Proceedings ofthe United StatesNational MuseumSMITHSONIAN INSTITUTION ? WASHINGTON, D.C.
Volume 115 1964 Number 3490SHRIMPS OF THE GENUS BETAEUSON THE PACIFIC COAST OF NORTH AMERICAWITH DESCRIPTIONS OF THREE NEW SPECIES
By Josephine F. L. Hart
IntroductionShrimps of the genus Betaeus are members of the section Carideaand the family Alpheidae (or Crangonidae in much of the NorthAmerican literature). Members of this genus are characterized bythe lack of a rostrum in the adult and by the inversion of the "hands,"with the result that the dactyls are on the lower side. The terms
"visored shrimps" and "hooded shrimps" are sometimes used becausethe carapace projects forward to overhang the eyes. Up to the presenttime four species have been recognized as occurring in the area fromMexico to Alaska: Betaeus harrimani Rathbun, B. longidactylusLockington, B. ensenadensis GlasseU, and B. harfordi (Kingsley).However, these species have not been well defined.The sole published record of Betaeus harrimani is the originaldescription by M. J. Rathbun (1904) based on a single female takenin southern Alaska. The examination of a series of both males andfemales from a number of more southerly locations has made it possibleto elaborate on this description. Such an elaboration is especially431693-433?63 1
432 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115important because of the overlap of the range of the closely alhed formB. longidactylus Lockington (1877) in California. These two species,as well as B. ensenadensis Glassell (1938) of southern CaUfornia andMexico, are figured and redescribed here in some detail.In the 1870's W. N. Lockington and J. S. Kingsley each describedtwo species of hooded shrimps from California. They engaged inconsiderable controversy in their papers as to whether or not Betaeuswas a valid genus, and occasionally they referred species of Betaeusto the genus Alpheus; as a result, in the literature both Alpheus andBetaeus are used in referring to these shrimps. Lockington (1877b,1878) described two species briefly, without figures, and, unfortunately,named one Betaeus equimanus, a name that had been used previouslyby Dana (1852, p. 560; 1855, pi. 35) for a shrimp from New Zealand.Early in 1878 Kingsley published a description of apparently thesame shrimp and called it Alpheus harfordi but indicated that if
"aequimanu^'^ should prove to be different, the shrimp would have tobe called ''aequalis.'' Later in the same year Lockington called itB. egualis and enlarged his description by incorporating Kingsley'sdescription of B. harfordi. He was convinced that they were identicalbut because of his use of a preoccupied name, he lost the honor ofnaming the species. Betaeus harfordi (Kingsley) is therefore thevalid name.Holmes (1900) gave a description with more details than thosegiven by the early writers, but unfortunately he added new habitatsthat perhaps have helped to mask the presence of four distinct forms:two commensal and two free-living.In 1934 several hooded shrimps were taken in tide pools on the westcoast of Vancouver Island by E. G. Hart. These agreed mth pub-lished descriptions of B. harfordi except for a few minor differences.Since shrimps of the family Alpheidae often show considerable vari-ation in relative proportions according to age and sex, no suspicion ofsignificant differences arose until individuals from California werecompared with the northern specimens. As a result of a detailedcomparative study, it now seems that those found commensal withinthe mantle cavity of abalones are referable to B. harfordi and thatthe remainder represent three undescribed species. The similar, butquite distinct, form that lives in association with sea urchins I proposeto call B. macginitieae in recognition of Nettie MacGinitie, who, withher husband Professor G. E. MacGinitie, has contributed so much toour knowledge of the genus. For the two free-hving forms I havechosen the names B. gracilis and B. setosus.In view of the fact that the published descriptions and figures ofthe known species are inadequate, an attempt is made herewith tosupplement them with more detailed descriptions and illustrations.
SHRIMPS OF THE GENUS BETAEUS?HART 433The figures are drawn to three magnifications : scales A, B (two timesA), C (ten times A). The whole animal, the chelipeds, and chelaeare scale A; the dactyli, scale C; and the remainder, scale B. Newkeys and comparative tables incorporating the new species are pre-sented. In this aspect the work of L. B. Holthuis (1952) on twospecies found in Chile has been very useful.In classification, emphasis in the past has been placed on the pro-portions of the segments of the antennular peduncle, and on the size,shape, and dentition of the large chelae. These features, however,have been found to differ with age, sex, and extent of regeneration.In this paper, therefore, an attempt is made to use as diagnosis thosecharacters that do not vary appreciably during the life of theindividual.Sexual dimorphism also is noted in this account. Differences dueto sex are most obvious in large males of B. longidactylus, in whichthe chelipeds are much larger in relative proportion to the rest of thebody than in smaller males or in females. All the other speciesdealt with here also show sexual dimorphism but only to a minorextent such as stouter appendages and narrower pleura on theabdomens of the males. AcknowledgmentsMy thanks are due to Dr. Waldo L. Schmitt and Dr. Fenner A.Chace, Jr., of the United States National Museum for the loan ofcomparative material, for making comparisons with the type of B.harrimani, and for much other help. In addition. Dr. Chace read themanuscript and gave invaluable advice. The late Dr. Belle A. Stevenswas most helpful in the earl}^ stages of the study. I wish also to thankboth Dr. Paul L. Illg of the Department of Zoology, University ofWashington, and Mrs. G. E. MacGinitie, retired from KerckhoffMarine Laboratory, for much help and encouragement, as well asfor specimens. Thanks also are due to Mr. Keith W. Cox, CaliforniaDepartment of Fish and Game, who has given me specimens andmuch useful data on Betaeus harfordi obtained in connection withhis work on Haliotis.For the use of material I am indebted to the following: Mr. T. H.Butler and the Fisheries Research Board of Canada; Dr. Joel W.Hedgpeth (Pacific Marine Station) ; Dr. Rolf Bolin and Dr. DonaldP. Abbott (Hopkins Marine Station) ; Dr. Olga Hartman (Universityof Southern California, Allan Hancock Foundation) for the loan ofthe late S. A. Glassell's collection; Dr. James E. Lynch (Universityof Washington) for collections by Dr. John S. Laurie; and Dr. R. U.Gooding of the University of Washington at the time.
434 PROCEEDINGS OF THE NATIONAL MUSEUM vol. usI am also indebted to Mr. C. B. Rowland, Victoria, British Colum-bia, whose patience and ingenuity resulted in successful photographsof the living shrimps. I am grateful also to the Provincial Museumof British Columbia for the use of certain facilities and to the NationalScience Foundation for a grant providing equipment and aid in thecompletion of the study. Finally, I want to thank my husband, Dr.G. Clifford Carl, for his help in so many ways.In addition to the above material I have examined nearly 200specimens (mainly B. longidactylus) from the collection of the AUanHancock Foundation. I hope to report on this material in a futm-epaper. Family AlpheidaeRostrum, if present, unarmed. Eyes usually covered by carapace.Mandibles with incisor process and palp of two segments. Firstpair of legs often with one chela, or both chelae, powerfully developed.Second pair of legs minutely chelate, long, slender, equal, with seg-mented carpus. Telson usually broad, rounded.Genus Betaeus DanaRostrum not present; front not spined, either emarginate betweeneyes or evenly rounded. Chelae usually similar, inverted so thatdactyls are on lower side. Telson broad. Sixth abdominal somitewith movable plate articulated at posterolateral angle. Epipods onat least first two pairs of leg's.Key to Species of Betaeus from West Coast of North America1. Dactyli of walking legs slender and simple 2Dactyli of walking legs stout and bifid 42. Chelae of first legs with fingers longer than palm.B. longidactylus LockingtonChelae of first legs with fingers not longer than palm 33. Blade of antennal scale broad distally. Fixed finger of first cheliped de-creasing in width evenly to sharp curved tip . . B. harrimani RathbunBlade of antennal scale narrow distally. Fixed finger of first cheliped truncatebefore sharp curved tip B. ensenadensis Glassell4. Front curved, not emarginate B. macginitieae, new speciesFront emarginate 55. Emargination shallow. Telson with posterolateral spines small or missing.B. harfordi (Kingsley)Emargination deep. Telson with posterolateral spines well developed . . 66. Peduncle of antennule less than one-half carapace length. Merus of chelipedMath lower inner ridge with long bristles, upper ridge ending in sharp tooth;chela with fingers subequal to palm; chela is three times as long as wide.B. gracilis, new speciesPeduncle of antennule subequal to carapace length. Merus of chelipedwith lower inner ridge usually tuberculate, upper ridge with tuft of hairs;chela Math fingers longer than palm; chela is twice as long as wide.B. setosiis, new species
SHRIMPS OF THE GENUS BETAEUS?^HART 435Betaeus harrimani RathbunFigures 1-26, 29-31, 37-39; Plate 1Betaeus harrimani Rathbun, 1904, pp. 108-110, fig. 49 (type locality, Sitka,Alaska; holotype female, USNM 25692).?Banner, 1953, p. 5.Female.?Carapace laterally compressed but without carina.Smooth, with minute setae sparsely scattered except where concen-trated ventral to hepatic region, where they form a pubesence (seenbest in cast skins of large individuals). Longer setae under frontand along posterior margin. Front (fig. 17) slightly curved anddepressed anteriorly. Anterior margin (fig. 18) with two shallowsinuses. No distinct anterolateral angle. Width of carapace in-creases to midlateral point and then decreases slightly with faintundulations on margin. Lateral margin joins posterior in smoothcurve. Posterior margin angles in sharply to deep cardiac notch.Abdomen (fig. 1) smoothly rounded with posteroventral marginof pleura of first to third segments rounded, those of fourth and fifthsegments angled. First and fifth pleura margined with setae ven-trally. Minute setae on smooth surface.Telson (fig. 26) longer than sixth abdominal segment, much longerthan wide, outer margins curved. Two pairs of movable spines ondorsal surface, two spines at each posterolateral angle; outer onessmall. Posterior margin deeply curved, plumose setae long.Eye round with short, sharp-pointed tooth on each stout eyestalkmedially.Antennule with stylocerite broad at base; outer margin slightlyconvex, inner concave, tip reaching almost to end of second segmentof peduncle. First segment of peduncle with sharp compressedtooth ventrally (fig. 2). Second joint usually longer than third.Outer flagellum with sensory part fused except for about three seg-ments that are free from slender terminal flagellum.Antenna with peduncle slightly longer than that of antennule.Distal margin of basis produced into sharp-pointed flat tooth ventrally.Scale (fig. 19) broad: wide spine exceeding blade, separated fromit by short slit distally, outer margin straight. Tip reaching pastmiddle of last segment of antennular peduncle. Flagellum longerthan carapace, ovoid in cross section.Mandibles, maxillules, maxillae, first and second maxillipeds(figs. 3-7) similar to those of Betaeus truncatus Dana, as described byHolthuis (1952), but appearing more setose, as illustrated.Third maxilliped (fig. 8) reaching to end of antennal peduncle.Ischiomeropodite flattened, twisted. Exopodite longer than ischio-meropodite but narrow, fragile-looking, with soft plumose hairsdistally and striations on cuticle that can be observed in cast skins.
