Rev. Biol. Trop., 44(2): 603-617,1996

Habitats and biodiversity of decapod in the SE Gulf of California, México

Michel E. Hendrickx Estación Mazatlán, Instituto de Ciencias del Mar y Limnología, UNAM. Apdo. Postal 811, Mazatlán 82000 Sinaloa, México.

(Rec. 24-1-1995. Rev. 24-V-1995. Accep. 2-VI-1995)

Abstract: Samples of benthic macro-fauna were obtained in different habitats along and off the coast of Southern Sinaloa, Gulf of California, , from 1978 to 1991. Qccurrence of species of decapod crnstaceanswas registered for six habitats, fromthe intertidal to depth of 1200 m. A total of 299 species were collected, belonging to 53 farnilies and including 17 species of Penaeoidea, 45 of Caridea, 6 of Thalassinidea, 5 of Palinura, 1 of Astacidea, 63 of Anomura, and 162 of Brachyura. Number of species varied considerably from one habitat to another. Highest biodiver­ sity was observed in the Bay of Mazatlán, with 121 species, followed by the continental shelf and the rocky intertidal (107 species each), the estuarine/coasta1 lagoons (48 species), the upper slope (18 species) and the sandy beaches (9 species). Qne species was found to be strictly insular-terrestrialand two are primarilyassociated with the flotsam. The results of this survey were compared with distribution data available for decapod crnstaceans fauna from the SE Gulf of California and the Eastern Tropical Pacific zoogeographic region (ETP). The fauna collected represents 82% of the species cited for the area for coastal and shallow subtidal habitats (to ca. 115 m depth) and 57.6% of deep-water (> 200 m) species known to occur in the Gulf of California. Except in two cases, similarity indices (SI) based on number of species common to any pair of habitats were all very low. Continental shelf and the Bay of Mazatlán have 57 species

in cornmon (SI = 0.50), while rocky shore habitat and the Bay of Mazatlán share 27 species (SI = 0.24). Comparative studies of decapod crustaceans cornmunities for the ETP arealmost lacking alltogether. Available data,however, indi­ cate that biodiversity observed in Southern Sinaloa is so far the highest on record for marine and brackish-water habi­ tats for a given section of this tropical zoogeographic region.

Key words: Decapod crustaceans, biodiversity, habitats, Gulf of California.

Subtropical and tropical marine inverte­ are found in the data". The same authors (op. brates communities are largely undescribed. cit.) emphasized the urgent need for an increase This is due fundamentally to the complexity of of comprehensive surveys and inventories species-rich habitats found in tropical seas as which would require, as a basic tool, closer well as to a lack of trained scientists and insti­ cooperation between major taxonomic institu­ tutional infrastructure in many countries bor­ tions and training of a large number of paratax­ dering them. Consequently, present knowledge onomists to collect and document specimens. of many tropical marine communities is far Taxonomic lists and biodiversity studies in spe­ from being adequate, and it might never be so cific habitats serve as points of departure for considering their present rate of alteration by (among others) studying the structure of food the increased frequency and intensity of anthro­ chains, the relative abundance of species, and pogenic damage (Hatcher et al. 1989), As stat­ number of species or total number of organisms ed by McNeely et al. (1990), "As more taxo­ of variousphysical sizes (May 1992). Although nomic and survey work is done [ ... ] more new invertebrate groups often include small and discoveries are made ... [and] ... more new gaps obscure species, they must be counted along 604 REVISTA DE BIOLOGIA TROPICAL with other living resources requiring conserva­ and sorne species penetrate into the Gulf of tion and careful management (Beattie 1994). California, along one or both coast. The This is of paramount importance when one endemic component of the Gulf, which considers that conservation practice in poorly accounts for 15.3% (Gulf of California proper) known areas of the tropics cannot be organized to 17.5% (Cortez Province, extended to or reinforced without r�liable inventory of Magdalena Bay) of the known species of deca­ resources and human activities (Hatcher et al. pod crustaceans, also represents an important 1989). Two thirds of world population inhabits factor as mai1y endemics have their southern­ coastal areas, and while most tropical countries most distribution limit elose to or at the Gulf of are still developing or testing their strategies of California southern limito Nevertheless, it is resources exploitation, their rapid development now clearly recognized that the SE Gulf of and population growth has already had a signif­ California macrofauna is predominantly influ­ icant effect on resources utilization and a nega­ enced by the highly diverse tropical fauna of tive impact on natural coastal ecosystems Panamic origin (Brusca 1980, van der Heiden (Birkeland 1987, Hatcher et al. 1989). and Findley 1989,Hendrickx 1992a,1993b). The Gulf of California,is now widely recog­ Considering the urgent need to develop a nized as a zoogeographic province of the strategy for conservation of shallow tropical Eastern Pacific tropical region (see Brusca and marine ecosystems from anthropogenic dam­ Wallerstein 1979, Hendrickx 1992a, 1993a). ages, and the necessity to establish a source of Information dealing with its benthic decapod information to be used as a reliable reference of crustaceans fauna has increased dramatically in data in future comparative studies, a long-term the last ten years (see Hendrickx 1993a, sampling program was developed in Southern 1994b). Recent biodiversity studies on this Sinaloa in 1979. Marine and estuarine shallow group of invertebrates indicate that: 1) As water systems were routinely sampled, inelud­ many as 580 species are found in the Cortes ing the continental shelf and the upper slope to Province (Gulf of California and SW tip of about 1200 m depth. This paper presents the Baja California Peninsula); 2) This is the high­ results of this survey, and it is believed to be est biodiversity recognized to date for any pre­ the first to document the repartition of a given viously defined zoogeographic unit in the group of marine invertebrates - the decapod whole eastern tropical region (Mexico to north­ crustaceans - in a multi-habitats tropical coastal ern Peru), where approximately 930 species of area from the West coast of America. decapod crustaceans are known to occur; 3) Southern Sinaloa appears as one of the richest coastal and marine ecosystems, with 350 DESCRIPTION OF THE AREA species (Hendrickx 1993a, 1993b, 1993c); 4) Deep-water (> 200 m) decapod crustaceans in The area designated herein as "Southern the eastern Pacific Ocean number 183 species Sinaloa" extends from Teacapan (22°30'N) to (Wicksten 1989), of which 100 occur within North of the Altata lagoon complex (24°40'N) the limits of the Eastern Tropical Pacific (Fig. 1) along approximately 280 km of coast­ (ETP); furthermore, review of Wicksten's data lineo It is located in the southeastern section of and of precise ranges of these deep-water the Gulf of California and ineludes major habi­ species in the Gulf of California allowed tats such as sandy and rocky shores, coastal Hendrickx (1990) to conclude that only 26 lagoons and estuaries, a large semi-protected species were known to be present off the coast bay (the Bay of Mazatlán) and a gentIy slopping of Sonora-Sinaloa. Seven species were recently soft and mixed bottom continental shelf added to that list (Hendrickx 1995c). (Hendrickx, 1986a, 1992a). The upper slope is From a zoogeographic stand point, the ben­ found further offshore. The sandy shore habitat thic macrofauna of the southern Gulf of dominates the coastal area,with long and narrow California is under the influence of three major stretches of beaches interrupted by rivers mouth faunistic components. The warm temperate or coastal-lagoons inlets. Of these, most corre­ Californian fauna partly extends south of the spond to seasonaIly overflowing coastaI lagoons recognized California Province southern or river heads that dry up from December to boundary (Magdalena Bay, Baja California) May-June and accumulate water during the HENDRICKX: Habitats and biodiversity of decapod crustaceans 605 rainy season. Several major rivers connect to the One of the most interesting ecosystem in the sea all year round. Major coastal lagoons also area is the Bay of MazatIán, a semi-enclosed connect permanently to the sea, sometimes body of water extending over approximately 40 tbrough artificially maintained channels. Smaller km2, with tbree major islands and two emerging coastal lagoons are connected permanentIy to rocks. The shore is mostIy sandy with several the sea via an estuary, but most present a closed rocky points and sorne large extension of rocky inlet that open only seasonally, generally during beaches found next to the hills and on the flooding periods, when accumulated rain water, islands. Below about 5 m depth and irnmediate­ high tides and heavy waves generated by tropi­ ly in front of the sandy beaches, the bottom is cal depressions off the Pacific coast of Mexico, made of coarse and medium sand which is pro­ combine to wash away the sand bar. gressively replaced offshore with finer sand mixed with variable amount of silt and clay (Orozco-Romo 1980). Maximum depths of 14- • OFFSHOAE S'.'IONS

