Hydrobiologia 453/454: 201–226, 2001.
R.M. Lopes, J.W. Reid & C.E.F. Rocha (eds), Copepoda: Developments in Ecology, Biology and Systematics.
© 2001 Kluwer Academic Publishers. Printed in the Netherlands.
201
A human challenge: discovering and understanding continental copepod
habitats
Janet W. Reid
Department of Systematic Biology-Invertebrate Zoology, National Museum of Natural History, Smithsonian
Institution, Washington, DC 20560-0163, U.S.A.
E-mail: reid.janet@nmnh.si.edu
Key words: Cyclopoida, Harpacticoida, habitats, leaf litter, phytotelmata, artificial habitats, review
Abstract
Copepods have invaded an astonishing variety of aquatic and humid continental environments and microhabitats.
The historical process of discovery and investigation of copepods in ephemeral, acid and thermal waters, subterranean waters and sediments, phytotelmata, humid soils, leaf litter, human-modified and artificial habitats, and
other situations extends over about 130 years. The methods developed to collect in and study these habitats range
from simple nets to elaborate pumping systems and diving techniques. Investigations of non-lacustrine continental
environments have contributed greatly to the understanding of aspects of copepod biology such as reproduction,
diapause and population dynamics. Questions regarding faunistics and biological diversity, biogeography, evolution, transport and introductions of alien species have also been informed by such studies. This article briefly
reviews these topics, and provides detailed lists of records from some of the less well-known kinds of habitats.
Introduction
Free-living copepod crustaceans are customarily
treated as aquatic in the general limnological literature; however, some 130 years of imaginative collecting have revealed that they are capable of invading and
surviving successfully in nearly every continental habitat and situation where sufficient moisture and organic
matter are present, however intermittently. Certain
kinds of habitats, such as phytotelmata, have received
a good deal of attention from ecologists, but their
copepod fauna has not been reviewed recently. Other
kinds, particularly habitats modified or constructed by
humans, or extreme habitats such as thermal waters,
have been barely treated in the literature pertaining to copepods. This article briefly discusses some
of the early history of investigations in the betterstudied environments such as ephemeral waters, and
discusses the kinds of information gained from studies
in semi-aquatic natural or artificial habitats, referring
to reviews where possible.
The fauna of acidic waters is normally investigated
by standard limnological methods, and is not treated
in detail here. Species of several genera of cyclop-
oids (e.g. Acanthocyclops, Diacyclops, Paracyclops),
and harpacticoids (Bryocamptus, Moraria) have been
found in waters of pH 4 or below (Rylov, 1948;
Lewis, 1974; Fryer, 1980, 1993). The physiological
adaptations that enable some species but not others
to thrive in acidic waters have been almost entirely
uninvestigated.
Historical process and development of methods
Following the first observation of free-living freshwater copepods, apparently by Blankaart (1688), investigations proceeded sporadically until the era of
the great naturalists and collector-explorers in the
late 19th century. Although most early investigations focused on plankton, Pratz (1866, cited in
Schmeil, 1892) described several cyclopoids from
wells in Munich, Germany. Brady (1868) provided
the first report of copepods from a deep subterranean, human-altered habitat, a coal mine in England.
Studies of copepods in caves began with the collections of Vosseler (1886) near Württemberg, Germany.
Scourfield (1903) provided the earliest report of copepods in phytotelmata, in bromeliads and a pitcher
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plant in botanical gardens in London. The potential for
copepods not only to invade man-made habitats but
also to survive in them successfully was apparently
first recognized by Kraepelin (1886), who collected
cyclopoids and calanoids from the city water system
of Hamburg, Germany.
In the initial investigations of subterranean waters, natural caves and springs, or relatively large
permanent artificial deep holes such as dug and driven
wells were sampled. The Karaman-Chappuis method
(Chappuis, 1942, 1956) of simply digging holes in
sand and gravel banks of streams allowed investigators to reach the top several dm of sediment, in
any locality desired. The Bou-Rouch pump (Bou &
Rouch, 1967), a simplification of an apparatus developed by Delamare Deboutteville (1954) for use
in marine sandy beaches, yielded semi-quantitative
water samples from permeable sediments 1–2 m or
more deep, and became the standard method for
sampling interstitial and phreatic fauna. A modification of the Bou-Rouch pump to avoid damaging the
specimens, by capturing them in a container rather
than passing them through the pump, was advocated
by Vigna Taglianti et al. (1969). The double-packer
sampler, developed by Danielopol & Niederreiter
(1987), provides simultaneous samples of the fauna
and the water. Perhaps the most exact quantitative
method is the freeze-coring method of Bretschko &
Klemens (1986), in which the meiofauna is first paralyzed by an electric field, and then the core is frozen
with liquid nitrogen. What may be the largest semiportable well-drilling system was used by Stanford
& Ward (1988), to reach depths of 30 m in alluvial riverine sediments of the Flathead River valley,
Montana, U.S.A. Deeper sediments are more customarily reached by sampling from permanent driven
wells.
Habitats
Ephemeral waterbodies
The faunas of vernal pools were reviewed by Wiggins
et al. (1980), and those of temporary lentic waters,
particularly in semi-arid lands, by W. D. Williams
(1985). Adaptive strategies, particularly diapause, of
crustaceans for surviving dry periods have been the
subject of many studies, and especially the literature
on diapause in copepods has been reviewed recently
by Dahms (1996), Hairston & Cáceres (1996) and
Williams-Howze (1997), among others.
The faunas of ephemeral streams have received
much less attention. D. D. Williams & Hynes (1976)
found that Attheyella nordenskioldii and Acanthocyclops vernalis were abundant in the hyporheic zone
of a temporary stream in southern Canada, and both
species showed habitat-specific adaptations, such as
timed reproduction and summer resting stages. Otherwise, copepods (and the meiofaunal community in
general) of ephemeral streams seem to have been
almost unstudied. Rouch (1992) described two species of harpacticoids of a previously unknown genus,
Psammonitocrella, from an ephemeral desert stream
in Arizona, U.S.A.
Rock hollows and rock or cliff faces
Water collected in natural hollows such as solution
holes on bare rocks may come to hold copepods. These
are usually common species such as A. vernalis and
Eucyclops agilis in England (Scourfield, 1939), and
Cryptocyclops bicolor linjanticus (Kiefer, 1928) and
Mesocyclops leuckarti (probably = M. aspericornis) in
Fiji (Laird, 1956). However, in Western Australia, the
calanoid Boeckella opaqua Fairbridge, 1945, is found
only in water-filled holes (‘gnammas’) on granite outcrops (Bayly, 1979, 1992). Allocyclops ritae is known
only from a small pool on granitic rock in the Ivory
Coast (Dumont & Lamoot, 1978).
Subterranean habitats
The copepod fauna of groundwater-related and cave
habitats has been extensively reviewed (e.g. Graeter,
1910; Chappuis, 1927; Bowman, 1986; LescherMoutoué, 1986; Rouch, 1986, 1994). Galassi (2000)
traced the changing concepts of subterranean habitats,
from the early idea that they are unusual or challenging
situations, to the growing realization that subterranean
habitats can support a diverse and successful copepod fauna. Indeed, certain groups such as members
of the cyclopoid genus Diacyclops have been more
successful in subterranean than in epigean situations
(Stoch, 1995, 2000), although the reasons for their
success remain mysterious. Two less-investigated subterranean habitats are discussed here: the interstitial,
including the stream-hyporheos; and the ‘pholeteros’
or assemblage in crayfish and crab burrows.
The diverse copepod community inhabiting the interstices among the sand grains of marine beaches, as
well as in loose flocculent mud, was discovered by
Wilson (1935). He pointed out the particular morphological adaptations, including reduction and uniformity in body size, vermiform shape, increased body
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flexibility, development of sense organs, shortening
and strengthening of swimming legs, shape and carriage of the egg sacs, reduction in egg number with
increase in egg size, and behavior, seen in interstitial
copepods.
Following the pioneering studies of the microscopic fauna of sandy beaches in European lakes by
Sassuchin et al. (1927) and others, copepods in the
psammon of freshwater lake beaches were intensively
investigated by Pennak (1939a,b, 1940), who also reviewed the early history of investigations of the psammon (Pennak, 1968). Chappuis (1946) called attention
to the special groundwater habitat associated with the
alluvial sand, gravel and ‘cailleurs’ sediments below
and beside watercourses. This habitat, now known as
the hyporheic zone, was apparently first investigated
by Leruth (1938), in the groundwater of gravel deposits associated with the Meuse River in Belgium. The
development of the Bou-Rouch pump allowed investigators to reach the deeper strata of streambeds and
other permeable phreatic sediments.
The concept of the hyporheic zone as an ecotonal
habitat was discussed by Rouch et al. (1997). Rouch
& Danielopol (1997) argued that hypogean habitats
including the sand-interstitial, hyporheic, phreatic,
karstic and hole and cave faunas are much richer in
species than is commonly estimated, although longterm studies are necessary to estimate faunal diversity
adequately. Certainly, in the past six decades, rich
hypogean-interstitial copepod faunas have been discovered wherever these habitats have been investigated (e.g. Lescher-Moutoué, 1986; Rouch, 1986).
