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nvans ~~ JOURNAL 
OF THE 
KENTUCKY 
ACADEMY OF 
SCIENCE 


Official Publication of the Academy 


SMITASOT “ee 
Volume 62 


Number 1 
Spring 2001 


The Kentucky Academy of Science 
Founded 8 May 1914 


GOVERNING BOARD 
EXECUTIVE COMMITTEE 
2001 


President: Ron Rosen, Department of Biology, Berea College, Berea, KY 40404 
President Elect: Jerry W. Warner, Department of Biological Sciences, Northern Kentucky University, 
Highland Heights, KY 41099 

Vice President: Robert Barney, Community Research Service, Kentucky State University, Frankfort, KY 
40601 

Past President: Blaine R. Ferrell, Department of Biology, Western Kentucky University, Bowling Green, 
KY42101 

Secretary: Stephanie Dew, Department of Biology, Centre College, Danville, KY 40422 


Treasurer: Kenneth Crawford, Department of Biology, Western Kentucky University, Bowling Green, 
KY 42103 
Executive Secretary (ex officio): Donald Frazier, Science Outreach Center, University of Kentucky, Lex- 
ington, KY 40536-0078 
Editor, JOURNAL (ex officio): John W. Thieret, Department of Biological Sciences, Northern Kentucky 
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Editor, NEWSLETTER (ex officio): Brent Summers, Department of Biology, Campbellsville University, 
Campbellsville, KY 42718-2799 
Chair, Junior Academy of Science (ex officio): Elizabeth K. Sutton, Department of Chemistry, Campbellsville 
University, Campbellsville, KY 42718 
Program Director (ex officio): Robert O. Creek, Department of Biological Sciences, Eastern Kentucky 
University, Richmond, KY 40475 . 
Editor, JKAS Webpage (ex officio): Claire Rinehart, Department of Biology, Western Kentucky University, 
Bowling Green, KY 42103 


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, 
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40292 

Abstract Editor: Robert Barney, Community Research Service, Kentucky State University, Frankfort, 
KY 40601 


Editorial Board: John P. Harley, Department of Biological Sciences, Eastern Kentucky University, 

Richmond, KY 40475 

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

J.G. Rodriguez, Department of Entomology, University of Kentucky, Lexington, KY 
40546-0091 

John D. Sedlacek, Community Research Service, Kentucky State University, 
Frankfort, KY 40601 

Gordon K. Weddle, Department of Biology, Campbellsville University, Campbellsville, 
KY 42718-2799 


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JOURNAL OF THE KENTUCKY ACADEMY OF SCIENCE 
ISSN 1098-7096 


Continuation of 
Transactions of the Kentucky Academy of Science 


Volume 62 


spring 2001 


Number 1 


J. Ky. Acad. Sci. 62(1):1-17. 2001. 


The genus Trifolium (Fabaceae) in Kentucky 


Michael A. Vincent 
W. S. Turrell Herbarium, Department of Botany, Miami University, Oxford, Ohio 45056 


ABSTRACT 


The scope and range of the genus Trifolium (Fabaceae) were examined for Kentucky. Review of literature 
and 910 herbarium specimens from 35 herbaria revealed 11 species of clover as part of the state’s flora: T. 
arvense, T. aureum, T. campestre, T. dubium, T. hybridum, T. incarnatum, T. pratense, T. reflexum, T. repens, 
T. resupinatum, and T. stoloniferum. Four species are rejected as a part of the flora: T. alexandrinum, T. 
ambiguum, T. hirtum, and T. medium. Descriptions, illustrations, and distribution maps are provided for 


each species accepted. 


INTRODUCTION 


The genus Trifolium (true clovers) is a near- 
ly cosmopolitan member of the papilionoid 
Fabaceae (Leguminosae) that contains 240 to 
250 species (Zohary and Heller 1984), though 
this number, rising as new species are de- 
scribed, may be closer to 300 (Gillett and 
Cochrane 1973). In the Old World, Trifolium 
is native to the Mediterranean region in south- 
ern Europe, Asia Minor, the Midele East, and 
northern Africa, extending into northern Eu- 
rope and east to northwestern China; in Afri- 
ca, the genus occurs through eastern regions 
to South Africa. In the New World, there is a 
wide diversity of species in western North 
America, with fewer species native to eastern 
portions, Central America, and South Ameri- 
ca. According to Zohary and Heller (1984), the 
genus may be subdivided into eight sections, 
six of which (Paramesus, Mistyllus, Vesicaria, 
Chronosemium, Trifolium, and Trichocephal- 
um) are native entirely to the eastern hemi- 
sphere; Lotoidea is native to both the eastern 
and western hemispheres; Involucrarium is 
endemic to the western hemisphere. The 
word “clover” is probably derived from the 
Dutch “klafer” or the Anglo-Saxon “cloefer,” 


meaning “club,” a reference to the three-part- 
ed leaf that supposedly resemble the three- 
lobed club of Hercules (Evans 1957; Haragan 
1991). 

North America is home to ca. 95 species of 
Trifolium, 65 native, 30 introduced. These 
numbers are higher than those of Isely (1998), 
since he was not aware of some of the less 
frequently encountered introduced species. 
Kartesz (1999) lists 96 species for North 
America. Of the native North American spe- 
cies, 43 belong in section Lotoidea; of these, 
six are found east of the Mississippi River; the 
remaining 37, mainly in the Rocky Mountains 
and along the Pacific coast. The native species 
in section Involucrarium are all found in west- 
ern North America. The introduced species, 
representing all sections of the genus except 
Paramesus, have been imported mainly for ag- 
ricultural purposes, though some introduc- 
tions appear to have been inadvertent (Isely 
1998). 

Clover species have been widely cultivated 
as forage crops for hundreds, if not thousands 
of years, and are of great economic impor- 
tance (Duke 1981). In North America, most 
of the species cultivated are perennials, among 
the most commonly grown of which are T. 


2 Journal of the Kentucky Academy of Science 62(1) 


praten ed clover), T. repens (white, Dutch, 
or Ll. » clover), and T. hybridum (Alsike 
clov: \nnual species sometimes cultivated 
inclide LT. inearnatum (crimson clover), T. hir- 

rose clover), and T. alexandrinum (ber- 


cin clover). Clovers are used as forage crops, 
as sources of nectar for honey production 
Pellett n.d.), for erosion control, and as nitro- 
gen sources for crop fields and pastures. Men- 
ke and Hillenmeyer (1886) considered “clo- 
ver” (species not specified) to be the most im- 
port mt crop then grown in Kentucky. 

There is a considerable body of folklore as- 
sociated with the clovers (Evans 1957). Four- 
leaf clovers have long ee considered a 
source of good luck and with having the ability 
to protect against witchcraft (RDA 1986). 
Four-leaf clovers are actually leaves with mu- 
tations resulting in the prolife ration of leaflets, 
with leaflet numbers ranging anywhere from 
the more commonly seen 4 to 24 or more 
(Ford and Claydon 1996; Jaranowski and Bro- 
da 1978) or rarely a single leaflet (Atwood 
1938). The multifoliolate condition is relativ ely 
common throughout Trifolium. Also of inter- 
est to many are the leaf marking patterns on 
the leaves of many Clover species. These can 
range from the more commonly encountered 

V-shape -d pattern, or chevron, to various types 
of dark to light, colored or white patterns 
(Corkill 1971; Ganders et al. 1980). These 
marks, although attractive, are very variable 
within a species; they can be inherited to dif- 
fering degrees, may sometimes vary according 
to growing conditions, and provide no infor- 
mation of taxonomic value. 

During the last 200 years, beginning with 
M’Murtrie’s Florula Louisvillensis (1819), 
many reports have been published on the flora 
of Kentucky. In those reports, various ac- 
counts of the clovers have appeared. The ear- 
liest report of Trifolium in the state was of T. 
arvense, T. pratense, and T. repens (M°’Murtrie 
1819). In the 20th century, McFarland (1942) 
reported 9 species, and Braun (1943) reported 
5. Wharton and Barbour (1971) mentioned 
only 1, Meijer (1992) listed 9, and Browne and 
Athey ( 1992) and Medley (1993) accepted 11. 
Most rece tly, Kartesz (1999) recognized 10 
species as part of the Kentucky flora. 

The purpose of this paper is to determine 
which species of Trifolium are documented for 


Kentucky and to clarify the known distribution 
for each species in the state. 


MATERIALS AND METHODS 


| examined 910 herbarium specimens from 
the following herbaria (acronyms from Holm- 
1990): APSC, BEREA, BH, BRIT, 
CM, DAO, DHL; EKY,-F-GA 
GH, KNK, KY, LLO, LSU, MDKY, MICH: 
MO, MU, NCU, NY, OKL, OSH, PH, SIU, 
US, VDB, WIS, WKU, and WVA. In addition, 
specimens were studied from the herbaria of 
Cumberland College in Wilmington, Kentucky 
(cumb), the Kentucky Agricultural Experi- 
ment Station in Lexington (kes), and the clo- 
ver herbarium of the Department of Agricul- 
ture, University of Kentucky, Lexington (uk). 
Distributional data were gathered from her- 
barium records only; no undocumented re- 
ports are included in distribution maps. Un- 
fortunately, specimens in the Athey Herbari- 
um (MEM) were unavailable. Distribution re- 
cords from books, papers, and theses were not 
included, and data in other published sources 
may not coincide with those presented here 
(e.g., Browne and Athey 1992). 


gren et al. 


CAN, CINC, 


RESULTS 


Eleven species of Trifolium are documented 
by herbarium specimens for Kentucky. Of 
these, nine are introduced and two are native 
to the state. Three clover species reported for 
Kentucky in literature could not be docu- 
mented by specimens and are excluded from 
the flora. For 13 of the 120 Kentucky counties 
(Adair, Boyd, Cumberland, Hancock, Hender- 
son, Ow sley, Robertson, Scott, Simpson, Tay- 
lor, Union, W. ayne, and Webster), I saw no clo- 
ver specimens at all. 


TAXONOMIC TREATMENT 
Trifolium L., Sp. Pl. 764. 1753. 


Annual, biennial, or perennial, glabrous to 
pubescent herbs with a taproot or fibrous 
roots. Stems simple to much-branched from 
the base and above. Leaves alternate, pal- 
mately trifoliolate to 5-7 foliolate; leaflets 
toothed; stipules adnate to the petiole. Inflo- 
rescences umbelliform, racemose, or capitate, 
axillary or terminal, long-peduncled to sessile, 
leafy or not; involucre absent or of small to 
large free to fused bracts; flowers pedicellate 


The genus Trifolium (Fabaceae) in Kentucky—Vincent 


to nearly sessile, with or without bracts; calyx 
tubular to campanulate, 5-lobed, lobes nearly 
equal in length or the lower one longer, each 
lobe entire to toothed; petals free to basally 
fused, white, pink, red, purple, or yellow, per- 
sisting in fruit, the petals clawed and often 
fused with the staminal column, the upper 
(banner) broad, oblong to obovate, the lateral 
pair (wings) narrow, usually longer than the 
lower pair (keel), which are fused into a boat- 
shaped structure; stamens diadelphous, fila- 
ments dilated below the uniform anthers: ova- 
ry sessile or stalked, style curved upward, stig- 
ma capitate to curved, ovules 1-12; fruit a 
straight legume enclosed by the persistent ca- 
lyx and corolla, dehiscent or indehiscent, seeds 
1—3(9), globular to reniform. 


KEY TO SPECIES OF TRIFOLIUM IN 
KENTUCKY 


1. Plants perennial, stoloniferous, rooting at the 
nodes; inflorescences held on axillary, upright 
stems, with or without a pair of bract-like leaves; 
petals white to pinkish; flowers reflexing with age. 

2. Peduncles 1-2, arising from upright axillary 
stems with a pair of bract-like leaves; calyx 4-7 
mm long, teeth subulate, twice as long as tube 
a as rere cic Mite ah 11. T. stoloniferum 

2. Peduncles 1, arising from leaf axil on stolon, 
without bract-like leaves: calyx 3-5 mm long, 
teeth triangular-lanceolate, lower about same 
lemethtasyitibers warksn. Wh) 2) aplalt. ase 9. T. repens 

1. Plants erect or decumbent, not rooting at the 
nodes. 

3. Petals yellow; banner petal obovate, straight or 
downcurved; fruit with an obvious stalk inside 
calyx; petioles mostly shorter than leaflets. 

4. Terminal leaflet sessile or nearly so; stipules 
nearly as long as to longer than the petiole; 
fruits 2 times the length of the style; seed 
OVOIG! sists bh eR Sr eee eee 

2. T. aureum 

4. Terminal leaflet stalked; stipules about half 
as long as the petiole; fruits 3-6 times the 
length of the style; seeds ellipsoid. 

5. Inflorescences 5-7 mm wide: flowers 2.5— 

3.5 mm long; terminal leaflet stalk about 

RminnMlon cme este yen? 4. T. dubium 

5. Inflorescences 7-12 mm wide: flowers 
3.5-7 mm long; terminal leaflet stalk 1-3 
mm long 


MS RHA eae 3. T. campestre 
3. Petals white, pink, red, or purple; banner petal 
oblong, upcurved; fruit not stalked or minutely 
so; petioles mostly longer than leaflets. 


(ow) 


6. Flowers sessile or nearly so, erect to 
spreading in fruit, in dense globose to 
elongate heads. 

7. Petals lavender to white: flowers re- 
supinate; calyx with a more densely 
pubescent region dorsally, becoming 
inflated in fruit, with obvious reticulat- 
ing veins 

7. Petals pink, red, or white; flowers not 
resupinate; calyx glabrous or with 
evenly distributed pubescence, not be- 


10. T. resupinatum 


coming inflated in fruit, veins not ob- 

viously reticulating. 

8. Flowers 10-20 mm long; corolla 
much longer than the calyx; leaflets 
broadly ovate to obovate; stipules 
broad, ovate. 

9. Perennial; heads sessile or nearly 
so, globose to ovoid; stipules 
abruptly narrowed into an awn- 
hikemtipg 5p teen eee eosin. | 


S Seaets, 2 - Meter to gee 7. T. pratense 
9. Annual; heads stalked, elongate- 
ovoid to cylindrical; stipules 
broadly rounded at the tip 
4 Bee Ebates 6. T. incarnatum 
8. Flowers 5-7 mm long; corolla 
scarcely longer than to shorter than 
the calyx; leaflets narrowly oblong 
to linear-lanceolate; stipules nar- 
rowly ovate to oblong .. 1. T. arvense 
6. Flowers with pedicels, sharply reflexed in 
fruit, in umbels. 

10. Annual or biennial: flowers 8-14 mm 
long; inflorescences 2-4 cm wide: 
pedicels 4-12 mm long; calyx 6-9 
mm long, teeth 2-3 times the length 
of the tube; leaflets 14.5 x 0.5-2 
cm; stipules broadly ovate, leaflike 
SERRA CAR ang: 8. T. reflexum 

10. Perennial; flowers 7-10 mm long, in- 
florescences 1-2.5 cm wide; pedicels 
1-5 mm long; calyx 34 mm long, 
teeth 1-2 times the length of the 
tube; leaflets 1-3.5 X 1-2 cm; stip- 
ules narrowly obovate to lanceolate, 
not leaflike ......... 5. T hybridum 


1. Trifolium arvense L., Sp. Pl. 2: 769. 1753. 
Rabbit-foot clover. (Figure 1) 


Annual, upright, 5-40 cm tall. Stems often 
much-branched, with short appressed to 
spreading hairs. Leaves petiolate below to 
nearly sessile above, longest petioles to 15 
mm, shorter than the leaflets. Stipules ovate 
to oblong, tips long attenuate, longer than the 


4 Journal of the Kentucky Academy of Science 62(1) 


Trifolium arvense L. 


Figure 1. Documented county-dis- 


tribution in Kentucky; plant (from Besette and Chapman 
1992). Bar = 10 mm. 


associated petioles. Leaflets 5-20 x 2-4 mm, 
sessile or nearly so, linear to narrowly lance- 
olate, base cuneate, apex acute to mucronate 
and slightly serrate. Inflorescence 8-30 X 8— 
10 mm, racemose, densely ovoid or cylindrical, 
on peduncles 5—30 mm, or nearly sessile: flow- 
ers 10-150, sessile. Calyx long- hairy, often sil- 
very to pinkish or purplish, none 1.5-2 mm, 
teeth subulate, nearly equal, 2.5-5 mm, plu- 
mose. Corolla white to pinkish, slightly shorter 
than the calyx lobes, 4 mm, the ste indard nar- 
row, oblong, obtuse. Fruit ovoid, 1.3°> mm. 
Seed 1, pi ale yellow, 0.9-1.3 mm. 2n = 14, 16, 
28. Flowering in Kentucky in June—July, fruit- 
ing August-September. 


Native to Europe, northern Africa, and 
western Asia, rabbit-foot clover is naturalized 
in many areas of the world and throughout 
much of North America. Zohary and Heller 
(1984) recognized two varieties, which differ 
somewhat in habit and pubescence density. 
This species was first reported for Kentuc 
by M’Murtrie (1819). The earliest Kentucky 
eae I saw was from 1835, Fayette Coun- 

(Short s.n., CINC, KY). 

” Trifolium arvense is sometimes cultivated as 
a winter annual (Henson and Hollowell 1960). 
It is adapted to infertile, dry, often sandy soil 
such as that found on roadsides, where is 
makes an attractive, silvery-pink display when 
in flower, and rose to buff when in fruit. 

This species is sometimes also called hare’s 
foot (M’Murtrie 1819), stone clover, old-field 
clover, and pussies (Delorit and Gunn 1986; 
Small 1933). 


2. Trifolium aureum Pollich, Hist. Pl. Palat. 
2: 344. 1777. Hop clover. (Figure 2) 

(Tf. agrarium L., a confused name [Dandy 
1958]) 

Annual or biennial, upright, 20-60 cm tall. 
Stems often much-branched, with short ap- 
pressed hairs. Leaves petiolate below to short- 
petiolate above, longest petioles to 12 mm, 
shorter than the leaflets. Stipules oblong-lan- 
ceolate, tips narrowly long-triangular, as long 
as or longer than the associated petioles, ad- 
nate to the petiole for half their length or 
more. Leaflets 15-25 < 6-8 mm, sessile or 
essentially so, oblanceolate to one or ellip- 
tic, base cuneate, apex obtuse to emarginate 
and mucronate, serrate in the upper half. In- 
florescence 10-25 X 12-14 mm, racemose, 
densely ovoid or cylindrical, with a flat top in 
age, on peduncles 10-50 mm; flowers 10— 
40(80). short-pedicellate. Calyx glabrous, tube 
1 mm, teeth narrowly triangular to subulate, 
lower teeth 2-3 times the length of the upper, 
1.2-1.5 mm. Corolla bright yellow, turning 
brown with age, 5-S mm, the aaaneed broadly 
obovate, obtuse-emar ginate, strongly parallel- 
veined, especially in age. Fruit oblong, 3-3.5 
mm, stalked. Seed 1, pale yellow green to yel- 
low brown, 1—-1.2 mm. 2n = 14, 16. Flowering 
in Kentucky in June-July, fruiting August— 
September. 

Native to Europe, hop clover is introduced 
in eastern and northern North America; 


The genus Trifolium (Fabaceae) in Kentucky—Vincent 5 


George Washington is known to have ordered 
seed of this species from Europe in 1786 (Pie- 
ters 1920). Zohary and Heller (1984) recog- 
nized two subspecies, which differ mainly in 
the leaf apices and style position. The earliest 
Kentucky collection I saw was from 1903, 
Boone County (Davis s.n., kes). 

Trifolium aureum has also been called large 
hop clover, yellow clover, and palmate hop- 
clover (Gillett and Cochrane 1973; Gleason 
and Cronquist 1991; Knight 1985b). It is 
sometimes cultivated (Knight 1985b). 


3. Trifolium campestre Schreb. in Sturm, 
Deutsch. Fl. Abt. 1, Band 4, Heft 16, t. 253. 
1804. Low hop clover. (Figure 3) 


(T. procumbens L., a confused name [Dan- 
dy 1958]) 

Annual, upright to ascending (rarely pros- 
trate), 5-40 cm tall. Stems often much- 
branched, with short appressed hairs to nearly 
glabrous. Leaves pinnate, long-petiolate below 
to short-petiolate above, longest petioles to 1.5 
times as long as the leaflets. Stipules ovate, 
tips acute to somewhat attenuate, shorter than 
the associated petioles. Leaflets 4-16 x 4-8 
mm, oblong-obovate, base cuneate, apex trun- 
cate to emarginate, slightly serrate in the up- 
per half, the terminal leaflet on a 1-3 mm long 
stalk, lateral leaflets nearly sessile. Inflores- 
cence 7-15 X 7-10 mm, racemose, densely 
globose to ovoid or cylindrical, on peduncles 
as long as or shorter than subtending leaves; 
flowers (10)20—40(50), short-pedicellate. Calyx 
glabrous to slightly pubescent, tube 0.5-1 mm, 
teeth narrowly triangular to subtlate, lower 
teeth 2-3 times the length of the upper, 0.6— 
1.3 mm, each tooth often tipped with 1-2 stiff 
hairs. Corolla pale to bright yellow, 3.5-6 mm, 
the standard obovate, with a slightly toothed 
margin, emarginate, more or less enveloping 
the wing and keel petals, strongly parallel- 
veined, especially in age. Fruit oblong, stalked, 
2-2.5 mm. Seed 1, shiny yellow, 1-1.5 mm. 2n 
= 14. Flowering in Kentucky in April-June, 
fruiting June—August. 

Native to Europe and widely introduced 
elsewhere, Trifoliwm campestre is widely dis- 
tributed in North America, often being found 
along roadsides, in lawns, and in other dis- 
turbed places. Haragan (1991) considers this 
species a weed in Kentucky. The earliest Ken- 
tucky collection I saw was from 1882, Jessa- 


Trifolium aureum Pollich. Documented coun- 


Figure 2. 
ty-distribution in Kentucky; plant (from Cost 1901 [right 
figure] and Hegi 1923 [left figures]). Bar = 20 mm (whole 
plant), 10 mm (branch), 2 mm (flower). 


mine County (Peter s.n., KY). This species is 
also called hop clover, pinnate hop clover, and 
small hop clover (Gillett 1985; Gleason and 
Cronquist 1991; Knight 1985b). 

Low hop clover and especially least hop clo- 
ver (the next species) are often confused with 
Medicago lupulina L. (black medic), a com- 
monly encountered annual or biennial, pros- 
trate to ascending species. It differs from 
these clovers by its usually obviously toothed 
stipules, deciduous corolla, and reniform, 
shiny black fruits. 

A. Trifolium dubium Sibth., Fl. Oxon. 231. 
1794. Least hop clover. (Figure 4) 


Annual, upright, 5-40 cm tall. Stems simple 
or branched, glabrous to slightly hairy. Leaves 


6 Journal of the Kentucky Academy of Science 62(1) 


Documented 


Schreb. 
county-distribution in Kentucky; plant (from Cost 1901 
and Hegi 1923 [lower right and left 
figures}). Bar = 15 mm (whole plant), 


2 mm 


Figure 3. Trifolium campestre 


[upper right figure 
10 mm (branch), 


flower). 


pinnate, petiolate below to nearly sessile 
above, longest petioles to 15 mm, mostly 
shorter than the leaflets. Stipules ovate, tips 
acute, slightly adnate to and shorter than the 
associated petioles, 3-5 mm. Leaflets 5-15 
4-7 mm, terminal stalked, lateral nearly ses- 
sile, obovate, base cuneate, apex rounded to 
slightly emarginate and slightly serrate. Inflo- 
rescence 5-10 X 6—S mm, racemose, loosely 
ovoid to obovoid, on peduncles much longer 
than associated leaves; flowers 3-20, pedicels 
short, reflexing dramatically with age. Calyx 
rane. 0.5-0.8 mm, teeth subulate, 
lower about twice as long as upper. Corolla 
pale yellow, 3-4 mm, the standard narrow, ob- 
long, obtuse. Fruit ovoid, nearly sessile, 1.5-2 
mm. Seed 1 (rarely 2), shiny tan to dark 


glabrous, 


Figure 4. Trifolium dubium Sibth. Documented county- 
distribution in Kentucky; plant (from Cost 1901 [upper 
figures] and Hegi 1923 figure]). Bar = 20 mm 
(whole plant), LO mm (branch), 4 mm (flower). 


[lower 


brown, 1-1.5 mm. 2n = 16; 28. Flowering in 
Kentucky in May—June, fruiting July-August. 
Least hop clover is native to . Europe and is 
now introduced throughout the world. It is 
widely distributed in North America. It is 
sometimes cultivated as a pasture plant (De- 
lorit and Gunn 1986). Trifolium dubium was 
reported as new to Kentucky by McFarland 
(1942). Mohlenbrock et al. (1966) again re- 
ine the species, but I could not locate the 
specimen cited in that paper. The species was 
also reported for Henry County in an unpub- 
lished thesis (Gentry 1963), but again the 
voucher could not be located. Medley (1993) 
accepted only reports from Lyon, Trigg, and 


The genus Trifolium (Fabaceae) in Kentucky—Vincent 


Rockcastle counties. The earliest Kentucky 
collection I saw was from 1855, without local- 
ity (C.W. Short s.n., PH). 

Trifolium dubium is also called little hop 
clover (Gillett and Cochrane 1973; Gleason 
and Cronquist 1991), small hop clover (De- 
lorit and Gunn 1986; Isely 1998) and sham- 
rock (Small 1933). It is often confused with T. 
campestre, but can be distinguished from it by 
the smaller inflorescences with fewer flowers: 
its standard is not striate or only faintly so, 
whereas that of T. campestre is strongly striate. 
It is also commonly confused with Medicago 
lupulina L. (black medic), but can be distin- 
guished as described in the entry for T. cam- 
pestre; more records for little hop clover iden- 
tified as black medic may lurk in herbaria. Tri- 
folium dubium is thought by some to be the 
“shamrock” of Irish folklore, but others claim 
that the shamrock may be one of several spe- 
cies of Trifolium, Medicago, or Oxalis (Colgan 
1896; Everett 1971; Nelson 1991). 


5. Trifolium hybridum L., Sp. Pl. 2: 766. 
1753. Alsike clover. (Figure 5) 


Perennial, upright to ascending, 15-60(80) 
cm tall. Stems often much-branched, nearly 
glabrous, often somewhat fleshy. Leaves peti- 
olate, longest petioles to 80 (sometimes even 
100) mm, longer than the leaflets, gradually 
reduced upward. Stipules obovate to lanceo- 
late, tips long attenuate, 10-30 cm, adnate to 
petioles for about one-third their length. Leaf- 
lets 10-35 X 10-20 mm, sessile or nearly so, 
ovate to elliptical or fermi! base cuneate, 
apex rounded to slightly emar ginate, serrate. 
Inflorescence 10-25 mm roads globose, 
short-racemose to nearly onbelllare. on pe- 
duncles 20-80 mm; flowers 20-80; pedicels 1— 
5 mm, reflexing ih age. Calyx glabrous ex- 
cept in the U-shaped sinuses, tube 1-2 mm, 
teeth subulate, nearly equal, as long as or lon- 
ger than the tube. Corolla white and pink, 6- 
11 mm, the standard ovate-oblong, obtuse, 
sometimes emarginate. Fruit oblong, 3-4 mm. 
Seeds 2-4, mottled yellow brown, red brown, 
to nearly black, 1-1.3 mm. 2n = 16. Flowering 
in Kentucky in May-July, fruiting July-Sep- 
tember. 

Alsike clover is native to Europe, probably 
in the Mediterranean region. It is introduced 
throughout temperate regions worldwide, and 
is often cultivated. The species, also called Al- 


Figure 5. Trifolium hybridum L. Documented county- 
distribution in Kentucky; plant (from Hermann 1966). Bar 
= 20 mm (whole plant), 5 mm (flower), 1.5 mm (calyx). 


satian clover and Swedish clover (Delorit and 
Gunn 1986; Small 1933), was apparently first 
cultivated in Sweden, and first cultivated in 

England about 1832 (Taylor 1975). It was first 
brought to the United States about 1839 (Tay- 
lor 1975). The earliest Kentucky collection I 
saw was from 1895, Rockcastle ‘County (n.C., 
CINC). 

Trifolium hybridum may cause dermatitis in 
sensitive humans (Hardin and Arena 1974). 
Alsike clover is said to cause photosensitivity 
and biliary fibrosis in horses (Fisher 1995), 
though the connection between these diseases 
and the clover is not conclusive (Nation 1989). 

Trifolium nigrescens Viv. (ball clover, a 


8 Journal of the Kentucky Academy of Science 62(1) 


Medite:ranean species) is found with increas- 
ing frequency in southeastern United States 
(Isely 1990, 1998), and has been documented 


from: iumerous sites in Tennessee. It is pos- 
sible that this species will be encountered in 
the southern tier of Kentucky counties, espe- 
cially since it can be cultivated in the state 

Taylor and Sigafus 1984). Ball clover is an an- 
nual, prostrate to ascending, glabrous to gla- 
brescent species, which can be distinguished 
from Alsike clover by its habit, as well as by 

\-shaped sinuses be ‘tween the calyx lobes ( U- 
sh aped in T: hybridum), white to cream or yel- 
low-white (rarely pale pinkish) corolla (gen- 
erally deep pink in T. hybridum), and stipule s 
with sharply recurved, black to dark maroon, 
subulate tips (straight, green tips in T. hybri- 
dum). 


6. Trifolium incarnatum L., Sp. Pl. 2: 769. 
1753. Crimson clover. (Figure 6) 

Annual, upright, 20-90 cm tall, hairy 
throughout. Stems simple to sparingly 
branched below. Leaves long-petiolate below 
to nearly sessile above, longest petioles 4-5 
times the le ngth of the leafle tS. Stipules broad- 
ly ovate to oblong, sheathing the stem at the 
base, white to pale | green with dark green to 
red purple veins Below: tips toothed and 
rimmed with dark red purple or green. Leaf- 
lets 10-30(40) X 10-20(30) mm, sessile or 
nearly so, broadly ovate-obovate to orbicular, 
base broadly cuneate, apex obtuse to emargin- 
ate. Inflorescence 20-60 X 10-20 mm, spi- 

cate, densely cylindrical, peduncles 10-60 
mm; flowers many, sessile or nearly so. Calyx 
long-hairy, ae 3-5 mm, teeth subulate, near- 
ly equal, 1 2 times as long as the tube. Corolla 
crimson, te white or pink, longer than the 
calyx lobes, 10-17 mm, the standard linear- 
oblong to elliptical, acute. Fruit sessile, ob- 
long, 3-4 mm. Seed 1, buff to brown, 1.9-2.3 
mm. 2n = 14. Flowering in Kentucky in 
April-May, fruiting June-July. 

Crimson clover (also called Italian clover 
(Small 1933] and many other common names 
[Knight 1985a; Nourse 1894]), is native to 
southern and western Europe and widely nat- 
uralized in other areas. The species has been 
cultivated since the 1700s in Europe and was 
introduced into the United States in 1818 
(Knight 1985a). Crimson clover is used exten- 
sively as a ground cover in crop rotations, for 


Trifolium incarnatum L. Documented county- 
distribution in Kentucky; plant (from Hegi 1923). Bar = 
40 mm (whole plant), 10 mm (flower), 5 mm (calyx). 


Figure 6. 


green manure, and as a nitrogen-fixing plant 
in fields (Taylor and Sigafus 1984); it is also 
used an annual hay crop (Nourse 1894). It is 
occasionally cultivated in Kentucky (Garman 
1902: Taylor 1986: Taylor and Sigafus 1984). 
The “Males! Kentucky collection I saw was 
from 1934, Jefferson County (Bishop & Bish- 
op s.n., DHL). 


7. Trifolium pratense L., Sp. Pl. 2: 768. 1753. 
Red clover. (Figure 7) 

Perennial, ascending to upright, 20-60(100) 
em tall. Stems much-branched, with ap- 
pressed to spreading hairs or glabrous. Leaves 
long-petiolate below to nearly sessile above, 
longest petioles 3-4 times the length of the 
leaflets. Stipules ovate to lanceolate, 10-30 


The genus Trifolium (Fabaceae) in Kentucky—Vincent 9 


Figure 7. Trifolium pratense L. Documented county-dis- 
tribution in Kentucky; plant (from Besette and Chapman 
1992). Bar = 20 mm. 


mm, adnate to the petioles for most of their 
length, the lower portion pale with dark green 
to red veins tips long mucronate. Leaflets 10- 
30(50) < 7-15(25) mm, sessile or nearly so, 
ovate to elliptic or obovate, base broadly cu- 
neate, apex rounded, rarely slightly emargin- 
ate, essentially entire. Inflorescence single or 
in pairs, 10-30 < 10-30 mm, head-like, glo- 
bose, dense, sessile or on peduncles to about 
4 mm, subtended by a pair of bract-like leaves; 
flowers 40-150, sessile. Calyx hairy, tube 2.5— 
4 mm, teeth subulate, lowest about as long as 
the tube, others nearly equal and much short- 
er than the lowest. Corolla red purple to white 
or pinkish, longer than the calyx lobes, 11-18 
mm, the standard oblong-oblanceolate, emar- 

inate. Fruit ovoid-oblong, 2-3 mm. Seed 
1(2), tan to brown, 1.5-2 mm. 2n = 14, 28, 


56. Flowering in Kentucky in April—October, 
fruiting June—November. 

Trifolium pratense is morphologically very 
variable, and many binomials have been 
coined for the various forms; Zohary and Hell- 
er (1984) recognized six varieties of the spe- 
cies. Red clover (also called purple clover 
[Small 1933]) is the grown in more areas of 
the world than any other species of Trifolium 
(Taylor 1975). It is native to southeastern Eu- 
rope and Asia Minor (Smith et al. 1985). Tri- 
folium pratense has been in cultivation since 
the 3rd and 4th centuries, probably beginning 
in Spain, from where it spread to Holland and 
Lombardy, then to Germany. This species was 
introduced into England about 1645, from 
where it was brought to the New World by 
1663 (Taylor and Quesenberry 1996). It is a 
very important forage crop but may also cause 
bloating in animals overeating its young 
growth; a diet high in red clover may cause 
infertility in sheep (Taylor and Quesenberry 
1996). 

Red clover has been cultivated in Kentucky 
since at least 1803 (Fergus 1931; Taylor et al. 
1997a), and is probably naturalized in every 
county; Fergus (1931) indicated that red clo- 
ver was cultivated in every county in Ken- 
tucky. M’Murtrie (1819) reported red clover 
in the Louisville area as early as 1819. The 
earliest Kentucky collection I saw was from 
1892, Fayette County (Terrill s.n., kes). Tri- 
folium pratense has been described as a “ubi- 
quitist,” which may occur in practically any 
plant community (Merkenschlager 1934). Red 
clover has been used in revegetation of strip 
mine coal spoil fields in western Kentucky 
(Powell et al. 1980). It is the state flower of 
Vermont. 

Red clover may be used for tea or as an 
ingredient in herbal cough syrup (Coon 1980; 
Gibbons 1962), to flavor vinegar (Coon 1980), 
and as a salve to treat eye and skin diseases 
(RDA 1984). There are even claims that red 
clover can be used in cancer treatments (Duke 
1985; Ritchason 1995). The young growth can 
be cooked as a vegetable (Coon 1980). Dried 
flower heads have been powdered and used in 
breads during times of famine (Millspaugh 
1974). 

Trifolium medium (zigzag clover) is a similar 
perennial species sometimes cultivated in 
Kentucky (see Excluded Species). It differs 


10 Journal of the Kentucky 


Figure S. 
tribution in Kentucky; plant (from Isely 1951). 


Trifolium reflecum L. Documented county-dis- 
Bar = 20 
mm (whole plant), 3 mm (flower). 


from T. pratense by its peduncled inflores- 
cences, narrowly elliptical leaflets, and rhizo- 
matous nature. Trifolium hirtum (rose clover), 
an annual species resembling red clover, has 
been reported from nearby states and is oc- 
casionally cultivated in Ke ntucky (see Exclud- 
ed Species). 

8. Trifolium reflexum L., Sp. Pl. 2: 766. 1753. 
Buffalo clover. (Figure 8) 

Annual to biennial, ascending-upright, 20- 
50 cm tall. Stems simple to branched from the 
base, densely pubescent to glabrous. Leaves 
petiolate, gradually reduced “upward, longest 
petioles 3-4 times the le ngths of the eaters 
Stipules broadly ovate, leaflike, tips long acu- 
minate, entire to serrate. Leaflets 10—30(45) 
6—20(25) mm, sessile or nearly so, ovate to ob- 
ovate, base cuneate, apex acute broadly 


Academy of Science 62(1) 


rounded, serrate. Inflorescence 20—35(40) mm 
wide, umbellate, nearly spherical in flower, on 
peduncle s 20-60(S0) mm; flowers 10-40; ped- 
icels 4S mm, reflexing dramatically and elon- 
gating to 7-12(15) mm in fruit. Calyx hairy to 
glabrous, tube 1-1.5 mm, teeth linear, nearly 
equal, 3-7 mm, with broad, U-shaped sinuses 
between. Corolla deep pink to white, longer 
than the calyx lobes, 8-14 mm, the standard 
oblong to e lliptic, obtuse, often slightly emar- 
ginate. Fruit ote to oblong, 3-5 mm, slightly 
stalked. Seeds (1)2-4, pale hast 1-1.5 mm. 
2n = 16. Flowering in Ke sntucky in May, fruit- 
ing June. 

Buffalo clover is native to eastern North 
America from Virginia and the Carolinas south 
into Florida, west to central Texas, north to 
eastern Kansas, Nebraska, and Iowa, and east 
to Ohio; there is also an old record from east- 
ern Pennsylvania. The earliest Kentucky col- 
lection I saw was from 1835, Fayette County 
(Short s.n., GH); the most recent collection 
was from 1990, Trigg County (Chester et al. 
90-210, APSC). This species is becoming very 
rare in Kentucky (Taylor and Campbell 1989) 
and is listed as endangered in the state 
(KSNPC 1996): four extant “occurrences” are 
recorded by the Kentucky State Nature Pre- 
serves Commission (D.L. White, KSNPC, 
pers. comm., 19 Jan 2000). 

According to Taylor et al. (1994), buffalo 
clover is autogamously self pollinated. Some 
authors have recognized two varieties, based 
on pubescence differents but this character 
is variable and probably clinal in nature, with 
the more glabrous forms in the northeastern 
part of the range of the species. Glabrous and 
pubescent forms may grow intermingled in 
some mid-south populations, with the most 
densely pubescent populations occurring in 
the deep south and the western portion of the 
range. The species was rediscovered in Ohio 
in 1990 at a site that had burned the previous 
fall (Vincent 1991), but there have been no 
further fires, and it has not reappeared there 
since. Populations of this species often reap- 
pear in sites after a burn, heavy logging, or 
some equally severe disturbance. (pers. obs.). 

Trifolium virginicum Small ex Small & Vail 
(Kate’s Micha ein clover) is a similar species 
found on exposed shale barrens from south- 
western Pennsylvania south through West Vir- 
ginia to the Shenandoah alley of Virginia 


The genus Trifolium (Fabaceae) in Kentucky—Vincent 1] 


Figure 9. Trifolium repens L. Documented county-dis- 
tribution in Kentucky; plant (from Besette and Chapman 
1992). Bar = 20 mm (whole plant), 15 mm (inflores- 
cence). 


(Linscott 1994). It is a perennial species, with 
a rosette of leaves with narrowly elliptical leaf 
lets: the corolla is white. If suitable habitat 
were found in eastern Kentucky, this species 
might be found there. 


9. Trifolium repens L., Sp. Pl. 2: 767. 1753. 
White clover. (Figure 9) 


Perennial, stoloniferous to rhizomatous, 
rooting at the nodes, 10-30 cm tall. Stems 
much-branched, glabrous to sparsely hairy. 
Leaves petiolate, petioles 10-200 mm. Stip- 
ules thin and membranous, whitish to brown- 


ish, often with darker reddish to greenish 
veins, ovate-lanceolate, fused into a tube, tips 
short-attenuate, 8-15 mm. Leaflets 6-30 
10-25 mm, sessile or nearly so, broadly elliptic 
to ovate, base broadly cuneate, apex obtuse to 
emarginate or obcordate, serrate. Inflores- 
cence 15-35 mm broad, umbellate to short- 
racemose, nearly globose, on peduncles as 
long as or longer than the associated leaves, 
arising from leaf axils on the stolons; flowers 
20-50(100); pedicels reflexing dramatically 
with age. Calyx glabrous, often whitish with a 
purplish to green apex, tube 1.8—3 mm, teeth 
triangular-lanceolate, unequal, upper shorter 
than the tube, lower about as long as tube, 
sinus sharply V-shaped. Corolla white to pink- 
ish, 7-12 mm, the standard elliptic-obovate, 
obtuse. Fruit linear-oblong, 3-5 mm. Seeds 3- 
4, yellowish tan to brown, 0.9-1.5 mm. 2n = 
16, 28, 32, 48, 64. Flowering in Kentucky in 
March—November, fruiting June—November. 

