Notopontonia Platycheles, New Genus, New Species

JOURNAL OF CRUSTACEAN BIOLOGY, 11(4): 607-628, 1991

NOTOPONTONIA PLATYCHELES, NEW GENUS, NEW SPECIES (DECAPODA: PONTONIINAE) FROM SOUTH AUSTRALIA, WITH REMARKS ON PONTONIA PINNOPHYLAX (OTTO), THE TYPE SPECIES OF PONTONIA LATRETTIJ,

A. J. Bruce

ABSTRACT Notopontoniaplatycheles new genus, new species (Decapoda:Pontoniinae), is describedand illustrated.Caught off Robe, South Australia,at a depth of 64 m, and of unknowncommensal associations,this species increasesto five the numberof pontoniineshrimps known from South Australia.Assessment of the systematicrelationships of Notopontoniainvolved study of Pon- tonia pinnophylax(Otto, 1821), the type species of the genus Pontonia, which is redescribed and illustrated,leading to the discovery of an interestingmorphological feature of the feeding mechanismnot previouslyreported in any palaemonidshrimps.

The casual examination of the alpheid Carapacelength refers to the postorbital shrimp collection of the South Australian carapacelength. Museum, Adelaide, led to the discovery of an unusual shrimp belonging to the Pon- SYSTEMATICACCOUNT toniinae, a subfamily very poorly repre- sented in southern Australian waters. Fur- Notopontonia,new genus ther search unfortunatelyfailed to provide Definition.-Commensal shrimps of un- additional specimens. Although closely re- known association. Carapace strongly de- lated to the genus Pontonia Latreille, 1829, pressed, smooth, with short broad trian- the new species presents several small but gular rostrum, dorsally concave, lacking distinctive features, sufficient to justify its dorsal and ventral carinae, unarmed, dis- exclusion from that genus. A new genus is, tally setose; supraorbital,epigastric, hepat- therefore,described for its reception. ic, and antennal spines absent, orbit obso- Comparison with specimens of the type lete, inferior orbital angle large, blunt, species of the genus, Pontonia pinnophylax anterolateralangle of branchiostegite pro- (Otto) (Opinion 378, 1956) confirmed the duced, angular; abdomen smooth, pleura presenceof several distinctive charactersof rounded;telson broad, with 2 pairs of large generic value, including a remarkablepar- dorsal spines, 2 pairs of small posterior agnathalfeature that has not been described spines;antennule with short stylocerite,up- in any previously reported pontoniine or, per flagellumfeebly biramous;antenna with indeed, palaemonid shrimp. This unusual basicerite unarmed, scaphocerite well de- feature is clearly lacking in the new genus. veloped, distolateral tooth enlarged, epi- The type species of the genus Pontonia La- stome normal, paragnathwithout proximal treille was first described by Otto (1821) as median fissure; mandible normal, without Palaemonpinnophylax, in an academicthe- palp; maxillula with biolobed palp, upper sis. Subsequently, an account as Alpheus lacinia broad, lower lacinia slender;maxilla pinnophylax,with colored illustrations,was with short simple palp, basal endite bilobed, provided (Otto, 1828). The shrimps pres- upper lobe greatly enlarged, dorsally con- ently included in the genus Pontonia show cave, coxal endite obsolete, scaphognathite a wide range of variation in many specific normal; first maxilliped with slender palp, characters,which probablyindicate a poly- basal and coxal endites fused, elongate,dor- phyletic origin for that genus. sally concave, medial fringe of short setae, A new description of P. pinnophylax is, exopod well developed, carideanlobe large, therefore,provided, which may be useful as narrow,epipod bilobed; second maxilliped a yardstickagainst which the systematic re- with dactylar segment reduced, propodal lationships of the other species of Pontonia segmentenlarged, dorsally concave, exopod may be assessed. well developed, epipod large, triangular, 607 608 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991

without podobranch; third maxilliped with antennal spine. These species usually have ischiomerus and basis completely fused, simply biunguiculate ambulatory dactyls, as broad, exopod well developed, lateral plate in Notopontonia. Other species, with ob- normal, arthrobranch rudimentary; first solete orbits, lacking antennal spines, gen- pereiopods slender, chela with fingers elon- erally have complex biunguiculate dactyls, gate, subspatulate; fourth thoracic sterite and are usually associated with ascidians. without fingerlike median process; second (4) The maxilla has a characteristic basal pereiopods with large similar subequal che- endite, bilobed, with the upper lobe greatly lae, strongly compressed, carried horizon- enlarged and dorsally concave. In Pontonia tally with dactyl laterally, ventral margin these lobes are generally subequal, small, strongly carinate, dorsal margin feebly car- frequently much reduced. The fused endites inate, fingers without molar process and fos- of the first maxilliped in Notopontonia are sa, feebly dentate, proximal segments un- centrally expanded, with a medial fringe of armed; ambulatory pereiopods with dactyl short setae, and the epipod is bilobed. The simply biunguiculate, propod with disto- second maxilliped has the dactylar segment ventral spines; endopod of first pleopod reduced and the propodal segment enlarged curved distomedially, endopod of second in comparison with species of Pontonia. pleopod with appendix masculina exceed- (5) The paragnath in Notopontonia is sim- ing appendix interna; uropod with protopod ply bilobed, without distinct accessory lobes distolaterally unarmed; exopod with small and proximal median fissure, as in P. pin- distolateral tooth, with larger mobile spine nophylax. medially. (6) The first pereiopods have elongate spatulate fingers in Notopontonia, whereas Type Species. -Notopontonia platycheles, in P. pinnophylax they are compressed, new species. nonspatulate. Systematic Position. -The genus Notopon- (7) The posterior margin of the telson is tonia is undoubtedly closely related to Pon- provided with two pairs of spines only in tonia Latreille. Notopontonia may be most Notopontonia, whereas species of Pontonia readily distinguished from Pontonia by the are usually provided with the typical pon- following features: toniine complement of three pairs. The ex- (1) The chelae of the second pereiopods opod of the uropod in Notopontonia bears are very strongly compressed and carried in a small distolateral tooth, as in most of the a horizontal plane, with the dactyl laterally, Pontoniinae, but which is usually absent with very strongly carinate ventral margins from species of Pontonia. and a feebly carinate lateral margin to the (8) Where information is available, the dorsal palm, with feebly armed fingers, appendix intera of the male second pleo- crossing only at the tips. In Pontonia, the pod distinctly exceeds the appendix mas- chelae generally have the palm oval in sec- culina in species of Pontonia. In Notopon- tion, although some may have a small ven- tonia, the appendix masculina exceeds the tral carina, particularly in species with dis- appendix intera. similar chelae, the chelae generally being Etymology.--From notos, Greek, south, and held in an oblique or near vertical plane, Pontonia, a generic name first used for a with the dactyl ventral, in an extended or pontoniine shrimp by Latreille (1829). semiextended rather than a retracted position. Notopontonia platycheles, (2) The rostrum is broad, dorsally con- new species 14E cave, completely lacking dorsal and ventral Figs. 1-6, carinae. In Pontonia, the rostrum is vari- Material Examined.-1 $, northwestof Robe, South able, frequently compressed and often mi- Australia,approximately 36?53'S, 139?53'E,in 64 m, nutely dentate distally, never dorsally con- September1985, collected by Sangster. cave. Description. -Medium-sized, robust pon- (3) The orbit is obsolete, with the anten- toniine shrimp, of strongly depressed body nal spine lacking. Many species of Pontonia form. Carapace smooth, glabrous, strongly have a small distinct orbit, with a definite depressed, rostrum well developed, broadly inferior orbital angle and well-developed triangular, about 1.5 times broader than BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 609

