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(ISSN 0015-4040)
FLORIDA ENTOMOLOGIST
(An International Journal for the Americas)
Volume 73, No. 4 December, 1990
TABLE OF CONTENTS
Announcement 74th Annual Meeting ..................................... .............. i
SYMPOSIUM: ORIGINS OF ARTHROPOD FAUNA OF THE
SOUTHEASTERN UNITED STATES
Preface ......................... ........ .. ... .. .. ... ..... ..... .................. 527
DEYRUP, M.-Arthropod Footprints in the Sands of Time ............................ 529
ALLEN, R. T.-Insect Endemism in the Interior Highlands of North America 539
CARLTON, C. E.-Biogeographic Affinities of Pselaphid Beetles of the Eastern
U united States ................................................................................. 570
STOETZEL, M. B.-Some Aphids of Importance to the Southeastern United States 580
HAMILTON, S. W., AND J. C. MORSE-Southeastern Caddisfly Fauna: Origins
and Affinities ................................................................................ 587
Research Reports
HULTING, F. L, D. B. ORR, AND J. J. OBRYCKI-A Computer Program for
Calculation and Statistical Comparison of Intrinsic Rates of Increase and
Associated Life Table Parameters .................................................... 601
MATTHEWS, D. L., D. H. HABECK, AND D. W. HALL-Annotated Checklist of
the Pterophoridae (Lepidoptera) of Florida Including Larval Food Plant
R records ......................................................................................... 613
SCHEFFRAN, R. H., N.-Y. Su, AND B. DIEHL-Native, Introduced, and Struc-
ture-Infesting Termites of the Turks and Caicos Islands, B. W. I. (Isoptera:
Kalotermitidae, Rhinotermitidae, Termitidae) .................................. 622
STOETZEL, M. B., AND D. J. HILBURN-The Aphids and Phylloxera of Bermuda
(Homoptera: Aphididae and Phylloxeridae) ....................................... 627
BARNES, J. K.-First Record of Genus Phymatopterella in the Nearctic Region
and Description of P. ovatimacula, A New Humpbacked Fly from Florida
(Diptera: Phoridae) ........................................................................ 644
WIRTH, W. W.-Biting Midges of the Subgenus Schizoforcipomyia of Forcipomyia
in North America (Diptera: Ceratopogonidae) ................................... 649
VEGA, F. E., P. BARBOSA, AND A. P. PANDURO-An Adjustable Water-Pan
Trap for Simultaneous Sampling of Insects at Different Heights ............ 656
SHARP, J. L.-Mortality of Caribbean Fruit Fly Immatures in Shrinkwrapped
Grapefruit .................................................................................... 660
SIMMONS, A. M., AND R. E. LYNCH-Egg Production and Adult Longevity of
Spodoptera frugiperda, Helicoverpa zea (Lepidoptera: Noctuidae), and
Elasmopalpus lignosellus (Lepidoptera: Pyralidae) on Selected Adult Diets 665
Scientific Notes
BARROWS, E. M.-Sex Ratio, Parasitism, and Hosts of an Everglades
Population of Prochalia pygmea (Lepidoptera: Psychidae) .......... 672
Continued on Back Cover
Published by The Florida Entomological Society
FLORIDA ENTOMOLOGICAL SOCIETY
OFFICERS FOR 1989-90
President .............................................. ................ J. F. Price
President-Elect ........................ ........ ... ...................... J. L. Knapp
Vice-President ....................................................... D. F. W illiams
Secretary ....................... ... ............... ..... J. A. Coffelt
Treasurer ........................................................... .................... A. C. Knapp
Other Members of the Executive Committee
J. E. Eger J. E. Pefia J. R. Cassani J. Hogsette
J. R. McLaughlin S. Valles M. Lara
PUBLICATIONS COMMITTEE
J. R. McLaughlin, USDA/ARS, Gainesville, FL ....................................... Editor
Associate Editors
Agricultural, Extension, & Regulatory Entomology
Ronald H. Cherry-Everglades Research & Education Center, Belle Glade, FL
Michael G. Waldvogel-North Carolina State University, Raleigh, NC
Apiculture
Stephen B. Bambara-North Carolina State University, Releigh, NC
Biological Control & Pathology
Ronald M. Weseloh-Connecticut Agricultural Experiment Sta., New Haven, CT
Book Reviews
J. Howard Frank-University of Florida, Gainesville
Chemical Ecology, Physiology, Biochemistry
Louis B. Bjostad-Colorado State University, Fort Collins, CO
Ecology & Behavior
John H. Brower-Stored Product Insects Research Laboratory, Savannah GA
Theodore E. Burk-Dept. of Biology, Creighton University, Omaha, NE
Forum & Symposia
Carl S. Barfield-University of Florida, Gainesville
Genetics & Molecular Biology
Sudhir K. Narang-Bioscience Research Laboratory, Fargo, ND
Medical & Veterinary Entomology
Arshad Ali-Central Florida Research & Education Center, Sanford, FL
Resumen
Omelio Sosa, Jr.-USDA Sugar Cane Laboratory, Canal Point, FL
Systematics, Morphology, and Evolution
Michael D. Hubbard-Florida A&M University, Tallahassee
Howard V. Weems, Jr.-Florida State Collection of Arthropods, Gainesville
Willis W. Wirth-Florida State Collection of Arthropods
Business Manager ...................... ...... ..................... A. C. Knapp
FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September, and De-
cember. Subscription price to non-members is $30 per year in advance, $7.50 per copy;
institutional rate is $30 per year. Membership in the Florida Entomological Society,
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Please consult "Instructions to Authors" on the inside back cover.
This issue mailed December 14, 1990
ANNOUNCEMENT OF THE 74TH ANNUAL MEETING
FLORIDA ENTOMOLOGICAL SOCIETY
The 74th annual meeting of the Florida Entomological Society will be held August
4-7, 1991 at the Ritz Carlton Hotel, 280 Vanderbilt Beach Road, Naples, Florida 33963;
telephone (813 598-3300. Registration forms and information will be mailed to members
and will appear in the Newsletter and the March, 1991 issue of Florida Entomologist.
SUBMISSION OF PAPERS
The deadline for submission of papers and posters for the 74th annual meeting of
the Florida Entomological Society will be MAY 15, 1991. The meeting format will be
similar to those in the past with eight minutes allotted for presentation of oral papers
(with 2 minutes for discussion) and separate sessions for members who elect to present
a Poster Exhibit. There will be student paper and poster sessions with awards as in
previous years. Students participating in the judged sessions must be members of the
Florida Entomological Society and registered for the meeting.
David F. Williams, Chairman
Program Committee, FES
USDA-ARS
Medical and Veterinary Entomology Research Laboratory
Gainesville, Florida 32604
(904) 374-5982 or 374-5903
Symposium 527
SYMPOSIUM:
ORIGINS OF THE SOUTHEASTERN ARTHROPOD FAUNA
S. W. HAMILTON AND J. C. MORSE, CONVENERS
A large part of understanding almost any biological phenomenon is discovery of its
historical context. Consequently, we were led to ask from where and when did the
arthropod fauna of our area, the southeastern United States, originate.
For the purposes of organizing this symposium, we arbitrarily define the Southeast
politically to include the states of the Southeastern Branch of the Entomological Society
of America, at whose 1990 annual meeting in Orlando the following papers were pre-
sented. These states include Alabama, Arkansas, Florida, Georgia, Louisiana, Missis-
sippi, North Carolina, South Carolina, and Tennessee. All participants of this symposium
recognize that these boundaries are artificial in terms of the geological and biological
history of this region and did not feel strictly confined to these political boundaries for
their presentations.
Within this "Southeast," one finds a wide range of terrestrial and aquatic habitats,
from sandy hurricane-prone keys, barrier islands, and beaches to lowland swamps and
sandy plains to rolling clay hills to granite and limestone mountains. Biologically, there
Fig. 1. Areas of endemism in southeastern North America.
528 Florida Entomologist 73(4) December, 1990
appear to be at least three major regions that include large numbers of species of plants
and animals peculiar to them. These so-called "areas of endemism" include the southern
end of the Eastern Highlands (EH; including the Southern Appalachian Mountains and
the Cumberland Mountains and Plateau), the Interior Highlands (IH; including the
Ozark Plateau and the Ouachita Mountains), and the Southeastern Coastal Plain (CP)
of the Atlantic and Gulf of Mexico Coasts (Fig. 1). Sandhills and Piedmont Regions are
intermediate among these three Regions in position and degree of relief and typically
are habitat for an intergrading mix of the species found in them.
The objective of this symposium has been to explore this diverse terrain with ar-
thropod examples in order to suggest general trends for the origins of all of the south-
eastern biota. By coming to an understanding of these trends, it is hoped that we will
be better equipped to regulate the potential introductions of modern pest species, to
introduce beneficial species from the most appropriate portions of the globe, and to
appreciate the unique and irreplaceable species that have evolved in situ on our corner
of the continent.
Deyrup: Symposium-Origins SE Arthropod Fauna
ARTHROPOD FOOTPRINTS IN THE SANDS OF TIME
MARK DEYRUP
Archbold Biological Station
P.O. Box 2057
Lake Placid, FL 33852
ABSTRACT
Arthropod distribution in Florida's sandy uplands shows some obvious patterns.
Arthropods endemic to Florida scrub are concentrated on the Lake Wales Ridge, appar-
ently a long-term refuge. Scrub areas to the north apparently lost diversity, as some
species that must have migrated through there from southwestern North America are
now only in southern scrubs. The more recently formed Coastal Ridge was apparently
colonized by the remnant fauna of north Florida, not by Lake Wales Ridge species.
North Florida scrubs have few known endemics, but there are probably many endemics
in sandhill habitats. Expansion and contraction of sandhill areas may explain some
faunal patterns. Rapid destruction of Florida's upland habitats demands prompt re-
search.
RESUME
La distribuci6n de artr6podos de las tirras altas arnesas de la Florida demuestran
ciertos evidentes patrons. Artr6podos end4micos en malezas en la Florida, estAn con-
centrados en Lake Wales Ridge, aparentemente un refugio de largo plazo. Areas de
maleza hacia el norte aparentemente han perdido diversidad, puesto que algunas es-
pecies deben de haber migrado por ahi del suroeste de Norte Am6rica, y las cuales se
encuentran ahora en malezas surefias. El Coastal Ridge que se form recientemente fue
aparentemente colonizado por la fauna remanente del norte de la Florida, y no por
species del Lake Wales Ridge. Malezas del norte de la Florida conoce poca species
end6micas, pero posiblemente hay muchas endemicas en habitaciones altas arenosas.
La expansion y contracci6n de Areas altas arenosas pudieran explicar algunos patrons
faunales. La rapida destruction de habitaci6n de tierras altas en la Florida demand
una pronta investigaci6n.
The sandy uplands of Florida harbor a large number of habitat-specific arthropods,
many of which occur within a small region of Florida, and nowhere else (Fig. 1). The
distribution of these species among the patches of Florida upland habitat should provide
"footprints" that allow us to track the march of geologic and climatic events through
the state. There is no group of organisms better suited to this goal than the marvelously
speciose arthropods, if only we knew enough about them. Unfortunately our knowledge
of the distribution and taxonomy of Florida's upland arthropods remains fragmentary
at best, even though the attention of entomologists was early directed toward these
arthropods by the late Theodore Hubbell (1932, 1961). Readers may be appalled at the
burdens of biogeographic speculation that I have piled onto a few struggling beetles,
grasshoppers and spiders. In some cases, I can ease this burden by spreading it among
plants and vertebrates.
The sandy uplands of present-day Florida are usually divided into the two categories
of Florida scrub and sandhill, the latter also called high pine. Florida scrub is charac-
terized by a dense to sparse cover of low, evergreen sclerophyllous oaks, often with
sand pine (Pinus clausa) or Florida roemary (Ceratiola ericoides). Sandhill has a dense
to sparse herbaceous groundcover, with scattered pines (Pinus palustris), oaks (Quer-
Florida Entomologist 73(4)
December, 1990
1 2 3 4 5 6 7 8 9
Floridobolus penned Causey +
Latrodectus bishop Kaston + + + +
Phidippus xeros Edwards + + + +
Zelotes ocala Platnick & Shadab + +
Zelotes florodes Platnick & Shadab +
Geolycosa xera McCrone + + + +
Lycosa ceratiola Gertsch & Wallace + + +
Lycosa ericeticolu Wallace +
Sosippus placidus Brady +
Arenivaga floridensis Caudell + + + +
Schistocerca ceratiola Hubbell & Walker + + + + +
Melanoplus insignis Hubbell +
Melanoplus tequestae Hubbell + +
Melanoplus puer (Scudder) + + + + + +
Melanoplus forcipatus Hubbell + +
Nannocois arenaria Blatchley + +
Cicindela highlandensis Choate +
Cicindela scabrosa Schaupp + + + +
Serica frosti Dawson +
Anomala eximia Potts +
Phyllophaga elizoria Saylor + +
Phyllophaga panorpa Sanderson +
Phyllophaga elongata (Linell) + + + + +
Phyllophaga okeechobeea Robinson +
Phyllophaga skelleyi Woodruff & Beck + +
Trigonopellastes floridana (Casey) + + +
Mycotrupes pedester Howden +
Peltotnrpes young Howden +
Onthophagus aciculatulus Blatchley + +
Ataenius saramanri Cartwright + + +
Psammodius n. sp. + +
Mecynotarsus sp. +
Pleotomodes needhami (Green) +
Lucidota luteicollis LeConte + + +
Nemomydas melanopogon Steyskal + + +
Nemomydas lara Steyskal +
Nemouria outina Ferguson + + +
Dasymutilla archboldi Schmidt & Mickel + +
Photomorphus archboldi Manley & Deyrup + +
Conomymna elegans Trager +
Conomynna flavopectus (M.R. Smith) + +
Odontomachus clara Roger +
Fig. 1. List and known ranges of arthropods confined to upland ridges. Numbers
refer to Fig. 3, except 9: small ridges in southwest Florida.
cus spp., esp. Q. laevis), and shrubs, but no dense understory or overstory. The struc-
ture of both habitats is maintained in part by fire, and exclusion of fire may lead to a
dense hardwood canopy. For more detailed descriptions of these habitats, their vari-
ants, and their ecology see Abrahamson 1984, Abrahamson et al. 1984, Myers 1985,
Myers & White 1987.
Many of the species of arthropods restricted to Florida scrub and sandhill have
cryptic habits, and are seldom recognized or collected except by specialists. The most
Deyrup: Symposium-Origins SE Arthropod Fauna 531
fundamental problem of this study, therefore, is obtaining satisfactory distribution re-
cords. The highly restricted or disjunct distribution records characteristic of specialized
residents of sandy uplands can be mimicked by inadequate collection records. It is
worthwhile to present a case study of a species to show the kinds of judgment involved.
There is no doubt that the cockroach Arenivaga floridensis Caudell (Polyphagidae)
(Fig. 2) is endemic to scrub and sandhill. It is the sole eastern member of a genus that
includes several western species, all of which inhabit semiarid sandy areas (Blatchley
1920). In all species the female is wingless, and one might expect their distribution to
be restricted by geographic barriers as well as by habitat requirements. Arenivaga
floridensis is known from sites in the central part of the state and along the west coast
(Fig. 2). It seems unlikely that this array of sites represents its complete range. Most
of the locality records are from single specimens, a highly suspicious feature in itself.
This is a cryptic insect that is always buried under sand, except when males are flying
about seeking females. The apparent lack of long-range mobility suggests that the
species might not occupy isolated patches of suitable habitat, but this is not necessarily
true: there are a number of widespread scrub and sandhill organisms with apparently
/
/"
i//d
Fig. 2. Arenviaga floridensis Caudell, Male, and distribution of species.
532 Florida Entomologist 73(4) December, 1990
poor mobility such as Melanoplus puer Scudder (Acrididae) (Hubbell 1932), sand pine,
and scrub hickory (Carya floridana) (Little 1978). Moreover, going by the distribution
of other species, it is unusual for a species to be widely distributed in scrubs of north-
central Florida, part of the west coast, and the Lake Wales Ridge, but absent from the
Atlantic Coastal Ridge. For these reasons, I would hesitate to propound detailed
biogeographic scenarios based on the distribution of this cockroach. This species is now
without its biogeographic utility, as it exemplifies the link between Florida uplands and
southwestern xeric areas.
Lake Wales Ridge Species
The best-known pattern of endemism is provided by species restricted to scrubs of
the Lake Wales Ridge and adjoining interior ridges (Fig. 3) (Deyrup 1989). Fourteen
arthropods are known only from the scrubs of this ridge; about eight of these can
confidently be listed as Lake Wales Ridge endemics, the remainder have not been
assiduously sought to the north. On the other hand, there are undoubtedly many ar-
thropods confined to the ridge that have not even been described. There are ten species
of plants that are also restricted to the Lake Wales Ridge, Ziziphus celata, Dicerandra
1. Brooksville Ridge
2. Mount Dora Ridge
S3. Lake Wales Ridge
1;\ 5 4. Crescent City Ridge
5. Deland Ridge
3
6. Atlantic Coastal Ridge
7. Lakeland Ridge
/ 8. Bombing Range Ridge
Fig. 3. Major ridges of the Florida Peninsula (modified from White 1970).
Deyrup: Symposium-Origins SE Arthropod Fauna
christmanii, D. frutescens, Eryngium cuneifolium, Conradina brevifolia, Hypericum
cumulicola, Liatris ohlingerae, Calamintha ashei, Polygonella basiramia and Clitoria
fragrans (Christman & Judd 1990). There are no species of vertebrates that are found
only on the ridge, but there is a subspecies of lizard (Eumeces egregius lividus) re-
stricted to the ridge, and a monotypic endemic lizard genus (Neoseps) whose range
extends to just north of the ridge (Ashton & Ashton 1985).
There is no question that the Lake Wales Ridge has the highest diversity of Florida
scrub species, and has the only real concentration of species not found in scrubs else-
where in the state. The question is whether this ridge represents the "mother lode" or
the last best refuge of scrub species. At first glance, the mother lode theory seems
attractive. The Lake Wales Ridge is a relatively old feature, its present form probably
determined in the Pliocene (McCartan 1990, pers. comm.). The relative scarcity of
endemics restricted to other scrubs is not easily explained by hypothetical past contrac-
tions of these scrubs into very small patches, as there is no obvious reason why localized
endemics would be more likely to die out than widespread species. There is a small
number of scrub species restricted to sites off the Lake Wales Ridge, but in general
one might derive almost the entire biota of Florida scrub by dilution of the Lake Wales
Ridge biota, as if the rest of the state had been colonized by some of the more effective
dispersers from the Ridge.
There may have been a certain amount of dispersal of species from the Lake Wales
Ridge, but a more detailed analysis suggests that this ridge is actually a refuge that
has contributed little to other areas of Florida scrub. In the first place, widespread
scrub endemic species are not obviously more mobile than species restricted to the
ridge. Secondly, although the Lake Wales Ridge probably has more of its own endemics
than all the rest of Florida scrubs combined, these other endemics cannot be ignored.
Thirdly, there is a concentration of scrub endemics at the south end of the Ridge
(Christman & Judd 1990, Deyrup 1989), as if, even on the ridge, there had been a
negative factor to the north, though this might just be an increase in the dominance of
sandhill on the northern ridge. Finally, there is good evidence that at least some of the
endemics found at the south end of the ridge are western in origin, and therefore
probably came through northern Florida, where they no longer occur. Odontomachus
clarus (Formicidae) is a particularly useful example. This species is conspecific with, or
very close to, populations now found in the southwestern U.S., but in Florida it does
not occur off the Lake Wales Ridge (Deyrup et al. 1985). A number of other species
with obvious western affinities, such as the Florida scrub jay (Aphelocoma coerules-
cens), the cockroach Arenivaga floridensis, and two species of Nemomydas (Mydidae)
(Steyskal 1956), have no known populations in north Florida, including the panhandle
scrubs.
Biogeographical analysis, therefore, suggests that the Lake Wales Ridge is the last
best refuge for scrub biota. A refuge from what? The simplest hypothesis might be that
a relatively high proportion of scrub species are sensitive to certian climatic conditions,
especially periods of colder and wetter climate. Palynological studies show that over
much of the last 40,000 years north Florida was dominated by oak-hickory forest com-
parable to that of modern Pennsylvania and New Jersey, while south Florida had sand
dune scrub (Delcourt & Delcourt 1981). Changes in coastal marine invertebrate com-
munities at about 300,000 and 150,250 YBP may indicate even more severe cooling
(McCartan 1990, pers. comm.).
Sandhill habitat on the Lake Wales Ridge, although hardly enough remains to be
sampled, seems to contain no species absent from more northern sandhill, except for
scrub species that overlap into sandhill. This is quite different from northern Florida,
where there are narrowly distributed sandhill endemics. The plant Eriogonum
floridanum (Polygonaceae), which extends north into Marion Co., is the only sandhill
533
Florida Entomologist 73(4)
species that comes close to being a ridge endemic. This species, incidentally, is very
closely related to, or a disjunct population of a species of the lower Midwest (Ward 1979).
Atlantic Coastal Ridge Species
The Atlantic Coastal Ridge parallels most of the Atlantic Coast of the peninsula
(Fig. 3). Sand pine scrub and scrubby flatwoods once occupied old dune fields on this
ridge, but much of the natural habitat has been destroyed by development. The dunes
of the Coastal Ridge extend south of the Lake Wales Ridge, and one might expect from
the previous discussion that these would have a high concentration of scrub endemics,
including autochthonous species and species shared only with the Lake Wales Ridge.
This pattern, if it occurs at all, is excessively weak. Two arthropods are known to be
confined to the coastal scrubs: Lycosa pseudoceratiola Wallace (Lycosidae) (Wallace
1982) and Melanoplus insignis (Acrididae), the latter possibly a distinctive form of the
more widespread upland endemic M. forcipatus Hubbell (Hubbell 1932). Two species
of plants (Dicerandra immaculate, Asimina tetrmera) are also known only from the
Coastal Ridge (Lakela 1963, Austin & Tatje 1979). The only arthropod that appears to
be shared exclusively by the Lake Wales and Coastal Ridges is Phyllophaga elizoria
Saylor (Fig. 4), although there is an additional record from Okeechobee, once the south-
ern terminus of a low scrub ridge (Woodruff & Beck 1989).
The southern Atlantic Coastal Ridge, therefore, seems to contradict the model pro-
posed for the southern Lake Wales Ridge. There is no concentration of endemics, no
increase in species to the south, no scrub relicts absent from north central scrub areas,
unless P. elizoria is such a relict. A major difference between the two ridges is that
the Coastal ridge is much younger. Various names have been applied to the episodes
of elevated sea level that produced the complex of parallel ridges known as the Atlantic
Coastal Ridge, and these have been variously associated with warming trends in the
Northern Hemisphere, but estimates center on a period of time about 75,000 to 100,000
years ago (Austin et al. 1986, Fernald 1989, Nilsson 1982). The precise age is unimpor-
tant at this preliminary stage of analysis; the important fact is that the Lake Wales
Ridge is roughly ten times as old.
The disparity between the biotas of the Atlantic coastal and Lake Wales Ridges is
explained if one makes two assumptions. The first is that there was never an efficient
habitat corridor between the southern Lake Wales and the Coastal ridges. The biota of
the Coastal ridge is what it seems, a somewhat diluted representation of north-central
ridges such as the Mt. Dora and Deland, with a small amount of secondary speciation.
The other assumption is that the reduction in diversity of north-central scrubs antedatd
the appearance of the coastal ridge. With about 900,000 years of fluctuating climate to
work with, this seems plausible.
This is not to say that the Atlantic scrubs are uninteresting or truly depauperate,
as they include about one quarter of known scrub endemic arthropods. As in all major
scrub areas, there are surely many more endemic arthropods to be discovered, if their
habitat persists long enough.
Southwestern Uplands Species
The uplands of southwestern Florida, which once included large patches of sandhill,
as well as coastal scrub and scrubby flatwoods, have been largely extirpated. There is
Fig. 4. Distribution of three scrub-inhabiting Phyllophaga (modified from Woodruff
and Beck 1989).
December, 1990
Deyrup: Symposium-Origins SE Arthropod Fauna
L.
P.
elizoria
okeechobe
panorpa
0
.*
Florida Entomologist 73(4)
no evidence that there were ever many endemics restricted to these uplands. There is
one plant (Chrysopsis floridana) restricted to Hillsborough County scrub (Ward 1979)
and a flightless scarab (Mycotrupes pedester) known only from scrubs in three south-
western counties (Woodruff 1973). There are no species of any group known only from
the Lake Wales Ridge and from uplands west of the ridge. Unfortunately, little is
known of the arthropods of the remaining bits of sandhill from Tampa south, as one
might expect southern extension of some of the species endemic to sandhills to the
north. All of Florida's uplands should receive some "salvage entomology" if we are to
understand the patterns of biogeography, but the case of the southwestern uplands
appears particularly urgent.
North Florida Uplands
Biogeographic patterns in the northern third of Florida are particularly elusive.
These patterns are obviously complex, having been effected by radical, and often rapid
climatic changes. The areas of endemism are relatively subtle, compared with the
dramatic endemism of the Lake Wales Ridge or the Appalachicola drainage. The
biogeography of north Florida, aside from the Appalachicola area, has received rela-
tively little attention. The only arthropod studies that provide many useful data points
are those of Hubbell (1932), Woodruff (1973), Woodruff & Beck (1989), Gross et al.
(1989), and some of the accounts in Franz (1982).
There are a few general patterns that are already evident. First, as mentioned
above, there are only a few sand pine scrub endemics restricted to northern Florida,
in spite of the considerable area of scrub in the north-central ridges and the relative
isolation of the Panhandle scrubs. Second, there are numerous sandhill endemics, often
narrowly distributed, assuming that studies of Melanoplus grasshoppers (Hubbell 1932)
and scarabs (Woodruff 1973, Woodruff & Beck 1989) are only indicators of more inclusive
endemic biotas. There are also a number endemic plants, often with restricted ranges
or small disjunct populations (Ward 1979, Hardin & White 1989). Third, there is no
corresponding group of narrowly endemic vertebrates. Fourth, the upland endemics
with narrow distributions are scattered about without (as far as we know) any striking
concentrations of endemics. Fifth, the upland endemics seem matched by a greater
number of wetland endemics occurring nearby than in south Florida (Hubbell 1961).
These general patterns could all be the result of climatic fluctuations. Episodes of
dwindling and expanding habitat types could lead to many extinctions, so that there
would be relatively few of the original species in most patches of relict habitat and a
number of species would have highly disjunct populations. Larger species, or those with
low populations per unit area would be particularly vulnerable to extinction. Highly
mobile species would readily redistribute themselves, and thus be restricted endemics
(or locally extinct) in proportion to the amount of habitat that could support breeding
populations. If xeric and wetland habitats have both been expanding and contracting,
as seems likely (Delcourt & Delcourt 1981), one would expect to see both upland and
lowland endemics through north Florida. Climatic changes can also isolate populations
and apply new selective pressures in such a way as to generate new taxa (Axelrod 1981).
Some north Florida species may be autochthonous endemics; a number of wingless
Melanoplus species seem like good candidates.
Entomologists! Head for the Uplands!