436 PROCEEDINGS OF THE NATIONAL MUSEUM
SHRIMPS OF THE GENUS BETAEUS?^HART 437
438 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115Penultimate segment short, less than j^ length of distal segment,which has rows of stout bristles medially.First leg large, usually nearly twice length of carapace, with chelaas long as, or longer than, carapace. Ischium short. Merus shorterthan chela, somewhat triangular in cross section, distal marginrounded; ventral with two tuberculate ridges separated by shallowsinus. Outer ridge protruding distally to form broad flat process.Outer surface of merus with broad, oblique sinus, devoid of tubercles,into which second leg may fit. On inner surface a longitudinal sinusand distally a deep transverse groove. Large membranous areaventrally into which flat, platelike toothed process of carpus fits.Scattered tubercles over most of surface. Carpus short, roundeddorsally, with transverse and longitudinal sinuses. Chelae (figs.11, 13, 14, 30, 31) finely tuberculate, spinulate, with fine pubescence,especially dorsally. Not compressed but somewhat flattened on innerside of pahn. Great variation in proportion, length, width, dentitionof palm and fingers. Fingers occasionally subequal to palm but usuallyabout Yz as long. Gaping, large-toothed type much more commonthan nongaping, finely denticulated form described by Rathbun(1904). (There can be no doubt that both types are found in B.harrimani, as a number of specimens have one chela of each type.One such, kept in the laboratory, had the chehpeds injured and theregenerated chelae both had nongaping fingers). Tips of fingerscurved, with chitinous parts interlocking.Second leg (fig. 37) very slender, with elongated ischium, nearlyas long as 5-jointed carpus, longer than merus. First joint of carpusequal in length to next three together, or to chela; second shghtlylonger than subequal third and fourth; fifth, twice fourth. Chelate,with palm longer than fingers, tips setose.Third and fourth legs (figs. 9, 38) slender, somewhat flattened.Ischium subequal to carpus, about ^ length of merus. Merusslightly inflated, with movable spine. Carpus with two ventralterminal spines. Propodus with ventral marginal spines and setae:terminal pair stout, and group of bristles dorsally. Dactylus (fig.39) curved, thin, sharp-pointed, with curved setae dorsally; morethan K length of propodus, which is nearly as long as merus.Fifth leg (fig. 10) similar in size and spinulation to third and fourth,but with transverse bands of setae forming brush on distal half ofpropodus.First pleopod with smaU, sparsely setose endopodite. Secondto fifth with endopodite and exopodite subequal. Appendix internastraplike.Uropod (fig. 26) with distal margin of peduncle scalelike, producedinto two subequal teeth, curved margin between armed with long
PROC US. NAT. MUS. VOL. 115 HART?PLATE 1
PROC. U.S. NAT. MUS. VOL 115 HART?PLATE 2
q;
SHRIMPS OF THE GENUS BETAEUS?HART 439bristles. Median to these another tooth. Distal angle of outermargin of proximal part of exopodite a sharp tooth, covering baseof stout spine; distal margins rounded, with bristles as well as plumosesetae. Bristles dorsally on margin of exopodite, scattered over dorsalsurface of endopodite. Exopodite and endopodite nearly equal insize, longer than telson but about same width.Male.?Similar to female except that carapace and abdomenslightly more slender, pubescence on carapace thicker, setae on mar-gin of first pleura sparser. Antennular peduncle with middle segmentusually distinctly longer than that of females of comparable size.Chelae usually longer than carapace, with same range of variation asfemales but often somewhat wider, stouter, as are walking legs.Second pleopod (fig. 16) with appendix masculina only shghtly longerthan appendix interna, bearing brush of terminal setae.Color.?-The living annual (pi. 1) is transparent, except for clu"0-matophores which occur in a distinct pattern. The color consists ofsmall red chromatophores usually surrounded by dark blue spots.The pigmented areas of the carapace are two broad bands on thedorsal part separated by a thin middorsal line that is unpigmentedexcept between the eyes. Two colored patches occur on the carapaceat the base of the antennae. In the female the green eggs in theovary may show through the integument. The abdomen also ispigmented dorsally and, except for the first segment, the segmentsare clear anteriorly and middorsally but deeply colored in a bandposteriorly at the joints. The lateral part of the sixth abdominalsegment often is pigmented as is the telson, which, however, has alight streak. There is a fine band of color near the anterior marginof the tergum of the first segment.The eyestalks, antennules, and antennae are heavily pigmented.The flagella are reddish. The mouth parts are clear except for thethird maxillipeds, which have scattered spots. The chelipeds are wellcolored, mainly red, but the tips of the claws are clear. The secondpair of walking legs is clear, but the rest have scattered red dendriticchromatophores. The uropods are mainly red but the setae arecreamy white.The color ends in a straight line midlaterally and the ventral partsare quite transparent. The coloration varies with the state of thechromatophores. By day, when the red chromatophores are ex-panded, the animal is reddish or purplish; by night it is distinctlyblue. One specimen, after preservation, turned a uniform pale green.The color recorded by Kathbun (1904) is "fight green."Size.?Carapace length of smallest ovigerous female 5.5 mm.;largest female 12.5 mm.; males 2.5-13 mm. Length of chela oflargest female 10.5 mm.; largest male 14 mm.
440 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115Range.?Newport Harbor, Calif., to Sitka, Alaska.Habitat.?Intertidal, possibly partly commensal in the burrows ofthe mud shrimp Upogebia pugettensis (Dana) or the ghost shrimpCallianassa californiensis Dana. Individuals have been collected frompools formed in the depressions left by boulders on a gravelly beach;the openings of the burrows of the mud shrimps occur in the sides ofthese pools. Others have been found by chance when excavationswere being made for mud or ghost shrimps, and one was found out ofwater between two slabs of sandstone. In only one locality have theshrimps been found in any numbers: in the oyster dike area of Oak-land Bay, Puget Sound, where they were under logs, shells, anddebris near the low tide mark, and in the trickle of water leaking fromdikes (Dr. J. S. Laurie).Material.?The following specimens were examined:California.?Newport Harbor, N. MacGinitie, 1 male.?Monterey Bay,Elkhorn Slough, Aug. 2, 1927, G. E. MacGinitie, 1 ovigerous female (HopkinsMarine Station).?Monterey Bay, in muddy sand with Upogebia, July 3, 1951,C. Hand, 1 male (USNM 92660)^Washington.?Willapa Bay, muddy sand, in burrow of Callianassa californi-ensis Dana, May 4, 1958, F. Clogston and R. U. Gooding, 1 male.?Puget Sound:Oakland Bay, near Shelton, in oyster dikes under wood debris, June 28, 1942,A. H. Banner, 1 ovigerous female; Oakland Bay, shore, Sept. 26, 1951, J. S.Laurie, 1 female, 1 male; Oakland Bay, from under board, Aug. 4, 1952, J. S.Laurie, 1 male; Oakland Bay, Aug. 19, 1952, J. S. Laurie, 2 males; Oakland Bay,State Reserve Dikes, Sept. 16, 1952, J. S. Laurie, 43 females (1 ovigerous), 26 males,1 juvenile; Oakland Bay, Ersudt's Ground, Nov. 2, 1952, J. S. Laurie, 9 females,7 males; Seabeck, Sept. 18, 1928, 1 female; Warm Beach, June 8, 1931, I male.
?
