PATTERNS AND PROCESSES Of G ROUNDWATER INVASION By 
COPEPODS IN THE INTERIOR LOW PLATEAUS Of THE UNITED 
STATES
VZORCI IN PROCESI NASELJEVANJA CEPONOŽCEV V 
PODZEMELJSKE VODE NA PLANOTI INTERIOR LOW PLATEAUS 
V ZDRUŽENIH DRŽAVAH
Julian J. LEWIS
1
 & Janet W . REID
2
Izvleček UDK 574.9:595.34 (7)
Julian J. Lewis & Janet W . Reid: Vzorci in procesi naseljevanja 
ceponožcev v podzemeljske vode na planoti Interior Low Pla-
teaus v Združenih državah 
Med ceponožnimi raki, vzorčevanimi v podzemeljskih habi-
tatih, v jamah, vrtinah in rečnem hiporejiku blizu ter na sami 
planoti Interior Low Plateaus (ZDA), prevladujejo ciklopoidi 
z 39 vrstami, sledijo jim harpaktikoidi z devetimi vrstami in 
kalanoidi z dvema vrstama. Skoraj vse identificirane harpakti-
koidne in kalanoidne vrste so splošno razširjene, površinski 
generalisti. Štirinajst ciklopoidnih vrst, predstavnikov rodov 
Diacyclops, Itocyclops, megacyclops in Rheocyclops, je stigobion-
tov ali prebivalcev hiporejika. Vrste, ki imajo močneje izražene 
prilagoditve na podzemeljsko življenje, najdemo v južnih pre-
delih, ki v času ledenih dob niso bili pokriti z ledenim pok-
rovom. Rezultati vzorčenja podpirajo hipotezo, da tiste vrste, 
ki so bolj specializirane na podzemeljske-intersticialne pogoje 
niso bile sposobne razširjenja v predhodno poledenele predele; 
medtem ko so nekatere, manj specializirane vrste ob umikanju 
ledenikov uspele prodreti iz površinskih v podzemeljske vode.
Ključne besede: Copepoda, Crustacea, biogeografija, polede-
nitev, Severna Amerika.
1
 Lewis & Associates LLC, Cave, Karst & Groundwater Biological Consulting, 17903 State Road 60, Borden, IN 47106-8608, 
U.S.A.; e-mail lewisbioconsult@aol.com
2
 JWR Associates, 1100 Cherokee Court, Martinsville, V A 24112-5318, and Research Associate, Virginia Museum of Natural  
History, U.S.A.; e-mail: jwrassociates@sitestar.net 
Received/Prejeto: 07.02.2007
COBISS: 1.01
ACTA CARSOLOGICA 36/2, 279-289, POSTOJNA 2007
Abstract   UDC 574.9:595.34 (7)
Julian J. Lewis & Janet W . Reid: Patterns and Processes of 
Groundwater Invasion by Copepods in the Interior Low Pla-
teaus of the United States
The copepod crustacean fauna collected from subterranean 
habitats, including caves, wells, and the hyporheos of streams 
in and near the Interior Low Plateaus of the United States is 
dominated by Cyclopoida, with 39 species, followed by Harpac-
ticoida with 9, and Calanoida with 2. Nearly all of the harpac-
ticoid and calanoid species are widespread, primarily surface-
dwelling generalists. f ourteen of the cyclopoids, members of 
the genera Diacyclops, Itocyclops, megacyclops, and Rheocyclops, 
are apparently obligate stygobionts or hyporheic. Several of the 
species that are more strongly modified for subterranean exis-
tence occur only in the more southern, unglaciated areas. Our 
sampling data support the hypothesis that the more specialized, 
groundwater-interstitial species have been unable to disperse 
into previously glaciated regions; whereas some, less-special-
ized species may have invaded groundwaters from surface habi-
tats as the glaciers receded.
Key words: Copepoda, Crustacea, biogeography, glaciation, 
North America.
INTRODUCTION
We review published information and present new geo-
graphical records for the copepod crustaceans collected 
from subterranean habitats in and near the Interior Low 
Plateaus (ILP) physiographic province of the United 
States. f enneman (1938) described the ILP as an area of 
relative lowlands between the Appalachians to the east 
and the Ozark Plateaus to the west. The ILP stretches 
from northern Alabama, northward through Tennessee 
and Kentucky into southern Indiana and Illinois. Much 
of this area is karst topography and contains thousands 

ACTA CARSOLOGICA 36/2 – 2007 280
of known caves. The northern boundary of the ILP is the 
terminus of the Illinoian glaciation. Herein we trace the 
present-day distribution of the copepod species in this 
region, and attempt to infer the historical patterns and 
routes of their re-entry into previously glaciated areas.
The Interior Low Plateau region has been the focus 
of many earlier studies relating to the fauna of caves and 
karst, because of the notoriety of some local sites, espe-
cially in the central Kentucky karst, including what is 
now Mammoth Cave National Park. In 1928, C. Bolívar 
and René Jeannel made collections of copepods from 
numerous caves in the eastern United States, resulting in 
the first descriptions (Chappuis 1929, 1931) of subterra-
nean copepod taxa from this country: megacyclops don-
naldsoni (Donnaldson Cave, Lawrence County, Indiana), 
Diacyclops jeanneli (Marengo Cave, Crawford County, 
Indiana), and Bryocamptus morrisoni (Horse Cave, Hart 
County, Kentucky; and Donnaldson Cave). Cyclops clan-
destinus (renamed Diacyclops yeatmani by Reid, 1988) 
was described from Big Mouth Cave, Grundy County, 
Tennessee and a drain tile in Vermilion County, Illinois 
by yeatman (1964). Barr (1967) summarized collections 
of 17 taxa reported from Mammoth Cave and other caves 
in the central Kentucky karst by his own field work, as 
well as that of Kofoid (1899) and Chappuis (1931). 
Lewis began sampling cave faunas in the eastern 
United States in the 1970s, but only started focusing at-
tention on the micro-crustacean fauna in the 1990s (Lew-
is, and Lewis et al. 1998-2006). The initial impetus for 
these collections was the desire to rediscover Diacyclops 
jeanneli in Marengo Cave and the surrounding area dur-
ing a survey conducted for The Nature Conservancy in 
Indiana. The results from this sampling made it clear that 
a significant assemblage of subterranean microcrusta-
ceans was present in the groundwaters of the midwestern 
United States. Presented herein are the results of Lewis’ 
approximately 15 years of sampling. Newly described 
copepod taxa resulting from these collections are Rheo-
cyclops indiana (Reid et al. 1999), Diacyclops salisae, D. 
lewisi, D. indianensis and D. conversus (Reid 2004), and a 
new species of Itocyclops (Reid in review). 