My impressionistic coverage of the upland endemic arthropods of north Florida
should convey the need for further research. Even on the Lake Wales Ridge new
species of scrub endemics appear regularly. Entomologists could hold the vast majority
536
December, 1990
Deyrup: Symposium-Origins SE Arthropod Fauna
of the clues needed for a detailed understanding of Florida's biogeographic patterns. A
number of approaches are especially useful:
1. We should mount field expeditions to areas of probable endemism. There we might
search for taxa that already include known endemics, and species that have poor
dispersal ability. Other targets are species that are probably dependent on edaphic
conditions or specialized host plants of uplands.
2. We should look for subspecific differences between populations. Some patterns may
be revealed by analysis of coloration or morphology, others by electrophoresis. The
recent paper by Gross et al. (1989) provides an admirable model.
3. We should compile distribution records, preferably in the form of maps that can be
overlaid.
The accelerating destruction of upland habitats lends a deplorable urgency to the
research. If we do not collect our information soon, we may lose the opportunity. In a
more positive vein, entomological evidence can help identify concentrations of endemics
and assist in efforts to preserve habitats.
The study of Florida's upland arthropods, therefore, has everything to recommend
it. To the thrill of the chase and the joy of discovery, it adds the goal of preservation
of biological history and habitat. So seldom will inclination and duty so clearly coincide.
ACKNOWLEDGMENTS
I am particularly grateful to Dr. Lucy McCartan of the U.S. Geological Survey for
sharing her most recent discoveries on Florida geology. Nancy Deyrup prepared the
distribution maps, and Marcia Moretto typed the manuscript.
REFERENCES CITED
ABRAHAMSON, W. G. 1984. Species responses to fire on Florida's Lake Wales Ridge.
Am. J. Bot. 71: 35-43.
ABRAHAMSON, W. G., A. F. JOHNSON, J. N. LAYNE, AND P. A. PERONI. 1984.
Vegetation of the Archbold Biological Station, Florida: an example of the south-
ern Lake Wales Ridge. Florida Scient. 47: 209-250.
ASHTON, R. E., JR., AND P. S. ASHTON. 1985. Handbook of Reptiles and Amphibians
of Florida. II. Lizards, Turtles and Crocodiles. Windward Publ. Inc., Miami. 191
pp.
AUSTIN, D. F., F. R. POSIN, AND J. N. BURCH. 1986. Scrub species patterns on the
Atlantic Coastal Ridge, Florida. J. Coastal Res. 4: 491-498.
AUSTIN, D. F., AND B. E. TATJE. 1979. Species account, pp. 5-6 in Ward, P. [ed.],
Rare and endangered biota of Florida. Vol. 5: Plants. Univ. Presses of Florida,
Gainesville. 175 pp.
AXELROD, D. 1981. Holocene climatic changes in relation to vegetation disjunction
and speciation. Amer. Natur. 847-870.
BLATCHLEY, W. S. 1920. Orthoptera of Northeastern America, with special reference
to the faunas of Indiana and Florida. Nature Publ. Co., Indianapolis. 784 pp.
CHRISTMAN, S. P., AND W. S. JUDD. 1990. Notes on plants endemic to Florida scrub.
Florida Scient. 53: 52-73.
DELCOURT, P. A., AND H. R. DELCOURT. 1981. Vegetation maps for eastern North
America: 40,000 year BP to the present. pp. 123-165 in R. Romans [ed.]
Geobotany II. Plenum, New York.
DEYRUP, M. 1989. Arthropods endemic to Florida scrub. Florida Scient. 52: 254-270.
DEYRUP, M., J. TRAGER, AND N. CARLIN. 1985. The genus Odontomachus in the
southeastern United States (Hymenoptera: Formicidae). Entomol. News 96:
188-195.
FERNALD, R. T. 1989. Coastal xeric scrub communities of the Treasure Coast Region,
537
Florida Entomologist 73(4)
Florida: a summary of their distribution and ecology, with guidelines for their
preservation and management. Nongame Wild. Prog. Tech. Rep. 6: 113 pp.
FRANZ, R. ed. 1982. Rare and endangered biota of Florida. Vol. 6: Invertebrates.
Univ. Presses of Florida, Gainesville. 131 pp.
GROSS, S. W., D. L. MAYS, AND T. J. WALKER. 1989. Systematics of Pictonemobius
ground crickets (Orthoptera: Gryllidae). Trans. Amer. Entomol. Soc. 115: 433-
456.
HARDIN, E. D., AND D. L. WHITE. 1989. Rare vascular plant taxa associated with
wiregrass (Aristida stricta) in the southeastern United States. Natur. Areas J.
9: 234-245.
HUBBELL, T. H. 1932. A revision of the puer group of the North American genus
Melanoplus, with remarks on the taxonomic value of the concealed male genitalia
in the Cyrtacanthacridinae. Univ. Mich. Mus. Zool. Misc. pub. 23: 1-64.
HUBBELL, T, H, 1961. Endemism and speciation in relation to Pleistocene changes
in Florida and the southeastern coastal plain. Eleventh Internat. Congr. En-
tomol., Wein 1960, 1: 466-469.
LAKELA, 0. 1963. Dicerandra immaculate Lakela, Sp. Nov. (Labiatae). Sida 1:
184-185.
LITTLE, ELBERT L. 1978. Atlas of United States Trees. Misc. Publ. 1361 U.S.D.A.
For. Serv.: 1-22, + 256 maps.
MYERS, R. L. 1985. Fire and the dynamic relationship between Florida sandhill and
sand pine scrub vegetation. Bull. Torrey Bot. Club 112: 241-252.
MYERS, R. L., AND D. L. WHITE. 1987. Landscape history and changes in sandhill
vegetation in north-central and south-central Florida. Bull. Torrey Bot. Club
114: 21-32.
NILSSON, T. 1982. The Pleistocene: geology and life in the Quarternary Ice Age. D.
Reidel Publ. Co. Dordrecht, Holland. 651 pp.
WARD, D. B., ed. 1979. Rare and endangered biota of Florida. Vol. 5: Plants. Univ.
Presses of Florida, Gainesville. 175 pp.
WALLACE, H. K. 1982. Order Aranae, species accounts. pp. 120-129 in Franz, R.
[ed], Rare and endangered biota of Florida. Vol. 6: Invertebrates. Univ. Presses
of Florida, Gainesville. 131 pp.
WHITE, W. A. 1970. The geomorphology of the Florida Peninsula. Fla. Dept. Nat.
Res. Geol. Bull. 51: 164 pp.
WOODRUFF, R. E. 1973. Arthropods of Florida and neighboring land areas. Vol. 8:
the scarab beetles of Florida (Coleoptera: Scarabaeidae). Part I. The Laparosticti
(Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae, Ochodaeinae, Geo-
trupinae, Acanthocerinae). Florida Dept. Agric., Div. Plant Indust. 220 pp.
WOODRUFF, R. E., AND B. M. BECK. 1989. Arthropods of Florida and neighboring
land areas. Vol. 13: the scarab beetles of Florida (Coleoptera; Scarabaeidae).
Part II. The May or June beetles (genus Phyllophaga). Florida Dept. Agric.,
Div. Plant Indust. 225 pp.
538
December, 1990
Allen: Symposium-Origins SE Arthropod Fauna
INSECT ENDEMISM IN THE INTERIOR HIGHLANDS
OF NORTH AMERICA
ROBERT T. ALLEN
Department of Entomology
University of Arkansas
Fayetteville, AR 72701
ABSTRACT
Sixty-eight species of insects are considered endemic to the Interior Highlands of
North America. The area encompassed by these species consists of the Ozark, Ouachita,
Arbuckle, and Wichita Mountains of Illinois, Missouri, Arkansas, and Oklahoma. County
maps are given for each species as well as maps showing the distribution of close
relatives, where available. The hypothesis that all endemism in the Interior Highlands
is the result of events associated with Pleistocene glaciation is questioned because the
area has been an above water land mass since the Pennsylvanian era. Based on taxon/
area cladograms of sixteen of the species, a biogeographic pattern is suggested. It is
further suggested that only two vicariant events were necessary to account for the
origin of the species represented in the taxon/area cladograms. The times at which these
events may have occurred is uncertain. The first event may have been in the early
cretaceous when the Interior Highlands was an isolated island surrounded by epiconti-
nental seas.
RESUME
Sesenta y ocho species de insects se consideran end6micos al interior montafioso
de Norte America. El Area circundada por estas species consiste de las Ozark,
Ouachita, Arbuckle, y las montafas Wichita de Illinois, Missouri,, Arkansas, y Ok-
lahoma. Se proven mapas de condados para cada espcie asi como mapas ensefando la
distribuci6n de parientes cercanos cuando estaban disponibles. La hip6tesis que todo el
endemismo en el interior montafioso es el resultado de events asociados con la glacia-
ci6n Pleistocena se pone en duda porque el Area ha estado sobre el nivel del agua desde
la era Pensilvinica. Basado en cladogramas del Area por taxa de dieciseis species, se
sugiere un patron biogeografico. Se sugiere ademAs que solo dos events vicarios fueron
necesarios para dar cuenta del origen de las species representadas por cladogramas
del Area por taxa. Son inciertos los moments en que estos events ocurriron. El primr
event pudiera haber ocurrido temprano en la era cretdcea cuando el interior montafoso
era una isla aislada rodeada de mares epicontinentales.
As knowledge about the basic taxonomy and distribution of the North American
biota has increased during this century, biogeographers have recognized a number of
geographical areas containing significantly high numbers of endemic taxa. Three of the
better-known North American areas of endemism are peninsular Florida, the southern
Appalachians (especially the Great Smoky Mountains), and the Pacific Northwest. A
less well-known area of endemism is the mid-continent, Interior Highlands occupying
portions of Illinois, Missouri, Arkansas, and Oklahoma. There are over 200 species of
plants and animals that appear to be endemic to the Interior Highlands (Allen, unpub-
lished ms). Among the endemic animals are 68 species of insects. These species are
distributed throughout the class Insecta, specifically in the following orders: Diplura,
Microcoryphia, Ephemeroptera, Odonata, Plecoptera, Orthoptera, Hemiptera, Coleop-
tera, Trichoptera, Lepidoptera. This represents a significant number of endemic forms
539
540 Florida Entomologist 73(4) December, 1990
and indicates that the Interior Highlands have played an important role in the evolution
of the North American insect fauna.
A few insect systematists have discussed various insect taxa known to have endemic
species in the Interior Highlands and suggested reasons for this endemism. Among
these authors are: Ross, Trichoptera (1956, 1965); Ross & Ricker, Plecoptera (1971);
McCafferty, Ephemeroptera (1977); McCafferty & Provonsha, Ephemeroptera (1978);
Freitag, Coleoptera (1969); Allen & Carlton, Coleoptera (1988). Many authors, noted
throughout this paper, have described endemic Interior Highlands species.
When insect systematists have discussed the possible causal factors that have led to
speciation in the Interior Highlands, the explanation has been the events associated
with Pleistocene Glaciation (Ross 1965, Ross & Ricker 1971, McCafferty 1977). This
Pleistocene explanation is certainly one possibility. However, the Pleistocene explana-
tion fails to take into account that the Interior Highlands, specifically the Ozark and
Ouachita Mountains, have been a positive feature on the North American continent
since the Pennsylvanian (320 million years agao, mya). Thus the Interior Highlands
have been subject to habitation by a land biota much, if not most, of this time, and this
biota has been alternately connected and isolated from other North American biotas on
numerous, separate occasions. One may conclude that, except for the Pleistocene, the
vast majority of the geological and biological history of the Interior Highlands has been
virtually ignored. If an accurate understanding of the origin and evolution of the Interior
Highlands biota is to be formulated, then it will be necessary to consider the possible
effect that such events as the late Tertiary rejuventaion of the Laramide revolution
(beginning in the Mocene) and the formation of the prairie wedge (Cole & Armentrout
1979, Dickinson 1979) and the climatic and geological changes during and after the
Cretaceous (135-63 mya) had on this Interior Highlands biota. Such a task will not be
easy, but it is certainly necessary.
SIGNIFICANCE AND IMPORTANCE OF THE INTERIOR HIGHLANDS FAUNA
The significance and importance of the endemic Interior Highlands biota has not
been fully realized for a number of reasons. The descriptions and discussions of the
Interior Highland endemic insects (as well as other animals and plants) have appeared
in a diverse number of journals and books over an extended period of time. Specialists
for one group, say the Coleoptera, may not be aware of the endemic species in other
taxa.
A second factor that may have hindered work on the Interior Highlands insect fauna
is that collecting in the area can be fortuitous if one does not visit the area at the "right
time of the year." Normally, the best collecting time for insects is from April until June,
and from late September to mid October. This is when the most rainfall occurs. Because
of the porous nature of the soils the surface dries rapidly, even after heavy rains in the
spring, and insects become increasingly scarce as dry conditions prevail. A collector
visiting the Interior Highlands in July or August normally would find a depauparate
insect fauna except along the larger streams and around lakes or springs.
Some groups of insects seem to have significantly smaller populations in the Interior
Highlands than in other areas of North America. Members of the ground beetle tribe
Cychrini can be readily collected in the spring in the Great Smoky Mountains. In the
Interior Highlands one is lucky to find more than one or two specimens after a hard
day's work of moving rocks, leaf debris, and rotting logs. The pselaphid genus Arianops
is another example of the low population phenomenon. Three years of intensive collect-
ing in the Interior Highlands has produced only six specimens belonging to this genus,
whereas members of the genus seem to be more abundant in the Appalachian Mountains
(Barr 1974).
Allen: Symposium-Origins SE Arthropod Fauna
All of the factors discussed tend to deter insect collectors from working intensively
in the Interior Highlands. However, recent papers by a number of authors indicate that
the insect fauna of the area can contribute a great deal to our overall understanding of
the origin and evolution of the insect fauna of North America. Examples of some of the
recent papers are those of Allen & Carlton (1988), Robotham & Allen (1988), Allen et
al. (1988), Stark et al. (1983), Chandler (1988), Mathis & Bowles (1989).
The recent papers pertinent to the entomofauna of the Interior Highlands indicate
that the area contains a rich, unknown group of endemic insect taxa. The endemism not
only exists at the species level but also at the generic level (Chandler 1988, Mathis &
Bowles,1989). As these new forms are described, it would be profitable if workers
would also analyze the cladistic/biogeographic affinities of the taxa. Such analyses would
allow us to better understand the origin and evolution of this biota.
To hasten our understanding of endemism in the Interior Highlands, this paper
summarizes what is currently known about the entomofauna. The biogeographic af-
finities of this fauna with other North American areas of endemism will also be dis-
cussed.
PHYSICAL COMPOSITION OF THE INTERIOR HIGHLANDS
The Interior Highlands consist of a number of distinct geological provinces found in
the states of Illinois, Missouri, Arkansas, and Oklahoma, Fig. 1. The Ozark Mountains
Fig. 1. Map showing the major geological divisions of the Interior Highlands. AKB
= Arkhoma Basin, ARB = Arbuckle Mountains, IOZ = Illinois Ozark Mountains,
OUA = Ouachita Mountains, OZK = Ozark Mountains, WIC = Wichita Mountains.
Florida Entomologist 73(4)
extend from southern Illinois, through Missouri, into northern Arkansas and a small
part of northeastern Oklahoma. The Ouachita Mountains lie mostly in southwestern
Arkansas but extend into eastern Oklahoma. The Ozark and Ouachita uplifts are sepa-
rated by the Arkansas River valley, known geologically as the Arkhoma Basin. Erosion
in the Arkhoma Basin has formed isolated, positive, prominent features including
Magazine Mountain, Petit Jean Mountain, and Mount Nebo.
Magazine Mountain, 2,753 feet, is the highest point in Arkansas and one of the
highest points between the Appalachian and Rocky Mountains. At least 12 species are
known to be endemic to Magazine Mountain including a tree, land snails, beetles, a
mayfly, and a caddisfly.
The far western element of the Interior Highlands consists of two geological features
in the southern part of Oklahoma. The Arbuckle and Wichita Mountains are now an
area of relatively low relief but are, geologically (and probably biologically also), a part
of the Interior Highlands uplift. They are on the southern edge of the formation and
have received even less attention from biologists than the Ozark and Ouachita Moun-
tains.
The geological areas composing the Interior Highlands have been positive features
on the North American continent since the Pennsylvanian Period, arising some 320
million years before the present (Allen & Cox, in preparation). The areas surrounding
the Interior Highlands have been periodically inundated by epicontinental seas or sub-
jected to more xeric conditions than the highlands and, most recently flooded by melt
water from glaciers. There is no evidence to suggest that the Highlands, including the
southern Illinois segment, have ever been glaciated. Thus the Interior Highlands have
been alternately isolated and reconnected with other land areas in North America. But
the Highlands have remained above water and free of ice during their entire history
(Allen & Cox, in preparation). The area was thus available for habitation by a diverse
flora and fauna of both land dwelling and freshwater, aquatic taxa.
METHOD
Ideally, one would like to examine the cladistic and area relationships of the taxa
endemic to the Interior Highlands and determine the distribution patterns exhibited by
these taxa. This usually is not possible, however, because cladograms for the large
majority of the taxa have not been constructed. An alternative, the one used here, is
to map the distributions of the endemic forms, the distributions of their nearest relatives
(if known), and use those cladograms where available in order to search for "suggested"
distribution patterns. It is anticipated that this method will suggest fertile areas for
future work as well as shedding some light on what we can now summarize about the
evolution of the Interior Highlands insect biota.
Fig. 2. Distribution of endemic members of the order Diplura (Japygidae) in the
Interior Highlands: Catajapyx ewingi (circle), Eojapyx pedis (triangle), Occasjapyx
carltoni (square), Occasjapyx n. sp. (diamond), (Camposeidae) Campodeida n. sp. 1 &
2 (cross).
Fig. 3. Distribution of the genus Occasjapyx (Diplura: Japygidae) in North America.
Fig. 4. Pedetontus n. sp., (Microcoryphia: Machilidae): distribution in the Interior
Highlands.
Fig. 5. Distribution of the genus Pedetontus (Microcoryphia: Machilidae) in North
America.
Fig. 6. Distribution of endemic Ephemerellidae (Ephemeroptera) in the Interior
Highland: Ephemerella (Dannella) provonshai (circle); Paraleptophlebia calcarica
(cross); Habrophlebiodes annulata (circles).
542
December, 1990
Allen: Symposium-Origins SE Arthropod Fauna
_- provonshai U
S simplex
lita 0
Fig. 7. Distribution of the Ephemerella subgenus Dannella (Ephemeroptera:
Ephemerellidae) in North America: E. (Dannella) provonshai (square); E. (Dannella)
simplex (diamonds); E. (Dannella) lita (circles).
Fig. 8. Cladistic relationships of members of the Ephemerella subgenus Dannella
(Ephemeroptera: Ephemerellidae).
544 Florida Entomologist 73(4) December, 1990
The following section includes taxa considered to be endemic to the Interior High-
lands. Distributions given here include both published and unreported county records
following the state abbreviations.
ENDEMIC INSECT TAXA
Diplura
Relatively little is known about this order in North America. A series of papers by
Smith recorded a number of new taxa in the family Japygidae, mostly from California.
A few eastern North American species were also described by Smith. One of these
species, Eojapyx pedis, is known only from Missouri. At the present time there are
four species of Diplura known from the Interior Highlands. These species may or may
not prove to be endemic to the area. The two new species in the family Campodeidae
represent the first record of the subgenus Podocampus north of the Rio Grande Valley.
Japygidae
Catajapyx ewingi Fox 1941:28. AR: Howard. Fig. 2.
Eojapyx pedis Smith 1960:262. MO: Stoddard. Fig. 2.
Occasjapyx carltoni Allen 1988:22. AR: Newton. Figs. 2, 3.
Occasjapyx n. sp. AR: Garland. Figs. 2, 3.
Campodeidae
Campodea (Podocampa) n. sp. 1 and 2. AR: Logan. Fig. 2.
MICROCORYPHIA
Machilidae
There are five known North American species in the genus Pedetontus. The Arkan-
sas species is most closely related to P. persquamosus, a species found in California,
Fig. 4.
Pedetontus n. sp. AR: Logan. Fig. 5.
Ephemeroptera
There are three endemic species of mayflies in the Interior Highlands. A cladogram
is available for only Ephemerella (Dannella) provonshai (McCafferty 1977). This species
is most closely related to E. simplex confined to localities east of the Mississippi River.
Ephemerellidae
Ephemerella (Dannella) provonshai McCafferty 1977:886. AR: Johnson. Figs. 6, 7, 8.
Leptophlebiidae
Paraleptophlebia calcarica Robotham & Allen 1988:318. AR: Logan. Fig. 6.
Habrophlebiodes annulata Traver 1934:199. AR: Johnson, Scott. OK. Fig. 6.
Odonata
Only one dragonfly species is a known endemic of the Interior Highlands, Gom-
Allen: Symposium-Origins SE Arthropod Fauna
phurus ozarkensis. This species has a wide distribution in the Interior Highlands. Its
cladistic and biogeographic affinities are unknown.
Gomphurus ozarkensis (Westfall) 1975:91. Fig. 9.
Plecoptera
Several stonefly genera have endemic species in the Interior Highlands. Recent
work in Arkansas indicates that there are still a number of undiscovered and unde-
scribed species. Cladistic analyses of the genera Neoperla, Alloperla, and Isoperla are
needed.
Capniidae
Ross & Ricker (1971 discussed, at length and in some detail, their ideas concerning
evolution and speciation in Allocapnia. Basically, these authors suggested that specia-
tion in the Interior Highlands was the result of dispersal into and out of the area at
various times during the Pleistocene. Populations in the Interior Highlands became
isolated from those in the east and northeast and subsequently speciation occurred in
some lineages. The Illinois Ozarks were seen as a corridor with suitable habitats through
which Allocapnia populations could move. Ross (1956, 1965) had already discussed this
corridor in reference to caddisfly (Trichoptera) dispersal. McCafferty (1977) also thought
the Illinois Ozarks served as a dispersal route for at least one group of mayflies
(Ephemeroptera).
The taxon most closely related to Allocapnia is the genus Capnia. Ross & Ricker
(1971) suggested that the Capnia vidua group, found in Europe, is the sister group of
Allocapnia. Thus Allocapnia species endemic to the Interior Highlands have a Euro-
pean/Interior Highlands/eastern North American distribution pattern.
Allocapnia mohri Ross & Ricker 1964:91. OK: Leflore. Figs. 10, 13.
A. warren Ross & Yamamoto 1966:265. AR: Washington. Figs. 11, 14.
A. peltoides Ross & Ricker 1964:91. AR: Scott. OK. Haskell, Leflore. Figs. 11, 14.
A. ozarkana Ross 1964:172. AR: Madison, Washington. Figs. 11, 15.
A. jeanae Ross 1964:171. AR: Carroll, Madison, Washington. Figs. 11, 16.
A. sandersoni Ricker 1952:165. AR: Washington. Figs. 12, 16.
A. oribata Poulton & Stewart. 1987:296. AR: Searcy, Van Buren. Fig. 12.
Leuctridae
Ricker & Ross (1969) studied the basic taxonomy, distribution, and cladistic relation-
ships of the genus Zealeuctra. The distribution of the members of the genus is curious.
All the species occur north of the ancient demarcation line of the Pennsylvanian age
(320 mya) Ouachita/Marathon uplift areas in Arkansas, Oklahoma, and Texas. The range
of the qenus now extends east of the Mississippi River. The distribution of the extant
species suggests that the evolution of this group may be associated with the long geolog-
ical history of this area.
Zealeuctra wachita Ricker & Ross 1969:1119. AR: Polk, Scott. Figs 17, 18.
Z. cherokee Stark & Stewart 1973:192. OK: Adair, Leflore, Pushmataha, Sequoyah.
Figs. 17, 18.
Strophopteryx cucullata Frison 1934:29. OK: Haskell, Latimer, Leflore, Pushmataha,
Sequoyah. Figs. 17, 18.
Perlidae
Neoperla harpi Ernst & Stewart, In Ernst et al., 1986:646. AR: Carroll, Clark, Craw-
545
Florida Entomologist 73(4)
Fig. 9. Gompurus ozarkensis (Odonata: Gomphidae): general distribution in the In-
terior Highlands.
Fig. 10. Allocapnia mohri (Plecoptera: Capniidae) distribution in the Interior High-
lands.
Fig. 11. Distribution of endemic Allocapnia (Plecoptera: Capniidae) species in the
Interior Highlands: A. warren (open square); A. peltoides (squares); A. ozarkana (dia-
mond); A. jeanae (circles).
Fig 12. Distribution of endemic Allocapnia (Plecoptera: Capniidae) species in the
Interior Higlands : A. sandersoni (circles); A. oribata (square).
December, 1990
Allen: Symposium-Origins SE Arthropod Fauna
1 b polemistis peltoides *
3b -recta -5 frisoni
loshada granulata 0
mohri 14b unzicker 0
-malverna warreni *
fumosa E
virginiana o
maria C
.minima 0 stannardi 0
15b pechumani 0 16--rickeri 0
curiosa 1 ---- sandersoni Ci
ozarkana cunninghami *
forbesi zo:a
illinoensis perplexa
jeanae U
Fig. 13a, b. Distribution (13a) and cladistic relationships (13b) of members of the
Allocapnia mohri (Plecoptera: Capniidae) lineage.
Fig. 14a, b. Distribution (14a) and cladistic relationships (14b) of members of the
Allocapnia warreni-A. peltoides (Plecoptera: Caphiidae) lineage.
Fig. 15a, b. Distribution (15a) and cladistic relationships (15b) of members of the
Allocapnia ozarkana (Plecoptera: Capniidae) lineage.
Fig. 16a, b. Distribution (16a) and cladistic relationships (16b) of members of the
Allocapnia sandersoni-A. jeanae (Plecoptera: Capniidae) lineage.
Florida Entomologist 73(4)
ford, Franklin, Howard, Nevada, Pike, Polk, Saline, Scott, Sevier, Stone, Van Buren,
Washington, Yell. OK: Delaware. Fig. 19.
N. robisoni Poulton & Stewart, In Ernst et al., 1986:648. AR: Benton, Clark, Hot
Spring, Izard, Johnson, Nevada, Ouachita, Pike, Sevier, Sharp, Yell. OK: McCurtain.
Fig. 20.
N. falayah Stark & Lentz 1988:371. AR: Montgomery, Washington. MO: McDonald.
OK: Delaware. Fig. 21.
N. osage Stark & Lentz 1988:372. AR: Montgomery, Washington, MO: Christian,
McDonald. OK: Adair, Delaware. Fig. 22.
Chloroperlidae
Alloperla ouachita Stark & Stewart, In Stark et al. 1983:56. AR: Montgomery. Fig. 23.
A. caddo Poulton & Stewart, 987:297. AR: Garland, Perry. Fig. 23.
Perlodidae
Isoperla szczytkoi Poulton & Stewart, 1987:298. AR: Logan. Fig. 23.
ORTHOPTERA
Acrididae
Only one species in the order Orthoptera appears to be exclusively endemic to the
Interior Highland. Its nearest relative is Eximacris superbum known only from ex-
treme southwest Texas.
Eximacris phenax Otte 1984:165. OK. Figs. 24, 25.
Hemiptera
Only one hemipteran species is known to be endemic to the Interior Highlands.
However, relatively little is known about the distribution of many of the taxa in the
order that are soil dwelling forms. This may be a fertile area for investigation.
Tingidae
Acalypta susanae Allen, Carlton, Tedder 1988:126. AR: Logan, Polk. Fig. 26.
