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TRANSACTIONS
OF THE ©
KENTUCKY
ACADEMY OF
SCIENCE
Volume 58
Number 1
March 1997
_ Official Publication of the Academy
The Kentucky Academy of Science
Founded 8 May 1914
Governinc Boarp For 1996
ExecutivE COMMITTEE
President: William S. Bryant, Department of Biology, Thomas More College, Crestview Hills, KY 41017
President Elect: Marcus T. McEllistrem, Department. of Physics, University of Kentucky, Lexington, KY
40506-0055
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Past President: Robert O. Creek, Department of Biological Sciences, Eastern Kentucky University, Rich-
mond, KY 40475
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42301
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tucky University, Highland Heights, KY 41099; (606) 572-6390
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Kentucky University, Highland Heights, KY 41099
MEMBERS, GOVERNING BOARD
Robert J. Barney 1999 J. G. Rodriguez 1998
Patricia K. Doolin 1996 AAAS/NAAS Representative
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Gerald L. DeMoss 1997
Vincent DiNoto 1996
Chair, KJAS
COMMITTEE ON PUBLICATIONS
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John P. Harley, Department of Biological Sciences, Eastern Kentucky University,
Richmond, KY 40475
Robert O. Creek, Department of Biological Sciences, Eastern Kentucky University,
Richmond, KY 40475
Larry P. Elliott, Department of Biology, Western Kentucky University, Bowling Green,
KY 42101
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TRANSACTIONS of the
KENTUCKY
Trans. Ky. Acad. Sci. 58(1):1-3. 1997.
ACADEMY of SCIENCE
March 1997
Volume 58
Number 1
Relationships Among Habitat, Cover, and Eastern Cottontails
(Sylvilagus floridanus) in Kentucky
William M. Giuliano! and Charles L. Elliott
Department of Biological Sciences, Eastern Kentucky University, Richmond, KY 40475
and
Jeffery D. Sole
Kentucky Department of Fish and Wildlife Resources, Frankfort, KY 40601
ABSTRACT
We examined relationships among habitat, cover, and eastern cottontails (Sylvilagus floridanus) on five
wildlife management areas in Kentucky in 1989. The pellet plot technique was used to determine habitat-
use patterns and preferences, and a vegetation profile board was used to determine foliage cover. Old fields
and fencerows contained the greatest foliage cover and were preferred by cottontails year-round. Forested
areas and agricultural fields were not high-use areas and were typically avoided.
INTRODUCTION
The eastern cottontail (Sylvilagus floridan-
us), the most widely distributed species of cot-
tontail, is one of the most sought after game
animals in America (Chapman and Feldhamer
1982). In Kentucky, the cottontail provides
recreation and sport for thousands of hunters
each year. However, its numbers have been
declining in the state over the last several
years (Sole 1989). Changes in land use may be
facilitating this decline (Bruna 1952).
Intensive agricultural practices (cc over-
grazed pastures, large monocultures) typically
result in loss of valuable rabbit habitat such as
fencerows and other shrubby, so-called waste
areas (Bruna 1952). Without areas such as
these, food and cover for rabbits may be lim-
' Current address: Department of Biological and Envi-
ronmental Sciences, California University of Pennsylvania,
California, PA 15419.
iting their numbers (Barbour and Davis 1974).
Succession of fields to forests may also reduce
quality and quantity of rabbit habitat. Our ob-
jective was to examine relationships among
habitat, cover, and eastern cottontails to pro-
vide information that may be key to conserv-
ing cottontails in regions such as Kentucky
with declining populations of these animals.
METHODS
We conducted our study on five wildlife
management areas (WMAs) operated by the
Kentucky Department of Fish and Wildlife
Resources: Central Kentucky WMA (Madison
County), Clay WMA (Nicholas County), Yel-
lowbanks WMA (Breckinridge County), Tay-
lorsville Lake WMA (Spencer County), and
Lloyd WMA (Grant County). We classified ar-
eas on each WMA into one of four habitat
es: (1) fencerows: shrub- and tree-domi-
nated (> 25% cover) strips of vegetation rang-
ing from 1 m to 30 m wide; (2) forests: areas
2 Transactions of the Kentucky Academy of Science 58(1)
Table 1. Foliage cover and habitat use by eastern cottontails on five wildlife management areas in Kentucky, 1989.
Foliage cover (%)"
Cottontail use (%)
Mar May Jul Sep Nov
42A 4A 43A 50A 42A
538A 44A SDA 49A 56A
5B 2B 2B 1B 2B
OB OB OB OB OB
Habitat type <lm <2 m Jan
Fencerow 88A> 76A 46A
Old field TTA 52B 49A
Agricultural field = — 4B
Forest 32B 30C 1B
‘Foliage cover determined August 1989.
' Means in a column followed by the same letter are similar (P > 0.05).
of closed hardwood-dominated forest, with
tree cover >25%; (3) old fields: habitat rang-
ing from pastures of tall grass to older succes-
sional stages containing abundant woody
shrubs and small trees, with tree cover =25%;
and (4) agricultural fields: primarily corn
fields, with some winter wheat planted in fall
and winter.
Two areas of each habitat type per WMA
were randomly selected for study. Habitat use
by cottontails was determined using the pellet
plot technique (Pietz and Tester 1983). We es-
tablished and marked with wooden stakes 25
plots, each 1 m?. Plots were 10 m apart along
randomly located transects running through
forest, old field, and agricultural field sample
areas. In fencerows, transects were located
along the midline (total = 50 plots/habitat type/
WMA). We counted and removed fecal pellets
from plots every 4 to 8 weeks between 20 Nov
1988 and 20 Dec 1989. We tested the null hy-
potheses that cottontail use did not differ
among habitat types, months, and WMAs with
a factorial ANOVA followed by a Fisher's LSD
test (SYSTAT 1992). We examined habitat pref-
erences using chi-square tests (Neu, Byers, and
Peek 1974; SYSTAT 1992) by testing the null
hypothesis that cottontail use of habitats was
random. All four habitat types were sampled
with the same intensity (ie., 50 plots, each 1
m?/WMA). If use was random, each habitat
type should have received 25% of the total use.
We considered cottontail preference of a hab-
itat type if use was greater than expected based
on a random habitat-use pattern, and avoidance
of a habitat type if use was less than expected
based on a random habitat use pattern.
A vegetation profile board was used to de-
termine horizontal foliage cover in each hab-
itat type in August 1989 (Nudds 1977). We
estimated the percent of the profile board ob-
structed by vegetation when viewed from a
distance of 10 m, every 25 m along each 250
m transect in each habitat type. Foliage cover
was compared among habitat types and
WMaAs using a factorial ANOVA followed by
Fisher's LSD test (SYSTAT 1992).
RESULTS
Cottontail use (Table 1) differed among
habitat types (P<0.05) but not among months
or WMAs (P>0.05). Habitat preferences did
not differ by month (P>0.05). Fencerow and
old-field habitats were preferred (P<0.05),
and forests and agricultural fields were avoid-
ed (P<0.05).
Foliage cover (Table 1) differed by habitat
type (P<0.01) but not WMA (P>0.05). Fo-
liage cover estimates of agricultural fields were
not used because of high variability in cover
due to differences in crop rotations and prob-
lems associated with frequent disturbance
(i.e., harvest and planting activities).
DISCUSSION
The cover that habitats provide is the most
influential feature determining use of an area
by rabbits (Anderson and Pelton 1976; Swihart
and Yahner 1982). Our findings agree with
those of Barbour and Davis (1974) and suggest
that fencerow and old-field habitats consistently
received the most use and were preferred be-
cause they had greater foliage cover than forest
habitats, at least in summer. The added cover
in old fields is important for escape and nesting
(Bruna 1952). The additional cover provided by
fencerows may have been important as a source
of protection, escape routes, and travel corri-
dors (Edwards et al. 1981).
In fall and winter, old fields usually lose
much of their value to cottontails, both as a
source of food and cover, due to desiccation
of herbaceous vegetation and loss of foliage
from woody vegetation. Because of this, rab-
Habitat, Cover, and Cottontails—Giuliano and Elliott
bits typically move to forested areas and other
areas with abundant woody vegetation that
provides adequate cover (Barbour and Davis
1974; Lewis 1972). During our study, howev-
er, cottontails continued to prefer old fields
and to avoid forested areas even in fall and
winter. We believe this may have been due to
an adequate amount of cover being provided
by old fields, even in fall and winter. In addi-
tion, the winter of 1988-1989 was mild, with
below average snowfall (USDC-NOAA 1988-
1989). Therefore, rabbits may not have need-
ed to move to areas containing more woody
vegetation in search of food and cover (Han-
sen, Bailey, and Siglin 1969).
We suggest that limited use and apparent
avoidance of agricultural fields was probably
due to the continuous disturbance associated
with agricultural practices (Friley 1955). Ed-
wards et al. (1981) reported similar patterns
of use and avoidance of agricultural fields by
cottontails in Illinois.
We have shown that fencerows and old
fields are preferred, year-round cottontail hab-
itats; the creation and maintenance of these
areas should be a high management objective.
Forests and agricultural fields are not high-use
areas but may be seasonally important as
sources of food and cover. We suggest that
management activities include (1) maintaining
and planting of shrubs and trees along fences
and field edges to enhance fencerow habitat,
(2) disking, mowing, and burning of fields to
prevent succession to forest, and (3) coordi-
nating agricultural practices to minimize dis-
turbance of fields and provide additional
sources of food.
ACKNOWLEDGMENTS
This study was funded by Pittman-Robert-
son Federal Aid in Wildlife Restoration mon-
ies in cooperation with the Kentucky Depart-
ment of Fish and Wildlife Resources, P-R Pro-
ject W-45-21, and the Wildlife Program, De-
partment of Biological Sciences, Eastern
Kentucky University.
ioe)
LITERATURE CITED
Anderson, B.F., and M.R. Pelton. 1976. Movements,
home range, and cover use: factors affecting the sus-
ceptibility of cottontails to hunting. Proc. Southeast. As-
soc. Game and Fish Commissioners 30:525-535.
Barbour, R.W., and W.H. Davis. 1974. Mammals of Ken-
tucky. Univ. Press of Kentucky, Lexington.
Bruna, J.F. 1952. Kentucky rabbit investigations. Ken-
tucky Fish and Wildlife Dep., Fed. Aid. Proj. 26-R.,
Frankfort.
Chapman, J.A., and G.A. Feldhamer. 1982. Wild mam-
mals of North America. Johns Hopkins Univ. Press, Bal-
timore, MD.
Edwards, W.R., S.P. Havera, R.F. Labisky, J.A. Ellis, and
R.E. Warner. 1981. The abundance of cottontails in
relation to agricultural land use in Illinois (U.S.A.)
1956-1978, with comments on mechanism of regula-
tion. Proc. World Lagomorph Conf. 1:761-789.
Friley, C.E. 1955. A study of cottontail habitat prefer-
ences on a southern Michigan farming area. Fed. Aid
Proj. W-48-R.
Hansen, J.C., J.A. Bailey, and R.J. Siglin. 1969. Activity
and use of habitat by radio-tagged cottontails during
winter. Trans. Ill. Acad. Sci. 62:294—302.
Lewis, H.S. 1972. Cottontail rabbit home range, habitat
use, and mortality in southern Wisconsin. M.S. thesis,
Univ. Wisconsin, Madison.
Neu, C.W., C.R. Byers, and J.M. Peek. 1974. A technique
for analysis of utilization-availability data. J. Wildlife
Managem. 38:541—545.
Nudds, T.D. 1977. Quantifying the vegetative structure
of wildlife cover. Wildlife Soc. Bull. 5:113-117.
Pietz, P.J., and J.R. Tester. 1983. Habitat selection by
snowshoe hares in north central Minnesota. J. Wildlife
Managem. 47:686-696.
Sole, J.D. 1989. Quail and rabbit roadside survey. P-R
Proj. W-45-21. Ky. Dep. Fish and Wildlife Resources,
Frankfort.
Swihart, R.K., and R.H. Yahner. 1982. Habitat features
influencing use of farmstead shelterbelts by the eastern
cottontail (Sylvilagus floridanus). Am. Mid]. Naturalist
107:411-414.
SYSTAT. 1992. SYSTAT for Windows: statistics, Version
5 ed. SYSTAT, Inc. Evanston, IL.
[USDC-NOAA] U.S. Department of Commerce-National
Oceanic and Atmospheric Administration. 1988-1989.
Climatological data: Kentucky. Vol. 83, No. 13; Vol. 84,
Nos. 1-3. ISSN 0145-0433. National Environmental
Satellite, Data and Information Service, National Cli-
matic Data Center, Asheville, NC.
Trans. Ky. Acad. Sci. 58(1):4-8. 1997.
Separation of Spawning Habitat in the Sympatric Snubnose Darters
Etheostoma flavum and E. simoterum (Teleostei, Percidae)
Jean C. Porterfield
Fish Division, Museum of Zoology, Ann Arbor, MI 48109!
ABSTRACT
Individuals of two snubnose darter species, Etheostoma flavum and E. simoterum, were observed in Whites
Creek, Tennessee, in April 1995 and April 1996. Both species were present and spawning in 1995, but in
1996 the stream had experienced increased siltation and algal growth, and no E. flavum individuals were
seen. Spawning behaviors of both species were similar to those known for other snubnose darters: E. flavum
and E. simoterum attached their eggs to rock surfaces, and the male followed the female as she searched
for appropriate substrate. Behavioral observations in 1995 suggested that these two species utilized different
sites for spawning, and that interspecific aggression was not the mechanism by which habitat isolation was
obtained. Since most closely related darters have allopatric distributions, studying instances of sympatry such
as this may contribute to understanding of distribution patterns of darters as a group.
INTRODUCTION
Theoretical works on species concepts and
speciation processes traditionally cite repro-
ductive behavior as a significant isolating
mechanism between sympatric species (Mayr
1963; Paterson 1993). Choice of spawning site
is one aspect of reproductive behavior that has
been proposed as an isolating mechanism in
some fishes (lamprologine cichlids, Ribbink et
al. 1983; Lake Baikal sculpins, Smith and Todd
1984), and it may be important in some spe-
cies of darters. Darters are one of the most
speciose groups of freshwater fishes in North
America, and it is common to find more than
10 species in a single stream. Although most
closely related darters are allopatric, some do
share overlapping distributions. Several stud-
ies have demonstrated the significance of mi-
crohabitat segregation in competition among
sympatric darters, although reproductive be-
haviors were not specifically addressed
(Greenburg 1988; Kessler and Thorp 1993).
The snubnose darters, members of the sub-
genus Nanostoma (sensu Page 1981, but see
Bailey and Etnier 1988) in the genus Etheos-
toma, are comprised of 23 described species,
most of them restricted to the southeastern
United States (Page and Burr 1991) and most
distributed allopatrically. Etheostoma simoter-
um, the snubnose darter, has a wide range
throughout the Tennessee and Cumberland
' Present address: Illinois Natural History Survey, 607
East Peabody Drive, Champaign, IL 61820.
river systems where it is often sympatric with
two other species of snubnose darters: Eth-
eostoma duryi, the black darter, and E. fla-
vum, the saffron darter. Etheostoma flavum is
sympatric with E. simoterum throughout
much of its range in the lower Cumberland
and Duck river systems and in some tributar-
ies of the lower Tennessee River (Etnier and
Stamnes 1993). Spawning behavior has been
described from aquarium observations for E.
simoterum (Page and Mayden 1981) and from
both aquarium and field observations for E.
flacum (Keevin, Page, and Johnston 1989);
both species are classified as egg attachers
(Page 1985). Descriptions of both species can
be found in Etnier and Starnes (1993). Here
I describe the spawning behavior of E. simo-
terum under natural conditions, present the
results of observations on spawning habitat
separation between E. simoterum and E. fla-
vum in Whites Creek, Tennessee, and discuss
the implications for reproductive isolation be-
tween the two species.
MATERIALS AND METHODS
I observed spawning E. simoterum and E.
flavum while snorkeling in Whites Creek at
Hwy 41A bridge, Davidson County, Tennes-
see, on 8, 9, and 16 April 1995, and on 11
April 1996. Observations were made in all
available habitat types including silted bedrock
pools, gravel/cobble runs and riffles, and al-
gae-covered boulders (rip rap) piled at stream
edges; depths ranged from 30 cm to 1 m. In
1995, behaviors of both species were filmed
Spawning Habitat in Snubnose Darters—Porterfield 5
on 9 and 16 April using a Sony CCD-FX310
8-mm video camera in a Sony MPK-TRA2 un-
derwater housing. Observations were record-
ed at the spawning sites and later when I
viewed the video tapes. I observed E. simo-
terum at the bridge for a total of 4 hours on
2 separate days, during which time four
spawning pairs were followed; additionally, 2
hours of observations including one spawning
pair were made over the 2 days in a bedrock
pool at a site about 300 m upstream. Three
hours of observations on six individuals of E.
flavum were made on the same 2 days at the
bridge. I observed the darters from late morn-
ing to mid-afternoon each day. Water temper-
ature during the observations remained
around 21° C.
On 11 April 1996 fish were observed in the
late morning and early afternoon. The water
temperature was 11° C. No darters were
filmed, but observations on species present,
behaviors of these species, and habitat condi-
tion were noted.
RESULTS
Although courtship behavior prior to pair-
formation was not observed in either E. sim-
oterum or E. flavum, the spawning behavior
after pair-formation was observed to be iden-
tical in both species. The male closely fol-
lowed the female as she swam over the sub-
strate searching for egg deposition sites. He
stayed above and a little behind her with me-
dian fins semi-erect; in some cases he left to
chase an approaching male. When the female
found a potential site for egg-deposition, she
visually examined the site, as her body formed
a 30-45 degree angle with the substrate sur-
face. She then made one to four sharp jabs at
the spot with her snout as the male moved to
position himself above her. The head jabs ob-
served were similar to those described for E.
davisoni by Bart (1992) and for E. zonale by
Walters (1994) and were a consistent indicator
of oviposition, with 100% of all spawns in both
species preceded by at least one head jab. Im-
mediately following the head jabs, the female
dragged her body over the chosen site as the
male hovered closely above her. When the
female’s genital papilla reached the site, she
pressed her papilla tightly against the rock and
oviposited, while the male arched his body
against her, pressed his caudal peduncle to-
wards the rock, and vibrated, releasing sperm.
This spawning behavior is identical not only
between E. simoterum and E. flavum but also
among the other species in the subgenus Na-
nostoma that I have observed spawning in the
field: E. barrenense, E. duryi, and E. rafines-
uei.
In 1995, E. simoterum was the most com-
mon fish near the Hwy 41A bridge and the
most common darter at the riffle/pool habitat
upstream of the bridge. Many individuals were
seen in the large cobble raceway near the cen-
ter of the stream; although individuals of both
sexes were observed here, there were more
females, and no E. simoterum were observed
forming pairs or spawning in this area. Fewer
E. simoterum were present among the large
algae-covered boulders piled near the bridge
on the north side of the stream; however,
there was a greater proportion of males seen
here, and the only spawning pairs of E. sim-
oterum at the bridge site were located in these
piles. I observed pairs attaching eggs to the
boulders, and spawning positions ranged from
slightly angled near the tops of boulders, to
near vertical positions on the sides of boul-
ders, to upside-down under an overhanging
boulder. No other darter species were ob-
served in this large-boulder area, although
some E. caeruleum were seen at its periphery.
In 1995, E. simoterum was common at the
upstream site near the sides of the stream in
areas of algae-covered flat bedrock with some
cracks and shelves. There were many individ-
uals in the area, but only one spawning pair
was observed. The pair was observed on a
bedrock shelf spawning in an algae-covered
crevice in a horizontal position. At both sites,
individuals spawned on surfaces covered with
algae; out of the total of nine spawns observed,
eight occurred on large algae-covered boul-
ders or stretches of bedrock in a flowing pool
environment at the side of the stream. The
other spawn took place on a piece of cobble
also covered with algae but located closer to
the raceway in faster water.
The six individuals of E. flavum observed in
1995 were found only in the cobble raceway
above the Hwy 41A bridge. This raceway av-
eraged 60 cm deep; the substrate consisted of
gravel and cobble. Two spawning pairs were
observed, and two other individuals were
seen. In all four spawning bouts the females
6 Transactions of the Kentucky Academy of Science 58(1)
chose to attach eggs to pieces of clean rock in
the fastest current of the raceway. Three
spawns took place on pieces of cobble, while
the fourth took place on a side of a rock ledge.
Spawning positions were most often at about
a 45 degree angle to the substrate surface as
females oviposited on the sides of the rocks.
Etheostoma simoterum and E. caeruleum were
the two most common fishes in the raceway,
but I did not observe individuals of either spe-
cies spawning there.
There were no interspecific interactions be-
tween spawning pairs of E. simoterum and E.
flavum individuals in Whites Creek, but this
observation is affected by the demography in
the stream; E. flavum was not present in the
boulders where E. simoterum spawned. How-
ever, spawning E. flavum pairs often encoun-
tered E. simoterum individuals. Encounters
are defined as fishes coming within one body
length of each other. Of 11 of the interspecific
encounters documented for the E. flavum
pairs, two were with E. simoterum males. In
one instance the E. flavum individuals ignored
an E. simoterum male who swam between
them; in the other instance the E. flavum male
swam ahead of the female to approach an E.
simoterum male who seemed to be feeding in
their path. None of the interspecific encoun-
ters between single individuals (i.e., not
spawning pairs) involved aggression. Of two
encounters between E. flavum pairs and con-
specific males, both involved aggressive inter-
actions; for E. simoterum all seven intraspe-
cific encounters involved aggression.
Observations in Apri! 1996 revealed that the
habitat in Whites Creek was different from
the habitat observed a year earlier. The clean
cobble in the faster run where E. flavum was
seen in 1995 was covered with algae, and the
whole area appeared to have experienced
more siltation. Both male and female E. sim-
oterum were common in the cobble runs, but
no E. flavum individuals were seen. Large E.
simoterum males were still common among
the algae-covered boulders at the side of the
stream, and some E. simoterum females were
present as well. No spawns were observed, but
three pairs of E. simoterum were seen: one
among the boulders and two in the cobble
area at the head of the raceway.
Fish stream associates in 1995 at the bridge
site were Campostoma anomalum, Etheosto-
ma caeruleum, E. crossopterum, E. flabellare,
E. smithi, Lepomis cyanellus, L. megalotis,
Luxilus chrysocephalus, Micropterus punctu-
latus, and Percina caprodes. Associates in 1995
at the upstream site were Campostoma an-
omalum, Etheostoma blennioides, Lepomis
megalotis, Luxilus chrysocephalus, Lythrurus
ardens, Micropterus punctulatus, and Pime-
phales notatus. Observed fish associates in
1996 at the bridge site were Campostoma an-
omalum, Catostomus commersoni, Etheostoma
caeruleum, E. flabellare, Lepomis cyanellus, L.
megalotis, Luxilus chrysocephalus, and Mi-
cropterus punctulatus.
DISCUSSION
The spawning behaviors of E. flavum and
E. simoterum observed in Whites Creek are
nearly identical to previously published de-
scriptions for these two species (Keevin, Page,
and Johnston 1989; Page and Mayden 1981).
I found the angle of the spawning position for
E. flavum to be around 45 degrees; Keevin,
Page, and Johnston (1989) reported a near
vertical spawning position for aquarium-held
fish. These differences may reflect the cobble
habitat in which E. flavum was found in
Whites Creek as opposed to the large flat
rocks preferred by aquarium-held Warren
Fork fish (Keevin, Page, and Johnston 1989).
Males of neither species defended stationary
territories; instead, females would roam a
large area of substrate searching for oviposi-
tion sites, and males defended the females
with which they were paired. These observa-
tions are similar to those reported in studies
of the spawning behaviors of other snubnose
darters (Carey and Burr 1989; Keevin, Page,
and Johnston 1989; Page and Mayden 1981;
Stiles 1974).
Observations of E. flavum and E. simoterum
in Whites Creek in 1995 suggest that in this
stream these two snubnose darters utilize dif-
ferent spawning sites in different areas of the
stream. Previous studies have shown that un-
der natural conditions, both species preferred
flowing-pools with a variety of substrates; in
aquaria they preferred to deposit eggs on large
stones (Keevin, Page, and Johnston 1989; Page
and Mayden 1981). However, Keevin, Page,
and Johnston (1989) added that when large
rocks are not available, E. flavum individuals
spawn on a cobble substrate. I found E. fla-
Spawning Habitat in Snubnose Darters—Porterfield i
vum to be utilizing the clean cobble in the
swifter runs, while E. simoterum spawning
pairs were found among the large, algae-cov-
ered boulders in slower current at the side of
the stream. These data support the hypothesis
that E. simoterum individuals in Whites Creek
occupy the boulder pool habitat, making it un-
available to E. flavum individuals. Etnier and
Stames (1993) stated that E. simoterum indi-
viduals in sympatry with E. flavum are more
often found in the swifter riffles. My obser-
vations in Whites Creek suggest that while
many E. simoterum were seen in the faster
raceway, this is not the area utilized for spawn-
ing. Spawning pairs of E. simoterum were
found only in the boulder pools, suggesting
that this species may use different stream hab-
itats for spawning than for other activities such
as feeding. In addition, the presence of E. sim-
oterum in the faster water is not a result of
interactions with E. flavum, as individuals of
the latter species were not seen in the pools.
Behavioral observations show that members
of the two snubnose darter species rarely in-
teract aggressively with heterospecific individ-
uals, while intraspecific interactions among
males usually involve aggression. The greater
number of interspecific encounters of E. fla-
vum pairs with E. simoterum females (nine,
versus two with E. simoterum males) reflects
the demography of the E. simoterum popula-
tion in Whites Creek; females were concen-
trated in the faster current of the raceway
where the E. flavum individuals were spawn-
ing. Similarly, E. simoterum spawning pairs
had no interspecific encounters because no E.
flavum individuals were present in the boulder
areas where they spawned. In part because of
this relationship to demography, the rarity of
direct aggression observed between species
does not reject the hypothesis that competi-
tion for spawning sites causes E. flavum to
spawn in the cobble raceway. However, the
encounter data do suggest that agonistic be-
havior is not the means by which habitat iso-
lation is maintained. Other possible mecha-
nisms for maintenance of the habitat segre-
gation observed should be considered, such as
more subtle behavioral cues or recognition of
male color patterns.
In 1995 there were low numbers of E. fla-
vum in this section of Whites Creek. This de-
mographic factor may affect and/or be affect-
ed by the patterns of habitat use seen in this
species as well as in E. simoterum. The rarity
of E. flavum in this stream increases the like-
lihood that a given observation of spawning
habitat in this species is due to chance and not
to substrate choice. However, the spawning
substrate used by this species may affect its
relative success in this stream if, for example,
there is differential success of eggs in the dif-
ferent habitats. In 1996, E. flavum individuals
were not observed in Whites Creek at Hwy
41A, and there were fewer species of fishes in
general. These observations might be ex-
plained by the difference in water tempera-
ture (21° C in 1995 versus 11° C in 1996), but
E. simoterum was observed forming pairs in
the 11° C water in 1996, so presumably the
water was not too cold for darters to be pres-
ent and active. Or the absence of E. flavum
might be explained by the increased levels of
siltation and algal growth since April 1995; the
species spawned on clean cobble in 1995, sub-
strate, which was no longer available in 1996.
The absence of E. flavum might also be a re-
sult of interactions with E. simoterum; greater
numbers of E. simoterum competing for food
and lower egg survivorship for E. flavum in
the faster water are possibilities. At this point,
hypotheses concerning which factors might be
responsible for the absence of E. flavum are
speculative.
Etheostoma simoterum is also sympatric
with E. duryi in the middle Tennessee River
system; these two species may also exhibit
spawning habitat segregation when sympatric
(pers. obs.). Snorkeling observations in April
1995 at Gray Branch (Cypress Creek system,
Alabama) revealed that E. duryi attached eggs
to clean cobble and bedrock at the head of an
island, while most E. simoterum pairs were
spawning on cobble in the edge of a run on
one side of the island. These observations are
consistent with those reported by Etnier and
Starnes (1993). Etheostoma flavum and E.
duryi are occasionally sympatric, particularly
in the upper Duck River system as discussed
by Etnier and Bailey (1989), but I have not
made snorkeling observations at any of these
localities.
In most cases, the species of darters found
in a given stream are not closely related (i.e.,
in the same subgenus) and do not exhibit sim-
ilar reproductive habits such as choice of mi-
8 Transactions of the Kentucky Academy of Science 58(1)
crohabitat or spawning substrate. The result-
ing pattern is that the darter species in most
streams coexist without interspecific competi-
tion for spawning sites. Field data on E. sim-
oterum and E. flavum in Whites Creek based
on limited observations made during one
spawning season suggest that there is some ev-
idence for competition for spawning sites be-
tween closely related sympatric darters. If cor-
roborated with experimental data, these re-
sults could be significant to studies of the re-
productive ecology and distributional history
of snubnose and other darter species. For ex-
ample, if competition is found to be a signifi-
cant factor in the existence and/or persistence
of darter species in a given area today, then its
role in the historical distribution of darters
could be explored.
ACKNOWLEDGMENTS
This research was supported by the Univer-
sity of Michigan Department of Biology, the
University of Michigan Museum of Zoology,
the University of Illinois Graduate College,
and the Illinois Natural History Survey. I
thank W.E. Browne, P.A. Ceas and A.L. Utz
for field help; A.K. Peters and G.R. Smith for
the use of video equipment; and P.A. Ceas and
L.M. Page for helpful comments on the manu-
script.
LITERATURE CITED
Bailey, R. M., and D. A. Etnier. 1988. Comments on the
subgenera of darters (Percidae) with descriptions of two
new species of Etheostoma (Ulocentra) from southeast-
ern United States. Misc. Publ. Mus. Zool. Univ. Mich.
175:1-48.
Bart, H. L., Jr. 1992. Spawning behavior of Etheostoma
davisoni Hay. Copeia 1992:537-539.
Camey, D. A., and B. M. Burr. 1989. Life histories of
the bandfin darter, Etheostoma zonistium, and the fi-
rebelly darter, Etheostoma pyrrhogaster, in western
Kentucky. Ill. Nat. Hist. Surv. Biol. Notes 134:1-16.
Etnier, D. A., and R. M. Bailey. 1989. Etheostoma (UI-
ocentra) flavum, a new darter from the Tennessee and
Cumberland river drainages. Occas. Pap. Mus. Zool.
Univ. Mich. 717:1-24.
Etnier, D. A., and W. C. Starnes. 1993. The fishes of
Tennessee. University of Tennessee Press, Knoxville.
Greenberg, L. A. 1988. Interactive segregation between
the stream fishes Etheostoma simoterum and E. rufili-
neatum. Oikos 51:193—202.
Keevin, T. M., L. M. Page, and C. E. Johnston. 1989.
The spawning behavior of the saffron darter (Etheos-
toma flavum). Trans. Ky. Acad. Sci. 50:55-58.
Kessler, R. K., and J. H. Thorp. 1993. Microhabitat seg-
regation of the threatened spotted darter (Etheostoma
maculatum) and closely related orangefin darter (E. bel-
lum). Canad. J. Fish. Aquatic Sci. 50:1084-1091.
Mayr, E. 1963. Animal species and evolution. Harvard
University Press, Cambridge.
Page, L. M. 1981. The genera and subgenera of darters
(Percidae, Etheostomatini). Occas. Pap. Mus. Nat. Hist.
Univ. Kans. 90:1-69.
Page, L. M. 1985. Evolution of reproductive behaviors in
percid fishes. Bull. Ill. Nat. Hist. Surv. 33:275-295.
Page, L. M., and B. M. Burr. 1991. A field guide to fresh-
water fishes of North America north of Mexico. Hough-
ton Mifflin, Boston.
Page, L. M., and R. L. Mayden. 1981. The life history of
the Tennessee snubnose darter, Etheostoma simoterum,
in Brush Creek, Tennessee. Ill. Nat. Hist. Surv. Biol.
Notes 117:1-11.
Paterson, H. E. H. 1993. Evolution and the recognition
concept of species. The Johns Hopkins University
Press, Baltimore.
Ribbink, A. J., B. A. Marsh, A. C. Marsh, A. C. Ribbink,
and B. J. Sharp. 1983. A preliminary survey of the cich-
lid fishes of rocky habitats in Lake Malawi. South Afr.
J. Zool. 18:149-310.
Smith, G. R., and T. N. Todd. 1984. Evolution of species
flocks of fishes in north temperate lakes. Pages 45-69
in A.A. Echelle and I. Kornfield (eds). Evolution of fish
species flocks. University of Maine at Orono Press,
Orono.
Stiles, R. A. 1974. The reproductive behavior of the
Green and Barren River Ulocentra (Osteichthyes: Per-
cidae: Etheostoma). Assoc. Southeast. Biol. Bull. 21:86-
87.
Walters, J. P. 1994. Spawning behavior of Etheostoma
zonale (Pisces: Percidae). Copeia 1994:818-821.
Trans. Ky. Acad. Sci. 58(1):9-19. 1997.
Scientists of Kentucky
Common Names of Vascular Plants Reported by C.S. Rafinesque in an
1819 Descriptive Outline of Four Vegetation Regions of Kentucky
Ronald L. Stuckey
Herbarium in the Museum of Biological Diversity, Department of Plant Biology, College of Biological Sciences,
The Ohio State University, 1315 Kinnear Road, Columbus, Ohio 43212
and
James S. Pringle
Royal Botanical Gardens, Box 399, Hamilton, Ontario, Canada L8N 3H8
In the flora of North America, knowledge is
scarce on the sources and origins of the com-
mon names applied to vascular plants. In bo-
tanical literature, however, a large number of
studies have been published with the purpose
of providing common names of plants for ei-
ther general or specific regions of the United
States. McAtee (1913-1933), in a series of five
papers published over a 20-year period, listed
410 common names with the geographic areas
in which they were used, and 60 sources of
information. His list, consisting mostly of
plants of aquatic and marsh habitats, is of
names learned during field work for the Bio-
logical Survey in the United States Depart-
ment of Agriculture. McAtee’s list, containing
an index to the plant names in each of his five
papers, provides an excellent working base for
obtaining local common names from literature
sources published during the first third of the
20th century or earlier.
Constantine Samuel Rafinesque (1783-
1840) (Figure 1), the controversial, eccentric,
sometimes considered erratic European bota-
nist who traveled in eastern North America
during the first third of the 19th century, pub-
lished articles and books totaling over 900 ti-
tles in botany. Many of these publications
were on taxonomic botany, including cata-
logues of floras (Rafinesque 1836, 1836-1838,
1840), geographical floras (Rafinesque 1817b,
1817c, 1819a, 1824), medical floras (Rafin-
esque 1828, 1830), and studies of genera (Raf-
inesque 1811, 1820). In his Medical Flora, Raf-
inesque (1828, 1830) provided an English,
French, and German name, one or more of-
ficinal names (i.e., names used in pharmacy),
and one to several “vulgar names” for each
species, the last being additional names in En-
glish used by members of the public but not
preferred for scholarly communication. Oth-
erwise, in these well-known publications, he
did not provide common names for the plants
listed or described, many of which were des-
ignated by him as new taxa. In a few of his
papers among this vast output of botanical in-
formation, he reported the common names of
vascular plants.
Rafinesque traveled and explored for plants
in Kentucky during the years 1818 to 1826 and
was Professor of Botany and Natural History
at Transylvania University in Lexington from
1819 to 1825. During this period he wrote a
short article (Rafinesque 1819c; Figure 2) con-
sisting of a descriptive outline of the vascular-
plant vegetation in four major regions of the
state. This paper was published in 1819 in the
second number of the first volume of The
Western Review and Miscellaneous Magazine,
edited by William Gibbs Hunt (Rafinesque
1819c). In addition to providing general re-
marks, Rafinesque characterized each of the
four botanical regions as to geographical lo-
cation, general topography, kinds of bedrock,
and unique features of the vegetation. He also
provided a short list of vascular plants that he
considered somewhat specific or “peculiar” to
each region. Along with each binomial scien-
tific plant name, he added the common name,
or in his terminology, the “vulgar” name, not-
ing, however, that “the vulgar names of the
plants .. . cannot claim to be generally under-
stood even in [Kentucky], many being merely
10 Transactions of the Kentucky Academy of Science 58(1)
| |
4M
~~
Figure 1. Portrait of C.S. Rafinesque by American artist William Birch (1755-1834). After a long succession of private
owners, this enamel miniature (2 4” diameter) was purchased by Transylvania University, Lexington, Kentucky, in 1938.
Photo courtesy of Transylvania University Special Collections.
local or personal. The botanical names are
alone to be relied on.”
Two years earlier, in his review of the first
edition of Amos Eaton’s Manual of Botany for
the Northern States . . . [1817], Rafinesque
(1817a) supported the idea of providing the
vulgar names in works on local taxonomic bot-
any, and he noted that their inclusion in Ea-
ton’s Manual was “valuable.” He commented
that most of them had been taken by Eaton
from other cited published sources, rather
than being “taken directly from the vulgar,”
and wrote further that “vulgar names are at all
times a valuable appendage to classical syn-
onymy, and indispensable in local botanical
writings.”
Rafinesque probably prepared the article on
vegetation of Kentucky for a popular audi-
ence, which would explain his use of the com-
mon names of plants, but he reminded his
readers that the botanical names were the in-
tended authentic ones to be used for scientific
accuracy. Botanists always have and, it is ex-
pected, always will recognize that the botani-
cal names should be used in authoritative flo-
ristic and vegetational studies. Botanists, how-
ever, must also communicate with and make
their studies useful to those who work in other
fields of study but who are not familiar with
Latin binomials. Among these fields are geol-
ogy, zoology, wildlife and fisheries studies, nat-
ural resources management, endangered spe-
cies preservation, conservation organizations,
environmental law, ecological consultation,
and other groups with environmental con-
cems.
The use of common names of plants is of
more importance now than it was 25 or 50
Rafinesque Common Names—Stuckey and Pringle 11
MISCELLANY, =;
—=+
BOTANY OF KENTUCKY.
‘On its principal features, by C. S. Rarinesque, Professor of
Botany and Natural History in Transylvania Universi-
ty.
THE state of Kentucky being situated in the-centre of the
western country, has a flora similar to the generality of the
western states, and participating in their peculiar features, while’
if offers in itself a complete specimen of the western botany.
The peculiarities of this botany consist principally in the
total want of the maritime and mountain regions, which form
sach remarkable sections in the local floras of the Atlantic states,
and abound with plants peculiar’ to themselves. Another
striking feature in the vegetation of Kentucky and the western
states is the propensity which many plants and trees exhibit of
growing in a-social state, to the almost total exclusion of
every other. There are many plants which grow crowded
together, allover the United States; such for instance as the
grasses, ferns, the ‘Comptonia, the Studfonia, &c. but they al-
low many other plants to grow with them; while, in-the western
country, many extensive spaces of ground are covered with one
or a few crow: ‘ed species, to the exclusion of many others, which
are found in their company elsewhere. The plants which may
be quoted as a striking 3 instance of this singular fact are not few,
among which I shall select the following:
Vernonia frrealta, Tron Weed,.
Bapttisia cerulea, - Blue. Wild Indigo, .
Cacalia reniformis,. ~ Kidney Weed,
Stedeoma pulegivides, ’Penny-aoyal,
Chenopodium SU Worm ‘Weed,
Elephantofius scaber, ‘Elephant’ s Fo ts ‘
Figure 2. First page of C.S. Rafinesque’s paper on “Bot-
any of Kentucky[:] on its principal features,”
published in 1819 in volume 1(2) of The Western Review
and Miscellaneous Magazine.
which was
years ago; it will become more and more prev-
alent and necessary as time progresses. With
increasing frequency, floristic manuals, such as
that of Gleason and Cronquist (1991) and the
ongoing Flora of North America North of Mex-
ico, are providing vernacular names for all or
most of the species described therein. The use
of common names will demand more accuracy
and stability to the extent that nomenclatural
guidelines may need to be established in the
future. These guidelines may develop in a
manner similar to those parameters provided
in the International Code of Botanical Nomen-
clature. The correct interpretation of earlier
papers in which common names were used re-
quires that the authors’ applications of such
names be accurately determined. Some names
may have been applied in senses different
from more recent usage, and some may have
fallen into disuse. Efforts to standardize com-
mon nomenclature require the selection of ap-
propriate and otherwise acceptable names that
will not perpetuate or introduce confusion. At-
taining these objectives will require consider-
ation of historic and regional applications of
vernacular plant names. For these reasons we
bring to the attention of the scientific public
the common plant names in Rafinesque’s early
19th century publication on the botany of
Kentucky.
Rafinesque’s paper on the botany of Ken-
tucky is transcribed below. His lists of plant
species are annotated, as indicated in the fol-
lowing paragraphs, to permit their interpreta-
tion using current botanical and common
names. Separate indexes to botanical and com-
mon names are provided in Appendix 1.
Rafinesque—owing, he said in the errata, to
his absence from town—had no opportunity
for proofreading until after the paper had
gone to press. Consequently, the original lists
of plant names are rife with typographical er-
rors, even among the common names. Rafin-
esque (1819d) published corrections of a num-
ber of these errors in a later issue of the same
journal, on page 128. Several others, however,
escaped his notice. “Tris” for “Iris,” and “Pin
week” for “Pin weed,” for example, are obvi-
ous typographical errors.
In some cases, however, minor alterations
presumably do represent Rafinesque’s intent.
From some of his other publications, it is ev-
ident that he preferred a simplified spelling
such as “cerulea” to the original “caerulea” or
“coerulea” and “catesbei” for “catesbaei,” and
that he frequently made such arbitrary abridg-
ments. Such changes, whether required or
proscribed by present rules of nomenclature,
are not regarded as new names, and they re-
main attributed to the original authors. Be-
cause of the circumstances discussed above, it
appeared best to retain Rafinesque’s original
format in the lists, which omitted the attri-
bution of the authorship of the botanical
names. As the lists appear in the present pa-
per, the botanical names used by Rafinesque
are in italic type; the common names, in ro-
man type. (At the left margin we have num-
bered, in parentheses, each species for index-
ing purposes.) Indented and in square brack-
ets and smaller type below the names we have
added, first, the corrected orthography of the
12 Transactions of the Kentucky Academy of Science 58(1)
botanical name used by Rafinesque, in italics
if it is not the currently accepted botanical
name, with authorship; second, where appro-
priate and in roman type, the currently ac-
cepted botanical name; and third, a common
name or names in current use. When the bo-
tanical name used (or intended) by Rafinesque
remains in use for the species, this name ap-
pears in roman type, with the orthography cor-
rected if necessary and the authorship indi-
cated. Curly brackets have been used to in-
dicate the corrections made by Rafinesque
(1819d) in the errata. In Appendix 2, notes,
mostly on nomenclature, have been added for
14 taxa.
The moder common names we include
have generally been obtained from Fernald
(1950), Gleason and Cronquist (1991), and the
popular field guides by Peterson and Mc-
Kenny (1968) and Newcomb (1977). The orig-
inal Illustrated Flora by Britton and Brown
(1896-1898), which provided common names
for nearly every species covered and also re-
corded many local or otherwise relatively ob-
scure vernacular names, was also consulted.
Rafinesque’s paper (Rafinesque 1819c) on
the principal features of the botany of Ken-
tucky is as follows:
Botany of Kentucky
On its principal features, by C.S. Rafinesque,
Professor of Botany and Natural History in the
Transylvania University.
The state of Kentucky being situated in the
centre of the western country, has a flora sim-
ilar to the generality of the western states and
participating in their peculiar features, while
it offers in itself a complete specimen of the
western botany.
The peculiarities of this botany consist prin-
cipally in the total want of the maritime and
mountains regions, which form such remark-
able sections in the local floras of the Atlantic
states, and abound with plants peculiar to
themselves. Another striking feature in the
vegetation of Kentucky and the western states
is the propensity which many plants and trees
exhibit of growing in a social state, to the al-
most total exclusion of every other. There are
many plants which grow crowded together, all
over the United States; such for instance as
the grasses, ferns, the Comptonia, the Stud-
fonia {Hudsonia}, &c. but they allow many
other plants to grow with them; while, in the
western country, many extensive spaces of
ground are covered with one or a few crowded
species, to the exclusion of many others, which
are found in their company elsewhere. The
plants which may be quoted as a striking in-
stance of this singular fact are not few, among
which I shall select the following:
(1) Vernonia prealta, Iron Weed,
[Vernonia praealta Michx.; V. gigantea (Walter) Trel.
ssp. gigantea; tall ironweed]
(2) Baptisia cerulea, Blue Wild Indigo,
[Baptisia coerulea Eaton & Wright; B. australis (L.)
R.Br. ex W.T. Aiton; blue false-indigo]
(3) Cacalia reniformis, Kidney Weed,
[Cacalia reniformis Muhl. ex Willd.; C. muhlenber-
gii (Sch. Bip.) Fern; Aroglossum muehlenbergii
(Sch. Bip.) H. Rob.; great Indian-plantain; see Ap-
pendix 2, entry 1, for discussion of nomenclature of
this species]
(4) Stedeoma pulegivides {Hedeoma_pule-
gioides}, Penny-aoyal {Penny royal},
[Hedeoma pulegioides (L.) Pers.; American penny-
royal, bastard pennyroyal, blue-curls]
(5) Chenopodium anthelminthicum, Worm
Weed,
[Chenopodium anthelminticum L.; C. ambrosioides
L. var. anthelminticum (L.) A. Gray; wormseed; see
Appendix 2, entry 2, for discussion of nomenclature
of this species]
(6) Elephantopus scaber, Elephant’s Foot,
[Elephantopus scaber L.; E. carolinianus Raeusch.;
leafy elephant’s-foot; see Appendix 2, entry 3, for
discussion of nomenclature of this species]
(7) Gillenia stipulacea, Indian Physic,
[Gillenia stipulata (Muhl. ex Willd.) Nutt.; Porter-
anthus stipulatus (Muhl. ex Willd.) Britton; Ameri-
can ipecac]
(8) Miagia arupedinaria {Miegia arundinar-
ia}, Cane, &c.&c.
[Miegia arundinaria Raf., nom. nud., presumably
intended orthography; Arundinaria gigantea (Wal-
ter) Muhl.; giant cane]
I consider the state of Kentucky as divided
into four natural sections, or botanical regions,
which are all distinguished by some peculiar-
ities in their vegetation. They are:
1. THe Fuuviatite Recion. This includes
all the valleys, and bottoms of the large rivers,
Rafinesque Common Names—Stuckey and Pringle 13
such as the Ohio, Mississippi, Tennessee,
Cumberland, Kentucky, &c. with their tribu-
tary streams. The bottoms of the valleys are
formed of an alluvial soil, or the washings from
the hills. They are level and often overflowed:
while the sides of the valleys are steep, craggy,
and composed of limestone, sandstone, or sla-
ty rocks. The following are some of the trees
and plants peculiar to this region, and giving
a decided character to its vegetation:
(9) Platanus occidentalis, Sycamore or But-
ton wood,
[Platanus occidentalis L.; sycamore, buttonwood]
(10) Hesperis pinnatifida, Ohio Wall Flower,
[Hesperis pinnatifida Michx.; lodanthus pinnatifidus
(Michx.) Steud.; purple rocket]
(11) Jeffersonia cinata {Jeffersonia binata},
Tavin Weed {Twin leaf},
[Jeffersonia binata Barton; J. diphylla (L.) Pers.;
twinleaf]
(12) Capraria multifida, Sand Ragweed,
[Capraria. multifida Michx., Leucospora multifida
(Michx.) Nutt.; see Appendix 2, entry 4, for discus-
sion of nomenclature of this species]
(13) Solanum Carolinianum, Sand Briar
[An orthographic variant of Solanum carolinense L.,
possibly preferred and originated by Rafinesque;
horse-nettle]
(14) Lupatorium calutinum {Eupatorium
coelestinum}, Sy-weed {Sky weed},
[Eupatorium coelestinum L.; mist-flower]
(15) Polanina {Polanisia} graveolens, Stink-
ing weed,
[Polanisia graveolens Raf.; Polanisia dodecandra
(L.) DC. ssp. dodecandra; clammy-weed]
(16) Heliotropium Indicum, Heliotrope,
{Heliotropium indicum L.; Indian heliotrope, turn-
sole; see Appendix 2, entry 5, for discussion of no-
menclature of this species]
(17) Catalpium cordata, Catalpa tree,
[Catalpium cordifolium (J.St.-Hil.) Raf.; Catalpa
speciosa (Warder ex Barney) Engelm.; northern ca-
talpa; see Appendix 2, entry 6, for discussion of no-
menclature of this species]
(18) Populus angulata, Cotton tree,
[Populus angulata Aiton; P. deltoides Marshall var.
deltoides; cottonwood]
(19) Porcelia tribuba {Porcelia triloba}, Pa-
paw tree,
[Porcelia triloba (L.) Pers.; Asimina triloba (L.)
Dunal; pawpaw]
(20) Synandra grandiflora, Cow mint,
[Synandra grandiflora Nutt.; S. hispidula (Michx.)
Baill.; see Appendix 2, entry 7, for discussion of no-
menclature of this species]
(21) Nelumbium pentapetalum, Swamp lily,
[Nelumbium pentapetalum (Walter) Willd.; Nelum-
bo lutea (Willd.) Pers.; American lotus, yellow lo-
tus]; see Appendix 2, entry 8, for discussion of no-
menclature of this species]
(22) Pancratium liviosone {Pancratium lir-
iosme}, Lily,
[Pancratium liriosme Raf.; Hymenocallis caroliniana
(L.) Herbert; spider-lily; see Appendix 2, entry 9,
for discussion of nomenclature of this species]
(23) Iris crocea, Red lily,
[Iris crocea Raf., nomen nudum; presumably I. ful-
va Ker Gawl.; copper iris]
(24) Houstonia fruticosa, Rock weed,
[Houstonia fruticosa Raf., nomen nudum; H. nigri-
cans (Lam.) Fernald var. nigricans; narrow-leaved
houstonia; see Appendix 2, entry 10, for discussion
of nomenclature of this species]
(25) Prunus pendula, Cliff plumb, &c.&c.
[Prunus pendula Raf., nomen nudum, non Maxim.
nec K.Koch; not definitely identified, probably P.
munsoniana W. Wight & Hedrick; wild plum; see
Appendix 2, entry 11, for discussion of nomencla-
ture of this species]
These two last are new species from the
cliffs of the Kentucky river.
2. THe Centra REcion. It is formed by
the limestone tract included between the val-
ley of the Ohio and the hilly ridges or knobs.
The ground is slightly broken, very fertile and
mostly under cultivation. This section is re-
markably poor in the number of botanical spe-
cies growing spontaneously; I conceive that its
flora hardly contains 500 species, including
trees, shrubs, and naturalized plants! There
are hardly any species peculiar to it; but the
following ones, rare elsewhere, are here very
common:
(26) Eupatorium urticefolium, White nettle,
[Eupatorium urticifolium Reichard, “urticaefolium”;
E. rugosum Houtt.; white snakeroot]
(27) Pavia muricata, Prickly Buck-eye,
[Pavia muricata Raf., nomen nudum; Aesculus gla-
bra Willd. var. glabra; Ohio buckeye]
14 Transactions of the Kentucky Academy of Science 58(1)
(28) Isanthus ceruleus, Blue Penny-royal,
[Rafinesque’s preferred orthography for Isanthus
coeruleus Michx.; I. brachiatus (L.) BSP.; False pen-
nyroyal |
(29) Polymnia uvedalia, Scented Sun flower,
[Polymnia uvedalia L.; large-flowered leafcup, yel-
low-flowered leafcup]
(30) Phlox glaberrima, Pink, &c.&c.
[Phlox glaberrima L.; smooth phlox]
It is also highly singular that in this region,
the woods are open as parks, without shrubs
and with very few plants, except grass or some
social weeds.
3. THe Hiniy Recion. It contains the hills
and ridges which divide the waters of the Ken-
tucky, Green, Licking, Cumberland and Sandy
rivers, &c. being spurs from the Cumberland
mountains. Those hills are often called knobs,
although they have not always the knobby or
rounded appearance. The rocks are limestone,
or sandstone, or slate. The vegetation approx-
imates exceedingly to that of Virginia and
Pennsylvania. On the Cumberland mountain
and the highest ridges, I am told that there is
a similarity with the Alleghany regions, and
that the Kalmia latifolia, Common Laurel, and
the Gaultheria procumbens, Mountain Tea,
grow there; but having not yet visited them, I
am unable to ascertain whether they ought to
form another distinct region, which might be
called the mountain region. The hilly region is
rich in plants; I shall mention a few of those
peculiar to it in Kentucky:
(31) Iris cristata, Crested Tris or Flag,
[Iris cristata Aiton; crested iris, crested dwarf iris]
(32) Stylvianthes {Stylosanthes} elatior, Yel-
low Pea-clover,
[Stylosanthes elatior Raf., nomen nudum; S. biflora
(L.) Britton et al.; pencil-flower]
(33) Orchis ciliaris, Yellow-bunch,
[Orchis ciliaris L.; Platanthera ciliaris (L.) Lindl.;
orange fringed-orchid, yellow fringed-orchid]
(34) Juniperus Virginiana, Red Cedar,
[Juniperus virginiana L.; red cedar]
(35) Vaccinium album, Wild Currant,
[Vaccinium album Pursh, non L. nec Lam.; V. stam-
ineum L.; deerberry]
(36) Pinus rigida, Pitch Pine,
[Pinus rigida Mill.; pitch pine]
(37) Lechea minor, Pin week,
[Lechea minor L.; small pinweed]
(38) Rudbeckia fulgida, Rough Wort,
[Rudbeckia fulgida Aiton, presumably var. fulgida;
orange coneflower, eastern coneflower]
(39) Gerardia glabrata, Yellow Wort,
[Gerardia glabrata Raf., nomen nudum; not defi-
nitely identified, probably Aureolaria laevigata (Raf.)
Raf.; Appalachian false-foxglove; see Appendix 2,
entry 12, for discussion of nomenclature of this spe-
cies|
(40) Asarum Virginicum, Heart-leaf, &c.&c.
[Asarum virginicum L.; Hexastylis virginica (L.)
