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Volume 58 
Number 1 
March 1997 

Official Publication of the Academy 

The Kentucky Academy of Science 

Founded 8 May 1914 

Governing Board for 1 996 

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 

Vice President: Patricia K. Doolin, Ashland Petroleum Company, P.O. Box 391, Ashland, KY 41114 
Past President: Robert O. Creek, Department of Biological Sciences, Eastern Kentucky University, Rich- 
mond, KY 40475 
Secretary; Peter X. Armendarez, Department of Chemistry and Physics, Brescia College, Owensboro, KY 

Treasurer: Julia H. Carter, Wood Hudson Cancer Research Laboratory, 931 Isabella Street, Newport, KY 

Executive Secretary (ex officio): J. G. Rodriguez, Department of Entomology, University of Kentucky, 

Uxington, KY 40546-0091 
Editor, TRANSACTIONS (ex officio): John W. Thieret, Department of Biological Sciences, Northern Ken- 
tucky University, Highland Heights, KY 41099; (606) 572-6390 
Editor, NEWSLETTER (ex officio): Maria K. Falbo-Kenkel, Department of Physics and Geology, Northern 

Kentucky University, Highland Heights, KY 41099 

Members, Governing Board 

Robert J. Barney 1999 J. G. Rodriguez 1998 

Patricia K. Doolin 1996 AAAS/NAAS Representative 

David E. Hogan 1996 Wimberly C. Royster 1997 

James P. Hopgood 1998 Robert H. Wombles 1998 

Gerald L. DeMoss 1997 

Vincent DiNoto 1996 

Chair, KJAS 

Committee on Pubucations 

Editor and John W. Thieret, Department of Biological Sciences, Northern Kentucky University, 

Chair: Highland Heights, KY 41099 

Associate Editor: James O. Luken, Department of Biological Sciences, Northern Kentucky University, 

Highland Heights, KY 41099 
Index Editor: Varley Wiedeman, Department of Biology, University of Louisville, Louisville, KY 

Editorial Board: Vincent DiNoto, Natural Science Division, Jefferson Community College, Louisville, 
KY 40201 
Jerry N. Baskins, School of Biological Sciences, University of Kentucky, Lexington, 

KY 40506 
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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March 1997 

Volume 58 
Number 1 

Trans. Ky. Acad. Sci. 58(1): 1-3, 1997. 

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 


Jeffery D. Sole 

Kentucky Department of Fish and Wildlife Resources, Frankfort, KY 40601 


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. 


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 
decHning in the state over the last several 
years (Sole 1989). Changes in land use maybe 
facihtating this decUne (Bruna 1952). 

Intensive agricultural practices (e.g., 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 hm- 

' Current address: Department of Biological and Envi- 
ronmental Sciences, Cafifomia University of Pennsylvania, 
California, PA 15419. 

iting their numbers (Barbour and Davis 1974). 
Succession of fields to forests may also reduce 
quaUty 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 dechning populations of these animals. 


We conducted our study on five wildhfe 
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 
types: (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 cottontiiils on five wildlife management areas in Kentucky, 1989. 


Foliase co\ei 

r {%)■' 


lil use (%) 


<1 111 

<2 111 
















Old field 










al field 


















' Foliage cover determined August 1989. 

' Means in a column followed bv the same letter are similar (P > 0.0.5). 

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 AN OVA followed by a Fishers 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 (i.e., 50 plots, each 1 
m-AVMA). 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 
WMAs using a factorial ANOVA followed by 
Fisher's LSD test (SYSTAT 1992). 


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). 

FoHage 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). 


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 Sigfin 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 Ilhnois. 

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. 


This study was funded by Pittman-Robert- 
son Federal Aid in Wildhfe Restoration mon- 
ies in cooperation with the Kentucky Depart- 
ment of Fish and Wildhfe Resources, P-R Pro- 
ject W-45-21, and the Wildlife Program, De- 
partment of Biological Sciences, Eastern 
Kentucky University. 


Anderson, B.F., and M.R. Pelton. 1976. Movements, 
home range, and cover use: factors affecting the sus- 
ceptibiHty of cottontails to hunting. Proc. Southeast. As- 
soc. Game and Fish Commissioners 30:.52.5-.5.3.5. 

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- 
tuck-y Fish and Wildlife Dep., Fed. Aid. Proj. 26-R., 

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. 111. 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 utiUzation-availability data. J. Wildlife 
Managem. 38:541-545. 

Nudds, TD. 1977. Quantifying the vegetative structure 
of wildlife cover. Wildlife Soc. Bull. 5:113-117. 

Pietz, PJ., 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, 

Swihart, R.K., and R.H. Yahner. 1982. Habitat features 
influencing use of farmstead shelterbelts by the eastern 
cottontail {Sylvilagus floridmius). Am. Midi. Naturalist 

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(l):4-8. 1997. 

Separation of Spawning Habitat in the Sympatric Snubnose Darters 
Etheostoma flavutn and E. simoterum (Teleostei, Percidae) 

Jean C. Porterfield 

Fish Division, Museum of Zoology, Ann Arbor, MI 48109' 


Individuals of two snubnose darter species, Etheostoma flaviim 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. flavttm 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. 


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 cichUds, 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 dtiryi, the black darter, and E. jia- 
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 
Stames 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. 
flavum (Keevin, Page, and Johnston 1989); 
both species are classified as egg attachers 
(Page 1985). Descriptions of both species can 
be found in Etnier and Stames (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 imphcations for reproductive isolation be- 
tween the two species. 


I observed spawning E. simoterum and E. 
flavum while snorkehng 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 

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. 


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. zonule 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 
females 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- 

In 1995, E. simatenim was the most com- 
mon fish near the Hwy 41 A 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 
shghtly 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 

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 simotenirn and E. caenileum 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. smoterum 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 April 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 caenileum, E. crossoptenim, 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 hlennioides, Lepom,is 
megalotis, Luxilus chrysocephalus, Lythrunis 
ardens, Micropterus punctulatus, and Pime- 
phales notatus. Observed fish associates in 
1996 at the bridge site were Campostoma an- 
oimdum, Catostomus commersoni, Etheostoma 
caeruleum, E. flabellare, Lepomis cyanellus, L. 
megalotis, Luxilus chrysocephalus, and Mi- 
cropterus punctulatus. 


The spawning behaviors of E. flavum, and 
E. simoterum observed in Whites Creek are 
nearly identical to previously pubUshed 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 (Carney and Burr 1989; Keevin, Page, 
and Johnston 1989; Page and Mayden 1981; 
Stiles 1974). 

Observations oi 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 

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 spawni- 
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 £. 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 
41 A, 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 

Etheostomn 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, whUe 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 
Stames (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 

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- 

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- 
oterurn 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. 


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 WE. 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- 


Bailey, R. M., and D. A. Etnier. 1988. Comments on the 
subgenera of darters (Percidae) with descriptions of two 
new species oi Etheostonui (Ulocentra) from southeast- 
em United States. Misc. Publ. Mus. Zool. Univ. Mich. 

Bart, H. L., Jr. 1992. Spawning behavior of Etheostoma 
davisoni Hay. Copeia 1992:537-539. 

Carney, 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. 111. Nat. Hist, Surv. Biol. Notes 134:1-16. 

Etnier, D. A., and R. M. Bailey 1989. Etheostonui {Ul- 

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. Stames. 1993. The fishes of 
Tennessee. University of Tennessee Press, Knoxville. 

Greenberg, L. A. 1988. Interactive segregation between 
the stream fishes Etheostoma simoterum and E. nifili- 
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. 50:5.5-58. 

Kessler, R. K., and J. H. Thorp. 1993. Microhabitat seg- 
regation of the threatened spotted darter {Etheostoma 
maculatum) and closely related orangefin darter (£. hel- 
Uim). 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. 111. Nat. Hist. Surv. 33:27.5-295. 

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 sinwterum, 
in Brush Creek, Tennessee. 111. 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. Komfield (eds). Evolution offish 
species flocks. University of Maine at Orono Press, 

Stiles, R. A. 1974. The reproductive behavior of the 
Green and Barren River Ulocentra (Osteichthyes: Per- 
cidae; Etheostonui). Assoc. Southeast. Biol. Bull. 21:86- 

Walters, J. P. 1994. Spawning behavior of Etheostonui 
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 


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, fisted 
410 common names with the geographic areas 
in which they were used, and 60 sources of 
information. His fist, 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 fiterature 
sources pubfished during the first third of the 
20th century or earfier. 

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- 
fished articles and books totafing over 900 ti- 
tles in botany. Many of these pubfications 
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 Engfish, 
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- 
gfish used by members of the public but not 
preferred for scholarly communication. Oth- 
erwise, in these well-known pubfications, 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 Kentucl^ 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 outfine of the vascular- 
plant vegetation in four major regions of the 
state. This paper was pubfished in 1819 in the 
second number of the first volume of The 
Western Review and Miscellaneous Magazine, 
edited by Wilfiam 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, lands of bedrock, 
and unique features of the vegetation. He also 
provided a short fist 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 


Transactions of the Kentucky Academy of Science 58(1) 

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 V4" 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 pubhshed 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 

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- 

The use of common names of plants is of 
more importance now than it was 25 or 50 

Rafinesque Common Names — Stuckey and Pringle 




On itB principal features, by C. S. Rafinesque, Professor of 
Botany and Natural History in Transylvania Universi- 

THE state of Kentucky being situated in the centre of the 
western country, has a flora similar to the generality of the 
tvestem 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 principally in the 
total want of the maritime and mountain regions, which form 
&Qch 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 whicl: 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, all over the United States; such for instance as the 
grasses, ferns, the Comfitonia, 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 instance of this singular fact are not few, 
among which I shall select the following: 

Vernonia firealta, - Iron, Weed, 

Bafitisia cerulea, Blue^Wild Indigo, , 

Cttcalia renifo'rmis, Kidney Weed, i 

Stedeoma fiulegivides, Penny-aoyal, 

C/ienofiodium anthelminthicum, Worm Weed, 
MUfihantojius scaber, Elephant's Foot, 

Figure 2. First page of C.S. Rafinesque s paper on "Bot- 
any of Kentucky[:] on its principal features," which was 
published in 1819 in volume 1(2) of The Western Review 
and Miscellaneous Magazine. 

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 stabihty 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 appHcations of 
vernacular plant names. For these reasons we 
bring to the attention of the scientific pubUc 
the common plant names in Rafinesque s early 
19th century publication on the botany of 

Rafinesques 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 fists 
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 Rafinesques intent. 
From some of his other pubfications, it is ev- 
ident that he preferred a simplified spelfing 
such as "cerulea" to the original "caendea" or 
"coendea" and "cateshei" 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. Re- 
cause of the circumstances discussed above, it 
appeared best to retain Rafinesques original 
format in the fists, which omitted the attri- 
bution of the authorship of the botanical 
names. As the fists 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 


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 modem common names we include 
have generally been obtained from Femald 
(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. 

Rafinesques 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 generahty of the western states and 
participating in their pecuhar features, while 
it offers in itself a complete specimen of the 
western botany. 

The pecuHarities 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 pecuHar 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) Vemonia prealta. Iron Weed, 

[Vemonia praealta Miclix.; V. gigantea (Walter) Trel. 
ssp. gigantea; tall ironweed] 

(2) Baptisia cendea, Blue Wild Indigo, 

[Baptisia coendea Eaton & Wright; B. australis (L.) 
R.Br, ex W.T. Alton; blue false-indigo] 

(3) Cacalia reniforrnis. Kidney Weed, 

[Cacalia reniforiyiis Miihl. ex Willd.; C. mtihlenber- 
gii (Sch. Bip.) Fern; Amoglossum muehlenbergii 
(Sch. Bip.) H. Rob.; great Indian-plantain; see Ap- 
pendix 2, entry 1, for discussion of nomenclature of 
tliis 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 

[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. Elephants Foot, 

[Elephantopus scaber L.; E. carolinianus Raeusch.; 
leafy elephants-foot; see Appendix 2, entry 3, for 
discussion of nomenclature of this species] 

(7) Gillenia stipulacea, Indian Physic, 

[Gillenia stipidata (Muhl. ex Willd.) Nutt.; Porter- 
anthus stipulatus (Muhl. ex Willd.) Britton; Ameri- 
can ipecac] 

(8) Miagia arupedinaria [Miegia arundinar- 
ia]. Cane, &c.&c. 

[Mieaia animlimiria 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 Fluviatile Region. This includes 
all the valleys, and bottoms of the large rivers. 

Rafinesque Common Names — Stuckey and Pringle 


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) Jejfersonia cinata {Jejfersonia binata), 
Tavin Weed {Twin leaf], 

[jejfersonia binata Barton; J. diphylla (L.) Pers.: 

(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; 

(14) Lupatorium calntinum [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, Hehotrope, 

[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, 

[Catalpiiim cordifiiliiini (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 trihuha [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, yeflow lo- 
tus]; see Appendix 2, entry 8, for discussion of no- 
menclature of this species] 

(22) Pancratium liviosone {Pancratium lir- 
iosme), Lily, 

[Pancratium liiiosme Raf.; Hymenocallis caroliniana 
(L.) Herbert; spider-lily; see Appendix 2, entry 9, 
for discussion of nomenclature of this species] 

(23) 7m crocea. Red hly, 

[Iris crocea Raf, nomen nudum; presumably I. ful- 
va Ker Gawl.; copper iris] 

(24) Houstonia fniticosa, Rock weed, 

[Houstonia fruticosa Raf., nomen nudum; H. nigri- 
cans (Lam.) Femald var nigricans; narrow-leaved 
houstonia; see Appendix 2, entry 10, for discussion 
of nomenclature of this species] 

(25) Pniniis pendida. Cliff plumb, &c.&c. 
[Prunus pendula Raf, nomen nudum, non Maxim, 
nee 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 Central Region. It is formed by 
the limestone tract included between the val- 
ley of the Ohio and the hilly ridges or knobs. 
The ground is sUghtly 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 naturahzed plants! There 
are hardly any species peculiar to it; but the 
foUowing ones, rare elsewhere, are here very 

(26) Eupatorium urticefolium. White nettle, 

[Eupatorium urticifolium Reichard, "urticaefolium"; 
E. rugosum Houtt.; white snakeroot] 

(27) Pavia muricata. Prickly Buck-eye, 

[Pavia nmricata Raf, nomen nudum; Aesculus gla- 
bra Willd. var. glabra; Ohio buckeye] 


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.) BSR; False pen- 

(29) Polymnia uvedalia. Scented Sun flower, 

[Polymnia uvedalia L.; large-flowered leafcup, yel- 
low-flowered leafcup] 

(30) Phlox glaberrirna. Pink, &c.&c. 
[Phlox glaberrirna 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 Hilly Region. 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 Umestone, 
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 Gaiiltheria 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 Alton; 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. nee 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 Alton, 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- 

(40) Asarum Virginicum, Heart-leaf, &c.&c. 