San Juan Archipelago: San Juan Island, Kanaka Bay, with Callianassa californi-ensis Dana, July 31, 1956, R. U. Gooding, 1 ovigerous female; San Juan Island,Garrison Bay, with Upogebia pugeltensis (Dana), Aug. 22, 1957, R. U. Gooding,I male; Brown Island, with Upogebia pugeltensis (Dana), July 13, 1961, J. F. L.Carl, 1 male.British Columbia.?Victoria: Cadboro Point, under boulder, April 7, 1950,J. F. L. Carl, 1 female; Cadboro Point, in pool formed while digging for mudshrimps, July 27, 1958, G. C. and J. F. L. Carl, 1 ovigerous female, 1 male.?Eastcoast of Vancouver Island: Departure Bay, north of Nanaimo, from stomach offlounder, Pleuronichthys coenosus Girard, July 9, 1934, E. Kuitenen, 1 ovigerousfemale (damaged); Departure Bay, under boulders, July 29, 1938, J. F. L. Carl,3 females (2 ovigerous); Departure Bay, Aug. 8, 1938, J. F. L. Carl, 1 female;Departure Bay, Aug. 10, 1938, J. F. L. Carl, 3 females, 2 males; Departure Bay,Aug. 24, 1938, J. F. L. Carl, 5 females, 2 males; Departure Bay, March 23, 1939,J. F. L. Carl, 1 female, 1 male; Hammond Bay, north of Nanaimo, muddy gravel,with Upogebia pugeltensis (Dana), June 22, 1962, J. F. L. Carl, 11 females (3ovigerous), 8 males; Hornby Island, between slabs of sandstone, July 7, 1959,A. D. Carl, 1 male.?West coast of Vancouver Island, Clayoquot Sound, nearKakawis, Meares Island, June 14, 1946, E. F. Ricketts, 1 male (USNM 84397);Kyuquot, mudflat pool, July 10, 1958, G. C. Carl, 1 ovigerous female.Notes.?This is a very agile shrimp, inclined to "play possum" andthen to move very quicldy. Even in clear water, to see the animal
SHRIMPS OF THE GENUS BETAEUS?HART 441
is difficult because of its protective coloration and transparency. Theshrimp is so striking in appearance, however?with its large chelipedsand lobster-like form?that specimens found incidentally when otheranimals are being collected are not likely to be discarded. Yet thereare very few preserved specimens even from areas where the fauna iswell kno\vn. Is it, therefore, really rare or only rarely found? Dr. J.S. Laurie, collecting in Oakland Bay, Puget Sound, seems to be thesole person to have obtained the shrimps in any number, and thiswas done only over a period of more than one year. Subsequentsearching by other workers in the same area has not been productive.Rathburn (1904) described Betaeus harrimani from a single femalefound at Sitka, Alaska. With the exception of Banner's comment(1953) on the far northern habitat, I am unaware of any other ref-erence to the species in the literature. The present specimens agi"eewith Rathbun's description except in a few minor details. The palmof the second leg seems to be appreciably longer than the fingers, notsubequal as she stated, and the posterolateral angles of the fourth andfifth abdominal plem'al segments are square, rather than rounded.The proportionate length and width of the palm, the presence of largeteeth on the fingers, and the gape differ radically from the finelydenticulated, nongaping type she described. There is little doubtthat all specimens in the present collections are B. harrimani because,although in many specimens both chelae are very different, othershave one chela denticulate and nongaping and the other chela toothedand gaping.Ovigerous females have been found in the field June-September andin the laboratory December-July. The eggs are a bright green whenfii'st extruded but become brownish and more transparent beforehatching. Betaeus longidactylus LockingtonFigures 20-22, 27, 32-34, 40-42Betaeus longidactylus Lockington, 1877a, p. 35 (type locality, San Diego, Cali-fornia; holotype not extant); 1878, p. 480.?Rathbun, 1904, p. 108.?Baker, 1912, p. 106.?Hilton, 1916, p. 67.?Schmitt, 1921, p. 80, pi. 12;1924, p. 387.?Johnson and Snook, 1927, p. 310, fig. 262.?MacGinitie, 1930,p. 68; 1935, p. 658, 660, 686, 705-706; 1937, p. 1035.?MacGinitie andMacGinitie, 1949, p. 279.?Ricketts and Calvin, 1952, p. 42, fig. 18.Alpheus longidactylus Kingsley, 1878a, p. 198.?Holmes 1900, p. 190.Female.?Carapace laterally compressed but without carina.Smooth, with very fine scattered setae somewhat concentrated overbranchial region. Longer setae on posterior margin. Front (fig.20) straight, depressed anteriorly, slightly swollen over eyes. Nodistinct junction with lateral margin (fig. 21). Width of carapaceincreases rapidly so that most of carapace subequal in width, but
442 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115lateral margin somewhat undulate. Posterolateral margin curved,posterior margin angled below deep cardiac notch.Abdomen smoothly rounded, with posteroventral margins of firstto third segments rounded, those of fourth and fifth angled. Firstand fifth pleura ventrally margined with plumose setae. Minutesetae on smooth surface.Telson (fig. 27) longer than sixth abdominal segment, nearly twiceas long as wide, outer margin slightly undulate. Two pairs of movablespines on dorsal sm'face, two spines on each posterolateral angle;outer spines very small. Posterior margin deeply curved; plumosesetae relatively short.Eye round. Sharp cone-shaped tooth on median part of short,stout eyestalk.Antennule with stout peduncle of three subequal segments. Broad-based scimitar-shaped stylocerite reaching almost to end of secondsegment. Outer flagella with stout fused portion before division intoshort sensory part and much longer, slender flagellum. Pedunclewith large tooth ventrally.Antenna with stout peduncle slightly longer than antennularpeduncle. Distal margin of basis produced into sharp tooth. Scale(fig. 22) broad; wide spine exceeding blade, and separated by distinctslit distally, outer margin straight. Scale reaches nearly to end ofantennular peduncle. Flagellum longer than carapace, flattened ovoidin cross section.Third maxilliped reaching to last segment of antennular peduncle.Ischiomeropodite flat, relatively narrow, curved over mouth parts,slightly shorter than exopodite, about twice as long as ultimate, fourtimes penultimate segment. Medial surfaces all armed with bristles,which are particularly dense on distal segment.First legs (fig. 32) usually similar in size and shape, with chela muchshorter than carapace. Ischium short. Merus slender, increasing insize distally, somewhat triangular in outline with scattered tubercles.Shallow groove on outer side where second leg fits. Carpus short,smooth, cup-shaped, with sHght dorsal and larger ventral flattenedprojection. Chela narrow elongate, covered with fine spines. Fingerslonger than palm, narrow gape, with one large flat tooth medially onfixed finger, another proximally, which meshes with similar one ondactylus. Curved corneous tips cross and intermesh. Chela sub-rectangular, but deepest at base of dactylus. There may not be anylarge teeth nor any gape when fingers are closed, particularly in smallindividuals (fig. 33).Second leg (fig. 40) slender, with ischium subequal to merus. Firstjoint of carpus slightly longer than next three together, subequal to
SHRIMPS OF THE GENUS BETAEUS?HART 443fifth plus palm. Second joint longer than subequal third and fourth.Fingers subequal to, or longer than, palm.Third leg (fig. 41) relatively stout, slightly flattened, reaching tofingers of cheliped. Ischium about % length of merus, which is slightlydilated, has movable spine. Carpus stout, nearly twice length ofischium, with two ventral terminal spines. Propodus elongate, sub-equal to merus, with double row of spines on ventral margin (terminalpair much stouter than others). Dactylus (fig. 42) stout, withcurved corneus tip and bristles on slight elevation dorsally.Fom'th leg very like third but somewhat smaller.Fifth leg more slender. Ischium about K length of merus. Undi-lated merus with movable spine, longer than propodus, twice as long ascarpus. Propodus with transverse rows of bristles distally on outerface as well as double row of spines on ventral margin.First pleopod with slender endopodite about % length of exopodite.Second pleopod with endopodite slightly shorter, narrower than ex-opodite. Appendix interna straplike.Uropod (fig. 27) with distal margin of peduncle scalelike, producedinto two teeth, outer of which stouter, longer, separated by curvedmargin armed with long bristles. Another tooth median to these.Proximal part of exopodite with large posterolateral tooth, dorsal tolarge spine, which is nearly as long as distal section. Dorsal surfaceof endopodite with bristles, all posterior margins fringed with longplumose setae. Exopodite slightly longer than endopodite, bothlonger than telson,Male.?Similar to female except carapace and abdomen slightlymore slender and setae on first pleuron sparse, not plumose. Smallindividuals have chelipeds very like those of females but proportion-ately somewhat larger. Large males develop very large chelipeds,with wide gape, big teeth, and fingers touching only at the tips (fig.34). When the carapace length is more than 9 mm., the hand may belonger than the carapace and, in some instances, even Iji this length.Appendix masculina of second pleopod straplike; terminating in abrush of bristles and somewhat longer than appendix interna.Color.?Published records give the color of the living animal asolive-green, olive-brown, uniform red-brown, or blue-green, with aHght middorsal stripe, reddish legs with clear white tips, and darktail fan with yellow setae.Size.?Carapace length of smallest ovigerous female 7.8 mm. ;largest female 14.5 mm.; males 4-16 mm. Length of chela of largestfemale 11.3 mm. and of largest male 23.5 mm.Range.?Tepoca Bay, Gulf of California, Mexico (about 30? N,113? W) to Elkhorn Slough, Monterey Bay, Calif.Habitat.?Tide pools, under rocks, in eelgrass, in crevices among
444 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115
materials on boat bottoms, and paired in burrows of echiuroid wormUrechis caupo Fisher and MacGinitie or mud shrimp Upogebia puget-tensis (Dana).Material.?The following specimens were examined:Mexico.?San Felipo, May 11, 1937, S. A. Glassell, 1 female.California.?San Diego, dredged, March 9, 1949, T. E. Bowman, 2 males(USNM 98059).?La Jolla, from kelp holdfasts on beach, Aug. 28, 1918, W. L.Schmitt, 2 females (USNM 53910).?La Jolla, Sept. 21, 1918, W. L. Schmitt,1 female, 2 males (USNM 53914).?La Jolla, tide pools, Sept. 22, 1918, W. L.Schmitt, 1 ovigerous female, 9 males (USNM 53925).?La Jolla, Bird Rock,Nov. 3, 1942, 1 female, 1 male (Pacific Marine Station, 846, 847).?La Jolla, kelpbeds, March 16, 1954, C. Limbaugh, 1 male (USNM 96415).?Laguna Beach,W. A. Hilton, 1 ovigerous female, 2 males (USNM 48982).?Laguna Beach, W. A.Hilton, 1 ovigerous female, 2 males (USNM 50582).?Laguna Beach, W. A. Hilton,2 ovigerous females (USNM 50586).?Laguna Beach, W. A. Hilton, 1 male(USNM 50590).)?Laguna Beach, under stones, W. A. Hilton, 1 ovigerous female,2 males, pale olive-green (USNM 52756).?Newport Bay, Jan. 20, 1929, Dec.16, 1930, G. E. MacGinitie, 1 female, 1 male, 12 juveniles (Hopkins MarineStation).?Santa Catalina Island, west shore of Catalina Harbor, Dec. 12, 1912,Anton Dohrn, 3 females (USNM 49981).?Santa Catalina Island, Nov. 28, 1913,Anton Dohrn, 5 females, 10 males (USNM 49980).?Catalina Island, IsthmusHarbor, Nov. 27, 1913, Anton Dohrn, 9 females, 8 males (USNM 49982).?CatalinaIsland, west shore of Catalina Harbor, Nov. 28, 1913, Anton Dohrn, 20 juveniles(USNM 50012).?Long Beach, H. N. Lowe, 2 males (USNM 53018).?SanPedro, C. F. Baker, 1 male (USNM 29309).?San Pedro, Rocky Point, Feb. 2,1912, Anton Dohrn, 2 males (USNM 49978).?San Pedro, Portuguese Bend,June 26, 1914, Anton Dohrn, 1 male (USNM 49979).?San Pedro, GovernmentBreakwater and Point Fermin, littoral, April 2, 1913, Anton Dohrn, 3 females,6 males (USNM 49983).?San Pedro, first rocks north of Port Los Angeles, Dec.5, 1911, Anton Dohrn, 3 females, 2 males (USNM 49993).?San Pedro, May4, 1919, E. P. Chace, 1 female (USNM 54048).?San Pedro, June 25, 1895,5. J. Holmes (?), Univ. of Cahfornia, 20 females (14 ovigerous, 5 parasitized),6 males (USNM 87439).?Santa Monica, near long wharf, Venice Marine Bio-logical Station, 1 male (USNM 46118).?Santa Monica, April 1923, E. P. Chace,1 male (USNM 57174).?Morro Bay, Feb. 2, 1939, S. A. Glassell, 1 male.?SantaMonica, 8 miles north, open beach in Upogebia burrow, Feb. 2, 1931, G. E.MacGinitie, 1 female (Hopkins Marine Station).?Monterey Bay, ElkhornSlough, April 3, 1930, G. E. MacGinitie, 1 female (Hopkins Marine Station).Notes.?Betaeus longidactylus is a stouter species than B. harrimaniand may be separated easily on the basis of the chelipeds. Theranges overlap in the region from Newport Bay to Monterey, butfew specimens of either species are present in collections from thisarea. Ricketts and Calvin (1952) state that B. longidactylus is veryplentiful in southern California, where it occurs on the outer coast,but in the northern part of its range the species is restricted to quietwaters. Berried females have been taken in January, June, August,and September.
