Cnidarians and Copepods: A Success Story

Author(s): Arthur G. Humes

Source: Transactions of the American Microscopical Society, Vol. 104, No. 4 (Oct., 1985), pp.
313-320
Published by: Wiley on behalf of American Microscopical Society
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 TRANSACTIONS
of the
American Microscopical
Society
VOL. 104 OCTOBER 1985 NO. 4

Cnidarians and Copepods: A Success Storyl 2

ARTHUR G. HUMES
Boston University Marine Program, Marine Biological Laboratory,
Woods Hole, Massachusetts 02543, U.S.A.

Abstract. Copepods often live in association with other marine invertebrates. More
than 400 species of copepods live with about 350 cnidarians, particularly in tropical areas.
The greatest numbers of these associates occur with Anthozoa, especially scleractinian
corals (172 copepods), although soft corals, gorgonians, and sea anemones combined have
a nearly equal number of associated copepods. The order Poecilostomatoida contains by
far the largest number of cnidarian associates, many of them with their bodies highly
modified or transformed. Biological aspects of these associated copepods, such as immunity
to nematocyst toxin, host specificity, effects on the host, feeding, larval development,
means of dispersal, possible use of microniches in multiple infestations, and the role of
the copepods in their ecosystems are either not well understood or unknown.

The Copepoda in their evolutionary history have developed more associa-

tions with other animals than any other group of Crustacea. Although these
small crustaceans are well known as parasites of fishes, many live in various
degrees of association with marine invertebrates (Gotto, 1979). Fresh-water
invertebrates such as bivalve mollusks seem to offer a suitable refuge, but
generally lack copepod associates.

1 Past-presidential address presented on 28 December 1984 at the Annual Meeting of the Amer-
ican Microscopical Society in Denver, Colorado.
My contribution to the study of copepods associated with cnidarians could not have been made
without gratefully acknowledged help from many friends, colleagues, and institutions. Over the
past 30 years, field work in the Caribbean, West Africa, Madagascar, Enewetak Atoll, New Cale-
donia, the Moluccas, Hawaii, Moorea, Panama, and Australia, comprising nearly four years in all,
has enabled me to learn much about these copepods. Without the hospitality and assistance of
many persons and organizations this field work would have been impossible. I particularly thank
those who, over the years, collaborated with me in the collection and study of these copepods:
Roger F. Cressey, Alfred de Maria, Masahiro Dojiri, Pierre Fourmanoir, Bruce W. Frost, Carlos
Goenaga, Richard U. Gooding, Roger Halverson, Ju-Shey Ho, Charles T. Krebs, Pierre Laboute,
George S. Lewbel, Victor McCauley, Wayne L. Smith, and Jan H. Stock.
2 Publication costs, in part, are
being met by a grant from the Spencer-Tolles Fund of the
American Microscopical Society.
TRANS.AM. MICROSC.Soc., 104(4): 313-320. 1985.
? Copyright, 1985, by the American Microscopical Society, Inc.

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314 TRANS. AM. MICROSC. SOC.