Crayfish tunnels may serve as refuges for
many species of benthic copepods, including rather
eurytopic, widely distributed ones such as Attheyella
dentata (Poggenpol, 1874), Attheyella trispinosa
(Brady, 1880), Halectinosoma abrau (Krichagin,
1877) and Paracyclops affinis in burrows of Astacus
fluviatilis in Europe (Chappuis, 1926; Kiefer, 1927b;
Jakubisiak, 1939). In Ontario, Canada, burrows of
Cambarus fodiens along a temporary stream harbored
large populations of A. vernalis and A. nordenskioldii
(D. D. Williams et al., 1974; D. D. Williams & Hynes,
1976). In Victoria, Australia, Diacyclops cryonastes
Morton, 1985, was found in burrow water of Engaeus
sp., and in Tasmania, Acanthocyclops sp. occurred in
burrows of Parastacoides tasmanicus (Lake & Newcombe, 1975; Lake, 1977). Even normally planktonic
species such as the large calanoid Osphranticum labronectum Forbes, 1882, have been found in water
in the burrows of Cambarus diogenes in Missouri,
U.S.A., during a dry spell (Creaser, 1931). The assemblage of animals living in the water in burrows
of land crayfish was termed the ‘pholeteros’ by Lake
(1977).
Several species of harpacticoids (Attheyella crassa,
Attheyella northumbrica (Brady, 1880) (= dentata),
Attheyella trispinosa, Bryocamptus minutus, Canthocamptus staphylinus (Jurine, 1820) and Nitocrella
hibernica (Brady, 1880)) have been found on crayfish gills, but may be only accidentals (Chappuis,
1926; Gurney, 1930), and may also use the burrows
as refuges.
This possible use of refuges may have led to
the development of various degrees of commensalism between certain harpacticoid species and crayfish.
Nitocrella divaricata (Chappuis, 1923) is found only
on the carapace or in the gills of A. fluviatilis, Astacus
astacus, Astacus leptodactylus and Austropotamobius
torrentium in Europe, and is either a commensal or
an obligate associate (Chappuis, 1923, 1926; Kiefer,
1937; Jakubisiak, 1939; Straskraba, 1956; Sterba,
1964; Boshko, 1976; Defaye, 1996; Subchev & Stanimirova, 1998). Two North American harpacticoids,
Attheyella pilosa Chappuis, 1929, and Attheyella carolinensis Chappuis, 1932, have an intermediate habit:
they occur free-living, but are more frequently found
on the bodies of several species of Cambarus and
Orconectes rusticus (Prins, 1964; Bowman et al.,
1968).
In the tropics, burrows of land crabs (Cardisoma
carnifex) containing fresh or slightly brackish water
regularly harbor Mesocyclops aspericornis and Paracyclops fimbriatus on the volcanic islands of French
Polynesia (Rivière & Thirel, 1981; Rivière et al.,
1987; Rivière, Klein, Duval et al., 1998). Laird
(1956) collected Thermocyclops operculifer Kiefer,
1930, in crab holes on Tarawa (Gilbert Islands). Yeatman (1983) reported several copepod species from
crab holes:Darcythompsonia inopinata Smirnov, 1934
(Western Samoa, Fiji), Ectocyclops phaleratus (Fiji),
Halicyclops septentrionalis Kiefer, 1935 (Fiji), Halicyclops thermophilus Kiefer, 1929, s. str. (Tonga,
Western Samoa, Fiji), H. thermophilus spinifer Kiefer,
1935 (Tonga), Mesocyclops leuckarti (possibly =
M. aspericornis) (Tonga, Fiji), Microcyclops microsetosus Yeatman, 1983 (Fiji), Nitokra lacustris
pacifica Yeatman, 1983 (Western Samoa, Tonga), Nitokra pseudospinipes Yeatman, 1983 (Tonga, Fiji),
Schizopera tobae Chappuis, 1931 (Fiji), and Tisbella
pulchella (Wilson, 1932) (Fiji). Mogi et al. (1984) reported that Thermocyclops sp. and Mesocyclops leuck-
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arti (the latter possibly misidentified) were common
in crab holes in the Ryukyus. Laird (1988) collected
Eucyclops serrulatus and M. aspericornis from freshwater, and H. thermophilus s. str. from brackish-water
crab holes in Western Samoa. Hobbs & Villalobos
(1958) reported the presence of harpacticoid copepods
on the exoskeleton of the freshwater crab Pseudothelphusa lamellifrons in Mexico. The possibility that use
of crab holes as refuges may have facilitated the development of copepod commensals remains a subject for
future investigations.
Phytotelmata
In humid climates, the recesses of plant structures
often contain enough water to support small aquatic
invertebrates, of which some species seem to have a
predilection for, and others may be specifically adapted for this habitat. The observed behavior of certain
species implies that they may climb actively into the
plants. These copepods can live in a film of water
and tend to climb the walls of glass containers, out of
the water, such as Bryocamptus pygmaeus (observed
by Gurney, 1932) and P. fimbriatus, P. affinis and E.
phaleratus (Schmeil, 1892; Scourfield, 1894).
In the tropics, the water pools in bromeliads have
been much investigated, following the discovery and
characterization of this habitat by the celebrated naturalist Fritz Müller, working in Brazil (Müller, 1879).
Frank (1980) and Janetzky (1997) listed a number of
copepod species known from bromeliads; Table 1A
extends their lists.
Pitcher plants in temperate acid bogs are also
convenient collecting sites for copepods (Table 1B).
The tendency of certain species such as Bryocamptus minutus, E. phaleratus and P. affinis to climb
out of water onto vertical, though still moist surfaces
(Scourfield, 1894; Graham, 1907) may account for
the fact that some of these are common in pitcher
plants (Hamilton et al., 2000). Unidentified cyclopids
occurred in about 40% of the pitcher plants at a site
in northern Florida, U.S.A. (Harvey & Miller, 1996).
Also in Florida, cyclopoid copepods showed an aggregated distribution in plants, and their occurrence
was strongly correlated with biotic factors (Harvey &
Miller, 1993).
Treeholes (Table 1C) are probably the best studied type of phytotelm in temperate climates. The
European beech (Fagus silvatica) is especially prone
to develop holes at the junction of the trunk and its
elevated limbs. The practice of pollarding trees in
Europe has also led to the creation of a large number
of treeholes (Scourfield, 1915). Apparently the first
person to investigate treeholes was Scourfield (1915),
who described a new species of Moraria. However,
it is unusual for investigations of treeholes to include
quantitative estimates of non-insect invertebrates. In
an extensive study in Germany, Rohnert (1951) reported only one species of copepod, Moraria sp., and
considered that copepods were accidental arrivals.
Nearly any plant recess may form a receptacle for
water and come to harbor copepods (Table 1D). However, it is clear from the list of recorded species that
certain groups favor phytotelmata: Attheyella (subgenus Canthosella) and some Elaphoidella in neotropical bromeliads, and certain Ectocyclops, Paracyclops
and Tropocyclops, plus the ubiquitous soil-dwellers
Epactophanes richardi and Phyllognathopus viguieri
in both the tropics and temperate zones. Species of
Canthosella are predominantly found in bromeliads,
and Tropocyclops jamaicensis invades terrestrial bromeliads quickly and may be an obligate phytotelm
dweller (Reid & Janetzky, 1996).
Mosses
The records of harpacticoids and small cyclopoids
from aquatic and terrestrial mosses in humid climates
are so numerous as to be almost impossible to review. Both aquatic mosses (Sphagnum, Hypnum) and
terrestrial mosses and liverworts in more humid situations harbor their own, sometimes distinctive, copepod faunas. Masses of aquatic mosses when squeezed
yield small species of Acanthocyclops, Diacyclops
and other cyclopid genera. Many species of Bryocyclops and Muscocyclops live in moss (e.g. Scourfield,
1939). Some 12 species of harpacticoids, mostly canthocamptids, were collected from terrestrial moss in
open or forested locations in New Zealand (Lewis,
1984). Seeps on rock outcrops where moss and algae
grow may also hold copepods, usually common species such as B. pygmaeus, but sometimes rarer ones
such as Speocyclops demetiensis (Scourfield, 1932)
(Gurney, 1932; Scourfield, 1932, 1939) and Stolonicyclops heggiensis Reid & Spooner, 1998. Wet
moss on rock faces was the home of several canthocamptids in Tasmania (Hamond, 1988).
Leaf litter
The slowly decaying mats of leaf litter in humid temperate forests create a moist habitat favourable to several species of cyclopoids and harpacticoids, mainly
canthocamptids (Table 2). Among the earliest reports
were those by Gurney (1932) and Remy (1932). The
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Table 1. Species of copepods reported from phytotelmata. Species are listed under the current name of the taxon, as far
as possible; synonymies of most species were listed by Dussart & Defaye (1985, 1990) for Cyclopoida and Harpacticoida,
respectively
Kind of phytotelmata/ Species
A. Bromeliads
Harpacticoida
Attheyella aliena Noodt, 1956
Attheyella jureiae Por & Hadel, 1986
Attheyella mervini Janetzky, Martı́nez Arbizu & Reid,
1996
Attheyella striblingi (Reid, 1990)
Attheyella vera Por & Hadel, 1986
Canthocamptus sp.1
Elaphoidella bidens (Schmeil, 1894)
Elaphoidella bromeliaecola (Chappuis, 1928)
Elaphoidella malayica (Chappuis, 1928)
Elaphoidella sewelli (Chappuis, 1928)
E. sewelli
Elaphoidella sp.
