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Shore Fishes and Biogeographic Subdivisions of the Tropical Eastern Pacific

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Shore Fishes and Biogeographic Subdivisions of the Tropical Eastern Pacific Vol. 380: 1–17, 2009 MARINE ECOLOGY PROGRESS SERIES Published April 7 doi: 10.3354/meps07925 Mar Ecol Prog Ser OPENPEN ACCESSCCESS FEATURE ARTICLE Shore fishes and biogeographic subdivisions of the Tropical Eastern Pacific D. Ross Robertson1,*,**, Katie L. Cramer2,** 1Smithsonian Tropical Research Institute, Balboa, Panamá 2Center for Marine Biodiversity and Conservation, Scripps Institution of Oceanography, University of California, San Diego, California, USA Address for correspondence: DRR, STRI, Unit 9100, Box 0948, DPO, AA 34002, USA ABSTRACT: We examined the geographic distributions of 1135 species of resident shore fishes to assess biogeo- graphic subdivision of the Tropical Eastern Pacific (TEP), which stretches from the Gulf of California to northern Peru. Using hierarchical clustering refined by Analysis of Similarity (ANOSIM), we determined geographic groupings in the distributions of the entire fauna, of re- gional endemics and of 3 functional (habitat) groups of species. We also examined the distributions of local en- demics throughout the TEP and how differences in fau- nal size versus faunal composition among sites con- tribute to the subdivision pattern. Our results indicate that: (1) the continental coast contains 2 provinces, the Cortez (Gulf of California and lower Pacific Baja) and the Panamic (southward), each of which has a peak in abun- dance of local endemics and of overall species richness; (2) the northern and southern boundaries of the TEP are located near Magdalena Bay on Baja California (~25° N) and the southern shore of the Gulf of Guayaquil (~4° S), respectively; and (3) the 5 oceanic islands/archipelagos collectively represent a third, Ocean Island Province. Relative to mainland areas, the fauna of the ocean is- Regional distributions of 1135 near-shore fish species (see lands is smaller, has a different functional-group compo- www.stri.org/sftep) indicate that the Tropical Eastern Pacific sition, and includes more transpacific species and more is divided into 3 biogeographic provinces. highly localized endemics. The 3-province pattern prob- Illustration: Ernesto Peña ably developed in response to the formation of the Gulf of California, the rise of the Isthmus of Panama, immigra- tion from the north, south and west to the TEP, and dif- INTRODUCTION fering environmental conditions between and within provinces. In contrast, barriers to dispersal within this In his pioneering analysis of global marine bio- geographically simple region are weak and likely had geography Briggs (1974) identified 4 major tropical much less influence. ‘regions’, which represent centers of high diversity and KEY WORDS: Tropical eastern Pacific · Zoogeography · endemism for shore-living organisms: the Indo-West Dispersal barrier · Gulf of California · Isthmus of Panama · Pacific, the East Pacific, the West Atlantic, and the East Shore fishes · Speciation Atlantic. His and earlier analyses of the distributions of shore fishes (e.g. Walker 1960, Rosenblatt 1967) and Resale or republication not permitted without written consent of the publisher other tropical organisms (Ekman 1953) on the Pacific * Email: [email protected] © Inter-Research 2009 · www.int-res.com **Equal authorship 2 Mar Ecol Prog Ser 380: 1–17, 2009 coast of the Americas indicated that the Tropical East- as part of a warm temperate ‘NE Pacific Biorealm’, ern Pacific (TEP) spans the continental shore between equivalent to the California Province of Briggs (1974). southern Baja California to northern Peru, and in- (2) The southern limit of the TEP followed the limit set cludes the Galapagos and the 4 other isolated ocean by a study of crustacean distributions (Boschi 2000, our islands and archipelagos—the Revillagigedo group, Fig. 1C), and was placed further south (7° S), than the Clipperton, Cocos, and Malpelo (Fig. 1A). limit set by Briggs (1974) and Hastings (2000) (3° to Briggs (1974) used information on the regional distri- 4° S). (3) The TEP was split into 2 provinces, the Gala- bution of local endemic reef fishes and arbitrarily set a pagos and the remainder of the region (see Fig. 1D). minimum endemism rate of 10% to distinguish 2 main- The northern and southern boundaries of the TEP as land provinces within the TEP: the Mexican and the defined by all of these studies are not ‘hard’ barriers Panamic. He placed the northern limit of the TEP and of that physically prevent dispersal of adult or larval his Mexican Province at ~25° N, not only on the Pacific fishes. Rather they relate mainly to relatively sharp coast of Baja but also inside the Gulf of California, and temperature gradients between tropical and temperate the lower limit of the latter at ~16° N, on the northern conditions. As environmental conditions around those edge of the Gulf of Tehuantepec, Mexico. He excluded boundary areas fluctuate considerably over time and the northern and central Gulf of California (above space, so do the range limits of TEP shore fishes and 