In 1880, Antonio Della Valle described for the first time the copepod Li-
chomolgus actiniae (now Critomolgus actiniae, see Humes & Stock, 1983)
from the sea anemone Actinia concentrica at Naples, Italy. Since then, interest
in copepods living with cnidarians has increased as the diverse tropical and
subtropical faunas have been investigated. The last 30 years have seen rapid
growth in the study of copepod associates, largely on account of greater ac-
cessibility to remote areas and the use of SCUBA for making under-water
collections. Although 50 or more years ago it was common to find associates
described "from washings of marine invertebrates" or from unidentified cni-
darians, recent work has usually included the exact name of the host. This has
promoted studies of host specificity and geographical distribution.
In the last 40 years, the number of copepods known to live with cnidarians
has grown more than twentyfold. This expansion has been influenced by sev-
eral factors. (1) Cnidarians from readily accessible shallow-water inshore areas
are easy to collect. Copepod associates are therefore best known from intertidal
and subtidal cnidarians. (2) Beginning about 1950, improved air travel and
increased financial support made it possible to do field work in little-studied
areas, particularly in the tropical Indo-Pacific. (3) Special techniques such as
snorkeling and SCUBA diving have made possible collections that previously
had been unattainable. (4) The necessity for exact host identification and the
need for careful methods of collection in order to record the correct host-
copepod relationship have been recognized.
Among various invertebrate groups, the Cnidaria have more copepod asso-
ciates than any other group, approximately 416 species at present count (Table
I). These copepods are most abundant in shallow-water reef-building herma-
typic corals belonging to the Scleractinia, where 172 species are known (Table
II). Recently, however, Stock (1978) has found a copepod associated with a
solitary ahermatypic coral in 1,000 m off Sri Lanka, indicating that deep-water
cnidarians also serve as hosts. Moderately large numbers of copepods occur
with the Alcyonacea, Gorgonacea, and Actiniaria.
Of the various orders of Copepoda, the Poecilostomatoida contains by far
the greatest number of species associated with cnidarians (Table III). The
Siphonostomatoida are much less frequent, and the Cyclopoida, Harpacticoida,
and Calanoida are represented by relatively small numbers.
Copepods are associated with all three classes of Cnidaria. They have de-
veloped associations especially with the Anthozoa, both octocorals (with 161
copepods) and hexacorals (with 227 copepods). The Scleractinia serve as hosts
for many more copepods than other groups (Table II).
The Hydrozoa have relatively few copepods, perhaps on account of their
often small size and relative fragility. These copepods occur on the polyp stages
only. Macrochiron is a well-known genus originally thought to be free living
but now known from several hydroid genera such as Lytocarpus and Aglao-
phenia in the Indo-Pacific (Humes & Stock, 1973).
The Scyphozoa have only a few copepod associates, represented by Para-
macrochiron from medusae such as Thysanostoma near Japan (Humes, 1970).
Another genus, Sewellochiron, lives on the medusa Cassiopea in Puerto Rico

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VOL. 104, NO. 4, OCTOBER 1985 315

TABLE I
Numbers of copepods associated with groups of invertebrates

Number of host Number of host Number of copepod Number of copepod

genera species genera species

Porifera 36 45 27 81
Cnidaria 160 350 141 416
Polychaeta 73 107 57 95
Echinodermata 119 216 106 236
Crustacea 119 206 70 179
Mollusca 202 371 91 202
Ascidiacea 63 227 82 237
Nemertina 1 1 1 1
Turbellaria 2 2 2 2
Bryozoa 4 4 4 4
Sipuncula 4 4 4 4
Echiuroida 1 1 1 1
Merostomata 1 1 1 1
Hemichordata 1 1 1 1
Pterobranchia 2 3 2 3
Thaliacea 3 9 2 9
Pogonophora 2 2 3 3
Total 793 1,550 595 1,475

(Humes, 1969). Perhaps the sedentary behavior of the larvae of invertebrate-

associated copepods has prevented them from successfully invading planktonic
hosts such as medusae.
The Anthozoa have a large number of copepod associates, perhaps resulting
from the larger colonies of the hosts providing more space for copepods, from
the relatively large gastrovascular cavities offering a protected environment,
and from the greater opportunity for the evolution of host specificity in these
very diverse cnidarians.
Among the Anthozoa, the Octocorallia have relatively unmodified associates
such as Acanthomolgus from gorgonaceans (Humes, 1974; Stock, 1975a), high-
ly modified forms such as Lamippe from alcyonaceans (Bouligand, 1966), and
highly transformed copepods like Magnippe from a gorgonacean off Florida
(Stock, 1978).
In the Hexacorallia, the sea anemones have 37 copepod associates, most of
them apparently inhabiting the gastrovascular cavities of the polyps. These
copepods range from only slightly modified forms like Lambanetes from the
large sea anemone Stichodactyla (see Humes, 1982) to more transformed forms
like Mesoglicola that lives in galls in the corallimorpharian Corynactis (see
Haefelfinger & Laubier, 1965) and Staurosoma found in galls in the anemones
Anemonia and Anthopleura (see Laubier & Schmidt, 1971). Ridgewayia foss-
hageni, the only calanoid copepod associated with an invertebrate, lives in
aggregations near the tentacles of the anemone Bartholomea in Panama (Humes
& Smith, 1974).
The Antipatharia, the black and whip corals, have relatively few copepods.

