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JOURNALOFCRUSTACEANBIOLOGY,27(2):351–369,2007 FRESHWATER SHRIMP OF THE GENUS MACROBRACHIUM (DECAPODA: PALAEMONIDAE) FROM THE BAJA CALIFORNIA PENINSULA, ME´XICO Luis Herna´ndez, Gopal Murugan, Gorgonio Ruiz-Campos, and Alejandro M. Maeda-Mart´ınez (LH,GM, AMM-M)Centro deInvestigaciones Biolo´gicas del NoroesteS.C., MarBermejo 195, Col. PlayaPalodeSantaRita, LaPaz, Baja California Sur,C.P. 23090,Me´xico ([email protected]; [email protected]); (GR-C) FacultaddeCiencias, UniversidadAuto´noma deBajaCalifornia Ensenada,BajaCalifornia, C.P. 22800,Me´xico ([email protected]) (correspondingauthor (AMM-M): [email protected]) ABSTRACT FreshwaterdecapodslikethepalaemonidrivershrimphavereceivedlittleattentionintheBajaCaliforniaPeninsula,Me´xico.Fromthe firstformalreportin1878tonow,onlythreeMacrobrachiumspecieshavebeenreported(M.americanum,M.digueti,andM.tenellum) fromthepeninsulainonlythreebasins.Wemadeataxonomicstudyoffreshwatershrimp,whichincludedanextensivefieldsurveyat81 sitesdistributedonboththePacificandtheGulfofCaliforniaslopesandarevisionofmaterialfromtheareathathadbeendepositedin scientificcollections.WereportsixspeciesofMacrobrachiumbyaddingM.hobbsi,M.michoacanus,andM.olfersii.Wealsodiscuss someaspectsofthediversityandconservationofthesespeciesinthepeninsulaandprovideanidentificationkeyfortheMacrobrachium shrimpofnorthwesternMe´xico. INTRODUCTION wide geographic distribution, and commercial importance (Holthuis, 1952; Villalobos, 1982). The species of Macro- TheBajaCaliforniaPeninsulainnorthwesternMe´xicoisthe brachium have a primary tropical distribution (Hedgpeth, second longest and believed to be the most geographically 1949;Jayachandran,2001).IntheAmericancontinent,they isolatedpeninsulaintheworld(DurhamandAllison,1960). have been reported from the Illinois River basin, USA SurroundedbytheGulfofCalifornia(SeaofCorte´s)andthe (Bowles et al., 2000) to Argentina (Rodr´ıguez De La Cruz, Pacific Ocean, it extends about 1500 km from the Me´xico- 1965;Acun˜a,2002)andfromtheGulfofCaliforniatoPeru´ U.S.A. border in the north to Cabo San Lucas in the south. (Holthuis, 1952; Wicksten and Hendrickx, 2003). Its average width is about 70 km. The peninsula is The genus Macrobrachium is distinguished from other politically divided into two Mexican states, Baja California palaemonid genera by the following morphological traits; and Baja California Sur, with the interstate border at the 288N parallel. The peninsula has a wide range of climatic carapace with a projecting rostrum, mandibles with molar conditions, though arid conditions are predominant. The process furnished with a triarticular palp, first pair of general physiography is characterized by outstanding pereiopods chelate and slender and as long as the carapace, mountainous chains, which extend along the peninsula, secondpairchelateandofteninmaleslongerthantheentire withnumerousbasinsandcoastalplainsonboththeGulfof body, posterior three pairs of pereiopods simple, telson California and the Pacific slopes. There are no permanent triangular, terminating in asingle tip (Bate, 1868; Holthuis, rivers,butfreshwatersprings,intermittentcreeks,andsmall 1952; Mossolini and Bueno, 2003). They also have hepatic water bodies occur forming unique oasis ecosystems and antennal spines and two pairs of spines on the dorsal (Arriaga and Rodr´ıguez-Estrella, 1997). surface of telson (Hedgpeth, 1949; Holthuis, 1952). Mayaetal.(1997)recognized184oasesinthepeninsula, Holthuis (1952) concluded that only a few characteristics butonly77wererecordedtohavesurfacewater.Besidesthe are available for identification of the species of Macro- presenceoffreshwater(surfaceorinterstitialsoilwater),the brachiumandthatfemalesoftendifferstronglyfrommales. oases are characterized by the assemblage of species Villalobos (1982) proposed that the best identification representingbiogeographicrelictsofsubtropicalmesophilic characteristics are the shape of the rostrum and the chelae communities surrounded by the typical vegetation of the of the male’s second pair of pereiopods. Sonoran Desert (Grismer, 1994). The vegetation and fauna, The presence of freshwater shrimp in the Baja California such as arachnids, amphibians, reptiles, birds, and mam- Peninsulawasfirstannotatedin1780bytheJesuitpriestDel mals,havebeenstudiedinrepresentativeoases(Arriagaand Barco (Leo´n-Portilla, 1988). However, the first formal Rodr´ıguez-Estrella,1997).However,freshwatercrustaceans record was given by Lockington (1878), who reported like the palemonid river shrimp have received little Macrobrachiumtenellum(Smith,1871)(citedasPalaemon attention. longipes) from the oasis of Mulege´. From the same oasis Palaemonids are among the most abundant and diverse Bouvier (1895) also reported M. americanum Bate, 1868 shrimpthat inhabit seawater andfreshwater inboth tropical (citedasPalemonjamaicensis)andM.tenellum(citedasP. and temperate waters (Wicksten, 1983). The genus Macro- forceps) and