Revista de Biología Tropical ISSN: 0034-7744 [email protected] Universidad de Costa Rica Costa Rica Lessios, H.A. Echinoids of the Pacific Waters of Panama: Status of knowledge and new records Revista de Biología Tropical, vol. 53, núm. 3, -diciembre, 2005, pp. 147-170 Universidad de Costa Rica San Pedro de Montes de Oca, Costa Rica Available in: http://www.redalyc.org/articulo.oa?id=44919815009 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Echinoids of the Pacific Waters of Panama: Status of knowledge and new records H.A. Lessios Smithsonian Tropical Research Institute, Apartado 0843-03092, Balboa, Panama; Fax: 507-212-8790; [email protected] Received 14-VI-2004. Corrected 09-XII-2004. Accepted 17-V-2005. Abstract: This paper is primarily intended as a guide to researchers who wish to know what echinoid species are available in the Bay of Panama and in the Gulf of Chiriqui, how to recognize them, and what has been published about them up to 2004. Fifty seven species of echinoids have been reported in the literature as occurring in the Pacific waters of Panama, of which I have collected and examined 31, including two species, Caenopedina diomediae and Meoma frangibilis, that have hitherto only been mentioned in the literature from single type specimens. For the 31 species I was able to examine, I list the localities in which they were found, my impression as to their relative abundance, the characters that distinguish them, and what is known about their biology and evolution. Not surprisingly, most available information concerns abundant shallow water species, while little is known about deep water, rare, or infaunal species. Rev. Biol. Trop. 53(Suppl. 3): 147-170. Epub 2006 Jan 30. Key words: Eastern Pacific, sea urchins, Bay of Panama, Gulf of Chiriqui, systematics. The eastern Pacific waters of Panama the dry season (December-April) to these strong extend roughly from 9o01’ N, 82o52’ W to winds. The winds push surface water to the 6o50’ N, 77o54’ W (Fig. 1). They comprise South, and cause upwelling of cold, nutrient two embayments, separated by the Azuero rich water, which greatly increases primary pro- Peninsula: (1) The Bay of Panama, which ductivity, but also reduces surface temperatures contains the Pearl Island Archipelago and (2) from an average of about 26o to as low as 16oC. the Gulf of Chiriqui, which contains various Possibly as a consequence of the low tempera- island complexes, with Islas Coiba, Cebaco, tures, certain ecologically important echinoderm Ladrones, Secas and Contreras, being the most species, such as Acanthaster planci (L.), though prominent. Though depths within the two bays present in the Gulf of Chiriqui, do not enter the are shallower than 200 m, the 1 000 m isobath Bay of Panama (Glynn 1974). passes within less than a kilometer from the The echinoid fauna of the Pacific coast shores of the Azuero and Burica Peninsulas and of Panama (and the rest of the eastern Pacific) the island of Jicaron (Fig. 1). Oceanographic has been sampled and described during the conditions of the area have been described past two centuries mostly as the result of by Forsbergh (1969), Glynn (1972, 1982), the “Albatross” (Agassiz 1898, 1904), “Zaca” D’Croz et al. (1991), D’Croz and Robertson (H.L. Clark 1940), and Velero III (A.H. Clark (1997), and Podesta and Glynn (2001). There 1939, H.L. Clark 1948) expeditions. Detailed are pronounced differences in the seasonality information about the distribution of echino- of thermal and trophic regimes of the two bays. derms of the region has been very ably synthe- Whereas the Gulf of Chiriqui is protected by sized and tabulated by Maluf (1988), who has high mountains from the trade winds out of the rendered a tremendous service to echinoderm North, the Bay of Panama is exposed during biogeographers by combing the entire literature Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (Suppl. 3): 147-170, December 2005 147 on eastern Pacific echinoderms and plotting systematic study of certain species that occur the range of each species. Mortensen spent off the southern shores of Panama, much of several months on the island of Taboga in the it motivated by the known time of the closure Bay of Panama in 1915, and, though he never of the Isthmus, there is no work that contains published a specific work on the echinoderms detailed information on which echinoid species of the region, he included the information he are likely to be found in any given locality of gathered on the distribution and larval devel- the Bay of Panama or of the Gulf of Chiriqui, opment of the local echinoids in other pub- or a summary of the systematic, phylogenetic, lications (e.g. Mortensen 1921, 1928-1951). and ecological information that exists for each