Zoology Series SE THE NATURAL HISTORY MUSEUM VOLUME 66 NUMBER1 29 JUNE 2000 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Zoology Series is edited in the Museum’s Department of Zoology Keeper of Zoology Prof P.S. Rainbow Editor of Bulletin: Dr B.T. Clarke Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. 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(Zool.) 66(1): 148 Issued 29 June 2000 Generic concepts in the Clytemnestridae (Copepoda, Harpacticoida), revision and revival RONY HUYS AND SOPHIE CONROY-DALTON Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS IER ENON Gl ATC EL Yee ares Reerce «ss eee cg satan co neat ntsc Sete au te ceases «sob ss wa bawigete vu Om PNat oc ob Petco D nae wee coeur Bee doe vapuE Ti evadevas sededeen ee LeSER 2 MACE TiAl Stam PWICI OCS acsecesas sie cocadae vaxcsenes ahs apavasasowncceataacdet st cans carseadaure rasdvgnesved senses Tacdsweisassdhpoess ciesasaadedvani visadensdeaeaitbevdcaastodiecacsbce 2 EAN COTTON CH LAS OL gar ryan s co aise reac te Np ace once Ree ae Me das occas aon aac pe eae ee pag ce ao ace ae MPRA a Saas stand see anondugeieh as cena aay -oegncaeeaunes hee 7 COL SICIIIE SUE ANIA, LOA (yacteys sess aay ste sacs acs feat o ta tsaoes Toe fount «eae Dea ea eS ae menae ecna ao eces canto aGas sé dus deste osw En eee cmanetaee 2 GOMIODSVIASIBTAGDY L583 oxcecsctcasncacea tence cen ereswaccatneecascntaaeseseatedsuacdaattawaspanscveite-scesssucenncdacesvakusecs Alasaug@Neseadedesseacdseatetacscvenusarwcss 3 SPF ITT CAT IUD Oe sete ecs haces taranats 600 m (stns 20, 28) TYPE MATERIAL. Neotype 2 dissected on 11 slides (BMNH 1999.996); designated from material labelled Clytemnestra scutellata (BMNH 1948.4.28.121); collected either on 20 October 1928 (stns 19, 20) or 23 November 1928 (stn 28) during the Great Barrier Reef Expedition 1928-29 (Farran, 1936). OTHER MATERIAL EXAMINED. One <4 dissected on 10 slides (BMNH 1948.4.28.121); sampling data as for neotype. REDESCRIPTION. FEMALE. Total body length from tip of rostrum to posterior margin of caudal rami: 1121 um. Maximum width (355 tm) measured at posterior margin of cephalic shield. Posterolateral angles of cephalothorax laterally expanded (Fig. 1A). Somites bearing P2—P4 successively decreasing in width posteriorly and bearing back- wardly produced alate processes. Genital double-somite (Fig. 5A) slightly constricted bilaterally; original segmentation marked by paired transverse chitinous ribs lateroventrally and laterally. Copulatory pore slit-like, located medi- ally between genital apertures; leading to short posteriorly directed, membranous duct connected to bilobate seminal receptacle. Genital apertures located far anteriorly; closed off by small opercula derived from vestigial P6; each with 1 vestigial seta at inner distal corner and anterior tube-pore near base. Urosomites without dorsal ornamentation (Figs 1A, 4E); penulti- mate and anal somites with multiple rows of spinules around ventral hind margin (Fig. 5A). Caudal rami (Fig. 4E) about twice as long as wide, parallel; slightly tapering towards rear margin, with stepped outer margin marking insertion sites of setae I, II and III; produced into conical 7 process bearing terminal pore; posterior third with ventral spinular patch (Fig. 5A). Setae -II minutely bipinnate, spiniform and strongly developed. Seta III bipinnate. Setae IV—V basally fused; about equally long and only slightly longer than caudal ramus; without fracture planes, multipinnate and spiniform. Seta VI minute, bare; seta VII small, biarticulate at base, bare. Rostrum (Fig. 1A) triangular with rounded anterior margin, com- pletely fused to cephalothorax; with numerous dorsal surface pores as figured, none on ventral surface; with minute lateral sensillae near apex. Antennule (Fig. 2A) slender, 7-segmented; segment 7 longest. Plumose setae present on segments 14. Segment 1 with small pore near seta and few short spinules along anterior margin. Armature formula: 1-[1 plumose], 2-[9 + 3 plumose], 3-[4 + 3 plumose + 1 transformed], 4-[1 + 1 plumose + (1 transformed + ae)], 5-[1], 6-[3], 7-[8 + acrothek]. Apical acrothek consisting of aesthetasc, long transformed seta and short bare seta. Transformed setae on segments 3, 4 and 7 long and aesthetasc-like, with rounded tip; those on segments 4 and 7 basally fused to aesthetasc. Rudimentary element present at base of acrothek. Antenna (Fig.3A) 4-segmented, comprising coxa, basis and 2- segmented endopod. Coxa well developed, bare. Basis and proximal endopod segment without ornamentation; unarmed. Exopod inserted in membranous area between basis and endopod; represented by small, well defined segment bearing 2 strong recurved setae apically; exopodal setae multipinnate with long setules in proximal third. Distal endopod segment (Fig. 3A, B) with several surface frills and minute spinules on outer surface and patch of long setules on medial surface; lateral armature consisting of | naked seta; distal armature consisting of 5 apical, non-geniculate, bipinnate or multipinnate elements, 2 of which spiniform, recurved and bearing long spinules proximally. Labrum (Fig. 3C) large, with 6 secretory pores on anterior sur- face; distal margin spinulose medially and with spinular patch on either lateral lobe. Mandible (Fig. 3D) reduced. Palp represented by single naked seta. Gnathobase long and narrow, stylet-like; produced into number of cuspidate processes apically and subapically; without dorsal seta(e). Paragnaths (Fig. 3C) well developed hirsute lobes. Maxillule (Fig.3E) reduced; represented by small bilobed seg- ment bearing 2 naked apical spines and raised seta along outer margin; posterior surface with distinct pore. Maxilla (Fig. 3F) 2-segmented, comprising elongate syncoxa and allobasis. Syncoxa with expanded basal portion and 2 endites; exit of maxillary gland large (arrowed in Fig. 3F), partly concealed under lobate extension; proximal endite represented by small cylindrical process bearing very long plumose seta, distal endite cylindrical, with | naked and 2 pinnate spines apically. Allobasis with large articulating claw distally, smaller inner pinnate spine and naked seta along outer margin. Maxilliped (Fig. 4A, B) very large, articulating with well devel- oped pedestal; 3-segmented, comprising syncoxa, basis and endopod. Syncoxa extremely elongate, longer than basis; without ornamenta- tion but with | anterior, plumose seta near membranous articulation with basis. Basis elongate; distal third of palmar margin with double spinule row (anterior spinules coarser than posterior ones) and 2 elements located closely to articulation with endopod; proximal element spiniform and bare (arrowed in Fig. 4B), distal element pad- like and spinulose. Endopod represented by short segment bearing short naked claw; accessory armature consisting of 3 anterior and 2 posterior elements. Swimming legs with wide, narrow intercoxal sclerites and well R. HUYS AND S. CONROY-DALTON Libr ayygi iii ‘ “as | = < j LEE, ( rr Fig. 2 Clytemnestra scutellata Dana, 1847. A, Antennule @, dorsal; B, antennule 4, ventral; C, antennulary segment 3 6, anterior; D, antennulary segments 4~7 d, anterior [distal portion of segment 7 and proximal portion of segment 4 omitted]; E, antennulary segments 5S—6 d, ventral; F, antennulary segment 7 2, distal portion, dorsal [arrow indicating rudimentary element]. [A based on neotype]. GENERIC CONCEPTS IN CLYTEMNESTRIDAE Fig. 3 Clytemnestra scutellata Dana, 1847 (). A, Antenna, outer; B, distal antennary endopod segment, inner; C, oral area showing position of labrum, paragnaths, mandibles, maxillules and right maxilla [position of maxilliped (Mxp.) indicated], ventral; D, mandible, posterior; E, maxillule, posterior; F, maxilla [exit of maxillary gland arrowed], posterior. [all based on neotype]. R. HUYS AND S. CONROY-DALTON BD 50 —S— — — — ——~ LSS) \ LNT wi — Ny. eS “ LS SS so Ape SSS Fig. 4 Clytemnestra scutellata Dana, 1847. A, Maxilliped , posterior; B, maxilliped 9 distal half of basis and endopod, anterior [proximal palmar element arrowed]; C, maxilliped d, anterior; D, maxilliped d, distal portion of basis and endopod [proximal palmar element arrowed], posterior; E, right caudal ramus @, dorsal; F, right caudal ramus 4, dorsal. [A, B, E based on neotype]. iil GENERIC CONCEPTS IN CLYTEMNESTRIDAE il i] anhth a Va = iy, 100 A ZZ We) SS BEE Za Ze = go aa [inset showing setae IV—V at full length]; C, P6 3, ventral. [A Fig.5 Clytemnestra scutellata Dana, 1847. A, Urosome 2 ventral; B, urosome 6, ventral based on neotype] 12 developed praecoxa; both without ornamentation. Rami 3-segmented except for P1 exopod. P1 (Fig. 6A) separated from maxillipeds by large membranous area. Coxa and basis prolonged along dorsoventral axis; without surface ornamentation. Basis with plumose outer spine. Exopod 1- segmented, represented by elongate segment bearing long setules along outer margin; with subapical pore and 1 outer, 2 apical and 1 inner setae. Endopod 3-segmented; segments decreasing in