described a new species, M. digueti (Bouvier, brachium is well known because of the number of species, 1895) (cited as Palaemon digueti). Holthuis (1952), 351 352 JOURNALOFCRUSTACEANBIOLOGY,VOL.27,NO.2,2007 Rodr´ıguez De La Cruz (1968), R´ıos (1989), Wicksten and Hendrickx(1992, 2003),andHendrickx(1994),listed from the Baja California Peninsula the three species already recorded by Bouvier (1895). Wemadeataxonomicstudythatincludedanextensivefield samplingonboththePacificandtheGulfofCaliforniaslopes and a revision of voucher specimens from the area that had beendepositedinscientificcollections.Inthisworkwereport sixspeciesofthegenusMacrobrachiumforthepeninsulaand discuss some aspects on the diversity and conservation of thesespeciesintheregion.Wealsoprovideanidentification keyforMacrobrachiumofnorthwesternMe´xico. MATERIALSAND METHODS Sampling Shrimpwerecapturedusingdifferentfishinggear(handnet,castingnet,gill net,andminnowtrapsbaitedwithfishmeat).Inmostofthesamplingsites, water characteristics such as temperature, total dissolved solids (TDS) (Hachmodel44600),andpH(ORION230A)weremeasuredatthetimeof sampling. Thegeographicpositionof eachsamplingsite wasdetermined using a GPS (Garmin 12XL). The specimens collected were placed in plasticbags,anaesthesizedwithice,andfixedwith100%ethanol. StudyArea TheBajaCaliforniaPeninsulawasseparatedfromthemainlandbytectonic movementsbetweenthePacificandtheNorthAmericanplatesalongthe SanAndreasfault,withthegradualseparationoccurringoverthelast4to5 million years (Stock and Hodges, 1989). Today the northern part of the peninsulaisdominatedbyhighgraniticmountains(SierraJua´rezandSierra SanPedroMa´rtir)andthecentralregionischaracterizedbyrepeatinglayers of volcano-clastic sandstones and conglomerates (Sierra de La Giganta). Graniticrocksandhigherelevations(SierradeLaLaguna)appearagainin thesouthernCaperegion(DurhamandAllison,1960).TheBajaCalifornia Peninsula has a heterogeneous array of landscape and vegetation, from coniferous and tropical deciduous forest in the mountains to xeric desert scrubinlowaridplains(Riddleetal.,2000).Onthewesternsideofthe peninsulathetopographicaltitudedecreasesgraduallyfromthemountains tothecoasttoextensivesedimentaryalluvialplains,whereasontheeastern sidethedistancefromthemountaintothecoastisshorter,oftenwithabrupt escarpments(Grismer,2002).Becauseofitshistoricgeologywithcomplex tectonicmovementsincludingupliftsandsubmergences,plustheecological transformations produced by the geographic isolation and desertification, thepeninsulanowhasapeculiarenvironmentaldiversitywiththeclimate being characterized by relatively high annual mean temperatures (19 to 228C)andalowannualrainfall(100to300mm)(Grismer,2002). Atotalof81sitesweresampled(seeAppendix)with71inthestateofBaja CaliforniaSur(Fig.1)anddistributedalong20basinsandfourhydrological regions.TheremainingsevensitesareinthestateofBajaCaliforniaand distributedalongfourbasinsandtwohydrologicalregions. IdentificationofMaterial Anatomical nomenclature used in this work is according to McLaughlin (1980).Additionaltermsaredefinedasfollows(Fig.2): Fixedfinger:Nonjointedprojectionofthepropodus(manusorpalm)ofthe cheliped. Gapingfingers:Cuttingedgesoffixedfingeranddactylusarearched,thus thereisaclearspacebetweenthem. Closedfingers:Cuttingedgesoffixedfingeranddactylusarestraight,thus thereisnoaclearspacebetweenthem. Spine:A stout,sharpprocess,foundmostlyonthecarapace,pereiopods, chelae,andtelson. Spinules:Slendersmallspines,foundmostlyonthepereiopodsandchelae. Tubercle:Smallroundedprominence,foundmostlyonthepereiopodsand chelae. Fig. 1. Distribution of Macrobrachium species in the state of Baja Pubescence:Smallandnumerousseta-likestructures. California Sur, Me´xico (Southern Baja California Peninsula). Macro- brachium americanum (triangle), M. digueti (eight tip-star), M. hobbsi ‹ (square),M.michoacanus(fivetip-star),M.olfersii(invertedtriangle),and sampling sites with no shrimp found. The thick line indicates the M. tenellum (rhombus). Black dots indicate approximate locations of separationofthePacificandGulfofCaliforniaslopes. HERNA´NDEZETAL.:MACROBRACHIUMFROMBAJACALIFORNIA,ME´XICO 353 Fig.2. SchematicrepresentationoftheappendagewiththelargestchelaofthesecondpairofpereiopodsofthegenusMacrobrachiumshowingthemain structuresusefulforspeciesidentification(modifiedfromVillalobosandNates,1990). Teeth: As described by McLaughlin (1980), but also as conspicuous andnumberofmalesandfemaleswithtotallengthmeasurements(whenthe protuberances found sometimes along the cutting edge of one or both specimen was complete). For comparative purposes, specimens of three fingersofthechela. Macrobrachium species not found in the Baja California Peninsula were Denticles:Smallteethfoundalongthecuttingedgeofoneorbothfingersof alsoexaminedandincludedinthesectionsoftheirgeminatedspecies. thechela. Chelaeunequal:Rightandleftchelaaredifferentinsizeand-orinshape. SYSTEMATICS Chelaesubequal:Rightandleftchelaaresimilarinsizeandshape. Palaemonidae Rafinesque, 