[Asanim virginicum L.; Hexastylis virglnica (L.) 
Small; heart-leaf little brown jugs; see Appendix 2, 
entry 13, for discussion of identity of this species] 

4. The Barren 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, 

[Solldago rigida L.; stiff goldenrod] 

(42) Polygala polygama. Nimble weed, 

[Polygala polygama Walter; racemed milkwort, bit- 
ter milkwort] 

(43) Rudbeckia purpurea. Purple Sun-flow- 

[Rudbeckia purpurea L.; Echinacea purpurea (L.) 
Moench; purple coneflower] 

Rafinesque Common Names — Stuckey and Pringle 


(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 

(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- 

(48) [{Petalosteum}] purpureum, Nimble 

[Petalostemon purpureum (Vent.) Rydb.; Dalea pur- 
purea Vent, var purpurea; purple prairie-clover] 

(49) Silphium therebinthaceum. Turpentine 

[Presumably an intentional abridgement of Sil- 
phium terebinthinaceum Jacq.; prairie-dock] 

(50) Silene catesbri {Silene catesbei}, Scarlet 

[Silene cateshaei Walter; S. virginica L.; fire-pink] 

(51) Gentiana atnarellvides {Gentiana amar- 
elloides). Yellow Gentian, 

[Gentiana amarelloides Michx.; Gentianella quin- 
quefolia (L.) Small van quinquefolia; stiff gentian; 
see Appendix 2, entry 14, for discussion of nomen- 
clature of this species] 

(52) Buchnera Americana, Black Wort, 


[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 pecuhar divisions or regions may be 

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 hable 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) 

Amoglossum 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 austrafis (L.) R. Br. ex W.T. Aiton (2) 
Baptisia cemlea (2) 
Baptisia coerulea Eaton & Wright (2) 
Buchnera americana L. (52) 
Cacalia muhlenbergii (Sch. Bip.) Femald (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 Miclix. 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 urticifofium 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) 
lodanthus pinnatifidus (Michx.) Steud. (10) 
Iris cristata Aiton (31) 
Iris crocea Raf (23) 


Transactions of the Kentucky Academy of Science 58(1) 

Iris fulva Ker Gawl. (23) 

Isanthus brachiatus (L.) BSP, (28) 

Isanthus caeruleus Michx. (28) 

Isanthus ceruleus (28) 

Jeffersonia binata (II) 

Jeffersonia cinata (II) 

Jeffersonia diphylla (L.) Pers. (11) 

Juniperus virginiana L. (34) 

Lechea minor L. (37) 

Leucospora multifida (Michx.) Nutt. (12) 

Lupatorium cakitinum (14) 

Miagia anipedinaria (8) 

Miegia arundinaria Raf (8) 

Nelumbium pentapetalum (Walter) Willd. (21) 

Nelumbo lutea (Willd.) Pers. (21) 

Orchis ciliaris L. (33) 

Pancratium liriosme Raf (22) 

Pancratium liviosone (22) 

Pavia muricata Raf (27) 

Petalostemon candidum (Michx. ex Willd.) Mickx. (47) 

Petalostemon purpureum (Vent.) Rydb. (48) 

Petalvitemon candidum (47) 

Phlox glaberrima L. (30) 

Finns rigida Mill. (36) 

Platanus occidentalis L. (9) 

Platanthera ciliaris (L.) Lindl. (33) 

Polanina graveolens (15) 

Polanisia graveolens Raf (15) 

Polanisia dodecandra (L.) DC. 

ssp. dodecandra (15) 
Polygala polygama Walter (42) 
Polymnia uvedalia L. (29) 
Populus angulata Alton (18) 
Populus deltoides Marshall 

var. deltoides (18) 
Porcelia tribuba (19) 
Porcelia triloba (L.) Pers, (19) 

Porteranthus stipulatus (Muhl. ex Willd.) Britton (7) 
Prunus munsoniana W Wight & Hedrick (25) 
Prunus pendula Raf (25) 
Ruellia caroHniensis (J.F. Gmel.) Steud. 

ssp. ciliosa (Pursh) R.W. Long (44) 
Ruellia oblongifolia Raf (44) 
Rudbeckia fulgida Aiton (38) 
Rudbeckia purpurea L. (43) 
Silene catesbaei Walter (50) 
Silene catesbei (50) 
Silene catesbri (50) 
Silene virginica L. (50) 
Silphium terebinthaceum (49) 
Silphium terebinthinaceum Jacq. (49) 
Solanum carolinianum (13) 
Solanum caroliniense L. (13) 
Solidago rigida L. (41) 
Sorghastrum nutans (L.) Nash (45, 46) 
Stedeoma pulegioides (4) 
Stylosanthes biflora (L.) BSP (32) 
Stylosanthes elatior Raf (32) 

Stylvianthes elatior (32) 
Synandra grandiflora Nutt. (20) 
Synandra hispidula (Michx.) Baill. (20) 
Vaccinium album Pursh (35) 
Vaccinium stamineum L. (35) 
Vemonia gigantea (Walter) Trel. 

ssp. gigantea (I) 
Vemonia praealta Miclix. (1) 
Vemonia prealta (1) 

B. Index to Common or Vulgar Names 

Appalachian false-foxglove (39) 

American ipecac (7) 

American lotus (21) 

American pennyroyal (4) 

Barren oats (45, 46) 

Bastard pennyroyal (4) 

Bitter milkwort (42) 

Black wort (52) 

Blue-curls (4) 

Blue-hearts (52) 

Blue penny-royal (28) 

Blue false-indigo (2) 

Blue wild indigo (2) 

Button wood (9) 

Buttonwood (9) 

Cane (8) 

Catalpa tree (17) 

Clammy-weed (15) 

Cliff plumb (25) 

Copper iris (23) 

Cotton tree (18) 

Cottonwood (18) 

Cow mint (20) 

Crested dwarf iris (31) 

Crested Hag (31) 

Crested iris (31) 

Crested tris (31) 

Deerberry (35) 

Eastem coneflower (38) 

Elephant's foot (6) 

False pennyroyal (28) 

Fire-pink (50) 

Flag (31) 

Giant cane (8) 

Great Indian-plantain (3) 

Hairy mellia (44) 

Heart-leaf (40) 

Heliotrope (16) 

Horse nettle (13) 

Indian grass (45, 46) 

Indian heUotrope (16) 

Indian physic (7) 

Iron weed (1) 

Kidney weed (3) 

Large-flowered leafcup (29) 

Leafy elephant's-foot (6) 

Lily (22) 

Rafinesque Common Names — Stuckey and Pringle 


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) 
Smafl 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) 
TaU ironweed (1) 
Tavin weed (11) 
Turnsole (16) 
Turpentine weed (49) 
Twinleaf (11) 
Twin weed (11) 
White netde (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 mAmoglossum 
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. atripUcifolia L. 
as the type, the correct name for this species will be Ca- 
calia inuehlenbergii (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 scaher L. remains the cor- 
rect name for an accepted species, but not for a species 
that occurs in Kentucky. Elephantopus scaher sensu Mi- 
chaux has been identified as the species now called E. 

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 tnie 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 
smaU 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. 


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) Engehn., 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 Rafinescjiiianus (Merrill 1949) on this 
point. It is probablv appropriate to assume from the sim- 
ilarity of the epithets, as Merrill evidently did, that Ca- 
falpium 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, Catalphim 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. bigmmioides (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 
cordifoliurn/C. cordatum from Kentucky. 

7. We have located no common name for Sijnamlra 
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 

9. The name Pancratium liriosnie 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- 
ijotis. 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 identit)' 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 Hexastijlis 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 presvunably 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 Kentvicky 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. 


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 


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 Universit}', Ja- 
maica Plain, MA. 

Newcomb, L. 1977. Newcombs 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 famifies of plants. I. Species of the genus Callitri- 
che. 11. 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. l(6):426-430. 

Rafinesque, C.S. 1817b. Flonila of the White Mountain 
of New-Hampshire. Am. Monthly Mag. & Crit. Rev. 

Rafinesque, C.S. 1817c. Florula Ludoviciana: or, a flora 
of the state of Louisiana. C. Wiley & Co., New York, 
NY. Facsimfle ed. 1967, with introduction by Joseph 
Ewan, Hafner Publishing Co., New York, NY 

Rafinesque, C.S. 1819a. Prodrome des nouveau.x genres 
de plantes observes en 1817 et 1818 dans finterieur des 
etats-Unis d'Amerique. J. Phys. Chim, Hist. Nat. Arts 

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, FL.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- 

Rafinesque, C.S. 1819d. Errata [to botany of Kentucky]. 
Western Rev. & Misc. Mag. 1(3):128. 

Rafinesque, C.S. 1820. Prodrome d'une monographic des 
rosiers del I'Amerique septentrionale, contenant la de- 
scription de quinze nouvelles especes et vingt varietes. 
Ann. Gen. Sci. Phys. 5(14):210-220. 

Rafinesque, C.S. 1824. Florida Kentuckensis. Catalogue 
of the principal trees, shrubs and plants of Kentuck)'. 
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. PubHshed 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, Arnold Arboretum, Janiiiica 
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 Hyinen- 
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 pecuhar- 

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 
famiHar 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 

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 

a Fluviatile Region does not fit most modem 
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 pecuHar 
to this region was Hmited. 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 spiderUly {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 


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. 
MuUer and McComb (1983) described the up- 
land vegetation of the Knobs as being domi- 
nated by oaks, none of which Rafinesque men- 

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 Hsted 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 IlHnois." He pointed out the 
sparsity of trees and the presence of licks. 

which tended to attract grazing animals (e.g., 
buffalo). Rafinesque's Hst 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 {Silphiiim terebinthinaceum) , and purple 
coneflower {Echinacea purpurea) are still 
found in barren remnants (Baskin and Basldn 
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 

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 Harmonic (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 


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. 


Basldn, J.M., and C.C. Basldn. 1978. Plant ecology of 
cedar glades In the Big Barrens region of Kentucky. 
Rhodora 80:.54.5-557, 

Braun, E.L. 19.50. 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. 44:44^9. 

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. 

Meijer, W. 1973. The contributions by Rafinesque to the 
early botanical exploration of Kentucky. Castanea 38: 

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 CaroUnas. 
Pages 110-306 in R.G. Thwaites. 1904. Early Western 
Travels 1746-1846. Arthur H. Clark Co., Cleveland, 

Middleton, A.R., W.R. Jillson, FT. McFarland, and WA. 
Anderson Jr. 1926. Kentucky. Pages 149-1.54 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. l(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(l):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 


Naturally infected snails, Helisoma trivolvis, releasing cercariae of Echinostonm trivolvis and Cephalogon- 
imus vesicaudus, were collected from Owsley Fork Reservoir in northern Jackson and southern 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. 


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 Hfe 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- 
tramoUuscan 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 hght 
(Smyth and Halton 1983). These exogenous 
factors either act directly on the parasite or 
are mediated indirectly through the snail host. 

That hght triggers or inhibits cercarial re- 
lease from the snail host in many species of 
digenetic trematodes is well known. Reversal 
of the 24 h hght: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- 

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 Hght or 
dark is of a constant magnitude (i.e., contin- 
uous hght 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 

Little information is available concerning 
the effect of light on emergence of cercariae 
of Echinostoma trivolvis (Echinostomatidae) 



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 Hght 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. 


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 fight (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 (i.e., releasing cercariae of E. tri- 
volvis or C. vesicaudus) were then immediate- 
ly accfimated for 24 h at 26° C to one of the 
following photoperiod regimens: (1) 12 h fight 
(0700-1859):12 h dark (1900-0659), (2) 12 h 
dark (0700-1859): 12 h light (1900-0659), (3) 
24 h fight, 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 

Cercarial release in the six 2 h fight 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 spfit 24 h photoperiods. In the 
12L:12D and 12D:12L analyses, the fight 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 fight due to the 
times selected to evaluate cercarial emer- 
gence. A Kruskal-Walfis 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. 


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 fight 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 lb). Continuous emergence 
was apparent under conditions of constant 
fight (Figure Ic; range = 12-187 cercariae/2 
h interval) and dark (Figure Id; 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 
fight and dark intervals. An average of 153.8 
± 57.5 and 2021 ± 349.7 cercariae was re- 
leased in the fight 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 fight time in- 
tervals, respectively. Peak emergence was ob- 

Cercarial Emergence — Rosen et al. 














fj C^ r*% r'^ 

fl f^ fl f^ fl 


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 Hght, 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 
Ught periods (Figures 2a and 2b). Continuous 
cercarial emergence was observed under con- 
ditions of constant Hght (Figure 2c; range = 
499-2298 cercariae/2 h interval) and dark 
(Figure 2d; range = 177-808 cercariae/2 h in- 

The long-term study showed that the mean 
daily time for peak cercarial emergence was 
significantly different for E. trivolvis (Kruskal- 
Walhs test: P = .0002), but not for C. vesi- 
caudus (Kruskal-WalUs test: P=.9482). Peak 
























Si II 








































■ T 





"7 ^ 









I\= - 








*— 1— 

— r- 

f^ f) f*^ fO c*^ f*^ 

f*^ fo p*^ ^ 
O r4 3 \o 

o o o o 


Figure 2. Effect of light on the mean number ± SE of 
Cephalooonimus vesicaudiis 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- 

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- 
caudiis remained fixed at 2230 (dark period) 
over the same time (Figure 4). 


Light significantly triggered the release of 
E. trivolvis cercariae and inhibited the emer- 
gence of C. vesicaudiis cercariae in our 1990 
and 1992 studies. Photoperiod inversion was 


Transactions of the Kentucky Academy of Science 58(1) 































' — 1 1 — 

— 1 — 

1 — 












_^ / 








L N 







D — 

u / 














O — 

1 1 — 

— 1 — 

1 ' 

Figvire 3. Comparison of the effect of light on the mean 
number ± SE of Echinostoma frivolvis 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- 


Figure 4. Comparison of the effect of light on the mean 
number ± SE of Cephalogoniinns vcsicatidus 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/ 

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 Ught: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. 


bach and Alldredge 1974). Such an "entrain- 
ment" of the endogenous rhythm to Ught: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 hfe cycle as an adult in the small 
intestine of the spiny softshell turtle Trionyx 
spinifenis, 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 hkely 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 tiian 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 Ufe 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 estabfish 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. 


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 EdUn, Jose Ilagan, 
Kiely Law, and Manuel San. 


Asch, H.L. 1972. Rhythmic emergence of Sclmto.soma 
man.soni cercariae from Biomphalaria glabrata: control 
by ilknnination. 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 F.J. 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: 

Lewis, M.C., I.G. Welsford, and G.L. Uglem. 1989. Cer- 
carial emergence of Proterometra macrostoma and P. 
edneiji (Digenea: Azygiidae): contrasting responses to 
light:dark cycling. Parasitology 99:21.5-223. 

Luttermoser, G.W, 1955. Studies on the chemotherapy of 


Transactions of the Kentucky Academy of Science 58(1] 

experimental schistosomiasis III. Harvest of Schistoso- 
ma 77uinsoni 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 Schistosonuitiitin douthitti cercariae from their snail 
host. J. Parasitol. 37:201-204. 

Petrie, J.L., E.F. Burg III, and G.D. Cain. 1996. Molec- 
ular characterization of Ecliinosfoina caproni and E. 
paraensei by random amplification of polymoiphic 
DNA (RAPD) analysis. J. Parasitol. 82:360-362. 

Riley, M.W., and G.L. Uglem. 1995. Proterometra ma- 
crostoina (Digenea: Azygiidae): variations in cercarial 
morphology and physiology. Parasitology 110:429^36. 

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 nmnsoni cercariae: ecological significance 
in transmission to human and murine hosts. J. Parasitol. 

Theron, A., and C. Combes. 1988. Genetic analysis of 
cercarial emergence rhythms of Schistosoma nuinsoni. 
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: 

Trans. Ky, Acad. Sci. 58(l):29-32. 1997 

Classifying Free Bieberbach Groups 

Raymond F. Tennant 

Department of Mathematics, Statistics, and Computer Science, 
Eastern Kentucky University, Richmond, KY 40475 


Let F be a free group on n letters and G be a finite group. To each epimorphism e : F — ^ G is associated 
a free Bieberbach group F/N' where N is the kernel of the epimorphism e 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. 


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)-l) + l, where F is the free 
group on k generators with k=mk(A) (Lyndon 
and Schupp 1977, p. 16). 

It is well known that B is an n-dimensional 
Bieberbach group if and only if U"/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 
exact sequence, 1— >N— >F— >G— >1. Let 
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, 
^N^F^G^l. 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. 

Two free Bieberbach groups F, and Fo are 
isomorphic if there exists a ZG-module iso- 
morphism a and a group isomorphism P such 
that the following diagram commutes: 

^ N, ^ F, ^ G ^ 1 

aX (Bvl =xl 
^ N, ^ F. ^ G ^ 1 

where Nj and No 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 (p: G ^ GL(n,Z). Since F=Z"XG (as 
sets), then we may define multipUcation on F 
by (n,g)(m,h) = (n+cp(g)m + f(g,h),gh), where 
n,meZ" (thought of as column vectors), 
g,h€G, and feH2(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 
normahzed set map and i: Z" — > F is the in- 
clusion map. It is known (Tennant and Turner 
1992) that F is torsionfree. For the case, 
G7^{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". 



Transactions of the Kentucky Academy of Science 58(1) 


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^Zg. 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 O^Z-^B-^Zo— >1 where 
[R7B is homeomorphic to the Klein bottle. 

In dimension n = 3, a free Bieberbach 
group (other than Z^ with trivial point group) 
must have G=Zo and mk(F)=2. A result of 
Linnell (1981) states that if two epimorphisms 
G,: F, ^ G and g,: Fo -^ G, with 
mk(F,) = mk(F2), are Nielsen equivalent, then 
their associated free Bieberbach groups are 
isomorphic. Since any two epimorphisms e i. 
Go-. F(a,b) -^ Zo are Nielsen equivalent, con- 
sider the epimorphism derived from the pre- 
sentation of Zo given by {a,b | a-, b>. This pre- 
sentation and the action of Zo on Z^ induces a 
monomorphism (p: Zo -^ GL(3,Z). Since 
det((p(a))= — 1, the flat manifold associated to 
this free Bieberbach group is non-orientable. 
The crystaUographers 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=Z3 and mk(F)=2. 
Since there is only one Nielsen equivalence 
class of epimorphisms e : F(a,b) -^ Z3, there is 
only one free Bieberbach group with point 
group G^Zj. Since 2 does not divide |G|=3, 
the induced monomorphism tp: Z3 — > GL(4,Z) 
must satisfy det((p(g)) = l for aU geG and so it 
follows that the flat manifold W/F is orientable. 


Let k be a positive integer. If k is odd, there 
is one free Bieberbach group of dimension 
n=k+ 1 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 g: F(a,b) -> 

G is an epimorphism. If G=Z|, or G=D^, the 
dihedral group with k elements and 9: G -^ 
GL(k+l,Z) is the monomorphism induced by 
the action of G on Z" then there exists geG 
with ord(g)=2 such diat det((p(g))=-l. It fol- 
lows that the associated flat manifold is non- 
orientable. For the point group, G=Z|„ aU epi- 
morphisms, G : F(a,b) -^ Zi, are Nielsen equiv- 
alent so all free Bieberbach groups with point 
group G=Zi, are isomorphic. On the other 
hand, there exist k such that not aU epimorph- 
isms G : F(a,b) -^ D,. are Nielsen equivalent, so 
we may not apply LinneUs result to conclude 
that there is only one (k+l)-dimensional free 
Bieberbach group with point group G=Di^. Let 
G : F), ^ G be an epimorphism where F^ is free 
on k generators. Suppose another generator is 
added to F,, and mapped to 1 under g to form 
a new epimorphism 6 *: F,,+ i -^ G. The foUow- 
ing theorem describes how the respective free 
Bieberbach groups are related. 