Epactophanes richardi Mrázek, 1893
E. richardi
Phyllognathopus viguieri (Maupas, 1892)
P. viguieri
P. viguieri
P. viguieri
P. viguieri
P. viguieri
P. viguieri menzeli (Chappuis, 1928)
canthocamptids
harpacticoids
Cyclopoida
Bryocyclops anninae (Menzel, 1926)
Location
Reference
Germany (greenhouse)
Brazil
Jamaica
Noodt (1956)
Por & Hadel (1986)
Janetzky et al. (1996),
Janetzky (1997)
Costa Rica
Reid (1990)
Brazil
Por & Hadel (1986)
Java
Menzel (1922, 1924, 1925b)
Puerto Rico
Reid (1993b)
Java
Chappuis (1928, 1931),
Thienemann (1934)
Java
Chappuis (1931), Thienemann
(1934)
Jamaica
Laessle (1961)
Puerto Rico
Maguire (1970)
Puerto Rico
Reid (1993b)
England (botanical garden)
Gurney (1932)
Jamaica
Janetzky et al. (1996)
Java
Menzel (1922, 1924, 1925b, 1926b)
England (botanical gardens, Gurney (1932), Lowndes
purchased pineapples)
1931), Scourfield (1903,
1906, 1939)
Brazil
Chappuis (1936)
Poland
Jakubisiak (1929)
Puerto Rico
Maguire (1970)
Jamaica
Janetzky et al. (1996)
Java
Chappuis (1928, 1931),
Thienemann (1934)
Australia (botanical gardens) Hamond (1988)
Brazil
Lopez et al. (1998)
Java
Bryocyclops bogoriensis (Menzel, 1926)
Java
Bryocyclops caroli Bjornberg, 1985
Cyclops sp.2
Diacyclops bisetosus (Rehberg, 1880)
Ectocyclops bromelicola Kiefer, 1935
Ectocyclops phaleratus (Koch, 1838)
E. phaleratus
Puerto Rico
Argentina
England
Brazil
Costa Rica
Jamaica
Ectocyclops strenzkei Herbst, 1959
E. strenzkei
Ectocyclops sp.
Fimbricyclops jimhensoni Reid, 1993b
Fimbricyclops sp.
Muscocyclops operculatus (Chappuis, 1917)
Brazil
Germany (greenhouse)
Brazil
Puerto Rico
Puerto Rico
Brazil
Menzel (1926a), Thienemann
(1934)
Menzel (1926a), Thienemann
(1934)
Maguire (1970), Reid (1999)
Torales et al. (1972)
Gurney (1933)
Kiefer (1935)
Picado (1913)
Laessle (1961), Reid &
Janetzky (1996)
Herbst (1959)
Herbst (1959)
Hadel & Carvalho (1988)
Reid (1993b)
Reid (1999)
Kiefer (1935), Rocha &
Bjornberg (1987)
Continued on p. 206
206
Table 1. contd.
Kind of phytotelmata/ Species
Location
Reference
Paracyclops bromeliacola Karaytug & Boxshall, 1998
Paracyclops fimbriatus (Fischer, 1853)
P. fimbriatus f. bromeliarum Herbst, 1959
Paracyclops punctatus Karaytug & Boxshall, 1998
Paracyclops reidae Karaytug & Boxshall, 1998
Paracyclops sp.
Tropocyclop jamaicensis Reid & Janetzky, 1996
Brazil
Brazil
Brazil
Brazil
Trinidad
Brazil
Jamaica
Tropocyclops prasinus (Fischer, 1860)
Tropocyclops schubarti Kiefer, 1935
Tropocyclops sp.
Cyclopoids
copepods
Puerto Rico
Brazil
Brazil
Brazil
Belgium?
Karaytug & Boxshall (1998a)
Ferreira (1985)
Herbst (1959)
Karaytug & Boxshall (1998a)
Karaytug & Boxshall (1998a)
Hadel & Carvalho (1988)
Reid & Janetzky (1996),
Janetzky (1997), Laessle
(1961)
Reid (1993b)
Kiefer (1935), Herbst (1959)
Hadel & Carvalho (1988)
Lopez et al. (1998)
Oye (1923); cited by Frank
(1980)
B. Pitcher plants (Sarracenia spp.)
Harpacticoida
Bryocamptus hiatus (Willey, 1925)
Phyllognathopus viguieri
Cyclopoida
Acanthocyclops parasensitivus Reid, 1998
Acanthocyclops venustoides pilosus Kiefer, 1934
Diacyclops harryi Reid, 1992
Diacyclops languidus (G. O. Sars, 1863)
Paracyclops canadensis (Willey, 1934)
P. canadensis
Cyclopidae
Canada
England
Laird (1988)
Scourfield (1903, 1939)
U.S.A.
Canada, U.S.A.
U.S.A.
U.S.A.
Canada
U.S.A.
U.S.A.
‘Entomostraca’
U.S.A.
Reid (1998)
Laird (1988)
Reid (1992)
Ishida (1992)
Willey (1934)
Hamilton et al. (2000)
Miller et al. (1994), Harvey
& Miller (1996)
Hegner (1926)
C. Tree holes or stump holes
Harpacticoida
Bryocamptus minutus (Claus, 1863)
Bryocamptus pygmaeus (Sars, 1863)
Epactophanes richardi
Moraria arboricola Scourfield, 1915
England, Ireland
England
Germany
England
Moraria varica (Graeter, 1911)
England
Moraria sp.
Elaphoidella taroi Chappuis, 1955
Phyllognathopus viguieri
Tachidius discipes Giesbrecht, 1882
Cyclopoida
Apocyclops sp.
Australocyclops australis Morton, 1985
Bryocyclops bogoriensis
Bryocyclops fidjiensis Lindberg, 1954
Cryptocyclops linjanticus (Kiefer, 1928)
Diacyclops bisetosus
Germany
Fiji
Tonga, Western Samoa, Fiji
England
Fiji
New South Wales, Australia
Fiji (Ivi, Inocarpus edulis)
Tonga, Western Samoa, Fiji
Fiji
England
Gurney (1932)
Gurney (1920, 1932)
Kiefer (1924)
Gurney (1920, 1932),
Scourfield (1915, 1939)
Gurney (1920, 1932),
Scourfield (1939)
Rohnert (1951)
Yeatman (1983)
Yeatman (1983)
Gurney (1920)
Yeatman (1983)
Morton (1985)
Yeatman (1983)
Yeatman (1983)
Yeatman (1983)
Gurney (1933), Hollowday
(1949)
Continued on p. 207
207
Table 1. contd.
Kind of phytotelmata/ Species
Location
Reference
Diacyclops spp.
Halicyclops thermophilus Kiefer, 1929
Mesocyclops aspericornis (Daday, 1906)
Stoch (2000)
Yeatman (1983)
Rivière & Thirel (1981);
Rivière et al. (1987)
Paracyclops fimbriatus
Paracyclops poppei (Rehberg, 1880)
Italy
Tonga, Western Samoa, Fiji
French Polynesia: holes in
buttresses of Inocarpus
fagifer
Fiji
U.S.A.
Cyclopidae
copepods
Nukunono, Tokelau Islands
Brunei (dipterocarp forest)
D. Other phytotelmata
Harpacticoida
Attheyella gessneri Chappuis, 1956
Attheyella inopinata Chappuis, 1931
Attheyella ruttneri Chappuis, 1931
Bryocamptus pygmaeus
Canthocamptus sp.
Elaphoidella bromeliaecola
Elaphoidella cornuta Chappuis, 1931
Elaphoidella elegans Chappuis, 1931
Elaphoidella taroi
E. taroi
E. taroi
Elaphoidella thienemanni Chappuis, 1931
Elaphoidella sp.
Epactophanes richardi
E. richardi
E. richardi
E. richardi menzeli Chappuis, 1931
Venezuela: leaf axils of
Heliconia bihai (Musaceae)
Sumatra: leaf axils of
Cyrtandra glabra
(Gesneriaceae)
Bali, Java: wet leaves of
Elatostema macrophyllum
(Urticaceae) and leaf axils of
Colocasia indica (Araceae)
Germany: leaf axils of Scirpus
silvaticus (Cyperaceae)
Java: leaf axils of C. glabra
Java, Sumatra: leaf axils of
Colocasia sp., C. glabra and
Pandanus sp. (Pandanaceae)
Sumatra: leaf axils of C.
glabra
Java: leaf axils of C. indica
Fiji: leaf axils of taro
(Colocasia esculenta) and
Cordyline terminalis
(Agavaceae)
Fiji, Western Samoa: fallen
coconut shells, leaf axils of
taro
Tahiti: fallen coconut shells
Sumatra: leaf axils of C.
glabra
Fiji: leaf axils of Colocasia
sp.and C. terminalis
Java: leaf-cups of Nepenthes
sp. (Nepenthaceae)
Sumatra: leaf axils of C.
glabra
Java, Sumatra: inflorescences
of Zingiber macradenia
(Zingiberaceae), leaf axils of
Colocasia sp. and C. glabra
Sumatra: leaf-cups of
Yeatman (1983)
Reid & Marten (1994),
Karaytug & Boxshall (1998a)
Laird (1956)
Kitching & Orr (1996)
Chappuis (1956)
Chappuis (1931), Thienemann
(1934)
Chappuis (1931), Thienemann
(1934)
Strenzke (1951)
Menzel (1924)
Chappuis (1931), Thienemann
(1934)
Chappuis (1931), Thienemann
(1934)
Chappuis (1931), Thienemann
(1934)
Chappuis (1955)
Yeatman (1983)
Rivière, Klein, Thirel & Chebret (1998)
Chappuis (1931), Thienemann
(1934)
Laird (1956)
Menzel (1921)
Chappuis (1931), Thienemann
(1934), Lang (1935)
Chappuis (1931)
Thienemann (1932, 1934)
Continued on p. 208
208
Table 1. contd.
Kind of phytotelmata/ Species
Parastenocaris incerta Chappious, 1931
Parastenocaris staheli
Phyllognathopus viguieri
P. viguieri
P. viguieri
P. viguieri
P. viguieri
P. viguieri
P. viguieri
P. viguieri
P. viguieri
P. viguieri
P. viguieri menzeli
P. viguieri menzeli
Harpacticoida
harpacticoid
Cyclopoida
Bryocyclops anninae
B. anninae
Bryocyclops bogoriensis
Bryocyclops chappuisi Kiefer, 1928
Bryocyclops fidjensis
B. fidjiensis
Location
Nepenthes ampullaria; also
C. glabra, Zingiber sp.
Sumatra: leaf-cups of N.
ampullaria in moss
Surinam: moss in old leaf
axils of Livistona (Palmae)
Algeria: debris of a decaying
banana tree
Java: leaf axils of C. glabra
Poland: leaf axils of Musa
encete (Musaceae)
Sumatra: leaf axils of
Colocasia sp.