25° N) from the TEP, asserting that it has a largely tem- other organisms. During El Niño events waters along perate fauna that is allied most strongly to the fauna of the continental shorelines to the north and south of the the Californian Province, which extends north from TEP warm considerably, and the ranges of many TEP 25° N along the Pacific coast of the Baja Peninsula. His endemics expand beyond, sometimes well beyond, Panamic Province stretched from ~16° N to ~3° S at the their normal regional limits. Chirichigno & Vélez (1998) border between Ecuador and Peru (see Fig. 1B). Based and Love et al. (2005) present summaries relating to on literature indicating high levels of local endemism such temporary southern and northern range exten- among algae, corals, mollusks, and crabs as well as sions by shore fishes. shore fishes (Walker 1966), Briggs (1974) recognized The only zoogeographic studies to date that have ana- the Galapagos Archipelago as a third province of the lyzed the faunal relationships between all 5 ocean islands TEP. and the mainland of the TEP are Hastings (2000) on Earlier, Walker (1960) defined 2 provinces in Mexico chaenopsid blennies and Glynn & Ault (2000) on stony based on the distributions of locally endemic reef corals throughout the TEP and in the central Pacific. fishes: a Cortez Province, consisting of Pacific coast of Here, we examine biogeographic partitioning of the Baja below 25° N, plus all of the Gulf of California, and entire TEP from the perspective of its shore fish fauna, a Mexican Province for the remainder. Later informa- using more comprehensive information on the distrib- tion (Rosenblatt 1974, Thomson et al. 1979) showed utions of those fishes than was available to previous that tropical reef fishes are more widely distributed in workers. We assess levels of faunal similarity and local the Gulf of California than Briggs (1974) realized, sup- endemism among different parts of the TEP to provide porting Walker’s view that most or all of that Gulf is answers to 3 questions: (1) Where do the northern and part of the TEP. southern boundaries of the TEP lie? (2) How many con- Within the TEP there are 2 major breaks in the distrib- tinental provinces are there in the TEP, and where do ution of shoreline reef habitats. These stretches of sand boundaries between provinces occur? (3) Should the 5 and mud shorelines are known as the Sinaloan Gap (370 ocean islands be considered as one or more provinces km of shoreline in the SE Gulf of California) and the of the TEP? Finally, we considered the historical ori- Central American Gap (~1000 km of shoreline from the gins of the provincial arrangement of the TEP indi- Gulf of Tehuantepec, southern Mexico, to El Salvador). cated by our analyses. These 2 gaps define the breakpoints separating the 3 mainland provinces (Cortez, Mexican, and Panamic) as delimited by Hastings (2000) (and see Springer 1959, MATERIALS AND METHODS and Fig. 1A here) based on reef fish faunal affinities. The most recent formal subdivision of the TEP, as Species database. The zoogeographic database in part of a general scheme for all global shorelines, by Robertson & Allen (2006) that provides data on the dis- Spalding et al. (2007), differs significantly in 3 ways tribution of 1261 shore fish species currently known in from all previous schemes: (1) The northern limit of the shallow water (0 to 100 m depth) in the TEP was used in TEP was set at ~20° N on the mainland, well south of the present study. That database is a synthesis of infor- the boundary recognized by Walker (1960), Briggs mation obtained from a bibliography of >1100 primary (1974), and Hastings (2000); all of Baja and the Gulf of and secondary sources from the scientific literature, as California were excluded from the TEP and included well as surveys by D.R.R. in Mexico, El Salvador, Costa Robertson & Cramer: Biogeography of the Tropical Eastern Pacific 3 Rica, Panama and all 5 oceanic islands. Further, under- Our database includes 1135 species of ‘TEP resi- standing of the limits of distributions of shore fishes dents’, species whose abundance and/or distributions throughout the Gulf of California and along the coast of indicate they have self-sustaining populations in the Baja has been greatly enhanced by 2 recent reviews: region (cf. Robertson et al. 2004). Of those, 897 are Findley et al. (2003) and Love et al. (2005). ‘TEP endemics’, species whose ranges are entirely or Island A Cortez Panamic Island B Mexican Californian and Peruvian Sinaloan and Central American Gaps N33 N33 30° N31b N31 N31b N31 N N29b N29 N29b N29 N27b N27 N27b N27 N25b N25 N25b N25 N23 N23 N25a N25a 20° N21 N21 N19 N19 Revillagigedos N16a N16a N17 N17 N15 N15 N13 N13 10° N11 N11 N9 N9 Clipperton N9b N9b N7 N7 Cocos N8a N8a Malpelo N5 N5 ? Gorgona N3 ? N3 0° N1 N1 Galapagos S2 S2 S4 S4 S6 S6 A Ekman 1953 / Walker 1960 / Hastings 2000 B Briggs 1974 S8 S8 10° S N33 N33 30° N31b N31 N31b N31 N N29b N29 N29b N29 N27b N27 N27b N27 N25b N25 N25b N25 N23 N23 N25a N25a 20° N21 N21 N19 N19 N16a N16a N17 N17 N15 N15 N13 N13 10° N11 N11 N9b N9 N9b N9 N8a N7 N8a N7 N5 N5 N3 N3 N1 N1 0° S2 S2 S4 S4 S6 S6 C Boschi 2000 D Spalding et al.
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