Small; heart-leaf, little brown jugs; see Appendix 2,
entry 13, for discussion of identity of this species]
4. THe BarrREN REGION, or rather the
open region. This has an extensive range in
Kentucky, particularly in the western and
southern parts of the state. The numerous
barrens and licks compose it, lying scattered
and irregularly among the central and hilly
regions. The barrens are tracts of ground des-
titute of trees, or with few scattered small
ones; but thickly covered with a luxuriant
growth of plants; while the licks are almost
destitute of them, and those that grow in
their immediate neighbourhood are all small,
which is owing to their poor, slaty or argilla-
ceous soil. Their vegetation is however simi-
lar to that of the barrens. Both have a growth
of plants very similar to the vegetation of the
prairies of Ohio, Indiana, and Illinois, and
more different from that of the Atlantic
states, than the three foregoing regions. The
plants peculiar to them are very numerous; I
shall mention only a few, among the most re-
markable and singular.
(41) Solidago rigida, Stiff Golden-rod,
[Solidago rigida L.; stiff goldenrod]
(42) Polygala polygama, Nimble weed,
[Polygala polygama Walter; racemed milkwort, bit-
ter milkwort]
(43) Rudbeckia purpurea, Purple Sun-flow-
eX,
[Rudbeckia purpurea L.; Echinacea purpurea (L.)
Moench; purple coneflower]
Rafinesque Common Names—Stuckey and Pringle 15
(44) Ruellia oblongifolia, Rough Bell,
[Ruellia oblongifolia Raf., nomen nudum, Raf. ex
Nees, pro syn., non Michx.; R. caroliniensis (J.F.
Gmel.) Steud. ssp. ciliosa (Pursh) R.W. Long; hairy
ruellia]
(45) Andropogon arenaceum, Barren Oats,
[Andropogon avenaceus Michx.; Sorghastrum nu-
tans (L.) Nash; Indian grass]
(46) [Andropogon] nutans, Barren Oats,
[Andropogon nutans L.; Sorghastrum nutans (L.)
Nash; Indian grass]
(47) Petalvitemon {Petalostemon} candidum,
Nimble clover,
[Petalostemon candidum (Michx. ex Willd.) Michx.;
Dalea candida Michx. ex Willd.; white prairie-clo-
ver|
(48) [{Petalosteum}] purpureum, Nimble
clover,
[Petalostemon purpureum (Vent.) Rydb.; Dalea pur-
purea Vent. var. purpurea; purple prairie-clover]
(49) Silphium therebinthaceum, Turpentine
weed,
[Presumably an intentional abridgement of Sil-
phium terebinthinaceum Jacq.; prairie-dock]
(50) Silene catesbri {Silene catesbei}, Scarlet
Pink,
[Silene catesbaei Walter; S. virginica L.; fire-pink]
(51) Gentiana amarellvides {Gentiana amar-
elloides}, Yellow Gentian,
[Gentiana amarelloides Michx.; Gentianella quin-
quefolia (L.) Small var. quinquefolia; stiff gentian;
see Appendix 2, entry 14, for discussion of nomen-
clature of this species]
(52) Buchnera Americana, Black Wort,
&c.&e.
[Buchnera americana L.; blue-hearts]
From the above a faint, but correct idea
may be formed of the display and peculiarities
of the wide range of vegetation in Kentucky
and throughout the western states, wherein
the same peculiar divisions or regions may be
traced.
The vulgar names of the plants above men-
tioned are such as I found used in some parts
of Kentucky; but they cannot claim to be gen-
erally understood even in this state, many be-
ing merely local or personal. The botanical
names are alone to be relied on, being uni-
versal and not liable to mislead.
APPENDIX 1: Indexes to Botanical and
Common Names of Plants as Originally Listed
and Those Added to Rafinesque’s 1819 Paper
on the Botany of Kentucky.
A. Botanical or Scientific Names
Aesculus glabra Willd.
var. glabra (27)
Andropogon arenaceum (45)
Andropogon avenaceus Michx. (45)
Andropogon nutans L. (46)
Aroglossum muehlenbergii (Sch. Bip.) H. Rob. (3)
Arundinaria gigantea (Walter) Muhl. (8)
Aureolaria laevigata (Raf.) Raf. (39)
Asarum virginicum L. (40)
Asimina triloba (L.) Dunal (19)
Baptisia australis (L.) R. Br. ex W.T. Aiton (2)
Baptisia cerulea (2)
Baptisia coerulea Eaton & Wright (2)
Buchnera americana L. (52)
Cacalia muhlenbergii (Sch. Bip.) Fernald (3)
Cacalia reniformis Muhl. ex Willd. (3)
Capraria multifida Michx. (12)
Catalpa speciosa (Warder ex Barney) Engelm. (17)
Catalpium cordata (17)
Catalpium cordifolium (J. St.-Hil.) Raf. (17)
Chenopodium anthelminthicum L. (5)
Chenopodium ambrosioides L.
var. anthelminticum (L.) A. Gray (5)
Dalea candida Michx. ex Willd. (47)
Dalea purpurea Vent.
var. purpurea (48)
Echinacea purpurea (L.) Moench (43)
Elephantopus carolinianus Raeusch. (6)
Elephantopus scaber L. (6)
Eupatorium celestinum (14)
Eupatorium coelestinum L. (14)
Eupatorium rugosum Houtt. (25)
Eupatorium urticefolium (26)
Eupatorium urticifolium Reichard (26)
Gentiana amarelloides Michx. (51)
Gentiana amarellvides (51)
Gentiana quinquefolia (L.) Small
var. quinquefolia (51)
Gerardia glabrata Raf. (39)
Gillenia stipulacea (7)
Gillenia stipulata (Muhl. ex Willd.) Nutt. (7)
Hedeoma pulegioides (L.) Pers. (4)
Heliotropium indicum L. (16)
Hesperis pinnatifida Michx. (10)
Hexastylis virginica (L.) Small (40)
Houstonia fruticosa Raf. (24)
Houstonia nigricans (Lam.) Femald
var. nigricans (24)
Hymenocallis caroliniana (L.) Herbert (22)
Iodanthus pinnatifidus (Michx.) Steud. (10)
Iris cristata Aiton (31)
Iris crocea Raf. (23)
16 Transactions of the Kentucky Academy of Science 58(1)
Iris fulva Ker Gawl. (23) Stylvianthes elatior (32)
Isanthus brachiatus (L.) BSP. (28) Synandra grandiflora Nutt. (20)
Isanthus caeruleus Michx. (28) Synandra hispidula (Michx.) Baill. (20)
Isanthus ceruleus (28) Vaccinium album Pursh (35)
Jeffersonia binata (11) Vaccinium stamineum L. (35)
Jeffersonia cinata (11) Vernonia gigantea (Walter) Trel.
Jeffersonia diphylla (L.) Pers. (11) ssp. gigantea (1)
Juniperus virginiana L. (34) Vernonia praealta Michx. (1)
Lechea minor L. (37) Vernonia prealta (1)
Leucospora multifida (Michx.) Nutt. (12)
Lupatorium calutinum (14)
Miagia arupedinaria (8)
Miegia arundinaria Raf. (8)
Nelumbium pentapetalum (Walter) Willd. (21) Ammericaublotas OD)
Nelumbo lutea (Willd.) Pers. (21) American pennyroyal (4)
Orchis ciliaris L. (33) Barren oats (45, 46)
Pancratium liriosme Raf. (22)
B. Index to Common or Vulgar Names
Appalachian false-foxglove (39)
American ipecac (7)
Bastard pennyroyal (4)
Pancratium liviosone (22) Bitter milkwort (42)
Pavia muricata Raf. (27) Black wort (52)
Petalostemon candidum (Michx. ex Willd.) Michx. (47) Blue-eurs(4)
Petalostemon purpureum (Vent.) Rydb. (48) Blue-hearts (52)
Petalvitemon candidum (47) Blue penny-royal (28)
Phlox glaberrima L. (30) Blue false-indigo (2)
Pinus rigida Mill. (36) Blue wild indigo (2)
Platanus occidentalis L. (9) Button wood (9)
Platanthera ciliaris (L.) Lindl. (33) Buttonwood (9)
Polanina graveolens (15) Cane (8)
Polanisia graveolens Raf. (15) Catalpa tree (17)
Polanisia dodecandra (L.) DC. Clammy-weed (15)
ssp. dodecandra (15) Cliff plumb (25)
Polygala polygama Walter (42) Copper iris (23)
Polymnia uvedalia L. (29) Cotton tree (18)
Populus angulata Aiton (18) Cottonwood (18)
Populus deltoides Marshall Cow mint (20)
var. deltoides (18) Crested dwarf iris (31)
Porcelia tribuba (19) Crested flag (31)
Porcelia triloba (L.) Pers. (19) Crested iris (31)
Porteranthus stipulatus (Muhl. ex Willd.) Britton (7) Crested tris (31)
Prunus munsoniana W. Wight & Hedrick (25) Deerberry (35)
Prunus pendula Raf. (25) Eastern coneflower (38)
Ruellia caroliniensis (J.F. Gmel.) Steud. Elephant’s foot (6)
ssp. ciliosa (Pursh) R.W. Long (44) False pennyroyal (28)
Ruellia oblongifolia Raf. (44) Fire-pink (50)
Rudbeckia fulgida Aiton (38) Flag (31)
Rudbeckia purpurea L. (43) Giant cane (8)
Silene catesbaei Walter (50) Great Indian-plantain (3)
Silene catesbei (50) Hairy ruellia (44)
Silene catesbri (50) Heart-leaf (40)
Silene virginica L. (50) Heliotrope (16)
Silphium terebinthaceum (49) Horse nettle (13)
Silphium terebinthinaceum Jacq. (49) Indian grass (45, 46)
Solanum carolinianum (13) Indian heliotrope (16)
Solanum caroliniense L. (13) Indian physic (7)
Solidago rigida L. (41) Iron weed (1)
Sorghastrum nutans (L.) Nash (45, 46) Kidney weed (3)
Stedeoma pulegioides (4) Large-flowered leafcup (29)
Stylosanthes biflora (L.) BSP. (32) Leafy elephant’s-foot (6)
Stylosanthes elatior Raf. (32) Lily (22)
Rafinesque Common Names—Stuckey and Pringle 17
Little brown jugs (40)
Mist-flower (14)
Narrow-leaved houstonia (24)
Nimble clover (47, 48)
Nimble weed (42)
Northern catalpa (17)
Ohio buckeye (27)
Ohio wall flower (10)
Orange coneflower (38)
Orange fringed-orchid (33)
Pawpaw (19)
Pawpaw tree (19)
Pencil-flower (32)
Penny-aoyal (4)
Penny-royal (4)
Pink (30)
Pin weed (37)
Pitch pine (36)
Prairie-dock (49)
Prickly buck-eye (27)
Purple coneflower (43)
Purple prairie-clover (48)
Purple rocket (10)
Purple sun-flower (43)
Racemed milkwort (42)
Red cedar (34)
Red lily (23)
Rock weed (24)
Rough bell (44)
Rough wort (38)
Sand briar (13)
Sand ragweed (12)
Scarlet pink (50)
Scented sun flower (29)
Sky-weed (14)
Small pinweed (37)
Smooth phlox (30)
Spider-lily (22)
Stiff gentian (51)
Stiff golden-rod (41)
Stiff goldenrod (41)
Stinking weed (15)
Swamp lily (21)
Sycamore (9)
Sy-weed (14)
Tall ironweed (1)
Tavin weed (11)
Turnsole (16)
Turpentine weed (49)
Twinleaf (11)
Twin weed (11)
White nettle (26)
White prairie-clover (47)
White snakeroot (26)
Wild currant (35)
Wild plum (25)
Wormseed (5)
Worm weed (5)
Yellow-bunch (33)
Yellow-flowered leafcup (29)
Yellow fringed-orchid (33)
Yellow gentian (51)
Yellow lotus (21)
Yellow pea-clover (32)
Yellow wort (39)
APPENDIX 2: Notes on the Botanical and
Common Names (by J.S.P.)
1. The nomenclature of this species is unsettled at the
time of this writing, not only because of differences of
opinion as to appropriate generic circumscriptions, but
also pending a decision on proposals to reject the name
Cacalia, or, alternatively, for conserved typification. If Ca-
calia is rejected or is typified otherwise than as noted be-
low, this species will presumably be placed in Arnoglossum
regardless of whether the genus is circumscribed so as to
include Synosma Raf. ex Britton & A. Brown; if Cacalia
is not rejected, but is conserved with C. atriplicifolia L.
as the type, the correct name for this species will be Ca-
calia muehlenbergii (Sch. Bip.) Fernald.
2. The use of “wormseed” in works by other authors
from the same period indicates that “worm weed” prob-
ably represents a typographical error rather than a version
used in Kentucky during Rafinesque’s time.
3. The name Elephantopus scaber L. remains the cor-
rect name for an accepted species, but not for a species
that occurs in Kentucky. Elephantopus scaber sensu Mi-
chaux has been identified as the species now called E.
carolinianus.
4. No common name has been located for Leucospora
multifida in recent references other than the generic
name, or the now-obsolete generic name Conobea, used
as such. If the use of “sand-ragweed” actually persists, it
might be revived for wider application, hyphenated be-
cause this species is not in the ragweed genus Ambrosia
(Asteraceae). If, however, this was a coinage that expired
with Rafinesque, a name that might seem to indicate a
relationship to the true ragweeds would not be ideal.
5. Heliotropium indicum L. is now extensively but spo-
radically adventive or naturalized in the southeastern
United States, and, despite Kentucky's inland location, it
is not inconceivable that Rafinesque might have found a
small population there as early as 1819. However, because
no other North American authors mentioned H. indicum
as a naturalized species until considerably later, and be-
cause to the present day H. indicum has remained spo-
radic in its North American occurrences, it hardly seems
credible that Rafinesque could have found this species in
such abundance that it was among those “giving a decided
character to [the] vegetation” of any part of Kentucky.
Rafinesque’s (1838) later description of the species he had
so identified, which he then called Elopia riparia Raf.,
appears to have been derived largely from published de-
scriptions of H. indicum, but inevitably one wonders if his
concept of the species was actually based on some other
boraginaceous species.
18 Transactions of the Kentucky Academy of Science 58(1)
6. Rafinesque presumably referred to the Catalpa spe-
cies that is native to Kentucky, viz. C. speciosa (Warder
ex Barney) Engelm., which he might have seen on his trip
to the mouth of the Ohio River in 1818. There seems to
be an error in Index Rafinesquianus (Merrill 1949) on this
point. It is probably appropriate to assume from the sim-
ilarity of the epithets, as Merrill evidently did, that Ca-
talpium cordatum Raf. (Rafinesque 1819c) was intended
to be the same as his C. cordifolium Raf. (Rafinesque
1819b), published slightly earlier, the second being either
a lapsus calami or an intentionally amended orthography
for the first. According to Merrill, Catalpium cordifolium
Raf. was tied nomenclaturally to Catalpa cordifolia J. St.-
Hil. Merrill apparently confused C. cordifolia J. St.-Hil.
of 1804 with C. cordifolia Moench of 1794, which is a
synonym of C. bignonioides (Rehder 1949). The illegiti-
mate homonym Catalpa cordifolia J. St.-Hil., however, is
a synonym of C. speciosa, according to Rehder; this fits
well with Rafinesque’s report of his Catalpium
cordifolium/C. cordatum from Kentucky.
7. We have located no common name for Synandra
hispidula in recent literature, except for the generic name
used as such. As “cow-mint” does not appear to be used
for any other species, it could be considered for this spe-
cies, which is in the mint family (Lamiaceae).
8. Some recent authors have advocated “lotus-lily” or
“water-lotus” for Nelumbo, presumably because Lotus is
the botanical name for a genus in the Fabaceae. With this
genus very generally being known simply as “lotus,” this
illustrates the recurrent question as to what extent stan-
dardized English-language names should vary from names
actually in common usage. In this case, one might also ask
whether “lotus-lily,” even if hyphenated, is all that much
preferable for standardization, since Nelumbo is not in the
Liliaceae.
9. The name Pancratium liriosme Raf., published 2
years earlier by Rafinesque (1817c), is the basionym of
Hymenocallis liriosme (Raf.) Shinners. The species thus
named is native to Arkansas, Louisiana, Oklahoma, and
Texas. According to Shinners (1951), Rafinesque (1817c),
in describing P. liriosme, “was relying on [Claude C.] Ro-
bin’s description of plants that the latter had observed wild
in Louisiana.” The only species of Hymenocallis native to
Kentucky is H. caroliniana (L.) Herb., formerly known as
H. occidentalis (Leconte) Kunth.
10. No satisfactory common name appears to be avail-
able for this species. In popular field guides, the generic
name Houstonia is used as a common name for species
of this genus, except for two species of different aspect
that are called “bluets.” Britton and Brown (1896-1898)
called this species “narrow-leaved houstonia,” when it was
known botanically H. angustifolia Michx., but in the pop-
ular field guide by Newcomb (1977), “narrow-leaved
houstonia” is used for H. tenuifolia Nutt. Complicating the
issue, some botanists prefer to include Houstonia in Hed-
yotis. The name madderwort has reportedly been applied
to Houstonia, but this may have been in a context extend-
ing to other genera in the Rubiaceae (madder family).
“Rock weed” has little to recommend it, having been ap-
plied to Asperula odorata L. and possibly other species.
11. From the habitat given by Rafinesque—‘cliffs of
the Kentucky River’—an anonymous reviewer has con-
cluded that Rafinesque’s Prunus pendula is probably P.
munsoniana W. Wight & Hedrick, a species that was not
otherwise recognized as distinct and given a scientific
name until 1911. Under the current rules of botanical no-
menclature, because Rafinesque provided no description
of his “new species,” and because another botanist had
used the name Prunus pendula for a different species 4
years earlier, this interpretation does not affect the no-
menclature of P- munsoniana.
12. On the basis of Rafinesque’s phytogeographic com-
ments, Aureolaria laevigata, “Appalachian false-foxglove,”
seems the most likely identity of this species, but A. flava
(L.) Farw., “smooth false-foxglove,” cannot be ruled out.
13. Rafinesque’s “outline of four vegetation regions of
Kentucky” was published before its author visited that
portion of Kentucky in which Hexastylis species occur; for
this reason it has been suggested that his Asarum virgin-
icum might not have been any species now referred to
Hexastylis, the alternative interpretation being that it was
actually A. canadense L. This seems unlikely because Raf-
inesque presumably would have been familiar with A. can-
adense under its correct name and because, in all of the
literature available prior to 1819 in which A. virginicum
is described, that species is said to have glabrous, mottled
leaves. (Similarly, Rafinesque had not at that time traveled
into areas of Kentucky where any species of Lechea is
native, but one would hardly assume that he had misiden-
tified some other genus as Lechea.) Even if Rafinesque’s
concept of the “Hilly Region” included the Knobs that are
east of Lexington but west of the Pottsville escarpment, it
seems likely that some of his information, at least in this
relatively brief section of the paper, was more or less sec-
ondhand, perhaps derived from a combination of accounts
from other naturalists, studies of specimens in their her-
baria, and extrapolations from his firsthand knowledge of
the flora of adjacent states.
14. The name Gentiana amarelloides Michx. is a het-
erotypic synonym of Gentianella quinquefolia (L.) Small
var. quinquefolia, but in Kentucky this species is repre-
sented by var. occidentalis. From the Medical Flora, it is
evident that Rafinesque (1828) considered the plants with
yellow corollas to be a different species from the more
widespread form with purple corollas.
LITERATURE CITED
Britton, N.L., and A. Brown. 1896-1898. An illustrated
flora of the northern United States, Canada and the
British possessions .... Charles Scribner's Sons, New
York, NY.
Femald, M.L. 1950. Gray's manual of botany. 8th ed.
American Book Company, New York, NY. Reprinted
1970, D. Van Nostrand Co., New York, NY.
Gleason, H.A., and A. Cronquist. 1991. Manual of vas-
cular plants of northeastern United States and adjacent
Rafinesque Common Names—Stuckey and Pringle 19
Canada. 2nd ed. New York Botanical Garden, New
York, NY.
McAtee, W.L. 1913-1933. Some local names of plants,
I_-V. Torreya 13:225-236 (1913); 16:235—-242 (1916); 20:
17-27 (1920); 26:3-10 (1926); 33:81-86 (1933).
Merrill, E.D. 1949. Index Rafinesquianus: the plant
names published by C.S. Rafinesque with reductions,
and a consideration of his methods, objectives, and at-
tainments. Arnold Arboretum of Harvard University, Ja-
maica Plain, MA.
Newcomb, L. 1977. Newcomb’s wildflower guide. Little,
Brown and Company, Boston, MA.
Peterson, R.T., and M. McKenny. 1968. A field guide to
wildflowers of northeastern and north-central North
America. Houghton Mifflin Company, Boston, MA.
Rafinesque, C.S. 1811. Botanical information concerning
two families of plants. I. Species of the genus Callitri-
che. II. North American species of the genus Potamo-
geton. Med. Repos., third hexade 2(4):407—409.
Rafinesque, C.S. 1817a. [Review of Amos Eaton’s] A
manual of botany for the northern states, comprising
generic descriptions of all phenogamous and cryptoga-
mous plants to the north of Virginia. Am. Monthly Mag.
& Crit. Rev. 1(6):426-430.
Rafinesque, C.S. 1817b. Florula of the White Mountain
of New-Hampshire. Am. Monthly Mag. & Crit. Rev.
1(6):440-442.
Rafinesque, C.S. 1817c. Florula Ludoviciana: or, a flora
of the state of Louisiana. C. Wiley & Co., New York,
NY. Facsimile ed. 1967, with introduction by Joseph
Ewan, Hafner Publishing Co., New York, NY.
Rafinesque, C.S. 1819a. Prodrome des nouveaux genres
de plantes observés en 1817 et 1818 dans l’intérieur des
états-Unis d’Amérique. J. Phys. Chim. Hist. Nat. Arts
89(2):96-107.
Rafinesque, C.S. 1819b. [Review of] The genera of
North-American plants and a catalogue of the species
to the year 1817. By Thomas Nuttall, F.L.S. &c. 2 vols.
12 mo. Philadelphia, PA. 1818. Am. Monthly Mag. &
Crit. Rev. 1(3):184—196.
Rafinesque, C.S. 1819c. Botany of Kentucky[:] on its
principal features. Western Rev. & Misc. Mag. 1(2):92-
95.
Rafinesque, C.S. 1819d. Errata [to botany of Kentucky].
Western Rev. & Misc. Mag. 1(3):128.
Rafinesque, C.S. 1820. Prodrome d’une monographie des
rosiers del Amérique septentrionale, contenant la de-
scription de quinze nouvelles espéces et vingt variétés.
Ann. Gén. Sci. Phys. 5(14):210-220.
Rafinesque, C.S. 1824. Florula Kentuckensis. Catalogue
of the principal trees, shrubs and plants of Kentucky.
Pages [12]-16 in First catalogues and circulars of the
Botanical Garden of Transylvania University at Lexing-
ton in Kentucky, for the year 1824. Printed for the Bo-
tanical Garden Company by John M. M’Calla, Lexing-
ton, KY.
Rafinesque, C.S. 1828. Medical flora; or, Manual of the
medical botany of the United States of North America
., volume the first. Atkinson & Alexander, Philadel-
phia, PA.
Rafinesque, C.S. 1830. Medical flora; or Manual of the
medical botany of the United States of North America
..., volume the second. Samuel C. Atkinson, Philadel-
phia, PA.
Rafinesque, C.S. 1836-[1838]. New flora of North Amer-
ica. Published by the author, Philadelphia, PA. Facsim-
ile ed. 1946, Arnold Arboretum, Jamaica Plain, MA.
Rafinesque, C.S. “1836” [1837-1838]. Flora Telluriana.
Published by the author, Philadelphia. Facsimile ed.
1946, Arnold Arboretum, Jamaica Plain, MA.
Rafinesque, C.S. 1838. Sylva Telluriana. Published by the
author, Philadelphia. 184 pp. Facsimile ed. 1943, Ar-
nold Arboretum, Jamaica Plain, MA.
Rafinesque, C.S. 1840. Autikon botanikon or botanical
illustrations .... Published by the author, Philadelphia,
PA. Facsimile ed. 1942, Armold Arboretum, Jamaica
Plain, MA.
Rehder, A. 1949. Bibliography of cultivated trees and
shrubs hardy in the cooler temperate regions of the
northern hemisphere. Arnold Arboretum of Harvard
University, Jamaica Plain, MA.
Shinners, L.H. 1951. The north Texas species of Hymen-
ocallis (Amaryllidaceae). Field & Lab. 19:102-104.
Trans. Ky. Acad. Sci. 58(1):20—22. 1997.
Some Comments on Constantine Rafinesque’s 1819 Description of
Botanical Regions in Kentucky
William S. Bryant
Department of Biology, Thomas More College, Crestview Hills, KY 41017
Perhaps of greatest ecological significance
in Rafinesque’s (1819a, 1819b) Botany of Ken-
tucky was his recognition of distinct botanical
regions or natural sections in the state. His
regions and plant associations were not so well
defined as those currently recognized (e.g.,
Braun 1950: Kiichler 1964; Quarterman and
Powell 1978), but at least they were a first at-
tempt. Meijer (1973) noted that Rafinesque
often had good ideas of natural relationships
of vegetation as distinguished by its peculiar-
ities.
Rafinesque (1819a) recognized four botan-
ical regions but suggested that a fifth, the
Mountain Region, was probable. He had nei-
ther collected in nor visited the eastern por-
tions of Kentucky. His entrance into Kentucky
was from Pennsylvania via the Ohio River in
1818. From 1819-1825, he was Kentucky's
first professional resident botanist, teaching at
Transylvania College in Lexington (Meijer
1973). It seems apparent that Rafinesque was
familiar with the literature concerning some
aspects of natural history on the frontier as
detailed by Drake (1815), Filson (1784), Imlay
(1797), and Michaux (1904). Kentucky was
only 26 years old as a state when Rafinesque
arrived in 1818 and only 44 years removed
from the establishment of its first permanent
settlement.
Rafinesque’s botanical regions of Kentucky
are listed and described as follows:
1. The Fluviatile Region included the val-
leys and bottoms of large rivers and their trib-
utaries. Rafinesque especially noted the allu-
vial deposits but also included the steep-sided
gorges bordering many of these streams. His
inclusion of the Cumberland and Kentucky
rivers might be expected, but his inclusion of
the Tennessee River, which prior to 1818
formed the state’s western border, and the
Mississippi River, which after the 1818 Jack-
son Purchase became the western border, sug-
gests that Rafinesque was abreast of happen-
ings in the Commonwealth. His recognition of
20
a Fluviatile Region does not fit most modern
regional designations; however, Middleton et
al. (1926) stated that “the Alluvial land of the
Mississippi River bottoms constitutes the fifth
distinct area of the state.” The Ohio River low-
lands was also designated as a distinct geo-
graphical region (Burroughs 1926). Cotterill
(1917) noted that early maps of Kentucky
“seemed a mere network of rivers” (Figure 1).
Rafinesque’s list of plants and trees peculiar
to this region was limited. He included syca-
more (Platanus occidentalis) and cottonwood
(Populus deltoides), which are widely distrib-
uted throughout the state, but his inclusion of
catalpa (Catalpa speciosa) and spiderlily (Hy-
menocallis caroliniana) suggests that he had
some knowledge of the western third of the
state where these two species are found in
lowland forests.
2. The Central Region is the Bluegrass Re-
gion, which is bordered by the Knobs. He not-
ed that this fertile limestone area was mostly
under cultivation even in 1819. He also rec-
ognized a low species diversity including na-
tive and naturalized plants. Meijer (1973)
pointed to Rafinesque’s interest in the flora of
introduced weeds and how this invasion co-
incided with the arrival of white settlers.
Rafinesque’s statement that “it is also highly
singular that in this region, the woods are
open as parks, without shrubs and with very
few plants, except grass or some social weeds,”
is significant. He was describing the blue ash-
oak savanna woodland, which was the char-
acteristic vegetation of the Bluegrass region
(Bryant 1983; Bryant et al. 1980). His list of
species in no way characterizes this region.
3. The Hilly Region included the Knobs as
well as extensions of broken ground from the
Big Sandy River in the northeast to the Green
River in the west. The geologic substrate men-
tioned by Rafinesque included sandstone,
limestone, or slate. He found that the vege-
tation approximated that of Pennsylvania and
Virginia, with which he was familiar. The
Comments on Rafinesque—Bryant Dil
u to
= SS Se
a
= SSS eee to
—
7
~
‘ CORSET),
¢() Ye Wenpeky
Bu SS
ULRs.
i
hy COW ay tes,
Figure 1. The Samuel Lewis 1814 map of Kentucky showing the principal features. Note the lack of information on
the mountains of eastern Kentucky but the detail of the many streams and rivers. After Clark (1979).
plants listed, especially redcedar (Juniperus
virginiana), pitch pine (Pinus rigida), and
deerberry (Vaccinium stamineum) are found
on various substrates in the Knobs Region.
Muller and McComb (1983) described the up-
land vegetation of the Knobs as being domi-
nated by oaks, none of which Rafinesque men-
tioned.
4. The Mountain Region was included with
the Hilly region, but Rafinesque acknowl-
edged that he had not visited this part of the
state. He had been told of the similarity of the
mountains to the Allegheny region. This re-
gion included the Cumberland Mountains and
the highest ridges. The presence of mountain
laurel (Kalmia latifolia) and mountain tea
(Gaultheria procumbens) and a few of the
plants listed for the Hilly Region are inade-
quate to describe this area. Braun (1950) char-
acterized this portion of eastern Kentucky as
the center of the mixed mesophytic forest, the
richest forest type in eastern North America.
5. The Barren Region encompassed the
open country in the western and southern
parts of the state. Rafinesque noted that the
vegetation consisted of “a growth of plants
very similar to that of the prairies of Ohio,
Indiana, and Illinois.” He pointed out the
sparsity of trees and the presence of licks,
which tended to attract grazing animals (e.g.,
buffalo). Rafinesque’s list of plants of this re-
gion is perhaps more representative than his
lists for the other regions he described. Blue-
stems (Andropogon spp.), Indian grass (Sor-
ghastrum nutans), prairie-clovers (Petaloste-
mon candidum and P. purpureum), prairie-
dock (Silphium terebinthinaceum), and purple
coneflower (Echinacea purpurea) are still
found in barren remnants (Baskin and Baskin
1978; Braun 1950). Rafinesque was most in-
terested in the Barrens; along with his collab-
orator, Dr. Charles Short, he collected there
extensively (Meijer 1973). Rafinesque’s Barren
Region is part of the Pennyroyal or Mississippi
Plateau.
As can be determined from the above de-
scriptions of Botanical Regions, Rafinesque in
1819 was more familiar with those portions of
Kentucky from Lexington westward. Since he
travelled to and kept an association with some
of the residents of Harmonie (later New Har-
mony) in southwestern Indiana (Thompson
1898), it seems probable that he botanized on
his trips.
In conclusion, it was indeed fortunate that
Rafinesque made reference to different botan-
ical regions as being present in Kentucky. He
noted some of the plants, although not always
22. Transactions of the Kentucky Academy of Science 58(1)
those that might best characterize a commu-
nity or region. He mentioned topography, geo-
logic substrate, and soils as being of impor-
tance in regional differentiations. It is unfor-
tunate that he did not continue his interest
and study of plant associations at a time prior
to their manipulation and fragmentation by
the onrush of settlement.
LITERATURE CITED
Baskin, J.M., and C.C. Baskin. 1978. Plant ecology of
cedar glades in the Big Barrens region of Kentucky.
Rhodora 80:545-557.
Braun, E.L. 1950. Deciduous forests of eastern North
America. Blakiston Co., Philadelphia, PA.
Bryant, W.S. 1983. Savanna-woodland in the Outer Blue-
grass of Kentucky. Trans. Ky. Acad. Sci. 44:44—49.
Bryant, W.S., M.E. Wharton, W.H. Martin, and J.E. Var-
ner. 1980. The blue ash-oak savanna-woodland, a rem-
nant of pre-settlement vegetation in the Inner Bluegrass
of Kentucky. Castanea 45:149-165.
Burroughs, W.C. 1926. Geography of the Knobs. Ken-
tucky Geological Survey, Frankfort, KY.
Clark, T.D. 1979. Historic maps of Kentucky. Univ. Press
of Kentucky, Lexington, KY.
Cotterill, R. 1917. History of pioneer Kentucky. Johnson
& Hardin, Cincinnati, OH.
Drake, D. 1815. Natural and statistical view, or picture
of Cincinnati and the Miami Country. Looker and Wal-
lace, Cincinnati, OH.
Filson, J. 1784. The discovery, settlement, and present
state of Kentucky. James Adams, Wilmington, DE.
Imlay, G. 1797. A topographical description of the west-
em territory of North America. Debrett, London.
Kiichler, A. W. 1964. Potential natural vegetation of con-
terminous United States. Am. Geogr. Soc. Spec. Publ.
64.
Meijer, W. 1973. The contributions by Rafinesque to the
early botanical exploration of Kentucky. Castanea 38:
261-265.
Michaux, F.A. 1904. Travels to the west of the Allegheny
Mountains in the states of Ohio, Kentucky and Ten-
nessee, and back to Charleston, by the Upper Carolinas.
Pages 110-306 in R.G. Thwaites. 1904. Early Western
Travels 1746-1846. Arthur H. Clark Co., Cleveland,
OH.
Middleton, A.R., W.R. Jillson, FT. McFarland, and W.A.
Anderson Jr. 1926. Kentucky. Pages 149-154 in V.E.
Shelford (ed). Naturalist’s guide to the Americas. Wil-
liams & Wilkins Co., Baltimore, MD.
Muller, R.N., and W.C. McComb. 1983. Upland forests
of the Knobs Region of Kentucky. Bull. Torrey Bot.
Club 113:268-280.
Quarterman, E., and R.L. Powell. 1978. Potential ecol-
ogical/geological natural landmarks on the Interior Low
Plateaus. U.S. Dept. of Interior, National Park Service.
Rafinesque, C.S. 1819a. Botany of Kentucky. On its prin-
cipal features. Western Rev. & Misc. Mag. 1(2):92-95.
Rafinesque, C.S. 1819b. Errata [to botany of Kentucky].
Western Rev. & Misc. Mag. 1(3):128
Thompson, M. 1898. Stories of Indiana. American Book
Co., New York, NY.
Trans. Ky. Acad. Sci. 58(1):23-28. 1997.
Effect of Light on Daily Emergence of Cercariae of the Trematodes
Echinostoma trivolvis (Echinostomatidae) and Cephalogonimus
vesicaudus (Cephalogonimidae) from Natural Infections of the Snail
Helisoma trivolvis (Planorbidae) at Owsley Fork Reservoir, Kentucky
Ron Rosen, Peter Blair, Jeff Ellington, and Jason Backu
Department of Biology, Berea College, Berea, KY 40404
ABSTRACT
Naturally infected snails, Helisoma trivolvis, releasing cercariae of Echinostoma trivolvis and Cephalogon-
imus vesicaudus, were collected from Owsley Fork Reservoir in northern Jackson and southem Madison
counties, Kentucky, in June or July 1990, 1992, and 1995. Emergence of cercariae from snails was assessed
at 26° C for 24 h under the following photoperiod regimens: (1) 12 h light:12 h dark; (2) 12 h dark:12 h
light (inverted photoperiod); (3) 24 h light; and (4) 24 h dark. In the split photoperiods, maximum cercarial
emergence of E. trivolvis and C. vesicaudus was significantly greater in the 12 h of light and 12 h of dark,
respectively. Photoperiod inversion resulted in a complete reversal in timing of cercarial release for both
species. Continuous light or dark for 24 h induced continuous release of cercariae of both species. No
significant change was observed in the timing of peak cercarial emergence of C. vesicaudus in the 1990,
1992, and 1995 studies using a 12 h light:12 h dark photoperiod, but a marked difference was noted for E.
trivolvis in 1995 when peak cercarial release was shifted to the dark phase of the 24 h photoperiod. These
results are discussed with regard to (1) exogenous vs. endogenous factors affecting cercarial release and (2)
the use of cercarial emergence patterns for identifying parasitic strains.
INTRODUCTION
The timing of the emergence of trematode
(subclass Digenea) cercariae from their snail
host has been linked to activity of the next host
in the life cycle in several studies (Lewis,
Welsford, and Uglem 1989; Theron 1984), but
the physiological triggers associated with cer-
carial release are poorly understood. It has
been suggested that in-vitro cultivation of in-
tramolluscan stages in the absence of snail tis-
sue will be necessary to clarify the exact me-
chanisms/triggers (Glaudel and Etges 1973).
In addition to a possible endogenous circadian
rhythm, these triggers may be influenced by a
number of environmental factors including
temperature, humidity, oxygen, pH, and light
(Smyth and Halton 1983). These exogenous
factors either act directly on the parasite or
are mediated indirectly through the snail host.
That light triggers or inhibits cercarial re-
lease from the snail host in many species of
digenetic trematodes is well known. Reversal
of the 24 h light:dark photoperiod (i.e., pho-
toperiod inversion) often results in complete
reversal in the pattern of cercarial release
(Asch 1972; Giovannola 1936; Glaudel and Et-
ges 1973; Luttermoser 1955; Oliver 1951;
Wagenbach and Alldredge 1974). Such a re-
23
versal provides evidence that the pattern of
release is not a true circadian rhythm but is
mediated by the “rhythm of the snail which is
controlled to some extent by illumination”
(Asch 1972). However, a number of these
studies did not assess whether cercarial emer-
gence ceases to be rhythmic if the light or
dark is of a constant magnitude (i.e., contin-
uous light or dark). This criterion must be met
to rule out the presence of an endogenous cir-
cadian rhythm within the parasite itself (Wag-
enbach and Alldredge 1974).
Distinct strains of digenetic trematodes
have been documented in several studies
based on photoperiod and cercarial emer-
gence patterns (Gumble et al. 1957; Riley and
Uglem 1995; Theron 1984), but, to our knowl-
edge, shifting patterns of cercarial emergence
in a species in the same habitat over time have
not been observed. Further studies of this na-
ture might provide (1) insight into use of cer-
carial emergence patterns as stable biological
characteristics for some species of digenetic
trematodes and (2) a method for identifying
different strains of a species from the same
habitat.
Little information is available concerning
the effect of light on emergence of cercariae
of Echinostoma trivolvis (Echinostomatidae)
24 Transactions of the Kentucky Academy of Science 58(1)
(Schmidt and Fried 1996), and no studies of
this nature have been conducted with cercari-
ae of Cephalogonimus vesicaudus (Cephalo-
gonimidae). Both species are parasites of the
snail Helisoma trivolvis (Planorbidae) at Ows-
ley Fork Reservoir in northern Jackson and
southern Madison counties, Kentucky (Rosen
et al. 1994). Thus, the objectives of our study
were to determine (1) the effect of light on
the release of E. trivolvis and C. vesicaudus
cercariae from natural infections of H. trivol-
vis and (2) whether different strains of these
species exist in the reservoir based on photo-
period and cercarial emergence patterns as-
sessed over several years.
MATERIALS AND METHODS
Specimens of H. trivolvis were collected
from Owsley Fork Reservoir in June or July
1990, 1992, and 1995. A brief description of
the study site is provided in Rosen et al.
(1994). Within 2 h of their collection, snails
were placed individually into 50 ml beakers
filled with 40 ml of filtered reservoir water,
incubated at 26° C with a 12 h light (0700-
1859):12 h dark (1900-0659) cycle, and
screened for cercarial emergence at 1400,
2030, and 0830 for 1 day. Snails with mature
infections (ie., releasing cercariae of E. tri-
volvis or C. vesicaudus) were then immediate-
ly acclimated for 24 h at 26° C to one of the
following photoperiod regimens: (1) 12 h light
(0700-1859):12 h dark (1900-0659), (2) 12 h
dark (0700-1859):12 h light (1900-0659), (3)
24 h light, and (4) 24 h dark. Only the 12 h
light:12 h dark cycle was used to compare
emergence patterns in the 1990, 1992, and
1995 studies.
Following the initial 24 h acclimation peri-
od, cercarial counts were made every 2 or 4
hours for 24 h. At each time interval, snails
were transferred into new beakers and im-
mediately placed back in their designated pho-
toperiod regimens. The number of cercariae
in the original beaker was determined with the
aid of a dissecting microscope by direct counts
or by averaging ten 0.1 ml aliquots taken from
a uniform cercarial suspension that was then
adjusted to the number of cercariae/40 ml.
The latter technique was necessary when cer-
carial emergence was copious in a sampling
interval.
Cercarial release in the six 2 h light periods
and six 2 h dark periods were separately
pooled and compared with a Mann-Whitney
test to determine if the median cercarial re-
lease for the two species was significantly dif-
ferent in the split 24 h photoperiods. In the
12L:12D and 12D:12L analyses, the light time
interval consisted of 11.5 h of light and 0.5 h
of dark, while the dark interval consisted of
11.5 h of dark and 0.5 h of light due to the
times selected to evaluate cercarial emer-
gence. A Kruskal-Wallis test was used to de-
termine if the timing of the mean peak for
cercarial release during a 24 h period varied
significantly among the years 1990, 1992, and
1995 for E. trivolvis and C. vesicaudus. A
probability of P < 0.05 was considered signif-
icant for all statistical tests. Means are report-
ed with associated standard errors.
RESULTS
A significant difference (Mann-Whitney
test; P = 0.0233) was found in the median
cercarial release for E. trivolvis between the
light and dark intervals. An average of 156.7
+ 65.5 and 5.0 + 2.1 cercariae were released
in the light and dark periods, respectively.
Similarly, a significant difference (Mann-Whit-
ney test; P=0.0233) was found in the mean
cercarial release for this species in the invert-
ed photoperiod. An average of 5.0 + 2.1 and
272.7 + 41.3 cercariae were released in the
dark and light time intervals, respectively.
Maximum emergence occurred at 1430 and
0030; little or no emergence was observed
during the 2 h intervals of the dark periods
(Figures la and 1b). Continuous emergence
was apparent under conditions of constant
light (Figure lc; range = 12-187 cercariae/2
h interval) and dark (Figure 1d; range = 7—
130 cercariae/2h interval).
A significant difference (Mann-Whitney
test: P = .0001) was found for the median
cercarial release of C. vesicaudus between the
light and dark intervals. An average of 153.8
+ 57.5 and 2021 + 349.7 cercariae was re-
leased in the light and dark periods, respec-
tively. A significant difference (Mann-Whitney
test: P = .0003) was also found for the median
cercarial release in the inverted photoperiod.
An average of 1713.6 + 249.5 and 52.07 +
135.9 emerged in the dark and light time in-
tervals, respectively. Peak emergence was ob-
Cercarial Emergence—Rosen et al. 25
AVERAGE # CERCARIAE RELEASED / SNAIL
Figure 1. Effect of light on the mean number + SE of
Echinostoma trivolvis cercariae released from the snail
Helisoma trivolvis at 2 h intervals over 24 h at 26° C under
the following photoperiods: (a) 12 h light (0700-1859): 12
h dark (1900-0659), (b) 12 h dark (0700-1859):12 h light
(1900-0659), (c) 24 h light, and (d) 24 h dark. An average
of five infected snails was used for each photoperiod.
served at 2230 and 1230 and little or no emer-
gence occurred during the 2 h intervals of the
light periods (Figures 2a and 2b). Continuous .
cercarial emergence was observed under con-
ditions of constant light (Figure 2c; range =
499-2298 cercariae/2 h interval) and dark
(Figure 2d; range = 177-808 cercariae/2 h in-
terval).
The long-term study showed that the mean
daily time for peak cercarial emergence was
significantly different for E. trivolvis (Kruskal-
Wallis test: P = .0002), but not for C. vesi-
caudus (Kruskal-Wallis test: P=.9482). Peak
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1S)
*
io)
o
<
4
i)
>
<
Figure 2. Effect of light on the mean number + SE of
Cephalogonimus vesicaudus cercariae released from the
snail Helisoma trivolvis at 2 h intervals over 24 h at 26°
C under the following photoperiods: (a) 12 h light (0700-
1859):12 h dark (1900-0659), (b) 12 h dark (0700-1859):
12 h light (1900-0659), (c) 24 h light, and (d) 24 h dark.
An average of 15 infected snails was used for each pho-
toperiod.
emergence time for E. trivolvis shifted to
2230 (dark period) in 1995 from 1430 (light
period) in 1990 and 1992 (Figure 3). By com-
parison, the peak emergence time of C. vesi-
caudus remained fixed at 2230 (dark period)
over the same time (Figure 4).
DISCUSSION
Light significantly triggered the release of
E. trivolvis cercariae and inhibited the emer-
gence of C. vesicaudus cercariae in our 1990
and 1992 studies. Photoperiod inversion was
26 Transactions of the Kentucky Academy of Science 58(1)
AVERAGE # CERCARIAE RELEASED / SNAIL
TIME
Figure 3. Comparison of the effect of light on the mean
number + SE of Echinostoma trivolvis cercariae released
from the snail Helisoma trivolvis in 1990, 1992, and 1995.
Naturally infected snails were collected in June or July
from Owsley Fork Reservoir, Kentucky, and assessed at 4
h intervals over 24 h at 26° C under a 12 h light (0700-
1859):12 h dark (1900-0659) photoperiod. An average of
10 infected snails was used for each summer/experiment.
accompanied by a complete reversal in the
timing of peak cercarial release for both spe-
cies. This suggested that cercarial emergence
of the species is subject to control by the ex-
ogenous photoperiod under conditions of con-
stant temperature. By contrast, Schmidt and
Fried (1996) found that light vs. dark did not
affect the intensity of E. trivolvis cercarial
emergence. Their study provided no acclima-
tion period for snails to the tested conditions
(i.e., snails were maintained in the dark at 12°
C and then placed in conditions of light or
dark at 28-29° C for an immediate 1 hour
evaluation of cercarial emergence). The dif-
AVERAGE # CERCARIAE RELEASED / SNAIL
TIME
Figure 4. Comparison of the effect of light on the mean
number + SE of Cephalogonimus vesicaudus cercariae
released from the snail Helisoma trivolvis in 1990, 1992,
and 1995. Naturally infected snails were collected in June
or July from Owsley Fork Reservoir, Kentucky, and as-
sessed at 4 h intervals over 24 h at 26° C under a 12 h
light (0700—1859):12 h dark (1900-0659) photoperiod. An
average of 10 infected snails was used for each summer/
experiment.
ference between our results and those of
Schmidt and Fried (1996) may be more at-
tributable to experimental design than to pos-
sible strain differences of E. trivolvis.
Wagenbach and Alldredge (1974) found
that cercariae of the digenean Plagiorchis mi-
cracanthos, which normally emerge during the
dark phase of a 12 h light:12 h dark cycle,
show a rhythmic emergence in continuous
dark. They indicated that this latter phenom-
enon provided evidence for an innate emer-
gence rhythm, “which is synchronized to pho-
toperiod by the inhibition of light” (Wagen-
Cercarial Emergence—Rosen et al. 7
bach and Alldredge 1974). Such an “entrain-
ment” of the endogenous rhythm to light:dark
cycles has also been suggested for emergence
of Proterometra macrostoma cercariae, which
normally emerge during the dark phase of a
light:dark cycle but show peaks of emergence
under conditions of continuous light (Lewis,
Welsford, and Uglem 1989). The lack of such
a distinct rhythm under conditions of contin-
uous light or dark in our study suggests that
no circadian rhythm exists for E. trivolvis or
C. vesicaudus in H. trivolvis, but longer term
studies (i.e., in excess of 24 h) will be required
to clarify this.
No change was observed in timing of peak
cercarial emergence of C. vesicaudus in the
1990, 1992, and 1995 studies, but a significant
shift was noted for E. trivolvis. Cephalogoni-
mus vesicaudus is an autogenic species that
completes its life cycle as an adult in the small
intestine of the spiny softshell turtle Trionyx
spiniferus, which is present at Owsley Fork
Reservoir. The timing of peak cercarial release
is apparently a stable biological characteristic
for the population of this species at this locale.
By comparison, mammals (e.g., muskrats) and
a variety of migratory waterfowl (e.g., Cana-
dian geese, mallards, etc.) present at Owsley
Fork likely serve as definitive hosts for E. tri-
volvis in this habitat. The migratory nature of
the avian hosts provides an opportunity for pe-
riodic introduction of new strains of E. trivol-
vis into the existing Owsley Fork population
of this species. This may be indirectly assessed
by observation of new cercarial release pat-
terns appearing in the population over time.
In a somewhat similar study, Theron (1984)
documented chronobiological variation (i.e.,
one strain experienced an earlier daily cercar-
ial release than the second strain) in two
Schistosoma mansoni populations from the
same area, but different ecological foci. He
linked this variation to the amount of murine
host participation in the life cycle. Further ex-
periments determined that the cercarial emer-
gence pattern of S. mansoni has a genetic basis
that is a consequence of selective pressures
exerted by different host species (Theron and
Combes 1988). It has also been noted that the
response of Schistosoma japonicum cercariae
to light varies with the geographical strain of
the species (Gumble et al. 1957). Both daily
and seasonal differences in cercarial emer-
gence have been recorded for eight strains of
P. macrostoma (Riley and Uglem 1995). De-
velopment of these strains has been linked to
the species composition of sunfish definitive
hosts in a specific geographical region (Riley
and Uglem 1995). In the present study, since
both potential mammalian and waterfowl
hosts of E. trivolvis exist in the same “ecolog-
ical focus,” the resulting emergence patterns
for this species at Owsley Fork likely represent
blending of patterns from several different
strains. The changing prevalence of these dif-
ferent strains over time might explain the vari-
ation observed in the long-term emergence
patterns for E. trivolvis cercariae at the res-
ervoir. Molecular techniques recently used to
differentiate and establish relationships of spe-
cies in the genus Echinostoma (Morgan and
Blair 1995; Petrie, Burg, and Cain 1996; Sloos
et al. 1995) may be utilized in future studies
to establish genetic differences between these
proposed strains of E. trivolvis.
ACKNOWLEDGMENTS
This study was supported by a grant from
Merck & Company, the Andrew Mellon Foun-
dation Trust, and the Appalachian College As-
sociation to R. Rosen and the Department of
Biology at Berea College. We acknowledge
previous Berea College student participants
who contributed to this study including Mar-
ichelle Asuncion, Melissa Edlin, Jose Iagan,
Kiely Law, and Manuel San.
LITERATURE CITED
Asch, H.L. 1972. Rhythmic emergence of Schistosoma
mansoni cercariae from Biomphalaria glabrata: control
by illumination. Exp. Parasitol. 31:350-355.
Giovannola, A. 1936. Inversion in the periodicity of emis-
sion of cercariae from their snail hosts by reversal of
light and darkness. J. Parasitol. 22:292-295.
Glaudel, R.J., and FJ. Etges. 1973. The effect of pho-
toperiod inversion upon Schistosoma mansoni cercarial
emergence from Biomphalaria glabrata. Int. J. Parasi-
tol. 3:619-622.
Gumble, A., Y. Otori, L.S. Ritchie, and G.W. Hunter.
1957. The effect of light, temperature and pH on the
emergence of Schistosoma japonicum cercariae from
Oncomelania nosophora. Trans. Am. Microsc. Soc. 76:
87-92.
Lewis, M.C., I.G. Welsford, and G.L. Uglem. 1989. Cer-
carial emergence of Proterometra macrostoma and P.
edneyi (Digenea: Azygiidae): contrasting responses to
light:dark cycling. Parasitology 99:215-223.
Luttermoser, G.W. 1955. Studies on the chemotherapy of
28 Transactions of the Kentucky Academy of Science 58(1)
experimental schistosomiasis HI. Harvest of Schistoso-
ma mansoni cercariae by forced nocturnal emergence
from Australorbis glabratus. J. Parasitol. 41:201—208.
Morgan, J.A.T., and D. Blair. 1995. Nuclear rDNA ITS
sequence variation in the trematode genus Echinosto-
ma: an aid to establishing relationships within the
37-collar-spine group. Parasitology 111:609-615.
Oliver, L. 1951. The influence of light on the emergence
of Schistosomatium douthitti cercariae from their snail
host. J. Parasitol. 37:201-204.
Petrie, J.L., E.F. Burg IH, and G.D. Cain. 1996. Molec-
ular characterization of Echinostoma caproni and E.
paraensei by random amplification of polymorphic
DNA (RAPD) analysis. J. Parasitol. 82:360-362.
Riley, M.W., and G.L. Uglem. 1995. Proterometra ma-
crostoma (Digenea: Azygiidae): variations in cercarial
morphology and physiology. Parasitology 110:429-436.
Rosen, R.B., J.M. Ilagan, J.S. Law, M. Asuncion, M.E.
Denton, and M.L. San. 1994. Seasonal prevalence of
three species of digenetic trematodes in the snail Hel-
isoma trivolvis at Owsley Fork Reservoir, Kentucky.
Trans. Kentucky Acad. Sci. 55:32-35.
Schmidt, K.A., and B. Fried. 1996. Emergence of cer-
cariae of Echinostoma trivolvis from Helisoma trivolvis
under different conditions. J. Parasitol. 82:674-676.
Sloos, B., J. Meece, M. Romano, and P. Nollen. 1995.
The genetic relationships between Echinostoma cap-
roni, E. paraensei, and E. trivolvis as determined by
electrophoresis. J. Helminthol. 69:243-246.
Smyth, J.D., and D.W. Halton. 1983. The physiology of
trematodes. 2nd ed. Cambridge University Press, Cam-
bridge, U.K.
Theron, A. 1984. Early and late shedding patterns of
Schistosoma mansoni cercariae: ecological significance
in transmission to human and murine hosts. J. Parasitol.
70:652-655.
Theron, A., and C. Combes. 1988. Genetic analysis of
cercarial emergence rhythms of Schistosoma mansoni.
Behav. Genet. 18:201—209.
Wagenbach G.E., and A.L. Alldredge. 1974. Effect of
light on the emergence pattern of Plagiorchis micra-
canthos cercariae from Stagnicola exilis. J. Parasitol. 60:
782-785.
Trans. Ky. Acad. Sci. 58(1):29-32. 1997.
