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11-1-1991 Revision of the Cumacean Family Leuconidae Les Watling University of Maine - Main, [email protected]
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Repository Citation Watling, Les, "Revision of the Cumacean Family Leuconidae" (1991). Marine Sciences Faculty Scholarship. 108. https://digitalcommons.library.umaine.edu/sms_facpub/108
This Article is brought to you for free and open access by DigitalCommons@UMaine. It has been accepted for inclusion in Marine Sciences Faculty Scholarship by an authorized administrator of DigitalCommons@UMaine. For more information, please contact [email protected]. JOURNAL OF CRUSTACEAN BIOLOGY, 11(4): 569-582, 1991
REVISION OF THE CUMACEAN FAMILY LEUCONIDAE
Les Watling
ABSTRACT The family Leuconidaecurrently contains 99 species. With the exception of Epileucon,all generain the family were establishedby 1907. All new species have been subsequentlyassigned to those genera,gradually producing genera distinguishable on the basis of single characters.In this paperEpileucon is reducedto a subgenusof Leucon as proposed by Bacescu(1988), and the species of Leucon are furtherapportioned among the subgeneraLeucon Kroyer,Macrau- loleucon,new subgenus,Crymoleucon, new subgenus,and Alytoleucon,new subgenus.In ad- dition, the new genera Ommatoleucon,Austroleucon, Nippoleucon, and Bytholeuconare pro- posed, and the genus CoricumaWatling and Breedy is added to this family. The new genus Americuma,with unclear family affiliations,is proposed for a species previously assigned to Heteroleucon.Complete diagnosesare provided for all generaand keys are given to all genera and species.
The Leuconidaeis one of the oldest of all leucon, suggestingthat the characterspro- cumacean families, having been established posed were not significantand did not define by Sars in 1878 for the genera Leucon and all species in the genus. Instead,he proposed Eudorella.Eudorellopsis was createdby Sars that Leucon should be divided into the sub- in 1882 to accommodatetwo previouslyde- genera Leucon and Epileucon, recognizing scribedspecies. Additional species from the that the species in this genus could be seg- North Atlantic and Mediterraneanregion regatedinto discrete groups. In the follow- graduallyswelled the contents of these gen- ing account, several other subgenera are era, due especially to the efforts of Sars (13 proposed, each of which are hypothesized species).In the early 1900's,as materialfrom to be a phylogenetic lineage within the ge- various expeditionsbecame available,many nus. more new species were described.However, With the exception of Epileucon, there only a few belonged to the new genera de- have been no new genera created for the scribed by Zimmer (Pseudoleucon)in 1903 60 + species describedsince 1907. For most and Calman (Hemileucon, Heteroleucon, of the shallow northernhemisphere species, and Paraleucon)in 1907. These genera ac- this has not been a problem, since they fit commodated all subsequent new species quite comfortably into the genera estab- until Jones (1956) proposedEpileucon. The lished earlier for species from this region. family presently contains 99 species as- For species from Asia and the South Pacific, signed (occasionally with doubt) to those Central America, and the deep sea, use of eight genera. the established genera has been problem- Bishop(198 la) describeda numberof new atic. In several cases the generic diagnoses species which he assigned to Epileucon. He have been so stretchedthat only one or two noted that the original diagnosis of Epileu- characterscontinue to hold a genus togeth- con was based on a character("absence of er. Occasionally, the few members of a ge- a serrated dorsal crest on the carapace of nus do not look even remotely alike. In this the female," Jones, 1956) that did not show paper, several new generaare created in or- stability even in the type species. However, der to deal with these problems and all gen- based on an extensive examination of deep era are rediagnosed. Atlantic leuconids, the genus was redi- a suite of chief agnosed using characters, KEY TO GENERAOF LEUCONIDAE among them being the presenceon pereion- ite 5 of at least one pair of anteriorlycurved 1. Distinct eye lens and/or pigment present...... 2 ventral teeth (Bishop, 1981a). All other - Eyelobewithout lens or pigment 3 characterswere to 2. Eyelobe extending to end of pseudorostrum, acknowledgedby Bishop uropod endopod 2-articulate Coricuma occur also in some or several species of Leu- - Eyelobe not reachingend of pseudorostrum, con. Bacescu (1988) rejectedthe genus Epi- uropod endopod uniarticulate...... Ommatoleucon
569 570 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991
3. Exopodson pereiopods 1 and 2 only in 6 and only slightlybeyond pseudorostrum 2 2 Heteroleucon 2. Pereionite 5 with ventral hook(s) ...... Epileucon - Exopodson pereiopods 1-3 in 9 and 1-4 in 8 - Pereionite 5 without ventral hook(s) 3 4 3. Antenna1 accessoryflagellum extending at least 4. Uropod endopod uniarticulate 5 to midlengthof main flagellumfirst article ...... - Uropod endopod 2-articulate 6 Crymoleucon 5. Pereiopod2 article 3
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Fig. 1. Leucon (Leucon) nasica (from Sars, 1900): a, carapace,side view; b, antenna 1; c, uropod. Leucon (Epileucon)galatheae (from Bishop, 198 la): d, carapace,side view; e, pereionites3-5 and pleonites 1 and 2; f, antenna 1; g, uropod. Leucon (Macrauloleucon)spinulosus (from Hansen, 1920): h, carapaceand antenna 1; i, uropod. Leucon (Crymoleucon)tener (from Hansen, 1920): j, carapace and antenna 1; k, uropod. Leucon (Alytoleucon)medius (from Bishop, 1982): 1, carapace,side view; m, antenna 1; n, uropod. (Drawingsnot to same scale.)
