removed from an initial bucket at the start of the field season. Individual workers were marked by
allowing them to feed on filter paper saturated with Nile blue A dye. After three days of feeding,
marked workers were returned to their bucket. After one week all wood "sandwiches" in the field
site were examined for termite presence. The data from repeated marking and recapture were
analyzed and average number of workers in colonies of each species calculated. Foraging dis-
tances were determined to be as much as thirty feet. Comparisons of the two sites, the experi-
mental farm at University of Delaware in Newark and the field station at Lewes, suggested that
colony size and relations among the three species are different in the dry pinewoods at Lewes.

Dr. Whitney described her plans to confirm these findings at other sites and to try reducing
colonies with toxic baits to test population recovery.

In entomological notes, Susan Whitney reported finding mole crickets in Delaware; Jon
Gelhaus noted AES treasurer Howard Boyd's new book on the New Jersey Pine Barrens; and
Roger Fuester reported on the low levels of gypsy moth in Delaware this year - no acreage over
30% defoliated.

W. J. Cromartie
Corresponding Secretary

NOTE RE PRIORITY OF NEW SPECIES

DESCRIBED IN JANUARY-FEBRUARY 1998

ISSUE OF ENTOMOLOGICAL NEWS

The January-February 1998 issue, Vol. 109, No. 1, of Entomological News was mailed on
December 1, 1997. Due to this early mailing date, and for purposes of priority, any new
species described in that issue must be cited with a 1997 date, even though it appeared in a
1998 issue.

H.P.B., ed.

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FOR SALE: Baltic amber pieces with insects, spiders, plants. Material for scientific work, as
well as better pieces for display and teaching. O. Holden, Junkerg. 37, S-126 53 Hegersten,
Sweden, fax: 01146-8-7268522.

VOL. 109

QL-

MAY & JUNE, 1998

US ISSN 0013-872X
NO. 3

ENTOMOLOGICAL NEWS

(Plecoptera: Perlidae), a new stonefly
species from Alabama A, D. Smith, B. P. Stark 153

Biology & hosts of Procecidochares atra (Diptera:

Tephritidae): evidence for cryptic species

T.K. Phillips, D.C. Smith 159
Survey of summer tiger beetles on Ohio River beaches in

Ohio & eastern Indiana

G. Kritsky, AJ. Savage, S. Reidel, J. Smith 165
Ticks of genus Amblyomma (Acari: Ixodidae) from white-

lipped peccaries in northeastern Bolivia, with

comments on host specificity

R.G. Robbins, W.B. Karesh, R.L.E. Painter, S. Rosenberg 172
New records of Japygoidea (Diplura) from Louisiana,

with notes on behavior MA. Muegg, C.E. Carlton 111

New records ofAllocapnia (Plecoptera: Capniidae) from

Mississippi & Louisiana, with scanning electron

micrographs Mac H. Alford 183

First U.S. record ofDyschirius sextoni (Coleoptera:

Carabidae) F.F. Purrington, J.A. Maxwell 189

Two new species of Hynesionella (Heteroptera: Gerridae)

from South Africa J.T. Polhemus 191

Reassignment of Isotoma louisiana (Collembola:

Isotomidae R.D. Waltz 195

New host record for Sphaeropthalma pensylvanica pensyl-

vanica (Hymenoptera: Mutellidae) D.G.Manley,T.P.Carithers 198
Odonata of south central Nearctic Region, including

northeastern Mexico J.C. Abbott, E.W. Stewart 201

SCIENTIFIC NOTES:
New distributions for Raptoheptagenia cruentata &

Ametropus neavei (Ephemeroptera)

R.D. Waltz, G.F. Edmunds, Jr., G. Lester 213
Cloeodes excogitatus (Ephemeroptera: Baetidae) in

northern California R.D. Waltz, Peter Ode, Jon Lee 215

First record ofAleiodes depanochora (Hymenoptera:

Braconidae) from Brazil A.M. Penteado-Dias 217

BOOK REVIEW 164

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SOCIETY 219

SOCIETY MEETING OF NOVEMBER 19, 1997 224

ANNOUNCEMENT 200

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Vol. 109, No. 3, May & June, 1998 153

NEOPERLA COOSA (PLECOPTERA: PERLIDAE),

A NEW STONEFLY SPECIES

FROM ALABAMA 1

Angela D. Smith, Bill P. Stark 2

ABSTRACT: Neoperla coosa, a new stonefly species, is described from male, female and egg
stages. The species resembles Neoperla osage from the Ozark Mountains, but the male aedeagus
is more similar to that of Neoperla clymene. The new species is known only from the Coosa and
Cahaba drainage systems of Alabama.

During a scanning electron microscopy study of chorionic variability among
Alabama and Mississippi Neoperla, several populations from the Coosa and
Cahaba systems with distinctive eggs were found. These eggs were quite un-
like those of known Neoperla from the southeastern United States (Stark and
Baumann 1978; Stark and Lentz 1988; Stark 1995) and subsequently the fe-
males were found to be associated at several localities with males misidentified
as N. clymene (Newman). Because the eggs closely resemble those of N. osage
Stark and Lentz, comparisons were made of the male and female genitalia of
these species. Results of this study suggest the Coosa-Cahaba specimens rep-
resent a previously unrecognized species of the N. clymene complex. Termi-
nology for the description follows Stark and Lentz (1988) and Stark (1995).
The holotype is deposited in the United States National Museum of Natural
History (USNM) and paratypes are deposited in the University of Alabama
(UA) or in the collection of the junior author (BPS).

Neoperla coosa, NEW SPECIES

Male.- Forewing length 9-11 mm. General color pale brown, head pale except for dark
ocellar and mesal clypeal areas. Wing membrane and veins brown. Legs brown, cerci pale. Pro-
cess of tergum 7 apically truncate, upturned in lateral aspect, armed ventrally with prominent
sensilla basiconica. Mesal sclerite of tergum 8 triangular. Hemiterga rounded apically; finger-
like process of hemiterga slender and relatively straight (Fig. 1 ). Tube of aedeagus slightly sinu-
ate, ca. four times as long as bulb width (Fig. 2); spicule patch relatively prominent along
dorsobasal to dorsomesal sides and margins of tube (Figs. 2, 10). Sac unarmed in basal third,
apical sac armature of scattered, large and small spines (Fig. 2). Apex of tube curved slightly
ventrad (Fig. 2).

Female.- Forewing length 11-13 mm. Color pattern similar to male. Posterior margin of
sternum 8 sinuate, mesal portion slightly produced to approximate level of adjacent lateral areas
(Fig. 3). Spermathecal stalk robust and irregularly armed with fine brown setae; apex of arma-
ture truncate with subapical basally directed extensions (Fig. 4).

Egg.- Length ca. 331.0 7.0 |im; equatorial width ca. 181.1 3.9 (im. Collar sessile.

Received August 22, 1997. Accepted September 29, 1997.
Biology Department, Mississippi College, Clinton, MS 39058.

ENT. NEWS 109(3) 153-158, May & June, 1998

2 ? 1998

1 54 ENTOMOLOGICAL NEWS

surrounded by two irregular rows of reticulation; collar diameter ca. 66.7 1.4 jam (Fig. 7).
Striae slender near poles, widening at equator to ca. 12.9 0.6 u.m (Fig. 5). Number of visible
striae from anterior aspect ca. 38. Most striae connect directly to posterior follicle cell impres-
sion walls (FCIs), some arise from stalks of FCIs (Fig. 8). Sulci punctate with 2-3 rows of aeropyles
(Fig. 6); equatorial width of sulci ca. 1.3 0.2 urn. Micropyles form an irregular row displaced
slightly from equator toward posterior pole; micropylar orifices simple, without raised lip. Poste-
rior pole covered with FCIs containing ca. 21.8 2.5 aeropyles. FCI walls smooth and slightly
thinner than adjacent striae (Figs. 5, 8).

'types.- Holotype Cf and 41 9 paratypes from Yellowleaf Creek, Jumbo, Chilton County,
Alabama, 5 July 1989, P. O'Neil, S. McGregor (Holotype and 1 9 paratype deposited at the
National Museum of Natural History). Additional paratypes, all from Alabama: Cherokee Co.
Spring Creek, Hwy 87, 27 June 1989, S. Harris, S. McGregor, 2 9 (UA). Chilton Co. Yellowleaf
Creek, 2.5 mi SE Mineral Springs, 6 June 1989, 4 9 (UA). Walnut Creek, N Refuge Church, 5
July 1989, P. O'Neil, S. McGregor, 1 Cf, 17 9 (BPS). Clay Co. Cheaha Creek, abv. Lake
Chinnabbee, 3 June 1978, B. Stark, K. W. Stewart, 1 Cf (BPS). DeKalb Co. Little River at Bear
Creek, 22 June 1987, 5 C?, 98 9 (BPS). Elmore Co. Fischer Creek, 3.5 mi SW Weako, 24 June
1 987, S.Harris, P. O'Neil, 2 Cf, 3 9 (UA). Jefferson Co. Cahaba River, Trussville, 24 May 1981,
S. Harris, P. O'Neil 3 9 (UA). Cahaba River, 1-59, 14 August 1984, S. Harris, P. O'Neil,
1 Cf, 1 9 (UA). Shelly Co. Camp Branch Creek, Hwy 42, 17 June 1984, S. Harris 2 Cf , 5 9
BPS).

Etymology.- The species name is based on the Coosa River.

Diagnosis.- The aedeagal tube of male N. coosa is generally similar to that
of the group of species in which an abrupt bend at tube midlength is lacking. In
the southeastern United States, N. clymene (Newman), N. coxi Stark, N. harrisi
Stark and Lentz, N. occipitalis (Pictet), and TV. steward Stark and Baumann are
included, but only N. clymene and N. steward, of this group, are known to
occur with N. coosa. In the most recently available species key (Stark and
Lentz 1988), male N. coosa are identified as N. clymene but differences in the
tube dimensions and armature permit separation of these species. The follow-
ing modification of "couplet 8" from the Stark and Lentz (1988) key is of-
fered:

8. Tube apex essentially straight along ventral margin N. occipitalis

Tube apex curved ventrad (Fig. 2) 9

9. Dorsobasal margin of tube weakly armed with spicules (Fig. 1 2); tube length ca. 6 times width

at bulb N. clymene

Dorsobasal margin of tube with prominent spicules (Fig. 10); tube length ca. 4 times width at
bulb (Fig. 9) N. coosa

Vol. 109, No. 3, May & June, 1998

155

\'-Y 1' -I ' "' '. I: ''// -I Hf*

sri'i 1 ''''''( ( i I t 'i *' '" i ' I- 'r- MB vi&
fv' ;v ' \' i '//!^yf 1^

3 ""' '

Figs. 1-4. Neoperla coosa, male and female genitalia. 1. Male terminalia, dorsal. 2. Aedeagal
tube, sac partially everted, lateral, ventral side directed to the left. 3. Female stema 7-9. 4. Vagina
and spermathecal stalk, dorsal. B - bulb; T - tube; S - sac; T7 - tergum 7; T8 - tergum 8; S8 -
sternum 8; Ss - spermathecal stalk; Sp - spicule patch; Ht - hemitergum; Htp - hemitergal
process.

156

ENTOMOLOGICAL NEWS

Figs. 5-8. SEM micrographs of Neoperla coosa eggs. 5. Egg, lateral. 6. Detail of striae. 7. Detail
of collar and anchor. 8. Detail of posterior pole with FCIs. C = collar; FCI = follicle cell impres-
sion; St = stria.

Vol. 109, No. 3, May & June, 1998

157

Figs. 9-12. SEM micrographs of Neope rla aedeagal tubes. 9. N. coosa. lateral (B = bulb; T =
tube). 1 0. N. coosa, detail of dorsolateral spicule patch. 1 1 . N. osage, detail of dorsolateral spi-
cule patch. 12. N. clymene, detail of dorsolateral spicule patch.

158 ENTOMOLOGICAL NEWS

The egg (Figs. 5-8) and female subgenital plate morphology (Fig. 3) sug-
gest N. coosa is most closely related to an Ozark Mountain species, N. osage
Stark and Lentz. However, females of these species can usually be distinguished
by examination of the spermathecal stalk lining. In N. coosa the lining re-
sembles that of N. robisoni Poulton and Stewart (Ernst et al. 1986) in display-
ing 2-3 irregular dark longitudinal folds (Fig. 4) but in N. osage, the lining is
more uniformly distributed over the apical third of the stalk (Stark and Lentz
1988). The aedeagal tube spicule armature is also similar for N. coosa and N.
osage (Figs. 10, 11) however, the tube dimensions differ for these species.
Neoperla osage has the aedeagal tube length about three times the bulb width
(Stark and Lentz 1988) whereas in N. coosa the tube is about four times the
bulb width.

Neoperla coosa is presently known from eight sites in the Coosa-Little
River basin and two sites in the upper Cahaba River basin. All sites are above
the Fall Line in the Piedmont Upland or Alabama Valley and Ridge Physi-
ographic Section (Harris et al. 1991).

ACKNOWLEDGMENTS

We thank S. C. Harris for providing specimens used in this study and R. W. Baumann and B.
C. Kondratieff for prepublication reviews. This study was supported in part by the Howard Hughes
Medical Institute, Undergraduate Biological Sciences Education Program Grant #71 195-538901 .

LITERATURE CITED

Ernst, M. R., B. C. Poulton and K. W. Stewart. 1986. Neoperla (Plecoptera: Perlidae) of the

Ozark and Ouachi.a Mountain region, and two new species of Neoperla. Ann. Entomol. Soc.

Amer. 79:645-661.
Harris, S. C., P. E. O'Neil and P. K. Lago. 1991. Caddisflies of Alabama. Geol. Surv. Ala.

142:1-442.
Stark, B. P. 1995. A new species of Neoperla (Insecta: Plecoptera: Perlidae) from Mississippi.

Proc. Biol. Soc. Wash. 108:45-49.
Stark, B. P. and R. W. Baumann. 1 978. New species of nearctic Neoperla (Plecoptera: Perlidae),

with notes on the genus. Great Basin Nat. 38:97-1 14.
Stark, B. P. and D. L. Lentz. 1988. New species of nearctic Neoperla (Plecoptera: Perlidae).

Ann. Entomol. Soc. Amer. 81:371-376.

Vol. 1 09, No. 3, May & June, 1 998 1 59

THE BIOLOGY AND HOSTS OF PROCECIDOCHARES

ATRA (DIPTERA: TEPHRITIDAE):
EVIDENCE FOR CRYPTIC SPECIES? 1

T. Keith Philips 2 , D. Courtney Smith 3

ABSTRACT: Galls of Proceci dochare s atra on Solidago in late spring contained an average of
15.3 5.1 gregarious larvae (n = 3). Previous records note only monothalamous galls for this
species. Twenty-one larvae were found in one gall whereas the previous maximum number of
larvae per gall in the genus was 13. Our spring record indicates that P. atra is bivoltine, uncom-
mon for a temperate species of fruit fly. We also note new records of Solidago canadensis and
Erigeron canadensis as hosts and a Eurytoma species (Hymenoptera: Eurytomidae) as a parasi-
toid of P. atra. The possibility that our collections represent undescribed cryptic species of
Procecidochares closely related to P. atra is discussed.

Most species of Procecidochares Hendel cause galls on composite plants.
Galls form either in the stems or in the flowers of hosts (Benjamin 1934). One
known exception is the non-galling P.flavipes Aldrich, which reproduces in
flower heads (Goeden et al. 1994). Eleven species have been described in this
genus in North America north of Mexico (Foote et al. 1993). Procecidochares
atra (Loew) is one of the largest and most commonly collected species in the
eastern United States and ranges from Florida to Nova Scotia and West to
Utah and Idaho. Only monothalamous galls (i.e., galls which contain single
larvae) in late summer are known in this species (Felt 1918, Phillips 1946,
Philips and Smith, unpublished). Published hosts include several species of
Solidago Linneaus and an undetermined species of Aster Linneaus (Wasbauer
1972) with the latter record questionable (Foote et al. 1993). Here, we report
the occurrence of galls where the larvae develop gregariously within a single
gall; our observations represent the largest number of larvae per gall reported
in the genus. We also discuss the probability of a bivoltine life cycle and note
two new hosts for, and a parasitoid of, P. atra.

METHODS

Galls were collected in Franklin County, Ohio. Two galls on an undeter-
mined Solidago species were collected on 8 May, 1 99 1 . One of these galls was
dissected. A third gall on Solidago canadensis Linneaus was collected two
years later on the 1 4 May, 1 993. A single gall growing on Erigeron canadensis
Linneaus was found in August 1 992. All larvae or pupae were allowed to com-

Department of Entomology, Museum of Biological Diversity, Ohio State University, 1315
Kinnear Road, Columbus, Ohio 43212.

1 Received July 30, 1997. Accepted October 5, 1997.

2 Department of Entomology, Museum of Biologic*
Kinnear Road, Columbus, Ohio 43212.

3 Department of Entomology, Ohio State University, 1735 Neil Avenue, Columbus, Ohio 43210.

ENT. NEWS 109(3) 159-164, May & June, 1998

1 60 ENTOMOLOGICAL NEWS

plete development to adult stage in the laboratory. Twelve galls on Solidago
canadensis were collected in early September, 1996. These galls were dis-
sected and number of puparia per gall counted. Pupae were also examined for
evidence of parasitism. The single parasitoid discovered was allowed to emerge
and then killed for identification. Voucher specimens from both plant hosts,
the parasitoid, and Solidago galls from one of the May collections and the
September collection are deposited in the Ohio State University Insect Collec-
tion and the United States National Museum. Adults which emerged from Sol-
idago are also in the authors' collections.

RESULTS

The three late spring Solidago galls were found two to four inches above
the surface of the ground at the apical meristem. The first two galls collected
in 1991 contained 14 and 21 pupae. Adults emerged from five to 1 1 days after
collection (13-19 May) with the majority (25 of 33 flies) emerging from the
14-16 May. Two flies failed to eclose. The third gall collected in 1993 con-
tained 1 1 pupae. All adults successfully eclosed (but dates of emergence were
not recorded). This third gall (in a dry state) measured 25-30 mm in length by
9 mm in width. The average number of larvae per spring Solidago gall was
15.3 5.1 (n = 3). Larvae develop gregariously in each gall, without separate
compartments for each individual.

Galls on Solidago canadensis collected in September were all mono-
thalamous. Each contained a single, empty puparium except for two of them.
One produced a parasitoid in the genus Eurytoma Illiger (Hymenoptera:
Eurytomidae) and the other held a partially eclosed, dead P. atra. All late sum-
mer Solidago galls were situated approximately midway or higher on the plant
stalk on apical meristems. Both the late spring and late summer Solidago galls
were composed of swollen tissue surrounded by leaves in a typical rosette
growth form.

A single gall collected on Erigeron canadensis in August contained only
one larva which eclosed in late August or early September. The Erigeron gall
was a swollen stem approximately two inches above the ground.

DISCUSSION

Several species of Procecidochares produce galls containing single larvae.
Examples include P. atra (Felt 1918, Phillips 1 946), P. minuta (Snow) (Novak
et al. 1967), P. stonei Blanc and Foote (Tauber and Tauber 1968) and various
species in the literature without precise identities (Silverman and Goeden 1980,
Wangberg 1 980). In other species, one to several larvae develop within a single
gall. Wangberg ( 1 980) usually found one to three larvae but recorded up to six
in his Procecidochares sp. "B" and from one to seven (recorded in table 1 ) for
Procecidochares sp. "C." Stegmaier (1968) found P. australis Aldrich galls

Vol. 109, No. 3, May & June, 1998 161

contained two to eight larvae per gall. Similarly, Phillips (1946) found this
species to have up to eight, but most often had only a single larva per gall.
Bess and Haramoto (1958) record an average of three larvae in each gall of P.
utilis Stone. The highest reported number of larvae in a single gall is 13 in P.
stonei (Green et al. 1993). But the average for this species was only 2.5 0.1
larvae per gall. By comparison, in spring galls we found up to 21 larvae of P.
atra in a single gall and an average of 15.3 5.1 (n = 3) larvae or pupae per
gall. Published biologies of P. atra are not explicit on the number of larvae per
gall or the life cycle. For example, Felt (1918) lists a small rosette gall contain-
ing a single larval cell on Solidago altissima and refers to two species, P. polita
(Loew) and P. atra (as Oedaspis polita and O. atra). He cites Stebbins (1910),
but she only mentions P. polita and not P. atra. Felt (1918) either had his own
records or may have listed both species because they are sympatric in Massa-
chusetts, where Stebbins (1910) based her study. Phillips (1946) studied two
larvae of P. atra taken from goldenrod galls, implying a single larva per gall.
Galls we collected on Solidago canadensis in late summer contained only single
puparia but, as previously mentioned, galls collected in late spring produced
an average of 15.3 5.1 (n = 3) larvae or pupae.

The number of larvae per gall can vary within a species depending upon
the host. Procecidochares stonei, when living in Virguiera laciniata Gray, pro-
duces up to 1 3 larvae per gall (Green et al. 1 993). When this same species uses
Virguiera deltoidea Gray var. parishii (Greene) the maximum number of lar-
vae per gall drops to three. For unknown reasons, the number of larvae varies
seasonally even within the same host in P. atra. The spring generation on Sol-
idago develops with large numbers of larvae per gall, whereas the fall genera-
tion, whether on Solidago or on Erigeron canadensis, occurs as a single larva
per gall.

Galls with more than one larva can have two different types of internal gall
structure. Polythalamous galls contain separate compartments for each larva
while others are without internal divisions. Species of Procecidochares are
known to form both types. Wangberg's (1980) Procecidochares sp. "C" forms
polythalamous galls. In contrast, P. stonei larvae develop gregariously within
a gall (Green et al. 1 993). Our dissections showed that P. atra develops in galls
gregariously in the spring, without separate compartments for each larva.

Most species of Procecidochares appear to have a limited number of hosts
(Foote et al. 1993). For those species with large numbers of reported hosts,
such as P. minuta (Snow), it appears as though sibling or cryptic species are
involved (Wangberg 1980, Foote et al. 1993). Previously recorded hosts for P.
atra are Solidago altissima Linneaus, S. nemoralis Ait., S. odora Ait. and one
questionable record from Aster (Wasbauer 1972, Foote et al. 1993). Our col-
lections add two additional hosts, Solidago canadensis and Erigeron canadensis.
Procecidochares anthracina (Doane) is the only species of this genus previ-

162 ENTOMOLOGICAL NEWS

ously recorded from a species of Erigeron. Like P. atra, this species has been
recorded from both Solidago and Erigeron. This is evidence that galling by
one tephritid species on both of these hosts may be common. The other no-
table aspect of our Erigeron record is that the gall was located on the stem near
the ground and not on an apical meristem as in Solidago. In light of the differ-
ences in both host plant and gall location, we initially thought the Erigeron
galler represented a different species. But we are unable to differentiate this
specimen morphologically from other specimens of P. atra.

Procecidochares atra adult emergence from galls in late spring and late
summer indicate a bivoltine life cycle, whereas most temperate species of
tephritids have only one generation per year (Bateman 1972, Christenson and
Foote 1960). There are other species of Procecidochares that are bivoltine or
even multivoltine. Procecidochares utilis, a species native to Mexico, has two
generations per year (Hoy 1960). The Procecidochares sp. of Silverman and
Goeden (1980) is bivoltine in Southern California, although it is sometimes
univoltine and conceivably even biennial if adequate rainfall, which triggers
necessary vegetative regrowth, does not occur. Huettel and Bush (1972) men-
tion both P. australis and an undescribed Procecidochares species as multi-
voltine. These two species emerge in the fall from galls in flower heads. The
adults then oviposit in small, overwintering rosette plants. Larvae and their
galls develop slowly over the winter and the spring generation emerges from
these plants. Our records of Procecidochares atra suggest a similar life cycle,
except that adults may overwinter and oviposit on the perenial Solidago as
new shoots emerge in the spring. Most temperate species of fruit flies over-
winter as diapausing pupae (Bateman 1972).

Biological records for some Procecidochares species are questionable
because of the need for systematic revision of the genus. Incomplete taxonomy
has resulted in species listed by letter designations (Wangberg 1980) or spe-
cies listed as near a described taxon (Dodson 1986, 1987). Even with P. atra,
which could be considered a well-known species, there is potential for cryptic
species. As evidence, Foote et al. (1993) mention that specimens of P. atra
from the western part of the range have only one pair of dorsocentral bristles.
The more typical pattern is two pairs, with one pair anterior to and the other
posterior to the transverse suture. Although two pairs is the more common
pattern, we found this character to be extremely variable in our specimens
reared from the late spring galls. Almost half of our specimens have three or
four setae anterior to the suture (in addition to the posterior pair) and more
rarely a single pair anterior and two pairs of setae posterior to the suture.

Regardless of the morphological variability in P. atra, it is clear that our
May records from Solidago are not for another described species. The only
Procecidochares that use Solidago as a host are P. anthracina, P. minuta, and
P. polita. Of these, only P. polita (Loew) is an eastern species, and although

Vol. 109, No. 3, May & June, 1998 163

about the same size as P. atra, both are relatively easy to differentiate (see
Foote et al. 1993). One possibility is that our May records are for a new cryptic
species temporally separated from P. atra. Our record on Erigeron may even
represent a second new species. Similar to the situation described by Huettel
and Bush (1972), the species may have differentiated in their host plant use,
but have not diverged morphologically, even though they exist sympatrically.
Genetic studies may be useful in determining if our records represent a single
bivoltine species with two hosts, two separate species with possibly different
hosts, or even three species.

ACKNOWLEDGMENTS

We foremost thank George Keeney for his collection of the spring galls and his information
on the host species and gall location. We also thank John Furlow for his assistance with plant
identification and his discussions on Solidago and Andrey Sharkov for the Hymenoptera identi-
fication. Our appreciation to two anonymous reviewers whose comments greatly improved the
manuscript.

LITERATURE CITED

Bateman, M. A. 1 972. The ecology of fruit flies. Ann. Rev. of Entomol. 17: 493-518.

Benjamin, F. H. 1 934. Descriptions of some native trypetid flies with notes on their habits. U.S.
Depart. Agric. Tech. Bull. 401. 95 pp.

Bess, H. A. and F. H. Haramoto. 1958. Biological Control of pamakani, Eupatorium adenopho-
rum, in Hawaii by a tephritid gall fly, Procecidochares utilis I. The life history of the fly and
its effectiveness in the control of the weed. Proc. X Int. Cong. Entom. Montreal 4: 543-548.

Christenson L. D. and R. H. Foote. 1960. Biology of fruit flies. Ann. Rev. Entom. 5: 171-192.

Dodson, G. 1986. Lek mating system and large male aggressive advantage in a gall-forming
tephritid (Diptera: Tephritidae). Ethology 72: 99-108.

Dodson, G. 1987. Host plant records and life history notes on New Mexico Tephritidae (Diptera).
Proc. Entomol. Soc. Wash. 89: 607-615.

Felt, E. P. 1918. Key to American insect galls. N. Y. State Mus. Bull. 200. 310 pp.

Foote, R. H., F. L. Blanc, and A. L. Norrbom. 1993. Handbook of the fruit flies (Diptera:
Tephritidae) of America north of Mexico. Cornell Univ. Press, Ithaca. 571 pp.

Goeden, R.D., D.H. Headrick, and J.A. Teerink. 1994. Life history and descriptions of imma-
ture stages of Procecidochares flavipes Aldrich (Diptera: Tephritidae) on Brickellia spp. in
southern California. Proc. Entomol. Soc. Wash. 96(2): 288-300.

Green, J. F., D. H. Headrick, and R. D. Goeden. 1993. Life history and description of imma-
ture stages of Procecidochares stonei Blanc & Foote on Viguiera spp. in southern California
(Diptera: Tephritidae). Pan-Pac. Entomol. 69: 18-32.

Huettel, M. D. and G. L. Bush. 1972. The genetics of host selection and bearing on sympatric
speciation in Procecidochares (Diptera: Tephritidae). Entomol. Exp. Appl. 15:465-480.

Novak, J. A., W. B. Stoltzfus, E. J. Allen, and B. A. Foote. 1967. New host records for North
American fruit flies. Proc. Entomol. Soc. Wash. 69: 146-148.

Phillips, V. T. 1946. The biology and identification of trypetid larvae. Amer. Entomol. Soc.
Mem. 12. 161 pp.

Silverman, J. and R. D. Goeden. 1980. Life history of a fruit fly, Procecidochares sp., on the
ragweed Ambrosia dumosa (Gray) Payne, in southern California (Diptera: Tephritidae). Pan-
Pac. Entomol. 56: 283-288.

Stebbins, F. A. 1910. Insect galls of Springfield, Massachusetts, and vicinity. Springfield Mus.
Nat. Hist. Bull. 2. 138pp.

1 64 ENTOMOLOGICAL NEWS

Stegmaier, C. E., Jr. 1968. Erigeron, a host plant genus of tephritids (Diptera). Fla. Entomol.

51:45-50.
Tauber, M. J. and C. A. Tauber. 1 968. Biology of the gall-former Procecidochares stonei on a

composite. Ann. Entomol. Soc. Amer. 61(2): 553-554.
Wangberg, J. K. 1980. Comparative biology of gall-formers in the genus Procecidochares

(Diptera: Tephritidae) on rabbitbrush in Idaho. J. Kans. Entomol. Soc. 53(2) 401-420.
Wasbauer, M. S. 1972. An annotated host catalog of the fruit flies of America north of Mexico

(Diptera: Tephritidae). Occasional Papers No. 19, Bureau of Entomology, Calif. Dept. Agric.,

Sacramento. 172 pp.

BOOK REVIEW

WORLD CATALOG OF ODONATA. VOLUME I, ZYGOPTERA; VOLUME
II, ANISOPTERA. Henrik Steinmann. 1 997. Walter de Gruyter, Berlin & New
York. 1025 plus pp. $870.00 US.

These two volumes comprise a straightforward, well-produced, and comprehensive catalog
of the worldwide odonate fauna. It attempts to list all known species, arranged according to the
taxonomic categories recognized in Davies and Tobin's earlier ( 1 984, 1 985) lists, except that the
arrangement of Gomphidae follows Carle ( 1 986). Taxa are listed alphabetically within each higher
category (i.e., genera are alphabetical within tribes, etc). Each entry includes a fairly extensive
synonomy, although not a complete bibliography of each taxon. This is especially helpful for
categories above genus, for which such information often does not come easily to hand. Unfortu-
nately, in a number of cases Steinmann cites references in Davies and Tobin as the first entry in
a species synonymy rather than citing the actual original description (e.g., - "1985 Gomphurus
ozarkensis Westfall, 1975 - loc. cit. Davies & Tobin, ..."). Taxa down to and including tribes are
diagnosed briefly, although these descriptions merely repeat those of Davies and Tobin, which
sometimes are not, in fact, adequately diagnostic. The type species is listed for each genus as
well as the location of the type (if known) and type locality for each species. Species entries also
give an indication of the geographic range, although this often is quite general or incomplete. At
the end of each volume is a complete index of all names within the corresponding suborder and
at the end of Vol. II a selected bibliography totaling about 600 entries (not all synonymic listings
appear in the collected bibliography). A certain number of errors have crept in. The type species
of Aeshna, e.g., is given as Libellula vulgatissima L. (it is actually L. grandis; L. vulgatissima is
later, correctly, cited as the type of Gomphus), and Pseudohagenius is placed as a subgenus of
Hagenius rather than of Sieboldius. Warts and all, this work is much as one would expect from a
catalog for a medium-sized order of insects and, because it contains substantially more informa-
tion than those it is intended to supersede (Davies and Tobin, 1984, 1985;Tsuda, 1 99 1 ), it repre-
sents a major improvement on them. Unfortunately, it has appeared about six years too late.

I say this because the present work seems to have been done without any reference to (or
awareness of?) the excellent catalog of Bridges, first published in 1991, with the third and final
edition appearing in 1994, shortly before the author's untimely death. Bridges' work was pri-
vately published, but it has been discovered by most serious students of Odonata and has been
widely used and cited. It thus seems astonishing that Steinmann's books should take no account
of Bridges'. Be that as it may, a comparison is certainly in order, since each work has the same
general objectives and each has its strengths and weaknesses.

Bridges' catalog organizes and lists family group, genus group, and species group names,
each alphabetically within separate sections. A fourth section lists species within genera; genera
are again listed alphabetically, but higher classification to the level of subfamily or, when pos-
sible, tribe, is indicated in compact form after each genus. Thus Steinmann's work provides a
more quickly and easily grasped overview of classification, but the process of finding entries for

(continued on page 188)

Vol. 109, No. 3, May & June, 1998 165

A SURVEY OF SUMMER TIGER BEETLES

ON OHIO RIVER BEACHES IN OHIO

AND EASTERN INDIANA 1

Gene Kritsky, 2 A J. Savage, 2 S. Reidel, 3 J. Smith 2

ABSTRACT: A survey of the riparian tiger beetles of the Ohio River Valley from eastern Ohio to
eastern Indiana was conducted during May through August of 1 995 and 1 996. It has provided the
first composite analysis of tiger beetle distribution along the river and, combined with the river's
recent history, suggested likely causes for declines in riparian tiger beetle populations. The cur-
rent distribution of C. repanda and C. cuprascens shows how water management by locks and
dams has affected tiger beetle diversity including the extirpation of C. hirticollis from the Ohio
River Valley and the reduction of C. marginipennis to one isolated population in eastern Ohio.
The survey also found 6 new county records including the easternmost record of C. cuprascens
and the first Ohio River record of C. marginipennis.

Three species of riparian tiger beetles, Cicindela hirticollis Say, C. cupra-
scens LeConte, and C. marginipennis Dejean, are currently listed in the spe-
cial interest category by the Ohio Department of Natural Resources Wildlife
listings, which means too little is known about their distribution to allow for a
true evaluation as to their abundance. These species are sensitive to habitat
destruction and in other states have declined or been extirpated (Graves and
Brzoska 1991). Previous efforts to map these species' distributions in Ohio
(Graves and Brzoska 1991) and in Indiana (Knisley et al 1987) relied prima-
rily on museum records and inland collecting efforts. To determine the current
status of these riparian species we conducted a beach survey of the Ohio River
and its tributaries from eastern Ohio to eastern Indiana.

MATERIALS AND METHODS

Because the targeted species, C. hirticollis, C. cuprascens, and C. margini-
pennis, are summer species, we concentrated our survey during May through
early August of 1995 and 1996. The scouting of potential sites was completed
by using 1 6 and 1 4 foot speedboats to provide a more accurate coverage of the
beaches, bars, and islands along the Ohio River. Once a beach was found, its
location was determined using a global positioning system and its tiger beetle
fauna was sampled using aerial nets. Voucher specimens were collected and
deposited in the Cincinnati Museum of Natural History and Science.

1 Received August 18, 1997. Accepted October 17, 1997.

2 Department of Biology, College of Mount St. Joseph, Cincinnati, OH 45233.

3 Department of Zoology, Miami University, Oxford, OH 45056.

ENT. NEWS 109(3) 165-171, May & June, 1998

166

ENTOMOLOGICAL NEWS

RESULTS

Nearly 300 miles of the Ohio River and its tributaries were surveyed. Tiger
beetles were found at 78 sites. The most common species found was C. repanda
Dejean (figure 1) which was found throughout the Ohio River Valley. C.
cuprascens was less common, occurring in 25 localities (figure 2) primarily in
western Ohio and eastern Indiana. C. marginipennis was found in two widely
separated locations (figure 3) in Hamilton and Meigs counties in Ohio. C.
hirticollis was not found anywhere along the Ohio River. Non-riparian tiger
beetles were occasionally collected. C. sexguttata Fabricius was collected on
narrow sandy beaches that bordered wooded areas and C. punctulata Oliver
was found on beaches that were adjoining farms or other disturbed habitats.
Localities and the species collected are provided as an appendix.

Indiana

West Virginia

Figure 1. C. repanda and its Ohio River distribution.

Indiana

West Virginia

Figure 2. C. cuprascens and its Ohio River distribution.

Vol. 109, No. 3, May & June, 1998

167

Figure 3. C. marginipennis and its Ohio River distribution.

DISCUSSION

This two year survey of tiger beetles along the Ohio river marked the first
time the Ohio River was so thoroughly surveyed for tiger beetles. C. repanda
was collected for the first time in Switzerland and Ohio Counties in Indiana.
C. cuprascens was collected for the first time from Switzerland and Ohio Coun-
ties in Indiana, Meigs County in Ohio, and from Pleasants County in West
Virginia. The West Virginia locality is the farthest east that C. cuprascens has
been reported (Acciavatti et al 1992). C. marginipennis was found in Meigs
County, Ohio, which was the first time the species has been reported from
along the Ohio River. The specific collection information for the new records
is presented in bold type in the appendix.

C. hirticollis had been collected in Hamilton County, Ohio in 1 9 1 1 (voucher
specimen in the Cincinnati Museum of Science and History) but was not found
during the two years of our survey. We believe it has been extirpated from
southwestern Ohio (Kritsky et al 1996).

Graves and Brzoska (1991) warned of declines in populations of C. hirti-
collis, C. cuprascens, and C. marginipennis and argued that these species should
be better protected. To better understand the specifics of the declines, we stud-
ied the history of the Ohio River from eastern Ohio to central Indiana, trying to
document changes along the river that would have impacted the abundance of
riparian tiger beetles.

The shores along the Ohio River have changed a great deal during the past
two centuries. The Navigator (Anon 1 8 1 4), a publication of river descriptions
created to help boat pilots course the Ohio River, indicated that in the early
nineteenth century, the Ohio River ran dry during the summer months. Sandy
beaches, sandbars, and willow islands were common along the Ohio River
from eastern Ohio through east central Indiana. These locations are ideal

168

ENTOMOLOGICAL NEWS

habitats for C. hirticollis, C. cuprascens, and C. repanda. Areas east of Ports-
mouth and along what is now Meigs County were apparently more rocky, which
is the preferred habitat for C. marginipennis.

During the latter part of the Nineteenth Century a series of locks and dams
were built on the Ohio River to facilitate river transportation. This construc-
tion continued into this century. As new locks were completed, older ones
were removed. The most recent locks and dams were completed during the
1990s. The impoundment of water and flood control has had a great effect
along the shores of the Ohio. The effect is most severe in eastern Ohio where
seven locks and dams are in use. The water impoundment submerged the sandy
shores and bars that were present in 1814 and replaced them with high banks.

Today, pockets of sandy beaches occur down river of a lock and dam due
to the generally lower water levels, and these beaches have dense populations
of tiger beetles. The banks five to ten miles up river from a lock and dam are
devoid of sandy beaches and thus have no tiger beetles. Figure 4 shows the
effects of the locks and dams on tiger beetle distribution. The current locations
of locks and dams are indicated by the black lines and the dots show the loca-
tions of tiger beetle inhabited beaches. Every lock and dam from eastern Indi-
ana to eastern Ohio showed the same effects; a few suitable tiger beetle locali-
ties immediately down river from the dam and a large span of up to ten miles
up river without suitable tiger beetle habitats.

The history of the locks and dams may therefore be the key to understand-
ing the recent tiger beetle history in the Ohio River Valley in southwest Ohio.
In 1916, there were seven locks and dams in western Ohio and eastern Indiana
that have been since removed, and during the time of their operation, western
Ohio and Eastern Indiana had higher river levels and fewer beaches for tiger
beetles. One victim of this circumstance was C. hirticollis. While the loss of

Figure 4. The Ohio River's distribution of the riparian tiger beetles with the locks and dams
indicated by black lines.

Vol. 109, No. 3, May & June, 1998 169

beaches would have destroyed the tiger beetles' habitat in the area, C. hirticollis
was not in a position to recover as were the other species. By comparison, C.
repanda is common along the smaller streams and creeks. If its populations
along the Ohio River were reduced or obliterated, it could have moved back to
the Ohio River from the streams after the removal of the seven locks and dams
later in this century. Similarly, C. cuprascens is common on the willow islands
found on the Ohio River. While the dams would have caused the submergence
of many beaches, the willow islands would have survived and served as refu-
gia for C. cuprascens populations to repopulate the reformed beaches after the
dams were removed.

C. hirticollis was not as fortunate. Unlike C. repanda and C. cuprascens,
this species is very sensitive to habitat alterations. In areas along Lake Erie
where C. hirticollis is still present, it is only found in areas where human activ-
ity is limited. In the late nineteenth century, the streams in southwestern Ohio
were heavily polluted, which would have likely destroyed any C. hirticollis
populations already present and blocked the dispersal of C. hirticollis upstream
as the construction of the many locks and dams in southwestern Ohio pro-
ceeded. Therefore, when the Ohio River beaches were lost to high river water,
so too was C. hirticollis. Subsequently, when the locks and dams were re-
moved, there were no C. hirticollis populations along the streams and creeks
that flow into the Ohio to repopulate the beaches of the Ohio River.

C. marginipennis presents a completely different scenario. Today, C.
marginipennis is found on cobblestone bars and beaches along the Scioto, Great
Miami, and Little Miami Rivers. The 1814 Navigator (Anonymous) indicates
that it may have had suitable Ohio River habitats east of Portsmouth and in
Meigs County. The presence of C. marginipennis in Meigs County may be a
remnant of this past habitat. Other cobble beaches along the Ohio River have
since been destroyed by the lock and dam systems.

CONCLUSIONS

The tiger beetle survey of the Ohio River Valley from eastern Ohio to
eastern Indiana yielded 6 new county records for three species including the
easternmost limit of C. cuprascens and the first Ohio River record of C.
marginipennis. The decline in the riparian tiger beetles of the Ohio River is
likely linked to the history of the locks and dams used to control flooding and
promote river traffic. Suitable tiger beetle habitats are more common immedi-
ately down river of locks and dams where river levels are lower. This suggests
that river management to protect these sandy beaches would have a positive
impact to protect and even promote tiger beetles in the Ohio River Valley.

1 70 ENTOMOLOGICAL NEWS

APPENDIX

Collection data for the Ohio River riparian tiger beetles. Data is divided by
state and presented with county, latitude in decimal degrees, longitude in decimal
degrees, species collected, and date of collection.

INDIANA: Dearborn, 39.13, 84.8, repanda, 7/2/96; Dearborn, 39.11, 84.83, repanda, 7/2/96;
Dearborn, 39. 1 , 84.84, repanda, 7/2/96; Dearborn, 39.07, 84.89, repanda, 7/2/96; Dearborn, 39.34,
84.51, punctulata, 8/5/96; Floyd, 38.28, 85.75, cuprascens, 7/29/95; Jefferson, 38.72, 85.23,
repanda, 7/3/96; Jefferson, 38.7, 85.46, repanda, cuprascens, 7/9/96; Jefferson, 38.62, 85.44,
repanda, 7/9/96; Ohio, 39.03, 84.88, repanda, 6/27/96; Ohio, 38.99, 84.84, repanda, 6/27/96;
Switzerland, 38.87, 84.79, repanda, 6/27/96; Switzerland, 38.78, 85. 0\, repanda, cuprascens,
6/28/96; Switzerland, 38.78, 84.98, cuprascens, 6/28/96; Switzerland, 38.75, 85.04, repanda,
7/3/96.

KENTUCKY: Boone, 39.09, 84.66, repanda, cuprascens, 6/9/95; Boone, 39.13, 84.72, repanda,
6/9/95; Boone, 39.13, 84.77, cuprascens, 6/27/95; Boone, 39.14, 84.74, repanda, cuprascens,
6/27/95; Boone, 39.09, 84.66, repanda, punctulata, 7/12/95; Boone, 38.9, 84.86, repanda,
6/27/96; Boone, 38.9, 84.81, repanda, 6/27/96; Boone, 38.96, 84.83, repanda, punctulata,
6/27/96; Boone, 39.1, 84.82, repanda, 7/2/96; Boone, 39.08, 84.85, repanda, cuprascens,
7/2/96; Bracken, 38.8, 84.19, repanda, cuprascens, 6/16/95; Bracken, 38.77, 84.08, sexgutatta,
6/30/95; Bracken, 38.77, 84.02, repanda, 6/30/95; Bracken, 38.77, 84.02, repanda, 6/30/95;
Bracken, 38.8, 84.19, cuprascens, 7/25/95; Campbell, 39.05, 84.42, repanda, 6/15/95; Campbell,
39.01, 84.31, repanda, 6/15/95; Campbell, 38.89, 84.24, repanda, 6/16/95; Carroll, 38.7, 85.12,
repanda, 7/3/96; Carroll, 38.69, 85.2, repanda, 7/3/96; Lewis, 38.7, 83.55, cuprascens, repanda,
7/13/95; Lewis, 38.65, 83.34, repanda, cuprascens, 7/14/95; Lewis, 38.62, 83.24, repanda,
cuprascens, II \ 4/95; Mason, 38.77, 83.9 1 , punctulata, repanda, cuprascens, 7/7/95; Mason, 38.76,
83.87, repanda, 7/7/95; Mason, 38.82, 83.94, repanda, 7/7/95; Timble, 38.6, 85.43, repanda,
cuprascens, 7/9/96; Timble, 38.55, 85.41, repanda, 7/9/96; Trimble, 38.73, 85.42, repanda,
7/9/96.

OHIO: Adams, 38.69, 83.58, repanda, cuprascens, 7/13/95; Adams, 38.65, 83.64, cuprascens,
punctulata, 7/13/95; Adams, 38.63, 83.25, repanda, punctulata, 7/14/95; Adams (Brush Cr. Is.),
38.67, 83.46, repanda, cuprascens, 7/14/95; Brown, 38.73, 83.83, repanda, 7/7/95; Brown, 38.65,
83.72, repanda, cuprascens, punctulata, 7/13/95; Clermont, 39.04, 84.35, repanda, 6/15/95;
Clermont, 38.8, 84.2, repanda, cuprascens, 7/25/95; Clermont, 39.1 1, 84.2, repanda, 8/31/96;
Hamilton, 39.08, 84.64, repanda, sexgutatta, 6/9/95; Hamilton, 39.08, 84.64, cuprascens,
7/12/95; Hamilton, 39.09, 84.64, repanda, 7/18/95; Hamilton, 39.08, 84.63, repanda, 7/18/95;
Hamilton (Great Miami), 39.27, 84.67, repanda, 7/20/95; Hamilton (Great Miami), 39.26, 84.69,
repanda, 7/20/95; Hamilton (Great Miami), 39.26, 84.69, repanda, marginipennis, 7/20/95;
Hamilton (Great Miami), 39.26, 84.69, repanda, 8/1/95; Hamilton (Whitewater), 39.13, 84.8,
repanda, 6/27/95; Hamilton (Whitewater), 39. 1 3, 84.8, repanda, II 1 8/95; Hamilton (Whitewater),
39.13, 84.8, repanda, punctulata, 7/18/95; Hamilton (Whitewater), 39.15, 84.8, repanda,
7/18/95; Hamilton (Whitewater), 39.15, 84.8, repanda, 8/1/95; Meigs, 38.97, 81.92,
marginipennis, cuprascens, 7/26/96; Meigs, 38.94, 81 .76, repanda, 7/26/96; Ross, 39.3, 82.94,
punctulata, repanda, 7/1 1/96; Scioto, 38.62, 83.22, cuprascens, 7/14/95; Warren (Little Miami),
39.37, 84.15, repanda, 7/27/95.

WEST VIRGINIA: Jackson, 38.94, 81.76 cuprascens, 8/1/96; Mason, 38.8, 82.21, repanda,
7/26/96; Pleasants, 39.34, 81.35, cuprascens, 8/1/96; Pleasants, 39.35, 81.32, cuprascens,
8/1/96; Tyler, 39.43, 81.18, repanda, 8/1/96; Wetzel, 39.6, 80.95, repanda, 8/2/96; Wood, 39.27,
81.6, repanda, 8/1/96.

Vol. 1 09, No. 3, May & June, 1 998 171

ACKNOWLEDGMENTS

We thank Brian Armitage and the Ohio Biological Survey, the Division of Wildlife of the
State of Ohio, the College of Mount St. Joseph, and the Indiana Academy of Science for support
of this project. This project was also supported with funds donated through the Do Something
Wild! state income tax checkoff from the Ohio Division of Wildlife. We also thank Robert Waltz,
Indiana Department of Natural Resouces, and two anonymous reviewers for their valuable criti-
cism of this paper.

LITERATURE CITED

Acciavatti, R. E., TJ. Allen, and C. Stuart. 1992. The West Virginia tiger beetles (Coleoptera:

Cicindelidae). Cicindela 24 (3-4): 45-78.
Anonymous. 1814. The Navigator containing directions for navigating the Monongahela,

Allegheny, Ohio and Mississippi Rivers. Pittsburgh; Cramer, Spear, and Eichbaum.
Graves, R. C. and D. W. Brzoska. 1991. The tiger beetles of Ohio (Coleoptera: Cicindelidae).

Bull. Ohio Biol. Survey (new series) 8(4): vi+42 pp.
Knisley, C.B., D. W. Brzoska, and J. R. Shrock. 1987. Distribution, checklist and key to adult

tiger beetles (Coleoptera: Cicindelidae) of Indiana. Proc. Indiana Acad. Sci. 97:279-294.
Kritsky, G., L. Horner, S. Reidel, and A. J. Savage. 1996. The status of some tiger beetles

(Coleoptera: Cicindela spp.) in southern Ohio. Ohio J. Sci. 96:29-30.

1 72 ENTOMOLOGICAL NEWS

TICKS OF THE GENUS AMBLYOMMA

(ACARI: IXODIDA: IXODIDAE)

FROM WHITE-LIPPED PECCARIES,

TAYASSU PECARI, IN NORTHEASTERN BOLIVIA,

WITH COMMENTS ON HOST SPECIFICITY 1

Richard G. Robbins, 2 William B. Karesh 3 R. Lilian E. Painter^ 5

Susan Rosenberg 3

ABSTRACT: Adults of the ixodid ticks Amblyomma cajennense, A. naponense, A. oblongo-
guttatum, and A. pecarium are reported from Bolivian populations of the white-lipped peccary,
Tayassu pecari. These are the first published records of A. naponense and A. pecarium from
Bolivia. Infestations of the three most numerous tick species on T. pecari are shown to be statis-
tically independent of host age and sex. It is suggested that in some species of Amblyomma, host
specificity may manifest itself chiefly at the preimaginal level, diminishing or disappearing in
adults.

It has often been noted that ticks of the genus Amblyomma parasitize all
classes of terrestrial vertebrates, but that among those specific to mammals,
adults are generally more common on large herbivores, while immatures in-
fest much smaller mammals (e.g., rodents) or even birds (Hoogstraal 1973,
Hoogstraal and Aeschlimann 1982, Hoogstraal 1985). Less often reported are
data on the frequency or extent of multi-species assemblages of adult Am-
blyomma on large mammal hosts (Fairchild et al. 1966, Matthysse and Colbo
1987, Walker and Olwage 1987). In February of 1996, and again in February
of 1997, one of us (WBK) traveled to the Lago Caiman research camp ( 1 3.35S,
60.54W), Noel Kempff Mercado National Park, in the northeastern corner of
the Department of Santa Cruz, Bolivia, to assess the health of three herds of
15-40 white-lipped peccaries, Tayassu pecari (Link, 1795), that had been caught
as groups in a 0.405 ha capture corral. Tayassu pecari occurs from southern
Mexico to northeastern Argentina, but because the large tracts of wilderness
on which it depends are rapidly being fragmented, this species has disappeared
or become rare in the northern and southern portions of its range and is now
listed in appendix 2 (species not necessarily threatened with extinction but
that may become so unless trade is subjected to strict regulation) of the Con-
vention on International Trade in Endangered Species of Wild Fauna and Flora

1 Received August 7, 1997. Accepted August 30, 1997.

2 Armed Forces Pest Management Board, Walter Reed Army Medical Center, Washington, DC
20307-5001.

3 Field Veterinary Program, Wildlife Conservation Society, 185th Street and Southern Boule-
vard, Bronx, NY 10460-1099.

4 Department of Psychology, University of Liverpool, United Kingdom.

5 Proyecto BOLFOR, Santa Cruz, Bolivia.

ENT. NEWS 109(3) 172-176, May & June, 1998

Vol. 109, No. 3, May & June, 1998 173

(Mayer and Wetzel 1987, Nowak 1991, Wilson and Reeder 1993). Accord-
ingly, we decided to collect all tick specimens found on our Bolivian peccaries
in order both to document the diversity of species parasitizing T. pecari at this
locale and to determine whether particular host attributes have a bearing on
parasitization. Continuing habitat loss will render such analyses impossible
within the working lifetime of contemporary investigators.

METHODS

Over a period of years, one of us (RLEP) developed the Lago Caiman
research camp and constructed the peccary capture areas. The lowland forest
of Noel Kempff Mercado National Park is broadly classified as subhumid but
comprises several forest types; one of these, which includes Lago Caiman, is
tall forest with canopy heights of 30-35 m. This part of Bolivia is character-
ized by a marked dry season in the austral winter, a mean daily temperature of
25C, and annual precipitation greater than 1500 mm (Killeen 1996).

On 25 February 1996, 13 captured peccaries were randomly targeted for
immobilization and tick collection at Lago Caiman. On 17 and 18 February of
the following year, an additional 27 peccaries were similarly selected. All pec-
caries were sedated using a combination of tiletamine hydrochloride and
zolazepam administered by projectile syringe dart. Their pelage was then care-
fully searched for ticks, and virtually complete collections were secured from
35 of the 40 animals. Following recovery from anesthesia, all peccaries were
released. Ticks were preserved in 70% ethanol and shipped to RGR for identi-
fication.

By means of contingency tests, it was possible to examine whether tick
infestations were dependent on particular attributes of individual peccaries,
such as age (adults vs. immatures) and sex. Weight was discounted as a test-
able attribute because of its dependence on numerous variables, including sex,
health and season. In all cases, the variety of contingency test used was the log
likelihood ratio or G-test, with Yates' correction for small sample sizes (Sokal
and Rohlf 1 973). Peccaries harboring only preimaginal ticks (larvae, nymphs),
which could not be identified to species, were excluded from this analysis.
Because all tests were 2x2, computed values of G were compared with a
critical value of the chi-square distribution of 3.841 (one degree of freedom) at
P = 0.05.

RESULTS AND DISCUSSION

Adults of four species of Amblyomma -A. cajennense (Fabricius, 1787)
(9C?), A. naponense (Packard, 1869) (35Cf , 27 Q ),A. oblongoguttatiim Koch,
1844 (36C\ 409 ). and A. pecarium Dunn, 1933 (359 ) - were found on 31 of
35 parasitized T. pecari at Lago Caiman (four peccaries harbored only Am-
blyomma nymphs or larvae). All are relatively common ticks that have previ-

174

ENTOMOLOGICAL NEWS

ously been reported from this host (Aragao and Fonseca 1961 , Hoffmann 1962,
Fairchild et al. 1966, Jones et al. 1972). Yet, to the best of our knowledge,
these are the first published records of A. naponense and A. pecarium from
Bolivia.

The Field Veterinary Program, Wildlife Conservation Society (formerly
New York Zoological Society), has assigned accession numbers WLP 2 through
WLP 14 and MED ARKS (Medical Archives) numbers 96-0422 through 96-
0435 to the tick collections made at Lago Caiman in 1996; those made in 1997
have received accession numbers WLP 15 through WLP 40 and MED ARKS
numbers 97-0546 through 97-0567. All collections are on long-term loan to
RGR.

Descriptive statistics for the prevalence and intensity of parasitization by
adults of each tick species appear in Table 1 . Typically, ectoparasites are con-
tagiously dispersed (overdispersed, clumped) on host populations, a condition
in which the zero class is often large (Robbins and Faulkenberry 1982). In the
case at hand, of 3 1 sampled peccaries, only 6 were infested by A. cajennense,
1 5 by A. naponense, 23 by A. oblongoguttatum, and 19 by A. pecarium. How-
ever, the range of parasitization was broad, as reflected in the disproportion-
ately large standard deviations and coefficients of variation of each tick spe-
cies. Also, sex ratios (males/females) differed dramatically among the four
species of Amblyomma: all male for A. cajennense, 1.29 for A. naponense,
0.90 for A. oblongoguttatum, and all female for A. pecarium, perhaps an indi-
cation that these species were sampled at different stages of their life cycles on
T. pecari, or that T. pecari is an incidental host for one or more of them. In this
regard, it should be noted that while males of A. cajennense and A. pecarium
differ markedly in facies, females of the two species are easily confused. There-
fore, throughout this study, no female specimen was accepted as A. pecarium
unless it met all the differential criteria of Jones et al. (1972): palpal segment II
about 2V 2 times as long as segment III; festoons ventrally rugose and relatively
poorly defined, first 4 on either side of the median festoon each with a well-
Table 1 . Descriptive statistics for adults of four species of Amblyomma parasitizing 3 1 individu-
als of T. pecari from Lago Caiman, Bolivia, 26 February 1996 and 17-18 February 1997, collec-
tor W. B. Karesh.

Tick

Range

Mean with

Standard

Coefficient of

Species

(Ticks/Peccary)

Standard Error

Deviation

Variation

A. cajennense

0-2

0.3 + 0.1

0.6

200.0

A. naponense

0-34

2.0+ 1.1

6.1

305.0

A. oblongoguttatum

0-13

2.5 + 0.6

3.1

124.0

A. pecarium

0-5

1.1 +0.3

1.4

127.3

Vol. 109, No. 3, May & June, 1998 175

developed tubercle at the posterointernal angle; and internal spur of coxa I
broad and blunt.

G-test results for the three most numerous tick species on T. pecari appear
in Table 2, where the observed statistical independence between tick infesta-
tion and host age or sex indicates that factors other than host attributes are
responsible for the spectrum of parasitization summarized in Table 1 . Because
53 of the world's approximately 104 Amblyomma species occur in the Neo-
tropics, host specificity is almost certainly one such factor. But while adults of
A. pecarium seem to be strict parasites of peccaries (Fairchild et al. 1966),
adults of A. cajennense, A. naponense and A. oblongoguttatum are known from
a variety of large- and medium-sized mammals (Jones et al. 1972). Clearly,
our results lend themselves to any number of explanations, yet we suggest that
the presence of four species of Amblyomma on but a single species of host may
be a sign that host specificity either diminishes or disappears in adults of some
amblyommines, manifesting itself instead chiefly at the preimaginal level.
However, as has often been stated (Fairchild et al. 1966, Jones et al. 1972,
Keirans 1992, Robbins et al. 1997), immature Amblyomma, especially in the
Neotropics, remain mostly unidentifiable. Until rearing or molecular genetic
studies enable us to associate the immatures found on one set of hosts with the
adults found on another, definitive explanations for ostensible instances of
host specificity will remain beyond our grasp.

Table 2. Tests of association between tick infestation and attributes of Tayassu pecari.
Attribute Tick Species Results

Age A. naponense Independent; G = 0.370; P 0.05

A. oblongoguttatum Independent; G = 0.308; P 0.05

A. pecarium Independent; G = 1 .066; P > 0.05

Sex A. naponense Independent; G = 0.0 1 6; P 0.05

A. oblongoguttatum Independent; G = 0.086; P 0.05

A. pecarium Independent; G = 0.348; P 0.05

ACKNOWLEDGMENTS

We thank the Bolivian National Secretariat for Protected Areas for permission to work in
Noel Kempff Mercado National Park, and the National Directorate for the Protection of
Biodiversity for help in acquiring the necessary permits. We are extremely grateful to Robert
Wallace, Marcela M. Uhart, Nicolas Tagua, Jose Chuvina and Walter "Tirana" Perez, whose
dedication and initiative were exemplary throughout this study. Thanks also to Damian Rumiz
and the staff of the Bolivian Sustainable Forestry Project (BOLFOR) and to F.A.N. (Fundacion
Amigos de la Naturaleza, Santa Cruz, Bolivia) for the logistical support they provided. For their
insightful comments on an earlier version of this work, we thank Thomas J. Daniels and Richard

176 ENTOMOLOGICAL NEWS

C. Falco of Fordham University's Louis Calder Center in Armonk, New York. And thanks to
Lance A. Durden and James E. Keirans, Institute of Arthropodology and Parasitology, Georgia
Southern University, Statesboro, for generously providing several important taxonomic refer-
ences and for critical assistance in distinguishing A. cajennense from A. pecarium. This study
was funded primarily through a grant from the Environmental Program of the Dutch Embassy in
Bolivia, by BOLFOR (which is financed by the U.S. Agency for International Development and
the Bolivian government), and by the Wildlife Conservation Society. The opinions and asser-
tions advanced herein are those of the authors and are not to be construed as official or reflecting
the views of the U.S. Departments of the Army or Defense.

LITERATURE CITED

Aragao, H. de B. and F. O. R. da Fonseca. 1961. Notas de ixodologia. VIII. Lista e chave para
os representantes da fauna ixodologica brasileira. Mem. Inst. Oswaldo Cruz 59: 1 15-130.

Fairchild, G. B., G. M. Kohls, and V. J. Tipton. 1966. The ticks of Panama (Acarina: Ix-
odoidea). pp. 167-219 In: R. L. Wenzel and V. J. Tipton (eds.). Ectoparasites of Panama.
Field Mus. Nat. Hist., Chicago.

Hoffmann, A. 1962. Monografia de los Ixodoidea de Mexico. I Parte. Rev. Soc. Mex. Hist. Nat.
23: 191-307.

Hoogstraal, H. 1973. Acarina (ticks), pp. 89-103 In: A. J. Gibbs (ed.). Viruses and invertebrates.
North Holland Publ. Co., Amsterdam.

Hoogstraal, H. 1985. Ticks, pp. 347-370 In: S. M. Gaafar, W. E. Howard, and R. E. Marsh
(eds.). Parasites, pests and predators. Elsevier Sci. Publ. Co., Amsterdam.

Hoogstraal, H. and A. Aeschlimann. 1 982. Tick-host specificity. Mitt. Schweiz. Entomol. Ges./
Bull. Soc. Entomol. Suisse 55: 5-32.

Jones, E. K., C. M. Clifford, J. E. Keirans, and G. M. Kohls. 1972. The ticks of Venezuela
(Acarina: Ixodoidea) with a key to the species of Amblyomma in the Western Hemisphere.
Brigham Young Univ. Sci. Bull. (Biol. Ser.) 17: 1-40.

Keirans, J. E. 1992. Systematics of the Ixodida (Argasidae, Ixodidae, Nuttalliellidae): an over-
view and some problems, pp. 1-21 In: B. Fivaz, T. Petney, and I. Horak (eds.). Tick vector
biology. Medical and veterinary aspects. Springer- Verlag, Berlin.

Killeen, T. J. 1 996. Historia natural y biodi versidad de Parque Nacional "Noel Kempff Mercado,"
Santa Cruz, Bolivia. Plan de manejo - components cientifico. Museo de Historia Natural
Noel Kempff Mercado.

Matthysse, J. G. and M. H. Colbo. 1987. The ixodid ticks of Uganda. Entomol. Soc. Am.,
College Park, MD.

Mayer, J. J. and R. M. Wetzel. 1987. Tayassu pecari. Mammalian Species (293): 1-7.

Nowak, R. M. 1991. Walker's mammals of the world. 5th ed. Johns Hopkins Univ. Press, Balti-
more.

Robbins, R. G. and G. D. Faulkenberry. 1982. A population model for fleas of the gray-tailed
vole, Microtus canicaudus Miller. Entomol. News 93: 70-74.

Robbins, R. G., W. B. Karesh, S. Rosenberg, N. Schonwalter, and C. Inthavong. 1997. Two
noteworthy collections of ticks (Acari: Ixodida: Ixodidae) from endangered carnivores in the
Lao People's Democratic Republic. Entomol. News 108: 60-62.

Sokal, R. R. and F. J. Rohlf. 1973. Introduction to biostatistics. W. H. Freeman and Co., San
Francisco.

Walker, J. B. and A. Olwage. 1987. The tick vectors ofCowdria ruminantium (Ixodoidea, Ixo-
didae, genus Amblyomma) and their distribution. Onderstepoort J. Vet. Res. 54: 353-379.

Wilson, D. E. and D. M. Reeder. 1993. Mammal species of the world. A taxonomic and geo-
graphic reference. 2nd ed. Smithson. Inst. Press, Washington, DC.

Vol. 1 09, No. 3, May & June, 1 998 1 77

NEW RECORDS OF JAPYGOIDEA

(HEXAPODA: DIPLURA) FROM LOUISIANA,

WITH NOTES ON BEHAVIOR 1

Mark A. Muegge? Christopher E. Carlton^

ABSTRACT: Previous published records of the hexapod order Diplura from Louisiana have
documented one species. We provide collection data for seven species of Japygoidea: five within
Japygidae and two within Parajapygidae. Two genera and four species of Louisiana Japygidae
are undescribed. Members of Japygidae are saprophagous and predatory in feeding behavior.
Cerci were not used during any observations of prey capture, but were employed defensively as
pincers.

Japygoids are a primitive group of blind, flightless hexapods belonging to
the class Diplura. The bionomics of this obscure group of hexapods remain
poorly known. Japygoids have been recorded primarily from mesic habitats
beneath rocks, rotting logs, leaf litter, humus, and soil. Some species occur in
xeric habitats and three species are obligate cavernicoles (Muegge, 1992; Pages,
1972, 1977).

Japygids are prone to extreme endemism (Allen, 1988; Muegge, 1992;
Muegge and Bernard, 1989; Smith, 1960). Their relatively small size, wing-
less and eyeless anatomy, and the occurrence of most species in edaphic habi-
tats contribute to this tendency toward endemism. Thus, while Diplura and
similar cryptic organisms could be useful in providing insight into biogeo-
graphic patterns, particularly in identifying areas of endemism, an almost total
lack of information about their distributions and regional diversity is a barrier
to accomplishing this goal. The information presented here is part of a con-
tinuing effort to document the distributional patterns and systematic status of
Japygoidea in North America and provide information about their biology.

Voucher specimens of taxa reported here are deposited in the Louisiana
State Arthropod Museum and the first author's collection.

BEHAVIORAL NOTES

Observations of behavior and examination of gut contents by one of us
(MAM) indicate that Japygoids are generally saprophagous and predatory,
pursuing and consuming live prey opportunistically. Examinations of the gut

1 Received August 18, 1997. Accepted September 22, 1997.

2 Texas A & M University, District 6 Extension Center, P.O. Box 1 298, Ft. Stockton, TX 79735
USA.

3 Christopher E. Carlton, Louisiana State University Agricultural Center, Department of Ento-
mology, Baton Rouge, LA 70803 USA.

ENT. NEWS 109(3) 177-182, May & June, 1998

1 78 ENTOMOLOGICAL NEWS

contents of numerous specimens of several species revealed an abundance of
arthropod body parts, primarily from springtails and mites, and a significant
amount of undigested, unrecognizable organic matter.

The japygid Metajapyx remingtoni Smith and Bolton was observed during
prey stalking and capture. The long musculated antennae were used to search
for prey. Once a potential prey item was detected, the animal slowly crept to
within striking distance, then lunged forward to capture, subdue, and consume
the prey. Individuals of M. remingtoni were observed capturing and consum-
ing entomobryid springtails. The feeding habits of Japyx sp. were studied by
Pages ( 1 95 1 ), who reported predation on an isopod, Platyarthrus hoffmanseggti
Brandt and the following gut contents: isopods, mites (mainly oribatids and
gamasids), Symphyla, japygids, Diptera larvae, adult beetles, vegetable de-
bris, and mycelia. Schaller (1968) described the use of forceps during prey
capture and depicted it in a pair of drawings, but provided no observational
data or references to support the description. Kuhnelt ( 1 976) published a simi-
lar drawing depicting the capture of a campodid dipluran using the forceps.
Finally, Conde and Pages (1991) reported that individuals of Heterojapyx sp.
were observed buried in soil with cerci exposed, waiting to capture small
arthropods that came within reach using the cerci. However, they did not state
whether they observed the cerci being used in this manner.

Thus, there are numerous reports in the literature suggesting that japygoids
use their cereal forceps during prey capture, but these reports are not sup-
ported by detailed observational data. We have observed forceps being used in
defensive behavior on several occasions, but never during prey capture. The
tenth abdominal segment and forceps are heavily sclerotized, musculated, and
quite powerful considering the size of the animal. Japygids under duress were
observed grasping and completely severing the bodies of other similar sized
arthropods using the forceps. These animals were never consumed during our
observations.

NEW RECORDS FROM LOUISIANA

(Fig. 1)
Family JAPYGIDAE

Mixojapyx tridenticulatUS (Fox). Specimens examined, 21. Avoyelles Parish, near
Hamburg, 2 July 1979, H. Lambert, habitat: in soil, 1 female. East Feliciana Parish, Idlewild
Research Station, 15 January 1989, M. A. Muegge, habitat: in moist soil at base of Quercus sp.,
mixed oak/pine forest, 1 male, 1 female. Same data, 3 June 1989, base of Pinus sp. 1 male. Same
data, 23 January 1990, 1 male. Same data, 1 March 1990, 1 female. Same data, 13 February
1 99 1 , 2 males, 4 females. Grant Parish, Kisatchie National Forest, 29 March 1 992, M. A. Muegge,
habitat: moist soil, mixed beech/magnolia forest, 1 male, 3 females. Natchitoches Parish, Kisatchie
National Forest, Red Dirt National Wildlife Management Area, 1 April 1989, M. A. Muegge,
habitat: in soil beneath rock, primarily long-leaf pine forest. Same data, 10 June 1991, M. A.
Muegge, 1 female. St. Landry Parish, Thistlewaite Wildlife Management Area, 14 January 1989,
M. A. Muegge, habitat: in moist sandy soil, pine/oak forest, 1 male, 1 female. Same data, 27

Vol. 109, No. 3, May & June, 1998 179

January 1990, 1 female. Washington Parish, near Southeast Research Station, 25 January 1990,
M. A. Muegge, habitat: about 25 cm deep in moist soil near Pinus sp. along roadside, 1 female.

Range. Gulf Coast States.

Comments. Mixojapyx tridenticulatus is the only species of this primarily
Mexican genus found in Louisiana. It has the widest distribution of any
Mixojapyx species north of Mexico, occurring in the gulf coastal region from
east Texas to Florida. The specimens examined from east of the Mississippi
River display slight, but consistent chaetotaxic differences from those west of
the Mississippi River. Upon further investigation, these two populations may
be found to represent distinct species.

Metajapyx undescribed species 1. Specimens examined. 26. East Feliciana Parish,
Idlewild Research Station, 1 March 1990, M. A. Muegge, habitat: moist soil, mixed oak/pine
forest, 2 males, 5 females. Same data, 27 February 1990, 3 males, 3 females. Same data, 1-3
March 1991, 4 males, 7 females. Same data, 13 February 1991, except habitat, under rotting log
near stream bank, 1 male, 1 female.

Range. Known only from the above locality.

Comments. Metajapyx is a widespread genus reported almost exclusively
from locations east of the Mississippi River (Reddell, 1983). Rathman et al.
(1988) reported an undescribed Metajapyx species from eastern Washington
State, but some characters needed to confirm the identification were not
described. Thus, further study will be required to determine its correct generic
placement. Fox (1941) and Smith (1960) reported M. subterraneus from
Oregon and Stoddard counties, Missouri, respectively, and these are the only
reliable records of the genus west of the Mississippi River. In Louisiana, speci-
mens of Metajapyx have only been found east of the Mississippi River.

Metajapyx undescribed species 2. Specimens examined, 17. East Feliciana Parish,
Idlewild Research Station, 3 June 1989, M. A. Muegge, habitat: in moist soil and litter at base of
Pinus sp, mixed oak/pine forest, 1 male, 16 females.

Range. Known only from the above locality.

Comments. This and the preceding species are being described in a revi-
sion of Metajapyx currently underway.

Undescribed genus and species 1. Specimens examined, 1 . Caddo Parish. Shreve-
port, 4634 Dixie Blvd., 12 December 1993, J. T. McBride and V. L. Moseley, habitat: under
rotting wood in urban backyard, 1 male.

Range. Known only from the above locality.

Comments. The discovery of this undescribed genus in an urban habitat
was unexpected. Chaetotaxy, structure of the subcoxal organs, and structure of
the cerci suggest that this individual represents a distinct and undescribed
genus.

Undescribed genus and species 2. Specimens examined, 1 . Webster Parish, Kisatchie
National Forest, 22 March 1990, M. A. Muegge, habitat: soil at base of Pinus sp., moist mixed
oak/pine forest, 1 male.

