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Shore Barnacles (Cirripedia, Thoracica) and a Revision of Their Provincialism and Transition Zones in the Tropical Eastern Pacific

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Shore Barnacles (Cirripedia, Thoracica) and a Revision of Their Provincialism and Transition Zones in the Tropical Eastern Pacific BULLETIN OF MARINE SCIENCE, 46(2): 406-424, 1990 SHORE BARNACLES (CIRRIPEDIA, THORACICA) AND A REVISION OF THEIR PROVINCIALISM AND TRANSITION ZONES IN THE TROPICAL EASTERN PACIFIC Jorge E. Laguna ABSTRACT The emergence of the Isthmus of Panama separated the Caribbean from the Tropical Eastern Pacific (TEP) and played a major role in the biogeographical development of the tropical and latitudinal marine provincialism seen today. Latitudinal provincialism in the Tropical Eastern Pacific has been assessed in terms of the distribution of a variety of taxa. A detailed study of the barnacles provides a basis for a refined model for the provincial associations observed. Closer attention needs to be paid to transition zones, and as it turns out, the GuifofCalifornia is not a province as strictly defined, but an extension of a transition zone. Provincial asso- ciations are defined in terms of the barnacles of the Tropical Eastern Pacific, and are compared with those known for other taxa. From the early Oligocene to the late Pliocene the Atlantic and Pacific Oceans were connected through the Panamic Strait and, by shallow water circulation, biotic exchange took place between them (Van Andel, 1979; Keigwin, 1982; Valentine, 1971; 1973). Because of the transport of planktonic larvae of sedentary forms by equatorial currents, the fauna of both oceans remained very similar until the late Miocene (Woodring, 1966). With the tectonic closure of the Panamic Isthmus, and an increase in the Caribbean temperature (about 5.7 my B P), a modem hydrographic setting and circulation were established (Emiliani et a1., 1972; Keigwin, 1982). The physical changes brought about by this geologic event separated the biota of both oceans, causing the extinction of some species, survival of others, and the evolution of new ones. This is indicated by the fact that some species are still found on both sides of the Isthmus and others have survived in either the Pacific or the Caribbean. The Tropical Eastern Pacific, a larger and more open body of water than the Caribbean, was less affected (had fewer extinctions) by the physical changes produced by this uplift (Woodring, 1966). These changes led to the present marine tropical provincialism found in these oceans (Ekman, 1953). The Tropical Eastern Pacific was defined by Ekman (1953) as the region ex- tending from the southern tip of western Baja California (23°N) and the Gulf of California to the Gulf of Guayaquil (3-4°S). Transitional subtropical faunas were defined between southern Baja California and San Diego (23-33°N) in the north, and between Ecuador and Peru in the south (3-6°S; Ekman, 1953). A number of modified definitions have been given (Cromwell and Bennett, 1959; Abbott, 1966; Zullo, 1966; Briggs, 1973), but these remain basically the same as Ekman's (1953). However, Valentine (1966; 1973) found that the northern transition zone between tropical and temperate molluscan species was further south (20-28°N, Valentine's Surian Province). Brusca and Wallerstein (1979) came to the same conclusion based on isopod crustacean data (24-28°N). This also seems to be the case for the barnacles. The Tropical Eastern Pacific has also been described as the region lying between the 20°C isotherms (Abbott, 1966; Valentine, 1966; 1973). The thermal tolerance 406 LAGUNA: BARNACLE PROVINCIALISM, EASTERN PACIFIC 407 of an organism is often related to temperature optima for enzyme systems, and temperature often regulates reproductive cycles (Newman and Abbott, 1980). Subtropical faunal breaks are mainly regions of rapid thermal change associated with discontinuity in the coastal surface currents (Abbott, 1966). These regions lead to changes from one faunal association to another and are referred to herein as transition zones. Similar circumstances can occur in the tropics. Off Guatemala for example, two thermally distinctive water masses converge (Cromwell and Bennett, 1959; Abbott, 1966) giving rise to appreciable biotic changes (Valentine, 1973). Until the work of Laguna (1985; 1987) there was no major unifying study of the balanomorph barnacles from the Tropical Eastern Pacific, as evident from the depauperate literature for this region. An earlier attempt to compile this knowledge was made by Zullo (1966), but at that time, there were only 57 species ofbalanomorph barnacles known from the entire eastern Pacific, seven of which were assigned to the Tropical Eastern Pacific. This number has now increased to more than 50 species for the TEP, excluding whale and turtle barnacles. Of the 57 tropical species, 28 range throughout the Panamic Province s.1.