Classifying Free Bieberbach Groups
Raymond F. Tennant
Department of Mathematics, Statistics, and Computer Science,
Eastern Kentucky University, Richmond, KY 40475
ABSTRACT
Let F be a free group on n letters and G be a finite group. To each epimorphism €: F — G is associated
a free Bieberbach group F/N’ where N is the kernel of the epimorphism € and N’ is the commutator
subgroup of N. In low dimensions, the free Bieberbach groups are described. In high dimensions, the free
Bieberbach groups are described for the case when G is cyclic or dihedral. A theorem is given describing a
method for factoring certain free Bieberbach groups as a semi-direct product of a lower dimensional free
Bieberbach group and the integral group ring ZG.
INTRODUCTION
Let G be a finite group. A group S is called
a crystallographic group (on the point group
G) if S contains a finitely generated maximal
abelian torsionfree subgroup A of finite index
so that S/A=G and G acts faithfully by con-
jugation on A. The rank of A is called the di-
mension of the crystallographic group. A tor-
sionfree crystallographic group is called a Bie-
berbach group (Charlap’s definition [1986,
p.74]). The dimension of the Bieberbach
group is given by Schrier’s formula
n=|G|(mk(F)—1)+1, where F is the free
group on k generators with k=rnk(A) (Lyndon
and Schupp 1977, p.16).
It is well known that B is an n-dimensional
Bieberbach group if and only if R’/B is an n-
dimensional flat manifold. Let (X | R) be a
presentation of G with finitely many genera-
tors, that is, G=F/N where F is the free group
on X and N is the normal closure of R in F.
This may be expressed in terms of the short
exaciasequence, 1 —> Nv" h > G —> Ihlet
N’=[N,N] be the commutator subgroup of N,
N=N/N’, and F=F/N’. The sequence above
induces a short exact sequence on N and F, 0
> N->F->G>2Q 1. The action by conju-
gation of G on N gives N a ZG-module struc-
ture. N is referred to as the relation module
of F/N (Linnell 1981). F is known to be a Bie-
berbach group and will be referred to as a free
Bieberbach group (Farkas’ definition [1981)).
It is noted that of the flat manifolds associated
to these free Bieberbach groups, some are or-
ientable and some are non-orientable.
29
Two free Bieberbach groups F, and F, are
isomorphic if there exists a ZG-module iso-
morphism a and a group isomorphism B such
that the following diagram commutes:
0>5N,>F,>°GQO>1
ene = (eu dh
0>N,>°F,>G > 1
where N, and N, are the relation modules as-
sociated to F, and F, respectively.
Since N is a ZG-module with free abelian
rank equal to n, it is convenient to write N as
Z when we are viewing F as an actual crys-
tallographic group. Further, Aut(N)=GL(n,Z)
so the action of G on N induces a represen-
tation @: G > GL(n,Z). Since F=Z"XG (as
sets), then we may define multiplication on F
by (n,g)(m,h)=(n+9(g)m+f(g,h),gh), where
n,meZ" (thought of as column vectors),
gheG, and feH*(G,Z"). The 2-cocycle f:
GXG — Z* may be constructed from
i(f(g,h))=s(g)s(h)s(gh)~!, where s: G > F is a
normalized set map and i: Z" > F is the in-
clusion map. It is known (Tennant and Turmer
1992) that F is torsionfree. For the case,
GA{1}, this is equivalent to the 2-cocycle f:
GXG ~ Z being nontrivial.
For a given dimension n > 0, there is a free
Bieberbach group isomorphic to Z" where the
point group is G={1}. Note that in this case,
the compact flat manifold associated to the
free Bieberbach group Z" is the n-torus, the
orbit space R'/Z”.
30 Transactions of the Kentucky Academy of Science 58(1)
FREE BIEBERBACH GROUPS IN
LOW DIMENSIONS
Let G be a finite group. In dimension n =
1, a free Bieberbach group (other than Z with
trivial point group) must have the point group
G=Z,. The compact flat manifold associated
to this group is the circle. In dimension n =
2, a free Bieberbach group must have point
group G={1} and mk(F)=2 so F=Z?. There-
fore, the first case of a Bieberbach group B
that is not a free Bieberbach group is defined
by the sequence 0 — Z? > B > Z, > 1 where
R’/B is homeomorphic to the Klein bottle.
In dimension n = 3, a free Bieberbach
group (other than Z? with trivial point group)
must have G=Z, and mk(F)=2. A result of
Linnell (1981) states that if two epimorphisms
€,; F, ~ G and e,: F, ~ G, with
mk(F,)=rnk(F,), are Nielsen equivalent, then
their ee ed free Bieberbach groups are
isomorphic. Since any two epimorphisms €,,
€,: F(a,b) > Z, are Nielsen equivalent, con-
sider the epimorphism derived from the pre-
>. b). This pre-
sentation and the action of Z, on Z° induces a
monomorphism g: Z, > GL(3,Z). Since
det(g(a))=—1, the flat manifold associated to
this free Bieberbach group is non-orientable.
The crystallographers refer to this particular
manifold (crystal) as belonging to the mono-
clinic class (Brown et al. 1978).
In dimension n = 4, a free Bieberbach
group (other than Z* with trivial point group)
must have point group G=Z, and mk(F)=2
Since there is only one Nielsen equivalence
class of epimorphisms €: F(a,b) > Z;, there is
only one free Bieberbach group with point
group G=Z,. Since 2 does not divide |G|=3,
the induced monomorphism ¢: Z,; > GL(4,Z)
must satisfy det(g(g))=1 for all geG and so it
follows that the flat manifold R‘/F is orientable.
SOME FREE BIEBERBACH GROUPS IN
HIGH DIMENSIONS
Let k be a positive integer. If k is odd, there
is one free Bieberbach group of dimension
n=k+ | where the point group is G=Z,. The
associated flat manifold is orientable by the
same reasoning as above. Suppose k is even, G
is a point group with |G|=k, and €: F(a,b) >
G is an epimorphism. If G=Z, or G=D,, the
dihedral group with k elements and g: G >
GL(k+1,Z) is the monomorphism induced by
the action of G on Z" then there exists geG
with ord(g)=2 such that det(g(g))=—1. It fol-
lows that the associated flat manifold is non-
orientable. For the point group, G=Z,, all epi-
morphisms, €: F(a,b) > Z, are Nielsen equiv-
alent so all free Bieberbach groups with point
group G=Z, are isomorphic. On the other
hand, there exist k such that not all epimorph-
isms €: F(a,b) > D, are Nielsen equivalent, so
we may not apply Linnell’s result to conclude
that there is only one (k+1)-dimensional free
Bieberbach group with point group G=D,. Let
e: F, > G be an epimorphism where F, is free
on k generators. Suppose another generator is
added to F, and mapped to | under € to form
a new epimorphism €*: F,,,, > G. The follow-
ing theorem describes how the respective free
Bieberbach groups are related.
Theorem
Let €: F, > G and e*: F,,, > G be epi-
morphisms such that € * (eel ), Sisk and
€*(x,,,)=1, then F* is isomorphic to Fx,ZG
where F and F* are the free Bicherback
groups associated to € and €* respectively
and x, represents a semi-direct product.
Proof
Recall that F may be expressed in terms of
the short exact sequence
jo N > he coat
The action by conjugation of G on N induces
a monomorphism ¢: G > Aut(N). Let s: G >
F be a normalized set map and define a 2-co-
cycle f: GXG > N by i(f(g,h))= s(g)s (h)s(gh)~1.
By viewing F as setwise equal to NXG we may
define multiplication on F by (i,g)(m,h)=
(n+ (g)m+f(g,h), gh). A theorem of Lyndon
(1962) states that N*=N/ZG as ZG-modules.
The isomorphism a: N* —> N/ZG induces an
isomorphism A: Aut(N*) — Aut(N/ZG) and
the projection p: N/ZG —> N induces a pro-
jection P: Aut(N/ZG) > Aut(N) so that the
action of G on N* induces a monomorphism
o*: G — Aut(N*) that satisfies P(A(p*(g))).
The free Bieberbach group may be expressed
in terms of the short exact sequence
Free Bieberbach Groups—Tennant 31
() CSI Sree esar
A set map s*: G — F* can be chosen so that
the 2-cocycle f*: GXxG — N* defined by
i*(f*(g,h))=s*(g)s*(h)s*(gh)~! satisfies p(a(f*
(g,h)))=f(g,h). Define the epimorphism é : N*
> N by €(n*)=p(a(n*)). Then 0 — ZG >
N* > N —> 1 is a short exact sequence of
ZG-modules. The group F* is equal to N¥ XG
(as sets) so define the epimorphism é: ha
F by €(n*, g)=(E(n*),g). The ker(é)=ZG
and we have the following commutative dia-
gram:
0 0
dt AL
07> ZG>~ ZG 1
y L L
0 > N* 59 F*¥ 5G 1
y A L
0O2>N> Fo Gol
t L L
1 It ]
Define the normalized set map §: N > N*
by §(n)=a—!(n/0) where 0 is the additive iden-
tity in ZG. The set map §: F > F* defined by
§(n,g)=(8(),g) is then a homomorphism and
so it follows that the sequence 0 ZG > F*
— F > 1 splits and F*=Fx,ZG.
REMARK
For the case where F*=Fx.ZG, if F has di-
mension n then F* has dimension n+|G| and
F* can be viewed as a free Bieberbach group
which has a subgroup, F, that is also a free
Bieberbach group. To crystallographers, F* be-
longs to the class of superspace groups [Janner
and Janssen 1979]. Equivalently, R'/F is an n-
dimensional flat manifold sitting inside of the
(n+|g|)-dimensional flat manifold R™*'C/F*.
Corollary 1
If Epi(F,,,,G) contains only one Nielsen
equivalence class and €: F, > G and €*: F,,,
— G are epimorphisms then F*=Fx,ZG.
As an illustration, the (k+1)-dimensional
free Bieberbach group (k odd) with point
group G=Z, can be factored as the semi-di-
rect product F=Zx,Z(Z,).
There are certain point groups with the fol-
lowing property (known as the “swap” prop-
erty [Tennant and Turner 1992)).
Swap Property
Given any two epimorphisms €,,€,: F, >
G, there exists a finite sequence of epimorph-
isms e€,,€, --, e, where e, F,,, > G for i odd
and e,;: F, > G for i even, 1Six<r such that
for each of the pairs of epimorphisms, (€ ,,e,),
(ei,e,,,) for 1SisSr—1, and (e,€,), the epi-
morphism on F, is isomorphic to the epi-
morphism on F,,, restricted to some subset
(of k basis elements) of F,,,,.
Corollary 2
Let G be a finite group, and €,,€,: F, > G
be two epimorphisms. If G possesses the swap
property then F,x,ZG=F.x,ZG where F, and
F, are the free Bieberbach groups associated
to €, and é€, respectively.
Example
The class of finitely generated abelian
groups possesses the swap property (Tennant
and Turner 1992). Since point groups must be
finite, we are concerned only with the subclass
of finite abelian groups.
Corollary 3
If G is a finite abelian group and €,,¢€,: F,
— G are two epimorphisms then F x.ZG=
F.x.ZG where F, and F, are the free Bieber-
bach groups associated to €, and €, respec-
tively.
When F,x,ZG=F’x.ZG is said to be one step
stably equivalent to F,.
LITERATURE CITED
Brown, H.R., R. Bulow, J. Neubuser, H. Wondratschok,
and H. Zassenhaus. 1978. Crystallographic groups of
four-dimensional space. J. Wiley & Sons, New York, NY.
Charlap, L.S. 1986. Bieberbach groups and flat mani-
folds. Springer Verlag, New York, NY.
Farkas, D.R. 1981. Crystallographic groups and their
mathematics. Rocky Mt. J. Math. 11:511-551.
Janner, A., and T. Janssen. 1979. Superspace groups.
Physica A 99:47-—76.
Linnell, P.A. 1981. Relation modules and augmentation
ideals of finite groups. J. Pure Appl. Algebra 22:143—-
164.
32 Transactions of the Kentucky Academy of Science 58(1)
Linnell, P.A. 1985. A cancellation theorem for lattices Lyndon, R.C., and P.E. Schupp. 1977. Combinatorial
over an order. J. Lond. Math. Soc. IH. 31:450-456. group theory. Springer Verlag, New York, NY.
Lyndon, R.C. 1962. Dependence and independence in Tennant, R.F., and E.C. Turner. 1992. The swap conjec-
free groups, J. Reine Angew. Math. 210:148-173. ture. Rocky Mt. J. Math. 22:1083-1095.
Trans. Ky. Acad. Sci. 58(1):33-34. 1997.
DISTINGUISHED SCIENTIST AND
OUTSTANDING TEACHER AWARDS 1996
Distinguished Scientist Award
Dr. D. Allan Butterfield—Professor of Chemistry and
Director of Center for Membrane Sciences, University
of Kentucky, Lexington, Kentucky
D. Allan Butterfield received the degree of bachelor of
arts with highest distinction from the University of Maine
in 1968. After a 3-year high-school teaching position in
Zimbabwe (then Rhodesia) he began his doctoral work at
Duke University where he received the Ph.D. in physical
chemistry in 1974. After a l-year NIH _ post-doctoral
fellowship in 1974, also at Duke, he joined the
Department of Chemistry at the University of Kentucky
in fall 1975 as an assistant professor. In 1983 he was
promoted to the rank of professor. Since 1986 he has been
the director of the University of Kentucky’s Center of
Membrane Sciences.
Dr. Butterfield has over 140 publications in refereed
journals or monographs, an additional 15 publications
either in press or submitted for publication, and about 80
published abstracts. He has presented papers at
professional meetings throughout the United States and
in England, Canada, and Australia. He serves as referee
for some 20 journals and organizations. He has been the
principal investigator or co-principal investigator of over
20 funded grants valued at nearly $13 million.
Dr. Butterfield’s research in the field of membrane
science as it pertains to Alzheimer’s disease is most
impressive. His research allows for a_ unified
understanding of the causes of Alzheimer’s disease,
contributing to the eventual development of methods for
either prevention of the disease or minimization of its
effects.
Dr. Butterfield has taught 18 different courses at both
the undergraduate and graduate levels and has given
invited seminars at universities and colleges. He has
directed 17 master’s-level and 14 Ph.D.-level students;
currently seven graduate students are working under his
direction. He has guided 18 undergraduates in NSF
sponsored summer research; eight of these students
co-authored referreed scientific publications based on
their research. Dr. Butterfield has been involved in the
placement of about 75 students in graduate and
professional positions.
Dr. Butterfield has served on numerous university and
department committees. He is a member of several
professional organizations and honorary societies, e.g.,
American Chemical Society, Society of Sigma Xi, Phi Beta
Kappa Honorary Society, and Phi Kappa Phi Honorary
Society. He serves on the editorial board of Journal of
Membrane Science. Dr. Butterfield has also been involved
with several community organizations, including Habitat
for Humanity.
It is obvious that D. Allan Butterfield’s research
33
program and teaching are having an immediate positive
impact on intellectual growth in the Commonwealth. This
in itself is a great accomplishment. However, it is also
quite likely that his research, which is providing scientists
with a better understanding of the underlying causes of
Alzheimer's disease, will lead directly to improvement in
the quality of life in the Commonwealth and throughout
the world.
Outstanding College/University
Science Teacher Award
Dr. Karan Kaul—Professor, Biology Principal Investigator
and Program Area Coordinator, KSUCRS, Kentucky
State University, Frankfort, Kentucky
Karan Kaul earned the degree of bachelor of science in
botany with honors in 1966 and the degree of master of
science in botany in 1968 from the University of Delhi.
He received a Ph.D. in biology from the University of
Kentucky (UK) in 1972. After receiving his doctorate, he
remained at UK as a postdoctoral fellow in the Depart-
ment of Biochemistry until 1977. Dr. Kaul spent 3 years
outside Kentucky in research-associate positions before
beginning his career at Kentucky State University (KSU)
in 1980. Early in his career at KSU most of his time was
spent in botanical research. As time passed, however, he
began to devote more and more time to teaching. He pro-
gressed from assistant professor to associate professor in
1990 and earned the rank of full professor in 1993. Al-
though Dr. Kaul is nationally known as a prominent re-
searcher in botanical and biological studies (with 14 pub-
lications since 1991), he has chosen to spend over half of
his time in teaching cell biology, vertebrate embryology,
histology, principles of biology, botany, and several other
biology courses.
Dr. Kaul spends many hours outside the classroom in
mentoring students, in coordinating pre-health clinics, and
in assisting students in national examinations, sometimes
at his own expense. He cares about the education of his
students, encouraging their learning through time spent
inside and outside the classroom; for this, he has the ut-
most respect of students and faculty members alike.
Dr. Kaul was listed in Who's Who Among America’s
Teachers this year. The Kentucky Academy of Science is
proud to present to Dr. Karan Kaul the 1996 Outstanding
University Science Teacher Award.
Outstanding Secondary School
Science Teacher Awards
Douglas C. Jenkins—Instructor, Warren Central High
School, Bowling Green, Kentucky
34 Transactions of the Kentucky Academy of Science 58(1)
Douglas C. Jenkins is co-recipient of the 1996 Kentucky
Academy of Science Outstanding Secondary School Sci-
ence Teacher Award. Mr. Jenkins received the degree of
bachelor of science in physics and mathematics from
Western Kentucky University (WKU) in 1968. For the
next few years, he worked as project engineer for the U.S.
Army Night Vision Laboratory in Fort Belvoir, Virginia.
During this time, in 1972, he received the Department of
Army Civilian Outstanding Performance Award. In 1973
he returned to Kentucky where he worked while continu-
ing his education. He began his career at Warren Central
High School in 1976. In 1978, he completed requirements
for the degree of master of arts in education from WKU;
in 1986 he obtained his Rank I certification.
During his spare time he became a part-time instructor
in the Department of Physics and Astronomy at WKU
and, more recently, a teacher for the Verbally and Math-
ematically Precocious Youth Program. At Warren Central
he has served as boy’s golf and tennis coach. He is an
active member in many professional organizations and has
served on multiple commissions and task forces. Mr. Jen-
kins was one of the original planners of the Barren River
Imagination Museum of Science, a hands-on science mu-
seum that serves surrounding school systems. Through his
leadership, most Warren Central High School physics stu-
dents were involved in preparing and constructing exhibits
in 1993. He continues to involve his students in maintain-
ing exhibits and participating in museum programs.
“Doctor J,” as Mr. Jenkins is known by students and
faculty, has increased interest in science at Warren Central
through innovative teaching practices. His teaching incor-
porates innovative, motivational practices that create op-
portunities for development of student interest and suc-
cess. His instruction focuses on learning outcomes. His
classes are active and emphasize cooperative and group
learning using real-life experiences.
Professional and cooperative, Mr. Jenkins certainly con-
tributes in many ways to the teaching profession. He has
long been recognized as an exceptional teacher. In 1983
he received the Presidential Award of Excellence in Sci-
ence Teaching; in 1986, the Warren County Teacher of
the Year Award; and in 1991, the Sallie Mae Teacher Trib-
ute Award.
The Kentucky Academy of Science is proud to present
to Douglas C. Jenkins the Outstanding Secondary Science
Teacher Award. He is an exemplary representative of the
many fine teachers of our Commonwealth.
Janet Yates Ward—Instructor, Reidland High School, Pad-
ucah, Kentucky
Janet Yates Ward is co-recipient of the 1996 Kentucky
Academy of Science Outstanding Secondary School Sci-
ence Teacher Award. Ms. Ward received from Murray
State University the degree of bachelor of science in 1986,
the degree of master of science in biology in 1988, and
Rank I certification in 1996. She graduated summa cum
laude from her class. In fall 1989 she began her career at
Reidland High School. Since that time, she has earned an
excellent reputation from students and faculty members
alike.
Ms. Ward has made a positive impact on the young
people at Reidland. Demanding of her students and cre-
ative in her teaching approach, she has a genuine love of
science, a love reflected in her attitude. She is instrumen-
tal in spurring her students to excellence; many have re-
ceived awards for their work. As sponsor of the biology
competition team, she was instrumental in her students’
accomplishments as exhibited by high finishes in compe-
tition. One of her students was awarded a full-tuition
scholarship to the University of Louisville, and another
was invited to present a project at the Kentucky Junior
Science and Humanities Symposium sponsored by the
University of Louisville. Reidland Science Club, which is
sponsored by Ms. Ward, won bronze and silver medals at
the State Science Olympiad.
Ms. Ward has been named in Who’s Who Among Amer-
ican Teachers and has been active on the McCracken
County Education Association and National Education
Association. She is a member of the Alpha Chi National
Honor Society and Kappa Delta Pi Honorary Education
Society. In 1994 she was awarded the Golden Apple
Achiever Award sponsored by Ashland Inc. Her enthusi-
asm, her skill at innovation, and her genuine concern for
biology education are exemplary. One of her students
wrote, “Through Mrs. Ward’s biology class, I began to see
that science was a great collection of discoveries. I feel I
owe to her my utmost gratitude and appreciation for her
instruction and ability to convey her love of science, which
seems to be contagious!” We hope that many more stu-
dents will catch the love of science from Janet Yates Ward.
Trans. Ky. Acad. Sci. 58(1):35-36. 1997.
NOTES
Human Myiasis in Kentucky Caused by Cuterebra
sp. (Diptera: Oestridae).—Species of Cuterebra (Dip-
tera: Oestridae) parasitize rodents and rabbits. The rabbit
Cuterebra can be found also in young dogs and cats (1,
2). Little is known about the host range of this genus in
Kentucky. The few documented infestation rates for nat-
ural hosts have been quite variable. Over 70% of the
trapped specimens of Peromyscus leucopus, the white-
footed mouse, on Kentucky Lake islands in western Ken-
tucky and Tennessee were infested with Cuterebra fonti-
nella fontinella (3). In another study, 36% of P. leucopus
individuals captured in eastern Tennessee harbored the
same oestrid (4). Only three adult specimens of this genus,
all identified as C. fontinella, are in the entomology col-
lection of the University of Kentucky. No biological infor-
mation is associated with the material.
The case of myiasis reported herein involved a 19-year-
old male. The larva was submitted to the Cooperative Ex-
tension Service for identification following its removal
from a lesion on the ankle. I determined it to be a late
second or early third instar Cuterebra sp. The larva was
sent to Dr. R. D. Hall, University of Missouri, who con-
firmed the identification (pers. comm.). Accurate identi-
fication of immatures of this group is best accomplished
with mature third instars so that cuticular spines and cau-
dal spiracles can be examined (5). Most documented cases
of human oestrid myiasis have involved the rabbit-infest-
ing species (6) C. buccata and C. abdominalis. These par-
asitize eastern cottontail rabbits and are the two members
of this group ranging into Kentucky (R.D. Hall, pers.
comm.).
The man answered a set of questions for me that de-
scribed the general circumstances of the incident at which
the infestation might have started and its course. This in-
formation is summarized as follows.
The most obvious opportunities for exposure appear to
have occurred on four occasions between 5 Sep and 10
Sep 1995 while the man was squirrel hunting in Greenup
County, Kentucky. He wore long pants, long socks, and
high-topped canvas tennis shoes on all trips. While hunt-
ing, he sat at the base of trees in wooded areas for | to 2
hours at a time. In most cases he cleared away fallen
leaves before sitting on the ground, but occasionally he
sat on a fallen tree.
The first indication of a problem was noticed on 14 Sep
1995. He experienced “ warm, itching sensation, similar
to the feel of a mosquito bite” and discovered a small red
bump on skin of the medial surface of an ankle. Two more
bumps appeared 2 days later. At that point, the irritation
was described as a “gnawing sensation as if the flesh were
being torn.” Ten days after the initial bump appeared,
there were five bumps spaced at about 2-cm intervals
along the ankle. At that time, the largest bump was about
2 cm in diameter and protruded about 1.5 cm above the
surface of the skin.
Anxious about the increasing size and numbers of le-
35
sions, the man visited a physician on 25 Sep 1995. By that
time, the largest bump had “come to a head like a boil.”
The wound was diagnosed as a spider bite; topical appli-
cation of a hydrocortisone cream was recommended for
the area. After returning home, the man squeezed a single
bot larva from the boil-like spot. The area itched for about
a week after the larva was removed but healed normally.
Following the anxiety associated with the removal of a
live maggot from the boil, the young man in this case was
relieved to learn the identification of the insect. He re-
membered that he had occasionally encountered mature
bot larvae in subdermal cysts or warbles when skinning
squirrels taken on previous hunting trips.
Eggs of Cuterebra spp. are laid near burrows or areas
frequented by the host (7). Mature eggs can hatch rapidly
in response to stimulation of the host, such as an increase
in air temperature and CO, concentration (8). These mag-
gots can complete a portion of their development in hu-
mans (9). Human infestations usually occur when an in-
dividual spends some time near a host lair and comes in
contact with eggs.
Some human subjects of oestrid myiasis are aware of a
“stinging” feeling. This probably occurs as the larvae use
their mouth hooks to enter the skin. Penner (10) observed
the activity of three Cuterebra larvae intentionally placed
on his skin. The stinging sensation developed within 9 to
17 minutes and as the larvae entered the skin. Complete
penetration took at least 18 minutes. Edema and redden-
ing of the area developed and persisted for 2 weeks even
though the larvae were removed immediately after pen-
etration.
The site of infestation, the ankle, is unusual in this Ken-
tucky case. Rice and Douglas (11) reviewed five previously
reported cases and documented two new ones. Four were
on the face in or around the nose or eyes. The remainder
were on the neck or torso. In two of the cases, the indi-
vidual was outdoors at the time and felt the initial pene-
tration. Baird, Podgore, and Sabrosky (6) summarized the
30 cases of human infestation by Cuterebra known to
1982; Baird, Baird, and Sabrosky (5) provided an update
7 years later. All but four infestations were on the torso
or head. The infestation site for three was unknown and
one was on the scrotum.
In this case from Kentucky, there was a delay of 4 days
between the last hunting trip and skin penetration by the
maggot. These trips were the most obvious times that this
individual could have come in contact with Cuterebra ovi-
position sites. However, a few cases indicate no history or
activity that places the person near oviposition sites. In
the Kentucky case, an egg may have been picked up and
remained on a shoe until development was complete or it
was stimulated to hatch.
LITERATURE CITED. (1) Hall, M., and R. Wall. 1995.
Myiasis of humans and domestic animals. Adv. Parasitol. (20C)
35:257-334. (2) Muller, G.H., R.W. Kirk, and D.W. Scott.
1989. Small animal dermatology. 4th ed. W.B. Saunders, Phil-
36 Transactions of the Kentucky Academy of Science 58(1)
adelphia, PA. (3) Kollars, T.M., Jr. 1995. Factors affecting
the distribution of bot flies (Diptera: Cuterebridae) on is-
lands in Lake Barkley in Kentucky and Tennessee. J. En-
tomol. Sci. 30:513-518. (4) Dunaway, P.B., J.A. Payne,
L.L. Lewis, and J.O. Storey. 1967. Incidence and effects
of Cuterebra in Peromyscus. J. Mammol. 48:38-51. (5)
Baird, J.K., C.R. Baird, and C.W. Sabrosky. 1989. North
American cuterebrid myiasis. J. Am. Acad. Dermatol. 21:
763-772. (6) Baird, C.R., J.K. Podgore, and C.W. Sabros-
ky. 1982. Cuterebra myiasis in humans: six new case re-
ports from the United States with a summary of known
cases (Diptera: Cuterebridae). J. Med. Entomol. 3:263—
267. (7) Catts, E.P. 1982. Biology of the New World bot
flies: Cuterebridae. Ann. Rev. Entomol. 27:313-338. (8)
Catts, E.P. 1967. Biology of a California rodent bot fly
Cuterebra latifrons Coquillett. J. Med. Entomol. 4:87—
101. (9) James, M.T. 1947. The flies that cause myiasis in
man. USDA Misc. Publ. 631. (10) Penner, L.R. 1958.
Concerning a rabbit cuterebrid, the larvae of which may
penetrate the human skin (Diptera, Cuterebridae). J. Kan-
sas Entomol. Soc. 67-71. (11) Rice, P.L., and G.W. Doug-
las. 1972. Myiasis in man caused by Cuterebra (Diptera:
Cuterebridae). Ann. Entomol. Soc. Am. 65:514-516.—
Lee H. Townsend, Department of Entomology, Univer-
sity of Kentucky, Lexington, KY 40546-0091.
Trans. Ky. Acad. Sci. 58(1):37-54. 1997.
Abstracts of Some Papers Presented at the
1996 Annual Meeting of the
Kentucky Academy of Science
AGRICULTURAL SCIENCES
Geographic differentiation and allozyme variation in
pawpaw (Asimina triloba, Annonaceae). HONGWEN
HUANG#* and DESMOND R. LAYNE, Land-Grant Pro-
gram, Atwood Research Facility, Kentucky State Univer-
sity, Frankfort, KY 40601.
The pawpaw (Asimina triloba) is the largest tree fruit
native to the United States and the only temperate mem-
ber of the tropical custard apple family. In 1994, Kentucky
State University was established as the USDA-ARS-Na-
tional Clonal Germplasm Repository for Asimina spp.
Seedling trees from 400 pawpaw accessions representing
70 distinct geographic regions from 17 states are currently
being grown at our research farm. In a preliminary study,
32 pawpaw cultivars or advanced selections were assayed
in 30 enzyme systems, using an isoelectric focusing poly-
acrylamide slab gel system of pH 4-9. Twelve enzymes
produced high resolution without tissue specificity and
were further used for evaluation of allozyme diversity of
geographic populations. We discuss the degree of genetic
diversity within populations and differentiation between
populations as evaluated by the expected heterozygosity
(He), the proportion of polymorphic loci (P), the average
number of alleles per locus (A), Chi-squared analysis of
allele frequency heterogeneity, Nei’s standard genetic dis-
tance (D), and identity (1). Dendrograms were generated
by cluster analysis using the unweighted pair group meth-
od to demonstrate the relationships of geographic popu-
lations in the 17 states evaluated. The strategy for germ-
plasm conservation and cultivar development through
breeding is discussed. Issues related to establishment of a
“core” collection are addressed.
Influence of shade and root-zone modification on early
growth and development of pawpaw (Asimina triloba, An-
nonaceae) seedlings grown in the greenhouse. DES-
MOND R. LAYNE, Land-Grant Program, Atwood Re-
search Facility, Kentucky State University, Frankfort, KY
40601.
This experiment was designed to determine the optimal
light level for growing pawpaw (Asimina triloba) seedlings
in the greenhouse. In addition, we wanted to determine
if modifying the root-zone would impact seedling growth
and development. We examined the individual and com-
bined effects of shade and root-zone modification on early
growth and development of seedlings. Experimental treat-
ments were imposed from sowing until the plants were
destructively harvested. The experimental design was a
split-plot where blocking was done by position in the
greenhouse. The main plot of the experiment was shade.
This was accomplished by growing seedlings under a
wooden frame covered with shade cloth to reduce inci-
37
dent light intensity received by the plant by 30, 55, 80, or
95%. The control treatment was 0% shade or ambient
greenhouse light level. The split plot was root-zone mod-
ification. Half of all growing containers were untreated
(control) while the other half were painted with Spin-
Out™, a commercially available product that stimulates
development of a finer root system. There were 40 rep-
licate seedlings per experimental treatment combination
per block. Seedling shoot length and unfolded leaf num-
ber was recorded 2/week from seedling emergence until
destructive harvest. At harvest, leaf samples were taken
for chlorophyll determination. Whole plant leaf area was
also determined. Leaves, stems, and tap and lateral roots
were separated and dried to determine biomass parti-
tioned to the respective organs. Based on the results of
this study, optimal greenhouse growing conditions for
pawpaw seedlings have now been refined and are dis-
cussed.
Pawpaw (Asimina triloba, Annonaceae) morphological
development during seed germination and seedling emer-
gence. C.L.H. FINNESETH# and DESMOND R.
LAYNE, Land-Grant Program, Atwood Research Facility,
Kentucky State University, 40601; R.L. Geneve, Depart-
ment of Horticulture and Landscape Architecture, Uni-
versity of Kentucky, Lexington, KY 40546.
Stratified pawpaw (Asimina triloba) seeds were germi-
nated in vermiculite at 25°C to study embryo develop-
ment. Seedlings were destructively harvested every 3 days
prior to radicle protrusion (day 12) and at 5-day intervals
thereafter. At each harvest date, 10 seedlings were ran-
domly chosen for length measurement (mm) and fresh
and dry weight (mg) determinations. Initial length of the
underdeveloped embryo was less than 2 mm, but by day
70 seedling length increased to more than 350 mm.
Twelve days after sowing, simultaneous development of
the radicle and cotyledons produced lengths of 3.4 and
3.0 mm, respectively. Neither hypocotyl nor epicotyl was
visible at this time. At radicle protrusion, radicle, cotyle-
don, and hypocotyl lengths were 4.4, 4.0, and 3.2 mm,
respectively. Endosperm was the greatest proportion of
dry weight (99.1%); radicle, cotyledons, and hypocotyl
represented the remainder. The first aboveground indi-
cation of germination was hypocotyl hook emergence,
which occurred at 27 days. At this time, average seedling
measurements (mm length/% dry weight) were 41.7/17.0,
16.7/5.7, and 11.9/1.2 for the radicle, hypocotyl, and cot-
yledons, respectively. Endosperm dry weight had de-
creased to 76.1% by this time. Cotyledons reached maxi-
mum length at 40 days. At day 45, the development of a
discernible epicotyl occurred and the percentage dry
weights for the radicle, hypocotyl, cotyledons, epicotyl,
and endosperm were 44.4, 12.7, 3.5, 0.4, and 39.0, re-
38 Transactions of the Kentucky Academy of Science 58(1)
spectively. At day 50, the seedcoat containing the cotyle-
dons and residual endosperm abscised; average radicle,
hypocotyl, and epicotyl measurements were 182.0/93.8,
16.0/6.0, and 7.3/0.2, respectively.
Using flood control reservoirs for paddlefish (Polyodon
spathula) production. R. ONDERS,* S. D. MIMS, and
C. WANG, Community Research Service, Kentucky State
University, Frankfort, KY 40601.
Kentucky has a large number of flood-control reservoirs
that may be used for paddlefish (Polyodon spathula; Po-
lyodontidae) production. Juvenile paddlefish stocked into
these reservoirs will feed primarily on zooplankton natu-
rally present in the water. Paddlefish grow rapidly up to
4 kg per year with no feed cost and little management.
Paddlefish can be harvested within 2 years for its quality
white meat, which is boneless and firm in texture. Poten-
tially this production system also will permit paddlefish to
reach sexual maturity and the female fish can be harvested
for its roe, which is processed into high value caviar. Roe
production will be greatly increased by implementing all-
female production technology currently being developed
at the Aquaculture Research Center, Kentucky State Uni-
versity. Both the meat and roe of paddlefish will provide
an alternative source of income for the landowners, com-
mercial fisherman, and food processors in Kentucky.
BOTANY & MICROBIOLOGY
Cryopreservation of seeds of endangered Ohio wetland
plants. LARRY A. GIESMANN,* Department of Biolog-
ical Sciences, Northern Kentucky University, Highland
Heights, KY 41099; VALERIE C. PENCE and NYREE
CONARD, Center for Research of Endangered Wildlife,
Cincinnati Zoo and Botanical Garden, 3400 Vine Street,
Cincinnati, OH 45220.
Cryopreservation of seeds in liquid nitrogen (LN,) has
been shown to be an effective way of preserving germ-
plasm of endangered plant species. With funding from the
Robert H. Reakirt Foundation and the L and L Nippert
Foundation, the Plant Division of the CREW facility at
the Cincinnati Zoo and Botanical Garden has been ex-
ploring the feasibility of cryopreserving seeds of endan-
gered Ohio wetland plants. Most wetland species have or-
thodox seeds and should be candidates for cryopreserva-
tion. Seeds of 14 species from nine families of flowering
plants were collected from several Ohio wetland sites and
tested in the CREW labs. All species are listed by the
Ohio Department of Natural Resources as endangered,
threatened, or potentially threatened in the state. In our
study, four treatments were used for each species with at
least 20 seeds in each treatment. In all treatments, seeds
were desiccated over silica gel for 7 d. Two groups of
seeds were then exposed to LN, for 1 hr and rehydrated
for 1 d. One group was planted in pots and placed in the
greenhouse immediately; seeds in the other group were
stratified at 5°C for 3 mo before planting. Seeds in the
remaining two treatments were not exposed to LN, and
served as controls. The seeds of eight species germinated
at rates ranging from 3% to 74%. Stratification increased
the germination rate in most species. Work continues on
16 additional wetland species collected in the past year.
Fine structure of Mycena leaiana (Tricholomataceae)
basidiospores. DONALD G. RUCH* and MARK C.
ROBBINS, Department of Biology, Ball State University,
Muncie, IN 47306.
The basidiospore wall of Mycena leaiana is composed
of a thin, single, electron transparent layer, which exhibits
some variations in thickness. No distinctive surface orna-
mentation or germ pore is present. The protoplasm is sur-
rounded by a typical membrane, which lacks distinct in-
vaginations. Centrally located nonmembrane-bound lipid
droplets comprise the bulk of the protoplasm. Spores are
uninucleate. Mitochondria with a few well-delineated
plate-like cristae are present. Endoplasmic reticulum is
scant. Ribosomes occur regularly attached to the ER and
outer mitochondrial membrane, as well as being densely
packed throughout the cytoplasm. Various sized single
membrane-bound vacuoles containing an electron dense
material are present. Microbody-like organelles are readily
observed. These microbodies are probably glyoxysomes,
since malate synthase assays are positive indicating the
presence of the glyoxylate cycle in the spore.
Photomorphogenic control of flower scape elongation
in dandelion (Taraxacum officinale; Asteraceae). MICH-
ELLE JO LEATHERS and DAVID LOWELL ROBIN-
SON,* Biology Department, Bellarmine College, Louis-
ville, KY 40205.
Dandelion (Taraxacum officinale) is a major weed of
turfgrass. One reason for this weediness is its ability to
produce a large number of seeds disseminated with the
assistance of a parachute-like pappus. The flower stalk
(scape) aids in this seed dispersal by carrying the seedhead
far above the canopy. Elongation of the dandelion scape,
therefore, is an important ecophysiological function.
Three experimental approaches were taken to study the
effect of light on dandelion scape growth. First, we ex-
amined scape elongation at ca. 70 different turfgrass sites.
Three measurements were taken at each location: the
heights of the dandelion flower and seedhead scapes, and
the height of the turfgrass surrounding the dandelion
plants. We found a significant, positive correlation be-
tween the height of the turf and that of the dandelion
scapes. We hypothesized that the turf canopy altered the
light microenvironment in such a way that it altered scape
growth. Our second experiment was done with dandelion
scape explants treated with various wavelengths of light.
Light inhibited scape growth, with flower scapes being
more sensitive than the seedhead scapes. In our third set
of experiments we examined whole-plant scape elongation
at different wavelengths. Far-red light treatment resulted
in longer flower scapes than did treatment with red, blue,
or clear filters, indicating a role for phytochrome. Seed-
head scapes were less sensitive to light. It appears, there-
Abstracts, 1996 Annual Meeting 39
fore, that dandelion scape elongation is influenced both
by light microenvironment and stage of development.
Taxonomic status of the varieties of Seneca snakeroot,
Polygala senega (Polygalaceae). AMY E. TRAUTH* and
ROBERT F. C. NACZI, Department of Biological Sci-
ences, Northern Kentucky University, Highland Heights,
KY 41099.
Seneca snakeroot, Polygala senega (Polygalaceae), is an
uncommon plant ranging throughout most of eastern
North America. Two varieties have been described for this
species: variety senega and variety latifolia. The taxonomic
merit of these varieties is controversial. In general, both
the reproductive and vegetative features of variety latifolia
are much larger than those of variety senega. The goals of
this research are (1) to more accurately define taxonomic
differences between the varieties and (2) to test the hy-
pothesis that the varieties senega and latifolia are separate
species with morphologic differences not merely induced
by ecology. Most of my work on the project has been in
the laboratory and field. In the laboratory, univariate and
multivariate statistical analysis of 13 vegetative and seven
reproductive characters has been completed on 165 spec-
imens from 11 herbaria. Among these morphologic char-
acters are length of the shoot and inflorescence, length/
width ratio of the leaves, length of the wing and keel, and
size of the seed. Field research involved the gathering of
specimens of both varieties and habitat data, including soil
samples. Preliminary results suggest significant differences
between the measured morphologic characters. These re-
sults suggest that the varieties of P. senega are better treat-
ed as separate species.
GEOGRAPHY
Effects of weather systems on migraine headaches: a
pilot study. L. MICHAEL TRAPASSO,* Department of
Geography and Geology, Western Kentucky University,
Bowling Green, KY 42101; KENNETH EMBRY, Embry
Clinic, 1733 Campus Plaza Court, Bowling Green, KY
42101.
Over a period of 9 months in 1992, 15 migraine head-
ache sufferers in the Bowling Green, Kentucky, area were
surveyed to document the onset, intensity, and duration
of their migraine headache attacks. Eighty-two cases were
analyzed with respect to the meteorological conditions of
temperature, relative humidity, precipitation, and baro-
metric pressure in effect during the migraine episodes.
Graphic analysis consisted of superimposing the migraine
attack, represented as a coded line drawn through the
time period of the attack, upon the meteorological re-
cording instrument charts. This analysis revealed promis-
ing relationships between the simultaneous onset of head-
aches from several subjects with a drop in barometric
pressure, a rise in temperature, and a rise in relative hu-
midity. Statistical analysis, however, failed to find these
relationships to be significant. In general it was found that
the absence of precipitation and the day of the week main-
tained the best relationships with the onset of migraine
headaches.
HEALTH SCIENCES
Differences in health and diet status: African-American
vs. Caucasian rural southern elderly. MARTHA MAR-
LETTE,* SUSAN TEMPLETON, and C. J. LEE, Hu-
man Nutrition Research, Kentucky State University,
Frankfort, KY 40601.
We surveyed 1554 rural elderly in nine southern states.
The sample was 84.2% female, 15.8% male, 37.7% Afri-
can-American, and 61.7% Caucasian; average age was
72.7. Occurrences of selected diet-related diseases were
examined using ANOVA (SPSS). Significant (p < .05) dif-
ferences between the African-American and Caucasian
groups were found in the prevalence of diabetes (22.8%
vs. 12.9%), hypertension (53.0% vs. 44.1%), osteoporosis
(1.8% vs. 10.2%), and atherosclerosis (2.5% vs. 6.1%) and
cancer (1.6% vs. 4.7%). Two non-consecutive 24-hour
food intakes were analyzed (Nutritionist IV) for average
daily nutrient intakes. A Dietary Status Index (DSI), in-
cluding Dietary Adequacy Status (DAS) and Dietary Mod-
eration Status (DMS) factors, was computed; all dietary
differences reported were significant at p < .05. African-
American participants had a significantly lower DSI than
Caucasian participants—a mean of 47.9 vs. 52.4. Though
no significant difference for DAS was found, African-
Americans did have lower intakes for vitamin E (111% vs.
136% RDA), calcium (65% vs. 73% RDA), and magne-
sium (68% vs. 82% RDA). African-Americans had a sig-
nificantly lower mean DMS (49.9 vs. 57.4); though their
sodium intake (2100 vs. 2205 mg) was lower, African-
Americans had significantly higher cholesterol intakes
(327 vs. 228 mg), and more calories from fat (33.1% vs.
32.0%) and saturated fat (12.6% vs. 12.0%). These find-
ings demonstrate the need for nutrition education for Af-
rican-Americans, especially programs focusing on the re-
lationship between fat intake and health problems.
Health status and dietary knowledge and intakes of
southern rural elderly. SUSAN TEMPLETON,* MAR-
THA MARLETTE, and C. J. LEE, Human Nutrition Re-
search, Kentucky State University, Frankfort, KY 40601.
Rural elders (n = 1554) in nine southern states report-
ed arthritis (63%), hypertension (47%), heart (26%), gas-
trointestinal (18%), diabetes (17%), respiratory (13%),
kidney (10%), osteoporosis (7%), atherosclerosis (5%),
anemia (5%), and cancer (4%) problems. Smoking status,
weight status, and chronic health problems were com-
bined into health scores, range 9-100, mean 77.2; this
correlated highly (r = .3832, p < .0001) with self-reported
health status (“poor” to “excellent”). Surveys assessed re-
spondents’ dietary knowledge; two non-consecutive
24-hour food intakes were analyzed for nutrient intakes
(Nutritionist IV) and food group servings. ANOVA (SPSS)
was used to determine significant (p < .05) differences
among participants with low (LHS), medium (MHS), and
high (HHS) health scores. The HHS group had signifi-
40 . Transactions of the Kentucky Academy of Science 58(1)
cantly greater fiber knowledge (72.6% of responses cor-
rect vs. 67.9%, LHS and 66.7%, MHS) and dietary fiber
intake (15 grams daily vs. 13, LHS and MHS). The HHS
group also had a significantly higher expectation about
fruit servings required: 2.3 daily vs. 2.1 for both LHS and
MHS. The HHS group consumed significantly more fruit
(1.3 servings daily) than the MHS group (1.0) but not the
LHS group (1.1); both MHS and HHS groups consumed
significantly more grain products (3.9 and 4.1 servings dai-
ly, respectively) than the LHS group (3.6). Fat and cho-
lesterol knowledge, fat calorie intake, cholesterol intake,
and sodium intake did not differ significantly among the
LHS, MHS, and HHS groups. Participants with the best
health appear to be those who recognize good fiber
sources and include fiber-rich items in their diet.
MATHEMATICS
Polynomial conservation laws of the generalized Em-
den-Fowler equation. B.D. VUJANOVIC, Faculty of
Technical Sciences, University of Novi Sad, 21121 Novi
Sad, Serbia; A.M. STRAUSS, Department of Mechanical
Engineering, Vanderbilt University, Box 1612 Station B,
Nashville, TN 37235; S. E. JONES, College of Engineer-
ing, The University of Alabama, Tuscaloosa, AL 35487-
0276; PETER P. GILLIS,* Department of Chemical and
Materials Engineering, University of Kentucky, Lexington,
KY 40506.
We consider the polynomial conservation laws of fourth
degrees with respect to x of the generalized Emden-Fow-
ler equation x + (b/t) x + yx*t! = 0. We demonstrate that
the existence of conservation laws depends upon the so-
lution of a system of partial differential equations, usually
termed the generalized Killing’s equations. The general
form of the fourth degree conservation laws of the Em-
den-Fowler equation is given; some concrete examples are
discussed.
Surface geometry of the Morehead radio telescope
and the optimization of operating parameters. BRIAN M.
LEWIS,* CHRISTOPHER L. SMITH,* and BENJA-
MIN K. MALPHRUS, Morehead Astrophysical Obser-
vatory, Morehead, KY 40351.
The operating parameters of the Morehead Radio Tel-
escope (MRT) at Morehead State University (Morehead,
KY) affecting its performance are the half power band-
width, the minimum detectable flux density, and the an-
tenna gain. The MRT is in the process of being resurfaced
to improve its radiation pattern and performance charac-
teristics by optimizing the reflector geometry. The surface
geometry has a direct effect on the radiation pattern,
which determines the operating parameters. To determine
these parameters, the precise surface area of the antenna
is needed. The purpose of the current research is to de-
termine the surface area and to compare the theoretical
parameters before and after the telescope is resurfaced.
The empirical and theoretical methods for approximation
of the original surface and the new surface are discussed.
The empirical method for the old surface is finite in na-
ture and consists of the approximation and summation of
planar areas. The empirical methods for finding the new
surface is based upon the fact that a curved surface is an
altered flat surface. Both theoretical methods involve in-
tegral approximations and manipulations. Values from the
empirical and theoretical surface areas of the old surfaces
are compared to determine the improvements in perfor-
mance characteristics.
MOLECULAR & CELL BIOLOGY
Alkylation of DNA by strain-induced electrophilic cyclic
rings. ELIZABETH M. THOMAS* and ARTHUR CAM-
MERS-GOODWIN, Department of Chemistry, Univer-
sity of Kentucky, Lexington, KY 40502.
Strain-induced electrophilic cyclic rings should alkylate
the minor groove of the DNA fragments containing AAAG
sites on the 5’ to 3’ strand. Alkylation of DNA would then
inhibit DNA translation, thus proving to be a possible
anti-tumor/anti-cancer drug agent. 3-phenylcyclobut-2-en-
l-one and 2-propanone-3-(3-phenylcyclobuten-1-diyl)
should exothermically alkylate DNA, releasing 2-3 kcal of
ring strain in a first chemical step and potentially releasing
approximately 27 kcal in a second irreversible step. The
research includes the syntheses of these two compounds
using zinc in an inert surrounding. DNA (15-mer) incu-
bated with 3-phenylcyclobut-2-en-1-one showed no signs
of alkylation by electrophoresis. This result needs to be
checked by nmr.
Alzheimer’s PS-1 mutation alters calcium homeostasis
and sensitizes neurons to death induced by A and trophic
factor withdrawal. BILL BARRETT,* QING GUO, and
MARK MATTSON, Department of Anatomy and Neu-
robiology, University of Kentucky, Lexington, KY 40502.
Mutations in the presenilin-1 (PS-1) gene on chromo-
some 14 are linked to autosomal dominant, early onset,
Alzheimer’s disease. The amino acid sequence of PS-1
predicts an integral membrane protein with a structure
suggesting functions as a receptor or ion channel, or in
protein trafficking. PS-1 is localized to the endoplasmic
reticulum (ER), and it is shown here that expression of a
PS-1 mutation (L286V) in cultured neuronal cells exag-
gerates Ca** responses to agonists (carbachol and brady-
kinin) that induce Ca** release of the ER. Cells expressing
L286V exhibit increased vulnerability to amyloid B-pep-
tide toxicity and apoptosis induced by trophic factor with-
drawal. The endangering action of L286V involves oxida-
tive stress and disruption of calcium homeostasis, and an-
tioxidants and calcium channel blockers counteract the ad-
verse consequences of this PS-1 mutation. By perturbing
Ca?* homeostasis, PS-1 mutations may sensitize neurons
to age-related accumulation of AB and reduced trophic
support.
Assessment of metal impact in the Robinson Forest
creek system using metalliothionein. KEVIN COMP-
TON,* and CHRISTER HOGSTRAND, Department of
Biology, University of Kentucky, Lexington, Ky 40502.
Abstracts, 1996 Annual Meeting 41
Metallothionein (MT) was chosen and analyzed as a
prospect for being a good bioindicator in the Robinson
Forest creek system (RFCS), Breathitt County, KY. MT
is a low-molecular-weight protein known to be induced by
heavy metals such as Cd, Cu, Hg, and Zn, and to also
positively correlate with hepatic concentrations of those
metals. In the RFCS, recent mining has posed the prob-
lem of possible stream contamination. It was, therefore,
important to obtain raw data on stream conditions to assist
in future monitoring efforts. All water metal concentra-
tions at the six sites sampled were found to be below EPA
published guidelines. Four species of fishes were sampled
but one prevalent species, the stoneroller (SR) (Campo-
stoma anomalum), was found to have the least amount of
fluctuation in MT concentrations between members at
each site. A significant variance (p < 0.05) was also found
between two sites when SR MT concentrations were com-
pared. The relative prevalence of the species combined
with the lack in fluctuation of their MT concentrations
between site members lead to the conclusion that they
could be a good species from which MT could be sampled
for metal exposure monitoring. Results on possible cor-
relations between hepatic metal and MT concentrations
of the species are forthcoming. If the concentrations are
shown to positively correlate then it could be concluded
that SR’s would be a good species from which to sample
MT to use as a bioindicator to show increasing metal pol-
lution in the RFCS. The presence of correlations could
also possibly indicate a metal of most concern.
Calcium homeostasis neurotoxicity hypothesis: possible
evidence through L-type calcium channel density In hip-
pocampal subregions. ALEXANDER COON,* ROSE-
MARIE BOOZE, and DAVID WALLACE, Department
of Anatomy and Neurobiology, University of Kentucky,
Lexington, KY 40536.
Alzheimer’s Disease (AD) has been diagnosed by mea-
suring neurofibrillary tangle and senile plaque densities.
In addition to these pathological markers, considerable
neuronal cell death occurs in the brains of patients with
AD. Considerable research has focused on determining
the mechanism by which selective neuronal cell death oc-
curs. An altered calcium homeostasis is believed to be an
integral part of this cell death mechanism. L-type calcium
channels are primarily responsible for the movement of
Ca?* ions across the neuronal membrane. However, the
relationship between channel density and the pathological
diagnosis of AD has yet to be fully elucidated. To further
examine this relationship we selected [*H]PN200-110, a
calcium channel antagonist, to perform radioligand bind-
ing experiments. First, rat brain homogenates were used
to obtain the channel affinity and B,,,,(density). Second,
in vitro receptor autoradiography was performed with
[7H]PN200-110 on 25 human hippocampus samples (AD,
n = 15; control, n = 10). Tissue was sectioned to 20 wm
using a cryostat and then incubated with 1.0 nM
[7H]PN200-110. Non-specific binding was binding in the
presence of 1.0 1m Nifedipine. The autoradiograms were
then analyzed with computer-aided densiometry to quan-
tify the density of the channels in the hippocampal sub-
regions. Preliminary data suggests that the binding of
PN200-110 is specific to the L-type channels. However,
the exact relationship between the channel density and the
AD pathology is still unclear at this time.
Characterization of AD-like pathology in normal aging.
CHRISTOPHER R. BRACKNEY,* LARRY SPARKS,
and STEVE SCHEFF, Department of Anatomy and Neu-
robiology, University of Kentucky, Lexington, Ky 40536.
Alzheimer’s disease is characterized by a progressive
cognitive deterioration coupled with marked personality
changes. The most notable pathological changes associat-
ed with Alzheimer’s disease are Senile Plaques (SP’s) and
Neurofibrillary Tangles (NFT’s). SP’s contain the B-amy-
loid protein, a 39-42 amino acid peptide fragment derived
from a transmembrane amyloid precursor protein (B-
APP). NFT’s consist of a mass of introcellular argento-
philic fibers, which are Paired Helical Filaments (PHF)
resulting from abnormally phosphorylated tau protein.
The present study assesses which B-APP peptide compro-
mise the B-amyloid immunoreactivity in the brains of non-
demented, non-heart disease, individuals 20-80 years of
life. By using specific antibodies we will be able to con-
clusively demonstrate which portion of the B-APP is ac-
cumulating in normal aging.