1815 Macrobrachium Bate, 1868 Inthelaboratory,thematerialwassortedaccordingtogender.Maleswere Macrobrachium americanum Bate, 1868 distinguished by the presence of the appendix masculina on the (Fig. 3) secondpleopods.Thefollowingmeasurementswereobtained:totallength (TL)(fromtipofrostrumtoposteriorendoftelson)andlengthofmerus, Macrobrachium americanum Bate, 1868; Holthuis, 1952; Rodr´ıguez De carpus,palm(lengthandheight),anddactylusofthelargerchelaofthesecond La Cruz, 1968; Wicksten, 1983, 1989; R´ıos, 1989; Wicksten and pairofpereiopods.Thenumberofteethonbothmarginsoftherostrumwas Hendrickx,1992,2003;Villalobos-Hiriartetal.,1993;Hendrickx, recorded.Adatabasewithallmorphometricandmeristicdatawasdeposited 1994. in the Crustacea collection at Centro de Investigaciones Biolo´gicas del PalemonjamaicensisHerbst;Bouvier,1895. Noroeste,S.C.,LaPaz,BajaCaliforniaSur,Me´xico(CIB).Specimenswere Type Locality.—Lago Amatitla´n, Guatemala. identified using the keys for the species of Macrobrachium proposed by Holthuis(1952)andWicksten(1989)andtheoriginaldescriptionsofthe Diagnosis.—Rostrum strong, arched over orbital margin, species.AllcollectedmaterialwasdepositedintheCrustaceacollectionat CIB. The material of Macrobrachium was also revised in the scientific curved upward at tip; it reaches the end joint of antennular collections at Universidad Nacional Auto´noma de Me´xico, Coleccio´n peduncle;dorsalmarginwith10to12teethofwhichthreeto Nacional de Crusta´ceos; Instituto de Biolog´ıa, Me´xico City (CNCR); four are placed behind orbital margin; ventral margin with Coleccio´n de Crusta´ceos, Instituto de Ciencias del Mar y Limnolog´ıa, two to four teeth. Second pair of pereiopods with subequal Mazatla´n(EMU);UniversidadAuto´nomadeBajaCalifornia,Facultadde Ciencias,Coleccio´ndeCrusta´ceos(UABC);andthecrustaceancollectionof chelae and gaping fingers; carpus almost twice as long as theMuse´umNationald’HistoireNaturelle,Paris(MNHN). high and shorter than merus. Whole chela covered with spinules.Palmelongated,fingersslightlyshorterthanpalm. PresentationoftheTaxonomicAccount Fingers with single strong tooth on cutting edge; tooth of The taxonomic account includes: 1) species name, author, and year of fixedfingeronfirstthirdofcuttingedge,dactyluswithtooth description;2)synonymy,restrictedtonamesusedforMexicanspecimens inthemiddle;twotofourproximaldenticlesonbothfingers. (for a complete synonymy see Holthuis, 1952); 3) type locality; 4) diagnosis;5)distributionintheBajaCaliforniaPeninsula,includingdataon Distribution in the Baja California Peninsula.—This water characteristics and on the co-occurrence of the species with other common species was previously reported from Mulege´, congenericforms;6)distributioninMe´xico(followinganorth-southorder); La Paz, and Cabo San Lucas (Bouvier, 1895; Holthuis, 7)generaldistribution;8)remarks;and9)materialexamined,indicatingthe country,state, site,collectiondate,seniorcollector’sname,catalogcode, 1952). We found it on the Pacific slope in the basins 354 JOURNALOFCRUSTACEANBIOLOGY,VOL.27,NO.2,2007 Fig.3. AdultmalesofMacrobrachiumamericanumBate,1868.A,Rostruminrightlateralview;B,Anteriorregionindorsalview;C,Largestchelaofthe secondpairofpereiopodsininnerview.AllfiguresfromCIB814.Scalebars¼10mm. Santa Rita, Las Pocitas, Todos Santos, and Plutarco El´ıas 1.5 g/L, pH 6.7 to 8.3, and temperatures 23.9 to Calles, and on the Gulf of California slope in the basins 34.58C. Macrobrachium americanum was found often of Mulege´, El Coyote, San Bartolo, and San Jose´ del co-ocurring with the other five species found in the Cabo. Specimens were found in waters with TDS 0.34 to peninsula. An assemblage of four species (M. americanum, HERNA´NDEZETAL.:MACROBRACHIUMFROMBAJACALIFORNIA,ME´XICO 355 M. digueti, M. hobbsi, and M. michoacanus) was recorded However Chirichigno etal. (1982) only mentioned that this at the Los Potreros site. species may be located in the fishery zone 77A, which comprises an enormous area from Baja California to Distribution in Me´xico.—BAJA CALIFORNIA SUR (see Oaxaca, but did not give any particular data from Isla above). SONORA: Guaymas (Holthuis, 1952), R´ıo Yaqui, Cedros. Records of this species from Mexican islands are and R´ıo Mayo (Rodr´ıguez De La Cruz, 1968). SINALOA: those from Isla Mar´ıa Magdalena and Isla Mar´ıa Cleofas, El Rosario, R´ıo El Fuerte, R´ıo Presidio, R´ıo Quelite Nayarit (Holthuis, 1952; Herna´ndez and Mart´ınez, 1992). (Holthuis, 1952; Rodr´ıguez De La Cruz, 1968; Hendrickx, 1994), and Mazatla´n. NAYARIT: Isla Mar´ıa Magdalena, Material Examined.