PATTERNS Of DISTRIBUTION AND HABITAT
Like any other puzzle, a modicum of the pieces have to 
be gathered and arranged before even a hint of the overall 
picture begins to emerge. Enough sampling from caves 
and other windows into groundwater in the Interior Low 
Plateaus of the east-central United States has now been 
conducted to permit a few observations about the pat-
terns of distribution, as well as to infer the processes by 
which copepods have invaded this realm.
One of the ecological fortes of cyclopoid copepods 
of the genus Diacyclops is their use of groundwater habi-
tats, and it appears that some species are indeed stygobi-
onts. The best-known of these in the United States is Dia-
cyclops jeanneli. f or many decades after its discovery in 
1928, this species remained obscure, with even its micro-
habitat in Marengo Cave uncertain. Using shrimp-baited 
jar traps, Lewis (1996) found D. jeanneli in a well, not 
SAMPLING METHODS
A variety of means were used to collect copepods from 
different windows into the groundwater. Plankton drift 
samples were collected by hanging a plankton net over 
the orifice of a spring, in a flowing cave stream, or at the 
mouth of a drain tile. In deeper streams, rimstone pools, 
or drip pools in caves, the net was drawn through the 
water. In shallow drip pools, fed by the epikarst, water 
was dipped with a cup and strained through the plank-
ton net. In cave streams with sufficiently deep gravel, the 
Karaman-Chappuis technique was used to gather fauna 
living in the interstices. In wells, copepods and other 
fauna came readily to jars equipped with perforated lids 
and baited with a single, uncooked dead shrimp; these 
were left overnight and then retrieved. Longer trapping 
intervals appeared to result in fouling of the water and 
decrease in the fauna that was captured. f inally, a Bou-
Rouch pumpwell was used to sample hyporheic inter-
stices of the Blue River in southern Indiana. 
All samples were transported alive to the laboratory 
for visual inspection. The copepods and other fauna were 
much easier to detect alive than preserved. The animals 
were removed from the sample with an eye dropper, and 
then preserved in 70% ethanol. Voucher specimens were 
deposited in the collections of the National Museum of 
Natural History (Smithsonian Institution), or the Vir-
ginia Museum of Natural History.
JULIAN J. LEWIS & JANET W . REID

ACTA CARSOLOGICA 36/2 – 2007 281
associated with caves or karst, on the edge of the Knob-
stone Escarpment, in floyd County, Indiana. This well 
had been hand-dug many decades earlier, penetrating al-
luvium and then the top of the New Albany Shale to a to-
tal depth of 21 feet (6.4 m) below the surface. Also found 
in the well was the phreatobitic isopod Caecidotea tere-
sae, previously known from a series of drain tiles across 
the street on the Indiana University Southeast campus in 
New Albany, associated with an undescribed groundwa-
ter amphipod Stygobromus sp. (Lewis 1982). 
At the time that D. jeanneli was first collected from 
Marengo Cave, a relatively small amount of passage had 
been explored, and none of the known passages con-
tained a significant stream. Since that time, not only one, 
but two, cave “rivers” have been connected by explorers 
to the historic section of Marengo Cave. Lewis (1998) 
took plankton drift samples in these streams, but did not 
find D. jeanneli. f urther sampling revealed that these 
copepods are fairly common in drip pools in the upper 
level of the cave, even in shallow pools on the concrete 
trails used by tourists visiting the cave. The area of the 
cave where D. jeanneli was found is highly decorated with 
stalactites, stalagmites, and rimstone pools. In contrast to 
the lower-level cave rivers, all of the water in this part of 
the cave is epikarstic in origin.
Subsequently, D. jeanneli was found in five other 
southern Indiana caves, in pools fed by ceiling drips. In 
several instances, the stygobiont ostracod Pseudocandona 
jeanneli or a related, undescribed species of Pseudocan-
dona were taken from the same samples. Most recently, 
D. jeanneli has been taken from a seep emerging from al-
luvium in Clark County, Indiana. It appears that D. jean-
neli is an inhabitant of groundwater interstices, whether 
in a cave or not. The cave records may well be incidental 
findings, where the copepods have fallen in drip water 
from the preferred epikarst habitat. Clearly, this species 
is at home in non-karst groundwater interstices as well. 
Based on the collection of fresh material as noted, Reid 
(2004) redescribed the species and summarized the lo-
calities from which it is now known.
In contrast, present data suggest that megacyclops 
donnaldsoni is a strict cavernicole. Now known from 
sporadically located collections in Indiana, Kentucky, 
and Tennessee, all records are from caves, and all but one 
from a cave river. Reid (unpublished data) identified a 
single female ascribable to m. donnaldsoni, collected from 
a stream in Cemetery Pit Cave in Dade County in north-
west Georgia, January 2001, by J. Jensen and W . Reeves 
(specimen in VMNH). The single non-stream habitat 
was a deep rimstone pool, probably near local base-level, 
in Dry Cave, Grundy County, Tennessee. Bunting (1973) 
did not mention the subhabitat of his record from a cave 
in Tennessee. 
Diacyclops yeatmani has now been recorded from 
the hyporheos of the Blue River in southern Indiana, a 
drain tile in glaciated Illinois, and three caves in Tennes-
see. This is remarkable because the species is distributed 
both south of the Illinoian glacial maximum, and a sig-
nificant distance north of it, in the till plains of Illinois. 
A somewhat similar distribution pattern is known for D. 
indianensis, which is recorded from the type-locality in 
a cave in glaciated southeastern Indiana, and from seven 
caves in Tennessee. In Indiana D. indianensis occurs in a 
cave about 20 km outside the Illinoian glacial boundary 
region, whereas D. yeatmani occurs perhaps 80-100 km 
inside this boundary. 
Several other species of Diacyclops are presently 
known from one or two sites in southern Indiana: D. 
lewisi, D. salisae, D. conversus. All display some degree of 
morphological adaptation to the groundwater environ-
ment, and are known only from groundwater habitats. 
Otherwise, little is known of the ecology of these spe-
cies. 
Of note, serial sampling over a three-year period 
produced four species of Diacyclops from the Bou-Rouch 
pumpwell placed in a gravel bar on the Blue River in 
southern Indiana. These were D. sororum in 1997, D. con-
versus and D. yeatmani in 1998, and D. nearcticus in 2000 
(Lewis 1998, Reid 2004).