Let G : F|, ^ G and g *: F^^+i ^ G be epi- 
morphisms such that G *(Xi) = G (x,), l<i<k and 
G*(x^+i) = l, then F* is isomorphic to Fx,ZG 
where F and F* are the free Bieberbach 
groups associated to g and g * respectively 
and Xj represents a semi-direct product. 


Recall that F may be expressed in terms of 
the short exact sequence 


The action by conjugation of G on N induces 
a monomorphism cp: 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)-^ 
By viewing F as setwise equal to N X G we may 
define multiplication on F by (n,g)(m,h) = 
(n + cp(g)iTT+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 
cp*: G -^ Aut(N*) that satisfies P(A(9*(g))). 
The free Bieberbach group may be expressed 
in terms of the short exact sequence 

Free Bieberbach Groups — Tennant 


A set map s*: G -^ F* can be chosen so that 
the 2-cocycle f*: GXG -^ 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 e : N* 
^N b>^e(n*)=p(a(n*)). Then ^ ZG ^ 
N*— >N— >lisa short exact sequence of 
ZG-modules. The group F* is equal to N*XG 
(as sets) so define the epimorphism g : F* ^ 
F by e(n*, g) = (e(n*),g). The ker(G)=ZG 
and we have the following commutative dia- 













i i 

^ N ^ F 

G^ 1 


I i 

1 1 

Define the normalized set map s: N — > N* 
by s(n)=a"'(n/0) where is the additive iden- 
tity in ZG. The set map s: F ^ F* defined by 
s(n,g) = (s(iT),g) is then a homomorphism and 
so it follows that the sequence -^ ZG — > F* 
^ F ^ 1 splits and F*=Fx,ZG. 


For the case where F*=Fx3ZG, 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 U"/F is an n- 
dimensional flat manifold sitting inside of the 
(n+|g|)-dimensional flat manifold R"+I^'l/F*. 

Corollary 1 

If Epi(F|,+ i,G) contains only one Nielsen 
equivalence class and g : F,, — > G and e *: Fi,+ i 
—> G are epimorphisms then F*=Fx5ZG. 

As an iUustration, the (k+l)-dimensional 
free Bieberbach group (k odd) with point 

group G=Z;, can be factored as the semi-di- 
rect product F=Zx,Z(Zi,). 

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 ^1,^2' ^u ~^ 
G, there exists a finite sequence of epimorph- 
isms ei,eo •••, e^ where e^: F,,+ i -^ G for i odd 
and Cj: Fi, ^ G for i even, l^i^r such that 
for each of the pairs of epimorphisms, (e i,ei), 
(ei,e,+,) for l<i<r— 1, and {e^,Go), the epi- 
morphism on F,, is isomorphic to the epi- 
morphism on F|,+i restricted to some subset 
(of k basis elements) of F,,+i. 

Corollary 2 

Let G be a finite group, and g 1,60: F,, ^ G 
be two epimorphisms. If G possesses the swap 
property then FiX^ZG^FsX^ZG where F, and 
F, are the free Bieberbach groups associated 
to G 1 and G 2 respectively. 


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 g i,G2: F,, 
— > G are two epimorphisms then F,X5ZG= 
F2X5ZG where F, and Fg are the free Bieber- 
bach groups associated to g 1 and g 2 respec- 

When F,x,ZG=F2X5ZG is said to be one step 
stably equivalent to F2. 


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. 11:511-5.51. 

Janner, A., and T. Janssen. 1979. Superspace groups. 
Physica A 99:47-76. 

Linnell, RA. 1981. Relation modules and augmentation 
ideals of finite groups. J. Pure Appl. Algebra 22:143- 

32 Transactions of the Kentucky Academy of Science 58(1) 

Linnell, P.A. 1985. A cancellation theorem for lattices Lyndon, R.C., and RE. Schupp. 1977. Combinatorial 

over an order. J. Lond. Math. Soc. II. 31:450^56. group theory. Springer Verlag, New York, NY. 

Lyndon, R.C. 1962. Dependence and independence in Tennant, R.F., and E.G. Turner. 1992. The swap conjec- 

free groups, J. Heine Angew. Math. 210:148-173. ture. Rocky Mt. }. Math. 22:1083-1095. 

Trans. Ky. Acad. Sci. 58(l):33-34. 1997. 


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 1-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 

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 

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 Anumg America's 
Teachers this year The Kentucky Academy of Science is 
proud to present to Dr Karan Kaul the 1996 Outstanding 
Universitv Science Teacher Award. 

Outstanding Secondary School 
Science Teacher Awards 

Douglas C. Jenkins — Instructor, Warren Central High 
School, Bowling Green, Kentucky 



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 — Instnictor, 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 

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' 
accomphshments 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(l):35-36. 1997 


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 leiicopiis, 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. abdo?ninalis . These par- 
asitize eastern cottontail rabbits and are the two members 
of this group ranging into Kentucky (R.D. Hall, pers. 

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 1 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 ex-perienced " 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- 

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 sldn. 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- 

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 sldn 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. Saimders, Pliil- 



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 Barldey 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 Peromysctts. 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 k-nown 
cases (Diptera: Cuterebridae). J. Med. Entomol. 3:263- 

267. (7) Catts, E.R 1982. Biology of the New Worid bot 
flies: Cuterebridae. Ann. Rev. Entomol. 27:313-338. (8) 
Catts, E.R 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, PL., and C.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(l):37-54. 1997 

Abstracts of Some Papers Presented at the 

1996 Annual Meeting of the 

Kentucky Academy of Science 


Geographic differentiation and allo2yme variation in 
pawpaw {Asimina triloba, Annonaceae). HONGWEN 
HUANG* and DESMOND R. LAYNE, Land-Grant Pro- 
gram, Atwood Research FaciUty, 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 (I). 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 

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 seedUngs. E.xperimental treat- 
ments were imposed from sowing until the plants were 
destructively harvested. The e.xperimental design was a 
split-plot where blocking was done by position in the 
greenhouse. The main plot of the ex-periment was shade. 
This was accomplished by growing seedlings under a 
wooden frame covered with shade cloth to reduce inci- 

dent fight 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 2X/week from seedfing 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- 

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 protnision (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 hvpocotyl 
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- 



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 (Pohjodon 
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 (Pohjodon 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. 


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 fisted 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 ex-posed 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 Mt/cena leaiana (Tricholomataceae) 
basidiospores. DONALD G. RUCH* and MARK C. 
BOBBINS, 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-hke 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- 
SON,* Biolog)' 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 Annucil Meeting 


fore, that dandelion scape elongation is influenced both 
by light microenvironment and stage of development. 

Taxonomic status of the varieties of Seneca snakeroot, 
Polijgala senega (Polygalaceae). AMY E. TRAUTH* and 
ROBERT F. C. NACZI, Department of Biological Sci- 
ences, Northern Kentucky University, Highland Heights, 
KY 41099. 

Seneca snakeroot, Pohjgala 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 sofl 
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. 


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 

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 


Differences in health and diet status: African- American 
vs. Caucasian rural southern elderly. MARTHA MAR- 
man Nutrition Research, Kentucky State University, 
Frankfort, KY 40601. 

We surveyed 1554 mral 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- 


Transactions of the Kentucky Academy of Science 58(1J 

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- 
Iv, 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. 


Polynomial conservation laws of the generalized Em- 
den-FowIer 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,* Depart:ment of Chemical and 
Materials Engineering, University of Kentucky, Lexington, 
KY 40506. 

We consider the polynomial conservation laws of fourth 
degrees wdth respect to x of the generahzed Emden-Fow- 
ler equation x + (b/t) x + 7x°'t'' = 0. We demonstrate that 
the existence of conservation laws depends upon the so- 
lution of a system of partial differential equations, usually 
tenned 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 

Surface geometry of the Morehead radio telescope 
and the optimization of operating parameters. BRIAN M. 
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. 


Alkylation of DN A by strain-induced electrophilic cyclic 
MERS-GOODWIN, Depart:ment of Chemistry, Univer- 
sity of Kentucky, Lexington, KY 40502. 

Strain-induced electrophilic cycHc 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- 
1-one and 2-propanone-3-(3-phenylcyclobuten-l-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-l-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 Ap 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 exliibit 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 Ap and reduced trophic 

Assessment of metal impact in the Robinson Forest 
creek system using metalHothionein. KEVIN COMP- 
TON,* and CHRISTER HOGSTRAND, Department of 
Biology, University of Kentucky, Lexington, Ky 40502. 

Abstracts, 1996 Annual Meeting 


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 
pubhshed 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 neuroto.xicity hypothesis: possible 
evidence through L-type calcium channel density In hip- 
pocampal subregions. ALEXANDER COON,* ROSE- 
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 
[^H]PN200-I10 on 25 human hippocampus samples (AD, 
n = 15; control, n = 10). Tissue was sectioned to 20 |i,m 
using a cryostat and then incubated with 1.0 nM 
[^H]PN200-110. Non-specific binding was binding in the 
presence of 1.0 ixm 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 stifl unclear at this time. 

Characterization of AD-like pathology in normal aging. 
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 
NeurofibriUary Tangles (NFT's). SP's contain the 3-amy- 
loid protein, a 39-42 amino acid peptide fragment derived 
from a transmembrane amyloid precursor protein ((3- 
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 P-APP peptide compro- 
mise the P-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 (3-APP, precludes the formation and deposition of 
SP's. The two immediate derivatives of the C-100 frag- 
ment are P-A42(43) and P-A40. It has been shown that 
p-A42(43) is found in greater accumulations then (3-40 in 
Alzheimer's disease. To examine the presence of 
P-A42(43) and P-A40 in normal aging we used specific 
antibodies to discrete regions of the (J-APP peptide se- 
quence. We found several individuals who had the 
P-42(43) with lower densities then that of Alzheimer's dis- 
ease. There were no accumulations of the ^-40 in normal 
aging. It should also be noted there was no correlation 
between the presence of (3-A42(43) and age. These results 
suggest that the (S-APP processing in normal aging is sim- 
ilar to that found in Alzheimer's disease. 

Chromosome 17 deletions in a subset of ovarian tu- 
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- 


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 
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 
v^dth antigen into the eye. Using an adoptive transfer 
mouse model vwth 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 
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. 
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 ex-pression, 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 nin23 proteins in the WR-082-01 cell line. 
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 


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 HI and H2 isoforms of mn23 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-Hl and H2 resulted in a 
general repression (< 2-fold) of all promoter constructs 
analyzed. While these nm23 isofonns exerted no addition- 
al inhibition upon vectors containing the 5 'SHS element, 
an additional two-fold repression was conferred by HI 
upon vectors containing the entire NRR. These data sug- 
gest that nm23-Hl 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. MARCl 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 microsateUite 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 
fines 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 

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 isoforni 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 vafidate 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 
BARAO BONDADA, Department of Microbiology and 
Immunology, University of Kentucky, Lexington, KY 

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% 


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 pro\'ided, sup- 
porting the concept that the poor response of neonates to 
polysaccharides is likely due to a deficiency in cytokine 

Identification of genes involved in splicing. ELIZA- 
BETH OTTE* and BRIAN RYMOND, Department of 
Biolog)', 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 P-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 (3-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 
stabihty, 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 stmctural 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 exliibited 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 carrjdng 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 ex-pression 
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-P Type II re- 
ceptor gene mutations in human pancreatic carcinoma. 
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 


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-p. This loss in regulation is 
often mediated by lack of expression of the TGF-P 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 RII 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 
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 
l-methyl-4-phenyl-l,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- 
Hne 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- 

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- 


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, 
n, 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, = 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 £. 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 

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 dnigs 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 jiM). 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- 
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 Idlodalton (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 grouch 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 


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-l/eIF-4E interaction 
and suppress malignant properties in oncogenic cells. 


Celestial seasons. RICHARD HACKNEY,* KAREN 
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- 

Comet Hale-Bopp: Opportunity for education and de- 
veloping student involvement. ROGER SCOTT,* KAR- 
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 
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- 
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 191.5 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 instmment for undergraduate faculty and student 
research in radio frequency astrophysics. BENJAMIN K. 
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 


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 laborator)' 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 instrimien- 
tation technology in a cutting-edge science, one that is in 
the midst of scientific revolution. This presentation pro- 
vides and overview of the M RT and its current operating 

NASA Kentucky Space Grant Consortium, Center for 
Automated Space Science, and EPSCoR opportunities for 
space-related science, technology, and education. KAREN 
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, Rowling Green, 
KY 42101; MICHAEL BUSBY Center for Automated 
Space Science, Center of E.xcellence 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 facult)' 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- 

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 laboratorv 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 ex-plore, 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 Laborator)' Ex-periences in Astronomy, 
with copies available upon request. A list of soft\vare 
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- 
em Kentucky University Astrophysical Observatory, Bowl- 
ing Green, KY. 

Photometry with CCDs: Optimizing the signal extrac- 

Abstracts, 1996 Annual Meeting 


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. 


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 

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 
Astnmotas ocellatiis (CichUdae). 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/stnictural damages caused by 
the gentamicin to the fish kidney. The experimental sub- 
jects, the teleost fish Astronotus ocellatiis, 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 |xCi/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 ocellatiis (Cichlidae). STEVEN M. 


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 ototo.xic 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 

Relationship between muscle lipid content and body 
weight of paddlefish (Pohjodon spathula). X. LOU, T L. 
WHITE,* C. WANG, and S. D. MIMS, Community Re- 
search Service, Kentucky State University, Frankfort, KY 

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 ^H-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 {Pohjodon 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 {Pohjodon 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 (TEARS) 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 TEARS 
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. 
SHIEER, 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 cofleges (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 


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 concerns if they are to produce the "finest" 
graduates for the 21st century. 

Student attitudes toward the study of biology. JOHN R. 
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 

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 ex'perimental 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 


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: II. 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. 


Analyses of size and density of Tectarius muricatus (Lit- 
torinidae) on San Salvador Island, Bahamas. MELINDA 
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 \dr- 
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 vWth a computer-based SIGNAL analysis 
system. Time, frequency signal, and spectrogram corre- 

Abstracts, 1996 Annual Meeting 


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 

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. imhricata (which 
lacks nectaries). We studied a set of 10 H. latispatha, 10 
H. imhricata, and 8 hybrids, all growing along the edge 
of the landing strip at Estacion 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. imhricata 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 o( Heliconia (Heliconiaceae) 
with respect to the presence of extrafloral nectaries. THO- 
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- 
hricata, 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 imhricata attracted 
significantly fewer insects than H. latispatha. In fact H. 
imhricata attracted so few insects that we conclude that 
it does not produce nectar on the buds. We wifl 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 whilfe it was occurring in the lab; we did find 
remains of six cannibalized victims (some stifl 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. 


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- 
surgerv, 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 anestnis 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 

Impacts of dam construction on densities and distri- 
butions of bald eagles, Haliacettis leucocephaliis, 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 constnic- 
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 constmc- 
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 i/4-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 Sciunis 
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 (Sciiints 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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B. Papers (in triplicate) may be submitted at any time to 
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John W. Thieret 

Biological Sciences 

Northern Kentucky University 

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Phone: (606) 572-6390 

FAX: (606) 572-5639 


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Transactions of the Kentucky Academy of Science 58(1) 

thors use name of the first author followed by et al. — 
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Lacki, M.J. 1994. Metal concentrations in guano from a 
gray bat summer roost. Transactions of the Kentucky 
Academy of Science 55:124—126. 


Ware, M., and R.W. Tare. 1991. Plains life and love. Pi- 
oneer Press, Crete, WY. 


Kohn, J.R. 1993. Pinaceae. Pages 32-50 in J.F. Nadel 
(ed). Flora of the Black Mountains. University of 
Northwestern South Dakota Press, Utopia, SD. 


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serts of Trans-Pecos Texas. 2nd ed. Ocotillo Press, 
Yucca City, TX. 



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KAS Annual Meetings 

The 1997 annual meeting of the Kentucky Academy of Science will be held 13-15 November 
at Morehead State University, Morehead; the 1998 meeting, at Jefferson Community College, 
Southwest Campus, Louisville. 


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, Uxington, KY 40508-4008; ISBN 0-8131-0898-5; price $17.95 
(+$3.50 shipping). 