" ::�: SAMPUNO 15 m are found just beyond the islands, although depthsof up to 20 m occur in the nav­ igation channel at the harbor entrance. The study area is under direct influence of the warm tropical Mexican Coastal Current in the summer (May to September). Shallow coastal water experience a considerable decrease in temperature during the winter (October to April) due to the infIuence of southwards flowing currents north of Cape Corrientes and local upwelling produced by northerly winds (Alvaréz-Borrego 1983, Hendrickx et al. 1984, Hendrickx 1995b). As in other tropicallsubtropical areas, there are only two typical seasons in the SE Gulf of California, which are influenced more by wind, 1M aAY Of' MAZAnAN rainfall and current patterns than temperature. el COASTAL LAGOON$

MI ItOCKY SHOftE Onshore water temperatures normally vary SS 'AHOY SHORE between 30-32° C in the surnmer and 18-20°C in the winter. Subtidal epibenthic water temper­ atures in the Gulf of California have been ana­ lyzed by Hendrickx (1992a).The Southeastern Gulf of California presents relatively stable environmental conditions in the winter, with outer (90-110 m) and midshelf (60-75 m) tem­ Rocky points are scarce. The major rocky perature of 13.2 and 15.4°C, and inner shelf shores, either intertidal or submerged, are (30-40 m) temperature reaching 14.2-17.2°C. located in the Bay of MazatIán. There are no Summer bottom water temperatures are much rocky shores south of MazatIán, and there are higher on the inner shelf (26.4-27.2°C) than in only three extensive rocky areas located north deeper w�ter (15.6-21.1oC). Below ca. 120 m of Mazatlán. Tidal amplitude is rather the epibenthic environment becomes hostile reduced (ca. 1.5 m) when compared to other with rapid dissolved oxygen depletion « 0.5 areas of the Gulf of California, where tides of mI O2/1). Anoxic conditions consistentIy occur up to 10 m have been observed. Tidal cur­ on the deeper part of the outer shelf and on the rents are usually weak, although tropical upper slope (Hendrickx 1995b) to ca. 500-750 depression and hurricanes occurring from m, depending on location. From 200 m and July to October off the SW coast of tropical deeper, bottom water temperature decreases Mexico can have dramatic effect on the sta­ steadily from 11 to ca. 4-5°C at 1200 m bility of beaches in the area (Montayo-Ley et (Hendrickx 1992b). Extension of the continen­ al. 1988, Hendrickx 1993b). tal shelf (0-200 m) is notably significant in the 606 REVISTA DE BIOLOGIA TROPICAL

Cortez Province, and it represents 12.3% of the obtained for distinct habitats were compared area covered by sea; comparative value for among themselves using Sorensen Similarity upper slope, to 1000 m, is 12.1%. This corre­ Index, SI = 2C/A+B, where C = number of 2 sponds to a total of ca. 172,000 km of off­ species in common, A and B = number of shore habitat (Hendrickx 1993a). species for habitat A and B (Sorensen 1948) and to existing lists found in the literature.

MATERIAL ANO METHODS RESULTS Since 1979, specimens of decapod crus­ taceans were obtained from sampling opera­ A total of 299 species were collected during tions in southern Sinaloa. Depending upon the the study period (Tables 1, 2). Of these, 287 habitat sampled, specimens were collected with were found in coastal habitats and on the shelf, dredges and grabs,otter trawls,cast nets,traps representing 82% of all decapod crustaceans and baited lines, hand nets and by hand. species cited for the area by Hendrickx (1993b), Intertidal species were mostIy collected by who included species found immediately north hand during low tides and rotenone fish-killer and south of Southern Sinaloa in addition to was occasionally used in rocky pools. In those species that had actually been collected coastal lagoons deepest channels,samples were within the study area boundaries. Slope species obtained with a small 25 x 50 cm bottom occurring in the Gulf of California were com­ dredge while dip-nets were used along channel puted by Hendrickx (1990, 1992b, 1995c) on edges or in dense patches of sea-grass.. Cast the basis of existing literature and new records. nets were also occasionalIy used in open areas Of the 33 known species,19 (57.6%) were actu­ and fish-baited traps were left overnight in ally collected during the study periodo All �ol­ deepest channels. Surf species were caught in lected species have been listed for each habttat depths of 1 to 2 m with a 25 x 50 cm bottom in an appendix at theend of this papero dredge. Soft bottom associated species of the The sandy shore and surf zone habitat: Bay of Mazatlán were captured using 5 and 10 Considering the 9 species cited herein, the 1 Van Veen grabs and a 3.5 m x 4.5 m otter­ sandy shore decapod community is trawl. The continental platform decapod crus­ . the less diversified of all. Except for Coenobaa taceans were collected with a 40 1 Van Veen compressus, which is a terrestrial species of grab, a 2.5 m wide oyster dredge and semi­ hermit-, and Ocypode occidentalis which is commercial or commercial otter trawls (12 to found in the supratidal zone, all species in this 28 m). Upper slope samples were obtained with habitat are dwelling organisms living buried or a modified Agassiz dredge. AlI collected mate­ semi-burled in sand beyond the water line and rial was kept in a 4-8 % formaldehyde solution well-adapted to the wave action; they are com­ for at least 48 hours after collection. Samples monly found in the surf zone and occasionally were then washed with freshwater and pre­ at the lower limit of the intertidal during neep­ served in 70 % ethanol. Preserved specimens tide (Albuneidae and Hippidae). were generally incorporated in the Invertebrates Reference Collection of the The rocky shore habitat: A total of 107 Estación Mazatlán UNAM (see Hendrickx species were collected, including 11 �pecies 1994a). Species are listed acc�rding to the that are not considered common or typlCal for habitats where they were collected: sandy this habitat. As expected, Penaeoids shrimps shore, including the surf zone; rocky shore, are not well represented in the rocky intertidal, from the upper intertidal and down to about 5 and only two species of were collect­ m depth; estuaries and coastal lagoons; 5-20 m ed there. These are caught infrequently in tidal shallow coastal water of the Bay of Mazatlán pool or trapped between rocks at low tide. In soft bottom; continental shelf beyond the 20 m turn, 17 species of Caridean shrimps were depth and to a maximum of 120 m depth; upper found, including 12 species of Alpheidae. The slope, from ca. 200 to 1200 m depth. Species three species of Panulirus collected in the area found indistinctly in two or more habitats were are generally found beyond 5 m depth, and included in corresponding lists. Lists of species occasionally occur in shallower water.· Two of HENDRICKX: Habítats and bíodíversíty ofdecapod crustaceans 607