Fig. 1. Notopontonia platycheles, new genus, new species, holotype male, off Robe, South Australia, lateral. Scale divisions = 3 mm.

long, slightly exceeding intermediate seg- 0.5 of anterior margin width, with 2 pairs ment of antennular peduncle, unarmed, of lateral spines only, small outer spine, with without distinct dorsal or ventral carinae, largerblunt spine medially, 1.75 times length strongly compressed dorsoventrally, slight- of lateral spine, 2.8 times longer than basal ly thickened centrally, dorsally concave, lat- width, central posterior margin with about eral margins convex, with pair of conspic- 30 long simple setae, 2 long plumose setae. uous setae distoventrally; supraorbital, Antennule with proximal peduncular seg- epigastric, hepatic, and antennal spines ab- ment broad, about 1.25 times longer than sent; orbital notch well developed, without proximal width, distolateral angle strongly defined orbit, inferior orbital angle large, produced, broadly acute, stylocerite short, slightly upturned, blunt; anterolateral angle broad, moderately acute, reaching to about of carapace strongly produced, acute; ven- 0.5 of segment length, statocyst normal with tral branchiostegite feebly angulate ventro- granular statolith; ventromedial margin with medially. small acute tooth at 0.5 of length; inter- Abdomen smooth, glabrous, subcylindri- mediate segment short, broad, about 1.2 cal; pleura of first 3 segments broadly times broader than long; distal segment 1.3 rounded, fourth and fifth rounded, small; times longer than broad, distal segments sixth segment about 1.2 times length of fifth, combined about 0.6 of proximal segment depressed, 1.8 times broader than long, 2.0 length; antennular flagella carried extended; times longer than anterior depth, postero- upper flagellum short, biramous, rami fused ventral angle large, subacute, posterolateral proximally for 11 segments, robust, shorter angle small, acute. Telson about 2.0 times free ramus with single segment, longer ra- longer than maximum width at about 0.1 mus about 0.6 of fused portion, slender, with of length, lateral margins convex, feebly 5 segments; with about 16 groups of aesthe- converging posteriorly, anterior margin tascs; lower flagellum short, slender, sub- about 0.83 of width, dorsal surface deeply equal to long ramus of upper flagellum, about concave, with 2 pairs of large subequal lat- 0.2 of carapace length, with 13 segments. eral dorsal spines, about 0.15 of telson Antenna with short, stout, unarmed basi- length, at 0.16 and 0.6 of telson length; pos- cerite, ischiocerite and merocerite normal, terior margin broad, feebly biconvex, about carpocerite about 4.5 times longer than dis- 610 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991

-,?1;?.??? ?':..:?????':?: "? .-.?..??? ?::??.:?i..?'-"'-????::: :.':1::?::?:??.:1:? .?.???:: ??.?':: ii?'-:?. ? '??;.:.?.? '`"'" .?:???.-.??.? ' ?...?.,??????; ???:????? ?????_:?-,?::? ?-'?1:?':??...v ? ?.:;.l'r:?.???"?'. ???" ???,.

Fig. 2. Notopontonia platycheles, new genus, new species, holotype male, off Robe, South Australia, dorsal. Scale divisions = 3 mm. BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 611

2.0mm I IV

1..Omm 2.0mm CD j ^G

i H D

I0.5mm HI

^ ?QJ)~~~

E A Fig. 3. Notopontonia platycheles, new genus, new species, holotype male. A, anterior carapace, eyes, left antennae, dorsal; B, anterior carapace, lateral; C, antennule; D, antenna; , buccal region, ventral;tsixth , abdominal segment, dorsal;G, telson; H, same, posteriorspines; I, uropod;J, right eye, dorsal.

tal width, uniform, compressed, distinctly of tip of rostrum, broad, about 1.5 times exceeding rostrum, flagellum short, slender, longer than wide, distal lamina broadly subequal to carapace length; scaphocerite rounded, lateral margin convex, with enor- well developed, lamina extending to level mous curved distolateral tooth, arising at 612 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991

I .Omm I

Fig. 4. Notopontonia platycheles, new genus, new species, holotype male. A, mandible; B, maxillula; C, maxilla; D, first maxilliped; E, second maxilliped; F, third maxilliped.