Fig. 17. Distribution of Zealeuctra (Plecoptera: Leuctridae) species in the Interior
Highlands: Z. cherokee (squares); Z. wachita (diamonds); Strophopteryx cuculata (cir-
cles).
Fig. 18 a, b, c. Distribution (18a, b) of Zealeuctra (Plecoptera: Leuctridae) species
in North America; cladistic relationships (8c) among species of Zealeuctra.
Figure 19. Neoperla harpi (Plecoptera: Perlidae): distribution in the Interior High-
lands.
Figure 20. Neoperla robisoni (Plecoptera: Perlidae): distribution in the Interior
Highlands.
Figure 21. Neoperlafalavah (Plecoptera: Perlidae): distribution in the Interior High-
lands.
Fig. 22. Neoperla osage (Plecoptera: Perlidae): distribution in the Interior High-
lands.
December, 1990
Allen: Symposium-Origins SE Arthropod Fauna
+
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o
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i 55
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7 I
- I T l ^
_, ^, ~ ^ ~ ~ J
S/ r i' -, \ -.- -
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549
Florida Entomologist 73(4)
Fig. 23. Distribution of endemic Plecoptera Alloperla (Chloroperlidae) and Isoperla
(Perlodidae) species in the Interior Highlands: A. ouachita (square); A. caddo (circles);
Isoperla szczytkoi (diamond)
Fig. 24. Eximacris phenax (Orthoptera: Acrididae): distribution in the Interior
Highlands.
Fig. 25. Distribution of the genus Eximacris (Orthoptera: [Family?]) in North
America.
December, 1990
Allen: Symposium-Origins SE Arthropod Fauna
Coleoptera
The beetles are represented in the Interior Highlands by a number of endemic forms
in several different families. As the beetle fauna is explored many additional endemic
taxa may be discovered. Those taxa for which reliable cladograms are available are
Scaphinotus, subgenus Nomaretus and the genus Evarthrus in the family Carabidae
and the genera Arianops and Ouachitychus in the family Pselaphidae.
Carabidae
Rhadine ozarkensis Sanderson & Miller !941:39. AR: Washington. Figs. 27, 28.
Scaphinotus (s. str.) parisiana Allen & Carlton 1988:130. AR: Logan, Washington.
Figs. 27, 29.
Scaphinotus (Nomaretus) infletus Allen & Carlton 1988:132. AR: Newton. Fig. 27.
There are five species in the subgenus Nomaretus. Four of these species occur west
of the Mississippi River, and the fifth species, S. bilobus, occurs both west of the
Mississippi River and in previously glaciated areas in the East, Fig. 29a. The cladogram
proposed by Allen & Carlton (1988) places infletus at the base of this monophyletic
cluster of species.
Gidaspow (1973) noted that the subgenus "occurs in the Ozark Uplift (Missouri),
except for S. bilobus which, following the retreating ice sheet, wandered to the region
of the Great Lakes and the mountains of New York and New Hampshire." She also
noted that S. liebeki occurs in the South, in Texas and in one locality in Louisiana. It
should also be noted that S. fissicollis and S. cavicollis are found outside the Interior
Highlands in Kansas and S. cavicollis in Oklahoma. Because of the sympatry in these
species it is not possible to suggest what events may have led to their isolation and
evolution. However, consideration of the sister taxa of the subgenus Nomaretus does
show a distinct disjunct distribution pattern in three monophyletic lineages.
Evarthrus whitcombi Freitag 1969:129. AR: Figs. 30, 31.
E. parasodalis Freitag 1969:150. AR: Figs. 30, 32.
Evarthrus incisus, occurring mostly north of the Arkansas River, is the sister
species of E. whitcombi and occurs mostly south of the Arkansas River. There are five
additional species in the monophyletic lineage that gave rise to E. incisus and E. whit-
combi. These additional species all occur east of the Mississippi River. The cladogram
of relationships proposed by Freitag (1969) would suggest that an original cosmopolitan
eastern population was divided into eastern and western segments and that the eastern
segment eventually gave rise to five species and the western segment evolved into E.
whitcombi and E. incisus.
Evarthrus parasodalis belongs to a monophyletic lineage composed of four other
species: E. sodalis (with three subspecies), E. furtivus, E. alternans, E. iowensis. The
cladogram of relationships among these species and the distribution patterns, Fig. 32,
Fig. 26. Acalypta susanae (Hemiptera: Tingidae): distribution in the Interior High-
lands.
Fig. 27. Distribution of endemic Carabidae (Coleoptera) in the Interior Highlands:
Rhadine ozarkensis (square); Scaphinotus (s. str.) parisiana (circle); Scaphinotus
(Nomaretus) infletus (diamond).
Fig. 28. Distribution of the genus Rhadine (Coleoptera: Carabidae) in North
America: R. eurepes (small stippling); R. ozarkensis (circle); R. lavalis (left slanted
diagonals); R. caudata (right slanted diagonals).
Fig. 29a, b. Distribution (29a) and cladistic relationships (29b) of three subgenera of
Scaphinotus (Coleoptera: Carabidae).
Florida Entomologist 73(4)
3.. sodalis z e
32b parasodalis 0
furtivus S
alternans c>
/ -iowensis 0
-convivus
-sinus A
-sigillatus 0
-blatchleyi @
-floridensis :
-incisus
-whitcombi *
Fig. 30. Distribution of endemic species of Evarthrus (Coleoptera: Carabidae) in the
Interior Highlands: E. parasodalis (squares); E. whitcombi (cirlces).
Fig. 31a, b. Distribution (31a) and cladistic affinities (32b) of members of the Evar-
thrus whitcombi (Coleoptera: Carabidae) lineage.
Fig. 32a, b. Distribution (32a) and cladistic affinities (32b) of members of the Evar-
thrus parasodalis (Coleoptera: Carabidae) lineage in North America.
Fig. 33. Distribution of endemic species of Dytiscidae (Coleoptera in the Interior
Highlands: Hoperius planatus (circle); Hydroporus sulfurius (square); Hydroporus
ouachitus (diamond).
suggests that this lineage had historically been divided into eastern and western popu-
lations by the Mississippi River Valley. These ancient eastern and western populations
subsequently gave rise to a number of species from each population, Fig. 31a, 32a.
Dytiscidae
Hoperius planatus (Fall) 1927:178. AR: Hempstead. Fig. 33.
Hydroporus sulfurius Matta & Wolfe 1979:287. AR: Benton. Fig. 33.
H. ouachitus Matta & Wolfe 1979:289. AR: Polk. Fig. 33.
December, 1990
Allen: Symposium-Origins SE Arthropod Fauna
Pselaphidae
Arianops sandersoni Barr 1974:21. AR: Logan. Figs. 34, 36.
A. stephani Carlton In Carlton and Allen 1989:62. OK: Latimer. Figs 34, 36.
These two species are known from isolated, single localities in the Ouachita Moun-
tains. Arianops sandersoni lives in leaf litter and beneath stones on the north and east
slopes of Magazine Mountain, Logan County, Arkansas. The genus Arianops occurs in
the Appalachian Mountains and seems to be confined to mountains in the Interior High-
lands south of the Arkansas River. This distribution pattern may indicate an ancient
connection between the southern Appalachians and the Ouachita Mountains across
northern Alabama and Mississippi.
Ouachitychus parvoculus Chandler 1988:160. AR: Logan. Figs. 34, 35, 36.
This endemic genus and species is also known only from Magazine Mountain, Logan
County, in western Arkansas. The sister qroup of Ouachtychus is the genus Cylin-
drarctus with 10 species in eastern North America. The biogeographic origins of
Ouachitychus are unclear.
Staphylinidae
Derops divalis (Sanderson) 1946:131. AR: Logan, Washington. Figs. 37.
Coccinellidae
Scymnus (P.) wingoi Gordon 1976:215. MO.
Bruchidae
Acanthoscelides schrankiae Horn 1873:331.
Chrysomelidae
Pachybrachis pinicola Rouse & Medvedev 1972:82. AR: Nevada. Fig. 38.
Lema maculicollis ab. inornata Rouse & Medvedev 1972:81. AR: Montgomery. Fig. 38.
Trichoptera
There are at least 22 known endemic species of caddisflies in the Interior Highlands.
These species are usually found in clear, fast-flowing streams or in isolated fresh water
springs. Some of the large streams, such as the Buffalo River and the White River,
flow throughout the year while many of the small streams have an intermittent flow-
heavy in the spring and decreasing or drying up by late June or early July.
The origin and evolution of many of the Interior Highland endemic caddisflies has
been discussed by H. H. Ross (1956). Ross thought that Pleistocene Glaciation played
an important role in the isolation of Ozark/Ouachita Mountain populations. These iso-
lated populations subsequently evolved into distinct taxa. This possibility will be dis-
cussed in greater depth in the discussion section of this paper.
Hydropsychidae
Cheumatopsyche rossi Gordon 1974:129. AR: Fulton, Johnson, Logan, Madison,
Washington. OK:. Haskell. Fig. 39.
In her treatment of the genus Cheumatopsyche, Gordon (1974) placed C. rossi in a
monophyletic lineage containing three other species. Two of the species in this lineage,
553
Florida Entomologist 73(4)
35 Ouachitychus
--Cylindrarctus
Custotychus
/4 Atychodea
IH
NAe
NAe
AS
-other Arianops
-sandersoni
-amplyponica
other amply. sp.
Paramaurops
Amaurops
Fig. 34. Distribution of endemic species of Pselaphidae (Coleoptera) in the Interior
Highlands: Arianops sandersoni (circle); Arianops stephani (diamond); Ouachitychus
parvoculus (square).
Fig. 35. General distribution and cladistic affinities among genera closely related to
Ouachitychus (Coleoptera: Pselaphidae).
December, 1990
554
Allen: Symposium-Origins SE Arthropod Fauna 555
C. smith and C. logani are found only in the west, while C. pettiti is transcontinental.
Since C. pettiti has a transcontinental distribution it is unclear where or when a once
cosmopolitan population was first divided. The intermediate position of C. rossi in the
cladogram does suggest that this Interior Highland endemic shares a close relationship
and possible and immediate common ancestor with two western species.
Ceratopsyche piatrix (Ross) 1938a:148. AR: Fulton. MO: Oregon. Fig. 40.
Psychomyiidae
Paduniella nearctica Flint 1967:310. AR: Crawford, Johnson, Washington. Fig. 41.
This Interior Highland endemic represents one of the most interesting and provoca-
tive distribution patterns among the endemic fauna. The genus Paduniella is found in
Ceylon, Java, Africa, India, Phillippines, Indonesia, and the southern Usuri region of
the Soviet Union. Paduniella nearctica is the sole representative of the subfamily
Paduniellinae in North America. This suggests a very ancient geographic relationship-
perhaps of pre-Cretaceous origin. Paduniella nearctica appears to be most closely re-
lated to P. sanghamittra Schmid from Ceylon (Flint 1967).
Philopotomidae
Wormalida strota (Ross) 1938c:11. AR: Johnson, Madison, Perry, Washington. Fig. 42.
Ross (1956) placed W. strota in a monophyletic lineage with 4 other species, fig. 42b.
The pattern suggested by the cladogram indicates that at least two independent, trans-
continental, ancient populations were isolated into eastern and western segments and
gave rise to a number of separate but closely related species. One population gave rise
to W. thyria and W. hamata, and the other population gave rise to W. occidea, W.
shawnee and W. strota.
Glossosomatidae
Agapetus artesus Ross 1938a:106. MO: Oregon, Phelps. Fig. 43.
A. illini Ross 1938a:106. AR: Benton, Carroll, Crawford, Franklin, Garland, Johnson,
Logan, Madison, Montgomery, Polk, Scott, Washington. IL: Hardin, Pope, Union. OK:
Cherokee. Fig. 43.
A. medicus Ross 1938a:107. AR: Benton, Clark, Hot Spring, Montgomery, Pike, Polk.
Fig. 44.
Rhyacophilidae
Rhyacophila fenestra Ross 1938a:102. IL: Hardin, Jackson, Johnson, Massac, Pope,
Tazewell, Union, Vermilion. Fig. 45.
R. kiamichi Ross 1944:37. AR: Crawford, Hot Spring, Johnson, Logan, Madison, Pike,
Polk, Washington. OK: Cherokee, Haskell, Johnston, Latimer. Fig. 45.
Fig. 36a, b. Distribution (36a) and cladistic affinities (36b) among North American
species of Arianops (Coleoptera: Pselaphidae).
Fig. 37a, b. Distribution of Derops divalis (Coleoptera: Staphylinidae) in the Interior
Highlands (37a); world wide distribution of the genus Derops (37b).
Fig. 38. Distribution of endemic Chrysomelidae (Coleoptera) in the Interior High-
lands: Pacybrachis pinicola (circle); Lema maculicollis var. inornata (square).
Florida Entomologist 73(4) December, 1990
556
Allen: Symposium-Origins SE Arthropod Fauna 557
There are two species in the genus Rhyacophila that are possibly endemic to the
Interior Highlands. Rhyacophilafenestra has been reported from two localities outside
of the Interior Highlands and is included here provisionally. Ross (1956) assigned these
two species to his "Branch 6" of the genus. The two species were placed in a monophyle-
tic lineage including R. ledra, R. carolina, and R. teddyi, all found in eastern North
America. The cladogram given by Ross (1956), Fig. 45b, suggests that the two Interior
Highland endemics were isolated from one of the other eastern species, possibly R.
kiamichi.
Brachycentridae
Micrasema ozarkana Ross & Unzicker 1965:254. AR: Fulton. MO: Carter, Oregon. Fig.
46.
Leptoceridae
Ceraclea nepha (Ross) 1944:230. AR: Benton, Washington, White. OK: Pushmataha.
Fig. 46.
Setodes oxapia (Ross) 1938b:88. AR: Benton, Washington. Fig. 46.
Trianodes smith Ross 1959:40. IL: Union. Fig. 46.
Helicopsychidae
Helicopsyche limnella Ross 1938a:179. AR: Benton, Carroll, Franklin, Garland, Hot
Spring, Johnson, Madison, Montgomery, Ouachita, Sebastian, Washington. Fig. 47.
Ross (1938a) suggested that H. limnella was so similar to its close relative H.
mexicana (known from the southwest and Mexico) that the Ozark populations might
only be a variation. However Bowles and Allen (in press) established the validity of H.
limnella and suggested a cladogram of relationships for the complex to which this
interior Highland endemic belongs. H. limnella represents a distinct type of Interior
Highlands/Mexico vicariance pattern.
Hydroptilidae
Ochrotrichia anisca (Ross) 1941:58. AR: Benton, Clark, Crawford, Garland, Johnson,
Madison, Polk, Washington. IL: Union. OK: McCurtain. Fig. 47.
0. contorta (Ross) 1941:60. MO: Oregon. Fig. 48.
0. unio (Ross) 1941:56. IL: Hardin, Jackson, Pope, Union. Fig. 48.
0. weddleae (Ross) 1944:274. AR: Hot Spring, Polk. OK: Latimer. Fig. 48.
0. edalis Ross 1941:62. OK: Johnson. Fig. 46.
Neotrichia kitae Ross 1941:60. MO: Taney. Fig. 48.
Paucicalcaria ozarkensis Mathis & Bowles 1989:188. AR: Logan. Fig. 49.
Fig. 39a, b. Cheumatopsyche rossi (Trichoptera: Hydropsychidae): distribution in
the Interior Highlands (39a); cladistic and biogeographic relationships of the C. pettiti
lineage (39b), C. smith (square); C. logani (closed circles); C. rossi (triangle); C. pettiti
(open circles).
Fig. 40a, b. Ceratopsyche piatrix (Trichoptera: Hydropsychidae) distribution in the
Interior Highlands (40a); distribution of C. piatrix and its nearest relative (40b).
Fig. 41a, b. Paduniella nearctica (Trichoptera: Psychomyiidae) distribution in the
Interior Higlands (41a); world wide distribution of the genus Paduniella (41b).
Florida Entomologist 73(4)
' J 42 --
^ 42
04Nt ^
strota
shawnee
occidea
hamata
thyria
AR,OK
IL
OR
NC
CA
kiamichi
45b fenestra
ledra
carolina
/teddyi
December, 1990
558
Allen: Symposium-Origins SE Arthropod Fauna 559
LEPIDOPTERA
Papilionidae
Papilio joanae Heitzman 1973:2 Fig. 50.
DISCUSSION
Sixty-eight species of insects are suspected to be endemic to the Interior Highlands
of North America. Table 1 lists these species and summarizes the available information
about close relatives occurring in eastern North America. Seven species have affinities
with western taxa. The nearest relatives of 4 species occur outside of North America,
and the close relative of 1 species is a boreal form. We do not know the close relatives
of 18 of the Interior Highland endemic species.
An analysis of the geographic patterns of the Interior Highland endemics using
taxon/area cladograms is difficult. The pattern that emerges is complex and not clearly
defined. Several factors obscure the biogeographic pattern including the following: (1)
the close relatives of the Interior Highland endemics occur in many different geographi-
cal areas in eastern and western North America; (2) the lack of cladograms for most of
the Interior Highland endemics; (3) the detailed geological history of small individual
areas in eastern North America is unknown in many cases. Nevertheless it may be
instructive for future workers to scrutinize as closely as possible the data that is avail-
able.
Using a method proposed by Allen (1983), area cladograms are changed into a linear
form (Fig. 51b) so that repetitious patterns may be discerned more readily and corre-
lated with possible vicariant events. This linear form was proposed because taxa seldom,
if ever, have relatives present in all the geographical areas under consideration. How-
ever, the cladistic/biogeographic relationships of the taxa and areas in which members
do occur may remain the same. For example, gven three taxa A, B, C that may or may
not have members in areas a, b, c, d, e, f, it is possible to detect a repetitious pattern
using linear relationships, Figure 51.
Using Figure 51 as a guide, a brief explanation may be helpful. Taxon A has mem-
bers in areas a, c, d, and f, but not in areas b and c. Taxon B has members in three of
the areas, b, e, and f, but not in a, c, and d. And taxon C has members in all of the
areas except c. An important point to note is that the cladistic/area relationships remain
the same in all of the taxon/area cladograms. That is, areas a and b share a common
node (a common ancestor) in taxon A and taxon C and, in turn, the a, b, lineage shares
a common node with d in both taxa. In taxon B, even though areas a, c, and d are
missing the ancestor/descendent relationships of the areas present, b, e, and f, are the
same as those elements (b, e, and f) in taxon A. This sharing of common nodes (or
Fig. 42a, b. Wormalida strota (Trichoptera: Philopotamidae) distribution in the In-
terior Highlands (42a); cladistic and biogeographic relationships of the lineage containing
W. strot (42b).
Fig. 43. Agapetus artesus (circle) and A. illini (squares) (Trichoptera: Glos-
sosomatidae) distribution in the Interior Highlands.
Fig. 44. Agapetus medicus (Trichoptera: Glossosomatidae) distribution in the In-
terior Highlands.
Fig. 45a, b. Distribution of endemic Rhyacophila (Trichoptera: Rhyacophilidae)
species in the Interior Highlands: (R. fenestra (squares); R. kiamichi (circles) (45a);
cladistic relationships of members of Branch 6 of the genus Rhyacophila (45b); see text
for distribution data.
Florida Entomologist 73(4)
limnella
mexicana
orealis
560
December, 1990
Allen: Symposium-Origins SE Arthropod Fauna 561
common ancestors), which is based on the principles of Hennigian phylogenetic systema-
tics, is what makes it possible to detect common patterns as shown in Figure 51d.
If the pattern shown in Figure 51d was not present, then the taxon/area cladograms
would have different ancestor/descendent relationships. For example, if we add taxon
D, Figure 51e, with a different set of ancestor/descendent relationships it is obvious
that taxon D belongs to a different pattern.
It should also be noted that terminal taxa (nodes) can be rotated 180 degrees such
that the taxon/area cladograms can be written a, b, or b, a. Such a rotation does not
affect the relationships of the taxa or the areas. Other nodes in the cladogram can also
be rotated.
It might be argued that rotating the cladogram at different nodes changes the pat-
tern. This is not true because the ancestor/descendent relationships at each node re-
mains the same regardless of how the nodes may be rotated. For example, in taxon C
rotating the a, b, d lineage so that it fits between e and f, Figure 51f, does not make f
and a closer relatives. Area a still shares its closest relationship with b followed by d,
followed by e, and then f. If such a rotation were done for taxon C, the same rotation
could be done for the two other taxa A and B. The results would be the same area
relationship pattern shown in Figure 51d, but written in a different sequence.
When using these taxon/area/linear diagrams it is important that the original data
be presented so that other workers will be able to accurately and independently assess
the validity of the conclusion, i. e. the biogeographic patterns.
Basically there appear to be three separate groups of species relationships: two
species with distant Asian affinities, Derops divalis and Ephemerella (Dannella) pro-
vonshai; several caddisfly lineages and the ground beetle subgenus Nomaretus with
distinct western North American relationships (Figures 52 through 57); and the major-
ity of the lineages with a complex eastern North American relationships (Figures 58
through 66).
The insect lineages with endemic Interior Highland species that appear to have
distinct western affinities are listed and shown in Figures 52 through 57. In these
lineages we see five possible areas of endemism having relationships with one another
(Figure 65): areas in western North America (NAw), the Interior Highlands (IH), areas
east of the Appalachian Mountains (app,e), several restricted areas west of the Appalac-
hian Mountains (app,w), western North American repeated (NAw), and widespread
taxa in North America between the western edge of the Appalachians and the Missis-
sippi River Valley (NAe,ws). One possible explanation for this pattern might be that
Fig. 46. Distribution of some endemic species of caddisflies (Trichoptera: Brachycen-
tridae and Leptoceridae) in the Interior Highlands: Micrasema ozarkana (Trichoptera:
Brachycentridae) (diamonds); Ceraclea nepha (Trichoptera: Leptoceridae) (circles);
Setodes oxapia (Trichoptera: Leptoceridae) (crosses); Trianodes smith (Trichoptera:
Leptoceridae) (square).
Fig. 47a, b. Helicopsyche limnella (Trichoptera: Helicopsychidae) distribution in the
Interior Highlands (47a); cladistic relationships of species closely related to H. limnella
(47b); see text for discussion of distribution data.
Fig. 48. Distribution of some endemic species of micro-caddisflies (Trichoptera: Hy-
droptilidae) in the Interior Highlands: Ochrotrichia anisca (closed circles); 0. contorta
(open circles); 0. unio (closed squares); 0. weddleae (closed diamonds); 0. edalis (open
square); Neotrichia kitae (cross).
Fig. 49. Paucicalcaria ozarkensis (Trichoptera: Hydroptilidae) distribution in the
Interior Highlands.
Fig. 50. Papilio joanae (Lepidoptera: Papilionidae) distribution in the Interior High-
lands.
Florida Entomologist 73(4)
December, 1990
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Florida Entomologist 73(4)
51a
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Taxon C
see text
a/d/-/f/-/b
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Fig. 51a, b, c, d. A hypothetical set of taxon/area cladograms and their linear equi-
valents; see text for discussion.
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b Taxon B
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564
December, 1990
Allen: Symposium-Origins SE Arthropod Fauna
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Florida Entomologist 73(4)
Allocapnia jeanae
Arianops sanderoni
Allocapnia malverna
Allocapnia vir./frisoni
Allocapnia vir./unzickeri
Allocapnia illinoensis
Evarthrus whitcombi
Ephemerella provonshai
Zealeutra warren
app/NAe,ws/IH/app,w/cp/app/cp/IH/cp/NAe,ws/MV,w
app/NAe,ws/IH/app,w/--/---/--/IH/--/------ /-
app/------/IH/app,w/--/app//--/--/--/ ---/--
---/--- /--/app,w/--/app/--/IH/--/- -----
---/------/I/app,w/--/---/--/--/--/NAe,ws/-
---/-----/IH/app,w/--/---/--/--/--/NAe,ws/--
---/----/--/app,w/--/---/--/IH/--/NAe,ws/----
---/------/--/app,w/cp/app/cp/IH/--/---- /MV,w
-------/IH/app,w/--/- -/--/--/--/---- ----
---/----/--/-----/--/---/--/IH/--/NAe,ws/MV,w
66
Fig. 65. The principal areas of endemism in North America showing relationships
with endemic species in the Interior Highlands.
Fig. 66. The general distribution pattern of lineages with endemic Interior Highland
representatives and affinities with eastern North America.
widespread, cosmopolitan species were first divided into eastern and western popula-
tions during the Cretaceous (100 mya) or possibly in the early Tertiary (60 mya). A
second division into eastern and western groups could have occurred during the Miocene
(12-25 mya) (Figure 57). This possibility is in contrast to explanations offered by Ross
(1956) suggesting a Pleistocene (1 mya) origin of the Interior Highland endemics.
Whether an old (Cretaceous, Miocene) vicariance pattern or a relatively young (Pleis-
tocene) explanation is correct will have to be settled by future research.
Turning to those insect lineages whose evolution occurred mostly in eastern North
America (Figures 58-64), we see a complex of biogeographic relationships. We cannot
hope to corroborate this pattern until many additional cladograms for other taxa are
established, and more is known about the geological history of eastern North America
and about areas of endemism within eastern North America. There are, however, a
number of interesting points that can be considered.
It is noted that most of the Interior Highland endemic insect species have close
relationships with species lying east of the Mississippi River Valley and west of the
December, 1990
566
Allen: Symposium-Origins SE Arthropod Fauna 567
Appalachian Mountains (app,w), (Figure 66). It is also noted that there were apparently
only two vicariance events that gave rise to Interior Highland endemics since this area
of endemism is repeated twice in the linear sequence (Figure 66). Whether either or
both of the events that isolated Interior Highland populations occurred during the
Pleistocene is unclear. Such Pleistocene explanations have been offered by several work-
ers as cited in the preceding text. We may also note that none of the lineages seem to
have close or distantly related sister groups east of the Appalachian Mountains as we
saw in the caddisfly and ground beetle taxa previously discussed.
Based on the cladistic/biogeographic data presented here and after a study of the
geological literature it is not possible to suggest specific earth events occurring in
eastern North America that may be correlated with the taxon/area/linear cladograms.
It is suggested, however, that from the meager data available, a tentative biogeographic
pattern is emerging (Figure 66). A great deal of additional work will be necessary
before this pattern can be corroborated, rejected, or modified.
END NOTE
Distribution ranges shown for eastern North America are approximate. The reader
is advised to consult the original citations for greater detail.
ACKNOWLEDGMENTS
I wish to thank Dr. C. E. Carlton and Dr. J. R. Phillips (University of Arkansas),
two anonymous reviewers, Dr. John Morse (Clemson University) and Susan Allen for
their thoughtful and helpful comments. Ms. Sherri Montani and Cecile Ueltschey typed
and corrected the manuscript. The Entomology Department, University of Arkansas,
the Arkansas Nature Conservancy, the Arkansas Natural Heritage Commission, and
the U. S. Forest Service, Ozark-St. Francis National Forest have provided support for
this work.
Published with the approval of the Director, Arkansas Agricultural Experiment
Station, University of Arkansas, Fayetteville, AR 72701.