6. Carapacewith dorsolateralspines on frontal - Pseudorostrallobes subquadrate(blunt) dis- lobe robustus tally, carapace with dorsal serrations - Carapacewithout dorsolateralspines on fron- throughoutlength 13 tal lobe serratus 11. Pereiopod 1 basis with large serrations...... 7. Pereiopod 1 article 5 longer than article 6 ...... 8 magnadentata - Pereiopod1 articles5 and 6 subequalin length - Pereiopod 1 basis without large serrations...... 12 12. Pereiopod 1 basis with several plumose setae 8. Uropod endopod broadened laticauda along margin subnasica Uropod endopod normally elongate 9 - Pereiopod 1 with few setae of any kind along 9. Pereiopod 2 article 3 present panamensis margin minor Pereiopod 2 article 3 not present 10 13. Uropod peduncle shorterthan endopod ...... 10. Pseudorostrallobes subtriangularto acutedis- armatus tally, carapacewith dorsal serrationsabsent - Uropod peduncle as long as endopod 14 from posteriorthird 11 14. Carapaceand following pereionites strongly 572 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991
convex dorsally nasicoides - Pseudorostrumrelatively short (< <0.5 times - Carapaceand followingpereionites not arched carapacelength) 3 dorsally kobjakovae 2. Pseudorostrallobes distally subacute 15. Antenna 1 peduncle article 3 at least as long tenuirostris as main flagellum 16 - Pseudorostrallobes distally bluntly rounded - Antenna1 pedunclearticle 3 shorterthan main ensis flagellum 19 3. Sterite of fifth pereionitewith at least 4 spi- 16. Uropod exopod extendingonly to distal mar- niform teeth 4 gin of endopod basal article profundus - Sterniteof fifth pereionitewith 1 or 2 spini- - Uropod endopod extending beyond distal form teeth 5 marginof endopod basal article 17 4. Peduncleof uropod with 3 or 4 setae on me- 17. Carapacewith dorsolateraltooth behindfron- dial margin galatheae tal lobe nathorsti - Peduncleof uropodwith 5-8 setae on medial - Carapacewithout this tooth 18 margin spiniventris 18. Uropod exopod inner marginwith numerous 5. Sterite of fifth pereionite with single mid- (> 10) setae nasicus ventral tooth kalluropus - Uropod exopod inner marginlightly (<9) se- - Sterniteof fifth pereionitewith pairedteeth tose affinis 6 19. Uropod peduncleinner marginheavily setose 6. Uropod exopod longer than endopod basal along length mediterraneus article 7 - Uropod peduncleinner margin sparsely setose - Uropod exopod shorterthan endopod basal 20 article 10 20. Pereiopod2 article4 with strongseta at distal 7. Pereiopod 1 dactylusabout one-halflength of corner assimilis propodus 8 - Pereiopod2 article4 with at most 1 plumose - Pereiopod 1 dactylusabout two-thirdslength seta at distal corner 21 of propodus 9 21. Uropod exopod with dense clusterof long se- 8. Carapaceanteroventral corner with protrud- tae distally homorhynchus ing tooth bengalensis - Uropod exopod sparselysetose distally ...... fulvus - Carapace anteroventral corner indistinct, markedby a sinus latispina 9. Pereiopod2 dactyluswith 3 or 4 plumose se- tae acclivis 1956 - Pereiopod 2 dactylus with more than 6 plu- Epileucon Jones, mose setae socius Fig. Id-g 10. Pereionite 4 pleural plate with ventrally di- Type Species. -Epileucon galatheae Jones, rected tooth pusillus - Pereionite4 pleuralplate ventrallyrounded 1956. 11 Diagnosis (emended). -Branchial siphon 11. Pereiopod 2 article 5 with several (>4) plu- antenna 1 at mose setae craterus normal; accessory flagellum - 2 least half of main first ar- Pereiopod2 article5 with 1 or plumosesetae length flagellum longirostris ticle; pereionite 5 with at least 1 pair of ventral teeth or single midventral tooth; uropod endopod terminal seta not fused to distal article. Macrauloleucon, new subgenus Fig. lh, i Additional Species. -L. (E.) acclivis (Bish- Type -Leucon Hansen op, 198 a); L. (E.) bengalensis Lomakina, Species. spinulosus 1920. 1967; L. (E.) craterus (Bishop, 1981a); L. (E.) ensis (Bishop, 198 la); L. (E.) kalluro- Etymology. -Combination of Greek mak- pus Stebbing, 1912; L. (E.) latispina Jones, ros, long, aulos, pipe tube, and leucon, the 1963; L. (E.) longirostris Sars, 1871; L. (E.) stem genus, referring to the unusually long pusillus (Bishop, 198 a); L. (E.) socius branchial siphon exhibited by these species. L. Han- (Bishop, 1981a); (E.) spiniventris Diagnosis. -Branchial siphon greatly elon- L. tenuirostris 1887. sen, 1920; (E.) Sars, gate; antenna 1 accessory flagellum extend- ing beyond midlength of main flagellum first article; pereionite 5 without ventral teeth; KEY TO FEMALESOF LEUCON uropod endopod terminal seta not fused with (EPILEUCON) distal article. (abbreviatedand emended from Bishop, 198la) Additional Species. -L. (M.) siphonatus 1. Pseudorostrumlong (<<0.5 times carapace Calman, 1905; L. (M.) stenorhynchus Gamo, length) 2 1988. WATLING: REVISION OF LEUCONIDAE 573
KEY TO SPECIESOF LEUCON 6. Carapaceantennal notch defined ventrallyby 2 teeth macrorhinus (MACRAULOLEUCON) large - Carapaceantennal notch indistinct,anteroven- 1. Pseudorostrumand frontallobe withlong spines tral marginwith very small serrations 2 serrulirostris - Pseudorostrumwith low spines, none on fron- 7. Uropod exopod extendingwell beyond endo- tal lobe siphonatus pod dayae 2. Pereionites2 and 3 with dorsal and dorsolat- - Uropod exopod equal to or shorter than en- eral spines spinulosus dopod 8 - Pereionites2 and 3 without spines stenorhynchus 8. Uropod exopod extending only to end of en- dopod basal article 9 Crymoleucon,new subgenus - Uropod exopod at least equal to entire endo- k pod (complexrepresenting sagitta, vanhoeffeni, Fig. lj, and kerguelensis) TypeSpecies. -Leucon tenerHansen, 1920. 9. Antennal notch strong;pereiopod 1 article 7 -From Greek cold, more than three-fourthslength article 6 ...... Etymology. krymos,icy septemdentatus for the waters in which the members of this - Antennal notch obsolescent; pereiopod 1 ar- subgenusare found or appear to have been ticle 7 about one-half length article 6 ...... bishopi derived, and leucon, the stem genus. Diagnosis.