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316 TRANS. AM. MICROSC. SOC.

TABLE II
Numbers of copepods associated with groups of Cnidaria

Number of host Number of host Number of copepod Number of copepod

genera species genera species

Hydrozoa 11 17 10 20

Scyphozoa 6 6 6 8
Anthozoa
Octocorallia
Stolonifera 3 5 6 11
Telestacea 2 2 3 3
Alcyonacea 25 93 25 85
Gorgonacea 27 44 15 47
Pennatulacea 8 14 6 15
Hexacorallia
Actiniaria 23 37 16 37
Corallimorpharia 3 3 4 5
Scleractinia 47 118 41 172
Zoanthidea 1 2 4 6
Antipatharia 3 8 4 6
Ceriantharia 1 1 1 1
Total 160 350 141 416

These include Calonastes from Stichopathes in Puerto Rico (Humes & Goe-
naga, 1978) and Vahinius from the same genus of whip corals in Madagascar
(Humes, 1967) as well as from Cirripathes in the Moluccas (Humes, 1979a).
Other copepods live with Stolonifera, Telestacea, Pennatulacea, Zoanthidea,
and Ceriantharia (Table II).
The Scleractinia, or hard corals, have the greatest number of copepod as-
sociates (Humes, 1979b; Table II), about half of them in one family, the Xa-
rifiidae (Humes, 1985). This copepod family is a well-defined group in which
the body is elongate and transformed, usually less than 3 mm in length, with
weakly defined segmentation and reduced appendages. The four genera in the
Xarifiidae (Xarifia, Orstomella, Lipochrus, and Zazaranus) are known only
from shallow-water hermatypic corals, where they live in the polyps. The
largest genus, Xarifia, with 75 species, lives in 93 species of corals from the
Red Sea-Madagascar area eastward to an arc formed through Japan-Enewetak
Atoll-New Caledonia. Xarifia is absent, as far as we know, from the Pacific
Ocean east of 166?; none of these copepods have been found in Hawaii, Moo-
rea, and Panama (Humes, 1985).
Xarifiidae are absent in corals in the Caribbean. Instead, various Caribbean
corals are parasitized by members of another family of highly transformed
poecilostomatoid copepods, the Corallovexiidae (Stock, 1975b). Although a few
species of several Indo-Pacific coral genera, such as Acropora, Favia, Porites,
and Tubastrea, live in the Caribbean, they have no xarifiid associates.

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VOL. 104, NO. 4, OCTOBER 1985 317

TABLE III
Distribution of 416 species representing five orders of copepods that live in association with Cni-
daria

Harpacti- Poecilo- Siphono-

Calanoida coida Cyclopoida stomatoida stomatoida

Hydrozoa - 1 12 7

Scyphozoa - 2 - 6
Anthozoa
Octocorallia
Stolonifera - - 11
Telestacea - - 3 -
Alcyonacea - - 4 80 1
Gorgonacea - - - 45 2
Pennatulacea - - -15 -

Hexacorallia
Actiniaria 1 1 - 32 3
Corallimorpharia - - 3 2
Scleractinia - 3 - 153 16
Zoanthidea - - - 6 -
Antipatharia - -- 6
Ceriantharia - - 1 -
Total 1 7 4 373 31