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569
570 Florida Entomologist 73(4) December, 1990
BIOGEOGRAPHIC AFFINITIES OF PSELAPHID BEETLES
OF THE EASTERN UNITED STATES
CHRISTOPHER E. CARLTON
Department of Entomology
University of Arkansas
Fayetteville, AR 72701
ABSTRACT
The family Pselaphidae in eastern North America consists of 67 genera and approx-
imately 352 species. Thirty genera are endemic to the region. Sixteen genera belong to
two generalized tracks of Laurasian ancestry. One of these is a poorly defined Holarctic
Track, the other a well defined Eastern Nearctic-European Track. Thirteen genera
belong to two Nearctic-Neotropical Tracks. The first extends from eastern North
America through Mexico to Central America. The second overlaps the first and also
extends from Central America and Mexico up the West Coast of the United States to
southwestern Canada. Biogeographic affinities were not determined for 34 genera,
primarily because of a lack of information about sister group relationships at either
generic or specific levels.
RESUME
La familiar PselAfida del este de Norte Am6rica consiste de 67 generos y ap-
roximadamente 352 species. Treinta g6neros son endemicos a la region. Dieciseis g6n-
ros pertenecen a dos trazas generalizadas de antecedentes Laurasianos. Uno de estos
es una traza HolArtica pobremente definida, el otro es una traza del este NeArtico-
Europeo muy bien definida. Trecce generos pertenecen a dos trazas Neirticas-Neot-
ropicales. La primera se extiende del este de Norte America a traves de Mexico hacia
Centro Am6rica. La segunda sobrelapa la primera y tambien se extiende desde centro
America y M6xico hacia la costa occidental de los Estados Unidos y el suroeste de
CanadA. No se determinaron las afinidades biogeograficas de 34 g6neros principalmente
por falta de informaci6n sobre relaciones de grupos hermanos a niveles gendricos o
especificos.
The Pselaphidae is a large (over 8000 species) family of small (0.5-5.0 mm in length)
beetles with close phylogenetic ties to the Staphylinidae. Pselaphids are distinguished
from staphylinids primarily by their more heavily sclerotized and less flexible integu-
ment combined with great proliferation of integumental foveae, especially on the
thoracic sterna. The vast majority of pselaphid species inhabit moist forest litter, and
a large number are facultative or habitual associates of ants or, less frequently, ter-
mites. With the exception of a few specialized myrmecophilous taxa, pselaphids are
active predators with large sickle-shaped mandibles. Their developmental stages are
poorly known. Park (1942) provided a comprehensive treatment of pselaphid morphol-
ogy and biology, and Newton and Chandler (1989) presented a classification of the
family at the generic level.
This paper is a first step in reconstructing the biogeographic history of the
Pselaphidae in the eastern United States. The distribution of most species of pselaphids
in the eastern United States is poorly known and it is premature to attempt a detailed
biogeographic analysis at the species level for the family as a whole. But, the generic
Carlton: Symposium-Origins SE Arthropod Fauna 571
distributional limits are reasonably well recognized, so I have concentrated my efforts
at that taxonomic level.
PREVIOUS WORK
Park (1942) discussed the biogeography of Western Hemisphere pselaphids in a
framework of tectonic stability. He relied primarily on ecological cycles and dispersal
to explain the distribution of the group. Jeannel (1948, 1950) allowed that the continents
were not fixed in their present positions, and discussed the origins of the Eastern
Hemisphere pselaphid fauna from a methodological perspective that approached vic-
ariance. In the only series of papers dealing exclusively with pselaphid distributions,
Reichle (1966, 1969) examined the unique pselaphid fauna of postglacial subboreal bogs
in the Great Lakes Region and offered some ideas about their origins as glacial refugia.
Barr (1974), as part of his revision of Arianops discussed the distributions of species
groups and topographic features that isolate them.
METHODOLOGY
My analytical approach is essentially that of a cladistic biogeographer: Similar pat-
terns of phyletic and geographic relationships are identified and depicted as generalized
tracks and area cladograms; these data are then compared with the known topographic
and geological histories of the regions concerned. The result is a synthetic hypothesis
comprising the sequence of isolating events that led to the independent evolution of the
various lineages (see Nelson & Platnick 1981, and references cited therein). Following
a general summary of pselaphid distributional patterns in the eastern United States, I
have assigned genera to four continental-scale generalized tracks. In the accompanying
figures, the thickest track lines represent taxa containing several common, widespread
species. Lines of intermediate thickness, or short lines, represent species of limited
distribution. The thinnest lines indicate disjunctions between related taxa. Unique
biological and distributional features of taxa within these tracks are discussed using
specific examples when available, and some exemplary cladograms are presented. Ten-
tative explanations are offered about the vicariant origins of some of the tracks. Finally,
I have grouped the genera that I was unable to assign to tracks and discuss some
problems peculiar to each group.
In practice my attempts to allocate many genera to specific tracks were thwarted
by inadequate data, particularly the lack of cladograms, or any form of phylogenetic
arrangements of related genera. But, this is a common dilemma and should not stop
one from taking the first step. I have relied heavily upon some less than reliable meas-
urements of phylogenetic relationships. I discuss genera as if they were monophyletic
units, which in many cases is probably true, and I have based some of my conclusions
on my own judgment about relationships.
DISTRIBUTIONAL OVERVIEW
The distribution of pselaphids of the eastern United States can be summarized at
three taxonomic levels (Table 1). The high levels of endemism at both generic and
specific levels is characteristic of substrate arthropod taxa such as pselaphids and
suggest that these groups are distributionally sedentary. Such groups are regarded by
some cladistic biogeographers as relatively more reliable in biogeographic reconstruc-
tion than more vagile taxa (see Noonan, 1988 for a relevant discussion). Figures for
introduced taxa are based on one rather well documented case involving the genus
Trichonyx (Park 1953, Cooper 1961), plus more circumstantial cases involving two
species of Euplectus (Wagner 1975).
Florida Entomologist 73(4)
TABLE 1. DISTRIBUTIONAL SUMMARY OF PSELAPHID BEETLES OF THE EASTERN
UNITED STATES.
Tribes Genera Species
Total 18 67 352
Worldwide 8 12 0
Holarctic 4 3 3
Nearctic-Neotropical 1 13 under 10
Endemic Nearctic 31 8 90%2
Endemic E. U.S. 31 30 80%2
Introduced 2 2 3
'Figures overlap
'Estimates
Holarctic Track (Figs. 1, 2)
The genera that I have assigned to the Holarctic Track typically have species scat-
tered throughout the eastern United States and extreme western North America, espe-
cially moist forests from northern California to British Columbia. They also have con-
Europe
Mayetia Mulsant and Rey Branhyglutn Thomson
Bibloporus Thomson Euboarhexius Grigarick
Batriandae Reitter and Schuster*
Rybaxil Saulcy* Piloplus Casey
'endemic to eastern United States
Fig. 1. Holarctic Track and pselaphid genera assigned to it.
December, 1990
Carlton: Symposium-Origins SE Arthropod Fauna 573
Western Europe E- or NW U.S.
Trocaster Trogasteropala Eubarhexlu
Fig. 2. Phylogeny and distribution of Euboarhexius and related genera (Carlton,
unpublished data, see Carlton and Allen, 1986).
generic species or, more often, related genera in either Europe or northeastern Asia.
The northeastern Asian connection is perhaps not as clearly defined as it should be
because of nomenclatural misalignments. I have included Brachygluta and Pilopius in
this track because, though they contain a small number of tropical species, they are
predominantly Holarctic in distribution.
The phylogeny of Euboarhexius and related genera (Fig. 2) does not include taxa
inhabiting regions comprising the complete Holarctic Track, but it is an interesting
example. Euboarhexius contains four species on opposite sides of North America, two
in the east and two in the west (see Carlton & Allen 1986). The four have not been
cladistically resolved and their apparent phenetic relationships run counter to what
appears logical from a biogeographic perspective. It suggests an east-west, east-west
relationship, not the expected east-east, west-west relationship.
This track appears to be Laurasian in origin, but of the relative sequence of such
events as epicontinental seaways, rain shadows, and Beringean connections I cannot
comment.
Southeastern United States-European Track (Figs. 3, 4)
This track includes certain genera in eastern North America and Europe, which are
concentrated in mountainous regions of both continents. This track is related to the
preceding in that it is of Laurasian origin. But, in contrast to the Holarctic Track it is
well defined, not only in terms of distributional and phylogenetic relationships, but by
certain common biological attributes of the included genera. The most outstanding dis-
tributional feature is the high degree of endemism within the track, and the concentra-
tion of forms in the mountains, where the species commonly occupy restricted and
largely allopatric ranges. Varying degrees of eye and wing loss are common in members
of this track, and many species are hypogean.
The origin of this tract may be traced to ancestral lineages that evolved in montane
habitats along the Ouachita-Appalachian-Hercynian Thrust Belts during middle Jurassic
to early Cretaceous time. The phylogeny of genera of the tribe Amauropsini (Fig. 4)
Florida Entomologist 73(4)
Eutyphlus LeConte*
Batriasymmodea Park'
Texamaurops Barr
and Steeves*
Arianops Brendel*
Nearctitychua Chandler'
Tychobythinua Ganglbauer
Machaeroden Brendel*
Spelechua Park*
Subterrochua Park*
*endemic to eastern United States
Fig. 3. Southeastern United States-European Track and pselaphid genera assigned
to it.
suggests independent divergence of European and North American lineages following
their isolation, which resulted from the Cretaceous vicariance of mountain ranges link-
ing North America and Europe (see Burke 1976 and Ziegler et al. 1983). The isolation
of three species of Arianops in Arkansas and Oklahoma is considered by Carlton and
Cox (1990) to be the result of the formation of the Mississippi Embayment during late
Cretaceous.
Nearctic-Neotropical Tracks (Figs. 5, 6, 7, 8)
Two major tracks linking the eastern pselaphid fauna with taxa in Mexico, Central
America, and, in some cases, South America can be identified. The first track (Fig. 5)
is confined to the east, the second (Fig. 6) comprises the western United States and,
for some taxa, southwestern Canada.
Distributional patterns within these tracks are well defined. Only a small number
of the genera involved are endemic to eastern North America. Typically, North Amer-
574
December, 1990
Carlton: Symposium-Origins SE Arthropod Fauna
575
Europe E United Statese
other
Ouachlta amplyoponica species
Amaurops Paramaurop lineage lineage groups
series series
Arlanps
Fig. 4. Phylogeny and distribution of the tribe Amauropsini (Carlton, unpublished
data).
ican species represent the northern-most members of widespread Neotropical genera.
Also, for the majority of genera within these tracks, at least one of the North American
representatives is a common, widespread form that occupies a variety of habitats. A
few southeastern genera, such as Eupsenius and Lemelba have closely related species
or even share species with the West Indies.
Phylogenetic hypotheses of Eutrichites (Fig. 7) and of Conoplectus and related gen-
era (Fig. 8) illustrate both tracks. Eutrichites consists of one common, widespread
eastern species, E. zonatus, which is the terminus of a clade that extends south to at
least northern Venezuela (Carlton 1989). A similar pattern is shown by Conoplectus.
In addition to the widespread C. canalicalatus, several species with more restricted
distributions occur in the southeastern United States (Carlton 1983). The western track
lineage is represented by the sister genera Rhexidius and Oropus, and the eastern
pattern is repeated at the base of the phylogeny by the Eurhexius-Rhexius lineage.
The development of these tracks is most probably related to the Cretaceous-mid
Tertiary orogenic cycles of the Laramide Revolution and the effect it had in fragmenting
forest corridors throughout North America and the northern Neotropics, as well as the
inundation of a large section of the midcontinent during much of this time. A less
informative alternative is that many of the taxa with Neotropical affinities in eastern
North America are the result of dispersal following the mid to late Tertiary consolidation
of Central America and South America. A dispersal component seems inescapable when
dealing with taxa that extend into the South Anierican mainland.
East-West Nearctic Genera
Sonoma Casey Lucifotychus Park and Wagner
Actizona Chandler Mipseltyrus Park
Actium Casey Adranes LeConte
Actiastes Casey
Florida Entomologist 73(4)
Triminplectua Brendel*
Trimlomrelba Casey'
Lemelha Park
Thelium Casey
Rhinoacepila LeConte
*endemic to eastern United Stati
Rhxeius LeConte
Anchylarthron Brendel
Eupsenlus LeConte
Cirnocerus Motschulsky
Fig. 5. Eastern Nearctic-Neotropical Track and pselaphid genera assigned to it.
The genera listed here do not represent a generalized track, but are simply those
which have species distributed in both the Eastern and Western United States. Most
of these genera are found in mountainous regions and several, including Actizona and
possibly Lucifotychus are subboreal. I have not been able to reach any conclusions
about sister group relationships for these genera.
Inconclusive Endemic Genera
Pycnoplectus Casey Briaraxis Brendel
Leptoplectus Casey Custotychus Park and Wagner
Acolonia Casey Cylindrarctus Schaufuss
Racemia Newton & Chandler Ouachitychus Chandler
Ramecia Casey Speleobama Park
Trigonoplectus Bowman Prespelea Park
576
December, 1990
Carlton: Symposium-Origins SE Arthropod Fauna
Dalmosella Casey
Pygmactium Schuster & Grigarick
Pseudactium Casey
Nisaxis Casey
Ceophyllus LeConte
Cedius Le Conte
Atinus Horn
Unfortunately, a majority of the pselaphid genera endemic to eastern North America
have not contributed any useful information to this analysis of biogeographic affinities.
In most cases, this is the result of my inability to reach any reliably informed decisions
about the sister group relationships of the genera. Thus, the endemics stand alone,
without biogeographic perspective. In several cases, an endemic genus is sympatric
with its sister genus. Such a situation is biogeographically uninformative, because no
historical isolating mechanism may be proposed without an additional assumed dispersal
event.
Worldwide Genera
Euplectus Leach Bibloplectus Reitter
Thesiastes Casey Tmesiphorus LeConte
Reichenbachia Leach Fustiger LeConte
Only one of these genera, Reichenbachia has what appears to be a truly worldwide
distribution. But, the remainder have species in far-flung places around the world and
Cnnnplectus Brendel Eutrichita LeConte
Decarthron Brendel Hamotus Aube
Fig. 6. Nearctic-Neotropical Track and pselaphid genera assigned to it.
577
Florida Entomologist 73(4)
Cent. America o Mexlco-+SW U.S.-- E U.S.
Scalenarthrus arlzonenals zonatus
XYbarldM parral
Sveracruzenal
busckI
V funiculls Vr
Eutrichites
Fig. 7. Phylogeny and distribution of Eutrichites and related genera (from Carlton,
1989).
so are listed here collectively as "worldwide". Again, I have not attempted to draw any
biogeographic conclusions from the distributions of these genera, but I believe many of
them have the potential to be tremendously informative, that information lying at the
species level. Careful taxonomic and cladistic analyses of these genera may reveal a
wealth of information concerning biogeographic patterns on a worldwide scale.
This preliminary discussion of the biogeographic affinities of the pselaphid fauna of
eastern North America points out that, although the distributions and relationships of
Florida
Mexico-* Calif.-* NW U.S.
Cent.
nConplectus
Fig. 8. Phylogeny and distribution of Conoplectus and related genera (from Carlton,
1983).
578
December, 1990
Carlton: Symposium-Origins SE Arthropod Fauna 579
litter inhabiting arthropods is still poorly known, hypotheses concerning the biogeog-
raphic patterns and their origins can be proposed based on the amount of available data.
These preliminary analyses open up new avenues of inquiry and provide comparative
data against which the bioeographic relationships of other groups may be measured.
ACKNOWLEDGMENTS
I thank Drs. Robert T. Allen, John C. Morse, William C. Yearian, and two anony-
mous reviewers for offering comments and suggestions that substantially improved the
quality of this manuscript.
Published with the approval of the Director, Arkansas Agricultural Experiment
Station, Fayetteville, AR 72701
REFERENCES CITED
BARR, T. C., JR. 1974. The eyeless beetles of the genus Arianops Brendel (Coleopt-
era:Pselaphidae). Bull. Am. Mus. Nat. Hist. 154: 1-51.
BURKE, K. 1976. Development of graben associated with the initial ruptures of the
Atlantic Ocean. Tectonophysics 36: 83-110.
CARLTON, C. E. 1983. Revision of the genus Conoplectus (Coleoptera:Pselaphidae).
Coleops. Bull. 37: 55-80.
CARLTON, C. E. 1989. Revision of the genus Eutrichites (Coleoptera:Pselaphidae).
Coleops. Bull. 43: 105-119.
CARLTON, C. E., AND R. T. ALLEN. 1986. Revision of the genus Euboarhexius
(Coleoptera:Pselaphidae). Coleops. Bull. 40: 285-296.
CARLTON, C. E., AND R. T. COX. 1990. A new species of Arianops from central
Arkansas and the biogeographic implications of the Interior Highlands Arianops
species (Coleoptera:Pselaphidae). Coleops. Bull. 44: 365-371.
COOPER, K. W. 1961. Occurrence of the European beetle Trichonyx sulcicollis
(Reichenbach) in New York state. Entomol. News 72: 90-92.
JEANNEL, R. 1948. Revision des Amaurops et genres voisins (Pselaphidae). Rev.
Francaise d'Ent. 15: 1-19.
JEANNEL, R. 1950. Faune de France, 53. Coleopteres pselaphides. Paris,
Lechevalier, iv+421 pp.
NELSON, G., AND N. PLATNICK. 1981. Systematics and biogeography. Columbia
University Press. xi + 567 pp.
NEWTON, A. F., JR., AND D. S. CHANDLER. 1989. World catalog of the genera of
Pselaphidae (Coleoptera). Fieldiana (Zoology) 53: iv + 93 pp.
NOONAN, G. R. 1988. Biogeography of North American and Mexican insects and a
critique of vicariance biogeography. Syst. Zool. 37: 366-384.
PARK, 0. 1942. A study of Neotropical Pselaphidae. Northwestern Univ. Stud. Biol.
Sci. Med. 1: ix+403 pp.
PARK, 0. 1953. A European pselaphid beetle collected in New York Natur. Hist.
Misc. 117: 1-3.
REICHLE, D. E. 1966. Some pselaphid beetles with boreal affinities and their distribu-
tions along the postglacial fringe. Syst. Zool. 15: 330-334.
REICHLE, D. E. 1969. Distribution and abundance of bog-inhabiting pselaphid beetles.
Trans. Ill. Acad. Sci. 62: 233-264.
WAGNER, J. A. 1975. Review of the genera Euplectus, Pycnoplectus, Leptoplectus,
and Acolonia including Nearctic species north of Mexico. Entomologica
Americana 49: 125-207.
ZIEGLER, A. M., D. B. ROWLEY, A. L. LOTTES, D. L. SAHAGIAN, M. L. HULVER,
AND T. C. GIERLOWSKI. 1985. Paleogeographic interpretation: with an exam-
ple from the mid-Cretaceous. Ann. Rev. Earth Planet Sci. 13: 385-425.
580 Florida Entomologist 73(4) December, 1990
SOME APHIDS OF IMPORTANCE TO THE SOUTHEASTERN
UNITED STATES (HOMOPTERA: APHIDIDAE)
MANY B. STOETZEL
Systematic Entomology Laboratory
Agricultural Research Service
U.S. Department of Agriculture
Beltsville, MD 20705
ABSTRACT
Despite their importance as pests of ornamentals and crops and as transmitters of
plant viruses, aphids have been poorly studied and collected throughout the United
States. Lists are available for only 31 of the 50 states; and of those lists, only 3 have
been published since 1970. In the southeastern United States, published lists exist for
only Louisiana and North Carolina. Of the aphids discussed, Acyrthosiphon kondoi
Shinji, Aphis nerii Boyer de Fonscolombe, Brachycorynella asparagi (Mordvilko),
Idiopterus nephrelepidis Davis, Myzus varians Davidson, and Toxoptera aurantii
(Boyer de Fonscolombe) are reported to occur in some southeastern states and may
occur throughout the area. Toxoptera citricida (Kirkaldy) and Pterochloroides persicae
(Cholodkovsky) represent species not known to occur in the United States but which
have the potential of becoming serious pests if introduced. For each species there are
sections on distribution in the southeastern United States, hosts, taxonomic characteris-
tics of apterous and alate viviparae, and general information.
RESUME
A pesar de su importancia como plagas de plants ornamentales y de cultivos, y
como transmisores de viruses, los afidos han sido pobremente estudiados y coleccionados
en los Estados Unidos. Solamente hay listas de 31 de los 50 estados, y de esas listas,
solo 3 se han publicado a partir de 1970. En el sudeste de los Estados Unidos, solo
existen listas publicadas de Louisiana y de Carolina del Norte. De los afidos que se
discuten, Acyrthosiphon kondoi Shinji, Aphis merii Boyer de Fonscolombe,
Brachycorynella asparagi (Mordvilko), Idiopterus nephrelepidis Davis, Myzus varians
Davidson, y Toxoptera aurantii (Boyer de Fonscolombe), se report que ocurren en
algunos estados del sudeste y que pudieran ocurrir en todo el Area. Toxoptera citricida
(Kirlady) y Pterochloroides persicae (Cholodkovsky) representan species que no son
conocidas en los Estados Unidos pero que tienen el potential de convertirse en series
plagas si se introdujeran. Para cada especie hay secciones sobre su distribuci6n en el
sudeste de los Estados Unidos, hospederos, caracteristicas taxon6micas de viviparos
apteros y alados, e informaci6n general.
Despite the importance of aphids as pests of ornamentals and crops and as transmit-
ters of plant viruses, the family has been poorly studied and collected throughout the
United States. Smith & Parron (1978) reported that 1,380 species in 277 genera occurred
in the United States. However, lists are available for only 31 of the 50 states. All but
3 of these lists of aphids occurring in various states or regions were published before
1970 and are considerably out-of-date. In 1970, Leonard and Bissell published a list of
the aphids of Washington, DC, Maryland, and Virginia. In 1974, Leonard published a
list of the aphids of Oregon. In 1983, Knowlton published a list of the aphids of Utah.
For the purpose of this paper, the southeastern United States contains the states
of Alabama, Arkansas, Florida, Georgia. Louisiana, Mississippi, North Carolina, South
_ __I_~
h ~___
Stoetzel: Symposium-Origins SE Arthropod Fauna
Carolina, and Tennessee. A major difficulty in assessing the distribution of any category
of aphids in the southeastern United States is our lack of knowledge of what aphids are
represented in the region. Our knowledge of the aphids in the southeastern United
States is less than that for any of the other geographical regions in the United States.
In the southeastern United States, published lists exist for only Louisiana and North
Carolina. In 1938, Elliott published a list of 37 aphid species found in Louisiana. Boud-
reaux continued Elliott's study of aphids and in 1951 published a list containing 121
aphid names.
In Wray's 1967 publication on the insects of North Carolina, 46 aphid names were
listed. In the 1978 publication by Smith and Parron on the aphids of North America,
approximately 300 species were listed as occurring in North Carolina. Smith and Parron
have in preparation a manuscript on the aphids of North Carolina, and it contains
approximately 390 species and 5 subspecies in 122 genera (C. F. Smith, personal com-
munication).
In the USDA, Beltsville, MD, correspondence files, there is in the archived files of
Mortimer D. Leonard an unpublished listing of Florida aphids. The list contains 109
aphid names associated with 325 plant names.
All aphids are plant feeders and are potential pests because they suck sap from the
plants, because they transmit plant viruses, or because plant owners find their presence
offensive. As with many insects, interest in aphids focuses on those species that are
pests. Much of our knowledge of the movement and ultimate distribution of aphids
results from the reports made on pest species.
Of the aphids discussed, Acyrthosiphon kondoi Shinji, Aphis nerii Boyer de
Fonscolombe, Brachycorynella asparagi (Mordvilko), Idiopterus nephrelepidis Davis,
Myzus varians Davidson, and Toxoptera aurantii (Boyer de Fonscolombe) are reported
to occur in some southeastern states and may occur throughout the area. These species
were chosen because they are still dispersing within the southeastern United States,
because they are species that continue to go undetected, or because they can be misiden-
tified easily. Toxoptera citricida (Kirkaldy) and Pterochloroides persicae (Cholod-
kovsky) represent two of the many species not known to occur in the United States but
which have the potential of becoming serious pests if introduced. These last two species
are included here because they are expanding their ranges. For each species there are
sections on distribution in the southeastern United States, hosts, taxonomic characteris-
tics of apterous and alate viviparae, and general information.
Acyrthosiphon kondoi Shinji
DISTRIBUTION IN SOUTHEASTERN US. Arkansas, Georgia, Louisiana, and North
Carolina.
HOSTS. Within the family Leguminosae, found mainly on Medicago, Melilotus, and
Trifolium but also on Astragalus, Dorycnium, Lens, and Lotus.
TAXONOMIC CHARACTERISTICS. Apterous vivipara: In life bluish green. Small to
medium aphids (1.8-2.9 mm). Antennae 6-segmented, base of antennal segment VI
approximately equal to length of hind tarsal II; 1-2 secondary sensoria on antennal
segment III near base. Cornicles long, pale to slightly dusky. Cauda elongate with 3
(rarely 4) pairs of lateral setae and 1-2 preapical setae, pale.
Alate vivipara: In life bluish green with a dark brown thorax. Small to medium
aphids (1.5-2.8 mm). Antennae 6-segmented, base of antennal segment VI approxi-
mately equal to length of hind tarsal segment II; 6-11 secondary sensoria on antennal
segment III and usually confined to basal half of segment. Cornicles long, pale to slightly
dusky. Cauda elongate with 3 (rarely 4) pairs of lateral setae and 1-2 preapical setae,
pale.
582 Florida Entomologist 73(4) December, 1990
DISCUSSION. Acyrthosiphon kondoi, the blue alfalfa aphid, probably originated in
Asia. It was collected in Arizona, California, Idaho, Nevada, and Utah in 1975, in
Kansas and New Mexico in 1976, in Nebraska, Oklahoma, Oregon, and Wyoming in
1977, in Texas and Washington in 1978, in Missouri in 1979, in Georgia, Kentucky, and
Louisiana in 1983, in Arkansas and North Carolina in 1985, and in Iowa in 1989. This
aphid has become established as a pest of alfalfa throughout most of the alfalfa-growing
regions of the United States. It is known to be an early and late season pest in alfalfa
fields. Reports of "unusually early" problems with Acyrthosiphon pisum (Harris), the
pea aphid, in alfalfa fields probably are erroneous reports of damage by A. kondoi. The
shorter base of antennal segment VI and the stouter, fewer caudal setae will separate
A. kondoi from A. pisum whose antennae are banded at the segmental junctions.
Aphis nerii Boyer de Fonscolombe
DISTRIBUTION IN SOUTHEASTERN US. Florida, Georgia, Louisiana, and Missis-
sippi.
HOSTS. Found mainly on plants in the Asclepiadaceae especially Asclepias spp., and
Apocyanaceae, especially Nerium oleander.
TAXONOMIC CHARACTERISTICS. Apterous vivipara: In life with aposematic colora-
tion of bright yellow or yellowish-orange body with black cornicles and cauda and usually
black antennae and legs. Small to medium aphids (1.5-2 6 mm), rounded. Antennae
6-segmented; no secondary sensoria on antennal segment III. Cornicles long, black.
Cauda elongate, with 3 pairs of lateral setae and 4-5 preapical setae, black. Legs and
antennae long, black.
Alate vivipara: In life with aposematic coloration of bright yellow or yellowish-
orange body with black cornicles and cauda and usually black antennae and legs. Small
to medium aphids (1.5-2.6 mm), rounded. Antennae 6-segmented; secondary sensoria
6-13 on antennal segment III and 0-5 on IV. Cornicles long, black. Cauda elongate, with
3 pairs of lateral setae and 4-5 preapical setae, black. Legs and antennae long, black.