1 80 ENTOMOLOGICAL NEWS

Range. Ouachita Highlands of eastern Oklahoma and Arkansas, south to northwestern
Louisiana.

Comments. Many specimens representing several species in this uncle-
scribed genus have been collected by the authors and others from Arkansas
and Oklahoma. The individual collected in northern Louisiana may represent
the southernmost limit of the genus' range. Further study is underway to deter-
mine the number of species in the genus.

Family PARAJAPYGIDAE

Parajapyx (Parajapyx) isabellae (Grassi). Specimens examined, 62, not sorted by
sex. Acadia Parish, 15 October 1964, W. Sonnier, habitat: soil from sweet potato field. Acadia
Parish, LSU Rice Research Station near Crowley, 12 June 1994, M. A. Muegge, habitat: soil
sample from rice field. Avoyelles Parish, near Hamburg, 2 July 1979, H. Lambert, habitat: soil, 1
specimen. Bienville Parish, near Lake Bistineau, 1 August 1990, M. A. Muegge, habitat: soil,
mixed oak/pine forest, 2 specimens. East Baton Rouge Parish, 11 March 1982, P. J. Barbour,
habitat: soil/leaf litter berlesate. East Feliciana Parish, Idlewild Research Station near Clinton,
28 May 1989, M. A. Muegge, habitat: sandy soil near stream, mixed beech/magnolia forest, 6
specimens. Same data, 30 July 1989, 1 1 specimens. Same data, 9 February 1991, 12 specimens
from sandy soil near stream, 9 specimens from soil near decaying log. Same data, 30 June 1 991 ,
soil near Pinus sp. St. Landry Parish, 15 July 1979, C. E. Eastman, habitat: soil. Same, except
near Port Barre, 22 June 1 982, 2 specimens. Webster Parish, Kisatchie National Forest, 22 March
1990, M. A. Muegge, habitat: in soil near Pinus sp., mixed forest, 2 specimens.

Range. Cosmopolitan.

Comments. The only previously published references to Japygoids from
Louisiana were made by Ingram (1931), Ingram etal. (1950), and Fox (1957).
These specimens were reported only as Japyx sp. Ingram ( 1 93 1 ) and Ingram et
al. (1950) reported that specimens of Japyx sp. were commonly found in sug-
arcane fields, and suggested that they could be potential pests, causing dam-
age by feeding on the root systems of the plant. Although we have not exam-
ined any specimens collected by Ingram, we agree with Reddell (1983) that
these records represent P. isabellae. This species is widespread and common,
and is the only japygoid that is commonly collected in agricultural monocul-
tures.

Parajapyx (Grassjapyx) grassianus "maiusculella" Silvestri. Specimens ex-
amined, 12. East Feliciana Parish, Idlewild Research Station near Clinton, 9 February 1989, M.
A. Muegge, habitat: sandy soil near stream, mixed beech/magnolia forest, 2 specimens. Same
data, 30 July 1989, 1 specimen. Same data, 30 July 1991, soil at base of Pinus sp., 2 specimens.
Same data 9 February 1991, 7 specimens.

Range. Coastal Louisiana to Florida.

Comments. The taxonomic status of the P. grassianus is uncertain. The
type locality for P. grassianus is Cordoba, Veracruz, Mexico (Silvestri, 1911)
and it has been recorded only from locations in Mexico. Silvestri (1948) sub-
sequently described one form, "forma vel mutans", and two variations of that
form, "maiusculella" and "robustior", from Florida These descriptions were
based primarily on cereal dentation. Based on cereal dentation, the specimens

Vol. 109, No. 3, May & June, 1998

181

Mixojapyx tridenticulatus
. Metajapyx undescribed sp 1
Metajapyx undescribed sp 2
Undescribed genus & sp 1
Undescribed genus & sp 2
Parajapyx isabellae
Parajapyx grassianus

Figure 1. Collecting localities of Japygoidea recorded from Louisiana.

1 82 ENTOMOLOGICAL NEWS

from Louisiana represent the variant "maiusculella". Further study is neces-
sary to determine if the Florida and Louisiana specimens represent undescribed
species, or geographical variants of P. grassianus.

ACKNOWLEDGMENTS

Publication of this manuscript was supported in part by a grant to the second author from the
Louisiana Education Quality Support Fund. We thank an anonymous reviewer for calling our
attention to several pertinent references.

LITERATURE CITED

Allen, R. T. 1988. A new species of Occasjapyx from the Interior Highlands (Insecta: Diplura:

Japygidae). Proc. Ark. Acad. Sci. 42: 22-23.
Conde, B., and J. Pages. 1991. Diplura. pp. 269-271. In. CSIRO (ed.), The Insects of Australia:

A Textbook for Students and Research Workers (2nd ed.). Vol. 1. Cornell Univ. Press, Ithaca,

NY.

Fox, I. 1941. New or little known North American Japygidae. Can. Entomol. 73: 28-31.
Fox, I. 1957. The insect family Japygidae (order Thysanura) in Puerto Rico. J. Agr. Univ. Puerto

Rico 41: 35-37.

Ingrain, J. W. 1931. Soil animals attacking sugarcane. J. Econ. Entomol. 24: 866-869.
Ingram, J. W., E. K. Bynum, L. J. Charpentier, and W. E. Haley. 1950. Chemical control of

soil insects and organisms attacking sugarcane. Sugar Journal 12: 13-14, 20.
Kuhnelt, W. 1976. Soil Biology. Faber and Faber, London. 483 pp.
Muegge, M. A. 1992. New species of cavernicolous japygid (Japygidae: Diplura) from Texas.

Ann. Entomol. Soc. Amer. 85: 40-412.
Muegge, M. A., and E. C. Bernard. 1990. Two new species of Me tajapyx (Diplura: Japygidae)

from Tennessee. Proc. Entomol. Soc. Wash. 92: 793-801.
Pages, J. 195 1 . Contribution a la connaissance des Diploures. Bull. Scientifique de Bourgogne

13, Supplement 9: 1-97, 12 pis.

Pages, J. 1972. Les Japygidaes cavernicoles de la francaise. Int. J. Speleol. 4: 61-66.
Pages, J. 1977. Dicellurata genavensia III. Japygides du Sud-Est. asiatique n 1. Rev. Suisse

Zool. 84: 687-698.
Rath man, R. J., R. D. Akre, and J. F. Brunner. 1988. External morphology of a species of

Metajapyx (Diplura: Japygidae) from Washington. Pan-Pacific Entomol. 64: 185-192.
Reddell, J. R. 1 983. A checklist and bibliography of the Japygoidea (Insecta: Diplura) of North

America, Central America, and the West Indies. Texas Mem. Mus., Pearce-Sellards Ser., no.

37. 41 pp.

Schaller, F. 1968. Soil Animals. The University of Michican Press, Ann Arbor. 144 pp.
Silvestri, F. 1911. Materiali per lo sstudiodeiTisanuri. XII. Un novo genere e undici specie nove

di Japygidae dell 'America settentrionale. Boll. Lob. Zool. Gen. Agr. Protici 5: 72-87.
Silvestri, F. 1948. Intorna ad alcune anomalie di Japygidae (Insecta, Diplura). Boll. Lab. Ento.

Agr. Portici 8: 209-213.
Smith. L. M. 1960. Japygidae of North America 7. A new genus in the Provalljapyginae from

Missouri. Proc. Biol. Soc. Wash. 73: 261-266.

Vol. 109, No. 3, May & June, 1998 183

NEW RECORDS OF ALLOCAPNIA

(PLECOPTERA: CAPNIIDAE) FROM MISSISSIPPI

AND LOUISIANA, WITH ACCOMPANYING

SCANNING ELECTRON MICROGRAPHS 1

Mac H. Alford 2

ABSTRACT: Seventy-one sites were surveyed for Alloc apnia (Plecoptera: Capniidae) in south-
western Mississippi and three of the Florida parishes of Louisiana. These surveys extended over
six major river drainages. New records of A. virginiana, A. recta, and A. aurora are reported, and
scanning electron micrographs of the male genitalia are provided for each species.

Small winter stoneflies (Capniidae) constitute the largest family of Plecop-
tera in North America (Borror, Triplehorn, and Johnson 1 989). They have cap-
tivated curious minds at least since the mid-nineteenth century (Prison 1929)
because they are generally less than 10 mm in length and, unlike most other
insects, emerge as adults during the winter or early spring (Ross and Ricker
1 97 1 ). Most capniids in eastern North America belong to the genus Allocapnia
Claassen (Borror, Triplehorn, and Johnson 1989). Allocapnia is typical of the
Capniidae and is distinguished from the other genera of the family by the pres-
ence of a straight RI of the fore wing just beyond the origin of R s (Stewart and
Harper 1996) and by the presence of a dorsal process on the eighth tergite and
a double epiproct in males (Ross and Ricker 1971). Differentiation of species
is chiefly based on characteristics of the male genitalia.

METHODS

Adult Allocapnia were collected from December 1996 to February 1997
by hand from bridges and by rod and beating sheet from streamside plants.
Nymphs were collected by browsing through leaf litter in streams. Specimens
were killed and stored in 80% ethanol and were examined and identified using
a dissecting microscope. The genitalia of representative male samples were
then viewed using an AMRAY 1810D scanning electron microscope and pho-
tographed. Specimens were deposited in the entomological collection of B. P.
Stark at Mississippi College.

Forty-eight sites were surveyed in the southwestern Mississippi counties
of Amite, Franklin, Lincoln, Pike, Walthall, and Wilkinson. Twenty-three sites
were surveyed in three of the Florida parishes of Louisiana, namely, East
Feliciana, St. Helena, and West Feliciana. The Florida parishes of Louisiana

1 Received July 18, 1997. Accepted September 8, 1997.

2 Department of Biological Sciences, Mississippi College, Clinton, MS 39058-4045. Current
address: Department of Botany, Duke University, Durham, NC 27708-0338.

ENT. NEWS 109(3) 183-188, May & June, 1998

184

ENTOMOLOGICAL NEWS

are those that occur in the southeastern part of the state east of the Mississippi
River. The surveys included collection sites from the Amite River, Bogue Chitto
River, Buffalo River, and Homochitto River drainages in Mississippi and the
Bayou Sara and Thompson Creek drainages in Louisiana. Also, one small stream
in Mississippi and two small streams in Louisiana which flow directly into the
Mississippi River were surveyed (Fig. 1).

Fig. 1 . Map of the study area: southwest Mississippi and three of the Florida parishes of Louisi-
ana. Sites with Allocapnia records in the winter of 1996-1997 are indicated by a H and negative
records are indicated by a A. Numbered collection sites are further described in the text.

RESULTS AND DISCUSSION

Fig. 1 is a summary map of the sample localities. Sites with Allocapnia are
numbered as follows:

1 . Wilkinson Co., MS, Clark Cr., Clark Cr. Natural Area, A. recta.

2. West Feliciana P., LA, Kimball Cr. at Pinckneyville Rd., unresolved 9 (likely A. recta).

3. West Feliciana P., LA, Dry Cr. at LA 421, A. recta.

4. West Feliciana P., LA, Mill Cr. at LA 421, A. recta.

5. West Feliciana P., LA, Thorn Cr. at LA 421, A. recta.

6. West Feliciana P., LA, un-named tributary to Middle Fork Thompson Cr. at LA 421, unre-

solved 9 (likely A. recta).

Vol. 109, No. 3, May & June, 1998 185

7. East Feliciana P., LA, Hurricane Cr. at Thompson Cr. Rd., A. recta, A. virginiana.

8. East Feliciana P., LA, Shady Grove Branch at Thompson Cr. Rd., A. recta, A. virginiana.

9. Wilkinson Co., MS, Buffalo R. at Hiram McGraw Rd., A. recta, A. virginiana.

10. Amite and Wilkinson Co., MS, Foster Cr. at MS 33, nymph.

1 1 . Amite Co., MS, Foster Cr. at MS 33, A. aurora, A. virginiana.

12. Amite Co., MS, Brushy Cr. at Cobb Rd., A. virginiana.

13. Amite Co., MS, Brushy Cr. at New Hope Rd., A. virginiana.

14. Amite Co., MS, un-named tributary to Birdman Branch at Fox Rd., A. recta.

15. Amite Co., MS, Caston Cr. at Oxford-Meadville Rd., A. virginiana.

16. Franklin Co., MS, Middleton Cr. at USFS 100, A. virginiana.

17. Franklin Co., MS, Porter Cr. at USFS 108, A. recta, A. virginiana.

18. Franklin Co., MS, Dry Cr. at USFS 145, A. aurora, A. recta, A. virginiana.

19. Franklin Co., MS, McGehee Cr. at low-water bridge, A. virginiana.

20. Franklin Co., MS, Cane Mill Branch near Little Springs, A. aurora, A. virginiana.

21. Franklin Co., MS, Goober Cr. at Little Springs, A. virginiana.

22. Amite Co., MS, un-named tributary to West Fork Amite River, A. recta.

Of the seventy-one surveyed sites, twenty-two (31%) yielded at least one
specimen of Allocapnia. The majority of these sites (82%) occur in the
Homochitto River drainage of Mississippi and the Thompson Creek drainage
of Louisiana. No specimens of Allocapnia were collected from the Bogue Chitto
River drainage. Specimens were collected from one site in the Amite River
drainage, one site in the Buffalo River drainage, one site in the Bayou Sara
drainage, and one site from a stream which flows directly into the Mississippi
River.

Overall, 188 adults (151 Cf , 37 Q ) and 14 nymphs were collected. Of the
identifiable specimens (some dehydrated individuals were collected from spi-
der webs and some females could not be resolved), 84 (56%) belonged to
Allocapnia virginiana Prison, 62 (41%) to A. recta (Claassen) Prison, and 4
(3%) to A. aurora Ricker. Scanning electron micrographs are provided in Figs.
2-7. A. virginiana (Figs. 2-3) is characterized by a large apical segment of the
upper limb of the epiproct and by a wide but short dorsal process with a small
anterior process. A. recta (Figs. 4-5) is characterized by a long, thin apical
segment and a dorsal process with a semicircular apical ridge. A. aurora (Figs.
6-7) is characterized by a slender apical segment and a dorsal process with
widely separated lobes.

Although the data imply a paucity of Allocapnia outside of the Homochitto
River and Thompson Creek drainages, this is somewhat misleading. Two of
the three streams which flow directly into the Mississippi River probably did
not produce Allocapnia individuals because occasional inundation from the
Mississippi River hinders nymphal establishment. The Bayou Sara drainage
experiences similar effects from the Mississippi River. Clark Creek, on the
other hand, has several high waterfalls in the loess bluffs region and is pro-
tected from periodic flooding and backwater.

The Buffalo River probably has more Allocapnia than reported. Of the
four sites investigated, three fall within company timberland, where the streams
have litter composed primarily of pine straw. The Amite River and Bogue Chitto

186

ENTOMOLOGICAL NEWS

Fig. 2. Allocapnia virginiana epiproct, dorsal aspect.
Fig. 3. Allocapnia virginiana epiproct, lateral aspect.
Fig. 4. Allocapnia recta epiproct, dorsal aspect.
Fig. 5. Allocapnia recta epiproct, lateral aspect.
Fig. 6. Allocapnia aurora epiproct, dorsal aspect.

Fig. 7. Allocapnia aurora epiproct, lateral aspect.

Vol. 109, No. 3, May & June, 1998 187

River, however, are somewhat different. Whereas the previous drainages con-
sist primarily of fast-flowing, shallow, sandy streams, the Amite River and
Bogue Chitto River drainages consist of deep, slow-flowing, mixed gravel-silt
streams. The single record from the Amite River drainage indicates that Allo-
capnia are present and suggests that they are piobably limited to localized
populations where stream habitat is suitable. Further searches in the Bogue
Chitto River drainage are necessary to determine the presence or absence of
Allocapnia there.

These data indicate a wide distribution of both A. recta and A. virginiana.
A. recta appears to predominate in Louisiana while A. virginiana appears to
predominate in the hilly Homochitto drainage of Mississippi. This apparent
discrepancy, however, may be due to collection and emergence timing. B. P.
Stark (pers. comm.) has previously collected both species in central Missis-
sippi and A. virginiana in the Homochitto drainage. Ross and Ricker (1971)
report neither species in Louisiana or southwest Mississippi, but this may be
due to little or no sampling in the area. They do, however, report collections of
A. virginiana from eastern Mississippi and note the abundance of A. recta in
the Coastal Plain (Ross and Ricker 1971).

Allocapnia aurora was only found in the Homochitto drainage in a few
disjunct locations. Stark (pers. comm.) has not collected A. aurora southwest
of Tishomingo Co., MS, despite intensive collecting in central Mississippi,
and Ross and Ricker (1971) report it no farther west than southwestern Ala-
bama. This interesting anomaly of distribution could be the result of dispersal
via the Pliocene Tennessee River entering the retreating Mississippi Embayment
near southwest Mississippi during a glacial advance. Stern (1976) provides
evidence for such an event based on unionid mussel distributions and sums up
the geologic evidence. Why no A. aurora occur between southwest Missis-
sippi and the southern Appalachians or southwestern Alabama, nevertheless,
remains unclear. Regardless, because distributional information is often inte-
grated with phylogeny to construct hypothetical dispersal paths and to specu-
late about selection pressures, these new records are valuable to an accurate
evolutionary understanding of Allocapnia.

ACKNOWLEDGMENTS

Mississippi College is thanked for use of the scanning electron microscope facilities, and
Dr. Bill Stark at Mississippi College is thanked for verifying species identifications and for help-
ful comments. Earl Alford of the Mississippi Forestry Commission is thanked for helping to
select appropriate collection sites.

LITERATURE CITED

Horror, D. J., C. A. Triplehorn, and N. F. Johnson. 1989. An Introduction to the Study of
Insects, 6th ed. Harcourt Brace Jovanovich, Fort Worth.

1 88 ENTOMOLOGICAL NEWS

Prison, T. 1929. Fall and Winter Stoneflies, or Plecoptera, of Illinois. Bull. 111. Nat. Hist. Surv.,

18:345-361,399-403.
Ross, H. H. and W. E. Ricker. 1 97 1 . The Classification, Evolution, and Dispersal of the Winter

Stonefly Genus Allocapnia. 111. Biol. Monog., 45: 1-166.
Stern, E. M. 1976. The Freshwater Mussels (Unionidae) of the Lake Maurepas-Pontchartrain-

Borgne Drainage System, Louisiana and Mississippi. Unpubl. Ph.D. Thesis, LA State Univ.
Stewart, K W. and P. P. Harper. 1996. Plecoptera. pp. 217-266 In: Merritt, R. W. and K. W.

Cummins, eds. An Introduction to the Aquatic Insects of North America, 3rd ed. Kendall/

Hunt, Dubuque, IA.

BOOK REVIEW

(continued from page 1 64)

given genera and species is not so transparent (the indices ameliorate the problem to a consider-
able extent, however). It should be noted that the classification of Odonata at the level of sub-
families and tribes is currently not very well-founded and is in a state of flux, although this is
certainly no fault of either author.

In general, the Steinmann volumes use a much more traditional format, so entomologists
accustomed to using catalogs for other groups may feel more at home with them, but I have
found that, with a little initial effort, Bridges' catalog is extremely easy to use and provides a
number of advantages over the common approach. On the other hand, Steinmann certainly is
physically easier to use, as it is printed as two compact volumes, while Bridges is a rather un-
wieldy tome. Steinmann also provides a slightly more extensive synonymy, and the full refer-
ences appear with the species to which they pertain, whereas Bridges' citations are numerically
coded and the full references appear only in the bibliography. The latter is by far the more exten-
sive and up-to-date, however, with over 6500 entries, and it is accompanied by indices that allow
cross-referencing by author and journal. This in itself is an outstandingly useful reference. Bridges
provides no indication of range nor does he include diagnoses of any taxa, but the last edition
does contain an appendix with figures of wings of most genera.

The major failing of Steinmann 's World Catalog, especially compared to Bridges', how-
ever, is that it simply is not current. It has used Davies and Tobin (1984, 1985) as its starting
point, but, although it has added substantially to their work, it has not moved beyond that as it
could and should have. Despite considerable, and accelerating, work on the taxonomy of the
Odonata over the last decade, almost no species described after 1990 are included here. Also
missed were, e.g., Lohmann's (1992) revision of the Cordulegastridae, and the major reanalysis
by Carle and Louton (1994) separating a new family, the Austropetaliidae, from the now mono-
typic Neopetaliidae, with a radically new understanding of the position of the latter. Thus, de-
spite the 1997 publication date, this catalog lags considerably behind Bridges' 1994 edition.
Coupled with its astronomical price (over $850 at current exchange rates), this makes it hard to
recommend as a practical tool for the study of Odonata, despite its several useful features.

LITERATURE CITED

Bridges, C. A. 1 994. Catalogue of the Family-group, Genus-group and Species-group Names of
the Odonata of the World (Third Edition). Urbana, IL, privately published.

Carle, F. L. 1986. The classification, phylogeny and biogeography of the Gomphidae (Anisop-
tera). I. classification. Odonatologica 15: 275-326.

Carle, F. L., & J. A. Louton. 1994. The larva of Neopetalia punctata and establishment of
Austropetaliidae fam. nov. (Odonata). Proc. Ent. Soc. Wash. 96: 147-155.

(concluded on page 194)

Vol. 109, No. 3, May & June, 1998 189

FIRST UNITED STATES RECORD OF DYSCHIRIUS
SEXTONI (COLEOPTERA: CARABIDAE) 1

Foster Forbes Purrington? Judith A. Maxwell^

ABSTRACT: We report the first United States capture of the ground beetle, Dyschirius sextoni,
at Fort McCoy Military Reservation, Monroe County, Wisconsin (16 July 1996). In addition we
newly record from Wisconsin the ground beetles, Harpalus indianus, H. indigens, Selenophorus
planipennis, Cymindis ptanipennis and Helluomorphoides praeustus bicolor.

Fort McCoy Military Reservation in northern Monroe County, Wisconsin
is characterized by glacial outwash plains, with extensive unconsolidated strati-
fied surface gravel and sand. Annual total mean precipitation (1937 to 1959)
in Monroe County is 71 cm (Barndt and Langton 1984). This area is located
within a circumscribed upper Midwestern drift-less region that remained ice-
free during the Wisconsinan glaciation (Curtis 1959).

Vegetational development at Fort McCoy is dominated by scrubby oak
barrens, a Quercus velutina Lam. and Q. ellipsoidalis E.J. Hill co-climax, with
associated forbs and grasses such as Lupinus perennis L., Andropogon gerardi
Vitman, Rudbeckia hirta L. and Baptisia leucophaea Nutt. Scars from military
tank traffic persist in many sand and gravel areas, and sand borrow pits present
colonization opportunities for several open ground specialist carabid species
like tiger beetles. The carabid fauna of tallgrass prairie in the physiographically
anomalous drift-less zone was studied by Purrington and Larsen (1996) in
northeastern Iowa.

MATERIALS AND METHODS

Water-filled yellow pan traps and automatic blacklight traps (BioQuip, Inc.)
were used from May through August in 1995 and 1996 to assay the insect
fauna of the unglaciated oak barrens habitat of Fort McCoy. Ground beetles
were taken as a consequence of this broad-based synoptic survey.

Voucher specimens of all ground beetle species obtained are held in the
University of Wisconsin Department of Entomology Research Collection in
Madison.

RESULTS
Sixty-seven species of ground beetles in 15 tribes were obtained in Mon-

1 Received 9 September, 1997. Accepted 29 October, 1997.

2 Department of Entomology, The Ohio State University, 1 735 Neil Avenue, Columbus OH 432 1 0.

3 Department of Entomology, University of Wisconsin, 237 Russell Laboratories, Madison WI
53706.

ENT. NEWS 109(3) 189-190, May & June, 1998

1 90 ENTOMOLOGICAL NEWS

roe County at Fort McCoy State Natural Area in 1995 and 1996. Six species
(Table 1) are unlisted for Wisconsin and one of these, Dyschirius sextoni
Bousquet, heretofore unrecorded for the United States (Bousquet and Larochelle
1993), is a rare clivinine described only recently (Bousquet 1987) that until
now has been known only from the type locality in Belleville, Ontario, about
1000 km east of Fort McCoy. The four specimens we found were taken on 16
July 1996 by pan trap at a sand quarry in oak savanna.

The five other ground beetles we report as new for Wisconsin (Table 1 ) are
known from various adjoining states and Canada. They are species whose
known habitat preferences are reflected in the sandy scrub biotope where they
were taken, characterizing much of the Fort McCoy area.

Table 1. Ground beetles new to Wisconsin, collected in Monroe County on the Fort McCoy
Military Reservation (1995 and 1996). Dyschirius sextoni is also new to the United States.

Clivinini Trap Date

Dyschirius sextoni Bousquet 16Jul 1996

Harpalini

Harpalus indianus Csiki 24Aug 1996

Harpalus indigens Casey 20 Jul 1 995

Sele nophorus planipennis LeConte 19Jun 1996

Lebiini

Cymindis planipennis LeConte 25 Aug 1 996

Helluonini

Helluomorphoides praeustus bicolor (Harris) 13 Jun 1996

ACKNOWLEDGMENTS

We thank Robert L. Davidson, Carnegie Museum of Natural History, Pittsburgh for his gen-
erous help with ground beetle determinations. The effort of Tim Wilder (Endangered Species
Section, Environment and Natural Resources Management Division, Fort McCoy Military Res-
ervation) in monitoring traps and curating insects is greatly appreciated.

LITERATURE CITED

Barndt, W.D. and J.E. Langton. 1 984. The soil survey of Monroe County, Wisconsin. U.S.D.A.

Soil Conservation Service, Res. Div. Coll. of Agric. and Life Sci., U. Wise., Madison; 206

pp.
Bousquet, Y. 1987. The carabid fauna of Canada and Alaska: range extensions, additions and

descriptions of two new species of Dyschirius (Coleoptera: Carabidae). Coleopt. Bull.

41 (2): 11 1-135.
Bousquet, Y. and A. Larochelle. 1993. Catalogue of the Geadephaga (Coleoptera: Trachy-

pachidae, Rhysodidae, Carabidae including Cicindelini) of America north of Mexico. Mem.

Entomol. Soc. Canada, No. 167.
Curtis, J.T. 1959. The vegetation of Wisconsin: an ordination of plant communities. U. Wise.

Press, Madison; 657 pp.
Purrington, F.F. and K.J. Larsen. 1997. Records of thirteen ground beetles (Coleoptera:

Carabidae) new to Iowa. J. Iowa Acad. Sci. 104(2): 50-51.

Vol. 109, No. 3, May & June, 1998 191

TWO NEW SPECIES OF HYNESIONELLA

(HETEROPTERA: GERRIDAE)

FROM SOUTH AFRICA 1

John T. Polhemus 2

ABSTRACT: Two new species of Hynesionella, H. karatara, and H. slateri, are described from
South Africa.

The following new species of Trepobatinae are described to make the names
available for completion of a world overview and checklist of the subfamily,
and a key to the known species of Hynesionella Poisson is provided. A rede-
scription of the genus Hynesionella was given by Polhemus & Polhemus ( 1 994),
thus is not repeated here. In that work, the two species described below were
listed as "two undescribed species from South Africa."

All measurements are in millimeters.

Hynesionella differs from the other two genera of Naboandelini,
Naboandelus Distant and Calyptobates J. Polhemus & D. Polhemus, in having
the posterior margin of the mesonotum definitely carinate medially as well as
laterally, the metanotum strongly declivant, the mesonotum excavated behind
posterolateral margins in both sexes, the posterolateral mesonotal plates large,
almost vertical. Hynesionella also differs in having the male fore femur thick-
ened, often distinctly bent, sculptured basally or medially, often with large
ventral protuberances, whereas in Naboandelus the male fore femur slightly
thickened with at most a slight basal protuberance set with very short setae,
and in Calyptobates the fore femur is slender, very slightly thickened basally
and unmodified. In most species sexual size dimorphism is pronounced, the
males much smaller. The metasternal scent gland orifice (omphalium) is vesti-
gial or absent, whereas it is present in the other two genera of Naboandelini,
although very poorly developed.

Hynesionella karatara NEW SPECIES

Figures 1 - 3

Length, apterous male 2.63 (mean, N - 7; min. 2.55, max. 2.77), maximum width 1.30
(mean, N = 7; min. 1 .28, max. 1 .39). Length, apterous female 3. 1 4 (mean, N - 3; min. 3.05, max.
3.27), maximum width 1.83 (mean, N - 3; min. 1.72, max. 1.89).

General color black, sides covered with grayish pruinosity, appearing yellowish in certain
light. Small elongate spot medially on pronotum brownish yellow. U-shaped marking on base of
head extending forward along inner eye margins, brown. Median quadrate region of mesonotum
glabrous; entire body set with short pubescence.

1 Received July 28, 1997. Accepted September 5, 1997.

2 Colorado Entomological Museum, 31 15 S. York St., Englewood, CO. 80110

ENT. NEWS 109(3) 191-194, May & June, 1998

1 92 ENTOMOLOGICAL NEWS

Structural characters. Apterous male. Head set with moderate length black setae, length
0.44, width 0.89; eye width (0.22), about half the width of the interocular space (0.43). Pronotum
short, lateral margins rounded, length 0.33, width 0.86; mesonotum long, broad, sides almost
straight, posterior margin almost straight, length 0.78, width 1 .33 (Fig. 1 ); pronotum, mesonotum
set with numerous black stout moderate length setae dorsally and laterally; metanotum fused
with first two abdominal tergites, all three moderately long, weakly indicated laterally, combined
length 0.50; abdominal tergites III-VI subequal in length (0.08-0.10), VII longer (0.17). Length
of antenna! segments I-IV: 0.89; 0.55; 0.33; 0.33; segment I long, weakly fusiform (Fig. 3).

Fore femur curved basally, notched, set with a thick brush of very short setae over most of
length, forming a pad on distal half (Fig. 2); fore tibia slightly curved, broader on distal half.
Measurements of legs as follows: femur, tibia, tarsal 1 , tarsal 2 of fore leg, 1.17, 0.78, 0.06, 0.33;
of middle leg, 2.36, 3.50, 1.11, 0.78; of hind leg (tarsal 1 and 2 fused), 2.66, 1 .03, 0.44.

Lateral arms of proctiger not prominent, blunt distally, captured between broad lateral mar-
gins of tergite VIII and stemite VIII.

Apterous female. Structure, dorsal setae, and coloration mostly as in male, except larger and
more robust, with unmodified fore legs; antennal segment I distinctly more slender than in male.
Length of antennal segments I-IV: 0.72; 0.55; 0.39; 0.42. Middle femur broad on basal 2/3,
flattened.

Type material. Holotype, apterous male, South Africa, Cape Prov., Karatara Pass, CL 968,
22 Feb. 1979, J. T. Polhemus (USNM). Paratypes (all apterous), 2 males, 2 females, same data as
holotype (JTPC); 4 males, 2 females, South Africa, Cape Prov., 15 mi. (24 km) NW of Knysna,
Phantom Pass Rd., 10 Feb. 1968, R. T. Schuh, J. & S. Slater, M. Sweet (JTPC, AMNH).

Etymology. The name karatara, a noun in apposition, refers to the type
locality, Karatara Pass.

Comparative notes. The lack of any gray pruinose markings on the
mesonotal dorsum separates karatara from the four other known species of
Hynesionella. The sculpturing of the fore femur of karatara (Fig. 2) also is
diagnostic. Hynesionella karatara sp. n. is closest to H. capensis (Poisson), as
both of these species have the median part of the mesonotum shining black; in
capensis, however, the median black area is narrow, flanked by gray pruinose,
whereas in karatara it is broad, without gray pruinose. In addition, capensis is
unique in having the posterior margin of the female mesonotum curved and
extended posteriorly, instead of straight as in all other species of the genus.

Hynesionella slateri, NEW SPECIES

Figures 4-5

Length, apterous male 2.42 (mean, N = 3; min. 2.39, max. 2.44), maximum width 1.28
(mean, N = 3; min. 1 .28, max. 1 .28). Length, apterous female 2.89 (mean, N = 2; min. 2.89, max.
2.89), maximum width 1.58 (mean, N = 2; min. 1.55, max. 1.61).

General color gray, sides covered with grayish pruinosity, appearing yellowish in certain
light. Broad depressed quadrate areas on pronotum and mesonotum densely clothed with decum-
bent, closely appressed whitish gray coarse pubescence underlain with whitish integument, at
least on pronotum. Head orange yellow to orange brown, with a dark median wedge on anterior
1/2 to 2/3. Entire body set with short gray pubescence, longer on abdominal tergites. Fore femur
orange except extreme base and extreme tip.

Vol. 109, No. 3, May & June, 1998

193

Structural characters. Apterous male. Head with moderate length black setae, length 0.33,
width 0.83; eye width (0.22), about half the width of interocular space (0.39). Pronotum short,
lateral margins rounded, length 0.28, width 0.78; mesonotum long, broad, sides slightly curved,
length 0.72, width 1 .27; pronotum, mesonotum with numerous black stout moderate length setae
laterally and on pleura; metanotum fused with first two abdominal tergites, all three moderately
long, weakly indicated laterally, combined length 0.39; abdominal tergites III-VI subequal in
length (0.06-0.10), VII longer (0.14). Length of antennal segments I-IV: 0.68; 0.44; 0.33; 0.33;
segment I long, very weakly fusiform (Fig. 4).

Fore femur stout, short, curved basally, set with a thick brush of very short setae over most
of length, forming a pad on distal half (fig. 5); fore tibia slightly curved, broader on distal half.
Measurements of legs as follows: femur, tibia, tarsal 1 , tarsal 2 of fore leg, 0.89, 0.67, 0.06, 0.33;
of middle leg, 2.77, 3.11,1.17, 0.72; of hind leg (tarsal 1 and 2 fused), 2.39, 0.94, 0.42.

Lateral arms of proctiger not prominent, tapered distally, captured between broad lateral
margins of tergite VIII and stemite VIII.

Apterous female. Structure, dorsal setae, and coloration mostly as in male, except larger and
more robust, median depressed area of mesontotum not clearly differentiated in color, dorsal
pubescence shorter; fore legs unmodified; antennal segment I about as thick as in male. Length
of antennal segments I-IV: 0.67; 0.39; 0.33; 0.33. Middle femur not broadened nor flattened.

Type material. Holotype, apterous male, South Africa, Cape Prov., 18 mi. (28 km) NW of
Kimberley, 18 Jan. 1968, R. T. Schuh, J. & S. Slater, M. Sweet (AMNH). Paratypes (all apter-
ous), 2 males, 2 females, same data as holotype (JTPC, AMNH).

Etymology. The name slateri honors James A. Slater in recognition of his
outstanding and voluminous contributions to the study of Heteroptera.

Comparative notes. Hynesionella slateri sp. n. is closest to H. aethiopica
Hoberlandt, but differs in having the fore femur orange with a less pronounced
basal notch.

Figures 1 - 3. Hynesionella karatara sp. n. 1 . Female mesonotum, dorsal view. 2. Male fore leg.
3. Antenna.

Figures 4 - 5. Hynesionella slateri sp. n. 4. Antenna. 5. Male fore leg. Scale bar = 1 mm.

194 ENTOMOLOGICAL NEWS

Key to the known species of Hynesionella

1 . Mesonotal dorsum without gray pruinose markings karatara sp. n.

Mesonotal dorsum with gray pruinose markings , 2

2. Fore femur with a prominent basal tubercle in both male and female . . . cobbeni Linnavuori
Male fore femur modified, but without a prominent basal tubercle; female fore femur

unmodified 3

3. Fore femur mostly orange in both male and female slateri sp. n.

Fore femur mostly dark colored, not orange 4

4. Male fore femur not excavated basally, not abruptly widened near middle; female
mesonotum with posterior margin curved posteriorly capensis (Poisson)

Male fore femur excavated basally, abruptly widened near middle; female mesonotum

with posterior margin almost straight aethiopica Poisson

ACKNOWLEDGMENTS

I am indebted to R. T. Schuh and D. A. Polhemus for their constructive reviews.

The invaluable assistance of Brian Stuckenberg and Dennis Brothers, Pietermaritzburg, South
Africa, is gratefully acknowledged. Both aided with guidance and logistics, and without their
help the field survey would not have been nearly as successful.

J. A. Slater and R. T. Schuh kindly permitted me to study specimens under their care; the
specimens, originally at the University of Connecticut, Storrs, have since been transferred to the
American Museum of Natural History, New York.

This research was sponsored by grant 1 806-77 from the National Geographic Society, Wash-
ington, D. C., and by grant DEB-9528025 from the National Science Foundation, Washington,
D. C. I thank these organizations for their continued support of research into the systematics and
zoogeography of aquatic Heteroptera. This research was carried out as a faculty affiliate of the
Entomology Department, Colorado State University, Fort Collins.

LITERATURE CITED

Polhemus, J. T. and D. A. Polhemus. 1994. The Trepobatinae (Gerridae) of New Guinea and
surrounding regions, with a review of the world fauna. Part 2. Tribe Naboandelini. Entomol.
Scand. 25: 333-359.

BOOK REVIEW

(concluded from page 188)

Davies, D. A. L., & P. Tobin. 1984. The Dragonflies of the World: A Systematic List of the

Extant Species of Odonata, Vol. 1 , Zygoptera, Anisozygoptera. Utrecht, S.I.O. Rapid Comm.

(Suppl.)3. 127pp.
Davies, D. A. L., & P. Tobin. 1985. The Dragonflies of the World: A Systematic List of the

Extant Species of Odonata, Vol. 2, Anisoptera. Utrecht, S.I.O. Rapid Comm. (Suppl.) 5.

151 pp.
I ohmunn, H. 1992. Revision der Cordulegastridae. 1. Entwurf einer neuen Klassifizierung der

Familie (Odonata: Anisoptera). Opusc. Zool. Flumin. 96: 1-18.
Tsuda, S. 1991 . A Distributional List of World Odonata. Osaka, privately published. 362 pp.

- Michael L. May
Department of Entomology
Rutgers University, NJ

Vol. 109, No. 3, May & June, 1998 195

REASSIGNMENT OF ISOTOMA LOUISIANA
(COLLEMBOLA: ISOTOMIDAE) 1

R.D. Waltz 2

ABSTRACT: The holotype specimen of Isotoma louisiana was examined and found to be as-
signable to the genus Isotomurus. The species is distinct among all Nearctic species based on the
tuberculate denies, mucro with a unique ventral tooth, and a unique chaetotaxy. Comparisons
and contrasts with other selected isotomid genera are made justifying its inclusion in Isotomurus.

The cosmopolitan hydrophilic genus Isotomurus Borner is readily distin-
guished among other Isotomidae by the combination of a quadridentate mu-
cro, presence of long, specialized sensory setae known as bothriotricha, greater
than ten ventral manubrial setae, and a characteristic maxillary outer lobe. The
genus in North America is comprised of eight nominal species (Christiansen
and Bellinger 1980), including this present species, and several species to be
described in forthcoming work. In preparation for a review of Nearctic Iso-
tomurus, I reviewed the holotype of Isotoma louisiana Scott and found it to be
assignable to Isotomurus. The species is herein redescribed and newly com-
bined as Isotomurus louisiana (Scott), n.comb. Some illustrations of the spe-
cies were provided by Scott (1962); further comparative illustrations will be
presented in the forthcoming revision of the Collembola of North America,
north of the Rio Grande, by Christiansen and Bellinger (in press) and a review
of Nearctic Isotomurus species (Waltz, MS). This species is unique among all
Nearctic species based on its possession of tuberculate dentes and a strongly
excavate ventrobasal projection ("ventral tooth" ) of the mucro.

Isotomurus louisiana (Scott), NEW COMBINATION

Description: Yellow or green body color in most specimens, with or without strongly contrast-
ing color patterns. Color pattern, when present, with medial and lateral longitudinal stripes and
with or without banded posterior abdominal terga. Prominently patterned individuals may also
possess a distinct ventral stripe medially on the ventral abdominal segments extending to Th II.
The holotype specimen possesses purple stripes, posterior abdominal bands and a ventral
purple stripe medially on a yellow body. Specimens studied from Arizona are light green and
without contrasting pattern. Specimens studied from Kansas are very similar to the type material
but lack the ventral purple stripe.

Head: PAO less than or subequal to nearest eye. Eyes G and H smaller than remaining eyes.
Outer lobe of maxilla with palp bifurcate and with four sublobal setae.

Thorax: Unguiculus without inner tooth. Unguis elongate, without teeth. Tibiotarsi of metatho-
racic legs without long, outstanding, exterior setae.

1 Received October 2, 1997. Accepted October 15, 1997.

2 IDNR, Division of Entomology and Plant Pathology, 402 West Washington, Room W-290,
Indianapolis, IN 46204.

ENT. NEWS 109(3) 195-197, May & June, 1998

196 ENTOMOLOGICAL NEWS

Abdomen: ABD. V-VI without coarse ciliate macrochaetae; common body setae numerous and
non-ciliate; bothriotricha of Abd. II-Abd. IV as 3+3+1 pairs; Abd. V S setae (terminology after
Deharveng and Lek 1993): accp I is absent, accp 2, accp 3, accp 4, and accp 5 present, as 1 and
as 2 are present and a third seta here designated as as X is present anterior to as 1 and 2; ventral
tube with 12-22 lateral distal setae; tenaculum quadridentate (not as illustrated by Scott 1962:
Fig. 2) with 12-25 setae; denies tuberculate; mucro with mucronal basal seta and lamella, ventral
mucronal tooth present, i.e., with excavate ventromucronal base.

Known distribution: Arizona, Kansas, Louisiana.

Material examined: (Academy of Natural Sciences, Philadelphia) Holotype (#107), on slide,
"taken on water, shore of Lake Pontchartrain, approximately 1 5 feet above mean sea level, Norco,
St. Charles Parish, Louisiana, 29-iii-1960, J.H. Eslinger." Label affixed ventrally: Isotomurus
louisiana (Scott) n. comb. Del: R.D. Waltz 5/97. Additional material examined (Presently in the
collection of K.A. Christiansen): AZ: NE of Parker, puddle at edge of Colorado River, 28-x-
1986, P. Bellinger (7051). AZ: Pima Co., Santa Rita Mtns, Florida Saddle, SE Tuscon, 7-iii-
1989, Olson (7193). KS: Leavenworth Co., 1 mile N Eudora, Kansas River, 8-ix-1988, (52-1-
7,12), D.S. Hammer and L.C. Ferrington. KS: Douglas Co., (7072).

Remarks: This species, first described by Scott (1962), was deposited at the
Collection of the Academy of Natural Sciences of Philadelphia (Scott 1962;
Roback 1981). Bellinger (1985) reviewed the holotype specimen and reported
the species as similar to "Agrenia (tuberculate denies) and Isotomurus
(abdominal bothriotricha)" but commented that it did not fit into any known
genus. The location of the remainder of the paratype material cited by Scott
(1962) is unknown.

This review of this "striking species" (Bellinger 1985) has resulted in reas-
signment of this species within Isotomurus rather than Agrenia Borner or an-
other genus due to the following symmorphic characters shared with Isotomurus
species: 1 ) maxillary outer lobe possesses a bifurcate palp and four sublobal
hairs (versus bifurcate palp and no sublobal hairs in Agrenia) (see Fjellberg'
1984); 2) denies lack the distal elongate setae found in all Agrenia species (see
Fjellberg 1986, 1988); 3) presence of abdominal bothriotricha (found also in
Hydroisotoma Stach, Archisotoma Linnaniemi, and other Isotomidae but not
reported in Agrenia); 4) the characteristic Isotomurus quadridentate mucro
(quite distinct from the mucro of Agrenia, Archisotoma, and Hydroisotoma);
5) claws that lack the distinct tunica of the ungues (present only in Agrenia);
and 6) a manubrium with many ventral setae (few only in Archisotoma). That
/. louisiana belongs in Isotomurus is strongly indicated by the above charac-
ters which are uniquely symmorphic with Isotomurus and not shared in com-
bination with other known genera characterized by the possession of abdomi-
nal bothriotricha (especially Archisotoma and Hydroisotoma) or with tubercu-
late denies (Agrenia). As noted by Bellinger (op cit) Isotomurus louisiana dif-
fers from other previously described species in the genus by the luberculale
denies (mosl Isotomurus species bear crenulale denies; some bear only par-
lially luberculate denies).

Vol. 109, No. 3, May & June, 1998 197

The combination of tuberculate denies, the excavate ventral base of the
mucro, and the Abd. V chaetotaxy is unique among European (see Deharveng
and Lek 1993) and Nearctic Isotomurus species and provides the basis for
diagnosis of this species.

Partially tuberculate dentes have been reported in at least one Palearctic
species, /. ciliatus Stach. Several new species are pending description in the
Nearctic including species with fully tuberculate dentes and partially tubercu-
late dentes.

Differences in dental structure have been found to be useful historically in
justifying generic or subgeneric status (e.g., subgenera ofProisotoma Borner).
For the present, elevation of this species and putatively related species bearing
tuberculate or partially tuberculate dentes to a subgenus grouping appears to
be clearly countered by the existence of intermediate dental types, and an ab-
sence of clearly autapomorphic chaetotactic states or other morphology rela-
tive to the type species of the genus, /. palustris (Muller).

ACKNOWLEDGMENTS

I thank D. Azuma, Academy of Natural Sciences, Philadelphia, for loan of the Holotype
slide of/. Louisiana. I thank P. Bellinger and K.A. Christiansen for loan of Nearctic collections
of this species and preliminary data and Arwin Provonsha, Purdue University, for providing
helpful comments on an early draft of this manuscript.

LITERATURE CITED

Bellinger, P.P. 1985. The identity of H.G. Scott's Collembola in the Academy of Natural Sci-
ences, Philadelphia, PA. Entomol. News 96: 78-82.

Christiansen, K.A. and P. F. Bellinger. 1980. The Collembola of North America, north of the
Rio Grande. Part 2. Families Onychiuridae and Isotomidae. Grinnell College, Grinnell, Iowa,
pp. 387-784.

Deharveng, L. and S. Lek. 1993. Remarques sur la morphologic et la taxonomie du genre
Isotomurus Borner, 1903 et description de deux especes nouvelles de France (Collembola:
Isotomidae). Ann. Entomol. Soc. France (N.S.) 29: 245-259.

Fjellberg, A. 1984. The maxillary outer lobe, an important systematic tool in Isotomidae
(Collembola). Annls. Soc. royale Zool. Belg. 114: 83-88.

Fjellberg, A. 1986. Revision of the genus Agrenia Borner, 1906 (Collembola, Isotomidae).
Entomol. Scand. 17: 93-106.

Fjellberg, A. \98S.Agrenia lamellosa, a new species of Collembola (Isotomidae) from Pennsyl-
vania. J. N. Y. Entomol. Soc. 96: 110-112.

Roback, S.S. 1981. Collembola described by Dr. H.G. Scott in the collections of the Academy of
Natural Sciences of Philadelphia. Entomol. News 92: 209-210.

Scott, H.G. 1962. Collembola from Louisiana. Entomol. News 73: 27-28.

198 ENTOMOLOGICAL NEWS

A NEW HOST RECORD FOR SPHAEROPTHALMA

PENSYLVANICA PENSYLVANICA
(HYMENOPTERA: MUTILLIDAE) 1 ' 2

Donald G. Manley, 3 T. Paige Carithers 4

ABSTRACT: Wasps belonging to the family Mutillidae are known to be parasitic on the resting
stages of other insects, including other Hymenoptera. However, host records are known for less
than five percent of the described species. The mutillid Sphae ropthalma pensylvanica pensylvanica
has been reported emerging from at least two different species and genera of sphecid hosts
(Hymenoptera: Sphecidae). Here we report the emergence of S. pensylvanica from yet another
species and genus of sphecid host, that being Isodontia mexicana. Details are described.

Sphaeropthalma (Sphaeropthalma) pensylvanica pensylvanica (Hymenop-
tera: Mutillidae) was first described as Mutilla pensylvanica by Lepeletier
(1 845), based on a male collected in Philadelphia. It is found from North Caro-
lina to central Florida, Louisiana, Missouri, Kansas and Texas. A second sub-
species, 5. pensylvanica scaeva, was described as M. scaeva by Blake (1871).
The female was described in the same publication asM. balteola (Blake 1871).
It is found from North Carolina to Massachusetts, Ohio, Illinois, Missouri,
Kansas and Texas. A third subspecies, Sphae rophthalma (!) pennsylvanica (!)
floridensis, was described by Schuster (1944). It is found only in southern
Florida. The females of all three subspecies are indistinguishable.

Members of the family Mutillidae are known to be parasitic on the resting
stages (usually the prepupa or pupa) of other insects, including aculeate Hy-
menoptera. Of more than 4000 described species of Mutillidae, host records
exist for less than five percent (Brothers 1972). Sphaeropthalma pensylvanica
is one of the species for which at least some hosts are known.

Rau (1922) reported a male specimen of S. pensylvanica scaeva emerging
from a cocoon of the mud dauber Sceliphron caementarium (Drury) (Hy-
menoptera: Sphecidae) in Missouri. He later (Rau 1928) reported males of 5.
pensylvanica scaeva having been reared from the cocoons of both Trypoxylon
(Trypargilum) politum Say (Hymenoptera: Sphecidae) and from Sceliphron
nests.

In 1984, a male mutillid was sent to Manley for identification. The mutillid

1 Received September 29, 1997. Accepted October 20, 1997.

2 Technical contribution no. 4344 of the South Carolina Agricultural Experiment Station, Clem-
son University.

3 Department of Entomology, Clemson University, Pee Dee Research and Education Center,
2200 Pocket Rd., Florence, SC 29506-9706.

4 Department of Zoology and Wildlife Science, 101 Cary Hall, Auburn University, Auburn, AL
36849.

ENT NEWS 109(3) 198-200, May & June, 1998

Vol. 109, No. 3, May & June, 1998 199

was determined to be S. pensylvanica pensylvanica. It had emerged from the
cocoon of a mud dauber (the host cocoon was not sent for determination) on
30 May 1984. The cocoon was collected by J. R. Brushwein in Clemson, South
Carolina in March of 1984.

Here, we report the emergence of a male S. pensylvanica pensylvanica
from a diapaused cocoon of another sphecid wasp, Isodontia mexicana
(Saussure). This North American wasp ranges throughout the United States
east of the Rocky Mountains, south into Mexico and Central America, and has
also become established in Hawaii and France (Bohart & Menke 1 963). Across
this wide range, this species has been observed to nest in such above-ground
cavities as hollow plant stems and twigs, rolled leaves, abandoned bee bur-
rows in logs, artificial trap-nests, and the tubular leaves of pitcher plants (Sar-
racenia) (Bequaert 1930). Isodontia mexicana is one of the "grass carrier (or
carrying) wasps" because the females are often seen carrying blades of grass
in their mandibles (Bohart & Menke 1976). The females will then proceed to
use these plant fibers to construct their nests.

Nesting by /. mexicana in the leaves of pitcher plants has been well docu-
mented (under synonym Isodontia philadelphica St. Farg.) (Hubbard 1 896,
Jones 1904, Fish 1976, Rymal & Folkerts 1982). Over 400 Isodontia nests in
pitcher leaves were reared for a study on the nesting biology of this wasp
(Carithers 1998). The male mutillid reported here was reared from an
/. mexicana nest built in the tubular leaf of the pitcher plant Sarraceniaflava.
The nest was collected on 9 November 1996 in the Apalachicola National For-
est in Liberty County, Florida. One intact Isodontia cocoon was contained in
the one-celled nest. Upon holding the cocoon up to a light source, it appeared
to contain a flacid Isodontia pre-pupa. A mutillid male emerged from the
Isodontia cocoon in an environmental chamber on 13 March 1997. Within the
Isodontia cocoon, the mutillid had spun its own cocoon. This is believed to be
the first report of a mutillid parasite from /. mexicana, although Bohart and
Menke (1976) reported that Sphaeropthalma has been bred from the nest of/.
elegans (F. Smith).

LITERATURE CITED

Bequaert, J. 1930. Nesting habits of Isodontia, a subgenus of Chlorion (Hymenoptera). Bull.
Brooklyn Entomol. Soc. 25: 122-3.

Blake, C. A. 1 87 1 . Synopsis of the Mutillidae of North America. Trans. Amer. Entomol. Soc. 3:
217-265.

Bohart, R. M. and A. S. Menke. 1963. A reclassification of the Sphecinae with a revision of the
nearctic species of the tribes Sceliphronini and Sphecini. Univ. Calif. Pub. Entomol. 30: 90-
182.

Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps of the World. Univ. Calif. Press, Berke-
ley, Calif. 695 pp.

Brothers, D. J. 1972. Biology and immature stages of Pseudomethocaf. frigida, with notes on
other species (Hymenoptera: Mutillidae). Univ. Kans. Sci. Bull. 50: 1-38.

200 ENTOMOLOGICAL NEWS

Carithers, T. P. 1 998. Nesting biology of the grass-carrier wasp Isodontia mexicana (Sphecidae)

in pitcher plants (Sarracenia). M. S. thesis, Auburn University, Auburn, AL.
Fish, D. 1976. Insect-plant relationships of the insectivorous pitcher plant Sarracenia minor.

Fla. Entomol. 59: 199-203.
Hubbard, H. G. 1 896. Some insects which brave the dangers of the pitcher plant. Proc. Entomol.

Soc. Wash. 3: 314-318.

Jones, F. M. 1904. Pitcher-plant insects. Entomol. News 15: 14-19.
Lepeletier, A. L. M. 1845. Histoire Naturelle des Insectes. Hymenopteres. Paris. 1: 14; 3: 516-

517,588-646.
Rau, P. 1 922. Ecological and behavior notes on Missouri insects. Trans. Acad. Sci. St. Louis 24:

3-8.
Rau, P. 1928. Field studies in the behavior of the non-social wasps. Trans. Acad. Sci. St. Louis

25: 325-489.
Rymal, D. E., and G. W. Folkerts. 1982. Insects associated with pitcher plants (Sarracenia:

Sarraceniaceae), and their relationship to pitcher plant conservation: a review. J. Ala. Acad.

Sci. 53: 131-151.
Schuster, R. M. 1944. Notes and records of the Eastern representatives of the Photopsidine

genera of Mutillidae with descriptions of new forms. Bull. Brooklyn Entomol. Soc. 39: 139-

155.

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Vol. 109, No. 3, May & June, 1998 201

ODONATA OF THE SOUTH CENTRAL NEARCTIC
REGION, INCLUDING NORTHEASTERN MEXICO 1

John C. Abbott, Kenneth W. Stewart 2

ABSTRACT: There has not been a concerted effort to document the extent of biodiversity, distri-
bution and geographic affinities of the Odonata of the south central United States and northeast-
ern Mexico. The area is an important boundary for some species representing eastern Nearctic
and subtropical faunas, and a mixing zone or dispersal corridor for other species. Since 1993, we
have done extensive collecting of all life stages in the seven biotic provinces of Texas, and com-
piled published and extensive unpublished records from the portions of these provinces of U.S.
and Mexican states that join Texas. Here we list 228 species for this region (196 in Texas), indi-
cate their distributions by biotic province, and discuss the regional biogeography and importance
of rare species.

Current emphasis on inventorying aquatic insects of the neotropics
Paulson 1982; (Gonzalez and Novelo 1991; Novelo and Gonzalez 1991;
Quintero and Aiello 1992; Baumann and Kondratieff 1996; Gonzalez and
Novelo 1996; McCafferty & Lugo-Ortiz 1996) and realization of the great risk
to aquatic invertebrate biodiversity in temperate regions (Franklin 1988;
Haffernik 1989, 1992), prompted us to investigate the status of the odonate
fauna of the biotic provinces of Texas and adjoining states of the United States
and northeastern Mexico that border the Rio Grande River. The odonate fauna
of this region is poorly known except from scattered publication records, un-
published reports of collectors, the general documentation and discussion of
Mexico's fauna by Gonzalez and Novelo (1996) and Johnson's (1972) work
on Texas Zygoptera. There has never been a concerted effort to document the
biodiversity and geographic affinities of the Odonata of this area. Over half of
the species of Odonata still unknown as larvae are from this region (McCafferty
etal. 1990).

The earliest major documentation of Odonata in Texas was by Hagen ( 1 861 ).
A number of species accounts from the state then appeared in Calvert (1901-
1 908) and Muttkowski (1910). Williamson (1914) listed numerous records for
Texas and Oklahoma. Several localized studies within Texas have supplemented
these works: Tinkham (1934) and Gloyd (1958) from West Texas and Tucker
(1908), Ferguson (1940, 1942), Harwell (1951), Donnelly (1978), Williams
(1982), and Laswell and Mitchell (1997) from north central and eastern parts
of the state. Abbott (1996) reported the following new species records for Texas:
Aeshnapsilus Calvert, Gomphus exilis (Selys), Somatochlora georgiana Walker,
Dythemis maya Calvert, Micrathyria didyma (Selys), Sympetrum illotum

1 Received July 18, 1997, Accepted September 20, 1997.

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

ENT. NEWS 109(3) 201-212 , May & June, 1998

202 ENTOMOLOGICAL NEWS

(Hagen), Tauriphilia azteca Calvert, and Tramea insularis Hagen. Johnson's
(1972) treatise on Texas Zygoptera remains the only comprehensive faunal
analysis of that group for the state. Young and Bayer (1979) compiled a list of
the dragonfly larvae in the Guadalupe River basin in Texas. Only Kennedy
(1921) has made a major effort to describe larvae of several species in the
region.

Species lists have been published for the peripheral states of Louisiana
(101 spp.; Bick 1957), Arkansas (133 spp.; Harp and Rickett 1977; Harp
1983a,b; Harp 1985; Harp and Harp 1996), Oklahoma (126 spp.; Bick and
Bick 1957) and New Mexico (97 spp.; Evans 1995). These lists have been
helpful additions to the knowledge of dragonfly and damselfly distributions in
this region, but generally have not involved the systematic sampling of stream
systems or vegetational or physiographic subregions (except Bick 1957), nor
association with abiotic and biotic parameters of occurrence, necessary for
biogeographic analysis.

STUDY AREA AND METHODS

The south central Nearctic Region, as we are defining it (Fig. 1), covers
approximately 560,000 km 2 , of which 412,000 km 2 are in Texas. It includes
the seven biotic provinces of Texas and the portions of those provinces from
Arkansas, Oklahoma, Louisiana, New Mexico and northeastern Mexico that
immediately join Texas. The Mississippi River forms the eastern boundary,
and the Navahonian biotic province bounds the western edge of the region.
Mean annual precipitation ranges from 25-147 cm/yr, most falling in March-
May. Major vegetation types include eastern pines and hardwoods, central prai-
ries and grasslands and western deserts. The seven distinct regional biotic prov-
inces differ in topography, temperature, vegetation, soil type, geology and cli-
mate as outlined by Blair (1950), Dice (1943) and Blair and Hubbell (1938).
They are: 1) Chihuahuan, 2) Navahonian, 3) Kansan, 4) Balconian, 5)
Tamaulipan, 6) Texan, and 7) Austroriparian (Fig. 1). Elevation ranges from
sea level to 2,667 meters (Guadalupe Peak, Culberson Co., TX) in the Guadalupe
Mountains National Park.

Intensive sampling, with emphasis on Texas, began in September, 1993
and included more than 30 expeditions transecting the biotic provinces or con-
centrating in such areas as the Guadalupe Mountains National Park of the
Navahonian Province. Collection of adults and larvae involved sampling all
traversed lotic habitats and selected lentic habitats. Documentation of the fauna
of states adjacent to Texas was based primarily on both published and unpub-
lished records of acknowledged collectors and museum holdings, and limited
sampling.

Adults were placed in glassine envelopes and submerged in 99% acetone
for overnight, then removed and allowed to dry completely before being per-

Vol. 109, No. 3, May & June, 1998

203

Oklahoma

NAVAHONIAN

BALCONIAN

f ,'~

AMAULIPA

Mexico

Figure 1. The six natural biotic provinces of the south central Nearctic Region (modified from
Blair 1950, Dice 1943 and Blair and Hubbell 1938).

manently stored in polyethylene envelopes with data cards. Larvae were col-
lected using a Wildco-type 46 \ 22.5 cm net with a 1 mm mesh size. Exuviae
were hand collected from emergence sites. Both exuviae and larvae were placed
in Kahle's solution and then rinsed and permanently preserved in 80% ethanol
upon returning to the laboratory. Numerous distribution records were based on
reared species; mature larvae were transported alive to the laboratory in por-
table styrofoam rearing chambers as described by Szczytko and Stewart ( 1 979),
and reared in a conditioned Frigid Units Living Stream maintained at collec-
tion or slightly raised temperatures.

Collections Examined

We examined all regional material from the extensive and previously undocumented Beatty
collection, housed at the Frost Entomological Museum of Penn State University (PSU) and from

204 ENTOMOLOGICAL NEWS

the Arkansas State University Museum of Zoology (ASUMZ), Florida State Collection of
Arthropods (FSCA), International Odonata Research Institute (IORI), Sul Ross State University
(SRSU) and the Texas A&M Insect Collection (TAMU). Individuals and institutions who do-
nated study material include T.W. Donnelly (Binghamton, New York), S.W. Dunkle (Collin County
Community College), J. Gelhaus (Academy of Natural Sciences of Philadelphia), S. Jasper (Texas
A&M University), B.C. Kondratieff (Colorado State University), D.R. Paulson (University of
Puget Sound) and C.R. Nelson (University of Texas, Austin).

RESULTS

Table 1 lists the 228 species of damselflies and dragonflies currently known
from the south central Nearctic biotic provinces (Fig. 1). The classification
used here follows that of Garrison (1991), except that in our view Epicordulia
and Tetragoneuria are considered subgenera of Epitheca (K.J. Tennessen pers.
comm.). The following four species represent new Texas species records, and
one, Neoneura amelia, is a new U.S. species record:

Neoneura amelia Calvert. TEXAS: Hidalgo Co., Rio Grande River nr. La Lomita Mission, W of
Granjeno, 12 May 1997, 3Cf , 29, Univ. of North TX Collection. New for the US.

N eoerythromma cultellatum (Hagen in Selys). TEXAS: Hidalgo Co., Rio Grande River nr. La
Lomita Mission, W of Granjeno, 12 May 1997, 2Cf , Univ. of North TX Collection. New for
Texas.

Aphylla williamsoni (Gloyd). TEXAS: Jefferson Co., 1 2 August 1989, 1 Cf , photograph taken by
R.A. Honig. TEXAS: Montgomery Co., pond @ E side of IH 45 and S of FM 1488, 11
August 1997, 1 9 , photograph taken by R.A. Behrstock. New forTX.

Somatochlorafilosa (Hagen). TEXAS: Trinity Co., Rt. 94, 1 .3 mi W of Angelina County line, 23
August 1995, 1 9 , B. Mauffray Collection. New for TX.

DISCUSSION

The south central Nearctic Region (Fig. 1) is important as a boundary
(Paulson 1982) for some species of the largely eastern fauna of central and
east Texas (Texan and Austroriparian Provinces) that represent a temperate
element, and those of south Texas and northeastern Mexico (Tamaulipan Prov-
ince), representing a subtropical element. However, for other species these
provinces are actually a mixing zone, and at least a short distance dispersal
corridor; for example, our records indicate that of the 177 species occurring in
the Austroriparian and Texan Provinces, 81 species (46%) also occur in the
Tamaulipan Province and 59 species (33%) cross the Rio Grande to the south,
into northeastern Mexico. Conversely, of the 108 species in the northeastern
Mexico states of Tamaulipas and Nuevo Leon, 80 species (74%) cross the Rio
Grande, entering Texas and 61 species (57%) occur in the Austroriparian and
Texan Provinces.

A similar boundary or mixing phenomenon exists near the Rio Grande
River between the more northern Kansan/Navahonian/Balconian and the more
southern Chihuahuan Province. Of the 142 species occurring in the Kansan,

Vol. 109, No. 3, May & June, 1998 205

Navahonian and Balconian Provinces, 84 species (59%) also occur in the
Chihuahuan Province and many of these cross the Rio Grande farther south,
into northeastern Mexico. Conversely, of the 50 species in the northern Mexico
states of Coahuila and Chihuahua, 41 species (82%) cross the Rio Grande,
entering Texas with 39 (46%) of them occurring in the Kansan, Navahonian
and Balconian Provinces.

The Balconian Province represents diverse species assemblages from the
northern and southern provinces bordering or near it. Fifty percent (99) of the
199 species occurring in the northern Navahonian/Kansan/Texan/Austroriparian
Provinces occur in the southern Chihuahuan/Tamaulipan Provinces. Conversely,
93 (69%) of the 1 35 species occurring in the southern Chihuahuan/Tamaulipan
Provinces occur in the northern Kansan/Texan/Austroriparian Provinces. These
examples and recent discoveries of dispersals across the Rio Grande (Abbott
1996), including Neoneura amelia Calvert, Neoerythromma cultellatum Hagen
in Selys, Aeshna psilus Calvert, Dythemis maya Calvert, Micrathyria didyma
(Selys), Tauriphilia azteca Calvert and Tramea insularis Hagen, from Mexico
into Texas suggest that the area is indeed a dispersal corridor and that future
dispersals across it may occur. These distinct species assemblages and mixing
zones undoubtedly relate to the climate and vegetational characteristics defin-
ing these provinces. The Rio Grande border is a more effective barrier among
certain groups than in others. The cordulegastrids and corduliids are poorly
represented in Mexico, Central and South America, while groups such as the
coenagrionids, aeshnids and libellulids are much more widely distributed and
are well represented in these areas (Paulson 1982).

The large number of Odonata larvae still unknown to science from the
southwestern U.S. is attributed in part to relatively less collecting effort
(McCafferty et al. 1990) and low species densities in combination with sparse
and patchy habitats (Provonsha and McCafferty 1973). The larval descriptions
of many of these regional species are insufficient for identification purposes
(McCafferty et al. 1990). The rarity and/or local distributions of many species
in the eastern part of this region account for the large number of unknown
larvae there.

Three species of Odonata in the south central Nearctic Region are listed as
"species of concern" by the United States Fish and Wildlife Service. This fed-
eral listing identifies species "for which information now in the possession of
the Service indicates that proposing to list as endangered or threatened is pos-
sibly appropriate, but for which persuasive data on biological vulnerability
and threat are not currently available to support proposed rules" (USFWS 1 996).
Two species, Somatochlora margarita Donnelly and Argia leonorae Garrison
have locally restricted distributions. Argia leonorae is known from only five
counties in south and west Texas and in northern Mexico. The larva of this
species, in its natural habitat, remains undiscovered.

206 ENTOMOLOGICAL NEWS

A status survey of Somatochlora margarita conducted by the Texas Parks
and Wildlife Department (Price et al. 1989) expanded its initial range, a five
mile radius in the Sam Houston National Forest, to a now estimated 10,000
square mile area in east Texas and western Louisiana. Somatochlora margarita
is endemic to the longleaf and loblolly pine forests of southeastern Texas. We
are in the process of describing its larva from the exuviae of reared material
(donated by S.W. Dunkle). We believe that rigorous sampling in the sandy
bottom streams of the Big Thicket of East Texas will lead to the discovery of
this larva in its natural habitat, and provide potential insights into its life
history.

A third species, Macromia wabashensis Williamson, is also listed as Cat-
egory 2 by the USFWS. Dubious records and the validity of M. wabashensis
as a distinct species are factors leading to its Category 2 status. Several speci-
mens referred to as M. wabashensis have been collected in McLennan and
Falls Counties, Texas (Williams 1982). These records are currently thought to
be exceptionally yellow forms of M. taeniolata Rambur or a hybrid between
M. taeniolata and M. pacifica Hagen (Dunkle pers. comm.; Garrison 1995). It
has thus been omitted from our list.

There is considerable taxonomic confusion concerning the Tetragoneuria
group of this region. We provisionally list Epitheca (T.) semiaquea (Burmeister)
from Texas and Oklahoma based on determinations made by K.J. Tennessen.
Further study including DNA analysis is needed on this group.

Bick (1983) reported 32 species of North American Odonata (8% of that
fauna) to be at risk, citing the loss of high-quality, undisturbed streams as the
most significant factor endangering odonates in North America. Four of these
(Neoneura aaroni Calvert, Gomphus ozarkensis Westfall, Somatochlora mar-
garita Donnelly and 5. ozarkensis Bird) occur in the south central U.S. and are
considered rare (Bick 1983). Effective conservation efforts depend on accu-
rate knowledge of the current distribution of each species (Moore 1991).
Continued effort is needed to indicate which strategies should be taken to
conserve particular species and which breeding sites should be given priority
for protection.

ACKNOWLEDGMENTS

We wish to thank all who provided data and support, especially S.W. Dunkle, T.W. Donnelly,
R.W. Garrison, B. Mauffray and G.L. Harp. K.J. Tennessen provided valuable input and verifica-
tion of specimens of Epitheca. We would like to thank S.R. Moulton, II and G.H. Beatty for
allowing us to examine the Beatty collection, Frost Entomological Museum at Penn State. R.A.
Behrstock and R.A. Honig generously furnished us with photographic records. Valuable field
help was provided by J.W. Chirhart, J.O. Martinez, K.V. Moore and M.V. Passanante. We also
thank S.W. Dunkle, C.R. Nelson and two anonymous reviewers for comments on earlier drafts of
this manuscript.

Vol. 109, No. 3, May & June, 1998

207

Table 1 . Distribution of the 227 species of Odonata currently known from the south central
Nearctic Region. Arkansas (AR), Austroriparian (AUST), Balconian (BALC), Chihuahua (CHI),
Chihuahuan (CHIH), Coahuila (COA), Kansan (KANS), Louisiana (LA), Navahonian (NAVA),
New Mexico (NM), Nuevo Leon (NLN), Oklahoma (OK), Tamaulipan (TAMA), Tamaulipas
(TAM), Texan (TEXA) and Texas (TX). Distributional annotations are noted by: (*) = new state
record, (**) = new U.S. record.