SHRIMPS OF THE GENUS BETAEUS?HART 445Betaeus ensenadensis GlassellFigures 23-25, 28, 35, 36, 43-45
Betaeus, new species, MacGinitie, 1934, pp. 173-174; 1937, pp. 1035-1036.Betaexis ensenadensis Glassell 1938, pp. 416-419, pi. 28 (type locality, Estero de laPunta Banda, Ensenada, Baja California, Mexico; holotype male, San DiegoSoc. Nat. Hist. No. 1121).?MacGinitie and MacGinitie, 1949, p. 279.Female.?Carapace laterally compressed but without carina;smooth, naked. Front (fig. 23) shghtly curved, depressed anteriorly.Anterior margin without sinuses or with shallow ones. Anterolateralangle obtuse (fig. 24). Width of carapace increases rapidly to base ofthird maxilliped, followed by slight undulation over base of first legs.Posterolateral region rounded. Blunt angle beneath cardiac notch.Abdomen smoothly rounded with posteroventral margins of pleuraof first four segments rounded, fifth bluntly angled. Ventral marginof first pleuron sparsely setose.Telson (fig. 28) subequal to sixth segment, nearly twice as long aswide, with outer margins slightly undulate. Two pairs of movablespines on dorsal sm'face, two spines at each posterolateral angle, outerones very small. Posterior margin slightly curved, armed with plu-mose setae.Eye somewhat oval, with small flat tooth on eyestalk.Antennule with stout scimitar-shaped stylocerite, reaching to aboutdistal third of second segment of peduncle. Subterminally first seg-ment of peduncle produced into thin tooth ventrally. Second segmentof peduncle longer than fu'st, which is longer than third. Flattenedfused part of outer flagella shorter than peduncle; distally dividedinto few free segments and slender flagellum subequal to fused portion.Inner flagellum considerably longer.Antennal peduncle subequal to antennular peduncle, with ventraldistal part of basis produced into sharp tooth. Scale (fig. 25) withspine much longer than narrow blade, slit between minute; outermargin almost straight. Scale reaching to middle of third segment ofantennular peduncle. Flagellum short, wide, much flattened, reachingslightly past posterior margin of carapace.Thu'd maxilhped reaching to middle of second joint of antennularpeduncle. Ischiomeropodite sparsely setose, slender, flattened, some-what curved, subequal to two distal segments together in length.Exopodite longer than ischiomeropodite, relatively stout. Penulti-mate segment shortest, sparsely setose. Distal segment bandedmedially with many short bristles.First legs (fig. 35) similar. Ischium short with ridge flatly tubercu-lated ventrally. Stout merus with deep groove into which second legfits. Two evenly toothed ridges ventrally, transverse sulcus distally.
446 PROCEEDINGS OF THE NATIONAL MUSEUM vol. usCarpus short, rounded dorsally, smooth, except for two tubercles orteeth on inner margin. Platehke ridge ventrally with somewhat roughedge. Chela covered with fine denticles, particularly on outer sur-faces. Inner side of chela flat, outer rounded. Dactylus slightlyshorter than palm, with three teeth: one small, near junction with palmand meshing with similar tooth on fixed finger; large conical toothnear middle; terminal tooth meshing with truncate terminal part offixed finger. Fixed finger stout with wide cutting surface bordered oneach side with denticulated, elevated edge; both fingers end in sharptips that cross when closed. Distinct gape.Second leg (fig. 43) slender. Merus and ischium subequal, slightlyshorter than 5-jointed carpus, which has fu-st joint longer than fifthand about equal to second, third, and foiu'th together, which aresubequal to each other. Fingers subequal to palm.Third (fig. 44) and fourth legs flattened with dilated merus. Is-chium nearly ji merus in length. Distinct movable spines on merusand ischium. Sparsely setose; single spine and setae on distal ventralpart of carpus, usually three equidistant spines and terminal pair onventral part of propodus. Few bristles dorsaUy. Needle-sharpcurved dactylus (fig. 45), more than % length of propodus.Fifth leg without movable spine on ischium, but one on undilatedmerus. Kather sparse brush of bristles on distal part of propodus.Dactylus similar to that of thu'd or fourth leg.First pleopod with round-tipped endopodite less than ji length ofexopodite. Second to fifth pleopods with exopodite slightly longerthan endopodite, both slender. Appendix interna stout and straplike.Uropod (fig. 28) broad and considerably longer than telson. Pe-duncle with straight margin; two small teeth separated by serratedspines graduated in length with longest medial. Proximal marginof exopodite with serrated spines of varying lengths forming thatchbetween small lateral tooth and large distal spine.Male.?With straight front. Cheliped (fig. 36) proportionatelyheavier, wider than that of female (median tooth on dactylus maybe missing). Appendix masculina on second pleopod only slightlylonger than appendix interna, blunt tip armed with bristles of variedlengths.Color.?"The carapace, abdomen and chelipeds are covered withlight tinted chromatophores in reds and blues, the fingers and telsonare tinted a light purple" (GlasseU, 1938). "Minute red and bluespots" (MacGinitie and MacGinitie, 1949).Size.?Carapace length of smallest ovigerous female 5 mm.; largestfemale 7 mm.; males 4.5-6.8 mm. Length of chela of largest female6 mm.; of largest male 5.6 mm.
SHRIMPS OF THE GENUS BETAEUS?^HART 447Range.?El Estero de la Punta Banda, Ensenada, Baja California,Mexico, to False Bay, San Diego, Calif.Habitat.?MacGinitie (1937), MacGinitie and MacGinitie (1949),and Glassell (1938) record the shrimps living in pairs in the burrowsof the ghost shrimp Callianassa californiensis Dana at Ensenadaand in the burrows of the mud shrimp Uiiogehia pugettensis (Dana)at San Diego,Material.?The following specimens were examined:Mexico.?El Estero de la Punta Bauda, Ensenada, Baja California, Dec. 19,1930, G. E. MacGinitie, 1 ovigerous female, 1 male.?El Estero de la PuntaBanda, Ensenada, Baja California, Jan. 21, 1932, G. E. MacGinitie, 7 females(1 ovigerous), 3 males.California.? Mission Bay, in burrow, December 1949, T. E. Bowman, 1 speci-men (USNM 102442).?False Bay, San Diego, with Upogebia, Nov. 18, 1937,G. E. MacGinitie, 3 females (1 ovigerous).Notes.-?A small, slight shrimp of the same type as Betaeusharrimani and B. longidadylus, but easily distinguished from these bythe shape of the antennal scale, the truncate tip of the fixed finger,the large teeth on the dactylus of the chela, tlie presence of a movablespine on the ischium of the third and fourth walkuig legs, and thecharacteristics of the uropod. Ovigerous females have been taken inNovember, December, and January.Betaeus harfordi (Kingsley)Figures 46, 47, 54, 58-61, 73, 74Betaeus eqidmanus Lockington, 1877b, p. 43 (type locality, Cataliua Island,California; types not extant).Alpheiis harfordi Kingsley, 1878a, pp. 198-199 (type localities, Santa Barbara andCatalina Island, California; types not located); 1878b, pp. 58-59; 1882,p. 124, pi. 2, fig. 4.Alpheus aequalis Kingsley, 1878a, p. 199.Betaeus aequalis Lockington, 1878, pp. 479-480.Alpheus (Betaeus) aequalis Holmes, 1900, p. 189, pi. 3, fig. 47 (part).Betaeus harfordi Rathbun, 1904, p. 108.?Schmitt, 1921, p. 79, fig. 55.?MacGinitieand MacGinitie, 1949, p. 279 (part).?Ricketts and Calvin, 1952, pp. 323-324(part)
.