The accumulation of the C-100, a fragment derived
from B-APP, precludes the formation and deposition of
SP’s. The two immediate derivatives of the C-100 frag-
ment are B-A42(43) and B-A40. It has been shown that
B-A42(43) is found in greater accumulations then B-40 in
Alzheimer’s disease. To examine the presence of
B-A42(43) and B-A40 in normal aging we used specific
antibodies to discrete regions of the B-APP peptide se-
quence. We found several individuals who had the
B-42(43) with lower densities then that of Alzheimer’s dis-
ease. There were no accumulations of the B-40 in normal
aging. It should also be noted there was no correlation
between the presence of B-A42(43) and age. These results
suggest that the B-APP processing in normal aging is sim-
ilar to that found in Alzheimer’s disease.
Chromosome 17 deletions in a subset of ovarian tu-
mors. LYNN CARRICO* and MAURA PIERETTI, De-
partment of Pathology and Laboratory Medicine, Univer-
sity of Kentucky, Lexington KY 40506.
Ongoing studies in the Pieretti laboratory indicate that
different alterations in cancer-controlling genes occur in
different histological types of ovarian tumors. The labo-
ratory has already compiled data concerning genetic mu-
tations on 145 tumors. However, in order to statistically
correlate this genetic information to patients’ survival and
recurrence, a larger number of cases needs to be evalu-
ated. For this purpose, 45 additional cases were identified
through the Kentucky Cancer Registry, which has a record
of clinical, demographic, and follow-up data from Ken-
tucky cancer cases after 1991. These tumors can be stud-
42 Transactions of the Kentucky Academy of Science 58(1)
ied for the genetic alterations that characterize the differ-
ent histological types: genetic deletions of chromosome
17, p53 mutation, and K-ras activation. Genetic deletions
of chromosome 17 were studied by polymerase chain re-
action of microsatellite markers and were identified as the
loss or partial loss of one allelic band.
The tumor tissue was removed from paraffin embedded
blocks by a “punching out” method, and the DNA was
phenol extracted and ethanol participated. The resultant
DNA was then studied for deletions using the previously
mentioned method. Preliminary results are available for
13 of the 46 total tumors. Five tumors showed no dele-
tions on chromosome 17. Six tumors showed deletions on
both arms of chromosome 17. Two tumors had deletions
confined to the short arm of chromosome 17. All genetic
deletions were observed in tumors of high grade. Further
studies will be conducted with these samples and the re-
sulting data will be evaluated for prognostic significance.
Determination of cytokine production following OVA
injection into the eyes of adoptive transfer mice. JASON
E. DELONG,* JEROLD W. WOODWARD, RITA
EGAN, and RICHARD BLACK, Department of Immu-
nology and Microbiology, University of Kentucky, Lexing-
ton, KY 40502.
The eye, along with the central nervous system, has
been regarded as an immune-privileged site where normal
immune responses such as inflammation are suppressed
and antibody production is favored. Anterior chamber-as-
sociated immune deviation (ACAID) is a stereotypic sys-
temic response to ocular antigens that involves dendritic
cells assisting in trafficking antigens to the spleen where
t-cells are activated and an immune response is elicited.
The ACAID phenomenon is characterized by a shift in
the T-helper 1/T-helper 2 balance of responses toward a
dominant Th2 response. This alteration should be evident
in the cytokine profiles of mice following immunization
with antigen into the eye. Using an adoptive transfer
mouse model with an immunoflourescent intracellular cy-
tokine staining procedure, we have been able to detect
and analyze individual cytokines secreted by antigen-spe-
cific cells. The data attained through this project demon-
strates successful isolation and recognition of individual
IL-2 and IL-4 cytokines using the staining procedure with
flow cytometry. This establishes the intracellular cytokine
staining procedure as a useful tool in analyzing the im-
mune response following antigen injection.
Development of a sensing system for antimonite based
on genetically engineered bacteria and green fluorescent
protein. JANET MERCER,* SRIDHAR RAMANA-
THAN, and SYLVIA DAUNERT, Department of Chem-
istry, University of Kentucky, Lexington, KY 40506.
A sensitive and selective sensing system for antimonite
based on genetically engineered bacteria will be de-
scribed. In this system, the selective binding of a bacterial
protein to antimonite was coupled with fluorescence.
When antimonite enters certain bacteria, it is effluxed out
of the bacterial cell by a selective protein pump. The
structural proteins ArsA, ArsB, and ArsC form the protein
pump. A set of five genes, called the ars operon, code for
the three structural proteins, in addition to two regulatory
proteins, ArsR and ArsD. In the absence of antimonite,
the ArsR binds to the promoter region of the ars operon
inhibiting the expression of the protein pump. ArsR has a
highly specific binding site for antimonite. Thus, when an-
timonite reaches the cytoplasm it binds to ArsR inducing
a conformational change, which releases the promoter re-
gion of the ars operon. This induces the expression of the
protein pump, which effluxes the antimonite out of the
cell. Bacteria were genetically engineered to introduce the
gene of Green Fluorescent Protein (GFP) downstream
from the ArsR gene. The concentration of GFP in cells
can be monitored easily by checking for fluorescence.
Thus, the concentration of the GFP can be related to the
concentration of the antimonite in the cell.
Effect of retinioc acid on TIMP and stromelysin in HT
1080 human fibrosarcoma cancer cells. KEITH D.
BRICKING* and RAYMOND E. RICHMOND, Depart-
ment of Biological Sciences, Northern Kentucky Univer-
sity, Nunn Drive, Highland Heights, Ky 41099.
Recent research indicates that stromelysin, a matrix me-
talloproteinase (MMP) expressed by a large percentage of
malignant cancer cells, is capable of degrading extracel-
lular matrix proteins. Matrix degradation is an important
first step for cancer cell invasion, which is a precursor of
metastasis. Stromelysin may be inhibited by a specific
class of naturally occurring polypeptide inhibitors known
as tissue inhibitors of metalloproteinases (TIMP). TIMP
appears to act by non-covalently binding to stromelysin.
Therefore any therapeutic agent affecting cellular expres-
sion of stromelysin or TIMP may alter invasion. We ex-
amined the hypothesis that retinoic acid (a vitamin A an-
alog believed to have anticancer effects) will increase
TIMP expression in a highly metastatic HT 1080 human
fibrosarcoma cell line. These alterations will decrease the
invasive properties of the cell. Stromelysin and TIMP ex-
pression in 10-° M retinoic acid and vehicle control treat-
ed cells were detected by polyacrylamide gel electropho-
resis and immunoblot analysis. Invasion of HT 1080 can-
cer cells after addition of retinoic acid and vehicle control
has also been analyzed quantitatively using Matrigel In-
vasion Chambers. Our preliminary data indicate that there
was no change in stromelysin expression, an increase in
TIMP expression, and a significant decrease in the rate of
invasion after the addition of retinoic acid. These results
suggest that future antimetastatic agents should focus on
TIMP expression as well as matrix metalloproteinases.
Effect on platelet-derived growth factor A-chain gene
transcription by nm23 proteins in the WR-082-01 cell line.
MATTHEW SINEX,* BIN LIU, JAMES REID, and DA-
VID KAETZEL, Department of Pharmacology, Univer-
sity of Kentucky Medical Center, Lexington, KY 40536.
Platelet-derived growth factor (PDGF), a dimeric gly-
Abstracts, 1996 Annual Meeting 43
coprotein comprised of two disulfide-linked polypeptide
chains (A and B), is a powerful mitogen found in normal
and malignant cells. This laboratory has previously re-
ported that a 1 kb region of the PDGF A-chain promoter
located between nucleotides — 1800 and —880 (relative to
the transcription start site, +1) exhibits potent transcrip-
tional repression. Within that negative regulatory region
(NRR), a 31 bp silencer element (5’SHS) was identified
at approximately — 1400. Screening of a cDNA expression
library derived from HeLa cells with a probe derived from
the 5’SHS sequence yielded clones encoding nm23, a pro-
tein previously implicated in cancer metastasis. Subse-
quent studies revealed that purified nm23 binds with high
affinity to the 5’SHS sequence. The purpose of this study
was to determine whether nm23 plays a role in mediating
repression of the PDGF A-chain promoter. WR-082-01
cells, which are deficient in nm23 production, were co-
transfected with DNA constructs directing overexpression
of the H1 and H2 isoforms of nm23 and various A-chain
promoter fragments. The NRR was inactive in these cells,
suggesting that nm23 was important for its repression ac-
tivity. Overexpression of nm23-H1 and H2 resulted in a
general repression (< 2-fold) of all promoter constructs
analyzed. While these nm23 isoforms exerted no addition-
al inhibition upon vectors containing the 5’SHS element,
an additional two-fold repression was conferred by H1
upon vectors containing the entire NRR. These data sug-
gest that nm23-H1 mediates repression of A-chain gene
transcription, probably via interactions with multiple si-
lencer elements dispersed throughout the NRR.
Effects of nucleotide excision repair mutants on mis-
match repair. MARCI ADKINS* and ISABEL MEL-
LON, Department of Pathology, University of Kentucky,
Lexington, KY 40502.
We have previously shown that mismatch repair mu-
tants are also deficient in transcription coupled repair.
These results are important because they imply a possible
correlation between what was previously thought to be
distinct DNA repair systems. Present work has been un-
dertaken to further explore the connection between these
two repair processes. Since microsatellite instability is a
good marker for defective mismatch repair, we are spe-
cifically looking for microsatellite instability in Xeroderma
Pigmentosum (XP) cell lines. XP patients have deficiencies
in one or more nucleotide excision repair genes, which
have been named XP-A through XP-G. We have examined
XP-A, which is involved in damage recognition, and XP-C,
which repairs non-transcribed DNA. Clones of these cell
lines have been examined using primers specific for cer-
tain dinucleotide repeat sequences in a Polymerase Chain
Reaction (PCR). The PCR products are then run on a 6%
polyacrylamide gel and viewed by autoradiography. No
microsatellite instability has been detected in either one
of these cell lines. Therefore, our results deny a connec-
tion between mismatch repair and transcription coupled
repair.
Electrophysiological studies of the SCN8a voltage-gated
sodium channel. BRIAN DELISLE* and JONATHAN
SATIN, Department of Physiology, University of Ken-
tucky, Lexington, KY 40502.
SCN8a is a voltage-gated sodium channel isoform ex-
pressed in motor neurons. Loss of expression of the alpha
subunit in transgenic mice results in Motor Endplate Dis-
ease (MED). Transgenic mice often display a phenotype
that mimics human disease. Symptoms of MED include
progressive paralysis and juvenile death. The alpha sub-
unit of voltage-gated sodium channels consists of four
highly conserved domains. Most variability among the iso-
forms occurs among the one-two linker of the alpha sub-
units. This region contains functional glycosylation and
protein kinase A phosphorylation sites. Utilizing RT-PCR
protocol, we cloned the functional sites of SCN8a. Chi-
meric channels are currently under construction. The pro-
cess involves ligating digested BR2a, an extensively stud-
ied isoform, and cloned SCN8a. The chimeras are ex-
pressed in Xenopus laevis oocytes. We use the 2-electrode
voltage clamp to study the kinetic properties of the ex-
pressed ion channels and express the BR2a alpha subunit
with and without the beta-1 subunit. Coexpression of the
beta-1 subunit with BR2a modulates the rate of current
decay and shifts voltage dependance of the channels. All
experiments were then performed in the presence of
8-bromo-cAMP, promoting the phosphorylation of the so-
dium channel. The alpha subunit plus and minus the
beta-1 subunit both demonstrate a modest change in cur-
rent amplitude. We use these experiments to validate a
system to study the chimeric channels. The study is being
used to extend how coexpression of the beta-1 subunit
further modifies the sodium channel's kinetics.
Identification of cytokines responsible for enhancing
neonatal immune responses to polysaccharides. NIKOLE
L. GILBERT,* RALPH CHEL VARAJAN, and SUB-
BARAO BONDADA, Department of Microbiology and
Immunology, University of Kentucky, Lexington, KY
40502.
Polysaccharide antigens elicit a protective immune re-
sponse in adults but not in neonatal individuals, thereby
making neonates especially vulnerable to pathogenic,
polysaccharide-bearing bacteria such as Streptococcus
pneumoniae. Past research has shown that the B cell an-
tibody response, which mediates immune protection in
adults, is dependent on the presence of certain cytokines.
It was thus hypothesized that the neonatal B cells may be
competent to respond to polysaccharide antigens, but fail
to do so due to a deficiency of cytokines required to stim-
ulate neonatal B cells. To obtain an adult-like B cell re-
sponse, murine neonatal splenocytes were cultured in
vitro for 4 days with the polysaccharide antigen TNP-Fi-
coll and various cytokines known to effect B cell activity.
The plaque-forming cell-assay technique was used to mea-
sure the number of antibody forming cells per culture.
Neonatal splenocytes cultured with IL-4 and IL-5 togeth-
er and IL-1, IL-5, and IL-6 separately resulted in 40-70%
44 Transactions of the Kentucky Academy of Science 58(1)
of the adult response, while individual cytokines IL-2,
IL-3, IL-4, IL-10, and GM-CSF provided no enhance-
ment of the neonatal immune response. Neonatal splen-
ocytes cultured with a combination of cytokines IL-1 and
IL-5 produced 139% of the adult response to TNP-Ficoll.
In conclusion, our studies have shown that the neonatal
B cells can respond to polysaccharides as well as adult B
cells if an adequate mixture of cytokines is provided, sup-
porting the concept that the poor response of neonates to
polysaccharides is likely due to a deficiency in cytokine
availability.
Identification of genes involved in splicing. ELIZA-
BETH OTTE* and BRIAN RYMOND, Department of
Biology, University of Kentucky, Lexington, KY 40506.
The removal of introns from mRNA precursor, a pro-
cess vital to eukaryotes, is carried out by the spliceosome.
This complex enzyme consists of five small nuclear RNA
molecules and 50-100 protein subunits. A genetic screen
hoping to identify one or more of these spliceosomal pro-
teins was performed. In this screen, a mutant intron was
imbedded within the lacZ gene to produce a yeast strain
in which production of the enzyme B-galactosidase was
dependent upon the proper removal of this intron. Since
this fusion gene’s intron was poorly excised, this strain was
pale blue when grown in the presence of the chromogenic
substrate, X-gal. My project was to characterize a set of
recombinant plasmids which, when present in 20-50 cop-
ies per cell, turned the colonies a much darker blue, ap-
parently having enhanced B-galactosidase expression. It
was hypothesized that such genes might suppress the in-
hibitory effects of the intron mutation and increase pre-
mRNA splicing efficiency. The ratio of pre-mRNA to ma-
ture mRNA and the level of B-galactosidase in each pu-
tative suppressor was assayed. The data obtained indicate
that, under the conditions of assay, none of the suppres-
sors enhanced splicing efficiency or enzyme production. I
conclude that the enhanced blueness of these strains is
due to either an indirect effect, such as enhanced per-
meability of the X-gal substrate, or to enhancement of
lacZ transcription, pre-mRNA splicing, mRNA or protein
stability, or enzyme activity when cells are grown on the
surface of an agar plate but not when grown (as assayed)
in liquid culture.
Innervation of immune tissues and the role of nerve
growth factor. MARK PARRISH* and SONIA CARL-
SON, Department of Anatomy and Neurobiology, Uni-
versity of Kentucky Medical School, Lexington, KY 40536.
The existence of a structural and functional relationship
between the nervous and immune systems has been elu-
cidated in detail over the last decade. Immune tissues are
innervated by the sympathetic nervous system and the re-
lease of neurotransmitter modulates a variety of immune
responses. This modulatory relationship is well docu-
mented, but the plethora of directional cues guiding these
neurons to their immune tissue targets is not well under-
stood. Increasing evidence suggests that a protein, Nerve
Growth Factor (NGF), plays a significant role in directing
the pattern of peripheral innervation resulting from the
dependence of neurons on a target derived supply of this
factor. Recent work with a line of transgenic mice, altered
to overexpress NGF in skin, has given evidence that this
growth factor may also be influential in guiding sympa-
thetic innervation to immune tissues. NGF transgenics
show dramatic changes in density and pattern of inner-
vation to spleen and peripheral lymph nodes. To deter-
mine if spleens from transgenic mice release a substance
that alters neurite outgrowth, sympathetic ganglia were
co-cultured with spleen from transgenic and non-trans-
genic animals in the presence/absence of NGF and out-
growth was measured. Cultured ganglia exhibited growth
away from spleen which was reduced when cultured with
transgenic tissue or in the presence of NGF suggesting
that transgenic spleen retains NGF capable of stimulating
outgrowth. However, ganglia paired with transgenic
spleen also demonstrated changes in neurite morphology
and a reduced total neurite area which we believe may be
the result of cytokine production.
Localization of pyrimidine biosynthesis enzymes with
fusion proteins. DANA R. WEAVER,* JIANYUAN LUO,
and JOHN M. RAWLS, School of Biological Sciences,
University of Kentucky, Lexington, KY 40502.
De novo pyrimidine biosynthesis is carried out by six
enzymes contained in three proteins in animals: the CAD
protein, DHOdehase, and the UMP synthase protein.
These proteins are encoded by the r. dhod. and r-l genes
respectively in Drosophila. Biochemical cell fractionation
studies have shown that the CAD and UMP synthase pro-
teins are cytosolic, whereas DHOdehase is located in mi-
tochondria. We are carrying out studies to determine the
distributions of these proteins within cells and among dif-
ferent tissues, using fusion proteins containing the Aequo-
rea victoria green fluorescent protein (GFP) and the he-
magglutinin epitope (HA). A series of recombinant con-
structs was created containing the GFP and HA open
reading frames fused to the open reading frame of the
dhod gene. Transgenic animals were produced that con-
tain those fusions and genetic complementation confirmed
in vivo function of the transgenes. Patterns of expression
of the transgene protein were studied by UV fluorescence
microscopy (GFP) and immunocytochemistry (HA epi-
tope) in cells known to abundantly express CAD and
DHOdehase: ovaries (i.e., nurse cells) and testes (i.e.,
elongated spermatids). Results of these experiments will
be presented.
Microsatellite instability (MIN) and TGF-B Type II re-
ceptor gene mutations in human pancreatic carcinoma.
CINDY HARP,* CAROL SWIDERSKI, and JAMES
FREEMAN, Department of Surgery, University of Ken-
tucky Chandler Medical Center, Lexington, KY 40536.
Tumor progression is a multi-step process involving an
increase in positive growth signals and a decreased re-
sponse to negative growth regulation. In pancreas cancer
Abstracts, 1996 Annual Meeting 45
this process often involves the mutational activation of the
ras cellular oncogene and a concomitant loss in function
of tumor suppressor genes. We found that a common
characteristic of pancreatic cancers is loss in negative
growth regulation by TGF-. This loss in regulation is
often mediated by lack of expression of the TGF-B type
II receptor gene (RII), a reported tumor suppressor gene.
Microsatellite instability (MIN) has been proposed as a
mechanism that targets repetitive sequences in the RII
gene for mutation, resulting in loss of RII expression and
function. The purpose of this study was to determine the
incidence of MIN in pancreas cancer and to determine
whether MIN targets the RII gene for mutation. Six loci
were analyzed for the occurrence of MIN in pancreas can-
cer specimens obtained from 16 different patients. The
results show that MIN (9 of 16, 56%) and loss of hetero-
zygosity (LOH), (7 of 16, 44%) are common in pancreatic
tumors. We further analyzed prospective and retrospec-
tive pancreas tumor specimens by *?’-SSCP for RIT mu-
tations at regions comprising 2 potential MIN target sites.
Mutations at potential MIN sites were detected in only 3
of 21 (14%) specimens all of which were in 5’ region of
the gene that included a polyadenine repetitive sequence.
RII mRNA was not detectable in 5 of 12 (42%) prospec-
tive samples. These studies suggest that there is a greater
incidence of MIN in pancreas tumors than reported for
other tumor types. MIN, LOH, and loss in RII expression
may play a role in the pathology of this disease. However,
MIN targeted RII mutations occur in only a small sub-
population of pancreas tumors. Therefore mechanisms
other than MIN must account for the lack of RII expres-
sion found in many pancreas tumors.
Neuroprotective effects of nicotine in MPTP-treated
C57B1/6 mice. ALAN NORTHINGTON+# and JAMES R.
PAULY, College of Pharmacy, University of Kentucky,
Lexington, KY 40536.
Parkinson’s disease (PD) is a neurodegenerative disor-
der characterized by a selective loss of CNS dopamine-
containing neurons. Epidemiological studies have consis-
tently demonstrated an apparent neuroprotective effect of
cigarette smoking on the development of PD, with smok-
ers having approximately half the risk of nonsmokers of
developing the disease. Mice injected with the neurotoxin
1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine (MPTP) de-
velop a syndrome resembling human PD with a loss of
nigrostriatal dopaminergic neurons. The purpose of the
present study was to examine whether nicotine pretreat-
ment reduces the neurotoxic actions of MPTP on striatal
dopaminergic neurons. Mice were injected with either sa-
line or nicotine (1.0 mg/kg, sc.) 10 minutes prior to ad-
ministration of MPTP (40 mg/kg, sc.) or saline. Locomo-
tor activity, motor coordination, and body weight were
monitored for 12 days following MPTP treatment. The
brains from each treatment group were then used for the
evaluation of [?H]-GBR12935 binding to the dopamine
transporter and binding to neuronal nicotinic receptor
subtypes (using [°H]-cytisine and [!*°I]-alpha bungarotox-
in). Animals treated with saline and MPTP demonstrated
significant behavioral toxicity and weight loss; these effects
were clearly attenuated in animals pre-treated with nico-
tine. In the striatum, MPTP caused a significant reduction
in the binding of [*H]-GBR12935 and [°H]-cytisine but
not [!*°I]-alpha bungarotoxin. The toxic effects of MPTP
on [°H]-GBR12935 and [*H]-cytisine were diminished by
nicotine pretreatment. Possible mechanisms of nicotine-
induced neuroprotection will be discussed.
Screening for mutations causing failures in the RNA
sorting system in spermiogenesis. NATHAN NOVOTNY,+*
JAN URSO, and JOHN RAWLS, School of Biological Sci-
ences, University of Kentucky, Lexington, KY 40506.
In spermatogenesis, post-meiotic differentiation of
spermatids is an extensive morphogenetic program that
takes place, largely or entirely, in the absence of transcrip-
tion. It utilizes RNAs synthesized during spermatocyte
growth, stored in an inactive form during meiosis, then
translated at various times during spermiogenesis. Sper-
matocytes have an active system that sorts RNAs to be
used immediately from those to store for later use; the
components of this RNA sorting system are largely un-
known. The objective of this research is to identify com-
ponents of this system among the numerous male-sterile
mutations that have been characterized in Drosophila mel-
anogaster, by seeking variant expression of dhod RNA in
mutant flies. A spermatocyte-specific promoter element of
the dhod gene leads to synthesis of RNA that, mediated
by a specific 5' RNA sequence (dTCE), is sequestered for
ultimate translation in elongated spermatids. In a primary
survey, spermatogenesis stages in over eighty male-sterile
strains have been examined for histochemical expression
of the dhod product dihydroorotate dehydrogenase
(DHOdehase). Most strains displayed essentially normal
expression patterns for DHOdehase, however a few
strains showed aberrant expression patterns that are pre-
dicted for mutations of the spermatogenesis gene control
system: zero or strongly reduced expression; expression of
DHOdehase in early stages (possibly precocious expres-
sion). As a secondary survey, mutants showing altered ex-
pression of DHOdehase are being tested for expression of
an array of lacZ reporter transgenes that contain various
elements that control dhod expression: the dhod promoter
or the promoter of another spermatocyte-expressed gene;
with or without the dTCE sequence that controls trans-
lational delay. Results of these experiments will be pre-
sented.
Site directed mutagenesis of a conserved phenylalanine
in cytochrome P450 2E1. ERIC DEDRICK* and TODD
PORTER, College of Pharmacy, University of Kentucky,
Lexington, KY 40536.
Cytochrome P450 2E1 is an enzyme that catalyzes the
oxidation of a wide array of substrates, xenobiotics, and
naturally occurring chemicals. The oxidative capabilities of
P450s arise from a heme group folded deep within the
molecule. Although it is not fully understood how an elec-
46 Transactions of the Kentucky Academy of Science 58(1)
tron donated from its reductase partner is able to make
the transition from the surface of the P450 molecule to
the heme group in the interior, it is thought that a highly
conserved phenylalanine at position 429 facilitates this
transfer. Previous mutagenesis experiments have suggest-
ed that hydrophobic compounds can substitute for this
phenylalanine and still provide a functional molecule. Fur-
thermore, because of its structure, proline may fit well at
this position, causing a minimal distortion of the P450;
notably, a proline has been discovered at this position in
a functional P450 enzyme from flax seed. In this experi-
ment, a proline codon was substituted for the phenylala-
nine codon at this position using polymerase chain reac-
tion. This mutation was then ligated into the expression
vector pJL2/3a and expressed in Escherichia coli. Once
positive mutants have been identified by sequencing, the
protein will be characterized in spectrophotometric and
catalytic assays.
Stage dependent induced hatching in the muskellunge,
Esox masquinongy. TAMARA SAPP,* SARAH M.
BLANK, and JOHN J. JUST, Department of Biological
Sciences, University of Kentucky, Lexington, KY 40506.
The embryos of all aquatic animals are encased in pro-
teinaceous egg cases from which they escape during de-
velopment. Hatching often involves digestion of the egg
case by a hatching enzyme secreted from hatching glands.
This study shows that hypoxic environments can induce
premature hatching in Esox masquinongy 9 days post-fer-
tilization, 5 days before normal hatching. Embryos 9, 10,
11, and 12 days old were subjected to environments with
various oxygen pressures by aerating vials containing em-
bryos with oxygen (PO, = 760 mm Hg), air (PO, = 160
mm Hg), or nitrogen (PO, = 0 mm Hg) for 1 hour. At
all stages tested, eggs aerated with nitrogen (n = 461)
exhibited an increased hatch rate compared to the control
embryos treated with air (n = 463), while those treated
with oxygen (n = 461) showed a greatly reduced hatch
rate. There was a progressive increase in percent hatch
with those embryos treated with nitrogen. Percent hatch
of 9, 10, and 11 day old embryos after 1 hour of nitrogen
treatment was 46%, 90%, and 99% respectively, while 9,
10, and 11 day old control embryos treated with air had
hatch rates of 0%, 1%, and 46%. It is believed that hatch-
ing can occur only after the development of hatching
glands and enzymes. Hatching medium was collected
from a mass hatching of E. masquinongy embryos and the
macromolecules (MW > 10,000) were concentrated via
ultrafiltration. Protease activity was demonstrated by di-
gestion of gelatin on photographic film. Work continues
to isolate the hatching enzyme and correlate its develop-
mental appearance with the ability to prematurely hatch.
TNF-alpha neuroprotection in opiate-mediated astrog-
lial toxicity. JENN JACKSON,* MARK P. MATTSON,
and KURT F. HAUSER, Department of Anatomy and
Neurobiology, University of Kentucky, Lexington, KY
40536.
Opiate drugs of abuse liability, such as heroin and mor-
phine, inhibit fetal brain development, in part by disrupt-
ing astroglial mitosis and also by inducing astroglial hy-
pertrophy. Although opioids affect growth, the mecha-
nisms by which opioids disrupt neurobehavioral develop-
ment are not understood. The mu-opioid receptor serves
as a primary target of opiate drugs of abuse. In astroglia,
mu-opioid receptor stimulation causes increases in intra-
cellular calcium. The resulting elevations in intracellular
calcium mediate morphine-induced alterations in astro-
glial growth. The present study tested the hypothesis that
a naturally occurring neuroprotective factor, tumor necro-
sis factor-alpha (TNF-alpha), will prevent the deleterious
effects of opiate drugs by blocking opiate-induced increas-
es in intracellular calcium. Astrocytes were isolated asep-
tically from the cerebral hemispheres of 1-day-old ICR
mice. Astroglial cultures were pretreated with TNF-alpha
at least 24 hours prior to the opioid insult to serve a neu-
roprotective role. Astrocytes were subsequently treated
with TNF-alpha in the presence or absence of morphine
(1 2M). Intracellular calcium was analyzed by ratiometric
analysis of the calcium indicator, fura-2. The study showed
TNF-alpha, at concentrations of 10 ng/ml and 100 ng/ml,
causes a concentration-dependent reduction in morphine-
induced activation of intracellular calcium. These findings
suggest that mu-opioid receptor and TNF-alpha signaling
pathways converge and have opposing actions in calcium
homeostasis. Studies in progress are assessing whether
TNF-alpha attenuates morphine-induced alterations in as-
troglial growth.
Translational regulation of mRNA in human breast can-
cer. BRAD SEGEBARTH* and STEPHEN ZIMMER,
Department of Microbiology and Immunology, University
of Kentucky, Lexington, KY 40502.
This proposal is based on the hypothesis that increased
initiation of messenger RNA (mRNA) translation in hu-
man breast cancer cells is a fundamental mechanism me-
diating increased tumor cell growth and the capacity to
invade and metastasize. The eukaryotic initiation factor 4E
(eIF-4E) is a 25 kilodalton (kDa) phosphoprotein involved
in the recruitment of messenger RNA to the polysomes.
This reaction is considered to be the rate-limiting step in
the synthesis of cellular proteins, with eIF-4E as the least
abundant subunit. Studies show that overexpression of
eIF-4E in human breast cancer cells is associated with
increased translation, cellular proliferation, transforma-
tion, and tumorigenesis. PHAS-1 is a 20 kDa protein
found to bind to eIF-4E. The binding seems to be regu-
lated by phosphorylation, with the complex dissociating
upon phosphorylation of PHAS-1. About 70% of mRNAs
containing extensive 5’-region secondary structure are on-
cogenes or important growth regulatory genes. One of
these, basic fibroblast growth factor (bFGF), is a multi-
functional molecule that affects cell growth and induces
angiogenesis. Due to the extensive 5’ region secondary
structure, bFGF, along with other malignant characteris-
tics, may be disproportionally inhibited in cells overex-
Abstracts, 1996 Annual Meeting 47
pressing PHAS-1. If this is found to be the case, potential
medical utility stems from being able to effectively devel-
op a drug that can affect the PHAS-1/eIF-4E interaction
and suppress malignant properties in oncogenic cells.
PHYSICS & ASTRONOMY
Celestial seasons. RICHARD HACKNEY,* KAREN
HACKNEY, ROGER SCOTT, CHARLES McGRUDER,
MICHAEL CARINI, RICHARD GELDERMAN, SAN-
DRA CLEMENTS, JOHNNY JENNINGS, JOHN NO-
BLE, and RICO TYLER, Western Kentucky University
Astrophysical Observatory, Department of Physics and As-
tronomy, Western Kentucky University, Bowling Green,
KY 42101.
Many have only a casual acquaintance with (and an in-
adequate understanding of the causal connection be-
tween) the seasons and related observable astronomical
phenomena. In our introductory astronomy courses, we
attempt to engage students’ minds through personal ob-
servation and interpretation of data to make the connec-
tions and to see the astronomical reasons for the seasons.
We describe a number of activities suitable for students
at various levels to illustrate through student participation
the primary astronomical observables related to the cycle
of the seasons. We acknowledge funding by the Eisenhower
Math and Science Program, the Kentucky PRISM-UG
Program, and the NASA Kentucky Space Grant Consor-
tium.
Comet Hale-Bopp: Opportunity for education and de-
veloping student involvement. ROGER SCOTT,* KAR-
EN HACKNEY, RICHARD HACKNEY, CHARLES
McGRUDER, MICHAEL CARINI, RICHARD GELD-
ERMAN, SANDRA CLEMENTS, and JOHN NOBLE,
Department of Physics and Astronomy, Western Kentucky
University, Bowling Green, KY 42101.
Comet Hale-Bopp, already visible with binoculars in
the early evening sky, promises to be an easy naked-eye
object in March and April 1997. Comets are notorious for
being unpredictable in brightness. However, Hale-Bopp
seems to be brightening about as expected and may put
on a show even better than Hyakutake. Bright comets of-
fer excellent opportunities for student projects and for ed-
ucational outreach to the general public. They are easily
photographed using any 35-mm camera capable of taking
a time exposure. We discuss simple student projects and
techniques for photographing the comet. Photographs of
other bright comets of recent decades are shown as ex-
amples. Charts and finding information are provided.
Enigmatic world of extragalactic radio jets. CASEY
WATSON,* ILDEFONSO GUILARAN, RYAN CULL-
ER, MONICA DECKARD, RICHARD GELDERMAN,
MICHAEL CARINI, JOHN NOBLE, SANDRA CLE-
MENTS, ROGER SCOTT, KAREN HACKNEY, and
RICHARD HACKNEY, Western Kentucky University As-
trophysical Observatory and Center for Automated Space
Science, Department of Physics and Astronomy, Western
Kentucky University, Bowling Green, KY 42101.
Radio galaxies, quasars, Seyferts, and blazars are all des-
ignated by the term Active Galactic Nuclei or AGN.
Among the most striking characteristics of AGN are the
collimated radio jets, up to millions of light years in
length, which emanate from their cores. The radio emis-
sion is due to synchrotron radiation, which arises when
charged particles from the active nucleus spiral around a
magnetic field. However, the precise physical nature of
the jets remains undetermined. Discovering the compo-
sition of the jets is crucial because that insight will lead
to greater understanding of the AGN’s central engine. It
is the primary goal of this project to make a contribution
to current jet theory. Our theoretical work will be com-
plemented by observations, including images from the
CCD camera at the WKU Astrophysical Observatory and
the Hubble Space Telescope. We thank the NASA Ken-
tucky Space Grant Consortium and the NASA Center for
Automated Space Science for support.
Gravitational lensing simulation for the classroom. MA-
RIA FALBO-KENKEL+# and JOE LOHRE, Department
of Physics and Geology, Northern Kentucky University,
Highland Heights, KY 41099.
Gravitational macro-lensing is a celestial phenomenon
that can be observed when two or more objects at differ-
ent distances from the earth lie along the same line of
sight in the sky. The observation of this effect, which pro-
duces distortions and apparent displacements of the ob-
ject furthest from earth in the alignment, was listed by
Einstein in his famous 1915 paper as one of three impor-
tant tests of general relativity. There are ways to optically
simulate gravitational lensing events. We present an in-
expensive and simple method to make a lens that can be
used to easily simulate gravitational lensing in the class-
room. We discuss two effects that can be studied with the
lens: (1) Einstein-ring formation and (2) multiple image
formation. This work was supported by funds from the
Kentucky Space Grant Consortium.
Morehead radio telescope: design and fabrication of a
research instrument for undergraduate faculty and student
research in radio frequency astrophysics. BENJAMIN K.
MALPHRUS,* BRIAN M. LEWIS, and CHRISTO-
PHER L. SMITH, Morehead Astrophysical Observatory,
Morehead, KY 40351.
Faculty and students of the departments of Physical
Sciences and Industrial Education and Technology at
Morehead State University (MSU) (Morehead, KY) have
designed and assembled the Morehead Radio Telescope
(MRT) to provide a research instrument for undergradu-
ate astronomy and physics students and an active labora-
tory for MSU undergraduate students and faculty in phys-
ics, engineering, and computer science and for under-
graduate students, faculty, and science teachers through-
out Kentucky. Goals of the MRT program are to enhance
the curricula in physics, physical science, electronics, and
48 Transactions of the Kentucky Academy of Science 58(1)
science education programs by serving to provide (1) a
research instrument for investigations in astronomy and
astrophysics; (2) an active laboratory in astronomy, phys-
ics, electrical engineering, and computer science; and (3)
a research instrument and laboratory for science teacher
education and inservice programs. The telescope incor-
porates a modular design in which components may be
easily removed for use in laboratory investigations and for
student research in design projects. The performance
characteristics of the telescope allow a varied and in-depth
scientific program. The sensitivity and versatility of the
telescope design facilitate the investigation of a wide va-
riety of astrophysically interesting phenomena. The MRT
provides hands-on experience in research and instrumen-
tation technology in a cutting-edge science, one that is in
the midst of scientific revolution. This presentation pro-
vides and overview of the MRT and its current operating
status.
NASA Kentucky Space Grant Consortium, Center for
Automated Space Science, and EPSCoR opportunities for
space-related science, technology, and education. KAREN
HACKNEY,* RICHARD HACKNEY, ROGER SCOTT,
CHARLES McGRUDER, MICHAEL CARINI, RICH-
ARD GELDERMAN, JOHN NOBLE, SANDRA CLE-
MENTS, and JOHNNY JENNINGS, Kentucky Space
Grant Consortium, NASA EPSCoR Program, and Center
for Automated Space Science, Department of Physics and
Astronomy, Western Kentucky University, Bowling Green,
KY 42101; MICHAEL BUSBY, Center for Automated
Space Science, Center of Excellence in Information Sys-
tems Engineering and Management, Tennessee State Uni-
versity, Nashville, TN 37203.
NASA has funded three programs in Kentucky to in-
volve university faculty and students in space-related re-
search, technology, and education. We describe (1) cur-
rent opportunities in the Kentucky Space Grant Consor-
tium for undergraduate scholarships and graduate fellow-
ships for students in mentored, space-related projects; (2)
funding opportunities for development of space-related
research projects and activities to support teaching of
space science and related disciplines; (3) progress of ex-
isting projects and new opportunities in the NASA
EPSCoR Program; and (4) special opportunities for mi-
nority students in the Center for Automated Space Sci-
ence.
Use of computers for the enhancement of introductory
astronomy for non-science majors. RAYMOND C. Mc-
NEIL, Department of Physics and Geology, Northern
Kentucky University, Highland Heights, KY 41099.
Within the sciences, introductory laboratory experienc-
es allow students a better understanding of scientific laws
and a better appreciation of the scientific method leading
to their discovery. At the heart of a laboratory course is
the assumption that students will participate in “hands-
on” activities that will provide experience in gathering
data, analyzing the data to find relationships, and drawing
conclusions from the experimental results. Traditionally,
introductory astronomy students have been limited in that
they generally cannot actively conduct experiments, con-
trolling one or more of the variables, as can students in
other sciences. The solution at Northern Kentucky Uni-
versity (NKU) and other institutions has been to utilize
laboratory computers to allow students to explore, then
investigate in detail, simulations of astronomical phenom-
ena. Observations may be repeated, and in some cases
variables changed, just as in the laboratories of other sci-
entific disciplines. The active nature of such experiences
can enhance learning and instill a sense of the excitement
of scientific discovery. The acquisition and initial use of a
dedicated set of computers for the astronomy laboratory
at NKU is described. Materials developed by the author
or adapted from other sources are discussed, including
Contemporary Laboratory Experiences in Astronomy,
with copies available upon request. A list of software
found to be most promising as the basis for laboratory
activities or other types of student involvement is also in-
cluded, as are future directions for this work. The author
gratefully acknowledges the support of the National Sci-
ence Foundation (Grant #9452219) and NKU.
Optical monitoring of very high redshift quasars. SAN-
DRA CLEMENTS, Western Kentucky University Astro-
physical Observatory and Center for Automated Space
Science, Department of Physics and Astronomy, Western
Kentucky University, Bowling Green, KY 42101.
The observations and results of a 1-year program to
monitor high redshift quasars is reported. Quasars are
among the most distant and energetic objects in the uni-
verse. Believed to be caused by activity in the centers of
galaxies, quasar behavior has been a challenge to under-
stand. After their discovery, monitoring campaigns quickly
noted variability in the radiation emitted by some quasars.
Subsequent variability studies have uncovered a wealth of
information about their underlying physics. However, due
to a lack of monitoring data, such studies have yet to be
performed on the most remote quasars, those with red-
shift = greater than 4. Since first discovered in the late
1980s, about 50 of these quasars have been found. Com-
paring the physical characteristics and behavior of quasars
in the very early universe with less remote quasars will
provide insight into their evolution. The variability studies
required to determine the underlying physics of these ob-
jects require monitoring data. To begin to satisfy this re-
quirement, a program to monitor quasars with z greater
than 4 was initiated in June 1995. to date, over a dozen
objects have been monitored at the University of Florida's
Rosemary Hill Observatory, Bronson, FL, and the West-
ern Kentucky University Astrophysical Observatory, Bowl-
ing Green, KY.
Photometry with CCDs: Optimizing the signal extrac-
tion. RYAN CULLER,* FONSIE GUILARAN, MONI-
CA DECKARD, CASEY WATSON, MICHAEL CARI-
NI, RICHARD GELDERMAN, JOHN NOBLE, RICH-
Abstracts, 1996 Annual Meeting 49
ARD HACKNEY, KAREN HACKNEY, ROGER
SCOTT, and SANDRA CLEMENTS, Western Kentucky
University Astrophysical Observatory and Center for Au-
tomated Space Science, Department of Physics and As-
tronomy, Western Kentucky University, Bowling Green,
KY 42101.
The CCD detector has revolutionized astronomical
photometry. Photometric precision better than 1% has
been achieved for bright sources. Faint sources, such as
Active Galactic Nuclei (AGN), can also be studied with
photometric precision approaching 1%. This is a signifi-
cant improvement over the 10% precision achieved with
photographic plates and the 5% precision usually achieved
with photomultiplier tubes. However, these improvements
can be reached only through the proper processing of the
CCD image and appropriate extraction of the brightness
information from the processed image. We discuss (1)
what we have found to be optimal methods for both the
processing of CCD images and the extraction of bright-
ness information and (2) application to the measurement
of brightness variations in AGN. We acknowledge funding
from the NASA Kentucky Space Grant Consortium and
the NASA funded Center for Automated Space Science
at Western Kentucky University.
PHYSIOLOGY & BIOCHEMISTRY
Effects of ovariectomy and dietary energy restriction on
the body weight of Fischer 344 rats. C. BROWN,* Y.
ZHANG, C. WANG, and C. J. LEE, Community Re-
search Service, Kentucky State University, Frankfort, KY
40601.
Thirty-two fischer 344 rats were randomly assigned into
four groups of eight rats each. Group 1 was sham-oper-
ated and fed the control diet (AIN-76A) ad libitum; Group
2 was ovariectomized and fed the control diet at the av-
erage intake of Group 1; Group 3 were sham-operated
and fed the energy-restricted diet at 60% of the average
intake of Group 1; Group 4 were ovariectomized and fed
the energy-restricted diet at 60% of the average intake of
Group 1. Sixty grams of the energy-restricted diet provid-
ed only 60% of the energy content, but equal amounts of
protein, minerals, and vitamins of 100 g of the control
diet. The experimental period was 8 weeks. There were
no significant changes in the body weight of Group 1,
whereas the body weight of Group 2 increased by 8% of
the initial body weight. Both Group 3 and 4 lost ca. 20%
of the initial body weight with no differences between the
two groups. These results indicate that ovariectomized
rats were more likely to gain weight than the sham-op-
erated rats when fed the control diet and that both ovar-
iectomized and sham-operated rats will lose weight when
fed energy-restricted diets.
Effects of ovariectomy surgery on urinary calcium ex-
cretion of Fischer 344 rats. D. HAMPTON,* Y. ZHANG,
C. WANG, and C. J. LEE, Community Research Service,
Kentucky State University, Frankfort, KY 40601.
Sixteen female Fischer 344 rats were fed AIN-76A for
1 week before they were placed in metabolic cages and
the urine excretion was completely collected for 3 days.
Half of the rats went through ovariectomy with the other
half through sham-operation. One week after the opera-
tion, rats were placed in metabolic cages with the urine
collected for 3 days. Urine samples were analyzed for cal-
cium by atomic absorption spectrophotometry. Dietary in-
take, thus, dietary calcium intake, was lower after surgery
than before surgery. Calcium intake post surgery was low-
er for ovariectomized than for sham-operated rats. Uri-
nary calcium excretion (mg/day) decreased by 20% for
sham-operated rats and by only 5% for ovariectomized
rats. These results suggest that surgery decreased calcium
absorption from the gut, leading to lower urinary calcium
excretion in sham-operated rats; increased calcium mo-
bilization from the bone may be responsible for the higher
urinary calcium excretion in ovariectomized rats.
Gentamicin-induced nephrotoxicity in the teleost fish
Astronotus ocellatus (Cichlidae). W. STEWART CURT-
SINGER* and HONG Y. YAN, School of Biological Sci-
ences, University of Kentucky, Lexington, KY 40506.
Aminoglycoside antibiotics (AGs) are effective in treat-
ing bacterial infections. The AGs also inflict ototoxic and
nephrotoxic effects on treated subjects. Nephrotoxicity of
gentamicin sulfate on fish was first described by Yan et
al.; edema caused by gentamicin was documented subse-
quently. We used histological and autoradiographical
methods to document temporal course of intake of gen-
tamicin as well as cellular/structural damages caused by
the gentamicin to the fish kidney. The experimental sub-
jects, the teleost fish Astronotus ocellatus, received one,
two, three, or four daily intramuscular injection(s) of 20
mg/kg or 120 mg/kg gentamicin and sacrificed 1 day after
the last injection. Fourteen fish also received one injection
of *H-labeled gentamicin (5 Ci/10 gm fish). Two fish as
a group were sampled 1, 2, 4, 8, 24, 36, 48 hours after
the injection. Kidney tissues were dissected and processed
with standard histological procedures. Kodak-NTB3 emul-
sion was applied to sections with *H-labeled gentamicin.
Exposure was carried out in a light-tight box up to 42 days
and developed. Edema was obvious after just one injec-
tion. The weight gain increased as fish received more in-
jections in both dosage groups. There was no significant
difference in weight gain between two treatments. An in-
dication that 20 mg/kg dosage was sufficient to illicit ne-
phrotoxic damages. The necrosis was evident in the prox-
imal tubular epithelial cells. Autoradiographical results
showed the maximal uptake of gentamicin occurred be-
tween 36 and 48 hours after injection. The present study
shows that fish kidney is a valid model for aminoglycoside
nephrotoxicity study. Supported by Deafness Research
Foundation, NIH-DC01729, and Howard Hughes Medi-
cal Institute.
Potential gentamicin-binding protein(s) in the ear of the
cichlid fish Astronotus ocellatus (Cichlidae). STEVEN M.
RIGDON,* W. STEWART CURTSINGER, BIN-TAO
50 Transactions of the Kentucky Academy of Science 58(1)
PAN, and HONG Y. YAN, School of Biological Sciences
and Department of Neurosurgery, University of Kentucky,
Lexington, KY 40506.
Evidence of aminoglycoside (e.g., gentamicin sulfate)
damages to the auditory sensory hair cells of the teleost
fish Astronotus ocellatus was previously demonstrated.
Continuation of the study further showed regeneration of
damaged ciliary bundles 10 days after maximal damage.
Our preliminary results using both immunocytochemical
and autoradiographical methods indicate that the incor-
poration of gentamicin into hair cells occurs between 36
and 48 hours after intramuscular injection. To better un-
derstand the ototoxic and regeneration mechanisms in-
volved, Western blotting was used to identify the presence
of potential gentamicin binding protein(s). The fish were
injected with gentamicin sulfate (120 mg/kg) and sacri-
ficed either 24 or 48 hours later. Control fish received only
saline injection. Three tissues (ear, kidney, and brain) were
dissected and processed following standard immuno blot-
ting protocols. Results indicated the presence of potential
gentamicin-binding protein(s) in ear and kidney tissues,
except tissues from control animals and brain tissues.
Higher degree of binding was observed in 48-hr tissues
than 24-hr samples. These results corroborated earlier im-
munocytochemical and autoradiographical findings on
temporal uptake sequences of gentamicin. On-going ex-
periments employing both biochemical and molecular
techniques are designed to characterize the nature of the
potential gentamicin-binding protein(s). It is hoped fur-
ther understanding of gentamicin-binding protein(s) could
lead to the development of prophylactic agent(s) against
both ototoxic and nephrotoxic damages caused by ami-
noglycoside antibiotics. Supported by Deafness Research
Foundation, NIH-DC01729 and Howard Hughes Medical
Institute.
Relationship between muscle lipid content and body
weight of paddlefish (Polyodon spathula). X. LOU, T. L.
WHITE,* C. WANG, and S. D. MIMS, Community Re-
search Service, Kentucky State University, Frankfort, KY
40601.
Lipid content of fish meat is important to processing
and marketing of fish meat. The objective of this study
was to determine the muscle composition of paddlefish
(Polyodon spathula: Polyodontidae) with various body
weight. Twenty-five paddlefish (body weight 4 to 20 kg)
were used for this study. Fillets with the red meat
trimmed off were prepared and analyzed for moisture,
total lipids, protein, and ash content. With increasing body
weight, muscle moisture and protein content tended to
decrease; muscle lipid content increased with ash content
remaining constant. Muscle lipid content increased dras-
tically when the body weight was 10 kg or more. These
data will have practical implications for the harvesting of
paddlefish and the ultimate use of paddlefish meat.
Skeletal compartmentalization and metabolism of cal-
cium in maturing male and female rats. D. L. DEMOSS*
and G. L. WRIGHT, Physiology Department, Marshall
University School of Medicine, Huntington, WV 25704.
The *H-tetracycline (*H-T) bone-labeling procedure
was employed to monitor bone resorption from urinary
loss curves in male and female rats of various ages. In
addition whole body dry skeletal mass and the loss of 3H-T
from individual bones was determined. It was found that
the dry skeletal mass/body mass ratio of females was sig-
nificantly greater than for males, indicating the impor-
tance of dry skeletal mass in quantitative assessments. The
urinary loss of *H-T was described by a double exponen-
tial equation (label loss from bone fluid and calcified skel-
etal compartments). The results indicate the presence of
two distinct and independent exchangeable bone com-
partments as determined by the *H-T method. Both com-
partments decrease in size with age but their label loss
activities are different. The label loss activity within the
rapidly depleted compartment, which we suggest affects
calcium loss from the bone fluid compartment, is “inert”
in the sense that it does not change with age or between
sexes. Label loss activity in the slowly depleted compart-
ment, which we suggest represents resorption of calcified
bone, is increased with aging. Despite a smaller skeletal
mass, resorptive activity in this compartment is higher in
female rats than in male rats by 24 weeks of age due to
a larger compartment size and a maintained higher rate
of resorptive activity within the calcified compartment.
Stability of paddlefish (Polyodon spathula) meat stored
in crushed ice. X. LOU,* C. WANG, Y. XIONG,! G. LIU,!
and S. D. MIMS, Community Research Service, Kentucky
State University, Frankfort, Kentucky 40601; Department
of Animal Sciences, University of Kentucky, Lexington,
KY 40546.
Fillets from six paddlefish (Polyodon spathula; Poly-
odontidae) were cut into seven sections for each fish. The
sections were stored in sealed plastic bags covered by
crushed ice. One section from each fish was taken on day
0, 1, 2, 3, 5, 7, 10. The samples were analyzed for thio-
barbituric acid-reactive substances (TBARS) as an indi-
cator of lipid degradation and protein solubility as an in-
dicator for protein degradation. The data were analyzed
with the GLM procedure of SAS and the storage time,
fish, and section were the independent variables. There
were no significant differences among the sections. There
were no significant changes in protein solubility or TBARS
during the 10 days of storage. These results indicate that
paddlefish meat stored in crushed ice is stable for at least
10 days.
Producing the finest community college graduates. J.G.
SHIBER, Division of Biological Sciences, Prestonsburg
Community College, Prestonsburg, KY 41653.
At Prestonsburg Community College. (Prestonsburg,
KY) 998 students completed a questionnaire on how com-
munity colleges (CC) can produce the “finest” graduates
for the 21st century, i.e., people well prepared for the
challenges ahead. Most said that the primary mission of a
Abstracts, 1996 Annual Meeting 51
CC should be teaching and learning, and that its basic
principle should be to provide a broad background so stu-
dents can further their education and/or enter the job
market. Successful graduates, they said, should be versa-
tile, willing to pursue varied career options, and able to
integrate disciplines. Nearly all agreed that to achieve this
end, the CC should offer courses and programs reflecting
basic economy and work force changes, draw upon all
parts of society's talent base to train students, and reassess
long-standing educational practices, especially in the sci-
ences and engineering. Are these students’ expectations
being met at their CC? When asked why enrollment is
down and what should be done about it, or how the CC
can best use its funds or improve its educational mission,
the most frequent response was always “Provide more
courses/degree programs/career options.” Many said that
their CC did not offer all the courses their majors re-
quired for entry into a university, that better facilities and
student services (e.g., daycare and tutoring) were needed,
and that more sensitivity to student needs and abilities,
with less emphasis on non-academic matters, is essential.
Of the 81% planning to attend a university, over half said
the most important thing they needed to experience at
the CC was improvement of personal discipline (study,
note-taking, attendance habits). The results indicate that
CC's may have to pay more attention to student academic
needs and concems if they are to produce the “finest”
graduates for the 21st century.
Student attitudes toward the study of biology. JOHN R.
MORRISON* and RUTH E. BEATTIE, T. H. Morgan
School of Biological Sciences, University of Kentucky,
Lexington, KY 40506.
A pre- and post-course attitudinal survey was adminis-
tered to students enrolled in freshman-level major and
non-major biology courses at the University of Kentucky
(Lexington, KY). The survey included questions on (1) the
perceived importance of biology in everyday life; (2) the
image of biology courses; and (3) factors that influenced
the student to take a biology course. The surveys were self
administered and the resulting data were analyzed using
the T-test and mean values. All surveys were administered
to students taught by the same instructor. Pre-course data
analysis indicated that non-biology-major students had a
less positive attitude than biology major students towards
the importance of biology in everyday life. Post-course
scores showed a significant increase in agreement with the
statement that “Everyone should study biology” and that
an understanding of biology is important in politics and
aids intelligent voting. All test groups agreed strongly with
the statement that “Biology involves a lot of memoriza-
tion.” An interest in the subject matter was the primary
factor in influencing students to take a biology course. The
majority of students surveyed had a positive experience in
their high school biology courses.
Student perspectives of field investigations: misconcep-
tions, problems, and educational benefits. TERRY L.
DERTING* and JILL H. KRUPER, Department of Bi-
ological Sciences, Murray State University, Murray, KY
42071.
Life is the theme of biology, yet all too often it is stud-
ied only in artificial laboratory environments. Despite the
inherent appeal of the outdoors, the idea of conducting
field investigations of biological phenomena is often met
with trepidation. We conducted a case study of a field
ecology class in which groups of students initiated and
completed field investigations. Many of these students had
no prior field experience. Throughout the course we mon-
itored the feelings, problems, and successes of each group.