TABLEI

Numbers ami % ( ) of species of decapod crustaceans collected in the area for each fami/y. Numbers of species cited for the

area ( = lOO %) are provided for comparative purposes ami include both species collected during this stÚdy ami species cited for the area (see text)o (*) This survey

Famíly Number ofspecíes Farnily Number ofspecíes Collected Cited Collected Cited

Benthesicyrnidae I (100) 1* Galatheidae 6 (lOO) 2+4* 7 (78) 9 Porcellanidae 34 (83) 41 Sicyoniidae 7 (87) 8 Drorniidae 2 (67) 3 2 (lOO) 2 Raninidae I (lOO) 1 Pasiphaeidae 1 (100) 1 Dorippidae 3 (100) 3 Palaemonidae 8 (62) 13 8 (89) 9 Gnathophyllidae 1 (100) 1 Leucosiidae 10(77) 13 Alpheidae 18(64) 28 Inachidae 6 (67) 9 Cmngonidae I (100) 1* Inachoididae 7 (100) 7 Glyphocrangonidae 1 (lOO) 1* Tychidae 2 (lOO) 2 Hippolytidae 7 (87) 7+ 1* 4 (80) 5 Nematocarcinidae 1 (lOO) 1* Pisidae 7 (lOO) 7 Ogyrididae I (lOO) 1 1 (lOO) 1 Processidae 3 (75) 3+ 1* Mithmcidae 11(92) 12 Pandalidae 3 (3 3) 8+1* Parthenopidae 7 (78) 9 Nephropidae 1 (lOO) 1* 1 (100) 1 Scyllaridae 1(100) I Atelecyclidae 1 (100) 1* Palinuridae 3 (lOO) 3 Portunidae 12(100) 12 Polychelidae I (lOO) 1 Cancridae 2 (lOO) 2 Axiidae 1 (25) 3 + 1* 34(87) 39 Callianassidae 2 (100) 2 Goneplacidae 9 (90) 10 Upogebiídae 3 (100) 3 3 (lOO) 3 Albuneidae 3 (60) 5 Gmpsidae 13(l00) 13 Hippidae 2 (lOO) 2 Pinnotheridae 9 (64) 14 Coenobitidae 1 (lOO) 1 Ocypodidae 8 (lOO) 8 Diogenidae 14(100) 14 Palicidae 1 (100) 1 Paguridae 3 (3 3) 9

these (P. inflatus and P. gracilis) are subject to TABLE 2 an intensive fishery in the area (pérez-González et al. 1992). Most Porcellanidae occurring in Number of species of decapod crustaceans collected in each habitat. Percentages were obtained considering (A) Southem Sinaloa are associated to the rocky total number of species foumiduring the study (299) ami habitat,and as many as 26 of the 34 species col­ (B) total number of species cited for the area (see text) lected during the study were found in this habitat while only 7 species of hermit- were found