One of the typical habitats of D. nearcticus is the 
hyporheos of streams (Strayer and Reid, 1999). In In-
diana it has been found, using a Bou-Rouch pumpwell, 
inhabiting the hyporheic zone of the Blue River, as well 
as the interstices of stream gravel in Bond Cave, in Or-
ange County. This species has been otherwise found over 
a broad range in the eastern United States, encompass-
ing an area from Massachusetts to the Great Lakes and 
south to florida. The distribution is sporadic and it has 
never been found in abundance at any site. In florida, it 
is found in the Everglades karst in seasonally dry solution 
holes (Bruno et al. 2000). 
Another very widespread species is D. chrisae, which 
in Indiana was found in White County just north of the 
Wisconsinan glacial maximum. It was previously known 
only from interstitial sediments in eastern Maryland and 
Ontario (Reid 1992). 
Second to Diacyclops, cyclopoids of the genus Acan-
thocyclops were the most frequently recorded copepod 
inhabitants of groundwaters. Acanthocyclops robustus is 
generally considered a species of surface waters, but in-
spection of Appendix 1 shows it to be the most frequently 
collected of any of the copepods encountered. Some of 
the collection sites are remote from any point of surface 
input. This “surface” species is apparently very comfort-
able in groundwaters. A total of seven species of Acan-
thocyclops were recorded, suggesting that the group as a 
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ACTA CARSOLOGICA 36/2 – 2007 282
whole has more than a passing relationship to ground-
water habitats. f or example, the rare but widespread spe-
cies Acanthocyclops parasensitivus was found in a seep in 
Indiana, where it emerges from saturated interstices of 
gravel, and from drip pools of epikarstic origin in a cave 
in Tennessee. Several of the species of Acanthocyclops are 
apparently stygophilic or perhaps phreatophilic.
Appendix 1 includes a number of species that are, 
without a doubt, common fauna of surface habitats. 
Some, such as Acanthocyclops brevispinosus, Acantho-
cyclops einslei, Eucyclops agilis, and Eucyclops conrowae, 
are reported herein primarily from springs, rather than 
windows into deeper groundwaters. These collections 
represent copepods that are visitors to groundwater, but 
may find the cool, environmentally stable spring pools to 
be suitable habitat. The calanoid Skistodiaptomus pallidus 
may well be only an accidental traveler through ground-
water conduits. The group consisting of Osphranticum 
labronectum, A. vernalis, m. albidus, and Orthocyclops 
modestus, found in a pool in Patton Cave, Tennessee, is 
typical of small, surface waterbodies, and may have en-
tered through seepage from a nearby lake.
Contrary to what might have been expected for the 
strictly benthic harpacticoids, most of the species found 
are very widespread and common in surface habitats in 
North America. Members of the genera Attheyella, Bryo-
camptus, and moraria are especially common in spring 
seeps and wet moss. The exception is Bryocamptus mor-
risoni s.str., which is known only from two cave-river col-
lections in Indiana and Kentucky.
f inally, of the genera in our list, Rheocyclops and 
Itocyclops remain the most enigmatic. Rheocyclops indi-
ana was first discovered in a drip pool in Ladder Cave, 
Washington County, Indiana, in the company of an un-
described species of the amphipod genus Stygobromus. 
This Rheocyclops was subsequently found in a drip pool 
in Elrod Cave, Orange County. The first impression was 
that this species was probably endemic to epikarstic habi-
tats in the south-central karst belt of southern Indiana. 
This notion was eliminated with the discovery by Reid 
(unpublished data) of R. indiana from two hyporheic 
habitats in the Great Smoky Mountains National Park. 
These finds suggest that this species is relatively wide-
spread, but rarely collected, probably because of its cryp-
tic nature and small size.
Itocyclops yezoensis was formerly known from Ja-
pan and Alaska, when it was subsequently reported 
from Great Smoky Mountains National Park, in Sevier 
County, Tennessee (Reid and Ishida 2000); and recently 
from a streambank in Henry County, southern Virginia 
(Reid 2006). Although we were surprised at the range 
extension of Rheocyclops from Indiana to the Smoky 
Mountains, this distance pales in comparison to that of I. 
yezoensis. A second North American species of Itocyclops 
was recently discovered in caves that lie in the path of 
an interstate highway project in eastern Kentucky (Lewis 
and Lewis 2006a), and also in a cave in north-central 
Tennessee (present report). The third known species of 
the genus was found in a spring seep in western Virginia 
(Reid, in review).
PROCESSES Of SUBTERRANEAN INVASION
In Indiana, where our sampling has been most concen-
trated, the availability of aquatic habitats can be estimat-
ed by the glacial history of the state. The glacial history of 
Indiana (summarized here from IGS 2007) is quite com-
plex and the record is incomplete, as successive ice sheets 
eroded the deposits of the previous glacial advances and 
created discontinuities in the geologic record. The present 
theory of glacial geologists is that the ice sheets initially 
progressed along valley systems during the Tertiary, in 
the area now comprising the boundary between the U.S. 
and Canada. As the glaciers moved southward, these val-
leys were carved into large basins that would become the 
present-day Great Lakes. The orientations of these basins 
in turn strongly influenced the direction and magnitude 
of glaciations in Indiana.
The climatic record derived from cores of extant 
glaciers in Greenland as well as ocean floor geochemical 
evidence suggest 12-18 periods over the last two million 
years that were cold enough to form continental glaciers. 
In Indiana, there is evidence that at least 11 distinct gla-
cial events occurred. These are divided into three major 
stages, each of which had several ice advances:
(1) pre-Illinoian Stage, which is the least under-
stood because of erosion and burial of the deposits. It 
comprises the period between initial glaciation and the 
Illinoian (700,000 - 300,000 years before present [BP]). 
The first Pleistocene ice sheet to reach Indiana appears 
to have entered the state from directly to the north (now 
Michigan). The glacier, thought to have been between 
a half mile and a mile (0.8-1.6 km) thick at the leading 
edge, moved slowly as it encountered bedrock hills and 
valleys that slowed its progress. The Teays River valley, 
the major drainage conduit for the eastern U.S., was 200-
400 feet (60-120 m) deep prior to glaciation. After bury-
ing this large stream valley, the first glacier stopped about 
mid-state, then slowly receded to the north. 
JULIAN J. LEWIS & JANET W . REID

ACTA CARSOLOGICA 36/2 – 2007 283
(2) Illinoian Stage, of which deposits are today ex-
posed on the surface in southeast and southwest Indi-
ana, lasted from about 300,000 to 140,000 years BP . The 
southeastern Indiana karst was completely glaciated 
during this stage. The area comprising the south-central 
karst area remained ice-free, but the limestone band con-
tinues to the north where it is buried under the till plain. 
