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Experimental Interbreeding Between Geographically Marine Biology 17, t49--157 (1972) by Springer-Verlag t972 Experimental Interbreeding between Geographically Separated Populations of the Marine Wood-Boring Isopod Limnoria tripunctata with Preliminary Indications of Hybrid Vigor* R. J. Menzies Oceanography Department, Florida State University; Tallahassee, Florida, USA Abstract larval stage, and are restricted by food requirements Populations of the marine wood-boring isopod Limnoria and habitat preference to submerged wood in the sea. tripunctata Menzies were cultivated and interbred in the Each piece of infested wood supports a more or less laboratory in order to determine whether geographically unique population. Within a restricted migratory or separated populations would interbreed. Two series of inter- swimming range, gene-flow is ]probably continuous and breeding studies were conducted. In the first, field populations from li Atlantic and Caribbean sites and 3 Pacific sites were a population in a given harbor or bay may be consid- interbred with a North Western Atlantic population from ered to constitute one "deme". Beaufort, North Carolina (USA). Viable crossings, to at least Between harbors, the amotmt of submerged wood F1, occurred in all but one case -- that of a Beaufort • Chat- available is usually slight, most having found its way ham (Massachusetts, USA) cross. In the second series of ex- periments, field populations from 9 Atlantic and Caribbean to the shore-line where silt, sand, and desiccation make sites were crossed with populations from St. Teresa, Florida, it unfit for Limnoria tripunctata. It was believed that Gulf of Mexico. In this ease also, all crosses except one pro- the crossing of populations from different harbors duced viable offspring, not only to the 2'I, but to the F~ gene- should yield some interesting information on the com- rations as well. The St. Teresa • Chatham cross was not successful. Geographic distance was not a factor in the success putability of geographically disjunct populations of of crossbreeding between disjunct populations. Chatham popu- the same species. Presumably gene-flow should be lations (mainly) are now believed t~) represent a valid but greatest (even though low, at best) between geograph- previously obscure species, Limnoria tuberculata Sowinski, ically close harbors, and least between more distant and not a variant of L. tripunctata Menzies. Enhanced vitality or vigor, as determined from rate of wood destruction, popu- harbors. lation growth rate and survival, was indicated in several crossings, but this was not necessarily correlated with distance Geographic Distribution between populations. Observations were made on the manner in which the species has achieved a worldwide distribution, and Limnoria tripunctata Menzies has a worldwide a hypothesis is put forward to explain the evolution of species distribution in the tropical, subtropical, and partly related to L. tripunctata. temperate seas. It has been collected between Boston, Massachusetts and Bahia Blanca, Argentina, in the Introduction Western Atlantic Ocean, including the Gulf of Mexico and Caribbean Sea, and between Plymouth, England The problem was to determine whether geograph- and Dakar, Senegal in the Eastern Atlantic Ocean, ically isolated populations of the marine isopod as well as the Mediterranean and Red Seas. In the Limnoria tripunctata were reproductively compatible. Pacific Ocean, it is distributed between San Francisco In major microscopic characteristics they had already Bay and Balboa, Panama, and among the islands of been determined to be phenotypica]ly conspecifie the Western and Central Pacific Ocean from Japan (Menzies, i957). This problem involves a number of to Northern Australia. In the Indian Ocean it has been sub-problems concerning the experimental methods recorded from Madras and Mombassa. This distribu- required for successful cultivation of marine species tion is a close parallel to subtropical-tropical tempera- from geographically distant localities. tm'e ranges (Menzies, 1959). The isopods are dioecious, lack a free swimming The contacts Limnoria tripunctata Menzies makes * Contribution from Zoology Department, Duke Universi- with other species at its northern range are of some ty, Durham, North Carolina, and Duke University Marine interest. On the Pacific coast, it meets Zimnoria Laboratory, Beaufort, North Carolina, and the Department of Oceanography, No. 289, Florida State University, Tal- quadripunctata Holthuis at San Francisco Bay, Cali- lahassee, Florida. Support in part from NSF Grant l~o. GB- fornia, where it is found sympatrically with that species 6898. (Menzies, 1958). North of Point Arenas, California, 20 MarineBiology, Vol. 17 t50 R.J. Menzies: Experimental Interbreeding