Habitat Number of As % oftotal there, usually in tidal pools. Brachyuran crabs species collected number ofspecies dominate the community: 19 spider crabs (A) (B) (Majoidea),22 Xanthidae and 8 Grapsidae make the bulk of the true crabs community, which also 2.6 Sandy shore 9 3.0 incIudes Aethra scutata and Cronius ruber, the Rocky shore 107 35.8 30.6 Est./Iagoons 48 16.1 13.7 latter an uncommon species for this habitat. Bay ofMazatlan 121 40.5 34.6 Shelf 107 35.8 30.6 The coastal lagoons/estuaries habitat: Upper slope 19 6.3 57.6 Coastal lagoons, estuaries and associated man­ Peculiar 3 1.0 0.9 grove forest of Southem Sinaloa harbour a rel­ atively high variety of species of decapod crus­ taceans (48 in total). In addition to the 4 species of Penaeus known for the area, 5 species of caridean shrimps were collected in these systems. Thalassinids were represented in 608 REVISTA DE BIOLOGlA TROPICAL samples by 4 species, including two undeter­ again, the Brachyura. As expected, it comprises míned Callianassidae. Anomuran crabs num­ mostly subtidal species Call Dorippidae, bered only 4 species,aH very characteristics for Calappidae, Leucosiidae, Portunidae, Parthe­ this habitat. Brachyuran crabs represent the most nopoidea, Goneplacidae, and Pinnotheridae; diversified component (30 species). Most species most Majoidea) and sorne Xanthidae atypical were never encountered in other habitats. for this habitat. Portunids are well-represented, as they ¡nelude 9 of the 15 species known fram Tbe Hay of Mazatlán soft bottom commu­ the Gulf of California. nity: Fourteen species of Penaoid shrimps were collected in the bay, and this represents almost The continental sheif: Although in most 2/3 of the 23 shelf species known from the places the upper slope cannot be clearly detect­ entire Gulf of California. It should be noted, ed before 150-160 m deep, the shelf fauna start however, that the two species of Metapenaeo­ vanishing much earlier. A sharp decrease in psis recorded from the Gulf and Solenoeera oxygen concentrarÍon 1S responsible for this, mutator are unknown or extremely rare in and from 90-110 rn onwards, only a few water shallower than 30 m, and that Sieyonia species adapted to low�oxygen concentratíon affinis and TraehypenaeusJaoe have been cited are to be found (see Hendrickx 1992a). only once off the coast of Sinaloa (Hendrickx Altogether 107 species of decapod crustaceans 1986b, 1995a). Caridean shrimps have been have been sampled. As expected, species typi­ relati vely well studied in southern Sin aloa cally associated with the subtidal environment (Hendrickx et al. 1983, Hendríckx and are well represented; this includes 26 species of Wicksten 1987) and the Bay of Mazatlán soft shrimps (13 Penaeoidea and 13 Caridea) and 9 bottom habitat hosts 22 species. Sorne,howev­ species of hermit crabs, which reach their hígh­ er, are rather associated with submerged isolat­ est diversity for the area. Brachyuran crabs ed rocky structure (e.g. Alpheus armillatus, again dominate the community as far as num­ Synalpheus sanjosei), and should therefore be ber of species (66) is concerned (particularly considered atypical for this habitat. It is note­ the Calappidae, Leucosiídae, Parthenopoidea, worthy that al! three species of Lysmata known Portunidae and Goneplacidae). Sorne families from the en tire Eastern Tropical Pacific occur of decapod also appear for the first time: in the Bay, and that the tiny Ogyrides Pandalidae, Galatheidae, Dromiidae and alphaerostris, unrecorded for this region until Cancridae, wíth 2 species each; Raninidae and 1987, is a dominant species of the soft bottom Palicidae with one species only. community (Hendrickx and Wicksten 1987, Wicksten and Hendrickx 1992). Except for the The upper slope (ca. 200 to 1200 m): In slippery lobster,Evibaeus prineeps, which typi­ addition to 7 species (So/enocera mutator, cally occurs on sandy bottom elose to shore, Sicyonia ingentís, S. pieta, Porcellana haneoeki, lobsters and mud-shrimps (Thalassinidea) are Pleuroneodes planipes, Platymeragaudieha udii, scarce. Anomura (20 species in total) ineludes and Stenocionops ovata) already collected in a small series of medium sized hermit crabs other habitats, 12 species were found exclusive­ species which feature an unbroken distribution ly on the upper slope. Of these, 3 species were pattern from inside the Bay towards the adja­ only recently reported for the area (Hendrickx cent shelf. Porcelain crabs are known to be 1995c). Altogether,these 19 species belong to as mostIy intertidal, sometimes extending their many as 16 families, 5 of which are exelusively habitat to the shallow subtidal where rocky bot­ found below 200 m (, tom and associated fauna and flora (e.g. Nematocarcinidae, Glyphocrangonidae, sponges and coralina) serve as refuges for Nephropidae, and Pylochelidae). many species of Petrolisthes, Paehyeheles or Megalobraehiunl. Exelusively subtidal porce­ lain occurring in the Gulf of California inelude DISCUSSION the genera Orthochela. Eueeramus, Poreellana, Minyoeerus, Ulloaia and Polyonix, all of them Fifty-three families of benthic decapod crus­ represented at least with one species in the Bay taceans (including the Majoidea sensu Drach of Mazatlán. The largest group of species is, and Guinot 1983, and the Pandalidae sensu HENDRICKX: Habitats and biodiversity ofdecapod crustaceans 609 lato, not sensu Christoffersen 1989) are known Hypnaea panosa, Colpomenia) (Sánchez­ to include southern Sinaloa in their distribution Vargas and Hendrickx 1987), which favor set­ range. Specimens of all these families were tlement of gastropods mollusks and majid collected during this study, and as many as 32 crabs. Coastal lagoons and estuaries represent were represented in samples by all known an adverse habitat for marine species. Turbidity species, thus providing a 100 % collecting rate is generaIly very high, water temperature may considering the number of species reported for il,)crease considerably when compare to adja­ the area by Hendrickx (1990, 1993b). For the cent coastal water, and many coastal lagoons rest, collecting rate varied from 25% (Axiidae) experience strong tidal or seasonal salinity to 92% (Mithracidae), with most families (18 variations (Hendrickx 1984, Alvarez del out of 21) scoring at least 60%. Families with Castillo et al. 1992). Hence, species found in few species sometimes present a relatively low this habitat are mostly species adapted to these collecting rate (60-80%), often corresponding environmental conditions, although temporal to a single missing species (e.g. Processidae invaders are occasionaIly spotted close to the and Dromiidae) (Table 1). The number of area connecting to the adjacent sea. Salinity upper slope species previously known for the within the lagoons systems can vary fromalmost area was occasionally equal to "O" (e.g. fresh water to hypersaline depending upon the N ematocarcinidae, G lyphocrangonidae). amount of rain and the exchange rate with adja­ However, for the sake of consistency, species cent sea water (Hendrickx 1984, Flores-Verdugo collected during the present survey were conse­ 1989). Biodiversity on the upper slope drops quently considered as present in the sampling dramaticalIy to 19 species only, although this area, and were therefore added to the number represents 57.6% of species presently known of "known" species. from deep-water offSonora-Sinaloa. As seen earlier, number of species recorded As expected, similarity between communi­ per habitat is highly variable. The minimum ties sampled in distinct habitats is very low. value (9 species) was found for the sandy shore Number of species in common for any given (3.0% of total for this survey and 2.6% of all pair of habitats variesfrom O to 57. When com­ species known for the area). The long, unpro­ paring intertidal with subtidal habitats, species tected beaches are under the infIuence of occurring in more than one habitat are essen­ waves, and occasional offshore atmospheric tialIy species that lives at the threshold of their disturbances results in strong displacement of normal depth range. Thus, several rocky shore sandy beaches that does not favor long term species are extending into the Bay of MazatIán settlement of species. Values for other habitats subtidal habitat, and 27 species are found in are much higher (Table 2), and the highest both habitats (Similarity index S = 0.24). number of species corresponds to the Bay of Highest similarity, however, is found between MazatIán (121 species, 40.5% of total for this the shelf fauna and the Bay of Mazatlán, with survey, 34.6% of all known species for the 57 species in common (S= 0.50) (Table 3). area). Rocky shore and continental shelf score Comparative data for the Bastern Pacific are similar values (both 107 species), while lacking almost all together. The only compre­ coastal-lagoons and estuaries are supporting a hensive, closely related study was published much lower diversity (48 species), as expected. almost twenty years ago by Abele (1976), and Rocky shores found in Southern Sinaloa are dealt with the Pacific (and Caribbean) decapod typical of tropical-subtropical region and they fauna of Pan ama associated with four major offer shelter and food to a high variety of habitats. Data were mostIy related to intertidal species of invertebrates, most of them present fauna. Sandy beach fauna of included and abundant all year-round. Well-protected, 16 species (none of Lepidopa, and 8 unidenti­ semi-protected and unprotected areas are fied species of "Callianassa") vs. only 9 for found. The lower section of the midlitoral and Southern Sinaloa. Twenty species were report­ part of the sublitoral fringe is often occupied by ed for the habitat in Panama, but no colonies of the Zoanthidae Palythoa and Caridea, Penaeus or Callinectes, all predomi­ Zoanthus. Sorne semiprotected areas are also nantIy subtidal species, were included; compar­ colonized by Padina durvillaei and other atively, Southern Sinaloa strictIy "mangrove" species of algae (e.g. Caulerpa sertularoides, fauna (no shrimp, no swimming-crabs) is made 610 REVISTA DE BIOLOGIA TROPICAL

TABLE 3

Numbers of species common ro pair of habirars. Similarity indexfor pair of habitats (0.00 ro 0.50) were obtained using Sorensen index SI (SS = sandy sJwre; CL = coastal lagoons - estuaries; RS = rockyshore; CS = continental shelf; BM = Bay of Mozatlán; SL= slope)

SS eL RS es BM SL

SL 00 00 00 06 02 BM 03 05 27 57 0.02 es 00 04 03 0.50 0.10 RS 00 07 0.03 0.24 0.00 CL 02 0.09 0.05 0.06 0.00 SS 0.08 0.00 0.00 0.04 0.00 of 41 species, which is still a much higher fig­ continental shelf for the en tire Gulf of ure than in Panama. The Pocillopora coral California (58% of all known species). Other community of Panama was made of 53 species, reports are surprisingly scarce. They often ineluding 8 obligate commensals. Pocilloporid appear to be uncompleted due to selective corals is an unknown habitat in Southern methods of sampling (e.g. large mesh-size Sinaloa, and a comparative analysis seems commercial shrimp-nets) or because they repre­ inappropriate. Still, excluding unidentified sent results of short period, isolated surveys. species cited by Abele (11 out of 53; op. cit.), Alvarez- León (1979) reviewed the marine obligate commensals (8 species), and Panamic species of commercial importance from the species not extending to the SE Gulf of Pacific coast of Colombia and Usted 27 species California (5 species), 20 out of the 29 species of decapod crustaceans, mostly captured by the left are also found in other habitats of Southern commercial fishing fleet. Méndez (1981) Sinaloa. The last habitat studied by Abele reported 69 marine and 13 fresh and brackish (1976) was the rocky intertidal, for which he water species shrimps from Peru; 30 species cited 78 species (23 identified only to genus or were associated with the continental shelf,and family,and 12 Panamic species not reaching 39 were slope species. Comparatively, 62 the SE Gulf of California). Southern Sinaloa species of Penaeoidea and Caridea were col­ rocky shore community ineludes 107 species, lected during this survey, out of 88 known ineluding 27 of the 43 properly identified,non­ species for the area (Table 1). During a exelusively Panamic species cited by Abele prospective survey of the Gulf of Tehuantepec,