United States

Mexico

Biotic Provinces

A L N O T
R A M K X

C C N T
H O L A
I A N M

A B C K N T T
U A H A A A E
S L I N V M X
T C H S A A A

ZYGOPTERA (73)

Calopterygidae (5)

Calopteryx dimidiata Burmeister

X X

C. maculate (Beauvois)

XX XX

XXX

Hetaerina americana (Fabricius)

X X X X X

X X X X

X X X X X X X

H. litia (Drury)

XX XX

X X

X X X X X X X

H. vulneraia Hagen in Selys

X X X X

XXX

Lestidae (9)

Archilestes grandis (Rambur)

X XXX

X XX

X X X X X X

Lestes alacer Hagen

XXX

X X X X X X X

L. disjunctus australis Walker

X X X X X

X X X X X X X

L.forficula Rambur

X XX

L. inaequalis Walsh

XX XX

L. rectangularis Say

X X

XXX

L. sigma Calvert

X X

X XX

L. unguiculatus Hagen

X X

XX X

L. vigilax Hagen

XX XX

Protoneuridae (3)

Neoneura aaroni Calvert

X X

N. amelia Calven**

Protoneura cara Calvert

X X

Coenagrionidae (56)

Acanthagrion quadratum Selys

X X

Amphiagrion abbreviatum (Selys)

X X

XXX

Argia alberta Kennedy

X X

A. apicalis (Say)

X X X X X

XXX XXX

A. barretti Calvert

X X

XX XX

A. bipunctulata (Hagen)

XX XX

X X

A. cuprea (Hagen)

X X

A.fumipennis Burmeister

X X X X X

X X X X X X X

A. hinei Kennedy

X X

XXX

A. immunda (Hagen)

X XXX

X XX

X X X X XX

A. leonorae Garrison

XX XX

A. lugens (Hagen)

XXX

XX X

A. moesta (Hagen)

X X X X X

X X X X

X X X X X X X

A. munda Calvert

X X

A. nahuana Calvert

XXX

X X

X X X X X X X

A. plana Calvert

X XXX

XXX

X X X X X X X

A. rhoadsi Calvert

X X

A. sedula (Hagen)

X X X X X

X X X X

X X X X X X X

A. tibialis (Rambur)

XX XX

XXX

A. translata Hagen in Selys

X XXX

X XX

X X X X X X X

Chromagrion conditum (Selys)

Enallagma antennatum (Say)

. aspersum (Hagen)

X XX

X X

E. basidens Calvert

X X X X X

X XX

X X X X X X X

. boreale Selys

XXX

E. civile (Hagen)

X X X X X

X X X X

X X X X X X X

. concisum Williamson

. cyathigerum (Charpentier)

XXX

. daeckii (Calvert)

XX XX

. divagans S61ys

XX XX

X X

. doubledayi (Selys)

. dubium Root

X XX

208

ENTOMOLOGICAL NEWS

Table 1 (Continued)

United States

Mexico

Biotic Provinces

A L N O T
R A M K X

C C N T
H O L A
I A N M

A B C K N T T
U A H A A A E
S L I N V M X
T C H S A A A

. durum (Hagen)

X X

X XX

. exsulans (Hagen)

XX XX

X X

X X X X XX

. geminatum Kellicott

XX XX

X X

E. ncvaehispaniae Calvert

X X

XX X

. praevarum (Hagen)

XXX

X X X X X

. signatum (Hagen)

XX XX

XX X XX

. traviatum westfalli Donnelly

XX XX

X X

E. vesperum Calvert

XX XX

X X

Hesperagrion heterodoxum (Selys)

X X

XXX

X X X X

Ischnura barberi Cmrie

XXX

X X X X X

/. damula Calvert

XXX

/. demorsa (Hagen)

XXX

X X

XXX

/. denticollis (Burmeister)

XXX

XXX X

/. hastata (Say)

X X X X X

X X

X X X X X X X

/. kellicotti Williamson

XX XX

/. perparva McLachlan in Selys

X X

XXX

/. posila posila (Hagen)

XX XX

X X X X XX

/. prognata (Hagen)

X X

/. ramburii (Selys)

XX XX

X XX

X X X X XX

/. verticalis (Say)

X XXX

Nehalennia integricollis Calvert

X XX

Neoerythromma cultellatum (Selys)

Telebasis byersi Westfall

XX X

T. salva (Hagen)

XXX

X XX

X X X X X X X

ANISOPTERAU55)

Petaluridae (1)

Tachopteryx thoreyi (Hagen in Selys)

XX XX

X X

Aeshnidae (16)

Aeshna constricta Say

A. dugesi Calven

X X

XX X

A. multicolor Hagen

XXX

X X X X X X

A. psilus Calvert

X X

A. umbrosa umbrosa Walker

X XX

X XXX

Anax amazili (Burmeister)

X X

A. junius (Drury)

X X X X X

XX X

X X X X X X X

A. longipes (Hagen)

XX XX

XX X

A. walsinghatni McLachlan

X X

X X X X

Basiaeschna Janata (Say)

XX XX

XXX X

Boyeria vinosa (Say)

XX XX

XX X

Coryphaeschna ingens (Rambur)

XX X

X XX

Epiaeschna hems (Fabricius)

XX XX

XX X XX

Gomphaeschnafurcillata (Say)

XX X

Gynacantha nen'osa Rambur

X X

Nasiaeschna pentacantha (Rambur)

XX XX

XX X XX

Gomphidae (38)

Aphylla angustifolia Garrison

X X

XX XX

A. protracta (Hagen)

X X

XX XX

A. williamsoni (Gloyd)

X *

Arigomphus lentulus (Needham)

X XX

X X

A. mamelli (Ferguson)

XX X

A. submedianus (Hagen)

XX XX

XX X

A. villosipes (Selys)

Dromogomphus armatus Selys

D. spinosus Selys

XX XX

XX X

D. spoliatus (Hagen in Selys)

XX XX

X X

XX X XX

Erpeiogomphus compositus Hag. in Selys

X X

X X X X X

. crotalinus (Hagen in Selys)

XXX

. designates Hagen in Selys

X X X X X

XXX

X X X X X X X

. eutainia Calvert

X X

. heterodon Garrison

X X

. lampropeltis Sampropeltis Kennedy

X X

Gomphus (Gomphurus) externus Hagen

X XXX

X X X X X X X

Vol. 109, No. 3, May & June, 1998

209

Table 1 (Continued)

United States

Mexico

Biotic

Provinces

A
R

L
A

N
M

T
X

C C N T
H O L A
I A N M

A
U

S
T

B
A
L
C

C
H

I
H

K N T
A A A
N V M

S A A

T
E
X

G. (G.) gomalezi Dunkle

G. (G.) hybridus Williamson

G. (G.) modestus Needham

G. (G.) ozarkensis Westfall

G. (G.) vastus Walsh

Gomphus (Gomphus) apomyius Donnelly

G. exilis Selys

G. graslinellus (Walsh)

G. lividus (Selys)

G. militaris Hagen

XXX

G. oklahomensis (Pritchard)

Hagenius brevistylus Selys

Ophiogomphus westfalli Cook & Daigle

Phyilogomphoides albrighti (Needham)

X X

P. stigmatus (Say)

X X

Progomphus borealis McLachlan

X X

P. obscurus (Rambur)

XXX

Stylogomphus albistylus (Hagen in Selys)

Stylurus intricatus (Hagen)

X X

5. laurae (Williamson)

S. plagiatus (Selys)

X X

Cordulegastridae (3)

Cordulegaster erronea Hagen in Selys

C. maculata Selys

C. obliqua obliqua (Say)

Corduliidae (24)

Macromiinae (6)

Didymops transversa (Say)

Macromia alleghaniensis Williamson

M. annulata Hagen

X X

M. illinoiensis georgina (Selys)

X X

M. pacifica Hagen

M. taeniolaia Rambur

Corduliinae(lS)

Epitheca (Epicordulia) princeps Hagen

X X

E. (Tetragoneuria) costalis (Selys)

E. (T.) cynosura (Say)

E. (T.) petechialis (Muttkowski)

X X

E. (T.) semiaquea (Burmeister)

E. (T.) spinosa Hagen

Helocordulia selysii (Hagen)

H. uhleri Selys

Neurocordulia alabamensis Hodges

N. molesta (WaJsh)

N. virginiensis Davis

N. xanihosoma (Williamson)

Somatochlora filosa (Hagen)

5. georgiana Walker

5. linearis (Hagen)

5. margarita Donnelly

5. ozarkensis Bird

S. lenebrosa (Say)

Libellulidae (73)

Brachymesia furcata (Hagen)

XXX

B. gravida (Calvert)

X X

B. herbida (Gundlach)

Brechmorhoga mendax (Hagen)

X X

XXX

Cannaphila insularis/unerea (Carpenter)

Celithemis amanda (Hagen)

C. elisa (Hagen)

C. eponina (Drury)

XXX

C.fasciata Kirby

C. ornata (Rambur)

C. verna Pritchard

210

ENTOMOLOGICAL NEWS

Table 1 (Continued)

United States

Mexico

Biotic Provinces

A L N T
R A M K X

C C N T
H O L A
I A N M

A B C K N T T
U A H A A A E
S L I N V M X
T C H S A A A

Dythemis fugax Hagen

X XXX

X X X X X X X

D. maya Calvert

X XX

X X

D. nigrescens Calvert

X X

XX XX

D. velox Hagen

X X X X X

X XX

X X X X XX

Erythemis collocata (Hagen)

X X

XXX

E. plebeja (Burmeister)

X XX

E. simplicicollis (Say)

X X X X X

X X

X X X X X X X

E. vesiculosa (Fabricius)

X X

XX X X X X

Erythrodiplax berenice berenice (Drury)

XX X

X X X X X X

E. connata (Burmeister)

E.funerea (Hagen)

X X

E.fusca (Rambur)

X X

E. minuscula (Rambur)

XX XX

XX X

E. umbrata (Linnaeus)

XX XX

X X

X X X X XX

Libellula auripennis Burmeister

XX XX

X XX

L. axilena West wood

X X

L. comanche Calvert

XXX

X X X X X

L. composite (Hagen)

X X

XXX X

L. croceipennis Selys

XXX

X X

X X X X XX

L. cyanea Fabricius

X XX

XXX X

L. deplanata Rambur

XX XX

X X

L. flavida Rambur

XX XX

XX X

L. forensis Hagen

XXX

L. incesta Hagen

XX XX

XXX X

L. luctuosa Burmeister

X X X X X

X X X X X X X

L. lydia Drury

X X X X X

X X X X X X X

L. needhami Westfall

XX X

X XX

L. pulchella Drury

X X X X X

X X X X X X X

L. saturata Uhler

XXX

X X X X X X X

L. semifasciata Burmeister

XX XX

X X

L. subornata (Hagen)

XXX

L. vibrans Fabricius

XX XX

X X

Macrodiplax balieata (Hagen)

XX X

X X X X XX

Macrothemis imitans leucoiona Ris

X X

X XX

M. inacuta Calvert

X X

M. inequiunguis Calvert

Miathyria marcella (Selys)

XX X

X X

XX XX

Micrathyria aequalis (Hagen)

X X

M. didyma (Selys)

M. hagenii Kirby

X X

XXX XX

Orthemis ferruginea (Fabricius)

X X X X X

X X X X

X X X X X X X

Pachydiplax longipennis ( Burm.)

X X X X X

X X

X X X X X X X

Paltothemis lineatipes Karsch

X X

X X X X X X

Pantalaflavescens (Fabricius)

X X X X X

X X X X

X X X X X X X

P. hymenaea (Say)

X X X X X

XXX

X X X X X X X

Perithemis domitia (Drury)

X X

P. lenera (Say)

X X X X X

XXX

X X X X X X X

Pseudoleon superbus (Hagen)

X X

XX XX

Sympetrum ambiguum (Rambur)

XX XX

XXX

S. corrupt urn (Hagen)

X X X X X

XX X

X X X X X X X

5. costiferum (Hagen)

5. illotum (Hagen)

X X

5. internum Montgomery

X X

S. occidentale fasciatum Walker

X X

XXX

S. vicinum (Hagen)

X XXX

XX XX X

Tauriphila azteca Calvert

Tholymis citrina Hagen

Tramea calverti Muttkowski

X XX

T. Carolina (Linnaeus)

XX XX

XX X

T. insularis Hagen

T. lacerata Hagen

X X X X X

X X X X X X X

T. onusta Hagen

X X X X X

X X

X X X X X X X

Totals

127 112 82 132 1%

37 24 66 62

157 109 92 95 76 106 139

Vol. 1 09, No. 3, May & June, 1 998 211

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Coop. Agreement No. 14-16-0002-86-925, Amndt. No. 7.
Pronvonsha, A.V. and W.P. McCafferty. 1 973. Previously unkown nymphs of western Odonata

(Zygoptera: Calopterygidae, Coenagrionidae). Proc. Entomol. Soc. Wash. 75:449- 454.
Quintero, D. and A. Aiello, eds. 1992. Insects of Panama and Mesoamerica.. Oxford Univ.

Press, Oxford, England. 720 pp.
Szczytko, S.W. and K.W. Stewart. 1979. The genus Isoperla (Plecoptera) of western North

America: holomorphology, systematics and a new stonefly genus Cascadoperla. Mem. Amer.

Entomol. Soc. 32:1-120.
Tinkham, E.R. 1934. The dragonfly fauna of Presidio and Jeff Davis Counties of the Big Bend

Region of trans-pecos, Texas. Can. Entomol. 66:213-218.
Thicker, E.S. 1908. Incidental captures of neuropterous insects at Piano, Texas. Psyche. 15:97-

100.
USFWS, Department of Interior. 1996. Endangered and threatened wildlife and plants; review

of plant and animal taxa that are candidates for listing as endangered or threatened. Federal

Odonatologica 1:157-168.
Williamson, E.B. 1914. Dragonflies collected in Texas and Oklahoma. Entomol. News. 25:41 1-

415,444-455.
Young, W.C. and C.W. Bayer. 1979. The dragonfly nymphs (Odonata: Anisoptera) of the

Guadelupe River basin, Texas. Texas J. Sci. 31:86-97.

Vol. 109, No. 3, May & June, 1998 213

SCIENTIFIC NOTE:

NEW DISTRIBUTIONS FOR

RAPTOHEPTAGENIA CRUENTATA AND AMETROPUS NEAVEI
(EPHEMEROPTERA: HEPTAGENIIDAE, AMETROPODIDAE)!

R.D. Waltz, 2 G. F. Edmunds, Jr?, Gary Lester 4

Large river habitats possess some of the least known mayfly species in North America
(McCafferty et al. 1990). Difficulty in sampling such habitats has undoubtedly contributed to the
report of widely disjunct distributions of large river species. Decline in the quality of large river
habitat has also possibly contributed to localized extirpations and further increased the apparent
disjunction of reported distributions (see Whiting and Lehmkuhl 1987, McCafferty et al. 1990).
Herein, two large river species, which are rarely collected, are newly reported from Montana.
One of these two species is also newly reported from Minnesota.

Raptoheptagenia cruentata (Walsh) has been reported previously from nine states or prov-
inces in North America based on available literature (see Whiting and Lehmkuhl 1987, Edmunds
and Waltz 1995). Reports of larval collections cited in the preceding papers include Arkansas,
Illinois, Indiana, Montana, Ohio, and Saskatchewan. McCafferty (1988) designated the neotype
of R. cruentata based on a larva in Indiana, which is housed in the Purdue Entomological Re-
search Collection (PERC), West Lafayette, IN. Adult collections have been reported from Illi-
nois, Indiana, Nebraska, Tennessee, and Manitoba.

Two R. cruentata larvae taken in the Powder River, by G. Romero, with the following col-
lection data: MT: Custer Co., Powder R., 11 -XI- 1976(1 larva), and same locale, 11 -VIII -1976
(2 larvae) were the source of the previously unpublished Montana record reported by Edmunds
and Waltz (1995). In addition a single, nearly mature larva of R. cruentata was recently taken in
collections made at the Montana state line in the Little Missouri River at MT: Carter Co., Little
Missouri River, Sec 1 2, T 6S, R 62E, 1 7- VII- 1 996. An additional new state distribution report for
R. cruentata is herein given for Minnesota as MN: Sibley Co., Minnesota R. (drift net), 30- VI-
1974, C.M. Haynes (1 young larva).

The more rarely reported species Ametropus neavei McDunnough, was also collected in the
Powder River from MT: Custer Co., Powder River, 1 1 -XI- 1976, G. Romero ( 1 larva). A. neavei
was previously known from the type locality in Alberta and from Saskatchewan, Canada (Allen
and Edmunds 1976) and Michigan (Steven and Hilsenhoff 1979).

The recording of distributions of rare or rarely collected species is important 1 ) in order to
provide data to adequately assess species rarity (e.g., see McCafferty and Edmunds 1997), and 2)
to provide local and regional regulatory personnel and ecologists with data potentially critical to
large stream management decision making.

The above habitats in Montana are large streams with firm sand substrates. We do not have
data characterizing the Minnesota habitat. However, detailed data from the Little Missouri River
site is available through monitoring activities reported to us by Warren Kellogg, Watershed Spe-
cialist, USDA NRCS, Helena, MT. Kellogg's data characterizes the Little Missouri at this site as

1 Received October 2, 1997. Accepted October 23, 1997.

2 IDNR, Division of Entomology and Plant Pathology, 402 West Washington, Room W-290,
Indianapolis, IN 46204.

3 University of Utah, Department of Biology, Salt Lake City, UT84112.

4 EcoAnalysts, Inc., P.O. Box 3103, Moscow, ID 83843-1906.

ENT. NEWS 109(3) 213-214, May & June, 1998

2 1 4 ENTOMOLOGICAL NEWS

an intermittent, warm water stream. Channel catfish and carp were present in the immediate area.
Physical characters available include: pH 8.6; temp 27 C; nitrates < 0.01 mg/1; TKN - 0.2 mg/1;
OP-0.014 mg/1; TP-0.03mg/l; TSS-26mg/l. The specimen of R. cruentata taken was collected by
means of a kick screen. The riparian vegetation includes patches of large cottonwoods (Populus
deltoides) and willows (Salix spp.) encroaching on the sand/silt bars. There was little or no shade
at the collection site.

We thank Warren Kellogg for providing this data characterizing the collection site.

Vouchers originating from the personal collection of George F. Edmunds, Jr., and the larva
from Little Missouri River are deposited at the Purdue University Entomological Research Col-
lection (PERC), West Lafayette, Indiana.

LITERATURE CITED

Allen, R.K. and G.F. Edmunds, Jr. 1976. A revision of the genus Ametropus in North America
(Ephemeroptera: Ametropodidae). J. Kansas Entomol. Soc. 49: 625-635.

Edmunds, G.F., Jr. and R.D. Waltz. 1995. Chapter 1 1 . Ephemeroptera. pp. 126-163. In: R.W.
Merritt and K. W. Cummins (Eds.), An Introduction to the aquatic insects of North America,
3rd Edition. Kendall/Hunt, Dubuque, IA.

McCafferty, W.P. 1988. Neotype designation for Raptoheptagenia cruentata (Walsh) (Ephe-
meroptera: Heptageniidae). Proc. Wash. Entomol. Soc. 90: 97.

McCafferty, W.P. and G.F. Edmunds, Jr. 1997. Critical commentary on the genus Siphlonisca
(Ephemeroptera: Siphlonuridae). Entomol. News 108: 141-147.

McCafferty, W.P., B.P. Stark, and A.V. Provonsha. 1 990. Ephemeroptera, Plecoptera, and
Odonata, p. 43-58. In: Systematics of the North American Insects and Arachnids: Status and
Needs, ed. M. Kosztarab and C.W. Schaefer. Virginia Agricultural Experiment Station Infor-
mation Series 90-1. Blacksburg: Virginia Polytechnic Institute and State University. 247pp.

Steven, J.C. and W.L. Hilsenhoff. 1 979. Ametropus neavei (Ephemeroptera: Ametropodidae) in
the Upper Peninsula of Michigan. Great Lakes Entomol. 12:226.

Whiting, E.R. and D. M. Lehmkuhl. 1987. Raptoheptagenia cruentata, gen. nov. (Ephe-
meroptera: Heptageniidae), new association of the larva previously thought to be Anepeorus
with the adult of Heptagenia cruentata Walsh. Can. Entomol. 119: 405-407.

Vol. 109, No. 3, May & June, 1998 215

SCIENTIFIC NOTE:

CLOEODES EXCOGITATUS (EPHEMEROPTERA:
BAETIDAE) IN NORTHERN CALIFORNIA 1

R.D. Waltz 2 , Peter Ode 3 , Jon Lee 4

The small minnow mayfly genus Cloeodes Traver is found in many different freshwater
habitats throughout the Neotropics, Afrotropics, Asia and southern temperate areas of North
America. Various revisionary works (Waltz and McCafferty 1987) and systematic works (Kluge
1991; Waltz 1993; Waltz and McCafferty 1994; McCafferty and Lugo-Ortiz 1995) have pro-
vided taxonomic data for the genus and added greatly to our knowledge of the world distribution
of the genus.

The genus was recognized as having tropical affinities by Waltz and McCafferty (1987),
McCafferty and Waltz (1990), and McCafferty et al. (1992). McCafferty and Lugo-Ortiz (1995)
and Lugo-Ortiz and McCafferty (1993, 1994, 1995) further added to the known distribution of
the genus in the Western Hemisphere, viz., southwestern North America, Mesoamerica, and South
America.

Herein, we report a remarkable, although possibly not unexpected, distribution record for
this primarily subtropical and tropical genus from northern California. Larvae of C. excogitatus
Waltz and McCafferty were taken in the following site in northern California: CA: Mendocino
Co., Williams Crk near Covelo, 29-IX-1996. The above site is characterized by fine silt bottoms
mixed with cobble. Stream temperatures are warm in the summer months. The above specimens
were taken in low flow conditions near the end of the summer dry season.

Lugo-Ortiz and McCafferty (1994, 1995) extended the known range of C. excogitatus from
its type locality in Oak Creek Canyon, Arizona, southward to Yavapai County (Arizona) and
Mexico. The type locale of C. excogitatus was previously the most northern report of the genus
in North America. This report extends the known range of this species northward into North
America by at least 600 miles reaching the Coastal Biotic Province of California (Usinger 1953)
at approximately 40 degrees north latitude. Other northward extensions of primarily tropical
genera into North America are also known in other mayfly genera, e.g., the baetid genera
Camelobaetidius Demoulin and Paracloeodes Day (see McCafferty et al. 1992).

Voucher specimens are deposited at the Purdue Entomological Research Collection, Purdue
University, West Lafayette, IN and the California Academy of Science, San Francisco.

LITERATURE CITED

Kluge, N. 1991 . Cuban mayflies of the family Baetidae (Ephemeroptera). 1 . Genera Callibaetis,
Cloeodes, and Paracloeodes. Zool. Zh. 12: 128-136. [in Russian].

Lugo-Ortiz, C.R and W.P. McCafferty. 1993. Genera of Baetidae (Ephemeroptera) from Cen-
tral America. Entomol. News 104: 191-195.

1 Received October 31, 1997. Accepted December 17, 1997.

2 1DNR, Division of Entomology and Plant Pathology, 402 West Washington, Room W-290,
Indianapolis, IN 46204.

3 Aquatic Biological Assessment Laboratory, California Department of Fish and Game, 2005
Nimbus Road, Rancho Cordova, CA 95670.

4 2250 Wilson Street, Arcata, CA 9552 1 .

ENT NEWS 109(3) 215-216, May & June, 1998

216 ENTOMOLOGICAL NEWS

Lugo-Ortiz, C. R. and W.P. McCafferty. 1994. New records of Ephemeroptera from Mexico.

Entomol. News 105: 17-26.
Lugo-Ortiz, C.R. and W.P. McCafferty. 1995. Annotated inventory of the mayflies

(Ephemeroptera) of Arizona. Entomol. News 106: 131-140.
McCafferty, W.P. and C.R. Lugo-Ortiz. 1995. Cloeodes hydation, n.sp. (Ephemeroptera:

Baetidae): An extraordinary, drought tolerant mayfly from Brazil. Entomol News 106: 29-

35.
McCafferty, W.P. , R.W. Flowers, and R.D. Waltz. 1992. The biogeography of Mesoamerican

mayflies, pp. 173-193. In: S.P. Darwin and A.L. Weldon (eds.). Biogeography of Mesoamerica:

proceedings of a symposium. Tulane Univ. Stud. Zool. Bot., Suppl. Publ. 1.
McCafferty, W.P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera)

of North and Middle America. Trans. Am. Entomol. Soc. 1 16: 769-799.
Usinger, R.L. 1953. Introduction to Aquatic Entomology. Stream and Lake Classifications, pp

7-8 in Aquatic Insects of California, with keys to North American genera and California

species. Univ. California Press, Berkeley. 508 pp.
Waltz, R.D. 1993. Cloeodes binocularis (Ephemeroptera: Baetidae) a new combination for a

Neotropical species of Pseudodoeon s. auctt. Entomol. News 104: 233-234.
Waltz, R.D. and W.P. McCafferty. 1 987. Revision of the genus Cloeodes Traver (Ephemeroptera:

Baetidae). Ann. Entomol. Soc. Am. 80: 191-207.
Waltz, R.D. and W.P. McCafferty. 1 994. Cloeodes (Ephemeroptera: Baetidae) in Africa. Aquat.

Insects 16: 165-169.

Vol. 109, No. 3, May & June, 1998 217

SCIENTIFIC NOTE

FIRST RECORD OF ALEIODES DEPANOCHORA
(HYMENOPTERA: BRACONIDAE) FROM BRAZIL

A. M. Penteado-Dias2

ABSTRACT: Aleiodes depanochora is recorded for the first time from Manaus, Descalvado and
Dourado, Brazil.

Aleiodes depanochora van Achterberg belongs to the Aleiodes dispar group in which males
have peculiar depressions on the second and third metasomal tergites that are absent in females
(van Achterberg, 1985). The Aleiodes dispar group includes eight species: three from the Old
World, A. excavatus (Telenga, 1941 ), A. takasuae van Achterberg, 1 985 and A. yasirae van Ach-
terberg, 1995, and five from the Neotropical region: A. elliptidepressus Penteado-Dias & van
Achterberg, 1995, from Brazil, A. longipendulatus van Achterberg, 1985, from Costa Rica, A.
rugosicostalis van Achterberg, 1995, from Peru, A. brevipendalatus van Achterberg, 1995, from
Ecuador and A. depanochora, van Achterberg, 1995, from Peru. The localities listed below are
the first records for A. depanochora from Brazil.

Diagnosis: Aleiodes depanochora can be recognized by the following characters: head (ex-
cept stemmaticum) yellowish-brown dorsally; side of pronotum dark brown or infuscate, paler
parts not strongly contrasting with darker parts; metasomal tergites 1-3 partly infuscate;
pterostigma evenly dark brown; hind tarsus infuscate or dark brown; length of malar space of
male 1.1-1.2 times basal width of mandible and 0.3-0.35 times height of eye; face transversely
rugose, dorsal face of pronotum in lateral view distinctly protruding; hind coxa densely trans-
versely striate dorsally; first metasomal tergite more coarsely rugose laterally than medially and
somewhat widened posteriorly; fourth tergite of male partly retracted and smooth; depression of
second metasomal tergite of male without median carina and slender ovoid; median carina of
second tergite of male 0.2-0.4 times length of depression; ratio of forewing veins 1-CUI: 2-CUI
= 1:8-12, vein 3-SR of fore wing 1 .0 1.3 times as long as vein 2-SR.

Specimens examined: 4 males, BRAZIL, Amazonas, Manaus, B. Klein, November 8,25,29,
1 984 (Instituto Nacional de Pesquisas da Amazonia); 1 male, BRAZIL, Sao Paulo, Descalvado,
Fazenda Escaramucas, M. M. Dias, 9.XI, 1985 (light trap); 1 male, BRAZIL, Sao Paulo, Dou-
rado, Fazenda Morro Chato, L. A. Joaquim, 26. V. 1994 (Departamento de Ecologia e Biologia
Evolutiva da Universidade Federal de Sao Carlos, SP).

The specimens from Manaus were collected in the Amazonian rainforest and those from
Descalvado and Dourado were collected in Atlantic forest areas (Fig. 1 ).

The discovery of the same species in the primary rain forest (Peru and Manaus) and in the
secondary Atlantic forest (Dourado and Descalvado) is specially interesting. The biology of the
Aleiodes dispar group species is unknown; the sparse data available on them indicate that they
are parasites of lepidopterous larvae living in low vegetation (van Achterberg, & Penteado-Dias,
1 995). Same as Aleiodes yasirae van Achterberg, 1 995 and A. elliptedepressus van Achterberg &
Penteado-Dias, 1995, we found A. depanochora in lowland vegetation, in a gallery forest close
to a river. Probably that is the distribution pattern for Brazilian Aleiodes dispar group species and
there we can find their hosts.

1 Received April 19, 1 996. Accepted September 8, 1997.

2 Departamento de Ecologia e Biologia Evolutiva, Universidade Federal de Sao Carlos. CP 676.
CEP 13 565-905, Sao Carlos, SP, Brasil.

ENT. NEWS 109(3) 217- 218, May & June, 1998

218

ENTOMOLOGICAL NEWS

30
Figure 1. Collection records of Aleiodes depanochora van Achterberg.

ACKNOWLEDGMENTS

I wish to thank the Institute Nacional de Pesquisas da Amazonia for the loan of the listed
specimens. This work was supported by Conselho Nacional de Desenvolvimento Cientifico e
Tecnologico and Funda9ao de Amparo a Pesquisa do Estado de Sao Paulo from Brazil.

I wish to thank also C. van Achterberg, R. A. Wharton and P. Marsh who provided sugges-
tions for improvements to the manuscripts.

LITERATURE CITED

Achterberg, C. van. 1985. The Aleiodes dispar group of the Palaearctic region (Hymenoptera:

Braconidae: Rogadinae) Zool. Med. Leiden 59: 178-187, figs. 1-20.
Achterberg, C. van & A.M. Penteado-Dias. 1 995. Six new species of the Aleiodes dispar group

(Hymenoptera: Braconidae: Rogadinae). Zool. Med. Leiden (1):1-18, fig. 1-48.

Vol. 109, No. 3, May & June, 1998

219

MEMBERS OF THE AMERICAN ENTOMOLOGICAL SOCIETY

December 18, 1997

The previous list of 3 1 3 members was published in the September-October
1984 issue of Entomological News. That issue celebrated the 125th Anniver-
sary of the Society. This present list contains 412 members from every state in
the union (and the District of Columbia), except Alaska, Montana, Oregon and
Rhode Island. The largest representation is from Pennsylvania (43). Interna-
tional members represent 16 countries on the five continents.

The format used in this list differs from the previous list in that all types of
members appear in a single alphabetical list. Names of both Honorary and Life
members are in bold face type. Names of Life members are further identified
with an asterisk (*). The date shown is the year of record in our office when
each member first joined the Society.

I thank Suzanne McElroy for her assistance in the preparation of this list.
Any corrections to this list can be sent to the Corresponding Secretary at the
A.E.S. address on the inside front cover of this issue. Corrections will be pub-
lished in a future issue of Entomological News.

William J. Cromartie,
Corresponding Secretary

State or

Surname

Name

Yr.

Country

Surname

Name

Yr.

Country

Abbott

John C.

1991

Bilby

Peter J.

1993

Ahlstrom

Kenneth R.

1996

Bilyj

Bohdan

1995

Canada

Ahn

Kee-Jeong

1996

Bohart

Richard M.

1967

Aime

Cathie

1996

Bolton

Michael J.

1987

Alexander

Kevin D.

1993

Boobar

Lewis

Allen

Robert T.

1988

Reginald

1996

Allred

Miriam L.

1987

Bouseman

John K.

1984

Amrine, Jr.

James W.

1983

Bowles

David E.

1985

Anderson

Robert S.

1968

Boyd

Howard P.

1939

Armitage

Brian J.

1980

Boys

Frank E.

1970

Baker

Charles W.

1980

Brigham, Jr.

John A.

1986

Ball

George E.

1963

Canada

Brown

John W.

1989

Balogh

George J.

1987

Brown

Kirby W.

1973

Barry

Donald W.

1986

Brown

Harley P.

1970

Bartlert

Charles R.

1992

Brown

Wendy S.

1993

Bartow

Dennis H.

1970

Brown

MarkW.

1997

Baumann

Richard W.

1970

Brown, Jr.

Keith S.

1970

Brazil

Baumgardner

David E.

1992

Brzoska

David W.

1987

Beal, Jr.

Richard S.

1970

Burbutis

Paul P.

1970

Beaman

Carol A.

1996

Burdick

Donald J.

1993

Beckemeyer

RoyJ.

1979

Burger

John F.

1980

Ntt

Belish

Timberley A.

1997

Burian

Steven K.

1997

Bell

Ross T.

1985

Burrows

Weldon L.

1983

Berk

Mary

1989

Bushey

Sonja Marie

1994

ENT. NEWS 109(3) 219-223, May & June, 1998

220

ENTOMOLOGICAL NEWS

State or

Surname

Name

Yr.

Country

Surname

Name

Yr.

Country

Butler

Linda

1979

Dunkle

Sidney W.

1983

Bystrak

Paul G.

1991

Durfee

Richard

1994

Cancellare

Joseph A.

1977

Earle

Jane

1993

Canterbury

Lawrence E.

1975

Edmunds

George F.

1948

Canton

Steven P.

1978

Edwards

Robert L.

1990

Carlton

Chris E.

1996

Elliott

Douglas E.

1984

Caron

Dewey M.

1983

Elmali

Meryem

1996

Turkey

Carter

Janet L. Stein

1986

Enns

Wilbur R.

1971

Cauble

Ronald L.

1993

Epler

John H.

1976

Chandler

Donald S.

1985

Erikson

JeffS.

1994

Chandler, Jr.

Jack H.

1996

Estes

William J.

1985

Chiba

Hideyuki

1988

Japan

Ettinger

William H.

1986

Chordas III

Steve W.

1997

Ettinger

William S.

1979

Ciborowski

Jan J. H.

1978

Canada

Evans

Howard E.

1973

Ciegler

Janet C.

1996

Fall

Louise H.

1985

Ciurlino

Randolph A.

1994

Fava

Jane F.

1997

Clausen

Philip J.

1983

Fee

Frank D.

1979

Coher

Edward I.

1994

Ferreira

Raul N.

1994

Conklin, Jr.

DonJ.

1988

Fisher

Stephen C.

1994

Conn

D. Bruce

1991

Fisher

Elizabeth

1945

Contreras-

Flemyng

E. Penryn

1986

Ramos

Atilano

1986

Mexico

Flint, Jr.

Oliver S.

1982

Cook

Jerry L.

1995

Floyd

Michael A.

1991

Cooper

Kenneth W.

1946

Forbes

Gregory S.

1977

Coovert

Gary A.

1997

Foye

Laurene

1994

Courtney

Gregory W.

1991

Franchine

Michael J.

1997

Covell, Jr.

Charles V.

1969

Frank

Kenneth D.

1979

Cromartie

William J.

1991

Frank

Susan E.

1979

Currie

Charles E.

1992

Fredrickson

Richard W.

1983

Cuthrell

David L.

1996

Froeschner

Richard C.

1970

Cutler

Bruce

1991

Fuester

Roger W.

1965

Dakin, Jr.

Matt E.

1971

Fullerton

Stuart M.

1990

Darlington

Mark B.

1994

Funk

David H.

1987

Davis

Jack R.

1991

Gage

1986

Davis

Link M.

1995

Galford

Jim

1980

Day

William H.

1966

Ganeo de

Francisco de

De Souza

Paulo

Mello

Assis

1991

Brazil

Moutinho

Roberto

1995

Brazil

Gayubo

Severiano F.

1993

Spain

DeWalt

R. Edward

1991

Gelhaus

JonK.

1987

Deyrup

Mark A.

1978

Giesecke

Martiin

1979

Dolan

Michael F.

1997

Gomez-Arias

Luis M.

1992

Donelan

Laurence W.

1995

Goodrich

Michael A.

1989

Dougherty

Veronica M.

1984

Gordon

Robert D.

1969

Dozier

Herbert L.

1958

Gottschalk

Steven C.

1996

Drecktrah

H. Gene

1971

Graham

Alan C.

1993

DuBois

MarkB.

1988

Grant

Peter M.

1984

Duffield

Richard M.

1995

Graves

Robert C.

1967

Duncan

Paul M.

1963

Greathouse

Zane B.

1997

Vol. 109, No. 3, May & June, 1998

221

State or

Surname

Name

Yr.

Country

Surname

Name

Yr.

Country

Griffith

Michael B.

1992

Kingsolver

John M.

1969

Grubbs

Scott A.

1991

Kirchner

Ralph F.

1976

Gusten

Robert

1995

Germany

Kirchenstein

Barbara B.

1989

Haack

Robert A.

1996

Kistner

David H.

1985

Halstead

Jeffrey A.

1985

Klubertanz

TomH.

1997

Hamilton

Robert W.

1989

Knight

JeffB.

1963

Hamilton

Steven W.

1987

Knisley

C.Barry

1983

Hanley

Rodney S.

1992

Knizeski, Jr.

Henry M.

1985

Hansen

Dean

1990

Koehn

Leroy C.

1996

Harrington

DonG.

1989

Kondratieff

Boris C.

1982

Harris

Steven C.

1982

Koski

Joseph T.

1962

Hastriter

Michael W.

1996

Krinsky

William L.

1987

Heck

Mary L.

1995

Kritsky

Gene

1978

Henry, Jr.

Brad C.

1984

Krombein

Karl V.

1972

Heppner

John B.

1997

Krotzer

Steve

1991

Herbst

David B.

1996

Krysan

James L.

1981

Herrmann

Scott J.

1985

LaBerge

Wallace E.

1961

Heth

Robert K.

1996

Lacey

MarkS.

1994

Hildebrandt

Drew A

1984

Lago

Paul K.

1981

Hill

Richard E.

1993

Lanteri

AnaliaA.

1991

Argentina

Hill

Russell E.

1987

Lantsov

Vladimir I.

1996

Russia

* Hilton

Donald F. J.

1986

Canada

Larsen

Eric

1995

Hodges

R.W.

1960

Larson

Omer R.

1982

Hoebeke

E. Richard

1988

Lasalle

MarkW.

1979

Hoffman

Kevin M.

1987

Ledin

Kathryn E.

1993

Holdeman

Steven J.

1996

Lee

Sharon D.

1995

Holzbach

John E.

1975

Leschen

Richard A. B

1986

Australia

Homer

Norman

1987

Lester

Gary T.

1993

Houghton

David C.

1997

Levesque

Claire

1985

Canada

Huckstep

E. Elgin

1990

Levy

Stephen D.

1992

Hudson

Patrick

1985

Lewis

Robert E.

1956

lanni

Charles

1985

Lewis

Carolyn N.

1997

Iftner

David C.

1993

Lillie

Richard A.

1997

Insley

Sandra

Lingafelter

Steven W.

1997

Yawetz

1995

Linsley

E. Gorton

1962

Jackson

John K.

1990

Lipinski

Daniel R.

1989

Jasper

Sharon

Livingston

Margot

1987

Knight

1995

Livingston

CarlF.

1993

Jennings

Daniel T.

1969

Livingston

Rosemary A.

1993

Johnson

Zane B.

1997

Lowe

Graeme

1988

Katovich

Kerry

1994

Mac Neill

C.Don

1995

Kavanaugh

David H.

1995

Mackay

William

1996

Kearns

Ruth S.

1994

Mackenzie

Allen H.

1987

Kedanis

Richard J.

1997

Majerowicz

Eugene I.

1991

Keiper

JoeB.

1996

Manley

GaryV.

1980

Kelley

Richard I.

1988

Marque/, Luna

Juan

1994

Mexico

* Kennedy

James H.

1980

Marshall

Brett D.

1997

Kidd

Kathleen A.

1992

222

ENTOMOLOGICAL NEWS

State or

Surname

Name

Yr.

Country

Surname

Name

Yr.

Country

Martinez

Humberto

Quiroz

1989

Mexico

Penteado-Dias

Angelica

Marx

Rainer Peter

1996

Germany

Maria

1994

Brazil

Mason

Charles E.

1976

Perez

Juan

Mathis

Wayne N.

1977

Dominguez

Francisco

1988

Mexico

Matsunaga

Wallace 0.

1985

Peters

William L

1998

Matta

James F.

1974

Pfadt

Robert E.

1988

May

Michael L.

1992

Philtower

Ramona

1993

McCabe

Tim L.

1997

Philips

James R.

1982

McCafferty

W. Patrick

1987

Pinter

Lawrence J.

1980

McCaleb

John E.

1996

Plan

Austin P.

1988

McCauley

Luana M. M.

1995

Plotnikoff

Robert W.

1995

McHugh

Joseph V.

1989

Plummer

John A.

1990

Michener

Charles D.

1963

Polhemus

John T.

1996

Miller

LeD.

1967

Porter

Charles H.

1968

Miller

William B.

1993

Preston

Floyd W.

1980

Miller

Jacqueline Y.

1989

Price

Michele B.

1997

Molnar

Steven A.

1997

Purrington

Foster F.

1997

Morrison

Michael W.

1993

Ranger

Christopher

Morse

John C.

1980

1997

Moulton 11

Stephen R.

1987

Rayburn

Brian S.

1997

Muchmore

William B.

1989

Rebollar-Tellez

Eduardo A.

1993

United

Muniz Velez

M.enC. Raul

1992

Mexico

Kingdom

Munro

James B.

1995

Rentz

David C.F.

1965

Australia

Murphy

Clint

1991

Rhodes

Howard A.

1996

Murvosh

Chad M.

1991

Richards

Austin Brady

1996

Myles

Timothy G.

1986

Canada

Rider

David A.

1987

Naczi

Robert F.C.

1983

Riley

Edward G.

1986

Namkaidorj

1996

Mongolia

Ringgier

Theodore G.

1992

Navarrete-

Risley

Lance S.

1989

Heredia

Jose Luis

1992

Mexico

Ritter

Karla S.

1979

Neff

Stuart E.

1988

Robbins

Richard G.

1988

Nelson

Charles H.

1969

Roberts

Richard H.

1971

Nelson

S.Mark

1991

Robinson

Jeffrey M.

1995

Nelson

C. Riley

1984

Robison

Henry W.

1990

Nelson

Harry G.

1982

Romig

Ronald F.

1979

Nielson

M.W.

1992

Roth

Louis M.

1957

Novinger

James S.

1991

Rothschild

MarkJ.

1987

O'Brien

Charles W.

1993

Ruesink

William G.

1988

O'Donnell

Sean

1992

Ruffin

JaneM.

1988

O'Keefe

Sean T.

1991

Ruiter

David

1976

Ocus

Warren G.

1992

Russell

DanaC.

1992

Omer

John R.

1990

Rust

Richard W.

1974

Otte

Daniel

1976

Sabourin

Michael

1993

Parrott

Rod

1977

Canada

Salmon

J.T.

1960

New

Payne

Randall G.

1993

Zealand

Pennington

Wendell L

1985

Sandridge

Paul T.

1991

Vol. 109, No. 3, May & June, 1998

223

State or

Surname

Name

Yr.

Country

Surname

Name

Yr.

Country

Santiago de

Tennessen

Kenneth J.

1996

Bueno

Silvia

1994

Mexico

Thompson

F.C.

1969

Sarver

Randy J.

1996

Tinerella

Paul P.

1996

Saunders

Robert D.

1990

Canada

Torres

Felix

1993

Spain

Scarbrough

Aubrey G.

1987

Torres-Miller

Laura

1997

Schaefer

Paul W.

1985

Trond

Andersen

1997

Norway

Schesser, Jr.

James F.

1978

Turnbow, Jr.

Robert H.

1983

Schiefer

Terence L.

1984

Turner

Alan R.

1994

N.C.

Schmidt

Justin 0.

1987

Valenti

Michael A.

1985

Schmude

KurtL.

1992

Ventre

Vincent

1985

Schroder

Robert F. W.

1993

Vogtsberger

RoyC.

1989

Schweitzer

Dale

1989

Wagner

Michael R.

1980

Seaborg

Norman G.

1963

Wahl

David B.

1984

Seltmann

Katja Chantre

1996

Walls

Jerry G.

1984

Shapiro

Arthur M.

1967

Waltz

Robert D.

1983

Sheldon

Joseph K.

1973

Webb

David K.

1996

Shepard

William D.

1979

Weber

Neal A.

1961

Short

Andrew E.

1996

Weber

Richard G.

1986

Shubeck

Paul P.

1971

* Weisse

Theodore H.

1987

Sibley

Paul K.

1989

Weissmann

Michael J.

1988

Sikes

Derek S.

1996

Wesson

Laurence G.

1995

Sissom

William

Wheatley

John B.

1995

David

1995

Wheeler

Jeanette N.

1970

Skelley

Paul E.

1991

White

Harold B.

1975

Slayback

Scot E.

1997

White

David S.

1976

Sleeper

Elbert L.

1971

White

Timothy M.

1996

Smith

David R.

1966

D.C.

Whitehead

V.B.

1976

S. Africa

Smith

Stephen M.

1993

Canada

Whitfield

James B.

1993

Sneen

Martin E.

1993

Whitney

Susan P.

1990

Snider

Richard J.

1985

Wiersema

Nick

1997

Sobat

Thomas

1996

Wiker

James R.

1990

Spangler

Hay ward G.

1990

Willemse

Per

1968

Netherlands

Spokony

Harvey

1997

Williams

David W.

1988

Spooner

John D.

1986

Williams

Roger N.

1991

Springer

Charles A.

1992

Williams

J. Logan

1995

Staines

Charles L.

1986

Willink

Abraham

1963

Argentina

Stallings

Viola N.

1988

Wilson

Stephen W.

1992

Stein

Raymond J.

1988

Wimmer

H. Peter

1979

Stewart

Kenneth W.

1994

Wojtowicz

John A.

1983

Stidham

John A.

1982

Wolinski

Jeffrey A.

1988

Strassmann

JoanE.

1983

Wright

David M.

1995

Strazanac

John S.

1984

Young

Daniel K.

1980

Sublette

James E.

1990

Young

OrreyP.

1980

Summerville

Keith S.

1996

Young

Wayne L.

1986

Surman

Michael A.

1976

Zeigler

David D.

1990

Tarter

Donald C.

1972

Zoidis

John

1996

Greece

Taylor

Steven J.

1988

Zuccaro, Jr.

Anthony E.

1981

224 ENTOMOLOGICAL NEWS

SOCIETY MEETING OF NOVEMBER 19, 1997

Dr. Jim Marden

Dept. of Biology
Perm State University

Dr. Marden began by pointing out the evolution of flight is a classic challenge for Darwinian
natural selection, for it is difficult to envision how gradual, incremental selection can result in
complex traits that function only in their fully developed form. The fossil record offers little help in
determining the origin of insect wings and flight, for at the time winged insects first appear in
fossils (325mya), they had already radiated and diversified into stem groups of all of the major
lineages present today. Thus, we are left to decipher the evolutionary history of insect flight from
our judgment of which traits of fossils and extant insects most closely represent the ancestral con-
dition. The emergent view from this body of research is that insect wings arose from moveable,
articulated gills of aquatic ancestors, and that the orders Ephemeroptera (mayflies) and Plecoptera
(stoneflies) have the most primitive morphology among extant winged insects.

Dr. Marden 's laboratory has recently proposed that surface-skimming, a form of nonflying
aerodynamic locomotion used by certain modern stoneflies, is a feasible intermediate stage be-
tween swimming and flying, and might be a remnant of the ancestral condition for all winged
insects. This hypothesis has been lauded as the most reasonable model for a transitional process
offered to date, but it has also been criticized for failing to consider the phylogenetic position of
skimming stoneflies. In his presentation, he gave an overview of work examining i) functional
aspects of various forms of surface skimming, ii) preliminary results of a molecular phylogeny of
stoneflies using 18S rDNA sequence data, and iii) the phylogenetic distribution of skimming be-
havior across the Plecoptera. These results suggest that surface skimming is a primitive trait among
stoneflies, and thus a likely candidate for a transitional stage in the evolution of insect flight. The
talk was illustrated with many striking slides and a video of surface-skimming flight.

In insect-related news, Jon Gelhaus circulated a specimen of Dermatobia hominis larva
(Diptera) recently excised from a member of the ANS staff forty-five days after a trip to Guyana.
The adult of this parasite catches a mosquito, to which it attaches its offspring, and the mosquito
passes it on to a human.

Mention was also made of impending trials of two Lyme disease vaccines by Philadelphia
area pharmaceutical firms.

- W. J. Cromartie
Corresponding Secretary

ENT. NEWS 109(3) 224, May & June, 1998

Study notices, want items and for sale notices are published in The Entomologist's
Market Place .

Reprints: (without covers) may be ordered when corrected page proofs are returned to the
editor. Schedule of reprint costs will appear on order form.

This publication is available
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Call toll-free 800-521-3044. Or mail inquiry lo
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The Entomologist's Market Place

The following books are available from Kendall/Hunt Publishing:

Immature Insects, Volumes I and II, edited by Frederick W. Stehr (Volume I: 1 987/768 pages/
casebound/$136.44*/ISBN 0-84034639-5 and Volume II: 1991/992 pages/casebound/
241.44*/ISBN 0-8403-4639-5). Immature Insects provides information on the biology and
ecology of the families and selected important species. The two volume set also gives you a
means to identify insects ranging from the most common to the extremely rare through use of
the abundant illustrations, descriptions, and/or keys to selected species. Immature Insects is
the only reference that extensively covers updated information necessary to identify imma-
ture insects. These books also describe techniques necessary for the collecting, rearing, kill-
ing, preserving, storing, and studying of insects. Both books also include an introduction
defining how terms are used in the book, a complete glossary, and an extensive index.

For more information or to place an order, call Jill Crow at 1-800-228-0564.
*A11 prices are subject to change.

FOR SALE: Baltic amber pieces with insects, spiders, plants. Material for scientific work, as

well as better pieces for display and teaching. O. Holden, Junkerg. 37, S-126 53 Hegersten,

Sweden, fax: 01146-8-7268522.
FOR SALE: Light traps, 12 volt DC or 1 10 volt AC with 15 watt or 20 watt black lights. Traps

are portable and easy to use. Rain drains and beetle screens protect specimens from damage.

For a free brochure and price list, contact Leroy C. Koehn, 207 Quail Trail, Greenwood, MS

38930-7315. Telephone 601-455-5498.

VOL. 109

SEPTEMBER & OCTOBER, 1998

US ISSN 0013-872X
NO. 4

ENTOMOLOGICAL NEWS

n Mexican species ofGastrisus (Coleoptera:

taphylinidae) J.L. Navarrete-Heredia, J. Marquez 225

_ Aj'T'"" aism in Neogerris hesione (Heteroptera:

Jerridae) in southern Illinois S.J. Taylor, J.E. McPherson 233

A new species of Paruroctonus (Scorpiones: Vaejovi-

dae) from Big Bend National Park, Texas W.D. Sissom, R. N. Henson 240
Comparison of sand nesting wasps (Hymenoptera) from

Frank E. Kurczewski 247

two pine barrens areas in upstate New York
The identity of Tachysphex acutus (Hymenoptera:

Sphecidae), an unsolved mystery
Character variability & a new synonym of

Acerpenna pygmaea (Ephemeroptera:

Baetidae) R.D. Waltz, D.E. Baumgardner, J.H. Kennedy

Reared association & equivalency of Baetis

adonis & B. caelestis (Ephemeroptera:

Baetidae) W. P. McCafferty, E. L. Silldorff

Additions & corrections to Ephemeroptera species of

North America & index to their complete

nomenclature W.P. McCafferty

Taxonomic notes on Evaniodini (Hymenoptera:

Braconidae), with redescription of Evaniodes

spathiiformis & description of a new species

SM. Barbalho, A.M. Penteado-Dias
Zorcadium Bergroth, an objective junior synonym

of Pseudobebaeus Fallou (Heteroptera:

Pentatomidae) DA. Rider, C. Fischer

Mass appearance of lady beetles (Coleoptera:

Coccinellidae) on North Carolina beaches

CA. Nalepa, K.R. Ahlstrom, BA. Nault, J.L. Williams
Biological & morphological notes on Dasyhelea pseudo-

incisurata (Diptera: Ceratopogonidae) Lawrence J. Hribar

Eastward range extension in Canada of the alderfly

Sialis velata (Megaloptera: Sialidae), & the

potential of the genus as a contaminant monitor /. Roy, L. Hare

Gynandromorph of Helicoverpa armigera

(Lepidoptera: Noctuidae)

A. Josephrajkumar, B. Subrahmanyam, V.V. Ramamurthy
A new species of Sweltsa (Plecoptera: Chloro-

perlidae) from eastern No. America B.C. Kondratieff, R.F. Kirchner
SCIENTIFIC NOTES
Incorporation of Batrachospermum gelatinosum

(Rhodophyta) into cases of Ochrotrichia wojcickyi

(Trichoptera: Hydroptilidae) J.B. Keiper, DA. Casamatta, BA. Foote
Additions to Iowa mayflies (Ephemeroptera) W.P. McCafferty, T. Hubbard
BOOK REVIEW

BOOKS RECEIVED & BRIEFLY NOTED
ANNOUNCEMENT

SOCIETY MEETING of March 25, 1998
SOCIETY MEETING of April 22, 1998

Frank E. Kurczewski 252

257

261

269

274

277

282

285

288
293

NOV 9 1998

THE AMERICAN ENTOMOLOG

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Vol. 109, No. 4, September & October, 1998 225

A NEW MEXICAN SPECIES OF GASTRISUS
(COLEOPTERA: STAPHYLINIDAE) 1

Jose Luis Navarrete-Heredia^, Juan Marquez^

ABSTRACT: Gastrisus newtonorum, new species, is described based on specimens from the
states of Mexico, Guerrero, Jalisco and Morelos. One specimen was examined by Bernhauer,
however it was misidentified as G. mimetes Sharp. The Mexican record of G. mimetes in Bemhauer
and Schubert's and Blackwelder's catalogs was probably based on this misidentification. The
two species are compared, and the aedeagi are illustrated. Distributional and biological data are
provided.

Gastrisus Sharp, 1876 is an American genus,with most species in South
America. At present, 19 species are recognized: Blackwelder (1944) cited 12
species; Scheerpeltz (1972) moved six species from Trigonopselaphus to this
genus; and one more is described here.

Gastrisus mimetes Sharp was described based on a single specimen from
Costa Rica (Sharp, 1884: 360). Years later, Bernhauer and Schubert (1914)
and Blackwelder (1944) recorded this species from Mexico. However, we do
not know if the Mexican record was ever published elsewhere. Bernhauer (1912:
39) in his description of G. venezolanus mentioned that he had identified the
holotype earlier as G. mimetes, and that he was comparing it to specimens of
G. mimetes from Colombia and Peru (country records not included in the cata-
logs cited above).

One specimen from Guerrero (at FMNH), identified by Bernhauer as G.
mimetes belongs to an undescribed species. This situation was first recognized
by A.F. Newton, Jr. who compared one specimen from Guerrero with the holo-
type of G. mimetes at BMNH (see material examined). Later, he received speci-
mens from G. Ruiz-Lizarraga and L. E. Rivera collected in Guerrero and Jalisco
respectively, that he identified as Gastrisus n. sp.

Ruiz-Lizarraga (1993), in her excellent contribution on carrion staphylin-
ids, provided descriptions of the species that she collected, and included one
for Gastrisus n. sp. From that date until now this species has awaited formal
description. Upon request, Dr. Newton kindly sent us specimens of the
undescribed species from FMNH and four G. mimetes from Costa Rica to
support this work. Also, Miss G. Ruiz-Lizarraga allowed us to describe this
species. The goal of this paper is to describe and provide biological data on
this new species.

1 Received September 12, 1997. Accepted November 19, 1997.

Entomologia, Centro de Estudios en Zoologia, CUCBA, Universidad de Guadalajara, Apdo.
Postal 234, 45100 Zapopan, Jalisco, Mexico.

Lab. de Morfofisiologia Animal, Fac. de Ciencias, UNAM, 04510, Mexico, D.F.
ENT. NEWS 109(4) 225-232, September & October, 1998

226 ENTOMOLOGICAL NEWS

MATERIALS AND METHODS

Specimens for this study were borrowed from: Field Museum of Natural
History, Chicago (FMNH); Coleccion Entomologica, Escuela Nacional de
Estudios Profesionales, Iztacala (ENEPI); and Coleccion Entomologica,
Universidad de Costa Rica (UCR); others were collected by the authors during
their research. Acronyms for collections where type material will be deposited
are: American Museum of Natural History, New York ( AMNH); British Mu-
seum, London (BMNH); Canadian National Collection, Ottawa (CNC);
Entomologia, Centre de Estudios en Zoologia, Universidad de Guadalajara,
Zapopan (CZUG); Institute de Biologia, UNAM, D.F. (IBUNAM); Snow En-
tomological Museum, Lawrence, Kansas (SEM); Laboratorio Especializado
de Morfofisiologia Animal, Fac. de Ciencias, UNAM, D.F. (LEMA); Juan
Marquez Luna Collection, D.F. (JML); Jose Luis Navarrete Collection, Zapopan
(JLN); and Museo de Historia Natural Ciudad de Mexico, D.F. (MHNCM).

Throughout this paper we refer to abdominal segments by their morpho-
logically comparable names and use roman numerals for these. The first fully
visible segment is segment III. Tergum II is usually narrowly visible. Total
length was measured from the anterior margin of the head to the apex of ab-
dominal segment IX.

Gastrisus newtonorum Navarrete and Marquez, NEW SPECIES

Figs. 1,2,4,5,8.

HOLOTYPE MALE: Length 1 5.6 mm. Black, except abdominal segments VII-IX, last seg-
ment of maxillary and labial palpi, and tarsi rufotestaceous. Surface covered with microsculpture
consisting of isodiametric meshes, mixed with scattered micropunctures; tempora with irregular
waves; neck and abdominal segments with dual microsculpture: isodiametrical meshes at base
and irregular waves on apical portion (more distinct on segments V1I-VIII).

Head subquadrate; setiferous punctures denser at posterior angles and along medial borders
of eyes, dorsal surface without setiferous punctures (Fig. 2). With subocular ridge. First antennal
segment slightly shorter than next two segments combined, second segment shorter than third
segment; fifth to eleventh transverse, large setae decreasing in number but short setae more
conspicuous. Second and third segments of maxillary palpi broader at apex, last segment elon-
gate, subcylindrical, as long as second segment. Last segment of labial palpi as broad as penultimate
segment and as long as last segment of maxillary palpi. Mandibles subequal in length to head
along midline. Right mandible with a tooth opposite an emargination of left mandible. Gular
sutures confluent at middle. Neck with oblique longitudinal line dorsolaterally on each side.

Pronotum slightly larger than head; narrowed toward base; anterior angles rectangular, basal
angles obtuse; setiferous punctures scattered, denser at sides, dorsal punctures 3:3, asymmetri-
cal; postcoxal process of the hypomeron translucent. Elytra opaque, with numerous setiferous
punctures, with two humeral macrosetae and one near scutellum. Two macrosetae on anterior
middle of prosternum. Tibiae with spines, denser on mesotibiae. First four segments of anterior
tarsi dilated, as broad as anterior tibiae, with modified pale setae ventrally; last segment as long
as previous three segments combined. Middle and hind tarsi similar: first segment as long as
following three segments combined, last segment as long as previous two segments combined.

Abdominal segments as shining as head and pronotum; tergites II1-IV with impressed line

Vol. 109, No. 4, September & October, 1998

227

1 mm

Figure I . Dorsal view of Gastrisus newlonorum Navarrete and Marquez, new species.

228

ENTOMOLOGICAL NEWS

1 mm

Imm

0.1mm

Figures 2-9. Morphology ofGastrisus spp. G. newtonorum: 2. Head and pronotum, 4. Aedeagus
(parameres removed), 5. Paramere, 8. Distribution of peg setae on paramere. G. mimetes. 3.
Head and pronotum, 6. Aedeagus (paramere removed), 7. Paramere, 9. Distribution of peg setae
on paramere.

Vol. 109, No. 4, September & October, 1998 229

on basal portion. One black macroseta on each side of posterior border of terga III-VI. Sternite

VII slightly emarginate at middle; VIII with conspicuous triangular emargination; sternites VII-

VIII with three black macrosetae on each side. Sternite IX emarginate, with two black macrosetae.
Aedoeagus as in Figs. 4-5, 8. Paramere with apex almost reaching apex of medial lobe.

ALLOTYPE FEMALE: Length 14.1 mm. Similar to holotype, except for: head narrower;
first four segments of anterior tarsi slightly less dilated; dorsal setiferous punctures on pronotum
4:4, asymmetrical; abdominal sternites VII-VIII not emarginate, sternite IX with two rufous
macrosetae on apex, in addition to two black ones.

Variation: Length 11.9-15.6 mm. Specimens from Guerrero are mostly paler, primarily on
prothorax where borders are reddish brown, and with abdominal segments VII-VIII yellow. Also,
one specimen examined by Ruiz-Lizarraga (1993) and five examined by us have abdominal
segments VII-VIII dark reddish brown. Aedeagi of these specimens are black, their dark color
pattern is likely caused by the mixture of liquids used as killing agents. A few specimens have the
pronotum and elytra apparently green in dorsal view. One specimen from Morelos has the last
segment of the right maxillary palpus black. Dorsal punctures on the pronotum are highly vari-
able: 2:3, 3:3, 3:4, 3:5, 4:2, 4:3, 4:4, 4:5; in some specimens, the last puncture is on the posterior
half of the pronotum. The peg setae on the paramere are slightly variable in number.

MATERIAL EXAMINED: Holotype: MEXICO: MORELOS, Yautepec, Carretera Mexico-
Cuautla, km. 19. Selva baja caducifolia. ex excremento. 16-Jun-1996. J. Marquez col. Allotype:
Tlayacapan, camino a Sta. Catarina. Selva baja caducifolia. Zona 4, 1534 msnm. 1 al 30- VI-
1996. ex NTP-80 (calamar). J. Marquez col.

Paratypes: MEXICO: MORELOS, Tlayacapan. 31-X-1992. ex excremento vacuno. J.
Marquez col. (ICf). Same data except for: 27-VIII-1995. I. Sanchez y J. Marquez cols. (2Cf).
Same data except for: zona 5, cultivo de temporal y selva baja caducifolia. 1 al 30-VII-1995, ex
NTP-80. J. Marquez col. (2Cf, 19). Same data except for: 1 al 30-IX-1996 (19). Same data
except for: 1 al 30-VI-1996 (8Cf). Tlayacapan, camino a Sta. Catarina, zona 4, selva baja
caducifolia. 1 -VI- 1 996, ex excremento vacuno (1 9 , ICf ). Same data except for: 9- VI- 1 996 (29,
4Cf). Same data except for: ex frutos podridos (2Cf). Same data except for: 10-VI-1996, ex
excremento vacuno (ICf ). Same data except for: 1 al 30-VII-1995, ex NTP-80 (ICf). Same data
exceptfor: 1 al 30-VIII- 1995 (ICf, 29 ). Same data except for: 1 al 30-IX- 1995 (29). Same data
except for: 1 al 30-VI-1996 (2Cf, 29 ) Tlayacapan, San Jose de los Laureles. Bosque mesofilo
de montana perturbado, zona 3. 1 al 30-X-1996, ex NTP-80. K. Villavicencio y J. Marquez cols.
(19). Same data except for: 1 al 30-XI-1995 (ICf). Cuemavaca, Col. del bosque. Bosque de
Pino-Encino. 23-VII-1995, ex excremento vacuno, J. Marquez col (1C?). Yautepec. Carretera
Mexico-Cuautla, km. 19. Selva baja caducifolia. 16- VI- 1996, ex excremento. J. Marquez col.
(ICf, 19). Tlayacapan, San Jose de los Laureles, BMM, 1751 m, 21.VII-24.VIII. 1991, J.L.
Navarrete y G.A. Quiroz, # 941 D, NTP-80, (ICf, 19); same data except for: 29. VI. 1991, J.L.
Navarrete, #677, ex Boletus edulis IV (ICf). GUERRERO, 2900 ft. 6 mi El Ocotito. VIII.30-
IX.5-1971/ human dung trap 380. A. Newton/ yellow card. Gastrisus cf mimetes Sharp, comp.
Holotype A. Newton, 1989/ Gastrisus n. sp. det. Newton 1 992 (1 Cf ). Same data: (1 9 ). 9 mi NE
Iguala. 1340 m. VIII. 29-IX.4- 1971 A. Newton coll., human dung trap 378. (3Cf, !<?). 3100 ft.
7.5 mi N El Ocotito. VIII.30-IX.5- 1971, human dung trap 381. A. Newton (4Cf, 49). 9 mi NE
Iguala. 4500 ft. VIII.29-IX.4- 1971, human dung trap 379. A. Newton (ICf ). 3200 ft. 10 mi N El
Ocotito. VIII.30-IX.5-1971 , human dung trap 382. A. Newton (1 Cf ). 1464 m. 4 mi W Mazatlan.
VIII.30-IX.5-1971. A. Newton coll., human dung trap 383 (29). 2014 m. 7 mi W Mazatlan.
VIII.30-IX.5- 1971. A. Newton coll., human dung trap 384 (19). Chilpancingo, Guerrero, 4600
ft. June. H.H. Smith/ Gastrisus mimetes Sharp/ det. Bernhauer. Godman-Salvin col. 1911. 345/
Sharp colln. By Exchange with Brit. Mus. (N. H.)/ F.M.N.H. [green card]/ Gastrisus cf. mimetes
det. A. F. Newton 1989. (1). Mochitlan, Acahuizotla, 650 m, tropical subevergreen forest. June-
July, 1986. L. Delgado collr. Carrion trap NTP-80, 2 (2Cf ). Sierra del Alquitran, BEP, 1670m,
20.VII.1990, ex NTP-80, J. Blackaller y L. Delgado cols. (3Cf); same data except for: Encinar
tropical, 21. VII. 1990, (4Cf). JALISCO, Sierra de Manantlan, El Tigre, 18- VII- 1988. Selva me-

230 ENTOMOLOGICAL NEWS

dia, nocturna, 3:pescado; L. Rivera./ Exchange ex University of Guadalajara, Mex. FIELD MU-
SEUM/ Gastrisusn. sp. del. Newton, 1996 (ICf). Sierra de Manantlan, Mpio. Casimiro Castillo,
El Tigre, 700 m, 18- VII- 1988. Bosque trop. subcad., L. Rivera leg./ Exchange ex University of
Guadalajara, Mex. Field Museum (1C?). Same data except for: G. Andrade col. (ICf ). ESTADO
DE MEXICO, Nanchititla, 6/Octubre/ 1 995 1540 msnm. Est. 2. Selva Baja Caducifolia. NTP-80-
3. A. Morales col. (ICf, 59)- Same data except for: l/Agosto/1995. 1110 msnm. Est. 1. Selva
Baja Caducifolia. NTP-80-3. A. Morales col. (19). Same data except for: I/Julio/ 1995. 1540
msnm. Est. 2. Selva Baja Caducifolia. NTP-80-2. A. Morales col. (3Cf, 29 ) Same data except
for: 26/Agosto/1995. 1110 msnm. Est. 1. Selva Baja Caducifolia. NTP-80-2. A. Morales col.
(19)- Same data except for: 1540 msnm. Est. 2. Selva Baja Caducifolia. NTP-80-1. A. Morales
col. (2Cf , 29 ) Same data except for: I/Julio/ 1995. 1540 msnm. Est. 2. Selva Baja Caducifolia.
NTP-80-1. A. Morales col. (2Cf, 19). Same data except for: 4/Junio/1995. 11 10 msnm. Est. 1.
Selva Baja Caducifolia. NTP-80-1. A. Morales col. (19)- Same data except for: 27/Octubre/
1995. 1110 msnm. Est. 1. Selva Baja Caducifolia. NTP-80-1. A. Morales col. (19). Same data
except for: 21/10/95. Est. 2. Selva Baja Caducifolia. NTP-80-2. A. Morales col. (ICf ). Same data
except for: 27/Oct/95. Est. 2. Selva Baja Caducifolia. NTP-80-2. A. Morales col. (29). Holo-
type, Allotype and some paratypes will be deposited at CZUG, other paratypes will be deposited
at FMNH, ENEPI, LEMA, JML, JLN, MHNCM, BMNH, CNC, AMNH, IBUNAM and SEM.

Etymology. We are pleased to dedicate this species to our friends and aca-
demic supervisors A. F. Newton, Jr., and M.K. Thayer, for their kind help and
support in the study of Mexican staphylinids.

Gastrisus newtonorum is similar to G. mimetes in color pattern, which is
probably the reason why Bernhauer misidentified the Mexican specimens as
G. mimetes. However, Gastrisus newtonorum is easily recognized because it
has more setiferous punctures on the head and thorax (Fig. 2-3); males with
abdominal sternite VII slightly emarginate, sternite VIII emarginate, and pri-
marily by difference in the aedeagus: the paramere is slightly bifurcate at the
tip in G. mimetes, entire in Gastrisus newtonorum (Figs. 4-7, 9). Females are
distinguished by the punctation on the head and thorax.

Our description has some important differences from that provided by Ruiz-
Lizarraga (1993): head and thorax black-copper; last two abdominal segments
red; elytra with conspicuous impressions; second segment of maxillary palp
broader and larger than remaining segments; dorsal pronotal punctures 3:3,
sublateral punctures 4:4; aedeagus different from Gastrisus mimetes. Speci-
mens we examined don't have that color pattern on the head, thorax and abdo-
men, and have the last two abdominal segments yellow or rufotestaceous. Also,
dorsal pronotal punctation is inconstant as already mentioned (see Variation).

Specimens of Gastrisus newtonorum have been found on several kinds of
decomposing organic matter, such as: carrion, fruits, mushrooms [misidentified
as Xenopygus analis (Er.) by Navarrete-Heredia, 1996], and human and cow
dung. As predators they seek prey there, probably maggots and larvae of other
beetles. This behavior is similar to that shown by species of Platydracus,
Belonuchus, Philonthus, Styngetus and other genera common in these habi-
tats.

Vol. 1 09, No. 4, September & October, 1 998 23 1

Most of the specimens were collected in tropical deciduous and sub-
evergreen forest during the rainy season (June-November). At present, the high-
est abundance is recorded from Acahuizotla, Guerrero (Ruiz-Lizarraga, 1 993).
She examined 173 specimens, 86 males and 87 females, from carrion traps. In
this locality this was the second most abundant staphylinid species, after
Belonuchus aff. xanthomelas Solsky. Specimens were collected between June
and October. Highest abundance was in July (Fig. 10).

Number of specimens
100

JUN JUL AUG SEP OCT NOV DEC

Months

Figure 10. Phenology of Gastrisus newtonorum at Acahizotla, Guerrero.

On the other hand, during a systematic survey of necrophilous staphylin-
ids (in prep, by Marquez) five localities were selected and designated ( 1 . pine-
Quercus forest, 1,874 m; 2. pine forest, 1,930 m; 3. disturbed cloud forest,
1,783 m; 4. tropical deciduous forest, 1,534; 5. corn field-tropical deciduous
forest, 1 ,634 m). Carrion traps were used for this research and were examined
each month. As in Acahuizotla, higher abundance was detected in tropical for-
est with 23 specimens, whereas in locality 3 only two specimens were col-
lected. No specimens were found at higher elevations in coniferous-2^rcM5
forest.

We suspect that, as in Acahuizotla, Gastrisus newtonorum is an important
predator in tropical forests and probably competes with other predatory sta-
phylinid species of genera such as Platydracus, Belonuchus, Philonthus,
Paederomimus and others whose phenology is similar.

It is clear that Gastrisus newtonorum is primarily distributed in tropical
forests, so its abundance is probably higher south to Guerrero and widespread
west to Jalisco and east to Oaxaca where tropical forests are common. More

232 ENTOMOLOGICAL NEWS

field data on this species and other staphylinids are necessary to get a better
understanding of the distributional pattern. The highest altitudinal record is
2030 m; the lowest is 700 m.

In Mexico and Central America, Gastrisus species are recorded from Costa
Rica (G. mimetes Sharp; Blackwelder 1944), Panama (G. opaculus Sharp;
Blackwelder 1944). It is interesting to note that at present we lack records of
any Gastrisus species between Mexico and Costa Rica, an unusual pattern for
a Xanthopygina species. Is this widely allopatric distribution for G. mimetes
and G. newtonorum, without records for Guatemala, Belize, Honduras, El Sal-
vador and Nicaragua, real? Possibly, but this may be the result of a lack of
intensive field work on staphylinids in this area.

Finally, records of Gastrisus mimetes from Colombia and Peru require re-
examination of the specimens to verify the identifications, or recognize pos-
sible misidentifications or new sibling species as was the case for G.
venezolanus.

Examined specimens of G. mimetes from Costa Rica are labeled as: COSTA
RICA: Osa Peninsula, Agua Buena, 1 km NW Boscosa centro,25. VII. 1990,
Banana plantation, pig dung, leg. K.Vulinec (1 Cf , 29 ) (FMNH); same data,
plus Gastrisus mimetes Shp. det Newton 1996 (FMNH); Puntarenas, R.F. Golfo
Dulce, 3 km SW Rincon, 10m, V- VI. 1992, P. Hanson, ex malaise, Gastrisus
sp. det. J.S. Ashe, 1996 (1 C?) (OCR).

ACKNOWLEDGMENTS

The authors thank A.F. Newton, Jr. and G. Ruiz-Lizarraga for permission to describe this
species. Special thanks are due to A.F. Newton, Jr. and M.K. Thayer for comments and informa-
tion that improved the manuscript when the new species lacked name, and to H.E. Fierros-Lopez
for the drawings. This paper is a contribution to the project "Estudios basicos de la fauna silvestre
del estado de Jalisco: diversidad, distribucion y habitat" supported by the Universidad de
Guadalajara.

LITERATURE CITED

Bernhauer, M. 1912. Zur Staphylinidenfauna von Sudamerika (10. Beitrag). Verh. Zool.-Bot.