about 0.5 of lamina length, far exceeding sixth to eighth unarmed, progressively distal margin of lamina, separated by deep broadening posteriorly. fissure. Mouthparts (right): mandible without Eyes small; cornea hemispherical, well palp, corpus normal; molar process robust, pigmented, with discernible accessary pig- obliquely truncate distally, with 2 blunt teeth ment spot, diameter about 0.9 of carapace dorsally, 2 rounded laminar plates ventral- length; stalk subcylindrical, subequal to cor- ly, single acute tooth posteriorly, with small neal diameter, about 1.2 times longer than tufts of setae anteriorly and posteriorly; in- broad. cisor process feeble, obliquely truncate dis- Epistome normal, unarmed; ophthalmic tally, with 5 small acute teeth, sublateral somite simple, without bec ocellaire. La- tooth smaller than others. Maxillula with brum without special features. Paragnath small weakly bilobed palp, lower lobe ta- about 1.25 times broader than long, alae pering distally, with small distal tubercle well developed, feebly lobulate, with deep with short simple seta; upper lacinia broad, hollow at proximal end of median fissure; dorsally concave, with double rows of about basal portion short, broad, with central em- 11 short small finely denticulate spines dis- inence, without distinct lateral ridge or deep tally, short plumose setae ventrally; lower median groove. First three thoracic sternites lacinia slender, tapering distally, with 3 dis- broad, unarmed, fourth narrow, unarmed, tal spines, numerous simple setae. Maxilla fifth narrow, with strong transverse carina with short simple palp, basal endite well posterior to coxae of second pereiopods, developed, dorsally concave, bilobed, distal BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 613

H G

Fig. 5. Notopontoniaplatycheles, new genus, new species, holotype male. A, first pereiopod;B, same, chela; C, second pereiopod, left; D, same, chela, dorsal; E, same, right chela, dorsal; F, third pereiopod; G, same, propod and dactyl; H, fifth pereiopod, propod and dactyl; I, same, dactyl and distal propod, medial; J, first pleopod; K, second pleopod. lobe greatly enlarged, broadened, both lobes setose palp, basal and coxal endites fused with dense fringe of long marginal setae, forming large rounded lobe, with dense minutely serrulate, coxal endite obsolete; fringe of minutely serrulate marginal setae, scaphognathite normal, about 3.0 times concave dorsally and laterally thickened; longer than broad, posterior lobe large, about exopod well developed, flagellum with nu- 1.25 times longer than broad, 0.3 of sca- merous plumose setae distally, caridean lobe phognathite length, anterior lobe strongly large, broad; epipod unequally bilobed, tapering distally, medial margin concave. proximal lobe larger. Second maxilliped with First maxilliped with slender, simple non- dactylar segment small, about 4.0 times lon- 614 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991

0.5mm (0.1mmI F I

t .ant B

X 1., C-:~1, I 0.5mm I I GUI/1~ 1 I

0.mm f[ H E E I BCD I C Fig. 6. Notopontoniaplatycheles, new genus, new species, holotype male. A, antennal peduncle, proximal segment, distolateralangle; B, mandible, molar process; C, same, incisor process;D, maxillula, palp; E, first pereiopod,finger tips, dactyluppermost; F, thirdpereiopod, dactyl; G, firstpleopod, endopod; H, secondpleopod, appendix masculinaand appendix intera; I, uropod, expod, distolateralangle.

ger than broad, medially densely fringed with tapering distally, densely setose medially; coarsely serrulate setae; propodal segment terminal segment about 0.9 of penultimate large, 2.0 times length of dactylar segment, segment length, about 3.5 times longer than deeply concave dorsally, distomedial mar- proximal width, tapering distally, with nu- gin broadly rounded, densely fringed with merous transverse rows of short finely setu- long minutely serrulate setae; carpus, is- lose spiniform setae ventromedially; coxa chiomerus normal; basis normal, feebly ex- robust, feebly produced medially, with large cavate dorsally, with well-developed exo- subrectangular lateral plate; with rudimen- pod, with numerous plumose setae distally; tary arthrobranch. coxa feebly produced medially, sparsely se- First pereiopods slender, distal end of tose, with large simple triangular epipod lat- merus exceeding antennular peduncle; chela erally, without podobranch. Third maxil- slender with short, subcylindrical, feebly liped with endopod extending to about compressed palm, about 1.6 times longer middle of carpocerite; ischiomerus and ba- than deep, fingers elongate, broad, subspat- sis completely fused, combined segment ulate, about 2.0 times palm length, dactyl broad, about 3.3 times longer than central slightly longer than fixed finger, with 2 small width, dorsally concave medially, medial teeth distally, entire laminar cutting edge margin with feeble convexities in ischial and laterally, fixed finger similar single tooth basal regions, with numerous slender feebly distally, fingers with numerous groups of setulose setae, lateral margins sparsely se- long serrulate setae; carpus slender, about tose, longer setae; penultimate segment 1.6 times chela length, 7.2 times longer than about 0.45 of proximal segment length, distal width, tapering proximally, proximal about 2.3 times longer than proximal width, width about 0.5 of distal width; merus fee- BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 615 bly bowed about 0.95 of carpuslength, uni- lobe, unarmed; merus 1.2 times propod form, about 6.0 times longer than width; length, 3.0 times longer than central depth, ischium about 0.45 of merus length; basis unarmed; ischium about 0.9 of propod about 0.33 of merus length, coxa robust, length, 2.3 times longer than distal width, with small setose ventral lobe. tapered proximally, unarmed; basis about Second pereiopods well developed, che- 0.6 of propod length; coxa robust, without lae carriedretracted, with carpomeraljoint specialfeatures. Fourth and fifthpereiopods flexed, merus and ischium lying alongside similar to third;fifth propod 1.1 times third carapace, chelae carried horizontally with propod length, about 6.0 times longer than dactyls laterally; chelae subequal, similar, proximaldepth, with transverserow of short about 1.3 times carapace length, palmar serrulatesetae distolaterally,distal and pre- portion about 1.6 times longer than proxi- terminal ventral spines medially. mal width, strongly compressed, smooth, First pleopod with basipodite about 2.6 tapering slightly distally, with weak carina times longer than distal width; endopod proximolaterally, ventral margin enlarged about 0.55 of exopod length, 3.4 times lon- into broad fixed tooth, very strongly com- ger than central width, with distal portion pressedwith feebly irregularlycarinate ven- curved medially, distal and lateral margins tral margin; dactylus slender, oval in sec- with about 20 short plumose setae; central tion, carinate,outer marginfeebly irregular, third of medial margin with 4 longer plu- feebly curved, about 0.8 of palm length, tip mose setae proximally, about 12 short sim- acute, not hooked, distal cutting edge con- ple slender spines distally. Second pleopod cave, entire, blunt with 3 small acute teeth with basipodite about 2.2 times longer than proximally on left, 2 on right; fixed finger distal width; endopod about 1.1 times ba- very broad,about 0.5 of palm length,as long sipodite length, 4.0 times longerthan broad, as deep, with small acute tip, distal half of appendices at about 0.3 of medial margin cuttingedge situateddorsally, convex, sharp, length, proximal margin convex, with plu- separatedby blunt concave diastema from mose setae; appendix masculina subcy- 3 small proximal teeth on left, 2 on right: lindrical,slightly swollen centrally, 5.5 times carpus stout, distally enlarged, tapering longer than central width, with 8 ventro- strongly proximally, about 0.45 of palm medial and 10 ventrolateralsimple setae, 1 length, length subequalto distal width, fee- longer, finely serrulatedistal seta; appendix bly excavate distally, unarmed, left carpus interna slender, with few distal cincinnuli, with small acute tooth proximolaterally; slightly shorter than appendix masculina. merus robust, about 0.55 of palm length, Uropod with protopodite short, stout, 1.6 times longer than central width, cen- unarmed; exopod not exceeding telson, trally swollen, unarmed;ischium unarmed, about 2.5 times longer than wide, lateral about 0.28 of palm length; basis normal, margin convex, with small acute tooth dis- coxa stout. tally, with largermobile spine medially;en- Ambulatory pereiopods moderately ro- dopod subequalto exopod length, 2.6 times bust, third exceedingrostrum by half carpus longer than broad. length, dactyl about 2.8 times longer than -The male is des- about 0.36 of Types. single specimen proximaldepth, unguis corpus ignated as the holotype and is held in the length, 3.0 times longer than basal width, collection of the South Australian 1.8 times Museum, feebly curved, corpuscompressed, catalogue number C.4240. longer than proximal depth, without basal process,scarcely tapering distally, with large Measurements (mm).-Total body length acute distoventralaccessory tooth, with sin- (approximate), 36; carapace and rostrum, gle short medial and lateral sensory setae 15.5; postorbital carapace, 13; second pe- distally; propod about 0.33 of carapace reiopod chelae, left, 17; right, 18. length, robust, 4.2 times longer than prox- Coloration.-No data. imal distally, ventral depth, feebly tapering Host. -Unknown. marginwith 2 small slender spines distally, with single largerstouter spine proximally; Etymology.-From platys, Greek, flat, and corpus about 0.6 of palm length, 2.4 times chele, Greek,a claw, a referenceto the shape longer than width, with small distodorsal of the second pereiopods. 616 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991