-Branchial siphon normal; an- tenna 1 accessoryflagellum extending at least Alytoleucon,new subgenus to midlength of main flagellumfirst article; Fig. 11-n 5 without ventral pereionite teeth; uropod - 1982. endopod terminal seta not fused to distal TypeSpecies. LeuconjonesiBishop, article. Etymology.-From Greek alytos, continu- Additional -L. antarcticus ous, unbroken, referringto the fused con- Species. (C.) dition of the terminal L. bishopi Bacescu, uropod endopod seta, Zimmer, 1907; (C.) and the stem 1988; L. (C.) dayae Ledoyer, 1988; L. (C.) leucon, genus. heterostylis Calman, 1907a; L. (C.) inex- Diagnosis.-Branchial siphon normal; ac- cavatusLedoyer, 1977; L. (C.) kerguelensis cessory flagellumrudimentary; pereionite 5 Zimmer, 1908; L. (C.) macrorhinusFage, without ventral teeth; uropod endopod ter- 1951; L. (C.) sagitta Hansen, 1920; L. (C.) minal seta fused to distal article. septemdentatusZimmer, 1902; L. (C.) ser- AdditionalSpecies. -L. (A.) mediusBishop, rulirostrisLedoyer, 1988; L. (C.) vanhoef- 1982; L. (A.) pallidus Sars, 1865; L. (A.) feni Zimmer, 1907. turgidulusBishop, 1982. KEY TO FEMALESOF LEUCON KEY TO SPECIESOF LEUCON(ALYTOLEUCON) (CRYMOLEUCON) 1. Pereiopod1 compact,article 4 1.5 times length 1. Antenna 1 well of article 3 pallidus accessoryflagellum extending - beyond main flagellumbasal article 2 Pereiopod1 elongate,article 4 > 3 times length - Antenna 1 accessoryflagellum equal in length article 3 2 or shorterthan main flagellumbasal article ...... 4 2. Pereiopod 1 exopod articlesnot reachingdistal 2. shorter than basal end of basis turgidulus Uropod exopod endopod - article heterostylis Pereiopod 1 exopod articlesextending beyond - Uropod exopod equal to or longer than en- distal end of basis 3 dopod basal article 3 3. Pereiopod 1 article 7 slightly shorterthan ar- 3. Antennal notch obsolete; carapacemiddorsal ticle 6; pereiopod3 articles4 and 5 subequal__. teeth not extendingto posteriormargin ...... medius inexcavatus - Pereiopod 1 articles6 and 7 subequal;pereio- - Antennalnotch strong; carapace middorsal teeth pod 3 article 5 > 4 jonesi continuousfrom eyelobe to posteriormargin Remarks.-The was antarcticus genus Epileucon orig- 4. Pseudorostrallobes greatlyelongated 5 inally createdby Jones (1956) for the species - Pseudorostrallobes of normal length 7 E. galatheae, which had the characters of 5. Pereiopod 1 elongate, article 6 at least twice the genus Leucon, but differed from all length of article 7; pseudorostrumwith fringe known species by the absence of a serrated of long setae 6 dorsalcrest. noted that some - Pereiopod1 stout,article 6 slightlyshorter than Bishop (198 la) article7; pseudorostrumwithout long fringeof specimens of E. galatheae in fact possessed setae tener several small dorsal teeth or serrations on 574 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991 the carapace.He stated "the supposed dis- female;uropod endopod uniarticulate;male tinction between Epileucon and Leucon is with 2 pairsofpleopods; pigmentedeye with thereforeeroded" (p. 354). Bishop then re- single lens present in both sexes. defined the genus Epileucon, using as the AdditionalSpecies.--No others. primary character the presence of ventral teeth on pereionite 5. In addition, many Remarks.-The presenceof a pigmentedeye other characterswere listed, the combina- with a lens distinguishesthis genus from all tion of which was considered to be unique other members of the family. Ommatoleu- was in the to species of Epileucon. However, any one con ocularis originallyplaced ge- or more, but not all, of these characterscould nus Leucon principally on the basis of its be found in some species of Leucon. Baces- generalbody shape and the presenceof two of in the male. Its and cu (1988) rejectedthe genus Epileucon, sug- pairs pleopods eye the also made gestingthat the additionalfeatures provided uniarticulateuropod endopod it the members of Leucon. by Bishop were not significantat the generic unique among level and were not to be seen on all the Hemileucon Calman, 1907 species presumed to constitute the genus. Fig. 2f-i Instead, he divided the species of Leucon -Hemileucon into two subgenera,Leucon and Epileucon, Type Species. uniplicatus 1907a. dependingon whetheror not they possessed Calman, ventralteeth on pereionite5. Bishop (1981a) Diagnosis.-Pseudorostrum projecting an- reexamined several species and the details teriorly, shorter in male; antenna 1 genic- mentioned above regardingthe presence or ulatebetween peduncle articles 1 and 2; male absence of ventral teeth are taken from his antenna 2 with brush of setae on anterior paper. On this basis, L. bishopi is moved margin of peduncle articles 4 and 5; male out of the subgenusEpileucon. In this paper, antenna 2 flagellum not extending beyond other subgeneraare proposed which reflect pereion;female with exopods on pereiopods the common possession of selected features 1-3; male with exopods on pereiopods 1- by groupsof species of Leucon. In each case, 4; pereiopod 2 article 3 distinct;uropod en- it is not felt that these features are of suffi- dopod 2-articulate;male without pleopods. cient significanceto warrantthe erection of Additional Species. -H. comes Calman, new genera, but ratherare hypothesized as 1907a. representing evolutionary patterns within the genus. However, it is possible that the KEY TO THE SPECIES OF featuresused to diagnoseeach subgenuswill HEMILEUCON be found to be homoplasious upon the dis- 1. Carapacewith lateralridge extendingforward covery of additional species. from near posterior margin to beginning of pseudorostrallobes; pseudorostrallobes not Ommatoleucon,new genus upturned uniplicatus Fig. 2a-e - Carapacewith oblique lateralridge extending dorsoventrally;pseudorostrum upturned in 9 comes TypeSpecies. -Leucon ocularisHale, 1945. Austroleucon,new Greek genus Etymology.--From ommatos, eye, Fig. 2j-m referringto the presence of a lens on the eyelobe, and leucon, the stem genus. TypeSpecies. -Hemileucon levisHale, 1945. 1945. Diagnosis.-Pseudorostrum projecting an- teriorly,slightly shortenedin male; antenna Etymology.-Austr-, from Australia, and 1 weakly geniculate between peduncle ar- leucon, the stem genus. ticles 1 and 2; male antenna 2 with brush Diagnosis.-Pseudorostrum projecting an- of setae on anteriormargin of peduncle ar- teriorly, equal in male and female; antenna ticles 4 and 5; male antenna 2 flagellum 1 slightly geniculate between peduncle ar- extendingwell along pleon; female with ex- ticles 1 and 2; male antenna 2 with brush opods on pereiopods 1-3; male with exo- of setae on anteriormargin of peduncle ar- pods on pereiopods 1-4; pereiopod 2 article ticle 5 and posterior margins of peduncle 3 much shorter than wide in male, lost in articles 4 and 5; male antenna 2 flagellum WATLING: REVISION OF LEUCONIDAE 575