White clover (also called Dutch clover and 
Ladino clover) may very well be the most im- 
portant temperate pasture plant (Baker and 
Williams 1987). It was introduced so early and 
was so widely grown in North America that it 
was known to Native Americans as “White 
man’s foot grass” (Strickland 1801); its culti- 
vation may have begun in the early 1700s, and 
it was widespread by the middle of that cen- 
tury (Isely 1998). Piper (1924) considered 
white clover “the most important perennial 
pasture plant in North America.” The species 
is widely grown in Kentucky (Rice et al. 1982) 
and was the earliest clover species cultivated 
in the state (Carrier and Bort 1916). 
M’Murtrie (1819) reported this species from 
the Louisville area. The earliest known Ken- 
tucky collection I saw was from 1890, Fayette 
County (Garman s.n., kes). The species is un- 
doubtedly to be found in every Kentucky 
county. 

Trifolium repens is extremely morphologi- 
cally plastic, and varies greatly in size of both 
leaves and flowers depending upon environ- 
mental conditions (Gillett and Cochrane 
1973). Zohary and Heller (1984) recognized 
nine intergrading varieties. Most North Amer- 
ican specimens appear to be T. repens var. re- 
pens. A recent monograph on the species COvV- 
ers in great detail many aspects of its taxono- 
my, morphology, and cultivation (Baker and 
Williams 1987). 


12 

White clover may have some medicinal 
uses, although human ingestion of powdered 
fresh heads resulted in “a sensation of 
fullness and congestion of the salivary glands 
with pain, and mump-like pain ... fol- 
lowed by copious flow of saliva” (Millspaugh 


1974). 

\ similar species is T. calcaricum Collins & 
Wieboldt, which differs from white clover in 
its terminal inflorescences. It is native to the 
cedar glades of central Tennessee and south- 
western Virginia (Collins and Wieboldt 1992). 
If similar cedar glade habitat exists in south- 

eastern Rentals it is possible that this spe- 

cies could be found in the state: it is found in 
Lee County, Virginia, within 10 miles of the 
Kentuc ky state line. 


10. Trifolium resupinatum L., Sp. Pl. 2: 771 
1753. Persian clover. (Figure 10) 

Annual, procumbent to ascending or up- 
right, 10-60 cm tall. Stems often much- 
branched, glabrous or nearly so. Leaves peti- 
olate Belon to nearly sessile above, longest 
petioles to 4-5 times the length of the lee ths 
Stipules lanceolate to lanceolate-ov ate, tubular 
at the base, tips long attenuate, shorter than 
the associated petioles. Leaflets 5-20(30) x 2— 
4 mm, sessile or nearly so, obovate, elliptic to 
lanceolate, or rhombic, serrate, base cuneate, 
apex rounded to acute. Inflorescence 8—15 
mm broad, capitate, densely hemispherical, on 
peduncles 20-50 mm; flowers 6-20, short-pet- 
iolate to sessile. Calyx glabrous except for a 
dorsal band of hairs, w hitish to pale green with 
a dark basal band, tube 1.5—2 mm, teeth su- 
bulate, unequal, shorter than the tube, often 
dark green; calyx becoming inflated and en- 
closing the fruit at maturity, the veins obvi- 
ously reticulating. Corolla resupinate, lavender 
to pink or rarely white, 4-9 mm, the standard 
oblong, emarginate. Fruiting head globose, 
looking star-like. Fruit ovoid- lenticular, 1.7- 
2.3 mm. Seed 1, yellow to tan or purple 
brown, 1.2-2 mm. 2n = 14, 16, 32. Flowering 
in Kentucky in May-June, fruiting June-July. 

The resupinate (inverted, with ine standard 
below and the keel above) corolla and inflated 
fruiting calyx makes T. resupinatum easy to 
distinguish ‘from other clovers. The presence 
of Persian clover in Kentucky was first report- 
ed by McFarland (1942), whose report was ac- 
ce pted by Browne and Athey ( 1992) and Med- 


2 Journal of the Kentucky Academy of Science 62(1) 


Trifolium resupinatum L. 


Figure 10. Trifolium resupinatum L. Documented coun- 
ty-distribution in Kentucky; plant (from Hegi 1923). Bar 
= 20 mm (whole plant), 3 mm (flower and fruiting calyx). 


ley (1993). It may be cultivated in the state as 
a winter annual (Taylor and Sigafus 1984). I 
located only one Kentucky collection, from 
1915, Metcalfe County (Salmon s.n., kes). A 
report by Browne and Athey (1992) for the 
Shawnee Hills was rejected by Medley (1993), 
and I was unable to locate a specimen to verify 
the report. Widespread in much of the south- 
eastern United States (Isely 1990), this species 
was recently documented from Ohio (Vincent 
and Cusick 1998). It is reported from “scat- 
tered stations” in northeastern U.S. by Glea- 
son and Cronquist (1991). 


Ll. Trifolium stoloniferum Muhl. ex A. 
Eaton, Man. Bot. 468. 1818. Running buffalo 
clover. (Figure 11) 

Perennial, stoloniferous, upright flowering 
branches 10—40 cm tall. Prostrate stems often 


The genus Trifolium (Fabaceae) in Kentucky—Vincent 13 


Trifolium stoloniferum 
Muhl. ex A.Eaton 


Figure 11. 


little branched, glabrous or nearly so, rooting 
at the nodes, forming extensive clones. Leaves 
from stolons long-petiolate, those on the up- 
right flowering stems in a pair, petioles as long 
as the leaflets or shorter. Stipules of stolons 
broadly lanceolate, membranous, tips attenu- 
ate, shorter than the associated petioles; those 
of the upright stems leaf-like, ovate-oblong, 
broadly triangular to attenuate, slightly to 
much-serrate. Leaflets 10-40 < 8-35 mm, 
serrate, on petiolules about 1 mm, obovate to 
obcordate, base broadly cuneate, apex round- 
ed to emarginate. Inflorescence 15-30 mm 
wide, umbellate, spherical, on peduncles 10- 
30 mm; flowers 25-45; pedicels 2-8 mm, re- 
flexing dramatically in age. Calyx glabrous or 
nearly so, tube 1.5-2.5 mm, teeth subulate, 
nearly equal, about twice the length of the 
tube. Corolla white, sometimes pinkish with 
age, 8-14 mm, the standard obovate to ob- 


Trifolium stoloniferum Muhl. ex A. Eaton. Documented county-distribution in Kentucky; plant (drawn by 
Ethel Hickey 1995, used with permission). Bar = 20 mm. 


long, rounded to emarginate. Fruit oblong, 
2.5-3 mm. Seeds 1-2, yellow to brown, 1.3-2 
mm. 2n = 16, 32. Flowering in Kentucky in 
April-May, fruiting May-June. 

In spite of the use of the citation “Muhl., 
Cat. Pl. Amer. Sept. 67. 1813.” for this name 
(Browne and Athey 1992; Medley 1993; Zo- 
hary and Heller 1984), the authorship should 
be given as “Muhl. ex A. Eaton” since Muhl- 
enberg’s (1813) publication of the name was a 
“nomen nudum” as stated by Merrill and Hu 
(1949) and Brooks (1983). 

Running buffalo clover once ranged widely 
over middle east-central North America from 
present-day West Virginia west to Kansas, and 
from Arkansas north to north-central Ohio 
(Brooks 1983). Its predominant range was 
montane West Virginia and the Ohio River 
drainage in Ohio and south to central Ken- 
tucky. It was once found in great stands in 


14 Journal of the Kentucky 
Kentuck: (Campbell et al. 1988) and Ohio 
(Cusick 1989). This species, once thought ex- 
tinct as rediscovered in 1983 by B: urtgis 
(1985). and was listed in 1987 by the US Fish 


and \Vildlife Service as an endangered species 

\nonymous 1987) under the federal Endan- 
gered Species Act. It was thought to be ve ry 
rare in Kentucky (Taylor and C ‘ampbell 1989), 
and is listed as a threatened species in the 
state (KSNPC 1996): 69 extant “occurrences” 
are recorded in the state by the KSNPC (D.L. 
White, KNSPC, pers. comm. 19 Jan 2000). 
The earliest Kentucky collection I saw was 
from 1834, Fayette County (Peter s.n., MICH, 
NY): the most recent is from 1995, Madison 
County (Vincent et al. 6959, MU). 

Taylor et al. (1994) stated that running buf- 
falo clover is an outcrossing species that sets 
fewer seeds if selfing, Bal that seed set in 
selfed plants was still high enough to maintain 
the species in the wild. They At: suggested 
that habitat loss and competition may comely 
ute more to the decline of the species than 
inbreeding. Hickey et al. (1991) found that ge- 
netic diversity was low among many popula- 
tions of the species, based on “alllorsc me band- 
ing patterns, and that many populations might 
actually represent clones. Crawford et zal 

(1998), however, in a study using RAPDs, 
found that most populations were not single 
clones, and that even the smallest populations 
contained unique § genetic information. 

Running apelin clover is sometimes con- 
fused aie T. repens, from which it differs by 
the bract-like pair of leaves below the inflo- 
rescence on the upright stems, and by the 
overall larger size of the former. Another sim- 
ilar species is T. calcaricum, which is discussed 
under the treatment of T. repens. 

EXCLUDED SPECIES 

Trifolium alexandrinum L. Berseem clover 

This species was reported from cultivation 
by Garman (1902) and more recently by Tay- 
lor and Sigafus (1984). Its presence in Ken- 


tucky outside of cultivation was rejected by 
There is a Fayette County 


Medle 2y 
specimen 


(1993), 
(Garman s.n., 


kes), 


but it is from 


cultivation. I saw no specimen of this clover 


from other than cultivation. 


Trifolium ambiguum L. Kura clover 
Kura clover 
and is a very hardy rhizomatous perennial 


can be cultivated in Kentucky 


Academy of Science 62(1) 


(Taylor 199la; Taylor et al. 1997b). Isely 
(1998) and Kartesz (1999) reported T. ambig- 
uum as an escape in Ohio, but the reports 
were based on cultivated material; it is not yet 
documented that the species will escape in 
North America. All Kentucky collections I ex- 
amined were of cultivated material. 


Trifolium hirtum L. Rose clover 


Rose clover was reported for Kentucky by 
Isely (1990, 1998) and was provisionally ac- 
cepte 4 by Medley (1993). Kartesz (1999) ac- 
cepted the species as part of the state flora. 
The only Kentucky specimens identified by Is- 
ely as T. hirtum were at NCU. Of those, all 
were T. pratense except one, a cultivated spec- 
imen of rose clover from Jefferson County 
(Gunn J150, NCU). If this species becomes 
widely cultivated, and since it is possible to 
grow it in Kentucky (Taylor and Sigafus 1984), 
it could very well become established in the 
state. 


Trifolium medium L. Zigzag clover 
~ © 


Zigzag clover was reported for Kentucky by 
Garman (1902) and Linney (1880). I saw no 
non-cultivated Kentucky specimens of this 
species, although it is known to be cultivated 
in the state (Tay lor 1991b). Medley (1993) re- 
jected the occurrence of this species in Ken- 
tucky. Gleason and Cronquist (1991) reported 
that T. medium occasionally escapes from cul- 
tivation in northeastern North America. I have 
seen specimens from escaped populations in 
North Carolina, Massachusetts, Maine, and 
eastern Canada. 

Other clover species which are known only 
from cultivation in Kentucky include T. vesi- 
culosum Savi (arrowleaf clover) and T. subter- 
ranean L. (sub clover) (Taylor and Sigafus 
1984). In addition, Dr. Norman Taylor has cul- 
tivated many other species in greenhouses and 
field plots in Lexington. 


ACKNOWLEDGMENTS 


I thank the curators of the herbaria for al- 
lowing me access to specimens for this study. 
I am very grateful to Dr. Ralph Jones, Dr. 
Norman daylon Dr. John W. Thieret, and Dr. 

Ralph Thompson for sharing information 
about Kentucky clovers with me. 


The genus Trifoliwm (Fabaceae) in Kentucky—Vincent 15 


LITERATURE CITED 


Anonymous. 1987. Endangered and threatened wildlife 
and plants: determination of endangered status for Tri- 
folium stoloniferum (running buffalo clover). Fed. Reg- 
ister 52(108):21478—21480. 

Atwood, S. S. 1938. A “one-leaved” white clover. J. Hered. 
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J. Ky. 


Acad 62(1):18—-25. 2001. 


The Role of Light in Regulating Dandelion (Taraxacum officinale; 
Asteraceae) Inflorescence Height 


David Lowell Robinson 


Department of Biology, Bellarmine University, Louisville, Kentucky 40205 


ABSTRACT 

This research examined the ecophysiological basis for extension of the stalks (scapes) supporting the 
inflorescences (heads) of the common turfgrass weed, dandelion (Taraxacum officinale Weber). In turf, a 
statistically significant positive correlation was observed between turfgrass height and height of dandelion 
heads occurring in it. Dandelion heads tended to extend to the top of the turfgrass canopy only, whereas 
seedheads extended, on average, an additional 11 em above that. Excised scape segments taken from scapes 
at the pre-flowering stage elongated significantly less in the light than in the dark; those taken after flowering 
elongated the same in the light as in the dark and significantly more than the illuminated pre-flowering 
scapes segments. In a whole-plant study, pre-flowering scapes grown in a far-red enriched microenvironment 
elongated significantly more than scapes grown in other light microenvironments. Scape elongation after 
flowering, however, was not statistically different in any treatment. Pre-flowering scape elongation in dan- 
delion, therefore, appears to be a phytochrome-mediated response, whereas elongation after flowering is 
not. These results suggest that dandelion scape elongation may be physiologically linked to the height of the 


turfgrass canopy. 


INTRODUCTION 


Dandelion (Taraxacum officinale Weber) 
possesses numerous weedy characteristics that 
make it one of the most common invasive 
plants of turf in urban areas (Longyear 1918). 
It is adapted to a wide range of environments, 


is a long-lived species, reproduces vegetatively 


from its taproot, and has few natural enemies 
(Crutchfield and Potter 1995: Mitich 1989: 
Roberts 1936). Dandelion is also a prodigious 
seed producer: up to 60,000 seed m? per sea- 
son (Roberts 1936). Taraxacum species can set 
seed apomictically, thus ensuring high seed 
production even under conditions unfavorable 
for fertilization, such as extreme temperatures, 
water stress, or lack of pollinators (Munn 
1919). Seed dispersal over long distances is as- 
sisted by the parachute-like pappus attached 
to each seed. 

Another characteristic contributing to the 
weedy nature of this species concerns the 
grow th habit of its reproductive stalk (scape). 

Dandelion inflorescences (heads) often do not 
extend far enough above the soil surface to be 
cut by turforass. mowers. Prior to flowering, 
dandelion scapes grow straight upwards, but 
immediately after ‘flowering elongation slows 
noticeably and the uppermost portion of the 
scape can become diagravitropic (Clifford and 
Oxlade 1989). As a result, dandelion heads, for 


the majority of the time it takes for the seed 
to mature, are kept low to the ground and be- 
low the lawnmower blade (Longyear 1918). 
During seed maturation the scape rapidly 
elongates upwards again, lifting the seedhead 
far lsat e the canopy and into a better position 
for dispersal of seed by wind (Chao 1947; 
Longyear 1918). Growth during the pre- oes 
ering and pre-shattering (post-flowering) stag- 
es may be partially controller by hormones 
(Clifford et al. 1985: Clifford and Oxlade 1989: 
Oxlade and Clifford 1981). 

My research concerns the ecophysiological 
mechanisms controlling elongation of dande- 
lion scapes. There have been numerous re- 
ports on how radiant energy controls the way 
plants develop and interact with one another 
(Ballare et al. 1992; Briggs 1996; Holt 1995; 
Koornneef and Kendrick 1994: Schmitt and 
Wulff 1993). One of the most important plant 
molecules involved in the detection of light is 
phytochrome. Numerous plant photomorpho- 
genic responses are mediated by phyto- 
chrome, including those regulating plant 
height, branching, leaf shape, photoperiodic 
Gar ering, photosynthate allocation, and seed 
germination (Ballare et al. 1988; Ballare et al. 
1990; Ballare et al. 1992; Novoplansky 1991; 
Sanchez 1971; Schmitt and Wulff 1993; Smith 
1982: Vierstra 1993). 

Phytochrome is a family of photoreceptor 


Dandelion Inflorescence Height—Robinson 19 


molecules containing a photoreversible pig- 
ment that can absorb either red (maximal at 
666 nm) or far-red (maximal at 730 nm) light. 
Plants grown completely in the dark synthe- 
size the form of phytochrome (designated P,) 
that absorbs red light. Following an exposure 
to red light the P, pigment converts to a form 
(designated P,,) that absorbs far-red light. Af- 
ter an exposure to far-red light the P,, can con- 
vert back to P, in a cyclical process. One of 
the ways that red:far-red ratios are modified in 
nature is by the presence of neighboring plant 
foliage (Ballare et al. 1988; Holmes and Smith 
1975; Smith 1994). This is due to the prefer- 
ential absorption of red light (vs. far-red) by 
chlorophyll. If a plant is growing by itself in 
uninterrupted sunlight, the microenvironment 
will contain more red light than far-red, and 
there will be relatively more P,, in the plant, 
while a plant growing in a dense vegetative 
canopy will be exposed to more far-red light 
than red and contain relatively more P,. The 
physiologically active form of phytochrome is 
P,,. When present, it induces the synthesis of 
a cascade of gene products involved in the 
photomorphogenic traits mentioned above 
(Smith 1994; Vierstra 1993). Generally, high 
levels of P;, inhibit elongation of plant cells 
and tissues. The magnitude of phytochrome- 
mediated responses is often a function of the 
P,.: P, ratio at any given time. 

In plants, exposure to lowered red:far-red 
ratios typically induces an increase in apical 
elongation (height) at the expense of lateral 
growth (Ballare et al. 1987, 1990; Schmitt and 
Wulff 1993; Smith 1982), and is the reason 
that plants growing beneath a plant canopy are 
generally taller than in an open field (Holmes 
and Smith 1975; Solangaarachchi and Harper 
1987). Even plants similar in height can have 
this influence on one another at distances up 
to 30 cm (Smith et al. 1990). The ecological 
consequence of this “neighbor effect” is that 
plants detect the presence of other plants in 
their vicinity before those plants become com- 
petitors for sunlight or pollinators (Ballare et 
al. 1988, 1991; Holt 1995). 

The objectives of my study were to examine 
the role of light in regulating dandelion scape 
elongation, and to describe the developmental 
changes in light sensitivity of the scape during 
the transition from flowering to seed dispersal. 


MATERIALS AND METHODS 

Field Observation 

Naturally occurring variation in dandelion 
scape height was evaluated in Louisville, Ken- 
tucky, in April-May 1996 and April 1997. Six- 
ty-six randomly selected turfgrass sites were 
surveyed in 1996; 81 sites in 1997. Survey sites 
were from public and residential turfgrass ar- 
eas that were in full sunlight and had not been 
recently mowed. Measurements taken at each 
site were height of open inflorescence (from 
soil level to the bottom of receptacle), height 
of shattering seedhead (from soil level to bot- 
tom of receptacle), and turfgrass canopy 
height. In both cases, the head was pulled ver- 
tically taut while measuring heights. Best-fit 
regression analysis was performed with these 
data (Jandel Scientific 1995). 


Excised-Scape Study 


Dandelion scapes at either the pre-flower- 
ing or pre-shattering (post-flowering) stages 
were collected in May 1996 from a single, uni- 
form turfgrass location on the Bellarmine Uni- 
versity campus (Louisville, Kentucky), excised 
into 1 cm segments, and floated in 100 x 15 
mm petri dishes containing a 20 ml sucrose 
solution (10 g I) and 30 uM _ 3-indoleacetic 
acid, as described by Chao (1947). It has been 
determined that indoleacetic acid is necessary 
to prevent the scape segments from splitting. 
Only the upper 4 cm of each scape were used 
for sampling, and segments were evenly dis- 
tributed into different treatments (dishes). 

Dishes containing the scape segments were 
immediately placed onto a glass shelf in an en- 
vironmental growth chamber and allowed to 
grow for 3 days at constant illumination and 
constant temperature (26°C). Sixteen 160-W 
fluorescent bulbs provided illumination from 
above, and two 100-W clear incandescent 
bulbs provided illumination from below. The 
photosynthetic photon flux density (PPFD) 
provided from above was 874 mol m~°s"'; 
the PPFD from below was 42 pmol m~?s"!, 
both measured by a LI-1800 spectroradiome- 
ter (LI-COR Inc). 

Transparent, colored cellophane filters cut 
to the size of the dishes were placed on the 
bottom to filter the light coming from the in- 
candescent bulbs placed below. Filters trans- 
mitted clear, blue, red, or far-red light. Far- 


20 Journal of the Kentucky Academy of Science 62(1) 


red filters were achieved by overlapping blue 
with red filters. The thickness of the colored 
filters was adjusted so that each had an equiv- 
alent amount of light passing through. Their 
ectral qualities are described in the next sec- 
ion. Two experimental controls were used: 
one in which the bottom light was blocked 
with paper and another in which the dish was 
completely wrapped in aluminum foil. Each 
experiment consisted of 12 dishes of excised 
dandelion segments: six from scapes collected 
prior to flowering, and six from scapes col- 
lected prior to shattering. Each dish held 14 
scape segments taken from 14 different plants. 
The segments in each dish were illuminated 
the same from above but were treated with 
one of the five different light treatments from 
below, while the sixth dish received no light at 
all. Three days after the start of the experi- 
ment the ler vgth of each segment was mea- 
sured and av eraged. There were five replicat- 
ed experiments fitecks) ), and results were sub- 
jected to two-way analysis of variance (Jandel 
Scientific 1995). 


Whole-Plant Study 


Dandelion seeds used in this study were 
collected in May 1995 from a single popula- 
tion in Afton, Minnesota. The peed: were 
planted into 216 cm® plastic containers filled 
with potting soil (Metro-Mix 510) and grown 
in the environmental growth chamber with an 
alternating 14-hr light period (26°C) and 10- 
hr dark period (20°C). The same growth 
chamber as described above was used except 
that six incandescent bulbs (100 W) were used 
in addition to the fluorescent ones and no il- 
lumination was provided from below. This 
generated a PPFD of 950 pmol m~?s~!. After 
2 wk, plants were thinned to three plants per 
container. Seven weeks after sowing, the 
plants were vernalized by placing them in a 
5°C coldroom and illuminated with fluores- 
cent lights. After 4 mo, the plants were taken 
out and acclimated. 

After acclimation at room temperature for 
| day, single pots were placed individually into 
the Gatton of single transparent plastic cylin- 
ders (10-cm daneian! Two windows were cut 
into the cylinders just above soil level so leaves 
could be pulled out through the windows and 
held horizontal. This was to prevent photosyn- 
thesis from being affected by the light treat- 


ments and to prevent the foliage from inter- 
fering with the light microenvironment around 
the scape. Transparent colored cellophane 
(described above) was wrapped around the 
cylinders above the windows extending 7 cm 
above the soil. Blue, red, and far-red filters 
were used. As done previously, the thickness 
of the colored filters was adjusted so that there 
was an equivalent amount of light passing 
through each. As experimental controls, clear 
plastic was wrapped around one cylinder in 
the same fashion, and opaque black plastic was 
wrapped around another. The transmission 
spectra for the filters were determined with 
the spectroradiometer. Spectral quality (R: FR) 
of the different filters was calculated as de- 
scribed by Smith (1994). The R:FR_ ratios 
were: clear = 1.54, opaque = 1.43, red = 1.48, 
far-red 1.37, and blue = 1.36. Although 
these values are higher than those reported by 
other researchers (because most of the radiant 
energy was emitted from fluorescent light 
bulbs) they are within the range of values 
known to induce phytochrome-mediated re- 
sponses (Holmes and Smith 1975; Smith 1994; 
Weller and Reid 1993). Each replication 
(block), of which there were six in all, consisted 
of five cylinders representing these five light 
treatments. 

Cylinders were placed at least 30 cm apart 
in the growth chamber. The long- -day condi- 
tions in the chamber induced the plants to 
flower. Height of the first six dandelion scapes 
to emerge fr om each pot was monitored daily, 
from ANS first day of flowering until seedhead 
opening. Heights were measured from soil 
level to bottom of receptacle. Two-way analy- 
sis of variance was used to test for significance 
(Jandel Scientific 1995). 


RESULTS 
Field Observation 


To illustrate the relationship between dan- 
delion scape height and height of the sur- 
rounding canopy, observations were made in 
dozens of different public and residential turf 
grass communities in Louisville in spring 1996 
and 1997. Similar results were observed in 
both years (Figure 1). The height of the turf- 
grass canopies ranged from 2 to 35 cm; the 
height of the dandelion heads growing in them 
ranged from 3 to 32 cm. Generally, the heads 


Dandelion Inflorescence Height—Robinson 21 


Seedhead -——_~_ 
O Oo 
© 


Spring 1996 


Kn. Inflorescence 


Scape Height (cm) 


Scape Height (cm) 


Turf Height (cm) 


Figure 1. The relationship between turfgrass canopy 
height and dandelion scape height at various turfgrass sites 
in Louisville, Kentucky, during spring 1996 (upper) and 
1997 (lower). In 1996 (n = 66), inflorescence (head) 
height was described by Y = 3.884 + 0.737X (° = 0.70), 
and seedhead height was described by Y = 13.429 + 
0.912X (12 = 0.65). In 1997 (n = 81) inflorescence (head) 
height was described by Y = 2.683 + 0.717X (x2 = 0.86), 
and seedhead height was described by Y = 10.716 + 
0.953X (x2 = 0.75). 


extended only to the top of the turfgrass can- 
opies. Linear regression analysis revealed a 
statistically significant (P < 0.001) correlation 
between head height and turf height with r 
values of 0.70 for 1996 and 0.86 for 1997. In 
1996, average head height was the same as the 
average canopy height, whereas in 1997 the 
average head was | cm shorter than the can- 
opy. Regression analysis indicated that heads 
were slightly taller than the surrounding turf 
at canopy heights below 15 cm in 1996 (9 cm 
in 1997). Above that height, dandelion heads 
tended to be slightly shorter than the canopy. 
In the tallest turfgrass communities, the dan- 
delion heads averaged 5 cm (1996) and 7 cm 
(1997) below the canopy. 


Seedhead heights, measured at the same lo- 
cations, ranged from 10 to 54 cm and were 
significantly taller than the height of the 
neighboring turfgrass (on average, 12 cm taller 
in 1996 and 10 cm taller in 1997). Seedhead 
height was significantly correlated (P < 0.001) 
with the height of the turf canopy with r° val- 
ues of 0.65 for 1996 and 0.75 for 1997. In 
1996, the slopes for the head and seedhead 
regression lines were not statistically different 
from one another, whereas in 1997 they were 
(P < 0.001). 

In this field study, the positive correlation 
of dandelion scape elongation with height of 
the neighboring turf canopy suggests that dan- 
delions adjust their scape growth to the veg- 
etative growth of the turfgrass species sur- 
rounding it. Since head and seedhead heights 
were measured at the same time and at the 
same place, the changes in growth of dande- 
lion scapes prior to flowering versus their 
growth after flowering were probably due to 
physiological shifts within the scapes them- 
selves and not to changes in the environment. 
The other experiments described in this paper 
were designed to explore the possibility that 
this regulation of growth is phytochrome me- 
diated. Since scapes at the flowering and seed- 
head stages appeared to respond to the mi- 
croenvironment differently, the effect of light 
at these two developmental stages was also ex- 
amined. 


Excised-Scape Study 


Pre-flowering scape segments (that were il- 
luminated) elongated an average of 38%, 
whereas segments from older scapes, carrying 
heads that were about to shatter, elongated an 
average of 50% (Table 1). This difference was 
statistically significant (P < 0.001) and indi- 
cates that scape growth, in response to light, 
shifts as it progresses from one developmental 
stage to another. The only statistical difference 
in elongation among the pre-flowering scape 
segment treatments was for those kept in 
complete darkness. This experimental control 
elongated significantly more (15%) than pre- 
flowering scape segments grown in light. In 
the dark, however, the pre-flowering scape 
segments elongated as much as the pre-shat- 
tering segments kept in either the light or the 
dark. No significant differences were observed 
for pre-shatter scape elongation in any of the 


22 Journal of the Kentucky Academy of Science 62(1) 


Table |. Length of pre-flower and pre-shatter dandelion 
scapt sments after 3 d treatment with supplemental 
light. Scapes were collected in May 1996 in Louisville, 
K ky and excised into 10 mm lengths before treat- 
it. Average of 5 replications, 14 segments per repli- 
cation (n = 70 segments per treatment). Means followed 
by the same letter, within a column, are not significantly 
different (P < 0.05) according to the Student-Newman- 
Keuls test. 
a Pre-flower scape Pre-shatter scape 
segment length segment haneti 
Filter color mim) * SE mm) = SE 
Clear 13.8 + 0.09 b 14.9 + 0.05 a 
Barrier 13.7 + 0.06 b 14.6 + 0.05 a 
Blue 13.7 + 0.07 b 15.1 = 0.05 a 
Red 14.1 + 0.06 b 15.2 + 0.05 a 
Far-Red 13.7 = 0.04 b 152 = 0:05" a 
Wrapped 15.9 + 0.02 a 15.0 = 0.05 a 


treatments. All pre-shatter scape segments 
grew the same whether they were placed in 
the dark, in the light, or were supplemented 
with specific wavelengths of light from be- 
neath. This experiment demonstrates that 
scapes have the same growth potential before 
flowering as they do aftenw ards, but that this 
potential is not realized prior to flowering be- 
cause of light sensitivity at that stage. 


Whole-Plant Study 


The flowering head heights in the two con- 
trol treatments (clear atl opaque) were not 
significantly different from one another and 
averaged 10.6 cm above soil level, which was 
3.6 cm above the top of the filters (Table 2). 
The blue and red treatments were not statis- 
tically different from the clear and opaque 
treatments. Plants treated with far-red light, 
however, produced heads that were 23% taller 
than any of the other treatments or controls 
(significant at P < 0.05). This differential re- 
sponse for far-red light, versus red light, in- 


dicates that phytochrome may have a role in 
regulating head elongation. 

It took an ave rage of 7.5 days for heads to 
develop into shattering see »dheads. There were 
no statistical differences for this time frame 
among the treatments. During this period the 
scapes elongated another 6 cm, on average, 
representing an increase of 52% (Table 2). As 
with the heads, there were no significant dif- 
ferences in seedhead scape height in the clear, 
opaque, blue or red treatments. Elongation of 
the far-red treated scapes, however, was sta- 
tistically greater. When head height was sub- 
tracted from seedhead height, no significant 
differences were apparent, indicating that 
post-flowering elongation may not be strongly 
influenced ee the light microenvironment. 
These results support the field observation 
that dandelion seedheads extend a fairly uni- 
form distance above the turfgrass canopy re- 

gardless of their height at flowering (Figure 1). 
It also supports the explant study showing that 
seedhead scape tissue was less sensitive to 
light than scape tissue at the flowering stage 
(Table 1). 

Another photomorphogenic response that 
occurred in this study involved the time it took 
for the plants to flower. It took 13.2 days, from 
the time the blue-treated plants were placed 
in the growth chamber, to the time they pro- 
duced six heads (Table 3). The far-red treat- 
ment completed blooming after only 9.5 days, 
significantly faster (P < 0.05) than the blue- 
meried plants. Accelerated flowering rates are 
associated with plants pursuing a shade-avoid- 
ance strategy, which is generally phytochrome 
mediated (Smith 1994). 


DISCUSSION 


Like many weedy-plant species, dandelions 
show large amounts of phenotypic plasticity 


Table 2. Head and seedhead elongation in dandelion plants treated with supplemental light. Prior to treatment, plants 


were induced to flower under enviornmentally controlled conditions. Average of 6 replications, 6 scapes per replication 


(n = 36 scapes per treatment). Means followed by the same letter, within a column, are not significantly different (P 


< 0.05) according to the Student-Newman-Keuls test. 


Filter color Head height (em) + SE 


Seedhead height (em) = SE 


Difference (em) = SE 


Clear 10.1 + 0.46 b 
Opaque 11.1 + 0.79 b 
Blue 10.5 = 0.62 b 
Red 11.0 + 0.48 b 
Far-Red 13.1 = 0.64 a 


16.0 1.47 b 5iGr == allo 
16.9 + 0.82 b 5.8 + 0.62 a 
15.0 = 1.00 be 45+ 0.8la 
Vip E2S ih 6.2 + 140a 
19.8 + 1.22; 6.7, eS ra 


Dandelion Inflorescence Height—Robinson 93 


Table 3. Time it took for the first 6 heads to emerge in 
dandelion plants treated with supplemental light. Prior to 
treatment, plants were induced to flower under enviorn- 
mentally controlled conditions. Average of 6 replications, 
6 scapes per replication (n = 36 segments per treatment). 
Means followed by the same letter, within a column, are 
not significantly different (P < 0.05) according to the Stu- 
dent-Newman-Keuls test. 


Filter color Days to 6th head + SE 
Clear 10.3 = 1.41 ab 
Opaque 12.3 + 0.49 ab 
Blue 13.2 + 1.08 a 
Red 10.3 + 0.92 ab 
Far-Red 9.5 + 0.76 b 


(Clifford and Oxlade 1996). These ecological 
adaptations are exhibited in their reproductive 
biology. Dandelion heads require light in or- 
der to open, and they must open if they are 
to set seed (Longyear 1918; Roberts 1936; Ta- 
naka et al. 1987). This means that heads ex- 
tending to the top of the turfgrass canopy 
maximize their potential for setting seed. 
However, the farther above the canopy dan- 
delion heads extend the greater their risk of 
being separated from the plant by mowing. Af- 
ter flowering, the tight control of scape elon- 
gation may not be as important because ex- 
tension of the seedhead above the canopy sur- 
face enhances seed dispersal by wind. This 
shift in elongation strategy can occur rapidly 
as both Longyear (1918) and Roberts (1936) 
observed that dandelions producewiable seed 
long before seedheads actually shatter. 

This research illustrates that control of dan- 
delion scape growth is due to an interaction 
between the physiological status of the plant 
and the surrounding microenvironment. In 
the field, dandelion plants produce heads that 
just reach the top of the turfgrass canopy. This 
linkage of dandelion head height to the height 
of the surrounding turf is maintained in com- 
munities ranging from only 2 cm in height up 
to more than 17 times that. The laboratory 
studies demonstrate that control over scape 
elongation is mediated by phytochrome. The 
far-red treatment (with a red:far-red ratio of 
1.37) caused scapes to grow significantly taller 
than light treatments with higher red: far-red 
ratios, like clear (1.54), red (1.48), and opaque 
(1.43). Due to the asymptotic relationship be- 
tween red:far-red ratios and the amount of P,, 


in plant tissues (relative to P,,,,;), small changes 
in the quality of light have been found to elicit 
major physiological changes in plants (Smith 
1994). Previous researchers have shown that 
phytochrome-mediated responses are corre- 
lated to relative amount of far-red light up to 
red:far-red values of 2.00 (Holmes and Smith 
1975: Smith 1994; Weller and Reid 1993). 

We have measured the red:far-red ratios in 
natural turfgrass canopy microenvironments to 
be 1.06, compared to 1.14 in full sun. As dan- 
delion scapes form and elongate, the lower 
red: far-red ratio caused by the preferential 
absorption of red light by turf foliage may en- 
courage elongation due to predominance of P, 
phytochrome, the form which does not inhibit 
elongation. When the inflorescence bud ap- 
proaches the top of the canopy, however, the 
red:far-red ratio may be high enough (due to 
predominance of the P,, form of phytochrome) 
to discourage any further scape elongation 
and, at the same time, induce head opening. 

The photomorphogenic control of scape 
elongation by phytochrome appears to be de- 
velopmentally controlled. Whereas all three of 
these studies indicate that dandelion scapes, 
up to the time of flowering, are sensitive to 
light, no evidence was found that scape growth 
after flowering was affected by light. Thus, 
major physiological changes must be occurring 
in this tissue in a relatively short period of 
time: 7.5 days in this growth-chamber study 
vs. 9 or 10 days in the field (Longyear 1918). 
Scape elongation rates between the flowering 
and shattering stages in the clear-plastic treat- 
ment of the whole-plant study averaged more 
than 8 mm d“!. This growth is rapid enough 
to carry the head above the turfgrass canopy 
at a faster rate than the canopy height itself 
increases. Chao (1947) reported that the ma- 
jority of this scape growth occurred in the up- 
per third of the scape and involved increases 
in epidermal cell length, fresh weight, and 
non-protein nitrogen content, as well as de- 
creases in dry weight, and protein nitrogen 
content. 

An important source of light perception 
might reside in the dandelion head itself. Ta- 
naka et al. (1987) showed that some type of 
photoreceptor must occur in the dandelion 
head as its opening is dependent on sunlight. 
Since the majority of the scape growth is due 
to cell elongation in the upper third of the 


24 Journal of the Kentucky Academy of Science 62(1) 


scape (Chao 1947; Oxlade and Clifford 1981) 
it is possible that a growth regulator (like gib- 
bey //in, auxin, or ethylene) is synthesized in 
the cad and moves down to that region (Clif- 

ci and Oxlade 1989: Clifford et al. 1985) to 
sodulate tissue growth. This would explain 
he discrepancy he tween the excised-scape ex- 
periment and the whole-plant experiment. 
Some factor important to the phytochrome re- 
sponse in dandelion scapes may not have been 
present in the excised segments, 

This research has implications for turfgrass 
managers interested in reducing the propaga- 
tion of dandelion by seed. Since the majority 
of dandelion seed production and dissemina- 
tion is in the spring (Gray et al. 1973), the 
simple act of keeping turf taller during the 
preceding winter months might force dande- 
lion heads to elongate enough in the spring so 
that a larger percentage of them could be 
mowed before they set seed. The same sce- 
nario might be applied during the second flush 
of flowering that occurs in the autumn. This 
paper sets the groundwork for studies of these 
types of management strategies. 


ACKNOWLEDGMENTS 


The cooperation of Edie Greer and Karan 
Kaul at the Atwood Research Facility, Ken- 
tucky State University, is much appreciated. 
Thanks also go to students Scott Farmer, Mik- 
ki Jo eather and Anton Clemmons for their 
work on the dandelion project. Joann Lau and 
Snake Jones provided helpful suggestions. 
This research was funded by the Bellarmine 
University Faculty Summer Stipend Program. 


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J. Ky. Acad. Sci 26-34. 2001. 


Distri>ution and Status of Freshwater Mussels (Bivalvia: Unionoidea) 
in the Cumberland River Basin Upstream from 
Cumberland Falls, Kentucky 


Ronald R. Cicerello and Ellis L. 


Nature Preserves Commission, 801 Schenkel Lane, 


Kentucky State 


Laudermilk 


Frankfort, Kentucky 40601 


ABSTRACT 


Freshwater mussels were sampled in the Cumberland River basin upstream from Cumberland Falls in 


southeastern Kentucky in 1987-1999 to determine their distribution and status. A total of seven species was 


found at 57 of 434 sampling sites compared to 11 taxa previously reported. Alasmidonta atropurpurea and 


Anodontoides denigratus, USFWS- 


and/or KSNPC-endangered species, are the most abundant taxa and 


exclusively inhabit tributaries. Lampsilis ovata was introduced from below the falls and is extirpated from 


the basin along with Toxolasma parvus and Villosa lienosa. We believe that records for Strophitus undulatus 


are based on misidentified Alasmidonta atropurpurea and Anodontoides denigratus. The Marsh Creek fauna 


is the richest and most abundant upstream from the falls, and it should be the focus of mussel conservation 


efforts in the basin. 


INTRODUCTION 


Despite considerable interest in the fresh- 
water mussels of Kentucky’s upper Cumber- 
land River basin (e.g., Miller et al. 1984: Neel 
and Allen 1964; Gelneter et al. 1989: Wilson 
and Clark 1914), little has been published re- 
garding the fauna upstream from Cumberland 
Falls. Wilson and Clark (1914) and Neel and 
Allen (1964) each sampled six sites, all but one 
in the mainstem, and found a depauperate 
fauna ee eight generally uncommon spe- 
cies (Table 1). Recent examination of 14 Cum- 
berland River tributaries revealed mussels 
only in Marsh Creek and added Alasmidonta 
atropurpurea to the fauna (Call and Parmalee 
1981; Harker et al. 1979, 1980; Layzer and 
Anderson 1992). Museum records for Toxolas- 
ma parvus and Villosa lienosa increase the 
fauna to 11 species, or ca. 15% of the taxa 
known from the basin below Cumberland 
Falls (Cicerello et al. 1991). 

The basin above the falls has changed con- 
siderably since the work of Wilson and Clark 
(1914) and Neel and Allen (1964). Commu- 
nities and infrastructure have expanded, and 
surface and underground coal mines occur 
throughout the watershed (Leist et al. 1982). 
Development has degraded water quality and 
aquatic communities throughout the basin and 
downstream from Cumbedend Falls (Carter 
and Jones 1969; Cicerello and Peale 
1997: Harker al. 1979: KDOW 1996). 
Nonetheless, ie area contains many of Ken- 


et 


tucky’s highest quality streams (KDOW 1997), 
most of ach have not been thoroughly sam- 
pled for mussels. We examined streams in the 
upper Cumberland River basin of Kentucky 
upstream from Cumberland Falls and muse- 
um collections to determine the distribution 
and status of the mussel fauna. 