Ges. Wien, 62:26-48.
Bernhauer, M. and K. Schubert. 1914. Staphylinidae IV. In: Coleopterorum Catalogus, Pars

57. Ed., S. Schenkling. W. Junk, Berlin, pp. 289-408.
Blackwelder, R. E. 1944. Checklist of the Coleopterous insects of Mexico, Central America, the

West Indies, and South America. Part 1 . Bull. U. S. N. M. No. 1 85, pp. i-xii + 1 - 1 88.
Navarrete-Heredia, J.L. 1996. Coleopteros micetocolos de Basidiomycetes de San Jose de los

Laureles, Morelos, Mexico. Tesis de Maestria (Biologia Animal), Fac. de Ciencias, UNAM.
Ruiz-Lizarraga, G. 1993. Contribucion al conocimiento de los Staphylinidae (Coleoptera)

necrofilos de Acahuizotla, Guerrero. Tesis profesional, Fac. de Ciencias, UNAM.
Scheerpeltz, O. 1972. Eine neue Art der Gattung Trigonopselaphus Gemminger-Harold, nebst

einer Dichotomik der jetzt zu dieser Gattung gehorigen Arten, Bemerkungen iiber die aus

dieser Gattung auszuscheidenden Arten und neue, zum Teil auf diesen .. Arten gegriindete

Gattungen. Mitt. Miinchner Entomol. Gesell. 62: 31-48.
Sharp, D. 1884. Fam. Staphylinidae, pp. 313-392. In: Biologia Centrali-Americana. Insecta.

Coleoptera. Vol. 1 (2). Taylor & Francis, London.

Vol. 109, No. 4, September & October, 1998 233

VOLTINISM IN NEOGERR1S HESIONE

(HETEROPTERA: GERRIDAE)

IN SOUTHERN ILLINOIS 1

Steven J. Taylor 2 , J. E. McPherson 3

ABSTRACT: Voltinism in Neogerris hesione was studied in southern Illinois during 1989 and
1990. This species apparently overwintered as eggs, which hatched during late April and early
May. First and second instars were found from late April through late September, third instars
from early May through late September, fourth instars from mid-May through late September,
fifth instars from mid-May through the third week of September, and adults from late May through
early November. The sequences of peaks for instars and adults strongly indicates that this spe-
cies is trivoltine in southern Illinois.

The waterstrider Neogerris hesione (Kirkaldy) occurs from New York,
Michigan, and Nebraska south to Florida and Texas; it also occurs in Cuba,
Panama (Smith 1988), and Central America (Nieser 1994). Smith (1988) re-
ported it from Alaska, but, judging from Andersen's (1982) world distribution
map of the genus (p. 366), this location probably is invalid. It is found through-
out the southern two thirds of Illinois (Taylor 1996) and, because it has been
collected in central and south central Wisconsin (Hilsenhoff 1986), probably
occurs in northern Illinois.

Little has been reported on this insect's life history. It occurs in a variety
of habitats but seems to prefer ponds, lakes, and pools of streams (Taylor 1 996).
It has been collected in September in Wisconsin (Hilsenhoff 1986) and in Oc-
tober in Missouri (Froeschner 1 962). Herring ( 1 950), in northern Florida, col-
lected adults from April through December, excluding June, and nymphs in
April and from September through November. Kittle (1980), in Arkansas,
collected adults from May to November; and Wilson (1958), in Mississippi,
collected nymphs as early as 1 April and as late as 15 November. Kittle (1977)
reported that of 60 mating pairs collected in northwest Arkansas, 1 pair (2%)
was taken in August, and 18 (30%) and 41 pairs (68%) were taken in Septem-
ber and October, respectively. Wilson (1958) observed mating as late as 15
November. Osborn and Drake (1915) believed this species overwinters as
eggs.

Drake and Harris (1934) stated that macropterous adults were rare in the
North and common in the South. This is supported by Hilsenhoff ( 1 986), who
reported apterous adults as common in Wisconsin (macropterous adults pre-

1 Received November 17, 1997. Accepted January 12, 1998.

2 Center for Biodiversity, Illinois Natural History Survey, 607 E. Peabody Dr., Champaign, IL
6 1820 US A

-^Department of Zoology, Southern Illinois University at Carbondale, Carbondale, IL 62901-
6501 USA.

ENT. NEWS 109(4) 233-239, September & October, 1998

234 ENTOMOLOGICAL NEWS

sumably being rare); and by Herring (1950), who reported macropterous adults
as common in northern Florida. Kittle, however, stated that of 1 36 adults from
northwest Arkansas (1977) and 182 adults from Arkansas (1980), 92 and 90%,
respectively, were apterous, the remainder macropterous. Similarly, Wilson
(1958) found that in Mississippi, about 95% of the adults were apterous, the
remainder macropterous.

During 1 989 and 1 990, we studied voltinism in a population of this species
at President's Pond on the Southern Illinois University at Carbondale campus,
Jackson County, Illinois (see Taylor [1996] for detailed description of pond).
The roughly triangular 0.29 hectare (0.71 acre) pond is connected at its north-
ern end to an adjacent lake by a narrow channel (approximately 2-5 m wide, 2
m deep). Water depth along the eastern margin (where the present study was
conducted) increased sharply between 1 and 2 m from shore and commonly
exceeded 2 m at 2.5 m from shore.

Floating, emergent, and shoreline vegetation associated with the pond was
diverse (Taylor 1996). The western margin was bordered by a narrow, dense
band of cattails (Typha angustifolia L.). The southern border consisted of a
riprap dam covered with soil and crossed by a paved road. The eastern margin
was bordered by overhanging trees and other vegetation. During the summer,
the pond filled with a dense growth of aquatic vascular plants and filamentous
algae. Near the shoreline and wherever aquatic plants reached the water sur-
face, duckweeds built up into dense mats. The duckweeds (i.e., Lemna minor
L., Spirodela polyrhiza (L.) Scheiden, and Wolffia papulifera Thompson) tended
to move around the pond because of air currents unless they were partially
anchored in the underlying aquatic vegetation.

This paper presents information on voltinism in N. hesione, including times
of occurrence of the adults and nymphal instars.

MATERIALS AND METHODS

Samples were collected weekly from 1 8 March to 25 November 1989, and
biweekly from 1 1 February to 2 December 1990. Sampling was limited to an
area along the eastern shore because (1) the cattails along the western shore-
line prevented use of the quadrat sampler (see below); (2) the riprap shoreline
of the southern border was unnatural and, often, disturbed by fishermen; and
(3) the water surface along the eastern shore, which was a mosaic of open
water, duckweeds, and emergent stems, supported a diverse gerromorphan
fauna.

Four 60 m transects were made parallel to a relatively uniform section of
the eastern margin at 0, 0.5, 1.0, and 1.5 m from the shoreline. Each sample
was collected with a floating quadrat sampler (0.25 x 0.25 x 0.05 m), with four
replicates placed randomly along each transect; the resulting 1 6 quadrat samples
were pooled, providing a broad sampling of the habitat. Prior to each sample,

Vol. 109, No. 4, September & October, 1998

235

the collector (SJT) stood for approximately three minutes to allow the insects
to acclimate to the disturbance; then, the sampler was placed on the surface of
the water. Specimens were removed with a fine mesh nylon net, preserved in
alcohol, and sorted in the laboratory. Nymphal instars were distinguished by
size of the dorsal sclerites on the thorax and abdomen, the overall degree of
sclerotization, and, to a lesser extent, body size. Fifth instars and adults were
distinguished from younger instars and from each other by the extent of devel-
opment of the external genital ia and by presence of wings in macropterous
adults.

so-

Figure 1. Percent of individuals in each stage per sample of N. hesione collected at President's
Pond, Southern Illinois University at Carbondale campus, Jackson County, during 1989. Begin-
ning and end points of each shaded area represent sample dates preceeding and following collec-
tion of specimens, respectively.

236

ENTOMOLOGICAL NEWS

RESULTS AND DISCUSSION

Neogerris hesione, which apparently overwintered as eggs in southern Illi-
nois, was active from late April through early November (Figs. 1 -4). First and
second instars were found from late April through late September, third instars
from early May through late September, fourth instars from mid-May through
late September, fifth instars from mid-May through the third week of Septem-
ber, and adults from late May through early November.

Figure 2. Percent in each sample of total individuals of same stage of N. hesione collected at
President's Pond, Southern Illinois University at Carbondale campus, Jackson County, during
1989. Beginning and end points of each shaded area represent sample dates preceeding and
following collection of specimens, respectively.

Vol. 109, No. 4, September & October, 1998

237

This species apparently is trivoltine in southern Illinois. Most fifth instars
of the first generation became adults between late May and mid-June. Most
first instars of the second generation were found in mid- to late June and be-
came adults during mid- to late July. The third generation was less clearly
distinguishable than the first and second, but fifth instars of this generation
apparently appeared in August and September and reached adults beginning in
late August - early September. Abundance of adults was greatest in the third
generation (Figs. 2, 4). These third generation adults, then, laid the overwin-
tering eggs.

50-

Figure 3. Percent of individuals in each stage per sample of N. hesione collected at President's
Pond, Southern Illinois University at Carbondale campus, Jackson County, during 1990. Begin-
ning and end points of each shaded area represent sample dates preceeding and following collec-
tion of specimens, respectively.

238

ENTOMOLOGICAL NEWS

Most individuals were apterous. Of 648 adults collected during this study,
only three (0.46%) were macropterous. All three were males collected in 1989,
1 in August and 2 in September. Also, three fifth instars (2Cf C? , 19) collected
in early June 1989 had well developed wing pads and, presumably, would
have become macropterous adults.

Riding behavior, typically associated with copulation and pre- and post-
copulatory mate-guarding in Gerridae (see literature review by Spence and
Andersen [1994]), was observed shortly after the first appearance of adults in
late May and again from mid-July through early November.

Figure 4. Percent in each sample of total individuals of same stage of N. hesione collected at
President's Pond, Southern Illinois University at Carbondale campus, Jackson County, during
1990. Beginning and end points of each shaded area represent sample dates preceeding and
following collection of specimens, respectively.

Vol. 109, No. 4, September & October, 1998 239

The life history observations of Osborn and Drake (1915), Wilson (1958),
and Kittle (1977) generally are supported by our data (Figs. 1-4). Differences
in the times of occurrence of nymphs and adults reported here and by Herring
( 1 950) and Wilson ( 1 958) probably reflect the geographic locations of the three
studies (i.e., southern Illinois, northern Florida, and Mississippi). Finally, the
percentages of macropterous and apterous adults reported here are similar to
those reported by Kittle (1977, 1980) and Wilson (1958).

ACKNOWLEDGMENTS

We thank the following for their critical review of this manuscript: R. A. Brandon, J. A.
Beatty, B. M. Burr, Department of Zoology; D. Ugent, Department of Plant Biology, Southern
Illinois University at Carbondale; and D. W. Webb and M. J. Wetzel, Center for Biodiversity,
Illinois Natural History Survey.

LITERATURE CITED

Andersen, N. M. 1982. The semiaquatic bugs (Hemiptera, Gerromorpha). Phylogeny, adapta-
tions, biogeography and classification. Entomonograph Vol. 3. Scandinavian Science Press
Ltd., Klampenborg, Denmark. 455 pp.

Drake, C. J. and H. M. Harris. 1934. III. The Gerrinae of the Western Hemisphere (Hemi-
ptera). Ann. Carnegie Mus. 23:179-241.

Froeschner, R. C. 1962. Contributions to a synopsis of the Hemiptera of Missouri, Part V.
Hydrometridae, Gerridae, Veliidae, Saldidae, Ochteridae, Gelastocoridae, Naucoridae,
Belostomatidae, Nepidae, Notonectidae, Pleidae, Corixidae. Am. Midi. Nat. 67:208-240.

Herring, J. L. 1 950. The aquatic and semiaquatic Hemiptera of northern Florida. Part I : Gerridae.
Fla. Entomol. 33:23-32.

Hilsenhoff, W. L. 1986. Semiaquatic Hemiptera of Wisconsin. Gt. Lakes Entomol. 19:7-19.

Kittle, P. D. 1 977. The biology of water striders (Hemiptera: Gerridae) in northwest Arkansas.
Am. Midi. Nat. 97:400-410.

Kittle, P. D. 1980. The water striders (Hemiptera: Gerridae) of Arkansas. Ark. Acad. Sci. Proc.
34:68-71.

Nieser, N. 1994. A new species and a new status mNeogerris Matsumura(Heteroptera: Gerridae)
with a key to American species. Storkia 3:27-37.

Osborn, H. and C. J. Drake. 1915. Additions and notes on the Hemiptera - Heteroptera of
Ohio. Ohio Nat. 15:501-508.

Smith, C. L. 1 988. Family Gerridae Leach, 1 8 1 5, pp. 1 40- 1 5 1 . In T. J. Henry and R. C. Froeschner
(eds.). Catalog of the Heteroptera or true bugs, of Canada and the continental United States.
E. J. Brill, New York. 958 pp.

Spence, J. R. and N. M. Andersen. 1994. Biology of water striders: Interactions between
systematics and ecology. Ann. Rev. Entomol. 39:101-128.

Taylor, S. J. 1996. Habitat preferences, species assemblages, and resource partitioning by
Gerromorpha (Insecta: Heteroptera) in southern Illinois, with a faunal list and keys to species
of the state. Ph.D. Dissertation, South. III. Univ. at Carbondale. xviii + 345 pp.

Wilson, C. A. 1958. Aquatic and semiaquatic Hemiptera of Mississippi. Tulane Stud. Zool.
6:115-170.

240 ENTOMOLOGICAL NEWS

A NEW SPECIES OF PARUROCTONUS
(SCORPIONES: VAEJOVIDAE) FROM
BIG BEND NATIONAL PARK, TEXAS 1

W. David Sissom^ , Richard N. Henson^

ABSTRACT. A new species of the baergi microgroup of the genus Paruroclonus is described
from sand dunes along the Rio Grande River in Boquillas Canyon, Big Bend National Park,
Texas. Variation in setal characteristics is summarized.

As a result of taxonomic surveys of scorpions in Big Bend National Park,
one of us (RNH) collected specimens of a Paruroctonus Werner species in the
short stretch of sand dune habitat at the end of the Boquillas Canyon Trail. The
specimens were very similar to P. utahensis (Williams), which has a wide dis-
tribution from southern Utah and northwestern Arizona, through much of New
Mexico to the northern portions of the Trans-Pecos in Texas and the Samalayuca
Dunes in extreme northern Chihuahua. The site in Big Bend represents a con-
siderable disjunction and, after study, it was ascertained that the specimens
represented a new species in the baergi microgroup that differs dramatically
from P. utahensis in several important characters. It is the purpose here to
describe this new species.

Accessible sand dune communities are found in only a few locations in the
Big Bend region and, thus far, we have found this species only in Boquillas
Canyon. Sandy habitats along the Rio Grande in the Big Bend Ranch State
Park (e.g., Arenosa, Grassy Banks, Madera Canyon) have been sampled, but
this species was not encountered. Dune systems are much more extensive on
the Mexican side of the Rio Grande, however, and the new species would be
expected to occur there.

Paruroctonus boquillas, NEW SPECIES

(Figs. 1-8)

Type Data. Adult male holotype from sand dunes in Boquillas Canyon
(N29. 1 2.02: W 1 02.55. 1 1 ), Big Bend National Park, Brewster Co., Texas, USA
on 20 May 1996 (R. N. Henson, et al.); deposited in the United States National
Museum (Smithsonian Institution), Washington, D. C.

Distribution. Known only from the type locality.

Etymology. The specific epithet is derived from the type locality,
Boquillas Canyon, and is used as a noun in apposition.

1 Received December 6, 1 997. Accepted January 12, 1998.

2 Dept. of Life, Earth, and Environmental Sciences, West Texas A & M University, WTAMU
Box 808, Canyon, Texas 79016.

3 Department of Biology, Appalachian State University, Boone, NC 28608.

ENT. NEWS 109(4) 240-246, September & October, 1998

Vol. 109, No. 4, September & October, 1998 241

Diagnosis. -- Haradon (1983, 1984a, 19845, 1985) established a number
of setal characters that he determined to be of great significance in species
recognition in the genus Paruroctonus. Utilizing these and other characters
listed below, P. boquillas clearly belongs in the baergi microgroup, a subgroup
of the boreus infragroup. The baergi microgroup was defined (Haradon 1 984a)
as lacking the mid-retrosuperior seta (mrs) on the basitarsus (tarsomere I) of
leg II; having pectinal tooth counts usually in excess of 22 in males and 18 in
females; having the pedipalp chela with well developed, granulose carinae in
adults of both sexes; and having 25-44 primary row denticles on the cutting
margin of the pedipalp chela fixed finger and 35-57 such denticles on the mov-
able finger (excluding the basal row in both cases).

In bearing four long retrosuperior setae of equal length on the telotarsus (=
tarsomere II) of leg III and an mrs seta on the basitarsus of the same leg, P.
boquillas is most similar to P. utahensis and P. baergi (Williams & Hadley). It
differs from both P. baergi and P. utahensis in having higher setal counts on
most metasomal carinae of segments I-IV (Table 1 summarizes counts for P.
boquillas) and four external medial setae on the pedipalp femur. Because it
has only one retromedial seta on telotarsus III, typically 2 inframedial setae on
the pedipalp femur, lower pectinal tooth counts in males (23-28 vs. 29-37) and
exhibits slightly more distinct scalloping in the chela fingers (in both sexes), it
is readily distinguished from P. utahensis. From P. baergi it can be further
distinguished by having four internal chelal macrosetae (not two) and more
subtle scalloping in the chela fingers of the male.

Paruroctonus boquillas is very similar to P. arenicola Haradon in terms of
its setation patterns. It may be distinguished from that species by having four
long retrosuperior setae on telotarsus III (rather than three, or three long and
one short setae) and by having fewer setae on the inner aspect of the pedipalp
chela fingers. From P. arenicola arenicola Haradon of the Amargosa Desert in
Nevada, it further differs by having 1, 1, 1-2, 2 setae on the dorsolateral meta-
somal carinae of I-IV (rather than 1, 3, 3, 3-4) and by having the chela fingers
yellowish (rather than orange, contrasting to color of palm). From P. arenicola
nudipes Haradon of the eastern Mojave Desert in California, it differs by hav-
ing the mrs seta present on telotarsus III and by the setal pattern of the dorso-
lateral carinae of I-IV (in P. a. nudipes there are 0, 1 , 1,2 pairs of setae).

Description: Based on the male holotype. Coloration: Dorsum, metasoma, pedipalps, and
legs light yellow; venter slightly paler; pectines whitish; dentition of pedipalp chela fingers,
cheliceral fingers, and tip of telson dark reddish brown to reddish black.

Prosoma. Anterior margin of carapace straight, set with four pairs of reddish setae. Entire
carapacial surface densely coarsely granular.

Mesosoma. Tergites I-VI: median carina on II-VI present on post-tergite, granular. Tergite
VII: median carina vestigial, granular; submedian and lateral carinae strong, irregularly serrated.
Pectinal teeth numbering 24-24. Stemites III-VI densely minutely granular laterally; smooth,
lustrous, and punctate medially. Sternite VII with pair of moderate crenulate lateral carinae.

242 ENTOMOLOGICAL NEWS

Metasoma (Fig. 1). Segment I slightly longer than wide, III 1.43 times longer than wide;
segment V 3.21 times longer than wide. Segments 1-IV: Dorsolateral and lateral supramedian
carinae on I-IV strong, serrate. Lateral inframedian carinae on I complete, strong, irregularly
serrate; on II represented by four serrated posterior granules; on III represented by three serrated
posterior granules; on IV absent. Ventrolateral carinae on I-II strong, smooth; on III-IV strong,
smooth with a few posterior serrations. Ventral submedian carinae on I weak, smooth; II moder-
ate, smooth; on III strong, smooth; on IV strong, smooth with posterior serrations. Carinal setation
of segments I-IV (L/R): dorsolaterals 0/1,1/1,1/1,2/2; lateral supramedians 1/0,2/2,2/3,3/3; lat-
eral inframedians 2/2,0/0,0/0,0/0; ventrolaterals 2/2,4/3,4/4, 5/5; ventral submedians 3/3,4/4,4/
4,4/5. Segment V: Dorsolateral carinae strong, feebly crenulate; lateromedian carinae present on
anterior one-half, strong, irregularly serrate; ventrolateral and ventromedian carinae strong, ser-
rate. Carinal setation of segment V: dorsolaterals 4/4; lateromedians 3/2; ventrolaterals 10/11.

Telson (Fig. 1 ). Elongate; dorsal surface flattened, smooth; ventral surface essentially smooth,
with 1 2 pairs of larger setae; aculeus weakly curved.

Chelicera (Figs. 2-3). Ventral aspect of cheliceral fixed finger with smooth carina extending
length of finger. Ventral margin of movable finger mostly smooth, with one small rounded den-
ticle near the base.

Pedipalp. Trichobothrial pattern Type C, orthobothriotaxic (Vachon 1974). Femur: Dorso-
internal, ventrointernal, and dorsoextemal carinae, strong, serrate to serratocrenulate; ventro-
external carina represented by a few spinoid denticles. Internal face with irregularly-spaced large
conical granules; three inframedial setae along ventrointernal carina (Fig. 4). External face with
four medial setae (Fig. 5).

Patella: All carinae strong, serratocrenulate. Internal face with strong basal tubercle and
serrated oblique internal carina. Internal face with two supramedial setae along dorsointernal
carina and two inframedial setae along ventrointernal carina.

Chela. Octocarinate, with all keels strong, granulose. Dentate margin of fixed finger with
primary granular row divided into six subrows by five larger granules; granules of subrows num-
bering 3, 6, 8, 9, 9, 22; six inner accessory granules. Dentate margin of movable finger divided
into seven subrows by six larger granules; granules of subrows numbering 1,5,8, 10, 11, 1 3, 9;
seven inner accessory granules. Scalloping of dentate margin of chela fingers moderate (Fig. 6).
Chela length/width ratio - 3.03. Fixed finger length/carapace length ratio - 0.77; movable finger
length/chela width ratio = 1 .87. Internal carinae of palm with 4 macrosetae; fixed finger with 1
macroseta; movable finger with 2 macrosetae.

Legs. Basitarsus II without mid-retrosuperior (mrs.) seta; basitarsus III with mrs. Telotarsus
III with four long retrosuperior setae and one long retromedial seta (Fig. 7).

Measurements of Male Holotype (in mm): total L, 49.3; carapace L, 5.6; mesosoma L (I-
VII), 11.8; metasoma L (I-V), 24.8; telson L, 7.1. Metasoma: I L/W, 3.3/3.1; II L/W, 4.1/3.1; III
L/W, 4.3/3.0; IV L/W, 5.4/2.7; V L/W, 7.7/2.4. Telson: vesicle L/W/D, 4.2/1.8/1.9; aculeus L,
2.9. Pedipalps: femur L/W, 4.8/1.5; patella L/W, 5.1/2.1; chela L/W/D, 9.4/ 3.1/4.1; fixed finger
L, 4.3; movable finger L, 5.8.

Measurements of Female Paratype (in mm): total L, 54.2; carapace L, 6.6; mesosoma L (I-
VII), 14.7; metasoma L (I-V), 25.1; telson L, 7.8. Metasoma: I L/W, 3.4/3.3; II L/W, 4.0/3.3; III
L/W, 4.3/3.2; IV L/W, 5.4/2.9; V L/W, 8.0/2.6. Telson: vesicle L/W/D, 4.5/2.6/2.2; aculeus L,
3.3. Pedipalps: femur L/W, 5.3/1.8; patella L/W, 5.7/2.5; chela L/W/D, 10.4/3.1/4.0; fixed finger
L, 4.8; movable finger L, 6.5.

Variation. Females differ from males as follows: (1) the carinae of the
pedipalp chelae are somewhat weaker, (2) the scalloping of the pedipalp chela
fingers is weaker (Fig. 8); (3) body size is somewhat greater; (4) the metasoma
is not quite as elongate; and (5) the pectinal tooth counts are lower (see be-
low).

Vol. 109, No. 4, September & October, 1998

243

Figures 1-8. Morphology of Paruroctonus boquillas, new species; illustrations are of the
holotype male, unless otherwise stated. 1, left lateral aspect of metasoma and telson, showing
carinae and setal pattern; 2, right chelicera, dorsal view; 3, right chelicera, ventral view; 4, inter-
nal aspect of right pedipalp femur, showing setal pattern (im = inframedial setae); 5, external
aspect of right femur, showing setal pattern (em = external medial setae); 6, external aspect of
right pedipalp chela fingers, showing degree of scalloping; 7, retrolateral aspect of right telotarsus
III, showing setal pattern (rss = retrosuperior setae; rms = retromedial seta); 8, pedipalp chela
fingers of female paratype

244 ENTOMOLOGICAL NEWS

Juveniles possess fairly narrow pedipalp chelae with weakly granulose
carinae and no scalloping of the chela fingers. Their setae tend to be finer than
in adults, a significant factor in getting counts from the pedipalp chela fingers.
The seta of the fixed finger and the distal one on the movable finger tend to be
small and fine even in adults, but distinctly pigmented. In subadults and smaller
juveniles, the distal diagnostic setae on the fingers tend to be microchaetes and
are not included in the counts described below. Other setae in subadults were
not problematic, easily determined as macrosetae.

Variation in setal counts is of critical importance to the taxonomy of
Paruroctonus. Setal counts were taken from 25 specimens, and results are given
in Tables 1 and 2. Modal counts for the various setal patterns are as follows:
Metasomal setae of I-IV: dorsolaterals 1, 1, 1-2, 2; lateral supramedians 0, 2,
3, 3; lateral inframedians 2, 0, 0, 0; ventrolaterals 2, 3, 4, 5; ventral submedians
3, 4, 4, 5. Metasomal setae of V: dorsolaterals 4; lateromedians 3-4;
ventrolaterals 10. Pedipalpal setae: femoral inframedians 2; femoral external
medians 4; chela palm internals 4; fixed finger internals 1 ; movable finger
internals 2. As the tables demonstrate, there was considerable variation in many
of these counts. Setal counts of the retrolateral aspect of tarsi III appear to be
invariable or virtually so, however, with all specimens examined having 4
retrosuperior setae and 1 retromedian seta.

In males, pectinal tooth counts ranged from 23 to 28, with the following
distribution: there were 3 combs with 23 teeth, 7 combs with 24 teeth, 17
combs with 25 teeth, 19 combs with 26 teeth, 4 combs with 27 teeth, and 1
comb with 28 teeth. In females, counts ranged from 1 7 to 23 teeth, distributed
as follows: there were 3 combs with 17 teeth, 20 combs with 18 teeth, 24
combs with 19 teeth, 15 combs with 20 teeth, 1 comb with 21 teeth, 1 comb
with 22 teeth, and 1 comb with 23 teeth.

Specimens Examined (Holotype and Paratypes). - USA: TEXAS: BREWSTER CO.: Big
Bend National Park, sand dunes in Boquillas Canyon (N29. 12.02: W 102.55. 11), 23 May 1992
(R. Henson, T. & J. Weseman, R. Soeder), 1 subadult male, 6 subadult females (USNM); 7 June
1992, 6 subadult males, 1 juv male, 2 subadult females, 1 juv female (USNM); 25 May 1994 (R.
N. Henson, J. Hosier, H. Husted, E. Kaiser), 6 subadult males, 1 female, 3 subadult females
(USNM), 6 subadult males, 1 female, 3 subadult females (CAS); 20 May 1996 (R. N. Henson, et
al.), 1 holotype male, 3 females (USNM), 1 male, 2 females, 2 subadult females (RNH), 2 sub-
adult males, 2 females (AMNH), 1 male, 1 subadult male, 2 females, 2 subadult females (WDS).
Depositories for materials are as follows: AMNH, American Museum of Natural History, New
York; CAS, California Academy of Sciences, San Francisco; RNH, collection of R. N. Henson;
USNM, United States National Museum, Washington, D.C.; WDS, collection of W. D. Sissom.

Vol. 109, No. 4, September & October, 1998 245

Comments. The junior author was stung on the tip of the middle finger
while making one of the collections. The pain was very intense and "hot" for a
period of about 10 minutes, which is not unlike many scorpion stings. How-
ever, the effects of this sting were somewhat prolonged, with edema develop-
ing in the finger that lasted for 3 days. Accompanying this was some stiffness
and a throbbing ache. All symptoms disappeared by the fourth day.

Table 1. Variation in setation of the metasomal carinae of segments I-IV in 25 specimens of P.
boquillas. DL = dorsolateral carinae; LSM = lateral supramedian carinae; LIM = lateral
inframedian carinae; VL = ventrolateral carinae; VSM = ventral submedian carinae.

No.
Setae

DL
II III IV

LSM LIM
I II III IV I II III IV

VL
I II III IV

VSM
I II III IV

15 23 24 25

19 13

10 2 1

4 12 25

22 7 24

3 18 23 1

2 15 8

10 17 6

5 21 19 5

4 6 16

Table 2. Variation in setal counts for metasomal segment V and the pedipalps (Fed.) in 25 speci-
mens of P. boquillas. DL = dorsolaterals; LM = lateromedians; VL = ventrolaterals; Int. IM =
internal inframedians; Ext. Med. = external medials; Int. Palm = internals of chela palm; Fix.
Fing. = internals of fixed finger; Mov. Fing. = internals of movable finger.

No. Metasomal V Fed. Femur Fed. Chela Fed. Chela Fed. Chela

Setae DL LM VL Int. IM Ext. Med. Int. Palm Fix. Fing. Mov. Fing.

16 9

1 15

9 1 3

4 20

24 20

5 5

246 ENTOMOLOGICAL NEWS

REVISED KEY TO THE PARUROCTONUS BAERGI MICROGROUP

Modified from Haradon (1984a)

1 . Telotarsus III with two retrosuperior setae P. marksi

Telotarsus III with three or four retrosuperior setae 2

2. Telotarsus III with three long, and possibly one shorter subdistal, retrosuperior setae

P. arenicola ... 3

Telotarsus III with four long retrosuperior setae 4

3. Basitarsus III with mrs seta present; paired dorsolateral metasomal setae 1 , 2, 2-3, 3-4

P. arenicola arenicola

Basitarsus III without mrs seta; paired dorsolateral metasomal setae 0, 1 , 1,2

P. arenicola nudipes

4. Pedipalp femur with four external medial setae; with 1 , 1 , 1-2,2 pairs of setae on

dorsolateral carinae of metasoma I-I V; with 0, 2, 2, 3 pairs of setae on the lateral
supramedians; with 2 pairs of setae on lateral inframedians of metasomal segment I;

with 2, 3, 4, 5 pairs of setae on the ventrolaterals of metasoma I-IV P. boquillas

Pedipalp femur with two or three external medial setae; with 0, 1 , 1,2 pairs of setae
on dorsolateral carinae of metasoma I-IV; with 0, 1 , 1,2 pairs of setae on the lateral
supramedians; with 1 pair of setae on lateral inframedians of metasomal segment I;
with 2, 3, 3, 4 pairs of setae on the ventrolateral carinae of I-IV. 5

5. Telotarsus III with one retromedial seta; closed pedipalp fingers in adult male form

wide proximal gap P. baergi

Telotarsus III with two retromedial setae; closed pedipalp fingers in adult male

form narrow proximal gap P. utahensis

ACKNOWLEDGMENTS

We would like to thank Mike Fleming, Resource Management Specialist of Big Bend
National Park, and the National Park Service for granting permission to conduct research in the
park (Permit #BIBE-96-008). We are also grateful to Texas Parks & Wildlife, and in particular
David Riskind (Director of the Natural Resources Program) and Luis Armendariz (Superinten-
dent of BBRSP), for permission to collect in Big Bend Ranch State Park (State Park Scientific
Study Permit No. 10-96). We are grateful to the following individuals for participating in the
field work: K. Adcock, A. Baldwin, H. Bowers, J. Hosier, H. Husted, E. Kaiser, M. Riddle, R.
Soeder, J. and T. Weseman, and A. Zamba. Kari J. Me West kindly reviewed the manuscript at our
request.

LITERATURE CITED

Haradon, R.M. 1983. Smeringurus, a new subgenus of Paruroctonus Werner (Scorpiones,
Vaejovidae). J. Arachnol., 11: 251-270.

Haradon, R.M. 1984a. New and redefined species belonging to the Paruroctonus baergi group
(Scorpiones, Vaejovidae). J. Arachnol., 12:205-221.

Haradon, R.M. 1984b. New and redefined species belonging to the Paruroctonus borregoensis
group (Scorpiones, Vaejovidae). J. Arachnol., 12:317-339.

Haradon, R.M. 1985. New groups and species belonging to the nominate subgenus Paruroctonus
(Scorpiones, Vaejovidae). J. Arachnol., 13: 19-42.

Sissom, W. D. and O. F. Francke. 1 98 1 . Scorpions of the genus Paruroctonus from New Mexico
and Texas (Scorpiones, Vaejovidae). J. Arachnol., 9: 93-108.

Vachon, M. 1974. Etude des caracteres utilises pour classer les families et les genres de scor-
pions. 1. La trichobothriotaxie en Arachnologie. Sigles trichobothriaux et types de tricho-
bothriotaxie chez les Scorpions. Bull. Mus. Nat. Hist. Nat. Paris, ser. 3, No. 140 (Zool. 104),
pp. 857-958.

Vol. 109, No. 4, September & October, 1998 247

COMPARISON OF SAND NESTING WASPS

(HYMENOPTERA) FROM TWO PINE BARRENS IN

UPSTATE NEW YORK 1

Frank E. Kurczewski^

ABSTRACT: Collections of sand inhabiting wasps from the Fort Drum Military Reservation and
the Rome sand plains indicate a common fauna with pine inhabited sandy areas in the nearby
Black River Valley and elsewhere in upstate New York. There were no pine barren indicator
species in the collections.

Pine barrens are an imperiled ecosystem in the northeastern United States.
In upstate New York, they comprise depauperate pine-oak woodlands growing
on fire swept or otherwise disturbed sandy soils. The nutrient impoverished,
water deficient, primarily acidic sandy soils supporting these woodlands his-
torically were often anthropogenically altered. The sparse interrupted canopy
is composed of white pine (Finns strobus L.) and/or pitch pine (P. rigida Mill.)
trees. A tall dense shrub layer dominated by scrub oak (Quercus ilicifolia
Wangenh.) or, sometimes, stunted white oak (Q. alba L.) grows beneath these
trees. A low shrub layer often composed of Ericaceae, forbs, and grasses grows
beneath the oaks and in interspersed openings (Schweitzer and Rawinski 1 988).

The species of sand nesting wasps that inhabit upstate New York pine bar-
rens is poorly known. Two pine barrens areas devoid of wasp collection records
are the Fort Drum Military Reservation in Jefferson County and the Rome
sand plains in Oneida County (Fig. 1). Fort Drum is situated on a site known
historically as the "pine plains." An abundance of white pine, some pitch pine
and white oak, and many deciduous tree species grew there on level, dry, sandy
soil in the 1 790s. These pine plains were located on droughty sands and loamy
fine sands north of the large bend in the Black River between the villages of
Great Bend and Deferiet. More than 10,000 ha of moderately well to exces-
sively drained sandy soils blanket this section of Fort Drum (USD A 1989).

The Rome sand plains contain approximately 1 ,200 ha of sandy soil. Half
this acreage is moderately well to excessively drained and suitable for psam-
mophilous wasp habitation (USDA 1997 pers. comm.). Although the Rome
sand plains contained extensive pine barren acreage at the turn of the last cen-
tury (Stephens [sic] and Barrus 1911), the predominant vegetation through
most of the millenium was a mesophytic deciduous-coniferous forest (Russell
1996).

The purpose of this paper is to: (1) present the results of solitary wasp

' Received September 11, 1997. Accepted November 30, 1997.

2 Environmental and Forest Biology, State University College of Environmental Science and
Forestry, Syracuse, New York 13210-2778.

ENT. NEWS 109(4) 247-25 1 , September & October, 1998

248 ENTOMOLOGICAL NEWS

collections from these two localities; (2) relate these results to collection records
for this group from other pine inhabited sandy localities in upstate New York;
and, (3) determine whether or not pine barrens were part of the ancestral veg-
etation landscape based upon wasp species.

METHODS

Eighteen trips were made to the Fort Drum Military Reservation during
August 2-October 18, 1996 and April 3-October 4, 1997 for the purpose of
collecting sand inhabiting wasps. Actual time spent collecting there totalled
70 hours. Collections were made on barren Plainfield sand.

Seven trips were made to the Rome sand plains on June 27, 1992, July 28,
1993, July 13, 1995, and June 2, July 26 and September 10 and 27, 1997 for
this purpose. Actual time spent collecting at this locality totalled 35 hours.
Collections were made on barren Windsor loamy fine sand.

RESULTS

One-hundred and nine species of Tiphiidae, Mutillidae, Scoliidae, Pompili-
dae, and Sphecidae were collected at Fort Drum (Table 1 ). Twenty-two (20.2%)
of these species were not sand nesters. Sixty-three species of Tiphiidae,
Scoliidae, Pompilidae, and Sphecidae were collected at the Rome sand plains
(Table 1). Only five (7.9%) of these species were not psammophilous. Sixty-
two species of Tiphiidae, Scoliidae, Pompilidae, and Sphecidae were common
to both localities. Forty-seven species of wasps found at Fort Drum were not
collected at Rome. Only one species collected at Rome, Tachytes validus
Cresson, was not found at Fort Drum.

Stictiella emarginata (Cresson), a species rarely found in upstate New York,
provisioned nests at Fort Drum with adult moths of the family Noctuidae (pers.
obs.). Philanthus albopilosus Cresson collected at Fort Drum was at the east-
ern extremity of its range (Evans 1975).

DISCUSSION

The species of sand inhabiting wasps from Fort Drum and the Rome sand
plains were similar to those from the Black River Valley in adjacent Lewis and
Herkimer Counties and other localities in upstate New York (Cayuga County:
Auburn, Sennett; Oswego County: Granby Center, Mallory Station, Selkirk
Shores State Park; St. Lawrence County: Parishville) (Kurczewski 1994;
Kurczewski and Acciavatti 1990; pers. obs.).

The absence of Tachysphex pechumani Krombein, a characteristic faunal
element in the New Jersey and lower Michigan pinelands, implies that the
New York State pine barren sites were anthropogenically induced. The over-
whelming dominance of pre-Euro-American settlement mesophytic decidu-

Vol. 109, No. 4, September & October, 1998

249

Parishville

Fort Drum
aumown

Fig. 1. Sand wasp collection sites in northcentral New York. + designates Fort Drum Military
Reservation and Rome sand plains, * localities from previous years.

250

ENTOMOLOGICAL NEWS

Table 1 . Species of solitary wasps collected and/or observed at the Fort Drum Military Reserva-
tion and Rome sand plains.

ROME

FORT

SAND

FORT

SAND

SPECIES

DRUM

PLAINS

SPECIES DRUM

PLAINS

TIPHIIDAE

Ammophila pictipennis Walsh

Tiphia sp.

Ammophila procera Dahlbom

Paratiphia sp.

Ammophila urnaria Dahlbom

Myzinum quinquecinctum (Fabricius)

Mimesa basirufa Packard

Methocha stygia (Say)

Mimesa cressonii Packard

Mi I / IHtl (v It* -InvH li

MUTILLIDAE

LJlOaOnlUS ft tint It Fititnil

(Krombein)

Timulla varans (Fabricius)
Pseudomethoca frigida (Smith)

X
X

*Pemphredon lethifer
(Shuckard)

SCOLI1DAE

*Passaloecus sp.

Campsomeris plumipes (Drury)

*Sligmus americanus Packard
Asiaia leuthstromi Ashmead

X
X

POMP1LIDAE

Liris argenluta (Beauvois)

*Priocnessus nebulosus (Dahlbom)

Tachytes obductus Fox

*Priocnenis (Priocnemissus) minoratu Banks

Tach\U's validus Cresson

Priocnemis (Priocnemis) cornica (Say)

Tachysphex acuius (Patton)

*Priocnemis {Priocnemis) germana (Cresson)

Tachysphex similis Rohwer

* Priocnemis (Priocnemis) scitula relicla Banks

Tachysphex larsatus (Say)

Calicurgus hvalinatus (Fabricius)

Tachysphex lerminatus (Smith)

*Dipogon papa go unomalus Dreisbach

Lyroda subita (Say)

*Dipogon sayi Banks

Plenoculus davisi Fox

*Auplopus architectus (Say)

Miscophus americanus Fox

*Auplopus mellipes variitarsatus (Dalla Torre)

Oxybelus bipunctatus Olivier

Evagetes crassicornis (Shuckard)

O.rybelus emarginalus Say

Evagetes hyacinlhinus (Cresson)

O.rybelus suhcornutus Cockerell

Evage tes parvus (Cresson)

Oxybelus subulatus Robertson

Episyron biguttatus (Fabricius)

Anacrabro ocellatus Packard

Episyron quinquenotatus (Say)

Lindenius buccadentis Mickel

* Anoplius (Lophopompilus) aelhiops (Cresson)

Lindenius columbianus (Kohl)

Anoplius (Lophopompilus) atrox (Dahlbom)

Crossocerus maculiclypeus (Fox)

* Anoplius (Lophopompilus) Carolina (Banks)

Crabro advena Smith

Anoplius (Arachnophroclonus) relativus (Fox)

Crabro argusinus R. Bohart

Anoplius (Arachnophroclonus) semirufus (Cresson) x

Crabro cribrellifer (Packard)

Anoplius (Pompilinus) cylindricus (Cresson)

Crabro latipes Smith

Anoplius (Pompilinus) marginatus (Say)

Crabro monticola (Packard)

Anoplius (Pompilinus) splendens (Dreisbach)

Alysson melleus Say

Anoplius (Pompilinus) subcylindricus (Banks)

Nysson daeckei Viereck

Anoplius (Pompilinus) tenebrosus (Cresson)

Ochleroptera bipunctuta (Say)

*Anoplius (Anoplius) illinoensis (Robertson)

Corytes canaliculatus Packard

* Anoplius (Anoplius) nigerrimus (Scopoli)

Goryles simillimus Smith

* Anoplius (Anoplius) ventralis (Banks)

Pseudoplisus phaleratus (Say)

"Anoplim (Anoplius) virginiensis (Cresson)

Sphecius speciosus (Drury)

Pompilus (Arachnospila) arclus Cresson

Bicyrtes quadrifasciata (Say)

Pompilus (Arachnospila) scelestus Cresson

Bicyrtes ventralis (Say)

*Pompilus (Anoplochares) apicalus Provancher

Microbembex monodonta (Say)

Aporinellus completus Banks

Bembix americana (Lepeletier)

"Ceropales maculata fraternu Smith

Bembix pallidipicla Smith

Sticticllo cffiumifliJtii (Crcsson)

SPHECIDAE

Philanthus albopilosus Cresson

*Chalybion californicum (Saussure)

Philanthus bilunatus Cresson

*Sceliphron caementarium (Drury)

Philanthus gibbosus (Fabricius)

Sphex ichneumoneus (Linnaeus)

Philanthus lepidus Cresson

Sphe x pensylvanicus Linnaeus

Philanthus politus Say

*Isodontia mexicana (Saussure)

Philanthus solivagus Say

Prionyx atratus (Lepeletier)

Philanthus ventilabris Fabricius

Podalonia luctuosa (Smith)

Aphilanthops frigidus (Smith )

Podalonia robusta (Cresson)

Cerceris clypeata Dahlbom

Eremnophila aureonolata (Cameron)

Cerceris fumipennis Say

Ammophila harli (Fernald)

Cerceris nigrescens Smith

Ammophila nigrkans Dahlbom

Cerceris robertsonii Fox

*Non-sand inhabiting species.

Vol. 1 09, No. 4, September & October, 1 998 25 1

ous and deciduous-coniferous forests on sandy soils in upstate New York
(Seischab 1990, 1992; Marks and Gardescu 1992) and the substantial annual
amount of evenly distributed precipitation in the region (USD A 1941 ; Dethier
1966; Garwood 1996) support this contention.

ACKNOWLEDGMENTS

Howard Evans, George Ferguson, Bert Finnamore, Arnold Menke, and Mark O'Brien aided
in the identification of some of the wasp species. Anne Johnson provided entry onto the Fort
Drum Military Reservation. Hugh Boyle, Tom Conklin, Bud Mayfield, and Ed Stanton collected
some of the wasps at Fort Drum and Rome. Tim McCabe and Rick Hoebeke permitted examina-
tion of collection records at the New York State Museum and Cornell University, respectively.

LITERATURE CITED

Dethier, B. E. 1966. Precipitation in New York State. Cornell Univ. Agric. Exp. Sta. Bull. 1009:

1-78.
Evans, H. E. 1975. Nesting behavior of Philanthus albopilosus with comparisons between two

widely separated populations. Ann. Entomol. Soc. Amer. 68: 888-892.
Garwood, A. N. (Ed.). 1996. Weather America. Toucan Valley Pub., Milpitas, Ca. 1412 pp.
Kurczewski, F. E. 1994. An aculeate wasp collecting trip through the Black River Valley of

upstate New York. Sphecos 28: 19-20.
Kurczewski, F. E. and R. E. Acciavatti. 1990. Late summer-fall solitary wasp fauna of central

New York (Hymenoptera: Tiphiidae, Pompilidae, Sphecidae). Great Lakes Entomol. 23: 57-

64.
Marks, P. L. and S. Gardescu. 1992. Vegetation of the Central Finger Lakes Region of New

York in the 1790s. Pp. 1-35 in Marks, P. L., S. Gardescu, and F. K. Seischab. Late eighteenth

century vegetation of central and western New York State on the basis of original land survey

records. N. Y. State Mus. Bull. 484.
Russell, E. W. B. 1996. Six thousand years of forest and fire history in the Rome Sand Plains.

Cent. N. Y. Chapt. TNC Report. 26 pp.
Schweitzer, D. F. and T. J. Rawinski. 1988. Element stewardship abstract for northeastern pitch

pine/scrub oak barrens. East. Herit. Task Force, TNC. 21 pp.
Seischab, F. K. 1990. Presettlement forests of the Phelps and Gorham Purchase in western New

York. Bull. Torrey Bot. Club 1 17: 27-38.
Seischab, F. K. 1992. Forests of the Holland Company in western New York, circa 1798. Pp. 35-

53 in Marks, P. L., S. Gardescu, and F. K. Seischab. Late eighteenth century vegetation of

central and western New York State on the basis of original land survey records. N. Y. State

Mus. Bull. 484.

Stephens [sic], J. W. and G. L. Barrus. 1911. Map of Oneida County showing forest types.
United States Department of Agriculture. 1941. Climates of the States. Pp. 749-1228 in Cli-
mate and Man. Yearbook of Agriculture. U. S. Govt. Print. Off., Washington, D. C.
United States Department of Agriculture. 1989. Soil survey of Jefferson County, New York.

Soil Cons. Serv. Coop. Cornell Univ. Agric. Exp. Sta. 376 pp.

PLEASE SEE ANNOUNCEMENT ON PAGE 276

252 ENTOMOLOGICAL NEWS

THE IDENTITY OF TACHYSPHEX ACUTUS

(HYMENOPTERA: SPHECIDAE),

AN UNSOLVED MYSTERY 1

Frank E. Kurczewski^

ABSTRACT: A reevaluation of the identities of Tachysphex acutus and T. similis is given based
upon the species' descriptions of Patton (1881), Fox ( 1 894), Rohwer ( 1 9 1 0), and Williams (1914)
and examination of museum specimens. Although T. similis may be a synonym of T. acutus, the
continued usage of the specific names, as presently known, is recommended.

The true identify of Tachysphex acutus (Patton) remains a mystery. Patton
(1881) described "Larra acuta" from three specimens collected at Waterbury,
Connecticut in August. He indicated that the species is related to T. terminates
(Smith), T. tarsatus (Say), and T. montanus (Cresson), which did nothing more
than designate it as a species of Tachysphex. Patton 's original description fits
several dozen all-black Nearctic species in the genus and is totally undiagnostic
at the species group level. In addition, his definition of the propodeal dorsum
and side as being "uniformly and finely granulated" does not correspond with
specimens of T. acutus as recognized today (Pulawski 1988). To complicate
matters the type specimens of this species, which were placed in the Boston
Museum of Natural History, were destroyed by fire around the turn of this
century. Although Kohl ( 1 885), Dalla Torre ( 1 897), Ashmead ( 1 899), H. Smith
( 1 908), and J. Smith ( 1 9 1 0) all mentioned T. acutus in their treatises on sphecid
wasps, none of these authors provided an adequate description of the species.

Fox (1894), in his treatment of the genus Tachysphex, first delineated diag-
nostic morphological characteristics for T. acutus. However, his description of
the species fits better what is currently being called T. similis Rohwer than T.
acutus, especially with regard to the least interocular distance (R. Bohart 1 962),
punctation of the head and thorax, and width of submarginal cells. Further-
more, Fox (1 894) listed T. acutus directly after T.fusus Fox, T. terminatus, and
T. apicalis Fox, all members of the terminatus species group. T. similis also
belongs to this group. Fox (1894) recorded Georgia and Florida as collection
"localities" for T. acutus but Pulawski (1988), who examined 184 specimens
of this species, found no Georgia locality and only a single, new northern Florida
locality (Alachua County: Gainesville) for what is now being called T. acutus.
The Georgia and Florida specimens listed in Fox's (1894) revision proved to
be T. similis. The following questions arise: Did Fox examine Patton 's type
specimens prior to his revision of the genus? If not, did he presume that what

1 Received September 11, 1997. Accepted February 8, 1998.

2 Environmental and Forest Biology, State University of New York College of Environmental
Science and Forestry, Syracuse, NY 13210-2778.

ENT. NEWS 109(4) 252-255, September & October, 1998

Vol. 109, No. 4, September & October, 1998 253

is now being called T. similis was Pattern's "Larra acuta"? A search for addi
tional information is in order.

In 1910 Rohwer described T. similis and T. similans from eastern Texas
specimens. The two species are clearly synonymous, with T. similis being the
first name put in print (G. Bohart 1 95 1 ). Rohwer (1910) provided no diagnos-
tic characteristics which would assist in separating these species from others
in the genus. Rohwer (191 1) also described T. bruesi from Milwaukee County,
Wisconsin, and this species has been synonymized with what is currently be-
ing called T. acutus (G. Bohart 1951 ). Rohwer was notorious for erecting spe-
cies names ad nauseam and, in the process, creating innumerable synonyms.
Therefore, one has to question whether T. similis is a good species or merely
another of Rohwer's synonyms, in this case of T. acutus, especially in light of
the above information and the fact that the type specimens of T. acutus were
destroyed by fire before Rohwer described T. similis.

Compounding this ambiguity is the fact that Williams (1914), who re-
described the species of Tachysphex occurring in Kansas, made no mention of
T. similis in his redescriptions yet the species was already described (Rohwer
1910) and is abundant in the eastern half of the state (pers. obs.). His represen-
tation of T. acutus fits the current conception of this species, but some of the
morphological characteristics he listed for T. sepulcralis Williams agree with
those of T. similis. [T. sepulcralis was designated a synonym of T. acutus by
Pulawski (1988)]. Williams (1914) recorded T. sepulcralis as being related to
T. apicalis which, like T. similis, would place it in the terminatus group. He
noted that specimens of T. acutus matched both Patton's (1881) and Fox's
(1894) descriptions of this species, although he admitted he had not seen the
type specimens. Williams (1914) remarked that there were at least two species
of Tachysphex in the series labeled "acMta" in the Academy of Natural Sci-
ences of Philadelphia collection! Although both Viereck (1916) and Mickel
(1917) delineated T. acutus morphologically in their definitions of species of
Tachysphex, neither worker mentioned T. similis or T. sepulcralis. Mickel (1917)
leaned heavily on Williams' (1914) redescription of T. acutus in his own defi-
nition. Robertson ( 1 928) recorded females of T. acutus taking nectar from nearly
a dozen species of flowering plants but did not mention either T. similis or T.
sepulcralis. Pulawski (1988) regarded these records as useless because of the
doubtful determination of the wasp species. The omission of T. similis from all
of these studies is noteworthy because it was described and was common in
collections.

Further confusing were several series of specimens that I examined, some
with associated handwritten notes, in trays labeled "T. acutus" in various mu-
seums. Ten specimens in the collection of the North Carolina Department of
Agriculture determined originally as T. acutus and T. maneei Banks by N. Banks
were examined by K.V. Krombein in 1950 and some of them proclaimed to be

254 ENTOMOLOGICAL NEWS

T. similis (D.L. Wray, pers. comm.). A tray of 10 males and 10 females from
Georgia in the Cornell University insect museum were identified as T. acutus
by J.C. Bradley in 1 923. These specimens were redetermined by me in 1963 as
T. similis and, in 1980, reidentified as T. similis by W.J. Pulawski. However,
there is a "legitimate" female of what is now being called T. acutus identified
as such in the same collection (Southampton, Mass.; July 1 4, 1 894; #429) with
a very old determination label: "This spcm. ident. as Tachysphex acutus
PattonQ"- A tray of "7^ acutus" at the U.S. National Museum in 1963 con-
tained specimens identified as both T. sepulcralis (= T. acutus) and T. similis
(Det. K.V. Krombein), both species having been determined earlier as T. acutus
by S.A. Rohwer. These determinations were made after Rohwer described T.
similis in 1910! There is a note in the tray in Rohwer 's handwriting (teste K.V.
Krombein) that the Florida specimen (T. similis) agrees with J.C. Bradley 's
manuscript notes on T. acutus and that (Phil) Rau's specimens (T. sepulcralis)
are an entirely different species. This note was made the year (1923) that Bra-
dley determined T. similis specimens in the Cornell University collection as T.
acutus. However, neither Bradley in 1963 nor I upon his death in 1975 could
locate these notes at Cornell University. If Bradley indeed studied Patton's
type specimens, then it would appear that T. similis must be a synonym of T.
acutus and the name T. bruesi should be resurrected to accommodate the speci-
mens now being called T. acutus. However, the correct interpretation will never
be known because Patton's type specimens were destroyed a century ago and
the hymenopterists involved in the name continuation or its discontinuity have
passed away.

What steps then should be taken to correct this misconception, if one ex-
ists? I propose that everything remain status quo. R.M. Bohart, H.E. Evans,
K.V. Krombein, A.S. Menke, W.J. Pulawski, and other prominent sphe-
cidologists all recognize T. similis as presently defined. This species was re-
described in detail by Pulawski ( 1 988) and has a moderate amount of literature
associated with its ecology and nesting behavior. Tachysphex acutus, as pres-
ently accepted, is, on the other hand, less well known, and has been the seat of
much past taxonomic controversy. It, too, has a sizeable scattering of literature
associated with its identification. Unfortunately, much of this literature prob-
ably pertains to what is now being called T. similis, but my paper serves to
bring this misapplication of information to light. Pulawski (1988) redescribed
T. acutus, as currently known, and, in the process, designated a neotype speci-
men. Taking everything into consideration, present usage should prevail over
chronology and S.A. Rohwer's probable error. I, therefore, recommend con-
tinued usage of the names T. acutus and T. similis as presently applied, with
my paper providing an interpretation of any misconstruances that occurred in
the early to mid-20th century.

Vol. 109, No. 4, September & October, 1998 255

ACKNOWLEDGMENTS

R.M. Bohart, University of California, Davis, K.V. Krombein, Smithsonian Institution, and
W.J. Pulawski, California Academy of Sciences, provided much assistance in identifying speci-
mens of Tachysphex acinus and T. similis. K.V. Krombein, L.L. Pechuman, Cornell University,
and D.L. Wray, North Carolina Department of Agriculture, permitted me to examine specimens
at their disposal and provided me with pertinent information on the two taxa. J.C. Bradley, Cornell
University, and E.C. Zimmerman, Honolulu, Hawaii answered some of my questions about the
location of T. acutus type specimens. J.C. Bradley interpreted Cornell University Lot and Sublet
listings for me. P.M. Carpenter and S. Shaw, Museum of Comparative Zoology, Harvard Univer-
sity, and B.R. Wiseman, Museum of Science, Boston aided in obtaining information about the
disposition of the T. acutus type specimens and the Boston Museum of Natural History fire,
respectively. R.M. Bohart, K.V. Krombein, A.S. Menke, and W.J. Pulawski reviewed earlier ver-
sions of the manuscript.

LITERATURE CITED

Ashmead, W.H. 1 899. Classification of the entomophilous wasps, or the superfamily Sphegoidea.
Canad. Entomol. 3: 145-155, 161-174, 212-225, 238-251, 291-300, 322-330, 345-357.

Bohart, G.E. 1951. Tribe Tachytini. Pp. 945-953 in C.F.W. Muesebeck, K.V. Krombein, and
H.K. Townes (Eds.). Hymenoptera of America North of Mexico. Synoptic Catalog. USDA
Agric. Monogr. 2: 1-1420.

Bohart, R.M. 1962. New species of black Tachysphex from North America (Hymenoptera,
Sphecidae). Proc. Biol. Soc. Wash. 75: 33-40.

Dalla Torre, C.G. DE. 1897. Catalogus Hymenopterorum hucusque descriptorum systematicus
et synonynimicus, Volumen VIII: Fossores (Sphegidae). Guilelmi Engelmann, Lipsiae. 749
pp.

Fox, W.J. 1 894 ( 1 893). The North American Larridae. Proc. Acad. Nat. Sci. Phila. 45: 467-55 1 .

Kohl, F.F. 1885 (1884). Die Gattungen und Arten der Larriden Autorum 'sic.' Verh. Zool. Bot.
Ges. Wien34: 171-267.

Mickel, C.E. 1 9 1 7. A synopsis of the Sphecoidea of Nebraska (Hymenoptera). Univ. Nebr. Stud.
17: 342-456.

Patton, W.H. 1881 (1880). List of North American Larridae. Proc. Boston Soc. Nat. Hist. 20:
385-397.

Pulawski, W.J. 1988. Revision of North American Tachysphex wasps including Central Ameri-
can and Caribbean species (Hymenoptera, Sphecidae). Mem. Calif. Acad. Sci. 10: 1-211.

Robertson, C. 1928. Flowers and Insects. Lists of visitors of four hundred and fifty-three flow-
ers. Privately published, Carlinville, 111. 221 pp.

Rohwer, S.A. 1910. Some wasps from New Jersey. Proc. Entomol. Soc. Wash. 12: 49-52.

Rohwer, S.A. 1911. Descriptions of new species of wasps with notes on described species. Proc.
U.S. Natl. Mus. 40: 551-587.

Smith, H.S. 1908. The Sphegoidea of Nebraska. Univ. Nebr. Stud. 8: 323-410.

Smith, J.B. 1 9 1 0. A report of the insects of New Jersey. Ann. Rep. New Jersey State Mus. 1 909:
14-880.

Viereck, H.L. 1916. Guide to the insects of Connecticut. Part III. The Hymenoptera, or wasp-
like insects, of Connecticut. Conn. Geol. and Nat. Hist. Surv. 22: 1-824.

Williams, F.X. 1914 (1913). Monograph of the Larridae of Kansas. Univ. Kans. Sci. Bull. 8:
117-213.

256 ENTOMOLOGICAL NEWS

SCIENTIFIC NOTE

INCORPORATION OF BATRACHOSPERMUM

GELATINOSUM (RHODOPHYTA) INTO CASES OF

OCHROTRICHIA WOJCICKYI (TRICHOPTERA:

HYDROPTILIDAE) 1

J. B. Keiper 2 , D. A. Casamatta 3 , B. A. Foote

Microcaddisfly larvae (Trichoptera: Hydroptilidae) often incorporate filaments of algae into
their cases (Wiggins 1996). Sheath et al. (1995) studied the incorporation of freshwater red algae
(Rhodophyta) into the cases of a variety of caddisfly species. They reported that hydroptilid
larvae of Dibusa, Hydroptila, and Ochrotrichia had Rhodophyta incorporated into their cases,
but no Ochrotrichia were identified to species. To supplement their study, we report the incorpo-
ration of the rhodophytan Batrachospermum gelatinosum (L.) De Candolle into the cases of
Ochrotrichia wojcickyi Blickle. Although B. gelatinosum has been associated with the cases of
midge larvae (Diptera: Chironomidae) (Sheath et al. 1 996), this is the first report of B. gelatinosum
used as caddisfly case material.

Larvae were collected from cobble substrata in an unnamed low order stream located in Salt
Fork State Park (4006'03 N,8128'41 W), Guernsey Co., Ohio, on 26 May 1997. They were
transported to the laboratory, placed in aerated rearing chambers (Keiper and Foote 1996) with
field-collected riffle rocks, and the adults reared for species determination. Larvae were never
observed to consume B. gelatinosum in the laboratory. Living algae covered approximately 5- 1 5%
of cases examined.

Sand grains often constitute most of the case material used by larvae of Ochrotrichia (Wiggins
1996). The incorporation of B. gelatinosum into the cases of the specimens collected suggests
that O. wojcickyi is not specific in its case material requirements. The fate of living Rhodophyta
incorporated into caddisfly cases remains unknown.

ACKNOWLEDGMENTS

We thank the Ohio Biological Survey for supporting the collecting trips of the senior author.
L. Boydeletour of the Ohio Department of Natural Resources, Parks and Natural Areas kindly
supplied the collecting permit for Ohio state parks. A. Collier and J. A. Johns (KSU) critically
reviewed the manuscript.

LITERATURE CITED

Keiper, J. B. and B. A. Foote. 1996. A simple rearing chamber for lotic insect larvae. Hydro-

biologia339: 137-139.
Sheath, R. G., K. M. Muller, D. J. Larson, and K. M. Cole. 1995. Incorporation of freshwater

Rhodophyta into the cases of caddisflies (Trichoptera) from North America. J. Phycol. 31:

889-896.
Sheath, R. G., K. M. Muller, M. H. Colbo, and K. M. Cole. 1996. Incorporation of freshwater

Rhodophyta into the cases of chironomid larvae (Chironomidae, Diptera) from North America.

J. Phycol. 32: 949-952.
Wiggins, G. B. 1996. Larvae of the North American caddisfly genera (Trichoptera). 2nd ed.

Univ. Toronto Press, Toronto. 457 pp.

1 Received September 27, 1997. Accepted November 22, 1997.

2 Dept. of Entomology, University of California, Riverside, CA 92521.

3 Dept. of Biological Sciences, Kent State University, Kent, Ohio 44242-0001,

ENT. NEWS 109(4) 256, September & October, 1998

Vol. 109, No. 4, September & October, 1998 257

CHARACTER VARIABILITY AND A NEW SYNONYM

OF ACERPENNA PYGMAEA
(EPHEMEROPTERA: BAETIDAE) 1

R. D. Waltz 2 , D.E. Baumgardner 3 ' 4 , J.H. Kennedy 4

ABSTRACT: An adult of Acerpenna harti was reared from a larva and compared with larvae of
A. macdunnoughi, and A. pygmaea. Acerpenna harti can not be consistently distinguished as
larvae or adults from A. pygmaea based on observed variabilities in adult and larval characters.
Acerpenna harti is reported as a junior subjective synonym of Acerpenna pygmaea, new syn-
onymy.

Acerpenna harti (McDunnough) has only rarely been reported by mayfly
workers since its description over seventy years ago. The only published keys
to incorporate this species in the adult stage are those of Traver (1935, as Baetis)
and Burks (1953, as Baetis). McDunnough (1924) first described A. harti (as
Baetis) based on specimens collected July 11,1 898, from Urbana, Illinois, and
housed in the Canadian National Collection, Ottawa. Burks (1953) revised the
male description and included additional Illinois records. Waltz and McCafferty
(1987) first characterized Acerpenna as a genus distinct from Baetis Leach but
they did not transfer the species B. harti McD. or B. akataleptos McD. until
1990 (McCafferty and Waltz, 1990). As of this writing, only the larva of A.
akataleptos remains unknown among the Nearctic Acerpenna species. We
strongly suspect, but do not have any data to conclude at this time, that A.
akataleptos is also a synonym of A. pygmaea.

McCafferty et al. (1993), although not able to assign a species name to
larvae of Acerpenna collected in Colorado, alluded to the possible occurrence
of A. akataleptos in Colorado due to the discovery of Acerpenna larvae with
gills similar in form to A. pygmaea (Hagen) but differing in having both the
anterior and posterior margins of gill 7 serrate. This condition is distinctly
different from the generally accepted concept of A. pygmaea larvae character-
ized by Ide ( 1 937) and Morihara and McCafferty ( 1 979), in which the gills are
asymmetric and only the posterior margins are serrate.

MATERIAL EXAMINED

One of us (DEB) reared one larva from which the adult was identified as
Acerpenna harti. Collection data for the reared specimen is as follows: OKLA-

' Received October 4, 1997. Accepted November 20, 1997.

2 IDNR, Division of Entomology and Plant Pathology, 402 West Washington, Room W-290,
Indianapolis, IN 46204.

3 Current address: David Baumgardner, USACHPPM, ATTN: MCHB-TS-OEN, 5 1 58 Blackhawk
Road, Aberdeen Proving Ground, MD 21010.

4 Department of Biological Sciences, University of North Texas, P.O. Box 5218, Denton, TX
76203-0218.

ENT. NEWS 109(4) 257-260 , September & October, 1998

258 ENTOMOLOGICAL NEWS

HOMA: Pushmataha Co., Pine Ck @ unnamed rd (old main highway) in
Wadena, 16- VII- 1993, DE Baumgardner, (DB 93-137), [1 reared male larval
exuviae (slide mounted: euparal: abs. ale.) and its adult].

Paratypes: Acerpenna harti McDunnough (2 Male adults): IL: Urbana, July 11,1 898, 1NHS tag
No. 24491, CNC Paratype No. 708, housed in Canadian National Collection, Ottawa. 4 speci-
mens labeled Ontario: Ottawa, 7- V- 1921, 8-5-1921, 10- V- 1920, 1 7- V- 1921, all collected by J.H.
McDunnough.

AL: Choctaw Co., Tucklebaum Cr. @ Hwy 17, 12 May 1989, S. Harris, (20 males).
CO: Moffat Co., Green River, Echo Park, Dinosaur NM, 19 August 1993, B. Kondratieff and R.
Durfee, (1 larva). CO: Moffat Co., Yampa River, Echo Park, Dinosaur NM, 3 September 1994,
B. Kondratieff and R. Durfee, ( 1 larva). CO: Routt Co., Yampa River, near Steamboat Springs off
Rt. 131, 28 July 1991, Kondratieff, Durfee, and Painter, (1 larva). CO: Yuma Co., Chief Creek
Co. Road CC North, 25 April, 1993, B. Kondratieff, R. Durfee, (1 reared male and 2 reared
females and associated larval exuviae, 1 larva).

MI: Emmet Co., Lake Michigan, Trail's End Bay, VI-6-1994, P. Hudson (1 larval exuviae).
OK: Le Flore Co.; Rock Ck. @ Railroad Road ca. 0.5 mi N of Talihina, 17-111-1993, D.E.
Baumgardner, (DB 93-46), (5 larvae). OK: LeFlore Co.; Bimey Ck. @ Hwy 63, ca. 5 mi SE of
Talihina, 16-V-1993, D.E. Baumgardner, (DB 93-90), (3 larvae). OK: Pushmataha Co.; Beaver
Ck. @ Coffee Ck. Rd., ca. 1.5 mi S of Hwy 3 jet, ca. 4 mi W of Antlers, 1 7-1 V- 1993, D.E.
Baumgardner, (DB 93-49), (1 larva). Same as preceding but, 14-111-1993, (DB 93-27), (2 lar-
vae). OK: Pushmataha Co.; Cedar Ck. @ Snow Ck. Rd. in Snow, 1 7-III- 1 993, D. E. Baumgardner,
(DB 93-47), (2 larvae). OK: Pushmataha Co.; Kiamichi R @ unnamed rd., 3 mi E of Albion, 18-
IV- 1993, (DB 93-27), (2 larvae). OK: LeFlore Co.; Unnamed 2nd order trib. of Buzzard Ck. @
unnamed rd., ca. 2 mi. E. of Talihina, 17-111-1993, DE Baumgardner (DB 93-45.5) (6 larvae);
OK: Pushmataha Co., Marble Fobb Ck. (?) at main rd. in Kiamichi Wilderness Area, 17-IV- 1 993,
DE Baumgardner, (DB 93-53) (2 larvae); OK: Pushmataha Co., Cole Ck. @ unnamed rd., 3.7 mi.
S. of Miller, 14-111-1993, DE Baumgardner, (DB 93-25) (6 larvae). OK: Pushmataha Co., Dry
Ck. @ low water crossing @ unnamed rd., ca. 1.5 mi. E. of Tuskahoma, 17-VII-1993, DE
Baumgardner, (DB 93-142), 2 larvae). OK: Pushmataha Co., Panther Ck. @ Hwy. 2, 3.1 mi. N. of
Hwy. 2-3 jet., 11 -IX- 1992, DE Baumgardner, (DB 92-101), (1 larva). OK: Pushmataha Co.,
Tenmile Ck. @ Hwy. 2 N. of Antlers, 5- VIII- 1994, DE Baumgardner, (DB 94-005), (1 larva).

Slide mounted voucher material (Euparal: abs. ale.) and alcohol preserved material is deposited
at Purdue University, Entomological Research Collection, West Lafayette, Indiana, and at Uni-
versity of North Texas, Denton, Texas.

DISCUSSION

A male larva from Oklahoma, for which the reared adult was readily iden-
tified as A. harti, possessed setulae on the anterior margin of the gill, thereby
giving the gill a distinctly serrated appearance so that one might conclude that
serration existed on both margins of gill 7. In all other respects this larva was
found to be morphologically identical to A. pygmaea. Our examination of ad-
ditional collections [above] of non-reared and reared larvae identified as A.
pygmaea from Colorado, Oklahoma, Alabama, and Michigan indicated that
development of the setulae on the anterior margin of gill 7 was extremely
variable within populations, ranging from well developed to absent.

In adult specimens, we found hindwing characters to be highly variable.
Previously published keys have used characters such as hindwing venation

Vol. 109, No. 4, September & October, 1998

259

(Burks 1953) and development of the costal process (Traver 1935) to separate
adults of A. pygmaea from A. harti. Adult specimens examined from Alabama
and Colorado showed variation in both of these characters. The costal process
varied from acute and well developed to blunt and poorly developed (Figs. 1-
4). Traver (1935) considered the presence of an acute well-developed costal
process to be typical of A. harti while A. pygmaea possessed a small, blunt
costal process. Burks (1953) separated A. pygmaea from A. harti by the pres-
ence or absence of marginal intercalary veins in the hindwing. The hindwing
of Acerpenna pygmaea was considered to possess 1-2 marginal intercalary
veins, whereas A. harti completely lacked intercalary veins in the hindwing.
Series of adult specimens from Alabama and Colorado varied from 0-2 mar-
ginal intercalaries in the hindwing, with the location of the intercalates being
highly variable (Figs. 2-4). Additionally, some adults possessed hindwings with
a weakly developed third longitudinal vein (Figs. 1 , 3), while other adults pos-
sessed a cross vein between the two longitudinal veins (Figs. 2, 4), or pos-
sessed intercalaries along the outer margin (Fig. 2-4), or intercalaries in both
the outer margin and between the longitudinal veins (Fig. 3); at least one speci-
men demonstrated intersected intercalaries (Fig. 4).

Figs. 1-4. Hindwing (hw) variations in A. pygmaea. 1) hw with blunt costal process and third
vein (Alabama specimen), 2) hw with two free marginal intercalaries (Colorado specimen), 3)
hw with basal, third vein and intercalaries (Alabama specimen), 4) hw with intersected intercalaries
(Colorado specimen).

260 ENTOMOLOGICAL NEWS

Similar to our findings of venational variations in the hindwings of this
species, Durfee and Kondratieff (1993) reported nearly identical venational
variations of hindwings in their study ofBaetis magnus McCafferty and Waltz
and B. tricaudatus Dodds. Based on these studies, workers should use caution
in separating at least some species of baetids traditionally discriminated on
hindwing characters alone,including species discriminations based on minor
differences in wing venation and color (e.g., Waltz 1995).

Based on the above data, including our examination of the reared male
compared with the adult paratypes of Acerpenna harti, we conclude that
Acerpenna harti is a junior subjective synonym of Acerpenna pygmaea, new
synonymy.

ACKNOWLEDGMENTS

The senior author especially thanks Ed C. Becker, Canadian National Collection, Ottawa,
for providing access to the types of Acerpenna harti and A. akataleptos as well as other non-type
material of these species. We also thank B. Kondratieff and W.P. McCafferty for loan of other
Acerpenna material. We thank W. P. McCafferty and A.V. Provonsha for comments on an early
draft of this manuscript.

LITERATURE CITED

Burks, B.D. 1953. The Mayflies, or Ephemeroptera, of Illinois. 111. Nat. Hist. Surv. Bull. 26, 216
pp.