Pontonia pinnophylax (Otto, 1821) rounded. Telson about 2.0 times length of Figs. 7-13, 14F sixth segment, about 2.3 times longer than broad, lateral margins convex, convergent, anterior margin about 0.8 of greatest width; Palaemon pinnophylax Otto, 1821: 12. with 2 dorsal about Alpheus pinnophylax Otto, 1828: 341, pl. 21, figs. pairs ofsubequal spines, 1, 2. 0.09 of telson length, at 0.4, 0.66 of telson Pontonia pinnophylax Holthuis, 1947: 319 (full syn- length, submarginal; posterior margin fee- onymy). bly concave, 0.4 of anterior margin width, Material Examined. -1 6, 1 Q. Straits of Messina, with 3 pairs of small spines, lateral spines 38?15'N, 15?36'E, depth ?, 8 December 1988, in Pinna short, about 0.4 of dorsal spine length, in- collected N. Calafiore and G. nobilis, by Constanzo, termediate spines more robust, broken, sub- Northern Territory Museum (NTM) Cr.006750. median spines more slender, broken. Description of Male. -Medium-sized pon- Antennule small, peduncle distinctly ex- toniine shrimp of normal subcylindrical ceeding rostrum, proximal segment about body form. Carapace smooth, glabrous, with 1.8 times longer than broad, tapering dis- well-developed rostrum extending anteri- tally, medial margin sparsely setose, with orly to slightly exceed intermediate segment minute ventromedial tooth at 0.5 of length, of antennal peduncle, triangular in dorsal lateral margin sinuous, distolaterally pro- view, about 2.3 times longer than broad, duced, subtruncate, with weak rounded dis- dorsally convex, unarmed, without dorsal tolateral tooth, stylocerite short, broad, carina, distal portion slender, compressed, acute, reaching to 0.5 of segment length, with distinct ventral carina, unarmed, tip statocyst normal, with granular statolith; in- acute, with pair of simple preterminal dor- termediate segment about 1.8 times longer sal setae; supraorbital, epigastric, hepatic than broad, 0.55 of proximal segment length, spines absent; orbit distinct, feebly devel- with small lateral lamella, obliquely artic- oped, with small blunt inferior orbital angle, ulated with distal segment; distal segment antennal spine well developed, slender, about 0.9 of intermediate segment length, acute, marginal; anterolateral angle ofbran- 1.3 times longer than broad; upper flagellum chiostegite not produced, broadly rounded. feebly biramous, proximal 9 segments fused, Abdomen smooth, glabrous; sixth seg- robust, shorter free ramus with single segment depressed, about 1.4 times longer than ment, longer free ramus with 6 slender seg- fifth, as long as wide, posteroventral and ments; about 6 groups of aesthetascs pres- posterolateral angles acutely produced; ent; lower flagellum short, subequal to longer pleura of first 3 segments broadly rounded, upper ramus flagellum, with 16 slender seg- fourth, fifth, slightly produced posteriorly, ments.

Fig. 7. Pontonia pinnophylax (Otto), male, lateral, Straits of Messina. Scale divisions = 3 mm. BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 617

Fig. 8. Pontonia pinnophylax (Otto), male, dorsal, Straits of Messina. Scale divisions = 3 mm. 618 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991

1.0mm ABCD

.Omm EFIL J

I 0.5mm JK

2.0mm I A H1

H Fig. 9. Pontonia pinnophylax(Otto). A, anteriorcarapace and rostrum,eye, antennae,right orbit, dorsal; B, anteriorcarapace, rostm, eye, right orbit, dorsal;C, D, anteriorcarapace and rostrum,lateral; E, antennule; F, antenna;G, eye, dorsal;H, sixth abdominalsegment, dorsal; I, telson;J, K, same, posteriorspines; L, uropod. A, C, E-J, male; B, D, K, female.