STUDY AREA 


Located in the Appalachian Plateaus Phys- 
iographic Province in southeastern Kentucky 
and Tennessee, the study area encompasses 
5120 km? of the upper Cumberland River ba- 
sin in Kentucky extending from Cumberland 

Falls es to the southent Virginia bor- 
der (Figure 1). Cumberland Falls is a 17 m 
high barrier to the upstream movement of 
aquatic organisms that has receded ca. 72 km 
from its hypothesized original location near 
Burnside, Kentucky (McGrain 1966). The Big 
South Fork Cumberland, Rockeastle, and 
Laurel rivers and Buck Creek formerly dis- 
charged into the river above the falls, but now 
they enter between its hypothesized original 
and present locations. The Cumberland River 
is formed by the confluence of the Poor and 
Clover forks; it meanders westerly 212 km to 
the falls. Major tributaries are Marsh, Jellico, 
Stinking, Straight, and Yellow creeks and 
Clear and Martins forks. The headwaters of 
Marsh and Jellico creeks and Clear Fork are 
in Tennessee. Most streams have moderate to 
high gradients, clear water, and alternating 


Mussels collected from the Cumberland River system upstream from Cumberland Falls, following prevailing nomenclature. 


Table 1. 


1s 


This study and 
museum record 


1987-1988* 


1978-19809 


1947-1949? 


1910-1911! 


A. pectorosa 


A. pectorosa 


Actinonaias pecterosa 


Alasmidonta atropurpurea 


A. viridis 


A, atropurpurea 
A. viridis 


A. atropurpurea 


Alasmidonta minor 


A. denigratus 
L. fasciola 


A. ferussacianus 
L. fasciola 


Anodontoides ferussacianus 


Freshwater Mussels—Cicerello and Laudermilk 7 


L. fasciola 


Lampsilis fasciola 


Lampsilis ovata 


L. cardium 


L. ventricosa 


L. o. ventricosa 


Strophitis edentulus 


Unio gibbosus 


E. dilatata 


E. dilatata 


E. dilatatus 


Elliptio dilatatus 


Toxolasma parvus 


Villosa lienosa 


Neel and Allen (1964); ° Harker et al. (1979, 1980) and Call and Parmalee (1981); * Layzer and Anderson (1992). 


1 Wilson and Clark (1914); 2 


pools and riffles underlain with substrate: 
ranging from exposed sandstone bedrock to 
mixed sand and silt. Mean annual discharge 
and 7-day 10-year low flow at Cumberland 
Falls are 89.7 m?/s and 0.65 m/s, respectively; 
major tributaries such as Jellico and Stinking 
creeks have summer low flows of zero (Ruhl 
and Martin 1991: USGS 1993). The watershed 
is mountainous and land use is ca. 84% forest, 
13% agriculture, 2.5% mining, and 0.5% ur 
ban and developed areas (MSE 1975). Wil- 
liamsburg, Barbourville, Pineville, Middles- 
boro, and Harlan, the largest communities in 
the basin, have a combined population of less 
than 30,000. Although water quality is im- 
proving, many streams continue to be impact- 
ed by pollutants associated with coal mining, 
domestic waste, highway construction, and 
poor land use (Harker et al. 1980: KDOW 
1996). 


MATERIALS AND METHODS 


We examined 434 upper Cumberland River 
basin sites for mussels in 1987-1999. All hab- 
itats at each site were sampled using a viewing 
bucket or while snorkeling during low flow 
when the water was clear. Mussel collections 
at the Academy of Natural Sciences of Phila- 
delphia (ANSP), Eastern Kentucky University 
(EKU), Harvard University (MCZ), National 
Museum of Natural History (NMNH), Ohio 
State University Museum of Zoology (OSU), 
and University of Michigan Museum of Zool- 
ogy (UMMZ) were examined. Species ac- 
counts are presented alphabetically following 
scientific names in Gordon (1995) and Tur- 
geon et al. (1998). Cumberlandian regional 
endemics and Cumberland River endemics 
(Gordon 1995; Gordon and Layzer 1989; Ort- 
mann 1924) are indicated. Each annotation in- 
cludes collection site numbers (Appendix A) 
followed in parentheses by the number of 
specimens collected (L = living specimen(s), 
F = freshly dead, R = relic) during each sam- 
pling visit. Results for sites visited more than 
once are in chronological order. Terms used to 
summarize distribution follow Smith (1965) 
and include “generally distributed” (any suit- 
able habitat should yield specimens with a rea- 
sonably thorough search), “occasional” (suit- 
able-appearing habitat may or may not yield 
specimens even after prolonged search), and 


28 Journal of the Kentucky Academy of Science 62(1) 


Richland 
55 Creek 
Big Sy : Stinking 
berland Indian Creek Poor Fork 
P Creek 
as ~ 
Patterson 33 57 
ones 34 st ie? — 
‘ ! eg . Clover Fork 
7, 
~— 3 
PP 5 
ST A 6 
4 ‘4 4 
4 
42 
4 
a 3 
ara 
oO 
; VA 
( Lo ee eee 
Marsh TN 
Creek N 
Y 
Jellico Clear Fork 0 Ki 
se 
Creek 47-50 
Figure 1. Freshwater mussel collection sites in the upper Cumberland River basin above Cumberland Falls, south- 


eastern Kentucky, 1987-1999. Site numbers are referenced in Appendix A. 


“sporadic” (encountering specimens cannot be 
predicted at all). 


SPECIES ACCOUNTS 

A brief discussion of the historical and pre- 
sent distribution and status of mussels report- 
ed upstream from the falls follows. Only gen- 
eralized distributional information is present- 
ed for Corbicula fluminea. 

Actinonaias pectorosa (Conrad). Pheas- 
antshell. Cumberlandian regional endemic. 
Sites: 1(0; 19L, 3R; 16L): 2(8L); 3(14F: 1L): 
4(1L); 5(7L); 6(1R); 7(1R); 8(1R); 9(1R); 
LO(UIF): 11(1L, 1R; 1L, 1R); 1301R; 0); 14(2 2/ 
2R; 0); 16(1R); 18(1F); 19(8L, 1R; 7L, 1F, 1B; 
41, 3F): 20(8L, LF): 21(3L); 2201F): 23(1L); 
46(1L). This is the most common species in 
the upper Cumberland River and lower Marsh 
Creek, where it is occasional to generally dis- 
tributed; it is sporadic in Clear Fork. It inhab- 
its sand among cobbles or boulders, and sand 
and coal fines in bedrock fractures. It was not 
collected by Wilson and Clark (1914), and 
Neel and Allen (1964) found only one speci- 
men (near Williamsburg) which “appeared to 
be a recent immigrant.” 

Alasmidonta atropurpurea (Rafinesque). 
Cumberland elktoe. Cumberland River en- 
demic. Sites: 19(5L, 3R: 0: 3L, 2R): 20(5L): 
DAA ATs): p22( Qiks) 23 (Qa)! SACL earl) 


26(2L); 27(2L); 28(4L); 29(4L, 5F); 31(8L, 
OF): 32(2L, IR): 331, 1E): 34S 
QF); 35(1L, 17F; 22L; several L; 5L; 42L); 
36(0; 1L); 37(2L; 0); 38(1F); 39(1F); 40(1F); 
47(0; 1L); 48(1L; 0; 0); 49(1L); 50(1L). Until 
recently, this mussel was synonymized under 
A. marginata (Call and Parmalee 1981; Clarke 
1981). It historically inhabited the Cumber- 
land River and its southern tributaries in Ken- 
tucky and Tennessee, including the Big South 
Fork of the Cumberland River, upstream from 
the hypothesized original location of Cumber- 
land Falls near Burnside, Pulaski County, 
Kentucky (Gordon 1991; Gordon and Layzer 
1993). It was not reported by Wilson and 
Clark (1914) or Neel and Allen (1964), but, as 
observed by Gordon (1991), historic records 
for A. marginata from the Cumberland River 
just above the falls, McCreary/Whitley coun- 
ties (UMMZ 63954), and at Williamsburg, 
Whitley County (Clarke 1981, MCZ 224076), 
are A. atropurpurea. Downstream from the 
present falls location, it occurred in the Laurel 
River, Laurel County, (as Strophitus undula- 
tus) (Neel and Allen 1964; UMMZ 172886), 
Lynn Camp Creek, Whitley County (Clarke 
1981), and in the Big South Fork of the Cum- 
berland River above Burnside, Pulaski County 
(as A. marginata) (Wilson and Clark 1914). 
Wilson and Clark’s (1914) record for A. mar- 


Freshwater Mussels—Cicerello and Laudermilk 29 


ginata from the Cumberland River near Burn- 
side could be this species. This endangered 
mussel (KSNPC 1996; USFWS 1998) is spo- 
radic in Laurel Fork of Clear Fork, and it is 
generally distributed and common in Marsh 
Creek, where it was discovered by Harker et 
al. (1980) and Call and Parmalee (1981). It 
also inhabits the Big South Fork of the Cum- 
berland River, Kentucky and Tennessee, the 
Rockcastle River, where it occurs sympatrical- 
ly with A. marginata (contra Gordon 1991: 
Gordon and Layzer 1993), and Rock Creek, 
Kentucky (Cicerello et al. 1991; Gordon 1991, 
EKU). It lives in sand and silt often among 
cobbles and boulders in relatively shallow 
pools and runs (Gordon and Layzer 1989). 

Alasmidonta viridis (Rafinesque). Slipper- 
shell mussel. Sites: 19(0; 0; IR); 22(1F); 
93(1F); 27(1L); 28(4L); 330L); 47(1L; 1L); 
48(0: 1R; 1R). Wilson and Clark (1914) col- 
lected specimens from the Cumberland River 
between the falls and Pineville and from the 
Clear Fork. Neel and Allen (1964) did not find 
this species and hypothesized that it was lost 
to acid coal mine drainage. This species is oc- 
casional in Marsh Creek, where it was discov- 
ered by Harker et al. (1980) and Call and Par- 
malee (1981), and it is sporadic in Laurel 
Fork, a Clear Fork tributary. In both streams, 
small numbers of this easily overlooked mussel 
inhabit sand or mixed sand, gravel, and silt 
near boulders. 

Anodontoides denigratus (Lea). Cumber- 
land papershell. Cumberland River endemic. 
Sites: 18(1R); 23(1L); 25(1L); 30(3L); 31(2L, 
3F); 32(1L, 1F); 33(1L, 1R); 34(2F, 2 3/2R; 
HME) 3535L, LE, 2R; 21F; 1L, 4 1/2F; 40L; 
73L, 3F); 36(9L; 6L); 37(15L, 3F; 1R); 38(1L, 
3F); 39(4F); 40(5L, 1F); 41(3L); 51(15L); 
52(8L, IR; 13L; 10L, 3F); 53(2L; 3L); 54(5L, 
IF, 2R; 18L, 1F); 55(10L, 1F, 1/2R; 8L); 
56(1R; 1L). This species was described by Lea 
(1852) and has been confused with A. ferus- 
sacianus and Strophitus undulatus. It is dis- 
tinctive from the allopatric A. ferussacianus 
according to Gordon (1995), who is redescrib- 
ing it. It is restricted to the upper Cumberland 
River on the Cumberland Plateau in Kentucky 
and Tennessee upstream from the original lo- 
cation of the falls (Gordon 1995). Historically, 
A. denigratus was collected from the Cum- 
berland River at Pineville and Orby, Bell 
County (Ortmann 1918; Wilson and Clark 


~ 


1914; UMMZ 105530) (as A. ferussacianus ). 
and from the Clear Fork (UMMZ 66141) (as 
S. undulatus). Downstream from the present 
location of the falls, it is known from Lynn 
Camp Creek, Whitley County (UMMZ 
105533) (as A. ferussacianus), and from the 
Laurel River, Laurel County (Neel and Allen 
1964: UMMZ 172886) (as S. undulatus). Wil- 
son and Clark’s (1914) record for A. ferussa- 
cianus from the Clear Fork, Jellico, Tennes- 
see, probably is this species (Gordon 1995). 
This species inhabits Marsh, Moore, Pine, and 
Rose creeks, Mills Fork, Billies Branch, 
Demps Hollow, and Rockcastle River tributar- 
ies in Kentucky (EKU, KSNPC), and Big 
South Fork of the Cumberland River tributar- 
ies in Tennessee (Gordon 1995). The best 
population of this endangered mussel 
(KSNPC 1996) is generally distributed in the 
middle segment of Marsh Creek, where it 
comprised 57% of 282 live mussels specimens 
collected in 1994. Its habitat is identical to that 
of A. atropurpurea, slowly flowing or still 

ools and runs underlain with silt and sand. It 
is locally abundant in Moore, Pine, and Rose 
creeks, Mills Fork, Billies Branch, and Demps 
Hollow, all first- and second-order streams in- 
habited only by A. denigratus. 

Elliptio dilatata (Rafinesque). Spike. Sites: 
SF(1F); 11(0; 1R); 19(5L, 1R; 1R; 1F); 20(1L, 
QF): 21(1L); 24(1R); 26(1L); 42(1R); 43(1R); 
44(1R); 45(1 1/2R). Historically, the spike was 
the most widely distributed and abundant spe- 
cies above the falls. Wilson and Clark (1914) 
collected specimens at all of their sampling 
sites and found that it comprised about 90% 
of the Clear Fork mussel population. In the 
late 1940s, it was the predominant species in 
the upper Cumberland (Neel and Allen 1964). 
Call and Parmalee (1981) considered the spike 
common in Marsh Creek, where Layzer and 
Anderson (1992) also noted its presence. The 
distribution and abundance of E. dilatata have 
declined greatly throughout the basin. It is 
sporadic in the Cumberland River, Jellico 
Creek, and Clear Fork, where we found main- 
ly relic specimens, and relatively common only 
in lower Marsh Creek. 

Lampsilis cardium Rafinesque. Plain pock- 
etbook. Sites: 3(1F; 0); 11(1R; 0); 12(1F); 13(3 
WIR BO) "14(O2 MEL) FSe/2R): SCE, awk): 
16(1R); 19(0; 1L, 1F; 1L, 1F); 20(1L); 22(1R); 
93(1L); 24(1R); 35(1L; 0; 0; 0; 1L); 45(2L, 3 


30 Journal of the Kentucky 


3/2R) The plain poc ‘ketbook is widely 


distribute a ue sporadic in the Cumberl: ind 
Ri rom the falls upstream to Harlan Coun- 
bh : in Marsh Creek, where it was collected 


lurker et al. (1980) and Call and Parmalee 

S1). It is sporadic in the Clear Fork, Whit- 

County. Marsh Creek supports the best 
population above the falls, but only seven liv- 
ing or freshly dead specimens were found at 
21 s: umpling sites in 1994. It forme rly was very 
abundant above and below the falls (Neel and 
Allen 1964), where Wilson and Clark (1914) 
failed to find specimens. The only ai we 
found from above the falls that pre-date Neel 
and Allen (1964) are an undated Bryant Walk- 
er collection from the Cumberland River at 
Williamsburg, Whitley County (MCZ 46747), 
and collections made in 1941 and 1945 by 
Clark from the Cumberland River at Molus, 
Harlan County (MCZ 123966), and 5 miles 
east of Pineville (UMMZ 165273), respective- 
ly. 
~ Lampsilis fasciola Rafinesque. Wavyrayed 
lampmussel. Sites: 1(1/2R: 0: 0): 9(1R): 10 
(1R): 11(1L, 2R; 1/2F): 12(2 F); 13 (4R; 1R); 
14(1 2/2R; 2L): 16(2/2F): 17(1L): 19(0; 1F, IR: 
2L); 20(1L); 22(1R); 46(1/2R); 48(0; 0; 2F); 
S7(1F). Although L. fasciola was not collected 
by Wilson and Clark (1914), Neel and Allen 
(1964) reported that it was very abundant. Call 
and Parmalee (1981) and Layzer and Ander- 
son (1992) collected specimens from Marsh 
Creek. Lampsilis fasciola is widely distributed 
but sporadic in the Cumberland River, Poor 
Fork, Marsh Creek, and Clear Fork, where it 
inhabits sand and/or sand, pebbles, and gravel 
often near cobbles or boulders in shallow 
pools or runs. 

Lampsilis ovata (Say). Pocketbook. Sites: 
none. Wilson and Clark (1914) collected a few 
dwarfed specimens from the Cumberland Riv- 
er and re ported that L. ovata was transplanted 
into the river above the falls, possibly by pearl 
collectors (Neel and Allen 1964). This appar- 
ent effort to develop a commercially valuable 
stock failed. Now considered endangered in 


Kentucky (KSNPC 1996), L. ovata apparently 


has not been collected subsequently; it is con- 
sidered extirpated from above the falls. 
Strophitus undulatus (Say). Creeper. Sites: 
none. Wilson and Clark (1914) reported spec- 
imens from the Cumberland River at Pine- 
ville, collected by their collaborator J. F. Boep- 


dilatatus, and A. pectorosa.” 


Academy of Science 62(1) 


ple, and from the Clear Fork, Tennessee. Neel 
and Allen (1964) listed it from the Cumber- 
land River at Pineville and at Wallins (near 
Harlan), but stated that “[iJn the present sur- 
vey only 4 forms occurred above the falls: L 
fasciola, L. ovata ventricosa (=L. cardium), E. 
We did not find 
S. undulatus and we were unable to locate 
specimens from the area for re-examination. 
Based on the following, we believe Wilson and 
Clark's (1914) records for S. undulatus from 
above the falls are actually A. denigratus. We 
re-identified two specimens of S. undulae col- 
lected by Hubbs from the Clear Fork, Whitley 
County, (UMMZ 66141) as A. denigratus. Wil- 
son and Clark (1914) and Nese and Allen 
(1964) also reported S. undulatus from the 
Rockcastle and Laurel rivers. Both S. undulatus 
and A. denigratus inhabit the Rockcastle River 
basin (KSNPC, MCZ), but we re-identified 
Neel and Allen’s (1964) S. undulatus speci- 
mens from the Laurel River at Lily (UMMZ 
172886) as A. denigratus and A. atropurpured. 
Wilson and Clark (1914) noted that their S. 
undulatus specimens were “exceedingly vari- 
able and presented many puzzling forms,” in- 
dicating difficulty in making identifications. Fi- 
nally, ae shell descriptions and the picture of 
S. undulatus in Wilson and Clark (1914) and 
Neel and Allen (1964) also could be inter- 
preted as A. denigratus. 

Toxolasma parvus (Barnes). Lilliput. Sites: 
none. Known from only one specimen col- 
lected by C. Goodrich from the Cumberland 
River northwest of Pineville, Bell County 
(UMMZ 99672). The lilliput can be over- 
looked because of its small size, but it proba- 
bly is extirpated. 

‘Villosa lienosa (Conrad). Little spectacleca- 
se. Sites: none. A specimen collected by H.D. 
Athearn from the Cumberland River near 
Barbourville, Knox County, (HDA 13857) in 
1966 is the only record of this rare Kentucky 
mussel (KSNPC 1996) from above the falls. 
This record is interesting because Richland 
Creek, a nearby tributary, lies in close prox- 
imity to C ollins Fork, a Kentucky River trib- 
utary that supports one of Kentucky's largest 
V. lienosa populations (Cicerello pers. obs.). 

Geological evidence and fish distribution pat- 
terns identify this area as a stream capture 
theater, with probable multiple faunal ex- 
changes between the drainages during recent 


Freshwater Mussels—Cicerello and Laudermilk 3] 


geological times (Burr and Warren 1986: 
Kuehne and Bailey 1961). Stream capture oc- 
curs when natural erosion cuts across a head- 
water drainage divide and a stream segment 
and its biota are diverted from one basin into 
another. Villosa lienosa is absent, extirpated, 
or rare in the adjacent Tennessee, Cumber- 
land (below the falls), and Big Sandy rivers 
(Beetle 1973; Cicerello et al. 1991; Starnes 
and Bogan 1988), and it is tempting to invoke 
stream capture and transfer of its fish host 
from the Kentucky River drainage as the 
source of this highly localized record. How- 
ever, even the most suggestive stream capture 
evidence may lead to invalid conclusions, and 
limited distributions may result from intro- 
ductions rather than natural factors (Jenkins et 
al. 1971). The origin of V. lienosa is unclear; 


the species probably is extirpated. 
INTRODUCED SPECIES 


Corbicula fluminea (Miiller). Asian clam. 
This introduced, exotic clam is the most com- 
mon and widely distributed mussel above the 
falls. It inhabits the mainstem from the falls 
to near the headwaters of the Poor Fork, the 
lower half of Marsh Creek, Jellico Creek near- 
ly to the Tennessee border, Clear Fork up- 
stream into Mud Creek and Laurel Fork, Big 
Indian Creek including Mills Fork, lower 
Stinking Creek, and Martins Fork upstream to 
Martins Fork Lake. Densities exceed 100/m? 
in Marsh Creek. 


DISCUSSION 


We found a total of seven species, all living 
or freshly dead, at 57 of the 434 sites sampled 
(Table 1). With the exception of Marsh Creek 
and other streams inhabited by A. denigratus, 
mussels generally are sporadic and restricted 
to the mainstem Cumberland River and to the 
Clear and Poor forks. Species richness is 
greatest in Marsh Creek (7 species), Clear 
Fork (6), and the mainstem Cumberland River 
(4), and several streams support only one spe- 
cies (e.g., Big Indian, Jellico, Patterson creeks, 
and the Poor Fork). Lampsilis fasciola is the 
most widely distributed native species, inhab- 
iting the Cumberland River from the falls up- 
stream ca. 216 river km into Poor Fork, Marsh 
Creek, and Clear Fork. 

Three species found mainly in tributaries 
comprised ca. 92% (613/667) of the living or 


freshly dead specimens encountered. Anodoi- 
toides denigratus, the most abundant specic: 
is restricted to Marsh Creek, Clear Fork, and 
segments of Big Indian, Patterson, Richland, 
and Stinking creeks. Alasmidonta atropurpu- 
rea is found primarily in Marsh Creek, as is 
A. pectorosa, which lives also in the Cumber- 
land River from the falls to above Yellow 
Creek. Elliptio dilatata and, to a lesser extent, 
L. cardium and L. fasciola, formerly were the 
dominant species in the mainstem Cumber- 
land River (Neel and Allen 1964; Wilson and 
Clark 1914). Their decline in abundance is a 
result of persistent and varied water quality 
problems (Charles 1966; Jillson 1927; KDOW 
1996, 1998). 

Eleven species have been reported above 
the falls, but only as many as nine are native 
to the area. Specimens previously reported as 
S. undulatus probably were mis-identified A. 
atropurpurea and A. denigratus. Lampsilis 
ovata was introduced into the basin (Wilson 
and Clark 1914). Toxolasma parvus and V. 
lienosa each were collected only once from the 
basin and are considered extirpated from 
there. 

Marsh Creek is the most important refuge 
for mussels in the upper Cumberland River 
basin. It is the most species-rich, and it sup- 
ports the best populations of all mussels ex- 
cept A. pectorosa. More than 70% (471/667) 
of all living and freshly dead specimens we en- 
countered were found in Marsh Creek. The 

resence of Phoxinus cumberlandensis 
(USFWS (1998) threatened species) and Eth- 
eostoma nigrum susanae (KSNPC (1996) 
threatened species), upper Cumberland River 
basin endemic fishes, increase the importance 
of Marsh Creek as an epicenter for recoloni- 
zation of degraded streams throughout the ba- 
sin. 

We note several challenges to the protection 
of the biological diversity and integrity of 
Marsh Gres that also threaten other basin 
streams. Marsh Creek has relatively good wa- 
ter quality, but it is being impacted by silt 
from farms and inactive coal strip mines 
(KDOW 1996). A National Resources Conser- 
vation Service (formerly the Soil Conservation 
Service) proposal to remove the silt by chan- 
nelizing upper Marsh Creek was rebuffed, but 
this proposal could resurface despite recom- 
mendations to identify and revegetate eroding 


32 Journal of the Kentucky Academy of Science 62(1) 


areas. | reviously mined areas also could be re- 
mine (o obtain formerly inaccessible or un- 
pro!able coal deposits. The development of 
oi) sources in uplands along the stream pos- 


renewed threat to the biota. A 1987 oil 
oill killed hundreds of A. atropurpurea and 
\. denigratus along an undetermined length 
/! Marsh Creek (Cicerello pers. obs.). Finally, 
a 1965 United States Army Corps of Engi- 
neers proposal to construct a 24+ m high dam 
on the Cumberland River 1.6 km upstream 
from Cumberland Falls was revived in 1995 
by local entities interested in hydroelectric 
generation and recreation. This project would 
embay the Cumberland River upstream into 
lower Marsh Creek and adversely impact the 
aquatic biota and water quality. 


ACKNOWLEDGMENTS 

Thanks to M. Evans, R. R. Hannan, M. 
Mays, S. E. McMurray, B. Palmer-Ball Jr., M. 
A. Patterson, D. Peak, M. Thomas, and B. 
Winters (former or present KSNPC) and R. 
G. Biggins (USFWS) for field assistance; H. 
D. Athearn (HDA), A. J. Baldinger (MCZ), E. 
Hartowicz, D. O’Foighil (UMMZ), and G. A. 
Schuster (EKU) for access to their collections, 
data, or specimens; and C. Moore (KDOW) 
and J. Kiser for sharing the Pine Creek record. 
Our work was supported in part by the Ken- 
tucky Department for Surface Mining Recla- 


mation and Enforcement, Frankfort, Ken- 
tucky. 

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781:1-63. 


APPENDIX A 


Upper Cumberland River basin mussel col- 
lection sites in southeastern Kentucky listed 
by and within sub-basins from down- to up- 
stream. Site numbers refer to numbers in Fig- 
ure l. 

CUMBERLAND RIVER MAINSTEM: 
(1.) Just above Cumberland Falls. McCreary/ 
Whitley cos. 28 Apr, 10 Sep 1987; 14 Sep 
1995. (2.) Between Cumberland Falls and 
Ryans Branch. McCreary/Whitley cos. 30 Jul 
1993. (3.) At Marsh Creek. McCreary/Whitley 
Cos. 3 Jul, 10 Aug 1993. (4.) Between Buck 
Shoals and Crow creeks. McCreary/Whitley 
cos. 10 Aug 1993. (5.) At Summer Shoals. 
Whitley Co. 29 Jul 1993. (6.) At Rough Shoals 
Creek. Whitley Co. 27 Jul 1994. (7.) At Inter- 
state 75. Whitley Co. 27 Jul 1994. (8.) KY 296 
at Williamsburg. Whitley Co. 27 Jul 1994. (9.) 
At Whetstone Creek. Whitley Co. 28 Jul 1994. 
(10.) Ca. 1 km downstream from Big Indian 
Creek. Knox Co. 30 Sep 1994. (11.) Ca. 1.9 
km upstream from Stinking Creek. Knox Co. 
14 Jul, 22 Sep 1993. (12.) At Pineville. Bell 
Co. 5 Aug 1993. (13.) Ca. 1.6 km downstream 
from KY 119. Bell Co. 2 Aug 1990; 29 Sep 
1994. (14.) KY 1344 at Calvin. Bell Co. 10, 23 
Sep 1993. (15.) At KY 987. Bell Co. 23 Sep 
1993. (16.) At Minton Branch. Bell Co. 29 Jul 
1994. (17.) At Fourmile Branch. Harlan Co. 8 
Jun 1994. MARSH CREEK: (18.) Ca. 0.8 km 
upstream from mouth. McCreary Co. 10 Aug 
1994. (19.) Ca. 0.2 km downstream from 
Brushy Creek. McCreary Co. 16 Aug 1989; 9, 
18 Aug 1994. (20.) At Hens Nest Creek. 
McCreary Co. 10 Aug 1994. (21.) Ca. 2.7 km 
upstream from Hens Nest Creek. McCreary 
Co. 11 Aug 1994. (22.) Ca. 3.3 km upstream 
from Hens Nest Creek. McCreary Co. 11 Aug 
1994. (23.) At trib. 3.6 km upstream from 
Hens Nest Creek. McCreary Co. 10 Aug 1994. 
(24.) At tributary ca. 1 km downstream from 
KY 679. McCreary Co. 3 Aug 1994. (25.) At 
KY 679. McCreary Co. 15 Aug 1989. (26.) At 
tributary ca. 0.5 km downstream from KY 679. 
McCreary Co. 3 Aug 1994. (27.) Ca. 0.4 km 
upstream from KY 679. McCreary Co. 3 Aug 
1994. (28.) Ca. 0.5 km downstream from Lau- 
rel Creek. McCreary Co. 8 Aug 1994. (29.) Ca. 


34 Journal of the Kentucky Academy of Science 62(1) 


1 km downstream from Duck Run, McCreary 
Co. 9 ‘ng 1994. (30.) Ca. 0.2 km downstream 
fron: jaylor Branch. McCreary Co, 3 Aug 
199-1. (31.) At KY 478. McCreary Co. 30 Aug 


los7. (32.) Ca. 0.3 km upstream from KY 478. 
McCreary Co. 11 Aug 1994. (33.) At Big 
Branch. McCreary Co. 12 Aug 1994. -(34.) 
Downstream from Kidd School Road Ford. 
McCreary Co. 15 Aug 1989; 4 Aug 1994. (35.) 


Ca. 0.3 km upstream from Kidd School Road 
Ford. McCreary Co. 30 Aug 1987; 28 Apr 
1993; 12 Jul 1994: 4, 17 Aug 1994. (36.) Ca. 


0.5 km upstream from Kidd er Road 
Ford. McCreary Co. 4, 17 Aug 1994. (37.) Ca. 
0.7 km downstream from KY 1044. Morea 
Co. 2, 12 Aug 1994. (38.) Ca. 0.2 km down- 
stream from KY 1044. McCreary Co. 2 Aug 
1994. (39.) At KY 1044. McCreary Co. 18 Jul 
1989. (40.) Ca. 0.1 km upstream from KY 
1044. McCreary Co. 2 Aug 1994. (41.) Ca. 0.2 


km downstream from C eae Creek. McCreary 


Co. 2 Aug 1994. JELLICO CREEK: (42.) Ca 
4.4 km downstream from KY 92. Whitley Co. 
6 Jul 1994. (43.) Ca. 2.0 km downstream from 
KY 92. Whitley Co. 6 Jul 1994. (44.) Down- 
stream from Shut-in Branch. McCreary Co. 12 


May 1993. CLEAR FORK: (45.) At Tackett 
Creek. Whitley Co. 7 Jul 1994. (46.) At Buck 
Creek. Whitley Co. 7 Jul 1994. (47.) Laurel 
Fork at TN border. Whitley Co. 27 Jul 1993; 
21 Sep 1996. (48.) Laurel Fork ca. 0.4 km up- 
stream from TN ce Whitley Co, 27 Jul 
1993; 11 Jan 1994; 26 Sep 1996. (49.) Laurel 
Fork ca. 0.7 km upstream from TN border, 
Whitley Co, 27 Jul 1993. (50.) Laurel Fork ca. 
0.9 km upstream from TN border. Whitley Co. 
27 Jul 1993. (51.) Pine Creek at KY 190. Bell 
Co. 28 ee 1999. PATTERSON CREEK: 
(52.) Rose Creek ca. 0.2 km upstream from 
sao: Creek. Whitley Co. 16 Jun, 15 Oct 
1993; 22 May 1997. BIG INDIAN CREEK: 
(53.) Demps Hollow ca. 1.3 km upstream 
from Big Indian Creek. Knox Co. 21 Jul 1993, 
20 May 1997. (54.) Mills Fork ca. 0.8 km up- 
stream from Big Indian Creek. Knox Co, 21 
Jul 1993, 21 May 1997. RICHLAND CREEK: 
(55.) Billies Branch ca. 3.2 km upstream from 
Richland Creek. Knox Co. 22 Jul 1993, 20 
May 1997. STINKING CREEK: (56.) Moore 
Creek ca. 10.5 km NW Pineville. Knox Co. 28 
Jan 1993, 21 May 1997. POOR FORK: (57.) 
0.6 km upstream from Middleton Branch. 
Harlan Co. 24 May 1994. 


J. Ky. Acad. Sci. 62(1):35-38. 2001. 


Morphometric Variation of Cotton Mice (Peromyscus gossypinus) anc. 
White-footed Mice (P. leucopus) in Kentucky 


Nell A. Bekiares' and George A. Feldhamer 
Department of Zoology, Mail Code 6501, Souther Illinois University, Carbondale, Illinois 62901-6501 


ABSTRACT 


We captured 151 white-footed mice (Peromycus leucopus) and 38 cotton mice (P. gossypinus) in Ballard 
and Carlisle counties, Kentucky, during 3600 trap nights. There were significant differences between the 
two species in body mass, hind foot length, condylobasal length, and length of the nasal bone for both adult 
males and females. Morphological characteristics often used to differentiate the two species were not always 
sufficient to do so accurately. Large white-footed mice may be misidentified as cotton mice. 


INTRODUCTION 


Cotton mice occur in southeastern United 
States from eastern Texas and Oklahoma east 
to Florida and north to Virginia (Jones and 
Birney 1988). With regard to body size, the 
largest of the three commonly recognized sub- 
species, Peromyscus gossypinus megacephalus 
(Rhoads 1894), is found at the northern pe- 
riphery of the range. Cotton mice are uncom- 
mon in Kentucky, Missouri (Hall 1981), and 
the southernmost five counties of Illinois 
(Feldhamer et al. 1998: Hoffmeister 1989). 
The preferred habitat of cotton mice, “coin- 
cident with the location of rivers, streams, and 
other lowland areas” (McCarley 1963:787), in- 
cludes swampy woodlands, bottomlands, low- 
land forests, and sites near swamps, sloughs, 
oxbow lakes, and areas with high water tables 
(Goodpaster and Hoffmeister 1952; Laerm 
and Boone 1994; Linzey et ak 1976; H. 
McCarley 1954a, 1954b, 1963; W.H. Mc- 
Carley 1964; Pournelle 1952). Woody debris is 
used extensively (McCay 2000). 

Cotton mice are sympatric throughout 
much of their range with white-footed mice 
(P. leucopus). The two species presumably di- 
verged recently (Hooper 1968) and can be dif- 
ficult to distinguish in the field. The purpose 
of our study was to compare morphometric 
characteristics of cotton mice from western 
Kentucky, where the species is considered to 
be threatened, with sympatric white-footed 
mice. 


! Present address: 7 East Lakeshore Drive #23, Cincin- 
nati, OH 45237. 


35 


METHODS 


Live trapping occurred from August 1998 
through April 1999. Twelve sites were selected 
in Ballard and Carlisle counties, Kentucky 
(Bekiares 2000). Two Sherman live traps were 
set at each station, with stations established 10 
m apart along a 500-m transect. Traps were 
set at two sites each week and checked be- 
tween 0600 and 1000. Traps were set close to 
fallen logs, brush piles, stumps, pond edges, 
tree trunks, and on floating debris whenever 
possible to optimize trap success for P. gos- 


sypinus. Traps were baited with cracked corn 


and sunflower seeds and were set for three 
consecutive nights at each site, for a total of 
300 trap nights per site. During summer, traps 
exposed to sunlight were covered with leaves 
to decrease the amount of heat absorbed by 
the trap prior to checking. During cold tem- 
peratures, traps contained polyester fiberfill 
bedding material. 

The sex, age class (juvenile or adult, deter- 
mined by pelage color), and wet body mass 
(nearest g) of captured animals were recorded. 
Tchednels were then checked for a previous 
capture mark. If the animal was new, a hind 
foot measurement (mm) was taken. New cap- 
tures were marked with a green permanent 
marker along the ventral surface (Schmid 
1998). Marks could be observed for the du- 
ration of the three trap nights at each site. 

Because P. leucopus and P. gossypinus are 
morphologically similar and difficult to distin- 
guish in the field, initial size criteria of Hoff- 
pent (1977, 1989) were used to separate 
the species. Individuals with hind foot length 
=22 mm or body mass =26 ¢ were tentativ ely 
identified as PB. gossypinus. These animals 


36 Journal of the Kentucky Academy of Science 62(1) 


were © thanized using cervical dislocation, 
placed on ice, and returned to the laboratory. 
Allovyme analyses (Bekiares 2000) were used 
to confirm species identification of animals 


ollected. Other rodents, birds, and reptiles of 


ion-interest were released at the capture site. 

For animals removed from the field, stan- 
dard external measurements (in mm) were 
made prior to dissection: total body length, a 
length, ear length, and hind foot length. In 
addition, wet body mass (g) was re ae da 
second time using a triple- beam balance. 
Measurements (mm) of cleaned skulls includ- 
ed condylobasal length, length of the nasal 
bone, and le ngth of Te maxillary toothrow. 

The computer program Statview was used 
to compute unpaired t-tests for comparisons 
of means, Z-tests for comparisons of propor- 
tions, and general descriptive statistics. Statis- 
tical tests were considered significant at a = 
0.01. 


RESULTS 


During 3600 trap nights we captured 197 
fadividueales 151 white-footed mice, 38 cotton 
mice, three rice rats (Oryzomys palustris), one 
Eastern chipmunk (Tamias striatus), and four 


leopard frogs (Rana sphenocephala). Seven of 


the P. leucopus were believed to be P. gossy- 
pinus upon capture and were removed from 
the field. 

Only data from adults were used in mor- 
phological analyses because of the differences 
in juvenile sizes. Another consideration is that 
females collected in late summer and autumn 
may be pregnant. We collected only six preg- 
nant females and chose to remove them from 
analyses. As expected, adult P. leucopus were 
significantly smaller than P. gossypinus for 
four of the eight measurements examined. For 
males, P. leucopus were significantly smaller 
than P gossypinus in body mass, hind foot 
length, con lobasal length, and nasal length 


(Table 1 Female P leucopus were eaiilier: 
than ee P. gossypinus for the same four 
measurements. 


DISCUSSION 
Most of the mean values for morphological 
characteristics of white-footed mice in our 
study represent a biased sample. Only those P. 
leucopus tentatively identified as P. gossypinus 
in the field were used for all measurements 


and represented the largest individuals. Only 
the hind foot and body mass measurements 
represent an unbiased sample because data 
were obtained on all animals captured in the 
field, not just those presumed to be P. gossy- 
pinus. We expect that the other four charac- 
teristics measured (total length, tail length, ear 
length, and maxillary toothrow length) also 
would be significantly smaller in an unbiased 
sample of P le UCOPUS. 

Cotton mice in this study were significantly 
larger than six specimens reported from 
Horseshoe Lake Conservation Area, Alexan- 
der County, Illinois (Feldhamer et al. 1998). 
It is possible that the Ilinois specimens were 
large P. leucopus misidentified as P. gossypi- 
nus. More likely, we suspect they may have 
been hybrids between the two species. Bar- 
bour and Davis (1974) suggested hybrid cot- 
ton mice occurred in Kentucky. Other inves- 
tigators have also noted hybridization between 
the two species (Lovecky et al. 1979: Mc- 

Carley 1954b; St. Romain 1976; although see 
Bradshaw 1968), with hybrids exhibiting inter- 
mediate-sized mor hological characters. 

Boone (1995) analyzed morphometric data 
for cotton mice from throughout their range. 
Using only adults, we compared his data to 
our data from Kentucky. For all morphological 
traits considered in both studies, Kentucky 
specimens, from the periphery of the range, 
were significantly larger (P < 0.01; see Beki- 
ares 2000). This is consistent with the clinal 
size relationship noted by Boone (1995) for 
cotton mice throughout their range. 

Hoffmeister (1977, 1989) created a scatter- 
gram based on morphological characters to 
distinguish between cotton mice and white- 
footed mice. He used hind foot length X nasal 
bone length on the x-axis, and condylobasal 
length X maxillary toothrow length on the y- 
axis. Measurements of cotton mice group to 
the right of a line running approximately 
through (0, 134) and (275, 0). In our study, 
specimens “on the line” were white-footed 
mice, based on allozyme data (Bekiares 2000). 

Use of morphological measurements may 
need to be more conservative for differenti- 
ating white-footed mice and cotton mice on 
the periphery of their range in Kentucky, Il- 
linois, and Missouri. That is, “questionable” 
specimens on or near the right side of the de- 


Cotton Mice and White-footed Mice—Bekiares and Feldhamer 


Table 1. Differences (t-test; * = P < 0.01) in mean values of morphological characteristics between specimens o 
adult Peromyscus gossypinus and P. leucopus from Ballard and Carlisle counties, Kentucky, — collected between Augus 


1998 and April 1999. Variation is reported as standard error. 