Relationships Among Habitat, Cover, and Eastern Cottontails (Sylvila- 
gus floridanus) in Kentucky. William M. Giuliano, Charles L. Elliott, 
and Jeff ery D. Sole 1 

Separation of Spawning Habitat in the Sympatric Snubnose Darters Eth- 
eostoma flavum and E. simoterum (Teleostei, Percidae). Jean C. Por- 
terfield 4 

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 9 

Some Comments on Constantine Rafinesque's 1819 Description of Bo- 
tanical Regions in Kentucky. William S. Bryant 20 

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 23 

Classifying Free Bieberbach Groups. Raymond F. Tennant 29 

1996 33 


Human Myiasis in Kentucky Caused by Cuterebra sp. (Diptera: Oestri- 
dae). Lee H. Townsend 35 



k^^ ■ THE 

Volume 58 
Number 2 
September 1997 

Official Publication of the Academy; 

The Kentucky Academy of Science 

Founded 8 May 1914 

Governing Board for 1997 

Executive Committee 

President: Marcus T. McEUistrem, Department of Physics and Astronomy, University of Kentucky, Lexing- 
ton, KY 40506-0055 
President Elect: Patricia K. Doolin, Research and Development, Ashland Petroleum Company, P.O. Box 

391, Ashland, KY 41114 
Vice President: Gordon K. Weddle, Department of Biology, Campbellsville University, Campbellsville, KY 

Past President: William S. Bryant, Department of Biology, Thomas More College, Crestview Hills, KY 

Secretary: Peter X. Armendarez, Department of Chemistry and Physics, Brescia College, Owensboro, KY 

Treasurer: Julia H. Carter, Wood Hudson Cancer Research Laboratory, 931 Isabella Street, Newport, KY 

Treasurer Elect: William E. Houston, 161 Morningstar Court, Bowling Green, KY 42103 
Executive Secretary (ex officio): J. G. Rodriguez, Department of Entomology, University of Kentucky, 

Lexington, KY 40546-0091 
Editor, TRANSACTIONS (ex officio): John W. Thieret, Department of Biological Sciences, Northern Ken- 
tucky University, Highland Heights, KY 41099; (606) 572-6390 
Editor, NEWSLETTER (ex officio): Maria K. Falbo-Kenkel, Department of Physics and Geology, Northern 

Kentucky University, Highland Heights, KY 41099 
Chair, Junior Academy of Science (ex officio): Vincent A. DiNoto Jr., Department of Physics, Jefferson 

Community College SW, 1000 Community College Drive, 
Louisville, KY 40272 
Program Coordinator (ex officio): Robert O. Creek, Department of Biological Sciences, Eastern Kentucky 

University, Richmond, KY 40475 

Members, Governing Board 

Robert J. Barney 1999 

Gerald L. DeMoss 1997 

James F. Hopgood 1998 

Bruce A. Mattingly 2000 

Committee on Publications 

Editor and John W. Thieret, Department of Biological Sciences, Northern Kentucky University, 

Chair: Highland Heights, KY 41099 

Associate Editor: James O. Luken, Department of Biological Sciences, Northern Kentucky University, 

Highland Heights, KY 41099 
Index Editor: Varley Wiedeman, Department of Biology, University of Louisville, Louisville, KY 

Editorial Board: William S. Bryant, Department of Biology, Thomas More College, Crestview Hills, KY 
John P. Harley, Department of Biological Sciences, Eastern Kentucky University, 

Richmond, KY 40475 
Marcus T. McEUistrem, Department of Physics and Astronomy, University of 
Kentucky, Lexington, KY 40506-0055 

All manuscripts and correspondence concerning manuscripts should be addressed to the Editor. 

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@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 

Barbara L. Rafaill 


J. G. Rodriguez 


AAAS/NAAS Representative 

Lee A. Roecker 


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September 1997 

Volume 58 
Number 2 

Trans. Ky. Acad. Sci. 58(2):.5.5-.59. 1997. 

Continuing Decline in the Freshwater Unionid (Bivalvia: Unionidae) 
Fauna in the Cumberland River Downstream from Cumberland Falls, 


Ronald R. Cicerello and Ellis L. Laudermilk 

Kentucky State Nature Preserves Commission, 801 Schenkel Lane, 
Frankfort, Kentucky 40601 


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 decUne 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. 


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 

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. Ry 1947-1949, Cijdonaias tuhercidata, 
ElUpsaria lineolata, Ohliquaria reflexa, and 
Ptijchobranchus subtentum were lost from the 
site, but Villosa iris had been found. Retween 
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. 


Cumberland Falls, a 17-m-high barrier to 
the upstream movement of aquatic organisms 
(Rurr 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 (Rower and Jackson 1981) is ca. 84% for- 



Transactions of the Kentucky Academy of Science 58(2) 

River km 




Figure 1. Cumberland River study area downstream 
from Cumberland Falls, McCreary and Whitley counties, 
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 mVsec, consid- 
erably less than the mean of 26.6 m Vsec for 
July-September, when streamflow is at or near 
the annual low, and the mean annual discharge 
of 89.7 mVsec (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, 


In 1987, 1989, and twice in 1993, we ex- 
amined a 0.5-km-long river segment from the 
base of Cumberland Fafls 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 
snorkehng and with waterscopes, and the 
shoreline was examined for shells. Nomencla- 
ture follows Hoeh (1990) and Turgeon et al. 


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, Ptij- 
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. Wflson 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 


Table 1. Freshwater unionids collected from the Cumberland River downstream from Cumberland Falls, Kentucky, 
in four studies: 1910-1911, 1947-1949, 1961, 1987-1993, 








Actinonaias ligamentina (Lamarck) 





Actinonaias pectorosa (Conrad) 





Cijclonaias tuberculata (Rafinesque) 


Ellipsaria lineolata (Rafinesque) 


Elliptio crassidens (Lamarck) 





Elliptio dilatata (Rafinesque) 





Laiupsilis cardium (Rafinesque) 


Lampsilis fasciola Rafinesque 




Lampsilis ovata (Say)^ 



Lasmigona costata (Rafinesque) 




Ligumia recta (Lamarck) 





Medionidus conradicus (Lea) 




Ohltquaria reflexa Rafinesque 


Potamilus alatus (Say) 





Ptychobranchus fasciola lis (Rafinesque) 





Ptijchobranchiis suhtentum (Say)'^ 


Qiiadnda piisttdosa (Lea) 





Tritogunia verrucosa (Rafinesque) 





Tnincilla tntncata Rafinesque 



Villosa iris (Lea) 



Villosa lienosa (Conrad)^ 



Villosa trabalis (Conrad)' 




Total species 





Total individuals 




ca. 88 

' Wilson and Clark (1914); - Neel and Allen (1964); ' Stansbery (1969); ' maxiniiini ninnber obsen'ed 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; = 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) o( 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 Bumside, 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 poflutants 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 


Transactions of the Kentucky Academy of Science 58(2) 

cold, turbulent water released from Lake 
Cumberland preclude unionid reproduction, 
and only Cumherlandia monodonta Say and 
Cijclonaias tuherculata 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. capsaefomiis, 
Ptijchobranchus 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. 


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. 


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. Drainag^ areas of 
streams at selected locations in Kentucky. Open File 
Report 81-61. Geological Survey, U. S. Depai|tment of 
the Interior. | 

Burr, B.M., and M.L. Warren Jr. 1986. A distdbutional 
atlas of Kentucky fishes. Kentucky State future 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 luiionids (Bi- 
vaKda: Unionoidea) of Kentucky. Am. Malacol. Bull. 8: 

Dennis, S.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 
State Universit)', Blacksburg, VA. 

Gordon, M.E., and J.B. Layzer. 1989. Mussels (Bivalvia: 
Unionoidea) of the Ciunberland River: review of life 
histories and ecological relationships. U. S. Fish and 
Wildhfe Service Biological Report 89(15). 

Harker, D.F, Jr., M.L. Warren Jr., K.E. Camburn, S.M. 
Call, G.J. Fallo, and P. Wigley 1980. Aquatic biota and 
water quality survey of the upper Cumberland River 
basin. Kentucky Nature Presen'es Comm., Tech. Rept., 
Frankfort, KY. 

Hoeh, W.R. 1990. Phylogenetic relationships among east- 
ern North American Anodonta (Bivahda: Unionidae). 
Malacol. Rev 23:63-82, 

Houslet, B.S. 1996. Age, growth, and stability of a mussel 
assemblage in Horse Lick Creek, Kentuck)'. Master's 
thesis, Tennessee Technological University, Cookexille, 

Kentucky Division of Water. 1994a. Cumberland River 
biological and water quality investigation. Wild River 
segment. Kentucky Di\dsion of Water, Frankfort, KY. 

Kentuck)' Di\asion of Water. 1994b. 1994 Kentucky re- 
port to congress on water quality. Kentucky l^ivision of 
Water, Frankfort, KY. 

[KSNPC] Kentucky State Nature Preserves Commission. 
1996. Rare and extirpated plants and animals of Ken- 
tucky. Trans. Kentucky Acad. Sci. 57:69-91. 

Mayes, Sudderth, and Etheredge, Incorporated. 1975. 
The river basin water quality management plan for 
Kentuck)'. Upper Cumberland River. Prepared for Ken- 
tucky Division of Water, Frankfort, KY. 

McGrain, P. 1966. Geology of the Cumberland Falls 
State Park area. Kentucky Geological Survey Series X, 
Spec. Publ. II. University of Kentucky, Lexington, KY. 

Miller, A.C., L. Rhodes, and R. Tippit. 1984. Changes in 
the naiad fauna of the Cumberland River below Lake 
Cumberland in central Kentucky. Nautilus 98:107-110. 

Neel, J.K., and W.R. Allen. 1964. The mussel fauna of 
the upper Cumberland basin before its impovmdment. 
Malacologia 1:427-459. 

Ortmann, A.E. 1924. The naiad-fauna of the Duck River 
in Tennessee. Am. Midi. Naturalist 9:18-62. 

Schuster, G.A., R.S. Butler, and D.H. Stansbeiy 1989. A 
survey of the unionids (Bivalvia: Unionidae) of Buck 
Creek, Pulaski County, Kentucky. Trans. Kentucky 
Acad. Sci. 50:79-85. 

Smathers, K.L. 1990. An analysis of a bivalve (Mollusca: 
Bivalvia) community in the Licking River, at Moores 
Ferry, Kentucky. Master's thesis. Eastern Kentucky 
University, Richmond, KY. 

Stansbery, D.H. 1969. Changes in the naiad fauna of the 

Cumberland Unionids — Cicerello and Laudermilk 


Cumberland River at Cumberland Falls in eastern Ken- 
tucky. Bull. Am. Malacol. Union 1969:16-17. 

Thompson, Y.L. 1985. The mussel fauna of the Rockcastle 
River system, Kentucky (Bivalvia: Unionidae). Masters 
thesis. Eastern Kentucky University, Richmond, KY. 

Turgeon, D.D., A.E. Bogan, E.V. Coan, W.K. Emerson, 
W.G. Lyons, W.L. Pratt, C.EE. Roper, A. Scheltema, 
F.G. Thompson, and J.D. Williams. 1988. Common 
and scientific names of aquatic invertebrates from the 
United States and Canada: Mollusks. Am. Fish. Soc. 
Spec. Publ. 16:1-277. 

[USFWS] United States Fish and Wildlife Service. 1994. 
Endangered and threatened wildlife and plants. De- 
partment of the Interior, Washington, DC. 

[uses] United States Geological Survey 1993. Water re- 
sources data — Kentucky. Water year 1993. Data Report 
KY-93-1, Louisville, KY 

Wilson, C.B., and H.W Clark. 1914. The mussels of the 
Cumberland River and its tributaries. United States 
Fish Commission, United States Bureau of Fisheries 
Document 781:1-63. 

Trans. Ky. Acad. Sci. 5S(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' 


Timothy D. Phillips 

Department of Agronomy, University of Kentucky, 
Lexington, KY 40546-0091 


Pre\aous research identified four tall fescue (Festuca antndinacea 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 
e.\|3lant 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 
mg/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 
groMh, 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. 


Tall fescue {Festuca anindinacea 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 (Neotijphodiiim coenophialum 
(Morgan- Jones & Gams) Glenn, Bacon, & 
Hanhn). 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 depeiidably 
from anthers. 

' To whom correspondence should be addressed. Present 
address: USDA-ARS-SPA, NRGEEC, PO. Box287, 2900 
Hwy 130 East, Stuttgart, AR 72160. 

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 


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. 


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 fxE/mVs 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), Idnetin (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 II, 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 a 3 X 4 X 2 factorial design with trial, 
genotype, and 2,4-D concentration (cone.) be- 
ing the factors. (Each petri plate was a unit of 
rephcation.) This analysis showed that the trial 
X 2,4-D cone, genotype X 2,4-D cone, and 
trial X genotype X 2,4-D cone, interactions 
were non-significant. Only the trial X geno- 
type interaction was significant, so the data 
were reanalyzed with only the trial X 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 cone, 
and Idnetin 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: X 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 (3-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-isoamyl 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 l%(w/v)SDS, and 
further purified with a phenol-chloroform ex- 

RAPD analysis was modified from Williams 
et al. (1990) for use on a Perkin-Elmer DNA 
Thermocycler 480 (PE Applied Biosystems, 


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 |xM of each dNTP 
(Boehringer Mannheim, Indianapohs, IN), 
37.5 ng decamer primer (Operon Technolo- 
gies, Inc., Alameda, CA), 1.0 U Taq polymer- 
ase (Boehringer Mannheim, IndianapoUs, IN), 
and buffer (10 mM Tris pH 8.3, 50 mM KCl, 
1.5 mM MgClg) in 25.0 |xL volume. The ther- 
mocycler was programmed for six cycles of 
94°C for 1 min, 35°C for 1 min, 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 
fxL of each sample, electrophoresed on 1.2% 
agarose gels stained with 0.06 fxg/fxL 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, 

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: OPE9, OPFIO, OPJ15, OPJ19, 
OPK19, OPTOl, 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 


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 Idnetin 
did not significantly affect the expression of 
these same variables. The lack of response to 
Idnetin 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 II, 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 


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 varialile 


Callns growth 

Hoot growth 

Albino shoots 

CJreen shoots 

Ceen plants 


responding per 

100 anthers 


0.25 ab- 

0.00 a' 

0.25 be' 

0.00 a- 

0.00 a- 


0.67 a 

0.33 a 

2.17 a 

0.33 a 

0.33 a 


0.00 b 

0.00 a 

0.00 c 

0.00 a 

0.00 a 


0.67 a 

0.00 a 

0.75 b 

0.00 a 

0.00 a 


0,33 b 

0.00 a 

0.17 a 

0.00 b 

0.00 b 


0.67 ab 

0.00 a 

0.00 a 

0.44 a 

0.67 a 


0.83 a 

0.17 a 

0.11 a 

0.11 ab 

0.17 ab 


0.00 a 

0.00 b 

0.00 a 

0.00 b 

0.00 b 


0.04 a 

0.00 b 

0.00 a 

0.00 b 

0.00 b 


0.33 a 

0.67 a 

0.04 a 

1.13 a 

0.88 a 


0.16 a 

0,00 b 

0.04 a 

0.00 b 

0.00 b 


Trial II 

Trial 111 

' Significant at the 0.05 level using a square root transformation of the least-square means. 


m pi 


^^^ ^^W 

25 27 

25 &27 

25 27 

25 27 

25 27 

25 27 


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. 


Transactions of the Kentucky Academy of Science 58(2) 

K25 Materal K27 

Regenerants plants Regenerants 

^12 3 4 


1636 bp 

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 tisstie rather than from microspores. Genomic 
DNA was amplified using the primer OPFIO. 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 II 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 K27. 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 Itrans- 
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 bivalerits, 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 
ACFH-1, GOT-2, GOT-3, 6-PGD-l 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 OPFIO primer (Figure 3). Thus, pan- 
icle morphology and meiotic, isozyme, and 

Tall Fescue Plantlets — Eizenga and Phillips 


RAPD analyses indicated there was no cul- 
ture-induced variation for these parameters. 


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 Cornelius 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 
regenerabihty 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 hicolor (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. 


The authors acknowledge the excellent 
technical assistance of Mrs. Etta Mae Thacker 
and the statistical advice of Dr. Paul L. Cor- 


Transactions of the Kentucky Academy of Science 58(2) 

nelius. This manuscript is Scientific Journal 
Series No. 96-06-114 of the Kentucl<y 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 


Guenter A. Schuster 

Department of Biological Sciences, Eastern Kentucky Universit)', 
Richmond, KY 40475 


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, Cheumatopsijche campyla and Ceraclea 
tarsipu aetata 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. 


The collection, evaluation, and monitoring 
of aquatic communities over periods of time 
is a tool used to evaluate water quahty 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, hght 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- 

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 o{ Hijdwpsyche 
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. 


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 



Transactions of the Kentucky Academy of Science 58(2) 

Dix River 

South Fork Kentucky River 
Middle Fork Kentucky River 

North Fork Kentucky River 

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. Kentvicky River caddisflies. Location of collect- 
ing stations along the river mainstem. 

1 Carroll Co., Lock and Dam (L&13) 1, at River 

Mile (RM) 4.0 (River Kilometer [Rkm] 6.4) 
upstream from Carrollton 

2 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 


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- 
hft 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. 
Pubhshed 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 


Table 2. Trichoptera collected along the Kentucky River 
mainstem from 1993 through 1996. 

Table 2. (continued). 