Female.?Carapace laterally compressed but without carina;smooth, naked. Front (fig. 46) shallowly emarginate, produced tocover eyes. Anterior margin of carapace (fig. 47) shallowly cmwed,meeting lateral margin at blunt angle just ventral to base of antenna.Width of carapace increasing to base of third maxillipeds, graduallydecreasing to curved posterolateral margin. Posterior margin pro-trudes slightly before decreasing ventral to cardiac notch.Abdomen smooth, evenly rounded. Naked except for plumosesetae on ventral margin of pleura of first segment. Pleura with693-433?63 2
448 PROCEEDINGS OF THE NATIONAL MUSEUM vol. usposterolateral margins evenly rounded except fifth, which is somewhatacute.Telson (fig. 58) wide, flat, shorter than uropods. One and one-halfto twice length of sixth segment. Lateral margins curved. Distallyabout % as wide as proximally. Two pairs of small movable spineson distal half of dorsal sm-face, but posterolateral spines vestigial ormissing. Posterior margin cm'ved, bearing long plumose setae.Eye large, round, on short stalk with small median tooth.Antennule with stylocerite narrow, sharp-pointed, reaching to distalquarter of second segment of peduncle. Ventrally a slender scimitar-shaped tooth on distal part of first segment of peduncle. Peduncleslender, about % carapace length. Second segment about twice lengthof subequal first and third segments. Flagella slender, short: outerflagella fused for most of length with only terminal tip of sensory partfree and remaining flagellmn short, slender.Antenna slender. Peduncle with wide tooth ventrally on margin offirst segment; slightly longer than antennular peduncle. Scale (fig.54) narrow, with stout spine longer than narrow blade, separated fromit with slit for about % of length. Flagcllum longer than carapace,ovoid in cross section.Third maxilliped reaching to end of second segment of antennularpedmicle. Relatively narrow, with flattened ischiomeropodite sub-equal to distal segment, sparsely bristled. Penultimate segmentsparsely setose, slightly shorter than distal, which is armed mediallywith bands of stout bristles and scattered longer setae. Exopoditeslightly longer than ischiomeropodite.First legs subequal, large. Chela slightly shorter than carapace.Ischimn short. Merus naked except for few setae on inner margin:triangular in cross section, wdth dorsal carina smooth, sharp, ending incurved tooth distally, outer ventral ridge smooth, inner one \vith someflattened tubercles. Carpus short, smooth, naked, cup-shaped withsmall ventral plate. Chela (fig. 60) smooth, flattened laterally: fewfine scattered setae on dorsal ridge, stifT setae near tips of fingers andon cutting surfaces. Palm and fingers subequal but fixed finger muchwider than dactjdus. Somewhat flattened teeth of cutting surfacemay mesh without any gape or may have hiatuses on both fingersresulting in a round foramen. Chelae usually alike in dentition.Dorsal ridge of fixed finger and palm compressed, so that marginknife-edged.Second leg (fig. 61) slender. Ischium longer than merus; both flat-tened laterally. Carpus with first joint longest, fifth next, othersshorter, subequal. Chela longer than first joint, fingers subequal topalm. Cutting smface of fingers spinulate.Third leg (fig. 73) stout, flattened laterally, with scattered fine
SHRIMPS OF THE GENUS BETAEUS?^HART 449
setae. Ischium and merus comprise }^ length of appendage. Largemovable spine on merus. Carpus, with minute terminal spines, sub-equal in length to propodus. Propodus armed ventrally with smallsetae and vestigial spines. Terminal pair larger. Dactylus (fig. 74)short, stout, bifid, but claws masked by two median bands of shortstiff bristles.Fourth leg similar to third, but all segments except ischium slightlysmaller.Fifth leg distinctly shorter than fourth. Subequal merus andpropodus each longer than carpus. Brush of 4-5 bands of short stiffsetae on distal part of propodus. Dactylus similar to that of thirdand fourth legs. No movable spine on merus.First pleopod with round-tipped endopodite, nearly ji length andwidth of slender exopodite. Second pleopod with subequal slenderendopodite and exopodite. Appendix interna straplike, nearly %length of endopodite.Uropod (fig. 58) with posterior margin of peduncle produced intotwo sharp subequal teeth, separated by concave margin with few smallsetae. Distal angle of proximal part of exopodite a sharp tooth,covering base of short stout spine. Uropods longer than telson.Male.?Similar to female except that chelae (fig. 59) are propor-tionately larger, subequal in length to carapace. Appendix masculinawith bristles on margin of distal half and on blunt tip; twice as longas appendix interna, nearly as long as endopodite.Color.?Dark purple (Lockington, 1878; Holmes, 1900), blue-black(specimen label, E. P. Chace), deep blue (specimen label, W. A.Hilton) and deep blue, occasionally reddish-brown on sides (Mrs.G. E. MacGinitie).Size.?Carapace length of smallest ovigerous female 6 mm.; largestfemale 10.7 mm; males 3.3-8 mm. Length of chela of largest female8.5 mm.; of largest male 8 mm.Range.?^Magdalena Bay, Mexico, to Fort Bragg, Calif, (personalcommunication, Mr. Keith W. Cox) ; intertidal to depth of 12 fathoms.This is apparently a very elusive shrimp that leaves its host with littleprovocation, with the residt that, unless precautions are taken toprevent its escape, it may be consistently missed when abalones arecollected. UntU Mr. Cox was consulted, no specimens had beenrecorded in any collections taken north of Santa Monica, even withMr. D. Montgomery checking approximately 350 specimens ofHaliotis from the vicinity of San Luis Obispo. On the other hand,Dr. Howard Teas, on April 4, 1953, collected at El Estero de PuntaBanda, Ensenada, Baja California, Mexico, and reported to the Mac-Ginities that about four out of every five Haliotis fulgens had aBetaeus in the mantle cavity.093-433?63 3
450 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii5Habitat.?Found living in mantle cavity of ab alone and occasion-ally free, latter probably temporary state, owing to distm-bance ofnormal host. The specimens examined have occm^red most commonlywith the pink ab alone, Haliotis corrugata Gray, but also with the redabalone, H. rufescens Swainson, the black abalone, H. cracherodiiLeach, and the green abalone, H. fulgens PhUlipi. Mr. Keith Coxreports them also from H. kamtschatkana Jonas, H. wallalensisStearns, H. sorenseni Bartsch, and H. assimilis Dall.Material.?The following specmiens were examined:Mexico.?Magdalena Bay, Belchers Point, on shore, Jan. 30, 1938, S. A.Glassell, 1 specimen, dried.California.?La Jolla Cove, July 14, 1942, 1 female (Pacific Marine Station845).?La Jolla, kelp beds, March 16, 1954, C. Limbaugh, 1 ovigerous female, Imale (USNM 96414).?La Jolla, with Haliotis rufescens Swainson, March 28,1952, E. Dodge, 1 ovigerous female.?San Clemente Island, living beneath mantleof Haliotis, January 1936, Nell Murbarger, 2 ovigerous females, 1 male (USNM77743).?San Clemente Island, from abalone. May 8, 1888, Albatross, 2 females(USNM 63453).?Laguna Beach, from abalone, Sept. 19, I9I8, W. A. Hilton,I female, deep blue (LTSNM 53598).?Laguna Beach, shale reef ]/i mile offshore,midway to Corona del Mar, 12 fathoms, from mantle cavity of Haliotis corrugataGray, April 25, 1958, Robert Given, from N. MacGinitie, I ovigerous female.
?