During the subsequent Sangamon interglacial, the flat-
tened topography of the southeastern karst was dissected 
to some extent by streams down-cutting through the till. 
The modern-day Ohio River was formed and became a 
major drainage conduit for the water from the melting 
glaciers.
 (3) Wisconsin Stage, from about 50,000 to 10,000 
years ago. The northern half of Indiana is covered with 
deposits of this age. The caves in the southeastern Indiana 
karst region lie under Illinoian till, but almost all occur 
south of the Wisconsinan glacial boundary. The occur-
rence of enterable caves ends near where the limestone 
strata meet the edge of the Wisconsinan glacial boundary 
in Decatur County, although other caves may certainly 
exist buried under the carpet of till.
 Thus, different areas of Indiana, including the 
two karst areas, have been free of glaciers for relatively 
well-established time periods. The south-central karst 
has been ice-free and suitable, at least in that respect, 
throughout the Pleistocene. The southern-most area 
of the southeastern karst, in Clark, Jefferson, Jennings, 
and Ripley counties, was available from the Sangamon 
onward. The sites sampled in Vermilion Co., Illinois and 
White Co., Indiana both lie on Wisconsinan -age till 
plains. 
This brings us to a point of controversy in the theo-
ries concerning the zoogeography and evolution of sub-
terranean crustaceans in glaciated North America. How 
did tiny, substrate-bound animals such as Diacyclops 
yeatmani, D. indianensis, D. salisae, D. lewisi, or the more 
heavily collected (because they are easily visible to the 
naked eye) amphipod Bactrurus mucronatus and isopod 
Caecidotea kendeighi get to their present-day ranges deep 
within the glaciated areas? Two hypotheses have been 
advanced to account for this distribution pattern (Hols-
inger 1978, 1981, 1986; Lewis 1983, Lewis and Bowman 
1981, Koenemann & Holsinger 2001): (1) survival under 
the ice; (2) dispersal from areas that remained ice-free 
after the retreat of the glaciers. 
These authors have recognized that some combina-
tion of the two processes might well have occurred. We 
elaborate on a refinement of this idea. The continental 
glaciers were essentially rivers of slowly flowing ice, mov-
ing southward from their sources. It is thought that af-
ter the initial glaciation of Indiana, subsequent glaciers 
moved relatively quickly due to abundant water at the 
base of the glacier that served to lubricate the ice sliding 
over previously glaciated terrain. At the front edge of the 
glacier, the ice was constantly melting. During periods 
when melt rates were slow, the glaciers advanced, but re-
ceded when melting outpaced deposition of new ice.
Aquatic habitat for cold-stenothermic crustaceans 
should have been abundant. Peripheral to the glaciers, 
suitable habitat might have been available in three areas: 
(1) streams and meltwater lakes in front (south) of the 
glaciers; (2) the leading edge of the glacier where melting 
was occurring; (3) underneath the glacier, particularly in 
a zone behind the leading edge.
If the glaciers were in fact moving on a lubricating 
base of liquid water, if other factors were not limiting 
(e.g., dissolved oxygen, nutrients, scouring), then we see 
no reason to think that crustaceans would not have been 
present, particularly in the zone under the edge of the 
glacier. Beyond the glacier, meltwater lakes should have 
provided ideal habitats. This meltwater zone would have 
traveled northward with the receding glaciers, and the 
community present in it would have migrated northward 
as well. This would account for the presence of stygobi-
ont species such as Diacyclops yeatmani and D. chrisae far 
north of the glacial maxima.  
As the present interglacial proceeded, the disap-
pearance of meltwater lakes in southern Indiana would 
have encouraged the invasion of the other, major aquatic 
habitat that remained available: groundwater. When 
lakes and streams draining from the receding glaciers 
dried, some of the cyclopoids followed the water into the 
interstices of the saturated glacial till. Evidence for this 
is the presence of Diacyclops salisae and D. lewisi in this 
habitat today. Robertson and Milner (2006) reported that 
Diacyclops spp. were the most characteristic members of 
the meiofauna in newly formed meltwater streams (ca. 
50 years post recession of the glacier) in Glacier Bay Na-
tional Park, Alaska. 
Eleven species of Diacyclops have now been found 
during sampling of groundwater habitats in Indiana. 
Reid and Strayer (1994) reviewed the morphological 
modifications seen in subterranean copepods, some of 
which are noted in Table 1. Of the species of Diacyclops 
known from Indiana, D. crassicaudis brachycercus and D. 
navus exhibit slight or no morphological adaptations to 
subterranean life, and are most often found in epigean 
habitats. Diacyclops lewisi and D. salisae are also rela-
tively unmodified, and occur in wells. Diacyclops nearcti-
cus and D. sororum occur in both epigean situations and 
hyporheic sediments. Diacyclops chrisae, D. conversus, D. 
indianensis, and D. yeatmani have been found in deep 
interstitial and hyporheic situations. Diacyclops jeanneli 
occurs mainly in cave pools, but as suggested above may 
well be an inhabitant of the epikarst. Most of these lat-
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ACTA CARSOLOGICA 36/2 – 2007 284
ter species exhibit at least some morphological modifica-
tions.  
f or subterranean cyclopoid copepods found in In-
diana, Table 1 summarizes a comparison of morphologi-
cal adaptation as a function of the length of time available 
for colonization of the region. Of the Diacyclops species, 
D. jeanneli and D. conversus are the most strongly modi-
fied for subterranean existence, and both occur only in 
the unglaciated area of south-central Indiana. Indeed, 
D. jeanneli has been found to the very edge of the limit 
of Illinoian glaciation on the sides of the Knobstone Es-
carpment in Clark and floyd counties. Strayer and Reid 
(1999) analyzed the distribution of hyporheic cyclopoids 
in glaciated versus unglaciated areas in the eastern Unit-
ed States, and noted a break in species composition as-
sociated with the Pleistocene glacial maxima. They sug-
gested that the more specialized interstitial species, such 
as Diacyclops jeanneli and members of Rheocyclops, have 
been unable to disperse into the glaciated regions. Our 
sampling data lend some additional evidence to this the-
ory, although the correlation is clouded by the differences 
in microhabitat preferences among the species. f urther 
evidence is provided by the near-absence of a specialized 
subterranean harpacticoid fauna in this region, where, 
apparently, mainly the most adaptable generalist species 
were able to reach the subterranean habitats.
Tab. 1. Species of Diacyclops, megacyclops, and Rheocyclops known to inhabit different regions in Indiana, according to the length 
of time available since the last glaciation in that region, and the degree of morphological modification of each species. The degree of 
modification is estimated according to the number of several characters (total given in parentheses): oligomerization of the antennular 
segments, loss of one or more antennal basipodite setae, oligomerization of segments of the swimming legs, shortening of the terminal 
endopodite segment of the leg 4 endopodite, and the anal operculum posteriorly produced.