From a study of the distribution of associated copepod genera among the

major groups of Cnidaria (Table III), three observations may be made. First,
the Poecilostomatoida have succeeded in developing the most associations with
cnidarians. Second, the Siphonostomatoida have a respectable number of as-
sociated copepods, but far fewer than the Poecilostomatoida. It should be
noted, however, that interest in siphonostomes has not paralleled interest in
poecilostomes. On the basis of many collections of siphonostomes still awaiting
study in my laboratory, these copepods appear to be common associates. The
relatively small number of siphonostomes reported from cnidarians is probably
due to neglect and is not real. Third, the Harpacticoida have been considered
to be only rarely associated with cnidarians, but the recent discovery of several
species from hydrozoans and corals, e.g., species of Tegastes and Alteuthel-
lopsis (see Humes, 1981, 1984), indicates that harpacticoids may be more
frequent associates than previously believed.
The effects of copepods on the cnidarians are difficult to assess. Lamippe
can be seen in soft corals moving about within the polyps, disappearing inside
the coenenchyme and appearing again in another polyp, two or three polyps
down the line (Patton, 1972). Xarifia crawls in and out of the polyps, tearing
the polyp tissue with its claws (Gerlach quoted in Humes, 1960). The presence
of Mesoglicola induces the formation of galls in sea anemones (Haefelfinger &
Laubier, 1965; Taton, 1934). According to Vader (1970), as many as 310

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318 TRANS. AM. MICROSC. SOC.

individuals of the large copepod Antheacheres duebeni may occur in one sea
anemone, Bolocera tuediae, but even in such large numbers the copepod ap-
parently does not interfere with reproduction of the host.
For the most part, copepods associated with cnidarians probably feed on
mucus secreted by the host. Briggs (1977) found that the gut of Paranthessius
contains mucopolysaccharides, as would be expected from a mucus feeder.
Antheacheres lacks a digestive tract and evidently absorbs nourishment by
diffusion through its body wall (Vader, 1970). Such diffusion may also occur
in Lamippe, a genus commonly found in octocorals (Bouligand, 1966).
Copepod associates of actiniarians apparently have an immunity to the toxin-
producing nematocysts found in the tentacles of these cnidarians. In experi-
ments on copepod immunity to Anemonia sulcata, Briggs (1976) found that
of several copepod species tested only Paranthessius anemoniae, a natural
associate of this anemone, moved freely among the tentacles, while other co-
pepods were either paralyzed or digested, or showed avoidance reactions. Ex-
tract of tentacles of Anemonia had little effect on Paranthessius anemoniae,
but the extract appeared to paralyze other copepods temporarily.
Host specificity among copepods of cnidarians is not well understood. Only
recently have collections been extensive enough and sufficiently well docu-
mented to provide information on this phenomenon. There are many instances
where a copepod species is known from only one species of host and is thus
stenoxenous. But in the absence of large numbers of collections from related
hosts, it is often difficult to say definitely that the copepod is truly monoxenic.
On the other hand, we know of cases where a copepod has a wide choice of
hosts and is euryxenous, as is Doridicola spinulifer from 13 species of alcyo-
naceans in Madagascar (Humes & Stock, 1973). Even here, however, a degree
of specificity exists, with 10 of the 13 host species belonging to the genus
Lemnalia. In several species of Doridicola on Californian sea anemones (An-
thopleura, Epiactis, and Tealia), Lonning & Vader (1984) found strict host
specificity.
One host cnidarian (individual or colony) may support more than one species
of copepod. Presumably, the various copepods have particular microniches
involving location, feeding habits, and reproductive behavior. Among exam-
ples of hosts with several species of copepods may be cited the sea anemone
Heteractis crispa (see Humes, 1982), the soft coral Nephthea cupressiformis
(see Humes, 1980), and the hard coral Galaxea fascicularis (see Humes, 1979c),
each with five copepod associates, and the hard coral Acropora florida with
seven species of Xarifia (see Humes, 1985).
In summary, the Copepoda have been extremely successful in developing
associations with Cnidaria, particularly with soft corals, gorgonians, and hard
corals in tropical and subtropical regions. In doing so, the copepods have under-
gone considerable morphological modification and adaptation. The majority
of copepods living with cnidarians are Poecilostomatoida, although Siphonos-
tomatoida (e.g., Stock, 1981) and Harpacticoida (e.g., Humes, 1981, 1984) are
probably much more frequent associates than presently recorded.