Durfee, R. and B. C. Kondratieff. 1993. Description of the adults of Baetis magnus
(Ephemeroptera: Baetidae). Entomol. News 104: 227-232.

Ide, P.P. 1937. Descriptions of Eastern North American species of baetine mayflies with particu-
lar reference to the nymphal stages. Can. Entomol. 69: 219-231, 235-243.

McCafferty, W.P., R.S. Durfee, and B.C. Kondratieff. 1993. Colorado mayflies (Ephe-
meroptera): An annotated inventory. Southwest. Nat. 38: 252-274.

McCafferty, W.P. and R.D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera)
of North and Middle America. Trans. Am. Entomol. Soc. 1 16: 769-799.

McDunnough, J. 1924. New Ephemeridae from Illinois. Can. Entomol. 56: 7-9.

Morihara, D.K. and W.P. McCafferty. 1 979. The Baetis larvae of North America (Ephe-
meroptera: Baetidae). Trans. Am. Entomol. Soc. 105: 139-221.

Traver, J.R. 1935. Part II. Systematic, pp. 237-739 in Needham, J.G., J.R.Traverand Y.C. Hsu
(eds.), The biology of mayflies with a systematic account of North American species. Comstock
Publ. Co., Ithaca, NY. 759 pp.

Waltz, R.D. 1995. Baetis ochris, a new synonym of Baetis flavistriga (Ephemeroptera: Baetidae).
Entomol. News 106: 75-76.

Waltz, R.D. and W.P. McCafferty. 1987. New genera of Baetidae for some Nearctic species
previously included in Baetis Leach (Ephemeroptera). Ann. Entomol. Soc. Am. 80: 667-670.

PLEASE SEE ANNOUNCEMENT ON PAGE 276

Vol. 109, No. 4, September & October, 1998 261

REARED ASSOCIATION AND EQUIVALENCY OF

BAETIS ADONIS AND B. CAELESTIS

(EPHEMEROPTERA: BAETIDAE) 1

W. P. McCafferty 2 , E. L. Silldorff 3

ABSTRACT: The examination of reared larvae and adults of a southwestern North American
small minnow mayfly species (Ephemeroptera: Baetidae) belonging to the rhodani group of the
genus Baetis indicates that larvae previously referable to B. caelestis are actually B. adonis. The
species names are synonymized, the adult description of B. adonis is expanded, and other data
associated with this previously poorly known species are updated.

Morihara and McCafferty ( 1 979), in their revision of the Baetis species of
North America, described the distinctive larvae of three species from western
North America that they did not name at the time, but designated simply as
species A, B, and C. Those authors were reluctant to assign formal names to
those distinctive larval forms because numerous Baetis species in western North
America remained known only as adults, and there was a reasonable chance
that the newly discovered larvae were associated with one of those nominal
species.

In the absence of any adult association, Allen and Murvosh (1983) ap-
plied the name B. caelestis Allen and Murvosh to B. sp. A Morihara and
McCafferty. Later, Allen and Murvosh ( 1 987) applied the names B. libos Allen
and Murvosh to B, sp. B Morihara and McCafferty, and B. notos Allen and
Murvosh to B. sp. C Morihara and McCafferty, both in the absence of adult
associations. In the meantime, however, McCafferty and Waltz (1986) had
already named B. sp. B as B. magnus McCafferty and Waltz because they had
associated (by time and location, not rearing) distinct adults with the species
that were apparently not previously described, thus suggesting the validity of
the species. Based on reared materials, Durfee and Kondratieff (1993) were
able to provide the first descriptions of the distinctive adults of B. magnus,
showing that indeed the species was not known under a prior nomen. Durfee
and Kondratieff ( 1 995) later also reared B. notos and provided a description of
the adult. There remains a slight possibility that B. notos is a synonym of the
prior name B. caurinus Edmunds and Allen (which was based only on adults).
Adult characterization associated with the latter two names is possibly within
an expected range of variation, and B. caurinus has yet to be associated with
any larvae.

Received December 4, 1997. Accepted January 22, 1998.

Department of Entomology, Purdue University, West Lafayette, IN 47907, USA.

Department of Ecology, Evolution, and Marine Biology, University of California, Santa Bar-
bara, C A 93106, USA.

ENT. NEWS 109(4) 261-265, September & October, 1998

262 ENTOMOLOGICAL NEWS

Baetis sp. A, as B. caelestis, has become better known with recent new
reports of larvae from Arizona (Lugo-Ortiz and McCafferty 1995), Chihua-
hua, Mexico (Lugo-Ortiz and McCafferty 1996), New Mexico (McCafferty et
al. 1997), and Texas (McCafferty and Davis 1992). However, because larvae
were never reared to adult, the question has remained as to whether or not the
name would prove to be a synonym.

Recently, eggs taken from Mission Creek, Santa Barbara County, Califor-
nia, were reared to the larval stage identifiable as B. caelestis, and also to the
adult stage. Examination of those reared adults has indicated, however, that
they are referable to B. adonis Traver. Baetis adonis is a species that has been
known only as adults and was originally described by Traver (1935) from
material collected in the San Gabriel Mountains of southern California. The
only substantiated subsequent report of that species was that of McCafferty et
al. (1997) from the Pecos River drainage of eastern New Mexico. Recent ex-
amination of alate material on which Day (1963) had based a report of B.
adonis from Sonoma County in northern California indicated that that mate-
rial was not referable to B. adonis, but probably was B. diablus Day.

Morihara and McCafferty (1979) had indicated that the larvae of B. sp. A
was a member of the rhodani species group of Baetis, and B. adonis, by hav-
ing the so-called moffatti type of genitalia (Traver 1935), would also by defini-
tion be placed in the rhodani group. Several species besides B. adonis were
initial candidates for association with B. sp. A based on 1) their belonging to
the rhodani group, 2) being unknown in the larval stage, and 3) being distrib-
uted in a reasonable vicinity of southern California. These included B. palisadi
Mayo, B. parallelus Banks, B. persecutor McDunnough, B. piscatoris Traver,
and possibly B. diablus Day, B. sulfurosus Day, and B. thermophilos
McDunnough (the species group relationship of the latter three remains in
some doubt). Baetis caurinus Edmunds and Allen, known only from Oregon
and unknown in the larval stage, was easily excluded as a possibility on the
basis that it is a member of Ihefuscatus group of species (Edmunds and Allen
1957, Morihara and McCafferty 1979). All others, except B. adonis could be
excluded on the basis of various characteristics, particularly male genitalia,
that were not within the expected range of variation of the characterization
expressed by the reared adults under question. The color pattern that Traver
(1935) described for B. adonis matched well with that of the reared adults.
The tone of colors and the body and forewing size did not precisely match, but
were within a range of expected variation. Although Traver (1935) did not
illustrate or adequately describe the male genitalia of B. adonis, identification
of reared B. adonis from southern California was confirmed by morphological
comparisons with type material of that species.

This new association necessitates the following nomenclature adjustment:
Baetis adonis Traver [= Baetis caelestis Allen and Murvosh], n. syn. An ad-

Vol. 109, No. 4, September & October, 1998 263

equate description of the larvae of B. adonis may be found in Morihara and
McCafferty (1979:160) as B. sp. A, and the basic adult color patterning and
hindwing venation is adequately described by Traver (1935:680). The adult
description of B. adonis is augmented below to include minor variation in size
and color traits and a first description of the male genitalia.

Baetis adonis Traver

Material examined. 1 2 larvae, 55 final-instar larval exuviae, 73 subimagos, 1 7 male adults, and
12 female adults (all alates reared), California, Santa Barbara County, Mission Creek, VI.VII-
1997, E. Silldorff, deposited in the Purdue Entomological Research Collection (PERC). Male
adult holotype, allotype, and one male and three female paratypes, California, Los Angeles County,
San Gabriel Mountains, VI-3-1932, C. D. Michener, deposited in Cornell University Collection
(all type material in alcohol and in very poor condition, with many types broken and deteriorat-
ing). One male adult and five female adults, New Mexico, San Miguel County, 5 mi north Pecos,
Dalton Fishing Site, VII- 13- 1969, Koss, McCafferty, Provonsha (PERC).
Male adult. Body length, 5.0-7.0 mm. Forewing length, 5.0-6.5 mm. Base color of head and
antennae light reddish brown to medium brown. Dorsal face of turbinate eyes vivid orange,
columnar basal area cream (in alcohol). Base color of thorax reddish brown to medium brown to
dark olive-brown. Abdominal segments 1-6 ranging from light yellow-brown to medium brown
to medium olive brown, varying from pale to intense; segments 7- 1 always slightly to consider-
ably darker shade of same colors of segments 1 -6, sometimes also appearing opaque. Male geni-
talia as in Figure 1; base of forceps segment 2 evenly expanding basally and apically into con-
spicuous more or less symmetrical medially bulbous region, in some bulbous basal area sepa-
rated from longer narrow portion of forceps segment 2 by somewhat darkened band, or in some
entire bulbous basal area of forceps segment 2 darker than remainder of segment; forceps seg-
ment 3 relatively short and with demarcation from segment 2 appearing more notched laterally
and more fused medially; distal margin between base of forceps segments straight to slightly
convex and not notched or emarginate medially.

Distribution. Arizona; Baja California, Mexico; Chihuahua, Mexico; southern California; New
Mexico; west Texas.

Remarks. Based on comparisons of male genitalia, it appears that B. ado-
nis is related to B. persecutor, both of which are very similar with respect to
the basal area of forceps segment 2. Baetis persecutor is known only from
Alberta and British Columbia and remains unknown as larvae. Color pattern-
ing in the male adult of B. persecutor is quite different than that of B. adonis,
as is the terminal area of the forceps. The terminal region of the forceps of B.
adonis is similar to that of B. diablus, which is known from the central Califor-
nia coastal area, but otherwise the two species are very distinct.

The mature larvae of B. adonis from Santa Barbara are slightly larger than
others that have been seen from the Southwest. This may be related to the fact
that they were reared for their entire life under ideal laboratory conditions, and
were provided with laboratory cultured algae at all times. This slight size in-
crease is possibly reflected in the adult size of approximately 7.0 mm vs. the

264

ENTOMOLOGICAL NEWS

Fig. 1. Baetis adonis, male adult, genitalia (ventral view).

5.0 mm of the original type material of this species. We do not know if the
predominate olive brown color of the Santa Barbara adults (vs. the yellow-
brown of the original material) may also have been related to rearing conditions.
Reared females were predominantly reddish brown (quite reddish), and although
Traver (1935) described most females of B. adonis that she had seen as being
yellow-brown or pale reddish brown, she did note that one paratype was wholly
reddish brown.

ACKNOWLEDGMENTS

We thank Richard Hobeke (Ithaca, New York) for the loan of type material, Arwin Provonsha
(West Lafayette, Indiana) for illustrating the male genitalia, and Carlos Lugo-Ortiz (West Lafayette,
Indiana) for his assistance. This paper has been assigned Purdue Agricultural Research Program
Journal No. 15610.

Vol. 109, No. 4, September & October, 1998 265

LITERATURE CITED

Allen, R. K. and C. M. Murvosh. 1983. Taxonomy and zoogeography of the mayflies
(Ephemeroptera: Insecta) of Baja California. Ann. Entomol. Soc. Am. 76: 425-433.

Allen, R. K. and C. M. Murvosh. 1987. New Baetidae from the southwestern United States and
northern Mexico (Ephemeroptera: Insecta), with notes. Can. Entomol. 119: 1095-1099.

Day, W. C. 1963. Ephemeroptera, pp. 79-105 in R. L. Usinger (ed.). Aquatic insects of Califor-
nia. Univ. Calif. Press, Berkeley.

Durfee, R. S. and B. C. Kondratieff. 1993. Description of adults of Baetis magnus
(Ephemeroptera: Baetidae). Entomol. News 104: 227-232.

Durfee, R. S. and B. C. Kondratieff. 1 995. Description of adults of Baetis notos (Ephemeroptera:
Baetidae). Entomol. News 106: 71-74.

Edmunds, G. F., Jr. and R. K. Allen. 1957. A new species of Baetis from Oregon. J. Kans.
Entomol. Soc. 30: 57-58.

Lugo-Ortiz, C. R. and W. P. McCafferty. 1995. Annotated inventory of the mayflies
(Ephemeroptera) of Arizona. Entomol. News 106: 131-140.

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996. New Central American and Mexican records of
Ephemeroptera species. Entomol. News 107: 303-310.

McCafferty, W. P. and J. R. Davis. 1992. New and additional records of small minnow may-
flies from Texas. Entomol. News 103: 199-209.

McCafferty, W. P. and R. D. Waltz. 1986. Baetis magnus, new species, formal new name for
Baetis sp. B of Morihara and McCafferty (Ephemeroptera: Baetidae). Proc. Entomol. Soc.
Wash. 88: 604.

McCafferty, W. P., C. R. Lugo-Ortiz, and G. Z. Jacobi. 1997. Mayfly fauna of New Mexico.
Gr. Basin Natural. 57: 283-314.

Morihara, D. K. and W. P. McCafferty. 1979. The Baetis larvae of North America
(Ephemeroptera: Baetidae). Trans. Am. Entomol. Soc. 105: 139-221.

Traver, J. R. 1935. Part II, Systematic, pp. 237-739 in J. G. Needham, J. R. Traver, and T.-C.
Hsu (eds.). The biology of mayflies. Comstock Publ. Co., Ithaca, New York.

BOOK REVIEW

SPINELESS WONDERS. STRANGE TALES FROM THE INVERTEBRATE
WORLD. Richard Conniff. 1997. Owl Books, Henry Holt & Co. NY. 232 pp.
$12.95ppbk.

In this small book, nature writer Conniff offers engaging, yet sophisticated discussions on
the natural histories of the invertebrate world, from earthworms, leeches, slime eels, and giant
squids, to tarantulas, dragon flies, ground beetles, fleas, moths, mosquitoes, house flies, and fire
ants.

For each subject, Conniff obtained the advice and counsel of well recognized authorities and
further prepared himself by participating in several ongoing, scientific, field research experi-
ences, often in remote, native environments around the world. Thus, in addition to being interest-
ingly and cleverly written, these are scientifically accurate discussions. Spineless Wonders takes
one directly to the creepy, crawly frontier of natural science, to the hazards of being around
invertebrates, to the bizarre adaptations that enable them to survive in this world, and to the
astonishing work they do - work that ultimately enables us to survive.

An adequate bibliography is provided. It is unfortunate the publisher did not see fit to pro-
vide a cover that does not curl back on itself when not weighted down.

H.P.B.

266 ENTOMOLOGICAL NEWS

ADDITIONS AND CORRECTIONS TO

EPHEMEROPTERA SPECIES OF NORTH AMERICA

AND INDEX TO THEIR COMPLETE

NOMENCLATURE 1

W. P. McCafferty 2

ABSTRACT: Heretofore unpublished additions, deletions, and changes to the main checklist of
valid North American Ephemeroptera species are provided as are corrections and additions to the
nomenclatural index of all names historically associated with them. Index modifications include
the addition of 44 new entries and corrections to 21 previous entries.

Subsequent to the appearance of "The Ephemeroptera Species of North
America and Index to Their Complete Nomenclature" (McCafferty 1996), cer-
tain omissions and minor errors have been discovered. These emendations have
come to light primarily as a result of preparing the database of Nearctic
Ephemeroptera reflected in McCafferty (1997a), which is strictly predicated
on original orthography of species, and of close scrutiny of the literature asso-
ciated with the Canadian fauna currently under study. Data revealed specific
synonyms that had not appeared previously as well as additional name usage
(spelling and combinations) found for certain species. In order to maintain the
highest degree of utility of the North American index to mayfly species no-
menclature (McCafferty 1996), these pertinent modifications are presented
below. Additional recent significant changes and corrections to the list and
index not covered below can be found respectively in the revision ofAmeletus
by Zloty (1996) and adjustments to names by McCafferty (1997b); however,
for all changes published since McCafferty (1996), particularly with regard to
occasional new species and synonyms being published, the regularly updated
"The Mayflies of North America" found at the Mayfly Central World Wide
Web cite <http://www.entm.purdue.edu/entomology/mayfly/mayfly.html>
should be consulted. Additions and corrections are given below both for the
main numerically sequenced checklist of species and for the alphabetical in-
dex. In the index, species names appear in italics only for the current valid
form of the name as per the McCafferty ( 1 996) format.

Received November 17, 1997. Accepted December 28, 1997.
Department of Entomology, Purdue University, West Lafayette, IN 47907.

ENT. NEWS 109(4) 266-268, September & October, 1998

Vol. 109, No. 4, September & October, 1998

267

MODIFICATIONS TO THE
NUMERICAL CHECKLIST

Delete

003. Choroterpes ferruginea Traver, 1934

004. Choroterpes fusca Spieth, 1938

005. Choroterpes hubbelli Berner, 1946
132. Brachyce rcus floridicola Soldan, 1986
137. Brachycercus pini Soldan, 1986

Corrections

016. Leptophlebia bradleyi Needham, 1932
363. Baetis moffatti Dodds, 1923
574. Heptagenia adaequata McDunnough, 1924
636. Rhithrogena notialis Allen & Cohen, 1977
671. Stenonema terminatum terminatum
(Walsh), 1862

MODIFICATIONS TO THE
ALPHABETICAL INDEX

Additions

Baetis curiosus (McDunnough), 1923 - 432
Baetis moffati Dodds, 1923 - 363
Baetis pallidula McDunnough, 1924 - 357
Baetis persecuta McDunnough, 1939 - 367
Blasturus cupidus (Say), 1823 - 018
Blasrurus grandis Traver, 1932 - 019
Brachycercus pallidus (Ide), 1930 - 133
Callibaetis tessellatus (Hagen), 1861 - 399
Campsurus manitobensis Ide, 1941 - 092
Cloeon chlorops McDunnough, 1923 - 355
Cloeon punctiventris McDunnough, 1923 - 371
Cloeon virilis McDunnough, 1923 - 378
Ecdyonurus fuscus (Clemens), 1913 - 672
Ecdyonurus peterseni Lestage, 1930 - 542
Ecdyonurus werestschagini (Tshernova),
1952-619

Ecdyurus hyalinus Esben-Petersen, 1916 - 542
Epeorus undulatus Banks, 1924 - 642
Ephemerella fuscata (Walker), 1853 - 179
Eurycaenis pallida Ide, 1930 - 133
Heptagenia abnorme Tshernova, 1949 - 542
Heptagenia abnormis Tshernova, 1949 - 542
Heptagenia adequata McDunnough,

1924 - 574
Heptagenia luridipennis (Burmeister),

1839 - misidentification of 667
Heptagenia verticis (Say), 1839 - 085

Heptagenia werestschagini Tshernova,

1952-619

Iron petulans Seemann, 1927 - 635
Isonychia annulata Traver, 1932 - 515
Isonychia aurea Traver, 1932-524
Isonychia thalia Traver, 1934 - 515
Leptophlebia concinna (Walker), 1853 - 018
Leptophlebia hebes (Walker), 1853 - 018
Leptophlebia ignava (Hagen), 1861 - 018
Leptophlebia pallipes (Walker), 1853 - 018
Oreianthus sp. 1 Traver, 1937 - 126
Paraleptophlebia compar Traver, 1934 - 033
Paraleptophlebia pallipes (Hagen), 1875 - 049
Potamanthus rufus Argo, 1927 - 082
Pseudocloeon virilis (McDunnough),

1923 - 378

Rhithrogena fusca (Walker), 1853 - 634
Rhithrogena imanica Bajkova, 1972 - 619
Siphlonurus bernice McDunnough, 1923 - 509
Siphloplecton basalis (Walker), 1853 - 477
Stenonema Carolina (Banks), 1914 - 646
Stenonema tessellata (Walker), 1853 - 672

Corrections

Ameletus mantis Mayo, 1952 - 313
Baetis moffatti Dodds, 1923 - 363
Brachycercus floridicola Soldan, 1986 - 129
Brachycercus pini Solda"n, 1986 - 134
Brachycercus sp. A Berner, 1950 - 129
Choroterpes ferruginea Traver, 1934 - 002
Choroterpes fusca Spieth, 1938 - 002
Choroterpes hubbelli Berner, 1946 - 002
Cinygma deceptiva McDunnough, 1924 - 555
Cloe mollis Hagen, 1861 - 051
Cloeon vicinum Hagen, 1861 - 471
Epeorus humeralis Morgan, 1911 - 573
Heptagenia adaequata McDunnough, 1924 - 574
Leptophlebia bradleyi Needham, 1932 - 016
Paraleptophlebia bradleyi (Needham),

1932 - 016

Pseudocloeon Carolina (Banks), 1924 - 335
Pseudocloeon cingulatum McDunnough,

1931 - 353

Rhithrogena hespera Banks, 1924 - 561
Rhithrogena notialis Allen & Cohen, 1977-636
Rhithrogena petulans (Seemann), 1927 - 635
Thraulus albertanus McDunnough, 1931 - 077

268 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

I would like to thank Manny Pescador (Tallahassee, Florida), Bob Waltz (Indianapolis, Indi-
ana), and Boris Kondratieff (Fort Collins, Colorado) for calling my attention to certain informa-
tion. This paper has been assigned Purdue University ARP No. 15537.

LITERATURE CITED

McCafferty, W. P. 1996. The Ephemeroptera species of North America and index to their com-
plete nomenclature. Trans. Am. Entomol. Soc. 122: 1-54.

McCafferty, W. P. 1997a. Ephemeroptera. Pages 89-117. In: R. E. Poole and P. Gentili (eds.).
Nomina Insecta Nearctica, a checklist of the insects of North America, Volume 4: Non-ho-
lometabolous orders. Entomological Information Services, Rockville, Maryland.

McCafferty, W. P. 1997b. Name adjustments and a new synonym for North American Ephe-
meroptera species. Entomol. News 108: 318, 320.

Zloty, J. 1996. A revision of the Nearctic Ameletus mayflies based on adult males, with descrip-
tions of seven new species (Ephemeroptera: Ameletidae). Can. Entomol. 128: 293-346.

SCIENTIFIC NOTE

ADDITIONS TO IOWA MAYFLIES
(EPHEMEROPTERA) 1 ' 2

W. P. McCafferty 3 , Todd Hubbard 4

Klubertanz (1995) recorded 66 nominal species of mayflies from the state of Iowa. Thirty-nine
of those records represented new state records. Herein we add 10 new Iowa records. Newly
reported materials are deposited at the Purdue Entomological Research Collection, West Lafayette,
IN (PERC) or the University of Iowa Hygienic Laboratory, Des Moines, I A (IHL).

New state records include Baetisca obesa (Say) [larvae, Black Hawk Co, falls access,
VI-9-1973 (PERC)]; Homoeoneuria ammophila (Spieth) [larvae, Ida Co, Maple R, VIII-5-1997
(IHL)]; Labiobaelis dardanus (McDunnough) [larvae, Ida Co, Odebolt Cr, VIII-5-1997 (IHL)];
Leucrocuta juno (McDunnough) [adults, Story Co, Ames, VIII-1-1991 (PERC)]; Procloeon
irrubrum Lowen and Flannagan [all larvae, all IHL: Dickinson Co, Little Sioux R, 1X-4-1996;
Webster Co, Lizard Cr, IX- 1 1 - 1 996; Kossuth Co, Black Cat Cr, VIII- 1 3- 1 997; Shelby Co, West
Nishnabotna R, VIII- 15-1 997; Warren Co, Whitebreast Cr, VIII- 1 - 1 997]; Procloeon rufostrigatum
(McDunnough) [larvae, Webster Co, Lizard Cr, IX-1 1-1996 (IHL)]; Procloeon viridoculare
(Berner) [all larvae: Buchanan Co, Bear Cr, VIII-8-1996; Winnishiek Co, Canoe Cr, IX-9-1997
(IHL)]; Rhithrogena jejuna Eaton [larvae, Winnishiek Co, Upper Iowa R nr Kendallville,
IX-22-1991 (PERC)]; Stenacron candidum (Traver) [adults, Black Hawk Co, Cedar Falls,
VII-1-1973 (PERC)]; and Stenacron Carolina (Banks) [adults, Black Hawk Co, Cedar Falls,
VI-10- 1973 (PERC)].

LITERATURE CITED

Klubertanz, T. H. 1995. Survey of Iowa mayflies (Ephemeroptera). J. Kans. Entomol. Soc. 68:
20-26.

1 Received April 3, 1998. Accepted April 4, 1998.

2 Purdue Agricultural Research Program Journal No. 15699.

3 Department of Entomology, Purdue University, West Lafayette, IN 47907.

4 Hygienic Laboratory, University of Iowa, H. A. Wallace Bldg., Des Moines, I A 50319.

ENT. NEWS 109(4) 268, September & October, 1998

Vol. 109, No. 4, September & October, 1998 269

TAXONOMIC NOTES ON EVANIODINI

(HYMENOPTERA: BRACONIDAE), WITH

REDESCRIPTION OF EVANIODES SPATHIIFORMIS

AND DESCRIPTION OF A NEW SPECIES 1

S. M. Barbalho 2 , A. M. Penteado-Dias 3

ABSTRACT: .The monobasic Pariodes (Hymenoptera: Braconidae, Doryctinae, Evaniodini) is
synonymized with Evaniodes Szepligeti, 1901, thus P. spathiiformis Szepligeti, 1901 is returned
to Evaniodes. Redescription and illustrations of male and female Evaniodes spathiiformis are
provided as well as a description of a new species, Evaniodes marshi.

The tribe Evaniodini (Hymenoptera: Braconidae, Doryctinae) was created
by Fischer (1981) for two species described by Szepligeti (1901) from Brazil,
and characterized by the elevation of the metasoma above the hind coxa. One
of these species, Evaniodes areolatus Szepligeti, is known only from the fe-
male while the second species, E. spathiiformis Szepligeti, is known from the
male and female. Szepligeti (1901 ) differentiated the two species primarily by
the shape of the first metasomal tergite (Tl) (4 times longer than wide in
spathiiformis; 3 times longer than wide in spathiiformis) and by color
(spathiiformis reddish-yellow with black head and antenna and bicolored wings;
areolatus more extensively black with wings almost hyaline). Roman (1924)
noted that both sexes of E. spathiiformis had a closed 1st subdiscal cell in the
fore wing and lighter body color; the female was larger and the male had a
more compact hind wing venation illustrated by him as from spathiipennis. He
also described areolatus as being smaller than spathiiformis; with body color
black with reddish-brown areas and the fore wing brachial cell more or less
open.

Because of the pronounxced differences between the two species, Fischer
(1981) described a new genus, Pariodes, with spathiiformis (male) as its type
and only included species. He did not mention the female used by Roman
(1924). His redescription of E. areolatus indicates that the propodeum has a
pentagonal cell and the hind wing has more complete venation compared to P.
spathiiformis. Differences between the two include eye height in comparison
to temporal height; clypeus height; notauli (rounded or straight, deep or not
deep); venation of hind wing; fore wing 1st subdiscal cell open in E. areolatus
and closed in P. spathiiformis; size of fore wing; Tl height versus apical width
and size of the body.

1 Received June 27, 1997. Accepted December 2, 1997.

2 Programa de Pos-Graduacao em Genetica e Evoluc.ao, Universidade Federal de Sao Carlos, CP
676, CEP 13 565-905, Sao Carlos, SP, Brasil.

3 Departamento de Ecologia e Biologia Evolutiva, Universidade Federal de Sao Carlos, CP676,
CEP 13 565-905, Sao Carlos.SP, Brasil.

ENT. NEWS 109(4) 269-273, September & October, 1998

270 ENTOMOLOGICAL NEWS

Belokobyl'skiy (1993) also treated these two species as being valid for the
tribe Evaniodini and shows the following characters for the tribe: elevated
abdomen; thorax short and swollen; notauli complete; sternauli not developed;
propodeum with or without areola; fore wing 1st subdiscal cell closed; hind
coxa with an antero-ventral basal tubercle; Tl pedicellate and T2 and T3 smooth.
Marsh (1993) incorrectly included both E. areolatus and P. spathiiformis in
the group of Braconidae with no antero-ventral basal tubercle on the hind coxa.

Based on the study of male and female specimens of E. spathiiformis and
the male of a new species from Brazil, we propose to synonymize Pariodes
Fischer with Evaniodes Szepligeti, thus returning spathiiformis to the genus
Evaniodes. We compared male and female Evaniodes specimens with the de-
scriptions in the literature and found the only difference between them was in
the hind wing venation (Figs 1 ,2) with M+CU and cu-a meeting at junction of
SC+R1, Rl, SR and 2-M, with 1-M and r-m absent in the male and veins
M+CU, 1-m and Ir-m distinct, SC+R1 reaching Rl before the middle of the
wing in the female.

For the morphological terminology used in this paper, see van Achterberg
(1993).The following abbreviations are used to indicate deposition of speci-
mens: (INPA), Institute Nacional de Pesquisas da Amazonia, Brasil, (INPA),
Institute Nacional de Pesquisas da Amazonia, Brasil, (MPEG), Museu Paraense
Emelio Goeldi, Brasil, ( DCBU) Departamento de Ecologia e BiologiaEvolutiva
da Universidade Federal de Sao Carlos, Brasil.

Evaniodes spathiiformis Szepligeti, 1901

(Figs 1-4)
Female

Head.- 2.3 times wider than long, 1 .5 times wider than mesonotum; occipital carina present;
gena smooth, face height and width equal, possessing transverse striation; clypeus transversely
striated, 2.5 times broader than height, rounded ventrally; face as broad as basal width of man-
dible; very large eyes occupying most of head; in lateral view, eye height 1 .35 times greater than
length and 3.6 times longer than gena; antenna length shorter than body length; antennae with 46
antennomeres; first flagellomere length 1 .7 times longer than second, succeeding antennomeres
gradually shorter and thinner, placodes evident.

Mesosoma.- Length 1.2 times longer than width; mesonotum 1.3 times wider than long; tegula
trapezoidal; deep and light-crenulate notauli touching each other in rugose area at base of
mesonotum, with accompanying sparse hairs; scutellar sulcus with 5 cross carinae; scutellum
smooth with setae in apical area; propodeum smooth with no pentagonal areola and sparsely
setose; pronotum smooth with posterior sulcus; mesopleuron smooth; sternaulus not deep,
epicnemial carina present extending to medium area of mesopleuron; metapleuron with sparse
hairs; metasternum large and long; hind coxa with weakly distinct antero-ventral basal tubercle;
first tarsomere about 2 times longer than second, second 1 .2 times longer than third plus fourth,
fourth 2 times longer than third, fifth 2.5 times longer than fourth; apical tibial spurs short ( 1/6 of
basitarsus) (Fig. 1).

Fore wings.- Not reaching end of metasoma; 6.4 mm in length; stigma moderately broad , 6.7
times longer than vein r; r almost as long as stigma width; 3-SR length 1 .4 times longer than 2-
SR vein; SRI straight and 2.5 times longer than 3-SR; SRI vein reaching end of wing; m-cu

Vol. 109, No. 4, September & October, 1998 271

straight and interstitial; 2b cell square-shaped; 2-CU1 length 2.8 times longer than m-cu; cu-a
vein postfurcal; 4a cell closed at apex, vein CUlb present and distinctly meeting 2-1 A; CUla
vein arising below middle of 4a cell (Fig. 2).

Hind wings.- Veins M+CU, 1-M and Ir-m distinct; SC+R1 reaching Rl before middle of wing;
M+CU as long as 1 -M; 1 -M 2 times longer than m-cu; m-cu almost reaching posterior side of the
wing (Fig. 3).

Metasoma.- Inserted high on the propodeum, distance between insertion of metasoma and hind
coxa about equal in length to hind coxa; Tl long and thin, 3.3 times longer than apical width,
apical width slightly longer than basal one; Tl striated only at apex; T2 striated and wider than
long as well as remaining tergite; T3 striated at base and rugose at apex; remaining tergites
smooth.

Ovipositor.-Much longer than body length.

Color.- Head black; mandibles yellow with apical area black; yellow palps; antennae brown;
thorax fully yellowish; legs brown (only mesocoxa is yellow); metasoma with basal third light
brown and apical area black; wing membrane infuscated, stigma and veins yellow.
Variation in female. -Head black with dark-brown gena and occiput; eye height greater (varying
between 3.4 to 3.6) than gena; notauli very deep; 2 sulci in pronotum, dorsal smooth, ventral
crenulated; apical width of Tl varying between 3.0 to 3.2 times longer than basal width; T2 and
T3 fully striated with no rugose area; body length between 7.0 and 8.9 mm.
Variation in male.- Clypeus 1.7 to 2.25 broader than its height; eye height 2.3 to 3.1 longer than
gena, 2.3 to 3. 1 ; wings (Fig. 4) slightly longer than body, between 6.4 to 7. 1 mm; in hind wing
veins 1 -M and r-m absent, M+CU and cu-a meeting at junction of SC+R 1 , R 1 , SR and 2-M; T 1
3.8 to 4.3 times longer than its apical width; body length 6.0 to 7.4 mm.

Distribution.- Known only from Brazil : Belem (Para State), Manaus (Amazonas State), and
Mato Grosso State.

Material examined. 1 female (INPA), "Est. Cemat. R. Humb. MT, 26.IX.1975"; 1 female
(MPEG), "Amazonas, Manaus, 1 Km W Taruma, Falls, 14.11.1981, 100 m G. Okis, primary for-
est"; 1 female ( DCBU), "Reserva Ducke, Manaus, Amazonas, E.V.Silva & A. Faustino, 3. V.
1968", 1 male (INPA), "Est. Am 1, Km 101, Mn.Am., Brasil, V. 1968, col. several"; 1 male
(MPEG) "Brasil, Pa, Belem, Mocambo, 31. VIII. 1986"; 1 male (DCBU) , "F. Esteio, Res. 1112,
Manaus, ZF3, Km 23, B. Klein, col., 3. 1. 1986".

Evaniodes marshi Barbalho & Penteado-Dias SPEC. NOV.

(Fig. 5)

Male

Head. - with many hairs; 1.5 times longer than wide, 1.5 times wider than mesonotum; occipital
carina present; gena smooth, face height and width equal, possessing transverse striation; clypeus
transversely striated, as broad as long , straight ventrally; face as broad as mandibular base width;
very large eyes occupying most of head; in lateral view, eye height 1.2 times greater than length
and 3.6 times longer than gena; first flagellomere length 1.9 times longer than second, succeed-
ing antennomeres gradually shorter and thinner.

Mesosoma.- Length 1.6 times longer than wide; tegula elliptical; shallow and light-crenulate
notauli not touching each other at base of mesonotum, with accompanying sparse hairs;
mesoscutum smooth and shining; scutellum smooth with setae in the apical area; propodeum
rugose with no pentagonal areola and with many hairs; mesopleuron smooth , with many hairs,
as well as middle coxae; sternaulus absent, epicnemial carina present extending to medium area
of mesopleuron; metapleuron with many hairs; hind coxa without an antero-ventral basal tu-
bercle; first tarsomere about 2 times longer than second, second as long as third plus fourth,
fourth 2.4 times shorter than third, fifth 2.1 times longer than fourth; apical tibial spurs short (I/
6 of basitarsus); hind femora with a large blister like swelling anteriorly near base (Fig. 5).

272

ENTOMOLOGICAL NEWS

.2-SC+R

Mi-CU 1 r -, m ( fi
1^+R_Z_1X51

cu-a

Figures 1- 4. Evaniodes spathiiformis .1, female, hind tarsus; 2, female, fore wing; 3, female,

hind wing; 4, male, hind wing.

Figure 5. Evaniodes marshi spec. nov. .male, hind leg.

Vol. 109, No. 4, September & October, 1998 273

Fore wings.-Not reaching end of metasoma; 5.7 mm in length; stigma moderately broad , 4.6
times longer than vein r; r shorter than stigma width; 3-SR as long as 2-SR vein; SR 1 straight and
2.5 times longer than 3-SR; SR 1 vein reaching end of wing; m-cu straight and joining 2M before
2 RS; 2b cell square-shaped; 2-CU 1 length 2.6 times longer than m-cu; cu-a vein postfurcal; first
subdiscoidal cell closed at apex, vein CUlb present and distinctly meeting 2-1 A; CUla vein
arising below middle of 4a cell .

Hind wings.- Veins 1 -M and r-m absent, M+CU and cu-a meeting at junction of SC+R 1 , R 1 ,SR
and 2-M.

Metasoma.- Inserted high on propodeum, distance between insertion of metasoma and middle
coxa about equal in length to middle coxa; Tl long and thin, 2.7 times longer than apical width,
apical width slightly longer than basal width; Tl striated only at apex; T2 striated and wider than
long ; T3 striated; remaining tergites smooth.

Color.- Head black; mandibles yellow with apical area black; yellow palps; antennae black;
thorax fully black; fore and middle legs yellow (except middle coxa brown), hind legs dark
brown; metasoma brown; wing membrane lightly infuscated, stigma and veins brown.
Material examined.- Holotype, Brazil, Amazonas, Jurua, Mineruazinho. 1 male , 25/1/96 (depos-
ited in INPA). Female unknown.

Etymology.- Named for Paul M. Marsh because of his contributions to
the knowledge of Doryctinae wasps .

Key to species of Evaniodes.

la- Propodeum with a pentagonal cell; 1st subdiscal cell open E. areolatus Szepligeti

Ib- Propodeum without a pentagonal cell; 1st subdiscal cell closed 2.

2a- Body fully black and with many hairs; hind femora with a blister like

swelling anteriorly near base (known only from male) E. marshi spec, nov

2b- Body fully yellow and with sparse hairs; hind femora without a blister

anteriorly near base E. spathiiformis Szepligeti

LITERATURE CITED

Achterberg, C. van 1993. Illustrated key to the subfamilies of the Braconidae (Hymenoptera:
Ichneumonoidea). Zool. Verb. Leiden 283: 1-189, figs 1-66, photos 1-140, plates 1-102.

Belokobyl'skiy, S. A. 1993. On the classification and phylogeny of braconid wasps of the sub-
family Doryctinae and Exothecinae (Hymenoptera, Braconidae). Part 1. On the Classifica-
tion, l.Entomol. Rev, 77: 109-137.

Fischer, M. 1981.Versuch einer systematischen Gliederung der Doryctinae, insbesondere der
Doryctini, und Redeskriptionen nach Material aus dem Naturwissenschaftlichen Museum in
Budapest (Hymenoptera, Braconidae). Polskie Pismo Entomol., 51: 41-99.

Marsh, P. M. 1993. Description of new Western Hemisphere genera of the subfamily Doryctinae
(Hymenoptera, Braconidae). Contrib. Amer. Entomol. Instil., 28 (1): 1-58.

Roman, A. 1 924. Wissenshaftliche Ergebnisse der schwedischen entomologishen Reise des Herrn
Dr. A. Roman in Amazonas. 1914-1915. Hymenoptera: Braconidae, Cyclostomi pro p. Arkiv
Fur Zool., 20(16): 1-40.

Szepligeti, G. V. 1901. Tropische Cenocoelioniden und Braconiden aus der Sammlung des
Ungarischen National-Museums. Term. Fiiz., 24: 353-405.

274 ENTOMOLOGICAL NEWS

ZORCADWM BERGROTH, AN OBJECTIVE JUNIOR

SYNONYM OF PSEUDOBEBAEUS FALLOU

(HETEROPTERA: PENTATOMIDAE) 1

D. A. Rider 2 , Christian Fischer'

ABSTRACT: Zorcadium Bergroth, 1918, is placed as an objective junior synonym of
Pseudobebaeus Distant, 1911 based on study of their respective type species, Pseudobebaeus
goyazensis Distant, 1911, a junior synonym of Euschistus truncatus Fallou, 1 888.

While working on a catalog of the Pentatomidae of the world, a number of
nomenclatural problems have been discovered. Many of these problems have
already been corrected (Rider & Rolston 1995, Rider 1997). The problem
dealt with in this paper concerns two monotypic genera that were at one time
placed in separate families, but their respective type species are here shown to
be conspecific.

Pseudobebaeus Distant, 1911

Pseudobebaeus Distant, 1911: 254; Kumar, 1974: 20; Fischer, 1996: 83-87. Type species:

Pseudobebaeus goyazensis Distant, 191 1, by monotypy.
Zorcadium Bergroth, 1918: 307-308; Rolston & McDonald, 1981: 259, 269; Rider, 1994: 194,

217-218. Type species: Euschistus truncatus Fallou, 1888, by monotypy. New Synonymy.

Description: See Rolston & McDonald (1981) and Rider (1994) for de-
tailed descriptions of this taxon under the name Zorcadium. Also, see Fischer
(1996) for descriptive notes on Pseudobebaeus.

Pseudobebaeus truncatus (Fallou, 1888), NEW COMBINATION

Euschistus truncatus Fallou, 1888: 36.
Euschistus trancatus [sic]: Bergroth, 1892: 263.

Thoreyella truncata: Lethierry & Severin, 1893: 176; Kirkaldy, 1909: 137.
Pseudobebaeus goyazensis Distant, 1911: 255; Fischer, 1996: 83-87, figs. 1 -6. New Synonymy.
Zorcadium truncatum: Bergroth, 1918: 308; Rolston & McDonald, 1981: 269, 270, figs. 32-39;
Rider, 1994: 218, figs. 136-148.

Description: See Rolston & McDonald (1981) for detailed description
under the name Zorcadium truncatum, and Fischer ( 1 996), for descriptive notes
under the name Pseudobebaeus goyazensis.

1 Received September 23, 1997. Accepted February 9, 1998.

2 Department of Entomology, Box 5346, University Station, North Dakota State University,
Fargo, North Dakota 58105, U.S.A.

3 Freie Universitat Berlin, Institut fur Zoologie, AG Entomologie, Konigin-Luise-Str. 1-3, D-
14195 Berlin, Germany.

ENT. NEWS 109(4) 274-276, September & October, 1998

Vol. 109, No. 4, September & October, 1998 275

Distribution: Brazil, Peru.

Specimens Examined: 20*0", 39 9. 1 O", Brazil, UZMH; ICf, Brazil: MatoGrosso: 1025'
S, 5928' W, 1 7-22-III, DAR; 1 9 , Brazil: Goias [lectotype of Pseudobebaeus goyazensis], BMNH;
1 9 PERU: Andres AvelinoCaceres: Satipo, 10-VIII, USNM; 1 9 , Peru: Andres AvelinoCaceres:
Tingo Maria 1km E of town, forested eastern foothills of the Andes, 5- VIII, BMNH.

DISCUSSION

Fallou (1888) described Euschistus truncatus from Minas Gerais, Brazil.
Although this species is brown and superficially resembles other species of
Euschistus Dallas, Bergroth (1892) suggested that it probably belonged in a
new genus related to Thoreyella Spinola. He later (1918) described Zorcadium
to hold this single species. This is the position in which the senior author
(DAR) treated this taxon in his (Rider 1994) conspectus of the tribe Procleticini.

Distant (1911) described Pseudobebaeus goyazensis apparently from a
single female specimen from Goias, Brazil. Although Distant did not specifi-
cally state that this species belonged with the acanthosomatids, it is obvious
that this was his intended placement. His paper does not contain any subhead-
ings below Pentatomidae; it does treat (in order) scutellerids, pentatomids
(pentatomines), pentatomids (asopines), and acanthosomatids. He placed
Pseudobebaeus between the asopines and the acanthosomatids and stated that
it was "allied to Bebaeus, Dall., in general appearance and structure of head
and lateral pronotal angles; ..." Bebaeus Dallas is properly a member of the
Acanthosomatidae. Kumar (1974) followed Distant in placing this species in
the family Acanthosomatidae and further placed it in the Blaudusinae: Lanopini.
The junior author (CF), after examining the female lectotype, presented con-
vincing evidence that this species did not belong in the Acanthosomatidae
(Fischer 1996), but its specific placement within the Pentatomidae remained a
mystery.

DAR, while reviewing the above paper by CF, noticed that the illustrations
of Pseudobebaeus goyazensis were similar to his own illustrations of Zorcadium
truncatum presented in the procleticine conspectus. Further discussion, and a
re-examination by DAR of the lectotype of Pseudobebaeus goyazensis con-
firmed that the two species were conspecific. Although the holotype of
Euschistus truncatus is apparently lost (Rolston and McDonald 1981 ), its iden-
tity has not been in question; a voucher specimen designated by Rolston and
McDonald ( 1 98 1 ) was examined. Pseudobebaeus has priority over Zorcadium,
but P. goyazensis will fall as a junior synonym of Z. truncatum. Thus the
proper combination is Pseudobebaeus truncatus (Fallou, 1888). Its proper
placement is within the tribe Procleticini in the Pentatomidae.

Note: The first line on p. 217 of the diagnosis for Zorcadium in Rider
(1994) should read "Juga contiguous anteriorly; ..." with the "not" omitted.

276 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

We thank the following individuals for their help in lending specimens pertinent to this
project (acronyms are those used in the text; DAR is senior author's collection): Janet Margerison-
Knight, British Museum (Natural History), BMNH; T. J. Henry, Systematic Entomology Labora-
tory, USDA, c/oUnited States National Museum of Natural History, USNM; A. Jansson, Zoo-
logical Museum, University of Helsinki, UZMH. We also thank J. E. Eger, DowElanco, Tampa,
FL, and L. H. Rolston, Baton Rouge, LA, for their reviews of an early draft of the manuscript.

LITERATURE CITED

Bergroth, E. 1892. Notes synonymiques. Revue d'Entomol. 1 1:262-264.

Bergroth, E. 1918. Hendecas generum Hemipterorum novorum vel subnovorum. Ann. Musei

Nationalis Hungarici 16:298-314.
Distant, W. L. 1911. Rhynchotal notes. 53 Neotropical Pentatomidae. Ann. Mag. Nat. Hist.

(8)7:242-258.

Fallou, J. 1888. Hemipteres nouveaux recueilles a Minas Geraes. Le Naturaliste (2)1:36.
Fischer, C. 1996. On the systematic position of Pseudobebaeus goyazensis Distant, 1911 within

the Pentatomoidea (Heteroptera, Pentatomoidea). Deutsche Entomol. Zeitschrift 43(1): 83-

87.

Kirkaldy, G. W. 1909. Catalogue of the Hemiptera (Heteroptera) with biological and anatomi-
cal references, lists of foodplants and parasites, etc. Vol. I. Cimicidae. Berlin, xl + 392 pp.
Kumar, R. 1974. A revision of world Acanthosomatidae (Heteroptera: Pentatomidae): Keys to

and descriptions of subfamilies, tribes, and genera, with the designation of types. Austr. J.

Zool., supplement 34: 1-60.
Lethierry, L. and G. Severin. 1 893. Catalogue general des Hemipteres. Bruxelles, Pentatomidae,

1: x + 286 pp.
Rider, D. A. 1994. A generic conspectus of the tribe Procleticini Pennington (Heteroptera,

Pentatomidae), with the description of Parodmalea rubella, new genus and species. J. N. Y.

Entomol. Soc. 102(2): 193-221.
Rider, D. A. 1997. Rolstoniellini, replacement name proposed for Compastini Distant, 1902, a

tribal name based on a generic junior homonym (Heteroptera: Pentatomidae: Pentatominae).

J. N. Y. Entomol. Soc. 103(4):40 1-403.
Rider, D. A. and L. H. Rolston. 1995. Nomenclatural changes in the Pentatomidae (Hemi-

ptera-Heteroptera). Proc. Entomol. Soc. Wash. 97(4):845-855.
Rolston, L. H. and F. J. D. McDonald. 1981. Conspectus of Pentatomini genera of the western

hemisphere - part 2 (Hemiptera: Pentatomidae). J. N. Y. Entomol. Soc. 88(4)[1980]:257-

272.

ANNOUNCEMENT

After a number of years without an editor, the Memoirs of the American Entomological
Society is now under the stewardship of Norman E. Woodley. The Society is therefore eager to
continue publication of this long-standing monographic series. Potential manuscripts or propos-
als for submissions are welcome at any time. Correspondence about manuscripts should be sent
to:

Norman E. Woodley
Systematic Entomology Lab-USDA
c/o Smithsonian Institution NHB-168
Washington, DC 20560-0168

Memoirs are normally taxonomic or faunistic in subject matter. Manuscripts and plates should be
substantial enough to produce a minimum of 75 printed pages. Authors will be expected to have
funding to pay for a substantial portion of the cost of publication.

Vol. 109, No. 4, September & October, 1998 277

MASS APPEARANCE OF LADY BEETLES
(COLEOPTERA: COCCINELLIDAE)
ON NORTH CAROLINA BEACHES 1

C. A. Nalepa 2 , K. R. Ahlstrom 2 , B. A. Nault 3 , J. L. Williams 4

ABSTRACT: A mass appearance of lady beetles on the North Carolina coast was investigated in
May of 1996. Six species of lady beetles were identified, with Hippodamia convergens and
Coccinella septempunctata predominating. It is suggested that the insects were first-generation
adults dispersing from senescing grain fields.

Several publications document the sudden appearance of hordes of lady
beetles (Coccinellidae) on the beaches of oceans and large lakes (Oliver, 1943;
Hagen, 1962; Rothschild, 1971;Yanet al., 1983; Majerus and Majerus, 1996).
These sporadic mass appearances are not associated with dormancy or aggre-
gation and are usually attributed to the weather. Wind patterns concentrate
masses of flying beetles and drop them into bodies of water; large numbers of
beetles subsequently wash up on beaches as the result of wind and tides. The
number of beetles involved can be staggering. Oliver (1943), for example,
described a drift line of dead Coccinella undecimpunctata L. at least 13 miles
long with 70,000 beetles per linear foot. In the United States the phenomenon
has been reported by Lee (1980) in the Great Lakes of the upper midwest, and
by Schaefer et al. (1987) along the coast of Delaware. Hagen (1962) reported
that masses of Hippodamia convergens Guerin-Meneville are occasionally
deposited in the Pacific Ocean.

We had the opportunity to investigate reports of a large number of
coccinellids washed up on a beach in the city of Kitty Hawk (36.07N, 75.72W)
on one of North Carolina's barrier islands. Local residents reported that the
lady beetles arrived in large numbers on 18 May 1996. On 25 May 1996 we
collected and identified 919 insects from debris east of the primary dune. Of
these, 96% were predaceous coccinellids, 3% were other Coleoptera, and 1%
were assorted Hemiptera and Diptera. Seven percent of the insects were alive
when collected, and all but the Diptera were identified to species (Table 1 ). Six
species of lady beetles were collected, with Hippodamia convergens and

1 Received November 8, 1997. Accepted January 22, 1998.

2 Beneficial Insects Laboratory, North Carolina Department of Agriculture, P.O. Box 27647,
Raleigh, NC 27611.

3 Eastern Shore Agricultural Research and Extension Center, Virginia Polytechnic Institute and
State University, 33446 Research Drive, Painter, VA 23420-2827.

4 Planning and Environmental Branch, North Carolina Department of Transportation, P.O. Box
25201, Raleigh, NC 27611.

ENT. NEWS 109(4) 277-28 1 , September & October, 1998

278 ENTOMOLOGICAL NEWS

Coccinella septempunctata L. dominating (55.8% and 41 .5% of Coccinellidae
collected, respectively).

To determine if this mass appearance was a localized anomaly, one week
later (1 June 1996) we visited Wrightsville Beach, NC (34.2 1N, 77.80W),
281 km south of Kitty Hawk. Seventy-two dead beetles were collected in de-
bris at this location: H. convergens (51.4%), C. septempunctata (44.4%), C.
munda (2.8%), and C. maculata lengi (1.4%). The three non-coccinellids col-
lected were identified as Chrysomela scripta (Fabricius). These beetles were
all dead and infrequently encountered. We suspect that the insects collected at
both Kitty Hawk and Wrightsville Beach were remnants of the same phenom-
enon, but most of the beetles at the latter location had washed or blown away
by the time we visited.

The species collected at Kitty Hawk are a common assemblage of lady
beetles in North Carolina and are reported in varying proportions from crops
and ornamental plantings (Kidd, 1996; Nault, unpublished data; Nalepa, un-
published data). All are aphidophagous to varying degrees, conforming to the
prevailing hypothesis that mass appearances of lady beetles on beaches are
related to the nature of their aphid diet (Hodek, 1973; Hodek et al., 1993).
Aphids rapidly increase in number under favorable conditions, but this abun-
dance is sporadic and ephemeral in most habitats (Hodek, 1973). Aphido-
phagous lady beetles, in turn, have evolved two traits that predispose them to
tracking prey of this nature. First, they are able to respond to an abundance of
prey with spectacular increases in population size (Hagen, 1962; Hodek, 1973;
Hodek and Honek, 1996; Majerus and Majerus, 1996). Dickson et al. (1955),
for example, estimated that nearly 54,000 adult coccinellids emerged from
one acre of alfalfa heavily infested by aphids. Second, aphidophagous lady
beetles are more nomadic than species that use other food sources, and may
switch among several habitats with suitable prey during one vegetational sea-
son. They are especially prone to fly when hungry (Ewert and Chiang, 1 966a,b;
Hodek et al., 1993; Hodek and Honek, 1996; Majerus and Majerus, 1996).

In explaining the mass shoreline appearances of aphidophagous coccinellids,
Hagen (1962) proposed a plausible chain of events subsequently echoed and
endorsed by other authors (Hodek, 1973; Hodek and Honek, 1996; Majerus
and Majerus, 1996). Favorable environmental conditions, i.e., massive aphid
populations and optimal weather, allow for a high fecundity of
female coccinellids and a low mortality of larvae and pupae. Juvenile popula-
tions build quickly, and when the young adult beetles of this generation emerge,
there is stiff competition for remaining prey. Hunger increases their mobility,
and a hot day may bring them into the air by the millions; beetles in flight may
be further concentrated by thermals and prevailing winds. The insects are
brought back to earth en masse by air currents at the coast and perhaps a reluc-
tance to cross expanses of water. Those that land in water are washed back
onto the coast by wave action and tides.

Vol. 109, No. 4, September & October, 1998 279

Hodek and Honek (1996) consider the species composition of these mass
appearances purely accidental "pseudo-communities" that may not resemble
coccinellid communities of any habitat in the vicinity; they cite Klausnitzer's
(1989, 1992) work on the German coast of the Baltic Sea. This researcher
compared relative abundance of coccinellid species from seashore collections
with those present in nearby pine forests and found little correlation. The tim-
ing and species composition of the mass appearance of coccinellids on North
Carolina beaches in 1996, however, suggests the possibility that these origi-
nated in grain fields prevalent in the eastern half of the state. First, nearly
700,000 acres of small grains were harvested in this area of North Carolina
during 1995; harvest typically begins in late May and early June (Meadows,
1996). Second, adults of the first generation of coccinellids emerge in late
May, at about the same time grain is senescing (Kidd, 1996; Nault, unpub-
lished data). Third, over most of North Carolina prevailing winds near the
earth's surface blow from the southwest. The direction may be interrupted and
reversed due to offshore storms or diurnal fluctuations (Hardy et al., 1967).
Fourth, two of the major species comprising the beach population were also
abundant in nearby grain fields. Lady beetles swept from wheat at the Tidewa-
ter Research Station near Plymouth in Washington County on 3 May 1996
consisted of 38.3% C. septempunctata, 30.4% H. convergent, and 31.3% C.
maculata (n = 240) (Kidd, 1996).

The presence of C. septempunctata and H. convergent at the beach is not
difficult to explain. C. septempunctata is primarily an aphid predator (Gordon,
1985) prone to population explosions (Hodek and Honk, 1996; Majerus and
Majerus, 1996), is a strong flier (Marriner, 1939), and is the dominant species
collected from mass aggregations on coastlines (Rothchild, 1971; Van et al.,
1983; Schaefer et al., 1987). During the breeding period, the most important
movements of C. septempunctata in Europe occur after aphids disappear from
cereal stands, when the emergence of new adults more or less coincides with a
decline in aphid populations in the fields (Hodek and Honek, 1996). H.
convergent represented a higher proportion (55.8%) of our beach collection
than has been reported in the past. In the coastal collection described by Schaefer
et al. (1987), for example, just 5.3% were identified as H. convergent. This
coccinellid is strictly aphidophagous, and can be the most abundant species
present in cereals (Gordon, 1985; Hodek and Honk, 1996: Table 5.16). If the
lady beetles that appeared on the North Carolina coast in 1996 indeed origi-
nated from small grain, then Coleomegilla maculata is conspicuous by its rela-
tive absence from the beach. Although its scarcity might be due to variation in
demographic parameters (i.e., adults of the first generations of C. septem-
punctata and H. convergent may have emerged and flown while C. maculata
were still pupae), we think a better explanation lies in host range differences
among species. While the primary food source of C. septempunctata and H.
convergent is aphids, C. maculata is perhaps the most polyphagous lady beetle

280 ENTOMOLOGICAL NEWS

known, feeding on aphids, other insect prey, insect eggs, fungi, and pollen
(Hodek, 1973; Hilbeck and Kennedy, 1996). Up to 50% of the diet of C.
maculata can be composed of pollen from various plants (Forbes, 1883). As
such, the life history of this species is not strongly tied to aphid demographics
(Ewert and Chiang, 1966b), and it is less prone to long distance movements
(Hodek and Honek, 1996). After the collapse of aphid populations in small
grain, new adults of C. maculata can support themselves on nearby alternative
food instead of undertaking a risky dispersal flight in search of aphid prey.
Voucher specimens have been deposited in the North Carolina Department of
Agriculture Insect Collection, Raleigh.

Table 1. Insect species collected from beach debris at Kitty Hawk, North Carolina, on 25 May
1996 (n = 919 insects; 7 Diptera were not identified).

Family

Species

No.

Coccinellidae

Hippodamia convergens Guerin-Meneville

493

Coccinella seplempunctata L.

367

Cycloneda nninda (Say)

Harmonia axyridis (Pallas)

Coleomegilla maculata lengi Timberlake

Anatis labiculata (Say)

Scarabaeidae

Macrodactylus anguslatus (Beauvois)

Chrysomelidae

Diabrotica undecimpunctata howardi Barber

Chrysomela (Microdera) scripla (Fabricius)

Leptinotarsa decemlineata (Say)

Calligrapha (Coreopsomela) californica

coreopsivora Brown

Saldidae

Saldula major (Provancher)

Pentatomidae

Neottiglossa (Texas) cavifrons Stal

Cydnidae

Sehirus cinctus (Beauvois)

ACKNOWLEDGMENTS

We thank Robin Goodson and George Kennedy for commenting on the manuscript, and
alert citizen Jenny Rand for reporting the coastal coccinellid convergence.

LITERATURE CITED

Dickson, R. C., E. F. Laird, Jr., and G. R. Pesho. 1955. The spotted alfalfa aphid (yellow
clover aphid on alfalfa). Hilgardia 24: 93- 1 1 8.

Ewert, M. A. and H. C. Chiang. 1966a. Dispersal of three species of coccinellids in corn fields.
Can. Entomol. 98: 999-1003.

Ewert, M. A. and H. C. Chiang. 1 966b. Effects of some environmental factors on the distribu-
tion of three species of Coccinellidae in their microhabitat. pp. 195-219. In: I. Hodek [ed.].
Ecology of Aphidophagous Insects. Dr. W. Junk, Publishers, The Hague. 360 pp.

Vol. 109, No. 4, September & October, 1998 281

Forbes, S. A. 1883. The food relations of the Carabidae and Coccinellidae. Bull. 111. State Lab.
Nat. Hist. 1: 33-64.

Gordon, R. D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. J. N.Y.
Entomol. Soc. 93: 1-912.

Hagen, K. S. 1962. Biology and ecology of predaceous Coccinellidae. Ann. Rev. Entomol. 7:
289-326.

Hardy, A.V., C. B. Carney, and H. V. Marshall, Jr. 1967. Climate of North Carolina research
stations. NC Agric. Exp. Stn. Bull. 443, 75 pp.

Hilbeck, A. and G.G. Kennedy. 1996. Predators feeding on the Colorado potato beetle in insec-
ticide-free plots and insecticide treated commercial potato fields in eastern North Carolina.
Biol. Control 6: 273-292.

Hodek, I. 1973. Biology of Coccinellidae. Dr. W. Junk Publishers, The Hague. 260 pp.

Hodek, I. and A. Honek. 1996. Ecology of Coccinellidae. Kluwer Academic Publishers,
Dordrecht. 464 pp.

Hodek, I., G. Iperti, and M. Hodkova. 1993. Long distance flights in Coccinellidae (Co-
leoptera). Eur. J. Entomol. 90: 403-414.

Kidd, K. A. 1996. Coccinellids in wheat, 1996. Annual Report of Activities, Beneficial Insects
Laboratory, NC Dep. Agric., Raleigh, p. 17.

Klausnitzer, B. 1989. Marienkaferansammlungen am Ostseestrand. Entomol. Nachr. Ber. 33:
189-194.

Klausnitzer, B. 1992. Coccinelliden als Pradatoren der Holunderblattlaus (Aphis sambuci L.) im
Warmefruhjahr 1992. Entomol. Nachr. Ber. 36: 185-190.

Lee, R. E., Jr. 1980. Aggregation of lady beetles on the shores of lakes (Coleoptera:
Coccinellidae). Am. Midi. Nat. 104: 295-304.

Majerus, M. E. N and T. M. O. Majerus. 1 996. Ladybird population explosions. Br. J. Entomol.
Nat. Hist. 9: 65-76.

Marriner, T. F. 1939. Movements of Coccinellidae. Entomol. Rec. 51: 104-106.

Meadows, B. C. 1996. North Carolina Agricultural Statistics, 1996. NC Dept. Agric., Raleigh,
132pp.

Oliver, F. W. 1943. A swarm of ladybirds (Coleoptera) on the Libyan desert coast of Egypt
between Hammam and Abusir. Proc. R. Ent. Soc. Lond. (A) 18:87-88.

Rothschild, M. 1971. A large migration of the seven-spot ladybird (Coccinella septempunctata
L.) at Deauville, France. Entomologist (London) 104: 45-46.

Schaefer, P. W., R. J. Dysart and H. B. Specht. 1 987. North American distribution of Coccinella
septempunctata (Coleoptera: Coccinellidae) and its mass appearance in coastal Delaware.
Environ. Entomol. 16: 368-373.

Yan, J. J., Y. C. Chang and X. M. Cat. 1983. Observations on the aggregation of Coccinella
septempunctata L. (Col.: Coccinellidae) in different coastal areas. Natural Enemies of In-
sects (KunchongTiandi) 5: 100-103 [Rev. Appl. Entomol. Ser. A. 72: 2771].

PLEASE SEE ANNOUNCEMENT ON PAGE 276

282 ENTOMOLOGICAL NEWS

BIOLOGICAL AND MORPHOLOGICAL NOTES ON
DASYHELEA PSEUDOINCISURATA
(DIPTERA: CERATOPOGONIDAE) 1

Lawrence J. Hribar^

ABSTRACT: Dasyhelea pseudoincisurata larvae were collected from a waste tire and a bucket.
Larvae presumably fed in algae in these containers, but after collection they fed only on dead
insects. Larval thoracic pigmentation is described. Pupae move rapidly across the substrate with
the aid of abdominal spicules. The mean pupal period was 2.1 days. Attempts to recover eggs
from reared adults were unsuccessful.

Little is known about the immature stages of many biting midges (Cerato-
pogonidae), and even basic details of life history, including larval habitat and
feeding behavior, are poorly understood for most species. During a survey of
containers for Aedes albopictus (Skuse), a number of biting midge larvae were
collected from an old tire and from a plastic bucket. These larvae were placed
into a plastic petri dish along with water from the larval habitat. Pupae were
collected and placed into separate dishes to await adult emergence. The adults
reared from these larvae were identified as Dasyhelea pseudoincisurata Waugh
&Wirth(1976).

Dasyhelea spp. larvae may be found in a number of different habitats, in-
cluding natural cavities. Dasyhelea pseudoincisurata has been collected from
treeholes and cavities in rocks (Waugh & Wirth 1 976). Wirth & Waugh ( 1 976)
found other Dasyhelea spp. larvae in cavities in tree stumps. Collections of
Dasyhelea larvae from artificial containers are not often reported, but they are
by no means unknown. For example, Remmert (1953) described Dasyhelea
tecticola from specimens collected in rain gutters.

When D. pseudoincisurata larvae were collected, their alimentary tracts
were filled with a green material. This material was believed to be algae,
however, microscopic examination of one larva's alimentary tract revealed
mostly unidentifiable debris, although fungal spores were present. During
rearing, larvae were fed dead mosquito, chironomid, and psychodid larvae,
mosquito pupae, and a gravid adult female bibionid. These were first killed by
crushing them with forceps, after which all were consumed by the Dasyhelea
larvae. The Dasyhelea larvae began feeding at the wound site, and one
Dasyhelea larva pulled a mosquito's alimentary tract outside of its body and
began feeding on its contents. Other Dasyhelea larvae ignored the prey's ali-
mentary canal and instead fed on fat body and muscle tissue of the mosquito.
The feeding behavior of the Dasyhelea larvae resembled that of some Beizia

1 Received September 12, 1997. Accepted October 29, 1997.

2 Indian River Mosquito Control District, 5655 41st. Street, Vero Beach, FL 32967. Present ad-
dress: Monroe County Mosquito Control District, 506 106th St., Gulf, Marathon, FL 33050.

ENT. NEWS 109(4) 282-284, September & October, 1998

Vol. 109, No. 4, September & October, 1998

283

larvae (Hribar & Mullen 1 99 1 ). Some Dasyhelea larvae used their anal crochets
to hold themselves in place while feeding. It appeared that only the third and
fourth instar Dasyhelea larvae fed on mosquitoes. Twice, younger larvae were
seen attempting to feed, but these were pushed aside by the vigorous feeding
of the older larvae. Younger larvae are probably not strong enough to compete
against older larvae. Dasyhelea larvae most often are said to feed on algae and
fungi (Mullen & Hribar 1988). However, Lee & Chan (1985) reported that
Dasyhelea ampullariae Macfie larvae fed on mosquito larvae in the same
habitat. Mosquitoes and Dasyhelea grisea (Coquillett) can coexist in the same
habitat with no apparent harm to either species (Baumgartner 1986).

Thoracic pigmentation of the larvae was a brown coloration present on the
dorsal aspect which extended toward the ventral surface on the prothoracic
and mesothoracic segments. On these segments the pigment was distributed
in an easily recognizable pattern (Fig. 1 ), but the prothorax was more strongly

Fig. 1. Thoracic pigmentation of Dasyhelea larvae collected in waste tire; P - prothorax, Ms -
mesothorax, Mt - metathorax.

284 ENTOMOLOGICAL NEWS

pigmented than were other thoracic segments, with less pigmentation present
on the abdominal segments. The crochets on the anal segment were large and
arranged in a starburst pattern. The larva of D. pseudoincisurata is undescribed,
and these characters may permit larvae to be distinguished from those of other
species.

Pupae were placed into separate plastic petri dishes lined with moist filter
paper at ambient room temperature, which varied from 23 C to 26 C (74 F to
78 F). Pupae were active and used their large distinctive spicules to move
rapidly across filter paper. Average length of the pupal stage was 2.1 days; 23
adults were reared. Adult males used their foretarsi to groom their antennae.
There are long spines at the base of the tarsus, and perhaps these also are used
for grooming. Adults were provided with 10% sucrose solution as an energy
source for possible mating and oviposition, but these attempts to collect eggs
were not successful.

While laboratory conditions do not duplicate natural conditions, D.
pseudoincisurata larvae will feed on a range of offered prey. The distinctive
pattern of the larvae may be useful for species identification. I thank W.L.
Grogan, Jr., Salisbury State University, for identifying the midges. D.A.
Shroyer, Indian River Mosquito Control District, and G.L. Miller, USDA, com-
mented on the manuscript

LITERATURE CITED

Baumgartner, D.L. 1986. Failure of mosquitoes to colonize teasel axils in Illinois. J. Amer.

Mosq. Control. Assoc. 2: 371-373.
Hribar, L.J. & G.R. Mullen. 1991. Predation by Bezzia larvae (Diptera: Ceratopogonidae) on

mosquito larvae (Diptera: Culicidae). Entomol. News. 102: 183-186.
Lee, K.M. & K.L. Chan. 1985. The biology of Dasyhe lea ampullariae in monkey cups at Kent

Ridge (Diptera: Ceratopogonidae). J. Singapore Nat. Acad. Sci. 14: 6-14.
Mullen, G.R. & L.J. Hribar. 1988. Biology and feeding behavior of ceratopogonid larvae

(Diptera: Ceratopogonidae) in North America. Bull. Soc. Vector Ecol. 13: 60-81.
Remmert, H. 1953. Dasyhelea tecticola n.sp., eine Ceratopogonidae aus Regenrinnen (Diptera:

Ceratopogonidae). Beitr. z. Entomol. 3: 333-336.
Waugh, W.T. & W.W. Wirth. 1976. A revision of the genus Dasyhelea Kieffer of the eastern

United States north of Florida. Ann. Entomol. Soc. Amer. 69: 219-247.
Wirth, W.W. & W.T. Waugh. 1976. Five new Neotropical Dasyhelea midges (Diptera:

Ceratopogonidae) associated with culture of cocoa. Studia Entomol. 19: 223-236.

PLEASE SEE ANNOUNCEMENT ON PAGE 276

Vol. 109, No. 4, September & October, 1998 285

EASTWARD RANGE EXTENSION IN CANADA OF

THE ALDERFLY SIALIS VELATA (MEGALOPTERA:

SIALIDAE), AND THE POTENTIAL OF THE GENUS

AS A CONTAMINANT MONITOR 1

Isabelle Roy2, Landis Hare^

ABSTRACT: We report on a range extension for Sialis velata (Megaloptera: Sialidae) in eastern
Canada and assess the potential of the genus as a metal biomonitor.

Sialis velata Ross is reported from the province of Quebec for the first
time since its original collections (Ross 1937). Ours is the first record of any
Sialis species from the province since 1937. All previous collections of the
five Sialis species known from the province (S. velata, S. iola, S. itasca, S.
mohri and S. vagans) were from the extreme southwestern corner near the city
of Montreal ( 44N, 73W) (Ross 1937; Whiting 1991). It is now possible to
extend the range of 5. velata, north to the 47th parallel, and east to the Quebec
City region (71W). Ours is the most easterly record for S. velata in Canada.
Only two Sialis species are known to occur in Canada east of Quebec City, i.e.,
5. mohri (New Brunswick) and S. vagans (New Brunswick and Nova Scotia)
(Ross 1 937, Whiting 1 99 1 ). Because we have found Sialis larvae in the major-
ity of the Quebec lakes that we have visited, the lack of collection records in
eastern Canada is likely a consequence of a lack of effort in collecting adults
and rearing larvae rather than the rarity of the genus in eastern Canada.

Adults of S. velata (five males, three females) were reared from a large
number of larvae collected on May 1 8 1 997, from soft mud at a depth of 5 m in
a Shield lake (Lake St- Joseph; 4655'N, 7140'W). Sialis larvae for rearing
were held individually in small plastic containers, in 50 ml of water that was
renewed weekly, and fed live chironomid larvae. Gut content analysis indi-
cated that larvae in the field fed largely on chironomids and oligochaetes, as
has been reported by previous investigators (Azam and Anderson 1969,
Pritchard and Leischner 1973, Canterbury 1978). When individual larvae ex-
hibited agitated swimming behavior at the water surface they were transferred
to a small plastic container filled with a mixture of sand and soil for pupation.
In the laboratory, pupation and adult emergence were highly successful at room
temperature (10% mortality), but not at 10C (100% mortality). Our results
are consistent with those of Elliott (1996), who found that pupation success
could be described by a quadratic equation with a threshold at approximately
C and an optimum at 15.

1 Received October 20, 1997. Accepted January 21, 1988.

2 INRS-Eau, Universite du Quebec, C./P. 7500, Sainte-Foy, Quebec, Canada GIV 4C7.

3 Author to whom correspondence should be addressed.

ENT. NEWS 109(4) 285-287, September & October, 1998

286 ENTOMOLOGICAL NEWS

Our interest in the larvae of Sialis relates to their potential as a contami-
nant biomonitor by virtue of the following properties (with quotations from
Phillips and Rainbow 1993):

1 . Widespread and abundant Biomonitors should be abundant throughout
the study area. The genus occurs throughout the Holarctic region. In our
studies, larvae were collected in 2 1 of 34 Quebec and Ontario lakes sampled
in a cursory manner. This ease of collection suggests that larvae are abun-
dant when present.