—Macrobrachium americanum: ME´X- and Isla Mar´ıa Cleofas (Holthuis, 1952; Herna´ndez and ICO:BAJACALIFORNIASUR:Mulege´ dam,28.02.2004, Mart´ınez, 1992), R´ıo Santiago, and Jalcocota´n. JALISCO: A.Maeda,CIB803,1female;Guadalupedam,C.Me´ndez, R´ıo Santiago, and R´ıo Ameca (Holthuis, 1952), Chamela, CIB 860, 2 specimens; Las Paredes, 30.05.2004, L. Cuitzmala, and Puerto Vallarta. COLIMA: R´ıo Armer´ıa Herna´ndez, CIB 804, 1 male (139.5 mm); San Juanito (Holthuis, 1952). MICHOACA´N: La Villita, and Mexcalti- Nuevo, 22.09.2004, L. Herna´ndez, CIB 866, 1 specimen; tla´n.GUERRERO:Bah´ıaPetatla´n(Holthuis,1952),andR´ıo Rancho Tres Pozas, 30.05.2004, L. Herna´ndez, CIB 805, 2 Murga. OAXACA: Pochutla, Salina Cruz, R´ıo Valdeflores, males(134and246mm);Merecuaco,16.05.1998,G.Ruiz- Tuxtepec, Mixtequita dam, and Tehuantepec. CHIAPAS: Campos, UABC 003, 1 male (115 mm); Corral de Piedra, Mal Paso dam, and R´ıo El Naranjo. 29.05.04, L. Herna´ndez, CIB 806, 1 male (121.3 mm); El Caracol,29.05.2004,L.Herna´ndez,CIB807,1male(151.1 General Distribution.—Baja California Peninsula to Peru´, mm) and 1 female (151 mm); Poza de La Matanza, Cocos Island, and Gala´pagos Islands (Holthuis, 1952; 29.05.2004, L. Herna´ndez, CIB 808, 1 male (152 mm); Wicksten and Hendrickx, 1992, 2003). Santa Fe, 26.02.2004, L. Herna´ndez, CIB 867, 1 specimen; Remarks.—Similar to Macrobrachium carcinus (Linnaeus, Las Vinoramas, 21.11.2003, A. Maeda, CIB 809, 1 female 1758), M. americanum is one of the largest species of the (59.2mm);LasVinoramas,03.12.2003,L.Herna´ndez,CIB genus and its size may help to distinguish it from other 810,2males(63and69.8mm);TodosSantos,26.02.2004, species(Holthuis, 1952). Thelargestspecimen inourstudy L. Herna´ndez, CIB 811, 1 male (145 mm); La Poza, was found at Rancho Tres Pozas (Santa Rita basin) at 246- 10.07.2005, L. Mercier, CIB 861; 1 specimen; Agua mmTL.SmallspecimensmaybeconfusedwithmalesofM. Caliente, 13.12.2003, A. Maeda, CIB 812, 1 female (70 occidentale or M. heterochirus (Wiegmann, 1836), but M. mm);RanchoSanAntonio,08.09.2004,L.Herna´ndez,CIB americanumcanbedistinguishedbyitssubequalchelae,the 813, 1 female (129.4 mm); Los Potreros, 06.10.2002, A. form and size of the carpus, and by the rostral length. The Maeda, CIB 814, 1 female (144.5 mm, ovigerous); Los Atlantic geminated species of M. americanum is the big Potreros 24.07.2003, L. Herna´ndez, CIB 815, 2 females claw river shrimp M. carcinus (Holthuis, 1952) that has (109.6 and 112 mm, ovigerous); Poza de Santa Rosa, been reported from the Florida to Brazil (Bowles et al., 14.12.2003, A. Maeda, CIB 816, 2 females (131.7 and 68 2000). Differences among specimens is not easy to detect, mm);SanJose´ delCaboestuary,26.11.2004,L.Herna´ndez, however Holthuis (1952) noted that M. carcinus is sligthly CIB862,1specimen.SINALOA:Mazatla´n,10.09.1973,D. larger than M. americanum and that the carpus length is Pela´ez, CNCR 199, 1 specimen; EMU 0268, 3 specimens. more than twice as long as high in M. carcinus, whereas in NAYARIT: R´ıo Santiago, 07.07.1991, C. Rosales, CNCR M. americanum the same characteristic is not more than 11499, 1 specimen; Jalcocota´n, 13.09.2003, L. Herna´ndez, twice as long as high. In our revision of 86 voucher CIB 853, 5 males (104 to 185.5 mm), 1 female (118 mm), specimens, we noted that proportions on the carpus as and 1 ovigerous female (132.9 mm). JALISCO: Puerto previously established by Holthuis are consistent, so this Vallarta, 17.09.1973, A. Jime´nez, CNCR 202, 1 specimen; characteristicmayhelpustoseparatebothspecies.Colorin Chamela, 31.08.1992,CNCR1619,1 specimen. COLIMA: life for M. carcinus is dark brown with lighter mottling on R´ıo Armer´ıa, 12.09.1973, CNCR 200, 2 specimens. the sides and tan stripes laterally. Chelipeds are dark green MICHOACA´N: La Villita, 28.09.1973, A. Villalobos, tobluewithdarktuberclesandorangearticulations(Bowles CNCR 205, 2 specimens; La Villita, CNCR 13685, 1 male et al., 2000). In M. americanum the body is yellow-brown (190 mm). GUERRERO: R´ıo Murga, 08.05.1984, A. withtanstripeslaterallyfromthecarapacetotelsonandthe Villalobos, CNCR 2624, 5 juveniles; R´ıo Murga, periopodswithbluearticulations.Inadults,thefingersofthe 21.05.1984; A. Villalobos, CNCR 2663, 3 specimens; R´ıo secondpairofchelipedsaredarkwithsomelittlespecimens Murga,23.09.1986;J.C.Nates,CNCR3116,2males(118.4 showing the terminal third of the fingers in white. and 95 mm), 1 female (ovigerous). OAXACA: R´ıo After the Holthuis (1952) review, there was only one Valdeflores,24.05.1961,A.Villalobos,CNCR198,1spec- recordofM.americanumforthepeninsulareportedbyR´ıos imen; Salina Cruz, 11.09.1973, I. Larios, CNCR 201, 1 (1989), who found it at Mulege´. The new records from the specimen;Tuxtepec,13.09.1973,A.Villalobos,CNCR204, Pacific slope basins such as Santa Rita, Las Pocitas, Los 4 specimens; Mixtequita dam, 21.12.1955, A. Villalobos, Potreros, and Todos Santos extend the geographical CNCR 208, 2 specimens. CHIAPAS: Mal Paso dam, distribution of the species to the west side of the peninsula. 