DISUSSION. Accepting that Myzus persicae (Sulzer), the green peach aphid, is the
most common of all the aphids now distributed throughout the southeastern US, then
the second most commonly found aphid is probably Aphis nerii, the oleander aphid,
because of the widespread occurence of oleander (Nerium oleander) and milkweeds
(Asclepias spp.) in the region.
Brachycorynella asparagi (Mordvilko)
DISTRIBUTION IN SOUTHEASTERN US. Alabama, Georgia, North Carolina, and
South Carolina.
HOSTS. Found on several species of Asparagus: densiflorus (Kunth) Jessop cv.
Sprengeri, officinalis (L.), and setaceus (Kunth) Jessop.
TAXONOMIC CHARACTERISTICS. Apterous vivipara: In life green or grey-green
and often covered with mealy wax. Small aphids (1.2-1.7 mm), convex or oval, elongate.
Antennae 6-segmented, unguis less than twice length of base of antennal segment VI;
no secondary sensoria on antennal segment III. Cornicles short and truncate, porelike,
about as long as wide, pale. Cauda elongate, with 3-4 pairs of lateral setae and 1
preapical seta, pale to almost white. Legs and antennae rather short and held close to
body.
Alate vivipara: In life green and sometimes covered with mealy wax. Small aphids
(1.2-1.7 mm), convex, elongate. Antennae 6-segmented, unguis less than twice length
of base of antennal segment VI; 7-10 secondary sensoria, varying in size with the smal-
lest half the size of the largest, on antennal segment III. Cornicles short and truncate,
Stoetzel: Symposium-Origins SE Arthropod Fauna
porelike, about as long as wide, pale. Cauda elongate, usually with 3-4 pairs of lateral
setae and 1 preapical seta, pale to almost white.
DISCUSSION. Brachycorynella asparagi, the asparagus aphid, originated in eastern
Europe and is not really tropical in origin but is included here to call attention to its
probable presence throughout the southeastern United States. This aphid was collected
first in 1969 in the United States in New York and New Jersey on plants of Asparagus
officinalis. By the end of 1973, B. asparagi had been collected in most of the states
along the eastern seaboard from Massachusetts south to North Carolina. This aphid
was collected in pan traps in Illinois in 1977 and on asparagus in Missouri and
Washington in 1979, in Alabama, Georgia, Indiana, and Michigan in 1980, in Idaho,
Ohio, Oklahoma, and Oregon in 1981, in California and North Dakota in 1984, and in
South Carolina in 1985. The aphid can probably be found on asparagus in every state,
but it remains undetected because the characteristic distortion of terminals called a
"witch's broom" occurs on the negative growth long after the gardener has quit cutting
spears.
Idiopterus nephrelepidis Davis
DISTRIBUTION IN SOUTHEASTERN US. Florida and North Carolina.
HOSTS. Found in greenhouses on various kinds of ferns, especially Adiantum spp.
TAXONOMIC CHARACTERISTICS. Apterous vivipara: In life body dull black with
dorsal setae pale; eyes red; and antennae and legs almost a colorless white. Small aphids
(1.0-1.5 mm), pyriform. Antennae 6-segmented, beyond basal segments colorless with
dark apices; 1-5 secondary sensoria on antennal segment III. Cornicles elongate, cylin-
drical and rugose, basal half black and apical half white. Cauda short, acuminate, with
2 pairs of lateral setae and 1 dorsal preapical seta, black. Legs almost colorless with
tarsi dusky.
Alate vivipara: In life body dull black with dorsal setae pale; eyes red; antennae and
legs almost a colorless white. Small aphids (1.3-1.7 mm). Antennae 6-segmented, beyond
basal segments colorless with dark apices; secondary sensoria 7-14 in a line on antenna
segment III, 0-5 in a line on IV, rarely 1-2 on V. Body with several short, capitate hairs
on each dorsal abdominal segment. Cornicles elongate, cylindrical and rugose, basal half
black and apical half white. Cauda short, acuminate, with 2 pairs of lateral setae and 1
dorsal preapical seta, black. Legs almost colorless with tarsi dusky. Wing veins black,
and bordered; radial sector touches media.
DISCUSSION. Neotropical in origin, I. nephrelepidis can be quite common on ferns
in the greenhouse, yet it often goes undetected despite the fact that it prefers young
fronds and heavy populations cause the fronds to turn brown. This aphid probably
occurs in greenhouses throughout the southeastern United States but remains the least
collected tropical aphid in this and other areas.
Myzus varians Davidson
DISTRIBUTION IN SOUTHEASTERN US. Florida and North Carolina.
HOSTS. Clematis spp. in the United States. Primary host is Prunus persica in Asia,
but it is not known to occur on peach in the United States
TAXONOMIC CHARACTERISTICS. Apterous vivipara: In life yellow with antennae
with black bands on apical portion of all segments; apical portion of cornicles black.
Small aphids (1.3-1.8 mm). Antennae 6-segmented; no secondary sensoria on antenna
segment III. Cornicles long and rugose; apical, black portion approximately equal to
length of hind tarsal II. Cauda elongate, with 3-4 pairs of lateral setae, dusky. Legs
with tarsi dusky.
583
Florida Entomologist 73(4)
Alate vivipara: In life yellow with antennae and cornicles dusky. Small aphids (1.3-
1.7 mm). Antennae 6-segmented; 8-16 secondary sensoria in a line on antennal segment
III. Cornicles long and rugose, dusky. Cauda elongate, with 3-4 pairs of lateral setae,
pale to dusky. Legs with tarsi dusky. Abdomen with dusky dorsal patch on segments
III-V.
DISCUSSION. Myzus varians was first described from specimens collected on
Clematis in California in 1911 (Davidson 1912), but the aphid is probably Asian in origin.
It was collected in Florida in 1969. Smith & Parron (1978) reported it from North
Carolina. In 1989, specimens collected on Clematis in Maryland were brought to me for
identification; and I subsequently collected it on this host in two other locations in
Maryland. While there are slides of M. varians collected on peach in California and on
Prunus sp. in Hawaii, the species is not reported to be a pest of peach in the United
States.
This aphid is probably more widely distributed throughout the United States, but
it goes undetected because homeowners ignore the curled leaves of their Clematis
vines. Because of its similarity to the ubiquitous Myzus persicae (Sulzer), it probably
has been collected but misidentified as that species. The longer and less clavate corni-
cles, that have dark tips in the apterae and are dusky in the alatae, and the distinctively
banded antennae of the apterae easily separate M. varians from M. persicae.
Toxoptera aurantii (Boyer de Fonscolombe)
DISTRIBUTION IN SOUTHEASTERN US. Florida, Louisiana, and North Carolina.
HOSTS. Has been reported on a wide range of host plants. It occurs on various
genera in the Rutaceae and is the best known of the Citrus aphids. It is also a pest of
cacao, coffee, gardenia, magnolia, tea, etc.
TAXONOMIC CHARACTERISTICS. Apterous vivipara: In life shiny, brownish-black
or black. Small aphids (1.1-2.0 mm), oval. Antennae 6-segmented; no secondary sensoria
on antennal segment III; segments white with black apical bands on all segments.
Cornicles elongate, black. Cauda elongate, with 9-26 setae, black.
Alate vivipara: In life shiny, brownish-black or black. Small aphids (1.1-2.0 mm),
oval. Antennae 6-segmented; secondary sensoria 2-8 on antennal segment III and none
on IV; segments white with black apical bands on all segments. Cornicles elongate,
black. Cauda elongate, with 8-16 setae, black. Forewing with a black pterostigma and
usually a once-branched media.
DISCUSSION. Toxoptera aurantii, the black citrus aphid, probably originated in
New Zealand. All species in this genus are unique in having a stridulatory apparatus.
When ridges on the venter of the abdomen are rubbed with conical setae on the hind
tibiae, a sound is produced. This species is distributed throughout the tropics and sub-
tropics and is known in the United States from Florida, Louisiana, Texas, and California
north as far as Maryland, New York, and Oregon. It is a vector of several plant viruses
including Citrus tristeza virus. This species can be confused with Toxoptera citricida
(Kirkaldy), but the banded antennae of T. aurantii, the dark antennal segment III of
the alatae of T. citricida, and the differences in the number of caudal setae will separate
the two.
Toxoptera citricida (Kirkaldy)
DISTRIBUTION IN SOUTHEASTERN US. Not known to occur in the United States.
The species has a high potential of being introduced into the United States and is
included here because it occurs in South America and because of the trade in rutaceous
fruits between South and North America.
584
December, 1990
Stoetzel: Symposium-Origins SE Arthropod Fauna
HOSTS. Limited largely to Rutaceae, especially Citrus.
TAXONOMIC CHARACTERISTIS. Apterous vivipara: In life shiny, dark brown or
black. Small to medium aphids (1.5-2.8 mm), oval. Antennae 6-segmented; no secondary
sensoria on antennal segment III; segments not banded, but may get progressively
darker toward the tip of the unguis. Cornicles elongate, black. Cauda elongate, with
19-54 setae, black.
Alate vivipara: In life shiny, brownish-black or black. Small to medium aphids (1.1-
2.6 mm), oval. Antennae 6-segmented; secondary sensoria 7-20 on antennal segment III
and 0-4 on IV; segment III completely dark, other segments may have apical bands.
Cornicles elongate, black. Cauda elongate, with 21-40 setae, black. Forewing with a
pale pterostigma and usually a twice-branched media.
DISCUSSION. Toxoptera citricida, the brown citrus aphid, was originally described
from specimens collected in Hawaii, and the aphid is not known to occur in North
America. However, there is real danger that it may extend its range into Central and
North America. It is largely restricted to hosts in the Rutaceae, and its distribution is
more limited than that of T. aurantii. This species is the principal vector of Citrus
tristeza virus and other citrus viruses. All species in this genus are unique in having a
stridulatory apparatus. When ridges on the venter of the abdomen are rubbed with
conical setae on the hind tibiae, a sound is produced. Alatae are easily identified because
the third antennal segment is black and the cauda has 21-40 setae. Apterae have 19-54
setae on the cauda, and their antennae get gradually darker towards the last two
segments.
Pterochloroides persicae (Cholodkovsky)
DISTRIBUTION IN SOUTHEASTERN US. Not known to occur in the United States.
The species has a high potential of being introduced into the United States and is
included here because it has been moving across northern Africa and because of the
trade in citrus between the Mediterranean countries and North America.
HOSTS. Various species of Prunus, especially armeniaca L. and persica (L.)
Batsch., but also amygdalus Batsch., cerasus L., domestic L., spinosa L. Known also
from Citrus, Cydonia vulgaris Pers., and Malus pumila Mill.
TAXONOMIC CHARACTERISTICS. Apterous vivipara: In life shiny, dark brown or
black with some white coloration; ventral surface silvery white. Medium to very large
aphids (2.7-4.5 mm), oval. Antennae 6-segmented, short; usually no secondary sensoria
on antennal segment III but some specimens may have 4-7, 1-3 on segment IV. Cornicles
large, truncated hairy cones, dusky. Cauda rounded, not well developed. Dorsum of
abdomen with a double row of large, spinal tubercles.
Alate vivipara: In life shiny, dark brown or black with some white coloration.
Medium to large aphids (2.7-3.6 mm), oval. Antennae 6-segmented. short; 10-14 second-
ary sensoria on antennal segment III, 1-5 on antennal segment IV. Cornicles large,
truncated hairy cones, dusky. Cauda rounded, not well developed. Dorsum of abdomen
with a double row of large, spinal tubercles. Wings with large pigmented areas along
veins.
DIScUSSION. Pterochloroides persicae, the "clouded peach bark aphid," is probably
Asian in origin. It is known from the Mediterranean area, the Middle East, and Asia
and has extended its range into Europe and northern Africa. It is a pest of various
species of Prunus (almond, apple, apricot, peach, etc.), and has been reported to occur
on Citrus. Large populations can cause fruit to fall prematurely or not to develop at
all. The aphids are tended by ants and produce copious amounts of honeydew which
serve as a substrate for sooty molds.
We know that aphids move by dispersion or migration. Various quarantine programs
try to check aphid movement from one region to another; however, with the increase
585
586 Florida Entomologist 73(4) December, 1990
in world commerce, this control effort is becoming more and more difficult. It is
virtually impossible to prevent the introduction of aphids carried by air currents into
an area with favorable environmental conditions and an abundance of host plants. Such
was the case in 1986 when Diuraphis noxia (Mordvilko), the Russian wheat aphid, was
detected in Texas (Stoetzel 1987). This aphid probably traveled by air currents from
Mexico into Texas and subsequently into those areas in the western half of the United
States where humidity is low.
ACKNOWLEDGMENTS
For their review of this paper, I thank the following: L. M. Russell and A. L.
Norrbom, Systematic Entomology Laboratory, Agricultural Research Service, U.S.
Department of Agriculture, in Beltsville, MD and Washington, DC, respectively; Clyde
F. Smith, Department of Entomology, North Carolina State University, Raleigh, NC;
and James B. Kring, University of Florida, Bradenton (P.O. Box 713, Anna Maria, FL).
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KNOWLTON, G. F. 1983. Aphids of Utah. Utah State Univ. Res. Bull. 509, 155 pages.
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LEONARD, M. D. 1974. A list of the aphids of Oregon (Homoptera: Aphididae). Oregon
Dept. Agric. Publ., 116 pages.
SMITH, C. F., AND C. S. PARRON. 1978. An annotated list of Aphididae (Homoptera)
of North America. North Carolina Agric. Exp. Stn. Tech. Bull. No. 255. 428
pages.
STOETZEL, M. B. 1987. Information on and identification of Diuraphis noxia
(Homoptera: Aphididae) and other aphid species colonizing leaves of wheat and
barley in the United States. J. Econ. Entomol. 80(3): 696-704.
WRAY, D. L. 1967. The Insects of North Carolina, Third Supplement. North Carolina
Dept. Agric., Div. Entomol., pp. 31-33.
Hamilton & Morse: Symposium-Origins SE Arthropod 587
SOUTHEASTERN CADDISFLY FAUNA:
ORIGINS AND AFFINITIES
STEVEN W. HAMILTON
The Center for Field Biology
Austin Peay State University
Clarksville, Tennessee 37044
JOHN C. MORSE
Department of Entomology
Clemson University
Clemson, South Carolina 29634
ABSTRACT
About 40% of the 1369 Nearctic caddisfly species occur in the southeastern portion
of North America, with 12% being endemic to it. Using techniques of cladistic historical
biogeography, generalized tracks were found among Trichoptera species that link the
Southeast with areas outside of the continent and with areas elsewhere in the Nearctic
Region. A few tracks are known internally among the major southeastern areas of
endemism, but none are sufficiently well corroborated to be considered generalized.
Intercontinentally, the most recent generalized tracks are with the Eastern Palearctic,
Neotropical, and Western Palearctic areas of endemism. At least 16 species occur ex-
clusively in both the southeastern part of the Nearctic Region and the Neotropical
Region, suggesting a greater modern bio-continuity between these areas than with
other intercontinental regions. Within North America, the Southeast is most properly
considered a subset of the Eastern Nearctic area of endemism or, on a more refined
scale, as a mosaic of at least three areas of endemism. The biota of the Southeastern
Coastal Plain, especially, probably has experienced many episodes of cosmopolitanism
and vicariance through time.
RESUME
Aproximadamente el 40% de las species Nearticas de friganeas ocurren en la por-
ci6n sudeste de Norte America, el 12% siendo end6micas. Usando las t6nicas de biogeog-
rafia hist6rica cladistica, se encontraron trazas generals entire species de Tric6pteros
que conectan el sudeste con Areas fuera del continent y con Areas de otros lugares de
la region Neartica. Algunas trazas se conocen internamente entire las principles Areas
de endemismo en el sudeste, pero ninguna esta suficientemente corraborada para con-
siderarla generalizada. Intercontinentalmente, las trazas generalizadas mas reciente se
encuentran en las Areas de endemismo Paleartico del Este, Neotropical, y Paleartico
del Oeste. Por lo meno 16 species ocurren exclusivamente en ambas la parte sudeste
de la Regi6n NeArtica y en la Regi6n Neotropical, lo que sugiere una bio-continuidad
mayor entire estas Areas, que con otras regions intercontinentales. Dentro de Norte
AmBrica, el sudeste es el mas propiamente considerado como un grupo de la region
Nedrtica del Este del endemismo, o en una escala mAs refinada, como un mosaico de
por lo menos tres Areas de endemimo. La biota de la Llanura Costera del Sudeste,
probablemente ha exprimentado muchos episodio de cosmpolitismo y vicarismo a traves
del tiempo.
The caddisflies, or Trichoptera, are a monophyletic group of holometabolous insects
generally considered to be the sister group of the moths, or Lepidoptera. Their eggs,
larvae, and pupae develop almost exclusively in freshwater environments, most abun-
Florida Entomologist 73(4)
dantly in ponds, lakes, creeks, and rivers, but also in small snowmelt pools, seeps,
springs, and even ocean surf. About a half dozen species worldwide have terrestrial, or
virtually terrestrial larvae. In freshwater habitats, eggs usually are found in gelatinous-
matrix masses attached to stable surfaces, either terrestrial or aquatic; larvae may be
found burrowing in the hyporheic zone beneath the aquatic substrate, clinging to stones
or other solid materials, climbing vegetation, sprawling on the stream or pond bottom,
or swimming; pupae typically are buried in the hyporheic zone beneath the aquatic
subtrate or attached to stable benthic materials (Wiggins, 1984, 1987).
In North America, there are about 1369 caddisfly species (Table 1) in 149 genera in
22 families. A total of 544 of these species, or about 40%, occur in the southeastern
United States (as defined in the Introduction to this symposium), 168 of them exclu-
sively (about 12% of all Nearctic species). Among families with more than 100 North
American species, Hydroptilidae, Hydropsychidae, and Leptoceridae are disproportion-
ately well represented in the Southeast (137, 84, and 76 species, representing 59%, 55%,
and 67% of their respective families) and also show high percentages of endemism (25%,
13%, and 19%, respectively). Limnephilidae and Rhyacophilidae are poorly represented
(29 and 25 species, 10% and 20%, respectively), with low percentages of endemism (2%
and 5%, respectively).
These data and the caddisfly distributions that follow are based on our working
checklist of Nearctic Trichoptera, which is too extensive to present here (currently
nearly 250 manuscript pages). The most recently published Nearctic checklist was pro-
vided by Ross (1944); the Trichopterorum Catalogus (Fischer, 1960-1973) gave distribu-
TABLE 1. NORTH AMERICAN TRICHOPTERA SPECIES PRESENT OR ENDEMIC IN
THE SOUTHEASTERN UNITED STATES, WITH FAMILIES RANKED BY
NUMBER OF SPECIES PRESENT IN THOSE STATES.
Total spp. Endemic spp.
in SE states/ in SE states/
Total spp. percent of percent of
Family in N. Amer. N. Amer. spp. N. Amer. spp.
Hydroptilidae 231 137/59% 57/25%
Hydropsychidae 154 84/55% 20/13%
Leptoceridae 113 76/67% 21/19%
Polycentropodidae 76 41/54% 10/13%
Lepidostomatidae 66 31/47% 13/20%
Limnephilidae 293 29/10% 5/ 2%
Glossosomatidae 77 26/34% 9/12%
Rhyacophilidae 126 25/20% 6/ 5%
Brachycentridae 36 19/53% 4/11%
Philopotamidae 47 19/40% 5/11%
Uenoidae 47 13/28% 6/13%
Sericostomatidae 14 11/79% 7/50%
Phryganeidae 28 9/32% 0/ 0%
Odontoceridae 13 8/62% 2/15%
Molannidae 8 5/63% 0/ 0%
Psychomyiidae 18 4/22% 1/ 6%
Helicopsychidae 8 3/38% 1/13%
Beraeidae 3 2/67% 1/33%
Calamoceratidae 5 2/40% 0/ 0%
Ecnomidae 1 0/ 0% 0/ 0%
Hydrobiosidae 3 0/ 0% 0/ 0%
Xiphocentronidae 2 0/ 0% 0/ 0%
TOTAL/MEAN % 1,369 544/40% 168/12%
December, 1990
Hamilton & Morse: Symposium-Origins SE Arthropod 589
tions for all world species through 1960; and subsequent records were made available
in scattered publications. G. B. Wiggins and O. S. Flint. Jr., (pers. cor.) are preparing
an updated North American caddisfly species checklist for publication.
HISTORICAL BIOGEOGRAPHY TECHNIQUES AND DEFINITIONS
To use cladistic methods in historical biogeography (e.g.. Nelson & Platnick, 1981,
Allen, 1984, Humphries & Parenti, 1986), one first identifies areas of endemism by
congruence of limits of distribution for many different taxa. These can be identified on
any of several scales, from the six principal intercontinental biogeographical regions for
terrestrial organisms (Sclater, 1858) to intracontinental biomes and communities (An-
drewartha & Birch, 1954). One then examines the phylogenies of these taxa to discover
their interested patterns of historical relationships. Phylogenetic patterns among taxa
in a minimum of three different areas of endemism suggest patterns of historical re-
lationship among those areas. Two areas of endemism that are more closely related to
each other than to any other such area are said to be connected by a "track." Several
congruent tracks constitute a "generalized track" whose probability of occurrence by
chance alone is very small, suggesting a common cause.
Areas of Endemism: Intercontinentally, Sclater's (1858) six biogeographic regions
have merit as areas of endemism for caddisflies (Ross, 1956, 1967). In addition, the
Palearctic Region appears to have eastern and western areas of endemism (Levanidova,
1982, Malicky, 1983). Concerning large-scale areas of endemism in North America,
caddisflies generally occur either east of the Rocky Mountains or in and west of the
Rocky Mountains (Ross, 1956, Allen, 1984).
As mentioned in the Introduction to this symposium, the three major areas of en-
demism occurring within "the Southeast" are the Eastern Highlands, the Interior High-
lands, and the Southeastern Coastal Plain (Introduction, Fig. 1). Obviously, although
much or most of each of these areas of endemism lies within "the Southeast," each also
extends beyond its arbitrary political borders. Substantial evidence for the southeastern
Nearctic areas of endemism based on the caddisflies has been presented for the Eastern
Highlands by Etnier & Schuster (1979), Unzicker et al (1982), Harris (1986), Lago &
Harris (1987), and Morse et al (1989); for the Interior Highlands by Unzicker et al (1970)
and Bowles & Mathis (1989); and for the Southeastern Coastal Plain by Blickle (1962),
Morse et al (1980), Unzicker et al (1982), Harris et al (1982a, 1982b, 1984), Holzenthal
et al (1982), Lago et al (1982), Harris (1986), and Lago & Harris (1987). These include
at least 104 species endemic for the Eastern Highlands, 13 species endemic for the
Interior Highlands, and 89 species endemic for the Southeastern Coastal Plain.
Tracks and Associations: Few cladistic analyses have been made for the Trichoptera
either on a worldwide scale or for the Nearctic species. Therefore, evidence for "tracks,"
as discussed above, is limited and of various levels of usefulness. For us to infer the
phylogenies of each of the caddisfly taxa cited below in order to substantiate "tracks"
is beyond the scope of this work. Therefore, in addition to tracks, we cite three classes
of data which provide evidence for recent and historical associations (panbiogeographic
"tracks" of Croizat, 1958). The relative generality of an association, as determined by
the number of taxa supporting it, provides evidence inversely for the effectiveness of
recent and historical ecological barriers between the areas of endemism; i.e., the more
general the association, the less effective has been the ecological barrier either recently
or at some time in the past. Unlike track analysis, analysis of associations provides no
evidence for their relative recency.
A track is recognizable by two sister lineages (either distinct species or monophyletic
species groups) in different areas of endemism; in the minimum three-taxon statement,
a sister lineage of these two lineages ideally occurs in a third area of endemism (Nelson
& Platnick, 1981, Humphries & Parenti, 1986). Redundancy (two or more successive
Florida Entomologist 73(4)
clades distributed in the same area of endemism) is resolved by reduction. Absence of
one or more areas in a given cladogram is recognized by congruence of other cladograms
that include those areas. Ambiguity from apparently widespread taxa inhabiting two
or more areas of endemism may result, for example, from dispersal or from failure to
speciate in response to an ecological vicariance event; these are resolved by admitting
that some of the components of the cladogram are false (Nelson & Platnick, 1981,
Nelson, 1982, Humphries & Parenti, 1986). The result of a track analysis is the discovery
that two areas of endemism are more closely (= recently) related to each other than
either is to a third area.
A Class A Association is recognizable by the distribution of one modem species in
two particular areas of endemism. A Class A Association represents a situation in which
these areas have become areas of endemism for many other species, but for which the
cited species has not responded with allopatric speciation. Some possible reasons that
it has not responded are either (a) because of homeostasis in that species following
ecological vicariance separating an original area of endemism into two such areas, (b)
because the ecological requirements of the cited species are sufficiently broad that, for
it, no ecological vicariance has occurred, or (c) because the distribution represents a
recent dispersal of the species across an ecological barrier, a dispersal too recent to
have allowed allopatric speciation to occur.
A Class B Association is one for which a monophyletic or presumably monophyletic
superspecific taxon is distributed in only two areas of endemism. In such cases, there
may actually be one or more tracks or Class A Associations. Phylogenetic analysis
within the taxon would reveal these, but such analysis is outside the scope of this
discussion.
A Class C Association is one for which a monophyletic or presumably monophyletic
superspecific taxon is distributed in three or more areas of endemism; the sister taxon
occurs ideally in some other area of endemism. In such cases, an actual track between
any two of these particular areas of endemism may occur, but has not yet been demon-
strated. With such a more nearly cosmopolitan distribution and with such poor
phylogenetic evidence, it can be said only that the probability of a track occurring
between any two of these particular areas of endemism is greater than between any
other, uninhabited areas.
EASTERN NORTH AMERICAN TRACKS
Tracks are evident for the following southeastern caddisfly taxa between Australian
(AUS), Eastern North American (ENA), Eastern Palearctic (EPL), Ethiopian (ETH),
Neotropical (NTR), Oriental (ORL), Western North American (WNA), and Western
Palearctic (WPL) areas of endemism. Phylogeny is indicated by interested parentheses
in which terminal sister taxa or areas are separated with a "+ and, in area cladograms,
areas of endemism occupied by a widely distributed taxon are separated by a comma.
For example, in cladogram 1, the species alagma and brevis and neph and protonepha
and tarsipunctata share one or more homologues (= synapomorphies) not shared with
any other taxa and thus are grouped as a single lineage; major shares with them one
or more additional homologues; species of the spinosa Group share with all six of the
above one or more additional homologues uniquely; the cladogram of the areas of en-
demism that each species inhabit is organized completely congruently with the taxon
cladogram. In cladogram 2, ancylus and kamonis are sister species which together form
a single lineage; flava and forcipta are independent lineages for which no homologues
are known by which to resolve the trichotomy with the ancylus + kamonis lineage.
Also in cladogram 2, the redundancy of the original area cladogram is reduced in a
refined area cladogram.
December, 1990
Hamilton & Morse: Symposium-Origins SE Arthropod 591
1. Ceraclea (Athripsodina) (((alagma + brevis + nepha + protonepha + tar-
sipunctata) + major) spinosa Group) (Morse, 1974; Yang & Morse, 1988) =
((ENA + ORL) ETH).