size distally, each with anterior pore; enp-1 and -2 with few setules along outer margin, enp-2 and -3 with posterior spinules; enp-1 with very long inner seta; ornamentation of inner elements typically (multi)pinnate, distal elements of enp-3 plumose. P2—P4 (Figs 6B; 7A, B) with transversely prolonged basis bearing short outer seta. Endopods distinctly longer than exopods. Exopodal outer spines setiform with flagellate tip. Exopod segments typically with pore near outer distal corner; without ornamentation; exp-2 outer distal corner linguiform. Endopods with long proximal seg- ment, particularly in P2—P3; segments with anterior pore, setules along outer margin and spinules on posterior surface; setal ornamen- tation typically combination of setular and spinular rows; inner seta of P2—P3 enp-1! short. P1 exp-2 without outer spine. Spine and setal formula of swimming legs as follows: Exopod Endopod Pl 121 1.1.220 P2 R228 2-220 P3 EES 28 e237 P4 1.1.323 P2221 P5 (Fig. 1C) uniramous, laterally displaced; 2-segmented; not extending beyond posterior margin of genital double-somite (Fig. 5A). Basis with short outer seta and anterior pore. Exopod about twice as long as basis, slightly curved inwards; outer margin with 4 pinnate setae; inner margin with long plumose seta; apex and inner margin each with 1 long pinnate seta; anterior surface with 3 pores and spinules near apex and in proximal third. MALE. Total body length from tip of rostrum to posterior margin of caudal rami: 1064 um. Maximum width (337 um) measured at posterior margin of cephalic shield. Body (Fig. 1B) with similar projections as in 2; urosome more slender with genital and first abdominal somites separate (Fig. 5B). Rostrum (Fig. 1B) more obtuse than in &. Antennule (Fig. 2B) slender, distinctly 7-segmented with ances- tral segment XIII completely incorporated into segment 4 (Fig. 2C); haplocer, with geniculation located between segment 6 and 7. Plumose setae present on segments 14. Segment | with small pore near seta and few tiny spinules along anterior margin. Armature formula: 1-[1 plumose], 2-[8 + 3 plumose], 3-[5 + 3 plumose + 1 pinnate + | transformed + ae], 4-[2 + 3 plumose + (1 transformed + ae)], 5- [1 + 1 spine], 6-[2], 7-[9 + 2 modified elements + acrothek]. Apical acrothek consisting of aesthetasc, long transformed seta and short bare seta. Transformed setae on segments 3, 4 and 7 long and aesthetasc-like, with rounded tip; those on segments 4 and 7 basally fused to aesthetasc. Rudimentary element present at base of acrothek (arrowed in Fig. 2F). Segment 6 with 2 patches of spinules on anterior surface (Fig. 2D—E). Segment 7 with 2 fused elements near geniculation (Fig. 2D). Maxilliped (Fig. 4C) much larger than in 2 articulating with well developed pedestal; 3-segmented, comprising syncoxa, basis and endopod. Syncoxa extremely elongate but not distinctly longer than basis; without ornamentation but with 1 short anterior seta near R. HUYS AND S. CONROY-DALTON membranous articulation with basis. Basis elongate; more swollen than in 9; middle and distal thirds of palmar margin forming longitudinal furrow bordered by single row of spinules on both anterior and posterior sides; with 2 elements located closely to articulation with endopod; proximal element spiniform and bare (arrowed in Fig. 4D), distal element pad-like and spinulose. Endopod represented by short segment produced into very long naked claw which in reflexed position typically fits in palmar furrow with the apical part closely adpressed onto the anterior surface of the basis; accessory armature consisting of 3 anterior and 2 posterior setae; claw with spatulate apex. P5 (Fig. 7C) very similar to that of 2, with identical proportions, pore pattern and setation. Sixth pair of legs (Fig. 5B) weakly asymmetrical, forming highly membranous midventral area covering single, large median genital aperture; each P6 produced into cylindrical process (Fig. SC) with 1 apical and 2 outer bare setae; few spinules along inner margin. Urosomites 4—5 and anal somite with spinules around ventral hind margin (Fig. 5B). Caudal rami (Fig. 4F) somewhat shorter than in 9; seta II rela- tively longer; seta III more slender and with longer pinnules; setae IV-V long (60% of urosome length; Fig. 5B) and plumose; seta VI much longer than in 2 and sparsely plumose. Spermatophore with very long, recurved neck. VARIABILITY. The right distal exopod segment of the male P2 has only 2 outer spines (Fig. 6C). REMARKS. There are very few published records of C. scutellata that can be verified absolutely. There is little doubt that the species described by Poppe (1891) under the name C. hendorffi is synony- mous with C. scutellata. Poppe’s detailed description shows similar posterolateral projections on the cephalothorax which are absent in the other species from the Great Barrier Reef. C. hendorffi also shows great consistency in body size (2: 1.09 mm; d: 1.07 mm), relative proportions of the caudal rami and P5, and the ventral view of the female urosome demonstrates the absence of spinular patches on the second abdominal somite. The only significant discrepancy is found in the armature of the P2 exopod which Poppe had figured with an outer spine on the proximal segment. The absence of this element is a generic character and we suspect that Poppe had assumed its presence to be the rule in clytemnestrids and had altered his figure accordingly. Poppe’s (1891) material came from two localities in the Indian Ocean (West Australian Basin, south of Madagascar), three localities in the southwest Atlantic off the coasts of Brazil and Argentina, and the Karimata Strait in the Java Sea. He also re-identified Thompson’s (1888) material of Goniopsyllus rostratus from the Maltese Sea as C. hendorffi, confirming its presence in the Mediterranean. From a zoogeographical point of view (see below) it appears more conceivable that Thompson had collected the species described by Claus (1891a) under the name Goniopelte gracilis, the description of which was unknown to Poppe (1891). We have been unable to confirm the presence of C. scutellata in the Atlantic or the Mediterranean and therefore suspect that Poppe’s records from the southwest Atlantic might have been based on another species, possibly C. gracilis. Poppe based his illustra- tions on specimens from the West Australian Basin, suggesting an Indo-Pacific distribution pattern for C. scutellata. The redescription by Giesbrecht (1892) has long been accepted as the basis for identification of C. scutellata even though his material was not from the type locality. However, from our revision it is clear that Giesbrecht had redescribed Goniopelte gracilis (see below). Both species are closely related, sharing the posterolateral projec- tions on the cephalothorax and the presence of 3 outer spines on GENERIC CONCEPTS IN CLYTEMNESTRIDAE Ki \ W AN \\ AN \\ WR | + \ \ KALA SANK AN iN " ry SSS SSSA o~ Ss SS S Fig. 6 Clytemnestra scutellata Dana, 1847. A, P1 9, anterior; B, P2 Q anterior; C, right P2 exp-3 d, anterior, aberrant setation. [A, B based on neotype]. 18 R. HUYS AND S. CONROY-DALTON 14 BRASS SY A 50 50 Fig. 7 Clytemnestra scutellata Dana, 1847. A, P3 @, anterior; B, P4 9, anterior; C, P5 3, anterior. [A, B based on neotype]. GENERIC CONCEPTS IN CLYTEMNESTRIDAE P2—P4 exp-3 and 6 elements on the P5 exopod in both sexes. They can be separated by body size, length of caudal ramus setae IV—V, length of the PS in both sexes and urosome ornamentation in the female (Table I). Clytemnestra gracilis (Claus, 1891a) comb. nov. Goniopelte gracilis Claus, 1891a: 1-10; Taf. III. Clytemnestra scutellata Dana, 1847 sensu Giesbrecht (1892): 568— 572; Taf. 1, fig. 9; Taf. 45, figs. 16-18, 21, 23-24, 27-30, 32, 34-38. Clytemnestra rostrata (Brady, 1883) sensu T. Scott (1894): 106— 107; Pl. XII, figs. 47-57; Pl. XII, figs. 1-3. Clytemnestra scutellata Dana, 1847 sensu Sars (1921): 100-101; PI. LXVIII. Clytemnestra scutellata Dana, 1847 sensu Vilela (1968): 44; Est. XVII, fig. la—c. Clytemnestra scutellata Dana, 1847 sensu Boxshall (1979): 232; Fig. 1SA-K. Clytemnestra scutellata Dana, 1847 sensu Huys et al. (1996): 301; Fig. 120H. TYPE LOCALITY. Claus (1891a) collected his material from an unspecified locality in the eastern Mediterranean. The neotype designation below redefines the type locality as follows: North-east Atlantic, south-west of Azores, 35°N 33°W, 0-1 m. TYPE MATERIAL. Claus’ (1891a) description was based on a single specimen of either sex. Since the type material no longer exists a neotype is designated here to secure stability of nomenclature: adult 2in alcohol (BMNH 1999.1024); collected during RRS Discovery Cruise 121 (5-26 June 1981), station 10379; 13 June 1981, at night; torpedonet; leg. Institute of Oceanographic Sciences. OTHER MATERIAL EXAMINED. (a) from type locality: 11 22 and 8 dd in alcohol (1 2 and 1 3 dissected in half, in separate vials), 1 2dissected on 6 slides (BMNH 1983.53); 2 2Pand 1 d on SEM stub; collection data as for