Sumatra: leaf-cups of N.
ampullaria
Java, Sumatra: Colocasia
indica, Cyrtandra sp.,
Zingiber sp.
England: leaf-cups of Crinum
sp. (Amaryllidaceae)
Fiji: leaf axils of C. terminalis
and Colocasia sp.
Madagascar: leaf axils of
Typhonodorum sp. (Araceae)
Tonga, Western Samoa, Fiji:
taro leaf axils, bamboo
Java, Sumatra: leaf axils of
Colocasia spp., C. glabra,
cups of Nepenthes sp., and
inflorescences of Z.
macradenia
Guam: leaf axils of Pandanus
Sulawesi: leaf axils of taro
Costa Rica: bracts of
Heliconia imbricata
(Musaceae)
New Hebrides: empty
coconut husks
Guam: leaf axils of Pandanus
sp.
Sumatra: leaf axils of
Colocasia sp.; Java: leaf axils
of Pandanus sp.
Java: leaf axils of Crinum
hybridum
Fiji: leaf axils of C.
terminalis, Freycinetia milnei
(Pandanaceae), taro
(Colocasia antiquorum)
Fiji, Tonga, Western Samoa,
Reference
Chappuis (1931); Thienemann
Chappuis (1932)
Menzel, 1916
Chappuis (1916)
Menzel (1924, 1925b, 1926b)
Jakubisiak (1929)
Chappuis (1931)
Thienemann (1932)
Thienemann (1934)
Scourfield (1939)
Laird (1956)
Dussart (1982)
Yeatman (1983)
Chappuis (1931), Thienemann
(1934)
Watkins & Belk (1975)
Mogi & Sembel (1996)
Naeem (1988)
Lowndes (1928a)
Watkins & Belk (1975)
Kiefer (1933), Thienemann
(1934)
Kiefer (1933), Thienemann
(1934)
Lindberg (1955)
Yeatman (1983)
Continued on p. 209
209
Table 1. contd.
Kind of phytotelmata/ Species
Bryocyclops muscicola (Menzel, 1926)
Bryocyclops sp.
Ectocyclops rubescens (as E. medius)
Tropocyclops schubarti dispar Herbst, 1962
Cyclopidae
cyclopoids
Cyclopoida
copepods
copepods
Location
Reference
Hawaii: bamboo, leaf axils
of taro and Pandanus sp.
Sumatra: leaf axils of
Pandanus sp.
Fiji, Tonga: leaf axils of
Colocasia sp., C. terminalis,
and Freycinetia milnei; and
Pandanus sp., respectively
Java: leaf axils of C. indica,
leaves of nettles
Brazil: shells of fallen Brazil
nuts, Bertholletia excelsa
(Lecythidaceae)
Tarawa, Gilbert Islands:
coconut shell
Singapore: tree roots
Sulawesi: leaf axils of taroMogi & Sembel (1996)
Puerto Rico: water in fallen
leaves
Singapore: cups of N.
ampullaria
Kiefer (1933), Thienemann
(1934)
Laird (1956)
Kiefer (1933), Thienemann
(1934)
Herbst (1962)
Laird (1956)
Laird (1988)
Maguire (1971)
Ghosh (1928)
1 Should be considered an unidentified canthocamptid.
2 Since Cyclops is not a tropical genus, this record should be considered an unidentified cyclopoid.
scanty literature on copepods in leaf litter was last reviewed for harpacticoids by Menzel (1946), and more
generally discussed by Fiers & Ghenne (2000). Most
studies have been done in Europe, where extensive
areas are covered by beech (Fagus sylvatica) forests,
and leaf litter from that and other trees now forms,
or historically has formed vast carpets. Beech leaves
tend to pack tightly together in a dense layer that retains water, and the animals live mainly in the deeper,
more humid layers, directly on top of the forest floor
(Nielsen, 1966; Schaeffer, 1991). In Denmark, populations of the harpacticoids E. richardi and Maraenobiotus vejdovskyi tenuispina reached substantial densities (40 and 1.5 ind/g of leaves respectively) (Nielsen,
1966). Fiers & Ghenne (2000), treating a modest number of litter samples which were sampled and extracted
using techniques more suitable for nematodes, listed
9 cyclopoid and 13 harpacticoid species, 9 of these
new records for Belgium, and a significant addition
to the list of 71 species previously known from that
country. The leaf-carpet may be the primary habitat for
certain harpacticoid species, such as Maraenobiotus
vejdovskyi var. truncatus (Scourfield, 1939) and some
members of Moraria, as implied by the Yorkshire
records of Fryer (1993). Michailova-Neikova (1973)
found that 8 of 9 harpacticoid species living in wet
moss beside waterbodies on a mountain in Bulgaria
also appeared in the leaf litter.
Outside Europe, leaf litter has been much less
investigated (Table 2). Dumont & Maas (1988) described five new species of harpacticoids from leaf
litter in Nepal, at altitudes from 1900 to 3900 m.
Moist rotting wood may also harbor copepods.
Elaphoidella cuspidata Chappuis, 1941, was found
by Chappuis (1954) in rotted wood in India. Phyllognathopus camptoides was described by Boz̆ić (1965)
from moist dead wood collected from a forest in
Gabon.
Moist soils
Reid (1986) reviewed qualitative and quantitative
studies reporting copepods from moist soils. In their
study in Belgium, Fiers & Ghenne (2000) also discovered a surprising number of species in soil samples,
particularly in fallow soils.
The ecological role of soil copepods is not understood. Birch & Clark (1953) classified Epactophanes
sp. among the bacterial feeding organisms in a study of
210
Table 2. Copepods recorded from leaf litter. Species are listed under the current name of the taxon, as far as possible; synonymies
of most species were listed by Dussart & Defaye (1985, 1990) for Cyclopoida and Harpacticoida, respectively
Species
Location
Source
Harpacticoida
Attheyella crassa (Sars, 1863)
Attheyella ilami Dumont & Maas, 1988
Attheyella wierzejskii (Mrázek, 1893)
Bryocamptus hoferi (Douwe, 1907)
Bryocamptus minutus
Bryocamptus pygmaeus
Bulgaria
Nepal
Bulgaria
Bulgaria
Bulgaria
England
B. pygmaeus
B. pygmaeus
Bryocamptus stouti Harding, 1958
Bryocamptus weberi (Kessler, 1914)
Bryocamptus zschokkei (Schmeil, 1893)
B. zschokkei tatrensis Minkiewicz, 1916
Cantocamptus clavifurcatus Hamond, 1988
Canthocamptus dedeckkeri Hamond,1988
Canthocamptus dumonti Hamond, 1988
Canthocamptus globulisetosus Hamond, 1988
Canthocamptus howardorum Hamond, 1988
Canthocamptus lacinulatus Hamond, 1988
Canthocamptus longifurca Hamond, 1988
Canthocamptus mammillifurca Hamond, 1988
Canthocamptus mortoni Hamond, 1988
Canthocamptus tasmaniae Hamond, 1988
Canthocamptus timmsi Hamond, 1988
Echinocamptus hypophyllus Defaye & Heymer, 1996
Elaphoidella jochenmartensi Dumont & Maas, 1988
Elaphoidella propedamasi Defaye & Heymer, 1996
Elaphoidella pseudocornuta Dumont & Maas, 1988
Epactophanes muscicolus (Richters, 1901)
Epactophanes richardi
France
Bulgaria
New Zealand
England
Corsica
Bulgaria
Tasmania
Victoria, Australia
Victoria, Australia
Victoria, Australia
Tasmania
Victoria, Australia
Victoria, Australia
Victoria, Australia
Tasmania
Tasmania
Tasmania
Congo (Zaire)
Nepal
Congo (Zaire)
Nepal
England
England, Ireland
E. richardi
E. richardi
E. richardi
E. richardi
E. richardi
Fibulacamptus gracilior Hamond, 1988
Fibulacamptus tasmanicus Hamond, 1988
Fibulacamptus victorianus Hamond, 1988
Maraenobiotus vejdovskyi anglicus Gurney, 1932
M. vejdovskyi tenuispina Roy, 1924
M. vejdovskyi tenuispina
M. vejdovskyi truncatus Gurney, 1932
M. vejdovskyi truncatus
Moraria arboricola Scourfield, 1915
Germany
Corsica
Denmark
Hawaii
Congo (Zaire)
Victoria, Australia
Tasmania
Victoria, Australia
England
Denmark
France
England
France
England
Moraria frondicola Klie, 1943
Moraria ilami Dumont & Maas, 1988
Moraria poppei (Mrázek, 1893)
Corsica
Nepal
Bulgaria
Michailova-Neikova (1973)
Dumont & Maas (1988)
Michailova-Neikova (1973)
Michailova-Neikova (1973)
Michailova-Neikova (1973)
Scourfield (1940), Fryer
(1993)
Klie (1943)
Michailova-Neikova (1973)
Harding (1958)
Scourfield (1940)
Klie (1943)
Michailova-Neikova (1973)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Defaye & Heymer (1996)
Dumont & Maas (1988)
Defaye & Heymer (1996)
Dumont & Maas (1988)
Scourfield (1940)
Gurney (1932), Scourfield
(1940)
Precht (1936)
Klie (1943)
Nielsen (1966)
Evenhuis & Preston (1995)
Defaye & Heymer (1996)
Hamond (1988)
Hamond (1988)
Hamond (1988)
Scourfield (1940)
Nielsen (1966)
Remy (1932)
Scourfield (1940)
Klie (1943)
Scourfield (1940), Fryer
(1993)
Klie (1943)
Dumont & Maas (1988)
Michailova-Neikova (1973)
Continued on p. 211
211
Table 2. contd.