2. Ceraclea (Athripsodina) ((ancysus + kamonis) and/or flava and/or forcipata)
(Morse, 1974; Yang Morse, 1988) =
((ENA + EPL) and/or ENA and/or ORL) =
((ENA + EPL) ORL).
3. Ceraclea (Athripsodina) ((diluta + perplexa) (annulicornis Group ripariaa
Group + marginata Group))) (Morse, 1974, 1975) =
((ENA + WPL) (ENA, WNA, EPL, WPL ((WPL, EPL, ORL) + (EPL,
ORL)))) =
((ENA + WPL) (ENA, WNA (WPL + EPL, ORL))).
4. Ceraclea (Athripsodina) ((tarsipunctata Group + spinosa Group) rest of Sub-
genus Athripsodina) (Morse, 1974) =
((ENA + ETH) ENA, EPL, ORL, WNA, WPL)=
((ENA + ETH) EPL, ORL, WNA, WPL).
5. Ceraclea ((Subgenus Ceraclea + Subgenus Pseudoleptocerus) Subgenus
Athripsodina) (Morse, 1974, 1975, 1978) =
((ENA, EPL, WNA, WPL + ETH) ENA, EPL, ORL, WNA, WPL) =
((ENA, EPL, WNA, WPL + ETH) ORL).
6. Dolophilodes (D.) ((distinctus + similis) (indica + japonica + orientalis))
(Ross, 1956; Levanidova, 1982) =
((ENA + EPL) ORL, EPL, ORL)=
((ENA + EPL) ORL).
7. Glossosoma (Eomystra) (((nigrior + dulkejti) lividum) and/or (ussuricum +
inops) and/or intermediumm + verdona)) (Ross, 1956) =
(((ENA + EPL) ENA) and/or (EPL + EPL) and/or (ENA, EPL, WNA, WPL
+ WNA)) =
((ENA + EPL) (WPL + WNA)).
8. Micrasema (((rusticum and/or moestum and/or (rickeri (minimum +
setiferum))) rest of rusticum Group) baitinum) (Chapin, 1978) =
(((ENA and/or WPL and/or (ENA (WPL + WPL))) ENA) ORL) =
((ENA + WPL) ORL).
9. Mystacides ((sepulchralis + bifida) alafimbriata) (Yamamoto & Ross, 1966) =
((ENA + EPL) WNA).
10. Oxyethira (Argyrobothrus) ((zeronia (velocipes + flagellata)) Subgenus
Dampfitrichia) (Kelley, 1982, 1984) =
((ENA (ETH + ETH)) AUS, ENA, NTR, ORL, WNA) =
((ENA + ETH) AUS, NTR, ORL, WNA).
11. Oxyethira (Dactylotrichia) ((kingi + rest of Subgenus Dactylotrichia) Subgenus
Loxotrichia) (Kelley, 1982, 1984: Holzenthal & Kelley, 1983) =
((ENA + NTR) ENA, NTR, WNA)=
((ENA + NTR) WNA).
12. Oxyethira (Dampfitrichia) ((((florida + simulatrix) aculea) ulmeri) (incana +
galekoluma)) (Kelley, 1982) =
((((ENA + NTR) NTR) NTR) (AUS + ORL)) =
((ENA + NTR) (AUS + ORL)).
13. Oxyethira (Dampfitrichia) (((verna + circaverna) (discaelata + campesina))
pallida Subgroup) (Kelley, 1982, 1983, 1984) =
(((ENA + NTR) (NTR + NTR)) ENA, NTR, WNA, Hawaii) =
((ENA + NTR) WNA, Hawaii).
14. Oxyethira (Holarctotrichia) (((elerobi (archaica + iglesiasi)) rest of Subgenus
Florida Entomologist 73(4)
Holarctotrichia) Subgenera Argyrobothrus, Dampfitrichia, Mesotrichia, Oxy-
trichia, Tanytrichia, Dactylotrichia, and Loxotrichia) (Kelley, 1982, 1984) =
(((ENA (WPL + WPL)) ENA, WPL) AUS, ENA, ETH, NTR, ORL, WNA) =
((ENA + WPL) AUS, ETH, NTR, ORL, WNA).
15. Oxyethira (Oxytrichia) ((sininsigne + maryae) (unispina (dualis (leonensis +
glasa)))) (Kelley, 1982, 1983) =
((ENA + NTR) (NTR (ENA, WNA (ENA + ENA, NTR)))) =
((ENA + NTR) (WNA (ENA + NTR))).
16. Phylocentropus ((auriceps + spiniger) orientalis) (Ross, 1965) =
((ENA + WPL) ORL).
17. Polycentropus ((confusus Group + nigriceps Group) (arizonensis Group +
gertschi Group)) (Hamilton, 1986, 1988) =
((ENA + NTR) (NTR + WNA)).
18. Rhyacophila ((glaberrima + stigmatica Group) (vagrita Group + montana
Group)) (Ross, 1956; Schmid, 1970) =
((ENA + WPL) (WNA + ENA)).
19. Rhyacophila ((torva (nigrocephala Group)) rest of divaricata Branch) (Schmid,
1970)=
((ENA (EPL + ORL)) ORL, WNA, EPL) =
((ENA (EPL + ORL)) WNA).
20. ((Theliopsyche + Martynomyia) and/or Crunoecia and/or Archaeocrunoecia
and/or Maniconeurodes) (Weaver, 1983) =
((ENA + WPL) and/or WPL, EPL and/or WPL and/or WPL) =
((ENA + WPL) EPL).
21. Wormaldia (Doloclanes) ((mohri + kisoensis) montana) (Ross, 1956) =
((ENA + EPL) ORL).
22. Wormaldia (W.) ((moesta + chinensis + relicta) gabriella) (Ross, 1956) =
((ENA + EPL + ORL) WNA).
EASTERN NORTH AMERICAN ASSOCIATIONS
Associations are evident for the following southeastern Trichoptera taxa between
the Eastern North American (ENA) and the Western Palearctic (WPL) areas of en-
demism:
Class A Associations, ENA + WPL:
None.
Class B Associations, ENA + WPL:
Ironoquia.
Class C Associations, ENA + WPL + one or more other areas:
Agapetus, Agraylea, Agrypnia, Apatania, Beraea, Brachycentrus (B.), Cerato-
psyche, Cheumatopsyche, Chimarra, Culoptila, Diplectrona, Dolophilodes, Goera,
Glossosoma, Helicopsyche, Heteroplectron, Hydatophylax, Hydropsyche, Hydro-
ptila, Lepidostoma, Leptocerus, Limnephilus, Lype, Molanna, Mystacides, Neurec-
lipsis, Oecetis, Orthotrichia, Palaeagapetus, Parapsyche, Phryganea, Polycen-
tropus Protoptila Psychomyia, Rhyacophila, Setodes, Stactobiella, Triaenodes,
Wormaldia, Ylodes.
Associations are evident for the following southeastern Trichoptera taxa between
the Eastern North American (ENA) and the Eastern Palearctic (EPL) areas of en-
demism:
Class A Associations, ENA + EPL:
None.
Class B Associations, ENA + EPL:
Potamyia.
Class C Associations, ENA + EPL + one or more other areas:
592
December, 1990
Hamilton & Morse: Symposium-Origins SE Arthropod 593
Agapetus, Agrypnia, Anisocentropus, Apatania, Arctopsyche, Beraea, Brachycen-
trus (B.), Brachycentrus (Sphinctogaster), Ceratopsyche, Cheumatopsyche,
Chimarra, Diplectrona, Goera, Helicopsyche, Hydatophylax, Hydropsyche, Hydro-
ptila, Lepidostoma, Limnephilus, Macrostemum, Micrasema, Molanna, Neo-
phylax, Neureclipsis, Nyctiophylax, Oecetis, Oxyethira, Palaeagapetus, Para-
psyche, Phryganea, Phylocentropus, Polycentropus, Pseudostenophylax, Psy-
chomyia, Rhyacophila, Setodes, Stactobiella, Triaenodes, Ylodes.
Associations are evident for the following southeastern caddisfly taxa between the
Eastern North American (ENA) and the Neotropical (NTR) areas of endemism:
Class A Associations, ENA + NTR:
Cernotina calcea, Cheumatopsyche lasia, Cyrnellus marginalis, Hydroptila an-
gusta, Mayatrichia ayama, Nectopsyche spiloma, Nectopsyche pavida, Neotrichia
vibrans, Ochrotrichia tarsalis, Oecetis cinerascens, Orthotrichia aegerfasciella,
Oxyethira (Loxotrichia) azteca, Oxyethira (Oxytrichia) glasa, Oxyethira (Loxo-
trichia) janella, Oxyethira (Dampfitrichia) maya, Triaenodes mephitus.
Class B Associations, ENA + NTR:
Cernotina, Cyrnellus.
Class C Associations, ENA + NTR + one or more other areas:
Anisocentropus, Chimarra, Culoptila, Dolophilodes, Helicopsyche, Hydroptila,
Leucotrichia, Macrostemum, Nectopsyche, Neotrichia, Nyctiophylax, Ochrotrichia,
Oecetis, Protoptila, Triaenodes, Wormaldia.
Associations are evident for the following southeastern caddisfly taxa between the
Eastern North American (ENA) and the Oriental (ORL) areas of endemism:
Class A Associations, ENA + ORL:
None.
Class B Associations, ENA + ORL:
Psilotreta.
Class C Associations, ENA + ORL + one or more other areas:
Agapetus, Anisocentropus, Apatania, Arctopsyche, Ceraclea, Ceratopsyche,
Cheumatopsyche, Dolophilodes, Goera, Glossosoma, Helicopsyche, Hyropsyche,
Hydroptila, Leptocerus, Lype, Macrostemum, Molanna, Mystacides, Nyctiophylax,
Oecetis, Orthotrichia, Oxyethira, Paduniella, Parapsyche, Phryganea, Phylocen-
tropus, Polycentropus, Pseudostenophylax, Setodes, Triaenodes.
Associations are evident for the following southeastern caddisfly taxa between the
Eastern North American (ENA) and the Ethiopian (ETH) areas of endemism:
Class A Associations, ENA + ETH:
None.
Class B Associations, ENA + ETH:
None.
Class C Associations, ENA + ETH + one or more other areas:
Anisocentropus, Cheumatopsyche, Chimarra, Dolophilodes, Helicopsyche, Hydro-
psyche, Hydroptila, Leptocerus, Lype, Macrostemum, Nyctiophylax, Oecetis,
Orthotrichia, Paduniella, Polycentropus, Triaenodes, Wormaldia.
Associations are evident for the following southeastern caddisfly taxa between the
Eastern North American (ENA) and the Australian (AUS) areas of endemism:
Class A Associations, ENA + AUS:
None.
Class B Associations, ENA + AUS:
None.
Class C Associations, ENA + AUS + one or more other areas:
Anisocentropus, Cheumatopsyche, Chimarra, Diplectrona, Dolophilodes, Helico-
psyche, Hydropsyche, Hydroptila, Leptocerus, Macrostemum, Nyctiophylax,
Oecetis, Oxyethira, Triaenodes.
594 Florida Entomologist 73(4) December, 1990
SOUTHEASTERN NORTH AMERICAN TRACKS
Tracks are evident for the following caddisfly taxa between the Southeastern North
American (SE), other parts of the Eastern North American (ENA), and the Western
North American (WNA) areas of endemism:
23. Arctopsyche (irrorata grandiss + inermis)) (Schmid, 1968) =
(SE (ENA, WNA + WNA)) =
(SE (ENA + WNA)).
24. Brachycentrus (Sphinctogaster) (((etowahensis numerouss + solomoni)) (ap-
palachia + spinae) lateralis Subgroup) (Flint, 1984) =
(((SE (ENA + ENA)) (ENA + SE)) ENA, WNA)=
((SE + ENA) WNA).
25. Cheumatopsyche (((gyra + vannotei) etrona) gracilis) (Gordon, 1974) =
(((SE + ENA) ENA) ENA, WNA) =
((SE + ENA) WNA).
26. Cheumatopsyche ((rossi (logani + smithi) pettiti) (Gordon, 1974)=
((SE (WNA + WNA)) ENA + WNA) =
((SE + WNA) ENA).
27. Lepidostoma (Mormomyia) ((((((excavatum + sommermanae) libum) (vernale
Group + mitchelli Group)) serratum Group) griseum Group) Subgenus
Nosopus) (Weaver, 1983) =
((((((SE + ENA) ENA) (ENA + SE)) ENA, SE) ENA, SE) ENA, NTR,
WNA) =
((SE + ENA) NTR, WNA).
28. Lepidostoma (.1., ... .. .g, ((((((((flinti + glenni) vernale) sackeni) mitchelli
Group) libum Group) serratum Group) griseum Group) Subgenus Nosopus)
(Weaver, 1983) =
(((((((SE + SE) ENA) ENA) SE) ENA, SE) ENA, SE) ENA, SE) ENA, NTR,
WNA) =
((SE + ENA) NTR, WNA).
29. Oxyethira (Holarctotrichia) ((((setosa + dunbartonensis) forcipata) obtatus)
michiganensis) (Kelley, 1982) =
((((SE + SE) ENA) ENA) ENA, WNA)=
((SE + ENA) WNA).
30. Oxyethira (0.) ((((novasota + grisea) (coercens + rivicola)) lumosa) (al-
lagashensis (frici + simplex))) (Kelley, 1985) =
((((SE + ENA) (ENA + ENA)) SE) (ENA (WPL + WPL))) =
((SE + ENA) (ENA + WPL)).
31. Oxyethira (Oxytrichia) ((abacatia + anabola) aeola) (Kelley, 1982) =
((SE + ENA) WNA).
32. Parapsyche ((cardis + elsis) ((((apicalis + almota) spinata) extensa) tur-
binata)) (Schmid, 1968) =
((SE + WNA) ((((ENA + WNA) WNA) WNA) WNA)) =
((SE + WNA) ENA).
33. Rhyacophila (((amicis + melita) atrata) (pellisa + valuma)) (Ross, 1956) =
(((SE + ENA) ENA) (WNA + WNA)) =
((SE + ENA) WNA); or
Rhyacophila ((amicis + melita) ((atrata + colona) (pellisa + valuma)))
(Schmid, 1970) =
((SE + ENA) ((ENA + WNA) (ENA + WNA)))=
((SE + ENA) (ENA + WNA).
34. Rhyacophila ((montana (milnei + vagrita)) glaberrima) (Schmid, 1970) =
Hamilton & Morse: Symposium-Origins SE Arthropod 595
((SE (WNA + WNA)) ENA) =
((SE + WNA) ENA).
35. Theliopsyche (((((epilonis + parva) melas) grisea) corona) Genus Martynomyia)
(Weaver, 1983) =
(((((SE + ENA) ENA) ENA) SE) WPL) =
((SE + ENA) WPL).
36. Wormaldia (W.) ((thyria + hamata) (shawnee + strota) and/or occidea) (Ross
1956) =
((SE + WNA) (ENA + ENA) and/or WNA) =
((SE + WNA) ENA).
SOUTHEASTERN NORTH AMERICAN ASSOCIATIONS
Associations are evident for the following southeastern caddisfly taxa with the rest
of Eastern North America:
Class A Associations, SE + ENA:
317 species.
Class B Associations, SE + ENA:
Agarodes, Goerita, Oligostomis, Oxyethira grisea Group (Kelley, 1985), Platycen-
tropus, Pycnopsyche, Theliopsyche.
Class C Associations, SE + ENA + one or more other areas:
Agapetus, Agraylea, Agrypnia, Anisocentropus, Apatania, Arctopsyche,
Banksiola, Beraea, Ceraclea, Ceratopsyche, Cernotina, Chimarra, Culoptila, Cyr-
nellus, Diplectrona, Dolophilodes, Goera, Glossosoma, Helicopsyche, Hes-
perophylax, Heteroplectron, Hydatophylax, Hydropsyche, Hydroptila, Ironoquia,
Leptocerus, Leucotrichia, Limnephilus, Lype, Macrostemum, Mayatrichia,
Molanna, Mystacides, Nectopsyche, Neophylax, Neotrichia, Neureclipsis, Nyc-
tiophylax, Ochrotrichia, Oecetis, Orthotrichia, Parapsyche, Phryganea, Phylocen-
tropus, Polycentropus, Protoptila, Pseudostenophylax, Psilotreta, Psychomyia,
Ptilostomis, Setodes, Stactobiella, Triaenodes, Wormaldia.
Associations are evident for the following caddisfly taxa between Southeastern
North America (SE) and the Western North American (WNA) area of endemism:
Class A Associations, SE + WNA:
Culoptila thoracica, Dolophilodes sisko.
Class B Associations, SE + WNA:
None.
Class C Associations, SE + WNA + one or more other areas:
Agapetus, Agraylea, Agrypnia, Apatania, Banksiola, Brachycentrus (Sphinctogas-
ter), Ceraclea, Ceratopsyche, Chimarra, Diplectrona, Goera, Glossosoma, Helicop-
syche, Hesperophylax, Heteroplectron, Homoplectra, Hydatophylax, Hydropsyche,
Hydroptila, Lepidostoma, Leucotrichia, Limnephilus, Macrostemum, Mayatrichia,
Molanna, Mystacides, Nectopsyche, Neophylax, Neotrichia, Neureclipsis, Nyctio-
phylax, Ochrotrichia, Oecetis, Orthotrichia, Paleagapetus, Phryganea, Polycen-
tropus, Protoptila, Pseudostenophylax, Psychomyia, Ptilostomis, Triaenodes.
INTRA-SOUTHEASTERN NORTH AMERICAN TRACKS
The number of shared species and genera among the three areas of endemism in the
Southeast is so great (very many Class A, B, and C Associations) that relative relation-
ships actually are masked. The number of Tracks is smaller, mostly because of the few
phylogenetic analyses that have been accomplished to date. Despite this selective nature
of the evidence, the following Tracks are noteworthy. Tracks for at least the following
596 Florida Entomologist 73(4) December, 1990
caddisfly taxa have been inferred between the Southeastern Interior Highlands (SIH),
Southern Eastern Highlands (SEH), and Coastal Plain (SCP) areas of endemism as
indicated:
((SIH + SCP) sister area):
37. Micrasema ((ozarkana + n. sp.) wataga) (Chapin, 1978; Chapin & Morse, in
prep.) =
((SIH + SCP) ENA).
38. Setodes (((oxapius + dixiensis) (incertus + stehri)) arenatus) (Holzenthal, 1982)
(((SIH + SCP) (ENA + SEH)) SCP).
((SIH + SEH) sister area):
39. Helicopsyche ((limnella + paralimnella) and/or borealis and/or mexicana)
(Morse et al, 1989) =
((SIH + SEH) and/or ENA, NTR, WNA and/or WNA, NTR) =
((SIH + SEH) ENA, NTR, WNA).
40. Rhyacophila ((((kiamichi + ledra) fenestra) teddyi) carolina) castanea Group)
(Ross, 1956; Schmid, 1970) =
((((SIH + ENA) ENA) SEH) ENA) ORL) =
((SIH + ENA) SEH) ORL).
((SCP + SEH) sister area):
41. Agarodes alabamensisis + tetron) griseus) Ross & Scott, 1974; Harris, 1987) =
((SCP + SEH) ENA).
CONCLUSIONS
The intercontinental tracks suggest that the eastern North American caddisflies
evolved with close relations in several different regions, most recently in the Neo-
tropical (cladograms 11, 12, 13, 15, and 17) and East Palearctic (cladograms 2, 6, 7, 9,
19, 21, and 22) regions. As might be expected, the Neotropical + Eastern North Amer-
ican fauna generally had its most recent relatives in Western North America (Allen,
1984, Noonan, 1986). Some of the Eastern Palearctic + Eastern North American fauna
also had its most recent relatives at least partly in Western North America (cladograms
7, 9, and 19), but some of this fauna (cladograms 2, 6, 21, and 22) had its most recent
relatives in the Oriental Region. The Western Palearctic relationships are also relatively
recent (cladograms 3, 8, 14, 16, 18, and 20), typically preceded by ancestors also living
in one or more of each of the world's biogeographc regions. Lineages whose extant
species suggest ancestral distributions involving Eastern North America and the Ethio-
pian Region (cladograms 4, 5, and 10) or the Oriental Region (cladogram 1) are less
common, and provide insufficient evidence for discovering distributions of their histori-
cal antecedents, implying that the relationships are much more ancient than for the
previous three regions.
The Southeastern species mostly evolved from widespread Eastern North American
ancestors (cladograms 24, 25, 27, 28, 29, 30, 31, 33, and 35), preceded by older transcon-
tinental populations or populations also living in the Neotropical or Western Palearctic
regions. Thee are several examples, however, of apparent evolution directly from more
widespread North American ancestors (cladograms 23, 26, 32, 34, and 36).
The less rigorously analyzed associations for Eastern North America with Western
Palearctic, Eastern Palearctic, and Neotropical Biogeographic regions are supported
nearly equally by the evidence from caddisflies. The major difference is that there are
at least 16 Class A Associations of southeastern species presently living in Eastern
North America and the Neotropical Region and none known in both Eastern North
America and Europe or Asia. This suggests to us that the biogeographic relationships
Hamilton & Morse: Symposium-Origins SE Arthropod 597
of Eastern North America have been well established historically with these other three
regions, but that either the Atlantic and Pacific oceanic barriers have been more effec-
tive in facilitating allopatric speciation than has the arid North American Southwest
and/or the relationship with the Neotropical Region has been more recent. The large
number of unresolved Class C Associations among these areas of endemism suggests
that there is yet much historical biogeographic evidence to be gleened from future
phylogenetic studies.
Within the Nearctic Biogeographic Region, there are many species endemic to the
Southeast (168 spp.), but nearly twice as many more (317 spp.) are shared with Eastern
North America outside the Southeast. Clearly, if the Southeast is to be treated as an
area of endemism, it can only be considered a sub-area of Eastern North America.
The number of caddisfly associations between the Southeast and Eastern North
America also is much greater than between the Southeast and Western North America,
especially for Class A Associations (317 spp. versus 2 spp.). The number of Class B and
C Associations between the Southeast and Western North America (0 genn. and 42
genn., respectively) is comparable to those between Eastern North America and Europe
(1 gen. and 40 genn., respectively) or Asia (1 gen. and 40 genn., respectively) or the
Neotropics (2 genn. and 16 genn., respectively). This fact seems to corroborate the
hypothesis that Eastern and Western North American areas of endemism are as distinc-
tive as are these other areas.
Although the number of endemic caddisfly species in the southeastern areas of en-
demism is large, especially for the Eastern Highlands and the Coastal Plain, the number
of cladograms that have been published for these species is small, such that the number
of tracks presently known among them is insufficient to consider any intra-southeastern
track generalized. It is interesting, however, that many of the Coastal Plain endemics
are more closely related to species further north, such as the Class B Association for
Neureclipsis melco Ross (Polycentropodidae) with the other five species of the genus
in North America. Very few species have sister lineages that are also Coastal Plain
endemics, although a few examples exist, such as sister species Cheumatopsyche morse
Gordon and C. virginica Denning (Hydropsychidae; Gordon, 1974). We thus infer that
the endemic Southeastern Coastal Plain caddisfly fauna had multiple origins, perhaps
the result of numerous peripheral isolations of ancestral taxa resulting from fluctuating
climates and possibly associated sea level oscillations.
Flint and Harp (1990) offered a similar explanation for the origin of several members
of the Lepidostoma modestum Group. Four of the species are widespread from the
Northeast to the southern Appalachians while four other members are more restricted:
L. compressum Etnier and Way and L. modestum (Banks) in the Cumberland Plateau
and Appalachian Mountains, L. weaveri Harris in the upper Coastal Plain of Alabama,
and L. ozarkensis Flint and Harp in the Ozarks of the Interior Highlands. The first
three are sister species according to Weaver (1983), the latter is the sister species of
the widely distributed northeastern L. ontario Ross (Flint & Harp 1990). These authors
suggested that the southeastern species may represent "relictual distributions resulting
from expanding and contracting ranges related to earlier advances and retreats of the
main range of the species group." For this reason, although the Southeastern Coastal
Plain is an area of endemism for many species, the causes for this endemism must be
viewed as multiple and complex, and determination of the timing of vicariance events
will be very difficult to correlate geologically, ecologically, and faunistically.
ACKNOWLEDGMENTS
We are grateful to R. N. Vineyard (Royal Ontario Museum) for translating impor-
tant parts of the volume by Levanidova (1982).
Florida Entomologist 73(4)
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Hulting et al.: Population Growth Comparisons 601
A COMPUTER PROGRAM FOR CALCULATION AND
STATISTICAL COMPARISON OF INTRINSIC RATES OF
INCREASE AND ASSOCIATED LIFE TABLE PARAMETERS
FRED L. HULTING'
Department of Statistics
DAVID B. ORR AND JOHN J. OBRYCKI
Department of Entomology
Iowa State University
Ames, Iowa 50011
ABSTRACT
An often crucial component in the study of insects is the determination of reproduc-
tive capabilities. While it is important to study reproduction at the level of the individ-
ual, it is often more desirable to develop a standardized estimate of the growth rate of
insect populations. One such estimator, the intrinsic rate of increase (rm), has been used
for many years by insect ecologists. This statistic is useful not only as a measure of
population growth potential but also as a bioclimatic index and a natural enemy rating
parameter. Recently introduced methods to calculate variances for rm values have in-
creased the utility of the statistic, enabling comparisons of growth rates from different
species or biotypes. Although these values are usually calculated with the help of a
computer, programs to perform the required computations have not been widely avail-
able. In this paper we describe a computer program, written in Pascal, designed to
simplify the calculation of intrinsic rates of increase and associated Jacknife estimates
of variance. The program also computes (1) approximate confidence intervals, (2) several
commonly used fertility table parameters with their standard errors, and (3) daily means
for some of these parameters, which can be used for graphing data.
RESUME
Un component crucial que ocurre a menudo en el studio de insects es la deter-
minaci6n de la capacidad de reproducci6n. Mientras que es important estudiar la repro-
ducci6n al nivel del individuo, a menudo es mAs deseable desarrollar un estimado patr6n
de la taza de crecimiento de las poblaciones de insects. Un tal estimador, la taza
intrinsica de aumento (rm), ha sido usada por ecologistas de insects por muchos afios.
Esta estadistica es Util no solo como una media del potential de crecimiento de la
poblaci6n, pero tambien como un indice bioclimAtico y un parAmetro natural de la taza
del enemigo. M4todos introducidos recientemente para calcular la varianza de los valores
rm han aumentado la utilidad de la estadistica, lo que ha permitido comparar la taza de
crecimiento entire diferente species o biotipos. Aunque estos valores son calculados
usualmente con la ayuda de una computadora, los programs para llevar a cabo las
computaciones requeridas no han estado comunmente disponibles. En este trabajo
nosotros decribimos un program de computadora escrito en Pascal, disefiado para
simplificar la calculaci6n de tazas de aumentos intrinsicos y los estimados asociados de
varianza Jacknife. El program tambien computa (1) intervalos apr6ximados de con-
fianza, (2) algunos parametros comunmente usados en tablas de fertilidad con su patron
de errors, y (3) promedio diario para algunos de esos parametros, los cuals se pueden
usar para hacer grAficos de los datos.
'Current address: Mathematics Department, General Motors Research Laboratories, Warren, MI 48090-4055.
Florida Entomologist 73(4)
An often crucial component in the study of insects is the determination of reproduc-
tive capacities. While it is important to study reproduction at the level of the individual,
it is often more desirable to calculate a standardized estimate of the growth rate of
insect populations (Southwood 1978). One such estimator, the intrinsic rate of increase
(rm), has been used for many years by insect ecologists.