f
Fig. 2. Ommatoleucon ocularis (from Hale, 1945): a, carapace, side view; b, antenna 1; c, pereiopod 2; d, uropod; e, 6 antenna 2 peduncle and proximal flagellar aricles. Hemileucon comes (from Calman, 1907a); f, 6 carapace, side view, and antenna 1 and 2; g, antenna 1; h, pereiopod 2; i, uropod. Austroleucon levis (from Hale, 1945): j, antenna 1; k, pereiopod 2; 1, uropod; m, 6 antenna 2. Nippoleucon hinumensis (from Gamo, 1967): n, antenna 1; o, pereiopod 2; p, uropod; q, 6 antenna 2 with detail of basal flagellar structure. (Drawings not to same scale.) not extending beyond pereion; female with reception of sensory information while the exopods on pereiopods 1-3; male with ex- male is swimming through the water. In opods on pereiopods 1-4; pereiopod 2 ar- Austroleucon the setal brush on the anterior ticle 3 much shorter than wide; uropod en- margin of peduncle articles 4 and 5 is aug- dopod uniarticulate; male without pleopods. mented by a strong brush on the posterior margin of peduncle article 5. A change in Additional -No others. Species. the function of antenna 2 during the mate Remarks. -Austroleucon levis was originally pursuit process seems likely. Additionally, placed in the genus Hemileucon on the basis Austroleucon differs from Hemileucon by its of its lack of pleopods and shortened an- possession of a uniarticulate uropod endo- tenna 2 flagellum. The form of antenna 2, pod. however, differs from that seen in the spe- new cies of Hemileucon. The latter have a typical Nippoleucon, genus male antenna 2 with a strong brush of setae Fig. 2n-q on the anterior margin of peduncle articles Type Species. -Hemileucon enoshimensis 4 and 5. Such a setal orientation assures the Gamo. 1967. 576 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991
Etymology.-Nippon-, fromJapan, and leu- projecting anteriorly over pseudorostral con, the stem genus. lobes. -No others. Diagnosis.-Pseudorostrum projecting an- AdditionalSpecies. teriorly, shorter in male; antenna 1 not ge- Remarks.- Since this species seemed to niculate; male antenna 2 without brush of have the charactersof both the Bodotriinae setae on pedunclearticles 4 and 5, peduncle and the Leuconidae, it was tentatively as- article 5 subdivided by several annulations, signed to the formerby Watlingand Breedy each division with graspingtooth and setal (1988). However, many of the features it bundle on posterior margin; male antenna exhibits are the result of reductions and I 2 flagellumnot extending beyond carapace have felt it necessaryto reevaluateits place- posterior margin; female with exopods on ment. In particular,the number of exopods pereiopods 1-3; male with exopods on pe- is reduced,the mandible has no molar, and reiopods 1-4; pereiopod 2 article 3 very there are only 6 articles in pereiopod 3. On short; uropod endopod 2-articulate; male the basis of the presence of a strongly de- without pleopods. veloped eyelobe and the reducednumber of the was in the Bo- Additional -N. hinumensis exopods, genus placed Species. (Gamo, dotriinae. On the other hand, the number 1967). of pleopods, the lack of an internal process Remarks.-The two species assignedto this on the pleopod inner ramus, and truncate genus were originally placed in the genus mandible (not elongate as originally de- Hemileucon. Continuing the modification scribed), could have justified its placement of the male antenna 2 seen in Austroleucon, in the Leuconidae. While the strongly de- in Nippoleuconthe pedunculararticles have veloped eyelobe remains a problem, and become modified into grasping-likestruc- certainly necessitates a reevaluation of the tures.The brushof setae is completely miss- charactersdefining the family, the high de- ing from peduncular articles 4 and 5; pe- gree of similarity of the male antenna 2 duncle article 5 is ringed with several graspingstructure to that seen in Nippoleu- annulations,each subdivision bearinga re- con suggestsstrong affinities to the Leucon- curved flexible tooth and a small cluster of idae. A reduced male antenna 2 is seen in setae; the flagellumis still present but con- other families, for example, in the Lampro- sists of less than 12 articles. pidae (e.g., Lamprops[see Sars, 1900]), and Bodotriidae Mancocuminae (Spilocuma KEY TO SPECIESOF NIPPOLEUCON Watling, 1977). In both Lampropsand Spi- locuma the antennal modifications are on 1. Uropod exopod innermargin with plumose se- the anterior whereas in tae; carapaceand appendagesgenerally devoid margin, Nippoleu- of scales enoshimensis con and Coricuma the grasping structures - Uropod exopod inner marginwith long simple are modifications of the posterior margin, setae;carapace and appendagesgenerally cov- suggestingthat this function for the anten- ered with scales hinumensis nae has arisen independently. CoricumaWatling and Breedy, 1988 HeteroleuconCalman, 1907a Type Species.-Coricuma nicoyensis Wat- Fig. 3a-d ling and Breedy, 1988. -Heteroleucon akaroensis Diagnosis (emended). -Pseudorostrum Type Species. projecting anteriorly, slightly shorter in Calman, 1907a. male; antenna 1 not geniculate;male anten- Diagnosis.-Pseudorostrum projecting an- na 2 peduncular articles 4 and 5 without terodorsally, shorter in male; antenna 1 brush of setae, article 5 annulatedand pos- weaklygeniculate between pedunclearticles sessing graspingteeth; male antenna 2 fla- 1 and 2; male antenna 2 with brush of setae gellum absent;female with exopods only on on anterior margin of peduncle articles 4 pereiopod 1; male with exopods only on and 5; male antenna 2 flagellum not ex- pereiopod 1; pereiopod 2 article 3 lost; uro- tending beyond posterior margin of cara- pod endopod 2-articulate;male with 2 pairs pace; female with exopods on pereiopods 1 of pleopods; carapacewith distinct eyelobe and 2; male with exopods on pereiopods 1 WATLING: REVISION OF LEUCONIDAE 577