Laguna Beach, off Salt Creek, 8 fathoms, from Haliotis corrugata Gray, May 21,1958, Dale Seemen, from N. MacGinitie, 20 females (7 ovigerous), 5 males;May 26, 1958, 7 females (5 ovigerous).?Laguna Beach, 8 fathoms, from Haliotiscorrugata (2 specimens from Haliotis rufescens), April 28, 1958, Dale Seeman,from N. MacGinitie, 4 females (I ovigerous), 10 males.?Catalina Island, com-mensal in shell of green abalone, Venice Marine Biological Station, 1 ovigerousfemale (USNM 46119).?Catalina Island, Little Harbor, Dec. 27, 1912, AntonDohrn, 2 males (USNM 49977).?San Pedro, Rocky Point, from mantle of blackabalone, Feb. 2, 1912, Anton Dohrn, 1 ovigerous female (USNM 49975).?SanPedro, in gill chamber of Haliotis cracherodii Leach, June 1930, E. P. Chace,1 ovigerous female, blue-black when alive (USNM 64087).?Santa BarbaraIsland, from Haliotis rufescens, 8 fathoms, Sept. 11, 1962, Keith W. Cox, 22ovigerous females, 7 males.?Santa Monica Bay, Malibu Point, under mantle ofHaliotis washed up on beach, Dec. 19, 1937, A. Camp, from R. Crocker, 1 female(Hopkins Marine Station).?Southern California, Nov. 27, 1913, Anton Dohrn,1 female (USNM 49976).Notes.?Mr. Keith W. Cox informs me that there seems to be adefinite correlation in size between the shrimps and their hosts:large shrimps in large abalones, small in small. The usual number isone shrimp per host. Exceptions do occiu* and on several occasionsup to four have been found in the cavity between mantle and shell.MacGinitie and MacGinitie (1949, p. 279) state:We have found this shrimp with its purplish-red color form living in the mantlecavity of the green abalone Haliotis fulgens. For over a year one has been livingunder the protectorate of an abalone in one of our aquariums. It usually stayswith its head near the region of the mouth of the abalone, and although it is byno means inactive, its host does not give any evidence of being disturbed by itsmovements. On rare occasions we have seen it make short excursions onto the
SHRIMPS OF THE GENUS BETAEUS?HART 451
shell, but it soon returns to its shelter. More frequently it may be seen on theupper surface of the frilly mantle edge of its host. At the least disturbance ithastens beneath the mantle and the abalone closes down over it in what resemblesa protective manner.In a letter, Mrs. G. E. MacGinitie states:When the abalone diver brought in so many live B. harfordi we dumped about20 of them in an aquarium with two H. corrugaia. The next morning practicallyall of them had found a niche somewhere in the two abalones. But they didnot all stay with the abalones very long. One per abalone is apparently normal,and soon the others were elsewhere. Six or more took refuge in the niches of theegg case of the horned shark. Another snuggled next to the third abalone in theaquarium?a specimen about an inch long, and the shrimp was at least as longas the "host."Ovigerous females have been taken in March, April, May, Septem-ber, and December. Mrs. MacGinitie reports that the eggs, whenfreshly laid, are nearly opaque and a pale yellowish green, becomingmore transparent with development.Betaeus macginitieae, new speciesFigures 48, 49, 55, 62-64, 75, 76Alpheus aequalis Holmes, 1900, p. 189 (part, specimens found on sea urchins).Betaeus harfordi MacGinitie and MacGinitie, 1949, p. 279 (part, purple ones withsea urchins).Female.?Carapace laterally compressed but without carina;smooth, naked. Front (fig. 48) produced to form hood, which iscurved slightly anteriorly but without emargination. Lateral margin(fig. 49) faintly curved, closely adhered around base of antenna with-out distinct anterolateral angle. Width of carapace increasingrapidly to base of maxilliped, then decreasing slightly to form curvedlateral margin. Posterolateral junction rounded; posterior marginnearly straight ventral to junction of cardiac notch.Abdomen smoothly rounded. Posterior margins of all pleurarounded but that of fifth somewhat acute. No surface setae and onlypleura of first segment margined ventrally with plumose setae.Telson (fig. 62) flattened, longer than sixth segment. About twiceas wide proximally as distally. Two pairs of movable spines dorsally.Posterior margin curved, with pair of small spines (outer smaller)externally, and long plumose setae.Eye oval, on short eyestalk with small tooth,Antennule with stylocerite narrow, reaching to last quarter ofsecond segment of peduncle. Peduncle with middle segment nearlytwice as long as either first or third; almost bare of setae. Peduncleabout /^ length of carapace. Ventrally near distal margin of firstsegment a stout laterally compressed tooth. Inner flagellum stout,elongate. Fused part of outer flagella stout but free tips short.
452 PROCEEDESTGS OF THE NATIONAL MUSEUM vol. 115Antennal peduncle slightly longer than antennular, with ventralmarginal tooth under base of scale. Flagellum flattened, somewhatlonger than carapace. Scale (fig. 55) narrow: blade shorter thanspine but slightly wider, separated by slit for ji length. Reaches tonearly middle of third antennular peduncle segment.Thkd maxilliped reaches to about middle of second segment ofantennular peduncle. Ischiomeropodite broad, flattened, curvedover mouth parts; slightly longer than other segments together.Penultimate segment shortest, armed with tufts of stiff bristles, distalsegment with bands of stiff bristles on inner margin.First legs subequal, large. Ischium short, stout. Merus stout,about twice as long as wide, triangular in cross section. Dorsalridge smooth but ventral ridges with poorly developed setae and teeth.Carpus short, cup-shaped, naked. Chela elongate, more than twicelength of merus, 2^ times as long as wide, wdth palm slightly longerthan fingers. Blunt tooth on proximal part of fixed finger, followedby hiatus before denticulate cutting siu-face, which meshes withsimilar area on dactylus, which also has similar proximal tooth. Cut-ting surfaces sparsely setose. Chela somewhat flattened lateraUy,dorsal and ventral margins rounded, not knifelike.Second leg (fig. 64) long, slender. Ischium slightly longer thanmerus, which is shorter than carpus. Carpus with fu'st segmentlongest, about 1}^ of fifth, which is longer than subequal second, third,and fom'th segments. Fingers and palm subequal with curvedbristles distally and spinulose cutting surface.Third leg (fig. 75) with stout ischium and slightly dflated merus.Merus slightly longer than propodus, which is 1% times length ofcarpus. Movable spine on proximal part of merus. Dactylus(fig. 76) short, }i length of propodus, with bifid tip somewhat hiddenby setae. Spines on ventral margin of carpus and propodus shortbut stout; one pair on carpus, many on propodus with distal pairlongest.Fourth leg similar but slightly smaller.Fifth leg with shorter ischimn. Spines on ventral margin ofpropodus very small, bristles of brush short. No movable spine onmerus.First pleopod with endopodite curved and ji length of exopodite.Second pleopod with slender subequal exopodite and endopodite.AppendLx interna stout, straplike.Uropod (fig. 62) with posterior margin of peduncle produced intotwo small teeth close together, without setae. Posterolateral toothwell developed on proximal part of exopodite; spine beneath thissomewhat short, stout. Uropods longer than telson.Male.?^Very similar to female, but chelipeds (fig. 63) slightly longer
SHRIMPS OF THE GENUS BETAEUS
?
^HART 453in proportion. In one specimen, left side of peduncle of uropods cutinto two small teeth, but right side with only one tooth. Appendixmasculina bristled on distal half and on tip; twice as long as appendixinterna and reaching midway between tip of appendix interna andtip of endopodite.Color.?"Dark purple . . . resembled the color of the sea urchinsin whose spines they were entangled when captured" (Holmes, 1900)."In life I have noticed no difference in the color of B. harfordi and thelast one we are sending you from the sea lu-chin. For the most part,they are mostly blue, but sometimes along the side the color is areddish brown. The claws and tail are almost always entirely blueregardless of the color along the sides of the abdomen. I could findno correlation between color and sex" (Mrs. G. E. MacGinitie,letter, 1958).Size.?Carapace length of smallest ovigerous female 6.8 mm.;largest female 10.5 mm. ; males 5.5-10 mm. Length of chela of largestfemale 10.3 mm.; of largest male 10.5 mm.Range.^?Santa Catalina Island to Monterey, Calif.Habitat.?Occurs in pairs in association with sea urchins,Strongylocentrotus franciscana (Agassiz) and S. purpuratus(Stimpson)
.
Material.?-The following specimens were examined
;
California.?Santa Catalina Island, Isthmus Harbor, Nov. 27, 1913, AntonDohrn, 1 ovigerous female (USNM 107546).?Laguna Beach, under Strongylocen-trotus purpuratus (Stimpson) in hole in rock, Nov. 18, 1937, N. MacGinitie, 1ovigerous female, 1 male.?Corona del Mar and Laguna Beach, shale reef mid-way between, ]i mile offshore, 5 fathoms, from beneath Strongylocentrotus fran-ciscana (Agassiz), Apr. 17, 1958, Robert Given, from N. MacGinitie, 1 ovigerousfemale.?Corona del Mar, Arch Rock, tide pool (may have been with purple seaurchin before being disturbed), Oct. 1932, N. MacGinitie, 1 female.?Corona delMar, off Ladder Rock, 4 fathoins. May 22, 1958, Robert Given, from N. Mac-Ginitie, 1 female holotype (USNM 108228).? Monterey Bay, China Point, June1908, 2 females (1 ovigerous), 1 male (Hopkins Marine Station).Notes.?Some specimens have the proximal part of the fingers ofthe chelipeds denticulate rather than with a distinct tooth. Oviger-ous females have been taken in April, June, October, and November.Mrs. MacGinitie reports that the eggs, when freshly laid, are nearlyopaque and a pale yeUowish-green, becoming more transparent withdevelopment. Betaeus gracilis, new speciesFigures 50, 51, 66, 65-67, 77, 78Betaeus harfordi Hilton, 1916, p. 67.Female.?Carapace laterally compressed but without carina.Smooth, with scattered fine short setae. Setae on posterior margin
454 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115
of carapace ventral to cardiac notch. Front (fig. 50) depressed,deeply indented medially, covering eyes with two hoods. Anteriormargin (fig. 51) evenly curved, rounded at junction with lateralmargin. Carapace increases in width to base of first legs, decreasingslightly to rounded posterior. Ventral part of posterior marginconcave, then angled sharply to cardiac notch.Abdomen smoothly rounded, bearing few scattered fine setae.Posteroventral margins of pleura of fu-st three segments rounded,those of fourth and fifth bluntly square. Ventral margin of fu-stpleura with plumose setae.Telson (fig. 65) wide, flattened, longer than sixth abdominal seg-ment. Two pairs of movable spines dorsally; posterolateral spineswell developed, outer pair smaller. Posterior margin curved, borderedwith bristles and plumose setae.Eye round, with sharp tubercle on stalk.Antennule short, stout. Stylocerite reaches nearly to end of secondsegment of peduncle; margins curved. Peduncle less than % carapacelength. Slender inner flagellum scarcely as long as carapace, outershorter, with fused portion about as long as peduncle, with four freesensory joints and slender terminal flagellum.Antenna with peduncle longer than that of antennule. Margin offu-st segment produced into sharp tooth ventrally. Scale (fig. 56)broad, with stout spine, which is longer than blade and separatedfrom it for distal thhd. Flattened flagellmn ahnost as long as carapace.Third maxilliped reaches to end of antennular peduncle. Ischio-meropodite subequal in length to last segments together (penultimateslightly shorter). Medially armed with long bristles, distal seg-ment with bands of bristles as well. Exopodite stout, longer thanischiomeropodite.First legs (fig. 66) subequal. Ischium short. Merus, short, wide,triangular in cross section. Smooth doreal ridge ending distally insharp curved tooth, with few bristles on margin. Outer ventral ridgesmooth, inner with ten long bristles. Carpus cup-shaped but some-what elongate; no setae. Chela elongate with few setae on cuttingedges and tips of fingers, which are crossed over each other. Dorsalmargin of hand evenly rounded, not knifelike. Palm longer thanfingers, considerably less than ji as wide as long. Little gape betweenfingers.Second leg (fig. 67) slender. Ischium slightly longer than merus,together longer than carpus, which has first joint longer than fifth,and about as long as second, third, and fourth together. Chelaslender, with fingers shorter than palm and with spinulate cuttingsurfaces.Third leg (fig. 77) stouter than second. Merus only slightly dilated.