Although the use of rm has been criticized (e.g., Hirose 1986, Laughlin 1965) for the
assumption of a stable age distribution (see Andrewartha & Birch 1954) in study popu-
lations, it has been demonstrated to be both a predictive and comparative measure of
population growth potential (e.g., Force 1974, Force & Messenger 1964). This parameter
has been used widely in the study of a variety of insect populations (see Gaston 1988).
In biological control, this statistic has been used as a bioclimatic index and a natural
enemy rating parameter (e.g., Messenger 1964, 1970). It is not only useful for comparing
between beneficial species, but also for comparing biotypes of the same species, and for
comparing natural enemies to their prey or host (e.g., Force & Messenger 1964, Or-
phanides & Gonzales 1971, Nechols et al. 1989).
Since r, is an estimate, it is important to recognize the uncertainty associated with
it. Meyer et al. (1986) introduced a Jacknife estimate of the variance of rm which can
be used to assess this uncertainty. However, the calculation of rm and the variance
estimate must be done on a computer, and programs to perform the required computa-
tions have not been widely available. Abou-Setta et al. (1986) described a program
written in BASIC which calculated fertility table parameters for arthropods, however,
the program did not include an estimate of the variance of rm. In this paper we describe
a computer program, written in Pascal, which provides such an estimate in addition to
approximate confidence intervals, fertility table parameters with their standard errors,
and daily means of progeny production and survivorship.
PROGRAM DESCRIPTION
In this section we describe program input and output, as well as the computational
algorithms used by the program. Mathematical notation is introduced to abbreviate and
simplify the discussion. No details on program operation are included.
Input and Output
Input to the program consists of data from on or more replications (cohorts) for a
particular population. Each line of input consists of: a replication number (i), animal
number within the replication (j), a pivotal age (x) (see Birch 1948, and example in this
paper), the number of female offspring of the jth animal at age x (rijx), and the number
of male offspring of the fh animal at age x (Nijx). After this data has been inputted, the
program will ask for the pre-imaginal survivorship (r.o) (see Fig. 1).
Output is produced for each replication, and for all replications combined. The follow-
ing quantities are included: intrinsic rate of increase (rm), a Jacknife estimate of r (?j),
an estimate of the standard error of ij (6j), a confidence interval for r, fertility table
parameters (longevity, total progeny production, net reproductive rate (Ro), mean gen-
eration time, doubling time, pre-ovipositional and post-ovipositional periods) and a table
of daily means (mean number of female progeny per adult female (mx), survivorship
(I,), mean number of male progeny per adult female, percent females). Additionally,
approximate standard errors for the estimates of total progeny production, Ro, longev-
ity, and pre- and post-ovipositional periods are included.
In the following two sections we describe the calculation of the quantities and com-
ment on the interpretation of the output. The program also accepts a table of x, mx and
lx values as input, but in that case, not all of the output quantities can be computed. In
particular, &j is unavailable.
602
December, 1990
Hulting et al.: Population Growth Comparisons
***** Population Growth Rate Analysis Program ****
Animal Name: Species A
Temperature: 70 F
Time Interval: 1 day
Preimaginal Survivorship: 0.90
Confidence Level for Intervals: 0.95
***** For All Cohorts Combined *****
Number of Individuals: 10
X I Mx I Lx I Males % Females
-------------------------------------------
10.5 I 3.80 I 0.90 I 1.60 I 70.37
11.5 2.90 0.90 I 1.10 I 72.50
12.5 2.40 0.90 1.10 68.57
13.5 1.40 0.90 1.00 58.33
14.5 0.44 1 0.81 I 0.67 I 40.00
15.5 0.43 0.63 0.57 42.86
16.5 I 0.29 I 0.63 I 0.43 1 40.00
17.5 0.00 0.27 0.33 0.00
18.5 0.33 0.27 I 0.33 I 50.00
19.5 0.00 0.09 0.00 0.00
Complete Data Estimate )f r: 0.199
Jacknife Estimate of r: 0.204
Std Error of Jacknife Estimate: 0.030
Interval Estimate for r: [ 0.136 0.271
Adult Longevity: 7.00 ( 0.615 )
Total Progeny Production: 16.02 ( 1.546 )
Net Reproductive Rate: 10.35 ( 1.100 )
Sex Ratio (% Females): 63.81
Mean Generation Time: 11.77
Doubling Time: 3.49
Pre-Ovipositional Period: 0.00 ( 0.000 )
Post-Ovipositional Period: 1.50 ( 0.373 )
Finite Rate of Increase: 1.22
***** For Cohort 1 *****
Number of Individuals: 5
X I Mx I Lx I Males I % Females
10.5 3.80 0.90 1.60 1 70.37
11.5 3.00 0.90 1.20 71.43
12.5 2.20 0.90 1.00 68.75
13.5 1.40 0.90 1.00 58.33
14.5 0.50 0.72 0.75 I 40.00
15.5 0.50 0.72 0.25 66.67
16.5 0.25 0.72 0.50 33.33
17.5 0.00 0.36 0.50 0.00
18.5 0.50 0.36 0.50 50.00
19.5 0.00 0.18 0.00 0.00
Fig. 1. Program output for data presented in Tables 1 and 2.
603
604 Florida Entomologist 73(4) December, 1990
Complete Data Estimate of r: 0.199
Jacknife Estimate of r: 0.200
Std Error of Jacknife Estimate: 0.015
Interval Estimate for r: [ 0.159 0.241 ]
Adult Longevity: 7.40 ( 1.030 )
Total Progeny Production: 16.20 ( 2.745 )
Net Reproductive Rate: 10.44 ( 2.084 )
Sex Ratio (% Females): 62.31
Mean Generation Time: 11.81
Doubling Time: 3.49
Pre-Ovipositional Period: 0.00 ( 0.000 )
Post-Ovipositional Period: 1.80 ( 0.735 )
Finite Rate of Increase: 1.22
***** For Cohort 2 *****
Number of Individuals: 5
X I Mx I Lx I Males I % Females
10.5 3.80 0.90 1.60 70.37
11.5 2.80 0.90 1.00 73.68
12.5 2.60 0.90 1.20 68.42
13.5 1.40 0.90 1 1.00 58.33
14.5 0.40 0.90 I 0.60 40.00
15.5 0.33 0.54 1.00 25.00
16.5 0.33 0.54 0.33 50.00
17.5 0.00 0.18 0.00 0.00
18.5 0.00 0.18 0.00 0.00
Complete Data Estimate of r: 0.198
Jacknife Estimate of r: 0.199
Std Error of Jacknife Estimate: 0.009
Interval Estimate for r: [ 0.175 0.223
Adult Longevity: 6.60 ( 0.748 )
Total Progeny Production: 15.84 ( 1.791 )
Net Reproductive Rate: 10.26 ( 1.050 )
Sex Ratio (% Females): 65.31
Mean Generation Time: 11.73
Doubling Time: 3.49
Pre-Ovipositional Period: 0.00 ( 0.000 )
Post-Ovipositional Period: 1.20 ( 0.200 )
Finite Rate of Increase: 1.22
Fig. 1. (continued)
Hulting et al.: Population Growth Comparisons
***** Population Growth Rate Analysis Program
Animal Name: Species B
Temperature: 70 F
Time Interval: 1 day
Preimaginal Survivorship: 0.70
Confidence Level for Intervals: 0.95
***** For All Cohorts Combined *****
Number of Individuals: 10
X I MX Lx Males % Females
20.5
21.5
22.5
23.5
24.5
25.5
26.5
27.5
28.5
29.5
0.00
0.20
2.50
3.00
2.78
1.88
1.00
0.80
0.20
0.00
0.70
0.70
0.70
0.70
0.63
0.56
0.49
0.35
0.35
0.14
Complete Data Estimate of r:
Jacknife Estimate of r:
Std Er-or of Jacknife Estimate:
Interval Estimate for r:
0.00
0.30
1.60
3.10
2.78
1.75
1.14
0.60
0.80
1.00
0.085
0.087
0.015
[ 0.053 ,
0.00
40.00
60.98
49.18
50.00
51.72
46.67
57.14
20.00
0.00
0.122 ]
)
)
)
Adult Longevity:
Total Progeny Production:
Net Reproductive Rate:
Sex Ratio (% Females):
Mean Generation Time:
Doubling Time:
Pre-Ovipositional Period:
Post-Ovipositional Period:
Finite Rate of Increase:
***** For Cohort 1 *****
Number of Individuals: 5
X I Mx I Lx I Males I % Females
20.5
21.5
22.5
23.5
24.5
25.5
26.5
27.5
28.5
29.5
0.00
0.20
2.40
2.80
2.75
2.00
1.33
1.50
0.50
0.00
0.70
0.70
0.70
0.70
0.56
0.56
0.42
0.28
0.28
0.28
Fig. 1. (continued)
605
7.60
15.05
7.63
51.65
24.04
8.20
1.70
0.30
1.09
0.670
2.021
1.056
( 0.153 )
( 0.153 )
0.00
0.40
1.60
2.80
2.50
1.50
1.33
1.00
1.00
1.00
0.00
33.33
60.00
50.00
52.38
57.14
50.00
60.00
33.33
0.00
Florida Entomologist 73(4)
December, 1990
Complete Data Estimate of r: 0.084
Jacknife Estimate of r: 0.085
Std Error of Jacknife Estimate: 0.009
Interval Estimate for r: [ 0.060 0.109 ]
Adult Longevity: 7.40 ( 1.166 )
Total Progeny Production: 14.56 ( 3.353 )
Net Reproductive Rate: 7.56 ( 1.804 )
Sex Ratio (% Females): 53.16
Mean Generation Time: 24.18
Doubling Time: 8.28
Pre-Ovipositional Period: 1.60 ( 0.245
Post-Ovipositional Period: 0.40 ( 0.245 )
Finite Rate of Increase: 1.09
***** For Cohort 2 *****
Number of Individuals: 5
X I Mx | Lx | Males I % Females
20.5 0.00 0.70 0.00 0.00
21.5 0.20 0.70 0.20 50.00
22.5 2.60 0.70 1.60 61.90
23.5 3.20 0.70 3.40 48.48
24.5 2.80 0.70 3.00 48.28
25.5 1.75 0.56 2.00 46.67
26.5 0.75 0.56 1.00 42.86
27.5 0.33 0.42 0.33 50.00
28.5 0.00 0.42 0.67 0.00
Complete Data Estimate of r: 0.085
Jacknife Estimate of r: 0.086
Std Error of Jacknife Estimate: 0.007
Interval Estimate for r: [ 0.067 0.105 ]
Adult Longevity: 7.80 ( 0.800 )
Total Progeny Production: 15.54 ( 2.651 )
Net Reproductive Rate: 7.70 ( 1.328
Sex Ratio (% Females): 50.13
Mean Generation Time: 23.90
Doubling Time: 8.12
Pre-Ovipositional Period: 1.80 ( 0.200 )
Post-Ovipositional Period: 0.20 ( 0.200 )
Finite Rate of Increase: 1.09
Fig. 1. (continued)
Hulting et al.: Population Growth Comparisons
Computational Details
In what follows we will drop the "i" subscript and assume we are working with a
particular replication (or a new "replication" formed from all individuals in all replica-
tions). Define, for a given replication:
n = number of individuals in the replication
x = preimaginal development time
f = maximum age among the n individuals,
1j = age at death for the f^ individual,
= 1 if f individual is alive at age x
Tjx ~0 otherwise,
x1) = age at which fh individual produces first progeny
and
x(2) = age at which jth individual produces last progeny.
Ta 1
a=1
j=1
( ,J=1 7xo j=1
In
adult longevity = -
l l l
total progeny production per adult female
=XT ax)
in (
Sj= X=X-
T j=1 X=xo
m x)
daily sex ratio
overall sex ratio =
j=1
mean generation time
doubling time
S,=1i +rjx
Za-i(+x + N)
-x=xo 1lax
ZX=(~7ax Nax)]
ln(R,)
ln(2)
nio mx
n
Vt
a=1
Gxo
X= X m
j=1
- -js l
608 Florida Entomologist 73(4) December, 1990
1n
pre-ovipositional period = (x x )
and j=1
1
post-ovipositional period = X- (j xj(2))
j=1
Note that adult longevity, Ro, total progeny production, and pre- and post-ovipositional
period can be written as
1 n"
n
j=1
for suitably defined "y" (e.g., for longevity, yj = = x Tj). An estimated standard
error for these quantities is calculated as
1n
n(n 1) j=
The intrinsic rate of increase, rm, is defined as the solution to
1 = e-rXlxmx
X=Xo
which is a nonlinear equation in r. We solve Equation 1 using the Dekker-Brent method
(Press et al. 1986), a general-purpose, derivative-free method for solving nonlinar equa-
tions in a single variable. The procedure requires two initial values for r which bracket
the solution-we use 0 and 1. The finite rate of increase is calculated as em.
The Jacknife (Tukey 1958) is a general nonparametric procedure for obtaining esti-
mated standard errors for statistics which are complex functions of the data. Meyer et
al. (1986) use the jacknife to obtain standard errors for intrinsic rates of increase. A
brief description of the technique follows.
Let rm be the solution to Equation 1 obtained using all the data, and let rmj(j =
1,...,n) be the solution to Equation 1 obtained after dropping individual j out of the
data set. Then the Jacknife "pseudo-values" are computed as
r = nrm (n l)rmj.
The pseudo-values are used to construct a new point estimate of r
I n
j =1 V- -j
j=1
and an estimate of the standard error of Pj
Jn(
r n(n 1) j=
Hulting et al.: Population Growth Comparisons 609
If we assume that the rj are independent draws from an approximately normal distribu-
tion with mean r, an approximate 100(1 a)% confidence interval for r is given by
J - tGS aj
where v = n-1 and t, is the upper- a point of the Students t-distribution with v
2 2
degrees of freedom. In this program, the value of t is obtained using the procedure
of Koehler (1983).
Interpretation of Output
The confidence interval provided by the program enables the user to draw conclu-
sions about the true r value for the particular species under consideration. Note that
the interval is approximate; an assessment of the performance of the interval for
cladoceran (Daphnia pulex DeGeer) populations is given in Meyer et al. (1986).
The output from the program can also be used to statistically compare rm values
for two or more populations (species, biotypes, etc). First consider the comparison of
two species. Let 1') and &j1) be the Jacknife estimates for species 1 and let *52) and
oj2) be the estimates for species 2 (all numbers would be taken from the program
output). Now if r(1) and r(2) are the true population growth rates for the species, then
an approximate 100(1 a)% confidence interval for r() r(2 is
1(i) _42) (41 2 /+2h2
rj tfj) a)2 (2)
where
nj -1 n2-1
and nI and n2 are the number of animals in the data sets from species 1 and 2, respec-
tively. The degrees of freedom approximationfis that proposed by Satterthwaite (1946).
Intervals of the form (2) are discussed by Snedecor and Cochran (1980, sec. 6.11). Note
that may not be an integer; it should be rounded off for use with a t-table. Conservative
intervals may be obtained by replacing f with 2 -1. An interval which does not
2
include 0 can be viewed as evidence of a difference between the r values of the two
species. As in the last section, this interval is based on assumptions of independence
and approximate normality.
The above interval should be used to make specific pairwise comparisons. If more
than two species are being considered, and it is desired to make all pairwise compari-
sons, a multiple comparisons procedure must be used. There are many such procedures;
one choice is the Newmann-Keuls sequential test. This test requires the calulation of a
Q-statistic for different sets of species (see Snedecor and Cochran 1980, sec. 12.13 for
details on the procedure). Here we indicate how to compute the Q-statistic for a given
set of species. Specifically, consider species 1,2,...,k. Let ')...,*k) be the Jacknife
estimates of the population growth rates. Further, let j(A) be the maximum and jSB)
be the minimum of fj'),..., (k) and let &(A) and &(B) be the associated standard errors.
Then the Q-statistic is calculated as
Florida Entomologist 73(4)
December, 1990
,(A) (B)
Q= -- 2'
/&AA -2 1 ((B
its associated degrees of freedom are 2
Example
In the following example, we compare two hypothetical species (A and B) of parasitic
wasps. Species A has a mean preimaginal developmental time of 10 days and preimaginal
survivorship of 0.90; species B has corresponding values of 20 days and 0.70, respec-
tively. Note that this data could be from previous studies or could be collected from the
r,, study population. Also note that although data on male and female progeny produc-
tion are presented in this example, where this information may be difficult or impossible
to collect it may be necessary to multipy progeny production values by the total popu-
lation sex ratio to obtain m. values.
Tables 1 and 2 present 'raw data' for two cohorts (or replications) for each species.
Program output is given in Figure 1. In this example, the population growth rate for
Species A is estimated (using both replications) to be 0.20, and an associated confidence
interval is [0.14, 0.27] (while the program reports 3 significant figures, we follow the
recommendation of Meyer et al. (1986), and report only two significant figures here).
The values for Species B are 0.09 and [0.05, 0.12]. These results indicate a difference
in values between the two species. A better comparison is made using the 95% confi-
dence interval for the difference in r values. It is
(0.20 0.09) (2.16)(0.03)2 + (0.015)2
.11 .072
[.038, .182]
which does not include 0. Thus we would conclude that the population r values are
different. For this interval we used degrees of freedom
TABLE 1. PROGENY PRODUCTION (9,d) BY TWO
HYPOTHETICAL PARASITOID SPECIES
FOUND DEAD).
COHORTS REPLICATIONSS) OF
A (D=DAY PARASITOID WAS
Adult Pivotal COHORT I COHORT II
Adult Pivotal
Age(d) Age(d) 91 92 93 94 95 91 92 93 94 95
1 10.5 5,1 6,2 2,2 3,1 3,2 4,2 3,1 5,2 3,2 4,1
2 11.5 4,1 3,1 1,2 4,1 3,1 3,1 2,1 4,1 3,1 2,1
3 12.5 2,0 3,2 1,1 3,1 2,1 3,1 3,1 3,2 2,1 2,1
4 13.5 2,1 2,2 0,OD 2,1 1,1 2,1 1,1 2,1 1,1 1,1
5 14.5 0,1 1,1 1,1 0,0 1,1 0,OD 1,1 0,1 0,OD
6 15.5 0,0 0,0 2,1 0,0 0,1 0,1 1,1
7 16.5 0,1D 0,0 1,1 0,OD 0,OD 0,OD 1,1
8 17.5 0,0 0,1 0,0
9 18.5 0,OD 1,1 0,OD
10 19.5 0,OD
Hulting et al.: Population Growth Comparisons
TABLE 2. PROGENY PRODUCTION (9,6) BY TWO
HYPOTHETICAL PARASITOID SPECIES
FOUND DEAD).
COHORTS REPLICATIONSS) OF
B (D=DAY PARASITOID WAS
COHORT I COHORT II
Adult Pivotal
Age(d) Age(d) 91 22 23 94 25 21 92 93 94 95
1 20.5 0,0 0,0 0,0 0,0 0,0 0,0 0,0 0,0 0,0 0,0
2 21.5 0,0 1,1 0,1 0,0 0,0 1,1 0,0 0,0 0,0 0,0
3 22.5 4,3 2,1 1,1 3,1 2,2 3,2 3,2 3,2 2,1 2,1
4 23.5 5,3 0,OD 3,3 2,3 4,5 5,4 4,4 2,1 2,4 3,4
5 24.5 0,1 4,2 4,4 3,3 4,3 1,1 0,1D 3,5 6,5
6 25.5 0,OD 2,2 5,2 1,2 2,2 1,2 2,1 2,3
7 26.5 1,2 2,1 1,1D 0,1 1,1D 1,1 1,1
8 27.5 1,1 2,1 1,1 0,0 0,0
9 28.5 0,1 1,1 0,OD 0,1D 0,1D
10 29.5 0,1D 0,1D
((0.030)2 +(0.015)2)2
f = (0.030)4 (0.015)4
9 9
= 13.23 m 13.
In order to test the accuracy of this program, we analyzed data tabulated by Birch
(1948) for Sitophilus oryzae (L.) (Coleoptera: Curculionidae). The computed values were
the same as those presented by Birch (1948), as follows: rm = 0.762; R, = 113.48;
T = 6.21.
This program can be run on IBM PC-compatible microcomputers or DEC VAX
minicomputers. A copy of the compiled program (for IBM compatibles) or the program
code will be provided along with a manuscript reprint if a formatted 51/4 in. diskette is
sent to any of the authors. The program is written in standard Pascal, so it can be
compiled on a wide variety of computers. The program has been run on IBM compatible
microcomputers, DEC VAX minicomputers and Sun workstations.
ACKNOWLEDGMENTS
The authors are grateful for the technical assistance provided by J. Robison-Cox
and A. E. Morse in preparation of this manuscript. This work was supported in part
by a Natural Sciences and Engineering Research Council of Canada Postdoctoral Fel-
lowship to DBO. Journal paper J-13821 of the Iowa Agriculture and Home Economics
Experiment Station, Ames, Iowa.
REFERENCES CITED
ABOU-SETTA, M. M., R. W. SORRELL, AND C. C. CHIDERS. 1986. Life 48: A BASIC
computer program to calculate life table parameters for an insect or mite species.
Florida Entomol. 69: 690-697.
ANDREWARTHA, H. G., AND L. C. BIRCH. 1954. The distribution and abundance of
animals. Univ. of Chicago Press, Chicago.
BIRCH, L. C. 1948. The intrinsic rate of natural increase of an insect population. J.
Anim. Ecol. 17: 15-26.
FORCE, D. C. 1974. Ecology of insect host-parasitoid communities. Science 184: 624-
632.
Florida Entomologist 73(4)
FORCE, D. C., AND P. S. MESSENGER. 1964. Fecundity, reproductive rates, and
innate capacity for increase of three parasites of Therioaphis maculata
(Buckton). Ecology 45: 706-715.
GASTON, K. J. 1988. The intrinsic rates of increase of insects of different sizes. Ecol.
Entomol. 14: 399-409.
HIROSE, Y. 1986. Biological and ecological comparison of Trichogramma and Tele-
nomus as control agents of lepidopterous pests. Z. ang. Entomol. 101: 39-47.
KOEHLER, K. J. 1983. A simple approximation for the percentiles of the t distribution.
Technometrics 25: 103-105.
LAUGHLIN, R. 1965. Capacity for increase: a useful population statistic. J. Anim.
Ecol. 34: 77-91.
MESSENGER, P. S. 1964. Use of life tables in a bioclimatic study of an experimental
aphid-braconid wasp host-parasite system. Ecology 45: 119-131.
MESSENGER, P. S. 1970. Bioclimatic inputs to biological control and pest management
programs, pp. 84-99. In R. L. Rabb, and F. E. Guthrie (eds.), Concepts of pest
management. North Carolina State Univ., Raleigh.
MEYER, J. S., C. G. INGERSOLL, L. L. MCDONALD, AND M. S. BOYCE. 1986.
Estimating uncertainty in population growth rates: Jacknife vs. Bootstrap tech-
niques. Ecology 67: 1156-1166.
NECHOLS, J. R., J. L. TRACY, AND E. A. VOGT. 1989. Comparative ecological studies
of indigenous egg parasitoids (Hymenoptera: Scelionidae; Encrytidae) of the
squash bug, Anasa tristus (Hemiptera: Coreidae). J. Kans. Entomol. Soc. 62:
177-188.
ORPHANIDES, G. M., AND D. GONZALES. 1971. Fertility and life table studies with
Trichogramma pretiosum and T. retorridum (Hymenoptera: Trichogram-
matidae). Ann. Entomol. Soc. Am. 64: 824-834.
PRESS, W. H., B. P. FLANNERY, S. A. TEUKOLSKY, AND W. T. VETTERLING.
1986. Numerical Recipes: The Art of Scientific Computing. Cambridge Univer-
sity Press, Cambridge, MA.
SATTERTHWAITE, F. E. 1946. An approximate distribution of estimates of variance
components. Biometrics Bulletin 2: 110-114.
SNEDECOR, G. W., AND W. G. COCHRAN. 1967. Statistical Methods (6th ed.) Iowa
State University Press, Ames, IA.
SOUTHWOOD, T. R. E. 1978. Ecological Methods. Chapman & Hall, London.
TUKEY, J. W. 1958. Bias and confidence in not quite large samples. Annals of
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December, 1990
612
Matthews et al.: Pterophoridae of Florida 613
ANNOTATED CHECKLIST OF THE PTEROPHORIDAE
(LEPIDOPTERA) OF FLORIDA INCLUDING
LARVAL FOOD PLANT RECORDS
D. L. MATTHEWS, D. H. HABECK
Department of Entomology and Nematology
D. W. HALL
Department of Natural Sciences
IFAS, University of Florida
Gainesville, Florida 32611
ABSTRACT
A checklist of the Pterophoridae of Florida includes 32 species with larval host plant
records for 27 species. Species changes and updated nomenclature since Kimball (1965)
are noted.
RESUME
Se da una lista de los Pterophoridae de la Florida que incluye 32 species con el
registro de sus plants hospederas para 27 species. Se incluyen los nuevos cambios de
species y de nomenclatura desde Kimball (1965).
The following list provides an update of the plume moth fauna of Florida with known
larval hosts for each species. Kimball (1965) reported 27 species in Florida but there
have been several additions to synonomies and changes in generic assignments as re-
flected in the most recent checklist of North American Pterophoridae by Munroe (1983).
Nine species are added to the Florida fauna: Sphenarches anisodactylus (Walker),
Trichoptilus pygmaeus Walsingham, Platyptilia carduidactyla (Riley), Stenoptilodes
auriga (Barnes & Lindsey), Exelastis sp., Oidaematophorus eupatorii (Fernald), O.
paleaceus (Zeller), 0. glenni Cashatt, and Oidaematophorus sp. C. Five names are
synonomized or omitted (see Table 1 for details). The present list includes 32 species
arranged according to Munroe (1983) with recent additions and generic assignments in
accordance with Buszko (1979) and Prola & Racheli (1984).
Information on hosts in Florida and outside Florida is included under the heading
"Larval hosts". Each host species is followed by its common name(s) and a literature
citation or place of specimen depository. Where no citation is given, the moth species
was either reared or collected by the authors on the host listed. Synonyms of host
species are included in parentheses if they appear on specimen labels or in the references
cited. Museum abbreviations are USNM, United States National Museum, Washington,
D. C.; MCZ, Museum of Comparative Zoology, Cambridge, Mass.; FSCA, Florida State
Collection of Arthropods, Gainesville, Fla.; CMNH, Carnegie Museum of Natural His-
tory, Pittsburgh, Pa. Specimens from other museum collections were examined in addi-
tion to many private collections (see acknowledgments). Aspects of the biology of each
species are included under the heading "Notes".
Four species, Exelastis sp., Oidaematophorus sp. A, B, and C are unnamed, new
to the North American fauna, and may be undescribed. Pending examination of addi-
tional material these species will be treated accordingly in future publications. Species
accounts including life histories, distributions, adult and larval descriptions, and keys
to Florida species are described and figured in Matthews (1989).
Florida Entomologist 73(4)
TABLE 1. SUMMARY OF CURRENT STATUS OF NAMES REPORTED IN KIMBALL (1965).