Antenna with stout unarmed basicerite, pressed, 4.0 times longer than distal width, with small antennal gland tubercle medi- flagellum short, slender, about 1.6 times ally; ischiocerite, merocerite normal; car- carapace length; scaphocerite well devel- pocerite subcylindrical, slightly com- oped, broad, 1.8 times longer than wide, BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 619

1.0mm B-G

1.0mm I All B F

Fig. 10. Pontonia pinnophylax (Otto), male. A, paragnaths, ventral; B, mandible; C, maxillula; D, maxilla; E, first maxilliped; F, second maxilliped; G, third maxilliped; H, same, coxal region, lateral. extending well beyond antennal peduncle, about 0.075 of carapace length, without ac- lateral margin distinctly convex, with small cessary pigment spot, stalk subcylindrical, acute distal tooth, slightly exceeded by feebly compressed, about 1.2 times longer weakly angulate distal margin of lamella. than broad, 1.1 times corneal diameter. Eye with small hemispherical cornea, Epistome and labrum without special fea- 620 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991

- 3.0mm I 1.0mm I 1, I M I

N J

B 0

F Fig. 11. Pontonia pinnophylax (Otto). A, first pereiopod; B, same, chela; C, major second pereiopod; D, same, major chela; E, same, fingers; F, minor second pereiopod, chela; G, same, fingers; H, major second pereiopod, chela; I, same, fingers; J, minor chela same; K, same, fingers; L, third pereiopod; M, same, propod and dactyl; N, first pleopod; 0, second pleopod. A-G, L-O, male; H-K, female.

tures. Paragnath well developed, about 1.25 proximal half narrower with deep median times longer than broad, deeply bilobed dis- groove, bordered laterally by raised setose tally, with large subcircular smooth nonse- carinae, most marked anteriorly. Anterior tose lobes, with 2 pairs of smaller subme- thoracic sternites broader, second with dian ventral lobes, with depressed centers; rounded transverse median carina, third BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 621

0.5mm A

0.2mm I IKLN I I I

B B

0.5mm 1 BDEFG I

0.2mm -'? C

.+ :1. I;: c''':

".- f .:."?tr,?-

.?L?'::?;i:, rr:''?'???-'

0.5mm E IM

Fig. 12. Pontonia pinnophylax (Otto). A, rostrum; B, C, antennule, distolateral angle of proximal peduncular segment; D, scaphocerite, distolateral tooth; E, mandible, molar process; F, same, incisor process; G, maxillula, palp; H, second to fourth thoracic sternites; I, first pereiopod, tips of fingers (setae omitted); J, second pereiopod, proximal tooth of fixed finger; K, L, third pereiopod, dactyl; M, first pleopod, endopod; N, second pleopod, appendix masculina, and appendix interna; 0, uropod, distolateral exopod. A, C, L, female; B, D, E-K, M-O, male. 622 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991

0.5mm EF

l.Omm B

2.0 D E

0.2mm AC

B Fig. 13. Pontonia pinnophylax (Otto), female, Ulnis an der Schlei. A, tip of rostrum; B, minor second pereiopod, chela; C, third pereiopod, dactyl and distal propod; D, telson; E, same, posterior spines; F, male, Straits of Messina, tip of rostrum. BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 623

0.5mm I CD I

.I mm ABEF D

Fig. 14. Macrobrachium rosenbergii (de Man), immature male, Northern Territory, Australia. A, oral region, = = ventral (A ala of paragnath, B peduncle of paragnath, C = origin of first maxilliped, D = corpus of mandible, = E incisor process of mandible, F = mandibular palp, G = labrum); B, paragnaths, ventral aspect. Palaemonella rotumana (Borradaile), male, Coburg Peninsula, Norther Territory. C, paragnaths. Pontonia katoi Kubo, oviger- ous female, Heron Island, Queensland. D, paragnaths. Notopontoniaplatycheles, new genus, new species, holotype male. E, paragnaths, ventral. Pontonia pinnophylax (Otto), male, Straits of Messina. F, same, left lateral. with 2 lateral carinae, fourth with 2 lateral anteriorly, upper lacinia feebly expanded, carinae, without median process, fifth with 1.7 times longer than broad, distally curved well-developed lateral plates posterior to dorsally, distal margin curved continuously second pereiopod coxae, sixth to eighth un- with ventral margin, with about 15 minute armed, broadening posteriorly. simple spines distally, largely obscured by Mandible (right) normal, corpus robust, numerous short simple setae, extending without palp; molar process stout, with 2 along ventral border, lower lacinia large, large blunt teeth dorsally, 2 rounded, sharp- markedly broadened, thickened, distally edged teeth ventrally, one subacute tooth acute, curved dorsally, densely covered with posteriorly; incisor process slender, distally short simple setae distally, ventrally; max- truncate, with 5 small acute teeth distally, illa with simple palp, expanded proximally, longest teeth ventrally; maxillula with fee- tapered distally, with short plumose setae bly bilobed palp, larger lower lobe with small along lateral margin, basal endite well de- ventral tubercle with single short recurved veloped, bilobed, distal lobe longer than simple seta, with longer slender simple setae proximal, with about 22 long slender setae, 624 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991 minutely serrulate distally, proximal lobe chela 1.5 times longer than deep, subcy- with about 15 similar, slightly shorter setae; lindrical, slightly compressed, dactyl about coxal endite obsolete, medial margin con- 1.25 times palm length, 4.5 times longer vex, nonsetose; scaphognathite large, 2.7 than proximal depth, slender, tapering, times longer than wide, posterior lobe large, compressed, with low entire lateral cutting 1.5 times longer than wide, anterior lobe edge, tip acute with single small spine, fixed 1.3 times longer than wide, medial margin finger similar, with 2 small distal spines, concave; first maxilliped with simple, non- both fingers with numerous transverse rows setose palp, closely adpressed around dis- of long densely serrate setae laterally; carpus tolateral margin of basal endite, basal and about 1.6 times chela length, 5.5 times lon- coxal endites completely fused, combined ger than distal width, tapering proximally, endite elongate, narrow, dorsally concave, unarmed; merus subequal to carpus length, with dense fringe of long finely plumose se- 6.0 times longer than central width, slightly tae along medial margin, with row of more compressed, uniform, unarmed; ischium 0.6 sparse, longer, simple submarginal setae of merus length, ventrally carinate, setose; ventrally; exopod well developed, flagellum basis 0.4 of merus length, sparsely setose broad, with numerous plumose setae dis- ventrally; coxa normal, without ventral pro- tally, caridean lobe large, elongate, narrow, cess. epipod large, suboval; second maxilliped Second pereiopods well developed, che- with normal endopod, dactylar segment lae large, robust, subequal, similar; major about 4.0 times longer than broad, densely chela (right) about 1.9 times carapace length, fringed with coarsely serrulate spiniform se- smooth, subcylindrical, slightly swollen tae medially, dactylar segment with disto- centrally, oval in section, without ventral medial lobe small, with numerous longer carina, 2.3 times longer than deep, dactyl finely serrulate spiniform setae, ischiome- slender, curved, with slightly hooked blunt rus, with setae along medial margin, basis tip, about 0.5 of palm length, about 5.0 times feebly excavate medially, exopod normal, longer than proximal depth, distal half of with numerous plumose setate distally, coxa cutting edge concave, blunt, with small notch without medial protuberance, with large proximally, proximal half of cutting edge suboval epipod laterally, without podo- with large acute compressed tooth, fixed fin- branch; third maxilliped extending to about ger subequal to dactyl length, 2.5 times lon- level of middle of carpocerite; ischiomerus ger than proximal depth, tip slightly hooked, almost completely fused to basis, junction blunt, distal half of cutting edge with large indicated by small protuberance on medial low acute tooth, separated by deep diastema margin at proximal end of ischium, com- from large convex tooth with about 7 small bined segment about 3.0 times longer than denticles on dorsal margin, medial aspect proximal width, tapering distally, flattened, of fixed finger with fossa for reception of anterior width about 0.4 of proximal width, dactylar tooth when fingers closed, tips ex- basal margin feebly convex, with long sim- tensively crossing with fixed finger diastema ple setae, ischiomeral medial margin simi- opposing small predental dactylar notch with lar, proximal lateral margin sparsely setose; dactylus crossing over fixed finger centrally: penultimate segment about 0.45 of proxi- minor chela (left) about 1.75 times carapace mal segment length, subcylindrical, with length, 0.87 of major chela length, palm numerous groups of long finely serrulate se- about 2.3 times longer than central depths, tae ventromedially, terminal segment about dactyl 0.5 of palm length, fingers as in major 0.8 of intermediate segment length, 4.5 times chela; carpus about 0.25 of palm length, as longer than proximal width, tapering dis- long as distal depth, strongly tapered prox- tally, with numerous groups of longer, imally, excavate distally, unarmed; merus coarsely serrulate spines distally; exopod about 0.45 of palm length, 2.1 times longer well developed, exceeding distal merus, with than central depth, feebly excavate disto- numerous plumose setae distally; coxa ventrally, unarmed; ischium about 0.3 of without medial process, with large oval lat- palm length, 1.6 times longer than distal eral plate, with rudimentary arthrobranch. width, tapered, compressed, proximally un- First pereiopod slender, exceeding car- armed; basis and coxa robust, without spe- pocerite by chela and distal third of propod; cial features. BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 625