! 4 F1_h I 7
7
e
Fig. 3. Heteroleucon akaroensis (from Calman, 1907a): a, 9 carapace, side view; b, antenna 1; c, pereiopod 2; d, d carapace, side view, and antennae 1 and 2. Americuma heardi (from Bacescu, 1979): e, antenna 1; f, pereiopod 2; g, uropod. Bytholeucon hiscens (from Bishop, 1981b): h, antenna 1; i, pereiopod 2; j, uropod. Paraleucon suteri (from Calman, 1907a): k, carapace, side view; 1, antenna 1; m, pereiopod 2; n, uropod; o, $ antenna 2. Pseudoleuconjaponicus (from Gamo, 1964): p, carapace, side view; q, antenna 1; r, pereiopod 2; s, uropod. (Drawings not to same scale.) and 2; pereiopod 2 article 3 short; uropod on article 4 and 6 rows on article 5. Each endopod uniarticulate; male without pleo- row extended only slightly from the anterior pods. onto the lateral margin of the peduncle ar- ticle and did not to be on Additional Species. -No others. appear present the medial margin. Remarks. -A male from the Natural History Museum, London (BMNH.1907. Bytholeucon, new genus viii.27.79), with fully developed antenna 2 Fig. 3h-j was examined. The setal brush was seen to Type Species. -Paraleucon (?) hiscens Bish- consist of setae arranged in five distinct rows op, 1981b. 578 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991
Etymology.-From Greekbythos, the depths endopod uniarticulate;male with 1 pair of of the sea, and leucon, the stem genus. pleopods. Diagnosis.-Pseudorostrum projecting an- AdditionalSpecies. -No others. shorterin antenna 1 teriorly, slightly male; Remarks. -Specimens on slides 1907. geniculate between peduncle articles 1 and viii.29.135 and 1907.viii.29.136 in the col- 2; antenna 2 of mature male unknown but lection of the Natural in immature male brush of History Museum, seta seen de- London, wereexamined. In neithercase was on anterior of veloping margin pedunclear- it possible to see the crease that Cal- ticles 4 and antenna 2 slight 5; flagellumelongate man suggested was present on the in immature female with on uropod male; exopods endopod. Consequently, the uropod endo- pereiopods 1-3; male with exopods on pe- pod must be consideredto be uniarticulate. reiopods 1-4; pereiopod 2 article 3 very short; uropod endopod distinctly 2-articu- PseudoleuconZimmer, 1903 late; subadult male with (?)0 or 1 pair of Fig. 3p-s pleopods. Type Species.-Pseudoleucon sorex Zim- Additional Species. -B. ultraabyssalis mer, 1903. (Gam6, 1987). Diagnosis.-Pseudorostrum extending an- Remarks.-Bytholeucon hiscens was tenta- terodorsally,(male ?); antenna 1 geniculate tively assigned to Paraleucon by Bishop between articles 1 and 2; (? male antenna 2 (1981b) on the basis of the late appearance brush of setae on anteriormargin of pedun- of a single pair of pleopods in the one male cle articles 4 and 5); (? male antenna 2 fla- known that is mature enough to show de- gellum elongate); female with exopods on finitive characters. Besides the very great pereiopods 1-3; male with exopods on per- geographicand depth differencesin the dis- eiopods 1-4; pereiopod 2 article 3 narrow; tribution of Paraleuconsuteri (shallow wa- uropod endopod 2-articulate;male with 2 ters of New Zealand)and the species of By- pairs of pleopods. tholeucon in (deep sea the Atlantic and Additional Species.--P. japonicus Gamo, Pacific),the latter are distinguishedalso by 1964. the antenna2 in the male and long flagellum Remarks. short pereiopod 2 article 3. When a mature -While both sexes are known for P. the male is described male is found, the relationshipof Bytholeu- sorex, incompletely con to Paraleuconcan be better assessed. and needs to be reexamined. KEY TO SPECIESOF BYTHOLEUCON KEY TO THE SPECIESOF PSEUDOLEUCON 1. Uropod peduncle shorterthan basal article of endopod; basal article of pereiopod 1 exopod 1. Carapacewith forwardlydirected oblique ridge with 2 separatedspines hiscens extendingfrom dorsal margin to posteriorlimit - Uropod peduncle longer than basal article of of pseudorostrallobe japonicus endopod;basal article of pereiopod 1 exopod - Carapacewithout oblique ridge sorex with 4 closely set spines ultraabyssalis Eudorella Norman, 1867 Type Species. -Eudora truncatula Bate, Paraleucon Calman, 1907a 1856. Fig. 3k-o -Pseudorostrum directed dor- -P. suteri 1907a. Diagnosis. Type Species. Calman, sally in both male and female; antenna 1 Diagnosis.-Pseudorostrum extending an- geniculate between peduncle articles 2 and teriorly, shorter in male; antenna 1 weakly 3; male antenna 2 with brush of setae on geniculate between articles 1 and 2; male anterior margin of peduncle articles 4 and antenna 2 with brush of setae on anterior 5; male antenna 2 flagellumextending near- margin of peduncle articles 4 and 5; male ly to end of pleon; female with exopods on antenna 2 flagellum short; female with ex- pereiopods 1-3; male with exopods on pe- opods on pereiopods 1-3; male with exo- reiopods 1-4; pereiopod2 article3 lost; uro- pods on pereiopods 1-4; pereiopod 2 article pod endopod 2-articulate;male with 2 pairs 3 distinct, nearly as long as wide; uropod of pleopods. WATLING: REVISION OF LEUCONIDAE 579