neotype; (b) Gulf of Guinea, Telegraph Steamer Buccaneer (BMNH 1999.1007—-1016): 9 22 (2 damaged) and 1 6 (damaged); misla- belled as Clytemnestra rostrata, January—February 1886; leg. J. Rattray, det. T. Scott. [body length of 7 29: 1381-1541 um, x= 1444 um]; (c) South Adriatic, Croatia: 1 2in alcohol (BMNH 1999.1071); leg. F, Krsinié. [body length: 1309 pm]. DESCRIPTION. (based on Discovery material) FEMALE. Total body length from tip of rostrum to posterior margin of caudal rami: 1330-1562 um (xX = 1450 um; n = 10). Maximum width (382 um) measured at posterior margin of cephalic shield. | Posterolateral angles of cephalothorax slightly expanded (Fig. 8A). | General body shape as in type species. Genital double-somite (Fig. 8B) slightly constricted bilaterally; original segmentation marked by paired transverse chitinous ribs | lateroventrally and laterally, joining medially forming continuous | but weakly defined rib. Copulatory pore slit-like, located medially | between genital apertures (arrowed in Fig. 27B); leading to short posteriorly directed, membranous duct connected to bilobate semi- ' nal receptacle. Genital apertures (Fig. 11D) separated by number of | rounded swellings (also present in type species: Fig. SA); closed off by small opercula derived from vestigial P6; each with 1 vestigial seta (coarser than in C. scutellata) at inner distal corner and anterior | tube-pore near base (arrowed in Fig. 11D). Urosomites without dorsal ornamentation; penultimate and anal | somites with multiple rows or patches of spinules around ventral 15 hind margin and lateroventral patches on second abdominal somite (Fig. 8B). Caudal rami (Fig. 8B) as in C. scutellata but setae IV distinctly shorter than seta V. Rostrum (Figs 8A; 10C) triangular with rounded anterior margin, completely fused to cephalothorax; with numerous dorsal surface pores; minute lateral sensillae flanking middorsal raised pore. Antennule 7-segmented, with armature formula as in type species. Antenna, mandible (Fig. 10A), maxillule and maxilla (proximal endite on syncoxa present) as in type species. Palmar elements of maxilliped as in Fig. 10B; proximal element fused to basis and with apical pore; distal element pad-like, forming barbed, linguiform extension posteriorly and bearing double spinule row and tube pore anteriorly. P2—P4 armature formula: exopod endopod Pp 1.0228 2221 P3 1.1.323 1.2.32] P4 1323 1.2.221 P5 (Fig. 8B) elongate, extending clearly beyond posterior margin of genital double-somite. Exopod about 2.4 times as long as basis, with 6 setae. MALE. Total body length from tip of rostrum to posterior margin of caudal rami: 1420-1531 um (X= 1479 um; n=8). Body with similar projections as in 2; urosome more slender with genital and first abdominal somites separate (Fig. 9A). Antennule with armature as in C. scutellata. Maxilliped much larger than in 9; middle and distal thirds of palmar margin forming longitudinal furrow bordered by single row of spinules on both anterior and posterior sides (Fig. 10D). P5 (Fig. 9A) very similar to that of 9, extending to distal margin of first abdominal somite. Sixth pair of legs (Fig. 9A) weakly asymmetrical, forming highly membranous midventral area covering single, large median genital aperture (Fig. 11A); each P6 produced into cylindrical process (Fig. 11B) with 1 apical and 2 lateral bare setae. Urosomites 4—5 and anal somite with spinules around ventral hind margin (Fig. 9A). Caudal rami (Fig. 9A—B) longer and more slender than in 9; setae III bare; setae IV—V long (68% of urosome length; Fig. 9A) and plumose; seta VI longer than in 2 and sparsely plumose. VARIABILITY. Some variability was noticed in the caudal ramus length of the Buccaneer females, the majority having a slightly longer ramus than in Fig. 8C. In the Adriatic 2 the spinular patches on the first postgenital somite are wider medially forming an almost continuous zone around the posterior margin. REMARKS. Claus (1891b) himself surmised that Goniopelte graci- lis was conspecific with Clytemnestra hendorffi which in turn became relegated to a junior subjective synonym of C. scutellata by Giesbrecht (1892). It is beyond any doubt that Giesbrecht’s excel- lent redescription of C. scutellata was based on C. gracilis. His illustrations were based on Naples material only, however, it is likely that he included specimens of C. scutellata from the Pacific (Giesbrecht, 18915) in his length measurements, possibly account- ing for the lower end of his size range (2: 1.05-1.2 mm; ¢: 1.07-1.3 mm). C. gracilis is distinctly larger than C. scutellata and can be distinguished from the latter by the slender caudal rami and the longer P5 which extends clearly beyond the posterior margin of the genital double-somite in the female and reaches to the rear margin of 16 R. HUYS AND S. CONROY-DALTON a as t \ Arte 4 =N x. Fig. 8 Clytemnestra gracilis (Claus, 1891a) comb. nov. (2) A, Habitus, dorsal; B, urosome, ventral; C, anal somite and right caudal ramus, dorsal. GENERIC CONCEPTS IN CLYTEMNESTRIDAE Fig.9 Clytemnestra gracilis (Claus, 1891a) comb. nov. (d) A, Urosome, ven dorsal. Clytemnestra farrani sp. nov. C, P2 exp-3 9, anterior; D, PS 9, anterior; E, P5 6, anterior. tral [inset showing setae [V—V]; B, anal somite and right caudal ramus, R. HUYS AND S. CONROY-DALTON Fig. 10 Clytemnestra gracilis (Claus, 1891a) comb. nov. SEM photographs. A, Mandibular gnathobase 9; B, maxilliped 9, palmar elements; C, rostrum 9, frontal; D, maxilliped ¢, palmar furrow. GENERIC CONCEPTS IN CLYTEMNESTRIDAE Fig. 11 Clytemnestra gracilis (Claus, 1891a) comb. nov. SEM photographs. A, Genital aperture and sixth legs 5; B, P6 d; D, genital field 9 [position of copulatory pore arrowed]. Goniopsyllus clausi sp. nov. C, Genital aperture and sixth legs 6. 20 the first abdominal somite in the male. Females of both species can be differentiated by the ventral ornamentation pattern of the urosome (C. gracilis has lateral spinular patches on the first postgenital somite) and the ventral transverse chitinous ridge (marking the original segmentation of the genital double-somite) which is more strongly developed in C. gracilis. Giesbrecht (1892) did not illus- trate the second abdominal somite in the female, however, stated in the text that spinules were present ventrally around the posterior margin of all three postgenital somites. Caudal ramus seta IV is distinctly shorter than seta V in females of C. gracilis (see also Giesbrecht (1892): Taf. 45, Fig. 27; Sars (1921): Plate LX VII), while both setae are equally long in the female of the type species. Both sexes of C. gracilis have a propensity for developing asymme- try in the caudal rami whereby one ramus is markedly narrower than the other (see also Claus (1891a): Taf. I, Figs 1-2; Giesbrecht (1892): Taf. 45, Fig. 27). Despite his own arguments to the contrary, T. Scott (1894) inexplicably identified his clytemnestrid material from the Gulf of Guinea as C. rostrata. A. Scott (1909) re-identified the material as C. scutellata. Re-examination of the Buccaneer material (BMNH 1893.4.22.268-275) has revealed it to be an amalgamate of two species, containing 9 22 and 1 6 of C. gracilis and 7 99 of a smaller Goniopsyllus sp. This might explain the discrepancy found between the body length reported by T. Scott (1.25 mm) and our measurements (X = 1.44 mm). Since males are usually larger than females (Giesbrecht, 1892) it is doubtful whether Marques’ (1973) male specimen (0.99 mm) of C. scutellata from S40 Tomé (Gulf of Guinea) belongs to C. gracilis. The only illustrated record of C. scutellata from northern Europe is that by Sars (1921) who found a single female in Oslofjord and described it in great detail. His specimen, 1.24 mm in length, agrees in all aspects with C. gracilis and represents a significant range extension for this species. Kasturirangan (1963) reproduced Giesbrecht’s (1892) and Sars’ (1921) drawings of C. gracilis in his identification key to the planktonic copepods of Indian coastal waters, however its presence in the Indo-Pacific has yet to be confirmed. Vilela (1968) reported two females of C. scutellata, measuring 1.24-131 mm, from the Portuguese coast off Lisbon. Her illustra- tions of the caudal rami and PS positively identify her material as C. gracilis. Clytemnestra farrani sp. nov. TYPELOCALITY. Great Barrier Reef, Queensland, Australia. Farran (1936) recorded a total of 5 specimens (4 belonging to C. farrani, 1 to C. longipes) from serial townettings (his stations 62, 65, 68) at 3 miles east of the laboratory on Low Island (off Port Douglas); depth 32 m. ETYMOLOGY. This patronym commemorates the late G.P. Farran for his comprehensive contributions to our knowledge of planktonic copepods. TYPE MATERIAL. Holotype 2 dissected on 6 slides (BMNH 1999.998); paratypes are 1 2 and 2 dd in alcohol (BMNH 1999. 