Ambulatorypereiopods normal; third pe- endopod 0.9 of exopod length, 2.1 times reiopodexceeding carpocerite by propodand longer than broad. with dactyl;dactyl unguisshort, stout, blunt, Female. 1.5 times than basal 0.28 of Description of -Generally very longer width, similar to male, with much stouter carpus carpuscompressed, 2.0 times body, length, more expanded pleura, smaller than proximal width, taper- relatively longer slightly second pereiopod chelae. ing distally,ventral border blunt, with single Rostrum with single minute distoventral small subacute distal accessary tooth, with tooth, lateral margins feebly constricted few short simple setae distally,medially, and proximally; distolateral angle of proximal laterally; propod 5.5 times dactyl length; segment of antennularpeduncle with more 0.38 of carapace length, 6.6 times longer stronglydeveloped acute tooth; mouthparts than proximal depth, slightly tapering dis- not removed, deep proximal tally, with pair of distoventral spines, single paragnathal small ventral dis- groove clearlyvisible; major second pereio- very preterminal spine, with toventral pod chela 1.6 times of carapacelength, margin sparsely setose; carpus minor chela 1.12 times of about 0.55 of 3.0 times lon- carapacelength, propod length, 0.68 of chela dentition as in ger than distal width, tapered prox- major length, slightly male; simi- imally, with small distal lobe, unarmed;me- ambulatory pereiopod dactyls lar, accessary tooth more acute, recurved; rus about 1.25 times propod length, 5.0 dactyl and propod more setose; telson times longer than central width, slightly broader,posterior marginal spines small, ir- swollenproximally, unarmed; ischium about regular,only slightly differentin size. 0.7 of propod length, 3.5 times longer than distal width; basis and coxa without special Measurements (mm).-Male, total body features. Fourth and fifth pereiopods simi- length (approximate) 31.5; carapace and lar to third. rostrum, 14.5; postorbital carapace 11.0; First pleopod with basipodite about 2.3 second pereiopod, major chela, 23.0; minor times longer than wide; exopod 1.25 times chela, 18.5. Female, total body length (ap- basipodite length; endopod 0.66 of basipo- proximate) 39.0; carapace and rostrum, dite length, 4.5 times longer than proximal 16.5; postorbitalcarapace, 12.5; second pe- width, slightly tapering distally, apically reiopod, major chela, 20.0; minor chela, rounded, without median lobule, lateral 14.0. margin with 12 short plumose setae over materialofPalaemon 0.6 of distal Types-The type pin- proximal length, margin gla- nophylax Otto, 1821, is apparentlyno lon- brous, medial margin with about 40 short extant commu- setae over whole ger (H.E. Gruner, personal simple spiniform length, nication). setae slightly longer distally. Second pleopod with basipodite about 2.5 times longer Remarks.-The most remarkablefeature of than wide; endopod about 1.1 times basipo- the present redescription of Pontonia pin- dite length;endopod 0.95 of exopod length, nophylax is the presence of the deep lon- appendicesat 0.33 of medial marginlength, gitudinal medial groove along the proximal appendixmasculina with shorttapering cor- half of the ventral aspect of the elongated pus, about 3.0 times longerthan basal width, paragnath,a featurethat can be quite easily seen without the removal of the mouth- with about 18 long simple spines ventrally, about 1.4 times parts. Its functional significanceis not im- apical spine corpus length, obvious. The setose ventral mar- proximal spines shorter; appendix interna mediately gins of the lips of the groove are contiguous slender, 2.0 times length of appendix mas- with the short setae fringing the endite of culina corpus, far exceeding distal corpus, the first when these with few distal cincinnuli. maxilliped, particularly are adducted. It is also noteworthythat the Uropod with protopodite stout, short, broad ischiomerus-basal segments of the distolateral angle bluntly rounded; exopod third maxillipeds are widely separatedand distinctlyexceeding telson, broad, 1.75 times nonoperculate. However, the first pereio- longer than wide, lateral margin convex, pods appear to be carried,when not in use nonsetose, without distolateral tooth, with for feeding, with the carpomeraljoint fully minute mobile spinule, diaeresis distinct; flexed, so that the chelae lie against the is- 626 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991 chia. The pair of first pereiopods dovetail could survive. It may be noted that P. pin- together to form an effective closure of the nophylaxhas also been reportedfrom Tur- spacebetween the proximalparts of the third key, among stones at 0-1-m depth, where maxillipeds, so that a quadripartiteoper- no Pinna were collected (Holthuis, 1961). cular appearanceis presented ventrally. Bruce (1978) has reported an apparently The present specimens of P. pinnophylax free-living specimen of Pontonia sibogae possess ambulatory dactyls that are mark- from Madagascar,as has Holthuis (1986) edly differentfrom the illustrationprovided for specimens for P. sibogae and P. styli- by Lagardere(1971) in which it appearsdis- rostris from Oman, found under boulders. tinctly triunguiculate,a featurethat may be However, it may be that removal of boul- possibly due to individual abnormality or ders causes sufficientdisturbance to an as- injury and repair. cidian host to cause it to suddenly contract A furtherspecimen of P. pinnophylaxhas and expel its associatedshrimps. This could also been examined. A nonovigerous female readily occur if the shrimps happened to be was collected from Ulnis an der Schlei, Ger- in the vicinity of one of the siphons when many, in July-August, 1976. The specimen the host was caused to contract. has a carapacelength of 6.5 mm, and cor- As noted by Holthuis (1952) the upper respondsclosely to the above redescription. antennular flagellum in P. pinnophylax is A few small points of differencemay be not- curved backwards,lying alongside the ven- ed. The dorsalrostrum bears a minute distal tral rostral carina, in contrast to Notopon- tooth. This tooth was obsolete in the Med- tonia platycheles,in which it is directed an- iterraneanspecimens. The chelae of the sec- teriorly. ond pair of pereiopodsare unequaland dis- similar, with the minor chela about 0.6 of the major chela length, subequalto the car- DISCUSSION apace length, with the dactyl about 0.85 of The discovery of Notopontonia platy- the palm length, with the unarmed cutting chelesraises the small numberofpontoniine edgeoccupying two-thirds of the cuttingedge shrimpsknown from SouthAustralia to five. distally and a small acute tooth at about 0.3 