Time of last glaciation Species Relative degree of modification
Unglaciated Megacyclops donnaldsoni Slight (1)
Diacyclops navus* None
Diacyclops crassicaudis brachycercus* Moderate (2)
Diacyclops nearcticus* Moderate (2)
Diacyclops yeatmani Moderate (2)
Diacyclops conversus Moderate (3)
Diacyclops jeanneli Moderate (3)
Rheocyclops indiana Advanced (4)
Illinoian glaciation, ice-free  
ca. 140,000 years BP
Diacyclops lewisi None
Diacyclops salisae None
Diacyclops indianensis Moderate (2)
Wisconsinan glaciation, ice-free  
ca. 10,000 years BP
Diacyclops chrisae* Moderate (2)
Diacyclops yeatmani Moderate (2)
* Also occurs in previously glaciated regions elsewhere in North America.
ACKNOWLEDGEMENTS
We are grateful to all of the collectors who helped gath-
er material over the years, particularly Salisa L. Lewis, 
Ronnie Burns, Diane Tecic, Heather Garland, and Cory 
Holliday. JJL gratefully acknowledges the support of The 
Nature Conservancy, Indiana Natural Heritage Program 
(Indiana Department of Natural Resources), Hoosier 
National f orest (U.S.D.A. f orest Service), Big Oaks Na-
tional Wildlife Refuge (U.S. f ish and Wildlife Service), 
and the Tennessee Wildlife Resources Agency, as well as 
funds from the environmental impact studies for federal 
interstate highway projects I-69 and I-66 administered 
by the Kentucky Transportation Cabinet and the Indiana 
Department of Transportation. JWR gratefully acknowl-
edges the receipt of material support from the Virginia 
Museum of Natural History and the National Museum of 
Natural History (Smithsonian Institution).
JULIAN J. LEWIS & JANET W . REID

ACTA CARSOLOGICA 36/2 – 2007 285
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Cave System of Kentucky I: The biota.- International 
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(florida): descriptions of two new species of Ela-
phoidella (Harpacticoida, Canthocamptidae), and 
supplementary description of Diacyclops nearcticus 
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1171-1204, Leiden.
Bunting, D. L., 1973: The Cladocera and Copepoda of 
Tennessee. II. Cyclopoid copepods.- Journal of the 
Tennessee Academy of Science, 48, 138-141, Mur-
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Chappuis, P . A., 1929: Copépodes cavernicoles de 
l’ Amérique du Nord (Note préliminaire).- Bulletin 
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Chappuis, P . A., 1931: Campagne spéologique de C. Bo-
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f enneman, N. E., 1938: Physiography of the eastern 
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amphipod genus Stygobromus (Crangonyctidae), 
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Holsinger, J. R., 1981: Stygobromus Canadensis, a troglo-
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the North American subterranean amphipod genus 
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descriptions of Caecidotea teresae, n. sp., and C. 
macropropoda Chase and Blair (Crustacea: Isopoda: 
Asellidae).- Proceedings of the Biological Society of 
Washington, 95, 338-346, Lawrence.
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brates of glaciated southeastern Indiana.- National 
Speleological Society Bulletin, 45, 34-40, Hunts-
ville.
Lewis, J. J., 1996: Inventory of the subterranean biota 
threatened by urbanization of Clark and floyd 
Counties, Indiana. f inal report, Non-game and En-
dangered Wildlife Program, Division of f ish and 
Wildlife, Indiana Department of Natural Resources, 
71 pages.
Lewis, J. J., 1998: The subterranean fauna of the Blue River 
area. f inal Report, The Nature Conservancy, Species 
at Risk Program (USGS), Indiana Natural Heritage 
Program and Indiana Non-game and Endangered 
Wildlife Program, 266 pages.
Lewis, J. J., 1999: The subterranean fauna of the Interior 
Low Plateaus. f inal Report, The Nature Conservan-
cy, 26 pages.
Lewis, J. J., 2001: A biological reconnaissance of the Rum-
bling f alls Cave System, Van Buren County, Tennes-
see. f inal Report, Tennessee Environmental Coun-
cil, National Speleological Society, World Wildlife 
f und, 21 pages.
Lewis, J. J., 2002: Status and distribution surveys for rare 
cave-dependent organisms recently identified from 
the Rumbling f alls Cave System, Van Buren County, 
Tennessee. f inal Report, U.S. f ish & Wildlife Ser-
vice, 48 pages.
Lewis, J. J., 2004: Interstate 66 karst faunal study. Envi-
ronmental Impact Statement. 
Lewis, J. J., 2005: Cave fauna of the I69 Corridor in Mon-
roe County, Indiana. Environmental Impact State-
ment, Baker Engineering, Inc. and Ozark Under-
ground Laboratory.
Lewis, J. J. & Bowman, T. E., 1981: The subterranean asel-
lids (Caecidotea) of Illinois (Crustacea: Isopoda: 
Asellidae).- Smithsonian Contributions to Zoology, 
335, 1-66, Washington, D.C.
Lewis, J. J., Burns, R. & Lewis, S., 2004: The subterranean 
fauna of the Hoosier National f orest. U.S. Depart-
ment of Agriculture, f orest Service, 190 pages.
Lewis, J. J. & Lewis, S. L., 2005: Bioinventory of the sub-
terranean fauna of Tims f ord State Park, f ranklin 
County, Tennessee. Tennessee Natural Heritage 
Program, 28 pages.
REf ERENCES
PATTERNS AND PROCESSES Of G ROUNDWATER INVASION By C OPEPODS IN THE INTERIOR LOW PLATEAUS ...

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Lewis, J. J. & Lewis, S. L., 2006a: Cave f auna in the Inter-
state 66 Somerset to London Corridor. In: Rea, G. 
T., editor, Proceedings of the 2005 National Cave & 
Karst Management Symposium, Albany, New york, 
15-20.
Lewis, J. J. & Lewis, S. L., 2006b: Subterranean fauna of 
the Buddha Karst Preserve, Lawrence County, In-
diana. Indiana Department of Natural Resources, 
Division of Nature Preserves, 19 pages.
Lewis, J. J. & Rafail, S. T., 2002: The subterranean fauna 
of the Big Oaks National Wildlife Refuge. U.S. De-
partment of the Interior, f ish & Wildlife Service, 
Indiana Department of Natural Resources, Natural 
Heritage Program, 77 pages.