999-1001). This material was originally registered as C. scutellata under reg. no. 1948.4.28.121. Collected during Great Barrier Reef Expedition 1928-29 on either 15 June (stn 62), 10 July (stn 65) or 18 July 1929 (stn 68). OTHER MATERIAL EXAMINED. From R. Béttger-Schnack: 1 @ in alcohol (BMNH 1999.1065); southern Red Sea, Meteor cruise 5/5, stn 703 (15°34.8' N, 41°54.9' E); 03 August 1987; multiple opening- R. HUYS AND S. CONROY-DALTON closing net, 0.055 mm mesh, vertical hauling, 0-50 m (total water depth 970 m). DESCRIPTION. FEMALE. Total body length from tip of rostrum to posterior margin of caudal rami: 927—946 um (x = 937 um; n = 2). Maximum width (252 um) measured halfway the cephalic shield length. Posterola- teral angles of cephalothorax rounded, not expanded (Fig. 12A). Backwardly produced alate processes of somites bearing P2—P4 distinctly shorter than in C. scutellata and C. gracilis. Genital double-somite (Fig. 13A) not constricted bilaterally; original segmentation marked by small, paired, chitinous patches lateroventrally. Genital field as in type species. Urosomites without dorsal ornamentation; penultimate and anal somites with multiple rows or patches of minute spinules around ventral hind margin and with lateroventral spinular patches on second abdominal somite (Fig. 13A). Caudal rami (Fig. 13A, C) shorter than in previous species; setae IV slightly shorter than seta V but both setae distinctly shorter than in C. scutellata (only slightly longer than ramus and as long as seta III) and minutely pinnate. Rostrum (Fig. 12A) rounded anteriorly, obtuse. Antennule 7-segmented, with armature formula as in type species. Antenna, mouthparts (proximal endite on maxillary syncoxa present) and maxillipeds as in type species. P2 exp-3 with only 2 outer spines (Fig. 9C). P2—P4 armature formula: exopod endopod P2 He222, 220 P3 1323 APRS) P4 E328 1.2.221 P5 (Fig. 9D) extending to posterior margin of genital double- somite. Basis short, exopod about 3 times as long as basis, with 5 setae (3 outer, 1 apical, | inner). MALE. Total body length from tip of rostrum to posterior margin of caudal rami: 939-945 um (x = 942 um; n=2). Maximum width (257 lum) measured at posterior margin of cephalic shield. Body (Fig. 12B) with similar projections as in 2; urosome more slender with genital and first abdominal somites separate (Fig. 13B). Antennule, antenna, mouthparts and maxilliped with armature as in C. scutellata. P5 (Fig. 9E) distinctly shorter than in 2, not extending to distal margin of first abdominal somite; exopod 1.9 times as long as basis, apical and inner setae shorter than in @. Sixth pair of legs (Fig. 13B) weakly asymmetrical; each P6 produced into short cylindrical process with 1 outer and 2 apical bare setae. Urosomites 4—5 and anal somite with spinules around ventral hind margin (Fig. 13B). Caudal rami (Fig. 13B) stubbier than in 2; setae I-II bare; setae IV—-V very long (95% of urosome length) and plumose; seta VI much longer than in 9. REMARKS. C. farrani can be readily distinguished from its conge- ners by the swimming leg setal formula, showing only 2 outer spines on P2 exp-3 but 3 outer spines on P3—P4 exp-3. It is closely related to C. asetosa which resembles it in the small size, the absence of posterolateral processes on the cephalothorax and the presence of only 5 setae on the P5 exopod. The number of endites on the syncoxa, the spinulation pattern on the female urosome and the GENERIC CONCEPTS IN CLYTEMNESTRIDAE v. A, Habitus 2, dorsal; B, habitus 3, dorsal [inset showing setae IV—V at full length]. isp. no Clytemnestra farran Fig. 12 22 R. HUYS AND S. CONROY-DALTON Say ayy AAA Oy =—— ie] eee 0 MUI agts ager i DRAK\ A} tH Hed Sinyr nae HAV ghd g lta tina SUH id Sa Sea S— 7 ae 4 eee Nyy 112 ‘3 All: x ea iN = ca x So \ : od A A o hae i ; o DUO ASE AULA Ae MLN GEAR H YE AM 1 A ALIAY AL EP ACL eT My, SHUNT Ferrin Tene arnt f one mit Sees Z SS SN 100 v PPA soy TD HY 1p 0. Wp we A Cay misting (ale AN I ee ee ia Ai wah MAMI ALY a is : WSS oF SAV QV AB Fig. 13 Clytemnestra farrani sp. nov. A, Urosome &, ventral; B, urosome 6 (excluding P5-bearing somite), ventral [inset showing setae IV—V at full length]; C, anal somite and right caudal ramus 9, dorsal. GENERIC CONCEPTS IN CLYTEMNESTRIDAE Zz Zz Zz ZZZZZ AAZAAVAAA 200 SDVZ te —<$<$<——s SS ZLDAZ-_ZZDZE SS ——— ae 2A = =a SESS SSS —=S Fig. 14 Clytemnestra longipes sp. nov. (3). A, Habitus, dorsal; B, urosome, ventral; C, P2 exp-3; D, P5, anterior; E, right P6. @ 24 relative length of the P5 exopod serve to distinguish both species. C. farraniis currently known only from two widely separated localities in the Indo-Pacific, suggesting that itis probably widespread through- out this oceanic basin. Clytemnestra longipes sp. nov. TYPE LOCALITY. Great Barrier Reef — see C. farrani sp. nov. ETYMOLOGY. The species name is derived from the Latin longus (long) and pes (foot), and refers to the very long male P5 and P6. TYPE MATERIAL. Holotype d in alcohol (BMNH 1999.997). This material was originally registered as C. scutellata under BMNH 1948.4.28.121. Collected during Great Barrier Reef Expedition 1928-29 on either 15 June (stn 62), 10 July (stn 65) or 18 July 1929 (stn 68). DESCRIPTION. FEMALE. Unknown. MALE. Total body length from tip of rostrum to posterior margin of caudal rami: 1211 um. Maximum width (362 um) measured at posterior margin of cephalic shield. Posterolateral angles of cephalothorax angular, weakly produced (Fig. 14A). Backwardly produced alate processes of somites bearing P2—P4 well developed. Urosome with genital and first abdominal somites separate (Fig. 14B). Urosomites without dorsal ornamentation; all postgenital somites with multiple rows of minute spinules around ventral rear margin, those on urosomites 3, 5 and 6 arranged in paired patches either side of ventral midline (Fig. 14B). Caudal rami (Fig. 14B) with bare seta II and minutely pinnate setae I and III; setae IV—V long (54% of urosome length) and plumose. Rostrum (Fig. 14A) rounded anteriorly, protruding. Antennule, antenna, mouthparts (proximal endite on maxillary syncoxa present) and maxillipeds as in type species. P2—P4 exp-3 with only 2 outer spines (Fig. 14C). P2—P4 armature formula: exopod endopod P2 1.1.222 12221 P3 1322 232i P4 S22 P2221 P5 (Fig. 14D) narrow and elongate, extending to distal margin of first abdominal somite (Fig. 14B); exopod 2.7 times as long as basis; with 3 outer seta and | long seta at apex and subdistal inner corner. Sixth pair of legs (Fig. 14E) forming very long cylindrical process with 1 apical and 2 outer bare setae. REMARKS. The male of this species differs from all known males in (1) the ventral ornamentation pattern of the urosome, displaying spinules on all postgenital somites, and (2) the extreme elongation of the P5 and P6 (the distribution pattern of the 3 elements on the latter indicate that allometric growth must have happened prima- rily in the apical portion of the cylindrical process). C. longipes has the same swimming leg setal formula as C. asetosa but, in addition to the characters listed above, differs from the latter in body size and the presence of the proximal endite on the maxillary syncoxa. R. HUYS AND S. CONROY-DALTON Clytemnestra asetosa sp. nov. TYPE LOCALITY. Suez Canal. Port Taufig, Bay of Suez (Egypt). ETYMOLOGY. The species name alludes to the absence of the proximal enditic seta on the maxillary syncoxa. TYPE MATERIAL. Holotype d dissected on 10 slides (BMNH 1999.1025). Paratypes in alcohol are 3 22,2 dd (1 damaged) and 1 cop. V 6 (BMNH 1999.1026-1031); collected during the Cam- bridge Expedition to the Suez Canal, 1924. This material was originally identified as C. scutellata by Gurney (1927) and Boxshall (i979): OTHER MATERIAL EXAMINED. From R. Bottger-Schnack: 3 copepodid II stages in alcohol (BMNH 1999.1066—1068); central Red Sea, Meteor cruise 5/5, stn 682 (21°13.9'N, 38°05.7' E); 25 July 1987; multiple opening-closing net, 0.055 mm mesh, vertical haul- ing, 10-50 m (total water depth 1890 m). DESCRIPTION. FEMALE. Total body length from tip of rostrum to posterior margin of caudal rami: 758-830 um (x = 801 um; n= 3). Maximum width (226 um) measured halfway down the cephalic shield. Posterola- teral angles of cephalothorax rounded, not produced. Backwardly produced alate processes of somites bearing P2—P4 distinctly shorter than in C. scutellata and C. gracilis. General body shape (Fig. 15A) very similar to that of C. farrani (Fig. 12A). Genital double-somite (Fig. 15B) weakly constricted bilaterally; original segmentation marked by minute, paired chitinous patches ventrally. Genital field as in type species. Urosomites without dorsal ornamentation; penultimate and anal somites with multiple patches of minute spinules around ventral hind margin (Fig. 15B). Caudal rami (Fig. 15C) with bare setae I and II; setae IV slightly shorter than seta V, both plumose. Rostrum (Fig. 15A) rounded anteriorly, not distinctly delimited from cephalic shield. Antennule (Fig. 16A) 7-segmented, with reduced armature on segments 2 and 3. Armature formula: 1-[1 plumose], 2-[9 + 1 plumose], 3-[3 + 3 plumose + 1| transformed], 4-[1 + 1 plumose + (1 transformed + ae)], 5-[1], 6-[3], 7-[8 + acrothek]. Antenna with weakly defined exopod (Fig. 17G); one seta fused basally to segment. Mandible (Fig. 16B). Palp represented by minute seta; gnathobase with large lateral tooth (arrowed in Fig. 16C). Maxillule (Fig. 16D) produced into distal lash (derived from armature element); with | lateral seta and 1 apical spine. Maxilla (Fig. 16E) as in type species except for absence of proximal endite on syncoxa (position in other species arrowed in Fig. 16E). Maxilliped as in C. scutellata. Pl (Fig. 17A) asin C. scutellata but setules along inner margin of enp-1 absent. P2—P4 (Fig. 17B—D) with only 2 outer spines on exp- 3. P2—P4 armature formula: exopod endopod PD 1.1.222 1.2.221 128} TEES 22, 12321 P4 1.1.322 12221 P5 (Fig. 17E) nearly extending to posterior margin of genital double-somite. Basis short, exopod about 2.5 times as long as basis, with 5 setae (3 outer, 1 apical, 1 inner). GENERIC CONCEPTS IN CLYTEMNESTRIDAE — ry YON aE A by Santa Wy T9979 vp, SOU — Wie oly, apc Ips B 25 Fig. 15 Clytemnestra asetosa sp. nov. (9). A, Habitus, dorsal; B, urosome, ventral; C, anal somite and right caudal ramus, dorsal. 