The only other species so far reportedfrom of the length, with similar changes on the South Australia are: Periclimenes aesopius fixed finger.The ambulatorydactyls have a (Bate),associated with anemones;Pontonia relatively longer, more slender, and acute minuta Baker, host unknown; Anchistus unguis, and the ventral corpus is more custos (ForskAl), associated with bivalve stronglysetose, with the setae largelyin bi- molluscs; and Periclimenes carinidactylus lateral pairs. The ambulatory propods are Bruce, associated with crinoids. provided with a well-developed pair of dis- It is unfortunate that the host of Noto- toventralspines only. The telson is generally pontonia platycheles is unknown. The similar, but the posterior marginal spines stronglydepressed body form and the hor- are better developed. The exopods of the izontal disposition of the second pereiopod uropods appearto lack discernibledistolat- chelae suggest a bivalve mollusc, such as eral spinules. one of the Pectinidae. How such a shrimp The most remarkablefeature, however, came to be caught in a trap set for crayfish, is the presence of this species in the Baltic Panulirus spp., is particularlyobscure. As Sea. Holthuis (1952) gave the distribution faras is known, bivalve shrimpcommensals of this species as the MediterraneanSea, such as Anchistus, Paranchistus,and Con- from the east coast of Spain to Greece, the chodytes spp. (Hipeau-Jacquotte, 1973) Azores, Gabon, and north Angola. Chace never leave their hosts to forage indepen- (1966) recordedthis species from Saint Hel- dently and it seems unlikely that N. platy- ena, South Atlantic. The present specimen cheles would do so. was collected alive, by U. Pettke, from pil- The single specimen of Notopontonia ings (M. Turkiy, personalcommunication). platycheles falls outside the normally ac- Since the normal host for P. pinnophylax cepted definition of the Pontoniinae, in does not occurin the Baltic Sea, it is difficult which the posterior margin of the telson is to imagine how this commensal shrimp armed with three pairs of spines. It must be BRUCE: NEW PONTONIINE COMMENSAL SHRIMP FROM AUSTRALIA 627 considered possible that this unique speci- basal portion is also distinctly relatively men is abnormal, a situation that will only largerand more clearly separatedfrom the be resolved when furtherspecimens can be alar region. Palaemonella rotumana (Bor- examined. The presenceof two pairs of pos- radaile) has an essentially similar parag- terior spines on the telson is also known in nath, which may be consideredto represent the case of Chernocarisplacunae, a rather the primitive pontoniine condition for this similar associate of bivalve molluscs, Pla- feature. The alae have the lobes less well cuna placenta, also of extremely depressed marked than in Macrobrachiumand there body form (Johnson, 1967). This strength- is no distinct division between alar and bas- ens the suggestion that N. platycheles may al parts. The basal part also has the central also be a bivalve associate. Indeed, Cher- eminence, with smallerlateral swellings and nocaris appears to bear a parallel relation- a pair of well-separatedfeeble ridgesare dis- ship to Conchodytesas does Notopontonia cernible laterally on the median eminence. to Pontonia. In C. placunae, the examina- The paragnathin Pontonia katoi Kubo is tion of numerous specimens confirms that also basically similar to that in Palaemo- the presence of only two pairs of posterior nella, but the alar lobes are more distinct, telson spines is the normal complement for the basal portion lacks the smaller lateral this species. swellings and the median eminence has a The morphology of the paragnathsin pa- pair of feeble lateral carinae. In Pontonia laemonid shrimps has attractedlittle atten- pinnophylax,the alar lobes are distinct, the tion. Borradaile(1917) illustrated the par- basal portions markedly elongate, with a agnath of Palaemon serratusin relation to distinct median channel bordered by long the labrum,mandible, and maxillula,which raisedsetose carinae,quite distinctfrom that largelyobscure it, but did not discuss it fur- of Notopontonia. The extent to which the ther. Patwardhan(1937), in dealingwith the morphology of the paragnath,as found in mouthpartsof the Indian river prawn,Mac- P. pinnophylax, occurs in the genus Pon- robrachiummalcolmsonii, omitted the par- tonia is presentlyunknown, but it is clearly agnath from consideration. More recently, not a characteristicof the genus as presently Alexander (1988) described the paragnath defined. However, the morphological fea- in Palaemon serratus(Pennant) and Cran- tures of Pontonia show such a rangeof pres- gon crangon (L.). In species descriptions, ent-or-absent variations as to suggest a mention of the paragnathsis usually omit- polyphyletic origin of the species now in- ted, even when the mouthparts are de- cluded under this name. In particular,three scribed, possibly because they are easily distinct types of ambulatorydactyl suggest damaged during the removal of the paired at least three separate species groupings. appendages. These are: (1) simply biunguiculate, as in The examination of the paragnath in all the American species, most of which are Macrobrachiumconfirms the accuracy of associated with molluscs, particularly bi- Borradaile'sfigure, and it may be noted that valves; (2) biunguiculate, with additional the distal spines of the upper lacinia of the acute proximal and intermediate teeth, as maxillulaare in the closest apposition to the in P. katoi and P. sibogae, associates of as- distomedian fissurebetween the alae of the cidians; and (3) biunguiculate, with series paragnath.The lower laciniae lie adjacent of blunt hooklike teeth distally, as in P. as- to the median eminence of the basal por- cidicolaBorradaile, P. anachoretaKemp, P. tions of the paragnathand are directed an- okai Kemp, and P. monnioti Bruce, all as- teriorly towards the median fissure. The sociates of ascidians. paragnathin Macrobrachiumshows a ba- similar structureto that reportedfor sically ACKNOWLEDGEMENTS Palaemon serratusby Alexander(1988), but the division of the alae into posterior, me- I am most gratefulto Dr. WolfgangZeidler for the dian, and anterior lobes is much less dis- opportunityto studythis interestingspecimen, and also with to Dr. N. Calafioreand Dr. G. Costanzofor providing tinct. The basal portion also is larger, the specimensof Pontoniapinnophylax described, and a distinct rounded median eminence, with to Dr. M. Turkiay,for the loan of the intriguingBaltic smaller lateral swellings proximally. The specimen of P. pinnophylax. 628 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991