Kimball # Kimball species name Remarks
6459
6460
6461
6471
6472,1
6483
6483,1
6488
6490
6503
6504
6505
6506
6508
6509
6516
6517
6521
6522
6546
6554,1
6563
6566
6568
6569
6581
6588
Trichoptilus parvulus
Trichoptilus defectalis
Trichoptilus californicus
Platyptilia pusillidactyla
Platyptilia brevipennis
Platyptilia brachymorpha
Platyptilia taprobanes
Platyptilia carolina
Platyptilia edwardsii
Exelastis cervinicolor
Marasmarcha pumilio
Stenoptilia rhynchosiae
Stenoptilia parva
Stenoptilia zophodactyla
Stenoptilia pallistriga
Pselnophorus belfragei
Adaina bipunctata
Adaina buscki
Adaina ambrosiae
Oidaematophorus inquinatus
Oidaematophorus sp.
Oidaematophorus stramineus
Oidaematophorus
venapunctatus
Oidaematophorus
lacteodactylus
Oidaematophorus balanotes
Oidaematophorus unicolor
Oidaematoohorus
monodactylus
There is little doubt that more species will be discovered in the state. Many larvae
feed only on very young foliage or flowers, so the seasonal occurrence of several plume
moths is closely correlated with the flowering phenologies of their hosts. Because some
Current genus is Megalorhipida.
Current genus is Lantanophaga.
Species record based on early
literature record. No specimens
in museum collections examined.
Figure in original description
very close to M. taprobanes,
probably a synonym.
Both brachymorpha Meyrick and
crenulata B & McD. are synonyms
of Mariana taprobanes.
Current genus is Mariana.
Current genus is Stenoptilodes.
Northern species, found principally
in Canada. Kimball's specimens in
MCZ labelled as this species are
actually M. taprobanes.
Current name is Lioptilodes parvus.
The USNM specimen Kimball
reported was identified as S.
pallistriga.
Included in present checklist as
Oidaematophorus sp. A.
The 3 specimens listed in Kimball
from Univ. Michigan collection
were examined and found to be
O. paleaceus.
Kimball's specimens were identified
as Oidaematophorus sp. B.
Form kellicottii is now separate
species.
Current name is Emmelina
monodactyla.
December, 1990
Matthews et al.: Pterophoridae of Florida 615
of the hosts flower for only short periods and are restricted to specific habitats, they
can be easily missed by the general collector. New species may also become established
as the result of agricultural importations. For example, Amblyptilia pica (Wlsm.) has
been intercepted at Florida ports on cultivated Geranium (FSCA records, see also
Valley et al., 1981). Much collecting still needs to be done in the Florida Keys, northern
Florida, and especially in the Florida panhandle where typically northern species may
be found as relict populations of the Appalachian fauna.
SUBFAMILY PLATYPTILIINAE
Genus Sphenarches Meyrick
1. S. anisodactylus (Walker)
Larval hosts: Florida, Thalia geniculata L. (alligator flag) [Marantaceae]; outside
Florida Dolichos lablab L. lablabb), Cajanus cajan (L.) Huth (pigeon pea)
[both Fabaceae], Caperonia sp. [Euphorbiaceae] (USNM), Lagenaria
leucantha Rosby var. clavata and gourda [Cucurbitaceae] (Yano 1963).
Notes: The life history of this species in Florida is described by Cassani et al.
(1990). Larvae feed primarily on the flowers of the hosts.
Genus Trichoptilus Walsingham
2. T. parvulus Barnes & Lindsey
Larval hosts: Florida, Drosera brevifolia Pursh (dwarf sundew), D. intermedia
Hagne (spoon-leaved sundew), D. filiformis Raf. (dew threads)
[Droseraceae]; outside Florida, Drosera filiformis Raf. var. tracyi Diels
(thread-leaved sundew) (USNM).
Notes: Larvae feed on glandular trichomes, leaves, and flowers of the host.
Larval behavior is described by Eisner & Shepherd (1965). Larvae in Florida
are heavily infected by a species of Cotesia (Braconidae).
3. T. californicus (Walsingham)
Larval hosts: Florida, unknown; outside Florida, Isocoma veneta (HBK.) Greene,
Grindelia sp. [both Asteraceae] (Lange 1939).
Notes: Life history in California is described by Lange (1939).
4. T. pygmaeus Walsingham
Larval hosts: Florida, Chrysopsis scabrella Torr. & Gray (rough-leaf golden
aster) [Asteraceae]; outside Florida, Arctostaphylos columbiana Piper
[Ericaceae] (USNM).
Notes: Larvae feed on young leaves and the bracts of unopened flowers of
Chrysopsis.
Genus Megalorhipida Amsel
5. M. defectalis (Walker)
Larval hosts: Florida, Boerhavia diffusa L. (red spiderling), Okenia hypogaea
Schlecht. & Cham. (burrowing four-o'clock) [both Nyctaginaceae]; outside
Florida, Acacia neovernicosa Isely (C. vernicosa Standl.) [Fabaceae]
(USNM), Boerhavia diffusa L. (B. coccinea Mill) [Nyctaginaceae] (Zimmer-
man 1958), Amaranthus sp [Amaranthaceae] (Wolcott 1936).
Notes: On Boerhavia, larvae feed on flower buds and bore into the fruits. Larvae
on Okenia feed only on leaves. In Florida, 0. hypogaea is an endangered
species restricted to a few locations on the southeast coast of Florida (Ward
1979).
616 Florida Entomologist 73(4) December, 1990
Genus Platyptilia Hiibner
6. P. carduidactyla (Riley)
Larval hosts: Florida, unknown; (outside Florida, Cynara scolymus L. (ar-
tichoke), C. cardunculus L. (cardoon), Cirsium edule Nutt., C. vulgare (Sari)
Ten. (C. lanceolatum Scop.), C. occidentale (Nutt.) Jeps., C. quercetorum
(Gray) Jeps., C. discolor (Muhl.) Sprengel, C. undulatum (Nutt.) Sprengel,
C. arvense (L.) Scop. (Canada thistle), C. callilepis (Greene) Jeps. (C.
americanum Daniels), Centaurea melitensis L. (Tocalote), Silybum
marianum (L.) Gaertn. (milk thistle) [all Asteraceae] (Lange 1950).
Notes: This species is known in Florida from two specimens, one collected at the
Archbold Biological Station by H. V. Weems, 25-IX-1978, the other by H. O.
Hilton in Ocean City Florida, 22-X-1961.
Genus Lantanophaga Zimmerman
7. L. pusillidactyla (Walker)
Larval hosts: Florida, Lantana camera L. (lantana) [Verbenaceae], Caperonia
sp. [Euphorbiaceae] (Kimball 1965), also reared from the seed heads of Lippia
(=Phyla) lanceolata Michx. (northern frogfruit) [Verbenaceae] by C. E.
Stegmaier (MCZ). The latter host species was probably Phyla nodiflora (L.)
Green (match-head) since lanceolata does not occur in Florida. Outside
Florida, Lantana camera L.
Notes: Larvae feed on flowers and fruits of the hosts.
Genus Lioptilodes Zimmerman
8. L. parvus (Walsingham)
Larval hosts: Solidago odora Ait. (sweet goldenrod) (MCZ), Erigeron strigosus
Muhl. (daisy fleabane), Aster subulatus Michx. (annual marsh aster), Aster
puniceus L. subsp. elliottii (Torr. & Gray) A. G. Jones (Elliott's aster) [all
Asteraceae]; outside Florida, none reported.
Notes: Larvae feed inside flower heads, ova are usually deposited on flower
bracts.
Genus Mariana Tutt
9. M. taprobanes (Felder & Rogenhofer)
Larval hosts: Florida, C. E. Stegmaier reared this species on sweet broom,
Scoparia dulcis L. [Scrophulariaceae] (MCZ) and collected a larva on Mercar-
donia acuminata [Scrophulariaceae] (USNM), a specimen collected by P.
Perun was reared on Hydrolea quadrivalis Walt. [Hydrophyllaceae]; outside
Florida, Ocimum sp. (basil) and Plectranthus sp. [both Lamiaceae], Anti-
rrhinum majus L. (snapdragons) [Scrophulariaceae] (Zimmerman 1958), in
fruits of Limnophila heterphylla Benth., in seeds of Penstemon sp., and un-
ripe fruits of Veronica anagallis-aquatica L. (cited as V. anagallis L.) Ver-
bascum coromandelianum (Vahl) 0. Kuntze, (Celisia coromandeliana Vahl)
[all Scrophulariaceae] (Lange 1950), Russelia equistiformis Schlecht. &
Cham. (firecracker plant) [Scrophulariaceae] (USNM).
Notes: Mariana taprobanes was reported by Kimball (1965) from Lippia
(= Phyla) lanceolata Michx. (northern frogfruit) but these are Lantanophaga
pusillidactyla, based on examination of the genitalia of these specimens in
the MCZ.
Matthews et al.: Pterophoridae of Florida
Genus Stenoptilodes Zimmerman
10. S. carolina (Kearfott)
Larval hosts: unknown.
Notes: This species is included with some reservation since no specimens from
Florida were examined. Kimball (1965) reported two specimens from Escam-
bia county.
11. S. auriga (Barnes & Lindsey)
Larval hosts: Florida, unknown; outside Florida: Gerardia sp. [Scrophulariaceae]
and various species of Asteraceae (Neunzig 1987).
Notes: The only specimen known from Florida, a male, was collected 29-III-1980
at Torreya state park by Charlie Stevens.
Genus Stenoptilia Hiibner
12. S. pallistriga Barnes & McDunnough
Larval hosts: unknown.
13. S. rhynchosiae (Dyar)
Larval hosts: Florida, Rhyncosia cinerea Nash (ashy rhyncosia) [Fabaceae]; out-
side Florida, none reported.
Notes: Larvae feed on young leaves and shoots.
Genus Exelastis Meyrick
14. E. cervinicolor (Barnes & McDunnough)
Larval hosts: unknown.
15. Exelastis. sp.
Larval hosts: unknown.
Notes: This species is known from Florida by a series of specimens collected
11-III-1986 on Key Largo by Linwood C. Dow.
Genus Marasmarcha Meyrick
16. M. pumilio (Zeller)
Larval hosts: Florida, Desmodium tortuosum (S.W.) DC. D. incanum DC.
[Fabaceae]; outside Florida, Desmodium sp.
Notes: Some larvae feed exclusively on flowers while other individuals
skeletonize young leaves and shoots. Barnes & Lindsey (1921) report, with
skepticism, a record for Ambrosia artemisiifolia L. (common ragweed) [As-
teraceae] which was also cited by Kimball (1965). This host is unlikely since
members of this genus are chiefly legume feeders.
SUBFAMILY PTEROPHORINAE
Genus Pselnophorus Wallengren
17. P. belfragei (Fish)
Larval hosts: Florida, no larvae have been field collected but several have been
successfully reared from eggs on Dichondra caroliniensis Michx. (pony-foot)
[Convolvulaceae]; outside Florida, none reported.
Notes: This species is the most commonly encountered plume moth in Florida.
617
618 Florida Entomologist 73(4) December, 1990
Genus Adaina Tutt
18. A. bipunctata (Moeschler)
Larval hosts: Florida, Conoclinium coelestinum (L.) DC. (ageratum or mist
flower) (Stegmaier 1973), Carphephorus paniculatus (J. F. Gmel.) Herb.
(hairy trilisa), C. ordoratissimus (J. F. Gmel.) Herb. (vanilla plant), and
Pluchea rosea Godfrey [all Asteraceae]; outside Florida, Eupatorium can-
nabinum L. (hemp agrimony) [Asteraceae] (Wasserthal 1970).
Notes: Larvae feed within the composite flower heads.
19. A. buscki Barnes & Lindsey
Larval hosts: Florida, Ipomoea indica (Burm.f.) Merr. (blue morning-glory)
[Convolvulaceae]; outside Florida, unknown.
Notes: Larvae skeletonize young leaves.
20. A. ambrosiae (Murtfeldt)
Larval hosts: Florida, Ambrosia artemisiifolia L. (common ragweed), Pluchea
rosea Godfrey (Godfrey's fleabane), Melanthera nivea (L.) Small (M. deltoidea
Michx. (cat-tongue)); outside Florida, Ambrosia acanthicarpa Hook., A.
chamissonis (Less.) Greene, A. confertiflora DC., A. dumosa (A. Gray)
Payne, A. eriocentra (A. Gray) Payne, A. cumanensis HBK. (A. psilostachya
DC.) (western ragweed), Helianthus annuus L. (annual sunflower), Xanth-
ium strumarium L. (cocklebur) (Goeden & Ricker 1976), Helianthus tuberosa
L. (Jerusalem artichoke), and Cynara scolymus L. (artichoke) (USNM) [all
Asteraceae].
Notes: Larvae skeletonize leaves, and when not feeding rest along the midrib of
the upper leaf surface in shallow, elongated depressions which they carve into
the leaves and to which they repeatedly return after feeding bouts.
Genus Oidaematophorus Wallengren
21. 0. eupatorii (Fernald)
Larval hosts: Florida, unknown; outside Florida, Eupatorium sp., E. purpuras-
cens Sch. Bip. [Asteraceae] (Forbes 1923), Epilobium sp. [Onagraceae] (Dyar
1898).
Notes: Larvae feed externally and are gregarious, feeding on and tying together
the terminal shoots of the host with webbing (Barnes & Lindsey 1921). This
species is known from Florida by three specimens collected 17-V-1970 at Tor-
reya State Park by H. V. Weems (FSCA).
22. 0. inquinatus Zeller
Larval host: Florida, Ambrosia artemisiifolia L. (common ragweed) [As-
teraceae], outside Florida, same species.
Notes: Larvae feed externally on young foliage.
23. 0. paleaceus (Zeller)
Larval hosts: Florida, Vernonia gigantea (Walt.) Trel. ex Branner & Coville
(ironweed) [Asteraceae]; outside Florida, V. noveboracensis (Barnes &
Lindsey 1921) and V. missurica (Godfrey et al. 1987).
Notes: Larvae feed on young foliage.
24. 0. balanotes (Meyrick)
Larval hosts: Florida, Baccharis halimifolia L. (saltbush, sea myrtle, or
groundsel bush) [Asteraceae]; outside Florida, same species.
Notes: First instar larvae are leafminers while older larvae bore into the stems
and form extensive galleries in the larger branches of the host. There are
records of this species on Myrica sp. (wax myrtle) [Myricaceae] (USNM),
__
_^^^
Matthews et al.: Pterophoridae of Florida
however, we feel these are most likely misidentified Baccharis. Both plants
frequently occur in the same habitats and to an untrained eye are superficially
quite similar.
25. 0. kellicottii (Fish)
Larval hosts: Florida, Solidago canadensis L. (Canada goldenrod), S. fistulosa
Mill. (hollow goldenrod), S. gigantea Ait. (S. leavenworthii Torr. & Gray)
(giant goldenrod), (Fontes 1985); outside Florida, Solidago sp. (USNM), Bac-
charis neglecta Britton (Palmer 1987).
Notes: Larvae are stemborers.
26. 0. lacteodactylus (Chambers)
Larval hosts: Florida, unknown; outside Florida, Solidago sp. (USNM),
Eupatorium perfoliatum L. (boneset) (Godfrey et al. 1987) [both Asteraceae].
Notes: Larvae are stemborers. Kimball (1965) reported this species from Bac-
charis halimifolia L. However, numerous FSCA larval specimens labelled as
0. lacteodactylus from B. halimifolia were examined and determined to be
young larvae of 0. balanotes.
27. 0. glenni Cashatt
Larval hosts: Florida, unknown; outside Florida Solidago canadensis L. (Canada
goldenrod) [Asteraceae] (Cashatt 1972).
Notes: Larvae are stemborers. This species is recorded in Florida from two
specimens, one from St. Lucie county, the other from Walton county
(CMNH).
28. 0. unicolor (Barnes & McDunnough)
Larval hosts: Florida, Eupatorium capillifolium (Lam.) Small (dog fennel) [As-
teraceae]; outside Florida, unknown.
Notes: Larvae are borers in the stems and roots of the host.
29. Oidaematophorus sp. A
Larval hosts: Florida, Eupatorium capillifolium (Lem.) Small (dog fennel), and
E. compositifolium Walt. (dog fennel) [Asteraceae]; outside Florida; un-
known.
Notes: Larvae feed externally on the host.
30. Oidaematophorus sp. B
Larval hosts: Florida, Haplopappus divaricatus (Nutt.) A. Gray (scratch daisy)
and Conyza canadensis (L.) Cronq. horseweedd) [both Asteraceae]; outside
Florida, unknown.
Notes: Kimball (1965) identified specimens of this species as 0. venapunctatus
which is similar in maculation. The genitalia of Kimball's specimens (MCZ)
were compared with paratypes of venapunctatus and are distinctly different.
Larvae are stemborers and are commonly attacked by a braconid wasp, Bra-
con sp.
31. Oidaematophorus sp. C
Larval hosts: unknown.
Notes: Adults of this species are similar in size and maculation to Adaina am-
brosiae. The male genitalia are typical of Oidaematophorus. This species is
known from specimens collected on Big Pine Key by M. Hennessey.
Genus Emmelina Tutt
32. E. monodactyla (Linnaeus)
Larval hosts: Florida, unknown; outside Florida, Calystegia sepium (L.) R. Br.
(hedge bindweed), C. spithamaea (L.) R. Br. (low bindweed), Convolvulus
arvensis L. (field bindweed), C. microphyllus, Ipmoea batatas (L.) Lam.
619
Florida Entomologist 73(4)
(sweet potato), I. hispida Parodi [all Convolvulaceae], Hyoscyamus niger L.
henbanee), Datura stramonium L. (jimson weed) [both Solanaceae] (Parrella
& Kok 1978), Ipomoea purpurea (L.) Roth (tall morning-glory) [Convol-
vulaceae] (Godfrey et al. 1978), Chenopodium sp. and Atriplex sp.
[Chenopodiaceae] (Buszko 1979).
Notes: On Convolvulaceae hosts, early instar larvae feed on terminal shoots
while older larvae tunnel into flowers and feed on the reproductive structures
within.
ACKNOWLEDGMENTS
The authors thank the following persons and institutions for the loan of museum
specimens: Douglas Ferguson, United States National Museum, Charles Vogt, Museum
of Comparative Zoology, Howard Weems, Florida State Collection of Arthropods, John
Rawlins, Carnegie Museum of Natural History, Frederick Rindge, American Museum
of Natural History, and Julian Donahue, Los Angeles County Museum. The following
people are also gratefully acknowledged for the loan of material from their private
collections: Linwood Dow, Dave Baggett, Charlie Stevens, Terhune Dickel, John Hepp-
ner, and Bryant Mather. Several individuals also provided us with many specimens
which were used in compiling this list: Steve Steinhauser, Lee Miller, Jacqueline Miller,
Paul Skelley, Terry Lott, Mike Hennessey, Rick Gillmore, Don Hall, James Trager,
and John Watts. Lee and Jacqueline Miller, Allyn Museum of Entomology/Florida
Museum of Natural History, assisted in acquiring and transporting specimens on loan
from major museums. Tomas Zoebisch provided the Spanish abstract. We thank also
Everett D. Cashatt, Illinois State Museum, and Jacqueline Miller, Allyn Museum of
Entomology, for reviewing the manuscript. Florida Agricultural Experiment Station,
Journal Series No. R-00822.
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Mexico. Contrib. Nat. Hist. Lepid. North America 4: 281-483.
BUSZKO, J. 1979. Klucze do oznaczania owadow Polski. Polskie Towarzystwo En-
tomologiczne. 27: 1-140.
CASHATT, E. D. 1972. Notes on the balanotes (Meyrick) group of Oidaematophorus
Wallengren with description of a new species (Pterophoridae). J. Lepid. Soc.
26(1): 1-13.
CASSANI, J., D. H. HABECK, AND D. L. MATTHEWS. 1990. Life history of a plume
moth Sphenarches anisodactylus (Lepidoptera: Pterophoridae). Florida En-
tomol. 73(2): 257-266.
DYAR, H. G. 1898. Six new or little known larvae of Pterophoridae. Psyche 8: 249-250.
EISNER, T., AND J. SHEPHERD. 1965. Caterpillar feeding on a sundew plant. Science
150: 1608-1609.
FONTES, M. G. 1985. The diversity of the insect fauna of four species of Solidago
(goldenrods) in Gainesville and its relationship to plant architecture. Ph.D. Dis-
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FORBES, W. T. M. 1923. Lepidoptera of New York and neighboring states. Mem.
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GODFREY, G. L., E. D. CASHATT, AND M. 0. GLENN. 1987. Microlepidoptera from
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Hist. Survey Spec. Publ. 7.
GOEDEN, R. D., AND D. W. RICKER. 1976. Life history of the ragweed plume moth
Adaina ambrosiae (Murtfeldt), in Southern California. (Lepidoptera:
Pterophoridae). Pan-Pacific Entomol. 52(3): 251-255.
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Matthews et al.: Pterophoridae of Florida
KIMBALL, C. P. 1965. The Lepidoptera of Florida an annotated checklist. Division of
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LANGE, W. H. 1939. Early stages of California plume moths (Lepidoptera:
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MATTHEWS, D. L. 1989. The plume moths of Florida (Lepidoptera: Pterophoridae)
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Florida Entomologist 73(4)
NATIVE, INTRODUCED, AND STRUCTURE-INFESTING
TERMITES OF THE TURKS AND CAICOS ISLANDS, B.W.I.
(ISOPTERA: KALOTERMITIDAE,
RHINOTERMITIDAE, TERMITIDAE)
RUDOLF H. SCHEFFRAHN AND NAN-YAO SU
Ft. Lauderdale Research and Education Center
University of Florida Institute of Food and
Agricultural Sciences
3205 College Ave., Ft. Lauderdale, FL 33314
BRADFORD DIEHL
Walkin Diversified Services, Ltd.
Box 330, Providenciales Island
Turks and Caicos Islands, B.W.I.
ABSTRACT
A survey of termites (Isoptera) from Providenciales and Grand Turk, Turks and
Caicos Islands, British West Indies, yielded twelve species from three families. The
Kalotermitidae included Incisitermes snyderi (Light), Incisitermes sp. nr. bequaerti
(Snyder), Incisitermes sp. nr. miller (Emerson), Neotermes sp. nr. mona (Banks), N.
castaneus (Burmeister), N. jouteli (Banks), Procryptotermes corniceps (Snyder), and
the introduced Cryptotermes brevis (Walker). Rhinotermitidae collected were
Heterotermes tenuis (Hagen) and the introduced Coptotermes havilandi Holmgren, and
the Termitidae included Nasutitermes nigriceps (Haldeman) and N. costalis
(Holmgren).
RESUME
Una encuesta de termes(Isoptera) en las islas Providenciales y Gran Turco, Indias
Occidentales BritAnicas, revel6 doce species en tres families. Las Kalotermitidae in-
cluy6 Incisitermes snyderi (Light), Incisitermes sp. nr. bequaerti (Snyder), Incistermes
sp. nr. miller (Emerson), Neotermes sp. nr. mona (Banks), N. castaneus (Burmeister),
N. jouteli (Banks), Procryptotermes corniceps (Snyder), y el terme introducido, Cryp-
totermes brevis (Walker). Las Rhinotermitidae colectadas incluyeron Heterotermes
tennis (Hagen) y el terme introducido, coptotermes havilandi (Holmgren). Las Ter-
mitidae incluyeron Nasutitermes nigriceps (Haldeman) y N. costalis (Holmgren).
The first taxonomic treatise on termites of the West Indies was compiled by Banks
(1919) who listed 22 species, 9 of which he described, from museum specimens of collec-
tions made in the Bahamas, Jamaica, Trinidad, and the Greater and Lesser Antilles
(see Araujo's 1977 synonymies for Bank's 1919 species). Adamson (1937, 1938, 1940,
1946) reviewed by his collections of termites from Trinidad, with its preponderance of
Neotropical mainland species described earlier by Emerson (1925) from Guyana
(synonymies in Araujo 1977). Adamson (1938, 1940, 1948) also reported on limited collec-
tions by himself and others from the Lesser Antilles and Tobago on which were found
more depauperate "island" faunas. Ramos (1946) collected only four kalotermitid species
(identified by Emerson) from his extensive insect survey of Puerto Rico's Mona Island.
Snyder's (1956) list and key to the termites of the West Indies and Bermuda is the
most recent compilation for this region. His list is based on museum specimens and
December, 1990
622
Scheffrahn et al.: Termites of the Turks and Caicos Islands 623
records from his descriptions of 12 Antillean species and from descriptions by his con-
temporary authors including N. Banks, 14 spp.; A. E. Emerson, 15 spp.; and S. F.
Light, one sp. Of the 62 named termite species in Snyder (1956), 20 are found only on
Trinidad and the South American mainland. The remaining 42 species occur on one or
more of the other islands of the West Indies or on broader mainland distributions.
No deliberate field survey has been published for termites on any island of the West
Indies. Preliminary collections of termites from the Turks and Caicos Islands (B. Diehl,
unpubl.) indicated that a thorough survey of termites from two of the archipelago's most
surface-accessible islands was justified. The termite fauna of the Turks and Caicos
Islands was previously unknown. This paper reports the results of our survey.
MATERIALS AND METHODS
The Turks and Caicos Islands, British West Indies, consist of an archipelago of eight
major islands and many small uninhabited cays situated in the Atlantic Ocean between
21-22 N latitude and 71-73 W longitude. Termites were collected on several occasions
since 1985 and intensively 5-8 February 1990, on the two most developed and populated
islands of the group, Providenciales and Grand Turk. Providenciales (98 km2) is located
on the western side of the archipelago and Grand Turk (18 km2) is the easternmost
island and most isolated of the group. Both islands are semi-arid all year. Native low-
lying shrubs, vines, and cacti cover much of the rocky limestone soil.
Termites were collected from both islands in all discernible native and introduced
plant communities, from fenceposts and other incidental lumber, and from wood in
structures. The intent was to sample the entire diversity of native and man-made ter-
mite habitats present which we accessed by foot, bicycle, and off-road vehicle. Termites
were removed from wood and/or soil and immersed in 85% ethanol. Identifications were
based on original descriptions (listed in Araujo 1977), Banks' (1919) and Snyder's (1956)
partial keys, and comparison with specimens in the first author's collection.
RESULTS AND DISCUSSION
Collections from the two islands totaled twelve species of termites from seven genera
and three families (Table 1). Our survey yielded 53 and 15 termite samples from Pro-
videnciales and Grand Turk, respectively. All samples contained at least the soldier and
pseudergate/worker castes. At least one sample of each of the kalotermitid species
contained primary reproductive and/or alates.
Kalotermitidae
Colonies of all but one species, Paraneotermes simplicicornis (Banks), in this primi-
tive family are confined to the wood member initially infested by founding reproduc-
tives. As wood is their sole nest and foraging substrate, kalotermitid colonies lend
themselves to natural (e.g. floating logs) and human-aided (e.g. imported lumber, furni-
ture) introduction. An extreme example is the cosmopolitan species, Cryptotermes bre-
vis, found only in structural lumber (Bacchus 1987). Eight species, two-thirds of the
species known from Providenciales and Grand Turk (Table 1), are kalotermitids even
though this family represents less than one fourth of the species described from the
New World (Araujo 1977).
The largest of the three Incisitermes spp. in this survey is Incisitermes snyderi and,
although not taken from a building on these islands yet, is likely to appear in structures
as noted in Florida (Scheffrahn et al. 1988). Two other Incisitermes, one very near in
appearance to I. bequaerti and the other to the smaller I. miller, were collected in
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