25 26 R. HUYS AND S. CONROY-DALTON Fig. 16 Clytemnestra asetosa sp. nov. (). A, antennule, ventral [inset showing acrothek at full length]; B, mandible, posterior; C, mandibular gnathobase, other view [secondary tooth arrowed]; D, maxillule; E, maxilla, posterior [small arrow: exit of maxillary gland; large arrow indicating position of proximal endite in other Clytemnestra species]. GENERIC CONCEPTS IN CLYTEMNESTRIDAE “a ZZ bs S 7 i Uf Fig. 17 Clytemnestra asetosa sp. nov. (9). A, P1, anterior; B exopod, anterior; D, P4, distal portion of basis and exopod, anterior; E, PS, anterior; F, rostrum, dorsal; G, antennary exopod. , P2, intercoxal sclerite, protopod and exopod, anterior; C, P3, distal portion of basis and 7] R. HUYS AND S. CONROY-DALTON dal ramus, dorsal. ntral; C, P5, anterior; D, P6; E, anal somite and right cau ome, ve v. (3). A, Habitus, dorsal; B, uros sp. no etosa Fig. 18 Clytemnestra as GENERIC CONCEPTS IN CLYTEMNESTRIDAE MALE. Total body length from tip of rostrum to posterior margin of caudal rami: 920 um (n= 1). Maximum width (232 um) measured at posterior margin of cephalic shield. Body (Fig. 18A) with similar projections as in 9; urosome more slender with genital and first abdominal somites separate (Fig. 18B). Antennule, antenna, mouthparts and maxilliped with armature as in C. scutellata. P5 (Fig. 18C) as in 2 not extending to distal margin of first abdominal somite (Fig. 18B). Sixth pair of legs (Fig. 18B, D) weakly asymmetrical; each P6 produced into short cylindrical process with 1 outer and 2 apical bare setae. Urosomites 4—5 and anal somite with spinules around ventral hind margin (Fig. 18B). Caudal rami (Fig. 18B, E) with bare setae I-II; setae [V—V very long (75% of urosome length) and plumose; seta VI much longer than in 9. REMARKS. The early copepodid stages from the central Red Sea were identified on the basis of the absence of the proximal endite of the maxilla and the shape of the cephalothorax. C. asetosa, origi- nally identified as C. scutellata by Gurney (1927), is the smallest species in the genus. It is similar to C. farrani in many respects but differs from it in the armature formula of the antennule, the loss of the proximal endite of the maxilla, the presence of only 2 outer spines on P3—P4 exp-3 and a different spinulation pattern on the female urosome. The species is thus far known only from the Red Sea and the Bay of Suez. Clytemnestra hendorffi var. quinquesetosa Poppe, 1891 Poppe (1891) distinguished this variety on the basis of the following characters: (1) female P5 exopod distinctly longer and bearing 5 setae; (2) urosome of both sexes less slender; (3) caudal rami relatively wider proximally. This variety was collected from two localities in the Java Sea. Most authors have followed Giesbrecht’s (1892) decision to discard this variety and regarded it as a synonym of C. scutellata. Our revision has revealed that only C. scutellata and C. gracilis display 6 setae on the P5 exopod and that there are at least three species in the Indo-Pacific which have only 5 setae. As far as we could ascertain from the collections examined P5 setation is never variable within populations and always identical between sexes. Since Poppe (1891) did not provide any figures it is imposs- ible to make any positive statement as to the identity of his material. Other records Chen et al. (1974) reported C. scutellata from the East China Sea (one of the areas where Dana originally recorded the species from). Unfortunately the few illustrations of the habitus and female PS are _of no help in determining the specific identity of their material. | Moreover, the extreme body size range (1.0-1.9 mm) strongly | Suggests the co-occurrence of more than one species in their sam- | ples. Cheng et al. (1965) also illustrated C. scutellata from the East China Sea but their species has only 5 setae on the PS exopod, lacks | posterolateral processes on the cephalothorax and has only 2 outer | spines on at least P3 (which was mislabelled as the P2) and P4. Their | reported size range ( 29: 0.86—-1.0 mm; 3d: 0.80-0.85 mm) strongly | Suggests that they had identified C. asetosa or possibly a related | species. Mori’s (1929) description of C. scutellata from the Sea of Japan is equally brief. Posterolateral projections on the cephalothorax | appear to be absent in his material (although they could be obscured | by excessive squashing of the figured specimen), indicating that | Mori was probably dealing with another species. Mori supple- 29 Kazmi & Muniza (1994) present sketchy figures of what they believe to be C. scutellata in their samples from the Arabian Sea. Nothing can be said about the real identity of their material other than that were dealing with a Clytemnestra. The Caribbean records of C. scutellata by Owre & Foyo (1967) and Campos Hernandez & Suarez Morales (1994) require further investigations. Both descriptions show the unique presence of lat- eral protrusions halfway down the cephalothorax which may suggest the occurrence of a distinct species in this region. It is impossible to decide from Legaré’s (1964) inadequate illustrations whether this modification also occurred in his Venezuelan material. Interestingly, Morales & Vargas (1995) show similar protrusions in aclytemnestrid from the Pacific coast of Costa Rica which they identified as C. rostratus but has 7 segments in the antennule. Genus Goniopsyllus Brady, 1883 Sapphir Car, 1890 [type species: S. rostratus Car, 1890 — by monotypy] DIAGNOSIS. Clytemnestridae. Body with dorsal pattern of denticles and spinules on urosomites. Antennule 6-segmented in &, indis- tinctly 7-segmented in d with segments 34 incompletely fused; 3 segmental homologies: 1—I, 2—(II—-VIII), 3-(IX—XI]), 4—XIII, 5- (XIV-XVII), 6-(XVIII-XX), 7-(XXI-XXVIII). Antenna with | lateral and 4 apical elements on distal endopod segment; exopod represented by membranous segment bearing | long seta. Maxillule represented by triangular segment with | apical spine. Maxillary syncoxa with | endite bearing 2 setae. P1 without outer seta on basis; exopod with 3 setae. P2 with outer spine on exp-1. P1—P4 armature formula: exopod endopod Pl 021 1.1.220 P2 1.1.222 1.2.221 P3 1.1.323 Le PeBVeil P4 L323 222 P5 exopod with 5 setae in both sexes. Genital apertures fused in 2 forming common medial slit; closed off by paired P6 bearing 1 well developed seta; copulatory pore located medially in large circular depression halfway the length of the genital double-somite; copulatory duct strongly chitinized. Male P6 asymmetrical, forming membranous opercula closing off single (sinistral or dextral) genital aperture; bearing | seta. Caudal rami convergent, relatively short and conical; not sexually dimorphic. TYPE SPECIES. Goniopsyllus rostratus Brady, 1883 [by monotypy] OTHER SPECIES. G. clausi sp. nov., G. brasiliensis sp. nov. SPECIES INQUIRENDAE. Goniopsyllus tenuis (Lubbock, 1860) comb. nov.; Sapphir rostratus Car, 1890 Since the type species is only known from the damaged female holotype and no other material was available for study, G. clausi sp. nov. is instead selected for the model description. Goniopsyllus clausi sp. nov. Clytemnestra rostrata (Brady, 1883) sensu Giesbrecht (1892): pp. 568-572; Taf. 45, Figs 22, 31. Clytemnestra rostrata (Brady, 1883) sensu Vilela (1965): p. 21; Est. IX, Fig. 2a—e; (1968): p. 44; Est. XVII, Fig. 2a—c. AB Fig. 19 Goniopsyllus clausi sp. nov. A, Habitus °, dorsal; B, habitus of ovigerous 9, lateral. R. HUYS AND S. CONROY-DALTON GENERIC CONCEPTS IN CLYTEMNESTRIDAE Sil Fig. 20 Goniopsyllus clausi sp. nov. A, Antennule 9, ventral; B, distal portion of antennulary segment 6 of 9, ventral [rudimentary element arrowed]; C, antennule 3, ventral; D, antennulary segments 3-6 of 4, anterior; E, antennulary segment 7 of 3, ventral [rudimentary element arrowed]. 