Student misconceptions regarding the projects included
the necessity of sophisticated knowledge and equipment
and the inability of peers to contribute useful input to
other groups projects. Problems encountered were diffi-
culty focusing on a specific testable hypothesis, difficulty
observing experimental subjects, lack of experimental con-
trols, difficulty drawing conclusions from data, disagree-
ments among group members, and time constraints.
Groups whose data did not support their hypotheses, or
whose data collection proceeded more slowly, viewed their
project as less good than groups whose data supported
their hypotheses or who gathered data more quickly. The
educational benefits realized included recognition of the
importance of flexibility and patience when conducting an
investigation, awareness of biases in methodology and data
presentation, advantages of working in a group, and rec-
ognition of students’ abilities to pose and investigate eco-
logical questions. The class consensus was that field in-
vestigations are frustrating yet exciting and rewarding. The
students agreed unanimously that continual monitoring of
each student's progress, thoughts, and feelings was of
great benefit.
Traditional vs. computerized education: a student sur-
vey. J.G. SHIBER, Division of Biological Sciences, Pres-
tonsburg Community College, Prestonsburg, KY 41653.
A survey of 873 students at Prestonsburg Community
College (Prestonsburg, KY) reveals their opinions about
learning via computer technology (including TV, videocas-
settes, Internet) versus the traditional approach of attend-
ing live lectures, taking notes, using textbooks, etc. The
majority said that they prefer the traditional style of learn-
ing and that computer learning is more expensive, less
humane, and even less efficient. Most said that computers
will be essential in their careers but believe that ultimate
success in career and life is best derived from a traditional
education, which they define as more well rounded than
a computerized one. Teacher-student feedback, the ma-
jority agreed, is necessary for quality learning, and pro-
fessors lecturing in class can give more information,
through elaboration, than computers. Most students large-
ly attributed our present social problems and inability to
solve them to the fast rate of our technological develop-
ment. They were almost equally split, however, as to
whether or not computer use would eventually cause the
demise of society as we know it, or if we can live without
52 Transactions of the Kentucky Academy of Science 58(1)
computers. The majority believed that humans are not
organically (physically and mentally) evolved enough to be
completely dependent on computers. Nor did they see
exceptional, near-future benefits from computers. Indeed,
the trend towards computerization of everything bothers
the majority of those surveyed. Despite their preferences
and reservations, however, most students seemed resigned
to the inevitability that computerized education will even-
tually replace the traditional.
Undergraduate course in science ethics for biology and
chemistry majors: I. development and content, FRANK
H. WILBUR, Department of Biology, Asbury College,
Wilmore, KY 40390.
In 1986 selected ethical dilemmas relating to the health
professions were incorporated into PHP 201, Introduction
to the Health Professions, a course offered at Asbury Col-
lege (Wilmore, KY) and open to any student interested in
exploring career options in health-related fields. In 1990,
PHP 201 was dropped in favor of requiring a one-credit-
hour course in Science Ethics (BIO 372) for all senior
biology and chemistry majors. This course, which is now
required of all junior science majors, has been modified
each of the 6 years it has been taught. An introduction to
ethics and ethical theory, as well as discussions on world
views, values, character development, and the prevailing
cultural moral climate, precede class exercises focusing on
solving ethical situations and dilemmas in scientific re-
search, medicine, and ecology. While the ethical dilemmas
presented have varied somewhat from year to year, those
dealing with aspects of information disclosure (truth tell-
ing and informed consent), allocation of health care, pro-
fessional conduct in scientific research, and environmental
stewardship have been regularly included. Course evalu-
ations submitted by all students enrolled in the course
have played a major role in course modifications. The
most recent evaluations reveal considerable student sat-
isfaction and enthusiasm for course format and content.
Undergraduate course in science ethics for biology and
chemistry majors: I. world view and values, FRANK H.
WILBUR, Department of Biology, Asbury College, Wil-
more, KY 40390.
In 1990 a one-credit-hour course in Science Ethics (Bio
372) required for all biology and chemistry majors was
added to the science curriculum at Asbury College (Wil-
more, KY). It readily became apparent that many students
taking the course lacked the foundational skills necessary
to properly address ethical situations and dilemmas. They
had little or no knowledge of basic ethical theory, had not
sufficiently formulated their world view, and had not clar-
ified their personal value system. The course, as currently
offered, addresses these concerns. Twenty-five percent of
the course is devoted to basic ethical theory and assess-
ment of the present cultural climate. Students are pre-
sented with examples of different world views. These may
include a Biblical world view from the Christian and the
Jewish perspective, a postmodern world view, a relativist
world view, and/or a humanistic world view. Students are
required (1) to formulate and articulate their own world
view and to use it as a foundation in the ethical decision-
making process, (2) to identify the sources of their moral
beliefs, and (3) to articulate and defend their personal
value system. Teaching objectives include consciousness-
raising and sensitizing; and values analysis, clarifications,
and criticism. Student response to this approach, as de-
termined by comments and student evaluations, has been
most positive. Also, the ability of students to more intel-
ligently apply appropriate ethical decision-making pro-
cesses to solving ethical situations and resolving ethical
dilemmas has shown marked improvement.
ZOOLOGY & ENTOMOLOGY
Analyses of size and density of Tectarius muricatus (Lit-
torinidae) on San Salvador Island, Bahamas. MELINDA
L. CRAWFORD,* JACQUELINE M. GRANESE, and
RUDOLPH PRINS, Department of Biology, Western
Kentucky University, Bowling Green, KY 42101.
Samples of Tectarius muricatus collected from transects
at nine sites on San Salvador Island in the Bahamas were
counted and measured with regard to vertical zonation on
their respective rocky shores. Analyses were conducted
using a two-way ANOVA with no interaction effects. Size
differences were found among the different sites. How-
ever, the experimental results for T. muricatus contradict
those of many other littorinids in that no significant ver-
tical size gradient existed in the populations. Densities of
the snails within and between the sites were analyzed to
determine if vertical gradients existed. No significant dif-
ferences in density were found among the sites, but a sig-
nificant differential was exhibited between the lowest two
zones of the transects when averaged across all sites.
These results are discussed with respect to several possi-
ble environmental and physical factors that may have af-
fected the size and density distributions of T. muricatus.
Quantification of patterns such as distribution, density,
and size are necessary in preliminary studies of this vir-
tually unknown littorinid so that its superior physical tol-
erance may be investigated.
Characterization of snapping action of the alpheid
shrimp Alpheus heterochaelis. JARED FIALKOW* and
HONG Y. YAN, School of Biological Sciences, University
of Kentucky, Lexington, KY 40506.
Alpheus heterochaelis is a alpheid shrimp with a large
snapping claw. When this claw is forcibly closed, it results
in a loud snap sound. Since underwater sound propagates
fast over long distance, it has been hypothesized that the
snapping sound may be used for underwater acoustic
communication. This work is designed to test the “acous-
tic communication” hypothesis. Individual shrimp were
housed in a rectangular tank and a plastic rod was used
to prod to initiate snapping action. A hydrophone was
used to record the snapping sound. The recorded sounds
were analyzed with a computer-based SIGNAL analysis
system. Time, frequency signal, and spectrogram corre-
Abstracts, 1996 Annual Meeting 53
lations were compared between sounds of different indi-
viduals (males, females). Playback of the snapping sound
was also performed on 10 shrimp to initiate snapping ac-
tion. Snapping activity of two encountering individuals un-
der various conditions (light, dark, amputation of anten-
nular and antennal flagellum) was also videotaped and an-
alyzed. Sound similarity analyses showed no significant
difference in acoustic signatures made by different ani-
mals. Playback of the snapping sound failed to initiate any
snapping action. Contacts of antennular or antennal fla-
gellum of two encountering shrimp before snapping oc-
curred were observed in 256 snappings (out of a total of
268). For amputated shrimp physical contact of body parts
was required before snapping actions were initiated. The
results show that the snapping sound is not used for acous-
tic communication function. It is likely that the strong
underwater jet currents created by the snapping action are
used by the shrimp to ward off intruders. The snapping
sound produced is the by-product of the snapping action.
Supported by Howard Hughes Medical Institute and a
University of Kentucky Undergraduate Research Creative
Grant.
Comparison of a hybrid (Heliconia latispatha X imbri-
cata; Heliconiaceae) with its parent species with respect
to extrafloral nectaries. C. TONY R. HAMPTON* and
THOMAS C. RAMBO, Department of Biological Sci-
ences, Northern Kentucky University, Highland Heights,
KY 41099.
As part of on-going studies of the role of extrafloral
nectaries on buds of Heliconia latispatha and of the in-
sects visiting these nectaries, we studied a hybrid of H.
latispatha (which has nectaries) and H. imbricata (which
lacks nectaries). We studied a set of 10 H. latispatha, 10
H. imbricata, and 8 hybrids, all growing along the edge
of the landing strip at Estacién Sirena in Parque Nacional
de Corcovado, Costa Rica. We recorded the insect visitors
to the buds hourly from 0530 to 0930 and from 1330 to
1730, the time of most insect visitation, for 5 days. Heli-
conia latispatha had significantly more insects (primarily
ants) on the buds; H. imbricata and the hybrid had few
to no insects on the buds. The buds of H. latispatha and
the hybrid are similar in color, so color is not primarily
responsible for the attraction of insects. We hypothesize
that the insect attraction by H. latispatha is chemical, and
that the genes for this attraction must be recessive.
Comparison of two species of Heliconia (Heliconiaceae)
with respect to the presence of extrafloral nectaries. THO-
MAS C. RAMBO and C. TONY R. HAMPTON, De-
partment of Biological Sciences, Northern Kentucky Uni-
versity, Highland Heights, KY 41099.
In previous studies we have shown that the platanillo,
Heliconia latispatha, produces nectar on the outside of
bracts of terminal buds in an inflorescence. The nectar,
secreted onto the bud’s surface, is utilized by a variety of
insects, including ants, mosquitoes and other dipterans
(Dolichopodidae, Richardidae, Neriidae), crickets, roach-
es, lygaeid bugs, and wasps. To clarify the function of
these extrafloral nectaries we compared the insect visitors
to Heliconia latispatha with visitors to the buds of H. im-
bricata, a species growing close to H. latispatha. We re-
corded the visitors to 15 buds of each species hourly from
0530 to 0930 and from 1330 to 1730, the time of most
insect visitation, for 5 days. Heliconia imbricata attracted
significantly fewer insects than H. latispatha. In fact H.
imbricata attracted so few insects that we conclude that
it does not produce nectar on the buds. We will discuss
possible reasons for this difference and their implications
for determining the function of the extrafloral nectaries
on H. latispatha.
Effects of cannibalism on population structure of the
marine cave isopod Bahalana geracei from Lighthouse
Cave, San Salvador Island, Bahamas. JERRY H. CAR-
PENTER and RONALD D. BITNER,* Department of
Biological Sciences, Northern Kentucky University, High-
land Heights, KY 41099.
Cannibalism by Bahalana geracei was observed once in
Lighthouse Cave, in July 1995: a 11.0 mm long female
was eating a 6.5 mm long female. We performed experi-
ments to investigate factors influencing cannibalism in this
troglobitic species. From August to November 1996, two
to four specimens were placed together in small plastic
bowls, maintained without food in incubators at cave tem-
perature (26°C), and observed daily. We never witnessed
cannibalism while it was occurring in the lab; we did find
remains of six cannibalized victims (some still alive)—
translucent exoskeletons of cannibals revealed remains of
victims inside. The following cannibal/victim combinations
show the variety of sizes: (1) 3.2 mm second-instar manca
(baby)/2.7 mm first-instar manca (sex of mancas could not
be determined), (2) 4.2 mm fourth-instar juvenile female/
2.7 mm first-instar manca, (3) 8.2 mm female/8.2 mm fe-
male slightly damaged in another predator-prey experi-
ment, (4) 14.3 mm female and 7.4 mm female/7.5 mm,
(5) 14.3 mm female/7.4 mm female (same individuals as
above), and (6) 15.7 mm female and 13.9 mm female/12.4
mm female. These cannibalisms occurred after specimens
were together several days, except case #3, which oc-
curred within 1 day. In at least 10 additional cases, two to
four specimens of various sizes and sexes were together
>30 days without cannibalism. We conclude that canni-
balism in B. geracei is unpredictable. It is more likely
when size difference is large. Cannibalism in B. geracei
seems less common than in many surface-dwelling arthro-
pods (e.g., spiders and other marine isopods). Cannibalism
is reduced by B. geracei’s starvation resistance and slow
metabolism. Nevertheless, cannibalism is probably fre-
quent enough in this long-lived species to significantly af-
fect the population structure.
Female preferences for stimulatory male odors in the
prairie vole (Microtus ochrogaster). JILL H. KRUPER*
and TERRY L. DERTING, Department of Biological Sci-
ences, Murray State University, Murray, KY 42071.
54 Transactions of the Kentucky Academy of Science 58(1)
Physical contact with conspecific pheromones or urine
of males is necessary for reproductive stimulation of fe-
male prairie voles. We investigated whether female prairie
voles (Microtus ochrogaster) of different ages and repro-
ductive status prefer odors from males who are most likely
to stimulate female reproductive activity. We compared
odors from intact and castrated adult males because urine
from intact males is known to be a more effective stim-
ulant of female reproductive activity than is urine from
castrated males. Preferences of females for male odors
were determined using a Y-maze olfactometer or nesting
apparatus. At 7-10 d post-surgery, young anestrus females
significantly preferred the odors of intact males as com-
pared to the odors of castrated males. At 2 months post-
surgery, no significant preference for odors of intact or
castrated males was exhibited by older anestrus females.
Older females in estrus, however, tended to prefer the
odor of intact males. Young and old anestrus females
showed no significant preference for the odors of castrat-
ed or intact males during nesting-preference tests; how-
ever, young females in estrus tended to discriminate
against the odors of castrated males. Our results indicated
that young females, and those in estrus, were most likely
to discriminate among male odors, exhibiting less prefer-
ence for odors from castrated males than intact males.
Male odors indicative of reproductive potential may,
therefore, play a role in female mate-choice in prairie
voles depending upon the age and reproductive status of
females.
Impacts of dam construction on densities and distri-
butions of bald eagles, Haliaeetus leucocephalus, Accipit-
ridae) along the Ohio River shoreline between Paducah,
Kentucky, and Cairo, Illinois. VINCENT EVIN STAN-
FORD* and TERRY L. DERTING, Department of Bi-
ology, Murray State University, Murray, KY 42071.
Shoreline densities and distributions of wintering bald
eagles (Haliaeetus leucocephalus) were compared to ex-
amine the impacts of an ongoing lock-and-dam construc-
tion project on an Ohio River segment 65 km long be-
tween Paducah, KY, and Cairo, IL. Low-altitude flights
were conducted once weekly for 10 weeks from mid-De-
cember through mid-March in winters of 1987/1988 (pre-
dam construction) and 1994/1995 (ongoing dam construc-
tion). Densities and distributions of bald eagles per river
quarter-mile were recorded during the survey flights. Pro-
portional densities of bald eagles were compared between
the two survey periods in a 1- and a 6-mile interval with
the site of dam construction centered in each interval.
Proportional densities of bald eagles in the 1-mile interval,
during the 1994/1995 surveys, were significantly reduced
from proportional densities that occurred during the 1987/
1988 surveys (P = 0.0024). A marginally significant re-
duction in proportional densities of bald eagles occurred
in the 6-mile interval during the 1994/1995 surveys when
compared with the 1987/1988 surveys (P = 0.07). A mar-
ginally significant shift in %-mile distributions along the
entire segment also occurred between the two survey pe-
riods (P = 0.06). These results indicated that bald eagles
were avoiding the area of dam construction and that over-
all bald eagle distributions may be shifting away from the
area of dam construction. We recommend that further
studies be conducted to examine future impacts on bald-
eagle densities and distributions as construction proceeds
and to determine long-term impacts of dam placement on
bald eagles.
Nest-site selection and leaf-nest composition of Sciurus
carolinensis (Sciuridae) in continuous and isolated woods
in western Kentucky. JAMES S. ARMSTRONG* and
TERRY L. DERTING, Department of Biological Sci-
ences, Murray State University, Murray KY 42071.
Location and composition of grey squirrel (Sciurus car-
olinensis) leaf-nests were analyzed in three different forest
habitats: old (116-125 years) continuous-forest (3 sites, 2
ha each); young (85-92 years) continuous-forest (3 sites,
2 ha each); and isolated woodlot (106 years; 1 site, 3 ha).
The isolated woodlot contained a significantly higher num-
ber of small (DBH 3-25 cm) and total number of trees
than either continuous-forest habitat. No significant dif-
ferences in vegetational cover existed among habitats. The
isolated woodlot had significantly higher nest density (12.5
nests/ha) than the old or young continuous-forest habitats
(3.2 + 0.9 nests/ha and 3.3 + 0.3 nests/ha, respectively).
The density of grey squirrels in the isolated woodlot (13.2
squirrels/ha) was also significantly greater than squirrel
densities in old and young continuous-forest habitats (1.9
+ 0.6 and 1.8 + 1.0 squirrels/ha, respectively). A signifi-
cant positive correlation existed between leaf-nests/ha and
grey squirrels/ha across the seven study sites. Grey squir-
rels chose significantly fewer small size trees (DBH <20
cm) and significantly more medium size trees (DBH 20-
50 cm) for leaf-nest placement than expected based on
tree availability. Grey squirrels chose large size trees
(DBH > 50 cm) for leaf-nest placement in accordance
with tree availability. Tree species selected and the pre-
ferred DBH size for nest location were positively corre-
lated in the young continuous-forest and isolated woodlot
habitats. The composition of leaf-nests did not differ
among habitats although the leaf-nests in the old contin-
uous-forests had the greatest dry masses.
Guidelines for Contributors to the Transactions
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the editor.
John W. Thieret
Biological Sciences
Northern Kentucky University
Highland Heights, KY 41099
Phone: (606) 572-6390
FAX: (606) 572-5639
E-mail: thieretj}@nku.edu
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Transactions of the Kentucky Academy of Science 58(1)
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NEWS
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PUBLICATION
Wildflowers of Mammoth Cave National Park, by Randy Seymour, has just been published.
A soft-cover book of 254 pages, it illustrates by excellent color photographs about 400 wild-
flowers, each with accompanying text. Appendices show flowering times, where in the park the
various flowers have been seen, and the number of species noted by Seymour along each of
the park’s trails. Order from bookstores or directly from The University Press of Kentucky, 663
South Limestone Street, Lexington, KY 40508-4008; ISBN 0-8131-0898-5; price $17.95
(+$3.50 shipping).
CONTENTS
Relationships Among Habitat, Cover, and Eastern Cottontails (Sylvila-
gus floridanus) in Kentucky. William M. Giuliano, Charles L. Elliott,
and: Jeffery: D: Sole) oe Se ee, ONG SS OEN Oia Nae tae Ne tn atic aa
Separation of Spawning Habitat in the Sympatric Snubnose Darters Eth-
eostoma flavum and E. simoterum (Teleostei, Percidae). Jean C. Por-
Fer field esc cs ee a ee Sa oo ak Wade eee cata Soya R Us PAPA ay ae
Common Names of Vascular Plants Reported by C.S. Rafinesque in an
1819 Descriptive Outline of Four Vegetation Regions of Kentucky. Ron-
ald L. Stuckey and James S. Pringle .............0ccccccececcnceccecssecceccevece
Some Comments on Constantine Rafinesque’s 1819 Description of Bo-
tanical Regions in Kentucky. William S. Bryant ................0ccccseeceeeees
Effect of Light on Daily Emergence of Cercariae of the Trematodes Echi-
nostoma trivolvis (Echinostomatidae) and Cephalogonimus vesicau-
dus (Cephalogonimidae) from Natural Infections of the Snail Helisoma
trivolvis (Planorbidae) at Owsley Fork Reservoir, Kentucky. Ron Rosen,
Peter Blair, Jeff Ellington, and Jason Backus .................0cceccseeseees
Classifying Free Bieberbach Groups. Raymond F. Tennant ...............
DISTINGUISHED SCIENTIST AND OUTSTANDING TEACHER AWARDS
OG ee RSA cas caueceea ea cnn een ceee ns dere COU se oda ral oun ne ean ne Meg
NOTES
Human Myiasis in Kentucky Caused by Cuterebra sp. (Diptera: Oestri-
dae): Lee Hs Townsend oo ee ee a ee Wea a Ree ae
ABSTRACTS OF SOME PAPERS PRESENTED AT THE 1996 ANNUAL MEET-
ING OF THE KENTUCKY ACADEMY OF SCIENCE ..................cecesceceeceees
TRANSACTIONS.
vx | THE
wee NIT LICK Y
ACADEMY OF
toi NCE
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Volume 58
Number 2
September 1997
Official Publication of the Academy —
The Kentucky Academy of Science
Founded 8 May 1914
GoverninGc Boarp For 1997
Executive COMMITTEE
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MEMBERS, GOVERNING BOARD
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Wimberly C. Royster 1997
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7 tne
TRANSACTIONS of the
KENTUCKY
ACADEMY of SCIENCE
September 1997
Volume 58
Number 2
Trans. Ky. Acad. Sci. 58(2):55-59. 1997.
Continuing Decline in the Freshwater Unionid (Bivalvia: Unionidae)
Fauna in the Cumberland River Downstream from Cumberland Falls,
Kentucky
Ronald R. Cicerello and Ellis L. Laudermilk
Kentucky State Nature Preserves Commission, 801 Schenkel Lane,
Frankfort, Kentucky 40601
ABSTRACT
The freshwater unionids in the Cumberland River downstream from Cumberland Falls in southeastern
Kentucky were sampled to determine the status of the fauna relative to previous studies. A comparison of
collections by Wilson and Clark in 1910-1911, Neel and Allen in 1947-1949, and Stansbery in 1961 revealed
the loss of 6 of the 22 species known from the site and reduced abundance in several others. We found only
10 live species, all less abundant than during previous studies, and no shells. The continuing decline in the
fauna is attributable to habitat degradation and loss associated with coal, mining, general watershed devel-
opment, and impoundment of the river to create Lake Cumberland.
INTRODUCTION
Long-term monitoring can provide status
and trend information needed for conserva-
tion of freshwater unionids. To determine
changes in the Cumberland River fauna down-
stream from Cumberland Falls in southeast-
ern Kentucky following the creation of Lake
Cumberland, Stansbery (1969) sampled the
unionids and compared his findings to those
of Wilson and Clark (1914) and Neel and Al-
len (1964). Wilson and Clark (1914) found 20
species, 10 of which were common or abun-
dant, at this site in 1910-1911 while evaluating
the Cumberland River as a source of unionids
to replace upper Mississippi River populations
overharvested for button manufacturing and
pearls. During 1947-1949, just prior to the
closure of Wolf Creek Dam and the formation
of Lake Cumberland, Neel and Allen (1964)
encountered 15 species at the site, 13 of which
were common or abundant.
Stansbery (1969) collected below the falls in
1961 and noted that changes in faunal com-
position prior to Neel and Allen (1964) were
50
attributable to almost 40 years of increased ac-
ids from coal mining and washing above the
falls. Changes after 1949 may have resulted
from continuing impacts from coal mining op-
erations and the influence of Lake Cumber-
land. By 1947-1949, Cyclonaias tuberculata,
Ellipsaria lineolata, Obliquaria reflexa, and
Ptychobranchus subtentum were lost from the
site, but Villosa iris had been found. Between
1949 and 1961, Lampsilis cardium and Trun-
cilla truncata were lost, 6 of the 16 taxa found
by Stansbery (1969) had decreased in abun-
dance, and the remainder were stable or had
increased in number. We resampled this site
to determine the present status of the fauna.
STUDY AREA
Cumberland Falls, a 17-m-high barrier to
the upstream movement of aquatic organisms
(Burr and Warren 1986; McGrain 1966), is lo-
cated at river km 904.9 in McCreary and
Whitley counties in southeastern Kentucky
(Figure 1). Land use in the 5120 km? water-
shed (Bower and Jackson 1981) is ca. 84% for-
56 Transactions of the Kentucky Academy of Science 58(2)
River km
898.6
Dog
Slaughter
Creek
—> Z
McCreary County
Whitley County
Figure 1. Cumberland River study area downstream
from Cumberland Falls, McCreary and Whitley counties,
Eagle
Creek
Cumberland Falls
~ River km 904.9
Kentucky. Inset shows study area location in the state.
Shaded areas were searched for unionids.
est, 13% agriculture, 2.5% mining, and 0.5%
urban and developed areas (Mayes, Sudderth,
and Etheredge 1975). Downstream from the
falls, the river flows through a forested and
boulder-lined gorge and is impounded by the
backwaters of Lake Cumberland at approxi-
mate river km 898.6, depending on reservoir
pool elevation. The Cumberland River was
impounded at river km 742 in 1950 to create
Lake Cumberland. At low flow, river width
ranges from ca. 5 to 24 m in riffles and pools,
respectively, and depth exceeds 5 m in pools
and 1 m in riffles. Pools and riffles are com-
posed of large boulders, cobbles, and exten-
sive deposits of coal fines and sand intermixed
with lesser quantities of gravel, pebbles, coal,
and trash in eddies and areas sheltered from
swift current. During low flow, huge deposits
of sand and coal fines are exposed in the
plunge basin below the falls. Mean discharge
on the sampling dates was 7.4 m*/sec, consid-
erably less than the mean of 26.6 m*/sec for
July-September, when streamflow is at or near
the annual low, and the mean annual discharge
of 89.7 m*/sec (USGS 1993). Water quality is
improving, but the upper Cumberland River
continues to be impacted by pollutants asso-
ciated with coal mining (e.g., acid mine drain-
age, silt, and metals), domestic waste, highway
construction, and poor land use (Harker et al.
1980; Kentucky Division of Water 1994a,
1994b).
MATERIALS AND METHODS
In 1987, 1989, and twice in 1993, we ex-
amined a 0.5-km-long river segment from the
base of Cumberland Falls to Eagle Creek for
unionids (Figure 1). We made additional col-
lections 0.5 km downstream from Eagle Creek
and near the mouth of Dog Slaughter Creek
to further gauge the status of the fauna. Dur-
ing each visit, one or two persons searched 2
or more hours for shells and live specimens by
snorkeling and with waterscopes, and the
shoreline was examined for shells. Nomencla-
ture follows Hoeh (1990) and Turgeon et al.
(1988).
RESULTS AND DISCUSSION
We found a total of 10 species, a loss of six
taxa when compared to Stansbery’s (1969) re-
sults and an overall 54% reduction in the fau-
na compared to earlier studies (Table 1). With
the exception of Elliptio crassidens, all taxa
were less abundant than in 1961. Among the
taxa lost are Villosa trabalis, a USFWS (1994)
endangered species, and Lampsilis ovata, Pty-
chobranchus subtentum, and Villosa lienosa,
which are rare in Kentucky (KSNPC 1996).
Others that have declined or have been lost
(e.g., Actinonaias ligamentina, Elliptio dilata-
ta, Lampsilis cardium, Potamilus alatus,
Quadrula pustulosa, Tritogonia verrucosa)
have broad habitat requirements (Gordon and
Layzer 1989) and often are dominant mem-
bers of unionid communities (Houslet 1996;
Smathers 1990). Corbicula fluminea, an intro-
duced exotic bivalve, was uncommon.
Observations by Wilson and Clark (1914)
and Neel and Allen (1964) provide anecdotal
evidence of the decline in unionid abundance
at the falls. Wilson and Clark (1914) reported
that unionids were “usually found crowded
about the base of large rocks” where they
were “easily accessible to their enemies, es-
pecially during low water, and many of them
are killed by muskrats, raccoons, mink, and oc-
casional otter.” Neel and Allen (1964) found
Cumberland Unionids—Cicerello and Laudermilk aa
Table 1.
in four studies: 1910-1911, 1947-1949, 1961, 1987-1993.
1910-1911!
Species
Actinonaias ligamentina (Lamarck) 194
Actinonaias pectorosa (Conrad) We
Cyclonaias tuberculata (Rafinesque) 2
Ellipsaria lineolata (Rafinesque) P
Elliptio crassidens (Lamarck) 57
Elliptio dilatata (Rafinesque) 122
Lampsilis cardium (Rafinesque) 0
Lampsilis fasciola Rafinesque 16
Lampsilis ovata (Say)? 49
Lasmigona costata (Rafinesque) 4]
Ligumia recta (Lamarck) 8
Medionidus conradicus (Lea) P
Obliquaria reflexa Rafinesque 8
Potamilus alatus (Say) 8
Ptychobranchus fasciolaris (Rafinesque) 81
Ptychobranchus subtentum (Say)° 12
Quadrula pustulosa (Lea) 49
Tritogonia verrucosa (Rafinesque) 32
Truncilla truncata Rafinesque 16
Villosa iris (Lea) 0
Villosa lienosa (Conrad)* P
Villosa trabalis (Conrad)* 4]
Total species 20
Total individuals 810
Freshwater unionids collected from the Cumberland River downstream from Cumberland Falls, Kentucky,
1947-19497 1961 1987-1993"
Number collected
A 39 2
A 161 >50
0 0 0
0 0 0
C (2) 2
A 113 i
A 0 0
C 20 0
0 10 0
0 1(?) 2)
A it 1
A 154 0
0 0 0
E 28 5
A 35 5
0 0 0
A 122 10
A 75 4
R 0 0
R Oi 0
0) 9 0
A vi 0
15 16 10
a= 810 ca. 88
' Wilson and Clark (1914); * Neel and Allen (1964); > Stansbery (1969); * maximum number observed alive on any sampling date during the present study;
> USFWS (1994) and/or KSNPC (1996) listed species. A = abundant; C = common; R = rare; P = present; 0 = none taken; (2) = subfossil remains only,
“great numbers of mussels, many of which
were quite large and very old” in sand pockets
among large stones and slabs. We found a con-
centration of live unionids only in a small area
near the mouth of Eagle Creek. No shells de-
posited by floodwater or left by predators
were found along the shoreline.
In the river 0.5 km downstream from Eagle
Creek and near the mouth of Dog Slaughter
Creek, we found limited numbers (in order of
abundance) of only Actinonaias pectorosa, Tri-
togonia verrucosa, Actinonaias ligamentina,
Elliptio dilatata, Potamilus alatus, Ptycho-
branchus fasciolaris, and Quadrula pustulosa.
These sites are within the river segment ex-
tending from “Anvil Shoals,” 1.6 km below the
falls, to Burnside, Kentucky, that mussel fish-
erman reported was full of mussels (Wilson
and Clark 1914).
The fauna of this last remaining free-flow-
ing segment of the Cumberland River below
the falls in southeastern Kentucky is now com-
prised largely of species with broad environ-
mental tolerances (Dennis 1984; Gordon and
Layzer 1989). The change in the fauna is at-
tributable to pollutants associated with coal
mining and to the impoundment of the river
(Stansbery 1969), and general watershed de-
velopment (e.g., road building, channelization,
urbanization). Recolonization is precluded by
Lake Cumberland, which isolates remnant
unionid populations in Buck Creek (Schuster,
Butler, and Stansbery 1989), the Rockcastle
River (Cicerello unpubl. data; Thompson
1985), and the Big South Fork Cumberland
River (Bakaletz 1991), and which acts as a bar-
rier to the movement of host fishes. Direct
tributaries to the area (e.g., Eagle and Dog
Slaughter creeks) do not support unionid pop-
ulations, and the Cumberland River above
Cumberland Falls supports a depauperate fau-
na comprised of 11 species that generally are
uncommon in the basin (Cicerello unpubl.
data; Cicerello, Warren, and Schuster 1991).
The unionid community in the river below
Cumberland Falls is the last and best remain-
ing vestige of the diverse fauna that histori-
cally inhabited the mainstem Cumberland
River in southeastern Kentucky. Downstream
from Wolf Creek Dam, fluctuating levels of
58 Transactions of the Kentucky Academy of Science 58(2)
cold, turbulent water released from Lake
Cumberland preclude unionid reproduction,
and only Cumberlandia monodonta Say and
Cyclonaias tuberculata persist (Cicerello per-
sonal observation; Miller, Rhodes, and Tippit
1984). Lake Cumberland backwaters in the
lower Big South Fork Cumberland and Rock-
castle rivers, and the mainstem are heavily silt-
ed and have yielded Leptodea fragilis Rafin-
esque, Potamilus ohiensis Rafinesque, Pygan-
odon grandis Say, and Utterbackia imbecillis
Say. Only Potamilus alatus and P. ohiensis in-
habit the lower embayed segment of Buck
Creek (Schuster, Butler, and Stansbery 1989).
Perhaps 20 (31%) of the 65 taxa that histori-
cally inhabited the mainstem Cumberland
River remain (Cicerello unpubl. data).
Monitoring in the river downstream from
Cumberland Falls in the future will provide
additional information about the impact of
Lake Cumberland and watershed land use on
the unionid fauna. However, the survival of
Kentucky's remnant Cumberland River union-
id community, which includes members of the
unique Cumberlandian fauna (Ortmann 1924)
(e.g., Epioblasma brevidens, E. capsaeformis,
Ptychobranchus subtentum, and Villosa tra-
balis) found nowhere else in the state, is de-
pendent upon the protection of Buck Creek,
the Rockcastle River, and the Big South Fork
Cumberland River.
ACKNOWLEDGMENTS
We acknowledge the field assistance of Ste-
phen McMurray (Eastern Kentucky Universi-
ty). G.A. Schuster (Eastern Kentucky Univer-
sity) critically reviewed the manuscript and
provided valuable suggestions. This study was
supported in part by the Kentucky Depart-
ment for Surface Mining Reclamation and En-
forcement, Frankfort, Kentucky.
LITERATURE CITED
Bakaletz, S. 1991. Mussel survey of the Big South Fork
National River Recreation Area. Master's thesis, Ten-
nessee Technological University, Cookeville, TN.
Bower, D.E., and W.H. Jackson. 1981. Drainage areas of
streams at selected locations in Kentucky. Open File
Report 81-61. Geological Survey, U. S. Department of
the Interior.
Burr, B.M., and M.L. Warren Jr. 1986. A distributional
atlas of Kentucky fishes. Kentucky State Nature Pre-
serves Comm. Sci. Tech. Ser. 4.
Cicerello, R.R., M.L. Warren Jr., and G.A. Schuster. 1991.
A distributional checklist of the freshwater unionids (Bi-
valvia: Unionoidea) of Kentucky. Am. Malacol. Bull. 8:
113-129.
Dennis, $.D. 1984. Distributional analysis of the fresh-
water mussel fauna of the Tennessee River system, with
special reference to possible limiting effects of siltation.
Ph. D. dissertation, Virginia Polytechnic Institute and
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histories and ecological relationships. U. S. Fish and
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Call, G.J. Fallo, and P. Wigley. 1980. Aquatic biota and
water quality survey of the upper Cumberland River
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assemblage in Horse Lick Creek, Kentucky. Master's
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TN.
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the naiad fauna of the Cumberland River below Lake
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the upper Cumberland basin before its impoundment.
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Acad. Sci. 50:79-85.
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Bivalvia) community in the Licking River, at Moores
Ferry, Kentucky. Master’s thesis, Eastern Kentucky
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Cumberland Unionids—Cicerello and Laudermilk 59
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River system, Kentucky (Bivalvia: Unionidae). Master's
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Document 781:1-63.
Trans. Ky. Acad. Sci. 58(2):60-66. 1997.
Characterization of Tall Fescue Plantlets Regenerated from Cultured
Panicle Segments/Anthers for Meiotic, Isozyme, and DNA Changes
Georgia C. Eizenga
USDA-ARS-MSA, Department of Agronomy, University of Kentucky,
Lexington, KY 40546-0091!
and
Timothy D. Phillips
Department of Agronomy, University of Kentucky,
Lexington, KY 40546-0091
ABSTRACT
Previous research identified four tall fescue (Festuca arundinacea Schreb.) genotypes (K8, K16, K25, K27)
out of 20 in “Kentucky 31° that regenerated plantlets from inflorescence culture. The objectives of our study
were to determine whether plantlets could be obtained from these same genotypes using anthers as the
explant and to characterize the anther-derived plantlets for ploidy, meiotic, isozyme, and DNA changes.
Panicle segments of each genotype were cultured on Schenk and Hildebrandt medium with 2.0 or 4.0
me/liter 2,4-dichlorophenoxyacetic acid in three separate studies. After 2 weeks, florets opened; then anthers
were removed from the florets, equally divided among genotypes and hormone treatments, and cultured, 25
anthers per petri plate, on the same medium as the original panicle segment. Variables measured were callus
growth, root growth, albino shoot regeneration, green shoot regeneration, and green plantlets transplanted
into soil. Significantly more green plantlets regenerated from genotype K25 in two of the three studies.
Thirty-two K25 and two K27 regenerants transplanted to the field were hexaploid (2n=6x=42) with normal
chromosome pairing and morphologically like non-cultured tall fescue. Isozyme and RAPD profiles were
identical to the maternal plants, indicating the regenerants were derived from somatic tissue, rather than
being doubled haploids.
INTRODUCTION
Tall fescue (Festuca arundinacea Schreb.,
2n=6x=42) is an open-pollinated, perennial,
cool-season pasture and turf grass. Of the ca.
7.0 million acres of grassland in Kentucky, 5.5
million acres have been planted to “Kentucky
31° tall fescue (Pedersen and Lacefield 1989;
Siegel et al. 1984). Kentucky 31 has poor for-
age quality, and nearly 80% of the tall fescue
fields in Kentucky are infected with an endo-
phytic fungus (Neot yphodium coenophialum
(Morgan-Jones & Gams) Glenn, Bacon, &
Hanlin). A diet of infected tall fescue may be
detrimental to animals. Efforts at the Univer-
sity of Kentucky to improve the forage quality
of tall fescue would be enhanced if non-in-
fected tall fescue plants were identified that
could regenerate haploid plantlets poets
from anthers.
' To whom correspondence should be addressed. Present
address: USDA-ARS-SPA, NRGEEC, P.O. Box 287, 2900
Hwy 130 East, Stuttgart, AR 72160.
60
Earlier research by Kasperbauer et al.
(1980) reported that tall fescue haploids were
obtained from inflorescence (panicle) culture;
to our knowledge these plants are no longer
extant. Using a similar technique, Eizenga and
Dahleen (1990) identified four genotypes
(plants) from 20 genotypes in non-infected
Kentucky 31 tall fescue that regenerated
plantlets from inflorescence culture. All 198
inflorescence-derived plantlets were morpho-
logically normal. Meiotic and isozyme analyses
of the 95 regenerants showed them to have 42
chromosomes, normal chromosome pairing,
and the same isozyme banding patterns as the
parent plants. The isozyme banding patterns
suggested that the regenerants were derived
from somatic tissue.
Bohanec et al. (1995) used differences in
RAPD (random amplified polymorphic DNA)
profiles among regenerants from onion (Alli-
um cepa L.) ovules to identify different ploidy
levels. Also, RAPD profiles have been em-
ployed to determine the phylogenetic relation-
Tall Fescue Plantlets—Eizenga and Phillips 61
ships between species of Festuca and Lolium
(Stammers et al. 1995).
The objectives of our study were (1) to de-
termine whether regeneration would occur
from the four tall fescue genotypes identified
by Eizenga and Dahleen (1990) using anthers
as the explant and (2) to characterize the re-
generated plantlets for possible ploidy, mei-
otic, isozyme, and RAPD variants.
MATERIALS AND METHODS
The four Kentucky 31 tall fescue genotypes
(K8, K16, K25, K27) we used were selected
from 20 individual plants (genotypes) as _re-
generating plantlets from cultured tall fescue
inflorescences (Eizenga and Dahleen 1990).
These genotypes were evaluated in three dif-
ferent studies.
Previously, the procedures used to harvest,
decontaminate, and culture the panicles were
reported (Eizenga and Dahleen 1990). Cul-
tures of panicle segments were maintained at
25 + 2° C in the dark for 2 weeks, during
which time the florets began to open. After 2
weeks the anthers were removed from the two
proximal florets of each spikelet and placed on
the same medium used to culture the panicle
segments. Twenty-five anthers were placed on
each plate. The cultured anthers were main-
tained at 25 + 2°C in a 12-hour photoperiod
using 40 W cool-white fluorescent lamps (ca.
20 wE/m?/s at the culture surface).
Both the panicle segments and anthers were
cultured on Schenk and Hildebrandt basal salt
medium (Schenk and Hildebrandt 1976) with
75 g/liter sucrose, 6 g/liter agar, and 2 or 4
mg/liter 2,4-D (2, 4-dichlorophenoxyacetic
acid). In the second study (trial II), kinetin (1
mg/liter) was included to determine its effect
on plantlet regeneration. A total of 4800, 5400,
and 9600 anthers were cultured in the three
studies, trial I, trial HI, and trial III, respec-
tively; these anthers were equally divided
among the genotypes and hormone treat-
ments. The cultured anthers were observed
every 3 to 4 weeks for callus, root, albino
shoot, and green shoot growth over the next
3 months. This growth was scored as the total
number of anthers per plate showing evidence
of callus, root, and/or shoot growth. The total
number of albino and green plants regener-
ated from the cultured anthers was recorded.
All green plantlets were placed on a growth
medium with no auxin and half-strength major
salts prior to transfer to Jiffy-7 peat pellets (Jif-
fy Products, Batavia, Illinois). Subsequently,
the green plantlets were transplanted to the
field for vernalization and to obtain inflores-
cences for meiotic analyses.
Data for cultured anthers were analyzed us-
ing a3 X 4 X 2 factorial design with trial,
genotype, and 2,4-D concentration (conc.) be-
ing the factors. (Each petri plate was a unit of
replication.) This analysis showed that the trial
xX 2,4-D conc., genotype X 2,4-D conc., and
trial X genotype X 2,4-D conc. interactions
were non-significant. Only the trial < geno-
type interaction was significant, so the data
were reanalyzed with only the trial < geno-
type interaction included to determine the
least-square means. Significance was reported
at the 0.05 level using a square root transfor-
mation of the least-square means. Similarly,
trial II data were analyzed using a 4 X 2 X 2
factorial design with genotype, 2,4-D conc.,
and kinetin concentration being the factors.
Mitotic analyses followed the procedures of
Kasperbauer et al. (1980). Meiotic analyses,
pollen stainability, and enzyme assays for acid
phosphatase (ACPH), glutamate oxaloacetate
transaminase (GOT), malate dehydrogenase
(MDH), 6-phosphogluconate dehydrogenase
(6-PGD), and phosphoglucoisomerase (PGI)
were according to Dahleen and Eizenga
(1990). The alcohol dehydrogenase (ADH) as-
say was modified according to Eizenga and
Dahleen (1990). The chiasma frequency was
calculated as follows: < 100.
Genomic DNA was extracted from lyophi-
lized ground leaf tissue with CTAB buffer (0.1
M Tris at pH 7.5, 0.7 M NaCl, 0.01 M EDTA
at pH 8.0, 0.14 M B-mercaptoethanol, and
0.03 M mixed alkytrimethylammonium bro-
mide [Saghai-Maroof et al. 1984]), then ex-
tracted against 1 volume phenol and 1 volume
of chloroform-isoamy] alcohol (24:1), followed
by ethanol precipitation and a series of three
ethanol rinses (15 mM ammonium acetate in
80%; 70%; 70%). The DNA pellet was dis-
solved in TE (10 mM Tris at pH 8.0, 1 mM
EDTA), proteinase K and 1%(w/v)SDS, and
further purified with a phenol-chloroform ex-
traction.
RAPD analysis was modified from Williams
et al. (1990) for use on a Perkin-Elmer DNA
Thermocycler 480 (PE Applied Biosystems,
62 Transactions of the Kentucky Academy of Science 58(2)
Figure 1. Tall fescue regenerants. Green plantlet regenerating from anther callus. Emerging roots also are visible.
Foster City, CA). The reaction consisted of 25
ng sample DNA, 200 4M of each dNTP
(Boehringer Mannheim, Indianapolis, IN),
37.5 ng decamer primer (Operon Technolo-
gies, Inc., Alameda, CA), 1.0 U Taq polymer-
ase (Boehringer Mannheim, Indianapolis, IN),
and buffer (10 mM Tris pH 8.3, 50 mM KCI,
1.5 mM MgCl.) in 25.0 wL volume. The ther-
mocycler was programmed for six cycles of
94°C for 1 mins 35°C for 1min:72°C for’ 2
min, followed by 35 cycles of 94°C for 1 min,
40°C for 1 min, 72°C for 2 min ending with a
6 min extension period at 72°C. Amplification
DNA samples were separated by loading 12
wL of each sample, electrophoresed on 1.2%
agarose gels stained with 0.06 pg/wL ethidium
bromide, run with 1 X TBE buffer (89 mM
Tris-base, 89 mM borate, 2 mL EDTA, pH
8.0) at 100 mV for 2 h, washed, and photo-
graphed under UV light. The marker was a 1
Kb DNA Ladder (Gibco-BRL, Gaithersburg,
MD).
One hundred twenty-two random decamer
primers were initially screened with the ma-
ternal plants (K25, K27) to identify those
primers that detected different polymor-
phisms among the maternal plants. The fol-
lowing nine primers identified differences and
were selected to screen the entire set\ of 34
samples: OPES OPFI0. OB]15; OFI19;
OPK19, OPTO1, OPT20, OPAM18, and
OPAM19. Two independent runs were per-
formed for each primer; photographs were
scanned for band presence/absence in the ma-
ternal plants but band absence/presence in the
regenerants.
RESULTS
Both green (Figure 1) and albino plantlets
were regenerated from cultured anthers. For-
ty green plants (4 from K16, 33 from K25, 3
from K27) from three studies were regener-
ated and transplanted to soil. Thirty-four (32
from K25: 2 from K27) were vernalized in the
field and induced to flower. The panicles of
the regenerants were morphologically like
non-cultured tall fescue.
Statistical analysis of the culture data indi-
cated that the 2,4-D concentrations tested had
no significant effect on the number of anthers
producing callus, roots, and/or shoots. Similar-
ly, analysis of trial II data indicated that kinetin
did not significantly affect the expression of
these same variables. The lack of response to
kinetin was expected based on other research
(Kasperbauer 1990).
Genotypes K16 and K27 gave significantly
more callus growth than K8 and K25 in trial
I (Table 1). In trial UH, significant differences
were noted in callus growth with K27 giving
the most callus followed by K25 and K16.
In trial I, genotype K16 regenerated signif-
icantly more albino shoots (zero to 10 anthers
produced shoots per petri plate) than other
genotypes. In subsequent studies there were
no significant differences among genotypes.
The number of anthers per petri plate pro-
ducing albino shoots ranged from zero to
Tall Fescue Plantlets—Eizenga and Phillips 63
Table 1. Comparison of tall fescue anther culture response variables (callus growth, root regeneration, albino shoot
regeneration, green shoot regeneration, green plantlets transplanted to soil) over three different studies, and four tall
fescue genotypes.
Response variable
Experiment Genotype Callus growth Root growth Albino shoots Green shoots Green plants
% anthers responding per 100 anthers
Trial I K8 0.25 ab* 0.00 a* 0.25 be* 0.00 a* 0.00 a*
K16 0.67 a 0.33 a alii! 0.33 a 0.33 a
K25 0.00 b 0.00 a 0.00 c 0.00 a 0.00 a
K27 0.67 a 0.00 a 0.75 b 0.00 a 0.00 a
Trial IT K16 0.33 b 0.00 a 0.17 a 0.00 b 0.00 b
K25 0.67 ab 0.00 a 0.00 a 0.44 a 0.67 a
K27 0.83 a 0.17 a O.lla 0.11 ab 0.17 ab
Trial II K8& 0.00 a 0.00 b 0.00 a 0.00 b 0.00 b
K16 0.04 a 0.00 b 0.00 a 0.00 b 0.00 b
K25 0.33 a 0.67 a 0.04 a lL13a 0.88 a
K27 0.16 a 0.00 b 0.04 a 0.00 b 0.00 b
* Significant at the 0.05 level using a square root transformation of the least-square means.
€
ae : a
_ #* :
25 27 25&27 25 27 25 27 25 27 25 27
ACPH ADH GOT MDH 6-PGD_ PGI
Figure 2. Tall fescue regenerants. Banding patterns of the maternal plants (K25, K27) for ACPH, ADH, GOT, MDH,
6-PGD, and PGI. Banding patterns of the regenerants were identical to those of the maternal plants, indicating that
the regenerants were derived from somatic tissue rather than from microspores via spontaneous doubling.
64 Transactions of the Kentucky Academy of Science 58(2)
K25
Regenerants
=)
M1234
|
1636 bp =
Materal K27
plants Regenerants
Sima:
Figure 3. Tall fescue regenerants. RAPD profiles of maternal plants (K25, K27) and their respective regenerants. The
identical patterns indicate that the regenerants were derived from somatic tissue rather than from microspores. Genomic
DNA was amplified using the primer OPF10. K25 is distinguished from K27 by an additional band at the 1636 bp
fragment. The size marker is a 1-Kb DNA ladder (Gibco-BRL).
three. Mitotic analyses of the albino plantlets
obtained from the shoots were unsuccessful.
For green shoot regeneration, K16 ranked
the highest in trial I, and K25 was the highest
in trials IT and III. This ranking was the same
for green plantlets transplanted to soil and
nearly the same for root growth. In trial I, four
green plants were regenerated from K16, but
all four died when transplanted to soil in the
greenhouse. In trial II, 14 of the 15 green
plants transplanted to soil survived. Twelve
were from K25 and two from KQ27. In trial III,
20 of the 21 green plantlets regenerated from
K25 survived transplanting to soil. One of the
20 plantlets produced some green and white-
striped tillers after being transplanted to soil.
All 34 green plantlets that survived trans-
planting to soil (32 from K25; 2 from \K27),
had 42 chromosomes and normal chromo-
some pairing at metaphase I. The overall
mean chromosome pairing of the regenerants
was 0.17 univalents, 3.03 rod bivalents, and
17.88 ring bivalents with a mean chiasma fre-
quency of 92.4%. The mean pollen stainability
of these same plants was 79.3%, which is with-
in the range normally found in tall fescue.
These meiotic analyses and pollen stainability
were similar to those previously reported for
the maternal plants and inflorescence-derived
regenerants (Eizenga and Dahleen 1990).
The maternal plants (K25, K27) had differ-
ent banding patterns for five of the six en-
zymes tested (Figure 2). The banding patterns
were evidence of different alleles being pres-
ent in the maternal plants for the isozymes
ACPH-1, GOT-2, GOT-3, 6-PGD-1 and
PGI-2. No variant patterns were identified
among the regenerants.
For the nine primers evaluated, differences
in band presence/absence were noted in
RAPD banding patterns between the maternal
plants (K25, K27). Comparing these patterns
to the regenerants, no variant patterns, band
absence/presence, were found as illustrated
with the OPF10 primer (Figure 3). Thus, pan-
icle morphology and meiotic, isozyme, and
Tall Fescue Plantlets—Eizenga and Phillips 65
RAPD analyses indicated there was no cul-
ture-induced variation for these parameters.
DISCUSSION
The fact that isozyme and RAPD banding
patterns of the regenerants were the same as
the parents implies that regeneration was from
somatic tissue, possibly anther wall, rather
than from microspores via spontaneous dou-
bling. If regeneration had been from micro-
spores, some of the isozyme phenotypes
should have differed from the parents similar
to the segregation of isozyme markers ob-
served for PGI-2 and GOT-2 in androgenetic
progenies of Lolium perenne L. (Hayward et
al. 1990). Also, if the plantlets had been dou-
bled haploids (via spontaneous doubling of the
chromosomes in the microspore), most likely
a maximum of three different alleles would
have been present because tall fescue is a hex-
aploid species. The PGI-2 banding patterns of
the K25 and K27 regenerants require four al-
leles (Eizenga and Comelius 1991). In addi-
tion, the tall fescue haploids obtained by Kas-
perbauer et al. (1980) had fewer bands for
ACPH-1 and PGI-2 than normally found in
hexaploid tall fescue (Eizenga, unpublished
data). Similarly, regenerants for onion ovules
showed differences in RAPD profiles among
regenerants of different ploidy levels, but
identical profiles were found among regener-
ants of the same ploidy (Bohanec et al. 1995).
Screening of 764 beet (Beta vulgaris L.) re-
generants for variant isozyme alleles and 60
regenerants for variant restriction fragments
using RFLP (restriction fragment length poly-
morphisms) analysis suggested a percentage of
ca. 0.05% variant isozyme alleles and 0.1%
variant restriction fragments (Sabir et al.
1992). Yamamoto et al. (1994) found that
RAPD profiles could differentiate 35 rice
(Oryza sativa L.) varieties, but detection of
minor genetic alterations among somaclonal
variants or mutants and their maternal varie-
ties was not feasible. These studies suggest
that neither isozyme nor random PCR tech-
niques can differentiate among the 34 tall fes-
cue regenerants. An additional limitation is the
small sample size and the limited number of
primers used.
Two anther culture studies of perennial rye-
grass (Lolium perenne) genotypes demonstrat-
ed a significant genotypic effect on regenera-
tion percentage from microspore-derived em-
bryos (Madsen et al. 1995; Opsahl-Ferstad et
al. 1994). Both studies suggested that regen-
erability was under a relatively simple genetic
control. Tall fescue is closely related to peren-
nial ryegrass (Sleper 1985); thus, tissue culture
regeneration in tall fescue probably is under a
simple genetic control, similar to that de-
scribed in perennial ryegrass. Genes for in-
creasing regenerability from anther culture
and green plant frequency have been mapped
to chromosome arms in wheat-rye addition
lines (Martinez et al. 1994) and to chromo-
somal segments in maize (Zea mays L.) using
RFLP markers (Armstrong et al. 1992).
No significant correlation was found be-
tween regenerability from cultured anthers
and immature embryos of doubled haploid
wheat (Triticum aestivum L.) lines, which sug-
gested separate genetic control of regenera-
tion (Agache et al. 1988). Similarly, lack of a
significant correlation among regenerability
from leaf discs, anthers, and protoplasts was
reported in Solanum phureja Juz. & Buk.
(Taylor and Veilleux 1992). Possibly in tall fes-
cue there exists a separate genetic control for
regenerability from somatic tissues and micro-
spores. This would explain why only somatic
tissues responded in our study.
By comparison, anther culture of regener-
ative lines of sorghum (Sorghum bicolor (L.)