Species
Location
Source
Moraria terrula Kikuchi, 1991
Moraria tsukubaensis Kikuchi, 1991
Moraria valkanovi Michailova-Neikova, 1973
Moraria varica
Japan
Japan
Bulgaria
England, Ireland
M. varica
M. varica
Parbatocamptus jochenmartensi Dumont & Maas, 1988
Phyllognathopus cf. camptoides Boz̆ić, 1965
Phyllognathopus viguieri
P. viguieri
P. viguieri
unidentified harpacticoid
Cyclopoida
Allocyclops silvaticus Rocha & Bjornberg, 1988
Bryocyclops anninae
Bryocyclops phyllopus Kiefer, 1935
Goniocyclops sylvestris Harding, 1958
Metacyclops hirsutus Rocha, 1994
Muscocyclops operculatus
Paracyclops bromeliacola Karaytug & Boxshall, 1998
Paracyclops chiltoni (Thomson, 1882)
France
England
Nepal
Congo (Zaire)
England
Agrihan Is., Marianas
Hawaii
England
Kikuchi (1984, 1991b)
Kikuchi (1984, 1991a)
Michailova-Neikova (1973)
Gurney (1932), Scourfield
(1940)
Boz̆ić (1966)
Fryer (1993)
Dumont & Maas (1988)
Defaye & Heymer (1996)
Scourfield (1940)
Kikuchi (1994b)
Evenhuis & Preston (1995)
Stout (1963)
Brazil
Hawaii
Congo (Zaire)
New Zealand
Brazil
Brazil
Brazil
Brazil
Rocha & Bjornberg (1988)
Evenhuis & Preston (1995)
Defaye & Heymer (1996)
Harding (1958)
Rocha (1994)
Rocha & Bjornberg (1987)
Karaytug & Boxshall (1998a)
Karaytug & Boxshall (1998b)
soil fauna in Australia. However, Epactophanes, like
P. viguieri, may be a predator on nematodes. Copepods are inefficiently sampled by the Tullgren funnel
and other extraction methods used for soil fauna (Fiers
& Ghenne, 2000), and their true numbers in soils are
probably underestimated.
Thermal waters
It is difficult to define ‘thermal’, but 30 ◦ C may
be a reasonable limit above which temperate-zone
copepods do not easily survive. There are relatively
few records of copepods, mostly cyclopoids, from
thermal waters. Menzel (1925a) reported a species of
Halicyclops (subsequently described as Halicyclops
thermophilus by Kiefer, 1929, 1933) from a hot saline spring on Java (37–45 ◦ C, pH 6.7–7.1, salinity
26.7). Many of the temperatures reported by Kiefer
(1933) from waters on Sumatra investigated by the
Sunda-Expedition were over 30 ◦ C: a young cyclopoid
was found in a warm acid spring (35.5 ◦ C, pH 2.68);
other warm springs (29.5–39 ◦ C, pH 7.3–7.6) harbored
Paracyclops eucyclopoides, P. fimbriatus, Mesocyclops aequatorialis, Microcyclops varicans s. str., M.
varicans subaequalis, and Thermocyclops decipiens.
Brehm (1936) mentioned a species of Eucyclops from
‘heißen Quellen’ near Cuzco, Peru; no temperature
or water chemistry data were available. Mesocyclops
‘leuckarti’ was reported by Lindberg (1942) from a
basin of a soap factory in Iran, water temperature
40 ◦ C. In Iceland, a thermal pool (34 ◦ C) harbored
the eurytopic species E. serrulatus and Megacyclops
viridis (Starmühlner, 1969). Dussart (1974) reported Cletocamptus deitersi, Cryptocyclops linjanticus
micrura and Mesocyclops aequatorialis from warm
springs in Ethiopia, without further habitat data. Lewis
(1974) described Paracyclops waiariki from thermal
springs in New Zealand, but at a relatively low temperature of 27 ◦ C (and pH 3.0). Reid (1994) found a
cyclopoid, Microcyclops sp., in a spring-fed thermal
(about 40 ◦ C) pond in Brazil.
Other harpacticoids besides C. deitersi have been
found in thermal waters. Chappuis (1931) reported
Elaphoidella bidens coronata, Schizopera tobae and
Parastenocaris longicaudis from a warm spring (29.5
◦ C) on Sumatra. Por (1964) described Nitokra balnearia from hot mineral springs (31 ◦ C) on the Dead
Sea shore in Israel, where it reached immense numbers. Thermomesochra reducta Itô & Burton, 1980,
212
Table 3. Copepods reported from thermal waters. Species are listed under the current name of the taxon, as far as possible;
synonymies of most species were listed by Dussart & Defaye (1985, 1990) for Cyclopoida and Harpacticoida, respectively
Species
Harpacticoida
Cletocamptus deitersi (Richard, 1897)
Elaphoidella bidens coronata (Sars, 1904)
Nitokra balnearia Por, 1964
Parastenocaris longicaudis Chappuis, 1931
Schizopera tobae Chappuis, 1931
Thermomesochra reducta Itô & Burton, 1980
Cyclopoida
Cryptocyclops linjanticus
Cyclops sp.1
Cyclops sp.1
Eucyclops serrulatus (Fischer, 1851)
E. serrulatus
Temp. (◦ C)
Location: habitat
Reference
29.5
31
29.5
29.5
38–58
Ethiopia: thermal springs
Sumatra: thermal spring
Israel: hot mineral spring
Sumatra: thermal spring
Sumatra: thermal spring
Malaysia: hot spring
Dussart (1974)
Chappuis (1931)
Por (1964)
Chappuis (1931)
Chappuis (1931)
Itô & Burton (1980)
37–46
Ethiopia: springs
India: spring
31–34
Nigeria: spring
Dussart (1974)
Jana (1978); Jana &
Sarkar (1971)
Egborge & Fagade
(1979)
Starmühlner (1969)
Ponyi (1992)
34
28.6–29.7
(surface)
37–45
Iceland: spring
Hungary: springfed lake
Java: saline spring
Hungary: spring-fed lake
Iceland: spring
Ethiopia: springs
Sumatra: springs
Iran: basin
Sumatra: springs
Hungary: thermal spring-fed lake
Starmühlner (1969)
Dussart (1974)
Kiefer (1933)
Lindberg (1942)
Kiefer (1933)
Ponyi (1992)
M. varicans subaequalis (Kiefer, 1928)
Microcyclops sp.
28.6–29.7
(surface)
34
29.5–39
40
29.5–39
28.6–29.7
(surface)
29.5–39
40
Menzel (1925a);
Kiefer (1929, 1933)
Ponyi (1992)
Kiefer (1933)
Reid (1994)
Paracyclops eucyclopoides Kiefer, 1929
Paracyclops fimbriatus
Paracyclops waiariki Lewis, 1974
29.5–39
29.5–39
27
Thermocyclops decipiens Kiefer, 1929
unidentified cyclopoid
29.5–39
35.5
Sumatra: thermal springs
Brazil: spring-fed thermal
pond
Sumatra: thermal springs
Sumatra: thermal springs
New Zealand: thermal
springs
Sumatra: thermal springs
Sumatra: thermal acid
spring
Halicyclops thermophilus
Macrocyclops albidus (Jurine, 1820)
Megacyclops viridis (Jurine, 1820)
Mesocyclops aequatorialis Kiefer, 1929
M. aequatorialis
Mesocyclops leuckarti (Claus, 1857)
Microcyclops varicans (Sars, 1863)s. str.
M. varicans
Kiefer (1933)
Kiefer (1933)
Lewis (1974)
Kiefer (1933)
Kiefer (1933)
1 These records should be considered as unidentified cyclopoids.
from a hot spring in Malaysia, holds the survival
record at temperatures between 38 and 58 ◦ C.
High altitude and high latitude habitats
There has historically been much interest in the
aquatic faunas of high-altitude regions, although of
course most studies have been carried out in lakes.
The non-lacustrine copepod fauna of the Alps is
comparatively well known (e.g. Thienemann, 1950;
Gaviria, 1998). Several surveys in montane regions
outside Europe, mainly by Heinz Löffler, have included samples from non-lacustrine ‘minor’ habitats.
Examples from the Andes include the reports of Harding (1955) and Löffler (1960) from Peru and Bolivia.
Löffler (1965, 1968a) reported on Maraenobiotus and
other copepods from the high mountains of eastern
Africa. In Asia, Löffler has reported on Maraenobi-
213
otus and other harpacticoids from Nepal (1968b) and
Borneo (Mt. Kinabalu; Löffler, 1973).
Although the plankton of lakes at extremely high
altitudes tends to be species-poor (except in the Alps;
see Löffler, 1968c), most of these surveys have indicated the existence of a rich semiterrestrial copepod
fauna in locations where there is some vegetation and
moisture, even in the presence of permafrost. Typical
altitudes reached by copepods are up to about 4100
m in Mexico and Central America (Löffler, 1972) and
the Colombian páramo region (Gaviria, 1989), 3800
m on Mt. Kinabalu, Borneo (Löffler, 1973), 2140 m
in the Great Smoky Mountains, eastern U.S.A. (J. W.
Reid & W. Reeves, unpublished data), and to the glacier line (2300–2600 m) in the Austrian Alps (Gaviria,
1998). Dumont & Maas (1988) described five species
of harpacticoids living in leaf litter in montane forests
of Nepal at moderate to high altitudes (1900–3900 m).
Two planktonic species of calanoids and an unidentified cyclopoid live in lakes in Nepal at altitudes from
4600 to 5460 m (Manca et al., 1994). The record for
altitude is probably held by Glaciella yalensis Kikuchi, 1994a, which inhabits melt (cryconite) pits on the
Yala Glacier in Nepal, at altitudes between 5100 and
5700 m (Kikuchi, 1994a).