Protopfila maculata (Hagen) 

Ceratopsyche sparna (Ross) 
Cheumatopsijche cainpijla Ross 
Cheumatopsijche mimiscida 

Cheumatopsijche oxa Ross 
Cheumatopsijche paseUa Ross 
Hijclropsyche betteni Ross 
Hydropsyche frisoni Ross 
Hydropsyche orris Ross 
Hydropsyche simulans Ross 
Potamyiaflava (Hagen) 

Hi/droptila ajax Ross 
Hydroptila angusta Ross 
Hydroptila armata Ross 
Hydroptila consimilis Morton 
Hydroptila hamata Morton 
Hi/droptila perdita Morton 
Hydroptila uaubesiana Betten 
Ochrotiichia tarsalis (Hagen) 
Ochrotrichia xena (Ross) 
Orthotrichia aegerfasciella 

Orthotiichia cristata Morton 
Oxyethira pallida (Banks) 

Ceraclea ancylus (Vorhies) 
Ceraclea cancellata (Betten) 
Ceraclea Jlava (Banks) 
Ceraclea punctata (Banks) 
Ceraclea tarsipunctata (Vorhies) 
Ceraclea transversa (Hagen) 
Mystacides sepulchralis (Walker) 
Nectopsyche Candida (Hagen) 
Nectopsyche exquisita (Walker) 
Nectopsyche pavida (Hagen) 
Oecetis avara (Banks) 
Oecetis cinerascens (Hagen) 
Oecetis inconspicua (Walker) 
Oecetis persimilis (Banks) 
Triaenodes ignitus (Walker) 
Triaenodes tardus Milne 

Ironoquia punctatissima 

Pycnopsyche lepida (Hagen) 

Chimarra obscura (Walker) 


Agnjpnia vestita (Walker) 
Ptilostomis ocellifera (Walker) 
Ptilostomis semifaciata (Say) 
Cy melius fraternus (Banks ) 





























































































































































1 2 


4 5 


Ncu reclipsis crepitsculaiis 




Polycentropus cinereus Hagen 




Polycentropus confusus Hagen 



Rhyacophila lobifera Betten 





32 36 


24 24 


tucky River basin, which included sites in the 


Adult caddisflies were collected from six 
stations (Figure 1) along the Kentucky River 
mainstem from 1993 through 1996. Initially, 
stations 2 and 3 were sampled weekly during 
1993-94; stations 1, 4, 5, and 6 were added in 
1994. Stations 3 and 5 represented open-river 
habitats; remaining stations were at L&D. Lo- 
cations of all stations are shown in Table 1. 

Collections were made using a plastic ice 
chest tray with about 500 ml of 70% ethanol, 
set at the base of a 12 volt Coleman lantern, 
equipped with ultra-violet (UV) bulbs. At 
L&D sites, a lantern was turned on and placed 
on top of the downstream lock gate mooring 
wall at dusk; it was turned off an hour after 
dark. Private docks were used at the open-riv- 
er sites in the same manner. Species identifi- 
cations were based mostly on males. 


This study began in early April 1993, with 
weekly collections at stations 2 and 3 (Figure 
1). The purpose was to document species 
composition and emergence periods from dif- 
ferent habitat types (i.e., from a L&D site and 
a typical open-river site). A year of weekly col- 
lections showed that species compositions be- 
tween stations 2 and 3 were similar (Table 1). 
However, the functional feeding requirements 
of some species reflected the types and 
amounts of different microhabitats between 
the stations. For instance, differences in num- 
bers and species of hydropsychids were nota- 
ble at the L&D site compared to the open- 
river site. So stations 1, 4, 5, and 6 were added 
in 1994 to increase the scope of the study. Sta- 


Transactions of the Kentucky Academy of Science 58(2) 

Table 3. Seasonal occurance of trichoptera collected during this stud\' from the KentncW Rher Mainstem. 



12 3 4 12'! 


Protoptila maculata 

Ceratopsyche sparna 

Cheumatopsyche campy la 
Cheumatopsyche minuscula 

Cheumatopsyche oxa 
Cheumatopsyche pasella 

Hydropsyche betteni 
Hydropsyche fi-isoni 

Hydropsyche orris 
Hydropsyche simulans 

Polamyia flava 

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 1 and 2 species at open- 
river sites (Table 1). Station 2 had eight of the 
10 hydropsychid species. Hydropsyche orris 
and Potami/ia 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 
ohscura 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 Pohjcentropus 
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 DLx 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 


Table 3. (continued). 




Hydroptila armata 
Hydroptila consimilis 
Hydroptila hamata 
Hydroptila perdita 
Hydroptila waubesiana 
Ochrotrichia tarsalis 
Ochrotrichia xena 
Orthotrichia aegerfasciella 
Orlhotrichia cristata 
Oxyethira pallida 
Ceraclea ancylus 
Ceraclea cancellata 
Ceraclea jlava 
Ceraclea punctata 
Ceraclea tarsipunctata 

. . . . m 


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 ancijhis and C.flava were col- 
lected only at stations 1 and 2, and Mijstacides 
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 

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 lohifera). 
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- 


Table 3. (contintied). 

Transactions of the Kentucky Academy of Science 58(2) 




Ceraclea transversa 
Mystacides sepulchralis 
Neclopsyche Candida 
Nectopsyche exquisita 
Nectopsyche pavida 
Oecetis avara 
Oecetis cinerascens 
Oecetis inconspicua 
Oecetis persimilis 
Triaenodes Ignitus 
Triaenodes tardus 

Ironoquia punctatissima 
Pycnopsyche lepida 

Chimarra obscura 





Table 3. (continued). 




Agrypnia vestita 
Ptilostomis ocellifera 
Ptilostomis semifaciata 

Cyrnellus fratern us 
Neureclipsis crepuscularis 
Polycentropus cinereus 
Polycentropus confusus 

Rhyacophila lobifera 

, ^mtmmimmtmmt . 

Kentucky River Caddisflies — Houp and Schuster 


sonal-occurrence data show that while the ma- 
jority of species emerge throughout the sum- 
mer some species, e.g., Rhijacophila lohifera, 
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 Cheumatopsij- 
che campijla. 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 


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. 


Beckett, D.C. 1982. Phenology of Hi/dropsyclic orris 
(Trichoptera: Hydropsychidae) in the Ohio River? 
changes in lan'al age structure and substrate coloniza- 
tion rates. En\aronm. 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: 

Beckett, D.C, CR. 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. Surv. 
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. 43:494^99. 

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, PRI024, PRI025, PR1026, 
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. Vamer. 
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 Kentuck)' University, Richmond, 

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., PA. 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. Emdronm. 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): 

Rafinesque, CS. 1820. Monographic des coquilles bi- 
valves fluviatiles de la Ri\dere Ohio, contenant douze 
genres at soixante-huit especes. Ann. Gen. 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. Sen'., Cooke\ille, TN. 

Williams, J.C 1974. Commercial fisheries investigations 
of the Kentucky River. Part I of 111: fish population 
studies and mussel bed surveys. Kentucky Dept. Fish 
and Wildhfe Res., Frankfort, KY. 

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 Kentuck)- University, 
Richmond, Kentucky 40475 


C. Tom Bloom 

1209 Glade, College Station, Texas 77840 


Nineteen species of rare vascular plants are documented from Kentucky. Two species {Jiincus validus 
Coville and Scirpus [Schoenoplectus] mucronatus L.) are reported as new to the state; S. mucronatus has 
serious potential as an invasive wetland weed. Three others {Calopogon tuberosiis (L.) BSP. var. tuherosus, 
Fohjgala pauclfolia Willd., and Rlujnchospora macrostachya Torr. var. rnacrostachya) represent the only 
knouai e.xtant Kentucky populations. New count)' distributional information for 14 other taxa of rare Kentucky 
plants is also presented. 


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 


Adiantum capiUus-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 

Significance. CranfiU (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.; CranfiU 1980; McCoy 
1938; Medley 1993). In addition, CranfiU 

(1980) cited a formerly adventive colony fron 
Nelson County. This collection represents a 
westward range extension for this epipetric 

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 CO.: 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 (j 1253 (EKY). CUMBERLAND CO.: 
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 7.5 
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 — Libbij, Mears, and Bloom 


bridge, steep, rocky riverbank, 14 Sep 1995, 
Libbij 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 o{ Aureola ria patula 
in Kentucky and indicate that this species may 
not be as rare as previously thought (Krai 
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 CO.: 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- 

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. fubero- 
.SU.S. 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 locafities 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, 

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. 


Transactions of the Kentucky Academy of Science 58(2) 

KSNPC Endangered (reproducing trees only). 
HARLAN CO.: Louellen 7.5 Quad., Hi-Lewis 
Pine Savanna, upper s-facing slopes of Hi- 
-Lewis Branch, near Hiram, 24 Jun 1995, Lib- 
hij, Lihhij, Libhij, and Evans 988 (BEREA, 

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 
bhght [Cnjphonectria parasitica (Murr.) Barr] 
(Hicks and Marks 1990). 

Cenchnis 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, 

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. 

Cypeni.s crocem 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. 

"^Cypeni.s dijformis 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 dijfor- 

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 Cypenis dijformis 
as one of the worlds 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. 

Epdobium ciliafum 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." 

Junciis 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 CO.: 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 
Barthell 7.5 Quad., ca. 2 miles se of Bald 
Knob at USES 575/USFS 6130 junction, 
USES 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), CampbeU (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, Hears, and Bloom 


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 CO.: 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. Turks-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 Liliiim superbiim, 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. 

"^Liniini usitatissinium L. MADISON 
CO.: Berea 7.5 Quad., growing in tire rut on 
roadside at the comer of Elm Street and Scaf- 
fold Cane Road, a European cultigen, escaped 
or adventive, 16 Jul 1993, Libby 343 (BEREA, 

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 coflection 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 CO.: 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; Campbefl n.d.; Medley 1993). Our col- 
lections document its occurrence in two ad- 
ditional counties. 

Polygala paucifolia WiUd. LAUREL CO.: 
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). 

RJiynchospora macrostachya Torr. var. ma- 
crostachya. KSNPC Endangered. WAYNE 
CO.: Pamell 7.5 Quad., Cooley's Meadow, ca. 
0.8 air mile ssw of Murl, wet meadow ne of 
Cooley's tobacco bam and e of Hack Denny's 
cornfield, 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 coflection may 
represent the only knowTi extant population of 
tall beak- rush in the state. 

"^Scirpus mucronatus L. [ = Schoenoplectus 
mucronatus (L.) Paflas] PULASKI CO.: Ano 
7.5 Quad., pond at the end of USES 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 

Significance. Scirpus mucronatus has not 


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 Cronquist (1991) did not list this 
species as occurring within northeastern Unit- 
ed States. This species is morphologically very 
similar to the native Sciiyus purshiaims 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). 


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 Wildhfe 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. 


Adams, R.M., II. 1981. A systematic study and mono- 
graph of the tiirk's-cap liUes of eastern North America 

(with introduction of some new cultivars). Ph.D. dis- 
sertation. Cornell University, Ithaca, NY. 

Adams, R.M., II, and W.J. Dress. 1982. Nodding Liliitm 
species of eastern North America (Liliaceae). Baileya 

Arnold, S.C, and E.O. Beal. 1981. A sur\'ey 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 Presences Com- 
mission, Scientific and Technical Series Number 5, 
Frankfort, KY. 

Braun, E.L. 1941. Notes on Kentucky plants V. Castanea 

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 Cyjjeraceae. Austral. 
Syst. Bot. 8:125-305. 

Biyant, W.S. 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, Cij- 
pents, and KyUinga (Cyperaceae) in Mississippi with re- 
cords of eight species previously unreported to the 
state. Sida 16:171-182. 

Campbell, JJ-N. n.d. Atlas of the flora of Kentucky. Lex- 
ington, KY. [Unpublished draft] 

Chester, E.W., B.E. Wofford, R. Krai, 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 aOies of Kentuckv. Ken- 
tucky Nature Presei"ves 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., 
Kentucks' State Nature Preser\'es Commission, Frank- 
fort, KY 

Gibson, D. 1961. Life-forms of Kentucky flowering 
plants, Am. Midi. Nat. 66:1-60. 

Gillett, J.M. 1959. A revision of Bartonia and Oholaria 
(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, 

Godfrey, R.K., and J.W. Wooten. 1979. Aquatic and wet- 

Kentucky Vascular Plant Discoveries — Libbij, Mears, and Bloom 


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] Kentuck)' State Nature Preser\'es Commission. 
1996. Rare and extirpated plants and animals of Ken- 
tucky. Trans. Kentucky Acad. Sci. 57:69-91. 

Krai, 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 R8-TP2. United States De- 
partment of Agriculture, Forest Sei-vice, 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 (Categor)' 2) plants 
for federal hsting. Tech. Rept., Kentucky State Nature 
Preserves Commission, Frankfort, KY. 

Lipscomb, B.L. 1980. Ci/penis diffonni.s L. (Cyperaceae) 
in North America. Sida 8:320-327. 

McCoy, T.N. 1938. The ferns and fern allies of Kentucky. 
Am. Fern. J. 28:41^6, 101-110. 

McFarland, F.T. 1942. A catalogue of the vascular plants 
of Kentucky. Castanea 7:77-108. 

Mears, R.L., and G.W. Libby 1995. New records of Cy- 
pent-s (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, Louis\dlle, KY. 

Pennell, F.W. 1935. The Scrophulariaceae of eastern 
temperate North America. Acad. Nat. Sci. Philadelphia 
Monogr 1:1-650. 

Strausbaugh, PD., and E.L. Core. 1978. Flora of West 
Virginia. 2nd ed. Seneca Books, Morgantown, WV. 

[USFWS] United States Fish and Wildfife 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. Kv. Acad. Sci. 58(2):80-84. 1997. 

Effects of Acidic Minesoil on Nodulation of the Legume Sericea 

Lespedeza (Lespedeza cuneata; Fabaceae) by Bradyrhizobiutn 

Nitrogen-fixing Bacteria 

Gary R. Cline and Anthony F. Silvernail 

Community Research Service, Kentucky State University; Frankfort, KY 40601 


Sericea lespedeza (Lespedeza cuneata (Dumont) G. Don; Fabaceae) is a N^-fixing legiune 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 
Brady rhizobium 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,(S04)j 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 
Al2(S04)3-amended minesoil at pH levels of 4.2 and 4.5, which tended to affect nodulation similarly. 
Bradyrhizohium 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. 


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 

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 

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 No 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 No-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 (CHne and Senwo 
1993; Vogel 1981), the tolerance of its No-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 


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 No 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. 


We conducted a pot experiment with four 
acidity treatments and four inoculation treat- 
ments using a completely randomized factorial 
design with eight replications. Inoc ilation 
treatments consisted of (1) a noninoculated 
control, which received inorganic N, (2) a 
commercially available mixture of Brady rhi- 
zobium sp. strains (strain numbers 4122, 3223, 
32H1, 150B1, and 176A22 from Liphatech 
Inc., Milwaukee, WI, USA), and (3, 4) strains 
41A and CRl, 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 
CaCOa (Ca-msoil), and minesoil adjusted to 
pH 4.5 (lAl-msoil) or 4.2 (2Al-msoil) with 4.0 
or 5.3 g/kg Al(SO,)3(H20)3, 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 suppUed 
with 20, 64, 32, and 5 mg/kg Ca, P, K, and 
Mo, respectively. Following amendment with 
CaCOj or Al2(S04)3, minesoil treatments were 
equilibrated for 8 weeks at a moisture content 
of 15 ml HoO/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- 

'Serala' sericea lespedeza was grown from 
seed in a greenhouse for 20 weeks under full 
sunhght. Additional supplemental light (300 
(xmol photon/mVs photosynthetically active ra- 
diation) was provided with hafide lamps to 
provide 16-hour photoperiods. Noninoculated 
plants received NH4NO3 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 
millihters of solution containing a total of 10'* 
colony-forming units of Brady rhizobium 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 
HoO g/soil) for 16 hours and centrifuged at 
3000 X g. Supematants 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 supematants 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 

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. 


Transactions of the Kentucky Academy of Science 58(2) 

Table 1. Mean values of acidity-related parameters (±SE, n = 8) measured in extracts of minesoil acidity treatments 
used to monitor growth of sericea lespedeza and nodulation by Bradijrhizobiiim nitrogen-fixing; bacteria. 




Acidih treatment 





1 Al-msoil 

2 Al-msoil 

6.41 ± 0.06 
5.33 ± 0.03 
4.46 ± 0.04 
4.18 ± 0.05 

5.96 ± 0.06 
5.15 ± 0.04 
4.54 ± 0.06 
4.24 ± 0.06 


12 ± 

85 ± 3 

178 ± 10 

6 ± 

25 ± 1 

186 ± 10 

310 ± 17 


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 |xM 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- 
dy rhizobium 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- 

trol. The fact that sericea lespedeza was able 
to produce significant yields in 2Al-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 CRl inoculation treatments. This sug- 
gested that sericea lespedeza inoculated with 
Brady rhizobium probably did not receive suf- 
ficient fixed No for maximum growth, even at 
the most optimal pH level (i.e., Ca-msoil). 


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 CRl formed similar numbers of 
nodules in both untreated minesoil and Ca- 
msoil, but nodule weight production was sig- 

Table 2. Mean values of sericea lespedeza mean shoot diy weight production (g ± SE, n = 8) obtained in minesoil 
acidity treatments using different strains of Bradijrhizobium nitrogen-fixing bacteria. Values in columns followed by 
different letters were significantly (F < 0.05) different. 



Acidit)' treatment 




Control + N 


1 Al-msoil 

2 Al-msoil 

2.94 ± 0.17a 
2.96 ± 0.14a 
2.69 ± 0.14a 
2.67 ± 0.17a 

3.26 ± 0.18a 
2.92 ± 0.14ab 
2.73 ± 0.17bc 
2.52 ± 0.14c 

2.98 ± 0.16ab 
3.30 ± 0.24a 
2.63 ± 0.08b 
2.87 ± 0.16ab 

3.53 ± 0.23a 
2.98 ± 0.40b 
2.26 ± 0.24c 
2.41 ± O.Uc 

Servicea Lespedeza and Minesoil Acidity — Cline and Silvemail 


Table 3. Mean values of nodule numbers (±SE, n = 8) obtained for sericea lespedeza grown in minesoil acidity 
treatments using different strains o{ Bradijrhizohium nitrogen-fixing bacteria. Values in columns followed by different 
letters were significantly (P ^ 0.05) different. 