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VOL. 104, NO. 4, OCTOBER 1985 319

The biology of these associated copepods is poorly known. The effects on

the host, the method of feeding, the means of protection against nematocyst
toxin, host specificity, larval development, method of dispersal, the use of
microniches in multiple infestations, and the role of these copepods in the
ecosystems in which they live are aspects still poorly known for most species.
Although copepods are well known as abundant members of marine plank-
ton and as parasites of fishes, they also have been highly successful in forming
associations with many marine invertebrates. The story of their success is best
told by observing their presence on so many cnidarians.
LITERATURE CITED

BOULIGAND,Y. 1966. Recherches recentes sur les copepodes associ6s aux anthozoaires. In Rees,
W. J., ed., The Cnidaria and their Evolution, Academic Press, New York, pp. 267-306.
BRIGGS,R. P. 1976. Biology of Paranthessius anemoniae in association with anemone hosts. J.
Mar. Biol. Assoc. U.K., 56: 917-924.
1977. Structural observations on the alimentary canal of Paranthessius anemoniae, a copepod
associate of the snakelocks anemone Anemonia sulcata. J. Zool. (Lond.), 182: 353-368.
DELLA VALLE, A. 1880. Sui coriceidi parassiti, e sull'anatomia del gen. Lichomolgus. Mitt. Zool.
Stat. Neapel, 2: 83-106.
GOTTO,R. V. 1979. The association of copepods with marine invertebrates. Adv. Mar. Biol., 16:
1-109.
HAEFELFINGER, H. R. & LAUBIER,L. 1965. D6couverte en Mediterran6e occidentale de Meso-
glicola delagei Quidor, copepode parasite d'actinies. Crustaceana, 9: 210-212.
HUMES,A. G. 1960. New copepods from madreporarian corals. Kiel. Meeresforsch., 16: 229-
235.
1967. Vahinius petax n. gen., n. sp., a cyclopoid copepod parasitic in an antipatharian coe-
lenterate in Madagascar. Crustaceana, 12: 233-242.
1969. A cyclopoid copepod, Sewellochiron fidens n. gen., n. sp., associated with a medusa in
Puerto Rico. Beaufortia, 16: 171-183.
1970. Paramacrochiron japonicum n. sp., a cyclopoid copepod associated with a medusa in
Japan. Publ. Seto Mar. Biol. Lab., 18: 223-232.
1974. Cyclopoid copepods (Lichomolgidae) from gorgonaceans in Madagascar. Proc. Biol. Soc.
Wash., 87: 411-438.
1979a. Poecilostome copepods associated with antipatharian coelenterates in the Moluccas.
Beaufortia, 28: 113-120.
1979b. Coral-inhabiting copepods from the Moluccas, with a synopsis of cyclopoids associated
with scleractinian corals. Cah. Biol. Mar., 20: 77-107.
1979c. Poecilostome copepods (Lichomolgidae) associated with the scleractinian coral Galaxea
in the Moluccas. J. Nat. Hist., 13: 507-528.
1980. Copepods (Cyclopoida, Lichomolgidae) associated with the alcyonacean Nephthea in
the Moluccas. Hydrobiologia, 68: 49-71.
1981. Harpacticoid copepods associated with Cnidaria in the Indo-West Pacific. J. Crustacean
Biol., 1: 227-240.
1982. A review of Copepoda associated with sea anemones and anemone-like forms (Cnidaria,
Anthozoa). Trans. Am. Philos. Soc., 72: 1-120.
1984. Harpacticoid copepods associated with Cnidaria in the tropical Pacific Ocean. Zool. Scr.,
13: 209-221.
1985. A review of the Xarifiidae (Copepoda, Poecilostomatoida), parasites of scleractinian
corals in the Indo-Pacific. Bull. Mar. Sci., 36: 467-632.
HUMES,A. G. & GOENAGA,C. 1978. Calonastes imparipes, new genus, new species (Copepoda,
Cyclopoida), associated with the antipatharian coral genus Stichopathes in Puerto Rico.
Bull. Mar. Sci., 28: 189-197.