2. Easily kept in the laboratory The organism used should be easy to sample
and hardy enough to survive under laboratory conditions. We have cap-
tured Sialis larvae using both grabs and diver-operated benthic nets. We
can keep Sialis larvae for periods of up to 3 months at 10C in the labora-
tory in water alone (no sediment) with weekly feeding of live chironomid
larvae. Given this ease of maintenance in the laboratory, Sialis larvae could
be used to probe mechanisms of contaminant accumulation and toxicity.

3. Large sizeThe organism should provide sufficient tissue for contami-
nant analysis. We have successfully measured several trace metals in
whole individual larvae as well as in various larval tissues (pooled samples
of several individuals; Hare et al. 1991).

4. Contaminant tolerant Contaminants should be accumulated without le-
thal impacts to the species employed. An effective biomonitor should be
found along the full range of contaminant concentrations encountered in
nature. Sialis species are found in lakes influenced by acid mine drainage
(e.g., Gatewood and Tarter 1983), or metal smelters (Hare and Tessier, un-
published), both of which tend to be characterized by low pH's (down to
pH 4) and high concentrations of trace metals (e.g., up to 14 nM total dis-
solved Cd). This is a clear indication of metal and hydrogen ion resistance
in Sialis.

5. Relation between animal and contaminant concentrations A simple
correlation should exist between the contaminant content of a biomonitor
and the average contaminant concentration in its ambient environment.
Initial analysis of data from 17 lakes suggests that Cd concentrations in
Sialis are directly related to those of the free metal ion, Cd 2 +, when the
competitive effect of hydrogen ions on biological uptake sites is taken into
account (e.g., Hare and Tessier 1996).

Given the above-described characteristics of Sialis larvae, their potential as
a contaminant biomonitor seems high. However, the effective use of these

Vol. 109, No. 4, September & October, 1998 287

larvae as metal biomonitors could be improved by acquiring more information
on their: (i) Feeding: we do not know how Sialis larvae obtain trace metals, i.e.
from the water in contact with their gills, or from the food they eat. This infor-
mation will help us to determine if food-related variables such as prey type,
trophic position and metal assimilation rates (at present largely unknown) should
be included in the development of predictive bioaccumulation models; (ii)
Burrowing: because Sialis larvae are burrowers (Charbonneau et al. 1997),
more information on the depth, form and rate of burrowing could be useful in
understanding their exposure to sedimentary metals.

ACKNOWLEDGMENTS

We thank Michael F. Whiting for the identification of S. velata. For their technical assis-
tance, we thank M.-R. Doyon, J.P. Baillargeon, B. Patry, and R. Rodrigue. Funding is acknowl-
edged from Fonds pour la formation de Chercheurs et 1'aide a la Recherche (F.C.A.R.) and Hu-
man Resources Development Canada.

LITERATURE CITED

Azam, K.M. and N.H. Anderson. 1969. Life history and habits of Sialis rotunda and S. californica

in Western Oregon. Ann. Ent. Soc. Am. 62:549-558.
Canterbury, L.E. 1978. Studies on the genus Sialis (Sialidae :Megaloptera) in Eastern North

America. Unpubl. Ph. D. thesis, Univ. of Louisville, Louisville, KY.
Charbonneau, P., L. Hare and R. Carignan. 1997. Use of X-ray images and a contrasting

agent to study the behavior of animals in soft sediments. Limnol. Oceanogr. 42: 1823-1828.
Elliott, J.M. 1996. Temperature-related fluctuations in the timing of emergence and pupation of

Windermere alder-flies over 30 years. Ecol. Ent. 21:241-247.
Gatewood, R.W. and D.C. Tarter. 1983. Life history and ecology of the alderfly, Sialis aequalis

Banks, from Flatfoot Creek, Mason County, West Virginia. Proc. W. Va. Acad. Sci. 2: 102-

113.
Hare, L., A. Tessier and P.G.C. Campbell. 1 991 . Trace element distributions in aquatic insects:

variations among genera, elements and lakes. Can. J. Fish. Aquat. Sci. 48: 1481-1491.
Hare, L. and A. Tessier. 1996. Predicting animal cadmium concentrations in lakes. Nature

380:430-432.
Phillips, DJ.H. and P.S. Rainbow. 1993. Biomonitoring of trace aquatic contaminants. Elsevier

Applied Science, NY.
Pritchard, G. and T.G. Leischner. 1973. The life history and feeding habits of Sialis cornuta

Ross in a series of abandoned beaver ponds (Insecta;Megaloptera). Can. J. Zool. 51:121-131.
Ross, H.H. 1937. Studies of Nearctic aquatic insects I. Nearctic alderflies of the genus Sialis

(Megaloptera: Sialidae), Bull. 111. Nat. History Surv. 21(3):57-78.
Whiting, M.F. 1991. A distributional study of Sialis (Megaloptera : Sialidae) in North America.

Ent. News 102:50-57.

288 ENTOMOLOGICAL NEWS

GYNANDROMORPH OF HELICOVERPA ARMIGERA
[LEPIDOPTERA : NOCTUIDAE] 1

A. Josephrajkumar, B. Subrahmanyam, V.V. Ramamurt h \ ~

ABSTRACT: A gynandromorphic moth was observed in a laboratory culture of the American
bollworm, Helicoverpa armigera, with external characters of female on the left and of male on
the right side. Dissection revealed the presence of a testis and an ovary on the trans-lateral posi-
tions of the moth, respectively.

A gynandromorph is an individual in which one part of the body is mascu-
line and the other is feminine. Among insects, bilateral gynandromorphs are
most frequent, in which the left and right halves are of different sexes (Mayr
and Ashlock, 1991). However, anteroposterior gynandromorphs and forms with
irregular mosaic-like distributions of sexual characters also are known (Richards
andDavies, 1977).

In Drosophila melanogaster of XX chromosomal constitution, a gynan-
dromorph arises through the loss of one X chromosome in one of the early
cleavage nuclei of the embryo, so that deficient (XO) cells form male tissue
while those with a full complement of sex chromosomes yield female tissues
(Wilbert, 1953). Gynandromorph also can result from the "double fertiliza-
tion" of abnormal eggs possessing two nuclei, one of which gives rise to male
and the other to female tissues (White, 1968). Some parasitic Hymenoptera
yield many gynandromorphs at unusually high temperatures (Bowen and Stern,
1966).

Lepidopteran gynandromorphs have been recognised in species with sexual
dimorphism where the male has coloration and/or pattern elements that typi-
cally differ from those of the female. Hence, notable sexual differences in
phenotype appear on the two halves of the adult. An extraordinary hybrid gy-
nandromorph containing wing-pattern genes from at least three subspecies of
Heliconius melpomene (Lepidoptera : Nymphalidae) was reported by Emmel
and Boender (1990). A rare gynandromorph of Nacophora quernaria (Lepi-
doptera : Geometridae) from Florida showed a perfectly bilateral division be-
tween the male and female with different antennae, the thorax, and the poste-
rior anal tufts of the abdomen (Kutis and Heppner, 1990).

At the Division of Entomology, Indian Agricultural Research Institute, New
Delhi, a gynandromorph of Helicoverpa armigera arose for the first time in a
laboratory colony reared on the artificial diet of Singh and Rembold (1992).
Incidentally, this has been observed in a routine experiment with plumbagin, a
napthaquinone of plant origin having insect growth regulatory activity, where
the larva received a dose of 100p,g g-' applied topically.

1 Received August 25, (1997). Accepted December 17, 1997.

2 Division of Entomology, Indian Agricultural Research Institute, New Delhi - 1 1 1 2 India.

ENT. NEWS 109(4) 288-292, September & October, 1998

Vol. 109, No. 4, September & October, 1998

289

Over 20 generations, the normal coloration of the moths in the laboratory
culture is unaffected by the diet and they perfectly resembled those collected
at light or those emerged from larvae collected from chickpea, pigeonpea and
cotton fields. The female moth of//, armigera is dull orange-brown with a ' V
shaped marking on forewing and dull black border on the hindwing. Forewing
of male is greyish-green and poorly marked beyond the transverse posterior
line (Hardwick, 1965).

The single gynandromorph specimen observed had an orange-brown col-
ored forewing on the left side and greyish-green colored forewing on the right.
The color pattern fits perfectly with that of a female on the left and male on the
right. The posterior anal tufts of the abdomen were prominent on both sides
[Fig. 1 ]. Dissection of the moth revealed the presence of a testis and accessory
glands on the left side and an ovary and colleterial glands on the right side
[Fig. 2]. The placement of sex organs is therefore opposite to that of the sexual
differences in the external color pattern. The specimen is unique in this regard.
Though the moth survived for over one week on 10% honey solution, it is not
known if the moth might have been reproductively viable. Examination of the
moths of this species from the National Pusa Collection of our Division has
not revealed the occurrence of any specimen with such a mosaic of external
coloration.

Fig. 1 Gynandromorph of Helicove rpa armigera (Hubner) Left side forewing : female pattern,
right side : male pattern ( Scale line = 0.5 cm ).

290

ENTOMOLOGICAL NEWS

Fig. 2 Reproductive organs of the gynandromorph ( Note the testis (T) on the left side and the
ovary (O) on the right side, the opposite placement of the wing pattern) (Scale line = 2.0 mm).

DISCUSSION

Females of Lepidoptera are heterogametic (ZW) whereas males are ho-
mogametic (ZZ). The chromosomal constitution of eggs, rather than the sperm
cells, determine the sex of the progeny. Gynandromorphs may arise due to loss
of the Z chromosome during early zygotic divisions or due to double fertiliza-
tion of binucleate (ZW) eggs. The loss of a Z chromosome in ZZ zygotes is a
major cause of gynandromorphism (Robinson, 1971). This results in an em-
bryo that is both female (ZO) and male (ZZ). The gynandromorph noticed in
this study is an interesting case and deserves careful analysis. We found no
reference to such a condition in the literature on gynandromorphs.

Mitotic spindle orientation at the first zygotic division is always random.
In D. melanogaster subsequent mitotic products of the first two daughter nu-
clei do not intermingle before they migrate to the cellular blastoderm. Conse-
quently, a half male/female gynandromorph shows large contiguous patches
of male/female tissue, whose pattern of distribution is governed by the orien-
tation of the first zygotic spindle. A left versus right symmetrical distribution
of male versus female cuticular tissue in this H. armigera gynandromorph

Vol. 109, No. 4, September & October, 1998 291

specimen demonstrates that early blastoderm cells in H. armigera do not inter-
mingle as in D. melanogaster.

Gynandromorphs are powerful tools for following the clonal history of
cells and producing fate maps of the blastoderm (Hotta and Benzer, 1972).
The adult wings are derived from wing imaginal discs while the male and
female somatic reproductive structures are mesodermal in origin. Ferrus and
Kankel (1981), in a mosaic analysis of cuticular and muscular tissues in D.
melanogaster, showed that wing epithelial cells often attach to clonally unre-
lated muscle cells. Even though the converse was not stated by Ferrus and
Kankel (1981), it implies that clonally unrelated cells come to occupy extrin-
sically-related positions due to extensive cell movement in Drosophila during
embryogenesis. Comparatively little is known about embryogenesis and mor-
phogenetic movements in Helicoverpa. Considering that male and female re-
productive cells of mesodermal origin occupy positions trans-lateral to clonally-
related epidermal cells in our//, armigera gynandromorph specimen, we con-
clude that similar extensive cell movements occur in H. armigera during me-
sodermal differentiation also.

The influence of pesticidal molecules, including plumbagin, on epistatic
sex determination has not been well-studied. Intersexes arise due to distur-
bances of the epistatic relationship between male and female determining genes
during development. Such forms are common in Aedes sp. exposed to higher
temperatures and in Lymantria dispar when genetically distinct strains are
crossed (Richards and Davies, 1977).

Therefore, the insect under study can at the best be regarded as a natural
gynandromorph occurring at an extremely low frequency (0.000125%, i.e.,
one out of 8000 insects reared so far), as also observed in N. quernaria by
Kutis and Heppner ( 1 990) and in //. melpomene by Emmel and Boender ( 1 990).

ACKNOWLEDGMENTS

The authors are extremely grateful to N. Ramakrishnan, Division of Entomology and Shanti
Chandrasekharan, Division of Genetics, I.A.R.I., New Delhi for reviewing the manuscript.

LITERATURE CITED

Bowen, W. R. and V. M. Stern. 1966. Effect of temperature on the production of males and

sexual mosaics in an uniparental race of Trichogramma semifumatum (Hymenoptera :

Trichogrammatidae). Ann. Entomol. Soc. Am. 59: 823 - 834.
Emmel, T. C. and R. Boender. 1990. An extraordinary hybrid gynandromorph of He liconius

melpomene subspecies (Lepidoptera : Nymphalidae). Trop. Lepid. 1: 33 - 34.
Ferrus, A. and D.R. Kankel. 1981. Cell lineage restrictions in Drosophila melanogaster. The

relationship of cuticular to internal tissue. Dev. Biol. 85 : 485 - 504.
Hardwick, D.F. 1965. The corn earworm complex. Memoirs of the Entomological Society of

Canada. No.40. 246pp.
Hotta, Y. and S. Benzer. 1972 . Mapping of behaviour in Drosophila mosaics. Nature 240 : 527-

535.

292 ENTOMOLOGICAL NEWS

Kutis, J. S. and J. B. Heppner. 1 990. Gynandromorph of Nacophora quernaria in Florida (Lepi-

doptera : Geometridae). Trop. Lepid. 1: 42.
Mayr, E. and P. Ashlock. 1991. Principles of Systematic Zoology. 2nd ed. McGraw Hill Inc.

London. 475 pp.
Richards, O. W. and R. G. Davies. 1977. Imms' General Textbook of Entomology. 10th ed.

Vol.1, Chapman and Hall Ltd., London. 418 pp.

Robinson, R. 1971. Lepidoptera Genetics. Pergamon press. Oxford 687 pp.
Singh, A. K. and H. Rembold. 1992. Maintenance of the cotton bollworm Heliothis armigera

(Hubner) (Lepidoptera : Noctuidae) in laboratory culture - I. Rearing on semisynthetic diet.

Insect Sci. Applic. 13 : 333 - 338.
White, M. J. D. 1968. A gynandromorphic grasshopper produced by double fertilization. Aust.

J. Zool. 16: 101 - 109.
Wilbert, H. 1953. Normales und experimental beeinflusstes Auftreten von Mannchen und

Gynandromorphen der Stabheuschrecke. Zool. Jb. (Allg. Zool.) 64: 470 - 495.

SOCIETY MEETING OF APRIL 22, 1998

Andrew Short

Glasgow High School/Univesity of Delaware

ENGINEERED STREAM SYSTEMS AS TOOLS
FOR MACROINVERTEBRATE RESEARCH

Mr. Short has done research on the effects of elevated nutrient levels on benthic macroinvertebrates.
He has designed and used artificial stream systems that are capable of maintaining extended
pristine conditions. He received the Society's Calvert Award in 1996 for his work.

In the first part of his talk, Short discusssed the uses of artificial streams in dealing with riparian
ecosystems as well as their applications in macroinvertebrate studies. Designs of his own stream
systems were shown along with designs from Stroud Water Research Center and other engi-
neered environments. Construction materials and methods for engineered environment construc-
tion were presented. Procedures for the collection of sediments and organisms, such as commu-
nity block injection, and system operation were also given. Short described the function and uses
of algal turf scrubber technology as an advisable alternative to other water filters and purifiers.

During the second part of this talk, Short presented his current research on the effects of elevated
nutrient levels on benthic macroinvertebrates in artificial streams. Short outlined the procedures
for data collection in the engineered systems. He discussed his results, in which certain popula-
tions of macroinvertebrates, primarily Trichoptera: Hypopsychidae, experienced significant de-
clines over control populations, immediately after nutrient elevation. The nutrient factors used in
the initial trials were nitrate nitrogen and soluble phosphorus, in concentrations consistent with
poultry manure runoff that might be experienced regionally as a consequence of current agricul-
tural practices. These population declines took place without the effects of eutrophication taking
place. In subsequent trials, nitrate nitrogen elevation alone showed no negative effect on similar
macroinvertebrate populations.

- W. J. Cromartie,
Correspnding Secretary

ENT. NEWS 109(4) 292, September & October, 1998

Vol. 109, No. 4, September & October, 1998 293

A NEW SPECIES OF SWELTSA (PLECOPTERA:

CHLOROPERLIDAE)
FROM EASTERN NORTH AMERICA 1

Boris C. Kondratieff 2 , Ralph F. Kirchner 3 ' 4

ABSTRACT: Sweltsa holstonensis, new species, is described from southwestern Virginia. The
new species is most similar to Sweltsa urticae. The male epiproct of both species are illustrated
for comparison.

The genus Sweltsa in the Eastern Nearctic is represented by eight species
(Kondratieff and Kirchner 1991). These species, with the exception of S. naica
(Provancher), form a monophyletic lineage based on male epiproct shape. Two
species of this group, 5. urticae (Ricker) and S. voshelli Kondratieff and
Kirchner, are related and constitute a distinctive clade within the group. Both
species have the epiproct broadly dorsolaterally flanged and occur in small
streams of the Southern Appalachian Mountains. A third member belonging to
this clade was recently discovered in southwest Virginia by Bill P. Stark and
Ralph F. Kirchner.

Sweltsa holstonensis NEW SPECIES

(Figs. 1-2)

Male.- Body length 7.5-8.0 mm. Macropterous, length of forewing 8.5-9.0 mm. General color
bright yellow in life, yellow-white in alcohol. Head with three dark ocellar rings. Pronotum with
black margins, no median stripe. Middorsal region of abdominal terga 1-9 with black dashes or
stripes. Terga 9 with transverse ridge (Figs. 1 -2). Epiproct erectile, inflated in dorsal view (Fig. 1 ),
forming a flange laterally for most of its length, covered with appressed golden hairs; in lateral
view deeply incurved on distal fourth forming a dorsally directed hook, apex projecting above
dorsal plane and not transversely compressed; ventral aspect convex in lateral view (Fig. 2).
Female.- Body length 9.5-10.5 mm. Macropterous, length of forewing 10-10.5 mm. General
color and pattern as male. Subgenital plate about as long as wide, lateral margins slightly incurved;
apex acutely rounded, about one-fifth width of plate.

Types.- Holotype male, Washington County, Virginia, Little Moccasin Creek, County Rt. 690,
15 May 1997, B. P. Stark and R. F. Kirchner. Paratypes, same data as holotype, 2 males; same
data as holotype but 19 May 1998, R. F. Kirchner, 3 males, 3 females. The holotype will be
deposited in the United States Museum of Natural History, Smithsonian Institution, Washington
D.C., and paratypes in the collection of R. F. Kirchner and the C.P. Gillette Museum of Arthro-
pod Diversity, Colorado State University.

1 Received May 4, 1998. Accepted August 13, 1998.

2 Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort
Collins, CO 80523.

- Department of the Army, Huntington District Corps of Engineers, Water Quality Section, P.O.
Box 9, Apple Grove, WV 25502.

4 The views of this author do not purport to reflect the position of the Department of the Army or
the Department of Defense.

ENT. NEWS 109(4) 293-295, September & October, 1998

294

ENTOMOLOGICAL NEWS

Etymology.- The specific epithet refers to the Holston Valley of the Great
Appalachian Valley of Virginia. In Virginia, the Great Valley is divided into
the Shenandoah, Roanoke, New, and Holston Valleys based on drainage
systems. Little Moccasin Creek flows into the North Fork of the Holston River
at Holston.

Diagnosis.- Males of S. holstonensis can be distinguished from S. urticae by
the hook of the epiproct lacking the transversely compressed apex (Fig. 3),
epiproct ventrally convex, hook of epiproct usually exceeding the level of the
dorsal plane, and flange of epiproct covered with appressed golden hairs.
Additionally, the male of S. holstonensis can be separated from the only other
similar species, S. voshelli, by an epiproct only 2 to 3 times as long as the
greatest width and not gradually tapering to the apex (Figs. 2 and 3, Kondratieff
and Kirchner 1991). The female is similar to both S. urticae and S. voshelli,
but can be distinguished from S. voshelli by the acutely rounded apex of the
subgenital plate (S. voshelli, the apex is broadly rounded), and from S. urticae
by the longer and narrower apex, about one-fifth width of subgenital plate.

Remarks.- Little Moccasin Creek is a high gradient mountain stream originating
at Low Gap (1 150.3 m) between the Clinch and Brumley Mountains. It flows

Fig. 1 .-3. Sweltsa holstonensis. 1 . male terminalia, dorsal. 2. epiproct, lateral view. Sweltsa urticae.
3. epiproct, lateral view; inset, tip.

Vol. 109, No. 4, September & October, 1998 295

south 8.04 km to its confluence with the North Fork of the Holston River
(438.9 m) at Holston, just northwest of Abingdon. Most of the original forest
was cleared for upland pasture. Virginia's threatened peltoperlid, Tallaperla
lobata Stark, was also collected from Little Moccasin Creek near Low Gap
(Kondratieff and Kirchner 1991). Other species collected with 5. holstonensis
included Alloperla usa Ricker, 5. lateralis (Banks), S. onkos (Ricker), and
Yugus n. sp.

ACKNOWLEDGMENTS

We thank Bill P. Stark, Mississippi College for making the material available. Lynn C. Bjork
produced the illustrations.

LITERATURE CITED

Kondratieff, B. C. and R. F. Kirchner. 1991. New Nearctic Chloroperlidae (Plecoptera). J.
N.Y. Entomol. Soc. 99: 199-203.

Kondratieff, B. C. and R. F. Kirchner. 1991. Stoneflies. Pp. 214-225. In: Virginia's Endan-
gered Species. K. Terwilliger (ed.). McDonald & Woodward Publ. Co., Blacksburg, VA.

BOOKS RECEIVED AND BRIEFLY NOTED

RECENT ADVANCES IN ARTHROPOD ENDOCRINOLOGY. G.M. Coast
& S.G.Webster, eds. 1998. Cambridge Univ. Press. 406 pp. $1 10.00 (hardcover).

Nearly fifty contributors describe current work in selected areas of arthropod endocrinology
and highlight directions future studies may take. Endocrine mechanisms are dealt with in the first
sixteen chapters, while the final two chapters are concerned with peptide processing and the
development of stable lipophilic peptidomimetics.

THE BUTTERFLIES OF WEST VIRGINIA AND THEIR CATERPILLARS.
Thomas J. Allen. 1997. Univ. of Pittsburgh Press. 388 pp. 50 plates in color.
$37.50 (cloth); $22.95 (trade paper).

Descriptions of 1 28 species of butterflies, along with their caterpillars and pupae, found in
West Virginia. Each species account provides a description and information on distribution, habitat,
life history, nectar sources, and larval host plants. Twenty of the fifty colored plates depict larvae
and pupae, many not published elsewhere. Included are chapters on studying butterflies and
butterfly gardening.

INSECT HORMONES. H. E. Frederick Nijhout. A 1998 paperback edition of
a 1994 edition. Princeton Univ. Press. 267 pp. ($19.95 (paper).

The emphasis in this book on insect endocrinology is on the biology of the organism and the
ways in which physiological and developmental regulatory mechanisms are integrated into the
insect's life cycle.

296 ENTOMOLOGICAL NEWS

SOCIETY MEETING OF MARCH 25, 1998

Dr. Joseph K. Sheldon and Mr. Jeff Erikson
Department of Natural Science, Messiah College

SERPENTINE BARRENS OF PENNSYLVANIA AND MARYLAND

Dr. Sheldon began by explaining the nature of serpentine barrens, where they are located,
and why they are important ecologically. It was pointed out that the unique soil conditions have
resulted in an unusual plant community. The goal of the research currently being conducted by
Dr. Sheldon and Mr. Erikson is to determine whether an equally unusual community of msects is
associated with the unique plant community in the Nottingham and Goat Hill Serpentine Bar-
rens.

Dr. Sheldon briefly described the survey work already completed on the Lepidoptera by The
Nature Conservancy: five species of butterflies and 1 1 species of moths from Nottingham are
listed as rare, threatened, or endangered. Seven endangered Lepidoptera have been recorded
from Goat Hill. Dr. Sheldon's work on the Acrididae has found 18 species within the two bar-
rens. Grasshoppers are seasonally active within the barrens from April until the first killing frosts
of October or November. No species of special concern (as monitored by The Nature Conser-
vancy) have been encountered to date in the study. It was pointed out that this is not surprising
since our current knowledge of Acrididae is not sufficient for most species to be evaluated in
terms of their rarity. Additional natural history work on most species across their ranges will be
required.

The purpose of Mr. Erikson 's study was to gather baseline macroinvertebrate community
data and to determine if a proposed trout hatchery could utilize Black Run's water. Black Run is
a three mile long tributary which flows over serpentine geology in Nottingham Barrens on its
way to Ocotaro Creek (a tributary to the Susquehanna River).

Jeff reported on two unique chemical characteristics at Black Run: a low calcium hardness
and a high pH. These parameters did not negatively affect the macroinvertebrate community
richness. Over the past nine months he collected more than 100 genera of macroinvertebrates
among four sites along this stream. He outlined the different macroinvertebrate orders found and
showed their spatial and temporal distributions.

Jeff reported a phoretic relationship between Chironomidae: Plecopteracoluthus and
Corydalidae: Anchytarsus at Black Run.

He concluded that Black Run has a diverse macroinvertebrate community, which could be
utilized by trout as food, but he has yet to determine water temperatures and pH levels during the
summer season to see if these parameters are too high for trout survival.

In notes of entomological interest. Dale Schweitzer reported that the past winter moth sea-
son was the worst ever in southern New Jersey, but that butterflies were present unusually early:
two or three species in January, as many as five species in an hour of searching in February.
Included were numerous male Celastrina "Azures." Butterfly numbers went down in March,
following a cold spell.

- W. J. Cromartie,
Correspnding Secretary

ENT. NEWS 109(4) 296, September & October, 1998

Study notices, want items and for sale notices are published in The Entomologist's
Market Place .

Reprints: (without covers) may be ordered when corrected page proofs are returned to the
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The Entomologist's Market Place

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An Introduction to the Aquatic Insects of North America edited by Richard W. Merrill and
Kennelh W. Cummins (1995/880 pages/wire coil7$69.95*/lSBN 0-7872-3241-6 or 1995/880
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VOL. !())

NOVI;MBI;K & m.c I:MIU:K,

I'S ISSN 0013-X72X
NO. 5

ENTOMOLOGICAL NEWS

Cerococcus michaeli (Homoptera: Cerococcidae): a

new species of false pit scale from New Zealand Paris Lambdin 291
Larva of Macrothemis inacuta (Odonata:

Libellulidae) R. Novelo-Gutierrez, A. Ramirez 301

A new species of Neotropical genus Bythonia

(Homoptera: Cicadellidae) and the female

ofB.consensa M. Felix, G. Mejdalani 307

Perlesta golconda (Plecoptera: Perlidae), a

new stonefly species from Illinois

R.E. DeWalt, B.P. Stark, MA. Harris 315
Two new species ofChorebus (Hymenoptera:

Braconidae) from Spain /. Docavo, J. Tormos 318

Dipterous parasitoids from adults of moths

(Lepidoptera) Timothy L. McCabe 325

Biology of Tanychela pilosa (Hymenoptera: Ichneumon-

idae), a parasitoid of aquatic moth Petrophila

confusalis (Lepidoptera: Pyralidae) W. Jamie son, V.H. Resh 329
New Ohio records of Corixidae

(Hemiptera) S.W. Chordas, III, B.J. Armitage 339

Two corrections: Aradidae, Tingidae

(Heteroptera) Richard C. Froeschner 343

New name for a generic homonym in Teloga-

nodidae (Ephemeroptera) W.P. McCafferty, T.-Q. Wang 344

A new North American genus of Baetidae

(Ephemeroptera) and key to Baetis

complex genera C.R. Lugo-Ortiz, W.P. McCafferty 345

A new species of the small minnow mayfly genus

Plauditus (Ephemeroptera: Baetidae)

from South Carolina W.P. McCafferty, R.D. Waltz 354

New species of Cloeon and Demoulinia

(Ephemeroptera: Baetidae) from

Madagascar C.R. Lugo-Ortiz, W.P. McCafferty 357

Aedes albopictus (Diptera: Culicidae) occurrence

throughout Tennessee, with biological notes James P. Moore 363
Distribution and habitat ofCurictapronotata

(Hemiptera: Nepidae) in southeastern

Arizona J.D. Hockstra, R.L. Smith 366

Hexamethyldisilazane, a chemical alternative

for drying insects
BOOKS RECEIVED & BRIEFLY
PUBLISHER'S STATEMENT
MAILING DATES - VOLUME 1
INDEX - VOLUME 109

aty

, D. Hawks 369
314, 324

375
375
376

TIIK AMERICAN ENTOMOLOGICAL SOCIETY

Officers for 1997-1998: President: Jon K. Gelhaus; Vice-President: Susan P. Whitney;
Recording Secretary: Jeffrey S. Erikson; Corresponding Secretary: William J. Cromartie;
Treasurer: Howard P. Boyd.

Publications and Editorial Committee: Howard P. Boyd, Chr., D. Otte, and Norman E.
Woodley.

Previous editors: 1890-1920 Henry Skinner (1861-1926); 1921-1944 Philip P. Calvert (1871-
1961); 1945-1967 R.G. Schmieder (1898-1967); 1968-1972 R.H.Arnett, Jr.: 1973-4/1974 R.W.
Lake.

Subscriptions: Private subscriptions for personal use of members of the Society, domestic and
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Vol. 109, No. 5, November & December, 1998 297

CEROCOCCUS MICHAELI (HEMIPTERA:

CEROCOCCIDAE): A NEW SPECIES OF FALSE PIT

SCALE FROM NEW ZEALAND 1

Paris Lambdin^

ABSTRACT: A new species of pit scale insect, Cerococcus michaeli, from New Zealand is de-
scribed and illustrated. Cerococcus michaeli represents the second indigenous species to New
Zealand and the eighth species from the Australian Region in the genus. This species is distin-
guished from other species in the genus by the presence of an anal shield with a dorsal opening,
spine-like apical setae on anal lobes, the occurrence of asteroform tubular ducts, and the absence
of multilocular pores in transverse abdominal rows on the ventrum. A modified key is provided
to separate C. michaeli from other known species in this taxon.

Species of false pit scales (Cerococcidae) are represented in all major zoo-
geographical regions, however, most are found in the tropical and subtropical
areas. Several species are considered important pests of agricultural crops and
ornamentals (Lambdin and Kosztarab 1977). Of the 56 species assigned to the
genus Cerococcus, only one species, C. corokiae (Maskell), has been described
from New Zealand. Recently, a new species was found while examining a
series of unidentified scale insect material from the coccoidea collection of the
National Museum of Natural History, Washington, D.C., USA. This species
represents the eighth species known from the Australian Region in this genus.

This new species is placed in the genus Cerococcus based on measure-
ments and observations on 37 morphological characteristics of the adult fe-
male. The structures evaluated are considered common to species in this genus
(Borchsenius 1959, Danzig 1980, Lambdin & Kosztarab 1977). Specifically,
the presence of dorsal 8-shaped pores, prominent anal lobes, a triangular anal
shield, and cribriform plates are consistent structures for this taxon. In addi-
tion, this species has a three-segmented labium, one segmented antennae, bilocu-
lar pores, submarginal spiracles and associated spiracular furrows lined with
quinquelocular pores on the ventrum. All measurements are rounded to the
nearest micrometer. Where possible, 10 measurements of each structure were
taken and are presented as an average followed by the range in parentheses.
Terminology follows Lambdin and Kosztarab (1977).

Cerococcus michaeli, NEW SPECIES

(Fig. la-k)

Type Locality. Coromandel, New Zealand.

Type-Material. Holotype adult (encircled, lower center) and 4 paratype
on 1 slide (No. 1651), on Dysoxylum spectabile, Coromandel, New Zealand,

1 Received: July 2, 1998. Accepted July 27, 1998.

2 Department of Entomology and Plant Pathology, University of Tennessee, Knoxville, TN. 3790 1 ,

ENT. NEWS 109(5) 297-300, November & December, 1998

298 ENTOMOLOGICAL NEWS

Description of Slide-mounted Adult Females

Body (Fig. la) pear shaped, membranous, 732 (650-820) long, 464 (410-540) wide; with
distinct anal cleft; anal lobes (Fig. Ib) elongate, with scale-like overlapping cells, 88 (78-90),
long, 49 (48-50) wide, each lobe with an outer (conical) and inner (needle-like) subapical seta,
each 7 (6-12), and an stout apical seta (needle-like) 11 (10- 12) long.

Dorsal Surface (Left half)

Anal shield (Fig. la,b) triangular, with enclosed anterior anal opening, plate 69 (65-75)
long, 54(52-55) wide. Anal ring (Fig. la,b) minute, 12(12-13) indiam., with 6 fleshy setae, each
18 (14-21 ) long. Cribriform plates (Fig. Ic) unevenly aerolated, 2 pairs, 1 pair each located on
segments 7-8 anterior to anal lobes and triangular plate; each 5 (4-7) in diam. 8-shaped pores
(Fig. Id) sparse, appearing to form a swirled pattern on derm with subcircular areas devoid of
pores, each 4 (3-5) long, 2 (2-3) wide. Setae (Fig. le) sparse, few submedial tack-like setae in
longitudinal rows, especially on 6th - 9th abdominal segments, each 2 (2-3) long, other setae
rare. Simple disc pores absent. Tubular ducts (Fig. 1 asteroform, inner ductule reduced, invagi-
nated inner end with 1 -3 minute teeth, outer ductule long and slender; scattered throughout derm,
more numerous in marginal areas, especially in posterior abdominal region; each 21 (18-25)
long, 1 (1-2) wide.

Ventral Surface (Right Half)

Antennae (Fig. Ig) one segmented, 10 (8-12) long, 9 (6-15) wide; with 6 (4-7) setae. No
associated quinquelocular pores (Fig. Ih) at base. Bilocular pores (Fig. li) subcircular, irregu-
larly spaced, most numerous in submarginal area of cephalothorax, especially around mouth-
parts, antennae, and spiracles; few occasionally on margin of abdominal segments, each 4 (3-5)
in diameter. Clypeolabral shield rectangular, 1 38 ( 1 34- 1 40) long, 115(112-118) wide. Marginal
band of 8-shaped pores (Fig. Id) extending around body to apex of anal lobes and in transverse
rows, one pore wide, on abdominal segments; each 4 (3-5) long, 2 (2) wide. Labium three-seg-
mented, triangular, 54 (5 1 -58) long, 55 (5 1 -58) wide; with 5 (5-6) pairs setae, each 5 (3-6) long.
Legs absent. Spiracles (Fig. Ij) located in submarginal area, 29 (28-31) long, 11 (11-12) wide,
atrial diameter 4 (3-5); spiracles and spiracular furrows with associated quinquelocular pores,
few 4-7 locular pores; anterior spiracle with lateral cluster of 9 (7- 1 1 ) pores, spiracular furrow
with 2 1 ( 1 5-30) pores extending from spiracle to margin; posterior furrows bifid, cluster of 8 (6-
10) pores associated with spiracle, 7 (3-1 1 ) pores in anterior branch of spiracular furrow and 3
(2-5) pores in posterior branch, each pore 4 (4-5) in diam. Multilocular pores absent. Setae (Fig.
Ik) sparse, tack-like, in segmental transverse rows, 3 pairs associated with vulva, 3 medial pairs
between antennae, and a seta associated with each spiracle, each 2 (2-3) long. Tubular ducts (Fig.
If) similar in shape and size to those on dorsum, but fewer on ventrum. Vulva large, ca. 50 in
diam., largest among known cerococcids.

Etymology. This species is named for my son, Michael, in tribute to his
interests in insects.

DISCUSSION

Morphological similarities of C. michaeli to other species in the genus
Cerococcus include: a similar body shape (pyriform), a long anal cleft and
triangular anal shield, cribriform plates, and 8-shaped pores on the dorsum.
Ventrally, this species has a three segmented labium, one segmented antennae,
and a pair of thoracic spiracles with a bifid posterior furrow lined with
quinquelocular pores. Also, the type of pores (bilocular, quinquelocular, and
8-shaped) and their arrangement on the ventrum are also similar to those of
other species in the genus. This species is easily distinguished from other
species by the presence of an asterolecaniid type of tubular duct. This is the
first cerococcid known to have such tubular ducts. The long outer ductile has

Vol. 109, No. 5, November & December, 1998

299

Fig. 1 . Cerococcus michaeli Lambdin, n. sp.: (a) dorsoventral view; (b) anal lobes and anal shield;
(c) cribriform plate; (d) 8-shaped pore; (e) dorsal seta; (0 tubular duct; (g) antenna; (h)
quinquelocular pore, (i) bilocular pore; (j) spiracle; (k) ventral seta.

300 ENTOMOLOGICAL NEWS

been reduced, but there are one to three teeth within the cup. Also, the anal
shield differs somewhat from those typically exhibited by other species of
false pit scales. There exists an opening near the anterior margin of the anal
shield for waste elimination and the shield curves ventrad, but does not join
medially. The minute anal ring is enclosed by the anal shield and has six slen-
der anal ring setae. Of the 56 known species in the taxa, only one other spe-
cies, C. gallicolus Mamet, from Madagascar is known to have six setae. Also,
there is a lack of quinquelocular pores at the base of the antennae and mul-
tilocular pores on the ventral abdominal segments. In addition, these segments
are distinguished by a transverse row of 8-shaped pores.

The other endemic species to New Zealand, C. corokiae (Maskell), is dis-
tinguished from this species by the presence of more numerous cribriform
plates, an anal ring with eight setae, an anal shield without an opening, two
sizes of 8-shaped pores, presence of a submarginal row of quinquelocular pores
extending from the antennae to the posterior spiracles, and multilocular pores
in transverse abdominal rows. C. michaeli possess some of the more primitive
traits observed in species assigned to this genus (scale-like anal lobes, the un-
evenly aerolated cribriform plates, and perhaps the tubular ducts).

This new species may be identified using a modified key (Lambdin and
Kosztarab 1977) to the adult females of Cerococcus as follows:

32. Without quinquelocular pores at base of each antenna 32a

With quinquelocular pores at base of each antenna 33

32a. Posterior spiracular furrows absent; anal shield without dorsal opening; 1 -segmented leg

stubs present; with multilocular pores in transverse abdominal rows .... indonesiensis
Posterior spiracular furrows present; anal shield with dorsal opening; 1 -segmented leg
stubs absent; without multilocular pores in transverse abdominal rows michaeli

ACKNOWLEDGMENTS

Grateful appreciation is extended to D. R. Miller, SELXARS USDA, Bldg. 046, BARC-W,
Beltsville, MD 20705, for the loan of the material studied. I also thank R. Pereira, D. Paulsen,
and Adrian Mayor, University of Tennessee, Knoxville, TN, for their reviews and comments on
the manuscript.

LITERATURE CITED

Borchsenius, N. 1959. Notes on coccid fauna of China. 7. A new family of soft scales

Lecaniodiaspididae. fam. n. (Homoptera: Coccoidea). (In Russian). Entomol. Obozr. 38: 840-

846.
Danzig, E. M. 1980. Coccoids of the Far East of USSR (Homoptera, Coccinea) with a phyloge-

netic analysis of the coccoid fauna of the world. (In Russian). Entomol. Obozr. 54:51-64.
Lambdin, P. L. and M. Kosztarab. 1977. Morphology and systematics of the adult females of

the genus Cerococcus (Homoptera: Coccoidea: Cerococcidae). Va. Polytech. Inst. State Univ.

Res. Div. Bull. 128. 252pp.

Vol. 109, No. 5, November & December, 1998 301

THE LARVA OF MACROTHEMIS INACUTA
(ODONATA: LIBELLULIDAE) 1

Rodolfo Novelo-Gutierrez^ , Alonso Ramirez^

ABSTRACT: A detailed description and illustrations of the larva of Macrothemis macula are
provided. Larva is similar to M. celeno but can be distinguished by stouter movable hook, and
larger lateral spines and dorsal portuberances on abdominal segments 8-9. Larvae were found
living in lentic environments, in muddy areas close to the shore, where emerging and floating
vegetation was present.

The Neotropical genus Macrothemis comprises 37 species described to
date (Garrison, in litt.) of which only nine occur in Mexico and Central America
(Paulson, 1982; Gonzalez-Soriano & Novelo-Gutierrez, 1996). Despite the fact
that it is a very speciose genus, its immature stages are poorly known; larvae of
only three species have been described: M. celeno (Selys) (Klots, 1932), M.
musiva Calvert (Santos, 1970), and M. pseudimitans Calvert (Limongi, 1989).
Here, we describe and illustrate the larva of M. inacuta Calvert. Terminology
of the labium follows Corbet (1953).

Macrothemis inacuta Calvert
(Figs. 1-9)

Description: Exuviae yellowish-brown, larvae brown; body short and robust; integument
covered with small spiniform setae.

Head: Two times wider than long, narrowed posteriorly, occipital margin slightly con-
cave, cephalic lobes poorly developed, not bulging (Fig. 1), covered with minute spiniform
setae and long delicate setae; compound eyes rather small, slightly more dorsal than lateral.
Antennae 7-segmented (Fig. 2), the third the longest, relative length of antennomeres: 0.5, 0.7,
1 .0, 0.6, 0.7, 0.8, 0.8; scape mostly pale with a dark dorsobasal spot, pedicel pale with a dark
ring on distal end; third antennomere reddish-brown, antennomeres 4-6 dark in basal half, pale
in apical half; last antennomere mostly pale with apex dark. Labrum bare, setose on distal
border; clypeus bare. Frons and vertex with abundant spiniform setae and long and delicate
setae. Mandibles biramous (Fig. 3), external branch with four cusps in both mandibles and a
small cusp at base of ventral cusp on right mandible, internal branch vestigial, represented only
by three, low, blunt protuberances slightly more developed on right mandible. Maxillae: Gale-
olaciniae with seven acute teeth (Fig. 4), four large and three smaller, palp with numerous,
long, stiff setae, ending in a robust spine. Labium: Prementum-postmentum articulation reach-
ing posterior margin of mesocoxae; prementum subrhomboid with 7+3 and 7+2 setae (Fig. 5a),
lateral margins with small spiniform setae, ligula prominent with distal margin very slightly
serrate and with a row of stout setae on dorsal surface, but very close to distal margin, its tip .

1 Received December 4, 1997. Accepted February 25, 1998.

2 Institute de Ecologia, A.C. Departamento de Entomologia. Apartado Postal 63, 91000 Xalapa,
Veracruz, MEXICO. E-mail: novelor@ecologia.edu. mx

-* Institute of Ecology, University of Georgia, Athens, GA 30602, U.S.A. E-mail: aramirez@
arches.uga.cc.uga.edu

ENT. NEWS 109(5) 301-306, November & December, 1998

302

ENTOMOLOGICAL NEWS

PLATE I. Figs. 1-4, Macrothemis inacuta larva. 1) Dorsal habitus of last instar larva (left legs
omitted); 2) Left antenna, dorsal view; 3) Mandibles, ventrointernal view: a, left mandible, b,
right mandible; 4) Ventral view of right maxilla (a), ventrointernal aspect of the dorsal teeth (b).

Vol. 109, No. 5, November & December, 1998 303

Labial palp with six long setae and a row of short, robust, spiniform setae on basal 0.70 of outer
margin (Fig. 5c); movable hook robust, incurved, suddenly sharply-pointed, as long as palpal
setae (Fig. 5a); distal margin of palp with seven crenations, notches between crenations deep,
diminishing in depth from outer margin to internal one, each crenation finely serrate and bearing
three stout setae, one large and two small, except the two dorsal crenations which bear only two
(Fig. 5c); inner margin of palp with a single row of stout and stiff setae; a group of 7-9 setellae at
base of palp close to articulation with prementum; surface of palp sprinkled with dark spots of
different sizes in an irregular pattern.

Thorax light brown; lateral and posterior margins of pronotum rounded; pronotal disk
yellow with an inverted, dark V-shaped spot on middle part (Fig. 1 ); anterior margin of
proepisternum with a tuft of long stiff setae; proepimeron with a longitudinal reddish-brown
stripe on its upper margin. Synthorax mostly dark, pale on sutures. Anterior and posterior wing
pads dark, their tips darker brown, reaching and surpassing posterior margin of abdominal
segment five respectively. Legs long (e.g. hind legs when fully extended reaching beyond the
level of anal pyramid); meso- and metacoxae with a distal, ventrolateral, digitiform process
(Fig. 6a); femora slightly compressed laterally, dorsal and external surfaces covered with long,
stiff setae intermingled with spiniform ones; femora and tibiae pale, with a clear pattern of
three transverse bands at basal, middle and distal portions; tarsi yellow, dark on apical 0.50 of
third tarsomere; claws simple with a pulvilliform empodium.

Abdomen reddish-brown, with a complex color pattern shown on Fig. 1; tergites 3-9 with
spine-like protuberances well developed, that on 3 almost vertical, remainder gradually di-
rected rearward (Fig. 7a); lateral margins of 2-3 slightly convex, straight on 4-7 and slightly
concave on 8-9; those of 2-7 covered with minute spiniform setae which increase in length and
robustness on 8-9; lateral spine on 8 0.65 as long as dorsal length of 8; that on 9 as long as or
longer than dorsal length of 9, reaching level of tips of cerci (Fig. 8). Gonapophyses indistin-
guishable, just a minute gonopore visible in male. Epiproct and paraprocts pyramidal, acutely
pointed, with small spiniform setae on margins; epiproct little longer than its basal width (ratio
1:0.8), in lateral view (Fig. 9), the basal 0.50 of its dorsal margin slightly concave then becom-
ing straight but slightly slanting. Cerci sharply pointed, shorter than remaining appendages.
Epiproct and basal 0.35 of cerci dark, distal 0.65 of cerci and all of paraprocts pale. Size pro-
portions: Epiproct 1 .0, paraprocts 1 .0, cerci 0.80.

Dimensions (mm): Total length including appendages 17-17.6; abdomen 10.2-10.6; hind
femur 4.2-4.4; maximum width of head 4.5-4.6; lateral spines on abdominal segment 8, 0.5; on
9, 0.6-0.7.

Material examined: 2 exuviae (Cf Cf ), reared, 5 last instar larvae (4Cf Cf, 1 9 ) MEXICO:
Veracruz; Emiliano Zapata, Miradores (Lagoon), 1000 meters above sea level (masl), 8-IX-1996,
R. Arce leg., (5Cf Cf , 1 9 ) as last instar larvae. Deposited at Institute de Ecologfa, A.C. Xalapa,
Mexico. COSTA RICA: Provincia de San Jose; Zona Protectora El Rodeo, 16-XI-1990, A. Ramirez
leg., (ICf ) as last instar larva. Deposited at Museo de Zoologia, Universidad de Costa Rica.

DISCUSSION

The larva of M. inacuta closely resembles that of M. celeno in the number
of palpal setae (6), size proportions on caudal appendages, and general stat-
ure. It differs by the larger and stouter movable hook on the labial palp, coxal
processes more developed, larger lateral spines on abdominal segments 8-9,
and larger dorsal protuberances on 3-9 (mainly those on 8-9). The larva of M.
inacuta differs from that of M . pseudinritans by the stouter movable hook on
the labial palp, the larger metacoxal digitiform process (cf. Fig. 6), and the
shape of the dorsal protuberances on 7-9 (cf. Fig. 7). The larva of M. musiva

304

ENTOMOLOGICAL NEWS

LATE II. Figs. 5-9. Morphology of Macrothemis larvae. 5) Dorsal view of prementum (a), detail
of ligula (b), and dorsointemal view of right palp (c) of M. inacuta, 6) Left metacoxae of M.
inacuta (a) and M . pseudimitans (b), showing metacoxal process (mcp), lateroventral view; 7)
Profile view of abdominal dorsal line of M. inacuta (a) and M. pseudimitans (b); 8) Partial figure
of abdominal segments 8-10 showing lateral spines on 8-9, dorsal view; 9) Left lateral aspect of

Vol. 109, No. 5, November & December, 1 998 305

differs in having only 8 premental setae on each side of middle (5+3) and only
4 palpal setae. The genus Macrothemis has not been clearly defined based
upon larvae; this is important in order to differentiate Macrothemis from the
closely related genus, Brechmorhoga. Needham and Westfall ( 1 955) tabulated
some features based exclusively on M. celeno, but this does not constitute a
generic characterization. At present, these genera can be differentiated by the
following combination of features (those of Brechmorhoga [cf. Novelo-
Gutierrez, 1995] in parentheses): Integument mainly setose (granulose), cepha-
lic lobes widely rounded, not bulging (bulging); pronotum with an inverted V--
shaped dark mark (without such a mark); inferior margin of proepimeron with
long and delicate setae (with short and robust setae); sides of abdomen convex
throughout (parallel on segments 5-7); abdominal segment 10 and anal pyra-
mid hidden laterally by the lateral spines of segment 9 (segment 10 and anal
pyramid not hidden [partially hidden in B. rapax] by spines of 9); posterior
margin of sternite 10 usually visible in dorsal view (not visible from above);
dorsal protuberances well developed on abdominal segments 3-9, often acutely-
pointed (well developed on 2-5 or 2-6, vestigial or lacking on posterior seg-
ments, often bluntly-tipped); cerci 0.75-0.80 as long as epiproct (cerci 0.50-
0.66 as long as epiproct).

Ecological notes.- Larvae of M. inacuta were found in a lagoon in open
pasture land, on the muddy bottom near shore where scattered aquatic vegeta-
tion grew. Mature larvae were caught on September 8, 1996. The lagoon was
visited one year later (September 21, 1997) but no M. inacuta larvae were
found, although teneral adults were captured in surrounding areas. These pre-
liminary observations suggest potential synchrony (or seasonality?) in emer-
gence of the larvae, probably starting at the end of August or beginning of
September. Available published information indicates that all Macrothemis
species dwell in lotic habitats (Garcia-Diaz, 1938; Limongi, 1989; Gonzalez-
Soriano, 1992). However, Santos (1970) mentioned that Macrothemis musiva
larvae probably inhabit intermediate lotic and lentic waters. M. inacuta is an-
other exception since larvae were collected in lagoons, and adults are seen at
streams and rivers. It is likely that when living in a lotic system the larvae
favor still waters in pools and areas near the margins.

ACKNOWLEDGMENTS

We thank Roberto Arce-Perez, M. Sc. for his invaluable help collecting and maintaining
alive larvae of M. inacuta. We also thank Sidney Dunkle (Piano, TX) and Michael May (New
Brunswick, NJ) for their invaluable criticism to improve the final manuscript. A. Ramirez was
supported by the National Science Foundation grant DEB95-28434 during preparation of the
manuscript.

LITERATURE CITED

Corbet, P.P. 1953. A terminology for the labium of larval Odonata. The Entomologist 86: 191-
196.

306 ENTOMOLOGICAL NEWS

Garcia-Diaz, J. 1938. An ecological survey of the fresh water insects of Puerto Rico. 1. The
Odonata: with new life-histories. J. Agric. Univ. Puerto Rico 22(1 ): 43-97.

Gonzalez-Soriano, E. 1992. Macrothemis ultima spec, nov., a new dragonfly from the state of
Jalisco, Mexico (Anisoptera: Libellulidae). Odonatologica 21(l):91-95.

Gonzalez-Soriano, E. y R. Novelo-Gutierrez, 1996. Odonata, pp. 147-167. In: J. Llorente-
Bousquets, A.N. Garcia-Aldrete y E. Gonzalez-Soriano (eds.). Biodiversidad, taxonomia y
biogeografia de artropodos de Mexico: Hacia una sintesis de su conocimiento. UNAM, Mexico.

Klots, E.B. 1932. Insects of Porto Rico and the Virgin Islands. Odonata or Dragon Flies. Scien-
tific Survey of Porto Rico and the Virgin Islands 14: 1-107.

Limongi, J.E. 1989. Estudio morfo-taxonomico de nayades de algunas especies de Odonata
(Insecta) en Venezuela (II). Memoria 49 ( 1 3 1 - 1 32):405-41 9.

Novelo-Gutierrez, R. 1995. La nayade de Brechmorhoga praecox (Hagen, 1 861 ), y notas sobre
las nayades de B. rapax Calvert, 1898, B. vivax Calvert, 1906 y B. mendax (Hagen,
1861)(Odonata: Libellulidae). Folia Entomol. Mex. 94:33-40.

Paulson, D.R. 1982. Odonata, pp. 249-277. In: S.H. Hurlbert & A. Villalobos-Figueroa (eds.),
Aquatic Biota of Mexico, Central America and the West Indies. San Diego State University,
San Diego, California.

Santos, N.D. 1970. Descricao da ninfa de Macrothemis musiva (Hagen, 1861) Calvert, 1898
(Odonata: Libellulidae). Atas Soc. Biol. Rio de Janeiro 13:157-158.

Vol. 109, No. 5, November & December, 1998 307

A NEW SPECIES OF THE NEOTROPICAL GENUS

BYTHONIA (HOMOPTERA: CICADELLIDAE) AND

THE FEMALE OF B. CON SENS A 1

Marcio Felix^, Gabriel MejdalanP

ABSTRACT: Bythonia ferruginea, new species, is described from the states of Bahia and Minas
Gerais, Brazil. The female of B. consensa is described for the first time based on specimens from
the states of Rio de Janeiro (new record) and Espirito Santo, Brazil. The presence of two sclero-
tized plates from the sternum VIII is reported in females of Bythonia for the first time. Notes on
specimens of B. kalypso from the Brazilian states of Santa Catarina (new record) and Minas
Gerais are added. A map showing the known distribution of the species of Bythonia is also pre-
sented.

Three species of the Neotropical genus Bythonia Oman were recorded by
Blocker and Webb (1990). Only a few specimens of this genus were cited in
the literature. B. rugosa (Osborn, 1923), the type-species, is known only from
the female holotype from Bolivia and a male from Peru (Linnavuori 1959).
This species was originally described in the genus Nionia Ball. B. kalypso
Linnavuori, 1959 and B. consensa Blocker and Webb, 1990 were, until the
present paper, known only from their male holotypes from Brazil. The affini-
ties of Bythonia to other leafhopper genera were briefly discussed by Blocker
and Webb (1990), who assigned this genus to the subfamily lassinae. The origi-
nal description by Oman (1936) and subsequent descriptions by Linnavuori
(1959) and Blocker and Webb (1990) should be consulted for characteristics
of the genus.

A new species of Bythonia from the states of Bahia and Minas Gerais,
Brazil, is herein described. The previously unknown female of B. consensa is
described for the first time. This species is newly recorded from the state of
Rio de Janeiro, Brazil. Notes on six additional male specimens of B. kalypso
Linnavuori are added and the species is newly recorded from the state of Santa
Catarina, Brazil.

Acronyms for collections in which the specimens herein studied are de-
posited are as follows: DZRJ (Departamento de Zoologia, Universidade Fe-
deral do Rio de Janeiro, Rio de Janeiro, Brazil), MNRJ (Museu Nacional, Rio
de Janeiro), and NHM (The Natural History Museum, London). In quotations
of label data, a virgule (/) separates lines on a label and a semicolon separates
information on different labels. Morphological terminology follows mainly

1 Received January 22, 1998. Accepted February 25, 1998.

2 Departamento de Zoologia, Institute de Biologia, Universidade Federal do Rio de Janeiro
(UFRJ), Caixa Postal 68044, 21944-970 Rio de Janeiro RJ, Brasil.

3 Departamento de Entomologia, Museu Nacional, UFRJ, Quinta da Boa Vista, Sao Cristovao,
20940-040, Rio de Janeiro RJ, Brasil. E-mail: mejdalan@acd.ufrj.br.

ENT. NEWS 109(5): 307-314, November & December, 1998

308 ENTOMOLOGICAL NEWS

Young (1977). The software package FishMap (Buckup 1995) was used for
producing a distribution map of the species of Bythonia.

Bythoniaferruginea, NEW SPECIES

(Figs. 1-7)

Diagnosis. - Males of B. ferruginea can be distinguished from the other
known species of the genus by the following features: pronotum with
arrow-shaped group of irregular black spots; sternum VIII with anterior and
posterior acute processes on lateral margins; pygofer without processes, vent-
rolateral portion with longitudinal fold; basal half of styles with blunt dorsal
process; aedeagus with pair of lateral processes on apical third and without
median spine on apodeme.

Description of male. - Length including forewings, 6.3-6.8 mm. Head (Fig. !) short, me-
dian length of crown approximately one-eighth interocular width, anterior margin rounded; sur-
face of crown with small punctures; face with pubescence; clypeus with inconspicuous muscle
impressions; clypellus in lateral view forming angle at transition with clypeus; antennal ledges
slightly protuberant; lora broad, slightly striate; genae with deep excavation below eyes. Thorax
(Fig. 1 ) with pronotal width approximately equal to transocular width of head; disc of pronotum
with pubescence, transversely striate; dorsopleural carinae complete; mesonotum transversely
striate. Forewings with clavus and adjacent regions of corium punctate. Legs pubescent. Ab-
dominal sternum VIII (Fig. 2) broad in ventral view, lateral margins with anterior and posterior
small acute processes, posterior margin broadly concave, ventral surface with microsetae uni-
formly dispersed. Remaining morphological characteristics of head and thorax as in the original
generic description of Oman (1936: 358) and the subsequent description of Linnavuori (1959:
13).

Color. - Reddish-brown marked with black. Crown (Fig. 1) almost entirely black; ocelli
yellowish. Pronotum (Fig. 1 ) with arrow-shaped group of irregular black spots; mesonotum (Fig.
1 ) with pair of subtriangular black maculae on anterior portion and median black stripe continu-
ous with pronotal arrow-shaped mark. Forewings hyaline, with irregular amber areas. Hindwings
hyaline. Clypeus and clypellus with median black stripe narrowing toward inferior portion; lora
with blackish spot. Legs with blackish markings.

Male genitalia. - Pygofer (Fig. 3) elongate in lateral view, with apex rounded, ventrolateral
portion with longitudinal fold, apical half of disc with numerous microsetae, apical margin with
macrosetae extending anteriorly along one-third of ventral and dorsal margins. Subgenital plates
(Fig. 4) in lateral view elongate, extending posteriorly beyond apex of pygofer, median portion
slightly enlarged, apex rounded, plates with dispersed microsetae. Styles (Fig. 5) in lateral view
with apical portion curved dorsally, bifurcate, C-shaped, with short triangular process and small
group of long microsetae, dorsal margin with long microsetae, basal half with dorsal blunt pro-
cess. Connective (Fig. 6) in dorsal view with very short, dorsally curved arms, dorsal area of
stalk with median keel. Aedeagus (Fig. 7) curved dorsally in lateral view, with apical digitiform
process, distal third with pair of thin lateral processes; aedeagal apodeme well developed, apex
bifurcate.

Female unknown.

Known distribution. - Brazilian states of Bahia and Minas Gerais (Fig.
1 4). The two known records of B. ferruginea are included in areas originally
covered by the Brazilian Atlantic Forest (see map in Warren 1996).

Vol. 109, No. 5, November & December, 1998

309

Figs. 1-7. Bythonia ferruginea, new species, male. 1, Head and thorax, dorsal view. 2, Abdomi-
nal sternum VIII, ventral view. 3, Pygofer, lateral view. 4, Subgenital plate, lateral view. 5,
Style, lateral view. 6, Connective, dorsal view. 7, Aedeagus, lateral view.

Etymology. - The species epithet ferruginea, is of Latin derivation and
refers to the reddish-brown color of its anterior dorsum.

Type material. - Holotype: Male, Brazil, "Encruzilhada-B A [state of Bahia, 1 5 3 1 1 S, 40
54' W]/ XI-1972/ Alvarenga/ 960 m", MNRJ. Paratypes: Two males, same data as holotype,
MNRJ and DZRJ; three males, Brazil, "Pedra Azul/ Minas [state of Minas Gerais, 16 00' S,
41 17' W], Brasil; Seabra &/ Oliveira/ XI-72", MNRJ.

Notes. - The color pattern in B. ferruginea is very similar to that of B.
consensa (see description below). The anterior dorsum (Fig. 1) in both species
is reddish-brown with black markings and the face has a median black stripe.
In terms of morphology, the sternum VIII and male genitalia of B. ferruginea

3 1 ENTOMOLOGICAL NEWS

are also similar to those of B. consensa. The sternum VIII (Fig. 2) in the new
species presents an anterior and a posterior pair of lateral processes, while in
B. consensa only a single median pair of processes is present. The pygofer
(Fig. 3) in B.ferruginea has a ventrolateral longitudinal fold, a feature that is
not observed in B. consensa. On the other hand, the ventral margin of the
pygofer in the latter species has a bifurcate process which is not present in the
former. The apical portion of the styles (Fig. 5) in both species is curved dor-
sally and bifurcate, but the basal half of this structure in B. ferruginea has a
blunt projection which does not occur in B. consensa. The aedeagus (Fig. 7) in
these species is curved dorsally and bears a pair of lateral processes. These
processes are longer in B. ferruginea. The aedeagal apodeme in B. consensa
has a spine that is not present in B. ferruginea.

Bythonia consensa Blocker and Webb

(Figs. 8-13)

Description of female.- Length including forewings, 7.6 mm. Morphological characteris-
tics of head and thorax as in B.ferruginea, new species, and also as in the generic descriptions of
Oman (1936: 358) and Linnavuori (1959: 13) and the original specific description of Blocker and
Webb (1990: 294).

Color. - Reddish-brown marked with black, ocelli yellowish. Pronotum (Fig. 8) with small
blackish areas and faint, longitudinal blackish-brown stripe on median portion; mesonotum (Fig.
8) with pair of black maculae on anterior portion and faint, longitudinal blackish-brown stripe on
median portion continuous with pronotum stripe, extending to transverse sulcus. Forewings red-
dish-brown with apex amber. Hindwings hyaline. Clypellus with small black maculae on central
portion. Legs with blackish areas.

Female genitalia.- Abdominal sternum VII (Fig. 9) narrowed posteriorly, ventral surface
strongly convex, with microsetae uniformly distributed, lateral margins without processes, pos-
terior margin shallowly emarginate on each side of median quadrangular concavity, lateral mar-
gins of emarginations with spiniform processes. Sternum VIII (Fig. 10) well developed, sclero-
tized, formed in dorsal view by pair of plates, each with deep posterior concavity. Pygofer (Fig.
1 1 ) elongate in lateral view, with apex truncate, apical half with microsetae uniformly distrib-
uted, ventroapical portion with small group of macrosetae. Ovipositor with first valvulae long
and narrow, apex blunt, dorsal and ventral sculptured areas with sinuate and almost vertically
aligned striae; second valvulae (Fig. 12) slightly expanded apically, without preapical promi-
nence, apex blunt, apical portion with broad, rectangular, sloping teeth, median portion (Fig. 13)
with small irregular denticles.

Known distribution. - B. consensa was originally described by Blocker
and Webb ( 1 990) from the state of Espirito Santo. This species is herein newly
recorded from the state of Rio de Janeiro (Fig. 14). The known records of B.
consensa are included in areas originally covered by the Atlantic Forest.

Material examined. - One female, two males, Brazil, "Angra dos Reis [state of Rio de
Janeiro, 23 00' S, 44 19' W]/ Japuhyba/ 2-1944/ Wygodzinsky L.; MNRJ", MNRJ. Two fe-
males, Brazil, "Corrego Ita [Corrego Ita]/ E. Santo, [state of Espirito Santo] Br./ X-1954/ W.
Zikan", MNRJ. Male holotype, "Tijuco Preto/ Esp. Santo [state of Espirito Santo, 20 17' S, 40
53' W]; Holotype/ Bythonia consensa/ Blocker + Webb/ 1990", NHM.

Notes. - The two above-mentioned males of B. consensa from the state of

Vol. 109, No. 5, November & December, 1998

311

Figs. 8-13. Bythonia consensa, female. 8, Head and thorax, dorsal view. 9, Sternum VII, ventral
view. 10, Sternum VIII, dorsal view. 1 1, Pygofer, lateral view. 12, Second valvula of ovipositor,
lateral view. 13, Area between arrows in figure 12 at a higher magnification, lateral view.

Rio de Janeiro were identified using the original description of Blocker and
Webb (1990) and through the examination of the species' male holotype in the
NHM. The reddish-brown color of the anterior dorsum of the females of B.
consensa is similar to that of the males. However, the latter have a longitudinal
black stripe on clypeus and blackish areas on crown and anterior portion of
pronotum that are not observed in the females. B. rugosa is the only other
species in the genus Bythonia for which the female has been described. B.
consensa can be distinguished from B. rugosa by the form of the sternum VII.
In the former species this sternum (Fig. 9) does not have processes on lateral
margins and a quadrangular concavity is present on posterior margin, while in
the latter species it has a pair of lateral spiniform processes and a posterior
trapezoidal process (see Blocker and Webb 1990).

3 1 2 ENTOMOLOGICAL NEWS

Females of B. consensa present two sclerotized plates at the base of the
ovipositor dorsad of the sternum VII (Fig. 10). These plates, which are here
reported in the genus Bythonia for the first time, are homologous to those
described by Nielson (1965) in the proconiine genus Cuerna Melichar and to
the sclerites of the genital chamber described by Young (1977) in several
cicadelline genera. They are derived from the sternum VIII, which is greatly
reduced in females of cicadellids (Nielson 1965, Mejdalani in press). Nielson
(1965) demonstrated that this modified sternum can provide useful character-
istics for distinguishing species in Cuerna. Comparative studies on females of
Bythonia are necessary in order to know if the sternum VIII is also of taxo-
nomic value in this genus. The sternum VIII in B. consensa is similar to the
derived type of modified sternum described by Nielson (1965) in Cuerna, which
is characterized by the presence of two distinct plates.

The first and second valvulae of the ovipositor of B. consensa are very
similar to those of B. rugosa. The valvulae of the latter species were described
by Hill (1970). The dorsal sculptured area of the first valvulae presents in both
species sinuate, almost vertically aligned striae. These striae differ greatly from
the alveolate sculpturing found in other lassinae (see Hill 1 970, Dietrich 1 993).
The shaft of the second valvulae in B. consensa (Fig. 1 2) and B. rugosa, unlike
those of other lassinae (see Hill 1970), does not present dorsal tooth-like promi-
nences. The apical portion of these valvulae is slightly expanded and bears
teeth in both species.

Bythonia kalypso Linnavuori

B. kalypso was described from a single male from the state of Minas Gerais
(Linnavuori 1959: 15). We have identified five additional male specimens of
this species from Minas Gerais in the MNRJ collection; one male specimen
from the Brazilian state of Santa Catarina was identified in the NHM collec-
tion (Fig. 14). This is the first record of B. kalypso from Santa Catarina. This
species also occurs in areas of Atlantic Forest. In terms of color and morphol-
ogy, these specimens agree fairly well with the original description of the spe-
cies. However, they are slightly smaller (7.5-8 mm) than the holotype (9 mm).
Males of this species can be recognized by the following characteristics: pygofer
with a triangular projection on median portion of ventral margin; apex of styles
acute; apex of aedeagus with an acute process directed anteriorly and a blunt,
weekly sclerotized process directed dorsally. The holotype of B. kalypso is
reportedly deposited in the Hungarian Natural History Museum but could not
be located in that collection (Blocker and Webb 1990).

Material examined. - Five males, Brazil, "Pedra Azul/ Minas [state of Minas Gerais, 16
00' S, 41 1 T W], Brasil; Seabra &/ Oliveira/ XI-72; MNRJ", MNRJ. One male, Brazil, "Colesao/
Campos Seabra; CorupaV S. Catarina [state of Santa Catarina, 26 25' S, 49 14' W] Brasil/
I-1954/A. Mailer", NHM.

Vol. 109, No. 5, November & December, 1998

313

Bytfionja rugosa

Bytnonja consensa
^ Bytfionja tcaJypso

Bytfionja ferruginea

Fig. 14. Known geographic distribution of the species of the South American genus Bythonia. B.
rugosa was also recorded from an unknown locality in Peru by Linnavuori (1959).

ACKNOWLEDGMENTS

The manuscript benefited from the useful comments of A. L. Carvalho, L. B. N. Coelho, L.
F. M. Dorville, M. W. Nielson, S. A. Vanin, M. D. Webb, and two anonymous reviewers. This
work was finished during a visit of GM to the NHM. Working facilities in that museum arranged
by M. D. Webb, J. Margerison-Knight, and J. Martin are greatly acknowledged. Fellowships
from Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior (CAPES, Brazil) to MF
and from Fundacao de Amparo a Pesquisa do Estado de Sao Paulo (FAPESP, Brazil) to GM are
acknowledged.

LITERATURE CITED

Blocker, H. D. and M. D. Webb. 1990. The leafhopper genus Bythonia (Homoptera:
Cicadellidae). Entomol. News 101: 293-296.

Buckup, P. A. 1995. FishMap, a software package for producing species distribution maps.
Version 1.8. User's guide. Published by the author, Rio de Janeiro.

Dietrich, C. H. 1993. A new genus of lassinae from southeastern Brazil (Homoptera: Cicadellidae).
Proc. Entomol. Soc. Wash. 95: 475-480.

Hill, B. 1970. Comparative morphological study of selected higher categories of leafhoppers
(Homoptera: Cicadellidae). Unpublished Ph. D. dissertation, North Carolina State Univer-
sity, Raleigh.

3 1 4 ENTOMOLOGICAL NEWS

Linnavuori, R. 1959. Revision of the Neotropical Deltocephalinae and some related subfami-
lies (Homoptera). Ann. zool. Soc. Zool. Bot. Fenn. 'Vanamo' 20: 1-370.

Mejdalani, G. In press. Morfologia externa dos Cicadellinae (Homoptera: Cicadellidae):
comparacao entre Versigonatia ruficauda (Walker) (Cicadellini) e Tretogonia cribrata Melichar
(Proconiini), com notas sobre outras especies e analise da terminologia. Revta bras. Zool.

Nielson, M. W. 1965. A revision of the genus Cuerna (Homoptera, Cicadellidae). Tech. Bull. U.
S. Dep. Agric. 1318: 1-48.

Oman, P. W. 1936. A generic revision of American Bythoscopinae and South American Jassinae.
Univ. Kans. Sci. Bull. 24: 343-420.

Osborn, H. 1923. Neotropical Homoptera of the Carnegie Museum. Part 2. Records and descrip-
tions of five new genera and sixty-five new species of the subfamily Jassinae. Ann. Cam.
Mus. 15: 27-79.

Warren, D. 1996. A ferro e fogo. A historia e a devastacao da Mata Atlantica brasileira. Com-
panhia das Letras, Sao Paulo.

Young, D. A. 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2.
New World Cicadellini and the genus Cicadella. Bull. N. Carol. Agric. Exp. Stn. 239: 1-1 135.

BOOKS RECEIVED AND BRIEFLY NOTED

BUTTERFLY CONSERVATION. (Second ed.) T.R. New. 1998. Oxford Univ.
Press. 248 pp. $27.95 paperback).

This book provides a broad survey of the science of insect and butterfly conservation and the
rationale for conserving both. Most of the book explores butterfly conservation and international
efforts to safeguard species, including steps individuals can take to encourage and document but-
terfly conservation.

TERMITES. BIOLOGY AND PEST MANAGEMENT. M.J. Pearce. 1998.
Oxford Univ. Press. 172 pp. 32 plates. $65.00 (cloth).

This book provides a general scientific introduction to termites, including their biology,
behavior, pest status, and control. It is directed to advanced students in entomology and pest man-
agement, as well as to professionals.

A FIELD GUIDE TO COMMON TEXAS INSECTS. B.M. Drees & J.A.
Jackman. 1998. Gulf Publishing Co., Houston, TX. 359 pp. 381 color photos
on 64 plates. $18.95 (paper).

A regional field guide describing insects, mainly by order and family, and, in the larger orders,
by genus, with specific examples.

Vol. 1 09, No. 5, November & December, 1 998 3 1 5

PERLESTA GOLCONDA (PLECOPTERA: PERLIDAE), A
NEW STONEFLY SPECIES FROM ILLINOIS 1

R. E. DeWalt 2 , Bill P. Stark 3 , M. A. Harris 4

ABSTRACT: The male, female, and egg of a new species of perlid stonefly, Perlesta golconda
are described. Most specimens were captured from the banks of the Ohio River at Golconda,
Illinois. This species is most similar to P. lagoi Stark, but may be differentiated by wing and body
coloration, the male internal genitalia, and egg morphology. The addition of P. golconda brings
to 16 the total number of known Perlesta species.