Lewis, J. J., Moss, P ., Tecic, D. & Nelson, M., 2002: A Con-
servation f ocused Inventory of the subterranean in-
vertebrates of the Southwestern Illinois Karst.- Jour-
nal of Cave and Karst Studies, 65, 9-21, Huntsville.
Reid, J. W ., 1988: Copepoda (Crustacea) from a seasonal-
ly flooded marsh in Rock Creek Stream Valley Park, 
Maryland.- Proceedings of the Biological Society of 
Washington, 101, 31-38, Washington, D.C.
Reid, J. W ., 1992: Redescription of Diacyclops nearcticus 
(Kiefer, 1934) and description of four similar new 
congeners from North America, with comments on 
D. crassicaudis (G.O. Sars, 1863) and D. crassicaudis 
var. brachycercus (Kiefer, 1927)(Crustacea: Copepo-
da).- Canadian Journal of Zoology, 70, 1445-1469, 
Ottawa.
Reid, J. W ., 2004: New records and new species of the ge-
nus Diacyclops (Crustacea; Copepoda) from subter-
ranean habitats in southern Indiana, USA.- Jefferso-
niana, 12, 1-65, Martinsville.
Reid, J. W ., 2006: Local recreation parks as hospitable 
habitats for small aquatic animals: examples of co-
pepod crustaceans in Virginia.- Banisteria, 27, 10-
15, Richmond.
Reid, J. W ., In review: Two new species of Itocyclops, and 
a new record of I. yezoensis (Copepoda: Cyclopoi-
da: Cyclopidae) from the eastern U.S.A.- Journal of 
Crustacean Biology, Lawrence.
Reid, J. W . & Strayer, D. L., 1994: Diacyclops dimorphus, 
a new species of copepod from florida, with com -
ments on morphology of interstitial cyclopine cyclo-
poids.- Journal of the North American Benthologi-
cal Society, 13: 250-265, Lawrence.
Reid, J. W ., Strayer, D. L., McArthur, J. V ., Stibbe, S. E. & 
Lewis, J. J., 1999: Rheocyclops, a new genus of co-
pepods form the southeastern and central U.S.A. 
(Copepoda: Cyclopoida: Cyclopidae).- Journal of 
Crustacean Biology, 19, 384-396, Lawrence.
Reid, J. W . & Ishida, T., 2000: Itocyclops, a new genus 
proposed for Speocyclops yezoensis (Copepoda: Cy-
clopoida: Cyclopidae).- Journal of Crustacean Biol-
ogy, 20, 589-596, Lawrence.
Robertson, A. L. & Milner, A. M., 2006: The influence of 
stream age and environmental variables in structur-
ing meiofaunal assemblages in recently deglaciated 
streams.-Limnology and Oceanography, 51, 1454-
1465, Lawrence. 
Strayer, D. L. & Reid, J. W ., 1999: Distribution of hypo-
rheic cyclopoids (Crustacea: Copepoda) in the east-
ern United States.- Archiv für Hydrobiologie, 145, 
79-92, Stuttgart.
yeatman, H. C., 1964: A new cavernicolous cyclopoid 
copepod from Tennessee and Illinois.- Journal of 
the Tennessee Academy of Science, 39, 95-98, Mur-
freesboro.
Appendix 1. Most of the following records of copepods 
from groundwater habitats in and near the Inte-
rior Low Plateaus are taken from the unpublished 
reports of Lewis, and Lewis et al. (1996-2006). Re-
cords from Kofoid (1899), Chappuis (1929, 1931), 
yeatman (1964), Barr (1967), Bunting (1973), Stray-
er & Reid (1999), and Reid (2004) have also been 
included.
ORDER COPEPODA
SUBORDER CYCLOPOIDA
FAMILY CYCLOPIDAE
Acanthocyclops brevispinosus (Herrick, 1884) 
 ILLINOIS: Monroe Co.: Camp Vandeventer Spring.
Acanthocyclops einslei Mirabdullayev & Defaye, 2004
 ILLINOIS: Monroe Co.: Bat Sump Cave; INDIANA: 
Greene Co.: Rankin Spring.
Acanthocyclops exilis (Coker, 1934)
 KENTUCKy: Edmonson Co.: Mammoth Cave 
(Barr 1967).
Acanthocyclops parasensitivus Reid, 1998
 INDIANA: Clark Co.: Seep spring, Burns Hollow; 
TENNESSEE: Grundy Co.: Big Mouth Cave; KEN-
TUCKy: Pulaski Co.: Buck Creek, sandbar (Strayer 
& Reid 1999).
 
Acanthocyclops robustus (G.O. Sars, 1863) 
 ILLINOIS: Monroe Co.: Bat Sump Cave; INDIANA: 
Harrison Co.: Baker Hollow Cave, Black Medusa 
Cave, Bussabarger’s Cave, Harrison Cave Spring, 
JULIAN J. LEWIS & JANET W . REID

ACTA CARSOLOGICA 36/2 – 2007 287
Harrison Spring, Stygian River Cave, Wallier Cave; 
Jefferson Co.: well, Grapevine Branch section 19, 
Big Oaks National Wildlife Refuge (BONWR); Law-
rence Co.: Buddha Cave, Burton Hollow Cave, Hen-
shaw Bend Cave; Orange Co.: Wesley Chapel Gulf 
Cave; Washington Co.: Endless Cave; KENTUCKy: 
Edmonson Co.: Mammoth Cave (Chappuis 1931); 
TENNESSEE: White Co.: Lost Creek Cave.
Acanthocyclops venustoides (Coker, 1934) 
 INDIANA: Crawford Co.: well at abandoned house 
on road to Mesmore Cliffs area; Harrison Co., Bus-
sabarger’s Cave; Jefferson Co.: well, NE quarter-sec-
tion 7, Big Creek drainage, BONWR; White Co.: 
f ield drain tile outlet, 5.3 km west of Brookston; IL-
LINOIS: Vermilion Co.: drain tile outlet near Jordan 
Creek (yeatman 1964).
Acanthocyclops vernalis (f ischer, 1853) 
 INDIANA: Clark Co.: Seep spring on stream bank, 
Burns Hollow; ILLINOIS: Monroe Co.: Bat Sump 
Cave; KENTUCKy: Edmonson Co.: Mammoth 
Cave (Chappuis 1931); TENNESSEE: Grundy Co.: 
Bigmouth Cave (yeatman 1964); Rhea Co.: Marler 
Cave; Rutherford Co.: Patton Cave, pool; Jackson 
Co.: Haile Cave, rimstone pools; ILLINOIS: Vermil-
ion Co.: drain tile outlet near Jordan Creek (yeat-
man 1964).