(op. cit.). Here again, 6 species cited for the Sosa-Hernández et al. (1980) reported only 28 Panamic rocky shore habitat have been found species of decapod crustaceans obtained by in other habitats in Southern Sinaloa. trawling below 25 m depth. One of the most Composition of decapod crustaceans communi­ intensive survey for the ETP was made in the ty from a coastal lagoon of SW Mexico was Gulf of Nicoya, Costa Rica, a tropical estuary analyzed by Alvarez del Castillo et al. (1992) of 10-20 km wide, with bottom water salinity who reported 74 species, ineluding 31 species ranging from 28 to 37%0, and temperature from atypical for this habitat and collected exelu­ 14 to 28°C. Benthic macro-fauna was sampled sively in the rocky intertidal of the lagoon per­ during three cruises, between 8 and 52 m manent inlet. The only consistent analysis of depth, resulting in a total of 48 collected decapod crustaceans associated to sandy beach­ species of decapod crustaceans (Maurer et al. es in the ETP was provided by Dexter (1974, 1984). Comparatively, nearly three times as 1976) who cited 7 species for beaches in many species were collected in the Bay of Mexico,Costa Rica,and Colombia. Mazatlán, the most elosely related habitat for Data related to subtidal soft bottom decapod Southern Sinaloa. More recently, Bianchi crustaceans community in the ETP are not (1991) reported only 11 shelf species of deca­ available, except for Hendrickx (1992a: 6) who pod crustaceans in samples obtained by trawl­ concluded that approximately 308 species of ing (50 to ca. 250 m depth range) from the decapod crustaceans were associated to the Gulf of Tehuantepec, Mexico, to the Gulf of HENDRICKX: Habitats and biodiversity of decapod crustaceans 611

Papagayo, . Although the author nas c?steras (4 8 especies) , la plataforma superior (18 especIes) y las playas arenosas (9 especies). recognized that "Species identification often . Una especie correspondIó estnctamente a un hábitat poses serious problems in tropical areas [and] insular terrestre y dos están primariamente asociadas con el flotsam. Los no guides [ ...] were available for the eastern resultados de este estudio fueron comparados con las central Pacific", it is surprising that large, informaciones de distribución disponibles para la fauna common and easily recognizable species such de crustáceos decápodos del sureste del golfo de California as Hepatus kossmanni, Euphylax robustus or y el Pacífico Este Tropical. La fauna recolec­ tada representa el 82% de las especies citadas para el E. dovii were not reported in this survey. área para los hábitats costeros y poco profundos (hasta Another ¡ist of species available for the conti­ ca. 115 m de profundidad) y 57.6% de las especies de nental shelf of the Eastern Tropical Pacific is aguas profundas (> 200 m) conocidas para el golfo de by Hendrickx and V ázquez-Cureño (1995), California. Exceptuando dos casos, el índex de similitud (SI) basado en el número de especies comunes a who reported 46 specíes of decapod crus­ cualquier par de hábitats fue siempre muy bajo. La taceans collected by trawling during two plataforma continental y la bahía de MazatIán tienen 57 CEEMEX cruises in the Gulf of Tehuantepec, especies en común (SI = 0.50), mientras que el hábitat de from 26 to 180 m depth. la zona rocosa y la bahía de Mazatlán comparten 27 especies (SI = 0.24). Upper slope data are again lacking. Son pocos los estudios compara­ tivos de comunidades de crustáceos decápodos para el Wicksten (1989) cited 183 deep-water species Pacífico Este Tropical. Sin embargo, la información for the eastern Pacific, of which only 26 were disponible, indica que la biodiversidad observada en el known to be present off the coast of Sonora­ sur de Sinaloa es por mucho la más alta registrada para Sinaloa. During this survey, 19 specíes were los hábitats de agua marina y salobre para una sección dada de esta región zoogeográfica tropical. obtained: 9 of the 26 species known to be pre­ sent, 3 additíonal deep-water species not cited earlier for the Gulf (Hendrickx 1995c), and 7 ACKNOWLEDGEMENTS species also found on the shelf. The only avail­ able comparative data related to slope decapod The author thanks aH students, crew rnem­ crustaceans fauna is again by Méndez (1981); bers and colleagues who took an active part of the 69 marine species of shrimps she report­ during sampling activities in the study area. ed from Peru, 39 were slope species. The author acknowledges Enrique MacPher­ On the basis of present survey, it can be son, ICM, Barcelona, and the Dirección Gene­ concluded that: 1) There is a dramatic lack of ral de Investigación Científica y Técnica, Ma­ data related to composition of decapod crus­ drid, Spain, for their help and support during taceans communities in the entire ETP; 2) The his sabbatical stay in the Instituto de Ciencias biodiversity observed in the study area is the del Mar, CSIC, in Barcelona. Final manuscript highest ever reported for the Pacific coast of was prepared with the help of Mercedes Cor­ Mexíco; 3) The environmental features of the dero R. Bay of Mazatlán combined with its rich fauna, alIow us to classify this system as one of the most diversified sizable body of water north REFERENCES of Banderas Bay.

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Hendrickx, M.E., A.M. van der Heiden ano A. Toledano Sánchez-Vargas, D.P. and M .E. Hendrickx. 1987. Granados. 1984. Resultados de las Campañas SIPCO Utilization of algae and sponges by tropical decorating (sur de Sinaloa, Mexico) a bordo del B/O "El Puma". crabs (Majidae) in lhe southeastem Gulf of California. Hidrología y composición de las capturas efectuadas en Rev. Biol. Trop. 35: 161-164. los arrastres. An. ¡nst. Cienc. Mal' y Limnol., UNAM. 11: 107-122. Sorensen, J. 1948. A method of establ¡shing groups of equal amplitud in plant sociology based on similarity Maurer, D., C. Epifanio, H. Dean, S. Howe, J. Vargas, A. Dittel and M. Murillo. 1984. Benthic invertebrates of a of species content. Danske Vidensk. SeIk, Biol. Skr. tropical estuary: Gulf of Nicoya, Costa Rica. J. Nat. 5: 1-34. Hist. 18: 47-61. Sosa-Hernández, P., J.L. Hernández Aguilera and J.L. May, R.M. 1992. How many species inhabit the earth? ViIlalobos-Hiriart 1980. Estudio prospectivo de los Scientific Amer.267: 18-24. crustáceos (D ecapoda y Stomatopoda) del golfo de Tehuantepec, México. Sec. Mar. Dir. Gral. Ocean. lnv. McNeely, J.A., K.R. Miller, W.V. Reíd, R.A. Mittermeier Ocean. B-80-1O: 1-50. and T.B. Werner 1990. The information requíred to conserve biological diversity. In Conserving the van der Heiden, A.M. and L.T. Findley. 1988. Lista de world's biologica! diversity. WRI, el, WWF-US, The peces marinos del Sur de Sinaloa, México. An. ¡nst. World Bank p.71-81. Cienc. del Mar y Limnol., Univ. Na!. Auton. Mexico. 15: 204-224. Mendez G., M. 1981. Claves de identificación y distribu­ ción de los langostinos y camarones (Crustacea: Wicksten, M-K. 1989. Ranges of offshore decapod crus­ Decapoda) del mary ríos de la costa del Perú. Bol. !nst. taceans in the ea�tern Pacific Ocean.Trans. San Diego Mar Perú. 5: 1-170. Soc. Nat. Hist. 21: 291-316.