Characteristic P. gossypinus 
Male body mass (g) 30.97 + 1.20 
n= 17 
Female body mass (g) 33.18 + 1.04 
n = 2) 
Male total length 170.82 + 3.61 
n= 17 
Female total length WOSy = lll 
n= 21 
Male tail length TA 47 + 2.53 
m= i 
Female tail length 77.90 = 1.03 
n = 21 
Male hind foot length WOT 22 O27 
m = If 
Female hind foot length 92.81 = 0.20 
n= 21 
Male ear length 18.76 = 0.31 
n= 17 
Female ear length 18.86 = 0.51 
n= 21 
Male condylobasal length 98.05 = 0.27 
n=17 
Female condylobasal length 98.39 + 0.14 
n= 21 
Male nasal length 11.41 + 0.21 
n= 17 
Female nasal length 11.22 + 0.16 
n = 21] 
Male maxillary toothrow 3.94 + 0.08 
n= 17 
Female maxillary toothrow 3.98 + 0.06 
n= 21 


P. leucopus T., value 
22.95 + 0.45 7.65* 
n= 057 
24.69 + 1.01 = 5 (a2 
im = 3 
159.00 + 5.21 1.49 
n=4 
174.00 += 7.77 1.42 
n=3 
66.25 + 206 1.52 
n= 4 
82.67 += 5.04 —1.48 
ne 
17.62 + 0.27 8.98* 
n = 68 
18.01 = 0.33 —9.34* 
n = 30 
18.00 + 0.41 1.14 
n=4 
17.00 + 3.06 1.09 
n=3 
25.56 + 0.59 4.00* 
n=4 
927.03 + 0.55 3.34% 
n=3 
9.61 + 0.28 3.99* 
n=4 
10.03 = 0.21 2.75 
n=3 
3.77 = 0.19 0.93 
n=4 
3.73 + 0.02 1.62 
n=3 


I 


marcation line of Hoffmeister (1977, 1989) 
should be considered P. leucopus, 


ACKNOWLEDGMENTS 


We thank D. Kevin Davie of the SIUC 
Morris Library GIS lab, and Valerie Barko and 
Dr. J. Scheibe for assistance in the field. This 
paper represents a portion of a thesis submit- 
ted by N. Bekiares for the degree of Master 
of Science in the Zoology Department at 
Southern Illinois University at Carbondale. 


LITERATURE CITED 


Barbour, R. W., and W. H. Davis. 1974. Mammals of Ken- 
tucky. Univ. Kentucky Press, Lexington, KY. 

Bekiares, N. 2000. Morphometric and allozyme variation 
in the cotton mouse (Peromyscus gossypinus) in south- 
em Illinois, southwestern Kentucky, and southeastern 


Missouri. M.S. Thesis. Southern Illinois Univ., Carbon- 
dale, IL. 


Boone, J. L. 1995. Morphological and genetic variation in 
the cotton mouse (Peromyscus gossypinus ): implications 
for population genetics, systematics, and conservation. 
Ph.D. Dissertation. Univ. Georgia, Athens, GA. 

Bradshaw, W. N. 1968. Progeny from experimental mating 
tests with mice of the Peromyscus leucopus species 
group. J. Mammal. 49:475-480. 

Feldhamer, G. A., J. C. Whittaker, and E. M. Charles. 
1998. Recent records of the cotton mouse (Peromyscus 
gossypinus) in Illinois. Am. Midl. Naturalist 139:17S— 
180. 

Goodpaster, W. W., and D. F. Hoffmeister. 1952. Notes 
on the mammals of western Tennessee. J. Mammal. 33: 
362-371. 

Hall, E. R. 1981. The mammals of North America, 2nd 
ed. John Wiley and Sons, New York, NY. [2:601—-1175} 

Hoffmeister, D. F. 1977. Status of the cotton mouse, Per- 
omyscus gossypinus, in southern Illinois. Am. Midl. 
Naturalist 97:222—224. 

Hoffmeister, D. F. 1989. Mammals of Illinois. Univ. Illi- 
nois Press, Urbana, IL. 


38 Journal of the Kentucky Academy of Science 62(1) 


Hooper. |. T. 1968. Classification. Pages 27-74 in J. A. 
Kins d). Biology of Peromyscus (Rodentia). Spec. 
Publ. Am. Soc. Mammal. 2. 

Jones, J. K., Jr, and E. C. Bimey. 1988. Handbook of the 

unmals of the north-central states. Univ. Minnesota 
Press, Minneapolis, MN. 

iaverm, J., and J. L. Boone. 1994, Mensural discrimination 
of four species of Peromyscus (Rodentia: Muridae) in 
the southeasterm United States. Brimleyana 21:107— 
123. 

Linzey, A. V., D. W. Linzey, and S. E. Perkins, Jr. 1976. 
The Peromyscus leucopus group in Alabama, J. Alabama 
Acad. Sci. 47:109-113. 

Lovecky, D. V., D. Q. Estep, and D. A. Dewsbury. 1979. 
Copulatory behaviour of cotton mice (Peromyscus gos- 
sypinus) and their reciprocal hybrids with white-footed 
mice (P. leucopus). Anim. Behay, 27:371-375. 

McCarley, H. 1954a. The ecological distribution of the 
Peromyscus leucopus species group in eastern Texas. 

McCarley, H. 1954b. Natural hybridization in the Pero- 
myscus leucopus species group of mice. Evolution 8: 
314-323. 


McCarley, H. 1963. Distributional relationships of sym- 
patric populations of Peromyscus leucopus and Pero- 
myscus gossypinus. Ecology 44:784-788. 

McCarley, W. H. 1964. Ethological isolation in the ceno- 
species Peromyscus leucopus. Evolution 18:331—342. 
McCay, T. S$. 2000. Use of woody debris by cotton mice 
(Peromyscus gossypinus) in a southeastern pine forest. 

J. Mammal. $1:527-535. 

Pournelle, G. H. 1952. Reproduction and early post-natal 
development of the cotton mouse, Peromyscus gossy- 
pinus gossypinus. J. Mammal. 33:1-20. 

Rhoads, S$. N. 1894. Descriptions of four new species and 
two new subspecies of white-footed mouse from the 
United States and British Columbia. Proc. Acad. Nat. 
Sci. Philadelphia 46:253-261. 

Schmid, $. 1998. The impact of patch characteristics on 
small mammal fauna: Peromyscus leucopus and associ- 
ated species in previously cut forest patches. Ph.D. Dis- 
sertation. Southern Illinois Univ., Carbondale, IL. 

St. Romain, P. A. 1976. Variation in the cotton mouse 
(Peromyscus gossypinus) in Louisiana. Southwest. Nat- 
uralist 52:290-300. 


J. Ky. Acad. Sci. 62(1):39-51. 2001. 


Woody Plants of Six Northern Kentucky Counties 


Ross C. Clark 
Department of Biological Sciences, Eastern Kentucky University, Richmond, Kentucky 40475 


and 


Ryan M. Bauer 


Department of Computer Information Science, Eastern Kentucky University, Richmond, Kentucky 40475 


ABSTRACT 


Our field work and herbarium surveys have documented the woody flora of Bracken, Fleming, Harrison, 
Mason, Nicholas, and Robertson counties, Kentucky. The presently known woody flora consists of 172 taxa; 
11% of the woody flora is exotic. We report Thuja orientalis L. for the first time as an introduced member 
of Kentucky's flora. There is a significant discontinuity of distribution between the northern Bluegrass and 
Knobs regions. Arundinaria gigantea is a much rarer plant in northern Kentucky than conventional wisdom 
indicates. We include an annotated listing of woody plant taxa and a discussion of the state of natural habitats 


and potential impact of weedy woody species. 


INTRODUCTION 


Narratives related to the natural features of 
Kentucky have typically attested to an abun- 
dance and variety of natural resources. A re- 
cent, statewide assessment of biodiversity in 
Kentucky (Taylor 1995) convincingly argues 
that plant diversity in the state is substantial. 
However, in spite of recent attempts to esti- 
mate vascular plant diversity in Kentucky 
(Browne and Athey 1992; Medley 1993), suf- 
ficient information is not yet available for an 
accurate estimate of which vascular plant taxa 
actually occur in the state. Contributing rea- 
sons for this situation include repeated anec- 
dotal and unvouchered reports of occurrences 
and a general paucity of vouchered collections 
in institutional herbaria (Jones et al. 1995). 

However, the situation we find ourselves in 
is beside the point. The point is, too little is 
known about the flora of Kentucky, land use 
trends are continually shrinking and modifying 
native habitats, and we are at risk of losing 
elements of the flora before we discover the 
full extent of its diversity and distribution. 
There is an urgent need for more thorough 
documentation of the flora of Kentucky. 


SCOPE AND JUSTIFICATION OF 
THIS STUDY 


Our study was undertaken to document the 
occurrence of woody plants that are native, 
naturalized, or spreading from cultivation in 


39 


six counties of northern Kentucky: Bracken, 
Fleming, Harrison, Mason, Nicholas, and 
Robertson. Our goal was to document the 
woody plant diversity in these counties at 
greater than the 90% level. 

One of the reasons this region was selected 
for study is because it is not well known bo- 
tanically. Guetig (1993) compiled a summary 
of all floristic work previously done in Ken- 
tucky. His summary indicated that none of the 
counties included in the present study has 
ever been the subject of an organized effort 
to document the flora. Also, we felt that thor- 
ough vouchering of the woody plants of these 
counties would make subsequent field work to 
voucher the herbaceous vascular plants easier. 
(The logistics of field collecting are simpler 
when one is not continually searching for all 
sizes of plants.) In addition, woody plants were 
selected as the focus of our study because they 
are keystone organisms in forest and savanna 
ecosystems and have considerable economic 
significance. We felt that documenting the oc- 
currence of this portion of the flora would be 
a connection to Kentucky's heritage that peo- 
ple would be more likely to appreciate. Final- 
ly, most of this area of the state is typified by 
an advanced state of habitat elimination and 
modification. Consequently, we concluded 
that, if efforts to document the flora were de- 
layed much longer, significant elements of the 
woody flora could possibly be lost entirely. 


40 Journal of the Kentucky 


THE STUDY AREA 


A’ of the counties lie within the drainage 
of Licking River, except for the northern 
Mason and Bracken counties, 
hich are drained by creeks downcutting di- 
‘ctly to the Ohio River. The six county study 
totals almost 1700 mi2 (438,000 ha). 
which is almost 4.3% of the land area of Ken- 
tucky (Anonymous 2000a). 

Elevations within the study area range from 
about 500 ft. (152 m) above mean sea level 
near Meldahl Dam on the Ohio River in 
northwestern Bracken County, to about 1200 
ft. (370 m) above sea level in extreme eastern 
Fleming County. The maximal elevation at- 
tained in the Bluegrass part of our study area 
is about 1000 ft. (305 m), in extreme south- 
western Harrison, near Leesburg (U.S. Geo- 
logical Survey 1976, 1978). 

All of the study area except eastern Fleming 


tions of 


ea 


County lies within the Bluegrass Region of 


Kentucky. The Bluegrass is atdertain by Or- 
dovician strata. Main geologic formations in- 
clude Lexington Limestone where exposed by 
downcutting of main tributaries of the w estern 
and northern Licking River, with Garrard Silt- 
stone and the Clays Ferry Formation com- 
prising most of the rolling uplands of the west- 
em part of the study area. Proceeding east- 
ward from the former exposures, one encoun- 
ters relatively narrow outcropping belts of the 
(Ordovician) Bull Fork Formation, Drakes 
Formation, and Crab Orchard Formation 
(both of these Silurian and marking the tran- 
sition between Knobs and Bluegrass), and fi- 
nally, Devonian and Mississippian strata of the 
extreme western edge of the Mississippian 
Plateau. (McDowell et al. 1981) Only extreme 
eastern Fleming County is within the physio- 
graphic province known as The Knobs, char- 
een’ (sequentially as one goes eastward) 
by outcropping Silurian, Devonian, and Mis- 
sissippian rocks. The Knobs are interpreted as 
the dissected remnant of the Mississippian 
(Interior Low Plateau) Province, known pa- 
rochially in Kentucky as the Pennyrile Plateau 
(Fenneman 1938: Karan and Mather 1977). 
Except over heavily organic Ohio shale in the 
Knobs, most upland sails of our study region 
have an alkaline pH. 

Estimates of the original vegetation in this 
region vary and are stall debated, possibly be- 


Academy of Science 62(1) 


cause originally there was a complex mosaic 
from hich fire has now been systematically 
excluded for 200 years. It may have included 
a preponderance of ash-oak-prairie savanna in 
the Inner Bluegrass and various facies of oak- 
chestnut and mixed mesophytic on the Knobs 
(Braun 1950; Bryant 1987; Campbell 1980; 
Kiichler 1964; Martin et al. 1978). 


MATERIALS AND METHODS 


Prior to field collecting, each county was 
surveyed to determine areas which might yield 
more habitat diversity. Because of the relative 
lack of geologic div ersity in most of the study 
area, this surve y relied on physiographic maps 
and county road maps (Anonymous 1997; 
Puetz n.d.; U.S. Geological Survey 1976, 
1978). By this method, we were able to iden- 
tify circumstances with a high likelihood of 
giving us efficient and repeated coverage of 
major habitats within each county. 

About two field collecting days were spent 
in each county. Field work began on 7 Jun 
1999 and ended 20 Jul 1999. Specimens were 
collected wherever we found them first, from 
roadsides and ruderal habitats to reasonably 
intact habitats. Specimens were temporarily 
held in wet newspaper within large polyeth- 
ylene bags and later accessioned by collector, 
pressed, dried, labeled, and determined to tax- 
on. The senior author determined the taxa of 
most specimens without consulting additional 
sources, except to verify nomenclatural au- 
thorship. The field work required 2441 miles 
of travel, an average of about 3 miles (4.8 km) 
for each taxon Bollecred! 

Nomenclature in this report is generally ac- 
cording to Gleason and Cronquist (1991), 
which also was a source consulted for deter- 
mination of some groups. The International 
Plant Name Index (Anonymous 2000b) and 
Rehder (1940) were consulted in a few cases, 
and we used Fernald (1950) as an aid to the 
determination specimens of specimens of Cra- 
taegus and Rubus. 

Following processing and determination of 
our field Bollesnons herbarium collections at 
EKY, UKY, MDKY, and KNK were surveyed 
to see if additional taxa had been vouchered 
in these counties by other collectors. These 
surveys yielded a few (<1%) additional re- 
cords. Travel to herbaria required 3 additional 
work days and about 500 miles (800 km) of 


Woody Plants—Clark and Bauer Al 


Table 1. 
Bracken Fleming 
Area!? 495! 351! 
Total number of taxa 109 140 
Number of native taxa 92, 124 


Number of non-native taxa 15 16 
% Non-native 
Human density/mi? ! 


‘in sq. mi. 
? Source: Anonymous 2000. 


additional travel. See Table 1 for statistical flo- 
ristic data. 


RESULTS 


The field effort resulted in more than 760 
collections from the study area. Our subse- 
quent regional herbarium survey yielded a few 
additional records. Overall, this study resulted 
in the documentation of 172 woody taxa from 
the six-county area. Fleming County, with 140 
documented taxa, has the most diverse woody 
flora. Robertson County, with 97 documented 
taxa, has the least diverse woody flora (Table 
1). Information from the Internet is cited here 
in accordance with the standard proposed by 
Walker (1995). 


ANNOTATED VOUCHERS OF THE 
WOODY FLORA 


Below is a list of vouchers for the woody 
plants of the six-county study area, followed 
by citations of voucher specimens and a short 
comment on the pattern of occurrence. To 
save space, names of counties of origin of 
voucher specimens (Bracken, Fléming, Har- 
rison, Mason, Nicholas, Robertson) are abbre- 
viated to first letter in parentheses after 
voucher numbers. When only a number is list- 
ed, it is the accession number of the senior 
author and the specimen has been deposited 
in EKY. Vouchers from other Kentucky work- 
ers are more completely cited. Nomenclature 
generally follows Gleason and Cronquist 
(1991), but in the final analysis reflects the 
judgment of the senior author. Abbreviations 
for herbaria are from Holmgren and Holm- 
gren (2000). 

Little (1971, 1977) included some addition- 
al distributional data for a few taxa within our 
study area. Presumably, the vouchers on which 
some of his data are based are in herbaria not 
consulted during this study. Therefore, his 


Comparative woody plant diversity data for six northern Kentucky counties. 


Harrison Mason Nicholas Robertson Totals 
309! QA)} 196! 100! 1692! 
110 100 96 97 2, 

95 82 78 §3 142 
15 18 18 14 19 
13.6 18.0 18.9 14.4 11 
52.6 40 34.3 22.2, 


vouchers are not referenced here. We conser- 
vatively estimate that the specimens collected 
during our study constitute more than 95% of 
the data now extant for woody plants in this 
six-county area of Kentucky. 


PINOPHYTA 
Cupressaceae 


Juniperus virginiana L.—24915 (B), 25273 
(a), Daley (aD), Dele TOM ie Wiallse! (i) 
25038 (R). Common throughout. This is ap- 
parently the only gymnosperm native in the 
Bluegrass. 

Thuja orientalis L.—25057 (R). Mature off- 
spring from nearby cultivated plants are es- 
tablished in a fencerow. Introduced; This is 
apparently the first report of plants of this 
taxon escaping in Kentucky (Brown and 
Athey 1992; Medley 1993). 


Pinaceae 


Pinus echinata Miller—25244 (F). Occasional, 
Knobs. 

Pinus rigida Miller—25227 (F). Frequent, 
Knobs. 

Pinus virginiana Miller—25275 (F). Common, 
Knobs. 


ANTHOPHYTA 

Aceraceae 

Acer negundo L.—24869 (B), 25282 (F), 
94628 (H), 24788 (M), 25147 (N), 25011 
(R). Common throughout. 

Acer nigrum Michaux f—24957 (B), 25322 
(F), 24611 (H), 24799 (M), 25091 (N), 
25024 (R). Mesic woods, usually on north- 
and east-facing slopes or in alluvium along 
streams. 

Acer rubrum L.—25310 (F). Frequent in var- 
ious habitats, Knobs. 


42 Journal of the Kentucky Academy of Science 62(1) 


Acer saccharinum L.—24956 (B), 25342 (F), 
94606 (H), 24770 (M), 25136 (N), 24964 
(i). Frequent; low ground along streams 

nd terraces. 

er saccharum Marshall—24901 (B), 25232 
F), 24651 (H), 24796 (M). 25128 (N). 
25020 (R). Frequent; various upland habi- 
tats. 


Anacardiaceae 
Rhus aromatica L.—25327 (F), 24613 (H). 
Xeric blufftops; rare. 

Rhus copallina L.—25277 (F), 24648 (H), 
25077 (R). Disturbed rights-of-way; rare. 
Rhus glabra L.—24912 (B), 25271 (F), 24644 

(H), 24831 (M), 25123 (N), 25029 (R). Var- 
ious habitats, most common in fencerows. 
Rhus typhina L.—24832 (B), 24774 (M). Me- 

sic woods and low ground; infrequent; con- 
fined to habitats near the Ohio River. 
Toxicodendron radicans (L.) Kuntze—24946 
(B), 25281(F), 24597 (H), 24756 (M), 25151 
(N), 25079 (R).Various habitats; common. 
Toxicodendron pubescens Miller—24874 (B), 
24707 (H). Xeric woods: rare. 


Annonaceae 

Asimina triloba (L.) Dunal—24889 (B), 25313 
(F), 24646 (H), 24758 (M), 25139 (N), 
24990 (R). Various mesic and alluvial habi- 
tats; frequent. 


Apocynaceae 

Vinca minor L.—25218 (F), 25047 (H), 24802 
(M), 25047 (R). Introduced: ruderal habi- 
tats and dump sites; infrequent. 


Araliaceae 

Aralia spinosa L.—25286 (F). Mesic and al- 
luvial habitats: Knobs. 

Hedera helix L.—24850 (B). 
dump site; rare. 


Introduced: 


Berberidaceae 

Berberis thunbergii DC.—24867 (B), 25048 
(H), 25094 (N), 25048 (R). Introduced. Var- 
ious disturbed and degraded habitats; infre- 
quent. 


Betulaceae 


Alnus serrulata (Aiton) Willd.—Thieret s.n.. 
30 Jan 1983 (KNK) (B), 25235 (F), 24807 


(M). Streambanks and low, open ground; in- 
frequent; apparently absent from Bluegrass. 

Betula nigra L.—25338 (F). Alluvial woods, 
rare; Knobs. 

Carpinus caroliniana Walter—24924 (B), 
25307 (F), 24701 (H), 24754 (M), 25095 
(N), 24991 (R). Various habitats, usually 
mesic: frequent to occasional. 

Corylus americana Walter—24879 (B), 25289 
(F), 24694 (H), 25025 (R). Bluffs and mesic 
habitats; infrequent. 

Ostrya virginiana (Miller) K. Koch—24909 
(B), 25203 (F), 24617 (HA). 24746 etre: 
25096 (N): common. 


Bignoniaceae 

Bignonia capreolata L.—25324 (F), 24619 
(H), 25025 (R). South- and west-facing 
bluffs along major streams; rare, but locally 
common. 

Campsis radicans (L.) Seemann—24897 (B), 
25241 (F), 24653 (H), 24798 (M), 25146 
(N), 25015 (R). Most habitats; common. 

Catalpa bignonioides Walter—24837 (M), 
25132 (N). Low woods, rare. In spite of its 
common name (southern catalpa), this spe- 
cies appears to be native along the Ohio 
River in the study area. However, both this 
species and the following one are cultivated 
and can be expected to produce adventive 
plants. 

Catalpa speciosa Warder—24586 (H). An ap- 
parent escape from cultivation; Cynthiana. 


Caesalpiniaceae 

Cercis canadensis L.—24903 (B), 25255 (F), 
24681 (H), 24767 (M), 25170 (N), 25039 
(R). Various habitats; common throughout. 

Gleditsia triacanthos L—24917 (B), 25207 
(F), 24661 (H),. 24755 (M), 325145 CK 
25002 (R). Various habitats; common 
throughout. 

Gymnocladus dioica (L.) K. Koch—24836 (B), 
24332 (F), 24624 (H), 24721 (M), 25160 
(N), 25073 (R). Disturbed habitats and me- 
sic woods; infrequent. 


Caprifoliaceae 

Lonicera japonica Thaunberg—24098 (B), 
25314 (F), 25052 (Hl), 253580 (Mi)s aailian 
(N), 25054 (R). Introduced; most habitats; 
common. 


Lonicera maackii (Rupr.) Herder—24940 (B), 


Woody Plants—Clark and Bauer AS 


95251 (F), 24632 (H), 24772 (M), 25166 
(N), 24976 (R). Introduced; most habitats; 
common in counties bordering the Ohio 
River. In our opinion, this is one of the po- 
tentially most ecologically disastrous terres- 
trial plants ever introduced into eastern 
North America. It eliminates or severely in- 
hibits the reproduction of most native East- 
ern Deciduous Forest plants. 

Sambucus canadensis L.—24833 (B), 25252 
(F), 24695 (H), 24778 (M), 25174 (N), 
24969 (R). Stream corridors, low ground, 
and mesic woods; common. 

Symphoricarpos orbiculatus Moench—24885 
(B), 25215 (F), 24664 (H), 24810 (M), 
95116 (N), 25033 (R). Various habitats; fre- 
quent. 

Viburnum acerifolium L.—25309 (F). Decid- 
uous woods; Knobs. 

Viburnum prunifolium L.—24937 (B), 25221 
(Eee 5051 (Eee 24747, (MD) 251020) (IN), 
25058 (R). Upland woods, blufftops, fence- 
rows, and disturbed habitats; frequent. 

Viburnum rufidulum Raf—24894 (B), 25336 
(F), 24682 (H), 24764 (M), 25124 (N), 
25036 (R). Upland woods and disturbed 
habitats; frequent. 


Celastraceae 


Celastrus scandens 1..—24914 (B), 25315 (F), 
24671 (H), 24829 (M), 25140 (N), 25037 
(R). Upland and mesic woods and disturbed 
habitats; common. 

Celastrus orbiculatus Thunberg—25339 (F), 
25104 (N). Introduced; ruderal habitats and 
rights-of-way. Presently uncommon in the 
study area and often confused with plants 
of the preceding taxon. In our area, this spe- 
cies may have the potential of eclipsing 
most other introduced woody plants in its 
eventual negative ecological impact. 

Euonymus alatus (Thunberg) Siebold—25193 
(N). Introduced; fencerow; rare, but with 
the potential of becoming much more com- 
mon. 

Euonymus atropurpureus Jacquin—24873 
(B), 25206 (F), 24670 (H), 24750 (M), 
25153 (N), 24992 (R). Upland woods and 
disturbed habitats, common in fencerows; 
frequent. 

Euonymus fortunei (Turez.) Hand.-Maz.— 
24852 (B); 25049 (H), 24777 (M), 25109 
(N), 25049 (R). Introduced; old dump sites 


and also establishing adventively (spread | 
birds). Not yet common in the study area 
but soon will be. When fully established. 
supplants the entire herbaceous stratum of 
deciduous forest. 


Clusiaceae 


Hypericum stragulum Adams and Robson 
(= Ascyrum hypericoides var. multicaule of 
some authors)—25276 (F). Other woody 
hypericums in Kentucky apparently are ab- 
sent from the study area; Knobs only. 


Cornaceae 

Cornus amomum Miller var. schuetzeana 
(Meyer) Rickett—24839 (B), 25234 (F), 
24809 (M). During this study, located only 
on the banks of the Ohio River and in Big 
Run Swamp, Fleming County. 

Cornus drummondii C.A. Meyer—24878 (B), 
25329 (F), 24643 (H), 25100 (N), 25001(R). 
Xeric woods, disturbed habitats, and fence- 
rows; common. 

Cornus florida L.—24890 (B), 25292 (F), 
24703 (H), 24739 (M), 25090 (N), 25023 
(R). Mesic woods and bluffs; widespread 
but nowhere common; being decimated by 
fungal pathogens (Harlow et al. 1996). 

Nyssa sylvatica Marshall—25299 (F), 24636 
(H), 24830 (M), 25068 (R). Mesic woods; 
occasional to rare in study area, except for 
Knobs of Fleming County. 

Nyssa sylvatica Marshall var. biflora (Walter) 
Sargent—25226 (F). Rare; Big Run Swamp. 


Ebenaceae 

Diospyros virginiana L.—24921 (B), 25262 
(F), 24600 (H), 24732 (M), 25129 (N), 
25006 (R). Frequent in fencerows; also 
widespread but not common in mesic 
woods. 


Elaeagnaceae 

Elaeagnus umbellata Thunberg—25250 (F), 
94702, (H), 24791 (M), 25107 (N), 24972 
(R). Introduced; disturbed habitats: infre- 
quent. 


Ericaceae 


Gaylussacia baccata (Wangenh.) K. Koch— 
Meijer, Setser, and Meade 1150 = MDKY 
#4630 (F). Xeric woods, infrequent; Knobs. 


44 Journal of the Kentucky Academy of Science 62(1) 


Kalmia latifolia L.—25 
rare: Knobs. 

Oxydendrum arboreum (L.) DC.—25303 (F). 
| pland woods; Knobs. 


345 (F). Upland woods, 


95344 (F). 
land Wane fifequcnt ‘Kuobs only. 
Vaccinium pallidum Aiton—25270 (F). Same 
habitats and localities as Gaylussacia bac- 

cata. 
Vaccinium stamineum L.—25266 (F 
woods, frequent; Knobs only. 


Up- 


Upland 


Fabaceae 

Amorpha fruticosa L.—24841 (B), 24808 (M). 
In our study area, apparently confined to 
the banks of the Ohio River. 

Lespedeza bicolor Turez.—25205 (F), 25186 
(N). Introduced; escaping from plantings in 
state-managed wildlife management areas. 

94920 (B), 25295 

(F). 24633 (Hi 24781. (M), 25172. (N). 

25080 (R). Very common in varied habitats 

and disturbed areas. 


Fagaceae 

Castanea mollissima Blume—W. Meijer, 15 
Sep 1974, UKY #35813 (F). Introduced. 

Fagus grandifolia Ehrhart—24854 (B), 25230 
(F), 25089 (N), 25013 (R). Ravines and me- 
sic slopes; rare in Bluegrass, common only 
in Knobs. 

Quercus alba L.—24899 (B), 25311 (F), 
94660 (H), 24713 (M), 25110 (N), 25004 
(R). Mesic woods; common throughout. 

Quercus bicolor Willd.—25201 (F). Upland 
swamp, rare; Knobs. 

Quercus coccinea Muenchh.—24856 (B), 
25265 (F). Extremely rare in Bluegrass; we 
discovered a population of only two mature 
trees in a pasture fencerow in Bracken 
County and noted no reproduction. Very 
common in xeric and submesic woods in the 
Knobs. 

Quercus imbricaria Michaux—24934 (B); 
25229 (F): 24588 (H), 24820 (M), 25177 
(N), 24966 (R). Widespread but rare 
throughout our study area. 

Quercus macrocarpa Michaux—25320 (F), 
24623 (H), 24823 (M), 25162 (N), 25056 
(R). Frequent in the south but progressively 
more rare northward in the area we studied. 

Quercus montana Willd. ( 
some authors )—25287 (F). Xeric south- and 


=Q. prinus L. of 


west-facing slopes and ridges; frequent in 
Knobs. The type of Q. prinus in the Lin- 
naean herbarium may be of the lowland 
chestnut oak (=Q. michauxii Nuttall) in- 
stead of the upland chestnut oak, and the 
two names have been confused in the lit- 
erature for many years (Hardin 1979). Since 
QO. michauxii and QO. montana, respectively, 
are unambiguous names, it is better to use 
them instead. 

Quercus muhlenbergii Engelm.—24880 (B), 
95195 (F), 24656 (H), 24757 (M), 25143 
(N), 25041 (R). Upland habitats throughout; 
frequent. 

Quercus palustris Muenchh.—24947 (B), 
95258 (F), 24698 (H), 24789 (M), 25106 
(N). Frequent only along the Ohio River 
and in Fleming County upland swamps; 
otherwise sporadic and occasionally escap- 
ing from cultivation. 

Quercus rubra L.—24906 (B), 25288 (F), 
24696 (H), 25102 (N), 24986 (R). Mesic up- 
lands, usually on east- and north-facing 
slopes; occasional to rare. 

Quercus shumardii Buckley—24870 (B), 
95294 (F), 24631) (Bl) °24734( Miao 
(N), 25012 (R). Various habitats; common. 

Quercus stellata Wangenh.—24871 (B), 25264 
(F), 24666 (H), 24825 (M), 24977 (R). Up- 
land woods and fencerows; occasional. 

Quercus velutina Lamarck—24910 (B), 25278 
(F). 24649 (EH), 247116 (Mi) 2a: 
25069 (R). Upland woods; frequent. 

Quercus Xwilldenowiana Zabel—24692 (H). 
A hybrid, found in a fencerow, between Q. 
velutina Lam. and Q. falcata Michaux. We 
were unable to find the latter parent in the 
vicinity; however, considering the advanced 
state of habitat loss, it easily could have for- 
merly occurred nearby. 


Grossulariaceae 


Ribes missouriense Nuttall ex Torrey and 
Gray—25357 (N). Canebrake, rare. 


Hamamelidaceae 


Hamamelis virginiana L.—25294 (F). Various 
wooded habitats; Knobs. 

Liquidambar styraciflua L.—25259 (F). Vari- 
ous wooded habitats; Knobs. Plants of this 
taxon we noted along the Ohio River ap- 
peared to have been planted. 


Woody Plants—Clark and Bauer £5 


Hippocastanaceae 

Aesculus flava Aiton—24835 (B); 25223 (F); 
24814 (M). Confined to ravines and north- 
facing slopes along the Ohio River and its 
direct tributaries; also in mesic woods of 
Knobs. 

Aesculus glabra Willd.—24855 (B), 25196 (F), 
94614 (H), 24722 (M), 25173 (N), 25021 
(R). Wooded slopes, disturbed habitats, and 
fencerows; common. 


Hydrangeaceae 

Hydrangea arborescens L.—24927 (B), 25291 
(F), 24720 (M), 25176 (N), 25061 (R). Nor- 
mally confined to mesic east- and north-fac- 
ing bluffs along major streams; infrequent. 

Philadelphus coronarius L.—25243 (F). Intro- 
duced; established and spreading on old 
roadcut; rare. Most other adventive popu- 
lations of this genus in Kentucky probably 
should be referred to this taxon. 


Juglandaceae 

Carya cordiformis (Wangenh.) K. Koch— 
94887 (B), 25202 (F), 24649 (H), 24763 
(M), 25142 (N), 25031 (R). Most wooded 
habitats; the most common hickory in our 
study area. 

Carya glabra (Miller) Sweet—24891 (UB). 
25269 (F), 24676 (H), W. Meijer, 13 Jun 
1969, UK #33801 (M); 25017 (R). Upland 
woods; infrequent to rare. 

Carya laciniosa (Michaux f.) Loud—24898 
(B), 25328 (F), 24668 (H), 24712 (M), 
25125 (N), 25014 (R). Various wooded hab- 
itats; frequent. 

Carya ovata (Miller) K. Koch—24892 (B), 
94635 (H), 25125 (N), Thieret 52747 in 
KNK (R). Infrequent to rare; most common 
near the border between Bluegrass and 
Knobs in Fleming County. Plants of this tax- 
on in our region do not fit well the recent 
description given by Stone (1997). The ap- 
parent reason is that, throughout most of 
central Kentucky, introgression may be oc- 
curring between plants of this taxon and the 
following one. 

Carya tomentosa (Poiret) Nuttall—24883 (B), 
95222 (F), 24640 (H), 24824 (M), 25087 
(N), 24989 (R). Upland woods and woods 
remnants; frequent to occasional. 

Carya ovata X Carya tomentosa—24733 (M). 
As mentioned above, introgression between 


these two taxa is common in our region. 
producing various degrees of intermediacy. 
This is a collection that appeared to us to 
be truly intermediate, so we cite it as an 
unnamed hybrid. 

Carya pallida (Ashe) Engler and Graebner— 
24860 (B), 25112 (N), 25062 (R). Upland 
woods and woods margins; infrequent. 

Juglans nigra L.—24896 (B), 25280 (F), 
24677 (H), 24730 (M), 25148 (N), 24923 
(R). Various habitats; common. We did not 
observe any specimens of J. cinerea L.., liv- 
ing or dead, in the study area. 


Lauraceae 

Lindera benzoin (L.) Blume—24876 (B), 
25293 (F), 24650 (H), 24762 (M), 25088 
(N), 25019 (R). Various disturbed, degrad- 
ed, and intact habitats; one of the most 
common shrubs in this area of Kentucky. 

Sassafras albidum (Nuttall) Nees—22419 (B), 
25285 (F), 24647 (H), 24718 (M), 25114 
(N), 25027 (R). Various habitats; common. 


Magnoliaceae 

Liriodendron tulipifera L.—24853 (B), 25305 
(F), 24627 (H), 24717 (M), 25018 (R). Me- 
sic woods; rare in Bluegrass, progressively 
more common eastward. 

Malvaceae 

Hibiscus syriaca L.—24761 (M). Introduced; 


escaped along roadside; rare, but to be ex- 
pected more commonly in the future. 


Menispermaceae 

Menispermum canadense L.—24877 (B), 
95246 (F), 24592 (H), 24760 (M), 25164 
(N), 24998 (R). Various habitats, more fre- 
quent in highly disturbed situations, such as 
fencerows and rights-of-way; common. 


Mimosaceae 

Albizia julibrissin Durazzini—25216 (F). In- 
troduced; possibly persistent after cultiva- 
tion, or possibly an escape. 


Moraceae 


Broussonetia papyrifera (L.) Ventenat—24719 
(M). Introduced; an escape into highly dis- 
turbed habitat. 

Maclura pomifera (Raf.) C.K. Schneider— 
24859 (B), 25337 (F), 24684 (H), 24768 


46 Journal of the Kentucky Academy of Science 62(1) 


(M). 25156 (N), 25076 (R). Various dis- 
turbed habitats; common. 

Morus alba L.—24948 (B), 25319 (F), 24669 

H), 24818 (M), 25182 (N), 25078 (R). In- 
troduced; roadsides and other disturbed 
habitats; occasional. 

Vlorus rubra L.—24904 (B), 25290 (F), 24679 
(H), 24779 (M), 25161 (N), 25008 (R). Me- 
sic slopes and alluvial terraces; infrequent, 
but more common eastward. 


Oleaceae 


Chionanthus virginica L.—25300 (F). Upland 
woods; occasional: Knobs. 

Fraxinus americana L.—24902 (B), 25296 (F), 
24667 (H), 2481] (M), 25171 (N), 25007 
(R). Woodlands and woodland remnants; 
common. 

Fraxinus pennsylvanica Marshall—24949 (B), 
25204 5B)" 2AbS6CH). 24723" (MI) 25137 
(N), 25060 (R). Alluvial terraces and stream 
margins; Common. 

Fraxinus quadrangulata Michaux—24938 (B), 
25325 (F), 24618 (H), 25141 (N), 24983 
(R). Various wooded habitats but most com- 
mon on south- and west-facing xeric wood- 
lands and stream bluffs: frequent. 

Ligustrum sinense Louriero—24851 (B), 
24602 (H), 24804 (M). Introduced; escaped 
to waste places, highly disturbed habitats 
and alluvial woods. Most common near the 
Ohio River. 

Ligustrum vulgare L.—24951 (B), 24785 (M), 
25190 (N). Introduced; escaped to highly 
disturbed habitats. Most common in the 
Ohio River corridor. 


Platanaceae 

Platanus occidentalis L.—24941 (B), 25253 
(F), 24630 (H), 24787 (M), 25149 (N), 
25044 (R). Most common in low ground, 
but also in other situations; frequent. 


Poaceae 


Arundinaria gigantea (Walter) Chapman 
25355 (N). Wooded ravine; extremely rare. 


Ranunculaceae 

Clematis virginiana L.—24954 (B), 25257 (F), 
24700 (H), 24742 (M), 25175 (N), 25070 
(R). Low ground along streams and wood- 
land margins; infrequent. 


Rhamnaceae 


Rhamnus caroliniana Walter—25197 (F), 
24743 (M). Deciduous woods and disturbed 
rights-of-way. Very rare in Bluegrass, infre- 
quent in Knobs. 

Rhamnus lanceolata Pursh—25200 (F), 24693 
(H). Only two populations noted: one on a 
wooded alluvial terrace (Bluegrass), the oth- 
er in highly disturbed upland habitat (Blue- 
grass-Knobs border). 


Rosaceae 


Amelanchier arborea (Michaux f.) Fernald— 
25297 (F). Deciduous woods, occasional: 
Knobs. 

Crataegus calpodendron (Ehrh.) Medikus— 
24863 (B), Meijer, Setser, and Meade 1190 
= MDKY #1190 (F), 24594 (H), 25097 (N), 
25063 (R). Upland roadsides, blufftops, and 
woodland margins; infrequent. 

Crataegus coccinea L.—25211 (F), 24612 (H), 
25121 (N). Woodland margins and dis- 
turbed habitats; infrequent. 

Crataegus crus-galli L—25238 (B), 24685 
(F), 24812 (M), 25126 (N). Upland dis- 
turbed sites and woodland margins; infre- 
quent. 

Crataegus flabellata (Bosc) K. Koch—25208 
(F), 24590 (H), 25113 (M), 25064 (R). Up- 
land disturbed sites and woodland margins; 
infrequent. 

Crataegus mollis (Torrey and Gray) Scheele— 
24863 (B), 24678 (H), 24817 (M), 25158 
(N). Mainly in fencerows and on blufftops; 
infrequent. 

Crataegus pruinosa (Wendland) K. Koch— 
24862 (B). Disturbed woodland; rare. 

Malus angustifolia (Aiton) Michaux—25341 
(F), 25118 (N). Only two populations noted: 
open field (Bluegrass); low woods along 
creek (Knobs). Rare. 

Malus coronaria (L.) Miller—24932 (B), 
25247 (F), 24683 (H), 24815 (M). Upland 
woodland margins and fencerows; rare. 

Malus sylvestris (L.) Miller (=M. pumila of 
some authors)—25318 (F), 24765 (M), 
25152 (N), 24974 (R). Introduced; fence- 
rows and other ruderal habitats; infrequent. 

Physocarpus opulifolius (L.) Maxim.—G.F. 
Buddell Il #2206, in KNK (B). 

Prunus americana Marshall—24857 (B), 
25245 (F), 24589 (H), 24714 (M), 25144 


Woody Plants—Clark and Bauer AT 


(N), 24984 (R). Forest remnants, disturbed 
woods, and fencerows; frequent. 

Prunus cerasus L.—25219 (F). Introduced; 
adventive, adjacent to road right-of-way. 
Prunus mahaleb L.—24935 (B), 24690 (H), 
94745 (M), 25133 (N), 25034 (R). Intro- 
duced; roadcuts and rights-of-way; infre- 

quent. 

Prunus mexicana S. Watson—24905 (B), 
24639 (H), 24715 (M), 25122 (N), 24981 
(R). Forest remnants, disturbed woods, and 
fencerows; frequent. 

Prunus munsoniana Wight and Hedrick— 
25214 (F). Ridgetop in deciduous woods, 
Mississippian Plateau (Knobs); rare. 