Moench.) and alfalfa (Medicago sativa L.)
have been unsuccessful in producing haploid
plants (Wen et al. 1991). Alternatively, as sug-
gested in the studies of sorghum (Wen et al.
1991), it is possible that the tall fescue albino
plantlets produced in this study were haploid.
In conclusion, our research showed that se-
lection for regenerability from cultured inflo-
rescences was a good indicator of regenera-
bility from cultured anthers. The regeneration
of green plantlets apparently was from somatic
tissues rather than from microspores, although
it is possible that the albino plantlets were
haploid. Future studies designed to obtain
green tall fescue haploids will need to focus
on regenerability from microspores and alter-
nate methods of obtaining haploid plants.
ACKNOWLEDGMENTS
The authors acknowledge the excellent
technical assistance of Mrs. Etta Mae Thacker
and the statistical advice of Dr. Paul L. Cor-
66 Transactions of the Kentucky Academy of Science 58(2)
nelius. This manuscript is Scientific Journal
Series No. 96-06-114 of the Kentucky Agri-
cultural Experiment Station. Mention of a
trademark, vendor, or proprietary product
does not constitute a guarantee or warranty of
the product by the U.S. Department of Agri-
culture or the University of Kentucky and
does not imply its approval to the exclusion of
other products that may also be suitable.
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Trans. Ky. Acad. Sci. 58(2):67-73. 1997.
Caddisflies (Insecta: Trichoptera) of the
Mainstem of the Kentucky River, Kentucky
Ronald E. Houp
Kentucky Department for Environmental Protection,
Division of Water, Ecological Support Section,
14 Reilly Road, Frankfort, KY 40601
and
Guenter A. Schuster
Department of Biological Sciences, Eastern Kentucky University,
Richmond, KY 40475
ABSTRACT
Collections of adult caddisflies from six Kentucky River sites were made from 1993 through 1996. Initially,
stations 2 and 3 were sampled weekly in 1993-94 to determine phenology and differences in species com-
position between open-river and lock-and-dam habitats. Notable differences in species composition of hy-
dropsychid caddisflies prompted the addition of stations 1,4,5, and 6 in 1994. Stations 3 and 5 were open-
river sites; the remaining stations were at locks and dams. Fifty species representing nine families and 23
genera were collected. Most species belonged to the families Leptoceridae (16), Hydroptilidae(12), and
Hydropsychidae (10). Emergences ranged from late April through late October with peak emergences in
late July and August. A total of 17 species was found at all stations, Cheumatopsyche campyla and Ceraclea
tarsipunctata being the most common. The open-river stations in comparison to the lock-and-dam sites had
distinctly different habitats; this was expressed as differences in species composition. Locks and dams, be-
cause of the abundance of hard substrates, provided more habitat for filter feeders, scrapers, and predators.
INTRODUCTION
The collection, evaluation, and monitoring
of aquatic communities over periods of time
is a tool used to evaluate water quality and to
protect aquatic life. However, biological eval-
uations are of limited value when collections
do not represent most or all available habitats.
It has become clear that the long-term moni-
toring (Kentucky Division of Water 1978—
1995) of invertebrates from the Kentucky Riv-
er, using multiplate samplers, does not ade-
quately represent the river's invertebrate com-
munity. Some species were missed due to bi-
ases of the sampler; other species, because of
limitations in larval taxonomy, could be iden-
tified only to the generic level. For that rea-
son, light trapping was initiated to collect adult
caddisflies, which in most cases can be iden-
tified to the species level. Resh’s (1975) list of
the caddisflies of Kentucky included records
for 175 species; of these, only two were re-
corded from the Kentucky River.
Mozley (1979) pointed out the lack of eco-
logical information regarding benthic macroin-
67
vertebrates of large rivers in North America.
Prior to that time only a few studies had been
conducted on these rivers. Mason et al. (1967)
and Fullner (1971) studied and compared the
efficiencies of various types of artificial sub-
strate samplers for collection of invertebrates
in large rivers. In the Ohio River, Beckett
(1982) studied the phenology of Hydropsyche
orris and Beckett and Miller (1982) investi-
gated colonization rates of larval invertebrates
on multiplate samplers in various current ve-
locities. Hall (1982) compared colonization of
invertebrates on basket and multiplate sam-
plers in the upper Mississippi River. Beckett
et al. (1983) studied distribution of larval ma-
croinvertebrates from four habitat types in the
lower Mississippi. Our study is the first to doc-
ument the caddisfly fauna from one of the
larger impounded rivers in Kentucky.
KENTUCKY RIVER DRAINAGE
The North, Middle, and South forks of the
Kentucky River originate in the mountains of
southeastern Kentucky. Each fork flows in a
northwest direction, joining near the town of
68 Transactions of the Kentucky Academy of Science 58(2)
- Kentucky River
N
(el
Dix River —
South Fork Kentucky River ~ :
Middle Fork Kentucky River Sy
North Fork Kentucky River ~ Z
Figure 1. Outline of the Kentucky River drainage area showing sites of collection of Trichoptera from 1993 through
1966. Inset shows area location within Kentucky.
Beattyville, to form the mainstem (Figure 1).
From there, the river continues to flow north-
west, joining the Ohio River at Carrollton,
Kentucky, a distance of 408 km (255 miles)
(Martin et. al. 1979). There are 14 locks and
dams (L&D) on the mainstem, some of which
have been in place for many years. For ex-
ample, L&D 1 near Carrollton was completed
in 1834, L&D 7 at High Bridge in 1895, and
L&D 14 at Heidelburg in 1917. Locks 5-14
have been closed for over a decade. The river
passes through the Cumberland, Allegheny,
Table 1.
ing stations along the river mainstem.
Kentucky River caddisflies. Location of collect-
Station Location
] Carroll Co., Lock and Dam (L&D) 1, at River
Mile (RM) 4.0 (River Kilometer [Rkim] 6.4)
upstream from Carrollton
Jessamine Co., L&D 7, at High Bridge
3 Jessamine Co., at RM 118.6 (Rkm 191), private
property 1.0 km upstream from confluence
with Dix River
4 Jessamine Co., at RM 157.5 (Rkm 253), down-
stream from Valley View
5 Estill Co., at RM 221.9 (Rkm 357), 2.4 km up-
stream from Ravenna
6 Lee Co., at L&D 14, RM 255.0 (Rkm 410), at
Heidelberg
bo
and Interior Plateau ecoregions (Omernik
1986). About midway along its course it flows
through a part of the Interior Plateau known
as the Inner Bluegrass. That segment, called
the Palisades, reflects major distinctions in ge-
ology and vegetation because of a geologic up-
lift late in the Paleozoic Era (McFarlan 1943).
Sheer limestone cliffs created by the down-
ward cutting of the river are as high as 122 m
(400 ft) above the river level, exposing the old-
est Ordovician limestone rocks in the state
(McFarlan 1943). The absence of other large
streams in the Inner Bluegrass and a rapidly
growing populace have greatly increased eco-
nomic and recreational demands on the river.
Published biological data from the Ken-
tucky River are few and mostly relegated to
studies of freshwater mussels that began with
Rafinesque (1820). Danglade (1922) and Wil-
liams (1974) documented the mussel fauna
and reported on commercial potentials of the
existing communities for freshwater pearls and
buttons. Tolin and King (1986) surveyed the
unionid mussels from L&D 4 at Frankfort,
downstream to the confluence of the Ohio
River. Most recently, Bradfield and Porter
(1990) presented a summary of investigations
related to surface water quality in the Ken-
Kentucky River Caddisflies—Houp and Schuster 69
Table 2. Trichoptera collected along the Kentucky River Table 2. (continued).
mainstem from 1993 through 1996.
Station
Station Taxa lie Ou awe
oo aE its OR ee Neureclipsis crepuscularis xX xX
Glossosomatidae (Walker)
Protoptila maculata (Hagen) xX Polycentropus cinereus Hagen X X X
Hydropsychidae Polycentropus confusus Hagen x
Ceratopsyche sparna (Ross) xX Rhyacophilidae
Cheumatopsyche campyla Ross =X X X X X X Rhyacophila lobifera Betten X X
be carecmeyee minuscula X X X TOTAL NUMBER OF
j SPECIES: 2 36 25 24 24 <
Cheumatopsyche oxa Ross X ec: ATTEN dicate
Cheumatopsyche pasella Ross XOX X XK X
Hydropsyche betteni Ross xX X xX X
Hydropsyche frisoni Ross X tucky River basin, which included sites in the
Hydropsyche orris Ross X X mainstem
Hydropsyche simulans Ross X xX X
Potamyia flava (Hagen) Xe OX METHODS AND MATERIALS
Hydroptilidae
Hydroptila ajax Ross x x xX x x x Adult caddisflies were collected from six
Hydroptila angusta Ross X X X X X _ stations (Figure 1) along the Kentucky River
ee ar Uae XX XX mainstem from 1993 through 1996. Initially,
ge cee ee nets Morton Xx Stations 2’ and 3'were sampled weekly dung
Hydroptila hamata Morton X X 199394 : ee penis REA
Hydroptila perdita Morton X X xX eae stations » &, O, an WETE a as m
Hydroptila waubesiana Betten X X X X XK X 1994. Stations 3 and 5 represented open-river
Ochrotrichia tarsalis (Hagen) X habitats; remaining stations were at L&D. Lo-
Ochrotrichia xena (Ross) Xx _ . cations of all stations are shown in Table 1.
Orthotrichia aegerfasciella Xe Ne xe XX , : se
(Cen een Collections were made using a plastic ice
One einaveristata Morton X X chest tray with about 500 ml of 70% ethanol,
Oxyethira pallida (Banks) XE EX UIOXG SO XOX IX set at the base of a 12 volt Coleman lantern,
Leptoceridae equipped with ultra-violet (UV) bulbs. At
Ceraclea ancylus (Vorhies) X X —— ~_—__—s L&D sites, a lantern was turned on and placed
Ceraclea cancellata (Betten) ee eon top of the downstream lock gate mooring
Ceraclea flava (Banks) X X leageaagle Meee: dot i ft
Ceraclea punctata (Banks) X X X wall at BUSK: it was turned om an hour a ct
Ceraclea tarsipunctata (Vorhies) X X X X X X dark. Private docks were used at the open-riv-
Ceraclea transversa (Hagen) X X X X X X __ er sites in the same manner. Species identifi-
Mystacides sepulchralis (Walker) X cations were based mostly on males.
Nectopsyche candida (Hagen) X X X K XK X
Nectopsyche exquisita (Walker) XG Xe NN OX ae Xe RESULTS AND DISCUSSION
Nectopsyche pavida (Hagen) 6 2G POS IKHOK
Oecetis avara (Banks) xX This study began in early April 1993, with
Oecetis cinerascens ie X X X X X X_ weekly collections at stations 2 and 3 (Figure
Oecetis inconspicua (Walker) XC Xe Ne NX: il The purpose was to document species
Oecetis persimilis (Banks) XG Xe XXX ). ae le d Hone B dif
Triaenodes ignitus (Walker) Rex xxx x 2 CoMmpositen zal emer oT nee |e Cae ra
Te Horbetariis Milne xt ferent habitat types (i.e., from a L&D site and
Limnephilidae a typical open-river site). A year of weekly col-
Ironoquia punctatissima X lections showed that species compositions be-
(Walker) : tween stations 2 and 3 were similar (Table 1).
a eg cepidaa Hagen) % However, the functional feeding requirements
hilopotamidae :
Chimarra obscura (Walker) ETE XE OX, XS Xe EX of some SPCCIe® reflected the types and
amounts of different microhabitats between
Phyrganeidae h ve eee diff, ‘
Agrypnia vestita (Walker) x the stations. On instance, dl erences 1m num-
Ptilostomis ocellifera (Walker) X bers and species of hydropsychids were nota-
Ptilostomis semifaciata (Say) X ble at the L&D site compared to the open-
Polycentropodidae t : river site. So stations 1, 4, 5, and 6 were added
Cyrnellus fraternus (Banks) SoS 2a in 1994 to increase the scope of the study. Sta-
70 Transactions of the Kentucky Academy of Science 58(2)
Table 3.
Seasonal occurance of trichoptera collected during this study from the Kentucky River Mainstem.
Hydropsyche frisoni
Hydropsyche orris
Hydropsyche simulans
Potamyia flava
Hydroptilidae
Hydroptila ajax
Hydroptila angusta
tion 5 was the other open-river site. Filter-
feeding species (mostly hydropsychids) from
all L&D sites ranged from 6 to 11 species in
1994, compared with | and 2 species at open-
river sites (Table 1). Station 2 had eight of the
10 hydropsychid species. Hydropsyche orris
and Potamyia flava occurred only at stations 1
and 2; Cheumatopsyche oxa, only at station 1;
Ceratopsyche sparna, only at station 2; and
Hydropsyche frisoni, only at station 6. Filterer
species common to all stations were Chimarra
obscura and Cyrnellus fraternus, though col-
lected in fewer numbers at the open-river
sites. Station 2 yielded three of the four spe-
cies of polycentropodids, with Polycentropus
confusus occurring only at station 2.
Beckett and Miller (1982) observed that
flow velocities contributed to different inver-
tebrate communities becoming established in
the Ohio River at locks and dams. In our study
of adult caddisflies, it was also evident that dif-
ferent sustained flow velocities were respon-
sible at least in part for the diversity of filterer
organisms at the L&D stations and organisms
associated with the slower currents at open-
river stations.
Hall (1982) found 31 larval taxa in basket
samplers and 21 taxa from multiplate samplers
in the upper Mississippi River. In comparison,
the Kentucky Division of Water's (KDOW),
Biological Monitoring Program (BMP) multi-
plate collections from four mainstem Ken-
tucky River sites (KDOW 1978-1995) ranged
from 6 to 31 larval taxa. In our study, adult
caddisflies alone ranged from 24 to 36 species
at six sites, for a total of 50 species throughout
the river (Table 2).
The augmented flow from Dix River (Her-
rington Lake, hydroelectric turbines) (see Fig-
ure 1) is thought to be the reason for the large
number of taxa, and especially a large number
of filter-feeding species, collected at station 2
(Figure 1). In addition, 14 of 16 leptocerid
species were collected there. Because of the
occurrences of leptocerid larvae at all stations
and habitat types throughout the river, the mi-
crohabitat requirements for them are appar-
ently less specific in relation to flow velocities
Kentucky River Caddisflies—Houp and Schuster Za
Table 3.
(continued).
or other physical attributes. Although the
numbers of individuals collected in open-river
sites were much fewer for some species, there
were also distributional differences. For ex-
ample, Ceraclea ancylus and C. flava were col-
lected only at stations 1 and 2, and Mystacides
sepulchralis and Oecetis avara and were col-
lected only at station 6.
Five of the 12 hydroptilid species were col-
lected from all stations; at station 2 nine of the
12 were common. Most of these species are
scrapers during the larval stages, and the L&D
provides attachment sites for them as well as
for the periphyton (algae and diatoms) they
consume. Ochrotrichia xena was collected
only at station 2; O. tarsalis, only at station 6;
and O. cristata, only at stations 3 and 5.
From the distribution and numbers of hy-
dropsychids, polycentropodids, and certain hy-
droptilids collected, it is apparent that the
L&D sites provided the microhabitats re-
quired by these species. These habitats includ-
ed a variety of flow regimens and firm sub-
strates for needed attachment. In contrast,
species such as Orthotrichia cristata and Neu-
reclipsis crepuscularis, which require slower
currents, were more abundant at open-river
stations.
It is likely that some of the species collected
at light traps flew in from nearby tributary
habitats. These included four species from sta-
tion 1 (Ironoquia punctatissima, Ptilostomis
semifaciata, and Ptilostomis ocellifera), and
three from station 6 (Agrypnia vestita, Protop-
tila maculata, and Rhyacophila lobifera).
These species are often associated with head-
water streams.
The flight patterns of all species are shown
in Table 3. Most caddisflies exhibit a univol-
tine life cycle; our data indicate that this is
true for all Kentucky River species. It is ap-
parent that two different flight patterns are
common among these caddisflies. The first is
a single cohort that emerges over a long pe-
riod of time such as in Cheumatopsyche cam-
pyla or a short, synchronous emergence peri-
od such as in Protoptila maculata. The second
pattern is shown by species such as Ochrotri-
chia tarsalis, which apparently has two cohorts
emerging at different times. In addition, sea-
72 Transactions of the Kentucky Academy of Science 58(2)
Table 3. (continued).
Nectopsyche exquisita
nneccanteesceecsseecscanceccnccasceececcescenerssscontenecsncsenssnscassenensccsscuaneusceccuerentensecs theo
Nectopsyche pavida
Oecetis avara
Sedeg canevarrerensccmsctcnatesacennbsensstcpsceassscuscrcrd=srerscrcnntefigiccenaacstcetetsaderasessesnes! use
Oecetis cinerascens
cecucennnenpeacescacsecsoosucnnnccccsseusesesncscserccsussereccesssscestessusrecsessevascarsceseessesses Pes
Oecetis inconspicua
scnwunsendecccseveccecarcnessccsecuvsseessecessaccccsrecvesersuccssseusesseascsrcersnenensaessacscenresse= bps
Oecetis persimilis
Triaenodes tardus
Limnephilidae
Tronoquia punctatissima
Pycnopsyche lepida
Philopotamidae
Table 3. (continued).
Phyrganeidae
be edesnacon OS CO ZOS CREE EREDROELERERn Cero ISRO SEC CE CEC CECE DISD CID EEOC REO ROE the
Agrypnia vestita
Cyrnellus fraternus
Poverpesantueescaraspesceesasaccpeesnsdanstnsnnssasatstscsssssaeswustansesccaceceserscseteesuressaceses He
Neureclipsis crepuscularis
Rhyacophila lobifera
Kentucky River Caddisflies—Houp and Schuster 73
sonal-occurrence data show that while the ma-
jority of species emerge throughout the sum-
mer some species, e.g., Rhyacophila lobifera,
have a synchronous emergence period in the
early spring and that other species, e.g., Iron-
oquia punctatissima and Pycnopsyche lepida,
have an emergence period restricted to a very
short period in the fall. The family Leptocer-
idae had the largest number of genera and
species active at the same time throughout the
river (Table 3).
The most abundant species at all stations
and throughout the study was Cheumatopsy-
che campyla. The sex ratio of males to females
is often skewed in light trap samples towards
females. For example, we collected no males
of Cheumatopsyche oxa. Oecetis inconspicua
and O. persimilis were also collected as phar-
ate adults attached to posterior ridges of fresh-
water clams (Unionidae) in areas below station
2. It is believed that unionids represent an im-
portant type of habitat for many large-river in-
sect species in that they provide firm sub-
strates for attachment.
Locks and dams are the largest stationary
objects in the Kentucky River. They, in con-
cert with the various flow velocities created by
these structures, provide unique habitats for
filter feeders, scrapers, and predatory insects
in the mainstem where such habitats are often
scarce.
ACKNOWLEDGMENTS
We thank the following Kentucky Division
of Water personnel: Scott Hankla, Terry An-
derson, Dru Hawkins, Lajuanda Maybriar, Ka-
thy Collins, and Karen Smathers. We acknowl-
edge the help of Katy, Wes, and Laura Houp
with travel and equipment.
LITERATURE CITED
Beckett, D.C. 1982. Phenology of Hydropsyche orris
(Trichoptera: Hydropsychidae) in the Ohio River:
changes in larval age structure and substrate coloniza-
tion rates. Environm. Entomol. 11:1154—-1158.
Beckett, D.C., and M.C. Miller. 1982. Macroinvertebrate
colonization of multiplate samplers in the Ohio River:
the effect of dams. Canad. J. Fish. Aquatic Sci. 39:
1622-1627.
Beckett, D.C., C.R. Bingham, and L.G. Sanders. 1983.
Benthic macroinvertebrates of selected habitats of the
Lower Mississippi River. J. Freshwater Ecol. 2:247-261.
Bradfield, A.D., and S.D. Porter. 1990. Summary of bi-
ological investigations relating to surface-water quality
in the Kentucky River Basin, Kentucky. U.S. Geol. Sury.
Water-Resources Invest. Rep. 90-4051.
Danglade, E. 1922. The Kentucky River and its mussel
resources. U.S. Bur. Fish. Doc. 934:1-8.
Fullner, R.W. 1971. A comparison of macroinvertebrates
collected by basket and modified multiple-plate sam-
plers. J. Water Pollut. Control Fed. 43:494-499.
Hall, T.J. 1982. Colonizing macroinvertebrates in the Up-
per Mississippi River with a comparison of basket and
multiplate samplers. Freshwater Biol. 12:211-215.
Kentucky Division of Water. 1978-1995. Biological Mon-
itoring Program, PRI023, PRIO24, PRIO25, PRI026,
1978-1995. Frankfort, KY.
McFarlan, A.C. 1943. Geology of Kentucky. University
of Kentucky, Lexington, KY.
Martin, W.H., W.S. Bryant, T.S. Lassetter, and J.B. Varner.
1979. The Kentucky River Palisades, flora and vegeta-
tion. An inventory report to the Kentucky Chapter of
the Nature Conservancy. Geographical Studies and Re-
search Center, Eastern Kentucky University, Richmond,
KY.
Mason, W.T., J.B. Anderson, and G.E. Morrison. 1967. A
limestone-filled, artificial substrate sampler-float unit
for collecting macroinvertebrates in large streams. Prog.
Fish-Cult. 29:74.
Mason, W.T., P.A. Lewis, and T.B. Anderson. 1968. Mac-
roinvertebrate collections and water quality monitoring
in the Ohio River Basin 1963-1967. Coop. Rep. Office
Techol. Prog., Ohio Basin Regions & Anal. Qual. Con-
trol Lab. Water Qual. Office. Environm. Protect. Agen-
cy, Cincinnati, OH.
Mozley, S.C. 1979. Neglected characters in larval mor-
phology as tools in taxonomy and phylogeny of Chiro-
nomidae (Diptera). Entomol. Scand. Suppl. 10:27-36.
Omernik, J.M. 1986. Aquatic ecoregions of the conter-
minous United States. Ann. Assoc. Am. Geogr. 77(1):
118-125.
Rafinesque, C.S. 1820. Monographie des coquilles bi-
valves fluviatiles de la Riviére Ohio, contenant douze
genres et soixante-huit especes. Ann. Gén. Sci. Phys.
5(13):287-322, 2 plates.
Resh, V.H. 1975. A distributional study of the caddisflies
of Kentucky. Trans. Kentucky Acad. Sci. 36:6-16.
Tolin, W., and M. King. 1986. A survey of the freshwater
mussel populations on the lower Kentucky River,
Franklin, Owen, Henry and Carroll Counties, Kentucky.
U.S. Fish and Wildl. Serv., Ecol. Serv., Cookeville, TN.
Williams, J.C. 1974. Commercial fisheries investigations
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Trans. Ky. Acad. Sci. 58(2):74-79. 1997.
Noteworthy Vascular Plant Discoveries from Kentucky
Gary W. Libby and Randy L. Mears
Department of Biological Sciences, Eastern Kentucky University,
Richmond, Kentucky 40475
and
C. Tom Bloom
1209 Glade, College Station, Texas 77840
ABSTRACT
Nineteen species of rare vascular plants are documented from Kentucky. Two species (Juncus validus
Coville and Scirpus [Schoenoplectus] mucronatus L.) are reported as new to the state; $. mucronatus has
serious potential as an invasive wetland weed. Three others (Calopogon tuberosus (L.) BSP. var. tuberosus,
Polygala paucifolia Willd., and Rhynchospora macrostachya Torr. var. macrostachya) represent the only
known extant Kentucky populations. New county distributional information for 14 other taxa of rare Kentucky
plants is also presented.
INTRODUCTION
Kentucky is relatively poorly known floris-
tically. Recent field studies have yielded the
state records and other noteworthy collections
reported here. Some of these uncommon spe-
cies are restricted to specific habitats or eco-
logical communities in Kentucky; others are
recently naturalized or represent significant
range extensions. The rarity of these species
in Kentucky is based on the listing of endan-
gered, threatened, special concern, and his-
torical plants and animals of Kentucky (Ken-
tucky State Nature Preserves Commission
[KSNPC] 1996) and follows the scientific
name where applicable. An asterisk preceding
the scientific name indicates a non-indigenous
taxon.
NOTEWORTHY SPECIES
Adiantum capillus-veneris L. KSNPC
Threatened. LOGAN CO.: Dot 7.5 Quad.,
wet limestone seeps along Red River, adjacent
to KY 102 (near state line), se of Keysburg,
rare, 14 Sep 1995, Libby and Mears 1229
(BEREA, EKY, KNK).
Significance. Cranfill (1980) stated that
“no known stations are presently extant” in
Kentucky. Southern maidenhair-fern has been
collected in seven Kentucky counties (Adair,
Boyd, Carter, Powell, Pulaski, Taylor, and
Wayne) (Campbell n.d.; Cranfill 1980; McCoy
1938; Medley 1993). In addition, Cranfill
74
(1980) cited a formerly adventive colony fron
Nelson County. This collection represents a
westward range extension for this epipetric
pteridophyte.
Aureolaria patula (Chapman) Pennell.
KSNPC Threatened. This species is also a can-
didate (Category C2) for federal listing (Unit-
ed States Fish and Wildlife Service [USFWS]
1993). ADAIR CO.: Cane Valley 7.5 Quad.,
along Green River Lake near Holmes, steep,
rocky lakeshore, 26 Sep 1995, Libby 1277
(EKY). BARREN CO.: Meador 7.5 Quad.,
along Barren River Lake tailwaters near Gag-
ing Station, open riverbank, 4 Oct 1995, Libby
1302 (EKY). CLINTON CoO.:: Frogue and Al-
bany 7.5 Quads., along Dale Hollow Lake near
Skull Bluff, steep, rocky lakeshore and edge of
woods, 20 Sep 1995, Libby, Bloom, and Evans
1252 & 1253 (EKY). CUMBERLAND CoO.::
Frogue 7.5 Quad., along Dale Hollow Lake
nw of Boys Island, rocky lakeshore, 6 Oct
1995, Libby 1312 (EKY). GREEN CO.: Sum-
mersville 7.5 Quad., along Green River at KY
61/70 bridge, just s of Greensburg, rocky,
open woods, 15 Sep 1995, Libby and Mears
1240 (EKY). TAYLOR CO.: Cane Valley 19
Quad., along Green River just below Green
River Lake Dam, near KY 55 bridge, steep,
forested streambanks, 26 Sep 1995, Libby
1282 (EKY). WARREN CO.:: Bowling Green
South 7.5 Quad., along Drakes Creek, ca. 1/4
mile upstream from Old Scottsville Road
Kentucky Vascular Plant Discoveries—Libby, Mears, and Bloom 75
bridge, steep, rocky riverbank, 14 Sep 1995,
Libby and Mears 1235 (EKY). WAYNE CO.:
Nevelsville 7.5 Quad., along Big South Fork
Cumberland River just downstream of conflu-
ence with Fanny Hollow, open, limestone
bluff, 23 Aug 1995, Libby 1186 (EKY).
Significance. This species was previously
known from only five Kentucky counties (Ed-
monson, Hart, Logan, McCreary, and Pulaski)
(Braun 1941; Campbell n.d.; Evans 1989:
Medley 1993). Cumberland false foxglove has
now been documented in an additional eight
counties (Adair, Barren, Clinton, Cumberland,
Green, Taylor, Warren, and Wayne) through
extensive field work (Libby 1996). These col-
lections expand the range of Aureolaria patula
in Kentucky and indicate that this species may
not be as rare as previously thought (Kral
1983: Pennell 1935; USFWS 1993).
Baptisia tinctoria (L.) R. Br. KSNPC
Threatened. MCCREARY CO.: Sharp Place
7.5 Quad., growing on roadside and oak-pine
upland forest margin, w side of KY 167, ca.
0.5 mile n of Kentucky/Tennessee line, 12 Jul
1995, Libby and Klein 1003 (EKY, KNK).
WAYNE CoO.: Sharp Place 7.5 Quad., growing
on roadside and oak-pine upland forest mar-
gin, w side of KY 167, ca. 0.6 mile n of Ken-
tucky/Tennessee line, 12 Jul 1995, Libby and
Klein 1002 (KNK).
Significance. Previously known from
Letcher, Harlan, and Bell counties (Campbell
n.d.; Medley 1993), yellow wild indigo has
now been documented in two additional coun-
ties.
Bartonia virginica (L.) BSP. KSNPC
Threatened. CLINTON CO.: Albany 7.5
Quad., Pyle’s Marsh, 1 mile sse of Abstons
Comer, situated e of Old Burkesville Road
and w of KY 1590, moss-covered areas around
tree bases in swampy area, rare, 15 Jun 1995,
Libby and Bloom 959 (EKY).
Significance. Virginia screwstem had been
collected in Calloway, Harlan, Laurel, Mc-
Creary, and Whitley counties (Campbell n.d;
Medley 1993). This collection represents an
additional county and the first collection of
this species from the Highland Rim of Ken-
tucky. The more widespread B. paniculata
(Michx.) Muhl. var. paniculata was found
about 10 meters away in the adjacent wet
woods. Gillett (1959) found that B. paniculata
and B. virginica are sympatric over a large
portion of their ranges and that occasional in-
termediates occur.
Calopogon tuberosus (L.) BSP. var. tubero-
sus. KSNPC Endangered. LAUREL CO.:
Vox 7.5 Quad., powerline right-of-way near
Grove Recreation Area, ca. 0.7 road mile ne
of large transmission line, diverse assemblage
of prairie species, rare, 15 Jun 1992. No col-
lection was made because of the rarity and low
relative abundance at the site; photographs are
on file at the Kentucky State Nature Preserves
Commission, Frankfort, Kentucky.
Significance. This species was formerly re-
ported from 12 counties according to Camp-
bell (n.d.) and Medley (1993). Prior to our re-
port, however, grass pink had not been seen
in Kentucky since the early 1980s (Campbell
n.d.; Medley 1993).
Carex crawei Dewey. KSNPC Special
Concern. WAYNE CO.: Parnell 7.5 Quad.,
McClown Mountain Glade, ca. 0.5 air mile nw
of US 90/KY 858 intersection, xeric limestone
glade, locally common, 13 Jun 1995, Libby
and Bloom 937 (EKY, KNK).
Significance. Crawe’s sedge was described
as “to be looked for” in Kentucky by Beal and
Thieret (1986). Since that time, Carex crawei
is known with certainty only from Bullitt,
Grayson, Hardin, Larue, and Nelson counties
(Campbell n.d.; Medley 1993). This collection
represents a range extension and outlier from
known localities in the Bluegrass and the
Nashville Basin of Tennessee (Chester et al.
1993). This disjunction is probably due to the
unusual occurrence of a flat, xeric limestone
glade in this part of the Highland Rim.
Carex stricta Lam. var. stricta. CLINTON
CO.: Albany 7.5 Quad., Pyle’s Marsh, 1 mile
sse of Abstons Corner, situated e of Old Bur-
kesville Road and w of KY 1590, forming large
tussocks in a wooded swamp, locally abundant,
15 Jun 1995, Libby and Bloom 960 (EKY,
KNK).
Significance. In Kentucky, this species had
been collected only in Calloway, Harlan, Hart,
McCreary, and Metcalfe counties (Campbell
n.d.; Medley 1993). Our collection represents
an additional county record for this marsh-
adapted sedge. This taxon should be strongly
considered as an addition to the endangered,
threatened, special concern, and _ historical
plants of Kentucky (KSNPC 1996).
Castanea dentata (Marshall) Borkh.
76 Transactions of the Kentucky Academy of Science 58(2)
KSNPC Endangered (reproducing trees only).
HARLAN CoO.: Louellen 7.5 Quad., Hi-Lewis
Pine Savanna, upper s-facing slopes of Hi-
-Lewis Branch, near Hiram, 24 Jun 1995, Lib-
by, Libby, Libby, and Evans 988 (BEREA,
EKY, KNK).
Significance. This collection of American
chestnut in flower, including evidence of last
years fruits (hulls only), indicates the presence
of sexually reproducing chestnut trees in Ken-
tucky. On a return visit in October 1995 we
collected 15 nuts. Some of these were for-
warded to J. Durrell (Kentucky Nut Growers
Association) to test for reproductive potential.
It is significant that 23 trees (usually multistem
clumps) were observed in flower at this loca-
tion. This multistem character suggests that
these trees have survived previous chestnut
blight [Cryphonectria parasitica (Murr.) Barr]
(Hicks and Marks 1990).
Cenchrus longispinus (Hack.) Fern. ES-
TILL CO.: Palmer 7.5 Quad., growing in
sandy riverbank downstream from Kentucky
River Lock and Dam No. 11, rare, 25 Aug
1994, Libby and Walker 454 (BEREA, EKY,
KNK).
Significance. In Kentucky, this species had
previously been reported from Ballard, Ful-
ton, Hickman, Jefferson, and McCracken
counties in the Coastal Plain and Bluegrass
regions (Medley 1993). Our collection from
the Knobs region represents a significant east-
ward range extension.
Cyperus croceus Vahl. MCCREARY CO.:
Barthell 7.5 Quad., growing in wet area along
railroad tracks adjacent to Roaring Paunch
Creek near Barthell, several clumps, 7 Jul
1995, Mears and Klein s.n. (EKY).
Significance. This species was recently re-
ported as new to Kentucky (Madison County)
(Mears and Libby 1995); ours is the second
collection for the state.
*Cyperus difformis L. SIMPSON CO.:
wet ditch in front of gas station just w of 1-65
at exit 6 (KY 100) near Franklin, locally abun-
dant, 14 Sep 1995, Libby and Mears 1219
(KNK, WKU).
Significance. This naturalized species was
recently reported as new to Kentucky (Mears
and Libby 1995) from Knott, Floyd, and Pike
counties. This collection represents an addi-
tional county record and westward range ex-
tension. In the United States Cyperus diffor-
mis is known from Alabama, Florida, Louisi-
ana, Mississippi, North Carolina, Pennsylva-
nia, Tennessee, and Virginia, and also from
Nebraska, Texas, and California (Bryson and
Carter 1994; Godfrey and Wooten 1979; Lips-
comb 1980). Godfrey and Wooten (1979)
speculated that the species would likely be
found elsewhere in the eastern United States.
Holm et al. (1991) ranked Cyperus difformis
as one of the world’s worst weeds. Bryson and
Carter (1994) described it as particularly per-
nicious and noted that it is spreading in the
United States especially along major water-
ways, through introduction at ports-of-entry,
and in rice production in California.
Epilobium ciliatum Raf. var. cilia-
tum. MERCER CO.: Wilmore 7.5 Quad.,
directly below Kentucky River Lock and Dam
No. 7, growing in gravel wash, rare, 21 Oct
1994, Libby and Walker 440 (EKY).
Significance. This is the second report of
this taxon from Kentucky. Medley (1993) cited
a specimen from McCreary County collected
by J. Campbell. We have seen no vouchers of
this species from Kentucky. Gleason and
Cronquist (1991) described the habitat and
nature of this taxon as “wet places, often in
unstable habitats, variable [morphology] and
often somewhat weedy.”
Juncus validus Coville. MCCREARY CO.:
Barthell 7.5 Quad., coal mining property ca.
0.5 mile se of KY 791 just n of Blue Heron
Scenic Railway, w of Coffee Branch, frequent,
5 Jul 1995, Mears and Klein s.n. (EKY).
MCCREARY CoO.: Bell Farm 7.5 Quad., ca.
0.9 mile s of Pleasant Ridge Church, roadside
of KY 1363, wet ditch, 13 Jul 1995, Libby,
Mears, and Klein 1116B (EKY, KNK); same
location, 8 Aug 1996, Libby and Libby 1403
(BEREA, EKY, KNK). MCCREARY CoO.::
Barthell 7.5 Quad., ca. 2 miles se of Bald
Knob at USFS 575/USFS 6130 junction,
USFS 6130 roadside, 19 Jul 1995, Libby and
Mears s.n. (KNK).
Significance. This species is not reported
in Beal and Thieret (1986), Browne and Athey
(1992), Campbell (n.d.), or Medley (1993).
Our collections document the presence of
Juncus validus in the Commonwealth.
Lilium superbum L. KSNPC Threatened.
LAUREL CO.: growing in edge of swamp
along w side of U.S. 25 ca. 1 mile s of inter-
section with KY 1006, 25 Jun 1993, Mears
Kentucky Vascular Plant Discoveries—Libby, Mears, and Bloom We
93-234 (EKY). LAUREL CO.: e of KY 30 be-
tween Raccoon and Atlanta, wet woods along
Raccoon Creek, 26 Jun 1996, Libby and Har-
towicz 1396 (EKY). MCCREARY CoO.: Bar-
thell 7.5 Quad., floodplain along e side of
South Fork Cumberland River just n of con-
fluence with Bear Creek, locally frequent, 11
Jul 1995, Libby, Mears, and Klein 994 (EKY).
Significance. Turk’s-cap lily has an odd
distribution in Kentucky. According to Camp-
bell (n.d.) and Medley (1993), this species has
been collected from Big Black Mountain in
Harlan County and also from wet wooded
seeps in Calloway, Hickman, and McCracken
counties on the Coastal Plain. According to
Adams (1981) and Adams and Dress (1982),
however, the Coastal Plain specimens must
represent L. michiganense Farwell. Medley
(1993) noted that the Coastal Plain specimens
lack the “typical many-flowered inflorescence”
of Lilium superbum, but the descriptions in
Adams (1981), Adams and Dress (1982), and
Gleason and Cronquist (1991) allow for vari-
ation in number of flowers. Clearly, a thorough
taxonomic review of this genus is needed in
Kentucky. These represent the only known
collections of this species from the Low Hills
Section and Cliff Section of the Appalachian
Plateau of Kentucky.
*Linum usitatissimum L. MADISON
CO.: Berea 7.5 Quad., growing in tire rut on
roadside at the corner of Elm Street and Scaf-
fold Cane Road, a European cultigen, escaped
or adventive, 16 Jul 1993, Libby 343 (BEREA,
EKY, KNK).
Significance. This species has been col-
lected in Fulton County from the Coastal
Plain. A collection at the University of Ken-
tucky cited by Browne and Athey (1992) and
Wharton and Barbour (1991) from Fayette
County may be from cultivation (Campbell
n.d.; Medley 1993). Our collection represents
the second report of cultivated flax naturalized
in Kentucky. Grown in southeastern United
States until the late 1800s (Strausbaugh and
Core 1978), this European cultigen is adven-
tive in fields and roadsides in the United
States and southern Canada (Gleason and
Cronquist 1991).
Muhlenbergia cuspidata (Torr.) Rydb.
KSNPC Threatened. LOGAN CO.: Dot 7.5
Quad., limestone outcrops along Red River,
adjacent to KY 102 (near state line), se of
Keysburg, several clumps, 14 Sep 1995, Libby
and Mears 1230 (EKY). WAYNE CoO.: Mill
Springs 7.5 Quad., steep, rocky cliffs at Dry
Branch/Lake Cumberland confluence (w
side), several large clumps on rock face, 24 Jul
1995, Libby and Bloom 1136 (EKY).
Significance. In Kentucky, plains muhly
had been collected from four counties (An-
derson, Estill, Garrard, and Pulaski) (Bryant
1973; Campbell n.d.; Medley 1993). Our col-
lections document its occurrence in two ad-
ditional counties.
Polygala paucifolia Willd. LAUREL CoO.:
London SW 7.5 Quad., growing along bluffs
on w side of Willie Green Road ca. 2.5 miles
n of its intersection with KY 192, 1 May 1993,
Mears 93-181 (BEREA, EKY).
Significance. The report of flowering win-
tergreen in Gibson (1961) was probably based
on McFarland (1942): the voucher for that re-
port was destroyed in the 1948 Norwood Hall
fire at the University of Kentucky (Medley
1993). Our collection represents the only
known extant population of this taxon in Ken-
tucky. This is another taxon that should be
added to the endangered, threatened, special
concern, and historical plants of Kentucky
(KSNPC 1996).
Rhynchospora macrostachya Torr. var. ma-
crostachya. KSNPC Endangered. WAYNE
CO.: Parnell 7.5 Quad., Cooley’s Meadow, ca.
0.8 air mile ssw of Murl, wet meadow ne of
Cooley’s tobacco barn and e of Hack Denny's
comfield, rare, 27 Jul 1995, Libby and Bloom
1160 (BEREA).
Significance. This is the second report of
this taxon from Kentucky. The species may be
extirpated from Hundred Acre Pond, Hart
County (Medley 1993). Our collection may
represent the only known extant population of
tall beak-rush in the state.
*Scirpus mucronatus L. [|=Schoenoplectus
mucronatus (L.) Pallas} PULASKI CO.: Ano
7.5 Quad., pond at the end of USFS 807, w
of “The Narrows” of the Rockcastle River, 26
Jul 1995, Mears and Klein s.n. (BEREA,
MICH). PULASKI CO.: same location but
found in two additional ponds, possibly origi-
nally established from contaminated ornamen-
tal water-lilies (Nymphaea sp.) planted in larg-
est pond, 10 Oct 1995, Libby and Mears 1317
(BEREA, EKY, KNK, PH, WIS).
Significance. Scirpus mucronatus has not
78 Transactions of the Kentucky Academy of Science 58(2)
been previously reported from Kentucky (Ar-
nold and Beal 1981; Beal and Thieret 1986;
Browne and Athey 1992; Campbell n.d.; Med-
ley 1993). This European bulrush is known
from very few locations in the eastern United
States (S. G. Smith, University of Wisconsin,
pers. comm., 21 Oct 1996). Britton and Brown
(1970) reported only one location; “a swamp
in Delaware County, Pennsylvania” and fur-
ther stated that S. mucronatus is widely dis-
tributed in the Old World. Gleason (1952) de-
scribed the distribution of S. mucronatus as
“waif on ballast at Camden, New Jersey,” but
Gleason and Cronguist (1991) did not list this
species as occurring within northeastern Unit-
ed States. This species is morphologically very
similar to the native Scirpus purshianus Fern.
(=Schoenoplectus purshianus (Fern.) M. T.
Strong). Scirpus mucronatus can be distin-
guished from S. purshianus by its sharply tri-
gonal culms, trifid styles, and absence of leaf
blades. Scirpus mucronatus may be estab-
lished at other ponds and lakes in Pulaski
County; based on the relative abundance
(dominant or codominant emergent aquatic)
at the known sites this bulrush should be con-
sidered a pernicious weed. This taxon has
been recently placed in the genus Schoeno-
plectus (Bruhl 1995).
ACKNOWLEDGMENTS
Funds to support this research were par-
tially provided by the Kentucky State Nature
Preserves Commission, Kentucky River Au-
thority, National Biological Service, and U.S.
Fish and Wildlife Service. We thank R. Klein,
S.A. Walker, E. Hartowicz, and T. Libby for
valuable field assistance; Dr. R.L. Jones (Jun-
cus validus), Dr. R.F.C. Naczi (Carex crawei,
C. stricta var. stricta), Dr. A.A. Reznicek (Scir-
pus mucronatus), Dr. A.E. Schuyler (Scirpus
mucronatus), Dr. S.G. Smith (Scirpus mucron-
atus), and Dr. J.W. Thieret (Baptisia tinctoria)
for checking determinations; Dr. J.J.N. Camp-
bell for sharing his unpublished Atlas of the
Flora of Kentucky; and Dr. R.L. Thompson
and the anonymous reviewers of this manu-
script for critical comments.
LITERATURE CITED
Adams, R.M., II.
graph of the turk’s-cap lilies of eastern North America
1981. A systematic study and mono-
(with introduction of some new cultivars). Ph.D. dis-
sertation. Cornell University, Ithaca, NY.
Adams, R.M., IH, and W.J. Dress. 1982. Nodding Lilium
species of eastern North America (Liliaceae). Baileya
21:165-188.
Amold, S.C., and E.O. Beal. 1981. A survey of Scirpus
in Kentucky with problem species-complex analyses.
Trans. Kentucky Acad. Sci. 42:16-28.
Beal, E.O., and J.W. Thieret. 1986. Aquatic and wetland
plants of Kentucky. Kentucky Nature Preserves Com-
mission, Scientific and Technical Series Number 5,
Frankfort, KY.
Braun, E.L. 1941. Notes on Kentucky plants V. Castanea
6:137-140.
Britton, N.L., and A. Brown. 1970. An illustrated flora of
the northern United States and Canada. 3 volumes. Do-
ver Publications, Mineola, NY. [Reprint of a work first
published in 1913]
Browne, E.T., Jr., and R. Athey. 1992. Vascular plants of
Kentucky: an annotated checklist. University Press of
Kentucky, Lexington, KY.
Bruhl, J. 1995. Sedge genera of the world: relationships
and a new classification of the Cyperaceae. Austral.
Syst. Bot. §:125-305.
Bryant, WS. 1973. An ecological investigation of Panther
Rock, Anderson County, Kentucky. Ph.D. dissertation.
Southern Illinois University, Carbondale, IL.
Bryson, C.T., and R. Carter. 1994. Notes on Carex, Cy-
perus, and Kyllinga (Cyperaceae) in Mississippi with re-
cords of eight species previously unreported to the
state. Sida 16:171-182.
Campbell, J.J.N. n.d. Atlas of the flora of Kentucky. Lex-
ington, KY. [Unpublished draft]
Chester, E.W., B.E. Wofford, R. Kral, H.R. Deselm, and
A.M. Evans. 1993. Atlas of Tennessee vascular plants.
Volume 1: Pteridophytes, gymnosperms, and monocots.
Center for Field Biology, Austin Peay State University,
Clarksville, TN.
Cranfill, R. 1980. Ferns and fern allies of Kentucky. Ken-
tucky Nature Preserves Commission Scientific and
Technical Series Number 1, Frankfort, KY.
Evans, M. 1989. A natural areas inventory of Logan,
Simpson, and Warren counties, Kentucky. Tech. Rept.,
Kentucky State Nature Preserves Commission, Frank-
fort, KY.
Gibson, D. 1961. Life-forms of Kentucky flowering
plants. Am. Midl. Nat. 66:1—60.
Gillett, J.M. 1959. A revision of Bartonia and Obolaria
(Gentianaceae). Rhodora 61:43-62.
Gleason, H.A. 1952. The new Britton and Brown illus-
trated flora of the northeastern United States and ad-
jacent Canada. 3 volumes. Hafner Publishing Co., New
York, NY.
Gleason, H.A., and A. Cronquist. 1991. Manual of vas-
cular plants of northeastern United States and adjacent
Canada. 2nd ed. New York Botanical Garden, Bronx,
NY.
Godfrey, R.K., and J.W. Wooten. 1979. Aquatic and wet-
Kentucky Vascular Plant Discoveries—Libby, Mears, and Bloom 79
land plants of the southeastern United States: mono-
cotyledons. University of Georgia, Athens, GA.
Hicks, G.F., Jr., and G.C. Marks. 1990. A survey: Amer-
ican chestnut (Castanea dentata) trees in northwest In-
diana. Castanea 55:291—293.
Holm, L.G., D.L. Plucknett, J.V. Pancho, and J.P. Her-
berger. 1991. The world’s worst weeds: distribution and
biology. Krieger Publishing Company, Malabar, FL.
[KSNPC] Kentucky State Nature Preserves Commission.
1996. Rare and extirpated plants and animals of Ken-
tucky. Trans. Kentucky Acad. Sci. 57:69-91.
Kral, R. 1983. A report on some rare, threatened, or en-
dangered forest-related vascular plants of the south.
Volume II. Aquifoliaceae through Asteraceae and glos-
sary. Technical Publication RS-TP2. United States De-
partment of Agriculture, Forest Service, Southeastern
Region, Atlanta, GA.
Libby, G.W. 1996. Inventory for Aureolaria patula
(Chapman) Pennell (Cumberland false foxglove) in
Kentucky. Pages 16-43 in D.L. White (project coordi-
nator). Inventory for five candidate (Category 2) plants
for federal listing. Tech. Rept., Kentucky State Nature
Preserves Commission, Frankfort, KY.
Lipscomb, B.L. 1980. Cyperus difformis L. (Cyperaceae)
in North America. Sida §:320-327.
McCoy, T.N. 1938. The ferns and fern allies of Kentucky.
Am. Fern. J. 28:41-46, 101-110.
McFarland, F.T. 1942. A catalogue of the vascular plants
of Kentucky. Castanea 7:77—108.
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perus (Cyperaceae) from Kentucky. Castanea 60:79-82.
Medley, M.E. 1993. An annotated catalog of the known
or reported vascular flora of Kentucky. Ph.D. disserta-
tion. University of Louisville, Louisville, KY.
Pennell, FW. 1935. The Scrophulariaceae of eastern
temperate North America. Acad. Nat. Sci. Philadelphia
Monogr. 1:1—650.
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Virginia. 2nd ed. Seneca Books, Morgantown, WV.
[USFWS] United States Fish and Wildlife Service. 1993.
Plant taxa for listing as endangered or threatened spe-
cies: notice of review. Fed. Reg. 58:51144-51190.
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and life: land character, plants, and animals of the Inner
Bluegrass region of Kentucky; past present and future.
University Press of Kentucky, Lexington, KY.
Trans. Ky. Acad. Sci. 58(2):80-84. 1997.
Effects of Acidic Minesoil on Nodulation of the Legume Sericea
Lespedeza (Lespedeza cuneata; Fabaceae) by Bradyrhizobium
Nitrogen-fixing Bacteria
Gary R. Cline and Anthony F. Silvernail
Community Research Service, Kentucky State University, Frankfort, KY 40601
ABSTRACT
Sericea lespedeza (Lespedeza cuneata (Dumont) G. Don; Fabaceae) is a N,-fixing legume used to
revegetate minespoils in Kentucky and elsewhere. A greenhouse experiment was conducted (1) to examine
the ability of sericea lespedeza to grow and form nodules in acidic minesoil and (2) to compare commercial
Bradyrhizobium sp. strains with strains isolated from minespoils regarding their abilities to form nodules
with lespedeza in acidic minesoil. Minesoil was adjusted to various pH levels with Al,(SO,), and CaCO,.
Although growth of sericea lespedeza was significantly reduced (P = 05) by soil acidification for some
inoculation treatments, the plants were able to grow relatively well under acid conditions even at a pH
level of 4.2. Nodulation was significantly (P = 0.05) suppressed by increasing acidity, particularly in
Al,(SO,),;-amended minesoil at pH levels of 4.2 and 4.5, which tended to affect nodulation similarly.
Bradyrhizobium sp. strains responded similarly to acidified minesoil with the exception that the com-
mercial mixture of strains tended to produce more numerous but smaller nodules than local strains isolated
from acidic minesoil.
INTRODUCTION
More than 30% of the total energy and over
half of the electricity produced in the United
States are generated by coal burning plants
(U.S. Bureau of the Census 1988). Tradition-
ally, Kentucky has been one of the three lead-
ing states in coal mining with more than
700,000 acres of surface-mined land having
been disturbed by 4900 mining operations in
the Appalachian and Eastern Interior Coal
Regions of the state (Office of Surface Mining
1988a, 1988b; U.S. Bureau of the Census
1988). Revegetation of surface-mined land is
necessary to reduce stream pollution, control
erosion and runoff, restore minesoil produc-
tivity, and improve wildlife habitats and for es-
thetic reasons (Roberts et al. 1988; Vogel
1981).
Minesoil acidity has been considered to be
the most important soil fertility factor limiting
revegetation of surface-mined land (Carpenter
and Hensley 1979; Vogel 1981). Minesoils be-
come acidic due to the oxidation of pyrite, a
process occurring when these geologic mate-
rials from lower depths are left exposed at the
surface after mining. Under acidic conditions
high levels of Al, Mn, and H-ions are consid-
ered to be the primary sources of toxicity to
plants, with Al toxicity generally considered
the most important (Foy 1984). Calcium and
80
phosphorus can become deficient in soils with
low pH levels and high concentrations of sol-
uble Al.
Nitrogen is normally deficient in newly-de-
posited minesoils (Reeder and Berg 1977;
Ebelhar et al. 1982), and effects of initial N
fertilization are short-lived (Carpenter and
Hensley 1979). Because continued fertiliza-
tion is often economically and physically im-
practical (Bender et al. 1985), plant produc-
tion becomes dependent on N, fixation by leg-
umes (Roberts et al. 1988). One of the major
deterrents to achieving desirable revegetation
rates on minesites is the slow establishment
and growth of N,-fixing legumes, which are
generally less tolerant of acidity than are non-
leguminous plants (Bender et al. 1988; Vogel
and Berg 1968).
Sericea lespedeza, Lespedeza cuneata (Du-
mont) G. Don (Fabaceae), is a warm-season,
perennial, forb-legume that has been used to
reclaim surface-mined lands in Kentucky. This
species is not planted so frequently as it has
been in the past, but it still has a place in rec-
lamation as wildlife cover. Although sericea
lespedeza is considered tolerant of soils with
pH levels as low as 4.5 (Cline and Senwo
1993; Vogel 1981), the tolerance of its N,-fix-
ing symbiosis to this pH range is unclear based
upon a limited number of qualitative obser-
Servicea Lespedeza and Minesoil Acidity—Cline and Silvernail 81
vations (Hyland 1938; Lyle and Evans 1979;
Plass and Vogel 1973; Vogel and Berg 1968).
Bacterial strains commonly used to inocu-
late lespedeza were developed to fix maximum
amounts of N, for a number of legume species
under more favorable growing conditions than
acidic minespoils. Regarding symbiotic N, fix-
ation and nodulation, it was reasoned that
strains isolated from sericea lespedeza growing
in acidic minesoil might be more tolerant of
such conditions than commercial strains. This
generally proved to be true of Bradyrhizo-
bium in pure culture (Cline and Senwo 1994).
The objectives of our study were (1) to ex-
amine the ability of sericea lespedeza to grow
and form nodules in acidic minesoil and (2) to
compare commercial strains with strains iso-
lated from acidic minesoils regarding their
abilities to form nodules in acidic minesoil.