Acidit)' treatment 




Control + N 


1 Al-msoil 

2 Al-msoil 

53.0 ± 6.9a 
29.2 ± 3.7b 
35.7 ± 5.9b 
36.7 ± 4.0b 

71.9 ± 18.3a 
42.3 ± 6.5b 
27.9 ± 7.0bc 
26.8 ± 4.6c 

52.4 ± 8.3a 
52.0 ± 11.5a 
24.2 ± 4.0b 
29.4 ± 3.5b 


nificandy (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 lAl-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 lAl-msoil and 2Al-msoil acidification 
treatments, indicating that nodule dry weight 
production was suppressed similarly by both 
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 lAl- 
msoil and 2 Al-msoil treatments, no significant 
differences were detected among nodule dry 

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 CRl, 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 41 A and CRl 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. 


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 lAl- 

Table 4. Mean nodule dry weight production (mg ± SE, n = 8) obtained for sericea lespedeza grown in minesoil 
acidity treatments using different strains of Bracltjrhizobium nitrogen-fixing bacteria. Values in columns followed by 
different letters were significantly (P ^ 0.05) different. 


Acidity treatment 




Control + N 


13.9 ± 2.0a 

26.1 ± 5.6a 

21.6 ± 2.2a 



13.5 ± 1.9a 

23.3 ± 3.1a 

14.2 ± 2.3b 


1 Al-msoil 

7.2 ± 1.3b 

8.0 ± 1.4b 

9.4 ± 1.1c 


2 Al-msoil 

8.9 ± 1.3b 

10.0 ± 1.2b 

9.3 ± 1.0c 


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 Bradi/rhizobium 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 


This research was funded by a USD A/ 
CSREES grant to Kentucky State University 
under agreement KYX- 10-89- IIP. 


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. Pi. 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 oi Lespedeza cuneata grown in minesoil. Pi. Soil 

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. 

CaqDenter, PL., 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- 
dijrhizobium symbiotic relationship. J. Pi. Nutr. 16: 

Chne, G.R., and Z. Ngewoh Senwo. 1994. Tolerance of 
lespedeza Bradyrliizobium 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 to.xicities 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 varsdng 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. 

Lyle, E.S., and E.M. Evans. 1979. Revegetation of Ala- 
bama coal surface mines for soil cover and forage pro- 
duction. Reclamation Rev. 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. 1988a. Kentucky profile. U.S. 
Dept. of the Interior. Washington, DC. 

Office of Surface Mining. 1988b. O.S.M. reclamation en- 
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 

Reeder, J.D., and WA. Berg. 1977. Plant uptake of in- 
digenous fertilizer from a Cretaceous shale and coal 
mine spoils. Soil Sci. Soc. Am. J. 41:919-921. 

Roberts, J.A., W.L. Daniels, J.C. Bell, and D.G. Martens. 
1988. Tall fescue production and nutrient status on 
southwest Virginia mine soils. J. Environm. Qual. 17: 

U. S. Bureau of the Census. 1988. Statistical abstract of 
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. 

Vogel, W.G., and W.A. Berg. 1968. Grasses and legumes 
for cover on acid strip-mine spoils. J. Soil Water Con- 
servation 23:89-91. 

Trans. Ky. Acad. Sci. 58(2):8.5-91. 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 


Three swamp forests on the Mississippi alhmal plain of the Jackson Purchase Region of Kentucky are 
described. One forest was dominated by Taxodium clistichum, one by Nyssa aquatica, and one by T. cUstichiiin 
and A', aquatica. In terms of basal areas and densities, these forests compared favorably to other httle 
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 Kentuck)'. These swamps are remnants of a once more 
abundant wetland system on the Mississippi alluxdal plain. 


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. 

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 fist 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 

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 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 



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 

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. 


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 baldc)'press, 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- 
gression 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: 

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 equitabihty (evenness) of the stand 
(J) was calculated using a method based on the 
Shannon-Wiener function (H'): J = HVHmax, 
where Hmax is the diversity under maximum 
equitabihty (H' = lognS), and S is the number 
of species in the sample. 


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. Number (N), relative density (RD), relative dominance (RDo), importance value (IV), and importance 
percentage (IP) for Black Slough, Carlisle County, Kentuckx. 






Taxodium distichum 






Planera aquatica 






Fraxi nus p rofu n da 






Salix nigra 






Forestiera acuminata 












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-/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 cijlindrica, Hypericum wal- 
teri, Lycopus ruhellus, Ciiscuta sp., and Saii- 
rurus cemuus. 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, Uttle 
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. 

Basal area was 85.3 m'/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, baldc)^ress 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 rnVha; and average converted basal 
area was 71.3 mVha. Those values are com- 
parable to other little disturbed swamp forests 
in the southeastern United States (e.g., An- 

Table 2. Number (N), relative density (RD), relative dominance (RDo), importance value (IV) and importance 
percentage (IP) for the Fish Lake Swamp Forest, Ballard County, Kentucky. 






Nt/ssa aquatica 
Taxodium distichum 






Quercus lyrata 












88 Transactions of the Kentucky Academy of Science 58(2) 

Table 3. Niiniber (N), relative densit)- (RD), relative dominance (RDo), importance value (IV), and importance 
percentage (IP) for the Oscar Bottoms Swamp Forest, Ballard CounK', Kentucky. 






Taxodhnn distichuni 






Nyssa aquatica 






Quercus lijrata 






Acer sacchaiinum 






Fraxinus pen nsi/hanica 






Ulmus americana 






Populus heterophtjUa 






Popiilus deltoides 






Planera aquatica 






Platanus occidentalis 






Salix nigra 






Forestiera acuminata 












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 

Diameter Size Classes (cm.) 

Figure 1. Size-class distributions for baldcypress ( 
diiun distichuni) 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 )T and 416 )t, 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. 


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 

30 60 90 

Diameter Size Classes (cm.) 

Figure 2. Size-class distribution for all trees of baldcy- 
press (Taxodiuni distichuni) and water tupelo (Nyssa 
aquatica) in the Oscar Bottoms Swamp, Ballard County, 
Kentucky. Cray data points = baldcypress; black data 
points = water tupelo. 

Kentucky Swamp Forests — Bryant 


swamps are covered by water throughout most 
of the year (Penfound 1952). Basal areas (mV 
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 nibnim) 
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 iUinoinensis), 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 Kentuck)'. 

Lilley Cornett Woods (Martin 1975) 

Sugar maple-Basswood-Tulip poplar 

30,0 m^/ha 


42.4 m-/ha 

Chestnut oak 

29,1 m^/ha 

Beech-White oak 

28.8 mVha 

Lillev Cornett Woods (Muller 1982) 

Old growth composite 

27.8 m"/ha 

Bonayer Forest (Bougher and 

Winstead 1974) 

Oak forest 

31.9 m-/lia 

Boone County Cliffs Nature Preserve 

(Biyant 1978) 

Total forest 

27.2 m^/ha 

Lloyd Wildlife Presen'e Forest 

' (Bryant 1985) 

Mixed forest 

35.5 nr/ha 

Dinsmore Woods Nature Preserve 

(Held and Winstead 1976) 

Oak-ash-maple forest 

28.1 m-/ha 

Myers Woods (Held 1980) 

Beech forest 

37.1 m-/ha 

Greenwood Forest (Chester 

et al, 1995) 

Wet woods 

37.8 m-/ha 

Upper Bottoms (Biyant 1991) 

Sugarberry forest 

35.6 m-/ha 

Martin Creek (Martin 1983) 

Southern red oak 

29.2 m-/ha 

Cvpress Swamp (Mitsch 

et al. 1991) 

Stagnant swamp 

35.9 mVha 

Semi-permanent flooded 

32.7 m-/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 


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 >l.m dbh were not unusual, but 
none as large as the 2.3 m dbh reported by 
DeFriese (1880) were found. 

In dieir 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- 


Adams, B., G.E. Hunter, D.F. Austin, and K.H. Kernick. 

1968. The flora of Murphy's Pond. Trans. Kentucky 

Acad. Sei. 29:25-28. 
Anderson, R.C., and J. White. 1970. A cypress swamp 

outlier in southern Illinois. Trans. Illinois State Acad. 

Sci. 63:6-13. 
Barry, J.M. 1980. Natural vegetation of South Carolina. 

Univ. South Carolina Press, Columbia, SC. 

Benson, K.B. 1986. Hydrologic patterns and phosphonis 
dynamics in forested wetlands of western Kentucky. 
Ph.D. dissertation, Univ. Louisville, Louisville, KY. 

Bougher, C.K., and J.E. Winstead. 1974. A phytosocio- 
logical study of a relict hardwood forest in Barren and 
Edmondson counties. Trans. Kentucky Acad. Sci. 35: 

Braun, E.L. 1950. Deciduous forests of eastern North 
America. The Blakiston Company, Philadelphia, PA. 

Bray, J.R., and J.T Curtis. 1957. An ordination of the 
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Trans. Ky. Acad. Sci. 5S(2):92-95. 1997. 

Planning a Trip 

John S. Spraker, Daniel C. Biles, Mark P. Robinson 

Department of Mathematics 

Western Kentucky University 

Bowling Green, KY 42101 


The problem of minimizing the driving time for a trip is considered. A hyjjothetical trip from Louisville 
to Memphis is used as an illustration. A mathematical model is constructed and a solution technique is 
devised and implemented. 


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. 


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. 


We now consider the general problem of 
minimizing the driving time for a trip. One has 
a set of acceptable departure times S = {f,,, t^, 
fc, . . .), where t^^ <t^ <tc, < . . . (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 
To. Of course, we require f,, < T, < To. 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) = fit, x(f)), where t is the time after de- 
parture and xit) is the distance travelled at 
time t. Finally, we let D he 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 


Table 1. Planning a trip. Calculation of the trip time for 
the trip starting at 1200. 

Figure 1. Planning a trip. Illustration oi the trip starting 
at noon. 

these assumptions force/ to be discontinuous. 
Removal of these restrictions is discussed in 
the last section. Letting t represent the de- 
parture time and T the arrival time, we thus 
obtain the following model: 

Choose f so that T — f is minimized subject to 

x'if) = fit, x{t)), t G [t, T], 
x{t) = 0, 
x{T) = D, 
T, <T < To, 

t G S,t< T. 

Since /is discontinuous, a differentiable solu- 
tion may be impossible to find. For example, 


0, if < f < 1 

f e [0, 2] 

J, if 1 < f < 2, 
x(0) = 0. 
One is tempted to propose 

fo, if < /^ < 1 

as a solution; however, x'(l) does not exist and 
hence x does not satisfy the differential equa- 
tion. As a result, we must modify the classical 
definition of solution. 

Definition 1: x: [t, T] — > K is a solution to 

x'it) = fit, xit)),t G [i, T] 
x(f) = Xo 

Ti me of day 

Distance tni\ellecl 















if and only if i) x is continuous on [t,T], ii) x'it) 
= fit,xit)) for all but a finite subset of [t,T], 
iii) x(f) = Xq. 

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 / is assumed to satisfy the 

Definition 2: / : [f,,, TJ X [0,D) -^ R is 
hlockwise-constant if and only if there exists a 
finite collection of disjoint sets A, = [it, x) : a, 
< t < hi, c, <x<d,} such that UA, = [fo. To) 
X [0,D) and/|\; is constant for each i. 

Example 1: Consider /: [0, 1) X [0, 30) -^ R 
defined by 

fit, x) = 

15 if- < f < -, 5 < X < 6 

4 2 

35 elsewhere 

This / could model a 30-mile stretch of high- 
way on which the speed limit is 35, except for 

Table 2. Planning 

trip. Output from travel problem 

Starting time 

.•\in\al time 

Traxel time 



6 hr 42 min 



6 hr 39 min 



6 hr 34 min 



6 hr 29 min 



6 hr 28 min 



6 hr 28 min 



6 hr 28 min 



6 hr 22 min 



6 hr 13 min 



6 hr 7 min 



6 hr 11 min 



6 hr 18 min 


= 6 hr 7 min 


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 = V4 to t = Vi. 

Before we consider numerical approaches, 
we must ensure that our model is well-posed. 

Theorem 1: Let/: [t„ To) X [0,D) ^ K be 
blockwise-constant and positive-valued. Let f 
G S, t < To. Then, one (and only one) of the 
following must hold: 

i) There exists T g (to, Tj, x : [t, T] -^ R 
such that X is a solution to x'{t) = fit, 
x{t)) on [t,Tl x{t) = and x(T) - D, or 

ii) there exists x : (tTo] -^ R such that x is 
a solution to x'{t) = /(f,x(f)) on [tj.^l, 
x{t) = and xiT.^ < D. 

The proof of Theorem 1 is straightforward 
and is left to the reader. 

Theorem 2: Let/: [f„, T.) X [0,D) ^ K be 
blockwise-constant and positive-valued. Let t 
G S, t < To. Suppose X is a function satisfying 
one of the two conclusions of Theorem 1 . Let 
X be continuous, x'{t) = f{t,xit)) for all but a 
finite subset of the domain of x and assume 
there exists some t such that xit ) = x{t). Then, 
x(^) = x{t) for all t in the intersection of the 
domains of x and x. 

Proof Claim 1: x and x agree to the right 
of f, that is, i{ t G [t,f), where T represents 
the right-hand endpoint of the intersection of 
the domains of x and x, then x{t) = xit) for all 

t G [tJl 

Proof of Claim 1: Suppose not. Then, with- 
out loss of generality, there exists some t e 
iif] such that x(t) > x(t). Let t* = su^{t e 
[t, t) I xit) = xit)}. Since x and x are contin- 
uous, we have xit*) = xit*) and xit) > xit) for 
all t G it*j]. There exists some f g (f*,T) such 
that it,xit)) and it,xit)) e interior of some A, 
for all t G (f*,f). Let k, represent the constant 
value of/ on A,. Then, for t g (f*,f), we have 
x(f) = xit*) + it - t*)k^ = xit*) + it - t*)k, 
= xit), a contradiction. 

Claim 2: x and x agree to the left off, that is, 
if f G it,T], where T represents the right-hand 
endpoint of the intersection of the domains of x 
and X, then xit) = xit) for all f g [t, i]. 

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 

t G S,t < To, 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) 

\x'it) = fit, x), a < t < b, 
Ix(fl) = a, 

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- 
crete points T^, where a = Tq < t, < . . . < t^, 
= b, by approximating the solution xit) locally 
by the first two terms in its Taylor series. Eu- 
ler's method produces an approximation u^ to 
the exact solution x(tP at the mesh points ac- 
cording to the iteration 

+ /i,/(t,_,, m,._,), 

j = 1, 2, 


where hj = t^ — t^.,. Euler s method is in fact 
exact if the true solution xit) 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 {T^}^i'(|(that is, the 
only changes in slope occur on some subset of 
the set of mesh points (T^}y=o)- Iri our travel 
problem, the function /(f,x) is blockwise con- 
stant, and therefore the exact solution xit) 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. 


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 


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) g 
{—d,0,c} or, x"{f) e [—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) 
e [0,fit,x{t))] rather thanx'{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 CeUina 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/ 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 fx-plane for this / is a trapezoid 
or a triangle. 


Avibin, 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, 

Trans. Ky. Acad. Sci. 58(2):96-l()(). 1997 


Blue Catfish (Ictalurus furcatus; Ictaluridae) 
Predation on the Zebra Mussel in the Ohio River 
near Paducah, Kentucky. — Blue catfish (Ictahtnis fur- 
catus) were examined to determine whether they are 
predators of the exotic zebra mussel {Dreissena pohjmor- 
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 caq3 {Cijprinus caiyio) are predators of ze- 
bra mussels (5, 6, 7). Other fishes with molariform pha- 
ryngeal teeth or chewing pads — e.g., freshwater drum 
(Aplodinotus gmnniens), red-ear sunfish {Lepomis micro- 
lophu.s), and smallmouth buffalo {Ictiobu.s huhahis) — 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 (8). Members of the Ictal- 
uridae (catfish family) can feed on bivalves by swallowing 
them whole. French (1993) noted a zebra mussel with 
valves intact in a brown bullhead {Ameiunis nehulosus). 
On 1 Oct 1995 a commercial fisherman on the Ohio River 
told us that he observed zebra mussel shells in the stom- 
achs of blue catfish. To confirm this obser\'ation 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 Nov 1995 from 
Three Rivers Fish Market in Paducah, Kentucky. The 
sample consisted of 150 individuals with lengths ranging 
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. Three species of mollusks were present: 
Asian clam {Corbicula fluminea), fawnsfoot (Tnincilla 
donacifonnis), and zebra mussel. 

Blue catfish witli lengtlis of 28 to 33 cm were foimd to 
contain tlie majority of biviilves and consisted of 58% (87) of 
the sample. Of the 87 stomachs, only 31.0% (27) of indi\id- 
uals of tliis size had fed on bivalves. Eighteen stomachs 
(66.7%) contained only Asian clams; six (22.2%) contiuned 
zebra mussels. One stomach diat had zebra mussels iJso had 
three fawnsfoot mussels widi lengths of 7, 11, imd 12 mm. 
Three (11.1%) blue catfish widi lengths of 29, 30, and 32 cm 
had bodi 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 die total number of zebra mussels. 
In die diree species of bivtJves die liinge was intact, die shells 
were whole, and some viscertJ dssue was present. A total of 
1433 Asian clams was found widi lengdis 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 die stom- 
achs. Tliirty-seven zebra mussels were collected with lengths 
between 3 and 15 mm. Zebra mussel lengdis of 3 to 7 mm 

comprised 78.4% (29) of die total number of zebra mussels 
found in stomachs. Our obsen'ations on zebra mussels in die 
Oliio River at Dam 52 provide a length range of 3 to 24 mm 
widi die 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 and 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 
Universit)- for laboratory facilities and equipment. 