Prunus persica (L.) Batsch—25213 (F), 24687 
(H), 24806 (M), 25178 (N), 24963 (R). In- 
troduced; fencerows and other ruderal hab- 
itats; infrequent. 

Prunus serotina Ehrhart—24919 (B); 25304 
(F), 24637 (H), 24775 (M), 25169 (N), 
95042 (R). Most habitats; common. 

Pyrus calleryana Decne.—24884 (B), 25209 
(1B), DANS) (TEDL BAS (MD), SHI UN) 
25059 (R). Introduced; roadsides, margins 
of blufftop woods; infrequent. 

Rosa carolina L.—24858 (B), 25274 (F), 
24659 (H), 24735 (M), 25099 (N), 25009 
(R). Upland deciduous woods and glades; 
frequent. 

Rosa multiflora Thunberg—24916 (B), 25283 
(F), 24658 (H), 24786 (M), 25159 (N), 
95035 (R). Introduced; most habitats, com- 
mon. At present, the worst woody weed in 
our study area. ‘ 

Rosa setigera Michaux—24895 (B), 24591 
(H), 24741 (M), 25092 (N), 25075 (R). 
Fencerows, low rights-of-way; common. 

Rosa virginiana Miller—24680 (H). This is ap- 
parently the first report of this species from 
the Bluegrass (Browne and Athey 1992). 
However, because Medley (1993) rejected 
the notion that the species occurs in Ken- 
tucky, this collection actually may represent 
the first verified occurrence of the species 
in the state. Fencerow; rare. 

Rosa wichuraiana Crepin—24953 (B), 24601 
(H), 24822 (M). Introduced; roadsides and 
fencerows; infrequent, but undercollected. 

Rubus allegheniensis T.C. Porter—25308 (F). 
Various habitats, Knobs. 

Rubus argutus Link—24881 (B), 25346 (F), 


94675 (H), 24803 (M), 25150 (N), 2499 
(R). Various habitats; common. 

Rubus flagellaris Willd. (incl. R. ensleni 
Tratt. )—24634 (H), 24995 (R). Deciduous 
woods; infrequent, but undercollected. 

Rubus occidentalis L—24866 (B), 25248 (F), 
94674 (H), 24784 (M), 25165 (N), 24994 
(R). Upland and lowland woods and various 
disturbed habitats; frequent. 

Rubus. trivialis Michaux—25331 (F), 25083 
(R). Fencerows; rare. 

Spiraea tomentosa L.—Meijer, Setser, and 
Meade #1223, in MDKY (F). Wooded up- 
land swamp. 


Rubiaceae 

Cephalanthus occidentalis L.—24944 (B), 
25236 (F). Margin of Ohio River backwater; 
upland swamp; rare. 


Rutaceae 

Ptelea trifoliata L.—25321 (F), 24607 (H), 
25185 (N). Bluffs along major streams; 
common in habitat but absent elsewhere. 

Zanthoxylum americanum Miller—24931 (B), 
95198 (F), 24641 (H), 24737 (M), 25085 
(N), 25000 (R). Xeric bluffs and slopes, usu- 
ally south- or west-facing; common in hab- 
itat, absent elsewhere. 


Salicaceae 

Populus alba L.—24865 (B), 25194 (F), 24626 
(H), 24725 (M), 25098 (N), 25065 (R). In- 
troduced; spreading from cultivation at old 
home sites, fencerows, and ruderal habitats; 
occasional. 

Populus deltoides Bartr. ex Marshall—24840 
(B), 25340 (F), 24686 (H), 24795 (M), 
24973 (R). Low ground; occasional to rare, 
nowhere common. 

Populus grandidentata Michaux—25242 (F). 
Old roadcut; rare. Apparently absent from 
Bluegrass area. 

Salix alba L.—25108 (N). Introduced; estab- 
lished along Brushy Creek. 

Salix caroliniana Michaux—24848 (B). Back- 
water of Ohio River, apparently absent from 
other parts of study area. 

Salix discolor Muhl—J. Campbell, 16 May 
1992, UKY (F). 

Salix exigua Nuttall—24846 (B), 25240 (F), 
24706: (H), 24797 (M), 25135 (N), 24971 
(R). Open stream corridors; common. 


48 Journal of the 


Salix nigra Marshall—24942 (B), ee i (F), 
94705 (H), 24821 (M), 25135 (N), 34971 
(R). Open stream corridors; Regen al. 

Salix sericea Marshall—25261 (F). Open, up- 
‘and swamp; rare. 


Simar¢ yubaceae 


Ailanthus altissima (Miller) Swingle—24955 
(B), 25212 (F), 24604 (H), 24780 (M), 
25130 (N), 25082 (R). Introduced: dis- 
turbed habitats (mature trees; seedlings are 
found in almost all habitats). Occasional, 
but obviously accelerating in its rate of nat- 
uralization. 


Smilacaceae 

Smilax glauca Walter—24861 (B), 25267 (F), 
24645 (H), 25101 (N). South-and west-fac- 
ing wooded slopes, fencerows; infrequent. 

Smilax hispida Muhl.—24958 (B), 25279 (F), 
24652 (H), 24759 (M), 25119 (N), 25071 
(R). Most habitats; common. 

Smilax rotundifolia L.—25302 (F). 
habitats; Knobs. 

Staphylea trifoli 24930 (B), 25323 (F), 
24608 (H), 24740 (M), 25086 (N), 25066 
(R). Mesic woods; frequent. 


Various 


Tiliaceae 

Tilia americana L. var. americana—24S888 (B), 
22S) 2465. CEl)s 24749 (Ni) ss 2bl 7 
(N), 24987 (R). Mesic woods; frequent. 


Ulmaceae 


Celtis occidentalis L—24907 (B), 25301 (F), 
24672 (H), 24773 (M), 25168 (N), 24999 
(R). Various habitats, less frequent in the 
Knobs; one of the most common trees in 
this part of Kentucky. 

Celtis tenuifolia Nuttall (= C. occidentalis var. 
georgiana of some authors)—25330 (F), 
3.4599 (H), 24965 (R). Upland xeric habi- 
tats, fencerows; infrequent. 

Ulmus americana L.—24960 (B), 25256 (F), 
24663 (H), 24805 (M), 25157 (N), 25043 
(R). Various upland and low habitats; com- 
mon. 

Ulmus pumila L.—24843 (B), 24792 (M). In- 
troduced; adventive in counties bordering 
the Ohio River. 

Ulmus rubra Muhl. 
24691 (H), 


—24900 (B), 25249 (F), 
24766 (M), 25192 (N). Mesic 


Kentucky 


Academy of Science 62(1) 


wooded and disturbed slopes, stream bluffs; 
frequent. 

Ulmus thomasii Sargent—24616 (H). Wooded 
stream bluff; rare. 


Viscaceae 
Phoradendron serotinum (Raf.) Johnst.— 
Thompson and Thompson #89-150, in KNK 
3), Thompson and McLaughlin #88-3217, 
in MDKY me 24620 (on Ulmus thomasii, 
see above)(H), Thompson #88-3223, in 
KNK (M), ee and Denton #89- 
3015, in MDKY (R). Less frequently en- 
countered northward. 


Vitaceae 

Ampelopsis cordata Michaux—24849 (B), 
25046 (H), 24727 (M), 25046 (R). Stream- 
banks, low open ground, and fencerows; 
sometimes frequent but rare or absent in 
Knobs. 

Parthenocissus quinquefolius (L.) Planchon— 
94922 (B), 25316 (F), 24673 (H), 24752 
(M), 25163 (N), 25016 (R). Most habitats; 
common. 

Vitis aestivalis Michaux—24911 (B), 25317 
(F), 25053 (H), 24997 (R). Roadsides and 
dry woods; common in Knobs, rare in Blue- 
grass. 

Vitis riparia Michaux—24794 (M). Low 
ground along Ohio River; rare or absent 
elsewhere. 


DISCUSSION 

Habitats 

Before embarking on this study, we were 
aware that most of the study region had a long 
history of intensive post- -settlement land use. 
However, we were frankly surprised at the 
present extent of its effects. Most natural hab- 
itats in this region have been completely de- 
stroyed or degraded beyond recognition. Even 
though one does not encounter extensive row 
crops in this part of the Bluegrass, most of the 
uplands in this rolling country have been com- 
pletely cleared, or only remnant, highly dis- 
turbed, young-aged woodlots remain. Down- 
slope from uplands, where steeper terrain of- 
ten militates against row cropping, land is 
most often relegated to pasture. Cattle (and, 
we suspect, very high populations of white-tail 
deer, Odocoileus virginianus) range through 
forested slopes and small creek bottoms, 


Woody Plants—Clark and Bauer 49 


fenced off only from dwellings and roadways. 
As a result, most of the native herbaceous for- 
est flora has been eliminated, woody plants do 
not regenerate naturally, and exotic species 
(such as Lonicera maackii and Rosa multiflo- 
ra) are favored over natives. Oak regeneration 
has practically been eliminated in most of this 
part of the northern Bluegrass; scarlet oak 
(Quercus coccinea) probably will disappear 
from this part of the Bluegrass within the near 
future. 

In situations such as this, highway rights-of- 
way and fencerows sometimes act as refugia 
for elements of the flora. However, in this part 
of Kentucky, this limited habitat is under fre- 
quent assault from the heavy application of 
herbicides along public roadways. Typically, 
this results in the elimination of most herba- 
ceous and seedling woody dicots, not only 
along the roadway but as far as the sprayer can 
reach into the forest, and the invasion and her- 
baceous stratum dominance of species such as 
Festuca spp. and Phalaris arundinacea are en- 
couraged. In other words, herbicide spraying 
is eliminating one of the very few habitats re- 
maining in the Bluegrass for native plants. It 
is a purposeful, all-out assault by the State on 
its native plants. 

As a result of these land-use patterns and 
prolonged exclusion of fire as an environmen- 
tal factor, habitats of high quality in our study 
area were few and far between. In our opin- 
ion, the remnant site we encountered (other 
than Quiet Trails Nature Preserve [Harrison 
Co.], within which we did not collect) with 
highest natural value was within Clay Wildlife 
Management Area in Nicholas County. In our 
opinion, portions of Clay Wildlife Manage- 
ment Area should be managed as a atrial 
area to protect what has been lost elsewhere 
in the region. In addition, we recommend that 
all remaining forested steep bluffs encoun- 
tered along the Licking River and its major 
tributaries should receive protection. Without 
these remnant habitats, a significant percent- 
age of the native woody species in this region 
would vanish entirely. 


Distribution Patterns 


We feel our study area is large enough to 
indicate some possible geographic distribu- 
tional patterns of woody plant occurrence but 
too small to produce definitive evidence. 


However, four distributional patterns perti- 
nent to the woody flora have emerged within 
the study area as a result of the evidence we 
compiled. These distributional categories are 
(1) an apparent distributional discontinuity be- 
tween the northern Bluegrass and northern 
Knobs; (2) plants whose distribution seems re- 
lated to the Ohio River corridor: (3) notewor- 
thy rarities; and (4) significant weeds. 
Category 1. Within the area we studied, it 
is apparent that there is a sharp distributional 
divide between the Bluegrass and Knobs. The 
following species occur in the Knobs but ap- 
pear to be entirely missing from the Bluegrass. 
Plants present in the rani and absent from 
the Bluegrass (during this study) include all 
species of Ericaceae (eit species), Acer rub- 
rum, Amelanchier arborea, Aralia spinosa, 
Betula nigra, Chionanthus virginica, Hama- 
melis virginiana, Liquidambar styraciflua, Hy- 
pericum stragulum, Nyssa sylvatica var. biflo- 
ra, Pinus (three species), Populus grandiden- 
tata, Prunus munsoniana, Quercus montana, 
Rosa palustris, Rubus allegheniensis, Salix ser- 
icea, Smilax rotundifolia, and Viburnum ac- 
erifolium. This means that more than 19% of 
native woody plant taxa occurring in the 
Knobs apparently do not occur in the adjacent 
northern Bluegrass. This is a significant dis- 
continuity; reasons for it are hypothetical. One 
possible explanation could relate to edaphic 
factors, which affect mycorrhizae; the latter 
are known to be favored by acid soil. Members 
of Ericaceae and Pinaceae will grow if planted 
in the northern Bluegrass, but perhaps ger- 
mination and establishment are selected 
against in Bluegrass soils. Also, it is well 
known to nurserymen that some species un- 
dergo mineral deficiency stress in dry alkaline 
soils; Acer rubrum is one of those species. We 
hypothesize that the combination of periodic 
drought, conditions unfavorable to mycorrhi- 
zae, Ane fire may have been major factors that 
prevented the establishment of seed sources 
in the Bluegrass, without which sustaining 
populations of some plants could not persist. 
One additional word about this distributional 
discontinuity is that, if one were to include the 
Cumberland Plateau (not far to the east of the 
study area), the floristic discontinuity becomes 
much more striking (includes Ilex spp., Mag- 
nolia spp., etc.). This issue might be a fruitful 
possibility for autecological investigations. 


50 


Category 2. A second recognizable group 
of woody plants is composed ae those that are 
either confined to the Ohio River corridor or 
are tound only in the Knobs and Ohio River 


corridor. As we define it here, the Ohio River 
eorridor includes the Ohio River bottom- 
lands, adjacent bluffs, and small streams 


draining directly into the Ohio River. Plants 
appar ently confined closely to the Olin River 
in our study area include Amorpha fruticosa, 
Physocarpus opulifolius, Rhus typhina, and 
Salix caroliniana. Those associated with the 
Ohio River and the Knobs but not found in 
the Bluegrass interior include Aesculus flava, 
Alnus serrulata, and Cornus amomum var. 
schuetzeana. This physiographic feature (i.e., 
the river corridor) may actually serve as a mi- 
gration pathway for some species around the 
apparent barrier of the northern Kentucky 
Bluegrass region. 

Category Plants that are rare in the 
study area include those that are confined to 
particular habitats, as well as those that are in 
an attenuated portion of their ranges. Plants 
we would single out in this category include 
several plants confined to steep or gladelike 
bluffs (Bignonia capreolata, Rhus aromatica, 
Ulmus thomasii); those near the edges of their 
ranges (Malus angustifolia, Nyssa sylvatica 
var. biflora, Prunus munsoniana, Quercus coc- 
cinea, Rhamnus lanceolata, Rosa virginiana, 
and Ulmus thomasii). ; 

Perhaps the most surprising rare taxon we 
encountered is Arundinaria gigantea. A few 
early informal accounts waxed eloquent on the 
extensive canebrakes in Kentucky, and these 
accounts seem to have been repeated endless- 
ly by later writers. Perhaps it was true at one 
time. However, at the present time in our 
study area, Arundinaria is among the rarest of 
woody plants. We extensively but fruitlessly 
searched for it and even offered rewards for 
its discovery. Eventually, Dr. Wendell King- 
solver of Nicholas County took the senior au- 
thor to a single population on his land. Ac- 
cording to him, cane in the northern Bluegrass 
was once widespread but now has been re- 
duced to a few small, isolated populations 
along main stem of the Licking River by a 
combination of gr azing by cattle and long-time 
use of the stems for “jig poles,” or throw-away, 
temporary fishing poles. Kingsolver a 


o 
comm.) stated chat protection of the small (< 


Journal of the Kentucky Academy of Science 62(1) 


ha) canebrake on his land from cattle grazing 
and pole collecting has allowed it to increase 
considerably in area, and ramets have been 
used to re-establish cane in another location 
where an extensive canebrake was completely 
extirpated. 

Category 4. On the whole, we determined 
that 11% of the woody plant taxa of this region 
is not native. (See Table 1.) Many of these 
plants are not especially noticeable weeds at 
the present time. One of them (Thuja orien- 
talis) is reported herein for the first time as an 
escape in Kentucky. Others (not included in 
our statistics) are plants whose native ranges 
are becoming obscured because they are both 
native and escaped from cultivation (e.g., Jug- 
lans nigra, Liriodendron tulipifera, Maclura 
pomifera, and Quercus palustris). 

However, there are some significant exotic 
weeds and, based on the senior author's per- 
sonal experience, few of these have reached 
their zeniths. The impact of most of these will 
increase with time, but some are more ag- 
gressive invaders than others. Non-native ex- 
otics escaped in our study area include the fol- 
lowing (those marked with an asterisk were 
vouchered from a minimum of five counties): 
Ailanthus altissima,* Albizia julibrissin, Ber- 
beris thunbergii, Broussonetia papyrifera, Ce- 
lastrus orbiculatus, Elaeagnus umbellata,* Eu- 
onymus alatus, Ewonymus fortunei,* Hedera 
helix, Hibiscus syriaca, Lespedeza bicolor, Li- 
gustrum SNCS, Ligustrum vulgare, Lonicera 
japonica,* © Lonicera maackii,* Malus sylves- 
tris, Morus alba,* Philadelphus coronarius, 
Populus alba,* Prunus cerasus, Prunus ma- 
haleb,* Prunus persica,* Pyrus calleryana,* 
Rosa multiflora,* Rosa wichuraiana, Salix 
alba, Ulmus pumila, and Vinca minor. From 
our field observations, Lonicera japonica and 
Rosa multiflora have the greatest negative eco- 
logical impact at present. However, Ailanthus 
altissima, Celastrus orbiculatus, Ewonymus 
fortunei, and Lonicera maackii are showing 
signs of rapid establishment and have the po- 
tential for very significant negative impacts 
within a few years. One omission readers may 
note is Paulownia tomentosa (Thunb.) Steu- 
del; though it appears not to be established in 
the study area yet, doubtless that will happen 
soon. 


Woody Plants—Clark and Bauer 51 


ACKNOWLEDGEMENTS 


We are grateful to the Eastern Kentucky 
University Research Committee for its finan- 
cial support of this project; to Brenda Clark 
and Adrienne Scott for their forbearance and 
support; to herbarium curators who made 
specimens in their care available for exami- 
nation; and to Dr. Robert Kingsolver for help- 
ing the senior author make contact with Dr. 
Wendell Kingsolver, who granted access to the 
Nicholas County Arundinaria population. 


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Yarmouth, ME. 

Anonymous. 2000a. “Kentucky Atlas & Gazetteer: Ken- 
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kentucky-counties.html (25 May 2000). 

Anonymous. 2000b. “International Plant Name Index.” 
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Bryant, W. S. 1987. Actual and potential vegetation of the 
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KY. 

Campbell, J. J. N. 1980. Present and presettlement forest 
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Guetig, R. G. 1993. The vascular flora of Estill County, 
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bariorum.” [8th edition, updated 28 Mar 2000] http:/, 
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herbariorum kentuckiensis HI. Trans. Kentucky Acad. 
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J. Ky. Acad. S 2(1):52-59. 2001. 


Effects of Fish on Zooplankton Community Structure in Chaney Lake, 


a Temporary Karst Wetland in Warren County, Kentucky 


Nicole Vessels and Jeffrey D. Jack' 


Department of Biology, Western Kentucky University, Bowling Green, Ke ntucky 42101-3576 


ABSTRACT 
Lake is an ephemeral karst lake in southern Kentucky (USA). 
Chaney often contains fish that can enter the lake through the underlying Lost River drainage system. The 


Chaney Unlike most ephemeral lakes, 
effects of introduction of golden shiners (Notemigonus crysoleucas) on zooplankton in Chaney were examined 
for a 2-week period in June and July 1997. Twelve fish were transplanted from an isolated region of the lake 
into each of three enclosures in an area of the lake where no fish had been observed. Three additional 
enclosures served as fish-free controls. Zooplankton samples and water chemistry and nutrient data were 
taken every 4 days. Water chemistry and nutrient data showed no significant differences in the measured 
parameters between enclosures with fish and those without fish. Bosmina and Acanthocyclops showed de- 
creases in population growth rates in the presence of fish. The fish had no effect on the growth rates of the 


smaller zooplankton present such as the rotifers. Vertebrate predation in systems like Chaney Lake may pose 


significant ecological challenges for organisms adapted to temporary habitats. 


INTRODUCTION 


Fish can be important determinants of zoo- 
plankton abundance, species structure, and 
productivity in aquatic systems. Since the early 
work of Hrbacek et al. (1961) there have been 


numerous studies fede the impacts of 
fish on zooplankton communities. Removal of 


zooplanktivorous fish from lakes has been 
shown to increase the densities of herbivorous 
zooplankton with concomitant effects on the 
phytoplankton (Carpenter et al. 1985, 1987; 
Vanni et al. 1990). This work has prompted 
research in the use of fish in “biomanipula- 
tion” to aid managers in controlling water 
quality in lakes (Shapiro and Wright 1984). 
There has been less research eouduered on 
the effects of fish on zooplankton in wetland 
or forested lake communities, although fish 
have been shown to have significant effects on 
zooplankton in shallow, eutrophic, lakes that 
may be similar in many respects to wetlands 
(Hanson and Butler 1990). In a study of semi- 
permanent Minnesota wetlands, Hanson and 
Riggs (1995) found that densities, biomasses, 
and taxa richness of zooplankton were signifi- 
cantly lower in wetlands that contained fish as 
opposed to similar ponds which did not con- 
tain any fish. In a study of another Minnesota 


' Current address: Department of Biology, University of 


Louisville, Louisville, KY 40292: to whom correspondence 
should be sent. 


prairie lake, a fish kill resulted in shift in zoo- 
plankton species composition from Bosmina 
and Chydorus to the larger Daphnia galeata 
and D. pulex (Hanson anal Butler 1994). 
Wetland areas may provide more refuges to 
zooplankton from fish predation than the open 
water column of a lake.In a pond-enclosure 
experiment where the density of vegetation 
was controlled, perch did not consume as 
much zooplankton biomass when vegetation 
was present in the enclosures as when vege- 
tation was absent (Diehl 1992). In their study 
of a coastal marsh along Lake Erie, Kreiger 
and Klarer (1991) found that some copepods 
and cladocerans were more abundant near the 
sediments or near macrophytes than in the 
open-water column, which is consistent with 
earlier work suggesting that vegetation can 
provide an important refuge for zooplanktors 
from fish predation (Timms and Moss 1984). 
While such biotic interactions between fish 
and zooplankton are important in structuring 
planktonic communities in permanent aquatic 
systems, in most temporary or ephemeral 
pond systems zooplankton are not subject to 
fish predation, although other vertebrates such 
as amphibians may have strong effects on z00- 
plankton assemblz ages (Wilbur 1997). Indeed, 
taxa such as the large branchiopod Crustacea 
are thought to be successful in temporary sys- 
tems because fish are often excluded from 
these habitats (Kerfoot and Lynch 1987). Or- 
ganisms inhabiting ephemeral water bodies 


Fish and Zooplankton Community Structure—Vessels and Jack 53 


typically have life histories and ecological 
strategies that are synchronized to the hydrol- 
ogy of their habitat (reviewed in Wiggins et al. 
1980) and that do not necessarily give these 
organisms any advantage in responding to pre- 
dation from fish. In most transient water hab- 
itats, the threat of fish predation would be ex- 
ceedingly small; in some ephemeral karst 
lakes, however, fish predation may be an im- 
portant force structuring the zooplankton 
community. 

Karst geology is characterized by extensive 
caves, sinkholes, sinking streams, and springs. 
Because of the many conduits leading into the 
subsurface in well-developed karst landscapes, 
surface water may be directed rapidly into the 
groundwater areas through a sinkhole, may 
rise again at a spring, and then sink into the 
subsurface again as a sinking stream. These 
points of exit and entrance of water are called 
estavelles. In many karst landscapes, standing 
surface water is uncommon, but variations in 
local geology can produce ephemeral karst 
lakes and wetlands such as Chaney Lake in 
south-central Kentucky. Chaney Lake is a 68- 
hectare state nature preserve located about 10 
km south of Bowling Green in Warren County, 
Kentucky. The lake area is a shallow depres- 
sion that is connected by fissures (estavelles) 
in the subsurface rock to the Lost River Cave 
system, which has a drainage basin of about 
933 km?. The lake is formed because of the 
Lost River Chert formation, which overlays 
the limestone under the lake and prevents the 
rapid return of water to the subsurface except 
where estavelles are located. When the capac- 
ity of the Lost River Cave system is exceeded 
during periods of high precipitation, ground- 
water enters Chaney Lake via the estavelles 
and also through one intermittent surface 
stream on the south edge of the preserve. As 
the water levels in the Lost River drop, water 
may leave the lake through the estavelles as 
well, leaving large numbers of smaller, isolated 
pools behind. The lake usually holds water 
from December through August, although 
from May on most of that water is in small 
pools (Jack, personal observation). 

Historically, karst wetlands such as Chaney 
Lake were very important aquatic habitats in 
western Kentucky because they provided an 
important source of standing water in a terrain 
that had little surface water. Chaney Lake and 


another nearby karst lake, Rich Pond, 
huge numbers of migrating birds in the spri:< 
and may be an important foraging area for a 
variety of waterfowl on their spring migrations 
(Mason, personal communication). Chaney 
Lake contains a variety of zooplankton species, 
including common ephemeral pond taxa such 
as the fairy shrimp Streptocephalus. However, 
the same estavelles that deliver water to sup- 
port these communities can also serve as con- 
duits for vertebrates such as the spring fish, 
Chologaster agassizi, to enter the lake. If fish 
can enter Chaney Lake, they may have a 
strong impact on zooplankton densities and 
community structure. 

In early June 1997, a population of golden 
shiners (Notemigonus crysoleucas), which may 
have entered Chaney Lake via estavelles, was 
found in an isolated pool in the northeast cor- 
ner of Chaney Lake. These fish were trans- 
planted to experimental enclosures in another 
section of the lake to assess the effect of fish 
on the zooplankton communities in Chaney. 
We hypothesized that the fish would select the 
largest zooplankton species in the water col- 
umn and, in turn, would cause a decrease in 
population growth rates of these larger spe- 
cies. We expected that the smaller zooplank- 
ton species such as rotifers would have no sig- 
nificant response to the fish or that they would 
increase in numbers if they are released from 
competition for resources with, or predation 
from, the larger macrozooplankton (see Gil- 
bert 1988; Jack and Gilbert 1997). 


MATERIALS AND METHODS 


The study was conducted from 19 Jun 1999 
to 1 Jul 1997 in a marsh area in the south- 
eastern portion of Chaney (see Kelley et al 
2000). This area was chosen as the study site 
because no fish had been observed there and 
it was unlikely to dry during the period 
planned for the experiment. The dominant 
vegetation in the marsh was buttonbush (Ce- 
phalanthus occidentalis), aquatic plants such 
as Polygonum sp. and a liverwort in the genus 
Riccia. The higher ground around the marsh 
area is ringed by tree species such as swamp 
white oak (Quercus bicolor), red maple (Acer 
rubrum) and sycamore (Platanus occidentalis ). 
Large trees are not present in the marsh itself, 
perhaps because this part of Chaney consis- 


54 Journal of the Kentucky Academy of Science 62(1) 


tently holds water for most of the year (>7 
months in 1995-1998). 
Six |-m* enclosures were constructed using 
P\C piping for frames and plastic for the 

des. The enclosures were then placed in the 
marsh, enclosing the water column and the as- 
sociated zooplankton. They were anchored 
into the sediment of the marsh and were open 
at the top and at the bottom. Three enclosures 
were randomly selected to hold 12 fish each, 
with the other three serving as controls. The 
fish were caught by sweeping a net in the pool 
containing the fish and moving the fish into 
the appropriate enclosures in the marsh. 
These stocking densities were about % the es- 
timated density of the fish in the original pool. 
This was determined by visually assessing the 
numbers of fish in the source pool and then 
taking transects through the pool to determine 
average depth and emclen which were then 
used to estimate pool volume. Invertebrate 
and water chemistry samples were taken on 
the first day of the experiment and every 4 
days afterward in each of the six enclosures 
for the duration of the project. Depth, turbid- 
ity, specific conductivity, temperature, pH, dis- 
solved oxy gen, and percent dissolved oxygen 
were Peed using a YSI 6250 multiprobe. 
One-liter nutrient grab samples were also tak- 
en in acid-washed bottles to measure nitrates, 
ammonia, and soluble reactive phosphorous 
using a Hach DREL 6000 water analysis kit. 
Nitrate was measured by the Cadmium Re- 
duction method: ammonia was determined us- 
ing the Nessler method; and soluble reactive 
phosphorus was measured using the Hach 
Phosver 3 method. Invertebrate sampling was 
conducted using a 7-cm-diameter coring de- 
vice. Two 5-liter core samples were taken from 
each enclosure on each sampling date and fil- 
tered through a 20-4~m mesh sieve. The ma- 
terial collected was then washed into a con- 
tainer and preserved in 90% ethanol. 

All samples were counted in the laboratory 
by using an Olympus SZH 10 dissecting mi- 
croscope. The samples were counted in their 
entirety and the dominant taxa (>95% of nu- 
merical abundance) were identified down to 
the lowest practicable taxon (usually genus). 
Population growth rates were calculated for 
the domaaanit taxa from the first and last ex- 
perimental dates using the equation, r = 
InN,-InN,t', where r is equal to the popu- 


lation growth rate, N, is equal to the final sam- 
pling date, No represents the beginning sam- 
pling date, and t stands for the total number 
of days in the experiment (Jack and G ilbert 
1993). A Student's t-test was used to compare 
growth rates in the fish and fishless enclosures. 
A repeated measures ANOVA was calculated 
using SYSTAT version 7.0 to assess changes in 
the physical parameters in the enclosures. 


RESULTS 


The data met the assumptions of ANOVA 
so transformation was not needed before the 
data could be analyzed. Water chemistry and 
nutrient analysis data indicated no significant 
differences (P > 0.07) in measured parame- 
ters between treatments over time. The aver- 
age temperature for all enclosures was 23.4 + 
0.84°C and the pH in the enclosures averaged 
5.93 + 0.57 (all data are presented as mean 
+ (standard errors). Turbidities were ee 
variable in all enclosures (39.4 + 23.2 and 
40.0 + 24.3 for fish and non-fish espera 
and were higher after storm events. Dissolved 
oxygen levels ranged from just under 1 mg li- 
ter! to 3.3 mg liter~! but there were no dif- 
ferences in oxygen levels between treatments. 
Average nitrate, ammonia and soluble reactive 
phosphorus concentrations were not different 
between treatments (P > 0.15; Figure 1). 

The fish added to the enclosures had an ini- 
tial average size (mouth to base of caudal fin) 
of 1.95 + 0.21 cm; the average length at the 
end of the experiment was 2.46 + 0.06 cm. 
The macrozooplankton assemblage in the 
marsh was dominated (>90%) by Bosmina sp. 
and Acanthocyclops sp., with smaller numbers 
of Cer ‘iodaphnia sp., Daphnia sp., and at least 
two species of ostracods. The ostracods were 
similar in size and were grouped together for 
the purposes of the analysis. One isopod (Cae- 
cidotea sp.), one amphipod (Hyalella azteca), 
and at least one water mite (Hydracrina) spe- 
cies occurred in the samples. The latter three 
groups were considered to be accidentals in 
the plankton and were not included in the 
analysis. The microzooplankton assemblage 
was primarily composed of rotifers, with Mon- 
ostyla sp., Euchlanis sp. and Ascomorpha sp. 
as the numerical dominants (>78%). There 
were also one species each of Branchinous, 
Keratella, and Lecane, and two species of Po- 
lyarthra species present in some samples. We 


Fish and Zooplankton Community Structure—Vessels and Jack 


On 
Ot 


Nutrients 


@ Fish 


0.4 
0.3 
0.2 
0.1 

0 


= 
OD 
S| 


Ammonia 


Figure 1. 


@ No Fish 


Nitrate SRP 


Ending concentrations of ammonia, nitrate, and soluble reactive phosphorous (SRP) in fish and fishless 


enclosures in Chaney Lake, Warren County, Kentucky (19 Jun 1999-1 Jul 1997). Bars show means and standard errors. 


did not find any identifiable protists in our 
samples, but our filtering and fixation proce- 
dure may have prevented adequate sampling 
and recognition of these small organisms. 

The presence of fish in the enclosures neg- 
atively affected the larger invertebrate taxa but 
showed no effect on the smaller taxa. The fish 
negatively affected the Bosmina sp. and the 
Anthocyclops sp. growth rates (P < 0.006; Fig- 
ure 2). Ending mean densities of Bosmina 
were significantly higher in the fish- less enclo- 
sures (43 individuals liter~') compared to the 
fish enclosures (5.3 individuals liter~!). Acan- 
thocyclops sp. growth rates and ending mean 
densities were significantly higher in enclo- 
sures without fish (33 individuals liter~') than 
in enclosures with fish (7.3 individuals liter~'). 
Ostracods as a group had a positive population 
growth rate when the fish were present and a 
negative growth rate without fish, but there 
was no significant difference between the two 
treatments (P = 0.07). 

The presence of fish had no significant ef- 
fect (P > 0.07) on the population growth rates 
of the three dominant rotifer species- 
Ascomorpha, Euchlanis, and Monostyla (Fig- 
ure 3); however the densities of both Euch- 
lanis and Ascomorpha decreased over time in 
all of the enclosures. Density of Monostyla sp. 


increased nearly 10 times over the same time 
period in all enclosures. 


DISCUSSION 


Our results confirmed that fish predation 
can affect aquatic invertebrate community 
structure in Chaney Lake. 

Physical factors and nutrients assayed were 
not significantly different, so these factors 
probably did not contribute to the response of 
the zooplankton to fish. In enclosure experi- 
ments the effects of fish are sometimes the 
result of indirect mechanisms. Ammonia ex- 
creted by fish, for example, may enhance algal 
growth or the fish may feed on the algae, com- 
peting with the zooplankton. The golden shin- 
ers, however, are zooplanktivorous at this size 
(R. Hoyt, pers. comm.), and we found no dif- 
ferences in nutrient concentrations between 
the two treatments. The population growth 
rates we reported for our organisms were 
somewhat low compared to the instantaneous 
growth rates of related taxa in other ephem- 
eral systems (Taylor et al. 1989), but this may 
be a reflection of a more constrained resource 
base in Chaney Lake. 

Our data support the conclusion that fish 
predation in the enclosures was driving the 
suppression of large zooplankton in our study. 


~p 


56 Journal of the Kentucky Academy of Science 62(1) 


Crustaceans 


@ Fish 


Bosmina 


th rates day’ 
Sa 
je) = NO 


Population grow 
Sy. i 
Oo N 


Figure 2. 
Warren County, Kentucky (19 Jun 1999-1 Jul 1997). 


Bosmina are not typically considered very vul- 
nerable to fish predation because of their 
small size es: to other crustaceans, and 
many cyclopoids have well-developed escape 
responses that they may use to avoid fish pre- 
dation. As the largest common zooplankton 
prey in the Boece however, one would 
expect that visually foraging fish would focus 
on Bosmina and the Acanthoc ylops. The os- 
tracods were not significantly affected even 
though their mean size in the enclosures was 
comparable to that of Bosmina (0.65 vs. 0.44 
mm, respectively). Ostracods are often found 
in association with vegetation or in benthic ar- 
eas, where the golden shiners are unlikely to 
forage Secesecilly on them. The ostracods 
present in our samples may not have been 
common in the plankton but may have been 
captured when the corer displaced them from 
the vegetation or the bottom. 

Schneider and Frost (1996) found that in 
some cases the suppression of Daphnia in 
temporary ponds was associated with an in- 
crease in rotifer densities and taxon diversity. 
We did not see the increase in rotifer densities 
that we expected would occur once the ma- 


Acanthocyclops 


™ No Fish 


Ostracods 


Taxa 


Population growth rates of the dominant crustacean species in fish and fishless enclosures in Chaney Lake, 
Bars show means and standard errors. 


crozooplankton numbers were reduced; in- 
creases and decreases in rotifer numbers oc- 
curred across all enclosures regardless of 
treatment (see Results). This may indicate that 
the larger zooplankton are not significant 
predators on or competitors with the rotifers 
during this period in Chaney or that the roti- 
fers were being more strongly limited by re- 
source levels or other physiochemical factors 
in the lake. Previous research has indicated 
that small cladocerans like Bosmina are gen- 
erally not effective predators on rotifers (re- 
viewed in Gilbert 1988). The high densities of 
cyclopoid copepods in our study were also 
lower than those reported by Schneider and 
Frost (1996) in their study systems, so the im- 
pact of these crustaceans on the rotifers may 
have been too low to elicit a “release re- 
sponse” when the cyclopoid densities were re- 
duced. The general increases and decreases in 
rotifer densities could also be the result of an 
unidentified “enclosure effect” affecting the 
rotifers. However, since this effect was ex- 
pressed across all of the enclosures indepen- 
dently of treatment it should not affect our 


Fish and Zooplankton Community Structure—Vessels and Jack 57 


Rotifers 


0.75 


0.5 


Population Growth Rates Day” 


mg Fish 


wo No Fish 


ot 
oa 
me 


Taxa 


Figure 3. Population growth rates of the dominant rotifer species in fish and fishless enclosures in Chaney Lake, 
Warren County, Kentucky (19 Jun 1999-1 Jul 1997). Bars show means and standard errors. 


interpretation of the fish effects in these en- 
closures. 

The introduction of fish can have a pro- 
found impact on a system like Chaney. Early 
in the year, the entire Chaney basin is full, but 
as the year progresses it often dries to nu- 
merous smaller and disconnected ponds. If 
fish like the golden minnows are trapped in 
these pools, they could conceivably remove all 
of the large zooplankton from a particular 
pool. Many of the rotifers and crustaceans in 
these small pools appear to have multiple gen- 
erations before they produce diapause eggs. If 
fish are introduced to a pool and severely re- 
duce macrozooplankton densities, this could 
constrain the next year s recruitment in that 
pool. These pools may be colonized by zoo- 
plankton from other areas during periods of 
high water or there may be diapause eggs 
from previous years that could hatch, but that 
would be dependent on the vagaries of water 
level and on the natural histories of the organ- 
isms involved. If there are localized ecotypes 
adapted for the particular conditions in a pool 


or a closely associated group of pools, any 
unique genetic information in that popula- 
tion’s gene pool may be lost as a result of fish 
predation. This information would probably 
not be replaced by colonists from other areas. 

The path by which the fish enter Chaney 
may also be important. It is clear that fish may 
enter Chaney by the estavelles, but it may also 
be possible that they could enter the lake via 
overland flow during flood events. The area 
around Chaney contains at least one farm 
pond that may become connected to the lake 
during very high rainfall events. This would 
greatly expand the pool of potential fish colo- 
nists as fish that would be unable to survive in 
or travel through the Lost River system could 
enter Chaney in this manner. Very high den- 
sities of fish could be introduced this way; they 
could have an impact on the waterfowl and 
other vertebrate groups (e.g., amphibians) that 
use Chaney as breeding or feeding ground. 
Previous studies have already documented 
that waterfowl and fish are competitors for in- 
vertebrate prey in other wetlands (Hanson and 


58 Journal of the Kentucky Academy of Science 62(1) 


Riggs 1995), and that the presence of fish may 


reduce the prey available to birds that rely on 
Chaney as a foraging area during their spring 
migrations. 


SUMMARY 


Hydrology has long been identified as an 
import: int ‘factor impacting organisms in 
ephemeral lakes and pools. In C ff aney, how- 
ever, fish may be present in even anal pools 
with short duration since they may colonize 
the lake during the high flood period when the 
whole basin is inundated. 

Our data indicate that fish predation has sig- 
nificant effects on zooplankton community 


structure in Chaney Lake, but the impact of 


these effects on ecosystem level processes 
such as nutrient cycling and microbial activity 
remains unexplored. Temporary habitats like 
Chaney provide tremendous natural laborato- 
ries for inv estigating ecological processes at a 
number of different scales. In karst terrenes 
in particular, these lakes probably have land- 
scape-level effects on water quality and trans- 
port as well as local importance as centers for 
aquatic biological production and biodiversity. 
With many Bi these lakes threatened by de- 
velopment it is crucial that we continue to 
study these unique systems so that better 
management and preservation strategies can 
be devised to preserve the ecological integrity 
of these remarkable habitats. 


ACKNOWLEDGEMENTS 


We gratefully acknowledge the assistance of 
the following mndivicale Randall Kelley 
helped in che: field; Joyce Bender and Debra 
White of the Kentucky State Nature Preserves 
Commission gave logistical assistance; Wayne 
Mason provided information about the birds 
at Chaney; Robert Hoyt identified the fish; 
and Doug McElroy, Michael Stokes, and Da- 
vid Jenkins made useful comments on earlier 
versions of this paper. This work was support- 
ed by a grant from the Kentucky State Nature 
Preserves Commission to JD] and by support 
from the Western Kentucky University Un- 
dergraduate Honors Program to NV. 


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A Historiography of Archaeological Research in the Mammoth Cave 
Area of Kentucky: 1824-2000 


Kenneth C. Carstens 


Department of Geosciences, Murray State University, Murray, Kentucky 42071 


ABSTRACT 


Archaeological interest in the Mammoth Cave 


area of Kentucky has been ongoing since the early 19th 


century, primarily because of the unique preservation offered at the underground cave sites. In this paper 


[| examine almost 200 years of archaeological research conducted in the area. The paper loosely adheres to 


historical divisions first presented by Schwartz and later by Willey and Sablof. 