MATERIALS AND METHODS
We conducted a pot experiment with four
acidity treatments and four inoculation treat-
ments using a completely randomized fectorial
design with eight replications. Inoc ilation
treatments consisted of (1) a noninoculated
control, which received inorganic N, (2) a
commercially available mixture of Bradyrhi-
zobium sp. strains (strain numbers 4122, 3223,
32H1, 150B1, and 176A22 from Liphatech
Inc., Milwaukee, WI, USA), and (3, 4) strains
41A and CRI, which were isolated from seri-
cea lespedeza growing on acidic minesoils
(Cline and Senwo 1994). Acidity treatments
consisted of minesoil collected from an aban-
doned minesite in Muhlenberg County, Ken-
tucky, with a natural pH level of 5.2 (untreat-
ed), minesoil adjusted to pH 6.4 with 1.6 g/kg
CaCO, (Ca-msoil), and minesoil adjusted to
pH 4.5 (1Al-msoil) or 4.2 (2Al-msoil) with 4.0
or 5.3 g/kg Al,(SO,);(H,O);, respectively.
Minesoil was collected to a depth of 15 cm,
sieved to a particle size <13 mm, and _air-
dried. A previous study indicated that minesoil
collected from this site was N-deficient (Cline
and Senwo 1993). The minesoil was supplied
with 20, 64, 32, and 5 mg/kg Ca, P, K, and
Mo, respectively. Following amendment with
CaCO, or Al,(SO,),, minesoil treatments were
equilibrated for 8 weeks at a moisture content
of 15 ml H,O/g soil. The equilibrated minesoil
was mixed with river sand (50:50 by volume),
and 160 g of the mixture were added to 165
ml conical containers (4 cm d X 21 cm Ray
Leach Supercells, Stuewe and Sons, Inc., Cor-
valis, OR). The mixture was used because les-
pedeza requires well-aerated soil, and prelim-
inary experiments indicated that the plant did
not grow well in pure minesoil due to aeration
problems associated with containers. Lespe-
deza grows well in minesoils under field con-
ditions.
‘Serala’ sericea lespedeza was grown from
seed in a greenhouse for 20 weeks under full
sunlight. Additional supplemental light (300
mol photon/m?/s photosynthetically active ra-
diation) was provided with halide lamps to
provide 16-hour photoperiods. Noninoculated
plants received NH,NO, at rates of 50 mg
N/kg minesoil after 4 weeks and 25 mg N/kg
minesoil after 10 and 15 weeks. All soils were
supplied with 5 mg K/kg minesoil and 1 mg
Mo/kg minesoil after 10 and 15 weeks. Three
milliliters of solution containing a total of 10°
colony-forming units of Bradyrhizobium sp.
were added to inoculated plants 2 and 4 weeks
after planting. Quantification of colony-form-
ing units was based upon relationships be-
tween optical density (600 nm) and colony
plate-counts determined for each bacterial
strain including individual strains in the com-
mercial mixture.
At harvest, minesoil was removed from
roots, and nodules were counted. Nodules and
shoots were dried separately at 65° C for 48
hours prior to weighing. Minesoil from each
pot was equilibrated with distilled H,O (0.6 ml
H,O g/soil) for 16 hours and centrifuged at
3000 X g. Supernatants were recentrifuged at
75,000 X g and filtered using Whatman No.
42 filter paper. Final pH values were mea-
sured prior to centrifugation. Toxic inorganic
Al in the supernatants was determined by the
pyrocatechol violet colorimetric method of
Kerven et al. (1988a, 1988b); phosphorus was
measured in the filtered extracts by the meth-
od of Murphy and Riley (1962). Atomic ab-
sorption was used to measure Mn, Mg, and
Ca.
Significance of main effects and interactions
regarding inoculation and acidification treat-
ments were determined by two-way analysis of
variance. Significant differences among treat-
ment means were defined by the Least Sig-
nificance Difference.
82
Table 1.
Mean values of acidity-related parameters (+SE,
Transactions of the Kentucky Academy of Science 58(2)
n = §) measured in extracts of minesoil acidity treatments
used to monitor growth of sericea lespedeza and nodulation by Bradyrhizobium nitrogen-fixing bacteria.
pH
Acidity treatment Initial
Ca-msoil 6.41 + 0.06 5.96
Untreated 5.33 + 0.03 5.15
1 Al-msoil 4.46 + 0.04 4.54
2, Al-msoil 4.18 = 0.05 4.24
RESULTS AND DISCUSSION
Soil Parameters
General characteristics of the minesoil have
been described by Cline and Senwo (1993);
parameters relevant to acidity treatments of
the present study are described in Table 1.
Values of initial and final pH remained rela-
tively stable throughout the experiment except
for Ca-msoil in which pH decreased by 0.4
units but remained near 6.0, an optimal pH
for plant and bacterial growth. Concentrations
of Mn and toxic Al increased with increasing
minesoil acidity. Extract concentrations of P
and Ca were ca. 2 uM and 5 mM, respectively,
in all treatments except that extracts of un-
treated minesoil contained only 1 mM Ca.
Thus, negative responses of lespedeza or Bra-
dyrhizobium to acidified treatments should
not be related to P or Ca deficiencies.
Shoot Dry Weights
Shoot dry weights (above-ground plant por-
tions) were generally greatest in Ca-msoil and
tended to decrease with increasing acidifica-
tion (Table 2). However, no chlorosis was ob-
served, and shoot dry weight differences be-
tween the 2Al-msoil (most acidic) and Ca-
msoil (least acidic) treatments were only sig-
nificant (P < 0.05) for strain 41A and the con-
2:
Table
Al Mn
Final (uM) (uM)
+ 0.06 <5 6+ 0
+ 0.04 Pha 1(0) 25) BE]
+ 0.06 85 + 3 186 + 10
+ 0.06 178 + 10 SOME 7
trol. The fact that sericea lespedeza was able
to produce significant yields in 2A]-msoil (pH
4.2) agrees with the findings of Cline and Sen-
wo (1993) but contrasts with those of Bengt-
son et al. (1973), who reported negligible les-
pedeza growth at pH 4.6. In Ca-msoil, dry
weight production in the N-fertilized control
treatment was highest and significantly (P =
0.05) greater than production in the commer-
cial and CR1 inoculation treatments. This sug-
gested that sericea lespedeza inoculated with
Bradyrhizobium probably did not receive suf-
ficient fixed N, for maximum growth, even at
the most optimal pH level (i.e., Ca-msoil).
Nodulation
Although large numbers of nodules were
formed (Table 3), individual nodules tended to
be relatively small, as indicated by their low
total dry weights (Table 4). No nodules were
observed on noninoculated control plants re-
ceiving inorganic N. Generally nodule num-
bers and dry weights from plants grown in
Ca-msoil were significantly (P = 0.05) greater
than numbers and weights obtained from the
three more acidic soil treatments (Tables 3
and 4). Strain CR1 formed similar numbers of
nodules in both untreated minesoil and Ca-
msoil, but nodule weight production was sig-
Mean values of sericea lespedeza mean shoot dry weight production (g + SE, n = 8) obtained in minesoil
acidity treatments using different strains of Bradyrhizobium nitrogen-fixing bacteria. Values in columns followed by
different letters were significantly (P < 0.05) different.
Acidity treatment Commercial 41A
Ca-msoil 9.94 + 0.17a 3.26 + 0.
Untreated 2.96 + 0.14a 9.92 + 0.
1 Al-msoil 2.69 + 0.14a eres ax: (0),
2. Al-msoil 2.67 + 0.17a Depa a (0),
Inoculation
CRI Control + N
18a 2.98 + 0.16ab Soon ana!
14ab 3.30 + 0.24a 2.98 + 0.40b
17be 2.63 + 0.08b 2.26 + 0.24c
14c 2.87 + 0.16ab 2.41 + 0.1lc
Servicea Lespedeza and Minesoil Acidity—Cline and Silvernail
Table 3.
83
Mean values of nodule numbers (+SE, n = 8) obtained for sericea lespedeza grown in minesoil acidity
treatments using different strains of Bradyrhizobium nitrogen-fixing bacteria. Values in columns followed by different
letters were significantly (P = 0.05) different.
OIE Sa NRE DFE OR Sep Ca UM ESA A 0 er a
Inoculation
Acidity treatment Commercial 41A CRI Control + N
Ca-msoil 53.0 + 6.9a 71.9 + 18.3a 52.4 83a 0.0a
Untreated PASO a Ht 49.3 + 6.5b 5210) Saliba 0.0a
1 Al-msoil Bont == 5.9b 27.9 + 7.0be 24.2 + 4.0b 0.0a
36.7 + 4.0b 26.5 + 4.6c 29.4 + 3.5b 0.0a
2 Al-msoil 9 ‘
nificantly (P = 0.05) higher in Ca-msoil (Table
3). Conversely, the commercial mixture of
strains and strain 41A produced similar nodule
dry weights in untreated minesoil and Ca-
msoil but formed significantly (P = 0.05)
greater numbers of nodules in Ca-msoil (Table
4). Thus, although nodulation was generally
decreased in untreated minesoil compared to
Ca-msoil, the response among strains differed
regarding nodule numbers and size.
In the more acidic 1Al-msoil and 2Al-msoil
treatments, nodule dry weight production was
significantly (P = 0.05) suppressed compared
to other treatments (Table 4). However, no-
dulation in these acidic treatments did occur,
agreeing with Hyland (1938) who observed
significant nodulation from pH 4.1 to 4.8. For
each inoculation treatment there were no sig-
nificant differences in nodule dry weights be-
tween 1Al-msoil and 2Al-msoil acidification
treatments, indicating that nodule dry weight
production was suppressed similarly by holt
treatments (pH 4.5 and 4.2, respectively).
Compared to nodulation at the more optimal
pH levels, nodulation in the more acidic treat-
ments was suppressed less than that reported
by Cline and Senwo (1993). For both 1Al-
msoil and 2Al-msoil treatments, no significant
differences were detected among nodule dry
Table 4.
Mean nodule dry weight production (mg + SE,
weights obtained from the three inoculated
treatments, suggesting that the strains were
equally tolerant of the acid conditions. How-
ever, in these treatments nodule numbers for
the commercial mixture of strains tended to
be significantly (P = 0.05) greater than num-
bers obtained for strains 41A and CRI, indi-
cating that the commercial strains tended to
form more but smaller nodules under these
acidic conditions. This could be important
since larger nodules would be expected to
contain greater amounts of nitrogenase per
gram of nodule due to surface area/volume re-
lationships. In previous pure culture experi-
ments, strains 41A and CRI appeared to be
more tolerant of acidic soil factors than com-
mercial strains (Cline and Senwo 1994),
whereas here they responded similarly in
terms of nodule dry weight production.
SUMMARY
Although growth of sericea lespedeza was
significantly reduced (P = 0.05) by acidifica-
tion in some inoculation treatments, the plants
were able to grow relatively well under the
acidic conditions even in 2Al-msoil at a pH
level of 4.2. Compared to Ca-msoil, nodula-
tion was significantly (P = 0.05) suppressed by
the more acidic treatments, particularly 1AI-
n = 8) obtained for sericea lespedeza grown in minesoil
acidity treatments using different strains of Bradyrhizobium nitrogen-fixing bacteria. Values in columns followed by
different letters were significantly (P = 0.05) different.
Inoculation
Acidity treatment Commercial 41A CR1 Control + N
Ca-msoil 13.9 + 2.0a 26.1 + 5.6a DAV Gy cuss Ae 0.0a
Untreated 13'5y221.9a 93.3, sei 3ella 14.2 + 2.3b 0.0a
1 Al-msoil e2) eb 8.0 + 1.4b 9.4 + l.le 0.0a
2, Al-msoil 8.9 + 1.3b 10.0 = 1.2b 9.3 + 1.0c 0.0a
84 Transactions of the Kentucky Academy of Science 58(2)
msoil and 2Al-msoil, which appeared to sup-
press nodulation similarly for individual inoc-
ulation treatments. In acidified minesoil, nod-
ulation was similar among Bradyrhizobium in-
oculation treatments with the exception that
the commercial mixture of strains tended to
produce more numerous but smaller nodules
than local strains isolated from acidic mine
spoils.
ACKNOWLEDGMENT
This research was funded by a USDA/
CSREES grant to Kentucky State University
under agreement KYX-10-89-11P.
LITERATURE CITED
Bender, D.A., R.D. Morse, J.L. Neal, and D.D. Wolf.
1985. Delayed inoculation and starter nitrogen for en-
hancing early growth and nitrogen status of Lespedeza
cuneata. Pl. Soil 84:311-321.
Bender, D.A., R.D. Morse, J.L. Neal, and D.D. Wolf.
1988. Field evaluation of starter N and delayed inoc-
ulation of Lespedeza cuneata grown in minesoil. Pl. Soil
109: 109-113.
Bengtson, G.W., S.E. Allen, D.A. Maya, and T.G. Zarger.
1973. Use of fertilizers to speed pine establishment of
reclaimed coal-mine spoil in northeastern Alabama: I.
Greenhouse experiments. Pages 199-225 in R.J. Hutnik
and G. Davis (eds). Ecology and reclamation of dev-
astated lands, Vol. 2. Pennsylvania State University
Press, University Park, PA.
Carpenter, P.L., and D.L. Hensley. 1979. Utilizing N,-
fixing woody plant species for distressed soils and the
effect of lime on survival. Bot. Gaz. 140:S76-S81.
Cline, G.R., and Z. Ngewoh Senwo. 1993. Inhibitory ef-
fects of acidic minesoil on the sericea lespedeza/Bra-
dyrhizobium symbiotic relationship. J. Pl. Nutr. 16:
1867-1880.
Cline, G.R., and Z. Ngewoh Senwo. 1994. Tolerance of
lespedeza Bradyrhizobium to acidity, aluminum, and
manganese in culture media containing glutamate or
ammonium. Soil Biol. Biochem. 26:1067—1072.
Ebelhar, M.W., R.I. Barnhisel, G.W. Akin, and J.L. Powell.
1982. Effect of lime N, P, and K amendments to sur-
face-mined coal spoils. Reclamation Rev. 1:327-336.
Foy, C.D. 1984. Physiological effects of hydrogen, alu-
minium and manganese toxicities in acid soil. Pages 57—
97 in F. Adams (ed). Soil acidity and liming. American
Society of Agronomy, Madison, WI.
Hyland, H.L. 1938. Comparison of legume growth in dif-
ferent soil types at varying acidity levels. J. Am. Soc.
Agron. 30:111-121.
Kerven, G.L., D.G. Edwards, C.J. Asher, P.S. Hallman,
and S. Kokot. 1989a. Aluminum determination in soil
solution. I. Evaluation of existing colorimetric and sep-
aration methods for the determination of inorganic mo-
nomeric aluminium in the presence of organic acid li-
gands. Austral. J. Soil Res. 27:79-90.
Kerven, G.L., D.G. Edwards, C.J. Asher, P.S. Hallman,
and S. Kokot. 1989b. Aluminum determination in soil
solution. II. short-term colorimetric procedures for the
measurement of inorganic monomeric aluminium in the
presence of organic acid ligands. Austral. J. Soil Res.
27:91-102.
Lyle, E.S., and E.M. Evans. 1979. Revegetation of Ala-
bama coal surface mines for soil cover and forage pro-
duction. Reclamation Rey. 2:55-61.
Murphy, J., and J.P. Riley. 1962. A modified single solu-
tion method for determination of phosphate in natural
waters. Anal. Chim. Acta 27:31-36.
Office of Surface Mining. 198Sa. Kentucky profile. U.S.
Dept. of the Interior. Washington, DC.
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forcement annual evaluation fiscal year 1987. U.S.
Dept. of the Interior. Washington, DC.
Plass, W.T., and W.G. Vogel. 1973. Chemical properties
and particle distribution of 39 surface-mined soils in
southern West Virginia. USDA Forest Serv. Res. Paper
NE-276.
Reeder, J.D., and W.A. Berg. 1977. Plant uptake of in-
digenous fertilizer from a Cretaceous shale and coal
mine spoils. Soil Sci. Soc. Am. J. 41:919-921.
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1988. Tall fescue production and nutrient status on
southwest Virginia mine soils. J. Environm. Qual. 17:
Do=O02:
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the United States. U.S. Dept. of Commerce, Washing-
ton, DC.
Vogel, W.G. 1981. A guide for revegetating coal mine-
spoils. USDA Forest Serv. Gen. Techn. Rep. NE-68.
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Trans. Ky. Acad. Sci. 58(2):85-91. 1997.
Structure and Composition of Three Swamp Forests on the
Mississippi Alluvial Plain of Kentucky’s
Jackson Purchase Region
William S. Bryant
Department of Biology, Thomas More College
Crestview Hills, KY 41017
ABSTRACT
Three swamp forests on the Mississippi alluvial plain of the Jackson Purchase Region of Kentucky are
described. One forest was dominated by Taxodium distichum, one by Nyssa aquatica, and one by T. distichum
and N. aquatica. In terms of basal areas and densities, these forests compared favorably to other little
disturbed swamps in the southeastern United States. Basal areas of 56.5, 72.8, 84.6 m*/ha were two to three
times greater than most mature upland forests of Kentucky. These swamps are remnants of a once more
abundant wetland system on the Mississippi alluvial plain.
INTRODUCTION
Bottomland forests, like other wetland types
in Kentucky and the southeastern United
States, have undergone great reductions in
area and changes in composition (Sharitz and
Mitsch 1993). Today, only 20% of Kentucky’s
originally estimated 650,000 ha of wetlands re-
main; only 20% of those remnants are forested
(KEQC 1992). The majority of forest rem-
nants are found in the Jackson Purchase Re-
gion, especially on the Mississippi alluvial
plain in Ballard, Carlisle, Fulton, and Hick-
man counties (KNPC 1986). Other remnants
occur eastward along the Ohio, Green, and
Tradewater rivers of the Western Kentucky
Coalfields Region (Braun 1950; Mitsch et al.
1983).
Bottomland forests, hardwoods, and deep-
water swamps were first reported from the
Jackson Purchase in William O. Henderson's
1820 General Land Office Survey (Bryant and
Martin 1988): however, since that time little
basic information on these vegetation types
has been added. Aside from a few early timber
surveys (DeFriese 1880, 1884; Loughridge
1888); county floras (e.g., Grubbs and Fuller
1991); a floristic list for Murphy’s Pond, a
swamp in Hickman County (Adams et al.
1968); and a report on nutrient dynamics for
that same swamp (King et al. 1989), the lit-
erature is limited. Here and elsewhere on the
Mississippi alluvial plain, few quantitative
studies of bottomland forests have been re-
ported (Mitsch et al. 1979). Specifically, infor-
mation is scarce on the relative abundance of
85
species, structural features, gradient analysis,
and diversity relations (Robertson et al. 1978).
In an attempt to fill in some of the information
gaps, Bryant (1991) analyzed and compared
several bottomland remnants in the Jackson
Purchase. Additionally, forest vegetation of the
loess bluffs, which form an eastern border
along the Mississippi alluvial plain, was re-
ported (Bryant 1993). Forested wetlands of
the Western Kentucky Coalfield have recently
been the subjects of ecological analysis (Ben-
son 1986; Cardamone 1985; Hill 1983; Mitsch
et al. 1983, 1991; Taylor 1985) and may be
better known.
The purpose of this paper is to describe and
quantitatively compare stand characteristics
for some deepwater swamp communities in
which baldcypress (Taxodium distichum) and/
or water tupelo (Nyssa aquatica) are major
contributors. Comparisons to other swamp
and bottomland forests are made, especially as
they relate to providing answers to the forest
structure and community composition.
THE ENVIRONMENT
The Mississippi alluvial plain is a division of
the larger Coastal Plain Province (Fenneman
1938). In western Kentucky, it is the broad
historic and recent floodplain of the Mississip-
pi and Ohio river systems. Most of the allu-
vium that covers the floodplain has been de-
posited since the Pleistocene. The Mississippi
alluvial plain’s aggraded surface is marked by
river meanders and oxbow lakes, natural lev-
ees and backswamp areas, and shallow sloughs
86 Transactions of the Kentucky Academy of Science 58(2)
and marshes (Powers 1966). In their natural
state the low gradient streams that flowed
through the bottomlands were sluggish and
widely meandering, carrying a large sediment
load.
As a result of microtopographical differ-
ences on the floodplain, a mosaic of forested
swamps and bottomland hardwoods occurs
across the bottomlands. Flooding frequency,
hydroperiod, and depth of flooding are factors
that work with the microtopography to sort
out the communities. The forested swamps
occur in long, narrow, often curving sloughs,
oxbow lakes, bayous, and stream margins. Lev-
ees, channelization, and drainage may have
modified flood plains and altered modem
flooding regimes.
Stands of baldcypress and water tupelo usu-
ally have a long hydroperiod; however, a fall
drydown of water often occurs (Wharton et al.
1982). Soils are generally saturated for most
of the year.
METHODS
Efforts were made to locate swamp forests
that showed few signs of recent human dis-
turbance (e.g., stumps). I sampled three
swamps during the fall drydown period in Oc-
tober 1989. At that time it was possible to walk
or wade into the interior of each swamp. Two
of the swamps were in Ballard County (Oscar
Bottoms area and Fish Lake area) and one was
in Carlisle County (Back Slough area). All
were in recognized wetland priority areas
(KNPC 1986). Circular plots (0.04 ha), spaced
at 30 m intervals, were located through the
interior of each swamp. No samples were
taken near the swamp edge. The number of
plots sampled per swamp (or the size of the
area sampled) was to equal or exceed that
sampled by Anderson and White (1970) in
their study of a baldcypress swamp in southern
Illinois. Based on species area curves the num-
ber of plots taken per stand was sufficient. Six
plots were taken at Fish Lake, 10 at Back
Slough, and 12 at Oscar Bottoms.
All trees =10 cm at 1.4 m above the ground
(dbh) were measured in each plot. Because of
the swollen buttresses of baldcypress, Ander-
son and White (1970) proposed measuring
these trees at 2.1 m to make the basal areas
more comparable to those in other forest com-
munities. They found a linear relationship be-
tween baldcypress diameters at 1.4 m (normal
dbh) and 2.1 m. The 2.1 m dbh was about
90% (Y = .90X — 2.9) of the 1.4 m dbh. All
1.4 m dbh values for baldcypress were con-
verted to the 2.1 m values based on the re-
eression formula of Anderson and White
(1970), and basal areas for both heights were
determined. Water tupelo has swollen bases
also, but no diameter conversions were. at-
tempted for that species.
Tree data were analyzed to relative density
(RD), relative dominance (RDo), importance
value (IV), and importance percentage (IP).
IP, determined as IV/2, was used for compar-
isons to 100. Density (trees/ha), basal area
(m?/ha), species diversity (H’), and evenness
(J) were determined for each swamp forest.
Species diversity was calculated using the
Shannon-Weaver function:
H' = > (p,)(log,p;)
where s is the number of species, and p, is the
proportion of the total sample belonging to
the ith species (Shannon and Weaver 1949).
The similarity coefficient (C; Bray and Curtis
1957) was used to compare the swamps of this
sample and to compare these swamps to oth-
ers in the literature. The equation for calcu-
lating similarity is C = (2w)/(a + b), where a
equals the sum of the IVs or IPs for Swamp
One, b equals the sum of the IVs or IPs for
Swamp Two, and w is the sum of the lower
values for the species that occur in both sam-
ples. The equitability (evenness) of the stand
(J) was calculated using a method based on the
Shannon-Wiener function (H'): J = H’/Hmax,
where Hmax is the diversity under maximum
equitability (H’ = log,S), and S is the number
of species in the sample.
RESULTS
According to the dominant species, the
swamps sampled were classified as baldcypress
(Back Slough), water tupelo (Fish Lake), and
baldcypress-water tupelo (Oscar Bottoms). In
terms of composition and stand structure (IP),
the three swamps showed low degrees of sim-
ilarity, C = <2% to >42% range. Similarity
was lowest between the baldcypress and the
Kentucky Swamp Forests—Bryant
Table 1.
87
Number (N), relative density (RD), relative dominance (RDo), importance value (IV), and importance
percentage (IP) for Black Slough, Carlisle County, Kentucky.
N RD
Taxodium distichum 17] 92.93
Planera aquatica 9 4.89
Fraxinus profunda 2 1.09
Salix nigra ] 0.54
Forestiera acuminata 1 0.54
Total 184 99.99
water tupelo communities, which were nearly
monospecific stands.
Of the five tree species in the baldcypress
swamp (Table 1), baldcypress accounted for
93% of the density and >99% of the basal
area. There were 454.5 trees/ha with a basal
area of 116.5 m?/ha. The converted basal area
(Anderson and White 1970) was 72.8 m?2/ha.
Because of the near monospecific nature of
this stand, diversity (H’ = 0.46) and evenness
(J = 19.99) were low. Baldcypress knees were
abundant, and fallen logs were occasional on
the floor of this swamp, but no standing dead
trees were observed.
Cephalanthus occidentalis (buttonbush) was
the most abundant shrub in the swamp. Herbs
that occurred on baldcypress buttresses, fallen
logs, and drydown soils included Bidens dis-
coidea, Boehmeria cylindrica, H ypericum wal-
teri, Lycopus rubellus, Cuscuta sp., and Sau-
rurus cernuus. Anderson and White (1970) re-
ported a similar but more extensive list of
herbs in southern Illinois. Because the sam-
pling timeframe was limited for my study, little
effort was made to thoroughly characterize the
herb layer of this and the other swamps. Hei-
neke (1987) presented a more complete listing
of plants from the middle portion of the Mis-
sissippi alluvial plain.
The water tupelo swamp contained only
three tree species (Table 2). Water tupelo con-
tributed 95% of the density and >94% of the
stand basal area. There were 526.9 trees/ha.
Table 2.
RDo IV IP
99.05 191.98 95.99
0.30 5.19 2.60
0.07 1.16 0.58
0.54 1.08 0.54
0.03 0.57 0.29
99.99 199.98 100.00
Basal area was 85.3 m2/ha; converted basal
area was 84.6 m?/ha. Species diversity (H’ =
0.36) and evenness (J = 22.71) were low in
this nearly pure stand. Buttonbush was the
only commonly encountered shrub.
Twelve tree species were present in the
baldcypress-water tupelo swamp (Table 3).
Baldcypress and water tupelo accounted for
70% of the 69.9 m?/ha basal area. The con-
version value was 56.5 m?/ha. There were
403.4 tress/ha; however, baldcypress and water
tupelo collectively accounted for only 35% of
the total density.
Other species, Quercus lyrata (overcup
oak), Fraxinus pennsylvanica (green ash), and
Acer saccharinum (silver maple) were major
associates, especially in regard to their com-
bined density contributions. The other seven
tree species reflected the hydric nature of the
site; their presence assured a higher diversity
(H’ = 2.84) relative to the other two swamps.
The contributions of the several species were
reflected by a greater evenness (J = 79.3). The
hydroperiod may differ from that of the other
two swamps sampled. Barry (1980) found that,
with decreases in flooding depth and duration,
more codominants appear.
Average density for the three swamps sam-
pled was 461.6 trees/ha; average basal area
was 90.6 m?/ha; and average converted basal
area was 71.3 m?/ha. Those values are com-
parable to other little disturbed swamp forests
in the southeastern United States (e.g., An-
Number (N), relative density (RD), relative dominance (RDo), importance value (IV) and importance
percentage (IP) for the Fish Lake Swamp Forest, Ballard County, Kentucky.
N RD
Nyssa aquatica 121 94.53
Taxodium distichum 3 2.34
Quercus lyrata 4 3.14
Total 128 100.00
RDo IV IP
94.30 188.83 94.42
1.56 3.90 1.95
4.13 7.26 3.63
99.99 199.99 100.00
88 Transactions of the Kentucky Academy of Science 58(2)
Table 3.
Number (N), relative density (RD), relative dominance (RDo), importance value (IV), and importance
percentage (IP) for the Oscar Bottoms Swamp Forest, Ballard County, Kentucky.
N RD RDo IV IP
Taxodium distichum 4] 20.92 40.39 61.31 30.66
Nyssa aquatica 28 14.29 29.61 43.90 21.95
Quercus lyrata 45 22.96 11.19 34.15 17.08
Acer saccharinum Bho 16.84 5.79 22.63 RES?)
Fraxinus pennsylvanica 25 12.76 6.43 19.19 9.60
Ulmus americana 8 4.08 0.40 4.48 2.24
Populus heterophylla 6 3.06 1.32 4.38 2.19
Populus deltoides 2 1.02 3.22. 4.24 2.12
Planera aquatica 4 2.04 0.12 2.16 1.08
Platanus occidentalis 2 1.02 0.92 1.94 0.97
Salix nigra ] 0.51 0.57 1.08 0.54
Forestiera acuminata ] 0.51 0.03 0.54 0.27
Total 196 100.01 99.99 200.00 100.02
derson and White 1970; Penfound 1952; Rob-
ertson 1987; Schlesinger 1978).
Size class distributions for baldcypress and
water tupelo followed bell-curves, suggesting
even-aged stands (Figures 1, 2). No trees were
cored for age determinations; however, rough
Number of Stems
0 30 60 90 120
Diameter Size Classes (cm.)
Figure 1. Size-class distributions for baldcypress (Taxo-
dium distichum) at Back Slough and for water tupelo
(Nyssa aquatica) at Fish Lake in the Jackson Purchase
Region of Kentucky. Gray data points = baldcypress;
black data points = water tupelo.
age estimates were obtained by applying av-
erage diameter growth rate (3.25 mm/yr) for
baldcypress at Hovey Lake, Indiana (Kozel et
al. 1989) to baldcypress trees at Oscar Bot-
toms and Back Slough. Maximum ages were
322 yr and 416 yr, respectively. Age estimates
using Mattoon’s (1915) “rule of thumb” that
12 inches (30.5 cm) in diameter equals 100
years gave similar, but somewhat greater ages.
Results of either estimate add support that
these are long-established stands.
DISCUSSION
Baldcypress and water tupelo, either singly
or in association, are clearly the dominant tree
species at the hydric end of the moisture gra-
dient on the Mississippi alluvial plain in Ken-
tucky’s Jackson Purchase Region. Deepwater
Number of Stems
Diameter Size Classes (cm.)
Figure 2. Size-class distribution for all trees of baldcy-
press (Taxodium distichum) and water tupelo (Nyssa
aquatica) in the Oscar Bottoms Swamp, Ballard County,
Kentucky. Gray data points = baldcypress; black data
points = water tupelo.
Kentucky Swamp Forests—Bryant 89
swamps are covered by water throughout most
of the year (Penfound 1952). Basal areas (m?/
ha) and densities (trees/ha) for the three
swamps sampled compared favorably to other
southeastern United States swamp forests
(e.g., Schlesinger 1978), but basal areas greatly
exceeded those reported for the Western
Coalfield Region (Mitsch et al. 1991). Swamp
forests tend to have significantly greater stand
densities (Nelson 1997) and basal areas than
most regional upland forests including such
highly productive forests as the coves of the
Smoky Mountains (Whittaker 1966). Basal ar-
eas, converted or not, are two to three times
greater than those of mature upland forests in
eastern North America (Held and Winstead
1975; Martin 1992) (Table 4). Tree species di-
versities are low (Robertson 1987). Mitsch et
al. (1991) noted that complexity indices of cy-
press-dominated systems were low.
Braun (1950) observed that bottomlands
from Louisiana northward to southern Illinois
show a great floristic uniformity. An ecological
comparison of the swamps in my study was
made to a swamp in Louisiana (Conner and
Day 1970) and one in southern Illinois (An-
derson and White 1970). Maximum similarity
values were only 58% among those locations.
Bryant (1991) reported a 51% similarity be-
tween a bottomland hardwood stand in Ken-
tucky and one in Louisiana. Those similarities
are rather low and indicate some differences
in composition but especially in the contribu-
tions (IVs or IPs) of species in common be-
tween sampling sites along this north-south
gradient. Other differences may reflect the lo-
cation of sampling plots. Anderson and White
(1970) reported red maple (Acer rubrum)
from a southern Illinois swamp, but in western
Kentucky red maple was a tree of swamp mar-
gins, not of their interiors (Bryant, unpub-
lished data). Guthrie (1989) also noted the ap-
parent absence of water tupelo from Reelfoot
Lake, Tennessee, a swamp just south of the
Jackson Purchase sites. In regard to hard-
woods, Bryant (1991) reported higher impor-
tances for pecan (Carya illinoinensis), other
hickories, and oaks in Kentucky than in bot-
tomlands further south. Braun (1950) may be
correct in regard to floristic similarity, but ve-
getationally there appears to be a greater vari-
ability in the bottomlands.
Even-aged, densely stocked monospecific
Table 4. Basal areas (m?/ha) for selected upland and
lowland forests in Kentucky for comparison to swamps of
the Jackson Purchase Region of Kentucky.
Lilley Cornett Woods (Martin 1975)
Sugar maple-Basswood-Tulip poplar
Hemlock
Chestnut oak
Beech-White oak
Lilley Cornett Woods (Muller 1982)
Old growth composite
30.0 m?/ha
42.4 m*/ha
29.1 m?/ha
28.8 m?/ha
27.8 m2/ha
Bonayer Forest (Bougher and
Winstead 1974)
Oak forest 31.9 m2/ha
Boone County Cliffs Nature Preserve
(Bryant 1978)
Total forest 27.2 m/ha
Lloyd Wildlife Preserve Forest
(Bryant 1985)
Mixed forest 35.5 m2/ha
Dinsmore Woods Nature Preserve
(Held and Winstead 1976)
Oak-ash-maple forest
Myers Woods (Held 1980)
Beech forest
28.1 m?/ha
37.1 m?/ha
Greenwood Forest (Chester
et al. 1995)
Wet woods 37.8 m?/ha
Upper Bottoms (Bryant 1991)
Sugarberry forest
Martin Creek (Martin 1983)
Southern red oak
35.6 m2/ha
29.2 m2/ha
Cypress Swamp (Mitsch
et al. 1991)
Stagnant swamp
Semi-permanent flooded
35.9 m?/ha
32.7 m2/ha
stands of baldcypress or water tupelo appear
to be most common in deep water (Anderson
and White 1970; Johnson and Shropshire
1983). Monospecific stands of water tupelo
have been attributed to past logging and re-
moval of baldcypress (Robertson 1987). That
was not confirmed in my study, although
stumps were present in areas adjacent to sam-
pling sites. Water depth and hydroperiod play
extremely important roles in determining spe-
cies composition on the alluvial plain. Bald-
cypress-water tupelo forests may support re-
cruitment of mixed bottomland species follow-
ing drought or draining (Sharitz and Mitsch
1993). Swamp sites with shallower and more
90 Transactions of the Kentucky Academy of Science 58(2)
variable water depths may reflect this but ar-
eas where flooding is relatively constant are
usually dominated by baldcypress (e.g., Back
Slough) or water tupelo (e.g., Fish Lake). The
low numbers of standing dead trees at the
three swamps suggests that their compositions
are relatively stable and not undergoing com-
positional shifts as was found in the Western
Kentucky Coalfield (Mitsch et al. 1991).
Estimated ages of selected trees in the
swamps indicate that they are remnants of
long-established swamp systems. Baldcypress
trees with >1.m dbh were not unusual, but
none as large as the 2.3 m dbh reported by
DeFriese (1880) were found.
In their study of bottomland forests at
Horseshoe Lake, southern Illinois, Robertson
et al. (1978) found increased dominance, low-
er equitability, and decrease in number of spe-
cies of intermediate importance from the me-
sic to the wet segment of the moisture gradi-
ent. They attributed that reduction in diversity
to increased flooding and poorer drainage.
Wharton et al. (1982) referred to this as an
anaerobic gradient. Robertson et al. (1978)
stated, “The complex gradient acts as a filter
which effectively screens a number of species.
This filtering action continues as flooding in-
creases, until swamp forests dominated by T.
distichum and N. aquatica prevail.” That ap-
parently is also the situation on the Mississippi
alluvial plain in the Jackson Purchase Region.
Ewell and Odum (1984) stated, “Under-
standing wetland ecosystems is a major goal of
contemporary ecological sciences.” Wetlands
continue to be lost in Kentucky. It has taken
over 175 years for baldcypress and water tu-
pelo swamp forests to be described for the
Jackson Purchase Region. More research on
Kentucky's diminishing wetlands is needed so
that the roles these valuable ecosystems play
in the land-water interface may be better un-
derstood.
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Planning a Trip
John S. Spraker, Daniel C. Biles, Mark P. Robinson
Department of Mathematics
Western Kentucky University
Bowling Green, KY 42101
ABSTRACT
The problem of minimizing the driving time for a trip is considered. A hypothetical trip from Louisville
to Memphis is used as an illustration. A mathematical model is constructed and a solution technique is
devised and implemented.
DESCRIPTION OF THE PROBLEM
We consider a hypothetical trip from Lou-
isville to Memphis. We suppose the driver
wishes to depart no earlier than 1100 and de-
sires to arrive between 1745 and 1945, say, to
attend a meeting or presentation scheduled
for 2000. The driver wishes to select a depar-
ture time that minimizes total travel time. We
assume the driver must travel 20 miles
through Louisville at a speed of 55 mph, ex-
cept for lunch hour from 1200 to 1300, when
speed is restricted to 45 mph, and for “rush
hour” from 1600 to 1800, when speed is held
to 30 mph. The motorist can then drive at 65
mph over the next 150 miles to Nashville, ex-
cept for a 5-mile stretch near Elizabethtown,
Kentucky, on which the speed is limited to 40
mph from 1600 to 1700 due, say, to traffic
from a plant closing. The driver then travels
25 miles through the Nashville area, with the
same restrictions as the Louisville area. There
follows a 155-mile stretch of highway on which
the motorist can proceed at 65 mph, except
for a 15-mile long construction area beginning
40 miles outside of Nashville. In this area, the
speed is restricted to 20 mph until 1700 after
which speeds are held to 55 mph. Finally, the
driver must proceed 20 miles through the
Memphis area, with the same speed limits as
in Louisville and Nashville.
Suppose we establish a departure time of
1200. Consider Figure 1. At that departure
time we are restricted to 45 mph because of
lunch traffic. At 1227 we have travelled for 20
miles and reached the outskirts of Louisville,
at which point we can travel at 65 mph for a
150-mile stretch. We then hit the Nashville
area at 1445 where we must slow to 55 mph.
Continuing in this fashion, we obtain the re-
sults in Table 1.
92
RESULTS
Wishing to obtain the starting time resulting
in the shortest travel time, we select possible
starting times spaced at regular intervals and
calculate the travel time for each. Proceeding
as above for each of these times would be pro-
hibitively time-consuming. Therefore, we have
written a computer program that calculates
the arrival time and travel time for each of the
possible starting times. (See Section 3 for de-
tails.) We thus generate the results in Table 2.
The output is restricted to those trips for
which the arrival times satisfy the given con-
straints. Thus we see that the optimal depar-
ture time is 1300.
THEORY AND IMPLEMENTATION
We now consider the general problem of
minimizing the driving time for a trip. One has
a set of acceptable departure times S = {t,, f,,
t,,...}, wheret, <t, <t, <. .. (For example,
S might be {1500, 1530, 1600 ...}.) One
would also typically have an earliest desired
arrival time T, and a latest desired arrival time
T,. Of course, we require t) < T, < T,. One
also has constraints on speed. These involve
not only the posted limits but traffic conges-
tion caused by factors such as rush hour in a
large town or restricted speed limits due to
school zones or construction. Note that these
factors are typically dependent on both time
and position. Using these considerations, we
can construct an ordinary differential equation
x'(t) = f(t, x(t)), where t is the time after de-
parture and x(t) is the distance travelled at
time t. Finally, we let D be the travel distance.
We assume that the acceleration/deceleration
times between speed zones are negligible. We
also assume that the driver travels at the max-
imum speed possible at all times. We note that
Planning a Trip—Spraker, Biles, and Robinson 93
Distance
oe
5 °
55
170
65
20 45
Time
1200 1227 1445
Figure 1. Planning a trip. Illustration of the trip starting
at noon.
these assumptions force f to be discontinuous.
Removal of these restrictions is discussed in
the last section. Letting f represent the de-
parture time and T the arrival time, we thus
obtain the following model:
Choose ¢ so that T — t is minimized subject to
ee) = ft, x@)),t € [¢, 7],
cae = (0)
Cae
1 Bess ISS he
blenSat — L
Since f is discontinuous, a differentiable solu-
tion may be impossible to find. For example,
consider
Oke aa:
: Fe 8)
if 2
x(0) = 0.
One is tempted to propose
0,
Ws — 1,
as a solution; however, x'(1) does not exist and
hence x does not satisfy the differential equa-
tion. As a result, we must modify the classical
definition of solution.
te ()) = == ol
ele pes
Definition 1: x: [¢, T] > R is a solution to
ct) =f <@) ete fe, Tl
(E)) = 4a
Table 1. Planning a trip. Calculation of the trip time for
the trip starting at 1200.
Time of day Distance travelled
1227 20
1445 170
1512 195
1549 935
1634 250
1806 350
1828 370
if and only if i) x is continuous on [f,T], ii) x’ (¢)
= f(t,x(t)) for all but a finite subset of [67],
iti) x(£) = Xp.
Actually the definition of solution that is
standard for discontinuous differential equa-
tions involves Lebesgue measure theory; the
interested reader can consult Chapter 2 in
(Coddington and Levinson 1955). Our velocity
constraint function f is assumed to satisfy the
following.
Definition 2: f : [t, T,) X [0,D) > R is
blockwise-constant if and only if there exists a
finite collection of disjoint sets A, = {(t, x) : a,
=¢ =D) c= x <d,) suchsthat UA, = ([tauls)
x [0,D) and f |,, is constant for each i.
Example 1: Consider f: [0, 1) x [0, 30) > R
defined by
il
15 foe ee
fit, x) — 4 2
35 elsewhere
This f could model a 30-mile stretch of high-
way on which the speed limit is 35, except for
Table 2. Planning a trip. Output from travel problem
code.
Starting time Arrival time Travel time
1UT2 1754 6 hr 42 min
1124 1803 6 hr 39 min
1136 1810 6 hr 34 min
1148 1817 6 hr 29 min
1200 1828 6 hr 28 min
1212 1840 6 hr 28 min
1224 1852 6 hr 28 min
1236 1858 6 hr 22 min
1248 1901 6 hr 13 min
1300 1907 6 hr 7 min
1312 1923 6 hr 11 min
1324 1942 6 hr 18 min
OPTIMAL TRIP LENGTH = 6 hr 7 min
94 Transactions of the Kentucky Academy of Science 58(2)
a school zone from mile 5 to mile 6 on which
the speed limit is 15 from t = 4 tot = %.
Before we consider numerical approaches,
we must ensure that our model is well-posed.
Theorem 1: Let f : [t,, T,) X [0,D) > R be
blockwise-constant and positive-valued. Let f
€ S, t < T,. Then, one (and only one) of the
following must hold:
i) There exists T € (,.T,|,.x: [& T] 3-R
such that x is a solution to x’(t) = f(t,
x(t)) on [é,T], x) = 0 and. x(T) = D; or
ii) there exists x : (&T,] — R such that x is
a solution to x’(t) = f(t,x(t)) on [£T,],
x(t) =.0-and x(F,)< D.
The proof of Theorem | is straightforward
and is left to the reader.
Theorem 2: Let f : [t), T,) X [0,D) > R be
blockwise-constant and positive-valued. Let t
e S,t < T,. Suppose x is a function satisfying
one of the two conclusions of Theorem 1. Let
x be continuous, <'(f) = f(éx(t)) for all but a
finite subset of the domain of < and assume
there exists some f such that x(f ) = £(f). Then,
x(t) = x(t) for all t in the intersection of the
domains of x and x.
Proof. Claim 1: x and % agree to the right
of -¢, that is, 1f-¢ e.[ET)swhere © represents
the right-hand endpoint of the intersection of
the domains of x and «, then x(t) = x(t) for all
Pelen:
Proof of Claim 1: Suppose not. Then, with-
out loss of generality, there exists some T €
(£,T] such that <(t) > x(t). Let t* = sup{t €
[t, t) | x(t) = <(t)}. Since x and & are contin-
uous, we have x(t*) = x(t*) and x(t) > x(t) for
all t € (t*,t]. There exists some 7 € (t*,T) such
that (t,x(t)) and (t,x(t)) € interior of some A,
for all t € (¢*,#). Let k, represent the constant
value of f on A,. Then, for t € (t*,+), we have
nee") = tk ee ei ee
= x(t), a contradiction.
Claim 2: x and ¥ agree to the left of t, that is,
if f € (ET], where T represents the right-hand
endpoint of the intersection of the domains of x
and «, then x(t) = x(t) for all t € [& f].
Proof of Claim 2: Analogous to the proof of
Claim 1.
Corollary 1: The function x guaranteed by
Theorem 1 is unique.
As a result of Theorems 1 and 2, for each
fe S,t < T,, the numerical procedure has one
and only one solution to approximate.
For each of the starting times, the numer-
ical method employed in this study is an adap-
tive version of Euler’s method. Given an initial
value problem (IVP)
Ke = fit, x),
@a=t=b,
x(a) =O;
Euler's method, discussed in any elementary
numerical analysis textbook (see, for example,
(Burden and Faires 1997)) generates an ap-
proximate solution to the IVP at a set of dis-
erete.points T,, where @ = t).— 7 =... Ty
= b, by approximating the solution x(t) locally
by the first two terms in its Taylor series. Eu-
ler’s method produces an approximation u, to
the exact solution x(t,) at the mesh points ac-
|
cording to the iteration
Uo ai Q,
We Wat Dg rae Wn)
PHIM
where h, = 1, — 1;_;. Euler's method is in fact
exact if the true solution x(t) is linear through-
out the interval [a,b]. It is an immediate ex-
tension of this fact that Euler’s method will
also produce the exact answer if the exact so-
lution is merely continuous and piecewise lin-
ear with respect to the mesh {1,},‘,(that is, the
only changes in slope occur on some subset of
the set of mesh points {7},%,). In our travel
problem, the function f(t,x) is blockwise con-
stant, and therefore the exact solution x(t) is
continuous and piecewise linear. As a conse-
quence, it is possible to obtain the exact so-
lution (aside from roundoff errors) from the
use of Euler’s method, so long as suitable ad-
justments are made. For example, a given
time step may need to be modified to avoid
overshooting a block.
SUMMARY
We have considered the problem of mini-
mizing the driving time of a trip by modeling
the problem with a discontinuous differential
equation. Although it is straightforward (yet
tedious) to solve the differential equation by
hand, efficient solution of the minimization
problem may be accomplished through re-
peated application of Euler’s method on a
Planning a Trip—Spraker, Biles, and Robinson 95
computer. We finally consider a few ideas for
further work. One could include the fact that
a car has maximum acceleration and deceler-
ation, say, by adding the constraint x"(t) €
{—d,0,c} or, x"(t) € [—d,c], where c,d are pos-
itive constants. Thus, one would need to con-
sider differential inclusions rather than differ-
ential equations. One would now require x'(t)
€ [0,f(¢,x(t))] rather than x’(t) = f(t,x(t)), since
with acceleration constraints one may be un-
able to always drive at the speed limit. The
subject of differential inclusions is not an easy
one to get into; a well-known resource is (Au-
bin and Cellina 1984). Also, one may wish to
consider problems in which the driver may
have multiple routes from which to choose. An
application of graph theory may be necessary.
This problem can be extended further by at-
tempting to choose the ideal route and depar-
ture time given certain driver preferences. For
example, the driver may be interested in the
best route and departure time to get from A
to B, but may wish to pass through C on the
way. The reader is invited to use his or her
imagination to add even more twists along
these lines. Finally, we mention that it would
be worthwhile to consider shapes other than
rectangles to subdivide the domain of f. For
example, consider a school zone that drops the
speed limit from 35 mph to 15 mph at 1530.
In the blockwise-constant model, the driver
who enters the school zone at 1529 at 35 mph
will immediately realize at 1530 that he or she
should slow to 15 mph as long as he or she
remains in the school zone. Realistically, the
warning that the speed limit has changed from
35 mph to 15 mph may only be given at the
start of the school zone. In such a case, the
driver mentioned above would continue at 35
mph rather than 15 mph, having no warning
of the speed limit change. The reader may
convince himself or herself that the proper
shape in the tx-plane for this f is a trapezoid
or a triangle.
LITERATURE CITED
Aubin, J.P., and A. Cellina. 1984. Differential inclusions.
Springer-Verlag, Berlin.
Burden, R.L., and J.D. Faires. 1997. Numerical analysis.
6th ed. Brooks/Cole, Pacific Grove, CA.
Coddington, E.A., and N. Levinson. 1955. Theory of or-
dinary differential equations. McGraw-Hill, New York,
NY.
Trans. Ky. Acad. Sci. 58(2):96-100. 1997.
NOTES
Blue Catfish (Ictalurus furcatus; Ictaluridae)
Predation on the Zebra Mussel in the Ohio River
near Paducah, Kentucky.—Blue catfish (Jctalurus fur-
catus) were examined to determine whether they are
predators of the exotic zebra mussel (Dreissena polymor-
pha) in the Ohio River. Predation on zebra mussels by
diving ducks, crayfish, and several fish species other than
blue catfish has been described (1, 2, 3, 4). Cyprinids such
as the roach (Rutilis rutilis), bull chub (Nocomis raneyi),
and common carp (Cyprinus carpio) are predators of ze-
bra mussels (5, 6, 7). Other fishes with molariform pha-
ryngeal teeth or chewing pads—e.g.,
eed ear Sch (Lepomis micro-
freshwater drum
(Aplodinotus , grunniens )
lophus), and ealilnoutls buffalo (Ictiobus bubalus )—are
considered predators of the mussels in North America (4).
A common species in large rivers of southeastern Unit-
ed States is the blue catfish. Blue catfish lack pharyngeal
teeth but do feed on molluscs (8S). Members of the Ictal-
uridae (catfish family) can feed on bivalves by swallowing
them whole. French (1993) noted a zebra mussel with
val
On 1 Oct 1995 a commercial fisherman on the Ohio River
ves intact in a brown bullhead (Ameiurus nebulosus).
told us that he observed zebra mussel shells in the stom-
achs of blue catfish. To confirm this observation we chose
to examine blue catfish from a fish market that processed
fish from the Ohio River.
Blue catfish were sampled on 4, 6, and 7 1995 from
Three Rivers Fish Market in Paducah, The
sample consisted of 150 individuals with lengths ranging
Nov
Kentucky.
from 25 to 50 cm. Esophagus, stomach, and intestine were
removed from each fish and examined for the presence of
zebra mussels. The fish had been caught in the Ohio River
near Paducah between Smithland Dam and Dam 52; exact
site information was not available from the commercial
fishermen. A total of 1473 bivalves was recovered from 27
of the 150 catfish.
Asian clam
Three species of mollusks were present:
(Corbicula fluminea), fawnsfoot (Truncilla
donaciformis), and zebra mussel.
Blue catfish with lengths of 28 to 33 cm were found to
contain the majority of bivalves and consisted of 58% (87) of
the sample. Of the 87 stomachs, only 31.0% (27) of individ-
Eighteen stomachs
2.2%
24.4
uals of this size had fed on bivalves.
) contained
zebra mussels. One stomach that had zebra mussels also had
(66.7%) contained only Asian clams; six (2
three fawnsfoot mussels with lengths of 7, 11, and 12 mm.
Three (11.1%) blue catfish with lengths of 29, 30, and 32 em
had both Asian clams and zebra mussels. This size grouping
of blue catfish had 72.6% (1040) of the total number of Asian
clams and 94.6% (35) of the total number of zebra mussels.
In the three species of bivalves the hinge was intact, the shells
were whole, and some visceral tissue was present. A total of
1433 Asian clams was found with lengths between 1 and 12
mm. Asian clam lengths of 3 to 7 mm comprised 92.9%
(1331) of the total number of Asian clams found in the stom-
achs. Thirty-seven zebra mussels were collected with lengths
between 3 and 15 mm. Zebra mussel lengths of 3 to 7 mm
96
comprised 78.4% (29) of the total number of zebra mussels
found in stomachs. Our observations on zebra mussels in the
Ohio River at Dam 52 provide a length range of 3 to 24 mm
with the most common lengths of 12 to 20 mm.
Our study confirms that blue catfish do feed on zebra
mussels. The degree of predation and effects of this pre-
dation on the Ohio River zebra mussel population are un-
known. However, our findings suggest that there may be
a size selection (3-7 mm) occurring when blue catfish feed
on zebra mussels and Asian clams. Also, there appears to
be a size range for blue catfish that feed on bivalves. Con-
sidering the high numbers of blue catfish in large rivers
and reservoirs in southeastern United States stad the im-
pact of the zebra mussel invasion, we recommend more
extensive research toward assessing predation by blue cat-
fish on zebra mussels and efforts toward distinguishing
whether predation is incidental or selective.
We thank Hancock Biological Station of Murray State
University for laboratory facilities and equipment.
LITERATURE CITED. (1) Wormington, A., and J. H.
Leach. 1992. Concentrations of migrant diving ducks at Point
Pelee National Park, Ontario, in response to invasion of zebra
mussels, Dreissena polymorpha. Canad. Field-Naturalist 106:
376-380. (2) De Leeuw, J., and M. R. Van Eerden. 1992.
Size selection in diving tufted ducks Aythya fuligula explained
by differential handling of small and large mussels Dreissena
polymorpha. Ardea 80:353-362. (3) Love, J., and J. F. Savino.
1993. Crayfish (Orconectes virilis) predation on zebra mussels
(Dreissena polymorpha). J. Freshwater Ecol. 8:253-259. (4)
French, J. R. P., HI. 1993. How well can fishes prey on zebra
mussels in eastern North America? Fisheries 18(6):13—19. (5)
Prejs, A., K. Lewandowski, and A. Stanczykowska-Poitrowska.
1990. Size-selective predation by roach (Rutilis rutilis) on ze-
bra mussel (Dreissena polymorpha): field studies. Oecologia
83:378-384. (6) Cloe, W. W,, II, G. C. Garman, and S. A.
Stranko. 1993. The potential of the bull chub (Nocomis ra-
neyi) as a predator of the zebra mussel (Dreissena polymor-
pha) in mid-Atlantic coastal rivers. Am. Midl. Naturalist 133:
170-176. (7) Stein, R. A., J. F. Kitchell, and B. Knezevic.
1992. Selective predation by carp (Cyprinus carpio L.) on
benthic molluscs in Skadar Lake, Yugoslavia. Fish Biol. 7(3):
391-399. (8) Robison, H. W., and T. M. Buchanan. 1984.
Fishes of Arkansas. Univ. Arkansas Press, Fayetteville, AR —
Jeffrey J. Herod, Tricia L. Frye, and James B. Sickel,
Department of Biological Sciences, Murray State University,
Murray, KY 42071.