12.09.1973, C. Beutelspacher, CNCR 203, 4 specimens; The collections from the Santa Rita basin represent its R´ıoElNaranjo, 06.09.2002,E.Soto,CNCR22021,1male northernmost records in the Pacific slope. A´lvarez-Ruiz (69.8 mm). Macrobrachium carcinus: ME´XICO: MNHN et al. (1996) cited Chirichigno et al. (1982) as reporting 1222, 1 specimen (172 mm); MNHN 989, 1 specimen; M. americanum from Isla Cedros, Baja California, Me´xico. CNCR 13332, 2 males (118.4 and 140.1 mm); CNCR 356 JOURNALOFCRUSTACEANBIOLOGY,VOL.27,NO.2,2007 Fig.4. AdultmalesofMacrobrachiumdigueti(Bouvier,1895).A,Rostruminleftlateralview;B,Anteriorregionindorsalview;C-D,Largestchelaofthe secondpairofpereiopodsinlateralview.AandCfromCIB801,BfromCIB817,DfromMNHN1235,andEfromBouvier(1895).Scalebars¼10mm. 13667, 2 females (133.9 and 141.4 mm); CNCR 13676, 1 Macrobrachium digueti (Bouvier, 1895) female(143.9mm);CNCR16494,1male(61mm);CNCR (Fig. 4) 17174, 1 female (110.6 mm); CNCR 17380, 2 females PalaemondiguetiBouvier,1895. (151.3 and 159.4 mm); CNCR 17401, 1 male (228.5 mm); Macrobrachium digueti (Bouvier, 1895); Holthuis, 1952; Rodr´ıguez De La Cruz, 1968; Wicksten, 1983, 1989; R´ıos, 1989; Wicksten and CNCR 17403, 1 male (216.3 mm); CNCR 18648, 3 males Hendrickx,1992,2003;Villalobos-Hiriartetal.,1993;Hendrickx, (125.7 to 206.2 mm) and 1 male (174.5 mm). 1994;Roma´netal.,2000. HERNA´NDEZETAL.:MACROBRACHIUMFROMBAJACALIFORNIA,ME´XICO 357 Macrobrachium acanthochirus Villalobos, 1967; Villalobos, 1968; Villa- doigts be´ants, enfin on n’observe pas de longs poils lobosandNates,1990;Wicksten,1989;WickstenandHendrickx, duveteux sur la face externe de cette pince et le e´pines de 1992,2003;Villalobos-Hiriartetal.,1993;Roma´netal.,2000. la face interne sont moins nombreuses que dans le P. spinimanus et plus irre´gulie`rement dispose´es’’ (its second Type Locality.—Mulege´, Baja California Sur. large chela is shorter, higher, and completely without large setae between the fingers, finally we do not observe large Diagnosis.—Rostrum straight, reaching last joint of anten- setaeovertheexternalfaceofthechelaandthespinesofthe nularpeduncle;dorsalmarginwith13to16teeth,fourtosix internal face are less numerous than in P. spinimanus and placed behind orbital margin; ventral margin with three to more irregularly disposed). According to Holthuis (1952), five teeth. Second pair of pereiopods with unequal chelae Palemon spinimanus is a synonym of Macrobrachium and gaping fingers; carpus and merus globose at middle faustinum(DeSassure,1857).L.Herna´ndezinParisrevised section. Largest chela with palm almost as long as high, a lot (MNHN 1235) labeled as Syntype of P. Digueti, with spines on dorsal margin and external side. Palm with Mulege´, 1895. This lot contained 16 specimens (39.5 to scarce pubescence, without setae. Fixed finger with up to 56.4 mm), most of them incomplete. One incomplete four teeth on proximal half of cutting edge. Dactylus with appendage still having carpus, propodus, and dactylus a stout tooth, and up to four denticles on proximal part of seems to correspond to that described and figured by cutting edge. Bouvier (1895) (Fig. 4E). Bouvier’s measurements were Distribution in the Baja California Peninsula.—Macro- -longueur de la pince 38 mm; largeur maximum 16 mm; brachium digueti, previously collected from Mulege´, La longueur de doigt mobile 22 mm- (length of the chela Paz, and Cabo San Lucas (Bouvier, 1895; Holthuis, 1952; 38 mm, maximum height 16 mm; length of the mobile Hendrickx,1994),wasfoundonlyattwosites,oneinBoca finger 22 mm). The chela examined by L. Herna´ndez had de la Sierra, Santiago basin in the Gulf of California slope similarvalueswithacarpuslengthof12.7mm,palmlength andtheotherinLosPotreros,PlutarcoEl´ıasCallesbasinon 20.1 mm, and dactylus length 20.3 mm. The remaining 16 thePacificslope.SpecimenswerefoundinwaterswithTDS specimens were determined by L. Herna´ndez as Macro- 0.18 to 1.5 g/L, pH 7.8 to 8.8, and temperature 25.3 to brachium hobbsi Nates and Villalobos, 1990. 33.78C. We found this species co-occurring with three Macrobrachium acanthochirus Villalobos, 1967 was congeners at Los Potreros. Previously, Bouvier (1895) described from two lots, one from specimens collected in reported this species along with M. americanum and M. Tecoma´n, Colima and the other from R´ıo Valdeflores, tenellum. Oaxaca.Villalobos(1968)wasnotsureaboutthevalidityof this species, even proposing that it could be better assigned Distribution in Me´xico.—BAJA CALIFORNIA SUR as a subspecies of M. digueti. Upon the morphological (see above). SONORA: R´ıo Yaqui and R´ıo Mayo examination of 28 voucher specimens (see below the (Rodr´ıguez De La Cruz, 1968). SINALOA: Laguna material examined), the similar proportion values of the Caimanero and R´ıo Baluarte (Wicksten and Hendrickx, carpus length/palm length, and palm length/palm high 2003). NAYARIT: Colomo´. JALISCO: Cuitzmala and R´ıo shown by specimens of both nominal taxa (Table 1), the Los Cuartones. COLIMA: Tecoma´n, and Puerto Jua´rez. overlap distribution of both forms along the Pacific slope, MICHOACA´N: Mexcaltitla´n, R´ıo Murga, and La Villita. we concur that M. acanthochirus should remain as GUERRERO: Acapulco (Wicksten and Hendrickx, 2003). a synonym of M. digueti. OAXACA: Valdeflores. Macrobrachium digueti is similar to M. olfersii (Wieg- mann, 1836), however in M. digueti there isno pubescence General Distribution.