INTRODUCTION 


Examination of the history of archaeological 
study in the Mammoth Cave region of Scie 
central Kentucky reflects par allel deve ‘lopment 
with the growth of archaeology in North 
America (Guinane 1967: Willey 
1993), but it also demonstrates why additional 
intensive and systematic archaeological study 
should to be conducted in this very “significant 
archaeological region. In this paper I “provide 
an historical overview of archaeological re- 
search that has taken place in and around 
Mammoth Cave National Park. The paper 
loosely adheres to historical divisions first pre- 
sented by Schwartz (1967) and expanded by 
Willey and Sabloff (1993). 


The Speculative Period: Pre-1915 

One of the earliest archaeological records 
pertaining to the central Kentucky karst area 
was written in 1824 by Constantine Samuel 

Rafinesque (1824). Rafinesque was deeply in- 
terested in prehistoric remains of the Ohio 
Valley (Stout and Lewis 1995:83-90). Accord- 
ing to Col. Bennett Young (1910:18), Rafin- 
esque claimed to have located 148 ancient 
sites (settlements) and 505 monuments in a 
41-county area of Kentucky, speculating as 
with other early 19th century naturalists about 
the origin of these “natural” curiosities. Raf- 
ine sque’s entry for the central Kentucky karst 
lists “shell mounds along Green River and 
mummies in caves.” 

Following Rafinesque’s initial inquiry into 
Kentucky's prehistory, there appears to be an 
absence of related literature about the antiq- 
uities of the area. This is not to say that inter- 
est in antiquities had died; it had not. Accord- 
ing to Col. Bennett Young (1910), increased 


‘and Sabloff 


60 


farming activity and, in general, disruption of 
the land due to population growth, caused an 
escalation in destruction and looting of pre- 
historic sites. By 1870, the collecting, selling, 
and smuggling of antiquities in Kentucky was 
a major profession. Although the Mammoth 
Cave area is known mostly for its large cave 
system, the archaeological contents of the area 
offered a variety of artifacts and desiccated 
human remains for collecting and selling. 
Finds, such as Fawn Hoof in 1813, Scudder’s 
Mummy in 1814, Little Al in 1875, and Lost 
John in 1935, helped make the Mammoth 
Cave area famous (Meloy 1968). During the 
mid-19th and early 20th centuries, many in- 
dividuals explored nearby cave systems looking 
for mummies and Indian relics to sell (see 
Young 1910). Unfortunately, this dilettantish 
pastime stopped only in those caves that came 
under the pr otection of the National Park Ser- 
vice (NPS) after 1940 (e.g., Salts, Mammoth, 
Longs, Bedquilt, and Lee caves); even then, 
infrequent looting of caves within the National 
Park still occurred. 

The earliest historic date known from inside 
Salts Cave is 1809 (Watson et al. 1969:7). 
Dates and names upon various signature rocks 
in Mammoth and Salts caves indicate that the 
majority of historic caving dates “from the last 
quarter or so of the 19th century to the first 
quarter of the 20th century” (Watson et al. 
1969:7: see also Watson 1974:21-23). The veg- 
etal antiquities (e.g., textile bags, cordage, san- 
dals) that could be found within the dry caves 
were not preserved normally in surface or 
“open” sites. Hence, those items were espe- 
cially sought for collecting, smuggling, and 
looting. As an example of smuggling, in 1874 
or 1875 Louis Vial and some friends explored 


Archaeological Research in the Mammoth Cave Area—Carstens 61 


extensively in Salts Cave using a “new side en- 
trance known only to themselves” (Watson et 
al. 1969:7). During one of those cave trips 
they found the * ‘Salts Cave Mummy,” nick- 
named incorrectly “Little Alice” (Robbins 
1971, 1974; K. Tankersley et al. 1994; Watson 
et al. 1969:7). More recent examinations by 
the late Louise Robbins (1971:200—206) iden- 
tified the sex and age of this individual to be 
those of a 9-year-old male. 

During the 1890s, men such as F. W. Put- 
nam of the Peabody Museum, as well as local 
Kentuckians such as Colonel Bennett Young, 
T. F. Hazen, and W. D. Cutliff, made extensive 
collections and/or purchased prehistoric ma- 
terial from Salts and Mammoth caves. Young 
(1910:300, 305) stated: 


In 1893 Mr. Theodore F. Hazen ... opened a new 
entrance into Salts Cave . . . [and] obtained many in- 
teresting relics ... about the present entrance [Salts 
Sink], numerous spalls, flakes of flint, pestles, axes, 
awls, and other implements have been found .. . 


Young went on to describe many artifacts 
taken from within Salts and Mammoth caves, 
such as cords of bark, hemp, cattail leaves, and 
grass; basketwork; half-burmed cane torches: 
corn cobs (probably modern); an aboriginal 
ladder; wooden digging implements; cups, 
Aches bowls, and water bottles made from 
gourds and squash rinds; tobacco leaves and 
seed pods (also probably modern); and many 
chert implements. The large collection of an- 
tiquities Young acquired was sold to the Mu- 
seum of the American Indian, Heye Founda- 
tion, New York (Schwartz 195S8e; Watson 
1974:167). Later, John M. Nelson, who was a 
cave guide from 1894 to 1907, extensively col- 
lected antiquities both from the caves and 
from surrounding surface sites (Carey 1942; 
Schwartz 1958f:3; Watson et al. 1969; Watson 
1974). With the exception of the John M. Nel- 
son collection, the other large private collec- 
tions were either given or sold to the Ameri- 
can Museum of Natural History, the Smith- 


sonian Institution, or the Peabody Museum of 


Archaeology and Ethnology. It was the Mam- 
moth Cave Estates collection, donated to the 
American Museum of Natural History in 1913, 
that prompted Nels C. Nelson (no relation to 
John M. Nelson) to engage in the “only sci- 
entific archaeological investigations’ ’ (Schwartz 
1958d) of the Mammoth Cave area up to that 


time and usher in the Classificatory Period of 
archaeological work (Schwartz 1967; Wille 
and Sabloff 1993). 

The Speculative Period focused initiall, 
upon discovery, with only meager attempts to 
offer explanations of deriv Aeon of the discov- 
ered sites. Once they were discovered and 
made known, the sites were vandalized and 
exploited for private purposes. Whether pots, 
“arrowheads,” or “mummies,” the market for 
trafficking in North American antiquities had 
begun. But it was from the seeds of site de- 
struction that the first museum acquisitions 
were made, ushering in professional archaeo- 
logical work of the Classificatory Period. 


The Classificatory Period: 1916-1960 


Nels C. Nelson worked in Mammoth Cave 
National Park during May and November 
1916 as an archaeological representative of the 
American Museum of Natural History. His 
1917 report described the materials found 
during his surface and cave reconnaissances 
and excavations in the Mammoth Cave area. 
Specifically, Nelson described and compared 
his surface finds from the Mammoth Cave and 
Eaton Valley fields to similar bifacial chipped 
stone materials then being found in the 
French Paleolithic (Nelson 1917:16—19, 1923). 
In total, Nelson examined, through excavation 
and/or other study, six of nine cave sites, six 
of seven open-surface sites, and one of four 
rockshelters (Nelson 1917:11).The latter num- 
ber refers to the category of site types he re- 
ported for the Mammoth Cave area. Douglas 
W. Schwartz (1958d:1—2) stated that Neleons 
main contribution was to “scientifically docu- 
ment the presence in the caves of some classes 
of material previously only reported by ama- 
teurs.” Nelson also drew substantial conclu- 
sions from his materials, despite the lack of 
published reports concerning antiquities of 
the area and the role of plant domestication 
(he found large quantities of charred sunflow- 
er seeds in his Mammoth Cave vestibule ex- 
cavations) in the central Kentucky region. He 
concluded that the Flint-Mammoth Cave sys- 
tem had an economic importance to the N Na- 
tive Americans, e.g., the quarrying of flint 
(e.g., from Flint Alley in Mammoth Cave, 
which has since been questioned [Munson et 
al. 1989: Prentice 1993]); Nelson was not 
aware that the Native Americans also exploit- 


62 Journal of the Kentucky Academy of Science 62(1) 


ed the caves for minerals (e.g., mirabilite, gyp- 
sum, s.tin spar, and selenite; see Munson et 
al. 1989: K. Tankersley 1996). 


‘lson’s archaeological excavations inside 
th vestibule of Mammoth Cave is his major 
ork in the Mammoth Cave area. Although 
this excavation was exploratory, it was ex- 
tremely extensive and thorough. Nelson sank 
a series of 10 test trenches that revealed mid- 
den in two places. One was near the west wall 
of the entrance; the other, some 40 feet from 
the first, extended 50 feet back over the entire 
entrance area. Although Nelson’s notes are at 
times ambiguous, he demonstrated a strong 
concern for the temporal and spatial location 
of artifacts excavated (personal observation in 
April 1975 of Nelson’s catalog record on file 
with the American Museum of Natural His- 
tory, New York). 

Nelson excavated almost all of the vestibule 
entrance, but the number of artifacts found 
was few. Douglas W. Schwartz (1958d) ex- 
plained that ANG was probably the result of ex- 
tensive looting that had occurred earlier 
throughout the 19th century. It also may be 
the result of Nelson’s recovery me sthods (no 
screens were used) and/or extensive subsur- 
face alterations resulting from cave commer- 
cialization or previous saltpeter mining oper- 
ations during the War of 1812 (Meloy 1968). 
Any of these reasons may explain the paucity 
of artifacts recovered from the Mammoth 
Cave vestibule. Nelson did find and recognize 
evidence of prehistoric diet in the form of an- 
imal bone, sunflower seeds, and freshwater 
molluses (Nelson 1917; Watson 1974:212). He 
also found prehistoric tools such as bone awls, 
bone flakers, antler points, tubes, stone pro- 
jectile points, scrapers, ground stone imple- 
ments, and items for personal adornment. 
Most important, Nelson (1917:69) concluded 
that two different cultures could be distin- 
guished within his vestibule excavations. The 
lowed or more “primitive” group was identified 
yi Nelson as as archeologists would define 
2 decades later (e.¢., Bitehie 1933) as the Ar- 
chaic culture ie 1960a:133). 

Only one other reference to Mammoth 
Cave area prehistory appeared in print during 
the first 2 decades of the 20th century. This 
was a fleeting mention of a series of rockshel- 
ter sites near what is now the western bound- 
ary of Mammoth Cave National Park. The ref- 


erence was made by C. B. Moore who visited 
the Indian Hill rockshelter complex in 1915 
(Moore 1916). Fortunately, shallow water con- 
ditions on Green River forced Moore to ter- 
minate his plunderous Green River expedition 
near Indian Hill as his boat, The Gopher, was 
too large to continue the journey upstream. In 
1935, a newly discovered desiccated burial 
within Mammoth Cave, known to the cave 
guides as “Lost John,” brought additional ar- 
Gree ological publicity to the area (Pond 1935, 
1937). Mienae Pond and George Neumann’s 
analysis of Lost John ( (Neumann 1938) consti- 
tuted the only professional archaeological in- 
quiry in the Mammoth Cave area between N. 
C. Nelson’s 1916 work and the formation * 
Mammoth Cave National Park (MCNP) i 
1940 although several additional references te 
caves and rockshelter sites in and around the 
present boundary of the park appeared in 
print intermittently (e.g., Fowke 1922; Funk- 
houser and Webb 1932). With the final acqui- 
sition of lands by the federal government on 
25 Apr 1940, it ‘became a federal offense to 
remove materials from cave interiors within 
MCNP. 

During the formation of MCNP, the Mam- 
moth Cave National Park Association pur- 
chased, from John M. Nelson, a collection of 
prehistoric, historic, and geological specimens 
that were subsequently donated to MCNP on 
15 Jan 1942 (Carey 1942: 1). Henry A. Carey, 
then of the Archaeology Department at the 
University of Kentucky, was placed in charge 
of cataloging the park’s new acquisitions. He 
was Seeuiel by a new NPS employee, Jesse D. 
Jennings (Carey 1942). Unfortunately, the ma- 
jority of the : 25,000 specimens in the John M. 
Nelson collection was without provenience. 
Items in the collection had been bought from 
the local area with no note made as to the 
exact collecting location. Furthermore, Nelson 
kept only “mental notes” for his more unusual 
specimens. From the John M. Nelson collec- 
tion, Henry Carey concluded (1) that the 
MCNP area was utilized for an extensive pe- 
riod of time by aboriginal peoples; (2) that a 
typological sequence could be worked out for 
the area by using the collection but extreme 
caution should be used in drawing definitive 
conclusions due to the lack of controlled lo- 
cational data; and (3) that scientific archaeo- 
logical excavations inside the caves and at se- 


Archaeological Research in the Mammoth Cave Area—Carstens 63 


lected surface sites should be started imme- 
diately. Unfortunately, due to the start of 
World War II, the Mammoth Cave collections 
heralded for study by Carey were not exam- 
ined again until 1957 when Douglas W. 
Schwartz, from the University of Kentucky, ex- 
amined the John M. Nelson materials and at- 
tempted to relocate some of the surface sites 
from which John Nelson had made his collec- 
tions (Schwartz 1958f). Schwartz also brought 
systematization to the study of Native Ameri- 
can sites in MCNP and visited several major 
museums in the East to study collections ac- 
quired from the Mammoth Cave area at the 
turn of the century (Schwartz 1958a—h). These 
activities culminated in a series of valuable de- 
scriptive reports (Schwartz 1958a—h) and oth- 
er interpretive and popular accounts about the 
archaeology of the area (Schwartz 1960a; 
1965). 

The Classificatory Period brought a logical, 
scientific inquiry to the archaeology of the 
Mammoth Cave area. With the first work of 
Nelson in 1916, to the discovery of a desic- 
cated individual (Lost John) in 1935, to the 
systematic reporting of archaeological sites 
above ground and below by Douglas Schwartz 
during the late 1950s/early 1960s, the archae- 
ology of the Mammoth Cave area yielded, 
ever so slowly, evidence of very significant in- 
formation about prehistoric cultural adapta- 
tions and cultural processes. These later stud- 
ies served as the foundation for investigations 
by Patty Jo Watson, the Cave Research Foun- 
dation, the National Park Service, and the II- 
linois State Museum (Watson et ar 1969) dur- 
ing the next period of archaeological devel- 
opment. 


The Explanatory/Interdisciplinary Period: 
1960 to the Present 


In 1942, Henry Carey emphasized the need 
for further surface investigations and excava- 
tions in MCNP, but little was accomplished 
until the recent research efforts by Patty Jo 
Watson and her associates (Brown 1977; Car- 
stens 1974, 1975, 1976 1980; Carstens and 
Watson 1996; Marquardt 1972a, 1972b, 1974; 
Marquardt and Watson 1976, 1983; Robbins 
1971; Wagner 1976; Watson et al. 1969; Wat- 
son 1974). Whether you call it explanatory ar- 
chaeology, processual archaeology, or even 
post-processual archaeology, the post-1960 era 


of interdisciplinary archaeological researc): in 
the central Kentucky karst began to answer 
many questions about the area’s prehistory anc 
the avenues of cultural change and adaptation 
through time and space. 

Watson's archaeological work in MCNP be- 
gan in 1962 when, in conjunction with the 
Cave Research Foundation, the Illinois State 
Museum, and MCNP, she initiated an archae- 
ological reconnaissance of the large caves 
within the Flint Mammoth Cave system (Car- 
stens and Watson 1996: Watson et al. 1969:v). 
Watson’s initial work was carried out primarily 
in Salts Cave, but later research expanded into 
other caves (e.g., Mammoth, Lee, and Bluff), 
and to archaeological surface reconnaissance 
(Carstens 1974, 1980). Watson’s reason for 
studying the cultural materials from within the 
caves was that data derived from those mate- 
rials were highly relevant to the discovery of 
dietary practices during the early agricultural 
Late Archaic-Early Woodland period. Wat- 
son’s research was expanded in April 1969 to 
include, “excavation in Salts Cave Vestibule, a 
search for and testing of possible surface sites 
near Salts Sink, and recording of prehistoric 
remains in other caves within the Park” (Wat- 
son et al. 1969:v). Between 1973 and 1980, 
Watson and her colleagues initiated compara- 
tive studies at caves outside the park (e.g., Wy- 
andotte Cave, Indiana, and Wolf River or Jag- 
uar Cave, Tennessee) (Crothers 1986; Munson 
and Munson 1990; Robbins et al. 1981; K. 
Tankersley et al. 1994; S. Tankersley 1993). 
She also obtained a more complete radiocar- 
bon sequence from Salts and Mammoth caves, 
excavated and floated a stratigraphic column 
from Salts Cave vestibule; obtained pollen and 
parasitological analyses from human paleofecal 
specimens found within the cave, and took 
pollen core samples from nearby sinkhole 
ponds. 

Aided in her research by scientists from 
many different fields of study, Watson was able 
to approach the archaeological problems of 
the MCNP in a scientifically integrated man- 
ner, a methodological approach she had 
learned as a University of Chicago graduate 
participating in Robert Braidwood’s_interdis- 
ciplinary studies of agricultural origins in the 
Near East. This approach led to some answers 
and to many new questions, particularly with 
respect to environmental changes and their 


64 Journal of the Kentucky Academy of Science 62(1) 


possible effects on the prehistoric inhabitants 
of the study area. Watson and her colleagues 
(e.c. Munson et al. 1989: K. Tankersley 1996: 


K. Tankersley et al. 1985) documented that 
historic people using the cave were mining 
cave for minerals (e.g., mirabilite, gypsum, 
selenite, and satin spar) and were simply ex- 
ploring the cave system. She noted similar pat- 
terns of cultural ‘eae s in portions of other 
caves located inside (Lee and Bluff) and out- 
side the park boundaries (e.g., Wyandotte Cave 
in Indiana, and Big Bone and Jaguar caves in 
Tennessee: Crothers 1986; Munson and Mun- 
son 1990; Watson 1986:109-116).Although 
Lee, Bluff, Wyandotte, and Jaguar caves are 
not comp: arable in size to either Salts or Mam- 
moth Cave, the data collected by Watson and 
her colleagues clearly indicate that cave min- 
ing and exploration were widespread activities 
in this karstic region that probably began dur- 
ing the Late Archaic (Crothers et al. n.d.; 
Munson et al. 1989: Watson 1986; Watson 
1974:221-232: Watson and Kennedy 1991). 
The 50+ radiocarbon dates now available for 
the Mammoth Cave archaeological project 
clearly demonstrate the widespread prehistor- 
ic use of caves over a very important and sim- 
ilar time horizon (Kennedy 1990, 1996): that 


of early agriculture in the ‘Late Archaic- Early 


Woodland period (ca. 4000 to 2000 B.P.). 

Watson's research between 1962 and 1980 
in the Mammoth Cave region is unique for 
two reasons: (1) it is the er time such sci- 
entifically integrated archaeological deep-cave 
research i been attempted in the eastern 
U.S.; and (2) it provides an aspect of prehis- 
toric ae process that was extremely im- 
portant (i.e., the domestication of native 
plants, evidence for which was not then being 
found in “open” surface sites in the eastern 
U.S.). Watson’s research continues in the 
Mammoth Cave region. 

As a part of the Watson research team be- 
tween 1973 and 1975, my job was to docu- 
ment the culture history evident in a sample 
of 83 surface sites in and around MCNP., ex- 


amining the techno-economies of several of 


those sites diachronically and presenting a cul- 
es historical context within which the sur- 
face dwellers of the central Kentucky karst ex- 
plored and exploited the large caves (Carstens 
1980). 


Between 1977 and 1987, MCNP witnessed 


only intermittent archaeological research on 
the park’s surface archaeology. Most of the 
work accomplished included small, unrelated 
cultural resource management surveys (e.g., 
Beditz 1979, 1981: Carstens 1977, 1978). Be- 
tween 1981 and 1989, few archaeological pro- 
jects were conducted on the surface of MCNP. 
Exceptions are the work of Philip J. DiBlasi, 
who investigated the 1920s homestead of 
Floyd Collins (DiBlasi 1957a), a famous local 
cave explorer, and George Crothers, who doc- 
umented material left in Sand Cave where 
Collins died in 1925 (Crothers 1981, 1983). 
DiBlasi (1987b, 1996), working with the Cave 
Rese on Foundation, also found in Salts Cave 
a series of prehistoric pictographs and glyphs 
previously undocumented. Other studies con- 
cerning the human use of the cave system, and 
of the people who were using the cave, fo- 
cused on determining the exact nature of pre- 
historic mineral procurement (K. Tankersley 
1996), forensic examinations of the historic 
findspots of mummies (K. Tankersley et al. 
1994: S. Tankersley 1993), and a new and ex- 
citing search for pathogenic microorganisms in 
prehistoric and historic human feces and bodi- 
ly fluids (Ruppert 1994; S. Tankersley 1993). 

The most systematic undertaking to inven- 
tory a representative sample of the park for 
both historic and prehistoric cultural resources 
was directed by NPS archeologist Guy Pren- 
tice (1993). Although much of Prentice’s pre- 
history is a summary of Nelson (1917), 
Schwartz (1958a-g), Carstens (1980), and 
Watson and Carstens’ (1982) site inventories, 
Prentice adds new prehistoric and _ historic 
sites to the overall resource inventory of the 
park. As a result, Prentice (1994, 1996) was 
able to offer a settlement synopsis of MCNP 
for his doctoral dissertation that includes a hy- 
pothetical seasonal round between the Big 
Bend shell mound area and the Mammoth 
Cave area. 

In 1993, the Science and Resource Man- 
agement Division at MCNP began a long- 
term cultural resource inventory of all artifacts 
(historic and prehistoric) within the main cave 
in Mammoth Cave. This project is co-spon- 
sored by the NPS and Earthwatch; the field 
work for this project has been directed by Ken 
Tankersley, Mary Kennedy, George Crothers, 
Christine Hensley, and Bob Ward (Kennedy 
1993: Crothers and Ward 1995). Using an 


Archaeological Research in the Mammoth Cave Area—Carstens 


65 


Table 1. Archaeological Research in the Area of Mammoth Cave National Park, 1824 to the Present (not an exhaustive 


list). 


Speculative-pre-1915 
Classificatory 
1916-1970 


Explanatory-Interdisciplinary 
1971 to the present 


Rafinesque (1824), Young (1910) 

Carey (1942); Fowke (1922): Funkhouser and Webb (1932); Hanson 
(1960); Meloy (1968); Moore (1916): Nelson (1917, 1923); Neumann 
(1938); Pond (1935, 1938): Schwartz (1958a—h: 1960a, 1960b, 1965, 
1967); Schwartz and Hanson (1961); Schwartz and Sloan (1958, 1960a, 
1960b); Schwartz, Sloan, and Hanson (1960). 

Beditz (1979, 1981): Carstens 1974, 1975, 1976, 1977, 1978. 1980): Car- 
stens and Watson (1996): Crothers (1981, 1983): Grothers et al. n.d.: 
Crothers and Ward (1995); DiBlasi (1987a, 1987b, 1996): Duffield 
(1974); Hensley (1995, 1996); Kennedy (1990, 1993, 1996); Kennedy and 
Watson (1997); Marquardt (1974); Molnar and Ward (1974); Munson et 
al. (1989); Prentice (1993, 1994, 1996): Robbins (1971, 1974, 1980): 
Robbins et al. (1981); Ruppert (1994); Schoenwetter (1974); K. Tank- 
ersley et al. (1994); S. Tankersley (1993); Wagner (1976); Watson (1974, 
1986, 1992); Watson et al. (1969); Watson and Carstens (1975, 1982): 


Watson and Kennedy (1993). 


Electronic Distance Measurement (EDM) 
system, the Earthwatch team records the exact 
location of every artifact noted within the sur- 
veyed areas of the cave system. This makes it 
possible to prepare density plots of aboriginal 
activity within the cave system and to deter- 
mine prehistoric use areas within the cave de- 
spite 200 years of historic cave use and cave 
disturbance to the aboriginal materials in 
Mammoth Cave. 

In 1992, Patty Jo Watson, Mary Kennedy, 
Kristen Gremillion, and Kristin Sobolik began 
a new study in Mammoth and Salts caves. This 
new arena emphasized the collection of hu- 
man paleofecal samples for radiocarbon dat- 
ing, parasitological analysis, macro- and mi- 
croethnobotanical studies, and _ biochemical 
(hormonal) analysis. These studies would al- 
low prehistoric fecal specimens to be sexed 
and thereby enable a better understanding 
about specific individuals who explored and 
mined prehistoric Mammoth Cave (Watson 
1992; Watson and Kennedy 1993). 

In 1994, Christine Hensley and Tom Sus- 
senbach, while working for the NPS in MCNP, 
conducted excavations at the stairway rock- 
shelter (Hensley 1995, 1996). The three radio- 
carbon samples from Hensley’s excavations of 
Feature 1 consistently place the site’s occu- 
pation in the Early Woodland period, ca. 2170 
to 2570 B.P., a date range quite comparable 
to the majority of aboriginal use of Mammoth 
and Salts caves (Hensley 1995:24, Table’ 1; 
Kennedy 1990, 1996). Further, Paul Gardner 
identified more than 5000 seeds of domestic, 


semi-domestic, and wild chenopodium from 
the nackeineliver occupation floor (Hensley 


1995, 1996). Similar contractual ar chaeological 


studies continue today through the supervision 
of Bob Ward, cultural resource specialist at 
MCNP and the assistance of Darlene Appel- 
gate, archaeologist at Western Kentucky Uni- 
versity (Appelgate pers. comm. 1 July 2000). 
Analytically, the Explanatory/Interdisciplin- 
ary (post-1960) era of archaeological research 
has answered many questions about the Na- 
tive Americans who explored and exploited 
the environment and resources above and be- 
low ground in the central Kentucky karst. Ini- 
tial studies by Watson and her colleagues in 
the early 1960s through 1980s focused upon 
time-space and environmental reconstruction 
sequences, then turned to more processual is- 
sues while unraveling the prehistory of the 
Mammoth Cave area. That work inspired oth- 
er archeologists in Tennessee and Indiana to 
test several of Watson’s observations and con- 
clusions about the prehistory of Mammoth 
Cave specifically, and prehistoric cultural pro- 
cesses in general, finding that prehistoric ab- 
original mining and exploration was a wide- 
spread cultural phenomenon; it was not lim- 
ited solely to MCNP or only to the Late Ar- 
chaic-Early Woodland transition (e. g., 
Faulkner 1986). Furthermore, not only had 
Watson and her associates confirmed and ex- 
panded N. C. Nelson's initial observations 
about the importance of plant cultivation in 
the Mammoth Cave area, but they also went 
far beyond, examining human paleofeces for 


66 Journal of the Kentucky Academy of Science 62(1) 


parasites and micro-organisms and determin- 
ing w) ther prehistoric caving activities were 
cearricd out by both sexes, chereen helping to 
and more accurately describe, 
Cave prehistory. 


envy nde rH we 


\iammoth ¢ 


DISCUSSION 


The history of archaeology in the Mammoth 
Cave area closely parallels the growth and de- 
velopment of f archae ‘ological trends in North 
America. From the Speculative to ie Classi- 
ficatory to the Explanatory and Interdisciplin- 
ary pe sriods, archaeological inquiry in the cen- 
tral Ke ntucky karst hee demonstrated that the 
uncommon preservation characteristics of the 
cave environment provides unique insight into 
human behavior (prehistoric and historic) that 
may be more accessible than from open “sur- 
face” sites. Table 1 also reflects that, along 
with an increase in the intensity of cave ar- 
chaeological investigation, a greater sophisti- 
cation began once “interdisciplinary research 
was initiated at the park (post-1960). That 
work was initiated by Patty Jo Watson and her 
associates and colleagues. 

Early turn-of-the-century interests in cave 
archaeology, primarily atheoretical, prompted 
work at surface sites in the Mammoth Cave 
area and led to speculation about the origins 
of prehistoric cultural materials, sometimes 
comparing them to the better-understood Eu- 
ropean record (e.g., Nelson 1917; Young 
1910).Assessing significance of and attempting 
to order archaeological sites from within the 
park area, both Anode ground and below, and 
assessing Mammoth Cave collections held out- 
side chee park, were the foci of the Classifica- 
tory Period between 1916 and 1970, culmi- 
nating in the summaries of Douglas W. 
Gahennts (195Sa—h, 1960a, 1960b, 1965). 
More recent work by Watson and her col- 
leagues brought a theoretical and interdisci- 
plinary framework to the archaeology of the 
pé ark often reflecting various Themis promi- 
nent in the “New Archaeology.” Within the 
last decade cave art studies (DiBlasi 1996) 
have added a cognitive, or post-processualist 
research slant to fhe efforts begun by Watson’s 
group. However, additional Pacenrch is still 
needed in the Mammoth Cave area, above 
ground and below. Only the tip of the prover- 
bial iceberg at Mammoth Cave has been stud- 
ied, whereas artifacts from surface sites and 


from within the caves continue to be damaged 
or stolen in spite of the security efforts of the 
NPS. New. insights and new energies are 
needed today to carry on the fascinating study 
of the aboriginal and Euro-American people 
who explore A and exploited the Kentucky un- 
derworld. 


ACKNOWLEDGEMENTS 


Earlier readings of a shorter version of this 
paper were made during the 1970s by my doc- 
toral committee at W ashington University, St. 
Louis. For their comments I am truly thank- 
ful. This paper also has benefitted from con- 
structive comments made by Kenneth Sassa- 
man at University of South Carolina and Patty 
Jo Watson at W ‘ashington University, St. Louis. 
I thank the officials at MCNP for their past 
and continued support of our archaeological 
research at the park and the Department of 
Geosciences at Murray State University. My 
wife, Nancy Son Carstens, read and com- 
mented on earlier versions of this paper, 
thereby greatly improving it; however, errors 
and omissions in this paper remain mine 
alone. 


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J. Ky. 


Acad. Sci. 62(1):70-76. 2001. 


Jsing Composts as Growth Media in Container Production 
of Tomatoes 


Brian D. Lacefield and Elmer Gray 


Department of Agriculture, Western Kentucky University, Bowling Green, Kentucky 42101 


ABSTRACT 
During the 1996 and 1997 growing seasons, soil (S), brush composts (B), leaf composts (L), N-Viro Soil 
(N), and 50:50 mixtures of these materials (S:B, S$:L, S:N, B:L, B:N, L 
for their effects on seasonal distribution and total production for four tomato (Lycopersicon esculentum) 


:N) by volume were compared 


cultivars. Each growth medium was replicated in four containers (55 cm diameter, 38 em depth; ca. 64 liters 
‘Celebrity’, Red Cherry’, and ‘Small Red Cherry.’ 
based upon weekly production of vine-ripened fruits by each plant from mid-July to mid-October. The growth 


capacity). The cultivars were ‘Patio’, ‘Large Results were 
media were similar in their effects on seasonal distribution and total fruit production. All growth media/ 
cultivar combinations gave continuous tomato production throughout the growing season. Cultivar differ- 
ences were exhibited in both plant and fruit characteristics, permitting the container gardener to practice 


individual preferences. Overall, the results support the practice of composting waste products for use in 


container gardening. 


INTRODUCTION 


Sustainability of society is enhanced when 
recurring Ww aste products are effectively sub- 
stituted for diminishing natural resources. The 
sustainable dimension of food production is 
receiving public attention and tangible govern- 
ment support (Hudson and Harsch 1991). 

Composting is an age-old process for con- 
verting organic residues into forms that are 
more aesthetically acceptable and more avail- 
able for plant utilization. Composting is re- 
ceiving renewed support as a means of waste 
disposal because the process reduces the vol- 
ume of yard waste by a factor of five times or 
more and results in a product suitable for gar- 
den or landscape utilization (Fine 1989). In 
1992. 85% of the 4.6 million tons of solid 
waste produced in Kentucky were disposed in 
landfills. The Environmental Protection Agen- 
cy set a goal to reduce municipal solid waste 
going into landfills by 25% (Environmental Al- 
manac 1994). Since yard waste accounts for 
ca. 18% of municipal solid waste, composting 
is critical to efficient waste management. 

Organic gardening is based upon the sub- 
stitution of organic sources for inorganic 
sources of nitrogen. Recent increases in de- 
mand for organically grown food have expand- 
ed markets for composted organic matter. Or- 
ganic gardening varies in scale from individual 
plant containers to full-sized family gardens 
(Lindgren et al. 1990). 


70 


Urbanites have become increasingly in- 
volved in gardening during recent years. They 
utilize small plots or containers to provide on- 
going supplies of fresh fruits and vegetable. In 
Seletieann: gardening serves as a hobby that for 
many people is a source of pleasure, pride, 
and satisfaction. These benefits are not depen- 
dent on garden size (Bartholomew 1981). 

Tomato (Lycopersicon esculentum) is the 
crop of choice by most urbanite gardeners. 
Research has shown that tomato yields from 
litter-enriched plots matured earlier and were 
larger than those grown in commercially fer- 
tilized plots (Brown et al. 1995). 

The present study was part of an ongoing 
program sponsored by the Department of Ag- 
riculture, Western Kentucky University 
(WKU), to convert local municipal waste into 
forms usable in gardening and landscaping. 
Our objectives were to compare different 
waste composts for their effects on total and 
seasonal distribution of production of diverse 
tomato cultivars grown in containers. 


MATERIALS AND PROCEDURES 
Composts 


Three locally available waste products were 
evaluated. Brush- and leaf- -composts were pro- 
duced from Bowling Green yard waste col- 
lected in 1994. In a tree inventory of Bowling 
Green, Martin (1994) identified a variety of 
common deciduous trees including ash, elm, 


Container Production of Tomatoes—Lacefield and Gray ral 


Table 1. Nutritive value of composts used in container 
production of tomatoes. 
% Carbon: 
“Nitrogen 
Compost N P K Ratios 
Brush (B) 0.99 0.08 0.48 30:1 
Leaf (L) 0.88 0.04 0.42 Soul 
N-Viro soil (N) 0.96 0.36 0.67 16:1 


dogwood, maple, mulberry, oak, redbud, and 
willow as well as numerous ornamental trees. 
The sewage compost, known by the trade 
name N-Viro Soil (Kovacik 1988), is prepared 
by mixing treated sewage sludge with cement 
kiln dust according to approved procedures. 
The resulting moist mixture is aerated and 
composted on an environmentally approved 
site. The Pembroke silt loam soil, obtained 
from the WKU farm, tested medium to high 
in both phosphorus and potassium. 


Containers 


The study was based on container-culture to 
permit greater experimental control of the 
compost mixtures and to extend the applica- 
bility of the results to container gardening. 
Forty plastic barrel sections (58 cm diameter, 
38 cm depth) were located in an unshaded 
area. Containers were embedded in the soil to 
reduce drying and were punctured in the bot- 
tom to permit drainage. They were spaced 1.5 
m apart in 5 rows and 8 columns. Each con- 
tainer received ca. 64 liters of compost mix- 
ture. 


Growth Media 


Soil and the three composts were usec 
make 10 growth media; four consisted 
100% each of soil (S), brush compost (B), lea’ 
compost (L), and N-Viro Soil (N), and six con- 
sisted of 50:50 combinations by volume of the 
soil compost materials (S:B, S:L, S:N, B:L, 
B:N, L:N). All growth media were supple- 
mented prior to transplanting with fertilizer at 
the rate of 56.0, 24.5, and 46.5 kg/ha-t of N, 
P, K, respectively. No additional fertilizer was 


applied. 


Cultivars 


Four diverse tomato cultivars— Celebrity’, 
‘Patio’", ‘Large Red Cherry’, and ‘Small Red 
Cherry —were studied. ‘Celebrity’ is an inde- 
terminate garden type that produces large 
plants and large fruits. ‘Patio’ exhibits a com- 
pact, determinate growth habit and is more 
suitable for urban or “patio” production. 
‘Large Red Cherry and ‘Small Red Cherry 
are characterized by indeterminate growth 
habit and smaller fruits. 


Production 


One plant, ca. 15 cm tall, was transplanted 
in mid-May to each container. Vine-ripened 
fruits were harvested twice a week beginning 
in mid-July and continuing to mid-October. 
Fruits from each plant were counted and 
weighed. Data from the two harvests per week 
were combined and reported as production on 
a weekly basis. After the last regular harvest, 


‘ 
Table 2. Season production of tomato fruits per plant in 1996, Bowling Green, KY. 


Cultivar? 
Growth 
Media! C iP Mean® C P Mean 
Number Weight (kg) 

S 100 o2 76 13.3 497 9.14 
B 64 68 66 9.55 6.96 8.26 
L 92 58 75 15.34 3.6 9.47 
N 87 By) 70 12.07 5.4 8.74 
S-B 87 65 76 13.41 5.92 9.66 
S-L 99 66 82 15.64 7.35 11.5 
S-N 74 68 fall 10.3 6.52 S41 
B-L 120 Bil 88 125 SUDIL 11.38 
B-N 136 64 100 17.25 5.94 11.6 
L-N 98 83 90 13.29 6.79 10.04 
Mean® 95.7a 63.3b 79.4 13.74a 5.89b 9.82 

Ԥ = soil, B = brush compost, L = leaf compost, N = N-Viro Soil: 

>C = ‘Celebrity’, P = ‘Patio’. 

3 Growth mixture means were not significantly different (P > 0.05); Cultivar fruit number and fruit weight means followed by the same letters are not 


significantly different (P > 0.05). 


72 Journal of the Kentucky Academy of Science 62(1) 
Fruits/Plant 
ane eh) Serer of eee ee 
14 es * ‘Celebrity’ 
= +' iat | | 
aes! |+'Patio' | | 
12 f 
e | 
_2— ¥ 
10 } - \ —— { 
| 4 o— ——— } 
8 } = vA \ \ 
of / \ 
6 J ax . ; 
4 | ° oe aT 
Ts ae 
2 es y rs ~} te 
hae 
0 a — = — —— 
1 2 3 4 5 6 7 8 9 10 11 12 
Week 
Figure 1. Season distribution of number of fruits/plant for ‘Celebrity’ and ‘Patio’ tomatoes in 1996, Bowling Green, 
Kentucky. 
remaining green fruits were counted and In 1997, the addition of “Large Red Cherry 


weighed. 
Supplemental water 
once or twice per week to prevent plant wilt- 


ing. Support stakes placed in the periphery of 


The containers were connected with loose-fit- 
ting twine to provide plant support. Plants 
were permitted to spread without any restric- 
tive pruning. 
Design and Analysis 

The 10 growth media were replicated four 
times in a randomized complete block design 
(Steel and Torrie 1980). In 1996, ‘Celebrity’ 


and ‘Patio’ were studied, resulting in two rep- 
lications of each cultivar per growth medium. 


Fruit/Plant (kg.) 


was applied usually 


and ‘Small Red Cherry’ resulted in one repli- 
cation for each cultivar per growth medium. 
Data analyses were directed toward both sea- 
sonal distribution and total production of 
number and weight of fruits. 


RESULTS 
Compost Analysis 


Nutritive values for the brush- and leaf- 
composts were about equal for N (ca. 1%), P 
(trace), and K (0.45%), whereas N-Viro Soil 
had a similar level of N (ca. 1%) but higher 
levels of P (0.56%) and K (0.67%) (Table 1). 
Variance among the C:N ratios was the most 
important difference in nutritive value of the 


-©'Celebrity’ 
+ 'Patio' 


‘ a) 
Figure 2. 
Kentucky. 


Week 


Season distribution of weight of fruits/plant for ‘Celebrity and ‘Patio’ tomatoes in 1996, Bowling Green, 


Container Production of Tomatoes—Lacefield and Gray 


Table 3. Season production of tomato fruits per plant in 1997, Bowling Green, KY. 


Cultivar 


Growth — ee See = 
Mixture! (e 1p LRC SRC Mean® Cc P LRC SRC Mean 
Number Weight (kg) 

S 60 ol 525 940 394 5.15 2.42 8.18 4.0] 4.94 
B dl 76 67 1082 319 7.19 3.2] 1.26 a2 4,22 
L 9] 34 383 710 304 9.68 4.44 5.94 4 6.02 
N 73 34 408 980 374 10.17 3.84 6.34 4.54 6.22 
S-B 45 60 362 765 308 7.06 6.08 6.06 4.13 5.96 
S-L 80 28 270 786 291 9.31 1.91 4.19 3.92 4.83 
S-N 47 Al 320 1017 356 5.2 SD) De, 4,92 4.7 
B-L 92 45 344 794 319 11.68 3.69 6.57 4.03 6.49 
B-N 39 40 457 1077 403 4.69 3.76 7.79 5.86 Day 
L-N 57 32 353 899 335 8.91 3.79 5.71 3.88 5.56 
Mean? 64c 44Ac 349b 905a 340 7.95a 3.66b 5.72ab 4.45b 5.45 

'S = Soil, B = Brush Compost, L = Leaf Compost, N = N-Viro Soil. 

2 ¢ = ‘Celebrity’, P = ‘Patio’, LRC = ‘Large Red Cherry’, SRC = ‘Small Red Cherry’. 

