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The community structure of the decapod crustacean fauna of 7 tropical, shallowwater, marine habitats (sandy beaches, mangrove swamps and rocky intertidal habitats on both the Pacific and Caribbean coasts of Panama, and Pocillopora damicornis coral habitat of the Bay of Panama) were examined and analyzed for species composition and relative abundances. Collections from the 7 habitats yielded 4361 individuals, representing 236 species. The number of species per habitat was (Pacific, Caribbean): sandy beach (16, 7); mangrove (20, 17); P. damicornis (53); rocky intertidal (78, 67). There were more species represented by more individuals in the Pacific habitats. An index of faunal similarity was calculated for each pair (Pacific-Caribbean) of habitats. This index is the number of ecologically similar congeneric species which occurred in both habitats expressed as a percentage of the total number of species present in the pair of habitats. For the sandy beach communities there were three Pacific species which were similar to three Caribbean species, a similarity of 6/23 or 26%. The index of similarity for the mangrove communities is 54% and for the rocky intertidal communities it is 37%. The P. damicornis community has affinities with the Pacific rocky intertidal community (18%), with that of the Caribbean rocky intertidal (16%) and with that of Indo-West Pacific pocilloporid corals (20%). A few specialized species dominated each of the communities. The habitats and the number of species accounting for over half of the individuals present are (Pacific, Caribbean): sandy beach (1, 1); mangrove (4, 4); P. damicornis (5); and rocky intertidal (3, 6). Most of the species in each community were represented by one or a few individuals.

In Costa Rica, there are 591 species of decapods, 8.1% of the known marine biodiversity of the country in both oceans (Vargas and Wehrtmann 2009; Wehrtmann et al. 2009). The families in this group with the greatest species richness in Pacific Costa Rica are Xanthidae (45 spp.), Porcellanidae (44 spp.), Majidae (43 spp.), Alpheidae (34 spp.), Ocypodidae (28 spp.) and Palaemonidae (23 spp.) (Vargas and Wehrtmann 2009). Similarly, of the 1688 marine species reported for Isla del Coco, 8.2% are decapods (Cortés 2012). Some decapod species are adapted for symbiotic behavior. Among the most well-known groups for establishing associations with other species are the families Palaemonidae, Alpheidae, Pinnotheridae and Porcellanidae (Baeza 2007). The decapods that live in association with other animals in Costa Rica have been scarcely studied and in most publications they are only mentioned in species lists with no indication of their association with other organisms. The objective of the present study is to present a compilation of species of decapod crustaceans associated with macroinvertebrates in the Pacific of Costa Rica.

One-hundred associations are reported, which include 74 species of decapods are guests of six phyla of hosts (Table 1). The phylum with the most associated decapods was Cnidaria, with 74 species i.e., 74% of the total), followed by Echinodermata with 15 species, and four each with Annelida and Mollusca, two with Chordata, and one with Porifera. The host order with the greatest diversity of associated decapods was Scleractinia (Table 1). The species that showed the most associations with different hosts was Zenopontonia soror, which was found in five species of seastars (Table 1). This species was recently reported new to Costa Rica by Vargas-Castillo and Cortés (2019). The shrimp Pseudocoutierea elegans was found in hosts from three groups (Octocorallia, Hydrozoa, and Echinoidea). On the other hand, Tetrias scabripes was found associated solely with polychaetes (Table 1).

In total, 74 species of symbiotic decapods have been discovered in Pacific Costa Rica, 13 anomurans, two axiidids, 34 brachyurans, 24 carideans, and one gebiidid (Table 2). The family with the most symbiotic species was Palaemonidae (20 spp.), followed by the families Pinnotheridae and Porcellanidae, with nine species each. The genus with the most species was Trapezia with four, followed by Periclimenes with three. Fourteen new reports of decapods were recorded for Isla del Coco along with the occurrence of four new decapod records for Pacific Costa Rica, Calyptraeotheres pepeluisi Campos and Hernández-Ávila 2010, Raytheres clavapedatus (Glassell, 1935), Tuleariocaris holthuisi Hipeau-Jacquotte 1965 and Pseudocoutierea elegans Holthuis 1951. This is the first time that the genus Calyptraeotheres is reported in Costa Rica (Table 2).