—Baja California Peninsula to Peru´ but scarce setae on the second large chela, whereas in M. (Holthuis,1952; Wicksten and Hendrickx, 1992, 2003). olfersii the chela characteristically has copious setae and Remarks.—Villalobos (1968) indicated that the description pubescence. Though M. digueti has only a few teeth on the ofthisspeciesgivenbyHolthuis(1952)doesnotcorrespond proximalpartofthecuttingedgeofbothfingers,M.olfersii totheoriginalspeciesofBouvier(1895)andweagreewith hasmorethanfiveteethalongthecuttingedgeofbothfingers. this assessment. Holthuis (1952, plate 26) described the Macrobrachium digueti is rare in the Baja California largest second chela of Macrobrachium digueti as ‘‘The Peninsula. We found no specimens at the type locality, the second legs are very unequal in shape and size in the adult oasis of Mulege´. R´ıos (1989) did not find any specimens at male.Thefingersinthelargerchelaareaslongasthepalm; Mulege´ during four years of monitoring. This suggests that they arecurved andgape.The cutting edges ofboth fingers the species has been extirped from the type locality. Our bear in the extreme proximal part one large tooth, behind collection at Los Potreros represents the first record of the which 1 or 2 much smaller teeth are placed. The cutting speciesonthePacificslopeoftheBajaCaliforniaPeninsula. edgedistallyofthelargeteethisprovidedwithteeth9to12 in number, which are placed up to the tips of the fingers. Material Examined.—Macrobrachium digueti: ME´XICO: Tuftsofhairareimplantedalongthecuttingedges.Therest BAJA CALIFORNIA SUR: Mulege´, 1894, L. Diguet, ofthefingersisnakedandisdenselycoveredwithspinules. MNHN 1235, 1 second largest chela of a male (Syntype); Thepalmhasadistinct,largethicklypubescentareaateach Mulege´, CNCR 10619, 4 specimens; Boca de la Sierra, ofthelateralsurfaces’’.Thesefeaturesdonotcorrespondto 08.09.2004, L. Herna´ndez, CIB 801, 2 males (78.6 and those of the original description given by Bouvier (1895), 70 mm); Los Potreros, 06.10.2002, A. Maeda, CIB 802, 1 who wrote ‘‘sa grande pince est plus courte, plus large et male (79.4 mm); Los Potreros, 06.10.2002, A. Maeda, CIB comple`tement de´pourvue de longues soies raides entre les 817,1female(81.3mm);SanJose´ delCabo,19.09.1946,I. 358 JOURNALOFCRUSTACEANBIOLOGY,VOL.27,NO.2,2007 Table1. MorphometricdataofthelargestchelaofthesecondpairofperiopodsandrostralteethofMacrobrachiumdiguetimalesfromseveralMexican locations.CaL,carpuslength;PL,palmlength;PH,palmhigh;DL,dactyluslength;CL,carapacelength(allinmm). Specimen CaL PL CaL/PL PH PL/PH DL Rostralteeth CL Reference HolotypefromMulege´,BajaCaliforniaSur — — — 16 — 22 14-16 28 Bouvier,1895 SyntypefromMulege´,BajaCaliforniaSur(MNHN1235) 12.7 20.1 0.63 16.2 1.2 20.3 — — thisstudy MalefromTecoma´n,Colima(citedasM.acanthochirus) 11.4 14 0.81 13.2 1.06 16.5 13(5)/4 28.9 Villalobos,1967 MalefromTecoma´n,Colima(CNCR328,labeledas holotypeofM.acanthochirus) 10.4 15.7 0.66 13.3 1.18 16.7 13(5)/4 20 thisstudy MalefromValdeflores,Oaxaca(citedasM.acanthochirus) 10.3 14.5 0.71 12 1.2 15 — 27 Villalobos,1967 MalefromLosPotreros,BajaCaliforniaSur(CIB802) 10.3 12.6 0.79 10.1 1.24 13.4 15(4)/3 27.3 thisstudy MalefromColomo´,Nayarit(CIB866) 14.1 20.6 0.68 18.9 1.08 23.7 14(6)/4 29 thisstudy MalefromR´ıoMurga,Guerrero(CNCR3538,labeledas M.acanthochirus) 10.7 18.1 0.59 17.2 1.08 19.9 14(5)/5 24.4 thisstudy Bonet, CNCR 235, 1 male (77 mm); San Jose´ del Cabo, Distribution in Me´xico.—Endemic to Me´xico from Guer- 10.09.1946,F.Bonet,CNCR236,3specimens.SINALOA: rero to Chiapas and Veracruz (Villalobos and Nates, 1990; R´ıo Baluarte, EMU 0840, 3 females (62 to 74 mm). Wicksten and Hendrickx,2003) andfrom thewest andeast NAYARIT: Colomo´, 13.08.2003 L. Herna´ndez, CIB 866, sidesoftheBajaCaliforniaPeninsula,andinNayarit.BAJA 5 specimens (75 to 91 mm). JALISCO: Cuitzmala, CALIFORNIA SUR (see above). NAYARIT: R´ıo Huayna- 04.02.1983, J.C. Nates, CNCR 2723, 1 male (65.1 mm, mota and Los Salazares. GUERRERO: R´ıo Murga (Villa- labeled as M. acanthochirus); Cuizmala, 20.02.1984, J.C. lobos and Nates, 1990). OAXACA: R´ıo Astuta (Villalobos Nates, CNCR 3121, 9 specimens (40 to 87 mm); R´ıo Los and Nates, 1990). CHIAPAS: R´ıo Lagartero, Arroyo Cuartones, 01.03.1992, A. Novelo, 1 male, CNCR 13372 Ocuilapa, R´ıo Cintalapa, R´ıo Chacamax (Villalobos and (labeled as M. acanthochirus). COLIMA: Tecoma´n, Nates, 1990), R´ıo Grande, and R´ıo Urbina. VERACRUZ: 18.01.1943,F.Bonet,CNCR328,1male(62mm)(labeled R´ıo La Palma (Villalobos and Nates, 1990). as holotype of Macrobrachium acanthochirus Villalobos, Remarks.—Large specimens of Macrobrachium hobbsi 1967); Puerto Jua´rez, 23.06.1986, R. Navarro, 1 female, may be confused with M. olfersii (Wiegmann, 1836). One CNCR 20804 (labeled as M. acanthochirus). MICHOA- CA´N: Mexcaltitla´n, 31.07.1984, J.C. Nates, 12 males, important difference between the two species noted by Villalobos and Nates (1990) is the absence of space in the CNCR 3536 (labeled as M. acanthochirus); R´ıo Murga, cutting edges of both fingers of the second largest chela in 28.07.1984, CNCR 3538 (labeled as Macrobrachium sp.); M. hobbsi. The second pair of chelae are subequal in shape La Villita, 15.12.1994, CNCR 12986, 2 specimens. butnotinsize,whereasinM.olfersiithesecondlargechelae GUERRERO: Acapulco, 13.08.1973, W. Lo´pez, CNCR are unequal. 