5 Growth mixture means were not significantly different (P > 0.05); Cultivar fruit number and fruit weight means followed by the same letters are not 


significantly different (P > 0.05). 


composts. Ratios of 30:1, as exhibited in the 
brush- and leaf-composts, immobilize avail- 
able N and result in N starvation of the plants. 
The C:N ratio of 16:1 for the N-Viro Soil 
should not create N deficiency in plant 
growth. The recommended level of fertilizer 
for tomato production was applied to all 
growth mixture to compensate for nutrient 
differences. 


1996 Season Production 

Number and weight of fruits per plant are 
given in Table 2. ‘Celebrity plants produced 
significantly more fruits (ca. 96) than did ‘Pa- 
tio’ plants (ca. 63). Average fruit weight per 


‘ 


plant was significantly greater for ‘Celebrity 
(13.74 kg) than for “Patio’ (5.89 kg). Thus, ‘Ce- 
lebrity’ plants produced significantly more and 
heavier fruits than did ‘Patio’ plants during 
1996. Although there was variability among 
the number and weight of fruits produced on 
the different growth media, yield differences 
associated with growth media were not signif- 
icant. Analyses comparing groups of means, 
i.e., those mixtures including soil vs. those that 
did not, failed to detect any significant differ- 
ences. Also, comparisons of growth media 
means at different harvest dates during the 
season failed to reveal any consistent differ- 
ences. 


Fruits/Plant 
(0) 


-@'Celebrity’ 
+'Patio' 


Figure 3. 
Kentucky. 


Week 


Season distribution of number of fruits/plant for ‘Celebrity’ and ‘Patio’ tomatoes in 1997, Bowling Green, 


74 Journal of the Kentucky Academy of Science 62(1) 


Fruits/Plant 
140, _——_—_ ———___—— 
a: @LRC’ 
|+'SRC’ 
120 7 +'SRC 
+— 
100 y ae 
7 
80 + ¥ ~ 
+ er ——+. + 
60 # 7 * ~ 4 
° a 
40 s 
2 
+ ~ A 
20 - \+ ~ ——— ss 
es e . So 
+ 
0 ~——--— + -—---- -— = ~ 
1 2 3 4 5 6 7 8 9 10 1 12 
Week 
Figure 4. Season distribution of numbers of fruits/plant for “Large Red Cherry’ (LRC) and ‘Small Red Cherry’ (SMC) 


tomatoes in 1997, Bowling Green, 


Kentucky. 


Since the growth media means did not dif- 
fer significantly, yields were combined for 
each cultivar in determining season distribu- 
tions (Figures 1, 2). Fruit ripening began in 
mid-July and continued until frost acoured in 
mid-October. Peak production for both num- 
bers and weights of fruits occurred in late Au- 
gust and poukaed into early September. ‘Ce- 
lebrity’ produced more and heavier fruits per 
plant: than did ‘Patio’ during the second half 
of the season. 


1997 Season Production 


Number and weight of fruits per plant for 
each of the four auley ars are presented in Ta- 


Fruit/Plant (kg) 


ble 3. Both the number and weight of fruits 
per plants of “Celebrity and ‘Patio’ were lower 
in 1997 than in 1996. For number of fruits per 
plant, “Small Red Cherry’ was highest (905), 
‘Large Red Cherry’ was intermediate (349), 
and “Celebrity and ‘Patio’ were lowest (64 and 
44. respectively). Growth mixtures had no sig- 
nificant effect on number of fruits. For weight 
of fruits per plant. ‘Celebrity was highest 
(7.95 ke), ‘Large Red Cherry was intermedi- 
ate (5 72 kg), and ‘Patio’ and “Small Red Cher- 
ry were lowest (3.66 and 4.45 kg, respective- 
ly). Differences among number or weight of 
fruits were not influenced significantly by 
growth mixtures when such comparisons were 


© 'Celebrity’ 
2 ea ae en ae + 'Patio' [a 


Figure 5. 
Kentucky. 


Week 


Season distribution of weight of fruits/plant for ‘Celebrity’ and “Patio” tomatoes in 1997, Bowling Green, 


Container Production of Tomatoes—Lacefield and Gray 


5 Fruit/Plant (kg) 


*LRC 
_ [SRC] 


Figure 6. Season distribution of weight of fruits/plant for “Large Red Cherry’ (LRC) and ‘Small Red Cherry’ 


tomatoes in 1997, Bowling Green, Kentucky. 


based upon either total season or individual 
harvest period yields. 

Growth mixture yields were combined for 
each cultivar in determining season distribu- 
tions (Figures 3-6). Fruit ripening continued 
from mid-July throughout mid-October. For 
‘Celebrity and ‘Patio’ the 1997 season distri- 
butions of both number and weight of fruits 
per plant were similar to those in 1996 with 
the exceptions that ‘Patio’ production peaked 
earlier in 1997 than in 1996. “Large Red Cher- 
ry and ‘Small Red Cherry’ produced an abun- 
dance of fruits throughout 1997. Peak produc- 
tion occurred at week 6 (end of August) for 
‘Small Red Cherry’, but no single period of 
maximum production was exhibited by ‘Large 
Red Cherry. For weight of fruits per plant, 
yields were rather consistent for ‘Small Red 
Cherry throughout the season. whereas 
weight of fruits per plant decreased during 
early September for “Large Red Cherry.’ 


End of Season Production 


Following the first frost each year the un- 
ripened fruits on each plant were counted and 
weighed. In 1996, ‘Celebrity and ‘Patio’ av- 
eraged 12.8 and 22.6 fruits weighing 0.68 and 
0.76 kg per plant, respectively. In 1997, “Ce- 
lebrity’, ‘Patio’, “Large Red Cherry’, and ‘Small 
Red Cherry averaged 13.4, 23.7, 60.8, and 
185.6 fruits weighing 0.72, 0.78, 0.48, and 0.51 
kg per plant, respectively. 


Week 


(SMC) 


DISCUSSION AND SUMMARY 


These results indicate that composted com- 
mon waste products, either alone or in mix- 
ture, are suitable for growing tomatoes in con- 
tainers. Since compost materials vary in nutri- 
tive value and C:N ratios, composted g growth 
media need to be supplemented with a com- 
plete fertilizer (N, P, K) for protection against 
nutrient deficiencies. 

Cultivar selection is an important consider- 
ation in container production of tomatoes. Our 
present results indicate that cultivar charac- 
teristics such as plant shape, plant size, and 
fruit number and weight are consistent wheth- 
er grown in an open garden or in containers. 
The four cultivars exhibited different qualities 
for container gardening. All cultivars produced 
throughout the season and supported unrip- 
ened fruits at the time of frost, thereby pro- 
viding a continuous supply of fruits. ‘Celebrity’ 
was highly productive of large fruits as desired 
for some purposes. However, its large, open 
plants could be unsightly and problematic in 
the urban landscape. ‘Patio’ was intermediate 
in productivity and fruit size. Its compact plant 
size would be desirable in a “patio” setting. 
‘Large Red Cherry’ produced large numbers 
of Aredia sized fruits throughout - the season. 
Its spreading, open plant shape could be a lim- 
itation in an urban setting. “Small Red Cherry 
was a prolific producer of small fruits through- 
out the season. However, smallness in fruits 


76 Journal of the Kentucky Academy of Science 62(1) 


and the vine-like plant type could limit its util- 
ity. 
“These preliminary results provide encour- 
ac ent for using municipal waste products 
beneficial purposes. The finding that the 
different composts alone and in mixtures were 
equally effective in producing tomatoes sug- 
gests that a variety of composted waste prod- 
ucts may be used successfully in gardening. 


LITERATURE CITED 


Bartholomew, M. 1981. Square foot gardening. Rodale 
Press, Emmaus, PA. 

Brown, I. E., C. H. Gilliam, R. L. Shumack, D. W. Porch, 
and I. O'Donald. 1995. Comparison of broiler litter and 
commercial fertilizer on production of tomato. J. Veg. 
Crop Prod. 1(1):53-62. 


Environmental Almanac. 1994. A. Hammond (ed). World 
Resource Institute, Houghton Mifflin, Boston, MA. 
Fine, S. 1989. Composting nature's garbage. World Watch 

2(1):5-6. 

Hudson, W. J., and J. Harsch. 1991. The basic principles 
of sustainable agriculture. Coop. States Res. Serv., 
U.S.D.A., Washington, DC. 

Kovacik, T. L. 1988. Sludge kiln dust makes fertilizer. Wa- 
ter Engin. & Managem. (December):1-2. 

Lindgren, D. T., D. H. Steinegger, F. P. Baxendale, and 
]. E. Watkins. 1990. Organic gardening in the backyard. 
Univ. Nebraska Coop. Ext. Serv, 681-548. 

Martin, J. A., 1994. Street trees for Bowling Green, Ken- 
tucky, Street Tree Inventory Program of the Kentucky 
Division of Forestry. Division of Forestry, Department 
for Natural Resources, Frankfort, KY. 

Steel, R. G. D., and J. H. Torrie. 1980. Principles and 
procedures of statistics, Ind ed. McGraw-Hill, New 
York, NY. 


]. Ky. Acad. Sci. 62(1):77-78. 2001. 


BOOK REVIEW 


Gary E. Dillard. 1999. Common Freshwater 
Algae of the United States: An Illustrated 
Key to the Genera (Excluding the Dia- 
toms). J. Cramer in der Gebr. Borntraeger 
Verlagsbuchhandlung, Berlin. 173 pages; il- 
lus. ISBN 3-443-50026-9. Price not indicat- 
ed. 


Professor Gary E. Dillard put together this 
book after many years of teaching courses on 
the biology of algae at Western Kentucky Uni- 
versity. His students were required to identify 
genus algae from field collections. He found 
that students had difficulty using published 
keys and descriptions because of their lack of 
familiarity with the technical terms used in 
these keys to describe algal morphology. He 
set out to produce this “user friendly” manual, 
which avoids as much as possible discipline 
specific language. His goal has not been fully 
accomplished; there is still much technical jar- 
gon in the book. The beginning student is un- 
likely to be familiar with terms such as lorica, 
epicone, hypocone, dendroid colony, and oth- 
ers found throughout this manual. A simple 
glossary would have been a most useful addi- 
tion. Yet, this is the first simplified key to 
freshwater algae to be published since Pres- 
cott’s How to Know the Freshwater Algae 
(1978), which today is hard to obtain. 

The book starts with a statement of purpose 
and a definition of “algae” as representing a 
heterogeneous assemblage of oxygenic photo- 
autotrophs that lack tissue differentiation and 
contain chlorophyll a. Based on this definition, 
algae include prokaryotic groups (the cyano- 
bacteria and chloroxybacteria) as well as a 
wide variety of phylogenetically unrelated eu- 
karyotic groups. In this section the traditional 
and modern systems of classification are brief- 
ly mentioned. Since the purpose of the book 
is to act as a key to identify algae only to the 
generic level, Dillard did not find it necessary 
to place the genera into higher categories (di- 
vision, classes, orders). By doing this he also 
avoids discussing the recent dramatic and 
sometimes confusing changes in algal classifi- 
cation. This first section of the book calls -at- 
tention to the extensive bibliography for those 
readers interested in phylogenetic relation- 


~l 
~l 


ships among the algae or in proceeding to spe 
cies identification. 

The next section, on algal habitats and col- 
lection methods, describes how to obtain qual- 
itative samples with, for example, plankton 
nets or artificial substrates, for the purpose of 
conducting a survey of algal forms. There is 
no description of standard quantitative meth- 
ods. 

The rest of the book is divided into nine 
sections where genera are grouped by artificial 
characteristics such as presence of flagella. By 
this system phylogenetically related taxa may 
not cluster together. The sections are I: Char- 
ales, plant-like genera; II: unicellular flagellat- 
ed genera; III: unicellular, non-flagellated gen- 
era; IV: colonial, flagellated genera; V: eolonmal 
non-flagellated genera; VI: unbranched fila- 
mentous genera; VII: branched filamentous 
genera; VIII: pseudoparenchymatous genera; 
and IX: coenocytic or sac-like genera. ‘Within 
each section, the dichotomous keys to the gen- 
era are easy to follow, and each genus is illus- 
trated with a detailed drawing. This manual 
excludes diatoms and many genera that occur 
largely in soil or aerial habitats. 

Although the title of the book indicates that 
it deals with algae “of the United States,” most 
genera of algae are cosmopolitan in distribu- 
tion and the book can find users worldwide. 
There is a growing global demand for identi- 
fication of freshwater algae. This demand is no 
longer restricted to academic circles and phy- 
cology classes. Identification of algae is a skill 
valued by drinking-water utilities whose op- 
erators are eaneeened with the presence of 
possible taste-, odor-, or toxin-producing spe- 
cies in their source water. It is also valued by 
environmental regulatory agencies that use al- 
gae as water quality nmgicatore Recently the 
U. S. Environmental Protection Agency (EPA) 
has added algae to its ~ Candidate Contami- 
nant List” (CCL). This will increase the de- 
mand for the identification of algae. Like the 
students enrolled in the author’s algae class, 
most people needing to identify common 
freshwater algae lack knowledge of the tech- 
nical jargon. Many of them currently use “pic- 
ture keys,” partic ularly color charts available 


78 Journal of the Kentucky Academy of Science 62(1) 


from t! EPA that even have species names timely. It will be useful to anyone needing to 


(not +s! genera) often associated with a pic- identify common freshwater algae to genus. 

ture. this commonly leads to misidentifica- Miriam So Kannan 

tioos since the important diagnostic features Department of Biological Sciences 
not learned when one does not follow a Northern Kentucky University 


vritten key. The publication of this manual is Highland Heights, Kentucky 41099 


]. Ky. Acad. Sci. 62(1):79-88. 2001. 


Abstracts of Some Papers Presented at the 2000 Annual Meeting of the Kentu. 4 
Academy of Science 


AGRICULTURAL SCIENCES 


Insecticides from wild tomato: phase I—breeding, tri- 
chome counts, and selection of tomato accessions. 
GEORGE F. ANTONIOUS, Department of Plant and 
Soil Science, Kentucky State University, Frankfort, KY 
40601. 

Among the thousands of secondary metabolites that un- 
derlie the characteristic properties of higher plants are a 
diverse assemblage of potentially toxic allelochemicals. Al- 
lelochemicals are those natural products that affect the 
growth, health, behavior, or population biology of mem- 
bers of other species. These compounds may impart a 
selective advantage to plants by inhibiting, repulsing, and 
even killing non-adapted organisms thar may feed upon 
or compete with the producing plant. Production of toxic 
chemical compounds is one method by which trichomes 
(leaf-hairs) can impart resistance. In the present investi- 
gation morphological and chemical characteristics of two 
glandular trichomes (type IV and type VI) of six wild to- 
mato accessions of Lycopersicon hirsutum ip hirsutum, 
four accessions of L. hirsutum f. glabratum, two acces- 
sions of L. pennellii, and density of type VI glandular tri- 
chomes of the commercial tomato L. esculentum cv. Fab- 
ulous, are reported. Densities of type IV and VI glandular 
trichomes varied among the accessions tested. Trichome 
counts on the leaf surface were correlated to three types 
of trichome exudates. Two methyl ketones (2-tridecanone 
and 2-dodecanone), two sesquiterpenes (zingiberene and 
elemine) and total glycolipids (sugar esters) in glandular 
trichomes have been quantified per unit leaf area. Iden- 
tifying the constituents of glandular trichomes was 
achieved using chemical methods. 


Residues of pyrethrins and piperonyl butoxide in soil, 
water and on potato leaves. GEORGE F\ANTONIOUS* 
and GAYATRI A. PATEL, Department of Plant and Soil 
Science, Kentucky State University, Frankfort, KY 40601; 
JOHN C. SNYDER, Department of Horticulture, Uni- 
versity of Kentucky, Lexington, KY 40546. 

Residues of pyrethrin- I (Py-1) and pyrethrin-II (Py-II), 
the major insecticidal components of the pyrethrum daisy 
(Tanacetum cinerariifolium), as well as residues of pipe- 
ronyl butoxide (PBO, a pyrethrum synergist), were deter- 
mined in soil and on potato foliage grown under field con- 
ditions. A pyrethrum formulation “Multi-Purpose Insec- 
ticide” containing the three active ingredients was sprayed 
twice at the rate of 6 Ibs per acre of formulated product 
(5.4 and 27.2 g ALI. of pyrethrins and PBO, respectively) 
on potato foliage during the growing season. In soil, three 
management practices (yard waste compost, grass filter 
strips, and a no mulch treatment) were used to study the 


impact of surface soil characteristics on the amount of 


pyrethrins and PBO retained in soil. Soil samples and po- 
tato leaves were collected at different time intervals after 


79 


spraying. Samples were purified and concentrated usii 

solid-phase extraction columns containing C,,-octadecy! 
bonded silica. Residues were quantified by high-perfor- 
mance liquid chromatograph equipped with a UV detec- 
tor. Following the first spray, the initial deposits were 0.18, 
0.40 and 0.99 g/g potato leaves for Py-I, Py-II, and PBO, 
respectively. Py-I and PBO residues in soil were higher in 
compost treatments compared to no mulch treatments. 


Effects of nitrogen fertilization on monoculture and bi- 
culture cover crops in vegetable production. G. R. 
CLINE,* A. F. SILVERNAIL, and K. KAUL, Community 
Research Service, Kentucky State University, Frankfort, 
KY 40601. 

A four year experiment examined how winter cover 
crops were affected by plus (+N) or minus (—N) 
inorganic nitrogen (N) 


previous 
fertilization of sweet com (Zea 
mays L.) and by kill dates associated with vegetable tillage 
methods. Hairy vetch (Vicia villosa Roth) yields and ni- 
trogen contents remained relatively constant in all years 
and were not affected by nitrogen treatments. In the +N 
treatment, yields of winter rye (Secale cereale L.) and a 
vetch/rye biculture normally exceeded vetch yields. Nitro- 
gen contents of vetch and biculture cover crops were gen- 
erally similar and were greater than those of rye. In con- 
trast to rye, vetch and biculture cover crop nitrogen con- 
tents were similar in +N and —N treatments. Delaying 
cover crop kill dates by eight days for no-till vegetables 
increased vetch yields but did not affect vetch nitrogen 
contents. In the biculture treatment, vetch competed bet- 
ter with rye as nitrogen availability decreased. Total yields 
of the biculture cover crop were generally greater in the 
+N than in the —N treatment. 


Assessment of genetic diversity within pawpaw (Asimi- 
na triloba) patches. SHERI B. CRABTREE,* TERA M. 
BONNEY, SNAKE C. JONES, and KIRK W. POMPER, 
Land Grant Program, Kentucky State University, Frank- 
fort, KY 40601- 9335. 

The pawpaw (Asimina triloba) is a native tree fruit 
found in the southeastern and midwestern United States: 
it has great potential as a new high-value crop in these 
areas. Kentucky State University is the site of the USDA 
National Clonal Germplasm Repository for Asimina spp.. 
and our long-term goal is to develop a sampling strategy 
to assess levels of genetic diversity in pawpaw across its 
native range. The objective of this study was to determine 
the level of genetic similarity among trees in pawpaw 
patches. Since pawpaws sucker profusely from the roots, 
our hypothesis is that pawpaw patches are at least partially 
clonal. Pawpaw leaf samples were collected from three 
different patches in Franklin County, Kentucky, in May 
1999. DNA was extracted from the leaves, then analyzed 
using either the inter simple sequence repeat PCR (ISSR- 
PCR) or the random amplified polymorphic DNA meth- 


SO Journal of the Kentucky Academy of Science 62(1) 


odology 1e ISSR-PCR primer (UBC 855) yielded poly- 
morphic markers in a subset of samples from patch #1 
and» }\ APD primer (OPA-11) yielded polymorphic mark- 
ers . subset of samples from patch #2. The identifica- 
ti { marker polymorphisms suggests that the pawpaw 


‘ches are not completely clonal. Primer screening and 
uch evaluation will continue. 


Influence of Cry9C Bacillus thuringiensis transformed 
corn kernels on two stored product moth pests in the lab- 
oratory. JEROME R. FAULKNER,* ANTHONY M. 
HANLEY, BRYAN D. PRICE, and JOHN D. SEDLA- 
CEK, Land-Grant Program, Kentucky State University, 
Frankfort, KY 40601. 

The Indian meal moth (IMM) and Angoumois grain 
moth (AGM) are global pests of stored grains. Growing 
concern regarding increasing resistance development to 
insecticides by insects, perceived risks associated with res- 
idues in foods, the Food Quality Protection Act of 1996, 
and Montreal Protocol have focused research efforts on 
alternative methods of pest control. Relatively recent de- 
velopments in pest management of corn utilize transgenic 
plants (rDNA-modified) containing Bacillus thuringiensis 
(Bt) toxin genes. Bt CrylA(b) or Cry9C delta-endotoxin is 
present in various tissues of transformed corn plants. It 
was found recently that corn kernels of some CrylA(b) Bt 
isolines cause reduced IMM and AGM emergence and 
egg production. Some Cry9C isolines have been examined 
for survivorship and development of IMM. However, fe- 
cundity of IMM and all life history parameters of AGM 
need to be examined. Thus, the objective of this research 
was to quantify the effects of Cry9C transformed corn 
kernels on IMM and AGM life histories. Experiments 
were conducted at 27 + 1°C and =60% RH using Star- 
link® Bt corn kemels (AgrEvo), its non-Bt isoline, and 
topically applied Bt to the non-Bt kernels. Fifty eggs were 
placed in ventilated pint jars containing 170 g of cracked 
or whole corn for IMM and AGM, respectively. Adult 
emergence and fecundity were lower for both IMM and 
AGM in the Cry9C corm kernels and Bt treated seeds than 
controls. Results are consistent with those obtained for 
IMM and AGM reared on CrylA(b) transformed grain. 


Impact of CrylAb transformed corn kernels on Indian 
meal moth and Angoumois grain moth populations in farm 
storage. ANTHONY M. HANLEY,* JOHN D. SEDLA- 
CEK, BRYAN D. PRICE, and MICHAEL R. TINSLEY, 
Community Research Service, Kentucky State University, 
Frankfort, KY 40601. 

Increased concern by consumers regarding chemical 
residues in food products, the development of resistance 
by stored grain insect pests to organophosphorus insecti- 
cides, the Food Quality Protection Act of 1996, and other 
federal regulations and international accords such as the 
Montreal Protocol, have generated a need to examine new 
control methods to assist or replace existing control man- 
agement tactics of stored product insects utilizing several 
biopesticides. Bacillus thuringiensis (Bt) is available for 


use in stored grain, Currently, Dipel® is the only product 
registered for use in stored grain environments against 
Indian meal moth (IMM). Transgenic crops with insect 
resistant characteristics used against insect pests have 
been successful. One such example is transgenic Bt corn. 
This study examines the efficacy of CrylAb transformed 
Dekalb corn on Indian meal moth and Angoumois grain 
moth (AGM) populations in on-farm bin storage. One 
hundred bushels of Dekalb 679BTY (Bt+) and Dekalb 
679 (Bt—) were added tol6 bins located at the Kentucky 
State University Agricultural Research Farm in Franklin 
County, KY. One thousand IMM and AGM eggs were 
added to the bins twice to ensure infestation. Tempera- 
tures were recorded weekly and adult moths were quan- 
tified biweekly using sticky traps. Data from the sixth week 
of observation indicated that populations of IMM were 
twice as large in Bt— bins as those containing the Bt+ 
corn kernels. All data will be compiled at the finish of the 
field season and results will be discussed relative to on- 
farm storage capability and farmer impact. 


Use of conservation tillage and cover crops for sustain- 
able vegetable production. H. J. HRUSKA,* G. R. 
CLINE, A. F. SILVERNAIL, and K. KAUL, Community 
Research Service, Kentucky State University, Frankfort, 
KY 40601. 

Research began in 1999 to examine sustainable pro- 
duction of bell peppers (Capsicum annuum) using con- 
servation tillage and legume winter cover crops. Tillage 
treatments included conventional tillage, strip-tillage, and 
no-tillage, and winter covers consisted of hairy vetch (Vi- 
cia villosa), winter rye (Secale cereale), and a vetch/rye 
biculture. Pepper yields following the rye winter cover 
crop were significantly reduced if inorganic nitrogen fer- 
tilizer was not supplied. However, following vetch, yields 
of peppers receiving no additional nitrogen were similar 
to yields obtained in treatments receiving the recom- 
mended rate of inorganic nitrogen fertilizer. Thus, vetch 
supplied sufficient nitrogen to peppers in terms of yields. 
Pepper yields following the biculture cover crop were in- 
termediate between those obtained following vetch and 
rye. When weeds were controlled manually, pepper yields 
following biculture cover crops were similar among the 
three tillage treatments, indicating that no-tillage and 
strip-tillage could be used successfully if weeds were con- 
trolled. With no-tillage, yields were reduced without weed 
control but the reduction was less if twice the amount of 
residual cover crop surface mulch was used. Without man- 
ual weed control, pepper yields obtained using strip-tillage 
were reduced regardless of metolachlor herbicide appli- 
cation. It was concluded that a vetch winter cover crop 
could satisfy nitrogen requirements of peppers and that 
effective chemical or mechanical weed control methods 
need to developed to grow peppers successfully using no- 
tillage or strip-tillage. 


Pesticide movement under field conditions: an over- 
view. CHRISTINE LEE* and GEORGE F. ANTO- 


Abstracts, 2000 Annual Meeting 


NIOUS, Community Research Service, Department of 
Plant and Soil Science, Kentucky State University, F rank- 
fort, KY 40601. 

Different management practices for growing vegetable 
crops on highly erodible land (10% slope) have been eval- 
uated by the Water Quality Project at Kentucky State Uni- 
versity. Studies were conducted to determine the influ- 
ence of landscape features and soil amendments on pes- 
ticide movement into runoff and infiltration water. Three 
soil treatments on a Lowell silty loam soil (pH 6.7, 2% 
organic matter) were used to reduce soil erosion and sur- 
face water runoff. Pesticides infiltration into the vadose 
zone were monitored using pressure-vacuum lysimeters (n 
= 27). Twelve tipping bucket metering units were used 
to collect runoff. In 1998, three soil treatments were com- 
pared: black plastic (BP) mulch, living fescue mulch (tall 
fescue), and no-mulch treatments (NM, rototilled bare 
soil). In 1999, living fescue mulch 30 cm wide was planted 
between every cropping row (pepper intercropped with 
tomato each in a row) to create multiple barriers to runoff. 
Turf was also planted every other cropping row and both 
treatments were compared with NM treatments. In year 
2000, yard waste compost was used as a soil amendment. 
Residues of pyrethrins (Py-I and Py-II) and piperonyl bu- 
toxide (PBO) were quantified in soil and runoff water fol- 
lowing spraying of a pyrethrum formulation “Multi- 
Purpose Insecticide” containing Py-I, Py-II and PBO. Ac- 
cordingly, the impact of the different soil mulches on the 
horizontal and vertical movement of clomazone, dacthal, 
endosulfan, and pyrethrins was measured and evaluated 
under field conditions. 


Recent advances in thin-layer chromatography of pes- 
ticides. IESSHA MOORE,* HUMERA TAGUI, and 
GEORGE ANTONIOUS, Department of Plant and Soil 
Science, Kentucky State University, Frankfort, KY 40601. 

Potential hazards to human health, food, soil, water, and 
wildlife may be created by residues from some long-lived 
pesticides that build up in the food chain ‘and cause wide- 
spread contamination by pesticides. Thin-layer chroma- 
tography (TLC) can be used widely to detect and quantify 
residues of pesticides in various kind of samples such as 
food, drinking water, environmental matrixes (soil, 
groundwater and wastewater), biological materials, and to 
detect low concentrations of active ingredients of pesti- 
cides in their commercial formulations. The objective of 
this study was to separate and detect pesticides and relat- 
ed compounds using TLC plates. Methods for residue 
analysis of carbofuran, pirimiphos-methyl, pyrethrins, and 
piperonyl butoxide have been developed by KSU/Envi- 
ronmental Toxicology research group using selected mo- 
bile phases on silica gel plates. Hexane-ethyl acetate (80: 
20) and hexane-benzene-acetone (80:20:10) were used as 
mobile systems for best resolution. Orthophosphoric-tan- 
nic acid reagent in acetone was used as chromogenic spray 
reagent for simultaneous detection of pyrethrins and pi- 
peronyl butoxide in environmental samples. Critical fac- 
tors of the TLC procedure and stepwise confirmation of 


positive results leading to flexibility of the methox 
presented. 


Sustainable soil management practices and quality 
potato grown on erodible lands. GAYATRI A. PATEL,* 
GEORGE F. ANTONIOUS, and CHRISTINE LEE, De- 
partment of Plant and Soil Science, Kentucky State Uni- 
versity, Frankfort, KY 40601; JOHN C. SNYDER, De- 
partment of Horticulture, University of Kentucky, Lexing- 
ton, KY 40546. 

Land productivity can decline when top soil is lost. In 
Kentucky, limited resource farmers often produce vege- 
table crops on highly erodible lands. The objectives of this 
study were 1) to quantify the impact of three soil man- 
agement practices (SMPs) on quantity of potato produced 
on erodible land, 2) to evaluate the performance of py- 
rethrin and azadirachtin insecticides on potato tuber qual- 
ity, and 3) to assess the impact of yard waste compost on 
the chemical composition (ascorbic acid, free sugars, and 
phenol contents) of potato tubers. Potatoes (Solanum tub- 
erosum L. cv. Kennebec) were grown in a silty loam soil 
of 10% slope. Plots (n = 18) were 3.7 m wide and 22 m 
long, universal soil loss equation (USLE) standard plots, 
with metal borders of 20 cm above ground level. Two bo- 
tanical insecticides, a multi-purpose insecticide (contain- 
ing pyrethrin 0.2%) and Neemix 4EC (containing 0.25% 
azadirachtin) were sprayed twice on potato foliage during 
each of two growing seasons (1997 and 1999) at the rec- 
ommended rates of 6 lbs and 2 gallons per acre, respec- 
tively. The SMPs were living fescue strips (FS) inter- 
cropped between every other potato row, soil mixed with 
yard waste compost (COM) and no mulch (NM) treat- 
ment (roto-tilled bare soil). The experimental design was 
a 2X 3 X 3 factorial with main factors of two insecticides 
and three SMPs replicated three times. Potato rows (ori- 
ented on the contour of the slope) and the three SMPs 
were used as barriers to runoff. Average potato yield was 
lowest in NM and FS and highest in COM treatments. 
Yield obtained from the bottom of the plots was greater 
than that obtained from the top of plots. Tuber defects 
(rot, scab, sun green, hollow heart, necrosis, and vascular 
discoloration) were significantly different between the two 
growing seasons. The two insecticidal treatments and the 
three SMPs used did not have much influence on tuber 
defects. 


The influence of fertilizer rate and application method 
on early growth and development of container grown paw- 
paw (Asimina triloba) seedlings. KIRK W. POMPER,* 
EDDIE B. REED, and SNAKE C. JONES, Land Grant 
Program, Kentucky State University, Frankfort, KY 40601- 
2350. 

The pawpaw (Asimina triloba) is a native American tree 
fruit with potential as a new fruit crop and as an orna- 
mental plant. Development of fertilizer rates and appli- 
cation methods to facilitate rapid container production of 
seedlings would be desirable to nurseries. A factorial 
greenhouse experiment was conducted with treatments 


82 Journal of the Kentucky 
that incliled four levels of slow release fertilizer (either 
0.3.8 ~.6 or 23 g of Osmocote per cubic foot) and three 
level. of liquid feed fertilizer (0, 250 or 500 ppm Peters 
20) 20P-20K). Seeds were sown in rootrainers with Pro- 
! rowing medium that contained the slow release fer- 
er treatment levels. After seedlings reached 2-3 leaves, 
edlings were fertigated at 0, 250 or 500 ppm Peters 
20N-20P-20K. The treatments were arranged in a split 
plot design in three replicated blocks, where the main plot 
effect was liquid feed, and subplot effect was slow release 
fertilizer. There were 20 replicate seedlings per experi- 
mental treatment combination. After 8 weeks, plants were 
destructively harvested. Both liquid and slow release fer- 
tilizer main effects significantly influenced pawpaw seed- 
ling growth characteristics; however, interactions between 
the main effects were also significant for all growth pa- 
rameters except leaf number. Overall, the seedlings sub- 
jected to the highest rate in both fertilizer methods 
showed the greatest total dry mass, about three fold great- 
er than control plants. 


Effect of several beetle-active Bacillus thuringiensis 
products on life history attributes of lesser grain borer, 
Rhyzopertha dominica. BRYAN D. PRICE,* JOHN D. 
SEDLACEK. and ANTHONY M. HANLEY, Community 
Research Service, Kentucky State University, Frankfort, 
KY 40601. 

The lesser grain borer (Rhyzopertha dominica) (LGB), 
causes extensive damage to stored grains, especially 
wheat, and is considered a primary insect colonizer, feed- 
ing on intact kernels and developing entirely inside the 
grain. Artificial wheat kernels (pellets) were prepared in 
our laboratory by combining ground wheat, corn starch, 
wheat gluten and water. Pelleting techniques are a con- 
venient way to examine the effects of various pesticides 
and other compounds on life history attributes of inter- 
nally feeding insects. The kernels were used to bioassay 
several Bt products, Novodor™, Raven® and Foil™, 
against the LGB, and were composed of 51% corn meal, 
26% cor starch, 9% wheat gluten and 14% water. The 
appropriate amount of each Bt product was added at a 
concentration of 500 ppm active ingredient. Ten kernels 
of each treatment, including control, were placed in five 
vials. Eight LGB adults were placed into each vial, then 
placed randomly in a small plastic container. The adults 
were removed from the kernels after seven days and adult 
mortality quantified. Surviving adults were then trans- 
ferred to another set of vials containing untreated kernels 
to determine any effect on fecundity. Both sets of vials 
were observed until all progeny emerged, then the num- 
ber of progeny and development time were determined. 
There was no significant difference in adult survivorship 
among the different Bt products, but preliminary data in- 
dicates emergence from Raven and Foil treatments were 
significantly reduced from control. Also, development 
times were longer with those treatments. Fecundity data 
will also be presented. 


Academy of Science 62(1) 


Influence of CrylAaAbAc2A Bacillus thuringiensis del- 
ta-endotoxin on a non-target parasitoid, Habrobracon he- 
betor LOUIE RIVERS I,* ANTHONY M. HANLEY, 
BRYAN D. PRICE, and JOUN D, SEDLACEK, Land 
Grant Program, Kentucky State University, Frankfort, KY 
40601. 

Interest in biological control of stored grain pests has 
increased with decreasing efficacy of organophosphorus 
insecticides, perceived risks of toxic residues, the Food 
Quality Protection Act of 1996, and the Montreal Proto- 
col, Bacillus thuringiensis (Dipel® [CrylAaAbAc2A]) has 
been found to be effective against the Indian meal moth 
(IMM) in stored grains. Habrobracon hebetor is an effi- 
cient parasitoid of IMM. However, no studies have been 
conducted to determine whether or not Bt and H. hebetor 
are compatible methods of control. Thus, the objective of 
this research is to quantify the impact of Bt on several H. 
hebetor life history attributes. Ten treated larvae were 
placed into three 20 ml plastic vials per treatment with 
three replicates. Two male and one female H. hebetor 
adults were added to each vial and allowed to oviposit for 
24 hours. then were removed. Number of progeny and 
longevity were determined. Progeny were placed in vials 
in the previous ratio and allowed to oviposit on ten ad- 
ditional larvae for 24 hours. Parasitoids were moved to 
vials of fresh larvae daily until female death. Total number 
of progeny and their development time were determined. 
All rearing and experiments were conducted in an envi- 
ronmental chamber at 27°C, =60% relative humidity and 
in total darkness. Progeny production and longevity of F, 
and F, individuals were the same between treatments. 
Thus, it appears as though Dipel does not have a negative 
impact on the parasitoids. Longer term experiments con- 
stituting multiple generations should be performed as 
should behavioral assays. 


Insect fauna of ear corn stored in cribs on small farms 
in Kentucky. JOHN D. SEDLACEK,* BRYAN D. 
PRICE, and ANTHONY M. HANLEY, Kentucky State 
University, Frankfort, KY 40601. 

Storing ear corn in corn cribs is practiced by many small 
and limited resource farmers in the central Kentucky area. 
Approximately 53% of all corn cribs in Kentucky are lo- 
cated in this region and as much as 30% of the total com 
acreage and 75% of those storing corn on-farm in individ- 
ual counties store it in this manner. Ear corn is more sus- 
ceptible to attack by insects and fungi for the duration of 
storage because it is completely exposed. This past storage 
season we sampled ear corm in 15 corn cribs on small 
farms in Casey, Christian, Franklin, and Metcalfe coun- 
ties. Thus far we have identified 18 species or species 
groups of beetles and 3 species of moth pests. The major 
species of beetles are maize weevil, foreign grain beetle, 
flat grain beetle, and hairy fungus beetles. The major moth 
species were Indian meal moth and Angoumois grain 
moth. Large numbers of parasitoids in the families Bra- 
conidae and Pteromalidae were also found. Results will be 


Abstracts, 2000 Annual Meeting 


discussed relative to bin stored corn pests and potential 
effects of transgenic Bt hybrids in storage. 


CELLULAR & MOLECULAR BIOLOGY 


Transcription factor isoform-specific regulation of pi- 
tuitary hormone gene expression. SCOTT E. DIA- 
MOND* and AMY L. FERRY, Department of Physiology, 
University of Kentucky College of Medicine, Lexington, 
KY 40536-0298. 

We do not yet fully understand the mechanisms by 
which related transcription factor isoforms with identical 
DNA sequence specificity mediate distinct transcription 
responses. Pit-1 and Pit-1b direct proper development of 
and hormone expression by the anterior pituitary, and 
constitute such a pair of transcription factor isoforms. 
While Pit-1 and Pit-1b share identical DNA binding do- 
mains, they differ by the splice-mediated insertion of the 
26 amino acid b-domain in the transactivation domain. 
Pit-1 and Pit-1b regulate prolactin gene expression in op- 
posite ways; Pit-1 activates the prolactin promoter, yet Pit- 
1b represses basal prolactin promoter activity as well as 
Ras signaling to the prolactin promoter. We have previ- 
ously demonstrated that the amino acid sequence of the 
b-domain dictates isoform-specific repression. Here, we 
utilize epitope- and alanine-scanning mutagenesis experi- 
ments to identify specific residues of the b-domain inser- 
tion that dictate repression of prolactin gene expression. 
We demonstrate by pharmacological and molecular ge- 
netic methods that b-isoform repression requires the ac- 
tion of a histone deacetylase complex. In addition, we uti- 
lize a chromatin immunoprecipitation assay to show that 
Pit-1b specifically alters the histone acetylation state of the 
prolactin promoter. These findings provide significant in- 
sights into the structural determinants and mechanism of 
an important example of transcription factor isoform-spe- 
cific regulation of gene expression. 


GEOLOGY 


Nature and origin of the Cane Run Bed, Lexington 
Limestone, central Kentucky. H. LISA JEWELL* and 
FRANK R. ETTENSOHN, Department of Geological 
Sciences, University of Kentucky, Lexington, KY 40506- 
0053. 

Much of the Middle Ordovician Lexington Limestone 
represents coarse, bioclastic deposition on a shallow-wa- 
ter, carbonate ramp subject to frequent storm perturba- 
tion. In fact, position of the Lexington area in a subtrop- 
ical, trade-wind belt on the southeast margin of Laurentia 
during Middle Ordovician time supports the predomi- 
nance of storm or tempestite deposition in the Lexington 
Limestone. In contrast to most of the Lexington Lime- 
stone, however, the Cane Run Bed, which occurs in the 
uppermost Grier Member, is composed of interbedded 
fine-grained limestones and shales, which exhibit second- 
ary, soft-sediment deformation. Moreover, the Cane Run 
grades laterally into coarser grained limestones of the Tan- 
glewood and Grier members, which at the approximately 


equivalent horizon are also locally deformed. Mapp 
distribution of the fine-grained Cane Run Bed show - 

it was deposited in a paleobathymetric low between 
structural trends that formed high areas during Lexingt 
deposition. Hence, the fine-grained nature of the Can Rui 
Bed reflects storm deposition in deeper water, distal en- 
vironments, while coeval parts of the Tanglewood and 
Grier represent storm deposition in shallower waters on 
adjacent, uplifted structural highs. The prominent soft- 
sediment deformation in the Can Run Bed and its equiv- 
alents not only provides correlation among different li- 
thologies and facies, but indicates the necessity of likely 
seismic activity for deformation (seismites) on one of the 
structures. Presence of probable seismites in proximity to 
local structures suggests reactivation of those structures 
and is additional support for the impact of Taconian far- 
field forces in the area during the ongoing Taconic tec- 
tophase to the east. 


Possible modern analogues to bryozoan bioherms in the 
Tanglewood and Grier members, Lexington Limestone, 
central Kentucky. JASON R. LAMBERT,* FRANK R. 
ETTENSOHN, and ANDREA L. HOLBROOK, Depart- 
ment of Geological Sciences, University of Kentucky, Lex- 
ington, KY 40506-0053. 