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Appendix 1

Species of decapod crustaceans collected in dijferent habitats in southern Sinaloa: 299 specíes. Because some species have been collected in more than one habitat. total number of entríes is 413. (*) Not a common species. (**) Sur:[zone, occa­ siona!. Specimens are deposíted in the Reference Col/eetion of the Estación Mazatlán, UNAM

Sandy shore, intertidal (9 species).

Anomura Petrolisthes hirtispínosus Lockington, 1878 * A/bunea /ucasia de Saussure, 1835 Petrolisthes /ewisi (Glassell, 1936) Lepidopa cf. deamae Benedict, 1903 Petrolisthes nobilii Haig, 1960 * Lepidopa esposa Efford, 1971 Petrolisthes ortmanní Nobili, 1901 Lepidopa mearnsi Benedict, 1903 Petrolisthes polymitus Glassell, ! 937 Lepidopa mexicana Efford, 1971 Petrolisthes robsonae Glassell, 1935 *Hippa strigillata (Stimpson, 1860) Petrolisthes sanfelipensis Glassell, 1936 Emerita rathbunaeSchmitt, 1935 Petrolísthes tonsorius Haig, 1960 Coenobita compressus H. Milne Edwards, 1837 Neopisosoma dohenyí Haig, 1960 Brachyura Neopisosoma mexicanum (Streets, 1871) Ocypode occidentalís Stimpson, 1860 Pachycheles ca/culosus Haig, 1960 Pachycheles panamensis Faxon, 1893 Rocky shore, intertidal (107 species). Pac/zyche/es setimanus (Lockington, 1878) Pac/zyeheles spinidactylusHaig, 1957 Penaeoidea Clastotoechus dijfracfus (Haig, 1957) *Sicyonia laevigata (Stimpson, 1871) * Euceramus panatelus Glassell, 1938 * Sicyonia martini Pérez-Farfante & Boothe, 1981 Pisídia magdalenensis (Glassell, 1936) Caridea Megalobrae/zium festai (Nobili, 1901) Gllathophyllum panamense Faxon, 1893 Mega/obrachíum sinuinumus (Lockington, 1878) Alp/zeusleviuscu/us Dana, 1852 Brachyura Alpheus schmitti Chace, 1972 Eucinetops /ucasi Stimpson, 1860 A/p/zeus armilla/us H. Milne Edwards, 1837 Eucinetops rubellula Rathbun, 1923 A/pheus cy/indricus Kingsley, 1878 Inac/zoideslaevis Stimpson, 1860 * Alp/zeus sp/endidus Coutiere, 1897 Pitho sexdentata Bell, 1835 Alpheus sulcatus Kingsley, 1878 Pitho pieteti (de Saussure, 1853) Alpheus mal/eator Dana. 1852 Acanthonyx petiveri Milne Edwards, 1834 Synalpheus digueti Coutiere, 1909 Epialtus minimus Lockington, 1877 Syna/p/zeussanjosei Coutiere, 1909 Epia/tus sulcirostris Stimpson, 1860 Synalpheus biunguiculatus (Stimpson, 1860) Eupleurodon trifurcatus Stimpson, 1871 Syna/pheus lockingtoni Coutiere, 1909 Pelia pacifica A. Milne Edwards, 1875 Synalpheus nobí/U Coutiere, 1909 Pelia tumida (Lockington, 1877) *Palaemon rilteri Holmes, 1895 Herbstia camptacantha (Stimpson, 1860) Thor algicola Wicksten, 1987 Herbstia tumida (Stimpson, 1871) Lysmata galapagensis Schmitt, 1924 Ala comuta (Stimpson, 1860) *Brachycarpus biunguicu/atus (Lucas, 1849) Mithrax armatus de Saussure, 1853 Palinura Mithrax spinipes (Bell, 1835) Panulirusgracilis Streets, 1871 Mithrax denticula/us Bell, 1835 Panulirus injlatus(Bouvier, 1895) Teleophryscristulipes Stimpson, 1860 Panulirus penicil/atus(Olivier, 1791) Microp/zrys platysoma (Stimpson, 1860) Anomura Thoe su/catasulca/a Stimpson, 1860 * Trizopagurus magnificus(Bouvier, 1898) Ae/hrascutata Smith, 1869 Calcinus californiensisBouvier, 1898 *Croniusruber (Lamarck, 1818) C/ibanarius a/bidigitus Nobili, 1901 Liomeracinc/imana (White, 1847) *Clibanarius digueti Bouvier, 1898 Platypodiella rotundata (Stimpson, 1860) Clibanarius panamensis Stimpson, 1859 Cataleptodius occidentalis (Stimpson, 1871) * Paguristes anahuacus Glassell, 1938 Xanthodius sternberghii Stimpson, 1859 Pagurus /epidus (Bouvier, 1898) Xanthodius stimpsoni (A. Milne Edwards, 1860) Petrolisthes agassizzi Faxon, 1893 Cycloxanthops vittatus (Stimpson, 1860) Petrolisthes armatus (Gibbes, 1850) Metopocarcinus truncatus Stimpson, 1860 Petrolisthes crellulatus Lockington, 1878 Panopeus bermudensis (1) Benedict & Rathbun, 1891 Petrolisthes edwardsU (de Saussure, 1853) Panopeus chilensis Milne Edwards & Lucas, 1844 Petrolisthesgracilis Stimpson, 1858 Panopeus mirajloresensis Abele & Kim, 1889 Petrolist/zes/zaigae Chace, 1962 Eurypanopeus confragosus Rathbun, 1933 Petrolisthes hians Nobili, 1901 HENDRICKX: Habitats and biodiversity of decapod crustaceans 615