LITERATURE CITED. (1) Wonnington, A., and J. H. 
Leach. 1992. Concentrations of mignint di\ing ducks at Point 
Pelee Nadontil Paik, Onttuio, in response to inviision of zebra 
mussels, Dreissena polijmorpha. 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 
pohpmnyha. Ardea 80:35;3-.362. (3) Love, J., and J. F Savino. 
1993. Crayfish (Orconectes virilis) predation on zebra mussels 
[Dreissena pohjmorpha). J. Freshwater Ecol. 8:25.3-259. (4) 
French, J. R. P., III. 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 iiitills) on ze- 
bra mussel (Dreissena pohjmorpha): field studies. Oecologia 
83:378^84. (6) Cloe, W. W, III, G. C. Carman, tuid S. A. 
Stranko. 1993. The potential of die buU chub (Nocomis ra- 
netji) as a predator of the zebra mussel (Dreissena pohpnor- 
pha) in mid-Adantic coastal rivers. Am. Midi. Naturalist 133: 
170-176. (7) Stein, R. A., J. F Kitchell, and B. Knezevic. 
1992. Selective predation by carp (Cypiinus caiyio L.) on 
bentiiic molluscs in Skadar Lake, Yugoslavia. Fish Biol. 7(3): 
391-^99. (8) Robison, H. W, and T M. Buchanan, 1984. 
Fishes of Arkansas. Univ. Arkansas Press, FayetteviUe, AR. — 
Jeffrey J. Herod, Tricia L. Frye, ;ind James B. Sickel, 
IDepai-tment of Biologiciil Sciences, Mun-a)' State Uni\'ersity, 
Murray KY 42071. 

Rare and Extirpated Plants and Animals of Kentucky: 
1997 Update. — The Kentuch' State Nahire Preseives Com- 
mission (KSNPC) published a list of rare and e.xtirpated 
plants and aniniiils of Kentucky in 1996 (1). The list, de\el- 
oped with assistance from many scientific autiiorities, wiis 
based on distributional tmd ecological data available as of 1 
Mai- 1995. KSNPC (1) commiti:ed to update the list imnutilly 




so that decision makers would have current infonnation on 
rare species in Kentucky. Herein we update die 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 CI), 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 hsted 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 

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 indi\aduals for sharing infor- 
mation and for their assistance: W.M. Andrews, Kentucky 
Division of Environmental Services; TC. 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 WD. Pearson, 

Table 1. Conservation status changes for rare and extirpated Kentucky plants and animals. 




Aureola ria patula 
Berchemia scanclens 




Elodea nuttaUii 



Halesia tetraptera 

Lonicera dioica var. orientalis 



Trepocarpus aefhusae 
Triplasis purpurea 
Viburnum lentam 





Anguispira rugodenna 

Alasmidonta atropurpurea 

Epioblasma brevidens 

Epioblasma capsaefonnis 

Cambanis ornatus 
Orconectes inennis 

Pseudanophthalmus catonjctos 
Tseudanophthahnus pholeter 

Amnwcrypta clara 
Etheostoma cinereiim 
Etheostoma tippecanoe 
Macrhtjbopsis gelid a 
Macrhybopsis meeki 
Percina evides 

Aquila chrysaetos 













































' S = Special Conceni. T = Threatened, E = Endangered, H = Historic, X = Extirpated or Extinct. 
- PE = Proposed Endangered, E = Endangered, C; = Candidate for Listing, CI = Candidate for Listing 


Transactions of the Kentucky Academy of Science 58(2) 

Table 2. Nomenclature changes for rare and extirpated Kentucky plants and animals. 

old nan K- New name 

Carex atlantica ssp. capillacea 

Howe sedge 
Gentiana alba 

Halesia tetrapera var. tctrapfera 
Heracleuin maximum 

Pseudanophthalmus abditus 

A cave beetle 
Pseudanophthabnus Jujpolithos 

Stone-dwelling cave beetle 
Pseudanophthabnus inexpectatus 

A cave beetle 
Pseudanophthabnus scholasticus 

Schoolhouse cave beetle 

Etheostoma clanini 
Etheostoma peUuciduni 
Etheostoma vivax 
Notropis sp. 
Palezone shiner 


Elanoides foificatus foijicatus 

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 

Ammocrijpta clara 
Ammocn/pta pellucida 
Ammocn/pta vivax 
Notropis alhizonatus 

Swallow-tailed kite 
Ardea alba 

University of Louis\alle; 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 
IlUnois-Chicago; S.P. Rice, Kentucky Transportation Cab- 

Table 3. Additions to the list of rare and extirpated Ken- 
tuck\' plants and animals. 






Viburnum molle 

Missouri arrow- wood 




Helicodiscus notius specus 

A snail 


Helicodiscus p u n eta tell us 

Punctate coil 




Celitheniis verna 

Double-ringed pennant 




Etheostoma percnurum 

Dusk-ytail darter 



' S = Special Concern, T = Threatened. E = Endangered. 
- E = Endangered, 

inet; J, Slapcinsky, Field Museum of Natural History; C.A. 
Taylor, Illinois Natural History Survey; and K.J. Tennes- 

LITERATURE CITED. (1) Kentucky State Nature 
Preser\'es Commission. 1996. Rare and extirpated plants 
and animals of Kentucky. Trans. Kentucky Acad. Sci. 57: 
69-91. (2) U.S. Fish and WildUfe Serxace. 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, 
pui-ple 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, WL. 1956. A review of the nearc- 
tic Viburnum. WL. 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 moUusks 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, WL. Pratt, G.F.E. 
Roper, A. Scheltema, E.G. Thompson, and J.D. Williams. 
1988. Common and scientific names of aquatic inverte- 



brates from the United States and Canada: molhisks. Am. 
Fish. Soc. Spec. Publ. 16. (9) Barr, T.C., Jr. 1996. Cave 
beetle status sun'ey and preHsting recover)' project. Re- 
port prepared for the U.S. Fish and WikUife Semce and 
the Kentuck)' Dept Fish VVildl. 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 1994:868-886. (14) 
American Ornithologists' 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 brachijcaqmm 
C. Presl (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, Universit)- 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 
oi 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. Vs mile s of 1-275 bridge, ca. 2 miles s of Latonia, 
23 Jul 1997, Thieret and Allen 60239 (KNK, MO). 

Figure 1. Epilobium brachijcarpum. Photocopy of 
pressed specimens, X %. 

LITERATURE CITED. (1) Raven, RH. 1976. Generic 
and sectional delimitation in Onagraceae, tribe Epilo- 
bieae. Ann. Missouri Bot. Card. 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, Northern Ken- 
tucky University, Highland Heights, KY 41099. 

Homed-Pondweed, Zannichellia palttstris (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. palustiis 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 northern 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 oi Zannichellia for Calloway, Lyon, MarshiJl, 
and Trigg counties (Kentucky Lake). 

In May 1994, while looking for plants along a proposed 


Transactions of the Kentucky Academy of Science 58(2) 

Figure 1. Zaunichellia palustris. Plant {XV4); lower right, 
achenes (X6). Drawing by John Myers; used by permis- 
sion of Flora of NortJi America. 

right-of-way for the reconstruction of KY 17, near Inde- 
pendence, Kenton County, Kentucky, we found horned- 
pondweed in a pond with Eleocharis ohtusa 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 

Figure 2. ZaniclieUia palustris. Documented county dis- 
tribution in Kentucky. 

species now includes the northern and central parts of the 
state (Figure 2). Additional field work wall 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 

LITERATURE CITED. (1) Heal, E.O., and J.W. 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. (Zannichelh- 
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 Larry A. Giesmann Thomas C. Rambo 

Richard I. Bamhisel Vernon Hicks Arthur Raymond 

David M. Brandenburg Ellis L. Laudermilk James Sickel 

Jerry Carpenter James O. Luken David Sleper 

Ronald R. Cicerello Max E. Medley Jerry W. Warner 

Michael A. Flannery Robert F.C. Naczi Stephen White 

Michael A. Floyd Debra K. Pearce Steven Wilkinson 


Trans. Ky. Acad. Sci. 58(2): 10.3-1 10. 1997 


Compiled by Varley Wiedeman 

A-chain gene transcription, 42—43 
Aesculus glabra var. glabra, 15 
Abstracts, 37-54 
Accipitridae, 54 

saccharinum, 87, 88 

rubrum, 89 
Acidic mines, oil effects on nodula- 

tion, 80-84 

ligamentina, 56, 57 

spectorosa, 57 
AD-like pathology, in normal aging, 

Adiantum capillus-veneris, 74 
Aesculus glabra var. glabra, 13 
African- American 

health and diet status, 39 

rural southern elderly, 39 
Agricultural sciences, abstracts, 37- 

Agrypnia vestita, 69, 71, 72 
Alasmidonta atropurpurea, 97 
Alkylaton of DNA, 40 
Allium cepa, 60 

Alpheus heterochaehs, 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 

clara, 97, 98 

pellucida, 98 

xdva-x, 98 

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 

Amoglossum muehlenbergii, 12, 15 
Arrow-wood, Missouri, 98 
Arundinaria gigantea, 12, 15 

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, 


for non-science majors, 48 

use of computers for, 48 
Astronotus ocellatus, gentamicin-in- 

duced toxicity in, 49, 50 


laevigata, 14, 15, 18 

patula, 74, 97 

Bahalana ;eracei, 53 

effecis of cannibalism on, 53 

from jan Salvador Island, Baha- 
mas, 53 

population structure of 53 
Bald eagles 

densities, 54 

distributions, 54 

impacts of dam construction on, 
Baldcypress, 85, 88 

austrahs, 12, 15 

cerulea, 12, 15 

coerulea, 12, 15 

tinctoria, 75 
Barren oats, 15, 16, 

paniculata var. paniculata, 75 

virginica, 75 
Bastard pennyroyal, 12, 16 

cave, 98 

concealed cave, 98 

scholarly cave, 98 

schoolhouse cave, 98 

stone-dwelling cave, 98 

surprising cave, 98 
Berchemia scandens, 97 
Beta voilgaris, 65 
Bidens discoidea, 87 
Bieberbach groups, classifying free, 


Biology, student attitudes toward, 51 

Birds, 97, 98 


Bitter milk-wort, 14, 16 

Bivalva, 55-59 

in the Cumberland River, 55-59 
Black darter, 4 
Black wort, 15, 16 
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 energ)' restric- 
tion, 49 

effect of ovariectomy on, 49 
Boehmeria cylindrica, 87 
Botanical regions in Kentucky, 20- 

Constantine Rafinesque's 1819 
description of, 20-22 
Botany and microbology abstracts, 

Botany of Kentucky, 12 
Bradyrhizobium, 80-84 
Breast cancer, human, 46-47 
Briar, sand, 17 
Brown bullhead, 96 
BROWN, C, 49 
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 

Cacaha, 17 

atriplicifolia, 17 

muehlenbergii, 12, 15, 17 

reniformis, 12, 15 
Caddisflies, of the Kentucky River, 

Calcium excretion, urinary, 49 



Transactions of the Kentucky Academy of Science 58(2) 


homeostasis, 40 
neurotaxicity, 41 
metabolism, in rats, 50 
tuberosus, 75 

tuberosus var. tuberosus, 74 
Cambanis ornatus, 97 

Campostoma anomalum, 6 
Cane, 12, 16 

giant, 12, 16 
Cannibalism, 53 

effects on population structure, 
Capraria multifida, 13, 15 

atlantica ssp. eapillacea, 98 
crawei, 75 

stricta var. stricta, 75 
CarjD, common, 96 
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 

cordata, 13, 15 
cordatum, 18 
cordifolium, 13, 15, 18 
Catfish, blue, 96 

predation on zebra mussel, 96 
Catostomus commersoni, 6 

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, photometiy with, 48^9 
Cedar, red, 14, 17 
Celestial seasons, 47 
Celithemis verna, 98 
Cenchrus longispinus, 76 
Center for automated space science, 

Cephalanthus occidentahs, 87 
Cephalogonimidae, 23-28 
Cephalogonimus vesicaudus, 23-28 

ancylus, 69, 71 

cancellata, 69, 71 

fiava, 69, 71 

punctata, 69, 71 

tarsipunctata, 69, 71 

transversa, 69, 72 
Ceratopsyche sparna, 69, 70 

ambrosioidaes var. anthelminti- 
cum, 12, 15 

anthelminthicum, 12, 15 

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 

Cichlidae, 49, 50 
Clam, Asian, 96 
Clammy-weed, 13, 16 
Cliff phuiib, 16 
Clover, nimble, 15 
Common caqi, 96 
Common laurel, 14 
Common names, of vascular plants, 

Community college graduates, 50- 

Computerized education, a student 

sun'ey, 51-52 
Computers, use for astronomy, 48 
Concealed cave beetle, 98 

eastern, 14 

orange, 14, 17 

purple, 14, 17, 21 
Conobea, 17 

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 
Crested dwarf iris, 14, 16 
Crested flag, 16 
Crested iris, 14, 16 
Crested tris, 14, 16 
Crustaceans, 97 
Cr)'opreser\'ation of seeds, of Ohio 

wetland plants, 38 
CULLER, RYAN, 47, 48 

Cultiued panicle segments, 60-66 
Cumberlandia monodonta, 58 
Currant, wild, 14, 17 
Cuscuta sp., 87 

abdominalis, 35 

buccata, 35 

fontinella, 35 

sp., 35-36 
Cyclonaias tuberculata, 55, 57-58 

croceus, 76 

difformis, 76 
Cy}3rinus carpio, 96 
Cyrnellus fraternus, 69, 70 
Cytochrome P450 2E1, 45-^6 
Cytokine production, 42 
Cytokines, and neonatal immune re- 
sponses, 43^4 


Candida, 15 

pui"purea var. purpurea, 15 

black, 4 

saffron, 4 

duskytail, 98 
Ddarters, snubnose, 4—8 

separation of spawning habit, 4-8 

spawning habit in, 4-8 
Deerberi^', 14, 16, 21 
DEMOSS, D.L„ 50 
DERTING, TERRY L., 51, 54 
Dietar)' energ)' restriction, effects of 

on rats, 49 
Diptera, 35-36 

Distinguished scientist award, 33 
]3ouble-ringed pennant, 98 
Drum, freshwater, 96 
Duskytail darter, 98 
Dwarf iris, crested, 16 

Eastern coneflower, 14, 16 
Echinacea puipurea, 14, 15, 21 
Echinostoma trivolvis, 23-28 
Echinostomatidae, 23-28 
Elanoides forficatus forficatus, 98 
Electrophysiological studies, of the 
SCN8a voltage-gated sodi- 
um channel, 43 
Elephants foot, 12, 16 

leafy, 12, 16 

caber, 12, 15, 17 

carohnianus, 12, 15, 17 

Index to Volume 57 


Ellipsaria lineolata, 55, 57 

crassidens, 56, 57 

dilatata, 56, 57 
Elodea nuttallii, 97 
Emden-Fowler equation, polyno- 
mial conservation laws of, 40 
Endangered Ohio wetland plants, 
cryoperservation of seeds of, 38 

brachycarpiim in Kentucky, 99 

ciliatum var. ciliatum, 76 

paniculatum, 99 

brevidens, 58, 97 

capsaeformis, 58, 97 
EPSCoR opportunities for space-re- 
lated science, technology, 48 
Esox masquinongy, 46 

barrenense, 5 

blennioides, 6 

caeruleum, 5, 6 

cinereum, 97 

clarum, 98 

crossopterum, 6 

davisoni, 5 

duryi, 4, 5, 7 

Habellare, 6 

fla\aim, 4-8 

flavum, spawning in, 4-8 

pellucidum, 98 

percnurum, 98 

rafinesquei, 5 

simoterum, 4—8 

simotenun, spawning in, 4-8 

smithi, 6 

tippecanoe, 97 

vdvax, 98 

zonale, 5 

development and content, 51 

science, 51 

world view and values, 51 

celestinum, 15 

coelestinum, 13, 15 

rugosum, 13, 15 

urticefolium, 13, 15 
Extirpated animals of Kentucky, 96- 

Extirpated plants of Kentucky', 96- 

Extragalactic radio jets, 47 

Fabaceae, 80-84 
False pennyroyal, 14, 16 
False-foxglove, Appalachian, 14, 16 
False-indigo, blue, 16 
Fawnsfoot, 96 
Fescue, tall, 60-66 

characterization of for DNA 
changes, 60-66 

characterization of for isozyme 
changes, 60-66 

characterization of for meiotic 

changes, 60-66 

Festuca arundinacea, 60 


Field investigations, 51 

educational benefits, 51 

misconceptions, 51 

problems, 51 

student perspectives of, 51 
Fire-pink, 15, 16 
Fishes, 97, 98 
Flag, 14, 16 

crested, 16 
Forestiera acuminata, 87, 88 

pennsylvanica, 87, 88 

profunda, 87 
Free Bieberbach groups, classifying, 

Freshwater drum, 96 

orange, 14, 17 

yellow, 14, 17 

Gaultheria procumbens, 14, 21 

INI, 47 
Genes involved in splicing, 44 
Gentamicin-binding protein(s), 49- 

in ear of cichlid fish, 49-50 

stiff, 15, 17 

yellow, 15, 17 

alba, 98 

amarelloides, 15, 18 

amerelKades, 15 

flavida, 98 

quinquefolia var. occidentalis, 18 

quinquefolia van quinquefolia, 15, 
Geography abstracts, 39 
Gerardia glabrata, 14, 15 
Giant cane, 12, 16 

stipulacea, 12, 15 

stipulata, 12, 15 
Glossosomatidae, 69, 70 
Golden-rod, stiff, 14, 17 
Goldenrod, stiff, 17 