In equivalent parts of the Middle Ordovician Grier and 
Tanglewood members of the Lexington Limestone, just 
below the Brannon Member, a horizon of bryozoan bio- 
herms is known from three localities. The bioherms are 
sitting on a hardground surface, surrounded by coarse, 
crossbedded, fossil-fragment calcarenites. The bioherms 
are no more than a meter high and at most a meter and 
a half in breadth. Upper portions of the bioherms are 
composed of massive, but intricately encrusting platy 
bryozoans, whereas lower parts typically show branching 
or ramose bryozoans arranged around the base of the 
bioherm. On the east-facing, oceanward side of San Sal- 
vador Island, Bahamas, we observed within the surf zone 
and in deeper shoreface waters red-algae-coral patch reefs 
with some characteristics similar to those of the bioherms. 
Although the patch reefs were up to a few tens of meters 
in breadth and up to five meters in height, they were also 
attached to a hardground and surrounded by moving skel- 
etal sands. Upper parts of the reefs in the surf zone, es- 
pecially where facing out to sea, were composed of en- 
crusting red algae. Lower parts of the reefs below the surf 
zone, and those reefs in deeper shoreface waters, were 
composed of branching corals. Water depth and exposure 
to the surf zone apparently controls the morphology (en- 
crusting vs. branching) and taxonomic makeup of various 
reef parts. Although sizes are an order of magnitude dif- 
ferent, comparison of the two occurrences suggests that 
the Ordovician bioherms occurred within an ancient surf 
zone and that the different morphology and taxonomic 
makeup of bryozoans on different parts of the bioherms 
may reflect a similar zonation by depth. Hence, somewhat 


similar modern analogues allow us to fully understand the 


84 Journal of the Kentucky Academy of Science 62(1) 


nature a lepositional environments of an unusual, bio- 


herma! izon in the Lexington Limestone. 
‘tonic control of Sunbury (Kinderhookian) and Bor- 
d Osagean) deposition in northeastern Kentucky. 
£ I ) 


(ARLES E. MASON,* Department of Physical Scienc- 
Morehead State University, Morehead, KY 4035), 
RANK R. ETTENSOHN, Department of Geological 
Sciences, University of Kentucky, Lexington, KY 40506. 
Deposition of the Sunbury and Borden formations in 
northeastern Kentucky occurred during the fourth and fi- 
nal tectophase ( (Sunbury Cycle of Ettensohn, 1994) of the 
Acadian Orogeny. This Sunbury Cycle began in the Kin- 
derhookian with an unconformity at the base of the Sun- 
bury Shale. This break in deposition was in response to 
convergence of a newly placed load (Avalon microplate) 
on the eastern craton margin (Virginia promontory). This 
event was followed by deformational loading throughout 
most of remaining early Kinderhookian time, which re- 
sulted in rapid subsidence of the Appalachian forland ba- 
(west). The Sunbury 
Shale was deposited during this time. By middle Kinder- 


sin and its migration cratonward 


hookian time, the loading-type relaxation phase of the 
Sunbury Cycle began with deposition of the Henley Bed 
of the F 
phase reflects a deepening of the foreland basin and a 
concomitant eastward shift of bulge migration. During lat- 
est Kinderhookian time, carbonate turbidites were depos- 


Farmers Member of the Borden Formation. This 


ited in the basin, probably derived from the eastwardly 
migrating bulge to the west. The clastic turbidites com- 
Farmers Member, deposited during early Os- 
agean time, had an eastern source as did sediments com- 
posing the Nancy and Cowbell members of the Borden 


posing the F 


Formation. These members document the progradational 
infilling of the forland basin by a coarsening-upward clas- 


tic sequence during the loading-type relaxation phase of 
eae 


the Sunbury Cycle. The topmost member of the Borden 
Formation, the Nada Member (upper Osagean) records 
the reduction and ultimate cutoff of silicilastic sediments 
from the highlands to the east, related to eastward bulge 
uplift and migration combined with a regional eustatic 
lowstand. Sediment reduction is most reflected in the 
glauconite-rich Floyds Knob Bed in the Nada and equiv- 
alent units. 


New observations on Brachiospongia digitata (Hexac- 
tinellidae, Porifera) from the Middle Ordovician of Cen- 
tral Kentucky. DANIEL J. PHELPS,* Kentucky Paleon- 
tological Society, Lexington, KY 40503; RICHARD 
TODD HENDRICKS, Bardstown, KY 40004. 

Recent discoveries of Brachiospongia digitata from the 
Curdsville Member of the Lexington Limestone (Middle 
Ordovician, Trentonian), shed light on the ecology and 
taxonomy of Brachiospongia. Brachiospongia specimens 
are present in an exposure of the Curdsville Member in 
outcrops adjacent to the Kentucky River Fault Zone. In- 
dividuals range from 8 to 35 cm in diameter and occur as 
complete or macerated specimens in fining-upward se- 


quences ranging from calcarenites and calcisiltites to or- 
ganic-rich, fissile, shales. The sponges are filled with lith- 
ified sediment; some are compacted and distorted. In one 
bed, the sponges are in life position, and relatively undis- 
torted. In other beds, sponges are associated with depres- 
sions adjacent to lithified and bored mounds on hardg- 
round surfaces. In these lenses, virtually all Brachiospon- 
gia, even the largest specimens, are overturned or other- 
wise displaced from life position. This suggests transport 
and mutilation of the sponges during storm events. Faunas 
associated with the sponges include bryozoans, dendritic 
graptolites, crinoids, a paracrinoid, and other echino- 
derms. Trilobites, including articulated specimens of Cer- 
aurus sp., have been found inside Brachiospongia, sug- 
gesting that trilobites interacted with sponges, perhaps for 
shelter. Some specimens of B. digitata display areas in 
which the body of the sponge is broken away to reveal an 
internal mold or an internal structure that is covered with 
tubercles and a thin layer of black materials. These tu- 
bercles are similar to those found on Brachiospongia tub- 
erculata (James). These details suggest that B. tuberculata 
specimens represent internal molds of B. digitata, and 
that B. tuberculata is synonymous with B. digitata. 


HEALTH SCIENCES 


Unintentional influence on diet quality of taste panel- 
ists. MARTHA A. MARLETTE* and SUSAN B. TEM- 
PLETON, Human Nutrition Program, Kentucky State 
University, Frankfort, KY 40601. 

A taste panel to determine the acceptance of popular 
African-American foods modified to improve nutrient con- 
tent was conducted for 10 weeks. Thirty-one African 
Americans, 58% female and 71% under age 30, served as 
panelists; 24 hr food recalls were collected from them at 
the initial training session and during the exit interview. 
Intakes were analyzed using Nutritionist V™. The panel- 
ists were given charts indicating the calories, total fat, sat- 
urated fat, cholesterol and dietary fiber content of the 
foods to be evaluated. Panelists reported the same num- 
ber of food items on both recalls. However, the quality of 
dietary intake had improved in several areas, based on 
Recommended Daily Values (RDV) for a 2000 keal diet. 
Energy intake decreased from 123% to 104% RDYV, total 
fat decreased from 161% to 115% RDYV, saturated fat de- 
creased from 172% to 121% RDV, cholesterol decreased 
from 118% to 79% RDYV, and fiber increased from 65% 
to 75% RDV. These results suggest that panelists became 
more aware of foods they were consuming and made bet- 
ter food choices, because they were given fact sheets that 
indicated the importance of a low fat, low cholesterol and 
high fiber levels provided by the modified recipes. This 
new knowledge on the need to reduce dietary fat and cho- 
lesterol apparently improved the food choices made by 
the panelists. However, these data do not indicate if these 
reported dietary changes would be sustained by them over 
time as no formal nutrition education was provided. 


Abstracts, 2000 Annual Meeting 85 


Lead and copper in tapwater of eastern Kentucky 
homes: a pilot study. JOHN G. SHIBER, Division of Bi- 
ological Sciences & Related Technologies, Kentucky Com- 
munity & Technical College System/KCTCS, Prestons- 
burg Community College, Prestonsburg, KY 41653. 

One hundred nine Prestonsburg Community College 
students, representing six eastern Kentucky counties, col- 
lected two samples each of tapwater from their homes for 
lead and copper analysis. The first sample was taken from 
their kitchen faucets (cold water tap), before anyone had 
used it that day (B), and the second from the same faucet, 
after the water had been used for a few hours (A). Sev- 
enty-three percent of the samples originated from city wa- 
ter supplies, 24% from private wells, and 3% from other 
sources (private river, reservoir or mine). Lead concentra- 
tions found in the (B) samples ranged from ND (Not De- 
tectable) to 10.20 ppb (¥ = 0.78 ppb), and the (A) samples 
from ND to 5.48 ppb (« = 0.60 ppb). Copper ranged in 
the (B) samples from ND to 0.84 ppm (¢ = 0.25 ppm) 
and in (A) samples from ND to 1.19 ppm (« = 0.06 ppm). 
No samples exceeded the EPA lead or copper “action lev- 
el” of 15.0 ppb and 1.3 ppm, respectively. Although no 
direct correlation between type of water source and lead 
concentrations was attempted, 30% of the well samples 
had over 1 ppb of lead, while only 12% of the city water 
samples did. Sixty-eight percent of the students whose wa- 
ter source was from city supplies drink from the cold wa- 
ter tap, and 44% of those whose water comes from private 
wells do also. Overall, 61% of the students’ families drink 
from the cold water tap, and 14% drink directly from the 
hot water tap. Seventy-seven percent cook with water 
from the hot water tap. 


African American taste panelists accept nutritionally en- 
hanced recipes for traditional foods. SUSAN B. TEM- 
PLETON* and MARTHA A. MARLETTE, Human Nu- 
trition Program, Kentucky State University, Frankfort, KY 
40601. 

African Americans face a disproportionate risk for hy- 
pertension, stroke, heart disease, diabetes, and some can- 
cers, conditions known to be influenced by dietary factors. 
Analysis of the Continuing Survey of Food Intakes of In- 
dividuals 1994-1994 and National Health and Nutrition 
Examination Survey III (1988-1994) data revealed many 
food items frequently consumed by African Americans 
were high in fat, sugar, sodium, or cholesterol or low in 
fiber. Our goal was to nutritionally enhance traditional rec- 
ipes for six commonly consumed items and to test their 
acceptability to African Americans using a multistage taste 
panel. Nutritionist Five was used to analyze the nutrient 
content of original and modified recipes. Blind ratings 
were made on taste alone; for Informed ratings, food 
items were identified as original or modified and the nu- 
trient content of each was provided; for Informed-Blind 
ratings, nutrition information was provided, but the ver- 
sions were not identified. Fifty-eight percent of the 31 
African-American panelists were female; 71% were 19-30 
years old. Based on a 1 (low) to 7 (high) scale, mean ac- 


ceptability ratings for original/modified food iten: 
5.2/4.6 (Blind), 5.5/4.7 (Informed), and 5.6/5.1 (Info: 
Blind) (P < 0.05). Although original items received hig! 
ratings, over 50% of Informed panelists indicated th 
were equally or more likely to consume the modified vei 
sion. Modified item ratings improved with each tasting; at 
the third tasting, Informed-Blind panelists could not dis- 
tinguish the modified food item from the original 35% of 
the time. Our findings suggest that these low fat/high fiber 
alternatives are acceptable to African-American panelists. 


PHYSICS & ASTRONOMY 

Generating solutions in Hinstein-Maxwell gravity. 
SHARMANTHIE FERNANDO, Department of Physics 
and Geology, Northem Kentucky University, Highland 
Heights, KY 41099. 

Solutions to Einstein-Maxwell gravity with cylindrical 
symmetry is generated. By taking the symmetries of the 
metric, the Lagrangian is written as a 1 + 0 sigma model. 
Then, SL(2,R) transformations are applied to the func- 
tions in 1 + 0 dimensions to generate new solutions in 3 
+ 1 dimensions. Well-known electrical and magnetically 
charged static solutions are taken as seed metrics to gen- 
erate new rotating charged solutions. 


NASA Kentucky Space Grant Consortium and NOVA 
Program opportunities for space-related research, tech- 
nology, and education. KAREN HACKNEY,* RICHARD 
HACKNEY, ROGER SCOTT, CHARLES McGRUDER, 
SANDRA CLEMENTS, MICHAEL CARINI, RI- 
CHARD GELDERMAN, TERRY WILSON, DON 
COLLINS, KATHI MATTHEW, and MIKE MAY, Ken- 
tucky Space Grant Consortium and NASA NOVA Pro- 
grams, Department of Physics and Astronomy, Western 
Kentucky University, Bowling Green, KY 42101. 

NASA has partnerships with Kentucky and its univer- 
sities for the purpose of involving faculty and students in 
space-related research, technology, and education. We will 
describe current opportunities in the Kentucky Space 
Grant Consortium for undergraduate scholarships and 
graduate fellowships for students in mentored, space-re- 
lated projects. Funding opportunities for the development 
of space-related research projects will be described. Ac- 
tivities of the NOVA and Space Grant programs that sup- 
port the teaching of space-science and related disciplines 
will be outlined. Progress of existing projects and new op- 
portunities will be discussed. 


Digital dynamism for enlivening astronomy in the class- 
room. RICHARD HACKNEY,* KAREN HACKNEY, 
ROGER SCOTT, MICHAEL CARINII, RICHARD 
GELDERMAN, and CHARLES McGRUDER, Depart- 
ment of Physics and Astronomy, Western Kentucky Uni- 
versity, Bowling Green, KY 42101. 

Digital technology is resulting in a wealth of available, 
inexpensive illustrative materials for enlivening astronomy 
presentations in the classroom. NASA and other sources 
provide animated views that dynamically illustrate the mo- 


S6 Journal of the Kentucky Academy of Science 62(1) 


tions and pliysical activity of objects in the universe. Stu- 


dents « ain a sense of personal involvement with, and 
the ir cdiacy of, activity in space through timely sharing 
of « at observational material on the web. Instructors 
ay adents can make and share direct observations of 


momical objects such as the sun and moon using sim- 

telescopes and relatively inexpensive digital cameras. 
\Ve present an assortment of sources, examples, and 
methods for using these resources to enliven the teaching 
and learning of astronomy. 


Demonstrating the Ptolemaic and Copernican systems. 
ROGER SCOTT,* RICHARD HACKNEY, KAREN 
HACKNEY, MIKE CARINII, RICHARD GELDER- 


MAN, and CHARLES MecGRUDER III, Department of 


Physics and Astronomy, Western Kentucky University, 
Bowling Green, KY 42101. 

Many astronomy courses include a historical perspec- 
tive of how the modern concept of the Solar System was 
developed, and the role that Galileo's observations of the 
phases of Venus played. This presentation will discuss the 
significance of Galileo's observations, and will illustrate a 
simple way of demonstrating the phases of Venus in both 
the geocentric Ptolemaic system and the heliocentric Co- 
pernican system. 


Gravitational lensing by black holes. BRIAN TALBERT 
and SEAN ROBERTS, Department of Physics and Ge- 
ology, Northern Kentucky University, Highland Heights, 
KY 41099. 

Lensing by charged black holes characterized by the 
mass and charge is studied. As a first step, we have cal- 
culated the deflection of light from a electrically charged 
black hole with the assumption that the gravitational field 
is weak and that the deflection angles of light rays are 
small. This approximation will lead to three images if the 
parameters in lensing such as the mass, the charge and 
the distances satisfy certain restrictions. We have done the 
calculations for a supermassive black hole and found the 
magnification of the images. The support from the Ken- 
tucky Space Grant Consortium is gratefully acknowledged. 


PHYSIOLOGY & BIOCHEMISTRY 

Evidence for dramatically increased bone turnover in 
spontaneously hypertensive rats. D. L. DeMOSS, De- 
partment of Biology, Morehead State University, More- 
head, KY 40351; G. L. Wright, Department of Physiology, 
Marshall University School of Medicine, Huntington, WV 
25704. 

Using the *H-tetracycline model, whole body skeletal 
bone resorption was compared among male and female 
spontaneously hypertensive and normotensive Wistar Kyo- 
to and Sprague-Dawley rats. Immature animals undergo- 
ing rapid skeletal growth and bone sculpting showed a 
tendency for decreased indices of skeletal resorption in 
females compared to males. By 24 weeks of age, the in- 
dices of rate of resorption and extent of metabolically re- 


active bone in male rats were decreased an average of 


68% and 74%, respectively, compared to values obtained 
at 8 weeks. By comparison, values for 24 week old females 
decreased only 26% and 56%, respectively, resulting in 
evidence of a significantly elevated level of resorptive ac- 
tivity in mature females compared to males in each of the 
three rat strains studied. Within sex comparisons of 24- 
week-old animals indicated that bone resorptive activity 
was similar between normotensive male and between nor- 
motensive female groups. By comparison, the resorptive 
activity of both male and fernale hypertensive rats was 
significantly increased compared to normotensive con- 
trols. This condition was exaggerated in female hyperten- 
sive rats, which showed an approximate 78% and 41% 
increase in indices of rate of resorption and extent of met- 
abolically reactive bone compared to normotensive con- 
trols. The results indicate a marked sexual dichotomy in 
the decline of skeletal bone resorptive activity following 
maturation and slowing of skeletal growth. They further 
indicate a significant elevation of whole skeleton bone 
turnover in male hypertensive rats and dramatically in- 
creased bone turnover in female hypertensive rats. 


PSYCHOLOGY 


Effects of observer type on social facilitation. TIM 
KRAEMER and BONNIE BOWERS, Centre College, 
Danville, KY 40422. 

This experiment, which concems the effects of observer 
type on social facilitation for a learning task, was modeled 
after the research of Henchy & Glass (1968, cf. Gore & 
Taylor, 1973) and Gore & Taylor (1973). The present ex- 
periment proposed that males’ and females’ response time 
and accuracy of answers on a learning task would be 
quicker and more accurate for students observed by a per- 
ceived nonexpert on the task than those observed by a 
perceived expert on the task. Forty Centre College stu- 
dents, 25 males and 15 females with a mean age of 19.83, 
completed the Zinbarg learning task on standard college 
computers, while being observed by a perceived expert/ 
nonexpert or not being observed (control condition). The 
hypothesis was partially supported, such that only females’ 
scores showed social facilitation in that their performance 
was quicker and more accurate in the nonexpert condition 
than in the expert condition. 


SCIENCE EDUCATION 


Fear of algebra. JOHN G. SHIBER, Division of Bio- 
logical Sciences & Related Technologies, Kentucky Com- 
munity & Technical College System/KCTCS, Prestons- 
burg Community College, Prestonsburg, KY 41653. 

Frequent comments by community college students, es- 
pecially non-traditional students, about disliking and fear- 
ing algebra, prompted a survey to learn the extent of the 
problem, and if there were similar fears of biology, chem- 
istry or physics. Three hundred seventy-seven community 
college students (40% non-traditional) responded. Most 
had taken algebra, and 53% disliked or feared it. Twenty- 
six percent and 22%, respectively, feared chemistry and 


Abstracts, 2000 Annual Meeting 87 


physics. Non-traditionals were more prone to fear all. Bi- 
ology was least feared. Six hundred sixty-seven Eastern 
Kentucky high school seniors completed the survey for 
comparison purposes. Only 14% feared algebra, but many 
feared calculus, i.e., 40% who had taken it and 48% who 
hadn't. Twenty-four percent feared chemistry and physics, 
30% of females vs. 18% of males feared physics, and bi- 
ology, again, was least feared. Students blamed their fears 
of algebra and calculus on a lack of confidence, e.g., “can’t 
understand it,” “too confusing,” “not good at math,” etc. 
Suggestions for improving the courses, though, reflected 
need for better instructional approaches, wanting teachers 
to go slower, explain more in depth, and have more hands- 
on and real-life applications. Many high school seniors 
wanted more knowledgeable instructors who are interest- 
ed in math and student learning. It is concluded that a 
perceptual problem among students that labels math in 
general, and algebra and calculus in particular, as too dif- 
ficult for the average person, does indeed exist, and is 
often perpetuated by teachers and counselors. A more 
positive attitude about math when dealing with students, 
and a more concerted effort by all educators to remedy 
the problem, is urged. 


Student survey on violence. JOHN G. SHIBER, Divi- 
sion of Biological Sciences & Related Technologies, Ken- 
tucky Community & Technical College System/KCTCS, 
Prestonsburg Community College, Prestonsburg, KY 
41653. 

The wave of concern in recent years about school vio- 
lence prompted a survey on the issue at Prestonsburg 
Community College. Sixty-three percent of 307 students 
responding have weapons in their homes; 59% have more 
than one. Eighty-eight percent of them have rifles, 70% 
handguns, and 43% large knives. The weapons are used 
chiefly for hunting (33%), personal protection (25%), or 
both (27%). Twenty-nine percent of the students have 
children at home, 51% of whom are under 15 years of 
age. Over half know where the weapons are kept, and 23% 
could get them if they wanted. Of all students surveyed, 
58% often watch movies with a lot of violence, and 50% 
believe children under 18 should be admitted to “R-rated” 
movies. Only 17% play video games that are violent in 
nature. When very angry, 48% resort to some form of 
violent behavior, e.g., slamming doors, throwing things, 
screaming, hitting something (or someone), etc. Thirty- 
three percent have been in serious physical fights, and 
64% have witnessed them in school. According to many, 
most school fights are over the opposite sex (36%) and 
kids teasing or spreading rumors about each other (25%). 
Twenty-six percent had seen students with weapons in 
school, but few told an adult about it, for fear of reper- 
cussions. While attributing teen violence to many factors, 
students most frequently mentioned the lack of parental/ 
guardian supervision and guidance. Parents of violent 
teens, said a majority, are partially, if not entirely to blame 
for their offsprings’ behavior. Temporary suspension or 
permanent expulsion, with some type of police involve- 


ment, was most commonly suggested as a punishme 
students bringing weapons to school. 


ZOOLOGY & ENTOMOLOGY 

Abiotic factors as predictors of terrestrial vertebrate 
species richness in Kentucky. MATTHEW L. COLE* and 
TERRY L. DERTING, Department of Biological Scienc- 
es, Murray State University, Murray, KY 42071. 

On a global scale, abiotic factors, particularly those as- 
sociated with climate, have significant impacts on species 
richness. Climate variables have been used successfully in 
models that predict large-scale variation in species rich- 
ness. We tested the utility of abiotic factors as predictors 
of species richness at the regional level. Using regression 
analysis, we developed models to predict the combined 
richness of terrestrial vertebrate species in Kentucky and 
the species richness of amphibians, breeding birds, mam- 
mals, and reptiles. The abiotic variables used in our mod- 
els were precipitation, temperature, elevation, topograph- 
ic variation, and road density. The resulting models were 
generally effective, accounting for ~50% or more of the 
variation of species richness in the state. Temperature, 
precipitation, and elevation were of primary importance 
in all models. Temperature alone was the best predictor 
of total species richness (1? = 0.56). Elevation and tem- 
perature together were the best predictors of the species 
richness of breeding birds (1? = 0.49), mammals (1? = 
0.48), and reptiles (1? = 0.75). Abiotic variables were less 
effective as predictors of the species richness of amphib- 
ians, however. Elevation and precipitation yielded the best 
predictive model but explained only 22% of the variation 
in amphibian species richness. As with global models, abi- 
otic factors accounted for a significant amount of regional 
variation in species richness. Through future modeling ef- 
forts we will determine the extent to which biotic factors 
improve predictions of species richness in Kentucky. 


Mate selection of male Poecilia reticulata. GREG DAR- 
NELL* and ZACHARY McCARTY,* Department of Bi- 
ology, Transylvania University, Lexington, KY 40508. 

Our experiment hypothesized that when male guppies 
(Poecilia reticulata) were given the chance to select a po- 
tential mate, they would prefer the larger of two female 
guppies. To run trials, a test tank was setup with three 
clear partitions to divide the tank into three equally sized 
chambers. Different sized females were placed in the out- 
er chambers and a male was released in the middle. Trials 
were run for 15 minutes and data were collected based 
upon how much time the male spent in the vicinity of 
each female. All guppies were measured after the com- 
pletion of each trial and new guppies being used in the 
following trial. Of the 16 males tested, 12 preferred the 
large female, 3 preferred the smaller female, and 1 
showed no preference. Using the sign test to analyze these 
data, the null hypothesis stating that the male would be 
indifferent and spend equal time with each female was 
rejected with 95% confidence. Therefore, when presented 
with the opportunity for mate choice, male guppies dis- 


88 Journal of the Kentucky Academy of Science 62(1) 


criminaied and chose the larger female guppy. This may 
relate ‘ the increased fertility of the large females, which 
pro a greater number of eggs than the smaller fe- 
n 


low pattern of fruiting in tropical trees affects territory 
size of white-winged trumpeters. PETER T. SHER- 
MAN,* Department of Biology, Transylvania University, 
Lexington, KY, 40508; PERRI K. EASON, Department 
of Biology, University of Louisville, Louisville, KY 40292. 

We investigated determinants of territory size in white- 
winged trumpeters (Psophia leucoptera), which are group- 
living, frugivorous birds that defend large, permanent ter- 
ritories in Amazonian rainforest. During a 7-month peri- 
od, we measured changes in food density at a site in Manu 
National Park, Peru, and conducted full-day focal samples 
on individuals that allowed us to record daily food intake 
of territorial birds. Both census and focal sample data sug- 
gested that food availability on trumpeter territories varied 
widely. To determine whether trumpeter territory size is 
related to food density, we conducted fruit removals dur- 
ing which we removed daily an amount of fruit from the 
territory that was equivalent to the amount of fruit in- 
gested daily by the focal group during a control period 
that preceded the removal experiment. Trumpeter food 
consumption did not change when fruit was removed dur- 
ing periods of resource abundance, but, when experimen- 
tal removals were conducted during times when food was 
scarce, total caloric value of food consumed decreased sig- 
nificantly. Our results suggest that food abundance on 
trumpeter territories sometimes exceeds and sometimes is 
close to or below the amount needed to meet the ener- 
getic needs of the groups. Based on data collected, we 


propose that territory size of trumpeter groups reflects 
some minimum size needed to provide the group with a 
baseline level of food intake during seasonal periods of 
decreased food abundance. 


Swamp rabbit habitat modeling using a geographic in- 
formation system. ADAM SMITH* and TERRY DERT- 
ING, Department of Biological Sciences, Murray State 
University, Murray, KY 42071. 

Using a geographic information system (GIS), we de- 
veloped three models predicting the occurrence of swamp 
rabbits (Sylvilagus aquaticus) in western Kentucky. The 
models were compared to the existing Kentucky GAP 
model. All models were successful in the prediction of 
confirmed swamp rabbit observations, both of presence 
(>80%) and absence (>90%), with only slight variation 
among models. After selection of the best model based 
on statistical performance and model construction criteria, 
the best model was applied to geographic areas of Ken- 
tucky contiguous to the species’ current Kentucky range 
to investigate the possibility of swamp rabbit reintroduc- 
tions. Only five areas for potential reintroduction were 
identified, with two of marginal quality and the remaining 
unsuitable for swamp rabbit reintroduction. Land own- 
ership and protection status of all habitat determined to 
be suitable by our best model was assessed; the vast ma- 
jority of all land was in private ownership. Wildlife Man- 
agement Areas protected the most swamp rabbit habitat 
(~=8%), but most suitable habitat remains at risk. These 
models can serve as a solid basis for future swamp rabbit 
management considerations. Field validation and frequent 
assessments and modifications of the models associated 
with increased data availability and improved GIS tech- 
nology are recommended. 


Guidelines for Contributors to the Journal 


. GENERAL 


. Original research/review papers in science will be con- 
sidered for publication in JKAS; at least the first author 
must be a member of the Academy. Announcements, 
news, and notes will be included as received. 

. Acceptance of papers for publication in JKAS depends 
on merit as evaluated by each of two or more review- 
ers. 

. Papers (in triplicate) may be submitted at any time to 
the editor. Do not send a disk file. 


John W. Thieret 
Biological Sciences 
Northern Kentucky University 
Highland Heights, KY 41099 
Phone: (859) 572-6390 
FAX: (859) 572-5639 
E-mail: thieretj}@nku.edu 


List in the cover letter your telephone number, your 
e-mail address, and the names, addresses, and tele- 
phone numbers of two persons who are potential re- 
vieWers. 

. Format/style of papers must conform to these guide- 
lines and also to practices in recent issues of JKAS, 
which are, in effect, a style manual. 

. Papers should be submitted in hard copy. Do not sta- 

ple pages together. 

. Indent the first line of each paragraph (but not the 

first line of entries in the Literature Cited). 


. FORMAT 


Papers should be in 12-point type on white paper 8.5 
x 11 inches, with margins at least 1 inch all around. 
Double-space throughout the paper (i.e., one full line 
of space between each two lines of text, literature cit- 
ed, or tabular data). Do not justify right margins. 

. Except for scientific names of genera and of infrage- 
neric taxa, which should be typed in italics, the same 
type (roman) should be used throughout (i.e., one type 
size only; bold only for paper title). 

. Sequence of sections in papers should, where appro- 
priate, be as follows: title of paper, name/address of 
author(s), abstract, body of paper, footnotes, table cap- 
tions, figure captions (all the preceding on consecu- 
tively numbered pages), tables, and figures. 

. The running head (top right) should give name(s) of 
author(s), a short version of paper title, and page num- 
ber of total. 

. The first page should include the running head and, 
centered near the top of the sheet, the papers title 
and the name and address of author(s). These should 
be followed immediately by the abstract. (The first 
page should look as much as possible like the first page 
of articles in JKAS.) 

. The abstract, not to exceed 200 words, should be con- 


89 


4. 
A. 


cise, descriptive, and complete in itself without r: 
ence to the paper. 


. The body of the paper should, where appropriate, in 


clude the following sections: Introduction, Materials 
and Methods, Results, Discussion, Summary, Acknowl- 
edgments, and Literature Cited. 


. No more than three levels of headings should be used: 


level 1, in capitals, centered; level 2, in capitals/low- 
ercase, flush left; level 3, in italics, a paragraph indent, 
with initial capital only (except proper nouns and ad- 
jectives), and followed by a period, the text then start- 
ing after one blank space. 

Personal communications (avoid if possible) should be 
indicated in the text as follows: (name, affiliation, pers. 
comm., date), e.g., (O.T. Mark, Wainwright College, 
pers. comm., 5 Jun 1995). 


STYLE 


In text, spell out one-digit numbers unless they are 
used with units of measure (four oranges, 4 cm) and 
use numerals for larger numbers; do not begin any 
sentence with a numeral. 


. Use no footnotes except those for title page and tables. 


Footnotes, identified by consecutive superscript num- 
bers, should be entered on a separate sheet. 


. Measurements should be in metric and Celsius units. 


Define lesser-known symbols and give the meaning of 
acronyms at first use. Express time of day in the 24- 
hour system. Dates should be written day, month (ab- 
breviated to three letters), year without intemal punc- 
tuation. Units with multiple components should have 
individual components separated by a virgule (e.g., g/ 
m? or g/m?/yr). 


. Names of authors of binomials may be included but 


only at the first mention of the binomial. Cultivar 
names are not italicized but are enclosed in single 
quotes. 


. Useful guides for contributors to JKAS are the follow- 


ing: Scientific style and format: the CBE manual for 
authors, editors, and publishers, 6th ed., Cambridge 
University Press, 1994; The Chicago manual of style, 
14th ed., University of Chicago Press, 1993; The ACS 
style guide, American Chemical Society, Washington, 
DC, 1986; and AIP style manual, American Institute 
of Physics, New York, 1990. 


IN-TEXT CITATION OF LITERATURE 


Cite publications in the text by author(s) and date— 
e.g., (Readley 1994); multiple citations should be in 
alphabetical order and separated by semi-colons—e.g.., 
(Ashley 1987; Brown 1994; Foster 1975); multiple ci- 
tations of works by one author(s) should be in chro- 
nological order—e.g., (Jones 1978, 1983); publications 
by one author(s) in the same year should be distin- 
guished by a, b, c, ete—e.g., (Smith 1994a, 1994b). 
For in-text references to works with one or two authors 


90) Journal of the Kentucky Academy of Science 62(1) 


use ‘s of both authors—e.g., (Jones and Williams 
19 x works with three or more authors use name 
( first author followed by et al—e.g., (Lee et al. 


) 


o not include any reference unless it has been pub- 
lished or accepted for publication (“in press”; see be- 


low). 


5. LITERATURE CITED 


A. List all authors of each entry. Do not abbreviate jour- 
nal titles; abbreviations for these will be supplied by 
the editor. 

B. The first line of each reference should be typed flush 
left; the remaining lines should be indented. 

C. Examples of common types of references are given 


below. 


JOURNAL ARTICLE 

Lacki, M.J. 1994. Metal concentrations in guano from a 
gray bat summer roost. Transactions of the Kentucky 
Academy of Science 55:124—126. 

BOOK 

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

PART OF A BOOK 


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. 


WORK IN PRESS 


Groves, S.J., 1.V. Woodland, and G.H. Tobosa. n.d. De- 
serts of Trans-Pecos Texas. 2nd ed. Ocotillo Press, 
Yucca City, TX. 


6. ILLUSTRATIONS 


FIGURES (LINE DRAWINGS, MAPS, GRAPHS, PHO- 
TOGRAPHS) 


Figures must be camera-ready, glossy, black-and-white 
prints of high quality or laser prints of presentation qual- 
ity. These should be designed to use available space ef- 
fectively: a full page or part of one, or a full column or 
part of one. They should be mounted on heavy white 
board and covered with a protective sheet of paper; pho- 
tographs to be grouped as a plate should have no space 
between them. Dimensions of plates must observe page 
proportions of the journal. Each illustration in a plate may 
be numbered as a separate figure or the entire plate may 
be treated as one figure. Include scale bars where appro- 
priate. Lettering should be large enough to be legible after 


reduction; use lowercase letters for sections of a figure. 
Figure captions should be self-explanatory without refer- 
ence to the text and should be entered on a page separate 
from the text. Number figures in Arabic numerals. Statis- 
tics presented in figures should be explained in the caption 
(e.g., means are presented + SE, n = 7). 


i) 


TABLES 


Each table and its caption must be double-spaced, num- 
bered in Arabic numerals, and set on a sheet separate 
from the text. The caption should begin with a title relat- 
ing the table to the paper of which it is a part; it should 
be informative of the table’s contents. Statistics presented 
in the table should be explained in the caption (e.g., 
means are presented + SE, n = 7). 


7. ETHICAL TREATMENT OF ANIMALS AS RE- 
SEARCH SUBJECTS 


If vertebrate or invertebrate animals are involved in a re- 
search project, the author(s) should follow those guide- 
lines for ethical treatment of animals appropriate for the 
subjects, e.g., for mammals or for amphibians and reptiles. 
Papers submitted to JKAS will be rejected if their content 
violates either the letter or the spirit of the guidelines. 


8. PROOFS 


Authors are responsible for correcting proofs. Alterations 
on proofs are expensive; costs will be assessed to authors. 
Proofs must be returned to the editor within 3 days after 
the author receives them; delay in return may result in 
delay of publication. 


9. REPRINTS 


Forms for ordering reprints will be sent to the author 
when the proofs are sent. They are to be returned directly 
to Allen Press, not to the editor. 


10. PAGE CHARGES 


A page charge covering the partial cost of publication will 
be assessed for all authors. However, acceptance or rejec- 
tion of an individual manuscript is determined solely on 
scientific merit. Individuals may contact the KAS office 
(Kentucky Academy of Science, Science Outreach Center, 
University of Kentucky, Lexington, KY 40536-0078) to re- 
quest an application form for full or partial remission of 
page charges. The application form is also posted on the 
KAS website. Awards are limited to available funds. 


11. ABSTRACTS FOR ANNUAL MEETINGS 


Instructions on style of abstract preparation for papers 
presented at annual meetings may be obtained from the 
editor. Copies will be available also at each annual meet- 
ing of the Academy. 


NEWS 


The Morehead Electronic Journal of Applications in Mathematics (MEJAM) is a new interdisciplinary 
journal sponsored by Morehead State University, Morehead, Kentucky. The goal of MEJAM is to provide a 
refereed outlet for undergraduate students in any discipline to publish quality papers and see the results quick- 
ly. MEJAM accepts papers that are outside the realm of the typical undergraduate curriculum and that 
emphasize the applications of mathematics while maintaining significant mathematical interest. Papers may 
be historical, expository, or completely original in nature but must adhere to strict academic standards and 
must emphasize some aspect of the applications of mathematics. Papers from all disciplines will be consid- 
ered for publication. More information about the journal and instructions for submissions can be found on 
the journal’s website at http://www.morehead-st.edu/ colleges/science/math/mejam/. 


The Kentucky Academy of Science is seeking to complete its set of Transactions of the Kentucky 
Academy of Science. Various issues prior to 1985 are needed. Anyone willing to donate back issues or to 
sell them at a reasonable price should get in touch with the editor at thieretj}@nku.edu. 


The 2001 annual meeting of the Kentucky Academy of Science will be held jointly with the Tennessee 
Academy of Science on Thursday—Saturday, 29-30 November and 1 December 2001, at Middle Tennessee 
State University (MTSU), Murfreesboro, Tennessee. There will be a reception and symposium at the Garden 
Plaza Hotel in Murfreesboro on Thursday evening. Friday technical sessions will be help on the MTSU cam- 
pus in the Keathley University Center and James Union Building. The annual awards banquet and president's 
reception will be held Friday evening at the Garden Plaza Hotel. Technical sessions will conclude on Saturday 
in the Keathley University Center. 


PUBLICATIONS 


The Sibley Guide to Birds from the National Audubon Society is now available. This 544-page work, written 
and illustrated by David Allen Sibley, covers North America north of Mexico. Families and, in some cases, 
genera are introduced with small figures for initial comparisons. Each group is followed by individual species 
accounts including figures of the different forms and phases that may be exhibited by each species and also 
of both perching and flying views. The book has ca. 6600 illustrations (paintings) and descriptions of 810 
species and 350 regional populations; it is more a reference source than a field manual. The introductory 
chapter briefly discusses classification and techniques of field ornithology. A second chapter discusses and 
illustrates the topography of birds. Sibley concludes with an index to common and scientific names of species. 
The book is a Chanticleer Press edition published in 2000 by Alfred A. Knopf, Inc.; ISBN 0-679-45122-6; 
$35.00 (soft cover). 


The National Audubon Society Field Guide to Wildflowers. Eastern Region is a revision of the first edition 
of the work (1979); the revising author is John W. Thieret. This 879-page book has all new photographs of 
638 species, two or three per page. The introductory pages discuss the arrangement of the color plates 
(mostly by color and type of flower cluster), flower parts, inflorescence type, leaves, and plant classification 
and names. For each included species the text gives a description, flowering times, habitat, range, and com- 
ments. Relevant families are briefly discussed. The book concludes with an index to common and scientific 
names of species. The book is a Chanticleer Press edition published by Alfred A. Knopf, Inc., in 2001; ISBN 
0-376-40232-2. $19.95 (soft cover). 


Wt 
3 9088 01304 3419 . 


CONTENTS 


The Genus Trifolium (Fabaceae) in Kentucky. Michael A. Vincent ........ 1 


The Role of Light in Regulating Dandelion (Taraxacum officinale; 
Asteraceae) Inflorescence Height. David Lowell Robinson. ..................++ 18 


Distribution and Status of Freshwater Mussels (Bivalvia: Unionoidea) in 
the Cumberland River Basin Upstream from Cumberland Falls, Kentucky. 
Ronald R. Cicerello and Ellis L. Laudermilk ...............ccscsecessesseseseseces 26 


Morphometric Variation of Cotton Mice (Peromyscus gossypinus) and 
White-footed Mice (P. leucopus) in Kentucky. Nell A. Bekiares and 
George Ax. Feldhaime nh sisson: cncckavasaedasecaecensusccneddunsecvuabsessassnnndupapantantiieeae ao 


Woody Plants of Six Northern Kentucky Counties. Ross C. Clark and 
Reyaant Mi Bauer cies istesastcicdtecdsensgndascdesdaduasqrakanwivnis<cnanee@ahan patina 39 


Effects of Fish on Zooplankton Community Structure in Chaney Lake, a 
Temporary Karst Wetland in Warren County, Kentucky. Nicole Vessels and 


Sefiveny Di Sache) sos lssencesans cicie cavewqhsn deve scdecessvsubaiace ceavewcon teense healt 52 
A Historiography of Archaeological Research in the Mammoth Cave Area 
of Kentucky: 1824-2000. Kenneth C. Carstens ...............c.secsescesereceeeces 60 
Using Composts as Growth Media in Container Production of Tomatoes. 
Brian D. Lacefield ‘and. Elmer Gray <..scscosesscccedeccostecsiuaseeuensscemsenpensnens 70 
Book Review ss cccoicoeccin ci ie nain ecgenanteckedanhibetpaucdeetoues sacduainpechadd orgs ian 77 


Abstracts of Some Papers Presented at the 2000 Annual Meeting of the 
Kentucky Academy ‘of SCienCe® .isccasieccd caedae cases adeecanenceaseneweussneaensuunenaneeens 79 


Guidelines for Contributors to the Journal ................0c...ecccececccccececceceees 89