Additional Species.--E. abyssi Sars, 1887; - Uropod endopod terminal seta fused to ter- minal article 14 E. aequiremis Hansen, 1920; E. arctica 2. Uropod rami equal in length aequiremis Hansen, 1920; E. dentata Lomakiha, 1955; - Uropod exopod at least slightly shorterthan E. emarginata (Kroyer, 1846); E. fallax endopod 3 Zimmer, 1909; E. gottliebi Bacescu, 1961; 3. Uropod exopod much shorterthan endopod E. E. basal article 4 gracilior Zimmer, 1907; gracilis Sars, - as E. E. Uropod exopod about long as or longer 1871; groenlandica Zimmer, 1926; than endopod basal article 6 hirsuta (Sars, 1869); E. hispida Sars, 1871; 4. Carapace anteroventral corer with strong E. hurleyi Jones, 1963; E. intermedia Han- downwardpointing tooth above sinus ...... dentata sen, 1920; E. minor Lomakina, 1952; E. - Carapaceanteroventral corer with smooth or microdentatelobe above sinus 5 monodon Calman, 1912; E. nana Sars, 1879; 5. Pereiopod 1 article 6, 5-6 times as long as E. pacifica Hart, 1930; E. parvula Hansen, wide, antennuleboth flagellawith nakedmar- 1920; E. pusilla Sars, 1871; E. rochfordi gins gracilior' Hale, 1945; E. similis Calman, 1907b; E. - Pereiopod 1 article 6, 2-3 times as long as sordida Zimmer, E. wide, antennuleaccessory flagellum with plu- 1907; spitzbergensis mose setae on fallax 1926; E. distally margin Zimmer, splendida Zimmer, 1902; 6. Antennuleaccessory flagellum at least as long E. tridentata Hart, 1930. as main flagellumbasal article 8 - Antennule shorter than Remarks. -There are several or accessory flagellum pairs, main flagellumbasal article 7 groups, of closely related species in this ge- 7. Carapacepseudorostral lobes with many long nus. While Bacescu (1988) considered E. setae; antennule accessory flagellum three- difficilis Blake, 1929, to be a good species, fourthslength of main flagellumbasal article it was hirsuta Watling (1979) suggested probably - Carapacepseudorostral lobes with few short synonymous with E. pusilla. Since the type setae;antennule accessory flagellum less than of E. difficilis has been lost, this issue may one-half length main flagellumbasal article . never be fully resolved. My own collections monodon from the have 8. Uropod exopod clearly longer than endopod type locality, however, pro- basal article 9 duced no that differ from the E. specimens - Uropod exopod only as long as endopodbasal pusilla collected elsewhere on the east coast article 11 of the United States. Other very closely re- 9. Uropod endopod terminal seta much longer lated species pairs include: E. gracilior and than (>2 times) terminalarticle; pereiopod 2 E. sordida South E. dactyl with setae only on distal one-third ...... (from Georgia); groen- hurleyi landica and E. spitzbergensis (Greenland, - Uropod endopodterminal seta equalin length and Kara and Laptev Seas, respectively); E. to terminal article; pereiopod 2 dactyl with gracilis and E. parvula (Spitsbergen and the setae on distal two-thirds 10 Davis E. and 10. Uropod pedunclewith long, slenderterminal Straits, respectively); splendida seta between insertion of and exo- E. similis and re- endopod (Antarctic Subantarctic, pod; exopod with at least 7 setae on inner spectively); and E. pacifica and E. tridentata marginin adult groenlandica2 (boreal Pacific). Several species appear to be - Uropod peduncle without long slender ter- very close to E. truncatula, namely, E. in- minal seta; exopod with only 3 or 4 setae on termedia Strait to of inner marginin adult gracilis3 (Davis Bay Biscay, 11. Antennulepeduncle article 2 at least 1.5 times 2,000-5,000 m), E. gottliebi (eastern Med- length article 3 12 iterranean, 49-238 m), and E. nana (Med- - Antennulepeduncle articles 2 and 3 subequal iterranean, 37-300 m). The distributions of in length 13 the latter three lie well within the 12. Uropod exopod narrow, elongate, without species setae from which E. truncatula submarginalsetae, marginal plumose.... geographic range abyssi has been recorded. Clearly, there is a prob- - Uropod exopod slightly broadened,submar- lem with closely related species in this ge- ginal setae present, marginal setae not plu- nus. Since the variability in none of these mose splendida4 has been the of the 13. Antennule main flagellumbasal article with species studied, validity plumose setae, pereiopod2 article5 subequal closely related species cannot be assessed. to article 4 minor - Antennulemain flagellumbasal articlenaked KEY TO FEMALES OF EUDORELLA on margins, pereiopod 2 article 5 about 1.5 times length article 4 pacifica5 1. Uropodendopod terminal seta articulated with 14. Uropod exopod distinctly shorter than en- distal article 2 dopod basal article 15 580 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991
- Uropod exopod about as long as or longer 3. Carapacewith single, nearly horizontalridge; than endopod basal article 16 terminalseta of uropodendopod 2 times longer 15. Antennulemain flagellumwith plumosesetae than distal article resima on lateral margin; anteroventralcorner of - Carapacewith 2 obliquelycurving ridges; uro- carapacewith single large sinus; pereiopod 2 pod endopodterminal seta as long as or shorter terminalarticle with many more than 12 setae than distal article 4 emarginata 4. Uropod exopod distal articleexpanded, widest - Antennule main flagellumnaked; ventrolat- midwayalong its length;endopod basal article eral corner of carapacewith 2 small sinuses; nearlydevoid of setae derzhavini pereiopod2 terminalarticle with less than 12 - Uropod exopod distal article of normal con- setae pusilla struction,gradually tapering distally; endopod 16. Uropod exopod extendingto end of endopod basal articlewith lateralsetae 5 distal article arctica 5. Carapacewith anterolateralhorns; pleotelson - Uropod exopod only slightly longerthan en- terminallytriangular 6 dopod basal article 17 - Carapacewithout anterolateralhorns; pleotel- 17. Body more or less beset with fine setae;pseu- sonic somite broadlyrounded posteriorly ...... dorostrallobes with numerouslong setae;pe- biplicata reiopod 2 article 5 about 1.5 times as long as 6. Uropod endopodwith few setae on inner mar- article 4 hispida gin, distal articlewith no marginalsetae; pseu- - Body more or less free of fine setae; pseudo- dorostrallobes with especiallylong setae ...... rostrallobes withoutnumerous long setae;pe- ushakovi reiopod 2 articles4 and 5 subequal.... truncatula6 - Uropod endopodwith setae alongentire length of inner margin, including on distal article; i Notes: Includes E. sordida. 