Eurypanopeus planus (Smith, 1869) *Paehygrapsus transversus (Gibbes, 1850) Micropanope cristimanus Stimpson, 1871 Scleroplax granulata Rathbun, 1892 Menippefronta/is A. Milne Edwards, 1879 Pinnixa valerii Rathbun, 1931 Pilumnus pygmaeus Boone, 1927 Ucides occidentalis (Ortrnann, 1897) Pilumnus reticu/atus Stimpson, 1860 Oeypode occidentalis Stimpson, 1860 *Pilumnus townsendi Rathbun, 1923 Vea prineeps (Smith, 1870) Heteractaealunata (Milne Edwards & Lucas, 1843). Vea latimanus (Rathbun, 1893) Ozius per/atus Stimpson, 1859 Vea musica musica Rathbun, 1914 Epixanthustenuidaetylus (Lockington, 1877) Vca crenulata crenulata (Lockington, 1877) Ozius verreauxii de Saussure, 1853 Vea zacae Crane, 1941 Eriphia squamata Stímpson, 1859 Vca vocator eeuadoriensis Maccagno, 1928 P/agusia immaeulata Larnarck, 1818 Plagusia depressa tubereulata Larnarck, 1818 Percnon gibbesi (H. Milne Edwards, 1853) (1) Probably P. mirajloresensis Abele & Kim, 1989 Grapsus grapsus (Linnaeus, 1758) Geograpsus lívidus (Milne Edwards, 1837) Pachygrapsus tranversus (Gibbes, 1850) Hay of Mazatlan soft bottom, 5-25 m dcpth *Planes eyaneus Dana, 1852 (122 species). Tetragrapsusjouyi (Rathbun, 1893) ( l )?=P. mirajloresensis Abele & Kim, 1989 Penaeoidea Penaeus brevirostris Kingsley, 1878 Coastal lagoons and estuaries (48 species). Penaeus califomiensis Holmes, 1900 Penaeus stylirostris Stimpson, 1871 Penaeoidea Trachypenaeus pacijicus Burkenroad, 1934 Penaeus vannamei Boone, 1931 Trachypenaeus brevisuturae Burkenroad, 1934 Penaeus stylirostris Stímpson, 1871 Xiphopenaeus riveti Bouvier, 1907 * Penaeus californiensis Holmes, 1900 Sieyonia disdorsalis (Burkenroad, 1934) *Penaeus brevirostris Kingsley, 1878 Sicyonia disedwardsii (Burkenroad, 1934) Caridea Sicyonia ingentis (Burkenroad, 1938) Palaemonetes hiltoni Schmitt, 1916 Sicyonia laevigata Stimpson, 1871 Palaemon gracilis (Smith, 1871) Sicyonia martini Pérez-Farfante & Boothe, 198 t Macrobrachium tenellum (Smith, 1871) Sieyonia picta Faxon, 1893 Alpheus mazatlanicus Wicksten, 1983 Solenoceraflorea Burkenroad, 1938 * Synalpheus biunguiculatus (Stimpson, 1860) Caridea Thalassinidea Leptochela serratorbita Bate, 1888 Callianassidae sp. A Automate doliehognatha de Man, 1938 Callíanassidae sp. B Automafe rugosa Coutiere, 1900 Vpogebia dawsoni Williams, 1986 Salmoneus serratidigitus Coutiere, 1896 Vpogebia thistlei Williams, 1986 * Alpheus armillatus H. Milne Edwards, 1837 Anomura Alpheusjloridanus Kingsley, 1878 Coenobita compressus H. Milne Edwards, 1859 *Synalpheus sanjosei Coutiere, 1909 Clibanarius albidigitus Nobili, 1901 * Synalpheus digueti Coutiere, 1909 Clíbanarius panamensis Stimpson, 1859 Neopontonides denfiger Holthuis, /951 Petrolisthes robsonae Glassell, 1945 Periclimenes infraspinis (Rathbun, 1902) Petrolisthes lindae GOfe & Abele, 1973 Ogyrides alphaerostris (Kingsley, 1880) Brachyura Lysmata californica(S timpson, 1866) Callinectes arcuatus Ofdway, 1863 Lysmata galapagensis Schmitt, 1924 Callineetes bellicosus (Stimpson, 1859) Lysmata intermedia (Kingsley, 1880) Callinectes toxotes Ordway, 1863 Latreufes antiborealis Holthuis, 1952 Panopeus bermudensis (1) Benedict & Rathbun, 1891 Trachycarisrestricta (A. Milne Edwards, 1878) Panopeus ehilensis Milne Edwards & Lucas, 1844 Thor algieola Wicksten, 1987 Panopeus purpureus Lockington, 1876 Brachycarpus biunguiculatus (Lucas, 1849) Panopeus mirafloresensis Abele & Kim, 1989 Pontonia nUlrgarita Smith, 1869 Hexapanopeus sinaloensis Rathbun, 1930 Processaperuviana Wicksten, 1983 Eurytium albidigitus Rathbun, 1933 Proeessa sp. Eurytium affi ne Streets & Kingsley, 1876 Ambidexter panamensis Abele, 1972 Cyrtoplaxschmitti Rathbun, 1935 Thalassinidea Malacoplax ealiforniensis (Lockington, 1877) Callianassidae sp. crassum Smith, 1870 Vpogebia jonesi WiIliams. 1986 quadratus de Saussure, 1853 Palinura Armases magdalenense Rathbun, 1918 Evibacus princeps Smith, 1866 Sesarma rhizophorae Rathbun, 1906 * Panulirus gracilis Streets, 1871 Sesarma sulcatum Smith, 1870 * Panulirus injlatus (Bouvier, 1895) Aratus pisoni (Milne Edwards, 1837) Anomura Goniopsis pulchra(Lockington, 1876) Albunea lucasia de Saussure, 1835 616 REVISTA DE BlOLOGIA TROPICAL

Lepidopa esposa Efford, 1971 Edwardsium lobipes (Rathbun, 1898) Lepidopa mearnsi Benedict, 1903 *Xanthodius stimpsoni (A. Milne Edwards, 1860) Dardanus sinistripes (Stimpson, 1859) * Panopeus bermudellsis (1) Benedict & Rathbun, 1891 Aniculus elegalls Stimpson, 1859 Hexapanopeus orcufti Rathbun. 1930 Paguristes sp. Microcassiope xanrusi (Stimpson, 1871) Paguristes digueli Bouvier, 1892 Pilumnus towllsendi Rathbun, 1869 Pagu/'lts g/adius (Benedict, 1892) *Heteractaea [¡mata (Milne Edwards & Lucas, 1843) Orthochela pumita Glassell, 1936 Euryplax polita Smíth, 1870 *Petrolisthes agassizii Faxon, 1893 Speocarcinus granulil11allus Rathbull, 1893 "Petrolisthes crenlllatlls Lockington, 1878 Oediplax granulata Rathbun, 1893 Petrolisthes sllnt'elipensis Glassell, 1936 Pinnixa sp. Euceramuspallafe/us Glassell, 1938 Pinnixa ajjinis Rathbun. 1898 Euceramlls trallsversilineatus (Lockington, 1878) Pinnixapemberto¡¡i Glassell. 1935 Minyocerus kirki Glassell, 1938 Pinnixa (l sclunilli Rathbun, 1918 Porcellana wncrisocialis Glassell, 1936 Pinnixa transversalis (H. Milne Edwards Pisidill magdalenensis (Glassell, 1936) & Lucas, 1844) * Megalobrachium fe stai (Nobili, 1901) Calyptraeotlieresgranri Glassell, 1933 UI/oaillperpusil/ia Glassell, 1935 ( l) Probab1y P. miraj/oresensis Abele & Kim, 1989 Polyonix quadriungulatus Gla-;sell, 1935 Brachyura