Copepods living in extreme north and south latitudes confront similar problems of a short growing
season and lack of organic matter. For instance, saline lakes in Antarctica support a limited assemblage
consisting of three species of harpacticoids and two
of calanoids (Wright & Burton, 1981); and freshwater Antarctic lakes harbor two calanoids, Boeckella
poppei (Mrázek, 1901) and Gladioferens antarcticus
Bayly, 1994, and one species of cyclopoid, Acanthocyclops mirnyi Borutzky & Vinogradov, 1957 (Bayly
& Burton, 1993; Bayly, 1994). The known copepod
fauna of Iceland is similarly species-poor (Starmühlner, 1969).
Students of biogeography have been much interested in the distribution and relationships of invertebrate montane faunas. The isolation of populations
within and between mountain ranges has contributed
to the development of a confusingly rich array of
morphologically distinct populations of certain species. This phenomenon is well studied in the genus
Maraenobiotus (Löffler, 1965, 1968a,b, 1973), and in
several canthocamptid species in Japan (e.g. Ishida,
1991, 1994). Husmann (1975) argued that the edges
of melting glaciers provided large areas of sand and
gravel layers, followed by the moss tundra, which may
have formed a ‘combined system of migration ways’
for interstitial organisms to follow from warmer regions. However, Strayer & Reid (1999) found that previously glaciated sites in the U.S.A. contained fewer
species of interstitial specialist cyclopoid copepods
(although equivalent numbers of generalist species)
than historically unglaciated sites. This suggests that
it was mainly the eurytopic species that were able to
follow these ‘migration ways’.
Unclassifiable natural habitats
Some of the odder locations where copepods have
been found include a sea shell in Fiji, which contained
Halicyclops septentrionalis Kiefer, 1935 (Yeatman,
1983), and ants’ nests (B. H. Dussart, mentioned by
Reid, 1986).
Human-modified or artificial habitats
Holes in the ground created by the multiplicity of human actions may easily be colonized by copepods and
other aquatic animals. Ditches and borrow pits are
common sites to find copepods. For instance, Brady
(1907) and Graham (1907) described several species
of harpacticoids and cyclopoids from railway borrow
pits and gold mine pits in the African Gold Coast.
Münchberg (1956) described the limnological conditions and aquatic faunas of ca. 10-year-old bomb
craters in Germany, which included mainly common, generalist species such as Cyclops cf. strenuus
Fischer, 1851, and Macrocyclops albidus. Some examples of species collected from road ruts include
Canthocamptus australicus (Sars, 1908) in Australia
(Hamond, 1988), Mesocyclops leuckarti (probably =
M. aspericornis) in Guam (Watkins & Belk, 1975),
and Metacyclops minutus (Claus, 1863) and several
other species by Maier (1992, 1998) and Maier et al.
(1998). Even footprints may serve as copepod microhabitats: Brehm (mentioned in Gurney, 1933: 221)
found Diacyclops bicuspidatus (Claus, 1857) in the
small pools formed by his own footprints in spongy
woodland soil. The ecological role of borrow ditches
and pits in providing refuges for the aquatic faunas in
arid regions might fruitfully be examined.
Copepods have invaded a variety of man-made
habitats (Table 4), for example the roof of a tunnel in
a coal mine (Brady, 1868), the filter beds of sewage
treatment plants, and deeply buried agricultural drainage tiles (D. D. Williams, 1976). Copepods have been
noticed in water systems of various sizes and uses,
especially in older municipal systems where much
of the water was drawn from subterranean sources,
with minimal treatment. It is probably no accident
214
Table 4. Copepods reported from man-made or altered habitats. Species are listed under the current name of the taxon,
as far as possible; synonymies of most species were listed by Dussart & Defaye (1985, 1990) for Cyclopoida and
Harpacticoida, respectively
Species
Harpacticoida
Attheyella nordenskioldii (Lilljeborg, 1902)
Bryocamptus aquaeductus Borutsky, 1934
Bryocamptus echinatus (Mrázek, 1893)
Bryocamptus luenensis (Schmeil, 1894)
Bryocamptus minutus
B. minutus
B. minutus
Bryocamptus pygmaeus
Location: habitat
Reference
Canada: agricultural drainage
tiles
Ciscaucasia: city water pipe
Germany: city water pipes
Hungary: city water pipes
England: coal mine
Germany: city water pipes
Romania: city water pipes
Romania: city water pipes
Williams, D. D. (1976)
Bryocamptus typhlops (Mrázek, 1893)
Canthocamptus sp.
Germany: city water pipes
Germany: moist decaying
mine timbers
Chappuisius inopinus Kiefer, 1938
Hungary, Romania: city water
pipes
Elaphoidella dubia Kiefer, 1931
Romania: city water pipes
Elaphoidella elaphoides (Chappuis, 1924)
Romania: city water pipes
Elaphoidella gracilis serrulata Damian & Botosaneanu, Romania: city water pipes
1954
Elaphoidella grandidieri (Guerne & Richard, 1893)
Western Samoa: old tires
Elaphoidella juxtaputealis Damian & Botosaneanu,
Romania: city water pipes
1954
Elaphoidella phreatica (Chappuis, 1925)
Romania: city water pipe
Elaphoidella putealis (Chappuis, 1925)
Romania: city water pipe
Elaphoidella taroi
Fiji: tin cans
Epactophanes richardi
Hawaii: leaf-litter filled cup
traps on trees
E. richardi
Germany: trickle filters,
activated charcoal filters
Nitocrella calcaripes Damian & Botosaneanu, 1954
Romania: city water pipes
Nitocrella chappuisi Kiefer, 1926
Nitocrella hibernica s. str.
N. hibernica hyalina (Jakubisiak, 1929)
Nitocrella hirta Chappuis, 1924
N. hirta bucarestiensis Damian & Botosaneanu, 1954
Nitocrella kosswigi Noodt, 1954
Nitocrella omega Hertzog, 1936
Nitokra psammophila Noodt, 1952
Nitokra reducta Schäfer, 1936 s. str.
Nitokra sewelli husmanni Kunz, 1976
Germany: city water pipes
Germany: activated charcoal
filters
Poland: city water pipe
Romania, Hungary: city water
pipes
Romania: city water pipes
Romania: city water pipes
France, Hungary: city water
pipes
Germany: activated charcoal
filters
Germany: activated charcoal
filters
Germany: activated charcoal
filters
Borutsky (1934)
Kiefer (1926a, 1927a)
Török (1961)
Brady (1868)
Kiefer (1927a)
Damian (1958, 1959)
Damian & Botosaneanu
(1954)
Kiefer (1926a, 1927a)
Mrázek (1893)
Török (1951), Damian (1958,
1959), respectively
Damian (1958, 1959)
Damian (1958, 1959)
Damian & Botosaneanu
(1954)
Yeatman (1983)
Damian & Botosaneanu
(1954)
Chappuis (1925)
Chappuis (1925)
Yeatman (1983)
Evenhuis & Preston (1995)
Husmann (1966, 1982)
Damian & Botosaneanu
(1954)
Kiefer (1926b, 1927a)
Husmann (1982)
Jakubisiak (1929)
Chappuis (1925), Török
(1951), respectively
Damian & Botosaneanu
(1954)
Damian (1958, 1959)
Hertzog (1938), Török
(1961), respectively
Husmann (1982)
Husmann (1982)
Husmann (1982)
Continued on p. 215
215
Table 4. contd.
Species
Location: habitat
Reference
Nitokra spinipes Boeck, 1864
Germany: activated charcoal
filters
France: city water pipes
Romania: city water pipes
France: city water pipes
Romania: city water pipes
Hungary: city water pipes
Romania: city water pipes
Hungary: city water pipes
Hungary: city water pipes
France: city water pipes
Romania: city water pipes
Romania: city water pipes
Romania: city water pipes
Husmann (1982)
Paracamptus schmeili (Mrázek, 1893)
P. schmeili biserialis (Micoletzky, 1912)
Parastenocaris aedes Hertzog, 1938
Parastenocaris aquaeductus Chappuis, 1925
Parastenocaris budapestinensis Török, 1935
Parastenocaris clujensis Chappuis, 1925
Parastenocaris entzii Török, 1935
Parastenocaris germanica Kiefer, 1936
Parastenocaris hippuris Hertzog, 1938
Parastenocaris jeanneli Chappuis, 1924
Parastenocaris karamani brevicauda Damian, 1958
Parastenocaris latisetosus Damian & Botosaneanu,
1954
Parastenocaris minuta Chappuis, 1925
Parastenocaris nana Chappuis, 1925
Parastenocaris pannonica Török, 1935
Hertzog (1938)
Chappuis (1925)
Hertzog (1938)
Chappuis (1925)
Török (1935)
Chappuis (1925)
Török (1935)
Török (1951)
Hertzog (1938)
Damian (1958, 1959)
Damian (1958, 1959)
Damian & Botosaneanu
(1954)
Romania: city water pipes
Chappuis (1925)
Romania: city water pipes
Chappuis (1925)
Hungary, Romania: city water Török (1935, 1961), Damian
pipes
(1958, 1959), respectively
Parastenocaris similis Török, 1935
Hungary: city water pipes
Török (1935, 1961)
Parastenocaris subterraneus Damian, 1959
Romania: city water pipes
Damian (1958, 1959)
Parastenocaris uncinatus Damian & Botosaneanu, 1954 Romania: city water pipes
Damian & Botosaneanu
(1954)
Parastenocaris spp.