SHRIMPS OF THE GENUS BETAEUS?HART 455bearing movable spine on proximal lower margin; sparsely setose.Merus and propodus subequal in length. Spines on carpus and pro-podus small, with fine seta beside each spine. Narrow bifid dactyliis(fig. 78) with inconspicuous setae at base of claws.Fourth leg similar but smaller than third.Fifth leg well developed, but ischium and merus slightly shorterthan those of third leg, no movable spine present. No spines oncarpus, few on propodus, but usual brush on distal part of propodus.Dactylus longer, more slender than others but similarly bifid.First pleopod with endopodite short and round at tip. Secondpleopod with endopodite subequal to exopodite, appendLx internastraplike.Uropod (fig. 65) longer than telson. Posterior margin of peduncleproduced into two teeth, with setae on curved margin between.Outer tooth longer. Lateral margin of exopodite ends in small toothcovering base of large spine, nearly as long as terminal portion.Male.?Similar to female except tip of stylocerite reaches to endof second segment of antennular peduncle. Left chela with decidedgape, large triangular tooth on fixed finger, intermeshing smallerteeth on both fingers. Right chela with gape composed of roundhiatuses. Second pleopod with appendix masculina bristled on distalhalf, terminally on blunt tip.Color.?Pale olive-green (Hilton, 1916). Specimen label: "greeneyes."Size.?Carapace length of smallest ovigerous female 6 mm.;largest female 8 mm.; male 6 mm. Length of chela of largest female8 mm. ; of male 6 mm.Range.?Laguna Beach to Monterey Bay, Calif.Habitat.?Kelp holdfasts (Hilton 1916).Material.?The following specimens were examined:California.?Laguna Beach, W. A. Hilton, 1 ovigerous female holotype(USNM 48933), 1 ovigerous female (USNM 108230).?Pacific Grove, January1928, G. E. MacGinitie, 1 specimen.?Monterey Bay, China Point, shore, June1908, 1 ovigerous female, 1 male (Hopkins Marine Station).Notes.?^This species may be confused with Betaeus setosus, but ithas much shorter antennules and antennae, is more slender and lesssetose. The chelae of the female from China Point are toothed likethose of the male: intermeshing, with the hiatuses coinciding to forma foramen. Ovigerous females have been taken in June. The eggsare translucent gi"een (HUton, 1916).Betaeus setosus, new speciesFigures 52, 53, 57, 68-72, 79, 80; Plate 2Alpheus aequalis Holmes, 1900, p. 189 Cpart, free-living form).
456 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115Betaeus harfordi MacGinitie and MacGinitie, 1949, p. 279 (part).?Ricketts andCalvin, 1952, pp. 323-324.Female.?Carapace laterally compressed but no carina. Smoothbut with scattered short setae. Tuft of longer setae under front, be-tween eyes; cardiac notch and lower posterior margin fringed withsetae. Front (fig. 52) deeply indented medially, slightly depressedbut produced laterally to form "blister" over each eye. Anteriormargin (fig. 53) smoothly curved. Anterolateral junction rounded.Lateral margin evenly rounded with widest part at base of first legs.Posterior margin somewhat truncate with distinct cardiac notch.Abdomen rounded, with posteroventral margins of pleura roundedin segments one to thi-ee but angled in four and five. Ventral marginof first segment with plumose setae. Smooth with scattered setae.Telson (fig. 68) longer than sixth segment and about 1% timesas long as widest part. Dorsal surface with scattered setae andtwo pairs of movable spines. Posterolateral angle with two spines:inner much larger than outer. Posterior margin rounded with bristlesand plumose setae.Eye large, round, and with sharp tooth on stout eyestalk.Antennule with well-developed stylocerite; slender, sharp-pointed,reaching nearly to end of second segment of peduncle. First segmentof peduncle produced ventrally into thin sharp tooth. Middle jointof peduncle nearly twice as long as first or third joints, whole pedunclemore than K as long as carapace. Inner flagellum slender, aboutas long as carapace. Outer flagella fused for )^ its length, with longsensory setae, and terminating in short free part and long slenderflagellum.Antenna with fu-st segment of peduncle produced ventrally intosharp tooth. Scale (fig. 57) slender; spine large, longer than blade,separated from slender blade for nearly }^ its length. Peduncle onlyslightly longer than that of antennule. Flagellum twice as long ascarapace: flattened oval in cross section.Third maxilliped with ischiomeropodite broad, flattened proximally,twisted. Inner margin heavily bristled, outer sparsely. Penultimatesegment K length of distal segment, with bristles mainly on inner side.Distal segment nearly as long as ischiomeropodite, armed mediallywith rows of stiff bristles; reaches to end of antennular peduncle.Exopodite longer than ischiomeropodite.Fu'st legs subequal in size. Ischium short, with sharp ridgeventrally. Merus short, stout, triangular in outlme: dorsal ridgesharp, covered with soft setae; outer ridge undulate with knobson proximal part, inner ridge tuberculate for most of its length.Broad oblique groove in which second leg rests. Distal margins ofmerus concave on inner side of median ridge. Joint membrane large,
SHRIMPS OF THE GENUS BETAEUS?HART 457making pocket into which swollen part of palm and projection ofcarpus fit, when "wrist" is bent. Carpus cup-shaped, with small plateventrally, covered with long soft setae dorsally. Chela (figs. 70, 71)large, as long as carapace and nearly as wide, much compressed later-ally with upper margin of base of fixed finger thin, sharp. Fixed fingertwice as wide at base as dactylus, which is longer than palm. Cutting-surfaces, tips of fingers, dorsal part of fixed finger setose. Left chelawith large tooth medially on fixed finger, wide gape, smaller teethintermeshing proximally and terminally before crossed corneous tips.Right chela with slight gape but most of small flat teeth intermeshing.Hands covered mth fine short setae.Second leg (fig. 72) setose with ischimn and merus subequal, shorterthan carpus, which has first and last joints subequal and twice as longas each of subequal median joints. Fmgers almost as long as palmand finely spinulate on cutting surfaces.Third leg (fig. 79) with stout ischium. Merus inflated with largemovable spine on proximal part. Merus and propodus subequal inlength. Carpus shorter than either, with pair of stout spines andsetae on distal ventral margin. Propodus with stout spines andsetae on ventral margin and two large spines terminally. Dactylus(fig. 80) short, broad, bifid, armed with few bristles. Whole ap-pendage setose dorsally.Fom"th leg similar to thu-d but somewhat smaller.Fifth leg subequal in length to third but without movable spine onmerus, no spines on carpus. Spines on propodus small, hidden bytransverse bands of setae forming brush on distal third of propodus.First pleopod with endopodite about K exopodite in length andwidth. Second pleopod with exopodite and endopodite slender,subequal. Appendix interna large, straplike.Uropod (fig. 68) with posterior margin of peduncle produced intotwo large teeth; outer distinctly larger than inner. Long bristles oncurved margin between teeth. Exopodite with long bristles dorsolat-erally and small tooth above stout spine on posterolateral angle ofproximal part. Endopodite and exopodite with scattered bristlesdorsally. Uropods longer than telson.Male.?Similar to female except that chelipeds (fig. 69) are largerin proportion, chela longer than carapace, antennular peduncleproportionately longer, abdomen somewhat narrower. First pleopodwith very small endopodite. Second pleopod with appendix masculinalarge and bristled throughout its length. Appendix interna small.Color.?"Nearly white" (Holmes, 1900) and red or brown withkelp, and green with sea lettuce and eelgrass (MacGinitie and Mac-Ginitie, 1 949) . These were identified as Betaeus harfordi but probablythey were B. setosus. Living specimens taken in British Columbia
458 PROCEEDINGS OF THE NATIONAL MUSEUM vol. u5