Diacyclops bicuspidatus (Claus, 1857)
 KENTUCKy: Edmonson Co.: Mammoth Cave (Ko-
foid 1899). This record may refer to D. thomasi (S. 
A. f orbes, 1882), which was long considered a sub-
species of D. bicuspidatus, and was frequently con-
fused with the former in the earlier North American 
literature. Both D. bicuspidatus and D. thomasi are 
widespread in North America; D. thomasi is much 
more common, especially in the plankton of lakes 
and ponds, and also occurs in smaller waterbodies 
such as rock pools. This record is not treated further 
herein.
Diacyclops chrisae Reid, 1992
 INDIANA: White Co.: f ield drain tile outlet, 5.3 km 
west of Brookston.
Diacyclops conversus Reid, 2004
 INDIANA: Crawford Co.: Bou-Rouch pumpwell on 
Blue River.
Diacyclops crassicaudis brachycercus Kiefer, 1927
 INDIANA: Harrison Co.: Hiser Spring Cave.
 
Diacyclops jeanneli Chappuis, 1929 
 INDIANA: Clark Co.: Seep spring in Burns Hollow; 
Crawford Co.: Marengo Cave; floyd Co.: well, New 
Albany near Indiana University Southeast; Harri-
son Co.: Devils Graveyard Cave; Orange Co.: Ap-
ple Cave, Murray Spring Cave, Tucker Lake Spring 
Cave; Washington Co.: f redericksburg Cave.
Diacyclops indianensis Reid, 2004 
 INDIANA: Jefferson Co.: Henry Dilk f alls Cave; 
TENNESSEE: f entress Co.: Redbud Cave; f ranklin 
Co.: Crownover Saltpeter Cave, Keith Cave, Little 
Crow Creek Cave; Grundy Co.: Big Mouth Cave, 
Dry Cave; Pickett Co.: Bunkum Cave.
Diacyclops lewisi Reid, 2004
 INDIANA: Jefferson Co.: well in Big Creek drainage 
section 7, BONWR: Jennings Co.: well in Graham 
Creek drainage section 18, BONWR.
Diacyclops navus (Herrick, 1882)
 INDIANA: floyd Co.: abandoned cistern near inter -
section of I-265 and Grant Line Road; Gibson Co.: 
well 5.6 km east of f ort Branch.
Diacyclops nearcticus Kiefer, 1934 
 INDIANA: Crawford Co.: Bou-Rouch pumpwell on 
Blue River; Orange Co.: Bond Cave; KENTUCKy: 
Pulaski Co.: Stab Cave; TENNESSEE: White Co.: 
Lost Creek Cave; DeKalb Co.: Smith f ork, gravel 
bar (Strayer & Reid 1999).
Diacyclops salisae Reid, 2004 
 INDIANA: Jennings Co.: well, Otter Creek drain-
age, section 31, BONWR.
Diacyclops sororum Reid, 1992 
 INDIANA: Crawford Co.: Bou-Rouch pumpwell on 
Blue River; Harrison Co.: Squire Boone Caverns; 
KENTUCKy: Pulaski Co.: Buck Creek, sandbar 
(Strayer & Reid 1999).
Diacyclops yeatmani Reid, 1988 
 INDIANA: Crawford Co.: Bou-Rouch pumpwell on 
Blue River; TENNESSEE: Grundy Co.: Big Mouth 
Cave (yeatman 1964, as Cyclops clandestinus); Van 
Buren Co.: Camps Gulf Cave, Swamp River Cave; 
ILLINOIS: Vermilion Co.: drain tile outlet near Jor-
dan Creek (yeatman 1964).
Diacyclops undescribed species
 KENTUCKy: Pulaski Co.: Blowing Cave.
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ACTA CARSOLOGICA 36/2 – 2007 288
Eucyclops agilis (Koch, 1838) 
 ILLINOIS: Monroe Co.: Haney Spring; INDIANA: 
Greene Co.: Rankin Spring; Orange Co.: Rise of Lost 
River at Orangeville; KENTUCKy: Edmonson Co.: 
Mammoth Cave [Kofoid 1899, Chappuis 1931 (as 
E. serrulatus), Barr 1967]; Pulaski Co.: Buck Creek, 
sandbar (Strayer & Reid 1999); TENNESSEE: Grun-
dy Co.: Bigmouth Cave (yeatman 1964); DeKalb 
Co.: Smith f ork, gravel bar (Strayer & Reid 1999); 
Rutherford Co.: Herring Cave.
Eucyclops conrowae Reid, 1992
 INDIANA: Harrison Co.: f iretail Spring; Orange 
Co.: Rise of Lost River at Orangeville.
Eucyclops elegans (Herrick, 1884) 
 INDIANA: Crawford Co.: Marengo Cave; Greene 
Co.: Rankin Spring; Monroe Co.: Harp Spring Cave; 
Orange Co.: Rise of Lost River at Orangeville; KEN-
TUCKy: Edmonson Co.: Mammoth Cave (Chap-
puis 1931); TENNESSEE: White Co.: Lost Creek 
Cave.
Itocyclops undescribed species 
 KENTUCKy: Pulaski Co.: Sheep Cave South, rim-
stone pool; Stab Cave, rimstone pool; Redbud Cave, 
rimstone pool; TENNESSEE: Jackson Co.: Dud 
Cave, shallow drip pool. 
macrocyclops albidus (Jurine, 1820) 
 ILLINOIS: Monroe Co.: Bat Sump Cave; INDIANA: 
Greene Co.: Ashcraft Cave; Lawrence Co.: Buddha 
Cave; Monroe Co.: Well Cave; Orange Co.: Rise of 
Lost River at Orangeville; KENTUCKy: Edmonson 
Co.: Mammoth Cave (Kofoid 1899); TENNESSEE: 
Rutherford Co.: Herring Cave, pool; Patton Cave, 
pool.
macrocyclops fuscus (Jurine, 1820) 
 INDIANA: Greene Co.: Rankin Spring; Harrison 
Co.: Stygian River Cave; KENTUCKy: Edmonson 
Co.: Mammoth Cave (Chappuis 1931, Barr 1967).
megacyclops donnaldsoni (Chappuis, 1929) 
 INDIANA: Crawford Co.: Hidden Spring Cave, 
Wildcat Cave; Lawrence Co.: Donnaldson Cave 
(Chappuis 1931); KENTUCKy: Edmonson Co.: 
Mammoth Cave, Roaring River (Barr 1967); TEN-
NESSEE: Grundy Co.: Dry Cave; McMinn Co.: un-
named cave (Bunting 1973).