32 R. HUYS AND S. CONROY-DALTON Le ~—_— Aa 3 a. SS —— ae L- —> el Fig. 21 Goniopsyllus clausi sp. nov. A, antenna 9, outer; B, distal endopod segment of antenna @, inner [rudimentary elements arrowed]; C, mandible 9; D, mandibular gnathobase 9; E, mandibular gnathobase of d specimen; F, maxillule 9, posterior; G, maxilla &, posterior [exit of maxillary gland arrowed]; H, oral area 2 showing position of antenna (A,), labrum, paragnaths (P), mandible, maxillule, maxilla and maxilliped (Mxp.); I, rostrum Q, dorsal. 33 GENERIC CONCEPTS IN CLYTEMNESTRIDAE WY quan rnannnsnnssssees PL Stl ely epee Mae SSQOOn~sasyy S WWMM yyy Sys epee” (o) medial; D, same, ? maxilliped 2 distal portion of basis and endopod, anterior; C, same 2 B maxilliped 9, anterior; posterior; E, maxilliped d, anterior; F, maxilliped d, distal portion and endopod, anterior; G, maxillipedal basis and endopod 6, medial; H posterior. > Fig. 22 Goniopsyllus clausi sp. nov. A same, > 34 Clytemnestra rostrata (Brady, 1883) sensu Huys et al. (1996): pp. 300-303, Figs 120A—G, 121A—D. Clytemnestra rostrata (Brady, 1883) sensu Boxshall & Huys (1998): p. 782, Fig. 13(a)—(b). Bay of Cadiz, 36°30'N 7°20'W (Spain). ETYMOLOGY. The species is named in honour of Carl Claus, one of the most prolific 19th century copepodologists, who first called attention to the distinctiveness of the clytemnestrid genera. TYPE MATERIAL. Holotype 2 dissected on 10 slides (BMNH 1999.1035). Paratypes are 2 dissected dd (on 2 and 5S slides, respectively), 2 dissected 2(on 1 slide each), and9 2° (1 damaged), 1 3, 4 copepodids (2 Cop V, 1 Cop IV, 1 Cop II) in alcohol (BMNH 1999.1036-1055). In addition, 2 29 and 1 d were prepared for SEM. Donated by J.M. Gee, collected by A. Lindley (Plymouth Marine Laboratory), 1984. TYPE LOCALITY. OTHER MATERIAL EXAMINED. 4 99, 2d: Adriatic Sea, Station CJ-008, Pelegrin, Hvar (Croatia), leg. F. KrSini¢é, ‘Bios’, 23 May 1998 (BMNH 1999.1072-1077). DESCRIPTION. FEMALE. Total body length from tip of rostrum to posterior margin of caudal rami: 979-1067 um (x = 1017 um; n= 8). Maximum width (306 um) measured at posterior margin of cephalic shield. Postero- lateral angles of cephalothorax only weakly expanded laterally but markedly produced posteriorly (Fig. 19A, B). Somites bearing P2— P4 successively decreasing in width posteriorly and bearing backwardly produced alate processes. Genital double-somite (Figs 23A; 27C) slightly constricted bilat- erally; original segmentation marked by two minute chitinous patches ventrally. Copulatory pore (Figs 23C, D; 27A, C) located medially in large circular depression, halfway the length of genital double- somite; leading to anteriorly directed, strongly chitinized duct which at level of P5-bearing somite enters median seminal receptacle. Genital apertures located far anteriorly; closed off by small opercula derived from vestigial P6; each with 1 well developed seta (Figs 23C; 27D). Urosomites with zone of small denticles around dorsal hind margin (not figured in Fig. 19A, but see Fig. 23B); penultimate and anal somites also with larger spinules around ventral hind margin (Fig. 23A). Caudal rami short (Figs 23B; 26A), convergent; conical in shape with stepped inner and outer margins marking insertion sites of setae I, 11 and [V—V; produced into conical process bearing terminal pore; with numerous ventral pores as illustrated in Fig. 26A. Setae LI bipinnate, spiniform and strongly developed; seta I 1.85 times as long as seta II, extending beyond apex of caudal ramus. Seta III minutely bipinnate. Setae IV—V basally fused, without fracture planes, multipinnate and spiniform; seta V about 2.1 times ramus length. Seta VI minute, bare; seta VII biarticulate at base, bare. Rostrum (Figs 19A; 211) triangular and well offset, completely fused to cephalothorax; with numerous dorsal surface pores as fig- ured, none on ventral surface; with minute lateral sensillae near apex. Antennule (Fig. 20A) slender, 6-segmented; segment 6 very long. Plumose setae present on segments 14. Segment | with small pore near seta and few long setules along anterior margin. Armature formula: 1-[1 plumose], 2-[6 + 1 plumose + 3 pinnate], 3-[5 + 2 plumose + | transformed], 4-[1 + 1 plumose + (1 transformed + ae)], 5-[1], 6-[11 + acrothek]. Apical acrothek consisting of aesthetasc, long transformed seta and short bare seta. Transformed setae on segments 3, 4 and 6 long and aesthetasc-like, with minutely spiniform tip; those on segments 4 and 6 basally fused to aesthetasc. Rudimen- R. HUYS AND S. CONROY-DALTON tary element present at base of acrothek (arrowed in Fig. 20B). Antenna (Fig. 21A, B) 4-segmented, comprising coxa, basis and 2-segmented endopod. Coxa well developed, bare. Basis and proxi- mal endopod segment with few surface denticles; unarmed. Exopod inserted in membranous area between basis and endopod; repres- ented by small, weakly chitinized segment bearing strong recurved seta apically; exopodal seta multipinnate, spinules in proximal third distinctly longer. Distal endopod segment with 3 surface frills and minute denticles on outer surface and patch of long setules on medial surface; lateral armature consisting of 1 pinnate seta; distal armature consisting of 1 subapical and 3 apical, non-geniculate, bipinnate or multipinnate elements, 2 of which spiniform, recurved and bearing long spinules proximally; distal margin with 2 rudimentary elements on inner surface (arrowed in Fig. 21B). Labrum (Fig. 21H) large, with 6 secretory pores on anterior surface; distal margin smooth medially and with spinular patch on either lateral lobe. Mandible (Fig. 21C—E) reduced. Palp represented by single na- ked seta. Gnathobase long and narrow, stylet-like; produced into number of cuspidate processes apically and subapically; without dorsal seta(e). Paragnaths (Fig. 21H) well developed lobes without any con- spicuous ornamentation. Maxillule (Fig. 21F) reduced; represented by small triangular segment bearing naked apical seta and raised pore along outer margin. Maxilla (Fig. 21G, H) 2-segmented, comprising elongate syncoxa and allobasis. Syncoxa with expanded basal portion; exit of maxil- lary gland large (arrowed in Fig. 21G), partly concealed under lobate extension; coxal endite cylindrical, with 2 naked setae apically. Allobasis with large articulating claw distally, smaller inner spine and unipinnate seta along outer margin. Maxilliped (Fig. 22A) very large, articulating with well devel- oped pedestal; 3-segmented, comprising syncoxa, basis and endopod. Syncoxa extremely elongate, longer than basis; without ornamenta- tion but with 1 anterior, plumose seta near membranous articulation with basis. Basis elongate; distal third of palmar margin with double spinule row and 2 elements located closely to articulation with endopod (Fig. 22B—D); proximal element spiniform and bare, distal element stubby and spinulose. Endopod represented by short seg- ment bearing naked claw; accessory armature consisting of 3 anterior setae and 2 posterior setae (Fig. 22B—D). Swimming legs with wide, narrow intercoxal sclerites and well developed praecoxa; both without ornamentation. Rami 3-segmented except for Pl exopod. P1 (Fig. 23E) separated from maxillipeds by large membranous area. Coxa and basis prolonged along dorsoventral axis; without surface ornamentation. Basis without inner or outer seta (spine). Exopod 1-segmented, represented by elongate segment bearing long setules along outer margin; with subapical pore and 3 setiform elements distally, outer one less than half the length of others. Endopod 3-segmented; segments decreasing in size distally, each with anterior pore and few spinules/setules along outer margin; enp- 1 with very long inner seta; ornamentation of inner elements typically (multi)pinnate, distal elements plumose. P2—P4 (Figs 24A, B; 25B) with transversely prolonged basis bearing short outer seta. Endopods distinctly longer than exopods. Exopodal outer spines setiform with distinct flagellate tip. Exopod segments typically with pore near outer distal corner; without ornamentation. Endopods with long proximal segment, particularly in P2—P3; segments with anterior pore, setules along outer margin and spinules (enp-2 and -3) or setular tuft (enp-1) on posterior surface; setal ornamentation typically combination of setular and 35 : GENERIC CONCEPTS IN CLYTEMNESTRIDAE x <> SE ara INN Se 50 (iA aa aE “SS. > Sac SR CR EN a. \\ 0 == o “0-6 V4 dif 25 ga y Minas: 8 genital field, ventral; D, genital field, Fig. 23 Goniopsyllus clausi sp. nov. (2). A, Urosome, ventral; B, anal somite and left caudal ramus, dorsal; C, lateral; E, P1, anterior. IN R. HUYS AND S. CONROY-DALTO: 36 \y ee, Fig. 24 Goniopsyllus clausi sp. nov. (2). A, P2, anterior; B, P3, anterior; C, P5, anterior; D, aberrant P5, anterior. 