141, 1 male (59 mm, labeled as M. acanthochirus); Macrobrachiumhobbsiisanamphiamericanspecieswith OAXACA: Valdeflores, 25.05.1962, A. Villalobos, CNCR 331, 2 specimens (labeled as M. acanthochirus). a wide distribution in Baja California Sur and occurs on boththePacificandGulfofCaliforniaslopes.OjodeAgua in the La Pur´ısima basin is the northernmost site of the Macrobrachium hobbsi Nates and Villalobos, 1990 speciesdistribution.Thereisanapparentdistributiongapof (Fig. 5) the species between the Baja California Peninsula and the MacrobrachiumhobbsiNatesandVillalobos,1990;VillalobosandNates, R´ıo Huaynamota, Nayarit. 1990; Villalobos-Hiriart et al., 1993; Wicksten and Hendrickx, 2003. Material Examined.—Macrobrachium hobbsi: ME´XICO: Type Locality.—R´ıo El Naranjo, Chiapas. BAJA CALIFORNIA SUR: Mulege´, 1895, MNHN 1235, 16 specimens (39.5 to 56.4 mm) (labeled as M. Digueti); Diagnosis.—Rostrum straight, reaching last joint of anten- Ojo de Agua, 01.07.2004, G. Ruiz-Campos, UABC 009, 4 nularpeduncle;dorsalmarginwith13to16teeth,fourtosix males (59 to 68 mm) and 9 females (69 to 91 mm); Ojo de placed behind orbital margin; ventral margin with two to Agua,14.07.2004,L.Herna´ndez,CIB818,3males(64.4to four teeth. Second pair of pereiopods with unequal chelae 79.3 mm) and 1 ovigerous female (61.5 mm); San Isidro and closed fingers. Carpus shorter than palm length, dam, 15.07.2004, L. Herna´ndez, CIB 819, 5 males (49.3 to subequal to merus length. Palm with spinules covered with 69.8 mm) and 4 specimens; La Pur´ısima, 01.07.2004, G. pubescence. Palm about two times or more long as high. Ruiz-Campos,UABC008,10specimens(56.2to73.2mm); Adult specimens of the species are small (, 100 mm). La Pur´ısima, 02.07.2004, G. Ruiz-Campos, UABC 006, 3 DistributionintheBajaCaliforniaPeninsula.—Firstrecords males (72.5 to 77.2 mm); San Juanico road, 14.07.2004, L. of the species in the peninsula. It was collected on the Herna´ndez, CIB 820, 1 male (53 mm); San Pedro de la Pacific slope at La Pur´ısima, Santa Rita, Las Pocitas, and Presa,30.05.2004, L.Herna´ndez,CIB821,3females (53.5 Plutarco El´ıas Calles basins, and on the Gulf of California to 68.4 mm) and 8 specimens; San Basilio, 30.05.2004, L. slope in the basin of Mulege´. Specimens were found in Herna´ndez, CIB 822, 3 males (47 to 72.8 mm) and 9 waters with TDS 0.29 to 1.09 g/L, pH 6.7 to 8.9, and specimens; Las Paredes, 30.05.2004, L. Herna´ndez, CIB temperatures 22.1 to 32.58C. We found this species co- 823,8males(46.2to56mm)and51specimensofjuveniles ocurring with the other five species found in the peninsula. and females; Merecuaco, 16.05.1998, G. Ruiz-Campos, HERNA´NDEZETAL.:MACROBRACHIUMFROMBAJACALIFORNIA,ME´XICO 359 UABC 002, 7 specimens (43.6 to 69.2 mm); Paso Iritu´, L. Herna´ndez, CIB 824, 3 males (42.1 to 52.1 mm); El Caracol,29.05.2004,L.Herna´ndez,CIB825,3males(50to 63.2mmTL),and7specimens;ElColorado,02.03.2004,L. Herna´ndez, CIB 826, 1 male (67.2 mm and 6 females; Los Potreros, 06.10.2002, A. Maeda, CIB 827, 7 males, and 2 females (52.3 and 63.6 mm); San Pedro de la Soledad, 25.11.2004, L. Herna´ndez, CIB 828, 1 female. NAYARIT: R´ıo Huaynamota, 03.07.1991, A. Cantu´, CNCR 13357, 1 male (69.2 mm); Los Salazares, 14.08.2003, L. Herna´ndez, CIB 854, 2 males (65.2 and 66.8 mm) and 4 ovigerous females. CHIAPAS: R´ıo Grande, 14.02.1985, J.L. Villalo- bos, CNCR 5605, 3 males (68.9 to 79.7 mm); R´ıo Urbina, 10.06.2002,CNCR13335,1female(54.3mm);ElNaranjo, J.C. Nates, CNCR 2939 (Holotype), 1 male (67.5 mm). Macrobrachium michoacanus Nates and Villalobos, 1990 (Fig. 6) MacrobrachiummichoacanusNatesandVillalobos,1990;Villalobosand Nates, 1990; Villalobos-Hiriart et al., 1993; Wicksten and Hendrickx,2003. Type Locality.—R´ıo Mexcalhuaca´n, Michoaca´n. Diagnosis.—Rostrum straight, reaching last joint of anten- nularpeduncle;dorsalmarginwith14to15teeth,fivetosix behindorbitalmargin;ventralmarginwiththreetofourteeth. Second pair of pereiopods with unequal chelae and closed fingers.Merusabout1.5timeslongerthancarpus.Palmwith spines distributed on lateral surface, scarce pubescence on lateralventralarea,andlessthantwotimesaslongashigh. DistributionintheBajaCaliforniaPeninsula.—Firstrecords of the species for the peninsula. Macrobrachium michoa- canus was collected on the Pacific slope in Las Pocitas and Plutarco El´ıas Calles basins. Specimens were found in waters with TDS 0.71 to 3.56 g/L, pH 7.4 to 9.3, and temperatures 22.1 to 31.38C. This species was co-occurring with the other five congeneric species. Distribution in Me´xico.