Germany: trickle filters
Husmann (1966)
Phyllognathopus viguieri
Germany: aquarium
Kessler (1914)
P. viguieri
Switzerland: aquarium
Chappuis (1916)
P. viguieri
Germany: indoor swimming Klie (1924)
pool
P. viguieri
Germany: water in coal mine Ziegelmayer (1923)
P. viguieri
England: ‘experimental
Gurney (1932), Scourfield
filters’
(1939)
P. viguieri
Romania, Germany: trickle
Damian-Georgescu (1966),
filters
Husmann (1966), respectively
P. viguieri
Tonga, Western Samoa, Fiji: Yeatman (1983)
old tires, tin cups, plastic
containers, old boats
P. viguieri
Hawaii: leaf-litter–filled cup Evenhuis & Preston (1995)
traps on trees
Spelaeocamptus spelaeus (Chappuis, 1925)
Romania: city water pipes
Chappuis (1925, 1927)
Unidentified Harpacticoida
U.S.A.: old tires
Nasci et al. (1987)
Cyclopoida
Acanthocyclops exilis (Coker, 1934)
U.S.A.: old tires
Nasci et al. (1987)
Acanthocyclops rhenanus Kiefer, 1936
France: city water pipes
Hertzog (1938)
Acanthocyclops robustus (Sars, 1863)
Romania: city water pipes
Damian (1958, 1959)
Acanthocyclops venustus (Norman & Scott, 1906)
France: city water pipes
Hertzog (1938)
Acanthocyclops vernalis (Fischer, 1853)
Canada: agricultural drainage Williams, D. D. (1976)
tiles
A. vernalis
U.S.A.: old tires
Nasci et al. (1987), Marten
(1989), Reid & Marten
(1994)
Bryocyclops anninae
Hawaii: leaf-litter–filled cup Evenhuis & Preston (1995)
traps on trees
Continued on p. 216
216
Table 4. contd.
Species
Location: habitat
Reference
Bryocyclops fidjiensis
Yeatman (1983)
Diacyclops crassicaudis (Sars, 1863)
Diacyclops languidoides hiberniae (Gurney, 1927)
Diacyclops navus (Herrick, 1882)
Fiji, Tonga, Western Samoa:
tin cans, bottles
Fiji: metal drums
Germany, Romania: city
water pipes
Germany: city water pipes
Romania, Hungary: city water
pipes
Romania: city water pipes
Wales: cistern
U.S.A.: old tires
Diacyclops ? stygius (Chappuis, 1924)
Romania: city water pipes
Ectocyclops phaleratus
Ectocyclops rubescens
U.S.A.: old tires
U.S.A.: old tires
Eucyclops agilis (Koch, 1838)
E. agilis
France: city water pipes
U.S.A.: old tires
Eucyclops elegans (Herrick, 1884)
U.S.A.: old tires
Eucyclops serrulatus
E. serrulatus
Germany: city water pipes
Romania: city water pipes
Graeteriella unisetigera (Graeter, 1908)
Germany: city water pipes,
trickle filters
Tonga, Western Samoa, Fiji:
old tires
U.S.A.: old tires
Cryptocyclops linjanticus
Cyclops sp.
Diacyclops clandestinus (Kiefer, 1926)
D. clandestinus
Halicyclops thermophilus s. str.
Macrocyclops albidus
M. albidus
Megacyclops gigas (Claus, 1857)
Megacyclops viridis
M. viridis
Mesocyclops affinis Van de Velde, 1987
Mesocyclops aspericornis
M. aspericornis
M. aspericornis
M. aspericornis
M. leuckarti
Mesocyclops longisetus (Thiébaud, 1912)
Mesocyclops ogunnus Onabamiro, 1957
Mesocyclops cf. pehpeiensis Hu, 1943
Mesocyclops ruttneri Kiefer, 1981
Romania: city water pipes
Germany: city water pipes
France: city water pipes
England: water tank
Vietnam: water containers
French Polynesia: barrels in
pigpen and old tires
Hawaii: containers
Colombia: mosquito rearing
containers
Vietnam: water containers
Iran: basin of soap factory
Venezuela: drinking tank
Vietnam: water containers
Vietnam: water containers
Yeatman (1983)
Kraepelin (1886), Damian
(1958, 1959), respectively
Kiefer (1926a, c, 1927a)
Damian (1958, 1959), Török
(1961), respectively
Damian (1958, 1959)
Gurney (1933)
Nasci et al. (1987; as
Thermocyclops dybowskii),
Marten (1989), Reid &
Marten (1994)
Damian & Botosaneanu
(1954)
Nasci et al. (1987)
Marten (1989), Reid &
Marten (1994)
Moniez (1889)
Nasci et al. (1987), Marten
(1989), Reid & Marten
(1994)
Nasci et al. (1987; as E.
speratus); Reid & Marten
(1994)
Kiefer (1927a)
Damian & Botosaneanu
(1954)
Kiefer (1926a, 1927a),
Husmann (1966), respectively
Yeatman (1983)
Marten (1989), Marten et al.
(1993), Reid & Marten
(1994)
Damian (1958, 1959)
Kiefer (1927a)
Moniez (1889)
Scourfield (1939)
Vu et al. (2000)
Rivière & Thirel (1981),
Rivière et al. (1987)
Marten (1984)
Suárez et al. (1984)
Vu et al. (2000)
Lindberg (1942)
Dussart (1984)
Vu et al. (2000)
Vu et al. (2000)
Continued on p. 217
217
Table 4. contd.
Species
Location: habitat
Reference
Mesocyclops thermocyclopoides Harada, 1931
Mesocyclops woutersi Van de Velde, 1987
Mesocyclops yenae Holynska, 1998
Metacyclops planus (Gurney, 1909)
Metacyclops tredecimus (Lowndes, 1934)
Orthocyclops modestus (Herrick, 1883)
Vietnam: water containers
Vietnam: water containers
Vietnam: water containers
Romania: city water pipes
Venezuela: drinking tank
U.S.A.: old tires
Paracyclops affinis (Sars, 1863)
Paracyclops fimbriatus
P. fimbriatus
P. fimbriatus
Senegal: laboratory cultures
of nematodes
England: percolating filters of
sewage works
France: city water pipes
Germany: city water pipes
Romania: city water pipes
Vu et al. (2000)
Vu et al. (2000)
Vu et al. (2000)
Damian (1958, 1959)
Dussart (1984)
Marten (1989), Reid &
Marten (1994)
Reversat et al. (1992)
P. fimbriatus
P. fimbriatus
Fiji: old tire
U.S.A.: old tires
P. fimbriatus s. lat.
Easter Island: stone basins
and mouth of stone statue
U.S.A.: old tires
Paracyclops chiltoni
Paracyclops poppei
Tropocyclops prasinus
T. prasinus
Calanoida:
‘Calaniden’
Karaytug & Boxshall (1998b)
Moniez (1889)
Kiefer (1926a, 1927a)
Damian & Botosaneanu
(1954)
Yeatman (1983)
Marten (1989), Reid &
Marten (1994)
Dumont & Martens (1996)
England: water tank
U.S.A.: old tire
Marten (1989), Reid &
Marten (1994), Karaytug &
Boxshall (1998a)
Scourfield (1939)
Reid & Marten (1994)
Germany: city water systems
Kraepelin (1886)
that most records predate the use of modern water
sanitation procedures. Kraepelin (1886) was apparently the first to report copepods (Cyclops sp. and
‘Calaniden’) in a municipal water system, in Hamburg, Germany. Shortly thereafter, Moniez (1889)
found several common species of cyclopoids in the
water pipes of Lille, France. Chappuis (1925), Török
(1935) and Hertzog (1938) described several new species of parastenocaridid harpacticoids from city water
pipes in Europe.
A human-created analogue to the natural psammon is the sand-filter beds (trickle filters) of water
treatment plants. Psammic or hypogean cyclopoids
and harpacticoids such as Graeteriella unisetigera,
E. richardi, P. viguieri, and species of Parastenocaris may colonize trickle filters (e.g. Husmann, 1961,
1966; Duncan, 1989). Even activated charcoal filters
may harbor copepods (Husmann, 1982). The literature
on copepods in various kinds of filters, much of it in
obscure local reports, is difficult to review; but some
examples are included in Table 4.
Discussion
Studies in the more cryptic habitats have contributed substantially to fundamental understanding of the
biology of copepods. These have provided information on developmental rates, adaptations to low food
supply and/or low temperatures, dispersal by natural
and human agencies, biogeography, evolution and
conservation.
Some of the fastest-developing copepods live in
temporary ponds. The most spectacular example is the
cyclopoid Metacyclops minutus, which in tiny ponds
in Germany may pass through 8 or more generations in
a single growing season. Under natural conditions they
may develop from egg to adult in 4.7–8.6 days, and in
218
the laboratory have matured in as little as 4 days at
30 ◦ C (Maier, 1992). In this species also, males differ
little from females in size, females and males mature
at about the same rate, and adults and subadults may
pass through dry phases in a state of quiescence, and
through the winter in diapause (Maier, 1992).
Adaptations to extreme conditions of low food supply and/or low temperatures include reduction in the
number (usually with enlarged size) of eggs, as in the
harpacticoid Pseudomoraria triglavensis (Brancelj,
1994). Some soil harpacticoids may produce only 2
eggs at a time, and drop them or actively lay them
in the leaf substrate (Nielsen, 1966); some small cave
cyclopoids also produce as few as 2 large eggs, and
drop them (Lescher-Moutoué, 1973).