3i)/ GENERIC CONCEPTS IN CLYTEMNESTRIDAE —THFALWY TREE (aa) caarsl| 1 BIOS \ | il \ f ey pe VALS LL eg Jee" — VAFL IT PSL Ba SZ . SOS < SI } G \ \ X Kt q SKK LIES == EES SSS SSS 7 S Le | { Oe (\ <— Es ae = SOS : WY ~~ x 2 = NN Se SSS SSS LAA: g Y \ = < — > “SS SSS ‘ Ry, ~ NEN Y , Y y/ Aan we } 3= 5 ie} vey oO ‘S) it ie) B=) 2 5 OF + a -Q iS an = ie} ue) *o n =) = s x < = (e} i=} a. an a Ss S S a SI S g BS) S 1) ire) a ob —_ ee R. HUYS AND S. CONROY-DALTON 38 SN Ss: ZZ a ze =i = Gs sp I al SS ventral. Goniopsyllus rostratus Brady, 1883 (holotype @). C, Antennule , maxilliped, distal portion of basis and endopod, anterior; F, P5, posterior; G, anal somite and left caudal ®, lateral; B, urosome 6, Fig. 26 Goniopsyllus clausi sp. nov. A, Caudal ramus 13) > (armature omitted); D, maxilliped, anterior; ramus, dorsal. GENERIC CONCEPTS IN CLYTEMNESTRIDAE Fig. 27 Goniopsyllus clausi sp. nov. (2). SEM photographs. A, Circular depression surrounding copulatory pore (position obscured by remnant of spermatophore neck); C, genital double-somite; D, genital aperture. Clytemnestra gracilis (Claus, 1891a) comb. nov. (?). B, Genital apertures and copulatory pore [arrowed]. 40 spinular rows; inner seta of P2—P3 enp-1 short. Spine and setal formula of swimming legs as for genus. P5 (Fig. 24C) uniramous, laterally displaced; 2-segmented, com- prising basis and 1-segmented exopod; not extending to distal margin of genital double-somite (Fig. 23A). Basis with short outer seta and pore near outer distal corner. Exopod about twice as long as basis, slightly curved inwards; outer margin with 2 pinnate setae and 3 pores; inner margin with long plumose seta; apex with 1 pinnate and 1 plumose seta. MALE. Total body length from tip of rostrum to posterior margin of caudal rami: 1021 um (n= 1). Maximum width (304 um) measured at posterior margin of cephalic shield. Body (Fig. 25A) with similar projections as in 9; genital and first abdominal somites separate. Rostrum (Fig. 25A) more pointed than in 2. Antennule (Fig. 20C) slender, indistinctly 7-segmented with seg- ment 4 only demarcated dorsally (Fig. 20D); haplocer, with geniculation located between segment 6 and 7. Plumose setae present on segments 1—5. Segment 1 with small pore near seta and few long setules along anterior margin. Armature formula: 1-[1 plumose], 2-[5 + 5 plumose], 3-[5 + 1 plumose + | pinnate + 1 transformed + ae], 4-[2 plumose], 5-[4 plumose + 1 pinnate+ (1 transformed + ae)], 6-[1 + 2 pinnate spines + 1 smooth spine], 7-[10 + 2 vestigial elements + acrothek]. Apical acrothek consisting of aesthetasc, long transformed seta and short bare seta. Transformed setae on segments 3, 5 and 7 long and aesthetasc-like, with minutely spiniform tip; those on segments 5 and 7 basally fused to aesthetasc. Rudimentary element present at base of acrothek (arrowed in Fig. 20E). Segment 6 with continuous patch of spinules on anterior surface (Fig. 20D). Segment 7 with 2 vestigial elements near geniculation. Maxilliped (Fig. 22E) very large, articulating with well devel- oped pedestal; 3-segmented, comprising syncoxa, basis and endopod. Syncoxa extremely elongate but not longer than basis; without ornamentation but with | anterior, plumose seta near membranous articulation with basis. Basis elongate; more swollen than in &,; middle and distal thirds of palmar margin forming longitudinal furrow bordered by multiple rows of spinules on both anterior and posterior sides; with 2 elements located closely to articulation with endopod; proximal element spiniform and bare, distal element stubby and spinulose. Endopod represented by short segment pro- duced into very long naked claw which in reflexed position typically fits in palmar furrow with the apical part closely adpressed onto the anterior surface of the basis (Fig. 22E, G); accessory armature consisting of 3 anterior setae and 2 posterior setae (Fig. 22F—H). P5 (Fig. 25C) very similar to that of 9, with identical proportions and setation but lateral setae of exopod slightly shorter. Sixth pair of legs (Figs 11C; 26B) asymmetrical, represented by highly membranous non-articulating flaps covering single, large genital aperture (Fig.11C); each lobe with 1 bare seta at outer distal corner. Urosomites 4—S and anal somite with spinules around ventral hind margin (Fig. 26B). Caudal rami (Fig. 26B) slightly more slender than in 9; conical projection wider and setae I-II relatively shorter. Spermatophore with very long, recurved neck (Fig. 26B). VARIABILITY. The left P5 of the holotype @ shows slightly different segmental proportions and pore pattern (Figs 23A; 24D). REMARKS. This species was illustrated by Huys et al. (1996) as ‘Clytemnestra rostrata’. Their brief description which was based on material from the Gulf of Cadiz contains some observational errors. R. HUYS AND S. CONROY-DALTON The most significant is the setation of the maxillule which was actually based on C. gracilis. The armature on the genital field was omitted in their Fig. 120B. The female P5 (their Fig. 121C) also appears shorter but this is to be regarded as the result of excessive squashing during mounting. The distribution of G. clausi is thus far restricted to the Portu- guese coast (Vilela, 1965, 1968) and the Mediterranean with confirmed records from the Bay of Cadiz, Naples and the Adriatic. Sapphir rostratus has also been recorded from the Adriatic but is probably not synonymous with G. clausi (see below). The Naples record refers to Giesbrecht (1892) who found 1 6 of ‘C. rostrata’ in this area but also attributed Pacific specimens (3 29, 2 53) to this species. Goniopsyllus rostratus Brady, 1883 Clytemnestra rostrata (Brady, 1883) Poppe (1891) TYPE LOCALITY. South Atlantic, off Argentinean coast; 42°32'S 56°29' W; net at 54 m depth. MATERIAL EXAMINED. Holotype 2 dissected on slide (reg. no. C.C.46); collected during Voyage of H.M.S. Challenger during the years 1873-1876 (station 318); 11 February 1876. The dissection is imperfect and incomplete (e.g. antenna and P1 are lacking), and the specimen is partly aberrant in the swimming leg setal formula. REDESCRIPTION. FEMALE. Genital double-somite (Fig. 28A) relatively short in comparison with other species, not constricted bilaterally; original segmentation marked by two minute chitinous patches ventrally. Copulatory pore (Fig. 28A) located medially in large circular de- pression, halfway the length of genital double-somite; leading to anteriorly directed, strongly chitinized duct which at level of P5- bearing somite enters median seminal receptacle. Genital apertures located far anteriorly; closed off by small opercula derived from vestigial P6; each with 1 well developed seta. Urosomites with zone of small denticles around dorsal hind margin; penultimate and anal somites also with larger spinules around ventral hind margin (Fig. 28A). Caudal rami short (Figs 26G; 28A), convergent; similar in shape to G. clausi but proportionally smaller. Setae II bipinnate, spiniform and strongly developed; seta I 1.7 times as long as seta II, extending beyond apex of caudal ramus. Seta III minutely bipinnate. Setae [V— V basally fused, without fracture planes, multipinnate and more setiform and distinctly longer than in G. clausi (compare Fig. 23B); seta V about 3 times ramus length. Seta VI minute, bare; seta VII biarticulate at base, bare. Antennule (Fig. 26A) slender, 6-segmented; segment 6 longer than in G. clausi (length ratio segment 6 : segment 5 being 6.0 in G. rostratus, 5.0 in G. clausi). Armature pattern as in G. clausi. Maxilliped (Fig. 26D) with similar armature as in G. clausi but with different spinular ornamentation on palmar margin (Fig. 26E). P2—P4 spine and setal formula of swimming legs as follows (left P3 exp-3 and right P4 exp-3 with aberrant outer spine number): Exopod Endopod Right Left P2 1.1.222 222 1.2.221 P3 Likey3) LES 22 L221 P4 1.1.322 1S 23 1.2.221 P5 (Fig. 26F) 2-segmented, comprising basis and 1-segmented exopod;relative lengths as in G. clausi. Exopod outer margin with 2 GENERIC CONCEPTS IN CLYTEMNESTRIDAE AVY 100 ns TY TL Nt i Ls Cie et i) a <= I Ay atadrnheS a eS —— 21778, Uj Roney i Fig. 28 Goniopsyllus rostratus Brady, 1883 (holotype ¢). A, Urosome (excluding P5-bearing somite), ventral [distorted due to excessive squashing]. Goniopsyllus brasiliensis sp. nov. (2). B, Urosome, ventral; C, genital field, ventral; D, antennule (armature omitted). 