megacyclops latipes (Lowndes, 1927) 
 INDIANA: Harrison Co.: Rands Spring; Monroe 
Co.: Well Cave; TENNESSEE: Grundy Co.: Big 
Mouth Cave, Trussell Cave; Rhea Co.: Marler Cave.
megacyclops undescribed species 
 INDIANA: Martin Co.: Bluff House Cave; Orange 
Co.: Campground Cave.
mesocyclops edax (S.A. f orbes, 1891) 
 ILLINOIS: Monroe Co.: Camp Vandeventer Spring, 
Camp Vandeventer Cave; INDIANA: Crawford Co.: 
Hidden Spring Cave; KENTUCKy: Edmonson Co.: 
Mammoth Cave (Chappuis 1931; Barr 1967).
microcyclops rubellus (Lilljeborg, 1901) 
 ILLINOIS: St. Clair Co.: Stemler Cave; INDIANA: 
Crawford Co.: Bou-Rouch pumpwell on Blue River; 
KENTUCKy: Pulaski Co.: Buck Creek, sandbar 
(Strayer & Reid 1999); TENNESSEE: DeKalb Co.: 
Smith f ork, gravel bar (Strayer & Reid 1999).
Orthocyclops modestus (Herrick, 1883) 
 ILLINOIS: Monroe Co.: Bat Sump Cave; INDIANA: 
Crawford Co.: well at abandoned house near Mes-
more Cliffs; Harrison Co.: Binkley Cave; Jefferson 
Co.: well, Grapevine Branch section 19, BONWR; 
TENNESSEE: Rutherford Co.: Patton Cave, pool; 
Grundy Co.: Bigmouth Cave (Bunting 1973).
Paracyclops canadensis (Willey, 1934) 
 INDIANA: Crawford Co.: Carnes Mill Spring Cave.
 
Paracyclops chiltoni (Thomson, 1882) 
 INDIANA: Harrison Co.: Binkley Cave, Rands 
Spring; Lawrence Co.: Henshaw Bend Cave; Wash-
ington Co.: Mill Cave; KENTUCKy: Edmonson 
Co.: Mammoth Cave [Chappuis, 1931, reported 
P . finitimus Kiefer, 1929, but Chappuis’ record was 
interpreted by Barr, 1967 as P . fimbriatus (f ischer, 
1853). Paracyclops finitimus is currently considered 
a junior subjective synonym of P . chiltoni, and P . fim-
briatus is not thought to occur in the Americas; see 
Karaytug & Boxshall 1998]; Pulaski Co: Buck Creek, 
sandbar (Strayer & Reid 1999). 
Rheocyclops indiana Reid, 1999
INDIANA: Orange Co.: Elrod Cave; Washington Co.: 
Ladder Cave.
Rheocyclops undescribed species
 ALABAMA: Jackson Co.: Russell Cave (Russell 
Cave National Monument).
JULIAN J. LEWIS & JANET W . REID

ACTA CARSOLOGICA 36/2 – 2007 289
Tropocyclops extensus Kiefer, 1931
 INDIANA: Harrison Co.: Bussabargers Cave, Squire 
Boone Caverns.
Tropocyclops prasinus (f ischer, 1860)
 KENTUCKy: Edmonson Co.: Mammoth Cave 
(Chappuis 1931). Chappuis’ record of T. prasinus 
may refer to the subspecies mexicanus later de-
scribed by Kiefer.
Tropocyclops prasinus mexicanus Kiefer, 1938 
 ILLINOIS: Monroe Co.: Camp Vandeventer Spring, 
Camp Vandeventer Cave; INDIANA: Washington 
Co.: Endless Cavern; KENTUCKy: Edmonson Co.: 
Mammoth Cave (Barr 1967).
SUBORDER HARP ACTICOIDA
FAMILY CANTHOCAMPTIDAE
Attheyella (Neomrazekiella) illinoisensis (S. A. f orbes, 
1882) 
 INDIANA: Crawford Co.: Megenity Peccary Cave; 
Monroe Co.: May Cave.
Attheyella (Neomrazekiella) nordenskioldii (Lilljeborg, 
1902) 
 ILLINOIS: Monroe Co.: Haney Spring, Metter Cave; 
INDIANA: Clark Co.: seep spring in Burns Hollow; 
Crawford Co.: Wymans Cave; Harrison Co.: Rands 
Spring; Jefferson Co.: well, NW quarter-section 18, 
Graham Creek drainage BONWR; Monroe Co.: 
Shirley Springs, Waterfall Cave; Washington Co.: 
Karens Pit Cave; Orange Co.: Garlow Spring Cave. 
Attheyella (Ryloviella) pilosa Chappuis, 1929
 INDIANA: Clark Co.: seep spring in Burns Hollow; 
Crawford Co.: Wymans Cave; Harrison Co.: Squire 
Boone Caverns; Monroe Co.: Waterfall Cave; Wash-
ington Co.: Baynes Spring Cave, Endless Cavern, 
Nevin Spring Cave; KENTUCKy: Edmonson Co.: 
Mammoth Cave (Chappuis 1931).
Bryocamptus (Bryocamptus) newyorkensis (Chappuis, 
1927)
 TENNESSEE: Jackson Co.: Dud Cave, shallow drip 
pool.
Bryocamptus (Limocamptus) morrisoni morrisoni (Chap-
puis, 1929)
 INDIANA: Lawrence Co.: Donnaldson Cave (Chap-
puis 1929, 1931).
Bryocamptus (Limocamptus) morrisoni elegans (Chap-
puis, 1929)
 KENTUCKy: Hart Co.: Horse (Hidden River) Cave 
(Chappuis 1929, 1931).
Bryocamptus (Limocamptus) nivalis (Willey, 1925) 
 INDIANA: Harrison Co.: Harrison Cave Spring.
Bryocamptus (Limocamptus) undescribed species 
 TENNESSEE: Overton Co.: Mill Hollow Cave.
moraria cristata Chappuis, 1929
 INDIANA: Lawrence Co.: Donnaldson Cave (Chap-
puis 1931).
SUBORDER CALANOIDA
FAMILY CENTROP AGIDAE
Osphranticum labronectum S. A. f orbes, 1882
 TENNESSEE: Rutherford Co.: Patton Cave, pool.
FAMILY DIAPTOMIDAE
Skistodiaptomus pallidus (Herrick, 1879) 
 INDIANA: Harrison Co.: Harrison Cave Spring, 
Stygian River Cave; KENTUCKy: Edmonson Co.: 
Mammoth Cave (Barr 1967)
PATTERNS AND PROCESSES Of G ROUNDWATER INVASION By C OPEPODS IN THE INTERIOR LOW PLATEAUS ...