Rare and Extirpated Plants and Animals of Kentucky:
1997 Update.—The Kentucky State Nature Preserves Com-
mission (KSNPC) published a list of rare and extirpated
plants and animals of Kentucky in 1996 (1). The list, devel-
oped with assistance from many scientific authorities, was
based on distributional and ecological data available as of 1
Mar 1995. KSNPC (1) committed to update the list annually
Notes 97
so that decision makers would have current information on
rare species in Kentucky. Herein we update the 1996 list
based on data available through 31 Dec 1996.
The methods and status categories used herein follow
KSNPC (1), with the following exceptions. The United
States Fish and Wildlife Service (USFWS) (2) discontin-
ued use of status review Category 2 (C2) and Category 3
(3A) to reduce confusion and to clarify that these species
were not candidates for listing as endangered or threat-
ened. The USFWS (2) now maintains a list of only those
taxa that are candidates for listing as endangered or
threatened (C, which supersedes category C1), and taxa
proposed for listing as endangered or threatened (PE and
PT). Those species, including three taxa subsequently list-
ed as endangered by the USFWS (3), are listed in Table
1 with other species whose conservation status is being
changed. Changes in nomenclature and additions to the
list are presented in Tables 2 and 3, respectively. Common
names are used only when there is a nomenclature change
Table 1.
Old
Plants
Aureolaria patula
Berchemia scandens
Elodea nuttallii
Halesia tetraptera
Lonicera dioica var. orientalis
Trepocarpus aethusae
Triplasis purpurea
Viburnum lentago
DAAC Des
Snails
Anguispira rugoderma S
Mussels
Alasmidonta atropurpurea =
Epioblasma brevidens —
Epioblasma capsaeformis —
Crustaceans
Cambarus ornatus S
Orconectes inermis AR
Insects
Pseudanophthalmus catoryctos
Pseudanophthalmus pholeter
Fishes
Ammocrypta clara X
Etheostoma cinereum T
Etheostoma tippecanoe S
Macrhybopsis gelida =
Macrhybopsis meeki
Percina evides
Birds
Aquila chrysaetos X
from KSNPC (1), to identify undescribed species, and
when a species is added to the list. Sources for plant
names are Gleason and Cronquist (4), McAtee (5), and
Wilbur (6). Sources consulted for animal names are as
follows: gastropods—Hubricht (7) and Turgeon et al. (8);
insects—Barr (9), Garrison (10), and The Dragonfly So-
ciety of the Americas (11); fishes—Robbins et al. (12) and
Warren, Burr, and Grady (13); and birds—American Or-
nithologists’ Union (14). We welcome any questions or
comments about this update or KSNPC (1).
We thank the following individuals for sharing infor-
mation and for their assistance: W.M. Andrews, Kentucky
Division of Environmental Services; T.C. Barr Jr, Uni-
versity of Kentucky; D.L. Batch, J. Kiser, G. Libby, G.A.
Schuster, and T. Weckman, Eastern Kentucky University;
H.D. Bryan, Eco-Tech, Inc.; B.M. Burr, Southern Illinois
University at Carbondale; S.M. Call and K.L. Smathers,
Kentucky Division of Water; C. Cook, Florida State Col-
lection of Arthropods; C.V. Covell Jr., and W.D. Pearson,
Conservation status changes for rare and extirpated Kentucky plants and animals.
KSNPC! US?
New Old New
S oes a
T wt soo
T sans wa
T pL ae
E ee ales
T Lins atk,
H = ae
delist — —
T es et
= PE E
— PE E
— PE E
delist — —
S ben a
E es pis
E ne ce
E we aay
S can iss
delist — —
= GI GC
= Cl C
delist — -—
delist —_ —
'§ = Special Concem, T = Threatened, E = Endangered, H = Historic, X = Extirpated or Extinct.
2 PE = Proposed Endangered, E = Endangered, C = Candidate for Listing, Cl = Candidate for Listing.
98 Transactions of the Kentucky Academy of Science 58(2)
Table 2.
Nomenclature changes for rare and extirpated Kentucky plants and animals.
Old name
New name
Plants
Carex atlantica ssp. capillacea
Howe sedge
Gentiana alba
Halesia tetrapera var. tetraptera
Heracleum maximum
Insects
Pseudanophthalmus abditus
A cave beetle
Pseudanophthalmus hypolithos
Stone-dwelling cave beetle
Pseudanophthalmus inexpectatus
A cave beetle
Pseudanophthalmus scholasticus
Schoolhouse cave beetle
Fishes
Etheostoma clarum
Etheostoma pellucidum
Etheostoma vivax
Notropis sp.
Palezone shiner
Birds
Elanoides forficatus forficatus
American swallow-tailed kite
Casmerodius albus
Prickly bog sedge
Gentiana flavida
Halesia tetraptera
Heracleum lanatum
Concealed cave beetle
Ashcamp cave beetle
Surprising cave beetle
Scholarly cave beetle
Ammocrypta clara
Ammocrypta pellucida
Ammocrypta vivax
Notropis albizonatus
Swallow-tailed kite
Ardea alba
University of Louisville; D. Dourson and J.R. MacGregor,
United States Forest Service; J.J. Lewis; L.E. Kornman,
B.D. Laflin, and K.W. Prather, Kentucky Department of
Fish and Wildlife Resources; T.L. Poulson, University of
Illinois-Chicago; S.P. Rice, Kentucky Transportation Cab-
Table 3.
tucky plants and animals.
Additions to the list of rare and extirpated Ken-
KSNPC'! US?
status status
Plants
Viburnum molle
Missouri arrow-wood IE —
Snails
Helicodiscus notius specus
A snail 0 att
Helicodiscus punctatellus
Punctate coil S —
Insects
Celithemis verna
Double-ringed pennant S —
Fishes
Etheostoma percnurum
Duskytail darter E E
Special Concern, T = Threatened, E = Endangered.
IS=S§
2>E = Endangered.
inet; J. Slapcinsky, Field Museum of Natural History; C.A.
Taylor, Illinois Natural History Survey; and K.J. Tennes-
sen.
LITERATURE CITED. (1) Kentucky State Nature
Preserves Commission. 1996. Rare and extirpated plants
and animals of Kentucky. Trans. Kentucky Acad. Sci. 57:
69-91. (2) U.S. Fish and Wildlife Service. 1996. Endan-
gered and threatened wildlife and plants; review of plant
and animal taxa that are candidates for listing as endan-
gered or threatened species; notice of review. Fed. Reg.
61:7596-7613. (3) United States Fish and Wildlife Ser-
vice. 1997. Endangered and threatened wildlife and
plants; determination of endangered status for the Cum-
berland elktoe, oyster mussel, Cumberlandian combshell,
purple bean, and rough rabbitsfoot. Fed. Reg. 62:1647—
1658. (4) Gleason, H.A., and A. Cronquist. 1991. Manual
of vascular plants of northeastern United States and ad-
jacent Canada. 2nd ed. The New York Botanical Garden,
Bronx, NY. (5) McAtee, W.L. 1956. A review of the nearc-
tic Viburnum. W.L. McAtee, Chapel Hill, NC. (6) Wilbur,
R.L. 1988. The correct scientific name of the pale, yellow
or white gentian of the eastern United States. Sida 13:
161-165. (7) Hubricht, L. 1985. The distributions of the
native land mollusks of the eastern United States. Field-
iana Zool. 24:1-191. (8) Turgeon, D.D., A.E. Bogan, E.V.
Coan, W.K. Emerson, W.G. Lyons, W.L. Pratt, C.F-E.
Roper, A. Scheltema, F.G. Thompson, and J.D. Williams.
1988. Common and scientific names of aquatic inverte-
Notes 99
brates from the United States and Canada: mollusks. Am.
Fish. Soc. Spec. Publ. 16. (9) Barr, T.C., Jr. 1996. Cave
beetle status survey and prelisting recovery project. Re-
port prepared for the U.S. Fish and Wildlife Service and
the Kentucky Dept Fish Wildl. Resources, Frankfort, KY.
(10) Garrison, R.W. 1991. A synonymic list of the New
World Odonata. Argia 3(2):1-30. (11) The Dragonfly So-
ciety of the Americas. 1996. Common names of North
American dragonflies and damselflies. Argia 8(2). (12)
Robbins, C.R., R.M. Bailey, C.E. Bond, J.R. Brooker, E.A.
Lachner, R.N. Lea, and W.B. Scott. 1991. Common and
scientific names of fishes from the United States and Can-
ada. 5th ed. Am. Fish. Soc. Spec. Publ. 20. (13) Warren,
M.L., Jr, B.M. Burr, and J.M. Grady. 1994. Notropis al-
bizonatus, a new cyprinid fish endemic to the Tennessee
and Cumberland River drainages, with a phylogeny of the
Notropis procne species group. Copeia 1994:S68—886. (14)
American Omithologists’ Union. 1995. Fortieth supple-
ment to the American Ornithologists’ Union check-list of
North American birds. Auk 112:819-830.—Kentucky
State Nature Preserves Commission, 801 Schenkel
Lane, Frankfort, KY 40601.
Epilobium brachycarpum (Onagraceae) in Ken-
tucky.—While collecting plants in a railroad yard in
northern Kentucky, we noted, among Hypericum perfor-
atum and Potentilla recta, a number of non-flowering in-
dividuals of a plant we did not recognize. We eventually
concluded that it was probably Epilobium brachycarpum
C. Pres] (E. paniculatum Nutt.) (Figure 1), a xerophytic
annual new to the flora of Kentucky. Our conclusion was
proved correct when, later, the plants came into flower.
In the conterminous U.S., the plant was known east of the
Mississippi only from Minnesota and Wisconsin. Further
west, it occurs in the Dakotas and from Montana south
to New Mexico and west to the Pacific; in Canada it is
known from Quebec to British Columbia (International
Vascular Plant Database, Biota of North America Pro-
gram, North Carolina Botanical Garden, University of
North Carolina, Chapel Hill, NC 27599).
The species is distinctive even in non-flowering condi-
tion, with (1) its elongate leaves that often have fascicles of
smaller leaves in their axils and are folded up along the
midrib (V-shaped), sharply acute, and entire to obscurely
toothed, the teeth low, gland-tipped, and mostly 1 to 4 mm
apart; and (2) its cracking and exfoliating epidermis at the
base of the stem in older plants. Its flowers have pink petals
about 6 mm long that are lobed to about the middle; the
pollen is shed as monads, not as tetrads as in most species
of Epilobium (1). Our plants are glabrous. Although infra-
specific taxa have been described for this polymorphic spe-
cies, none of these was recognized by Raven (1).
We cite here a voucher specimen for the presence of
E. brachycarpum in Kentucky: KENTUCKY. Kenton Co.,
in gravelly soil along tracks in railroad yard along Licking
River ca. % mile s of 1-275 bridge, ca. 2 miles s of Latonia,
23 Jul 1997, Thieret and Allen 60239 (KNK, MO).
Figure 1. Epilobium brachycarpum. Photocopy of
pressed specimens, X ¥%.
LITERATURE CITED. (1) Raven, P.H. 1976. Generic
and sectional delimitation in Onagraceae, tribe Epilo-
bieae. Ann. Missouri Bot. Gard. 63:326-340.—John T.
Kartesz, Biota of North America Program, North Caro-
lina Botanical Garden, University of North Carolina,
Chapel Hill, NC 27599; Peter Allen and John W. Thier-
et, Department of Biological Sciences, Norther Ken-
tucky University, Highland Heights, KY 41099.
Horned-Pondweed, Zannichellia palustris (Zanni-
chelliaceae) from Northern and Central Kentucky.—
Zannichellia, a genus of aquatic monocots of the family
Zannichelliaceae, is represented in North America by a sin-
gle species, Z. palustris L. (Figure 1). Beal and Thieret (1)
located no Kentucky voucher specimens for the species but
indicated that it is “possibly to be expected” in the state.
They also noted that a population of this species formerly
occurred just across the Ohio River from northem Ken-
tucky in Hamilton County, Ohio, but was destroyed by ex-
pansion of the city of Cincinnati. In 1987 Patricia Haragan
found the species in Fayette County but did not report the
find. Two years later Webb and Chester (2) published Ken-
tucky records of Zannichellia for Calloway, Lyon, Marshall,
and Trigg counties (Kentucky Lake).
In May 1994, while looking for plants along a proposed
100
Figure 1.
achenes (X6). Drawing by John Myers; used by permis-
sion of Flora of North America.
Zannichellia palustris. Plant (<%); lower right,
right-of-way for the reconstruction of KY 17, near Inde-
pendence, Kenton County, Kentucky, we found horned-
pondweed in a pond with Eleocharis obtusa and Lemna
minor; in May 1995 we found Zannichellia at two sites in
Bourbon County (in a roadside pool along US 68, ca. 0.5
miles northeast of Paris, and in a pond north of Millers-
burg); and in July 1996 we discovered a population in a
pond off KY 356, ca. 2 miles east of Cynthiana, Harrison
County. The documented Kentucky distribution of this
Transactions of the Kentucky Academy of Science 58(2)
Figure 2. Zanichellia palustris. Documented county dis-
tribution in Kentucky.
species now includes the northern and central parts of the
state (Figure 2). Additional field work will probably yield
more Kentucky records.
Our voucher specimens are as follows: KENTUCKY.
Bourbon Co., US 68, ca. 0.5 miles ne of Paris, roadside
pool, 22 May 1995, Meade 2169 (MOKY, Morehead State
University Herbarium); Bourbon Co., n end of Millers-
burg, farm pond, 23 May 1995, Meade 2229 (MOKY,
Morehead State University Herbarium); Fayette Co., 8
miles e of Lexington, plants covering spring-fed farm
pond, 11 Jun 1987, Haragan 1528 (Univ. Ky. Coll. Agric.
Herbarium: KNK): Harrison Co., off KY 356, ca. 2 miles
e of Cynthiana, farm pond, 3 Jul 1996, Meade 2284
(MOKY, Morehead State University Herbarium); Kenton
Co., nr. Independence, farm pond, 23 May 1994, Meade
1511 (MOKY, Morehead State University Herbarium).
We acknowledge aid from Kathy Stewart and Nancy
Hopkins.
LITERATURE CITED. (1) Beal, E.O., and TW. Thier-
et. 1986. Aquatic and wetland plants of Kentucky. Kentucky
Nature Preserves Comm., Sci. Tech. Ser., 5:1-314. (2)
Webb, D.H., and E.W. Chester. 1989. Distribution of the
homed pondweed, Zannichellia palustris L. (Zannichelli-
aceae), in Tennessee and Kentucky. Trans. Kentucky Acad.
Sci., 50:158—160.—Les Meade and Brian Binion, De-
partment of Biological and Environmental Sciences, More-
head State University, Morehead, KY 40351; Peggy Mea-
sel and Brenda Hamm, T.H.E. Engineers Inc., Lexington,
KY 40509; and Patricia Dalton Haragan, Herbarium,
University of Louisville, Louisville, KY 40294.
Trans. Ky. Acad. Sci. 58(2):101. 1997.
List of Reviewers for Volume 58 of
Transactions of the Kentucky Academy of Science
John R. Baird
Richard I. Barnhisel
David M. Brandenburg
Jerry Carpenter
Ronald R. Cicerello
Michael A. Flannery
Michael A. Floyd
Larry A. Giesmann
Vernon Hicks
Ellis L. Laudermilk
James O. Luken
Max E. Medley
Robert F.C. Naczi
Debra K. Pearce
101
Thomas C. Rambo
Arthur Raymond
James Sickel
David Sleper
Jerry W. Warner
Stephen White
Steven Wilkinson
Trans. Ky. Acad. Sci. 58(2):103-110. 1997.
INDEX TO VOLUME 58
Compiled by Varley Wiedeman
A-chain gene transcription, 42-43
Aesculus glabra var. glabra, 15
Abstracts, 37-54
Accipitridae, 54
Acer
saccharinum, 87, 88
rubrum, 89
Acidic mines, oil effects on nodula-
tion, 80-84
Actinonaias
ligamentina, 56, 57
spectorosa, 57
AD-like pathology, in normal aging,
4l
Adiantum capillus-veneris, 74
ADKINS, MARCI, 43
Aesculus glabra var. glabra, 13
African-American
health and diet status, 39
rural southern elderly, 39
Agricultural sciences, abstracts, 37-
38
Agrypnia vestita, 69, 71, 72
Alasmidonta atropurpurea, 97
Alkylaton of DNA, 40
ALLEN, PETER, 99
Allium cepa, 60
Alpheus heterochaelis, snapping ac-
tion of, 52-53
Alzheimer’s PS-1 mutation, 40
Ambrosia, 17
Ameiurus nebulosus, 96
American ipecac, 12, 16
American lotus, 13, 16
American pennyroyal, 12, 16
American swallow-tailed kite, 98
Ammocrypta
clara, 97, 98
pellucida, 98
vivax, 98
Andropogon
arenaceum, 15
avenaceus, 15
nutans, 15
spp., 21
Anguispira rugoderma, 97
Animals of Kentucky, rare and extir-
pated, 96-99
Annonaceae, 37
Antimonite, sensing system for, 42
Aplodinotus grunniens, 96
Appalachian false-foxglove, 14, 16
Aquila chrysaetos, 97
Ardea alba, 98
ARMSTRONG, JAMES S., 54
Arnoglossum muehlenbergii, 12, 15
Arrow-wood, Missouri, 98
Arundinaria gigantea, 12, 15
Asarum
canadense, 18
virginicum, 14, 15, 18
Ashcamp cave beetle, 98
Asian clam, 96
Asimina triloba, 13, 15, 37
Asperula odorata, 18
Asteraceae, 17, 38
Astroglial toxicity
opiate-mediated, 46
TNF-alpha neuroprotection in,
46
Astronomy
for non-science majors, 48
use of computers for, 48
Astronotus ocellatus, gentamicin-in-
duced toxicity in, 49, 50
Aureolaria
flava,18
laevigata, 14, 15, 18
patula, 74, 97
BACKUS, JASON, 23
Bahalana ‘eracei, 53
effects of cannibalism on, 53
from san Salvador Island, Baha-
mas, 53
population structure of, 53
Bald eagles
densities, 54
distributions, 54
impacts of dam construction on,
54
Baldcypress, 85, 88
Baptisia
australis, 12, 15
cerulea, 12, 15
coerulea, 12, 15
tinctoria, 75
Barren oats, 15, 16,
BARRETT, BILL, 40
Bartonia
paniculata var. paniculata, 75
virginica, 75
Bastard pennyroyal, 12, 16
BEATTIE, RUTH E., 51
Beetle
cave, 98
concealed cave, 98
scholarly cave, 98
schoolhouse cave, 98
stone-dwelling cave, 98
surprising cave, 98
Berchemia scandens, 97
Beta vulgaris, 65
Bidens discoidea, 87
Bieberbach groups, classifying free,
29—32.
BILES, DANIEL C., 92
BINION, BRIAN, 100
103
Biology, student attitudes toward, 51
Birds, 97, 98
BITNER, RONALD D., 53
Bitter milkwort, 14, 16
Bivalva, 55-59
in the Cumberland River, 55-59
Black darter, 4
Black wort, 15, 16
BLACK, RICHARD, 42
BLAIR, PETER, 23
BLANK, SARAH M., 46
BLOOM, C. TOM, 74
Blue catfish, 96
predation on zebra mussel, 96
Blue false-indigo, 12, 16
Blue penny-royal, 14, 16
Blue wild indigo, 12, 16
Blue-curls, 12, 16
Blue-hearts, 15, 16
Bluestems, 21
Body weight in rats
effect of dietary energy restric-
tion, 49
effect of ovariectomy on, 49
Boehmeria cylindrica, 87
BONDADA, SUBBARAO, 43
BOOZE, ROSE-MARIE, 41
Botanical regions in Kentucky, 20-
22
Constantine Rafinesque’s 1819
description of, 20-22
Botany and microbology abstracts,
38-39
Botany of Kentucky, 12
BRACKNEY, CHRISTOPHER, 41
Bradyrhizobium, 80-84
Breast cancer, human, 46-47
Briar, sand, 17
BRICKING, KEITH D., 42
Brown bullhead, 96
BROWN, C., 49
BRYANT, WILLIAM S., 20, 85
Buchnera americana, 15
Buck-eye, prickly, 13, 17
Buckeye, Ohio, 13, 17
Buffalo, smallmouth, 96
Bull chub, 96
Bullhead, brown, 96
Button wood, 13, 16
Buttonbush, 87
Buttonwood, 16
Cacalia, 17
atriplicifolia, 17
muehlenbergii, 12, 15, 17
reniformis, 12, 15
Caddisflies, of the Kentucky River,
67-73
Calcium excretion, urinary, 49
104 Transactions of the Kentucky Academy of Science 58(2)
Calcium
homeostasis, 40
neurotaxicity, 4]
metabolism, in rats, 50
Calopogon
tuberosus, 75
tuberosus var. tuberosus, 74
Cambarus ornatus, 97
CAMMERS-GOODWIN, ARTHUR,
40
Campostoma anomalum, 6
Cane, 12, 16
giant, 12, 16
Cannibalism, 53
effects on population structure,
53
Capraria multifida, 13, 15
Carex
atlantica ssp. capillacea, 98
crawei, 75
stricta var. stricta, 75
CARINI, MICHAEL, 48
CARLSON, SONIA, 44
Carp, common, 96
CARPENTER, JERRY H., 53
CARRICO, LYNN, 41
Carya illinoinensis, 89
Casmerodius albus, 98
Castanea dentata, 75
Catalpa, 18, 20
bignonioides, 18
cordifolia, 18
northern, 13, 17
speciosa, 13, 15, 18, 20
tree, 13, 16
Catalpium
cordata, 13, 15
cordatum, 18
cordifolium, 13, 15, 18
Catfish, blue, 96
predation on zebra mussel, 96
Catostomus commersoni, 6
Caucasian
health and diet status, 39
rural southern elderly, 39
Cave beetle, 98
ashcamp, 98
concealed, 98
scholarly, 98
schoolhouse, 98
stone-dwelling, 98
surprising, 98
CCDs, photometry with, 48-49
Cedar, red, 14, 17
Celestial seasons, 47
Celithemis verna, 98
Cenchrus longispinus, 76
Center for automated space science,
48
Cephalanthus occidentalis, 87
Cephalogonimidae, 23-28
Cephalogonimus vesicaudus, 23-28
Ceraclea
ancylus, 69, 71
cancellata, 69, 71
flava, 69, 71
punctata, 69, 71
tarsipunctata, 69, 71
transversa, 69, 72
Ceratopsyche sparna, 69, 70
Chenopodium
ambrosioidaes var. anthelminti-
cum, 12, 15
anthelminthicum, 12, 15
Cheumatopsyche
campyla, 69—71, 73
minuscula, 69, 70
oxa, 69, 70, 73
pasella, 69, 70
Chimarra obscura, 69, 70, 72
Chromosome 17 deletions, in ovar-
ian tumors, 41—42
Chub, bull, 96
CICERELLO, RONALD R., 55
Cichlidae, 49, 50
Clam, Asian, 96
Clammy-weed, 13, 16
CLEMENTS, SANDRA, 47-49
Cliff plumb, 16
CLINE, GARY R., 80
Clover, nimble, 15
Common carp, 96
Common laurel, 14
Common names, of vascular plants,
9-19
Community college graduates, 50-
51
COMPTON, KEVIN, 40
Computerized education, a student
survey, 51-52
Computers, use for astronomy, 48
CONARD, NYREE, 38
Concealed cave beetle, 98
Coneflower
eastern, 14
orange, 14, 17
purple, 14, 17, 21
Conobea, 17
COON, ALEXANDER, 41
Copper iris, 16
Corbicula fluminea, 56, 96
Cotton tree, 13
Cottontails, eastern, 1-3
cover, 1-3
habitat, 1-3
in Kentucky, 1-3
Cottonwood, 13, 16, 20
Cow mint, 13, 16
CRAWFORD, MELINDA L., 52
Crested dwarf iris, 14, 16
Crested flag, 16
Crested iris, 14, 16
Crested tris, 14, 16
Crustaceans, 97
Cryopreservation of seeds, of Ohio
wetland plants, 38
CUILARAN, FONSIE, 48
CULLER, RYAN, 47, 48
Cultured panicle segments, 60-66
Cumberlandia monodonta, 58
Currant, wild, 14, 17
CURTSINGER, W. STEWART, 49
Cuscuta sp., 87
Cuterebra
abdominalis, 35
buccata, 35
fontinella, 35
sp., 35-36
Cyclonaias tuberculata, 55, 57-58
Cyperus
croceus, 76
difformis, 76
Cyprinus carpio, 96
Cyrnellus fraternus, 69, 70
Cytochrome P450 2E1, 45-46
Cytokine production, 42
Cytokines, and neonatal immune re-
sponses, 43-44
Dalea
candida, 15
purpurea var. purpurea, 15
Ddarter
black, 4
saffron, 4
duskytail, 98
Ddarters, snubnose, 4-8
separation of spawning habit, 4-8
spawning habit in, 4-8
DAUNERT, SYLVIA, 42
DEARTING, TERRY L., 53
DECKARD, MONICA, 47, 48
DEDRICK, ERIC, 45
Deerberry, 14, 16, 21
DELISLE, BRIAN, 43
DELONG, JASON E., 42
DEMOSS, D.L., 50
DERTING, TERRY L., 51, 54
Dietary energy restriction, effects of
on rats, 49
Diptera, 35-36
Distinguished scientist award, 33
Double-ringed pennant, 95
Drum, freshwater, 96
Duskytail darter, 98
Dwarf iris, crested, 16
Eastern coneflower, 14, 16
Echinacea purpurea, 14, 15, 21
Echinostoma trivolvis, 23-28
Echinostomatidae, 23-28
EGAN, RITA, 42
EIZENGA, GEORGIA C., 60
Elanoides forficatus forficatus, 98
Electrophysiological studies, of the
SCN8a_ voltage-gated _ sodi-
um channel, 43
Elephant'’s foot, 12, 16
leafy, 12, 16
Elephantopus
caber, 12, 15, 17
carolinianus, 12, 15, 17
ELLINGTON, JEFF, 23
ELLIOTT, CHARLES L., 1
Ellipsaria lineolata, 55, 57
Elliptio
crassidens, 56, 57
dilatata, 56, 57
Elodea nuttallii, 97
EMBRY, KENNETH, 39
Emden-Fowler equation, polyno-
mial conservation laws of, 40
Endangered Ohio wetland plants,
cryoperservation of seeds of, 38
Epilobium
brachycarpum in Kentucky, 99
ciliatum var. ciliatum, 76
paniculatum, 99
Epioblasma
brevidens, 58, 97
capsaeformis, 58, 97
EPSCoR opportunities for space-re-
lated science, technology, 48
Esox masquinongy, 46
Etheostoma
barrenense, 5
blennioides, 6
caeruleum, 5, 6
cinereum, 97
clarum, 98
crossopterum, 6
davisoni, 5
duryi, 4, 5, 7
flabellare, 6
flavum, 4—S
flavum, spawning in, 4—8
pellucidum, 98
percnurum, 98
rafinesquei, 5
simoterum, 4-8
simoterum, spawning in, 4—S
smithi, 6
tippecanoe, 97
vivax, 98
zonale, 5
Ethics
development and content, 51
science, 51
world view and values, 51
Eupatorium
celestinum, 15
coelestinum, 13, 15
rugosum, 13, 15
urticefolium, 13, 15
Extirpated animals of Kentucky, 96-
99
Extirpated plants of Kentucky, 96—
99
Extragalactic radio jets, 47
Fabaceae, 80-84
FALBO-KENKEL, MARIA, 47
False pennyroyal, 14, 16
False-foxglove, Appalachian, 14, 16
False-indigo, blue, 16
Fawnsfoot, 96
Fescue, tall, 60-66
Index to Volume 57
characterization of for DNA
changes, 60-66
characterization of for isozyme
changes, 60-66
characterization of for meiotic
changes, 60-66
Festuca arundinacea, 60
FIALKOW, JARED, 52
Field investigations, 51
educational benefits, 51
misconceptions, 51
problems, 51
student perspectives of, 51
FINNESETH, C.L.H., 37
Fire-pink, 15, 16
Fishes, 97, 98
Flag, 14, 16
crested, 16
Forestiera acuminata, 87, 88
Fraxinus
pennsylvanica, 87, 88
profunda, 87
Free Bieberbach groups, classifying,
29-32
FREEMAN, JAMES, 44
Freshwater drum, 96
Fringed-orchid
orange, 14, 17
yellow, 14, 17
FRYE, TRICIA L., 96
Gaultheria procumbens, 14, 21
GELDERMAN, MICHAEL CAR-
INI, 47
GELDERMAN, RICHARD, 47, 48
Genes involved in splicing, 44
Gentamicin-binding protein(s), 49—
51
in ear of cichlid fish, 49-50
Gentian
stiff, 15, 17
yellow, 15, 17
Gentiana
alba, 98
amarelloides, 15, 18
amerellvides, 15
flavida, 98
quinquefolia var. occidentalis, 18
quinquefolia var. quinquefolia, 15,
18
Geography abstracts, 39
Gerardia glabrata, 14, 15
Giant cane, 12, 16
GIESMANN, LARRY A., 38
GILBERT, NIKOLE L., 43
Gillenia
stipulacea, 12, 15
stipulata, 12, 15
GILLIS, PETER P., 40
GIULIANO, WILLIAM M., 1
Glossosomatidae, 69, 70
Golden-rod, stiff, 14, 17
Goldenrod, stiff, 17
105
Graduates, of community colleges,
50-51
GRANESE, JACQUELINE M., 52
Grass, Indian, 15, 16, 21
Gravitational lensing simulation, 47
Great Indian-plantain, 12, 16
Green ash, 87
GUILARAN, ILDEFONSO, 47
GUO, QING, 40
HACKNEY, KAREN, 47-49
HACKNEY, RICHARD, 47-49
Hairy ruellia, 15, 16
Halesia tetrapera, 97, 98
Haliaeetus leucocephalus, 54
densities, 54
distributions, 54
impacts of dam construction on,
54
HAMPTON, C. TONY R., 53
HAMPTON, D., 49
HARAGAN, PATRICIA DALTON,
100
HARP, CINDY, 44
Hatching, in the muskellunge, 46
stage dependent induced, 46
HAUSER, KURT F., 46
Health and diet status, differences
in, 39
Health sciences abstracts, 39-40
Heart-leaf, 14, 16
Hedeoma pulegioides, 12, 15
Helicodiscus
notiusspecus, 98
punctatellus, 98
Heliconia, 53
latispatha imbricata, 53
extrafloral nectaries, 53
Heliconiaceae, 53
Heliotrope, 13, 16
Indian, 13, 16
Heliotropium indicum, 13, 15, 17
Helisoma trivolvis, 23-28
Heracleum
lanatum, 98
maximum, 98
HEROD, JEFFREY J., 96
Hesperis pinnatifida, 13, 15
Hexastylis, 18
virginica, 14, 15
Hippocampal subregions, 41
HOGSTRAND, CHRISTER, 40
Horned-pondweed, 99-100
Horse nettle, 16
HOUP, RONALD E., 67
Houstonia, 18
angustifolia, 18
fruticosa, 13, 15
nigricans var. nigricans, 13, 15
tenuifolia, 18
Houstonia, narrow-leaved, 13, 17
Howe sedge, 98
HT 1080 human fibrosarcoma can-
cer cells, 42
106 Transactions of the Kentucky Academy of Science 58(2)
HUANG, HONGWEN, 37
Human breast cancer, mRNA in,
46-A7
Human myiasis in Kentucky, 35-36
Human pancreatic carcinoma, 44—
45
Hydropsyche
betteni, 69
frisoni, 69, 70
hetteni, 70
orris, 67, 69, 70
simulans, 69, 70
Hydropsychidae, 69, 70
Hydroptila
ajax, 69, 70
angusta, 69, 70
armata, 69, 71
consimilis, 69, 71
hamata, 69, 71
perdita, 69, 71
waubesiana, 69, 7
Hydroptilidae, 69, 7
Hymenocallis
caroliniana, 13, 15, 18, 20
liriosme, 18
occidentalis, 18
Hypericum
perforatum, 99
walteri, 87
oe
Ictaluridae, 96
predation on zebra mussel, 96
Ictalurus furcatus, 96
predation on zebra mussel, 96
Ictiobus bubalus, 96
Immune responses, neonatal, 43-44
Immune tissues, innervation of, 44
role of nerve growth factor, 44
Indian grass, 15, 16, 21
Indian heliotrope, 13, 16
Indian physic, 12, 16
Indian-plantain, great, 16
Indigo, blue wild, 16
Induced hatching, stage dependent,
46
Innervation
of immune tissues, 44
role of nerve growth factor, 44
Insecta of the Kentucky River, 67—
13
Insects, 97, 98
lodanthus pinnatifidus, 13, 15
Ipecac, American, 12, 16
Iris
cristata, 14, 15
crocea, 13, 15
fulva, 13, 16
Iris
copper, 16
crested, 14, 16
crested dwarf, 14, 16
Iron weed, 16
Ironoquia punctatissima, 69, 71-73
Ironweed, 12
tall, 17
Isanthus
brachiatus, 14, 16
caeruleus, 16
ceruleus, 14, 16
coeruleus, 14
Isopod, marine cave, 53
JACKSON, JANN, 46
Jackson Purchase Region, of Ken-
tucky, 85-91
Jeffersonia
binata, 13, 16
cinata, 13, 16
diphylla, 16
JENNINGS, JOHNNY, 48
JONES, S.E., 40
Juncus validus, 74, 76
Juniperus virginiana, 14, 16, 21
JUST, JOHN J., 46
KAETZEL, DAVID, 42
Kalmia latifolia, 14, 21
KENTUCKY STATE NATURE
PRESERVES COMMISSION,
99
Kentucky vascular plants, 74-79
Kentucky's Jackson Purchase Re-
gion, 85-91
Kidney weed, 12, 16
Kite
American swallow-tailed, 98
swallow-tailed, 98
KRUPER, JILL H., 51, 53
L-type calcium channel density, 41
Lampsilis
cardium, 56, 57
fasciola, 57
ovata, 56, 57
Large-flowered leafcup, 14, 16
Lasmigona costata, 57
LAUDERMILK, ELLIS L., 55
Laurel
common, 14
mountain, 2]
LAYNE, DESMOND R., 37
Leafcup
large-flowered, 14, 16
yellow-flowered, 14, 17
Leafy elephant’s-foot, 12, 16
LEATHERS, MICHELLE JO, 38
Lechea, 18
minor, 14, 16
LEE, C.J., 39, 49
Lepomis
cyanellus, 6
megalotis, 6
microlophus, 96
Leptoceridae, 69, 71
Leptodea fragilis, 58
Lespedeza
cuneata, 80-84
sericea, 80-84
Leucospora multifida, 13, 16, 17
LEWIS, BRIAN M.., 40, 47
LIBBY, GARY W., 74
Ligumiarecta, 57
Lilium
michiganese, 77
superbum, 76
Lily, 13, 16
red, 13, 17
swamp, 17
Limnephilidae, 69, 72
Linum usitatissimum, 77
List of reviewers, 101
Little brown jugs, 14, 17
Littorinidae, 52
LIU, BIN, 42
LOHRE, JOE, 47
Lolium perenne, 65
Lonicera dioica var. orientalis, 97
Lotus, 18
American, 13, 16
yellow, 13, 17
LOU, X., 50
LUO, JIANYUAN, 44
Lupatorium calutinum, 13, 16
Luxilus chrysocephalus, 6
Lycopus rubellus, 87
Lythrurus ardens, 6
Macrhybopsis
gelida, 97
meeki, 97
MALPHRUS, BENJAMIN K., 40,
47
Maple, red, 89
MARLETTE, MARTHA, 39
Mathematics abstracts, 40
MATTSON, MARK P., 40, 46
McGRUDER, CHARLES, 48
McGRUDER, MICHAEL CARI-
NI, 47
McNEIL, RAYMOND C., 48
MEADE, LES, 100
MEARS, RANDY L., 74
MEASEL, PEGGY, 100
Medicago sativa, 65
Medionidus conradicus, 57
MELLON, ISABEL, 43
MERCER, JANET, 42
Metal impact in the Robinson For-
est, using metalliothionein, 40—
4]
Metalliothionein, assessment of
metal impact, 40-41
Miagia
arupedinaria, 12
arupendinaria, 16
Micropterus punctulatus, 6
Microsatellite instability and TGF-b
Type II receptor gene muta-
tion, 44-45
Miegia arundinaria, 12, 16
Migraine headaches, effects of
weather systems on, 39
Milkwort
bitter, 14, 16
racemed, 14, 17
MIMS, S.D., 38, 50
Mink, 56
Mint, cow, 16
Mismatch repair, 43
Mississippi alluvial plain, swamp for-
ests of, 85-91
Missouri arrow-wood, 98
Mist-flower, 13, 17
Molecular and cell biology abstracts,
40-47
Morehead radio telescope, 47-48
optimization of operating param-
eters, 40
surface geometry of, 40
MORRISON, JOHN R., 51
Mountain laurel, 21
Mountain tea, 14, 21
MPTP-treated C57B1/6 mice, 45
mRNA in human breast cancer,
translational regulation of, 46—
47
Muhlenbergia cuspidata, 77
Muskrats, 56
Mussel, zebra, 96
in the Ohio River, 96
Mussels, 97
Mutagenesis
of a conserved phenylalanine, 45—
46
site directed, 45-46
Mutations, in the RNA sorting sys-
tem, 45
Mycena leaiana, fine structure of
bacidiospores, 38
Mystacides sepulchralis, 69, 71, 72
NACZI, ROBERT F.C., 39
Nanostoma, 4
Narrow-leaved houstonia, 13, 17
NASA Kentucky space grant consor-
tium, 48
Nectopsyche
candida, 69, 72
exquisita, 69, 72
pavida, 69, 72
Nelumbium pentapetalum, 13, 16
Nelumbo, 18
lutea, 13, 16
Neotyphodium coenophialum, 60
Nephrotoxicity, gentamicin-induced,
Nettle
horse, 16
white, 13, 17
Neureclipsis crepuscularis, 69, 71,
1
Nicotinein MPTP-treated C57B1/6
mice, 45
Nicotine, neuroprotective effects of,
45
Nimble clover, 15
Nimble weed, 14
Index to Volume 57
Nimbleweed, 17
Nitrogen-fixing bacteria, 80-84
nm23 proteins, in the WR-082-01
cell line, 42-43
NOBLE, JOHN, 47, 48
Nocomis raneyi, 96
Nodulation, effects of acidic mine
soil, 80-84
Northern catalpa, 13, 17
NORTHINGTON, ALAN, 45
Notes, 35-36, 96-100
Notropis
albizonatus, 98
sp., 98
NOVOTNY, NATHAN, 45
Nucleotide excision repair mutants,
43
Nymphaea sp., 77
Nyssa aquatica, 85, 87, 88, 90
=
Oaecetis
avara, 72
cinerascens, 72
inconspicua, 72
persimilis, 72
Oats, barren, 16
Obliquaria reflexa, 55, 57
Ochrotrichia
aergerfasciella, 69, 71
cristata, 69, 71
tarsalis, 69, 71
xena, 69, 71
Oecetis
avara, 69, 71
cinerascens, 69
inconspicua, 69, 73
persimilis, 69, 73
Oestridae, 35-36
Ohio buckeye, 13, 17
Ohio wall flower, 13, 17
Onagraceae, 99
ONDERS, R., 38
Optical monitoring, of quasars, 48
Orange coneflower, 14, 17
Orange fringed-orchid, 14, 17
Orchis ciliaris, 14, 16
Orconectes inermis, 97
Oryza sativa, 65
OTTE, ELIZABETH, 44
Otter, 56
Outstanding teacher award
college/university, 33
secondary school, 33-34
Ovariectomy, effects of on rats, 49
Ovariectomy surgery
effects of on rats, 49
effects on urinary calcium excre-
tion, 49
Overcup oak, 87
Owsley Fork Reservoir, 23-28
Oxyethira pallida, 69, 71
Pa-paw tree, 13
Paddlefish, 50
body weight, 50
107
muscle lipid content, 50
stability of meat stored in crushed
ice, 50
using flood control reservoirs for,
38
Palezone shiner, 98
PAN, BIN-TAO, 49
Pancratium
liriosme, 13, 16, 18
liviosone, 13, 16
Pancreatic carcinoma, 44—45
Panicle segments, cultured, 60-66
PARRISH, MARK, 44
PAULY, JAMES R., 45
Pavia muricata, 13, 16
Pawpaw, 13, 17
allozyme variation in, 37
geographic differentiation in, 37
influence of shade on, 37-38
morphological development, 37—
38
root-zone modification, effect on,
37-38
seed germination, 37-38
seedling emergence, 37-38
Pawpaw tree, 17
Pea-clover, yellow, 14, 17
Pecan, 89
PENCE, VALERIE C., 38
Pencil flower, 14, 17
Pennant, double-ringed, 98
Penny royal, 12
Penny-aoyal, 12, 17
Penny-royal, 17
blue, 14, 16
Pennyroyal
American, 12, 16
bastard, 12, 16
false, 14, 16
Percidae, 4—8
Percina
caprodes, 6
evides, 97
Peromyscus leucopus, 35
Petalostemon
candidum, 16, 21
moncandidum, 15
purpureum, 15, 16, 21
Petalvitemon candidum, 15, 16
Phenylalaninein cytochrome P450
2E1, 45-46
PHILLIPS, TIMOTHY D., 60
Philopotamidae, 69, 72
Phlox glaberrima, 14, 16
Phlox, smooth, 14, 17
Photometry with CCDs, 48-49
optimizing signal extraction, 45—
49
Photomorphogenic control, in dan-
delion, 38
of flower scape elongation, 38
Phyrganeidae, 69, 72
Physic, Indian, 12, 16
108 Transactions of the Kentucky Academy of Science 58(2)
Physics and astronomy abstracts,
47-49
Physiology and biochemistry ab-
stracts, 49-52
PIERETTI, MAURA, 41
Pimephales notatus, 6
Pin weed, 14, 17
Pine, pitch, 14, 17, 21
Pink, 14, 17
scarlet, 15, 17
Pinus rigida, 14, 16, 21
Pinweed, small, 14, 17
Pitch pine, 14, 17, 21
Plagiorchis micracanthos, 26
Planera aquatica, 87, SS
Planning a trip, 92-95
Planorbidae, 23-28
Plants, 98
Plants of Kentucky, 96-99
rare and extirpated, 96-99
Platanthera ciliaris, 14, 16
Platanus occidentalis, 13, 16, 20, 88
Platelet-derived growth factor, 42—
43
Plum, wild, 13, 17
Polanina graveolens, 13, 16
Polanisia
dodecandra ssp. dodecandra, 13,
16
graveolens, 13, 16
Polycentropodidae, 69, 72
Polycentropus
cinereus, 69, 72
confusus, 69, 70, 72
Polygala
paucifolia, 74, 77
polygama, 14, 16
senega, 39
senega var. latifolia, 39
senega var. senega, 39
Polymnia uvedalia, 14, 16
Polynomial conservation laws, of the
generalized Emden-Fowlser-
equation, 40
Polyodon spathula, 38, 50
Polysaccharides, responses to, 43-—
44
Populus
angulata, 13, 16
deltoides, 13, 20, 88
deltoides var. deltoides, 16
Porcelia
tribuba, 13, 16
triloba, 13, 16
PORTER, TODD, 45
Porteranthus stipulatus, 12, 16
PORTERFIELD, JEAN C,, 4
Potamilus
alatus, 56-58
ohiensis, 58
Potamyia flava, 69, 70
Potentilla recta, 99
Prairie vole, 53-54
female preferences, 53-54
male odors in, 53-54
Prairie-clover
purple, 15, 17
white, 15, 17
Prairie-clovers, 21
Prairie-dock, 15, 17, 21
Prickly bog sedge, 98
Prickly buck-eye, 13, 17
PRINGLE, JAMES S., 9
PRINS, RUDOLPH, 52
Proterometra macrostoma, 27
Protoptila maculata, 69-71
Prunus
munsoniana, 13, 16, 18
pendula, 13, 16, 18
Pseudanophthalmus
abditus, 98
catoryctos, 97
hypolithos, 98
inexpectatus, 98
pholeter, 97
scholasticus, 98
Ptilostomis
ocellifera, 69-72
semifaciata, 69-72
Ptychobranchus
fasciolaris, 57
subtentum, 55-58
Punctate coil, 98
Purple coneflower, 14, 17, 21
Purple prairie-clover, 15, 17
Purple rocket, 13, 17
Purple sun-flower, 14, 17
Pycnopsyche lepida, 69, 72, 73
Pyganodon grandis, 58
Pyrimidine biosynthesis enzymes
localization of, 44
with fusion proteins, 44
Quadrula pustulosa, 56, 57
Quasars
optical monitoring of, 48
very high red shift, 48
Quercus lyrata, 87, 88
Raccoons, 56
Racemed milkwort, 14, 17
Radio frequency astrophysics, 47-48
Radio jets, extragalactic, 47
Radio telescope, 47-48
Rafinesque’s 1819 description of bo-
tanical regions, 20-22
Ragweed, sand, 13, 17
RAMANATHAN, SRIDHAR, 42
RAMBO, THOMAS C., 53
Rare animals of Kentucky, 96-99
Rare plants of Kentucky, 96-99
Rats
effect of dietary energy restric-
tion, 49
effect of ovariectomy on, 49
effects on urinary calcium excre-
tion, 49
RAWLS, JOHN M., 44, 45
Red cedar, 14, 17
Red lily, 13, 17
Red maple, 89
Red-eared sunfish, 96
Redcedar, 21
REID, JAMES, 42
Retinioc acid
effect on stromelysin, 42
effect on TIMP, 42
Rhyacophila lobifera, 69, 71-73
Rhyacophilidae, 69, 72
Rhynchospora macrostachya_ var.
macrostachya, 74, 77
RICHMOND, RAYMOND E., 42
RIGDON, STEVEN M., 49
RNA sorting system, in spermiogen-
esis, 45
Roach, 96
ROBBINS, MARK C., 38
ROBINSON, DAVID LOWELL,
38
ROBINSON, MARK P.,, 92
Rock weed, 13, 17
ROSEN, RON, 23
Rough bell, 15, 17
Rough wort, 14, 17
Rubiaceae, 18
RUCH, DONALD G., 38
Rudbeckia
fulgida, 14, 16
fulgida var. fulgida, 14
purpurea, 14, 16
Ruellia
caroliniensis ssp. ciliosa, 15, 16
oblongifolia, 15, 16
Ruellia, hairy, 15, 16
Rutilis rutilis, 96
RYMOND, BRIAN, 44
Saffron darter, 4
Salix nigra, 87, 88
San Salvador Island, Bahamas, Tec-
tarius muricatus on, 52
Sand briar, 13, 17
Sand ragweed, 13, 17
SAPP, TAMARA, 46
SATIN, JONATHAN, 43
Saururus cernuus, 87
Scarlet pink, 15, 17
Scented sun flower, 14, 17
SCHEFF, STEVE, 41
Schistosoma
japonicum, 27
mansoni, 27
Schoenoplectus
mucronatus, 74, 77
purshianus, 78
Scholarly cave beetle, 98
Schoolhouse cave beetle, 98
SCHUSTER, GUENTER A., 67
Science ethics
development and content, 51
for biology and chemistry majors,
Sl
world view and values, 51
Scirpus
mucronatus, 74, 77, 78
purshianus, 78
Sciuridae, 54
Sciurus carolinensis, 54
in woods of western Kentucky, 54
leaf-nest composition, 54
nest-site selection, 54
SCN8a voltage-gated sodium chan-
nel, electrophysiological studies
of, 43
SCOTT, ROGER, 47-49
Sedge
Howe, 98
prickly bog, 98
SEGEBARTH, BRAD, 46
Seneca snakeroot, taxonomic status
of, 39
Sensing system, for antimonite, 42
Sericea lespedeza, 50-84
SHIBER, J.G., 50, 51
Shiner, palezone, 98
Shrimp alpheid, snapping action of,
52
Silene
catesbaei, 16
catesbei, 15, 16
catesbri, 15, 16
virginica, 15, 16
Silphium
terebinthaceum, 16
terebinthinaceum, 16, 21
therebinthaceum, 15
Silver maple, 87
SILVERNAIL, ANTHONY F., 80
SINEX, MATTHEW, 42
SISKEL, JAMES B., 96
Skeletal compartmentalization, in
rats, 50
Sky weed, 13
Sky-weed, 17
Small pinweed, 14, 17
Smallmouth buffalo, 96
SMITH, CHRISTOPHER L., 40,
47
Smooth phlox, 14, 17
Snailnatural infection of trematodes,
23-28
Snails, 97, 98
Snakeroot, white, 13, 17
Sodium channel, SCN8a_voltage-
gated, 43
Solanum
carolinianum, 13, 16
caroliniense, 13, 16
phureja, 65
SOLE, JEFFERY D., |
Solidago rigida, 14, 16
Sorghastrum nutans, 15, 16, 21
Sorghum bicolor, 65
Southern rural elderly dietary
knowledge, 39
health status, 39
intakes of, 39
Index to Volume 57
SPARKS, LARRY, 41
Spearmiogenesis, RNA sorting sys-
tem in, 45
Spider-lily, 13, 17
Spiderlily, 20
Splicing, genes involved in, 44
SPRAKER, JOHN S., 92
STANFORD, VINCENT EVIN, 54
Stedeoma
pulegioides, 16
pulegivides, 12
Stiff gentian, 15, 17
Stiff golden-rod, 14, 17
Stiff goldenrod, 17
Stinking weed, 13, 17
Stone-dwelling cave beetle, 98
Strain-induced electrophilic cyclic
rings, 40
STRAUSS, A.M., 40
Streptococcus pheumoniae, 43
STUCKEY, RONALD L., 9
Student attitudes, toward study of
biology, 51
Stylosanthes
biflora, 14, 16
elatior, 14, 16
Stylvianthes elatior, 14, 16
Sun flower
scented, 14, 17
purple, 14, 17
Sunfish, red-eared, 96
Surprising cave beetle, 98
Swallow-tailed kite, 98
American, 98
Swamp forests
composition of, 85-91
of Mississippi alluvial plain, 85-91
structure of, 85-91
Swamp lily, 13, 17
SWIDERSKI, CAROL, 44
Sy-weed, 13, 17
Sycamore, 13, 17
Sylvilagus floridanus, 1-3
cover, 1-3
habitat, 1-3
in Kentucky, 1-3
Synandra
grandiflora, 13, 16
hispidula, 13, 16, 18
Tall fescue, 60-66
characterization of for DNA
changes, 60-66
characterization of for isozyme
changes, 60-66
characterization of for meiotic
changes, 60-66
Tall ironweed, 12, 17
Taraxacum officinale, 38
Tavin weed, 13, 17
Taxodium distichum, 85, 87, 88, 90
Taxonomic status, of varieties of
Seneca snakeroot, 39
Tea, mountain, 14, 21
109
Tectarius muricatus
on San Salvador Island, Bahamas,
52
size and density of, 52
Teleostei, 4—S
TEMPLETON, SUSAN, 39
TENNANT, RAYMOND F., 29
TGF-b Type II receptor gene mu-
tations, 44-45
THIERET, JOHN W., 99
THOMAS, ELIZABETH M.., 40
TNF-alpha neuroprotectionin opi-
ate-mediated astroglial toxicity,
46
Traditional education, a student sur-
vey, 51-52
TRAPASSO, L. MICHAEL, 39
TRAUTH, AMY E., 39
Trematodes
daily emergence, 23-28
effect of light on, 23-28
Trepocarpus aethusae, 97
Triaenodes
ignitus, 69, 72
tardus, 69, 72
Tricholomataceae, 38
Trichoptera, of the Kentucky River,
67-73
Trip planning, 92-95
Triplasis purpurea, 97
Tris, crested, 14, 16
Triticum aestivum, 65
Tritogonia verrucosa, 56, 57
Truncilla
donaciformis, 96
truncata, 55, 57
Tupelo, water, 85, 88
Turnsole, 13, 17
Turpentine weed, 15, 17
Twin leaf, 13
Twin weed, 17
Twinleaf, 17
Ulmus americana, 88
Unionid, freshwater, 55-59
in the Cumberland River, 55-59
Unionidae, 55-59
in the Cumberland River, 55-59
URSO, JAN, 45
Utterbackia imbecillis, 58
Vaccinium
album, 14, 16
stamineum, 14, 16, 21
VARAJAN, RALPH CHEL, 43
Vascular plant discoveries, from
Kentucky, 74-79
Vascular plants
common names of, 9-19
in Kentucky, 9-19
reported by C.S. Rafinesque, 9-
19
Vernonia
gigantea ssp. gigantea, 12, 16
praealta, 16
110 Transactions of the Kentucky Academy of Science 58(2)
prealta, 12, 16
Viburnum
lentago, 97
molle, 98
Villosa
iris, 57
lienosa, 57
trabalis, 56-58
Vole, prairie, 53-54
female preferences, 53-54
male odors in, 53-54
VUJANOVIC, B.D., 40
Wall flower, Ohio, 13, 17
WALLACE, DAVID, 41
WANG, C., 38, 49, 50
Water tupelo, 85, 88
WATSON, CASEY, 47, 48
Weather systems, effects on mi-
graine headaches, 39
WEAVER, DANA R., 44
White nettle, 13, 17
White prairie-clover, 15, 17
White snakeroot, 13, 17
WHITE, T.L., 50
WILBUR, FRANK H., 51
Wild currant, 14, 17
Wild indigo, blue, 16
Wild plum, 13, 17
WOODWARD, JEROLD W,, 42
Worm weed, 12, 17
Wormseed, 12, 17
WR-082-01 cell line, 42-43
WRIGHT, G.L., 50
YAN, HONG Y., 49, 50, 52
Yellow fringed-orchid, 14, 17
Yellow gentian, 15, 17
Yellow lotus, 13, 17
Yellow pea-clover, 14, 17
Yellow wort, 14, 17
Yellow-bunch, 14, 17
Yellow-flowered leafcup, 14, 17
Zannichellia palustris, 99-100
Zannichelliaceae, 99—100
Zebra mussel, 96
in the Ohio River, 96
ZHANG, Y., 49
ZIMMER, STEPHEN, 46
Zoology and entomology abstracts,
52-54
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