Fifty-six species of decapods, distributed in five orders, 23 families and 50 genera, were found associated with 21 species of cnidarians. Opecarcinus crescentus has only been found in Pavona gigantea, while Alvarado and Vargas-Castillo (2012) reported 16 species of decapods associated with Pocillopora damicornis, all of which are typically found with this host. Six additional species are reported associated to Pocillopora sp. Seven species were found living on Porites lobata and one, Platypodiella rotundata, exclusively on Tubastraea coccinea. Fifteen species were found associated with Antipatharia. The two species associated with Antipathes sp. were also found in Myriopathes panamensis. Only one associated deep-water decapod, Coralaxius galapagensis, was found on Lillipathes ritamariae. In Octocorallia, 22 associated species were found. Eugorgia mutabilis was the host with the greatest diversity of decapods, six. The two most common decapods in octocorals were Neopontonides henryvonprahli and Pseudoveleronia laevifrons; four crustaceans could not be identified to species. In the hydrozoans, symbionts have only been collected from Stylaster marenzelleri, where Munida sp. and Pseudocoutierea elegans were found.

Four species of decapods, distributed in two orders, two families and three genera, were found associated with four species of molluks. Symbionts have been found primarily in bivalves. In specimens of the pearl oyster Pinctada mazatlanica, pairs of the shrimp Pontonia margarita have been found living inside the oyster on numerous occasions; P. simplex was found in Pinna rugose. In the oyster, Saccostrea palmula, the pinnotherid crab Austinotheres angelicus has been reported as a guest with a prevalence of 38% (Mena et al. 2014). Only pairs of Calyptraeotheres pepeluisi were found living in the interior of the gastropod Crepidula sp. on the mangrove roots at Punta Morales.

Four species of decapods, distributed two orders, two families and four genera, were found associated with three species of polychaetes. The tubes of one species of Onuphidae, one species of Terebellidae and the tube of an unidentified family were inspected in the intertidal zone of Punta Morales, Gulf of Nicoya. The Pinnotheridae crab, Glassella costaricana was found associated with the polychaete Lanicola sp. The species Pinnixa longipes and Polyonyx quadriungulatus were found in the tube of the onuphids.

Nine species of decapods, distributed in two orders, four families and nine genera, were found associated with 12 species of echinoderms. These species were observed living as epibionts in four species of echinoids and five species of asteroids (Table 1). On the sea star Asteropsis carinifera, three species were found: Pachycheles biocellatus, Zenopontonia soror and Calyptraeotheres sp., while in the sea star Pentaceraster cumingi and the sea urchin Diadema mexicanum, several individual symbionts of both sexes and in different stages of development were found in a single host individual. Symbiotic decapods were found both in solitary and in aggregated echinoderms, such as Astropyga pulvinata and D. mexicanum in reef sites in Bahía Culebra, the sea star Nidorellia armata in rocky reefs close to Playa Rajada, Bahía Salinas, the sea star Pentaceraster cumingi on soft bottoms near reefs in Golfo Dulce. Aggregates of P. cumingi are common in the rhodolith beds of Isla del Coco, however, no decapods were found associated.

Only one species of decapod, A. pusilla, was found associated with two species of ascideans. In 1970, Ascidonia pusilla was collected from specimens of the recently described ascidian Rhopalaea birkelandi from Playas del Coco, Bahía Culebra (Fujino 1972), and was described as Pontonia spighti. A specimen of A. pusilla was found in association with an unidentified sea squirt (Ascidiacea) from Isla Bolaños, northern Pacific Costa Rica.

In total, 28 associations of 24 species of decapods, in five ordens and 17 families, were found at Isla del Coco. Of the species found, 10 belong to the infraorder Brachyura and 10 to the infraorder Caridea, associated with nine orders distributed among four phyla (Tables 1 and 2).

In Costa Rica, few studies have focused on symbiotic decapods, with the majority carried out in the Pacific and only one in the Caribbean (Azofeifa-Solano et al. 2014). Most of these studies were focused on reproductive aspects of decapod guests. Fifty percent of the studies deal with pea crabs (Pinnotheridae), 35% are about the shrimp family Palaemoniidae, and the remaining 15% are derived from studies of the diversity of organisms associated with the coral P. damicornis (Cabrera-Peña and Solano-López 1996; Cabrera-Peña et al. 2001; Alvarado and Vargas-Castillo 2012; Azofeifa-Solano et al. 2014; Mena et al. 2014; Salas-Moya et al. 2014). There is a need for more detailed studies of decapod crustaceans associated with macroinvertebrates. 2b1af7f3a8