Stark (1989) recognized Perlesta placida (Hagen) as a species complex
nearly a decade ago. He described seven new species and revised and keyed
another five species from eastern North America. Shortly thereafter, Poulton
and Stewart (1991) added P. fusca from Arkansas. Recently, Kirchner and
Kondratieff (1997) described P. teaysia from Virginia, and Stark and Rhodes
(1997) added P. xube from Nebraska. During statewide monitoring of sensi-
tive aquatic insects in Illinois, this new species was collected from the banks
of the Ohio River near Golconda, Illinois.

Perlesta golconda DeWalt and Stark, NEW SPECIES

Male. Forewing length from 10.5 to 11 mm. General color pale yellow-brown. Ocellar
patch diffuse mesally, but with dark bars almost connecting lateral ocelli to anterior ocellus
(Fig. 1). Wing membrane pale amber with intercostal margin paler, veins light brown. Femora
pale yellow on anterior face, but dark brown on dorsum (Fig. 3). Paraprocts short, subapical
spine prominent (Figs. 2, 5). Tergum 10 sensilla basiconica patch sparse (Fig. 2). Penis tube
and sac moderately long and sinuate (Fig. 6); dorsal patch broad and with narrow lateral bands
of larger seta-like spines giving a distinctly darker aspect (Fig. 7). Caecum poorly developed;
ventral base of sac with a small nipple (Fig. 6). Ventral extension of aedeagus was only par-
tially everted (Fig. 6), but appeared to be long and narrow.

Female. Forewing length from 13 to 14 mm. Body and wing coloration similar to the
male. Subgenital plate lobes rounded, separated by a V-shaped notch; lobes with about 10
scattered large bristles each (Fig. 4).

Egg. The two females produced only poorly sclerotized eggs, but they were sufficiently
developed to demonstrate a smooth, to slightly granular chorion (Fig. 8). The egg collar was
sessile and the anchor fibrous.

Nymph. Unknown.

Types. The holotype male and paratypes consisting of one male and two females were
deposited at the Illinois Natural History Survey (INHS), Champaign, Illinois.

Type locality. The holotype male and a male and female paratype were collected from the
Ohio River at Golconda, Pope County, Illinois, on 25 June 1997. Universal transverse mercator
coordinates are zone 16, 368,710 m easting, and 3,600,000 m northing (1927 North American

1 Received January 22, 1998. Accepted February 25, 1998.

2 Center for Biodiversity, Illinois Natural History Survey, 607 E. Peabody Dr., Champaign, IL
61820.

3 Biology Department, Mississippi College, Clinton, MS 39058

4 United States Geological Survey, 221 N. Broadway Ave., Urbana IL 61801

ENT. NEWS 109(5): 315-317, November & December, 1998

316

ENTOMOLOGICAL NEWS

Scale, see legend

Fig. 1-8. Perlesta golconda : 1. Male head and pronotum. 2. Male tergum 10, sb - sensilla
basiconica, p=paraproct. 3. Male right fore femur. 4. Female sternum 8. 5. Male paraproct, lat-
eral, s=spine. 6. Aedeagus, lateral, c=caecum, dp=dorsal patch, n=nipple, v-ventral extension. 7.
Aedeagus, dorsal. 8. Egg. Scales: 0.6 mm (1), 0.3 mm (2-4), 0.15 mm (5-8).

Vol. 1 09, No. 5, November & December, 1 998 317

Datum). They were collected from a black light trap (2030-21 15 CST, 26.1C air temperature)
by R. E. DeWalt and L. J. Peraino. Another paratype female was taken from along the Ohio
River in Elizabethtown, Hardin County, Illinois, 30 June 1993, by M. A. Harris and H. E.
Kitchel.

Etymology. Perlesta golconda is named for the town along the Ohio River
where the majority of specimens were collected.

DISCUSSION

Perlesta golconda most closely resembles P. lagoi Stark. It may be differ-
entiated from the latter by its lighter wing and body coloration, by the dorsal
setal patch of the aedeagus forming two prominent, lateral bands (Fig. 6, 7),
by the absence of egg follicular cell impressions, and by a sessile egg collar
(Fig. 8). Perlesta lagoi has the typical dark brown wings of the genus, a dorsal
patch composed of a single, wide row of spines, and the egg has a button-like
collar and wide, circular, follicular cell impressions (Stark 1989). Additional
specimens of P. golconda may demonstrate the usefulness of the pigment pat-
terns of the ocellar patch and fore femora for identification. No nymphs of P.
golconda were available for comparison to its congeners.

Associated species in the light trap collection were the stonefly Isoperla
bilineata (Say), the mayflies Hexagenia bilineata (Say), Stenacron
interpunctatum (Say), Leucrocuta sp., Stenonema sp., the caddisflies Hydroptila
waubesiana Betten, Potamyia flava (Hagen), Neureclipsis crepuscularis
(Walker), Hydropsyche orris Ross, Nectopsyche Candida (Hagen), Ceraclea
sp., and the megalopteran Neohermes sp. Two species of Perlesta are now
known from Illinois. The other species, P. decipiens (Walsh), occurs statewide
and was reported by Harris and Webb (1995).

ACKNOWLEDGMENTS

We thank D. W. Webb, S. J. Taylor, G. A. Levin, all of the 1NHS, for reviewing earlier
drafts. Appreciation is extended to C. E. Warwick for scanning the original drawing and adding
labels. Travel expenses and publication costs were provided by the Illinois Department of Natural
Resources, Critical Trends and Assessment Project, a statewide monitoring program.

LITERATURE CITED

Harris, M. A., and D. W. Webb. 1995. The stoneflies (Plecoptera) of Illinois revisited. J. Kan.

Entomol. Soc. 67: 340-346.
Kirchner, R. F., and B. C. Kondratieff. 1997. A new species of nearctic Perlesta (Plecoptera:

Perlidae) from Virginia. Proc. Entomol. Soc. Wash. 99: 290-293.
Poulton, B. C., and K. W. Stewart. 1991. The stoneflies of the Ozarks and Ouachita Mountains

(Plecoptera). Mem. Am. Entomol. Soc. 38: 1-116.
Stark, B. P. 1989. Perlesta placida (Hagen), an eastern nearctic species complex (Plecoptera:

Perlidae). Entomol. Scand. 20: 263-286.
Stark, B. P., and H. A. Rhodes. 1997. Perlesta xube, a new stonefly species from Nebraska

(Plecoptera: Perlidae). Entomol. News 108: 92-96.

3 1 8 ENTOMOLOGICAL NEWS

TWO NEW SPECIES OF CHOREBUS (HYMENOPTERA:
BRACONIDAE) FROM SPAIN 1

I. Docavo , J. Tonnes-*

ABSTRACT: Chorebus pseudometallicus and C. pseudoasini, two new species from Spain, are
described and compared with allied species of the genus.

The subfamily Alysiinae is one of the most distinctive subfamilies of the
Braconidae because all members possess the exodont condition and are
endoparasitoids of cyclorrhaphous Diptera.

This subfamily is subdivided traditionally into two tribes: Alysiini and
Dacnusini. Chorebus Haliday, whose species are endoparasitoids of Agro-
myzidae and Ephydridae Diptera (there exists only one exception, a species
that attacks Psila rosae [F.]), is the largest genus of Dacnusini with approxi-
mately 215 Holarctic species. Many of its species are characterized morpho-
logically by having a densely setose metapleuron (metapleural rosette) and
usually a sculptured sternaulus. We have discovered two new species, described
below, which were obtained netting on Papilionaceae, in Alcira (province of
Valencia), Spain. The Dacnusini have been dealt with, both at the morphologi-
cal and biological levels, by Griffiths (1964, 1966, 1968, 1984) and Tobias
(1986, summary of the Palearctic taxa with keys to genera and species, trans-
lated into English 1995).

Terms for body morphology and wing venation follow Griffiths (1964)
and Wharton( 1977, 1986).

Chorebus pseudometallicus NEW SPECIES

Female: Head (Figs. 1, 2, 3) - Transverse, 1.87 times wider than long, 1.25 times higher than
long; occiput bare; vertex with scattered pubescence; base of the mandibles with tenuous pu-
bescence; eyes in lateral view 0.87 times as long as the temples; temples bulging beyond eyes
in dorsal view; eyes strongly converging below; face 1.4 times as wide as high; antennae with
23 antennomeres, apical flagellomeres ca 2.5-3.2 times as long as wide; mandibles 3-tooth,
1st relatively small, weakly expanded, blunt, 2nd tooth very long and pointed, 3rd tooth short,
expanded, slightly pointed; maxillary palpi long.

Mesosoma (Figs. 1, 2, 4) - 1.28 times as long as high, 1.71 times as long as width between
tegulae; pronotum bare and shining, only setose along anterior oblique suture; mesonotal disc
extensively bare, with only scattered setae on its anterior half and along each notaular line;
midpit shallow, narrow, extending from about posterior 1/3 of disc nearly to posterior margin;
notauli scarcely visible, represented by smooth fine line that seems reach midpit; sternaulus
extending to posterior border of mesopleuron, narrow, shiny, practically smooth; posterior

1 Received October 21, 1997. Accepted February 25, 1998.

2 Departamento de Biologia Animal, Biologia Celular y Parasitologia. Facultad de Biologia.
Universidad de Valencia. C/Dr. Moliner, 50. Burjasot (Valencia). Spain.

3 Unidad de Zoologia. Facultad de Biologia. Universidad de Salamanca. 37071 -Salamanca.
Spain.

ENT. NEWS 109(5): 318-324, November & December, 1998

Vol. 109, No. 5, November & December, 1998

319

1 mm

3 0.4

0.4 mm

1 mm

0.4 mm

6 02

FIGURES 1-6. Chorebus pseudometallicus sp. nov. (female).- 1, Body (except legs and wings)
in dorsal view; 2, in side view; 3, Head in lateral view; 4, Propodeum in lateral view; 5, Ante-
rior right wing; 6, Petiole in dorsal view.

320 ENTOMOLOGICAL NEWS

mesopleural furrow smooth; mesopleuron smooth, shiny, bare, only with a few long setae near
the ventral border; metapleuron and propodeum with pubescence only moderately densely se-
tose, which allows its rugose sculpture to be clearly seen; posterior coxae with a setae tuft little
differentiated on its posterior margin.

Wings (Fig. 5) - Pterostigma quite narrow and little darkened, 1.2 times longer than the
metacarpus; 1 st radial segment shorter than the length between its insertion and the parastigma,
and about as long as the pterostigma wide; remainder of radius evenly curved; n. rec. antefurcal;
3rd discoidal segment represented only by a shadow, so that cell B is open at its lower distal
corner.

Metasoma (Figs. 1, 2, 6) - Petiole 1.4 times longer than wide apically, glabrous, grooved,
with a pronounced central ridge; ovipositor sheath setose, robust, extending slightly beyond last
tergite in resting position.

Color and size - Head, mesosoma and metasoma black; face black, clypeus and labrum
darkish; maxillary and labial palpi dark brown - the labial palpi a little lighter-; antennae black,
with yellowish brown pedicel and base of the scape; centre of mandibles reddish-yellow;
mesopleuron black shiny; legs reddish-brown, with middle and posterior coxae, tibiae and tarsi
more infuscated (darker); wings hyaline, with dark pterostigma. Body length: 2.1 mm.

Male: unknown. Host: unknown.

Material examined: [deposited in the Fundacion Entomologica "Torres-Sala" (Docavo Col-
lection) (Valencia, Spain)]: Holotype: female, SPAIN: Valencia: Alcira, 30-11-1963 (leg. I.
Docavo). Paratype: female, SPAIN: Valencia: Alcira, 30-11-1963 (leg. I. Docavo).

Etymology: The specific name of this species makes reference to C.
metallicus Griffiths [Griffiths, 1968], to which the new species is very similar.

Chorebus pseudoasini NEW SPECIES

This new species appears very close to C. pseudometallicus, but can be
distinguished by the following characters:

Female: Head - Weakly transverse, 1 .6 times wider than long, 1 .4 times higher than long;
base of the mandibles with well differentiated pubescence (Fig. 7).

Mesosoma - Sternaulus pointed in its anterior part; posterior coxae with a tuft of setae well
differentiated on its posterior margin (Fig. 8).

Wings (Fig. 9) - Pterostigma narrow, imperceptibly joining the metacarpus; cell R nar-
rower and shorter, finishing before the tip of the wing; 3rd discoidal segment decolored; cell B
incompletely closed.

Metasoma - Petiole 2.1 times longer than wide apically, with the tubercules of the spiracles
very pronounced (Fig. 10); last tergite narrow and prolonged covering the ovipositor of which
only its apex can be appreciated.

Color and size - anterior and middle coxae a very yellowish reddish brown; posterior very
dark. Body length: 1.9 mm.

Male: unknown. Host: unknown.

Material examined: [deposited in the Fundacion Entomologica "Torres-Sala" (Docavo Col-
lection) (Valencia, Spain)]: Holotype: female, SPAIN: Valencia: Alcira, 28-11-1960 (leg. I.
Docavo). Paratypes: 2 females, Valencia: Alcira, 28-11-1960 (leg. I. Docavo).

Vol. 109, No. 5, November & December, 1998

321

0.4 mm

0.5 mm

' 0.5 mm

FIGURES 7-10. Chorebus pseudoasini sp. nov. (female). - 7, Mandible in lateral view; 8,
Sternaulus and lateral view of the right posterior coxae; 9, Anterior right wing; 10, Petiole in
dorsal view.

Etymology: The specific name makes reference to the closely related C.
asini Docavo [Docavo, 1 965].

Notes: These new species belong to a group which Griffiths ( 1 964) has described as the "affinis
complex". They are similar to C. metallicus and C. asini, from which they differ in the following
respects: a) number of antennomeres; b) pubescence from the base of the mandibles, sides of
pronotum, mesonotal disc, metapleuron (rosette), posterior coxae (setae tuft) and propodeum; c)
coloring of the legs (particularly of the coxae); d) morphology of the pterostigma, cell B (bra-
chial cell), sternaulus and metasoma (specially petiole morphology); e) body length.

322 ENTOMOLOGICAL NEWS

C. pseudometallicus can be differentiated from:

C. metallicus: a) By the clearly differentiated metapleuron pubescence (ro-
sette) (Figs. 2, 4) [less differentiated in C. metallicus]; b) coxae reddish-brown
[in C. metallicus gold yellow]; c) anterior oblique suture of the pronotum pu-
bescent (Fig. 2) [in C. metallicus the sides of the pronotum are bare and shiny];
d) mesonotal disc slightly pubescent (Fig. 1 ) [C. metallicus has only a few
setae along the notauli]; e) pubescence on propodeum more dense than in C.
metallicus (Figs. 1, 2, 4).

C. asini: a) Antennae with 23 antennomeres [C. as//?/ has 25 antennomeres];
b) mesonotal disc more shiny and less pubescent (Fig. 1 ) [in C. asini dull and
much more setose]; c) legs lighter; d) sides of pronotum pubescent (Fig. 2) [in
C. asini they are practically glabrous]; e) sternaulus narrow, pointed anteriorly
(Fig. 2) [in C. asini completely smooth]; f) pterostigma narrower (Fig. 5).

C. pseudoasini: Due to the morphological characters previously explained
in the description of this species. The differences between this species and C.
asini are the same as those of C. pseudometallicus except the number of
antennomeres which is 23.

The most important characteristic for recognizing these species lie in well
differentiated metapleural rosette for C. pseudometallicus (Figs. 2, 4), and in
well differentiated setae tuft in the posterior coxae for C. pseudoasini (Fig. 8).

The four species: C. pseudometallicus, C. metallicus, C. pseudoasini and
C. asini, can be distinguished in table I.

Table I - Morphological characterization of C. pseudometallicus, C. metallicus, C. pseudoasini
and C. asini [differentiated character (*); scarcely differentiated or missing (-)]. Number of
antennomeres (1). Pubescence well differentiated in: base of mandibles (2), sides of pronotum
(3), mesonotal disc (4), metapleuron (rosette) (5), propodeum (6); coxae dark (7), pterostigma
narrow (8); cell B completely open (9); sternaulus completely smooth (10); posterior coxae
(setae tuft) (11), Tubercles of the spiracles greatly differentiated (12).

Species

1 2 3

4 5 6 7 8 9 10 11 12

C. pseudometallicus

23 *

***.*_...

C. metallicus

21/22 -

.-*-*--

C. asini

* ***__*__

C. pseudoasini

23 * *

***_ **_ * #

Vol. 109, No. 5, November & December, 1998 323

These species can be inserted in the keys of Tobias (1995: 340) as follows:

Females

(475) (476)

(475a) (475b)

Antennae with 23 antennomeres. Mesonotal disc more or less shiny, with a few setae along
the notauli and central anterior part. Pronotum shiny, with a few setae, fine and scattered, along
the anterior oblique suture. Petiole 1.6 to 2.1 times longer than wide, shiny, practically bare,
with a well differentiated central ridge and longitudinal grooving. Propodeum scarcely pubes-
cent, more differentiated at its sides, although allowing the sculpture situated underneath to be
perceived. Pubescence of the metapleuron (rosette) and propodeum well differentiated and
characteristic of the genus Chorebus.

(475b) (475c)

Anterior coxae reddish brown, middle and posterior a darker brown with reddish over-
tones. Petiole almost bare, grooved, with a well differentiated central ridge. Metasoma oval,
elongated, less rounded on the apex than in C. pseudoasini, without the last tergite being elon-
gated and narrow; ovipositor, therefore, more protruding than in mentioned species. Pterostigma
wider than in C. pseudoasini, not so imperceptibly joining the metacarpus. Cell B somewhat
open in its lower distal corner, with just a trace of the 3rd discoidal segment. Length of body: 2
mm. Host: unknown.

C. pseudometallicus sp. nov.

(475c)(475)

Anterior and middle coxae, a more yellowish reddish brown, but the posterior, partly,
darker. Petiole shorter, but with extremely protruding stigmatipherous tubercules. Metasoma
rounder, with the last tergite narrow and prolonged, covering the ovopositor, of which only the
apex can be seen from above, while in lateral view the ovopositor appears curved upwards.
Pterostigma narrow, imperceptibly joining the metacarpus. Cell R narrower and shorter, finish-
ing beyond the extremity of the wing. Cell B completely open, as 3rd discoidal segment is
completely missing. Length of body: 1 .9 mm. Host: unknown.

C. pseudoasini sp. nov.

(475) (476)

ACKNOWLEDGMENTS

We are much indebted to Cees van Achterberg (Nationaal Natuurhistorisch Museum, The
Netherlands) and Max Fischer (Naturhistorisches Museum Wien, Osterreich), for their com-
ments on the manuscript. Financial support for this paper was provided from the Junta de
Castilla y Leon, project SA 18/96.

LITERATURE CITED

Docavo, I. 1965. Nuevas aportaciones al conocimiento de los Dacnusini de Espana (Hym.,

Braconidae). Graellsia, XXI: 25-39.
Griffiths, G.C.D. 1964. The Alysiinae (Hym., Braconidae) parasites of the Agromyzidae

(Diptera). I. General questions of taxonomy, biology and evolution. Beitr. Entomol., 1 4:

823-914.
Griffiths, G.C.D. 1966. The Alysiinae (Hym., Braconidae) parasites of the Agromyzidae

(Diptera). II. The parasites of Agromyza Fallen. Beitr. Entomol., 16: 551-605.

324 ENTOMOLOGICAL NEWS

Griffiths, G.C.D. 1968. The Alysiinae (Hym., Braconidae) parasites of the Agromyzidae
(Diptera). VII. The parasites of Cerodontha Rondani s.l. Beitr. Entomol., 18: 63-152.

Griffiths, G.C.D. 1984. The Alysiinae (Hym., Braconidae) parasites of the Agromyzidae
(Diptera). VII. Supplement. Beitr. Entomol., 34: 343-362.

Tobias, W.I. 1986. Identification key for the insects of the European part of the URSS. Vol. III.
Part V. Hymenoptera, Braconidae. pp. 100- 105 (key for genera of Alysiinae), 163-221
(Dacnusini). Akademia Nauk: Leningrad (in Russian, transl. 1995 in English).

Wharton, R.A. 1977. New World Aphaerela species (Hymenoptera: Braconidae) with a dis-
cussion of terminology used in the tribe Alysiini. Ann. Ent. Soc. Am., 70: 782-803.

Wharton, R.A. 1986. The braconid genus Alysia (Hymenoptera): a description of the subgen-
era and a revision of the subgenus Alysia. Syst. Ent., 1 1 : 453-504.

BOOKS RECEIVED AND BRIEFLY NOTED

INVERTEBRATE SURVEYS FOR CONSERVATION. T.R. New. 1998.
Oxford Univ. Press. 240 pp. $35.00 (paperback).

A comprehensive guide to the ecological methods used to survey invertebrate animals in ter-
restrial, freshwater, and marine environments. It describes how to select taxonomic groups for
study, how to collect and analyse samples, and how to set priorities for protection.

MELANISM. EVOLUTION IN ACTION. M.E.N. Majerus. 1998. Oxford
Univ. Press. 338 pp. $105.00 (cloth); $45.00 (paperback).

Placing melanism into its historical and scientific context, the author considers the diversity of
melanism in the animal and plant worlds, and its physical and genetic properties. Examining mela-
nism in moths and ladybeetles in detail, he explores the diversity of evolutionary reasons for mela-
nism and the complexities underlying this phenomenon.

THE BIRDER'S BUG BOOK. G. Waldbauer. 1998. Harvard Univ. Press. 290
pp. $27.95 (hardcover).

This book is an interesting introduction into the many fascinating relationships between birds
and insects. As past eons have come and gone, birds and insects have become increasingly en-
meshed in a complex web of interrelationships: birds eating insects, bloodsucking insects feeding
on birds, parasitic insects infesting birds, and birds struggling to rid themselves of the parasites. In
this book, the author describes these and many other interactions between birds and insects.

Vol. 109, No. 5, November & December, 1998 325

DIPTEROUS PARASITOIDS FROM ADULTS
OF MOTHS (LEPIDOPTERA) 1

Timothy L. McCabe 2

ABSTRACT: An adult moth, Cucullia lucifuga (Lepidoptera: Noctuidae), was parasitized by
the maggot of Sarcophaga aldrichi (Diptera: Sarcophagidae). Circumstances suggest that the
fly, which is a scavenger and a larval and pupal parasitoid of Lepidoptera, also attacks the
adults of moths. This is the first report of a sarcophagid fly from the adult of a moth. The
maggot of a phorid fly, Megaselia rufipes, is reported from the adult of another noctuid moth,
Amphipyra glabella.

It is unusual to have the adult stage of a moth or butterfly parasitized.
Adult lepidopterans with dipterous parasitoids of the families Tachinidae and
Phoridae are known. Tachinid observations were presumed to be examples of
larval or pupal parasitoids that carried over into the adult (Cockayne, 1911;
De Vries, 1 979; Smith, 1 98 1 ). Flemyng (1918) reported an adult sphingid para-
sitized by a phorid fly, Megaselia rufipes (Meigen). Flemyng 's original obser-
vation was too casual to ascertain whether the living moth or the carcass had
been attacked. Borgmeier (1965) considers M. rufipes maggots to be polypha-
gous; Robinson (1971) documents a wide range of hosts from lepidopteran
pupae to bat guano. Flemying's report, however, may have some credence.

In July of 1997, I held a captive female Amphipyra glabella (Morrison)
(Lepidoptera: Noctuidae), collected near Albany, New York, in a tightly sealed
container for a week. Soon after the moth's death maggots could be observed.
Two female flies emerged after ten days. These proved to be M. rufipes. De-
spite frequently having held moths for eggs, this was my first observation of
phorid parasitoids killing a moth. Whole abdomens of moths are treated chemi-
cally when lepidopterists prepare specimens for dissection, and I have only
twice encountered unidentified phorid maggot exoskeletons in the course of
three thousand dissections.

The following represents the first report of a sarcophagid parasitoid of an
adult lepidopteran.

Sarcophaga aldrichi Parker is an important internal parasitoid of the pupa
of tent caterpillars. Hodson (1939) proved unequivocally that tent caterpillar
larvae were not attacked, but that the pupae were. The fly has been reported as
a parasitoid of the larva of the satin moth (Lejeune and Silver, 1961) and the
gypsy moth (Hodson, 1939). Gypsy moth, satin moth, and tent caterpillars all
make an exposed cocoon. Sarcophaga aldrichi is viviparous. A female lays its
first instar maggot on the silk of the host's cocoon. The maggot finds its way
through the silk to the pupa and feeds internally for 3 to 5 days. After this

1 Received January 22, 1 998. Accepted March 13, 1998.

2 Biological Survey, New York State Museum, Albany, New York 12230.

ENT. NEWS 109(5): 325-328, November & December, 1998

326 ENTOMOLOGICAL NEWS

period the maggot may stay within the pupal cadaver for more than a month,
eventually dropping to the ground to pupate (Hodson, 1939). Sarcophaga
aldrichi also will breed in decaying organic material and carrion (Hodson,
1939) as do most Sarcophaga species (Sanjean, 1957).

On July 22, 1987, an adult female of Cucullia lucifuga [Denis and
Schiffermiiller] was captured nectaring at milkweed blossoms on the pine bar-
rens near Albany, New York. The moth was held for oviposition in a plastic
cup with a tight fitting lid. The moth died without ovipositing on July 24th.
The following day the moth carcass had the legs, wings, and head detached.
No internal organs remained. A third (ultimate) instar Sarcophaga (Diptera:
Sarcophagidae) maggot was present. It had been an internal parasitoid of the
moth. The maggot was offered additional fresh, decapitated, live moths and
fresh cadavers of moths, but refused to feed. The maggot was kept in a moist
container. It lived 51 days, until September, but died. Its carcass was preserved
and identified as Sarcophaga aldrichi.

The caterpillar of C. lucifuga is a flower feeder. When fully grown, the
caterpillar moves to the ground and enters the soil. It pupates in an under-
ground cocoon. My pupation boxes have 15 centimeters of peat and C. lucifuga
typically pupates at the bottom. The species is triple brooded in the north and
the summer brood has a pupal stage that lasts for 10 days or more. Therefore,
the sarcophagid loses its opportunity to parasitize C. lucifuga once the cater-
pillar enters the soil.

The fly is probably not a pupal parasitoid here, although it would have
had the opportunity to parasitize the larva. Hodson (1939) did not observe
larval parasitism by S. aldrichi in 100 field collected last-instar larvae of tent
caterpillars. Sixty percent of the tent caterpillar pupae from Hodson 's experi-
mental site had been parasitized by 5. aldrichi.

The sarcophagid maggot completes its development in 3 to 5 days (Hodson,
1939). Sarcophaga aldrichi is a large fly, larger than a house fly, and there is
no chance of a parasitized host the size of C. lucifuga surviving. A large satyrine
caterpillar has been reported as surviving despite having been parasitized by a
tachinid maggot (DeVries, 1984); the caterpillar survived the exiting of the
maggot and ultimately produced an adult. The maggot was not carried over to
the butterfly's adult molt.

In the present case the adult may have been the stage initially parasitized.
The reasoning for this is as follows: 1 ) I estimate the moth was 4 -7 days of age
at death (age determination based on my observations of many captive and
bred lepidopterans); 2) maggot development time is 3-5 days; 3) the maggot
was large enough to kill an organism the size of C. lucifuga at any stage; 4) the
pupa was probably not parasitized because pupation takes place deep under-
ground; 5) the larva was probably not parasitized because 5. aldrichi seldom
attacks larvae and the maggot development time of 3-5 days is too quick; 6)

Vol. 109, No. 5, November & December, 1998

327

the moth is active at early dusk, while there is still light, and both fly and moth
can be found visiting milkweed blossoms simultaneously.

Given the relatively constant development time for the maggot, one would
expect a small to mid-size imago to be killed within 3 to 5 days of attack.
When I collected the moth, which I determined to be mated based on clasper
marks, I estimated it to be 2-5 days of age. When the parasitoid emerged the
moth would have been 4-7 days of age, sufficient time for a first stadium mag-
got to have entered the imago and completely matured.

The preserved maggot agrees well with the description and illustration of
5. aldrichi given in Greene (1925). The maggot has anterior spiracles with
multiple rows of lobes. This is a rare condition in Sarcophaga. The lack of
spines around the posterior pocket and the prominent anal tubercles are all
concordant with S. aldrichi morphology. The mouthparts of the preserved 5.
aldrichi maggot are illustrated (Fig. 1). Mouthparts were not illustrated in
Greene's (1925) paper and prove to be valuable for species recognition. Sub-
sequent captures of adult Cucullia in the last ten years have not rediscovered
the parasitoid and it may prove to be a rare event.

Fig. 1 . Sarcophaga aldrichi. Cephalo-pharyngeal skeleton, showing mouthhooks and associated
structures, third instar. Scale line = 0.25 mm.

ACKNOWLEDGMENTS

I thank Christine Weber and Charles Sheviak for reviewing the paper and for helpful com-
ments. Specimens are vouchered in the New York State Museum. This is published as Contri-
bution number 738 of the New York State Science Service.

LITERATURE CITED

Cockayne, G. A. 191 1. A dipterous parasite bred from the imago of Nyxsia lapponaria. Ento-
mologist 44: 253.

328 ENTOMOLOGICAL NEWS

Borgmeier, T. 1965. Revision of the North American Phorid flies. Part III. The species of the

genus Megaselia, subgenus Megaselia (Diptera: Phoridae). Stud. Entomol. 8: 1-160.
DeVries, P. J. 1979. Occurrence of fly maggots in adult Morpho theseus (Lep.: Morphinae)

females from Costa Rica. Brenesia 16: 223.
DeVries, P. J. 1984. Butterflies and Tachinidae: does the parasite always kill its host? J. Nat.

Hist. 18: 323-326.

Flemyng, W. W. 1918. Sphinx convolvuli attacked by larvae of Diptera. Irish Nat. 27: 13.
Greene, C.T. 1925. The puparia and larvae of sarchophagid flies. Proc. U. S. Natl. Mus., Vol. 66,

Art. 29: 1-26.
Hodson, A. C. 1939. Sarcophaga aldrichi Parker as a parasite of Malacosoma disstria Hbn. J.

Econ. Entomol. 32(3): 396-401
Lejeune, R. R. and G. T. Silver. 1961 . Parasites and hyperparasites of the Satin Moth, Stilpnotia

salicis Linnaeus, (Lymantriidae) in British Columbia. Can. Entomol. 93: 456-467.
Robinson, W. H. 1971. Old and new biologies of Megaselia species (Diptera, Phoridae). Stud.

Entomol. 14:321-348.
Sanjean, J. 1957. Taxonomic studies of Sarcophaga larvae of New York with notes on the adults.

Cornell Univ. Agric. Exp. Stat. Mem. 349: 1-115.
Smith, K. G. V. 1981. Atachinid (Diptera) larva in the abdomen of an adult moth (Geometridae).

Entomol. Gaz. 32: 174-176.

Vol. 109, No. 5, November & December, 1998 329

BIOLOGY OF TANYCHELA PILOSA (HYMEMOPTERA:

ICHNEUMONIDAE), A PARASITOID OF THE AQUATIC

MOTH PETROPH1LA CONFUSALIS (LEPIDOPTERA:

PYRALIDAE) 1

Wanda Jamieson^, Vincent H.

ABSTRACT: Tanychela pilosa, a solitary endoparasite of the aquatic moth Petrophila confusalis,
has hymenopteriform eggs that are oviposited in late summer or early fall, in the Clearwater
River drainage of western Montana, USA. The first instar is a caudate-mandibulate type. Su-
perparasitism may result from polyembryony but only a single wasp larva survives. Larvae
undergo heteromorphosis as a third and final instar. Pupation occurs in the dry cocoon of the
moth. Adult sex ratios approach 1:1. 72.2% of sites examined in the Clearwater River drainage
had P. confusalis and 61.5% of these contained T. pilosa. Emergent rocks may enable oviposit-
ing females to enter streams and search for caterpillars. Parasitization of P. confusalis reported
from Idaho, Washington, and California may be by T. pilosa.

Hymenoptera parasitizing aquatic insects are rarely encountered in collec-
tions of stream insects. In an earlier study, we discovered that a population of
the aquatic lepidopteran Petrophila confusalis (Walker) was parasitized by an
aquatic wasp (Resh and Jamieson 1988) in Owl Creek, Missoula County,
Montana (Fig. 1). The rate of parasitism in the moth population ranged from
55% at the outlet of Placid Lake into Owl Creek to 0% less than 5 km down-
stream from the lake outlet. Wasp pupae were identified as Tanychela pilosa
Dasch, the type specimen of which is from Mexico (Dasch 1979).

In our original study, we indicated that parasitism occurred during the pu-
pal stage of the aquatic moth, as has been reported for the agriotypid (now
generally considered a subfamily of the Ichneumonidae) wasp Agriotypus
armatus Curtis that parasitizes the caddisfly Silo pallipes (Fabricius) in Eu-
rope (Elliott 1982). In retrospect, if this supposition were correct we would
have expected to see some type of activity indicating that moth pupae were
being parasitized, such as adult wasps near the pupal patches that occurred on
rocks in the streambed, or wasp eggs or larvae inside the moth pupae or moth
cocoons.

Because of the lack of detailed information on hymenopteran parasites of
aquatic insects (Hagen 1996), the present study was initiated to (1) describe
the biology of the parasitoid T. pilosa and (2) document the distribution of T.
pilosa beyond Owl Creek through surveys conducted in the Clearwater River
drainage system of western Montana.

1 Received October 11, 1997. Accepted January 12, 1998.

z Rathead Lake Biological Station, University of Montana, 31 1 Biostation Lane, Poison Mon-

tana 59860.

* Division of Insect Biology, 201 Wellman Hall, University of California, Berkeley, California
94720-31 12. Please send reprint requests to this author.

ENT NEWS 109(5): 329-338, November & December, 1998

330 ENTOMOLOGICAL NEWS

Fig. 1 . Petrophila confusalis and its endoparasite Tanychela pilosa.

METHODS AND MATERIALS

In May 1989, lepidopteran larvae were collected from Owl Creek near the
outlet of Placid Lake (Fig. 2). During the summers of 1993, 1994 and 1995,
samples of approximately 100 P. confusalis larvae and pupae were collected
randomly from various sites along the Clearwater River and its tributaries (Fig.

2).

Head widths of the moth larvae and prepupae were measured to determine
instar classification; the caterpillars were then dissected to determine rates of
parasitism. Wasp cocoons were dissected to determine stage of development
and sex of the wasps. Life cycle observations were based on examinations of
more than 1 400 aquatic wasp larvae and pupae, the dissections of more than
1900 aquatic moth larvae and prepupae, and the examination of more than
2000 empty wasp and moth cocoons.

BIOLOGY OF TANYCHELA PILOSA

Egg stage: T. pilosa wasps are solitary endoparasites of P. confusalis cat-
erpillars. The biology of the host (sometimes referred to as Paragyractis
confusalis) is described by Tuskes ( 1 977, 1 98 1 ), Me Auliffe & Williams ( 1 983),
Bergey (1995), and summarized by Lange (1996). In late summer or early fall,
the female wasp deposits an egg within the abdominal cavity of the moth larva.
The pale yellow, jellybean-shaped eggs, about 0.5 mm long and 0.2 mm wide,
are the hymenopteriform type (Hagen 1964, Fig. 14a).

Larval stage: Parasitism could occur during the moth's free-living first
instar or during its other larval stages that are spent beneath silken retreats that

Vol. 109, No. 5, November & December, 1998

331

Swan R.

Swan Rs'
Cygnet L.

'Clearwater L.
I Clearwater R.

Fig. 2. Sampling locations in the Clearwater Drainage, western Montana, USA.

332 ENTOMOLOGICAL NEWS

provide shelter (Lange 1996, Fig. 19.1). The free living, first-instar caterpillar
is more vulnerable to predation and to dislocation (i.e., drift), and the duration
of this stage is brief, e.g. first instars of P. confusalis occur in Owl Creek (Fig.
2) during a brief two-week period (McAuliffe & Williams 1983). In contrast,
second, third, and fourth instars occur from mid-August to mid-October
(McAuliffe & Williams 1983), and the female wasp could penetrate the silken
covering of these instars with her ovipositor, laying her eggs within the host
caterpillar.

The newly hatched wasp larva is less than 2 mm long. This primary or first
instar is of the caudate-mandibulate larval type described by Hagen (1964,
Fig. 18a, b, c). It has a brownish sclerotized head capsule with a black man-
dible. Its long, slender, segmented body is enclosed in a colorless, transparent
integument; the internal body mass is bright yellow. A long "tail" that is al-
most equal in length to the rest of its body extends posteriorly from the dorsal
portion of the wasp larva. Ullyett (1944) suggests the long tail may serve as
one and/or all of the three following functions: an egg burster, a balancing
organ to compensate for its large head, or for food absorption. Although the
first instar wasp has mandibles, we found no evidence in any of the host larvae
that internal tissues or organs had been damaged, nor did we ever observe any
tissue attached to the mandibles of the wasp larva that would indicate it had
been feeding. Apparently, the wasp larva absorbs food through its very thin
cuticle from its host's hemolymph. Strands of fat bodies were noticeably re-
duced or absent in the hemocoels of parasitized caterpillars. Because there are
no spiracles on the first instar larva, respiration is probably cutaneous.

T. pilosa overwinters in the first instar within its host. The wasp larva grows
very slowly from October to June and growth appears to be relatively constant
from year to year. For example, wasp larvae in samples collected from Owl
Creek and the Lake Inez outlet (Fig. 2) on 23 Oct 1993 were very similar in
size to the wasp larvae in samples collected during June 1994 and June 1995
from the same streams.

In areas where the rate of parasitism is high, two or more parasitoids may
occur in one host (i.e., superparasitism). For example, of 44 moth larvae that
were collected 23 Oct 1993 from Lake Inez (Fig. 2), 32 were parasitized. Of
these, six (19%) caterpillars had been parasitized more than once; three of
them had two larval parasites and each of the other three had three, four, and
five parasitic larvae, respectively.

Several genera of Ichneumonidae that parasitize the eggs and larvae of
Lepidoptera exhibit polyembryony (Hagen 1964, Chapman 1982) and this could
account for the multiple T. pilosa parasitoids within the host P. confusalis cat-
erpillar. We have not determined how long the different larvae within a single
host live, but we have observed that eventually all but one die. For example, in
a sample taken 15 Aug 1995 from the Lake Inez outlet (Fig. 2), one late-instar
moth larva had 10 wasp larvae within its hemocoel. All wasp larvae were the
same size, and had sclerotized heads and transparent body coverings, but only

Vol. 109, No. 5, November & December, 1998 333

one was wiggling; apparently nine of the wasp larvae had just died. Thus, this
moth larva had been host to 10 wasp larvae for 9 -10 months.

The dead wasp larvae can be found in the last abdominal segment; per-
haps, the wiggling motion of the caterpillar forces these dead larvae to the
posterior end of the moth's abdominal cavity where they remain (and subse-
quently can be counted). We noted that when host caterpillars contained only
one wasp larva, that larva was usually alive. The bodies of the dead wasp lar-
vae become white and opaque; their sclerotized head capsules enabled us to
distinguish them from the fatty tissues of the caterpillar.

Hagen ( 1 964) reported that the number of larval instars among hymenopter-
ous parasites is variable. However, there appears to be a tendency for ecto-
parasitic larvae to have five instars and endoparasitic forms to have fewer than
five (and often three).

During the fifth instar of the caterpillar, the wasp larva grows rapidly and
undergoes heteromorphosis (sensu Chapman 1982; Hagen 1964 refers to this
as hypermetamorphosis). The sclerotized head capsule splits along the dorsal
line and the wasp larva emerges from its exuvium with a new body form. The
new segmented body is covered with a loose transparent membrane. The body
internally is bright yellow. During this metamorphosis, the wasp larva abruptly
changes from a slender 2-mm long caudate-mandibulate type larva to a fat,
globular, grub-like larva that eventually grows to fill the thorax and abdomen
of the 10-mm long caterpillar within four weeks. This grub-like larva may
represent the third instar.

We examined the head-capsule width of approximately 100 caterpillars
from sites D in 1993, and N in 1993 and 1995 (Fig. 2). No size-class differ-
ences were observed in parasitized and unparasitized caterpillars.

Pupal stage: Prior to its pupation, the P. confusalis caterpillar modifies its
silken retreat on a rock surface to form a thicker, oval patch. This patch is
referred to as the external pupal case in Resh and Jamieson (1988, Fig. 1 ). The
patch has semicircular openings at the edge [as described by Lloyd 1 9 1 4, 1 9 1 9
for Petrophilafulicalis (Schaus)] and the caterpillar scores a C-shaped slit at
the upstream end of the patch. Beneath the patch, the caterpillar spins a water-
proof cocoon that provides a dry environment for the pupal stage. When it has
completely enclosed itself within a cocoon, the caterpillar becomes an immo-
bile prepupa. When examined externally, there is no indication whether or not
the caterpillar prepupa is host to a parasite.

If parasitization has been successful (i.e. sometimes the wasp larva dies
before heteromorphosis occurs), the wasp larva emerges from its host during
the caterpillar's prepupal stage. During emergence, the thoracic and abdomi-
nal skin of the caterpillar appears to disintegrate within the cocoon. The head
sclerites and other tissues of the moth fall to the posterior end of the cocoon as
the wasp larva emerges from its host within the dry environment of the host's
cocoon. At this time, spiracles are very prominent on the wasp larva.

The wasp larva immediately begins to spin its own cocoon within the

334 ENTOMOLOGICAL NEWS

caterpillar's cocoon and the wasp's cocoon is completed within a few hours.
Of the 5,000+ specimens examined for this study, <0. 1 % had wasp larvae that
were outside the host but not yet enclosed in their own pupal cocoons.

The structure of the wasp cocoon indicates that it is spun in a circular (or
sideways) manner back and forth from one end of the cocoon to the other, until
it is about three layers thick. During this process the wasp larva is visible be-
tween the strands of the cocoon. When the cocoon is completed, the wasp
larva is no longer visible. The cocoon is dark brown, leathery, oval-shaped,
and about 8 mm long. When the cocoon is turned inside out and placed under-
water, it takes on the metallic sheen characteristic of an unwettable surface.
During the construction of the cocoon, the wasp larva alternately faces anteri-
orly and posteriorly within the cocoon. But in the final phase of its prepupal
stage, the wasp larva always faces anteriorly within the cocoon, i.e., it points
its head in the same upstream direction that the caterpillar would have pointed
its head.

Because the stage of development cannot be determined until the cocoon
is dissected, all specimens within wasp cocoons have been described as pupae.
However, within this category, we have identified four phases of development
from what we found when the anterior end of the cocoon was opened. First, in
the "early" phase, which is actually still part of the larval stage, the larva is still
mobile within the cocoon and responds to touch when the cocoon is opened.
Morris (1937) referred to this as the eonymphal phase. As development pro-
ceeds, the wasp larva begins to transform into the pupal form. The colorless,
transparent membranous skin becomes white and opaque. Another change in
the larva is the formation of red spots where the compound eyes will form.
Differentiation of body regions begins. Morris (1937) referred to this final,
immobile, phase as the pronymphal stage.

During the second (or white) phase, three distinct body regions have formed;
the head is white and the compound eyes are red. Long legs and long antennae
have formed and in females the ovipositor is very well developed. In the later
part of this phase, although the head is still white, the thorax becomes black,
the eyes dark brown, and wing pads start to form.

The third (or black) phase is characterized by the pupa having a black
head, black antennae, black thorax, black legs, a brown and white striped ab-
domen, and very prominent wing pads. Both the white phase and the black
phase are enclosed within a transparent membrane. A meconium is attached
posteriorly to the developing pupa (Resh and Jamieson 1988, Fig. 10). These
three phases each require about one week. During the fourth (or pre-adult)
phase, the wasp pupa has shed the transparent membrane within the cocoon,
the meconium has become detached, and the wings are fully expanded.

Adult stage: To emerge from its cocoon, the adult wasp chews a hole dor-
sally through its cocoon and through the lepidopteran's external pupal case; it
does not use the C-slit made by the caterpillar for its own emergence. When
the newly emerged wasp comes in contact with the water, an air bubble from

Vol. 109, No. 5, November & December, 1998 335

the cocoon surrounds its hairy body. We observed that when an adult female
wasp emerged underwater (e.g. in a dish that contained moss and pebbles; we
have not observed this in nature), she grasped a sprig of moss with her hind leg
and groped in the dish with her other five legs. In another cocoon that was
opened underwater, we observed that after a female emerged from the water
she gripped the wooden handle of the teasing needle; then, with her hind legs,
she wiped down her abdomen and ovipositor and, with her front legs, wiped
her head and antennae. A few minutes after drying off, she flew away.

Based on the observations of 575 wasp pupae, the sex ratio of the wasps
approaches 1 : 1 (290 females: 285 males). Some streams, however, had collec-
tions favoring females (e.g. Fig. 2, site B, 43:11) or males (e.g. Fig. 2, site F,
8:22; site L, 7:22).

Emergence of wasps begins about two weeks after the unparasitized moths
have emerged. Once the moths have emerged from their cocoons, decomposi-
tion of their pupal patches begins and lasts for approximately one additional
month. The extended duration of the lepidopteran silken patch until the wasps
emerge is crucial to the wasps' existence and survival.

Parasitization of the moth larvae varies even on a single rock. For ex-
ample, 18 lepidopteran patches were collected 23 Jul 1995 from one rock at
the mouth of the Clearwater River (Fig. 2, site I). Among these patches: five
contained cocoons from which moths had emerged; three held moth larvae,
one of which was decomposing and another was parasitized; three were moth
pupae, one in an early stage of pupal development and the other two in a later
stage with scaled wings; seven specimens were wasp pupae, with three in the
early phase, three pre-adults (one male and two females), and one in a dam-
aged condition.

DISTRIBUTION AND PARASITISM

Of 18 sites sampled in the Clearwater River Drainage, P. confusalis was found at 1 3 sites: B,
C, D, F, G, H, I, J, K, L, M, N and Q (Fig. 2). T. pilosa parasitized P. confusalis at eight of those
sites: B (55% parasitism 29 Jul 1987, 44% 26 Jun 1988, 5% 3 Jul 1988, 41% 10 Jul
1988, 36% 22 Jul 1993, 45% 14 Jul 1993, 44% 28 Jul 1993, 50% 3 Aug 1993, 35% 11 Aug
1993, 13% 23 Oct 1993, 13% 25 Jul 1994, 29% 5 Jul 1995); C (24% 9 Aug 1987, 5% 26 Jun
1988,4% 16 Jul 1993); F (58% 22 Jul 1993, 43% 27 Jul 1994); G (41% 28 Jul 1993, 41% 3 Aug
1993, 48% 27 Jul 1994, 47% 23 Jul 1995, 74% 15 Aug 1995, 80% 22 Oct 1995); H (29% 3 Aug
1993, 20% 27 Jul 1994, 50% 5 Jul 1995); I (73% 3 Aug 1993, 64% 27 Jul 1994, 54% 23 Jul
1995); L (35% 22 Jul 1993, 46% 25 Jul 1994); N (75% 22 Jul 1993, 71% 11 Aug 1993, 67% 23
Oct 1993, 48% 12 Jul 1994, 47% 19 Jun 1995, 57% 15 Aug 1995). Variations in rates of
parasitism at individual sites may have been caused by sampling of different microhabitats at a
site.

What were the characteristics of the sites where either P. confusalis or T.
pilosa were absent? No lepidopterans were found in the cold water (10C) of
the West Fork of the Clearwater River on 28 Jul 1993. Algal patches on the
rocks of the warm Marshall Lake outflow (Fig. 2, site O) suggested a lepi-
dopteran population may have been present earlier in the season but had

336 ENTOMOLOGICAL NEWS

emerged. The Cleanvater River at Highway 83 (R) was sampled on 2 Aug
1993; the water was very cold (11C) and no moths were found. On 3 Aug
1 993 the Blackfoot River at the Russell Gates Fishing Access (J) was sampled.
The river was wide, shallow, and swift, and the rocks in the streambed were
large, round, smooth and very slippery. The lepidopteran population was very
sparse and only 1 1 specimens could be collected; none were parasitized. The
Blackfoot River at the Roundup Fishing Access (K) also had no parasitism in
the 49 moths examined. The river here was very deep, wide, and swift. Sam-
pling was limited to a rocky area near the shore.

Samples outside the Clearwater Drainage were also collected during sum-
mer 1993. No parasitism was found in a sample taken on 26 Jul 1993 from the
outflow of Loon Lake on the Fisher River in Lincoln County, Montana, nor in
a sample of 145 specimens taken on 27 Jul 1993 from the Lower Crow Creek
above the Lower Crow Reservoir near Pablo in Lake County. Although moth
pupae and larvae were abundant in Lower Crow Creek, the water was deep
and swift.

Ashley Creek in Flathead County was sampled 30 Jul 1993 at its outflow
from Ashley Lake and near the bridge between Lake Monroe and Lone Lake;
no parasitism of the moth larvae or pupae present at either site was evident.
The patches were extremely calcified and this may have prevented wasps from
chewing their way out of their cocoons even if parasitism had occurred. Three
sites (S, T, U) in the Swan River drainage system adjacent to the headwaters of
the Clearwater River (Fig. 2) sampled on 2 Aug 1 993 did not have lepidopteran
populations. Algae mats were thick on rocks in the streambed and the water
was very slow moving in Holland Lake Creek (T). Although the water was
warm at the Lake Lindbergh outlet (U), it was very deep and swift. At site S,
the Swan River was deep and wide. There was no parasitism in a sample of 65
specimens collected from Swan River below the dam near Bigfork on 10 Aug
1993. The water there was also deep and very swift.

FACTORS AFFECTING T. PILOSA DISTRIBUTION

From the above information, it is apparent that parasitization by T. pilosa
does not occur in all streams that had P. confusalis populations. Reasons un-
derlying this distribution are not readily evident. For example, Lake Inez and
Lake Alva (Fig. 2, sites N and Q) are two very similar lakes in the Clearwater
River system, with Lake Alva just to the north of Lake Inez in the upper por-
tion of the drainage. Both are warm-water lakes and are surrounded by sum-
mer homes. Wooden plank structures dam the outlets of each lake, and the
outflow streams have rocky bottoms and banks lined with vegetation. Each
stream supports a population of P. confusalis. However, parasitization at the
Lake Alva outflow is 0% while parasitization at the Lake Inez outflow is 75%.

Why is there such a difference in parasitism when these lake outlets are
less than 5 km apart? Water in the Lake Alva outlet flows smoothly (i.e. no

Vol. 109, No. 5, November & December, 1 998 337

emergent rocks or riffles) over a streambed of uniformly sized rocks that are
densely covered with lepidopteran pupal patches. When the water level goes
down in late summer, the stream still flows smoothly. However, at Lake Inez,
the outflow stream does not have a streambed of uniform composition and the
water does not flow smoothly; instead, riffles form from emergent rocks and
boulders interspersed among sand and gravel bars. Lepidopteran patches can
be found on the large rocks and boulders as well as on small rocks and pebbles.
In late summer, the larger rocks project above the water; if the female wasp
requires a landing place before she enters the water for oviposition, then the
Lake Inez outflow would be a possible habitat whereas Lake Alva would not.
Given that the above-described habitats differ in terms of emergent rocks or
riffles, and that this was often a feature lacking in sites where moths occurred
but parasitism was absent, we suggest that this may be a key feature in deter-
mining the local distribution of T. pilosa. Perhaps the wasp adult requires emerg-
ing rocks to enter the water and remain attached while it searches for a poten-
tial host. Our laboratory observations of adults holding on to objects suggest
that this is the case.

Owl Creek has characteristics of both Lake Inez and Lake Alva outflows.
Riffle regions alternate with pool regions. No parasitism was found in 5 1 speci-
mens collected 23 Jul 1995 at Owl Creek (Fig. 2, site B) from a pool area
located behind a line of rocks across the stream. However, in a riffle area at
this site, 22 of the 57 specimens collected were parasitized. The rocks in this
riffle area project above water in late summer.

Gustin (personal communication) found a species of Tanychela parasitiz-
ing populations of P. confusalis in the Potlatch River and several of its tribu-
taries in Latah Co, Idaho and in the Palouse River system in Latah County,
Idaho, and Whitman County, Washington. The immature ichneumonids found
in his study and the unnamed immatures previously reported by Tuskes ( 1 977)
in California may also be T. pilosa. If the distribution of T. pilosa follows that
of P. confusalis, it would include central California, north to British Columbia
and east to Nevada, Idaho, and Montana (Monroe 1972). However, because
the type specimen of T. pilosa is from Mexico, other species of moths may
also be parasitized.

Local factors may also affect distribution. Cool summers may favor the
growth and development of the wasp larvae whereas warm summer tempera-
tures may favor the rapid growth of fifth instar caterpillars and, consequently,
earlier pupation and emergence. If T. pilosa cannot follow the rapid transfor-
mation of its host, then it will not be ready for its own pupation.

Because the larval wasp occurs within the caterpillar's body and the moth
occurs within an air-filled cocoon (Lloyd 1919), the only aquatic portions of
T. pilosa s life cycle are when the wasp adult emerges underwater and then
returns to water to lay its eggs. This life history fits even the most narrow
definition of an aquatic insect (Hagen 1996). However, like many other para-

338 ENTOMOLOGICAL NEWS

sitic insects that occur in aquatic environments, once inside its host the biol-
ogy of T. pilosa more resembles that of a terrestrial insect than an aquatic one.

ACKNOWLEDGMENTS

We thank: Stuart Neff, Temple University, and Brett Merritt, Michigan State University,
for assistance in collecting specimens; Jack Stanford and the staff of the Flathead Lake Bio-
logical Station of The University of Montana for their assistance and use of facilities; Powder
River High School (Broadus, Montana) for use of facilities; the students at Powder River High
School for their encouragement to continue this project; Kenneth Hagen and Leo Caltagirone,
University of California, Berkeley, for their comments on the manuscript; and Jim Gustin for
permission to use his observations. We dedicate our efforts on this study to the late Professor
Kenneth Hagen.

LITERATURE CITED

Bergey, E. A. 1995. Local effects of a sedentary grazer on stream algae. Freshwat. Biol. 33:401-

409.
Chapman, R. F. 1982. The insects - structure and function. 3rd ed. Harvard University Press,

Cambridge.
Dasch, C. E. 1979. Ichneumon-flies of America North of Mexico: 8. Subfamily Cremastinae.

Mem. Amer. Entomol. Inst. 29: 1 -702.
Elliott, J. M. 1982. The life cycle and spatial distribution of the aquatic parasitoid Agriotypus

armatus (Hymenoptera: Agriotypidae) and its caddis host Silopallipes (Trichoptera: Goeridae).

J.Anim.Ecol. 51:923-941.

Hagen, K. S. 1964. Developmental stages of parasites p. 168-246. In: DeBach, P. (ed.). Biologi-
cal control of insect pests and weeds. Reinhold Publ. Corp., N.Y.
Hagen, K. S. 1996. Aquatic Hymenoptera. p. 474-483. In: R. W. Merritt and K. W. Cummins

(eds.). An introduction to the aquatic insects of North America. 3rd ed. Kendall/Hunt Publ.

Co., Dubuque, IA.
Lange, W. H. Aquatic and semiaquatic Lepidoptera. p. 387-398. In: R. W. Merritt and K. W.

Cummins (eds.). An introduction to the aquatic insects of North America. 3rd ed. Kendall/

Hunt Publ. Co., Dubuque, IA.

Lloyd, J. T. 1914. Lepidopterous larvae from rapid streams. J. N. Y. Entomol. Soc. 2: 145-152.
Lloyd, J. T. 1919. An aquatic dipterous parasite, Ginglymyia acrirostris Towns, and additional

notes on its lepidopterous host, Elophilafulicalis. J. N. Y. Entomol. Soc. 27:263-265.
McAuliffe, J. R. and N. E. Williams. 1983. Univoltine life cycle of Paragyractis confusalis

Walker (Lepidoptera: Pyralidae) in the northern part of its range. Am. Midi. Nat. 110:440-

443.
Morris, K. R. S. 1937. The prepupal stage in Ichneumonidae, illustrated by the life history of

Exenterus abruptoriusThb. Bull. Entomol. Res. 28:525-534.
Munroe, E. G. 1972-1973. Pyraloidea. Pyralidae (in part), p. 1-134, Fasc. 13.1, A-C. In: R. B.

Dominick (ed.). The moths of America north of Mexico. E. W. Classey Ltd., London.
Resh, V. H. and W. Jamie-son 1 988. Parasitism of the aquatic moth Petrophila confusalis (Lepi-
doptera: Pyralidae) by the aquatic wasp Tanychela pilosa (Hymenoptera: Ichneumonidae).

Entomol. News 99: 185- 188.
Tuskes, P. M. 1977. Observations on the biology of Paragyractis confusalis, an aquatic pyralid.

Can. Entomol. 109:695-699.
Tuskes, P. M. 1981. Factors influencing the abundance and distribution of the aquatic moths of

the genus Paragyractis (Pyralidae). J. Lepid. Soc. 35: 161-168.
Ullyett, G. C. 1944. On the function of the caudal appendage in primary larvae of parasitic

Hymenoptera. J. Entomol. Soc. S. Afr. 7:30-37.

Vol. 109, No. 5, November & December, 1998 339

NEW OHIO RECORDS OF CORIXIDAE (HEMIPTERA) 1

Stephen W. Chordas, III 2 , Brian J. Armitage 2

ABSTRACT: One new genus and five new species of corixids have recently been recorded from
Ohio as part of a comprehensive survey of adult aquatic insects. The total number of corixid taxa
now known from Ohio is 25. All five taxa were collected using black light traps. The newly
reported genus, Corisella, is known mainly from western North America. Corisella inscripta has
heretofore only been found west of the Mississippi River. The other four new species records for
Ohio represent three genera and include Palmacorixa buenoi, Sigara dejecta, S. hubbelli, and
Trichocorixa kanza. All five new taxa were collected on one or more of the following State
Wildlife Areas: Big Island, Killdeer Plains, and Resthaven.

There are almost 80 State Wildlife Areas in Ohio. These are managed by
the Ohio Division of Wildlife (the Division) for a variety of hunting, fishing,
recreational, and watchable wildlife functions. However, few of these wildlife
areas have been surveyed for their aquatic insect fauna. Recently the Division
funded the Ohio Biological Survey (the Survey) to conduct a three year study
of the state-listed, special category species for Killdeer Plains Wildlife Area.
The Survey is also conducting a multi-year comprehensive survey of the adult
aquatic insects in Ohio co-funded by the Division, the Partnerships for Wild-
life Foundation of the U.S. Fish and Wildlife Service, and the Procter and
Gamble Co., Inc. Finally, the Wisconsin Department of Natural Resources,
with funding from Partnerships for Wildlife, has contracted with the Survey to
conduct a six-state prairie Lepidoptera study. All of these projects generated
aquatic insect collections from state wildlife areas, especially Big Island (Marion
County), Killdeer Plains (Wyandot County), and Resthaven (Erie County).

The aquatic hemipteran fauna within Ohio, including the family Corixidae,
is poorly known. There have been no previous studies pertaining specifically
to the aquatic members of this order in Ohio. This void is evident when com-
prehensive aquatic Hemiptera works, such as Truxal's (1953) revision of the
genus Buenoa (Hemiptera; Notonectidae), contained no Ohio material. How-
ever, efforts such as that of Williams et al. (1996), Chordas (in review), this
paper, and the current survey of adult aquatic insects in Ohio, are addressing
this deficiency. Until recently, only 20 species of the family Corixidae (In-
secta: Hemiptera) were known from Ohio. These included the genera (# spe-
cies): Hesperocorixa (9); Palmacorixa (3); Rhamphocorixa (1); Sigara (5);
and, Trichocorixa (2). In contrast, there have been 49 species, representing 9
genera, of water boatmen reported from Wisconsin (Hilsenhoff, 1984).

1 Received February 6, 1998. Accepted May 10, 1998.

2 Ohio Biological Survey and Environmental Science Program, 1735 Neil Avenue Room 103,
Columbus, OH 432 10.

ENT. NEWS 109(5): 339-342, November & December, 1998

340 ENTOMOLOGICAL NEWS

In this paper we report five new Ohio records of water boatmen, represent-
ing one new genus and five new species. This is one of a series of papers about
this family and order which will culminate in a comprehensive diagnostic at-
las for aquatic Hemiptera in Ohio.

METHODS

Adult corixids were collected from May through October using black light
traps. Specimens from Killdeer Plains and Resthaven Wildlife Areas were col-
lected during a survey of prairie moths. Specimens from Big Island Wildlife
Area were collected during a study of aquatic insect dispersal to proposed
reconstructed wetland areas. Identifications were made using keys and descrip-
tions in Hungerford ( 1 948) and Sailer ( 1 948). Polhemus et al. ( 1 988) was used
as the reference for known distributional records. Specimens are preserved in
75% ethanol and deposited in the Ohio Biological Survey's Aquatic Insect
Collection.

RESULTS

All species collected on Big Island, Killdeer Plains, and Resthaven Wild-
life Areas are presented in Table 1 . The new genus and species records for
Ohio are indicated by symbols. Additional county records from sites outside
of the three wildlife areas are provided in table footnotes.

Table 1. Corixids found on three state wildlife areas.

Wildlife Areas

Species Big Island Killdeer Plains Resthaven

T* Corisella inscripta (\Jh\er), 1894 a XXX

Hesperocorixa atopodonta (Hungerford), 1927 X

H. obliqua (Hungerford), 1925 XXX

* Palmacorixa buenoi Abbott, 191 3 b X

Ramphocorixa acuminata (Uhler), 1897 X X

Sigara alternate (Say), 1825 XXX

* S. dejecta Hungerford, 1948 C X X
5. grossolineata Hungerford, 1 948 X

* 5. hubbelli (Hungerford), 1928 d X

5. modesta (Abbott), 1916 X

Trichocorixa calva (Say), 1832 XXX

* T.kanza Sailer, 1948 e XXX
T. sexcincta (Champion), 1901 X X

t = Denotes a previously unreported genus for Ohio * = Denotes a new state record for Ohio

a Also identified from Madison County

" Also identified from Clermont County

c Also identified from Athens, Clermont, and Greene counties

d Also identifed from Franklin and Jackson counties (kicknet samples)

e Also identified from Ashtabula, Clermont, Morgan, Pickaway, Richland, Tuscarawas,
Washington, and Williams counties.

Vol. 109, No. 5, November & December, 1998 341

DISCUSSION

The five new state records increase the total state list of corixids by 25% to
25 species. This relatively large increase in the faunal list, from a small num-
ber of sites, supports the notion that Ohio is undercollected. Given the distri-
butions published by Polhemus et al. ( 1 988), approximately 1 additional taxa,
primarily in the genus Sigara, could possibly be found in Ohio. Furthermore,
Chordas (in review) specifically discusses potentially resident species belong-
ing to the genus Hesperocorixa that may ultimately be found in Ohio. The Till
Plains physiographic province in western Ohio contains remnants of midwestem
prairies which once were common. Killdeer Plains and Big Island Wildlife
Areas contain some of these relict areas. Additional collecting in these and
other similar areas could reveal other taxa whose distribution is normally con-
sidered more typical of the prairie states west of the Mississippi River. Ohio
has portions of four other physiographic provinces (Lake Plains, Bluegrass,
Glaciated Allegheny Plateau, and Unglaciated Allegheny Plateau) which also
could contain taxa typical to these regions in other states but heretofore not
collected or identified from Ohio.

Sigara defecta: Ohio lies along the very southern edge of this species'
midwestern range (Hungerford, 1 948; Polhemus et al. , 1 988). This species has
been previously reported only from two states bordering Ohio (Michigan and
Pennsylvania). However, its occurrence in two widely spaced wildlife areas
suggests that it is likely to be found in additional Ohio locations.

Sigara hubbelli: This species has previously been reported for all of the
states bordering Ohio (Indiana, Kentucky, Pennsylvania, and West Virginia)
except Michigan to the north. Based on the distributional data presented by
both Hungerford (1948) and Polhemus et al. (1988), Ohio lies well within its
known range and it is no surprise that it has been found within the State. A
single male specimen was taken from Killdeer Plains Wildlife Area in Wyandot
County. Although the single male specimen was taken by black light sam-
pling, several specimens taken by dipnet sampling, from two additional locali-
ties within Ohio (Table 1 ), are in the first author's private collection. This fur-
ther serves to establish the presence of this species within Ohio.

Trichocorixa kanza: The report of this species in Ohio extends the north-
em distribution of this primarily southern species. Of the five new species
reported from Ohio, this species was taken in the largest numbers and was one
of the least anticipated. It has been taken from only one state, Pennsylvania,
bordering Ohio. In addition to its historical distribution in the southeastern
United States (Polhemus et al., 1988), the northern records of this species in-
clude the District of Columbia, Delaware, Maryland, Pennsylvania, and Wis-
consin (Hilsenhoff, 1984; Polhemus et al, 1988).

Palmacorixa buenoi: This species has previously been reported for all states
bordering Ohio (Indiana, Michigan, Pennsylvania, and West Virginia), except

342 ENTOMOLOGICAL NEWS

Kentucky to the south. Its reported distribution extends from Ontario south to
Florida and west to Iowa (Polhemus et al., 1988). This vast distribution, and
records for bordering states, clearly place Ohio within its range.

Corisella inscripta: Historical records for the genus Corisella are prima-
rily from western North America (Hungerford, 1948). Two species of this ge-
nus have been recorded from states east of the Mississippi River: Corisella
edulis (Champion), 1901 and Corisella tarsalis (Fieber), 1851 (Hilsenhoff,
1984; Hungerford, 1948; Polhemus et al., 1988). Prior to this paper, the east-
ernmost records of Corisella inscripta were from Missouri (Polhemus et al.,
1988) and a single male specimen from Arkansas (Cochran and Harp, 1990).
Corisella inscripta represents not only a new genus for Ohio, but also a con-
siderable eastern geographical range extension.

ACKNOWLEDGMENTS

Funding for projects from which this work is derived came from the Ohio Division of Wildife's
"Do Something Wild!" Income Tax Check-off Program, U.S. Fish & Wildlife Service's Partner-
ships for Wildlife Program, and the Procter & Gamble Co., Inc. Special thanks to George L.
Harp, Arkansas State University and Benjamin A. Foote, Kent State University for preliminary
review of this manuscript. Suggestions by two anonymous reviewers further improved the paper.

LITERATURE CITED

Chordas, S.W. III. 1998. First report ofHesperocorixa semilucida (Hemiptera: Corixidae) from

Ohio withnotes on distribution, habitat, and morphology. Ohio Biol. Survey Notes Series (in

review).
Cochran, B.C. and G.L. Harp. 1990. The aquatic macroinvertebrates of the St. Francis Sunken

Lands in northeast Arkansas. Proc. Ark. Acad. Sci. 44:23-27.

Hilsenhoff, W. L. 1984. Aquatic Hemiptera of Wisconsin. Great Lakes Entomol. 17( 1 ):29-50.
Hungerford, H.B. 1948. The Corixidae of the Western Hemisphere (Hemiptera). Univ. Kans.Sci.

Bull. 32:1-827.
Polhemus, J.T., R.C. Froeschner, and D.A. Polhemus. 1988. Family Corixidae Leach, 1815

the water boatman. Pages 93-120. In: T.J. Henry and R.C. Froeschner (eds.), Catalog of the

Heteroptera, or true bugs, of Canada and the continental United States. E.J. Brill. New York.

958 pp.
Sailer, R.I. 1948. The genus Trichocorixa (Corixidae, Hemiptera). Pages 289-407. In: H.B.

Hungerford (ed.), The Corixidae of the Western Hemisphere (Hemiptera). Univ. Kans. Sci.

Bull. 32:1-827.

Truxal, F.S. 1953. A revision of the genus Buenoa. Univ. Kans. Sci. Bull. 35:1351-1523.
Williams, R.N., M.S. Ellis, and D.S. Fickle. 1996. Insects in the Killbuck Marsh Wildlife Area,

Ohio: 1994 survey. Ohio J. Sci. 96(3):34-40.

Vol. 109, No. 5, November & December, 1998 343

TWO CORRECTIONS:
ARADIDAE, TINGIDAE (HETEROPTERA) 1

Richard C. Froeschner^

ABSTRACT: (1) Makes modification of captions and text to correct transposed captions for
Chelonoderus stylatus and Isodermus gayi in world list of flat bugs. (2) Rewrites couplets 1 9-22
of key to world genera of lace bug tribe Phatnomatini to properly place genus Minitingis.

( \ ) Kormilev and Froeschner 's ( 1 987) "Flat Bugs of the World" [Entomogr.,
5:1-246] contains a transposition of names of the figures on pages 220 and
221. The illustration on page 220 should be labeled "Fig. 2. Chelonoderus
stylatus Usinger. SEE p. 122."; and the illustration on page 221 should be
labeled "Fig. 3. Isodermus gayi (Spinola). SEE p. 94." This necessitates the
following changes in the text: on p. 94 the reference to the figure for Isodermus
gayi should read "Fig. 3 (p. 221 )"; and on p. 1 22 the reference for the figure of
Chelonoderus stylatus should read "Fig. 2 (p. 220)."

(2) In Froeschner,s (1996) "Lace Bug Genera of the World" [Smithson.
Contr. Zool, 574:i-iv, 1 -44] the genus Minitingis is misplaced within the key to
the world genera of the tribe Phatnomatini (pages 17-19). Couplets 1 9-22 must
be modified to read as follows:

19. Abdomen ventrally on basal half or more with a distinctly impressed groove

along midline 20

Abdomen ventrally not distinctly impressed along midline 21

20. Paranotum narrowest opposite humerus, thence widened cephalad to 4 or

more row of cells Phatnoma Fieber

Paranotum anteriorly not or only slightly (1-2 cells) widened Minitingis Barber

2 1 . Head with a clypeal spine 22

Head without a clypeal spine 25

22. Occipital spines nearly or quite as long as horizontal diameter of an

eye Gonycentrum Bergroth

Occipital spines absent or much shorter than horizontal diameter of an eye 23

Couplets 23-25 as published.

ACKNOWLEDGMENTS

Thanks are extended to T. J. Henry (U.S.D.A. -Systematic Entomology Laboratory) and P. J.
Spangler, Smithsonian Institution for reviews of this paper.

1 Received May 29, 1998. Accepted June 13, 1998.

2 Department of Entomology, MRC- 105, National Museum of Natural History, Washington, D.C.,
20560.

ENT. NEWS 109(5): 343, November & December, 1998

344 ENTOMOLOGICAL NEWS

NEW NAME FOR A GENERIC HOMONYM IN
TELOGANODIDAE (EPHEMEROPTERA) 1 * 2

W. P. McCafferty 3 , T.-Q. Wang 4

McCafferty and Wang ( 1 997) proposed the name Nadinella McCafferty
and Wang for two species of South African mayflies (family Teloganodidae)
that represented a new distinctive genus with numerous diagnostic features,
including, in the larvae, possession of two rows of denticles on the claws and
lamellate gills on abdominal segments 2-5. These mayflies had previously been
considered in the genus Ephemerellina Lestage. John Page of Great Britain
has kindly informed us by letter, however, that the name Nadinella is preoccu-
pied by a fossil protozoan described in 1 899 by Eugene Penard (Penard 1 899).
That name was more recently listed in Moore (1964), but we had missed the
citation in the Zoological Record and are not schooled in general invertebrate
paleontology.

We therefore propose Nadinetella McCafferty and Wang, nomen novum
for Nadinella McCafferty and Wang, 1 997: 399, nee Nadinella Penard, 1 899: 82.
The name Nadinetella incorporates the given name of Nadine McCafferty, for
whom it is named and who was of considerable aid in collecting specimens of
the new genus in South Africa.

LITERATURE CITED

McCafferty, W. P. and T.-Q. Wang. 1 997. Phylogenetic systematics of the family Teloganodidae
(Ephemeroptera: Pannota). Ann. Cape Provincial. Mus., Natural Hist. 19: 387-437.

Moore, R. C. (ed.). 1964. Treatise on invertebrate paleontology. Part C Protista 2. Sarcodinia,
chiefly 'thecamoebians' and Foraminifera. Volumes 1 & 2. Geol. Soc. Am., Lawrence, Kan-
sas.