2 Includes E. spitzbergen- pseudorostrallobes with few short setae ...... sis.3 IncludesE. parvula.4 IncludesE. similis.5 Includes longirostris E. tridentata.6 E. intermedia,E. gottliebi,and E. nana E. may all be variantsof truncatula.The key also does INCERTAESEDIS not consider E. rochfordias the details of its uropod endopod are not known. Americuma, new genus Fig. 3e-g EudorellopsisSars, 1882 Type Species. -Heteroleucon heardi Baces- Type Species.--Leucon deformis Kr0yer, 1979. 1846. cu, from and -Pseudorostrum directed dor- Etymology.--Amer-, America, Diagnosis. cuma, the stem group. sally in both male and female; antenna 1 geniculate between peduncle articles 1 and Diagnosis. -Pseudorostrum projecting an- 2; male antenna 2 with brush of setae on terodorsally, branchial siphon long; antenna anterior margin of peduncle articles 4 and 1 not geniculate; female with exopods on 5; male antenna 2 flagellum extending well pereiopods 1 and 2; pereiopod 2 article 3 along pleon; female with exopods on pereio- distinct; uropod endopod uniarticulate; male pods 1-3; male with exopods on pereiopods unknown. 1-4; pereiopod 2 article 3 lost; uropod en- Additional Species. -No others. 2-articulate; male with 2 pairs ofple- dopod Remarks. -This species, described from a opods. manca, was originally assigned to the genus Additional Species. -E. biplicata Calman, Heteroleucon. However, it differs from the 1912; E. derzhavini Lomakina, 1952; E. in- only other species of Heteroleucon, H. aka- tegra (Smith, 1879); E. longirostris Given, roensis from New Zealand, in several im- 1961; E. resima Calman, 1907b; E. uschako- portant respects: its antenna 1 peduncle ar- vi Lomakina, 1955. ticle 1 is very elongate, being as long as the remaining articles combined; it has a dis- KEY TO FEMALES OF EUDORELLOPSIS tinct, elongate pereiopod 2 article 3; there is no antennal notch on the carapace; and 1. Carapacewith lateral horizontal or obliqueridge 3 article 3 of maxilliped 3 is greatly enlarged. - Carapacewithout lateralridge 2 Further, it was obtained half a world away 2. Uropod exopod rectangularwith no setae on and at considerably greater depth, thus re- outer margin; pseudorostrallobes extending quiring the relating of two species in a man- somewhat forward,not strictly vertical;cara- ner which would be almost more than ...... unique among pace elongate,longer high integra the Cumacea. There is no evidence - Uropod exopod tapering,with setae along out- strong er margin;pseudorostral lobes strictlyvertical; that this species is, in fact, a member of the carapaceboxlike, as long as high deformis Leuconidae; it has been dealt with here as WATLING: REVISION OF LEUCONIDAE 581 a means of maintaining order within the . 1967. Studies on the Cumacea (Crustacea, genera of this family. Malacostraca)of Japan.Part 1.-Publications of the Seto MarineBiological Laboratory 15: 133-163. 1987. Two new cumaceancrustaceans, Para- ACKNOWLEDGEMENTS leucon?ultraabyssalis sp. nov. and Leptostylis?spi- nescens sp. nov. from the Japan Trench.-Bulletin This study was conducted during the tenure of a of the of 42: 43-48. SmithsonianSenior Postdoctoral for which BiogeographicalSociety Japan Fellowship, 1988. A new abyssal cumacean crustacean, the support of the Smithsonian Institution is greatly Leucon nov. from I thank Dr. T. E. Bowman for stenorhynchussp. (Leuconidae) appreciated. especially the Japan Trench.-Bulletin of the Biogeographical criticallyreading an earlierversion of this paper and of 43: 25-28. for Society Japan helping me to understandthe broaderconcepts of Given, R. R. 1961. The cumacean fauna of the The illustrationswere S. Bund. taxonomy. preparedby SouthernCalifornia continental shelf. No. 1, Family Leuconidae.-Bulletin of the Southern California LITERATURECITED Academy of Sciences 60: 129-146. Hale, H., 1945. AustralianCumacea, No. 10. The Bacescu, M. 1961. Contributiona l'etude des Cu- Family Leuconidae.-Transactionsof the Royal So- maces de la M6diterraneeet particulierementdes ciety of South Australia69: 86-95. c6tes d'Israel.- Rapportset Proces-verbauxdes Re- Hansen, H. 1920. Crustacea Malacostraca. IV.- unions de la C.I.E.S.M.M.16: 495-502. Danish Ingolf-Expedition3: 1-86. 1979. Heteroleuconheardi n. sp. from the Hart,J. F. L. 1930. Some Cumaceaof the Vancouver MexicanGulf.-Revue Roumainede Biologie (serie Island region.-Contributions to CanadianBiology Biologie animale) 24: 95-97. and Fisheries6: 25-40. 1988. CumaceaI (Fam. Archaeocumatidae, Jones, N. S. 1956. Cumaceafrom the west coast of Lampropidae, Bodotriidae, Leuconidae).-Crusta- Africa.-Atlantide Report 4: 183-212. ceorumCatalogus Part 7, Pp. 1-173. SPB Academic 1963. The marine fauna of New Zealand: Publishing,The Hague, The Netherlands. crustaceansof the Order Cumacea.-New Zealand Bate,C.S. 1856. On the BritishDiastylidae.-Annals OceanographicInstitute Memoir No. 23: 1-81. and Magazineof NaturalHistory (2)17: 449-465. . 1969. The systematics and distribution of Bishop, J. D. D. 198 a. A revised definition of the Cumaceafrom depths exceeding200 meters.- Gal- genus Epileucon Jones (Crustacea,Cumacea), with athea Report.Scientific Results of the Danish Deep- descriptions of species from the deep Atlantic.- Sea ExpeditionRound the World 1950-52, 10: 100- PhilosophicalTransactions of the Royal Society of 180. London (B) 291: 353-409. Kr0yer,H. 1841. Fire nye arteraf SlaegtenCuma.-- 1981b. Two new leuconids (Peracarida,Cu- NaturhistoriskTidsskrift 3: 503-534. macea)of widespreadoccurrence in the deep Atlan- 1846. Carcinologiskebidrag.-Naturhisto- tic.-Crustaceana 40: 144-159. risk Tidsskrift,Ser. 2, 2: 123-211. 1982. Threenew species of the genusLeucon Ledoyer,M. 1977. Cumac6s(Crustacea) des Iles Ker- Kr0yer, 1846 (Crustacea:Cumacea) from the con- guelenrecueillis par le N. O. "LaJaponaise" en 1972 tinental slope off Surinam.