Graduates, of community colleges, 

Grass, Indian, 15, 16, 21 
Gravitational lensing simulation, 47 
Great Indian-plantain, 12, 16 
Green ash, 87 

GUO, QING, 40 

Hair)' nielfia, 15, 16 
Halesia tetrapera, 97, 98 
Haliaeetus leucocephalus, 54 

densities, 54 

distributions, 54 

impacts of dam construction on, 
HAMPTON, D., 49 

Hatching, in the muskellunge, 46 

stage dependent induced, 46 
Health and diet status, differences 

in, 39 
Health sciences abstracts, 39^0 
Heart-leaf, 14, 16 
Hedeoma pulegioides, 12, 15 

notiusspecus, 98 

punctatellus, 98 
Heliconia, 53 

latispatha X imbricata, 53 

e.xtrafloral nectaries, 53 
Heliconiaceae, 53 
Heliotrope, 13, 16 

Indian, 13, 16 
Heliotropium indicum, 13, 15, 17 
Helisoma trivolvis, 23-28 

lanatum, 98 

maximum, 98 
Hesperis pinnatifida, 13, 15 
Hexastylis, 18 

virginica, 14, 15 
Hippocampal subregions, 41 
Horned-pondweed, 99-100 
Horse nettle, 16 
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 


Transactions of the Kentucky Academy of Science 58(2) 

Human breast cancer, mRNA in, 

Human myiasis in Kentucky, 35-36 
Human pancreatic carcinoma, 44- 


betteni, 69 

frisoni, 69, 70 

hetteni, 70 

orris, 67, 69, 70 

simulans, 69, 70 
Hydropsychidae, 69, 70 

ajax, 69, 70 

angusta, 69, 70 

armata, 69, 71 

consimilis, 69, 71 

hamata, 69, 71 

perdita, 69, 71 

waubesiana, 69, 71 
Hydroptilidae, 69, 70 

caroliniana, 13, 15, 18, 20 

liriosme, 18 

occidentalis, 18 

perforatum, 99 

walteri, 87 

Ictaluridae, 96 

predation on zebra mussel, 96 

Ictalunis 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, 


of immune tissues, 44 

role of nerve growth factor, 44 
Insecta of the Kentucky River, 67- 

Insects, 97, 98 

lodanthus pinnatifidus, 13, 15 
Ipecac, American, 12, 16 

cristata, 14, 15 

crocea, 13, 15 

fulva, 13, 16 

copper, 16 

crested, 14, 16 

crested dwarf, 14, 16 
Iron weed, 16 

Ironoquia punctatissima, 69, 71-73 
Iron weed, 12 

tall, 17 

brachiatus, 14, 16 

caeruleus, 16 

ceruleus, 14, 16 

coeruleus, 14 
Isopod, marine cave, 53 

Jackson Purchase Region, of Ken- 
tucky, 85-91 

binata, 13, 16 

cinata, 13, 16 

diphylla, 16 
JONES, S.E., 40 
Juncus validus, 74, 76 
Juniperus \arginiana, 14, 16, 21 
JUST, JOHN J., 46 

Kalmia latifolia, 14, 21 
Kentucky vascular plants, 74-79 
Kentucky's Jackson Purchase Re- 
gion, 85-91 
Kidney weed, 12, 16 

American swallow-tailed, 98 
swallow-tailed, 98 
KRUPER, JILL II., 51, 53 

L-tyj3e calcium channel density, 41 

cardium, 56, 57 

fasciola, 57 

ovata, 56, 57 
Large-flowered leafcup, 14, 16 
Lasmigona costata, 57 

common, 14 

mountain, 21 

large-flowered, 14, 16 

yellow-flowered, 14, 17 
Leafy elephants-foot, 12, 16 
Lechea, 18 

minor, 14, 16 
LEE, C.J., 39, 49 

cyanellus, 6 

megalotis, 6 

microlophus, 96 
Leptoceridae, 69, 71 
Leptodea fragilis, 58 

cuneata, 80-84 

sericea, 80-84 
Leucospora multifida, 13, 16, 17 

LEWIS, RRIAN M., 40, 47 
Ligumiarecta, 57 

michiganese, 77 

superbum, 76 
Lily, 13, 16 

red, 13, 17 

swamp, 17 
Limnephilidae, 69, 72 
Linum usitatissimum, 77 
List of reviewers, 101 
Little browTi jugs, 14, 17 
Littorinidae, 52 
LIU. BIN, 42 
Lolium perenne, 65 
Lonicera dioica var orientalis, 97 
Lotus, 18 

American, 13, 16 

yellow, 13, 17 
LOU, X., 50 
Lupatorium calutinum, 13, 16 
Luxilus chrssocephalus, 6 
Lycopus rubellus, 87 
Lythrurus ardens, 6 

gelida, 97 
meeki, 97 


Maple, red, 89 


Mathematics abstracts, 40 

MATTSON, MARK R, 40, 46 


NI, 47 


MEADE, LES, 100 



Medicago sativa, 65 

Medionidus conradicus, 57 



Metal impact in the Robinson For- 
est, using metalliothionein, 40- 

Metalliothionein, assessment of 
metal impact, 40-41 


arupedinaria, 12 
arupendinaria, 16 

Micropterus pimctulatus, 6 

Microsatellite instability and TGF-b 
Type II receptor gene muta- 
tion, 44^5 

Miegia arundinaria, 12, 16 

Migraine headaches, effects of 
weather systems on, 39 


Index to Volume 57 


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, 

Morehead radio telescope, 47-48 
optimization of operating param- 
eters, 40 
surface geometn' of, 40 
Mountain laurel, 21 
Mountain tea, 14, 21 
MPTP-treated C57B1/6 mice, 45 
mRNA in human breast cancer, 
translational regulation of, 46- 
Muhlenbergia cuspidata, 77 
Muskrats, 56 
Mussel, zebra, 96 

in the Ohio River, 96 
Mussels, 97 

of a conser\'ed phenylalanine, 45- 

site directed, 45—46 
Mutations, in the RNA sorting sys- 
tem, 45 
Mycena leaiana, fine structure of 

bacidiospores, 38 
Mystacides sepulchralis, 69, 71, 72 


Nanostoma, 4 

Narrow-leaved houstonia, 13, 17 
NASA Kentucky space grant consor- 
tium, 48 

Candida, 69, 72 

exquisita, 69, 72 

pavida, 69, 72 
Nelumbium pentapetalum, 13, 16 
Nelumbo, 18 

lutea, 13, 16 
Neotyphodium coenophialum, 60 
Nephrotoxicity, gentamicin-induced, 


horse, 16 

white, 13, 17 
Neureclipsis crepuscularis, 69, 71, 

Nicotinein MPTP-treated C57B1/6 

mice, 45 
Nicotine, neuroprotective effects of, 

Nimble clover, 15 
Nimble weed, 14 

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 
Notes, 35-36, 96-100 

albizonatus, 98 

sp., 98 
Nucleotide excision repair mutants, 

Nymphaea sp., 77 
Nyssa aquatica, 85, 87, 88, 90 


avara, 72 

cinerascens, 72 

inconspicua, 72 

persimilis, 72 
Oats, barren, 16 
Obliquaria reflexa, 55, 57 

aergerfasciella, 69, 71 

cristata, 69, 71 

tarsalis, 69, 71 

xena, 69, 71 

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 
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 urinaiy 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 

muscle lipid content, 50 

stability of meat stored in crushed 
ice, 50 

using flood control resen'oirs for, 
Palezone shiner, 98 

liriosme, 13, 16, 18 

liviosone, 13, 16 
Pancreatic carcinoma, 44-45 
Panicle segments, cultured, 60—66 
Pa\da muricata, 13, 16 
Paw]^5aw, 13, 17 

allozyme variation in, 37 

geographic differentiation in, 37 

influence of shade on, 37-38 

morphological development, 37- 

root-zone modification, effect on, 

seed germination, 37-38 

seedhng emergence, 37-38 
Pawpaw tree, 17 
Pea-clover, yellow, 14, 17 
Pecan, 89 

Pencil flower, 14, 17 
Pennant, double-ringed, 98 
Penny royal, 12 
Penny-aoyal, 12, 17 
Penny-royal, 17 

blue, 14, 16 

American, 12, 16 

bastard, 12, 16 

false, 14, 16 
Percidae, 4-8 

caprodes, 6 

evides, 97 
Peromyscus leucopus, 35 

candidum, 16, 21 

moncandidum, 15 

purpureum, 15, 16, 21 
Petalvitemon candidum, 15, 16 
Phenylalaninein cytochrome P450 

2E1, 45-46 
Philopotamidae, 69, 72 
Phlox glaberrima, 14, 16 
Phlox, smooth, 14, 17 
Photometry with CCDs, 48^9 

optimizing signal extraction, 48- 
Photomorphogenic control, in dan- 
dehon, 38 

of flower scape elongation, 38 
Phyrganeidae, 69, 72 
Physic, Indian, 12, 16 


Transactions of the Kentucky Academy of Science 58(2) 

Physics and astronomy abstracts, 

Physiology and biochemistry ab- 
stracts, 49-52 
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, 88 
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 occidentals, 13, 16, 20, 88 
Platelet-derived growth factor, 42- 

Plum, wild, 13, 17 
Polanina graveolens, 13, 16 

dodecandra ssp. dodecandra, 13, 

graveolens, 13, 16 
Polycentropodidae, 69, 72 

cinereus, 69, 72 

confusus, 69, 70, 72 

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- 


angulata, 13, 16 

deltoides, 13, 20, 88 

deltoides van deltoides, 16 

tribuba, 13, 16 

triloba, 13, 16 
Porteranthus stipulatus, 12, 16 

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 

purple, 15, 17 

white, 15, 17 
Prairie-clovers, 21 
Prairie-dock, 15, 17, 21 
Prickly bog sedge, 98 
Prickly buck-eye, 13, 17 
Proterometra macrostoma, 27 
Protoptila maculata, 69-71 

mimsoniana, 13, 16, 18 

pendula, 13, 16, 18 

abditus, 98 

catoryctos, 97 

h)polithos, 98 

inexpectatus, 98 

pholeter, 97 

scholasticus, 98 

ocellifera, 69-72 

semifaciata, 69-72 

fasciolaris, 57 

subtentum, 55-58 
Punctate coil, 98 
Purple coneflower, 14, 17, 21 
Purple prairie-clover, 15, 17 
Puqjie rocket, 13, 17 
Puiple sun-flower, 14, 17 
Pycnopsyche lepida, 69, 72, 73 
Pyganodon grandis, 58 
Pyrimidine biosynthesis enzymes 

localization of, 44 

with fusion proteins, 44 

Quadnila pustulosa, 56, 57 

optical monitoring of, 48 

very high red shift, 48 
Quercus lyrata, 87, 88 

Raccoons, 56 

Racemed milkwort, 14, 17 

Radio frequency astrophysics, 47^8 

Radio jets, extragalactic, 47 

Radio telescope, 47-^8 

Ralinesque's 1819 description of bo- 
tanical regions, 20-22 

Ragweed, sand, 13, 17 



Rare animals of Kentucky, 96-99 

Rare plants of Kentucky, 96-99 


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 hly, 13, 17 
Red maple, 89 
Red-eared sunfish, 96 
Redcedar, 21 
Retinioc acid 

effect on stromelysin, 42 

effect on TIMR 42 
Rhyacophila lobifera, 69, 71-73 
Rhyacophilidae, 69, 72 
Rhynchospora macrostachya var. 

macrostachya, 74, 77 
RNA sorting system, in spermiogen- 

esis, 45 
Roach, 96 


Rock weed, 13, 17 
Rough bell, 15, 17 
Rough wort, 14, 17 
Rubiaceae, 18 

fulgida, 14, 16 

fulgida var. fulgida, 14 

purpurea, 14, 16 

caroHniensis ssp. ciliosa, 15, 16 

oblongifolia, 15, 16 
Ruellia, hairy, 15, 16 
Rutilis rutilis, 96 

Saffron darter, 4 

Salix nigra, 87, 88 

San Salvador Island, Bahamas, Tec- 

tarius muricatus on, 52 
Sand briar, 13, 17 
Sand ragweed, 13, 17 
Saururus cemuus, 87 
Scarlet pink, 15, 17 
Scented sun flower, 14, 17 

japonicum, 27 

mansoni, 27 

mucronatus, 74, 77 

purshianus, 78 
Scholarly cave beetle, 98 
Schoolhouse cave beetle, 98 
Science ethics 

development and content, 51 

for biology and chemistry majors, 

world \iew and values, 51 

Index to Volume 57 



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-i9 

Howe, 98 

prickly bog, 98 
Seneca snakeroot, taxonomic status 

of 39 
Sensing system, for antimonite, 42 
Sericea lespedeza, 80-84 
SHIBER, J.G., 50, 51 
Shiner, palezone, 98 
Shrimp alpheid, snapping action of, 


catesbaei, 16 

catesbei, 15, 16 

catesbri, 15, 16 

virginica, 15, 16 

terebinthaceum, 16 

terebinthinaceum, 16, 21 

therebinthaceum, 15 
Silver maple, 87 

Skeletal compartmentalization, in 

rats, 50 
Sky weed, 13 
Sky-weed, 17 
Small pinweed, 14, 17 
Smallmouth buffalo, 96 

Smooth phlox, 14, 17 
Snailnatural infection of trematodes, 

Snails, 97, 98 
Snakeroot, white, 13, 17 
Sodium channel, SCN8a voltage- 
gated, 43 

carolinianum, 13, 16 

caroliniense, 13, 16 

phureja, 65 
Sohdago rigida, 14, 16 
Sorghastnmi nutans, 15, 16, 21 
Sorghum bicolor, 65 
Southern rural elderly dietary 
knowledge, 39 

health status, 39 

intakes of, 39 

Spearmiogenesis, RNA sorting sys- 
tem in, 45 
Spider-lily 13, 17 
Spiderlily 20 

Splicing, genes involved in, 44 

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 
Student attitudes, toward study of 

biology, 51 

biflora, 14, 16 

elatior, 14, 16 
StyKdanthes 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 allmdal plain, 8.5-91 

stnicture of 85-91 
Swamp lily, 13, 17 
Sy-weed, 13, 17 
Sycamore, 13, 17 
Sylvilagus floridanus, 1-3 

cover, 1-3 

habitat, 1-3 

in Kentucky, 1-3 

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 

Tectarius muricatus 

on San Salvador Island, Bahamas, 

size and density of, 52 
Teleostei, 4-8 

TGF-b T)'pe II receptor gene mu- 
tations, 44—15 
TNF-alpha neuroprotectionin opi- 
ate-mediated astroglial toxicity, 
Traditional education, a student sur- 
vey 51-52 

daily emergence, 23-28 

effect of light on, 23-28 
Trepocarpus aethusae, 97 

ignitus, 69, 72 

tardus, 69, 72 
Tricholomataceae, 38 
Triehoptera, of the Kentucky River, 

Trip planning, 92-95 
Triplasis purpurea, 97 
Tris, crested, 14, 16 
Triticum aestivum, 65 
Tritogonia verrucosa, 56, 57 

donaciformis, 96 

trimcata, 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 imbeciOis, 58 


album, 14, 16 

stamineum, 14, 16, 21 
Vascular plant discoveries, from 

Kentucky, 74-79 
Vascular plants 

common names of, 9-19 

in Kentucky, 9-19 

reported by C.S. Rafinesque, 9- 

gigantea ssp. gigantea, 12, 16 

praealta, 16 


Transactions of the Kentucky Academy of Science 58(2) 

prealta, 12, 16 

lentago, 97 

moUe, 98 

iris, 57 

lienosa, 57 

trabalis, 56-58 
Vole, prairie, 53-54 

female preferences, 53-54 

male odors in, 53-54 

Wall Hower, Ohio, 13, 17 
WANG, C, 38, 49, 50 
Water tupelo, 85, 88 

WATSON, CASEY, 47, 48 
Weather systems, effects on mi- 
graine headaches, 39 
White nettle, 13, 17 
White prairie-clover, 15, 17 
White snakeroot, 13, 17 
WHITE, T.L.. 50 
Wild currant, 14, 17 
Wild indigo, blue, 16 
Wild plum, 13, 17 
Worm weed, 12, 17 
Wormseed, 12, 17 
WR-082-01 cell line, 42-43 
WRICHT, C.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 

ZannicheUia palustris, 99-100 
Zannichelliaceae, 99-100 
Zebra mussel, 96 

in the Ohio River, 96 
ZHANG, Y, 49 
Zoology and entomology abstracts, 


OCT 99 

n„„„j .T„.Pi„c^ N. MANCHESTER.