Ethusa panamensis Finnegan, 193 1 Continental shelf, 25 • 115 ro depth (107 species). Hel'atus kossl1lanni'Neumann, 1878 Hepatella amica Smith, 1869 Penaeoidea Osachí/a acuta Stimpson, 1871 Penaeus brevirostris Kingsley, 1878 Iliacal!!ha hancocki Rathbun, 1935 Penaeus califomiensis Holmes, 1900 Persel'!uma townsendi (Rathbun. 1893) Penaeus vannamei Boone, 1931 LellcosiUajurinei (de Saussure, 1853) Trachypenaeus paClji'c:us Burkenroad, 1934 Randallia agaricias Rathbun, 1898 Xiphopenaeus riveti Bouvier, 1907 RandaUia americana (Rathbun, 1893) Sicyollia aliajJinis (Burkenroad, 1934) Randallia bulfigera Rathbun, 1898 Sicyonia disdursalis (Burkenroad, 1934) Col/odes ¡;ibbosus (Bell, 1835) Sicyonia disedwardsii (Burkenroad, 1934) Col/odes tenuirostris Rathbun, 1893 Sicyonia ingentis (Burkenroad, 1938) Pyromaia tuberculata (Lockington, 1877) Sicyonia martilli Pérez-Farfante & Boothe, 1981 Erileptus spinosus Rathbun, 1893 Sicyonia picla Faxon, 1893 lnachoides laevís Stimpson, 1860 Solenocera florea Burkenroad, 1938 Podochela latiman!!s (Rathbun, 1893) So/enocera mUlator Burkenroad, 1938 Podochela veleronis Gmth, 1958 Caridea Stenorhynchus debilis (Smith, 1871) Auromatedo/iclwgnatha de Man, 1938 Pi/ho pie/eli (de Saussure, 1853) Automate rugosa COlltiere, 1900 *Acanthonyx petiveri Milne Edwards, 1834 Salnwlleus serratidi¡;itus Comiere, 1896 Epialtus ¡nÍlúmus Lockington, 1877 Alpheus tloridanus Kingsley, 1878 * Ep ialtus su/cirostris Stimpson, 1860 Alpheopsis cortesiana Wicksten & Hendrickx, 1986 Pelia pacifica A. Milne Edwards, 1875 Pantomus qtfinis Chace, 1937 Neodoclea boneti Buitendijk, 1950 Plesionik(j mexicana Chace, 1937 Notolopas lamellatus Stimpson, 1871 Trachycaris restricta (A. Milnc Edwards, 1878) Notolopas mexicanus Garth, 1940 Lysmata califomica (Stimpson, 1866) *Maiopsis pUl/amensis Faxon, 1893 Lysmata intermedia (Kingsley, 1880) Mithrax armatus de Saussure, 1853 Latreutes anti[¡orealis Holthuis, 1952 Milhraxdenticularus Bell, 1835 Processa perul'iana Wicksten, ¡ 983 Microphrys platysoma (Stimpson, 1860) Processa sp. Hemus sp. Palinura Hemusfinneganae Garth, 1958 Evibacus princeps Smith, 1866 Leio/amb¡'uspUllctatissimus (Owen, 1839) Anomura Hererocrypta macrobrachia Stimpson, 1871 Petrochirus calijíir/úensis(St impson, 1859) Mesorhoea belli (Milne Edwards, 1878) Dardanus sinistripes (Stimpson, (859) Callinecfes arcumus Ordway, 1863 Anicu[us e/egans Stimpson, 1859 Portunus acuminatus (Stimpson, 1871) Paguristes sp. Portunus asper (A. Mílne Edwards, 1861) Paguristes bakeri Holmes, 1900 Portunus iridescens (Rathbun, 1893) Pagllristes digueti Bouvier, 1892 Portunus xantusii xantusíí (Stimpson, 1860) Paguristes praedilfor GlasseU, í 937 Arenaeus mexicanus (Gerstaecker, 1856) Pagurus gladius (Benedíct, 1892) Ellphylax dovii Stimpson, 1860 Pagurus smithi (Benedict. 1892) Euphylax robustus A. Milne Edwards, 1861 Pleuroncodes planipes Stímpson. 1860 Cronius ruber (Lamarck, 1818) MUllida mexicanu (Benedict, 1903) Actaea angusla Rathbun, 1898 Porcellalla cancnsocialis Glassell, [936 HENDRICKX: Habitats and biodiversity of decapod crustaceans 617

Porcellana hancocki Glassell, 1937 Edwardsium lobipes (Rathbun, 1898) Polyonix nitidus Lockington, 1878 Hexapanoeus orcutti Rathbun, 1930 Brachyura Microeassiope xantusi (Stirnpson, 1871) Cryptodromiopsis larraburei (Rathbun, 1910) Pilumnus fownsendi Rathbun, 1869 Hypoconcha panamensis Smith & Verrill, 1869 Quadrella nitida Smith, 1869 Raninoides benedicfi Rathbun, 1935 Trizocarcinus dentatus Rathbun, 1893 Efhusa ciliatifrons Faxon, 1893 Euryplax polita Srnith. 1870 Ethusa lata Rathbun, 1893 Panoplax mundata Glassell, 1935 Calappa convexa de Saussure, 1853 Sp eocarcinus granulimanus Ratlibun, 1893 Calappa saussurei Rathbun, 1898 Oediplax granulata Rathbun, 1893 Platymera gaudichaudii Milne Edwards, 1837 Chasmocarcinus lafipes Rathbun, 1898 Cycloes bairdi Stimpson, 1860 Chasmophora macrophfalma (Rathbun, 1898) Hepafus kossmanni Neumann, 1878 Chacellus pacificus Hendrickx, 1989 Osachila lata Faxon, 1893 Pinnixa sp. Lithadia cumingii Bell, 1855 Pinnixa transversalis (H. Milne Edwards Ebalia crisfafa Rathbun, 1898 & Lucas, 1844) Iliacantha hancocki Rathbun, 1935 Palieus fragilis (Rathbun. 1893) Iliacantha schmiffi Rathbun, 1935 Persephona edwardsii Bell, 1855 Persephona cf orbicularis Rathbun, 1893 Upper slope, ca. 200 to 1200 rn depth (19 species). Persephona townsendi (Rathbun, 1893) Randallia americana (Rathbun, 1893) Penaeoidea Randallia bulligera Rathbun, 1898 Benthesicymus fanneri Faxon. 1893 'Euprognafha bifida Rathbun, 1893 Sicyonia ingentis (Burkenroad, 1938) Collodes gibbosus (Bell, 1835) Sicyonia picfa Faxon, 1893 Col/odes granosus Stimpson, 1860 Solenocera mufatorB urkenroad, 1938 Collodes tenuirosfris Rathbun, 1893 Caridea Paradasigyius depressus (Bell, 1835) Acanfephyra brevicarinara Hanamura, 1989 Pyromaia fuberculafa (Lockington, 1877) Nemafocareinus agassizií Faxon, 1893 Erilepfus spinosus Rathbun, 1893 Lebbeus scrippsi Wicksten & Mendez, 198: Inachoides laevis Stimpson, 1860 Heferocarpus alfinis Faxon, 1893 Podochela larimanus (Rathbun, 1893) Paraerangon areolara Faxon, 1893 Stenorhynchus debilis (Smith, 1871) Glyphocrangon spinulosa Faxon, 1893 Stenocionops ovara (Bell, 1835) Astacidea Mithrax tuberculafus Stimpson, 1860 Nephropsis occidenralis Faxon, 1893 Parthenope hyponca (Stimpson, 1871) Palinura Parfhenope exilipes (Rathbun, 1893) SfereonUlsfis nana (S.Srnith, 1884) Parthenope excavara (Stirnpson, 1871) Anomura Solenolambrus arcuafUS Stimpson, 1871 Porcellana hancocki Glassell, 1938 Leiolambrus puncfafissimus (Owen, 1839) Munidopsis depressa Faxon. 1893 Heterocrypta macrobrachia Stimpson, 1871 Munidopsis diomedeae (Faxon. 1893) Mesorhoea belli (Milne Edwards, 1878) Pleuroncodes planipes Stimpson, 1860 Callinectes arcuafus Ordway, 1863 Brachyura Portunus acuminafus (Stirnpson, 1871) Platymera gaudiehaudii Milne Edwards. 1837 Porfunus asper (A. Milne Edwards, 1861) Trachycarcinus corallinus Faxon, 1893 Portunus iridescens (Rathbun, 1893) Sfenocionops ovara (Bell, 1835) Portunus xanfusií affinis (Faxon, 1893) Portunus xantusii xantusii (Stirnpson, 1860) Peculiar habitat (3 species). Euphylax robustus A. Milne Edwards, 1861 Cancer amphioefus Rathbun, 1898 Brachyura Caneer johngarfhi Carvacho, 1989 Gecarcinus planafus Stimpson, 1870 (Insular) Actaea angusta Rathbun, 1898 Plagusia immaculara Lamarck. 1818 (Flotsam) *Medaeus spinulife r (Rathbun, 1898) Planes cyaneus Dana. 1852 (F1otsam)