in Limnoria Mar. Biol. L. quadripunctata is replaced by Limnoria lignorum conditions in order to reach compatible thermal con- Rathke. A similar situation occurs in England, but ditions. there all three species are found together (Jones, 1963). The two southern species are replaced in Nor- Reproduction and Temperature way by L. lignorum Rathke. On the Atlantic coast of The relationship between reproduction and tem- North America, L. tripunctata meets L. lignorum at perature is a strong one in Limnoria tripunctata, and Chatham, Massachusetts. On this coast, however, appears to be related to the duration of temperature L. quadripunctata is absent. Kussakin (1963) has des- favorable for reproduction throughout the year. cribed species of Limnoria from the low Arctic. It Beckman and Menzies (i960) found that the species seems likely that his Arctic "species" are closely re- would breed and reproduce under laboratory condi- lated to L. lignorum. This contention might be subject tions between 15~ and 30 ~ This range of temperature to simple testing through interbreeding experiments coincides almost exactly with the temperature range such as those reported here. How a pantropical distri- encountered by the species in nature. The optimal bution has been realized in L. tripunctata has not yet temperature, at which reproduction rate was highest, been determined. It is commonly believed that the was between 20 ~ and 25 ~ Although the correlation isopods were transported on the infested hulls of between distribuSion and a temperature range of wooden ships and, although not the orrly one, that i5 ~ to 30 ~ is great, a better correlation is established seems a possibility. The chances are equally good that between the duration of temperatures favorable for the species had a worldwide thermal distribution reproduction and distribution. during the Eocene, and is now restricted to the modern limits of the subtropics and tropics. Although the possibility of occasional introductions Mode of Reproduction into a given locality from elsewhere must be admitted, it is reasonable to believe that the closer one location Limnoria, like many other marine isopods, is dioe- is to another, the better the chance of gene inter- cious, and requires the presence of a male and a female change. It has been demonstrated that populations for successful reproduction. Transfer of sperm to the of Limnoria tripunctata (Menzies, 1968) can withstand female has not been observed due to the secretive transit through the fresh water of the Panama Canal, nature of the animals. The male sexual apparatus and in the days of wooden ships, such transport might consists of a pair of genital apophyses located on the have been one mode of dispersal. By the time She canal midline at the caudal end of the seventh peraeonal was built, however, wooden ships were not in wide use. (thoracic) somite (Fig. t B). Sperm transfer is assisted The possibility of transisthmian transport of specimens by the appendix masculinum attached to the inner in the ballast tanks of ships cannot be excluded, but margin of the endopodite of the second male pleopod. iS is a remote possibility due to the foul conditions usually found in ship ballast and bilges. Transport of driftwood around Cape Horn and the Cape of Good Phenomena Associated with Reproduction Hope from one ocean to another would be difficult. On old infested wood, gravid females are usually Infested pieces of wood would have to originate in the found in the blind end of a burrow, with the male near tropics or subtropics and enter boreal and antiboreal the entrance (Fig. 2). Usually new burrows are started ~ haart ~mite $omite tu"num Fig. ~. Limnoria tripunctata. Sections showing'reproductiveapparatus. (A) Female; (B) male Vol. 17, No. 2, 1972 R. J. Menzies: Experimental Interbreeding irt Limnoria 151 ~_l arva, Fig. 2. Limnoria tripunctata. Section of infested wood showing position of male, female, and larva by the male. The female enters a newly started burrow Experimental Design as soon as it is long enough and large enough for All of the foregoing characteristics of reproduction double occupancy. The female enters a burrow in a and ecology have a bearing on the desigaa of the pregravid state, in which the oSstegites are small experiments. The most significant variable, ff ecologic and the internal ova are ripe for fertilization. Copula- factors remain favorable, is the reproductive behavior tion occurs in the burrow and is apparently simul- of the species. Limnoria tr@unctata lacks any sperm r with a moulting of the female because, at this storage mechanism, hence pairing of a male with a time, ~he large o6stegites are produced, the huge eggs "virgin" female is a prerequisite to successful repro- pass through the oviducts, are fertilized during passage
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