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320 TRANS.AM. MICROSC.SOC.

HUMES, A. G. & SMITH, W. L. 1974. Ridgewayia fosshageni n. sp. (Copepoda; Calanoida)

associated with an actiniarian in Panama, with observations on the nature of the association.
Caribb. J. Sci., 14: 125-139.
HUMES, A. G. & STOCK, J. H. 1973. A revision of the family Lichomolgidae Kossmann, 1877,
cyclopoid copepods mainly associated with marine invertebrates. Smithson. Contrib. Zool.,
127: 1-368.
1983. Redefinition of the genus Doridicola Leydig, 1853, synonymy of Metaxymolgus Humes
& Stock, 1972, and establishment of a new genus, Critomolgus (Copepoda, Poecilostoma-
toida, Lichomolgidae). Bull. Zool. Mus., Univ. Amst., 9: 93-96.
LAUBIER,L. & SCHMIDT,H. 1971. Le genre Staurosoma Will (copepode parasite d'actinies) en
Mer Rouge. Crustaceana, 21: 52-56.
LONNING, S. & VADER,W. 1984. Sibling species of Doridicola (Copepoda: Lichomolgidae) from
California sea anemones: biology and host specificity. J. Exp. Mar. Biol. Ecol., 77: 99-135.
PATTON, W. K. 1972. Studies on the animal symbionts of the gorgonian coral, Leptogorgia
virgulata (Lamarck). Bull. Mar. Sci., 22: 419-431.
STOCK, J. H. 1975a. On twelve species of the genus Acanthomolgus (Copepoda Cyclopoida:
Lichomolgidae) associated with West Indian octocorals. Bijdr. Dierkd., 45: 237-269.
1975b. Corallovexiidae, a new family of transformed copepods endoparasitic in reef corals.
Stud. Fauna Curaqao other Caribb. Isl., 47: 1-45.
1978. Magnippe caputmedusae n. gen., n. sp. (Copepoda: Lamippidae), a highly transformed
endoparasite in octocorals of the genus Thesea from the Gulf of Mexico. Mem. Hourglass
Cruises, 3: 1-11.
1981. Associations of Hydrocorallia Stylasterina with gall-inhabiting Copepoda Siphonosto-
matoidea from the south-west Pacific. Part II. On six species belonging to four new genera
of the copepod family Asterocheridae. Bijdr. Dierkd., 51: 287-312.
TATON, H. 1934. Contribution a l'etude du copepode gallicole Mesoglicola delagei Quidor. Trav.
Stat. Biol. Roscoff, 12: 53-68.
VADER, W. 1970. Antheacheres duebeni M. Sars, a copepod parasitic in the sea anemone, Bo-
locera tuediae (Johnston). Sarsia, 43: 99-106.

Baltimore Meetings
The annual meeting of the American Microscopical Society will be held in
conjunction with meetings of the American Society of Zoologists, Animal Be-
havior Society, The Crustacean Society, International Association of Astacolo-
gy, and Society of Systematic Zoology at the Baltimore Convention Center,
Baltimore, Maryland, 27-30 December 1985. The host institution is Towson
State University with Donald C. Forester and Philip D. Creighton chairing the
Local Arrangements Committee. Some 13 symposia are scheduled. Also, sev-
eral workshops will be offered, including one on Application of Histochemical
Techniques to Microscopy. I ..., sponsored by the American Microscopical
Society (see "Special Announcement," p. 377, this issue).
Housing and advance registration forms will be available in October with a
deadline of 20 November. Additional planned features of the meetings include
socials, special programs, commercial exhibits, and a job placement service.
For further information, contact Mary Adams-Wiley, ASZ Executive Officer,
Box 2739, California Lutheran College, Thousand Oaks, California 91360;
Telephone: (805) 492-3585, or Frederick W. Harrison, AMS Secretary, De-
partment of Biology, Western Carolina University, Cullowhee, North Carolina
28723; Telephone (704) 227-7244.

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