—Endemic to Me´xico, from Jalisco to Oaxaca (Villalobos and Nates, 1990; Wicksten and Hendrickx, 2003). Also found on the west side of the Baja California Peninsula. BAJA CALIFORNIA SUR (see above). JALISCO: R´ıo Cuitzmala (Villalobos and Nates, 1990), Chamela, and R´ıo Las Aletas. MICHOACA´N: Morelos dam, R´ıo Chucatitla´n, and R´ıo Papoyutla (Villa- lobos and Nates, 1990). GUERRERO: R´ıo Murga, R´ıo Aguas Blancas, Mexcalhuaca´n (Villalobos and Nates, 1990), R´ıo Salitrera, and La Villita. OAXACA: R´ıo Valdeflores (Villalobos and Nates, 1990; Wicksten and Hendrickx, 2003), and R´ıo Galva´n. Fig. 5. Adult males of Macrobrachium hobbsi Nates and Villalobos, Remarks.—This species is similar to M. digueti, but it can 1990.A,Rostruminleftlateralview;B,Anteriorregionindorsalview;C, be distinguished by the shape of the cutting edges of the Largestchelaofthesecondpairofpereiopodsinlateralview.AandBfrom fingerofthesecondmayorchelaasnotedbyVillalobosand CIB824,andCfromCIB828.Scalebars¼10mm. Nates (1990). Macrobrachium michoacanus has closed fingers on the second large chela, whereas M. digueti has the second species has spinules and pubescence on the gaping fingers. The differences between M. michoacanus palm and the palm length is at least two times longer and the other species with closed fingers, M. hobbsi, are than high. in the palm. The first species has spines on the palm and LasCuevasinLasPocitasbasinofthePacificslopeofthe the length of the palm is less than twice the high, whereas BajaCaliforniaPeninsulaisthenorthernmostboundaryofthe 360 JOURNALOFCRUSTACEANBIOLOGY,VOL.27,NO.2,2007 MaterialExamined.—Macrobrachiummichoacanus:ME´X- ICO: BAJA CALIFORNIA SUR: San Basilio, 15.05.1998, G. Ruiz-Campos, UABC 001, 5 males (51.5 to 75.2 mm) and 2 females; Rancho Las Cuevas, 30.05.2004, L. Herna´ndez, CIB 829, 2 males (57.9 and 74.2 mm); Paso Iritu´, 16.05.1998, G. Ruiz-Campos, UABC 004, 9 males (39.7 to 70.3 mm) and 13 females (34 to 65 mm); El Cardalito, 19.11.2003, A. Maeda, CIB 830, 1 ovigerous female (59.4 mm); Los Potreros, 24.07.2003, A. Maeda, CIB 831, 1 male (45.6 mm) and 2 other males. JALISCO: Chamela, 25.07.1998, C. Sa´nchez, CNCR 191, 2 speci- mens; Chamela, 19.06.1991, C. Sa´nchez, CNCR 192, 3 specimens; Chamela 12.09.1991, G. Casas, CNCR 196, 1 specimen; Puerto Vallarta, Las Aletas stream, 17.09.1973, A. Jime´nez, CNCR 195, 1 specimen. GUERRERO: La Villita,28.09.1973,A.Villalobos,CNCR209,3specimens; Mexcalhuaca´n,31.06.1984,J.C.Nates,CNCR3550,1male (56mm)(Holotype);R´ıoSalitrera,Zihuatanejo,22.07.1987, J.P.Gallo,CNCR13292,1male(17.1mm)and1ovigerous female. OAXACA: R´ıo Galva´n, 31.05.1995, J.P. Gallo, CNCR 13293, 1 male (84.7 mm). Macrobrachium olfersii (Wiegmann, 1836) (Fig. 7) PalaemonolfersiiWiegmann,1836. Macrobrachium olfersii (Wiegmann, 1836); Hedgpeth, 1949; Villalobos, 1968; Villalobos-Hiriart et al., 1993; Herna´ndez-Aguilera et al., 1996;WickstenandHendrickx,2003. Macrobrachium olfersi (Wiegmann, 1836); Holthuis, 1952; Villalobos, 1968. Type Locality.—Not assigned. The original material was reported from a Brazilian shore. Diagnosis.—Rostrum straight, reaching third joint of antennular peduncle; dorsal margin with 12 to 16 teeth, three to six placed behind orbital margin; ventral margin with one to five teeth. Second pair of pereiopods with unequalchelaeandgapingfingers;carpusshorterthanpalm lengthandaboutaslongasthemerus;palmabout1.5times long as high, with setae and pubescence on the lateral side; dactylus length slightly shorter than palm length; cutting edges of fingers with numerous denticles and dense hairs. Distribution in the Baja California Peninsula.—Our collec- tions represent the first records of the species for the peninsula. Macrobrachium olfersii is the second amphia- merican species recorded in the peninsula. It was found mainly on the Pacific slope in the basins of La Pur´ısima, SantoDomingo,SantaRita,LasPocitas,TodosSantos,and Plutarco El´ıas Calles. On the Gulf of California slope the Fig. 6. Adult males of Macrobrachium michoacanus Nates and species was found only in the Alfredo V. Bonfil and San Villalobos 1990. A, Rostrum in left lateral view; B, Anterior region in Jose´ delCabobasins.Specimenswerefoundinwaterswith dorsal view; C, Largest chela of the second pair of pereiopods in lateral TDS0.26to0.61g/L,pH8.1to8.4,andtemperatures19.1 view.AfromCIB829,andBandCfromCIB831.Scalebars¼10mm. to 34.88C. This species was co-occurring with other four congeners. speciesdistribution.ThespecieswasnotfoundalongtheGulf Distribution in Me´xico.—BAJA CALIFORNIA SUR (see ofCaliforniaslope.Thereisanapparentgapinthedistribu- above).SINALOA:R´ıoBaluarte(WickstenandHendrickx, tion of the species between the Baja California Peninsula 2003).NAYARIT:LosSalazares,R´ıoSantiago,andArroyo andthenorthernmostrecordatR´ıoCuitzmala,Jalisco. Bogadero. MICHOACA´N: Mexcaltita´n, and San Nicola´s

Description:
such as arachnids, amphibians, reptiles, birds, and mam- mals, have .. rero to Chiapas and Veracruz (Villalobos and Nates, 1990;. Wicksten and
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