Penard, E. 1899. Les Rhizopodes de faune profonde dans le Lac Leman. Rev. Suisse Zool.
7: 1-142.

' Received and accepted August 10, 1998.

2 Purdue Agr. Res. Prog. J. No. 16784.

3 Dept. of Entomology, Purdue University, West Lafayette, IN 47907.

4 23 Marsh Creek Dr., Mauldin, SC 29662.

ENT. NEWS 109(5): 344, November & December, 1998

Vol. 109, No. 5, November & December, 1998 345

A NEW NORTH AMERICAN GENUS OF BAETIDAE
(EPHEMEROPTERA) AND KEY TO BAETIS COMPLEX

GENERA 1 ' 2

C. R. Lugo-Ortiz, W. P. McCafferty 3

ABSTRACT: Plauditus, n. gen. (Ephemeroptera: Baetidae) is established for certain North
American Baetis complex species that lack hindwingpads and a developed median caudal fila-
ment in the larval stage and that were recently transferred from Pseudodoeon to Baetis or
Barbaetis. Thus, P. alachua, n. comb., P. armillatus, n. comb., P. bimaculatus, n. comb., P. cestus,
n. comb., P. cinctutus, n. comb., P. dubius, n. comb., P. elliotti, n. comb., P.futilis, n. comb., P.
punctiventris, n. comb., P. rubrolateralis, n. comb. P. veteris, n. comb., and P. virilis,
n. comb, are contained in the new genus. Plauditus cestus is the type species. The larval stage
of Plauditus is differentiated from other genera of the Baetis complex by numerous character-
istics, and the adult stage is differentiated from other North American baetids also having double
marginal intercalaries in the forewings and lacking hindwings (certain Acentrella, Apobaetis,
and Paracloeodes). A new, simplified, and illustrated generic key to the North American Baetis
complex larvae is provided.

Small minnow mayflies (Ephemeroptera: Baetidae) are relatively well
known in North America north of Mexico, with 143 species among 20 genera
currently reported (McCafferty 1996, 1997a; see also McCafferty and Silldorff
1998, Wiersema 1998, Wiersema and McCafferty 1998). Larvae are distin-
guished by having the initial lateral branches of the epicranial suture located
anterior to (below) the lateral ocelli (Wang and McCafferty 1996: Figs. 1-6)
and the femoral apices with a ventrally oriented dorsal lobe (Wang and
McCafferty 1996: Figs. 13-16). Adults are distinguished by having forewing
veins IMA, MA 2 , IMP, and MP 2 basally detached, and three-segmented mid-
and hindtarsi. Male adults are further distinguished by the presence of mem-
branous penes and turbinate compound eyes (the South American genus
Aturbina Lugo-Ortiz and McCafferty [1996] lacks turbinate compound eyes).
Despite being easily recognized at the family level, several North American
baetid species have been difficult to assign to genera. This situation is prima-
rily due to the fact that reductive trends and relative high frequency of ho-
moplasy within the family have limited the number of reliable diagnostic char-
acteristics of use at the genus level.

North American species previously assigned to Pseudodoeon Klapalek
are examples that have required revised generic placement as our knowledge

1 Received January 22, 1998. Accepted March 13, 1998.

2 Purdue Agricultural Research Program Journal No. 15576.

* Department of Entomology, Purdue University, West Lafayette, IN 47907.

ENT. NEWS 109(5): 345-353, November & December, 1998

346 ENTOMOLOGICAL NEWS

of genus level systematics has improved. North American species once as-
signed to Pseudocloeon have the following combination of characteristics:
absence of hindwings (and hindwingpads), presence of double marginal
intercalaries in the forewings, and presence of a highly reduced medial caudal
filament in the larval stage. As a result of the restriction of the concept of
Pseudocloeon to its Oriental type (Waltz and McCafferty 1985, 1987), all North
American species once considered to belong to Pseudocloeon were transferred
toApobaetis Day (Waltz and McCafferty 1986), Acentrella Bengtsson (Waltz
and McCafferty 1987), Baetis Leach (McCafferty and Waltz 1990), and
Barbaetis Waltz and McCafferty (McCafferty and Waltz 1990).

Our research indicates that all 1 1 species of Pseudocloeon that were provi-
sionally assigned to Baetis by McCafferty and Waltz (1990) and also Barbaetis
cestus (Provonsha and McCafferty) (originally described in Pseudocloeon)
represent a new Baetis-comp\c\ genus distinguishable by numerous charac-
teristics. We herein describe the new genus and provide a revised generic key
to the North American larvae of the Baetis complex.

Plauditus Lugo-Ortiz and McCafferty, NEW GENUS

Larva. Head: Antennae as long as head capsule or longer; scapes without distal notch,
subequal in length in length to pedicels (Fig. 1 ). Labrum (Fig. 2) broadly rounded anteriorly,
with anteromedial notch. Hypopharynx (Fig. 3) with lingua and superlinguae broadly rounded
apically. Left mandible (Fig. 4) with incisors fused; prostheca apically broad and denticulate.
Right mandible with incisors apically fused (Fig. 5) or slightly cleft; prostheca slender, apically
denticulate. Maxillae (Fig. 6) with palps two segmented. Labium (Fig. 7) compact; glossae
slightly shorter than paraglossae, apically narrower than base; paraglossae broad, broadly rounded
apically; palps three segmented; palp segment 1 slightly longer than segments 2 and 3 com-
bined; palps segment 2 as long as or slightly longer than segment 3; segment 3 relatively broad
apically, approaching truncate, sometimes appearing slightly concave, medial margin straight
to slightly convex distally (never convergent apically from base). Thorax: Hindwingpads ab-
sent. Legs (Fig. 8) with femora with villopore present (Fig. 9) and dorsal row of long, robust
setae. Tarsal claws (Fig. 10) somewhat elongate and with only weak curvature, with one row of
long, sharp denticles. Abdomen: Gills on abdominal segments 1 -7, broadly rounded, untracheated
or poorly tracheated, marginally smooth. Terga (Fig. 1 1 ) creased, with minute, sharp, triangular
spines, or with minute, fine, simple setae scattered over surface. Cerci with abundant fine,
simple setae medially. Medial caudal filament subequal in length to abdominal segment 10.

Adult. Head: Male compound eyes circular, somewhat enlarged. Male foretibiae 1.3-1.5x
length of femora. Mesoscutum with small, rounded anterior process in lateral view (Fig. 12).
Forewings with paired marginal intercalaries. Hindwings absent. Genital forceps (Fig. 13) three
segmented; segment 1 distomedially produced; segment 2 slender, strongly arched; segment 3
slender, ellipsoidal.

Type species. Pseudocloeon cestum Provonsha and McCafferty.

Vol. 109, No. 5, November & December, 1998

347

Figs, l-l I. Plauditus punctiventris. 1. Antennal scape and pedicel. 2. Labrum (dorsal). 3. Hy-
popharynx. 4. Left mandible. 5. Right mandible. 6. Left maxilla. 7. Labium (left-ventral; right-
dorsal). 8. Right foreleg (posterior face). 9. Villopore. 10. Tarsal claw. 1 1. Detail of tergum 4.
12. Adult mesoscutum (lateral). 13. Male genitalia.

348 ENTOMOLOGICAL NEWS

Included species. (In addition to the recombined species listed below, a
new species is presently being described by McCafferty and Waltz [1998], and
one other is in manuscript [R. D. Waltz, pers. comm.]).
Plauditus alachua (Berner), n. comb.

Pseudocloeon alachua Berner 1940:58 (larva; male, female adults).

Baetis alachua (Berner): McCafferty and Waltz 1990:775.
Plauditus armillatus (McCafferty and Waltz), n. comb.

Pseudocloeon parvulum McDunnough 1932:210 (larva; male, female adults),
[secondary homonym].

Baetis armillatus McCafferty and Waltz 1990:775. [renamed].
Plauditus bimaculatus (Bemer), n. comb.

Pseudocloeon bimaculatum Berner 1946:79 (larva; male, female adults).

Baetis bimaculatus (Bemer): McCafferty and Waltz 1990:775.
Plauditus cestus (Provonsha and McCafferty), n. comb.

Pseudocloeon cesium Provonsha and McCafferty 1982:28 (larva; male, female adults).

Barbaetis cestus (Provonsha and McCafferty): McCafferty and Waltz 1990:777.
Plauditus cinctutus (McCafferty and Waltz), n. comb.

Pseudocloeon cingulatum McDunnough 1931:85 (male, female adults); Ide 1937:236
(larva), [secondary homonym].

Baetis cinctutus McCafferty and Waltz 1990:776. '[renamed].
Plauditus dubius (Walsh), n. comb.

Cloeon dubium Walsh 1 862:380 (male, female adults).

Pseudocloeon dubium (Walsh): McDunnough 1924b: 115; Ide 1937:237 (larva).

Pseudocloeon chlorops McDunnough 1923:45.

Baetis dubius (Walsh): McCafferty and Waltz 1990:775.
Plauditus elliotti (Daggy), n. comb.

Pseudocloeon elliotti Daggy 1945:392 (male, female adults).

Baetis elliotti (Daggy): McCafferty and Waltz 1990:775.
Plauditus futilis (McDunnough), n. comb.

Pseudocloeon futile McDunnough 1931 :86 (male, female adults).

Baetis futile (McDunnough): McCafferty and Waltz 1990:775.

Baetis futilis (McDunnough): McCafferty 1997b:318.
Plauditus punctiventris (McDunnough), n. comb.

Pseudocloeon punctiventris McDunnough 1923:45 (male, female adults); Ide 1937:237
(larva).

Pseudocloeon anoka Daggy 1945:391.

Pseudocloeon edmundsi Jensen 1969: 14.

Pseudocloeon myrsum Burks 1953: 139.

Baetis punctiventris (McDunnough): McCafferty and Walt/. 1990:776.
Plauditus rubrolateralis (McDunnough), n. comb.

Pseudocloeon rubrolaterale McDunnough 1931:86 (male, female adults).

Baetis rubrolaterale (McDunnough): McCafferty and Waltz 1990:776.

Baetis rubrolateralis (McDunnough): McCafferty 1997a:318.
Plauditus veteris (McDunnough), n. comb.

Pseudocloeon veteris McDunnough I924a:8 (male, female adults).

Baetis veteris (McDunnough): McCafferty and Waltz 1990: 776.
Plauditus virilis (McDunnough), n. comb.

Cloeon virile McDunnough 1923:46 (male, female adults).

Pseudocloeon virile (McDunnough): McDunnough I924b: 1 16; Ide 1937:239 (larva).

Baetis virile (McDunnough): McCafferty and Waltz 1990:776.

Baetis virilis (McDunnough): McCafferty 1997b:3l2.

Vol. 109, No. 5, November & December, 1998 349

Distribution. Species of Plauditus are presently known from the conter-
minous USA and the lower tier of provinces of Canada. There remains the
possibility that the genus may eventually be found in Mexico because it is
presently well represented in Texas (Lugo-Ortiz and McCafferty 1995,
Wiersema and McCafferty 1998). It is also possible that Plauditus is repre-
sented in the eastern Palearctic by certain species that otherwise may have
been variously regarded as Acentrella, Baetis, or Pseudocloeon. We have no
evidence of this at the present.

Etymology. The generic name is an arbitrary combination of Latin letters
having a transliteral meaning of "little applause."

Diagnosis. The presence of the larval femoral villopore indicates that
Plauditus is a member of the Baetis complex of genera, represented in North
America north of Mexico by Acentrella, Baetis, Barbaetis, Heterocloeon
McDunnough, and Labiobaetis Novikova and Kluge. Within that complex,
larvae of Plauditus are distinguished by the following combination of charac-
teristics: lacking a medial field of setae dorsally on the labrum (Fig. 2); having
the right mandible with an apically denticulate prostheca (Fig. 5); lacking a
protuberance on the apex of maxillary palp segment 3 (Fig. 6); having glossae
shorter than paraglossae, relatively broad paraglossae, and a subquadrate palp
segment 3 (Fig. 7); lacking dorsal setae on the apical half of the glossae (viz.
well-developed medial aspect) (Fig. 7); lacking hindwingpads; lacking procoxal
osmobranchia; lacking clavate setae on the dorsal margin of the tibiae and tarsi
(Fig. 8); and having a highly reduced medial caudal filament (Provonsha and
McCafferty 1 982: Fig. 8). A more detailed diagnosis of Plauditus larvae in
relation to those of other Baetis-comp\e\ genera can be performed using the
identification key provided below.

Adults of Plauditus are similar to hindwingless Acentrella, Apobaetis Day,
and Paracloeodes Day. Adults of Plauditus differ from those of Acentrella in
having a small, rounded anterior process on the mesoscutum (Fig. 12), and
from those of Apobaetis by lacking a subconical process between the male
genital forceps (Fig. 1 3). Adults of Plauditus differ from those of Paracloeodes
in that segment 2 of the male genital forceps is slightly produced distomedially
and tends to be more slender and strongly arched (Fig. 13).

350 ENTOMOLOGICAL NEWS

KEY TO LARVAE OF NORTH AMERICAN BAETIS-COMPLEX GENERA

1 . Both mandibles lacking tuft of setae between incisors and prostheca
(Figs. 4, 5); villopore usually apparent on femora (Fig. 9); claws never

approaching or exceeding length of tarsi (Figs. 8, 10) Baetis complex, 2

One or both mandibles with tuft of setae between incisors and prostheca

(Fig. 14); villopore absent from femora; claw length variable other Baetidae

2. Antennal scapes with distal notch (Fig. 15); maxillary palp segment 2
with subapical excavation (Fig. 16); hindwingpads present; median caudal

filament developed Labiobaetis

Antennal scapes without distal notch (Fig. 1); maxillary palp segment 2

without subapical excavation (Fig. 6); hindwingpads present or absent;

median caudal filament developed or highly reduced 3

3. Median caudal filament developed; hindwingpads present 4

Median caudal filament reduced; hindwingpads present or absent 5

4. Antennae approximately twice length of head capsule Baetis

Antennae subequal in length to head capsule Barbaetis

5. Procoxae with (Fig. 19) or without gills; claws with two rows of denticles
(second row minute and sometimes conspicous only under high

magnification) Heterocloeon

Procoxae without gills; claws with one row of denticles 6

6. Hindwingpads absent 7

Hindwingpads present, sometimes minute 8

7. Femora, tibiae, and tarsi without row of long setae (Fig. 8); labial palp
segment 3 subquadrate, with medial margin almost straight (not receding

from base) and sometimes appearing slightly convex apically (Fig. 7) Plauditus

Femora, tibiae, and tarsi with row of long setae (Fig. 18); labial palp
segment 3 apically rounded, with medial margin receding from base
(Fig. 17) Acentrella

8. Labial palp segment 2 with well-developed distomedial thumb (Fig.

20); abdominal tergal scales present (Fig. 21) Baetis

Labial palp without well-developed distomedial thumb (Fig. 17);

abdominal tergal scales absent Acentrella

Vol. 109, No. 5, November & December, 1998

351

Figs. 1 4-2 1 . 1 4. Fallceon quilleri, left mandible (pointer towards setal tuft) (modified from Lugo-
Ortiz et al. [1994]). 15-16. Labiobaetis propinquus 15. Antennal scape and pedicel (pointer to-
wards notch) (modified from McCafferty and Waltz [1995]). 16. Left maxilla (pointer towards
excavation) (modified from McCafferty and Waltz [1995]). 17-18. Acentrella turbida. 17. La-
bium (modified from McCafferty et al. [1994]). 18. Right foreleg (modified from McCafferty et
al. [1994]). 19. Heterocloeon berneri, forecoxal gill (pointer towards gill) [modified from Miiller-
Liebenau [1974]). 20-21. Baetis bicaudatus, labial palp (pointer towards thumb) (modified from
Morihara and McCafferty [1979]). 21. Detail of tergum 4.

352 ENTOMOLOGICAL NEWS

ACKNOWLEDGMENTS

We thank R. D. Waltz (Department of Natural Resources, Indianapolis, Indiana) for discuss-
ing this paper, and A. V. Provonsha and R. P. Randolph (Purdue University, West Lafayette,
Indiana) for preliminary review of the manuscript.

LITERATURE CITED

Berner, L. 1940. Baetine mayflies from Florida (Ephemeroptera). Fla. Entomol. 23: 33-45, 49-
62.

Berner, L. 1946. New species of Florida mayflies (Ephemeroptera). Fla. Entomol. 28: 60-82.

Burks, B. D. 1953. The mayflies, or Ephemeroptera, of Illinois. Bull. 111. Nat. Hist. Surv. 26: 1 -
216.

Daggy, R. H. 1945. New species and previously undescribed naiads of some Minnesota may-
flies (Ephemeroptera). Ann. Entomol. Soc. Am. 38: 373-396.

Ide, F. P. 1937. Descriptions of eastern North American species of baetine mayflies with particu-
lar reference to the nymphal stages. Can. Entomol. 69: 219-231, 235-243.

Jensen, S. L. 1969. A new species of Pseudocloe on from Idaho. Pan-Pac. Entomol. 45: 14-15.

Lugo-Ortiz, C. R. and W. P. McCafferty. 1995. The mayflies (Ephemeroptera) of Texas and
their biogeographic affinities. Pp. 151-169. In: L. D. Corkum and J. J. H. Ciborowski (eds.),
Current directions in research on Ephemeroptera. Canadian Scholars' Press, Toronto.

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996. Aturbina georgei gen. et sp. n.: a small minnow
mayfly (Ephemeroptera: Baetidae) without turbinate eyes. Aq. Insects 18: 175-183.

Lugo-Ortiz, C. R., W. P. McCafferty, and R. D. Waltz. 1994. Contribution to the taxonomy of
the Panamerican genus Fallceon (Ephemeroptera: Baetidae). J. N. Y. Entomol. Soc. 102:
460-475.

McCafferty, W. P. 1996. The Ephemeroptera species of North America and index to their com-
plete nomenclature. Trans. Entomol. Soc. Am. 122: 1-54.

McCafferty, W. P. 1997a. Ephemeroptera. Pp. 89-117. In: R. W. Poole and P. Gentili (eds.),
Nomina Insecta Nearctica: a checklist of the insects of North America, Vol. 4: Non-holom-
etabolous orders. Entomol. Info. Services, Rockville, Maryland.

McCafferty, W. P. 1997b. Name adjustments and a new synonym for North American
Ephemeroptera species. Entomol. News 108: 318-319.

McCafferty, W. P. and E. L. Silldorff. 1998. Reared association and equivalency of Baetis
adonis and caelestis (Ephemeroptera: Baetidae). Entomol. News 109: 261-265.

McCafferty, W. P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera)
of North and Middle America. Trans. Am. Entomol. Soc. 1 16: 769-799.

McCafferty, W. P. and R. D. Waltz. 1995. Labiobaetis (Ephemeroptera: Baetidae): new status,
new North American species, and related new genus. Entomol. News 106: 19-28.

McCafferty, W. P. and R. D. Waltz. 1998. New species of the small minnow mayfly genus
Plauditus (Ephemeroptera: Baetidae) from South Carolina. Entomol. News 109: 354-356.

McCafferty, W. P., M. J. Wiggle and R. D. Waltz. 1994. Systematics and biology ofAcentrelta
turbida (McDunnough) (Ephemeroptera: Baetidae). Pan-Pac. Entomol. 70: 301-308.

McDunnough, J. 1923. New Canadian Ephemeridae with notes. Can. Entomol. 55: 39-50.

McDunnough, J. 1924a. New Ephemerida from Illinois. Can. Entomol. 56: 7-9.

McDunnough, J. 19245. New Canadian Ephemeridae with notes II. Can. Entomol. 56: 90-98,
113-122, 128, 133.

McDunnough, J. 1931. New species of Canadian Ephemeroptera. Can. Entomol. 63: 82-93.

McDunnough, J. 1932. New species of North American Ephemeroptera, II. Can. Entomol. 64:
209-215.

Morihara, D. K. and W. P. McCafferty. 1979. The Baetis larvae of North America
(Ephemeroptera: Baetidae). Trans. Am. Entomol. Soc. 105: 139-221.

Vol. 109, No. 5, November & December, 1998 353

Miiller-Liebenau, I. 1974. Rheobaetis: a new genus from Georgia (Ephemeroptera: Baetidae).

Ann. Entomol. Soc. Am. 67: 555-567.
Provonsha, A. V. and W. P. McCafferty. 1982. New species and previously undescribed larvae

of North American Ephemeroptera. J. Kansas Entomol. Soc. 55: 23-33.
Walsh, B. D. 1862. List of the Pseudoneuroptera of Illinois contained in the cabinet of the writer,

with descriptions of over 40 new species, and notes on their structural affinities. Proc. Acad.

Nat. Sci. Phila. 2nd ser., pp. 361-402.
Waltz, R. D. and W. P. McCafferty. 1985. Redescription and new lectotype designation for the

type species of Pseudocloeon, P. kraepelini Klapalek (Ephemeroptera: Baetidae). Proc.

Entomol. Soc. Wash. 87: 800-804.
Waltz, R. D. and W. P. McCafferty. 1986. Apobaetis etowah (Traver), a new combination in

Nearctic Baetidae (Ephemeroptera). Proc. Entomol. Soc. Wash. 88: 191.
Waltz, R. D. and W. P. McCafferty. 1987. Systematics of Pseudocloeon, Acentrella, Baetiella,

and Liebebiella, new genus (Ephemeroptera: Baetidae). J. N. Y. Entomol. Soc. 95: 553-568.
Wang, T.-Q. and W. P. McCafferty. 1996. New diagnostic characters for the mayfly family

Baetidae (Ephemeroptera). Entomol. News 107: 207-212.
Wiersema, N. A. 1998. Camelobaetidius variabilis (Ephemeroptera: Baetidae), a new species

from Texas, Oklahoma and Mexico. Entomol. News 109: 21-26.
Wiersema, N. A. and W. P. McCafferty. 1998. A new species of Pseudocentroptiloides

(Ephemeroptera: Baetidae), with revisions of other previously unnamed baetid species from

Texas. Entomol. News 109: 110-116.

354 ENTOMOLOGICAL NEWS

A NEW SPECIES OF THE SMALL MINNOW MAYFLY

GENUS PLAUDITUS (EPHEMEROPTERA: BAETIDAE)

FROM SOUTH CAROLINA 1

W. P. McCafferty 2 , R. D. Waltz 3

ABSTRACT: A newly discovered species of Baetidae (Ephemeroptera), Plauditus gloveri, n.
sp., is described from larvae taken in South Carolina. Characterization or combinations thereof
involving color pattern, antennal length, labral setation, maxillary palp shape and size, distal
shape of the labial palps, claw curvature, and tergal spines distinguish the new species. Antennal
and claw characterization suggests a possible close relationship with P. cestus.

Our examination of larval material of small minnow mayflies recently taken
from streams in South Carolina revealed a distinctive new species. Generic
characterization of the new species place it within the recently erected genus
Plauditus Lugo-Ortiz and McCafferty (1998). We are honored to name the
new species after James B. Glover (Columbia, South Carolina), who collected
the original material.

Plauditus gloveri NEW SPECIES

(Figs. 1-10)

Larva. Body length, 4.5-5.0 mm; antennae length, ca. 1.0 mm; cerci length 1.8-2.0 mm.
Base color pale yellow; markings light to medium brown. Head: Head capsule with submedian
rows of short, irregular, transverse dashes on either side of medial trunk of epicranial suture,
with pair of short transverse dashes below each compound eye, and with oblique pair of spots
above each antennal base. Antennae not marked. Dorsal chetotaxy of labrum as in Fig. 1 , with
pair of long, simple, hairlike, submarginal setae located approximately half way between lat-
eral margin and median line of labrum, and extending beyond distal labral margin for about
half length of seta (in some, extending setae slightly more proximate to each other than shown
in Fig. 1). Maxillae as in Fig. 2, with maxillary palp narrow and extending beyond apex of
galealacinia. Mandibles as in Figs. 3 and 4. Terminal segment of labial palp (Fig. 5) slightly
broadening apically, with very slight distolateral point; apical margin slightly concave in lat-
eral half, and rounded and somewhat bulbous in medial half. Thorax: Pronotum with three
prominent spots (one medial, two submedial) near anterior margin. Mesonotum with mostly
scattered, lateral light brown spots. Metanotum with pair of prominent submedial spots.
Hindwingpads absent. Legs not generally marked; anterior face of femora with pair of dorsal
and ventral dashes as in Fig. 6, dorsal dash sometimes appearing as two partially connected
elongate spots. Claws (Fig. 7) relatively straight. Abdomen: Abdominal segment 7 slightly
darker than other segments (possibly indicating darker segment band in other individuals),
more noticeable ventrally, and perceptible on specimens only when using black background
and low magnification; abdominal terga 1 -9 (Fig. 8) each with pair of dark submedian spots
(lateral spots also usually present); tergum 2 with conspicuous medial V-shaped mark at ante-
rior margin; tergal surfaces with weak creases and minute, simple setae; posterior margin of

1 Received January 22, 1998. Accepted March 13, 1998.

2 Department of Entomology, Purdue University, West Lafayette, IN 47907, USA.

3 IDNR, Division of Entomology and Plant Pathology, 402 West Washington, Rm. W-290,
Indianapolis, IN 46204, USA.

ENT. NEWS 109(5): 354-356, November & December, 1998

Vol. 109, No. 5, November & December, 1998

355

terga with slightly separated, triangular shaped spines slightly longer that basal width. Ab-
dominal sterna (Fig. 9) with submedian pair of diffuse brown spots; darker medial maculation
present on sterna 2-8 (prominent and larger on stema 3-5 and overlapping sterna somewhat at
anterior and posterior sternal margins); lateral subdermal striations evident on sterna 1-6. Gills
(Fig. 10) with margins rounded apically, smooth, and with only very few scattered, short, simple
setae; gill tracheal trunk not extending much beyond midlength of gill and with lateral branches
only weakly developed (details of tracheation evident only under high magnification). Cerci
with three light brown bands (not always perceptible). Median caudal filament highly reduced,
subequal in length to mid-dorsal length of tergum 10.

Adult. Unknown.

Material examined. Holotype: Larva, SOUTH CAROLINA, Cherokee County, Kings Creek @
S-l 1-209, 3 miles west of Smyrna, VI-25-1995, J. Glover (deposited in the Purdue Entomologi-
cal Research Collection, West Lafayette, Indiana). Paratypes: 1 larva (broken, some pans miss-

Figs. I -10. Plauditus gloveri. 1. Labrum (dorsal). 2. Maxilla. 3. Right mandible. 4. Left man-
dible. 5. Labial palp. 6. Midfemora (anterior face). 7. Claw. 8. Abdomen (dorsal). 9. Abdomen
(ventral). 10. Gill 5.

356 ENTOMOLOGICAL NEWS

ing), same data and deposition as holotype; 2 larvae (mounted on slides, medium Euparol), SOUTH
CAROLINA, Fairfield County, Little River S-20-60, 3.1 miles southwest of Jenkinsville, VI-
28-1995, J. Glover (same deposition as holotype).

Remarks. Larvae of Plauditus gloveri are easily distinguished from all
other known larvae of Plauditus on the basis of morphological and color pat-
tern characterization given above. Seven other species of Plauditus have been
reported from the southeast: P. alachua (Berner), P. armillatus (McCafferty
and Waltz), P. bimaculatus (Berner), P. cinctutus (McCafferty and Waltz), P.
dubius (Walsh), P. punctiventris ((McDunnough), and P. rubrolateralis
(McDunnough). On the basis of its relatively straight tarsal claw, P. gloveri
may be closely related to P. cestus (Provonsha and McCafferty). Plauditus
gloveri and P. cestus have antennae that are clearly shorter than other known
species of Plauditus, although those of P. gloveri are not nearly as short as
those off. cestus (Provonsha and McCafferty 1982). Plauditus gloveri, how-
ever, differs from P. cestus in numerous other characteristics, including the
position of the pair of extending dorsal labral setae, the relatively much longer
maxillary palps, femoral markings, the longer and somewhat more pointed
tergal spines, and patterning of the abdomen and cerci.

Abdominal segment banding on segment 7 is evident but not well devel-
oped on the specimens of P. gloveri that we have examined. It is probable that
segment 7 banding will be more pronounced in larger series. All of our speci-
mens were female larvae, and it is also possible that the banding as well as
other patterning will be more developed in males. This possibility is based on
the fact that such sexual dimorphism has been documented for certain other
species in the genus Plauditus (e.g., P. dubius and P. virilis [Ide 1937]). Ab-
dominal segment banding occurs on segment 5 in most specimens of P. cestus,
but has never been seen on segment 7.

Although all material of P. gloveri has been taken from the Broad River
Basin in South Carolina, no specific ecological data are yet associated with
this new species.

ACKNOWLEDGMENTS

We thank Jim Glover (Columbia, South Carolina) for collecting the new species and ini-
tially recognizing that it did not match other known species. We especially thank Arwin
Provonsha (West Lafayette, Indiana) for the illustrations used herein. This paper has been as-
signed Purdue Agricultural Research Program Journal No. 15602.

LITERATURE CITED

Ide, F. P. 1937. Descriptions of eastern North American species of baetine mayflies with particu-
lar reference to the nymphal stages. Can. Entomol. 69: 219-231, 235-243.

Lugo-Ortiz, C. R. and W. P. McCafferty. 1998. A new North American genus of Baetidae
(Ephemeroptera) and key to Baetis complex genera. Entomol. News 109: 345-353.

Provonsha, A. V. and W. P. McCafferty. 1982. New species and previously undescribed larvae
of North American Ephemeroptera. J. Kans. Entomol. Soc. 55: 23-33.

Vol. 109, No. 5, November & December, 1998 357

NEW SPECIES OF CLOEON AND DEMOULINIA

(EPHEMEROPTERA: BAETIDAE) FROM

MADAGASCAR 1 ' 2

C. R. Lugo-Ortiz, W. P. McCafferty 3

ABSTRACT: Cloeon emmanueli, new species, and Demoulinia insularis, new species, are de-
scribed from larvae from Madagascar. Cloeon emmanueli represents the first bonafide report of
Cloeon from the island. The species is distinguished by the setation of the labrum, abdomi-
nal color pattern, tergal armature, and irregular paraproctal spines. Cloeon cambouei, C. durani,
and C. irretitum are considered nomina dubia because they were described from subimagos
only. Demoulinia insularis is the first species of Demoulinia to be reported from the island. The
species is distinguished by the relatively wide anteromedial emargination of the labrum, eden-
tate tarsal claws, and numerous paraproctal spines.

The faunal composition of the small minnow mayflies (Ephemeroptera:
Baetidae) of Madagascar has recently received considerable attention (Lugo-
Ortiz and McCafferty 1997abef, 1998a), but requires additional study. Baetid
genera that were shown in those works to occur in Madagascar include
Afroptilum Gillies, Cheleocloeon Wuillot and Gillies, Dabulaman-ia Lugo-
Ortiz and McCafferty, Dicentroptilum Wuillot and Gillies, Edmulmeatus Lugo-
Ortiz and McCafferty, Herbrossus McCafferty and Lugo-Ortiz, Labiobaetis
Novikova and Kluge, Mulelocloeon Gillies and Elouard, and Xyrodromeus
Lugo-Ortiz and McCafferty. Lugo-Ortiz and McCafferty ( 1 998a) confirmed
that Nesoptiloides, first described from Madagascar by Demoulin (1973), was
a valid genus. Reports of adults of Centroptilum Eaton from Madagascar are,
however, highly tenuous because species previously assigned to that genus in
Africa have been shown to represent diverse evolutionary lineages not includ-
ing Centroptilum (Gillies 1990, Wuillot and Gillies 1994, Lugo-Ortiz and
McCafferty 1996abc, 1997cd, 1998ab, McCafferty et al. 1997). We expect
such adults in Madagascar to be members of the Centroptiloides complex (see
Lugo-Ortiz and McCafferty 1998a). Madagascar species described as Cloeon
Leach are based on subimagos (Navas 1 926, 1 930, 1 936) that cannot be placed
to genus with any reliability.

In this paper, we describe one species of Cloeon and one species of
Demoulinia Gillies based on larvae collected from Madagascar. The new spe-
cies of Cloeon represents the first substantiated report of the genus from Mada-
gascar, and the new species of Demoulinia is the first of that genus to be de-
scribed from the island. Examined specimens are housed in the Purdue Ento-
mological Research Collection, West Lafayette, Indiana.

1 Received March 11, 1998. Accepted April 4, 1998.

2 Purdue Agricultural Research Program Journal No. 15659.

- Department of Entomology, Purdue University, West Lafayette, IN 47907.

ENT. NEWS 109(5): 357-362, November & December, 1998

358 ENTOMOLOGICAL NEWS

Cloeon emmanueli Lugo-Ortiz and McCafferty, NEW SPECIES

Larva. Body length: 4.3-5.2 mm. Caudal filaments length: 3.0-4.7 mm. Head: Coloration
medium brown to medium yellow-brown, with no distinct markings. Antennae approximately
3.0x length of head capsule. Labrum (Fig. 1) with numerous long, fine, simple setae scattered
over surface. Hypopharynx as in Figure 2. Left mandible (Fig. 3) with seven denticles; prostheca
apically denticulate; tuft of long, fine, simple setae between prostheca and mola. Right man-
dible (Fig. 4) with outer set of incisors with four denticles and inner set with two denticles; tuft
of long, fine, simple setae between prostheca and mola. Maxillae (Fig. 5) with irregular row of
long, fine, simple setae submedially in midregion; palps three segmented; palp segment 1 ap-
proximately 0.63x length of segment 2 and 3 combined; segment 2 approximately 1.45x length
of segment 3; segment 3 poorly defined. Labium (Fig. 6) with glossae slightly longer than
paraglossae; glossae basally broad, apically narrow, with minute, fine, simple setae ventrally
near base; paraglossae broadly rounded apically, with three rows of long, fine, simple setae
ventrally and few minute, fine, simple setae scattered dorsally near base; palp segment 1 ap-
proximately 0.82x length of segments 2 and 3 combined; segment 2 subequal in length to
segment 3, with row of five to six minute, fine, simple setae dorsally; segment 3 with numerous
long, somewhat robust, simple setae scattered over surface. Thorax: Coloration medium yel-
low-brown, with no distinct markings. Hindwingpads absent. Legs (Fig. 7) pale yellow-brown;
femora with two rows of 10-12 robust, apically pointed setae dorsally and numerous short,
stout, simple seta ventrally; tibiae with few long, fine, simple setae and one long, robust,
simple setae dorsally near apex, and two rows of 15-17 robust, apically pointed, simple setae
ventrally; tarsi with few, long, fine, simple setae dorsally and two rows of robust, apically
pointed, simple and pectinate setae ventrally; tarsal claws (Fig. 8) with two rows of 15-20
minute to small denticles each. Abdomen: Coloration medium yellow-brown and medium
brown; segments 1 and 10 medium yellow-brown; segments 2 and 3 medium yellow-brown,
medium brown sublaterally; segments 4-9 medium yellow-brown, with small medium brown
markings anterolaterally. Sterna medium yellow-brown; sterna 4-9 with faint medium brown
broad band medially. Terga (Fig. 9) with numerous scale bases and few minute, fine, simple
setae scattered over surface; posterior triangular spination irregular. Gills 1-6 with two lamel-
lae, gill 7 single. Paraproct (Fig. 10) with 10-12 marginal spines and numerous minute, fine,
simple setae scattered over surface. Caudal filaments pale yellow-brown, with medium brown
annulations every three to four segments; terminal filament approximately 0.60x length of
cerci.

Adult. Unknown.

Material examined. Holotype: Larva, MADAGASCAR, Antsiranana Prov., Djabala R.,
1 1 km N W of Hell- Ville, Nosy Be, 25-X- 1 97 1 , G. F., C. H. Edmunds, and F. Emmanuel. Paratype:
Larva, same data as holotype [mouthparts, left foreleg, tergum 4, gills 4, and paraproct of one
larva mounted on slide (medium: Euparal)]. Additional material: Four larvae, same data as
holotype.

Etymology. We name this species after F. Emmanuel (Madagascar), who
assisted in its collection.

Discussion. Although no other species of Cloeon from Madagascar are
known from the larval stage, we expect that the setation of the labrum (Fig. \),
abdominal color pattern, tergal armature (Fig. 9), and paraproctal spination
(Fig. 10) will be diagnostic features of C. emmanueli.

The genus Cloeon has been reported from much of the world, although

Vol. 109, No. 5, November & December, 1998

359

reports of its presence in South America are considered incorrect (McCafferty
1998), and it is only adventive in North America (McCafferty 1996). Its pres-
ence in southern Africa and the Orient suggested that it could be present in
Madagascar. However, species previously assigned to Cloeon in Madagascar
include only the highly dubious C. durani Navas (1926), C. cambouei Navas
(1930), and C. irretitum Navas (1936). Cloeon durani and C. cambouei were
originally described from female subimagos, and C. irretitum was originally

Figs. 1-10. Cloeon emmanueli, larva. 1. Labrum (dorsal). 2. Hypopharynx. 3. Left mandible. 4.
Right mandible. 5. Right maxilla. 6. Labium (right- ventral; left-dorsal). 7. Right foreleg. 8.
Tarsal claw. 9. Tergum 4 (detail). 10. Paraproct.

360 ENTOMOLOGICAL NEWS

described from male and female subimagos. Because the three species are so
poorly known and because their taxonomic status cannot be corroborated, we
place the three names as nomina dubia. Thus, C. emmanueli is the only bona
fide species of Cloeon known from Madagascar at this time.

Demoulinia insularis Lugo-Ortiz and McCafferty, NEW SPECIES

Larva. Body length: 7.3 mm. Caudal filaments length: 3.5 mm. Head: Coloration me-
dium yellow-brown, with no distinct pattern. Antennal length unknown. Labrum (Fig. 1 1 ) with
numerous long, fine, simple setae scattered over surface. Hypopharynx as in Figure 12. Left
mandible (Fig. 13) with outer set of incisors with three denticles and inner set with two den-
ticles. Right mandible (Fig. 14) with outer set of incisors with three denticles and inner set with
two denticles. Maxillae (Fig. 15) with row of 10-12 long, fine, simple setae near crown of
galealaciniae and row of six to seven long, fine, simple setae submedially in midregion; palps
two segmented; palp segment 1 approximately 0.70x length of segment 2. Labium (Fig. 16)
with glossae subequal in length to paraglossae; glossae broadly rounded apically, with minute,
stout, simple setae dorsally; paraglossae acute apically, with numerous long, fine, simple setae
ventrally and three rows of long, fine, simple setae dorsally; palp segment 1 approximately
O.SOx length of segments 2 and 3 combined, with numerous long, fine, simple setae basomedially;
segment 2 approximately 2.25x length of segment 3, with numerous long, robust, simple setae
on distomedial process; segment 3 slender and elongate, with numerous long, robust, simple
setae scattered over surface. Thorax: Coloration medium yellow-brown, with no distinct mark-
ings. Hindwingpads absent. Legs (Fig. 17) pale yellow-brown; femora with numerous minute,
fine, simple setae and minute, stout, simple setae dorsally and ventrally, setae more abundant
ventrally; tibiae and tarsi with numerous minute, fine, simple setae dorsally and numerous
minute, stout, simple setae ventrally; tarsal claws approximately 0.63x length of tarsi, eden-
tate. Abdomen: Coloration medium brown to yellow-brown; tergum 1 medium brown, with no
markings; tergum 2 medium brown, with anteromedial pair of round, medium brown spots;
terga 3-9 medium brown anteriorly, yellow-brown posteriorly, with anteromedial pair of me-
dium brown dashes; terga 7-9 with submedial pair of round, medium brown spots in midregion;
tergum 10 yellow-brown. Sterna pale yellow brown, with no distinct pattern. Terga (Fig. 18)
with numerous scale bases; posterior triangular spines approximately 1.2x basal width. Gills
subtriangular, poorly tracheated, marginally smooth. Paraproct (Fig. 19) with numerous mar-
ginal spines, increasing in size apically. Caudal filaments pale yellow-brown; terminal fila-
ment subequal in length to cerci.

Adult. Unknown.

Material examined. Holotype: Larva, MADAGASCAR, Antananarivo (= Tananarive)
Prov., Ankeniheny R., 28C, 4 km S of Manjakatompo Forest Station, 1 -XI- 1 97 1 , G.F., C. H.
Edmunds, and F. Emmanuel [mouthparts, forelegs, tergum 4, and paraproct on slide (medium:
Euparal)].

Etymology. The specific epithet is a Latin word meaning "from an is-
land."

Discussion. Demoulinia has been known previously only from South Af-
rica (Demoulin 1 970, Gillies 1 990). Demoulinia insularis is distinguished from
the southern African species D. crassi (Demoulin) by the relatively wide
anteromedial emargination of the labrum (Fig. 11), edentate tarsal claws (Fig.
17), and numerous small spines of the paraproct (Fig. 19).

Vol. 109, No. 5, November & December, 1998

361

The presence of D. insularis in Madagascar is significant because it indi-
cates that Demoulinia was well established in at least West Gondwanaland
before the island began to separate from the African landmass approximately
100 million years ago.

\rwv~www

Figs. 1 1 -19. Demoulinia insularis, larva. 1 1. Labrum (dorsal). 12. Hypopharynx. 13. Left man-
dible. 14. Right mandible. 15. Right maxilla. 16. Labium (left-ventral; right-dorsal). 17. Right
foreleg. 18. Tergum 4 (detail). 19. Paraproct.

ACKNOWLEDGMENTS

We thank G. F. Edmunds, Jr. (Salt Lake City, Utah) for the donation of the material used in
this study.

362 ENTOMOLOGICAL NEWS

LITERATURE CITED

Demoulin, G. 1970. Ephemeroptera des faunes ethiopienne et malgache. S. Afr. Anim. Life 14:

24-170.
Demoulin, G. 1973. Ephemeropteres de Madagascar. III. Bull. Inst. R. Sci. Nat. Belg. 49: 1-

20.
Gillies, M. T. 1990. A revision of the African species of Centroptilum Eaton (Baetidae,

Ephemeroptera). Aq. Insects 12: 97-128.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1996a. The Bugilliesia complex of African Baetidae

(Ephemeroptera). Trans. Am. Entomol. Soc. 122: 175-198.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1996b. Crassabwa: a new genus of small minnow

mayflies (Ephemeroptera: Baetidae) from Africa. Ann. Limnol. 32: 235-240.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1996c. The composition of Dabulamanzia, a new

genus of Afrotropical Baetidae (Ephemeroptera), with descriptions of two new species.

Bull. Soc. Hist. Nat. Toulouse 132: 7-13.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1997a. Edmulmeatus grandis: an extraordinary new

genus and species of Baetidae (Insecta: Ephemeroptera) from Madagascar. Ann. Limnol.

33: 191-195.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1997b. Labiobaetis Novikova & Kluge

(Ephemeroptera: Baetidae) from the Afrotropical region. Afr. Entomol. 5: 241-260.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1 997c. Contribution to the systematics of the genus

Cheleocloeon (Ephemeroptera: Baetidae). Entomol. News 108: 283-289.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1997d. Maliqua: a new genus of Baetidae

(Ephemeroptera) for a species previously assigned to Afroptilum. Entomol. News 108: 367-

371.

Lugo-Ortiz, C. R. and W. P. McCafferty. 1997e. New Afrotropical genus of Baetidae (In-
secta: Ephemeroptera) with bladelike mandibles. Bull. Soc. Hist. Nat. Toulouse 133: 41-

46.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1997f. New species and first reports of the genera

Cheleocloeon, Dabulamanzia, and Mutelocloeon (Insecta: Ephemeroptera: Baetidae) from

Madagascar. Bull. Soc. Hist. Nat. Toulouse 133: 147-53.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1998a. The Centroptiloides complex of Afrotropical

small minnow mayflies (Ephemeroptera: Baetidae). Ann. Entomol. Soc. Am. 91: 1-26.
Lugo-Ortiz, C. R. and W. P. McCafferty. 1998b. Cheleocloeon falcatum (Crass): a new

combination for a southern African species previously assigned to Afroptilum Gillies

(Ephemeroptera: Baetidae). Afr. Entomol., in press.
McCafferty, W. P. 1996. The Ephemeroptera species of North America and index to their

complete nomenclature. Trans. Am. Entomol. Soc. 122: 1-54.
McCafferty, W. P. 1 998. Ephemeroptera and the great American interchange. J. N. Am. Benthol.

Soc., 17: 1-20.
McCafferty, W. P., C. R. Lugo-Ortiz, and H. M. Barber-James. 1997. Micksiops, a new

genus of small minnow mayflies (Ephemeroptera: Baetidae) from Africa. Entomol. News

108: 362-366.

Navas, L. 1926. Algunos insectos del Museo de Paris (3a serie). Broteria Zool. 23: 95-1 15.
Navas, L. 1930. Insectos del Museo de Paris (5a serie). Broteria Zool. 24: 5-24.
Navas, L. 1936. Comunicaciones entomologicas. 19. Insectos de Madagascar. Rev. Acad. Ci.

Exac. Fisquim. Nat. Zaragoza 19: 100-1 10.
Wuillot, J. and M. T. Gillies. 1994. Dicentroptilum, a new genus of mayflies (Baetidae,

Ephemeroptera) from Africa. Aq. Insects 16: 133-140.

Vol. 109, No. 5, November & December, 1998 363

AEDES ALBOPICTUS (DIPTERA: CULICIDAE)

OCCURRENCE THROUGHOUT TENNESSEE,

WITH BIOLOGICAL NOTES 1

James P. Moore^

ABSTRACT: Aedes albopictus, the Asian tiger mosquito, was recorded for the first time in 87 of
Tennessee's 95 counties. Continued occurrence of this species in the remaining 8 Tennessee
counties was confirmed. Notes are provided on other mosquito species sharing larval habitats
with Ae. albopictus, as well as the observed photoperiod- induced egg diapause of Ae. albopictus
in Montgomery County, Tennessee.

A survey of Tennessee's 95 counties was conducted during 1997 to deter-
mine the presence of Aedes albopictus (Skuse), the Asian tiger mosquito. This
survey, primarily of water-containing tire habitats, established the first record
of Ae. albopictus in 87 Tennessee counties and confirmed the continued pres-
ence of the species in the remaining 8 counties with previously reported infes-
tations (Chet Moore, CDC, personal communication). These 8 counties are
Anderson, Coffee, Davidson, Gibson, Henderson, Lawrence, Montgomery, and
Shelby.

Mosquito larvae were collected from 108 sites in 86 counties from July to
October 1997. These sites consisted of new and waste automotive tires located
at roadside dumps (12), county waste collection points (2), and commercial
tire businesses (91). Other sites included outdoor plastic and masonry contain-
ers at residences (2) and roadside dumps (1). Larvae were collected directly
from the tire or other container using a siphon, transported to the laboratory,
and identified using standard light microscopy and the taxonomic references
of Darsie and Ward ( 1 981), Darsie (1986), and Reinertetal. (1997). At 51% of
the larval collection sites, Ae. albopictus was the only culicid species collected
(Table 1 ). Aedes albopictus shared the habitat with other mosquito species at
an additional 31% of the sites, making this species the predominant culicid
resident of tire habitats in Tennessee.

Adult Ae. albopictus were collected from 50 sites in 50 counties during
this survey. An aspirator was used to collect the adults (both sexes) resting at
26 sites and feeding on man at 24 sites. Adult specimens were examined using
a dissecting microscope and identified using the taxonomic references of Darsie
and Ward ( 1 98 1 ) and Darsie ( 1 986). Collection data for each collection site is
preserved and available for anyone wishing to visit the collection sites for
control purposes.

Aedes triseriatus (Say) was found as the sole inhabitant at 5% of the sites.

1 Received February 6, 1998. Accepted April 19. 1998.

2 The Center for Field Biology, Austin Peay State University, Clarksville, TN 37044.

ENT. NEWS 109(5): 363-365, November & December, 1998

364 ENTOMOLOGICAL NEWS

Aedes triseriatus was found with other culicid species at an additional 6% of
the sites; Ae. albopictus was always one of its cohabitants. This indicates that
the Asian tiger mosquito has not totally excluded Ae. triseriatus from tire habi-
tats in Tennessee, as had been predicted by Livdahl and Willey (1991).

No specimens of Aedes aegypti (Linnaeus) were collected during this sur-
vey. However, Ae. aegypti larvae were collected previously by the author from
tire habitats in Montgomery County, TN, during July 1996 and April 1997.
This may indicate that Ae. albopictus is competitively displacing Ae. aegypti
from tire habitats in Tennessee. However, most of Tennessee is in the region
identified by Darsie and Ward (1981) as the extreme range for Ae. aegypti.

Some populations of Ae. albopictus exhibit a photoperiod-induced egg
diapause (Hawley et al. 1989, Estrada-Franco and Craig 1995). As the species
adapts to new locations, the photoperiod response may be altered, providing
researchers with a measure of the length of time the species has inhabited a
region (Chet Moore, CDC, personal communication). There are no historical
data on the photoperiod ism of Ae. albopictus in Tennessee. In Montgomery
County, TN, during 1997, the hatching of Ae. albopictus eggs in outdoor arti-
ficial containers was observed as early as 1 7 April (13.2 hours daylight) and as
late as 2 October (1 1.8 hours daylight).

Table 1. Species composition of larval mosquito collections from Tennessee (July - October
1997), showing percentage of 108 collection sites with a species alone (Solo) and percentage of
collection sites with cohabitant culicid species (With others).

Percentage of 108 larval collection sites

Mosquito species Solo With others Total %

Aedes albopictus (Skuse) 51 31 82

Aedes atropalpus (Coquillett) 2 24

Aedes triseriatus (Say) 5 6 11

Anopheles punctipennis (Say) / perplexens Ludlow 1 1

Anopheles quadrimaculatus sensu stricto Say 1 1

Culex pipiens Linnaeus/ quinquefasciatus Say 6 11 17

Culex restuans Theobald 5 7 12

Culex territans Walker 22

Toxorhynchites rutilus septentrionalis (Dyar & Knab) 3 47

Vol. 109, No. 5, November & December, 1998 365

ACKNOWLEDGMENTS

Grateful appreciation is extended to The Center for Field Biology and the Department of
Biology, Austin Peay State University, Clarksville, TN, for their support and laboratory facili-
ties, and to Chet Moore, Division of Vector-Borne Infectious Diseases, National Center for
Infectious Diseases, Centers for Disease Control and Prevention, Fort Collins, CO, for his ad-
vice and encouragement; and to Steve Hamilton, Department of Biology, Austin Peay State
University, Clarksville, TN, and C. Steven Murphree, Department of Biology, Belmont Univer-
sity, Nashville, TN, for providing helpful suggestions with the manuscript.

LITERATURE CITED

Darsie, R.F., Jr. 1986. The identification of Aedes albopicius in the Nearctic Region. J. Am.
Mosq. Control Assoc. 2(3): 336-40.

Darsie, R.F., Jr., and R.A. Ward. 1981. Identification and geographical distribution of the
mosquitoes of North America, north of Mexico. Mosq. Syst. Suppl. 1, 313 pp.

Estrada-Franco, J.G., and G.B. Craig, Jr. 1995. Biology, disease relationships, and control
of Aedes albopictus; Pan Amer. Health Org. Tech. Paper No. 42. Wash., DC, 49 pp.

Hawley, W.A., C.B. Pumpuni, R.H. Brady, and G.B. Craig, Jr. 1989. Overwintering survival
of Aedes albopictus (Diptera: Culicidae) eggs in Indiana. J. Med. Entomol. 26(2): 122-9.

Livdahl, T.P., and M.S. Willey. 1991. Prospects for an invasion: competition between Aedes
albopictus and native Aedes triseriatus. Science 253: 189-91.

Reinert, J.F., P.E. Kaiser, and J.A. Seawright. 1997. Analysis of the Anopheles (Anopheles)
quadrimaculatus complex of sibling species (Diptera: Culicidae) using morphological, cy-
tological, molecular, genetic, biochemical, and ecological techniques in an integrated ap-
proach. J. Am Mosq. Control Assoc. 13(Suppl.): 1-102.

366 ENTOMOLOGICAL NEWS

DISTRIBUTION AND HABITAT OF CURICTA

PRONOTATA (HEMIPTERA: NEPIDAE)

IN SOUTHEASTERN ARIZONA 1

Jon D. Hoekstra 2 , Robert L. Smith 3

ABSTRACT: Curicta pronotata is known from western Mexico and southeastern Arizona. Pre-
vious accounts of the species' Arizona distribution draw from only a few collections and locali-
ties. Distributional records clarifying the range and habitat of C. pronotata in Arizona are re-
ported here.

The genus Curicta (Hemiptera: Nepidae) is primarily Neotropical with spe-
cies entering the United States in Arizona, Texas, and Louisiana (Menke, 1 979).
Curicta pronotata 's principal range is in Mexico along the Sierra Madre Occi-
dental; it extends from Nayarit north to eastern Sonora and western Chihuahua
(Keffer 1996). C. pronotata has previously been reported from two different
canyons in the Huachuca Mts. in southern Arizona (Dubois 1 978, Keffer 1 996),
and Sites and Polhemus (1994) reported the species from Sabino Canyon near
Tucson on the basis of an individual collected in 1937 (full record given here).

This report extends the range of the species in Arizona. Curicta pronotata
has now been recorded from the Huachuca, Santa Catalina, and Galiuro moun-
tain ranges in southern Arizona. Based on our own findings and habitat infor-
mation from previous records, the habitat of C. pronotata in Arizona appears
to be small shaded streams with woody debris above about 1200m.

NEW RECORDS
Hemiptera: Nepidae: Curicta pronotata Kuitert 1949.

Arizona: Pima County: 1 male, Santa Catalina Mts., Sabino Canyon, 29- VII- 1937, E.D. Ball,
UAIC; Pima County: 1 nymph, Santa Catalina Mts., Sabino Canyon, 3660' (11 15 m), 3222'00"
N, 11047'10" W, 6-VI-1997, J.D. Hoekstra, UAIC; Pima County: Santa Catalina Mts., Bear
Canyon, 5530' (1685 m), 322r45" N, 11042'30" W, 19- VII- 1997, J.D. Hoekstra and C.
Creighton, 1 male, 1 female, 12 nymphs, UAIC, 1 female, 5 nymphs, JTPC; Cochise County: 1
female, Galiuro Mts., Wildcat Canyon, 4120' (1255 m), 3222'00" N, 11015'30" W, 20-VIII-
1996, J.D. Hoekstra and D.A. Lytle, JTPC; Cochise County, 1 male, same locality data as
preceding, 21 -IX- 1996, J.D. Hoekstra, JTPC.

Abbreviations: JTPC = J. T. Polhemus Collection, Englewood, CO. UAIC = University of Ari-
zona Insect Collection, Tucson, AZ.

DISCUSSION
In Sabino Canyon, a single nymph was collected from mud under a rock

1 Received December 29, 1997. Accepted February 8, 1998.

L Center for Aquatic Ecology, Illinois Natural History Survey, 607 E. Peabody Dr., Champaign,

IL61820.
-* Department of Entomology, University of Arizona, Tucson, AZ 85721.

ENT. NEWS 109(5): 366-368, November & December, 1998

Vol. 109, No. 5, November & December, 1998 367

along the margin of a drying pool. The nymph was inactive and presumed
dead until it was placed in ethanol, whereupon it revived for a short time. The
nymph's behavior may have been an example of "death-feigning" (thanatosis)
as reported in Nepa and Ranatra, or it may have indicated the initiation of
estivation, which has been documented in Nepa (Sites and Polhemus 1994).
This nymph is the first evidence of C. pronotata in Sabino Canyon since it was
collected in 1937. The authors have sampled the habitat in which it was found
extensively over two seasons and have found only this single nymph. We sus-
pect that it was a "stray" which had drifted down from an upstream source
population in Sabino Canyon or a tributary.

In Bear Canyon, Santa Catalina Mts., adults and especially nymphs were
very abundant among woody detritus in shallow bedrock-lined pools. The stream
was narrow (less than 1 m wide) and shallow (average maximum depth
30 - 40 cm). Curicta pronotata shared this habitat with Abedus herberti (Hemi-
ptera: Belostomatidae), which was also abundant. Of the twenty C. pronotata
collected on July 19, 3 were adults, 10 were F-l instar nymphs, and 7 were F-
2 instar nymphs. In Curicta scorpio, the fourth and fifth stadia lasted an aver-
age of 18.56 and 18.87 days for artificially reared individuals from two popu-
lations in Texas (Keffer et al. 1 994). If C. pronotata has a similar developmen-
tal rate, most of the individuals in the Bear Canyon population probably eclosed
to the adult stage by September.

Arizona collections of Curicta pronotata have been very few, despite the
species' fairly wide range as indicated by the records reported here. The spe-
cies probably has been overlooked because of its cryptic appearance and ten-
dency to feign death when captured (Sites and Polhemus 1994). In addition,
spatial and temporal components of the species' occurrence in Arizona may
have contributed to its rarity in collections.

Arizona populations of C. pronotata appear to be highly localized. Such
local populations could be relicts of a previously continuous distribution. The
Sonoran Desert region has aridified over the past 11,000 yr., with attendant
restriction of previously widespread mesic biotic communities to high eleva-
tions (Hall et al., 1989). This process may have reduced the number and ex-
tent of suitable habitats for C. pronotata, such that it currently persists in only
a few favorable habitat refuges.

Densities of C. pronotata probably fluctuate seasonally with changes in
streamflow, as noted by Keffer ( 1 996) for Texas populations of Curicta scorpio.
Local populations of C. pronotata may also be unstable from year to year in
Arizona. The species is probably capable of dispersal by flight, which has
been reported for C. scorpio (Sites and Polhemus 1994). Thus the disjunct
Arizona populations could be transient "sink" populations in a metapopulation
with a "core" to the south in the Sierra Madre Occidental. Such a metapopulation
dynamic has been reported for several southeastern Arizona butterfly species

368 ENTOMOLOGICAL NEWS

(Bailowitz and Brock 1991).

Additional surveys and long term monitoring of the Arizona populations
will be required to evaluate these alternative hypotheses about the population
dynamics and biogeography of C. pronotata at the northern limits of its range.

ACKNOWLEDGMENTS

We thank J.T. Polhemus for making the specific determinations on individuals from the
populations reported. J.D. Hoekstra also thanks the UA Department of Entomology and the
ARCS (Acheivement Rewards for College Scientists) Foundation for support during the time
in which this research was conducted.

LITERATURE CITED

Bailowitz, R.A. and J.P. Brock. 1991. Butterflies of Southeastern Arizona. Sonoran Arthropod

Studies, Inc., Tucson, Arizona.
DuBois, Mark B. 1978. A recent record of Curicla pronotata Kuitert (Hemiptera:Nepidae) from

its type locality. Entomol. News 89: 207.
Hall, W.E., C.A. Olson, and T.R. VanDevender. 1 989. Late Quaternary and Modern Arthropods

from the Ajo Mountains of southwestern Arizona. Pan-Pacific Ent. 65: 322-347.
Keffer, S.L. 1996. Systematics of the New World waterscorpion genus Curicla Stal (Heteroptera:
Nepidae). J. New York Entomol. Soc. 104(3-4): 117-215.
Keffer, S.L. 1991. Taxonomic revision of the neotropical genus Curicta Stal (Insecta:

Heteroptera: Nepidae). Ph.D. dissertation, S. I. U., Carbondale.
Keffer, S.L., SJ. Taylor and J.E. McPherson. 1994. Laboratory rearing and descriptions of

immature stages of Curicta scorpio (Heteroptera: Nepidae). Ann. Entomol. Soc. Am. 87( 1 ):

17-26.
Kuitert, Louis C. 1949. Some new species of Nepidae (Hemiptera). J. Kansas Entomol. Soc. 22:

60 - 68.
Menke, A.S. 1979. Family Nepidae. Pp. 70-75 in A.S. Menke [ed.],The semiaquatic and aquatic

Hemiptera of California (Heteroptera: Hemiptera). Bull. Calif. Ins. Surv. 21: 1-166.
Polhemus, J.T. 1976. Notes on the North American Nepidae. Pan-Pacific Ent. 52: 204-208.
Sites, R.W. and J.T. Polhemus. 1994. Nepidae (Hemiptera) of the United States and Canada.

Ann. Entomol. Soc. Am. 87( 1 ): 27-42.

Vol. 109, No. 5, November & December, 1998 369

HEXAMETH YLDISILAZANE - A CHEMICAL
ALTERNATIVE FOR DRYING INSECTS 1

John Heraty, David Hawks^

ABSTRACT: Two methods of chemically drying softbodied Chalcidoidea (Hymenoptera) are
compared: critical-point drying (CPD) and hexamethyldisilazane (HMDS). For three groups of
Eulophidae, Encyrtidae and miscellaneous Chalcidoidea, the CPD specimens were of consis-
tently higher quality for all groups, although the overall differences between CPD and HMDS
specimens were marginal.

Soft-bodied insect specimens have long been the bane of systematics.
Freshly killed and air-dried specimens (Fig. 1 ) undergo partial to complete
collapse of body parts, whereas specimens initially preserved in EtOH fare
even worse when subsequently removed from the liquid and air dried (Fig. 2).
This is not only a problem of obtaining quality museum specimens but in the
past has deterred some systematists from bothering with samples preserved in
alcohol, such as those taken in malaise or pan traps. Critical-point drying (CPD)
of specimens through a liquid CO 2 intermediate (Gordh & Hall 1979) pro-
vides a means of retrieving large numbers of soft-bodied specimens from EtOH
and is being widely used for some taxa, especially Chalcidoidea. The primary
advantage of using CPD is little or no collapse of soft body parts, including
internal muscles and nerves. Secondarily, the structure of muscles, nerve tis-
sue and other internal body parts is maintained, allowing for later survey of
these structures from museum specimens (Heraty et al. 1997). The disadvan-
tages with the CPD are that it 1 ) is relatively expensive to buy the initial equip-
ment ($2,000-8,000), 2) is necessary to obtain specialized CO 2 tanks that must
be maintained above 900 psi, 3) is labor intensive, 4) can cause abnormal
swelling or occasional bursting of some body parts, and 5) may leave surface
residues on specimens.

Several alternatives to air drying or CPD have been proposed, some of
which are freeze drying, Peldri II (Brown 1990), acetone vapor (van Noort
1995), xylene (R. Carlson pers. comm.), and hexane (D. Hawks, pers. comm.).
A new chemical method involving hexamethyldisilazane (HMDS) has been
proposed as a simple and cost-effective means of retrieving high-quality speci-
mens from collections preserved in EtOH (Nation 1983, Brown 1993). Only
the CPD and HMDS methods are regularly applied for the retrieval of large
collections of Chalcidoidea initially preserved in alcohol, and here we com-
pare the two methods.

METHODS
All specimens were initially killed and preserved in 70-75% EtOH at 4C.

1 Received October 22, 1997. Accepted March 31, 1998.

2 Department of Entomology, University of California, Riverside, CA 92521.

ENT. NEWS 109(5): 369-374, November & December, 1998

370 ENTOMOLOGICAL NEWS

Evaluations were of separate collections made from 1990 to 1996 in southeast
Asia, the Galapagos Islands and California. Lots that had a high proportion of
soft-bodied Chalcidoidea were chosen, and all specimens were scored from
each lot. Fourteen separate collections (362 specimens) were evaluated for the
CPD method and 5 collections (347 specimens) for HMDS. Overlap in collec-
tion time and country for each method occurred only for the southeast Asian
collections. Additional specimens of a new species ofdrrospilus (Eulophidae)
from California were examined as representatives of very soft-bodied Chal-
cidoidea.

Specimens were scored on a scale of 1 to 5, with 5 being a nearly perfect
specimen suitable for scanning electron microscopy (SEM). Scores were based
only on the softer body parts. Cirrospilus are almost entirely soft-bodied and
represent an extreme; in other taxa, for example pteromalids, the head and
mesosoma are well-sclerotized and do not collapse under any treatment, but
the antennae and gaster will partially or completely collapse. A score of 1
would be typical of air-dried eulophids taken from alcohol: completely shriv-
elled and collapsed (Fig. 2). A score of 2 was assigned to specimens that had
extensive collapse of the softer body structures (head, antennae and gaster)
(Fig. 1 ). A score of 3 was given to specimens with partial collapse of all softer
body parts (Figs 4, 5). Freshly killed and air-dried specimens would usually be
given a score between 1 and 3, with a score of 3 bordering on acceptable for
museum collections or SEM (at least partly shrivelled or collapsed). A score of
4 was given for very minimal collapse of not more than one body part or a
slight distortion (wrinkling or bloating) of the gaster (Figs 4, 5). The Cirrospilus
were not scored for comparative analysis. All material is deposited in the En-
tomology Research Museum, University of California, Riverside.

CPD method. The liquid vapor interface is the primary destructive force
in air-drying specimens, and if not about equal, results in the breakdown of
cell walls and collapse of tissue. For CO 2 the identical vapor pressure as a
liquid or gas, the critical point, is reached at 3 1 .0C and 1093 psi (Burstyn and
Bartlett 1975). Specimens were dried as outlined by Gordh & Hall (1979) by
1) dehydrating the specimens to 100% EtOH, 2) exchanging fluids through
liquid CO 2 under high pressure (900 psi) and low temperature (11-15C)
through a series of soaks and purges until the exhausted dry CO 2 did not leave
a liquid residue (complete sublimation), 3) drying under high pressure (1 100-
1200 psi) until the chamber temperature reached 41-43C, and then 4) slowly
exhausting the gaseous CO 2 to room atmospheric pressure.

HMDS method. HMDS ([(CH 3 ) 3 Si] 2 NH) reacts with water to produce
hexamethyldisiloxane ([(CH 3 ) 3 Si] 2 O) and ammonia (NH 3 ), both of which
evaporate from the specimen (Dave Jordon, Polysciences Inc., pers. comm.).
Specimens were dried in the manner outlined by Brown (1993) by 1) dehy-
drating the specimens to 100% EtOH, 2) replacing the alcohol with HMDS for
two soaks of 1/2 hour each in a covered glass vial or dish, and 3) after the

Vol. 109, No. 5, November & December, 1998

371

second soak, pouring off most of the HMDS and allowing the remaining HMDS
to evaporate in a fume hood (or outdoors in a well-ventilated area). Samples
can be soaked and dried in glass vials or dishes. Gas buildup in the vials may
cause the release of liquid HMDS while being uncapped, but this can be avoided
by using smaller volumes of HMDS (less than half of vial) or slowly unscrew-
ing the vial top. We prefer to line the bottom of a glass dish with a fine brass
screen and cover each sample with a screen lid during evaporation, thus pre-
venting dried specimens from disappearing into the exhaust. HMDS is a skin
irritant, and gloves and eye protection are recommended.

All specimens were card-mounted for examination following Noyes ( 1 982).

RESULTS

Cirrospilus (Eulophidae) was used as an example of a very soft-bodied
species that does not fare well under any of the drying methods (Figs 1 -6). Air-
drying (Fig. 1) resulted in collapse of the antennae, femora, and gaster dor-

Figures 1-6. Cirrospilus sp. (Eulophidae): 1, freezer killed and air dried. 2-6, killed and pre-
served in 70% EtOH and then: 2, air-dried; 3&5, CPD dried; 4&6, HMDS dried.

372

ENTOMOLOGICAL NEWS

sally and laterally; the mesosoma was relatively undistorted. The specimen
illustrated would receive a score of 2, which would be marginally acceptable
for use in collections. Air-drying from alcohol (Fig. 2) was disastrous, with
general collapse of all body parts (score 1). CPD Cirrospilus (Figs 3, 5) showed
slight collapse of the scape and scrobes, and distortion but not collapse of the
gastral tergites. Such a specimen (Fig. 3) would be scored as a 4 (less than
perfect). HMDS Cirrospilus (Fig. 4, 6) exhibited a greater degree of collapse
of the scape, head and metasoma, with the specimen receiving a score of 3.
For extremely soft-bodied specimens, the CPD method was consistently bet-
ter than the HMDS method, and both were better than air-drying.

Seven families of Chalcidoidea were encountered in the 19 collections
evaluated (Table 1). Each family presents a different problem with respect to
how they were affected by improper drying. Even when CPD- or HMDS-
treated, soft-bodied Eulophidae generally had some collapse or distortion of
all body parts (cf. Figs. 3-5). Using either CPD or HMDS, 51% of the

Table 1. Quality of soft-bodied Chalcidoidea dried using critical-point drying (CPD) or
hexamethyldisilazane (HMDS). Ranking based on a scale of 1 -5, with 5 indicating a near-perfect
specimen. Mean values were significantly higher for CPD specimens for all groups (Chi-square,
P=0.01). Data were pooled for Aphelinidae, Mymaridae, Pteromalidae, Torymidae and
Trichogrammatidae.

rank

1 n mean rank

Eulophidae

CPD

126

114

247 4.48

HMDS

169 4.38

Encyrtidae

CPD

71 4.56

HMDS

61 4.48

Aphelinidae

CPD

"""

44 4.82

HMDS

117 4.51

Pteromalidae

CPD

HMDS

Mymaridae

CPD

HMDS

Trichogrammatidae

CPD

HMDS

Torymidae

CPD

HMDS

Vol. 109, No. 5, November & December, 1998 373

Eulophidae treated had a score of 5, and, although a much higher proportion
than the CPD method, only 13.0% of the specimens received a score of 3, and
none received a 1 or 2. Many Eulophidae are reasonably well-sclerotized and
do not have problems similar to those of Cirrospilus. Often the most notice-
able artifact was a slight wrinkling of the gastral tergites (score of 4), which
was common in both treatments. Pteromalidae generally have a well-sclero-
tized head and mesosoma, but the gaster of males is particularly susceptible to
collapse. Both Trichogrammatidae and Aphelinidae are soft-bodied but re-
sponded well to either technique except for some collapse of the antennae,
which occurred with use of either method. Other than Eulophidae, all of the
chalcidoid groups responded well to either technique, with consistent scores
of 4 or 5, both of which are acceptable for museum collections.

For statistical comparisons, Eulophidae and Encyrtidae were common in
all samples and were treated separately; results for Aphelinidae, Pteromalidae,
Mymaridae, Torymidae and Trichogrammatidae were pooled. In all three com-
parisons, the CPD specimens were of significantly higher quality (rank) than
the HMDS specimens (Chi Square, P=0.01), although the differences in the
mean rank scores for each treatment were marginal (Table 1 ). The CPD method
after ethanol fixation also ranked better than HMDS in a study of pre- and
post-fixation techniques in four taxa (Swearingen et al. 1997). In contrast to
the techniques used by Swearingen et al. (1997), we have not found fixation in
osmium tetroxide to be a necessary step in preparation for either museum or
SEM specimens.

CPD and HMDS methods left little or no residue on the specimens, as
noted by Swearingen et al. (1997). Specimens treated by HMDS appeared to
be slightly cleaner, but we could see no way to quantify this characteristic
accurately. HMDS also works as a good degreasing agent for some insects
such as tiger beetles and robber flies. We also found various labels and ink
types (including laser-printed labels) to be unaffected by HMDS, allowing
their inclusion during processing. The same is possible for the CPD method,
although processing is usually in small capsules making inclusion of larger
labels impossible. At $30 U.S. per 400 ml of HMDS and 5 ml per large lot of
about 100 chalcidoids, we estimate a cost of about 37.5 cents per run, or 0.4
cents per specimen. We have tried HMDS on a variety of insects, including
Collembola, flies, beetles and other Hymenoptera (Perdita and Bombus), with
generally excellent results. Heavily sclerotized individuals processed using
HMDS are as good as CPD specimens. Internally, muscles and nerve tissue
are preserved in the same manner as using the CPD process. For larger speci-
mens, wings are often crumpled in smaller CPD capsules, but this was not a
factor with HMDS. In addition to improved specimen quality, it is also note-
worthy that mitochondrial DNA was successfully extracted from dried CPD
and HMDS specimens of Ichneumonidae and Encyrtidae (Austin & Dillon
1997).

374 ENTOMOLOGICAL NEWS

In summary, the use of HMDS is a viable alternative to use of CPD for
retrieving soft-bodied insects from alcohol. CPD specimens are marginally
better in quality than those treated with HMDS, but HMDS is cost-effective
and less labor intensive than CPD. If the equipment is not available, HMDS
may be the preferred technique.

ACKNOWLED

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