(the tropics). The rest are restricted to members of northern and southern overlapping sub- tropical provinces (Californian, Mexican, and Peruvian Provinces; Table 1, Fig. 1). METHODS All collections used for this study are housed in the Benthic Invertebrate Collections of the Scripps Institution of Oceanography, La Jolla, California, except the Jorge Laguna collection which is at the National Museum of Natural History, Washington, D.C. (Laguna, 1985). The major lots are: Scripps Institution of Oceanography-Smithsonian Tropical Research Institute (SIO-SI 1971-1973) collection: a survey of intertidal barnacles on both coasts of Panama, mostly in areas adjacent to the Panama Canal (Laguna, 1985, appendices I, II). Tropical Eastern Pacific Expedition (TEPE-1978) collection: a general survey of 47 stations found between the Gulf of Panama and the Gulf of California (Laguna, 1985, appendix II). Kathy DeReimer collection (1980): miscellaneous collections on the Pacific coast of Central America, and some sites on the Atlantic coast (Laguna, 1985, appendix II). Jorge Laguna collection (1984): a survey of the intertidal barnacles from the Pacific and Atlantic coasts of Panama (Laguna, 1985, appendix I). These collections make Panama the best sampled region for barnacles in the Tropical Eastern Pacific. In addition to the collections, available literature on cirripeds from this region was utilized for de- termining species ranges (Darwin, 1851; 1854; Pi!sbry, 1916; 1918; Rogers, 1948; Zullo, 1966; Henry and McLaughlin, 1975; Newman and Ross, 1976; Southward and Newman, 1977). The ranges of species of Tropical Eastern Pacific Cirripedia were plotted (Table I, Fig. I). Ranges of some non- cirriped taxa known from the literature are available for comparison with those of thoracic Cirripedia in the Tropical Eastern Pacific (Table 2, Fig. 2). These latitudinal ranges have been used to determine provincial associations in the TEP. Barnacle species cluster latitudinally. The regions between cluster of species constitute faunal breaks. The faunal breaks have regions of overlap referred to as transition zones, and form the basis for defining the faunal provinces. Each province has a group of endemic species, and wide-ranging species from other provinces. The number of species mentioned as belonging to each province corresponds to the endemic portion found there. Many authors have previously used the concepts of "province" and "transition zone" (Valentine, 1966; 1973; Briggs, 1974; Newman, 1979b; Brusca and Wallerstein, 1979). Some confusion on their use exists due to the lack of a unifying general definition. A marine province as envisaged in this paper, is a relatively large region encompassing a variety of habitats and a diversity of communities. A province has a characteristic high number of endemic species (45% or more; derived from the percent endemic versus the total number of species found in each province) that overlap to varying degrees those of adjacent provinces; but the substantial endemic portion distinguishes it. The region constituting a province is apparently determined primarily by physical factors (Valentine, 1966; 1973; Dolan et aI., 1972), although biological factors may be im- portant (Newman, 1979b; Brusca and Wallerstein, 1979). A latitudinal transition zone is the region of overlap between two or more faunal provinces; it may 408 BULLETIN OF MARINE SCIENCE, VOL. 46, NO.2, 1990 Table I. Cirripedia of the Tropical Eastern Pacific grouped according to Provinces. For ranges of distribution see Figure I and Laguna, 1985 Group Provincial Associations Californian 1. Tamiasoma regalis (Newman and Foster, 1987) 2. Oxynaspis rossi Newman, 1972:202 3. Membranobalanus orcutti (Pilsbry, 1916:233) 4. Tetraclita rubescense (Darwin, 1854:462) 5. Megabalanus californicus (Pilsbry, 1916:65) 6. Chthamalus fissus (Sowerby, 1833:74) 7. Balanus pacificus pacificus (Pilsbry, 1916:104) 8. Pollicipes polymerus Sowerby, 1833:74 9. Balanus nubilus Darwin, 1854:253 10. Concavus aquila (Pilsbry, 1907: 199) 11. Balanus glandula (Pilsbry, 1916: 180) II Mexican 12. Chthamalus "cortezianus" (undescribed; Hedgecock, 1979) 13. Megaba/anus peninsu/aris (Pilsbry, 1916:66) 14. Tetraclita confmis (Pilsbry, 1916:255) 15. Balanus inexpectatus inexpectatus Pilsbry, 1916:97 16. Chthamalus anisopoma Pilsbry, 1916:317 17. Concavus panamensis eyerdami (Henry, 1960) 18. Membranobalanus n.sp. Newman, pers. comm. 1985 19. Acasta n.sp. Newman, pers. comm. 1985 20. Heteralepas n.sp. Newman, pers. comm. 1985 21. Balanus parkeri Zullo, 1967 III Amphitropical 22. Pollicipes elegans (Lesson, 1830:441) IV Panamic s.1. 23. Conopea galeata (Linnaeus, 1771:544) 24. Octolasmis californiana Newman, 1960b 25. Megabalanus coccopoma (Darwin, 1854:196) 26. Hexacreusia durhami Zullo, 1961 27. Balanus poecilus Darwin, 1854 28. B. pacificus mexicanus Henry, 1941: 109 29. Euraphia imperatrix Pilsbry, 1916:320 30. E. eastropacensis (deOliveira,
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