-Zoological Journal of et 1974 et parle M. S. "Marion-Dufresne"en 1974.-- the Linnean Society 74: 345-357. Comit6 National Frangaisdes RecherchesAntarc- Blake, C. H. 1929. New Crustaceafrom the Mount tiques 42: 193-213. Desertregion. -Biological Surveyof the MountDes- 1988. Cumac6s (Crustacea)profonds de la ert Region, Part 3, Pp. 1-34. Wistar Institute of region de l'ile de Mayotte, Canal de Mozambique, Anatomy and Biology, Philadelphia,Pennsylvania. Oc6an Indien (CampagneBenth6di, 1977).-Meso- Calman, W. T. 1905. The marine fauna of the west g6e (Bulletin du Museum d'Histoire Naturelle de coast of Ireland,Part IV, Cumacea.-Scientific In- Marseille)48: 131-172. vestigationsof the Fisheries Branch,Ireland, 1904, Liljeborg,W. 1855. Om hafs-crustaceervid Kulla- Appendix No. I: 3-52. berg i Skane.-Ofversigt af Kongliga Vetenskaps- 1907a. On new or rareCrustacea of the Order AkademiensForhandlingar 12: 117-138. Cumaceafrom the collectionof the CopenhagenMu- Lomakina,N. B. 1952. Novye interesnyev zoogeo- seum.-Transactions of the Zoological Society of graficheskomotnoshenii nakhodki kumovykh rakov London 18: 1-56. v dal'neyvostochnykhmoryakh. [New zoogeograph- 1907b. Crustacea. II-Cumacea.-National ically interestingfindings of Cumaceain Far Eastern AntarcticExpedition 1901-1904, NaturalHistory 2: Seas.]-Zoologicheskii Zhurnal 30: 244-248. [In 1-6. Russian.] 1912. The Crustaceaof the OrderCumacea 1955. Kumovye raki (Cumacea) dal'nye- in the collection of the United States National Mu- vostochnykh moryakh.-Trudy Zoologicheskogo seum.-Proceedings of the United States National InstitutaAkademia Nauka S.S.S.R. 18: 112-165. [In Museum 41: 604-674. Russian.] Fage, L. 1951. Cumac6s.-Faune de France 54: 1- 1967. Novye vedyi kumovye(Crustacea, Cu- 136. macea) iz sborov Sovetckoye Antarcticheskoiek- Gam6, S. 1962. Leucon simanensis sp. nov. and L. spedichii(1956-1958) u yogo-vostochnoyeAustralii varians sp. nov.-Zoological Magazine (Dobutsu- i v severnoyechasti Indiiskogookeana [New species gaku Zasshi) 71: 256-261. of Cumacea collected by the Soviet Antarctic ex- 1964. On threenew species of Cumaceafrom pedition at south-easternAustralia and in the north the southern Sea of Japan.-Crustaceana 7: 241- of the Indian Ocean].-Trudy ZoologicheskogoIn- 253. stituta, Leningrad43: 99-108. 582 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 11, NO. 4, 1991
Norman, A. M. 1867. On the Crustacea,Echinoder- Stebbing,T. R. R. 1912. The Sympoda.-Annals of mata, Polypora, Actinozoa and Hydrozoa. Report the South Africa Museum 10: 129-176. of the committee appointed for the purpose of ex- Watling,L. 1977. Two new genera and a new sub- ploringthe coasts of the Hebridesby means of the family of Bodotriidae (Crustacea:Cumacea) from dredge.-Report of the British Association for the easternNorth America.- Proceedingsof the Biolog- Advancementof Science 36: 193-206. ical Society of Washington89: 593-598. 1879. CrustaceaCumacea of the 'Lightning,' 1979. Marine flora and fauna of the north- 'Porcupine,' and 'Valorous' Expeditions.-Annals eastern United States. Crustacea:Cumacea.-Na- and Magazineof Natural History (5)3: 54-73. tional Oceanic and Atmospheric Administration Ohlin, A. 1901. ArcticCrustacea collected during the Technical Report, National Marine Fisheries Cir- Swedish Arctic Expeditions 1898 and 1899 under cular 423: 1-23. the direction of ProfessorA. G. Nathorst. I. Lep- , and 0. Breedy. 1988. A new cumacean(Crus- tostraca, Isopoda, Cumacea.-Bihang till Kongliga tacea)genus from beachesof Golfo de Nicoya, Costa SvenskaVetenskaps-Akademiens Handlingar 26: 1- Rica.-Revista Biologia Tropica 36: 527-533. 54. Zimmer,C. 1902. Cumaceen.- Ergebnisseder Ham- Sars, G. 0. 1865. Om den aberrantekrebsdygruppe burgerMagalhaensischen Sammelreise, 1892/93, 2: Cumacea og den nordiske arter.-Forhandlingar i 1-18. Videnskabs-Selskabeti Christiania1864: 128-208. 1903. Die Cumaceendes Museums fur Na- 1869. Unders6gelserover Christianafyord turkundein Berlin.-Zoologische Jahrbiicher,Ab- dybvandsfauna.-Nyt Magazin for Naturvidenska- theilung fur Systematik, Geographieund Biologie berne 11: 1-58. der Thiere 18: 664-694. 1871. Nya arteraf Cumaceasamlade under 1907. CaliforniaCrustacea of the OrderCu- K. Svenska Korvetten Josephines Expeditioni At- macea.-Proceedings of the United States National lantiska Oceanen ar 1869 af F. A. Smitt och A. Museum 83: 423-439. Ljungman.-Ofversigt afKongligaVetenskaps-Aka- 1908. Die Cumaceender "DeutschenTief- demiens Forhandlingar1: 71-81. see-Expedition".- WissenschaftlicheErgebnisse der . 1878, 1879. Nye Bidragtil Kundskabenom Deutschen Tiefsee-Expedition auf dem Dampfer Middelhavets Invertebratfauna.II. Middelhavets "Valdivia" 1898-1899, 8: 158-196. Cumaceer.-Archiv for Mathematikog Naturviden- 1909. Die Cumaceender SchwedischenSiid- skab 3: 461-512; 4: 1-144. polarexpedition.-WissenschaftlicheErgebnisse der 1882. OversigtafNorges Crustaceermed fo- SchwedischenSudpolar-Expedition 1901-1903, 6: rel0bige bemaerkningerover de nye eller mindre 1-31. bekjendeArter. 1.-Ferhandlingar VidenskabsSel- 1926. Northernand Arctic invertebratesin skabetChristiania 1882: 1-124. the collection of the Swedish State Museum (Riks- 1887. Report on the Cumaceacollected by museum).-Kungliga Svenska Vetenskaps-Akade- H.M.S. Challengerduring the years 1873-1876.- miens Handlingar3: 1-88. Reporton the ScientificResults of the Voyageof the . 1943. CumaceendesStillenOzeans.-Archiv H.M.S. Challengerduring the Years 1873-1876 un- fur Naturgeschichte12: 130-174. der the Command of George S. Nares and Frank Tourle Thomson 19(55): 1-78. RECEIVED:20 MARCH1991. 1900. Cumacea.-An account of the Crus- ACCEPTED:28 May 1991. tacea of Norway 3: 1-115. BergenMuseum, Bergen, Norway. Address:Department of Oceanography,Darling Ma- Smith, S. I. 1879. The stalk-eyedcrustaceans of the rine Center, University of Maine, Walpole, Maine Atlantic coast of North America, north of Cape 04573. Cod.-Transactions of the ConnecticutAcademy of Arts and Science 5: 27-136.