LITERATURECITED Johnson, D. S. 1967. On some commensal decapod crustaceansfrom Singapore(Palaemonidae and Por- Alexander,C. G. 1988. The paragnathsof some in- cellanidae).-Journal of Zoology 153: 499-526. tertidalcrustaceans. -Journal of the MarineBiolog- Lagardere,J.-P. 1971. Les crevettesdes c6tes du Ma- ical Association of the United Kingdom 68: 581- roc.-Travaux de l'InstitutScientifique Cherifien et 590. de la Facult6de Science, serie Zoologie 36: 11-139. Borradaile,L. A. 1917. On the structureand func- Latreille,P. A. 1829. Crustac6s,Arachnides et partie tions of the mouth parts of palaemonid prawns.- des Insectes.-In: G. Cuvier, Le regne animal dis- of the Proceedings Zoological Society of London tribu6d'apres son organisation,pour servir de base 1917: 31-71. a l'histoirenaturelle des animaux et d'introduction A Bruce,A. J. 1978. report on a collection of pon- a l'anatomiecomparee, edition 2,4: 1-584. toniine from shrimps Madagascar and adjacent Opinion 378. 1956. Designations,under the Plenary seas. -Zoological Journalof the LinneanSociety 62: Powers, of a type species in harmony with current 205-209. usage for the genus Pontonia Latreille, 1829 (Class F. A. 1966. Chace, Decapod crustaceansfrom Saint Crustacea,Order Decapoda) a genus based upon a Helena Island, South Atlantic.-Proceedings of the misidentified type species.-Opinion of the Inter- United States National Museum 118: 623-662. nationalCommission for ZoologicalNomenclature, Hipeau-Jacquotte,R. 1973. Etudedes crevettesPon- 11:411-420. toniinae(Palaemonidae) associ6es aux molluscsPin- Otto, A. W. 1821. Conspectusanimalium quorun- nidae a Tulear (Madagascar).3. Morphologie des dam maritimarum nondum editorum (academic pieces buccales.-Tethys, suppl. 5: 95-116. thesis, Bratislava). Holthuis, L. B. 1947. Nomenclatorialnotes on Eu- 1828. Beschreibungeinige neuen,in den Jah- ropean macrurous Crustacea Decapoda.--Zoolo- ren 1818- und 1819 im MitteliindischenMeere ge- gische Mededelingen27: 312-322. fundeneCrustacea. -Nova Acta AcademiaeCaesa- 1952. The Decapoda of the Siboga Expedi- reae Leopoldino Carolinae germanicae naturae tion. Part XI. The Palemonidaecollected by the Si- curiosorum,14: 331-354. boga and Snellius Expeditionswith remarkson the Patwardhan,S. S. 1937. Palaemon.-In: K. N. Bahl, other species II. Subfamily Pontoniinae.-Siboga The Indian ZoologicalMemoirs 6: i-xi, 1-100. ExpeditionMonograph 393: 1-252. 1961. Report on a collection of Crustacea RECEIVED:1 October 1990. Decapoda and Stomatopodafrom Turkey and the ACCEPTED:30 April 1991. Balkans.-Zoologische Verhandelingen47: 1-67. 1986. Some Pontoniinae(Crustacea: Decap- Address: Division of Natural Sciences, Northern oda: Palaemonidae)from southernOman.--Zoolo- TerritoryMuseum, P.O. Box 4646, Darwin,Australia gische Mededelingen60: 263-272. 0801.