Boone (1928) listed observations on specimens of them. My own work over the past 25 years of echinoderms from Islas Perlas, along with on life histories and evolution of various spe- those from the Sea of Cortez, but occasionally cies of echinoids has led me to seek popula- confused the two areas, stating at one point tions and specimens in Panamanian waters. that the type specimen of Arbacia incisa (A. Here I attempt to summarize the information Agassiz) comes from “Guaymas, Panama”. I have gathered about systematics and micro- A.H. Clark (1946) also treated specimens that distributions of Panamanian echinoid species came into his hands from Islas Perlas, but con- in the course of this work, along with informa- centrated mostly on Encope , and described a tion that can be gleaned from the literature. new species, E. wetmorei A. H. Clark from the This paper is primarily intended as a guide to area. Chesher (1972) provided a list of east- researchers who may wish to know what echi- ern Pacific echinoids with presumed Atlantic noid species are available in the Bay of Panama “geminates”, i.e. species that were formed by and in the Gulf of Chiriqui, how to find them, the rise of the Isthmus of Panama. However, how to recognize them, and what has been pub- despite rather intense ecological and molecular lished about them up to 2004. Fig. 1. Pacific waters of Panama: depth contours and main features. Localities are identified in the text in relation to the main features and by indicating their coordinates the first time they are mentioned. Fig. 1. Aguas del Pacífico de Panamá: contornos de profundidad y características principales. Se identifican las localidades en el texto en relación con las características principales e indicando sus coordenadas en la primera vez que son mencionadas. 148 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (Suppl. 3): 147-170, December 2005 Records in the literature (L.), it seems to be confined to the outer islands of the eastern Pacific, without having established Table 1 is excerpted from the information populations close to the mainland (Lessios et al. presented by Maluf (1988) for the entire east- 1996, 1998). ern Pacific, so as to include only those species In addition to the preceding species about whose range includes Panama. As the Table which there may be doubt as to their presence indicates, 40 echinoid species are expected to in Panama, there are two species, the absence occur in depths of < 100 m, and 17 species of which in Panama is almost certain. Maluf’s in deeper water, for a total of 57 species. For (1988) records of Lytechinus pictus (Verrill), biogeographical purposes, it is important to and its synonym, L. anamesus H.L. Clark, in distinguish those species for which there are Panamanian latitudes are based on H.L. Clark’s actual records of specimen collection within (1948) mention of one specimen from Bahia Panamanian waters, from those for which collec- Honda and one specimen from Gorgona Island, tion records straddle Panamanian latitudes, but Colombia. Clark states that these may well be have not been documented as occurring in the specimens of L. panamensis Mortensen, and area. There are seven species that fall in the lat- this is very likely the case. Mitochondrial DNA ter category. Of these, Kamptosoma asterias (A. sequences have shown that, though L. pictus Agassiz), Plesiodiadema horridum (A. Agassiz), cannot be distinguished from L. anamesus and Cystocrepis setigera (A. Agassiz), Pourtalesia though L. panamensis cannot be distinguished tanneri A. Agassiz, and Homolampas fulva A. from L. semituberculatus (Valenciennes), the Agassiz are deep water species, difficult to col- clade composed of the first two is quite dis- lect, and thus could well be located off Panama tinct from the clade of the last two (Zigler and in future sampling. However, for two additional Lessios 2004). There is, therefore, no reason to species, the inference that they are likely to assume the alternative explanation, that L. pic- occur in Panama would not necessarily be cor- tus, L. anamesus and L. panamensis are all one rect. Centrostephanus coronatus (Verrill) is a species. The second case of mistaken literature subtropical species, preferring the colder waters reports concerns Caenocentrotus gibbosus (L. of California and of the Galapagos Archipelago. Agassiz and Desor). Maluf (1988) indicates It has not been found in Panama, though it is this species as occurring at 8oN, but none of apparently present in Costa Rica (J. J. Alvarado the references she cites places it in any locality pers. com.). Echinometra oblonga (Blainville) above the equator. was found North of Panama in Clarion and Maluf ’s (1988) compilation indicates that Socorro Islands (H.L.