Fiers & Ghenne (2000) found Graeteriella unisetigera, previously supposed to be a strict stygobiont, in
several beech litter samples, documented the striking
coincidence between its known distribution and the
historical limits of beech forests, and advanced the
hypothesis that the animals are able to travel along
the ground, rather than being restricted to particular
drainage basins. They pointed out that not only might
this be a convenient dispersal route, but would prevent
physical and genetic separation of populations in different drainage basins, thus reducing speciation. Fiers
& Ghenne (2000) noted that while larger stygobionts
tend to be restricted within drainage basins (examples
given by Notenboom et al., 1996), the tiny (about 0.5
mm long) copepods are usually more widely distributed. They may be not only less limited by such factors
as sediment permeability, but be more eurytopic than
previously estimated and able to exploit continuous
epigean highways of leaf litter, mosses, and moist
soils. Remy (1932) early remarked on the relative
uniformity of the leaf-carpet in space and time, and
suggested a possible role of humid forest habitats as
a refuge for specialists, particularly trogloxenes or
troglophiles, as well as for less specialized generalist
species.
A similar idea was advanced by Frey (1980), who
developed a concept of a worldwide rainforest habitat
and predicted that endemic or uniquely adapted species would be found there. Lewis (1986) remarked
on the analogous distribution of one harpacticoid
genus of the damp forest, Loefflerella, which occurs in Patagonia, Chile and New Zealand, to that of
the genus of semiterrestrial cladocerans, Bryospilus,
known from Puerto Rico, Venezuela and New Zealand
(Frey, 1980). The canthocamptid harpacticoid genus
Fibulacamptus is confined to wet temperate southern
Australia and Tasmania, mainly in semiterrestrial situations: a river, muddy gravel and dead leaves, and wet
moss and leaf litter (Hamond, 1988). Antipodiella is
endemic to New Zealand (Lewis, 1986). Among cyclopoids, several genera are endemic to humid forests,
and the known ranges of some are apparently restricted, e.g. Cochlacocyclops, Goniocyclops and Psammocyclops in Madagascar (Kiefer, 1955); Fimbricyclops in Puerto Rico (Reid, 1993b); Goniocyclops in
New Zealand (Harding, 1958); Muscocyclops mainly
in the Atlantic coastal forest of South America, with
outliers in continuous wetlands in central Brazil (Reid,
1987; Rocha & Bjornberg, 1987).
Lewis (1984) pointed out that in the harpacticoid
fauna of New Zealand, isolated since the Mesozoic,
only 2 of 16 genera are endemic but endemism at
the species level is high. The general picture of the
biogeography of the Canthocamptidae presented by
Lewis (1986) has not much changed with the addition of further data. Some exceptions are that a few
genera are now known to occur on more than one
continent, e.g. Gulcamptus in North America as well
as eastern Asia (Reid & Ishida, 1996). On the other
hand, advances in systematic understanding may alter
this picture for some groups; for instance, the New
World species of Attheyella (Canthosella) form a distinct group from the Asian species, and likely should
be considered a separate genus (Janetzky et al., 1996).
Future collections in semiterrestrial habitats outside Europe may reveal that some genera or species
are more widely distributed than presently thought.
For instance, although the genus Bryocamptus s. lat.
is widely distributed in the temperate northern hemisphere, in the southern hemisphere species have been
found in New Zealand (Lewis, 1986) and a wet campo
(hillside flush marsh) in central Brazil (Reid, 1993,
1994). The known distribution of Itocyclops yezoensis
(Ito, 1953) was extended by recent collections from
Japan and Alaska to the Great Smoky Mountains in
the eastern U.S.A. (Reid & Ishida, 2000).
Passive transport is usually invoked as an
effective distribution mechanism for microcrustaceans. Maguire (1971) described the colonization of
phytotelmata as “the result of a series of interlinked
events – dispersal, immigration and establishment.”
Janetzky (1997) suggested the possibility that disseminules might be blown by the wind, since P.
viguieri appeared in a rain gauge in Jamaica. Obviously protective resting stages have been observed in
few species of harpacticoids, although the eggs of at
least some leaf litter- and soil-dwelling species resist
219
desiccation to some degree (e.g. Bryocamptus (Arcticocamptus) spp., see Borutsky, 1952; E. richardi, see
Nielsen, 1966), and adults of a few others (e.g. A. nordenskioldii, see D. D. Williams & Hynes, 1976) form
drought-resistant cysts.
There are few papers that discuss natural transport, even by implication. Rouch (1972) described two
harpacticoids, Phyllognathopus bassoti and Nitocrella balli, which had established good populations on
a small sandy island that appeared in Wisdom Lake,
a shallow crater lake on Long Island, Papua New
Guinea. The copepods were collected 20 months after
formation of the small island.
Copepods are apparently easily transported by
various human activities. They may be carried in
plant parts and moist soil, and exotic tropical copepods have been found in greenhouses or associated
with tropical aquatic or terrestrial plants in several
parts of the world: the East Asian Sinodiaptomus sarsi
(Rylov, 1923) in California, U.S.A. (Light, 1939);
the Brazilian Attheyella aliena in Germany (described
by Noodt, 1956); the Asian Mesocyclops ruttneri
in Austria (reported and described by Kiefer, 1981),
Louisiana, Mississippi and Washington, D.C., U.S.A.
(Reid, 1993a, 1996); and Bryocyclops muscicola,
which was first described from Indonesia, in Florida, U.S.A. (Reid, 1999). Harpacticoids of the genus
Phyllognathopus are frequently found in moist soils,
and are probably so easily carried as to make determination of the origin of a particular population
nearly impossible. Phyllognathous viguieri has been
reported from greenhouse soils and the recesses of
tropical plants (Lang, 1948; Borutsky, 1952; Lehman
& Reid, 1993; Table 1). Hitchhiking with commercially cultured fish and crustaceans is more often
invoked as a transport mechanism (reviewed by Reid
& Pinto-Coelho, 1994).
The subterranean realm has been a hospitable situation for copepods (Stoch, 1995, 2001; Galassi, 2001).
Attempting to account for the relatively high species
and genus richness in certain hypogean areas, Stoch
(1995) proposed the adaptive zone model, involving
steps of colonization, speciation events and radiation,
to describe the process of invasion of epigean species
and their subsequent evolution in hypogean environments.
Copepods in general are an ancient group (Huys
& Boxshall, 1991; Stock, 1991), and they may evolve
extremely slowly in stable habitats, that is in habitats that do not themselves change much or allow easy
physical access to other kinds of habitats. An example
is the allopatric species-pair Acanthocyclops sensitivus (Graeter & Chappuis, 1914) in Europe and A.
parasensitivus Reid, 1998, in eastern North America.
These are identical in gross morphological characters
such as appendage segmentation and setation, and differ only in certain proportions and finer ornamentation
(Pospisil, 1999). Since both are rare and found only
in groundwater-related habitats, we can assume that
gene flow between the European and North American
populations is nil. These two taxa have diverged only
slightly since the continents separated.
However, where they have invaded complex habitats such as large lakes, particularly epibenthic and
infaunal copepods have radiated to a sometimes surprising degree, sometimes over a relatively short time.
The most complex species-flocks exist in Lake Baikal,
where cyclopoids (3 flocks of 3, 4 and 13 species) and
harpacticoids (6 flocks of 3–21 species) account for a
large proportion of the 120 or more copepod taxa in
the lake (Boxshall & Evstigneeva, 1994). The Baikal
basin complex has existed (as one or several basins)
since the Tertiary, and this long period has apparently
allowed the development of a high degree of trophic
specialization and mouthpart structure among the cyclopoids, and extreme sexual dimorphism in the caudal
rami among the harpacticoids (Boxshall & Evstigneeva, 1994). The more recent formation (during the
Pleistocene) of large subterranean lakes or cenotes in
the Yucatan Peninsula apparently allowed primarily
epigean benthic cyclopoids in the genera Diacyclops
and Mesocyclops to radiate into small flocks of 2 and 3
species, respectively, including true planktonic forms
(Fiers et al., 1996).
Many rarely collected copepods live preferentially
in and are adapted to ephemeral waters. In Germany,
where naturally created temporary habitats are disappearing, tiny pools maintained in early stages of
succession by vehicular activity or other factors harbor several rare or endangered species of copepods,
cladocerans and branchiopods (Herbst, 1982; Maier,
1998; Maier et al., 1998). Herbst (1982) listed 6 species of calanoids, 16 cyclopoids and 37 harpacticoids
that are at some degree of risk in Germany because
they inhabit special habitats, including small perennial
plant-rich waterbodies, oligotrophic to mesotrophic
ponds, acid bogs and moorland waters, cave waters,
moss, saline waters and lake beaches, in addition to
tiny ephemeral pools. Because the highly organic soils
of the wet campo, a kind of hillside flush marsh in
Brazil which harbors an extremely diverse copepod
fauna (Reid, 1994), are valuable for truck farming, wet
220
campos that are not in protected areas are at risk of being drained and destroyed. Bayly (1992, 1997, 1999)
reviewed the highly diverse fauna in gnammas (rock
pools) and seepage films on rock outcrops in Western
Australia, and argued strongly for their protection.
“Sehr zu Unrecht sind die Kleingewässer und
ihre Lebewelt bisher von der Wissenschaft etwas
vernachlässigt worden” (Kiefer, 1925). Hollowday
(1949) also urged that attention to neglected habitats
would result in valuable new information: “It is by the
investigation of such habitats as these that peculiar and
specialized forms are often met with”. Further pursuit
of imaginative collecting, and ecologically oriented
studies of copepods living at the natural extremes permissible to these basically aquatic forms are bound to
provide additional insights on many aspects of their
fundamental biology.
Acknowledgements
My gratitude is extended to Carlos E. F. Rocha for suggesting this topic. I thank Sam Lake, Gerhard Maier,
Giuseppe L. Pesce, David L. Strayer, and Nicole A. C.
Zyngier for providing or suggesting references, Frank
Fiers for a pre-publication copy of the manuscript by
Fiers & Ghenne, and Rubens Mendes Lopes, Carlos
Rocha, and David Strayer for their constructive comments. The facilities of the C. B. Wilson Copepod
Library and the Natural History Library, National Museum of Natural History, Smithsonian Institution were
invaluable.
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