were a uniform white or yellow in the light but became pink or orangein the dark. The change in color is due to the presence of tiny redchromatophores thicldy dotted over the dorsal half of the animal(except over the eyes) and on the chelae. These expand in the dark.In some specimens there is a pale blue tinge to the chelae and theflagella. The cast skin is a very pale blue dorsally, with yellow setaeon the antennal blade and on the tail fan. The chitinous tips of thedactyli of the legs are yellow.Size.?Carapace length of smallest ovigerous female 4.8 mm.;largest female 8.5 mm.; males 3-7 mm. Length of chela of largestfemale 9 mm.; of largest male 9 mm.Range.?Morro Bay, Calif., to Hecate Strait, Queen CharlotteIslands, British Coliunbia.Habitat.?The ''B. harfof^di" fomid under a rock at low tide(Holmes, 1900), under rocks and among seaweed (MacGinitie andMacGinitie, 1949), and on pilings (Ricketts and Calvin, 1952), wereprobably Betaeus setosus. The one specimen taken in Washingtonwas found intertidally under a rock on the exposed west side of SanJuan Island. In British Columbia, one specimen was dredged at10 fathoms, some were found in tide pools, and others with Pachy-cheles rudis Stimpson in cavities mider the holdfasts of kelp (usuallyPterygophora californica Euprecht) or eelgrass roots (Phyllospadixspecies). They often occur in pairs.Material.?The following specimens were examined:California.?Morro Bay, under rocks, Feb. 1, 1939, S. A. Glassell, 2 ovigerousfemales.?Monterey Bay, China Point, shore, June 1908, 2 males (HopkinsMarine Station).?Monterey Bay, HMS Line Sui-vey, No. 86, June 2, 1931,G. E. MacGinitie, 2 males (Hopkins Marine Station).?Monterey Bay, J. C.Brown, 1 ovigerous female, 1 male (USNM 23932).?Tomales Point, Dec. 26,1939, E. F. Ricketts, 2 males.?Tomales Point, reef, Laviinaria, June 9, 1948,1 female, 1 male (Pacific Marine Station).?Tomales Point, ocean side, Nov. 28,1947, 1 female, 1 male (Pacific Marine Station).?Campbell's Cove, inside en-trance to Bodego Lagoon, Feb. 11, 1949, Lee O. Miles, 1 ovigerous female (PacificMarine Station).-?Dillon Beach, rocks, north, June 30, 1946, 1 ovigerous female(Pacific Marine Station).Washington.?San Juan Island, west side, under rocks, intertidal, July 10,1960, P. L. lUg, 1 male.British Columbia.?Vancouver Island, west coast: Clayoquot Sound, FrankIsland, Cox Bay, Tofino, July 10, 1960, J. F. L. Carl, 3 ovigerous females, 1 male;Clayoquot Sound, July 29, 1961, J. F. L. Carl, 1 female holotype (USNM 108229),5 females (3 ovigerous), 3 males; Clayoquot Sound, Round Island, June 30, 1945.E. F. Ricketts, 2 ovigerous females (USNM 80630); Clayoquot Sound, May 30,1946, E. F. Ricketts, 1 ovigerous female, 1 male (USNM 84395); ClayoquotSound, Deadman's Island, June 27, 1945, E. F. Ricketts, 1 ovigerous female,1 male (USNM 80635); Esperanza Inlet, High Island, near Catala Island andTatchu Point, June 28, 1934, E. G. Hart, 2 females (1 ovigerous), 1 male; Esper-anza Inlet, Nootka Island, tide pool, July 25, 1934, E. G. Hart, 2 ovigerous fe-
SHRIMPS OF THE GENUS BETAEUS?HART 459
males, 1 male.?Hecate Strait, Queen Charlotte Islands, 4 miles northeast RosePoint, 10 fathoms, June 24, 1956, T. H. Butler, 1 female.Notes.?There are a number of variations to be seen in this species.The inner ridge of the merus of the chelipeds may be almost smooth,or there may be a few tubercles, or there may be a row of uniformbeadlike knobs. The dentition of the chelae varies considerablyfrom flat intermeshed teeth, with no gape, to others with a foramenand/or a large square tooth on the fixed finger, but with the tips ofthe fingers always crossed (figs. 70, 71). Most have similar subequalchelae. To judge from specimens kept in the laboratory, regeneratedchelae may be without teeth and a nmnber of moults may be requiredto regain normal size.Ovigerous females have been taken February-June in Californiaand May-August in British Columbia. The ovaries and freslilylaid eggs are a pale olive-green, becoming lighter and more trans-parent with development. Literature CitedBaker, B. M.1912. Notes on the Crustacea of Laguna Beach. 1st Ann. Rep. LagunaMarine Lab., pp. 100-117, figs. 53-64.Banner, Albert H.1953. The Crangonidae, or snapping shrimp, of Hawaii. Pacific Sci., vol.7, no. 1, pp. 2-147, 1 pL, 50 figs.Dana, James Dwight1852; 1855. Crustacea. Vol. 13 in United States Exploring Expeditionduring the years 1838, 1839, 1840, 1841, 1842 under thecommand of Charles Wilkes, U.S.N. Philadelphia, part 1,685 pp. (1852); atlas, 27 pp., 96 pis. (1855).Glassell, Steve A.1938. New and obscure decapod Crustacea from the West American coasts.Trans. San Diego Soc. Nat. Hist. vol. 8, no. 33, pp. 411-454, pis.27-36.Hilton, W. A.1916. Crustacea from Laguna Beach. Journ. Ent. Zool., Pomona College,California, vol. 8, pp. 65-73, figs. 1-19.Holmes, Samuel J.1900. Synopsis of California stalk-eyed Crustacea. Occas. Pap. CaliforniaAcad. Sci., vol. 7, 262 pp., 4 pis.Holthuis, Lipke B.1952, The Crustacea Decapoda Macrura of Chile. No. 5 in Reports ofthe Lund University Chile Expedition 1948-49. Lunds Univ.Arsskr., new ser., avd. 2, bd. 47, no. 10, 110 pp., 19 figs.Johnson, M. E., and Snook, J. H.1927. Seashore animals of the Pacific coast, 659 pp., figs.
460 PROCEEDLNGS OF THE NATIONAL MUSEUM vol. 115KiNGSLEY, J. S.1878a. A synopsis of the North American species of the genus Alpheus,VII. BuU. U.S. Geol. Geogr. Surv. Terr., vol. 4, no. 1, pp.189-199.1878b. List of the North American Crustacea belonging to the suborderCaridea. Bull. Essex Inst., vol. 10, pp. 53-70.18S2. Carcinological notes, no. V. Bull. Essex Inst. vol. 14, nos. 7-12,pp. 105-132, 2 pis.LOCKINGTON, W. N.1877a. Remarks on the Crustacea of the Pacific coast, with descriptions ofsome new species. Proc. California Acad. Sci., vol. 7, pp. 28-36.1877b. Description of seventeen new species of Crustacea. Proc. CaliforniaAcad. Sci., vol. 7, pp. 41-48.1878. Remarks on some new Alphei, with a synopsis of the North-Americanspecies. Ann. Mag. Nat. Hist., ser. 5, vol. 1, no. 6, pp. 465-480.MacGinitie, G. E.1930. Notice of extension of range and of new species of various inverte-brates. Ann. Mag. Nat. Hist., ser. 10, no. 6, p. 68.1934. The natural history of Callia7iassa californiensis Dana. Amer.Midi. Nat., vol. 15, no. 2, pp. 166-177, pis. 5-6.1935. Ecological aspects of a California marine estuary. Amer. Midi.Natur., vol. 16, no. 5, pp. 629-765, figs. 1-21.1937. Notes on the natural history of several marine Crustacea. Amer.Midi. Natur., vol. 18, no. 6, pp. 1031-1037, 1 pi.MacGinitie, G. E., and MacGinitie, Nettie1949. Natural history of marine animals, 473 pp., 282 figs.Rathbun, M. J.1904. Decapod crustaceans of the northwest coast of North America.Harriman Alaska Expedition, vol. 10, 210 pp., 10 pis, 95 figs.Ricketts, Edward F., and Calvin, Jack.1952. Between Pacific tides, 3rd ed., 502 pp., 134 figs., 46 pis.Schmitt, Waldo L.1921. The marine decapod Crustacea of California. Univ. CaliforniaPubl. Zool., vol. 23, 470 pp., 165 figs., 50 pis.1924. Crustacea (Macrura and Anomura). In Expedition of the CaliforniaAcademy of Science to the Gulf of Cahfornia in 1921. Proc.California Acad. Sci., ser. 4, vol. 13, pp. 381-388.
SHRIMPS OF THE GENUS BETAEUS?HART 461
Figures 1-16.
?
Betaeus harrimani Rathbun: 1, lateral view; 2, antennular peduncle, lateralview; 3, mandible; 4, maxillule; 5, maxilla; 6, first maxilliped; 7, second maxilliped; 8,third maxilliped; 9, fourth leg; 10, fifth leg; 11, right chela, female; 12, left chela, male;13, left chela, female; 14, right chela, female; 15, first pleopod, male; 16, second pleopod,male.
462 PROCEEDINGS OF THE NATIONAL MUSEUM
Figures ll-l'i.?Betaeus harrimani Rathbun: 17, frontal region, dorsal view, female; 18,same, lateral view; 19, antennal scale; 26, tail fan. Betaeus longidactylus Lockington:20, frontal region, dorsal view, female; 21, same, lateral view; 22, antennal scale; 27, tailfan. Betaeus ensenadensis Glassell: 23, frontal region, dorsal view, male; 24, same, lateralview; 25, antennal scale; 28, tail fan.
SHRIMPS OF THE GENUS BETAEUS?HART 463
Figures 29^S.?Betaeus harrimani Rathbun: 29, right cheliped, male; 30, left chela,female; 31, right chela, same; 37, second leg; 38, third leg; 39, dactylus of third leg.Betaeus longidactylus Lockington: 32, right cheliped, female; 33, right chela, female;34, left chela, male; 40, second leg; 41, third leg; 42, dactylus of third leg. Betaeusensenadensis Glassell: 35, right cheliped; 36, right chela, male; 43, second leg; 44, thirdleg; 45, dactylus of third leg.
464 PROCEEDINGS OF THE NATIONAL MUSEUM
Figures ^G-S1.?Betaeus harfordi (Kingsley): 46, frontal region, dorsal view, female; 47,same, lateral view; 54, antennal scale. Betaeus macginitieae, new species: 48, frontalregion, dorsal view, female; 49, same, lateral view; 55, antennal scale. Betaeus gracilis,new species: 50, frontal region, dorsal view, female; 51, same, lateral view; 56, antennalscale. Betaeus setosus, new species: 52, frontal region, dorsal view, female; 53, same,lateral view; 57, antennal scale.
SHRIMPS OF THE GENUS BETAEUS?HART 465
Figures 58-72.
?
Betaeus harfordi (Kingsley): 58, tail fan; 59, right cheliped, male; 60,right chela, female; 61, second leg. Betaeus macginiiieae, new species: 62, tail fan; 63,right cheliped, male; 64, second leg. Betaeus gracilis new species: 65, tail fan; 66, rightcheliped, female; 67, second leg. Betaeus setosus, new species: 68, tail fan; 69, rightcheliped, male; 70, right chela; 71, right chela; 72, second leg.
466 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lis
Figures 73-80.
?
Betaeus harfordi (Kingsley): 73, third leg; 74, dactylus of third leg.Betaeus inacginitieae, new species: 75, third leg; 76, dact)'lus of third leg. Betaeus gracilis,new species: 77, third leg; 78, dactylus of third leg. Betaeus setosus, new species: 79,third leg; 80, dactylus of third leg.
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