42 pinnate setae and 3 pores; inner margin with long plumose seta; apex with | pinnate and 1| plumose seta. MALE. Unknown. REMARKS. Upon re-examination Boxshall (1979) concluded that the holotype, identified by Brady (1883) as a male, was in reality female. The true sexual identity however, had already been noted by both Poppe (1891) and Claus (1891a—b) who based their conclusion on the 5-segmented urosome and the female facies of the antennule and maxilliped. This opinion was also confirmed by Giesbrecht (1892) but not by Car (18915) who continued regarding it as a male on the basis of the internal spermatophore drawn by Brady. The most plausible explanation is that Brady (1883) had misinterpreted the strongly chitinized copulatory duct, a suspicion reinforced by in- spection of the holotype. Giesbrecht (1892: 573) pointed out the discrepancy between the size mentioned in Brady’s text and that inferred from his habitus figure reproduced at x80 magnification. According to Brady the holotype is only 0.65 mm long (*1-40th of an inch’) but Giesbrecht considered 1.16 mm a more realistic figure. Re-examination of the slides strongly suggests that Brady must have made a morphometric error of at least a factor 2. The urosome (excl. P5-bearing somite) which is mounted intact measures 0.43 mm. Extrapolation by using the urosome/body length ratio found in its congeners G. clausi and G. brasiliensis (about 0.3) gives an estimated total body length of 1.43 mm. This large size rules out possible conspecificity with G. brasiliensis (x = 0.96 mm). Brady (1883) assumed all four swimming legs to be similar, having 3-segmented rami and resembling the leg illustrated in his Fig. 15 (i.e. the P2). His lateral habitus view suggests that the P1 possesses 3-segmented exopods and endopods, however Poppe (1891) suspected that Brady had overlooked the exopod and instead had superimposed both left and right endopods. For some unknown reason he assumed the P1 exopod to be 2-segmented but failed to confirm this against the holotype due to the absence of the P1 on Brady’s slide. G. rostratus can be readily identified from the other South- American species G. brasiliensis by the large body size (compare urosomes in Fig. 28A—B drawn at the same scale), the elongate caudal ramus setae [V—V, the long seta I clearly extending beyond the distal margin of the ramus, and additional differences in the ornamentation of the maxilliped (spinule pattern on palmar margin). Brady (1883) also illustrated well developed posterolateral exten- sions on the cephalothorax which are completely absent in G. brasiliensis. Goniopsyllus brasiliensis sp. nov. ? Clytemnestra rostrata (Brady, 1883) sensu Ramirez (1966): 291; Lam. II, figs 12-15. TYPE LOCALITY. Rio Grande do Sul (Brazil); outside opening of Lagoa dos Patos to ocean; 32°11'S 52°7'W. ETYMOLOGY. The species name refers to the type locality. TYPE MATERIAL. Holotype @ dissected on 8 slides (BMNH 1999.1056). Paratypes are 8 2° in alcohol (BMNH 1999.1057— 1064). Collected by G.A. Boxshall, February 1996, plankton haul. DESCRIPTION. FEMALE. Total body length from tip of rostrum to posterior margin of caudal rami: 892-1057 um (x = 958 um; n= 8). Maximum width (265 zm) measured at posterior margin of cephalic shield. Postero- R. HUYS AND S. CONROY-DALTON lateral angles of cephalothorax rounded, virtually not expanded laterally (Fig. 29A). Rostrum (Fig. 29A) rounded and less pro- nounced than in G. clausi. Backwardly produced alate processes of somites bearing P2—P4 distinctly shorter and less pointed than in C. clausi. Integument generally less chitinized than in G. clausi. Genital double-somite (Fig. 28B) not constricted bilaterally and relatively wider than in G. clausi; original segmentation marked by minute, paired, chitinous patches ventrally. Genital field as in G. clausi but with additional pores flanking copulatory pore (Fig. 28C). Urosomites with zone of small denticles around dorsal hind margin (Fig. 29B); penultimate and anal somites also with larger spinules around ventral hind margin (Fig. 28C). Caudal rami (Figs 28B; 29A—C) short, convergent. Setae III bipinnate, spiniform and strongly developed; seta I 1.2 times as long as seta II, not extending beyond apex of caudal ramus. Seta III minutely bipinnate. Setae [V—V basally fused, multipinnate and about as long as in G. clausi but seta IV more resilient (compare Fig. 23B); seta V about 1.5 times ramus length. Seta VI extremely small; seta VII biarticulate at base, bare. Antennule (Fig. 28D) slender, 6-segmented; segment 2 shorter than in G. clausi but armature pattern identical. Mandible and maxillule (Fig. 29D) somewhat more slender than in G. clausi. Maxilliped (Fig. 29E—-F) with similar armature as in G. clausi but with different spinular ornamentation on palmar margin (Fig. 29F). P1—P4 with setal formula as for genus. P5 (Fig. 28B) markedly longer than in G. clausi, extending beyond distal margin of genital double-somite. MALE. Unknown. REMARKS. Although many South-American authors have recorded specimens that they attribute to C. rostrata, there is good reason to believe that in fact often they have mistaken G. brasiliensis for this species. In general, with the discovery of G. brasiliensis many of the Brazilian records of G. rostratus are rendered doubtful (Bjérnberg, 1963; Bjornberg et al., 1981; Campaner, 1985; Carvalho, 1944; Gaudy, 1963; Monti, 1980; Monti & Gloeden, 1986; Monti & Cordeiro, 1988; Santos, 1973; Vega-Perez, 1993). The same applies to Legaré’s (1961, 1964) records of C. rostratus from Venezuelan coastal waters. The species illustrated by Ramirez (1966) as C. rostrata from Mar del Plata in Argentina differs from the one figured in his later paper (Ramirez, 1970) by the complete absence of posterolateral projections on the cephalothorax and is almost cer- tainly conspecific with G. brasiliensis. The author described the female antennule as 7-segmented but this clearly contradicts his illustration which shows only 6 segments as in other species of Goniopsyllus. The only anomaly remaining is the body size which according to Ramirez (1966) is 1.8 mm for the female and 1.5 mm for the male. Based on his illustrations and the accompanying scale bars the female only measures 0.74 mm and the male 0.77 mm. It is not clear whether Carvalho’s (1952) material of C. rostrata, consisting of 5 males from the Bay of Santés (Sao Paulo State), also belongs to C. brasiliensis. His size range (0.50—0.85 mm) precludes possible identity with C. rostratus but the illustrations accompany- ing the brief description are completely worthless and erroneous. The caudal rami are exceptionally long for this genus, the PS exopod has only 4 elements, and the antennule is 8-segmented. The speci- mens reported from Guaratuba (Parana State) in an earlier paper (Carvalho, 1944) are also very small (0.5 mm) and their fragmentary description is equally useless for identification purposes. Finally, there is no possibility of identifying any specimens from Campos-Hernandez & Suarez-Morales’ (1994) illustrations of C. rostrata from the Gulf of Mexico. GENERIC CONCEPTS IN CLYTEMNESTRIDAE 4 iv Vy ovvwo NW o yan vy WV Vi te y yury yy WY, Yoye ¥ Wyvavyy vr vy Fig. 29 Goniopsyllus brasiliensis sp. nov. (@). A, Habitus, dorsal; B, anal somite and caudal rami, dorsal; C, caudal ramus, lateral; D, mandible and maxillule; E, maxilliped, posterior; F, maxilliped, distal portion of basis and endopod, anterior. 43 44 Goniopsyllus tenuis (Lubbock, 1860) comb. nov. Clytemnestra tenuis Lubbock, 1860 Lubbock’s (1860) description is very incomplete and based on a single specimen. The antennule was figured as 7-segmented but comparison with other clytemnestrid descriptions indicates that the author had erroneously shown the second segment as subdivided into two distinct segments. The segmentation of the distal half of the antennule conforms with the Goniopsyllus pattern, justifying its placement in this genus. Giesbrecht (1892) regarded C. tenuis as a likely synonym of G. rostratus but in the light of the discovery of several closely related species we regard this course of action premature. Conversely, Marques (1973) listed C. tenuis in the synonymy of C. scutellata. Although Lubbock doubted the sexual maturity of the holotype female this is contradicted by his state- ments that the specimen was ovigerous and that the second and third abdominal somites had almost completely coalesced (this being in conflict with his illustration of a 6-segmented urosome lacking any trace of a genital double-somite). With the scanty information available it is extremely unlikely that C. tenuis will ever be recog- nized; it is ranked here as species inquirenda. Sapphir rostratus Car, 1890 Conspecificity between S. rostratus, described from Trieste (North Adriatic), and G. clausi, recorded from the South Adriatic (this paper), seems conceivable on zoogeographical grounds. The rela- tive lengths of the distal antennulary segments in both sexes and the length of caudal ramus seta II, however, do not agree with those of G. clausi. It is questionable whether these discrepancies are real or reflect observation bias since Car’s (1890) illustrations contain other, more significant errors such as the P5 which is shown with only 3 setae and the P4 which allegedly lacks an outer spine on the distal endopod segment. A final obstacle to conspecificity is the small size of S. rostratus which, based on the dorsal view of the male, measures only 0.58 mm. Rather than proposing a new replace- ment name in anticipation of potential secondary homonymy with the type species, we maintain this species as species inquirenda under its current name. If S. rostratus and G. clausi are conspecific then the former becomes a invalid senior synonym of the latter. Other records Monard’s (1928) description of “C. rostrata’ from Banyuls-sur-Mer contains several inconsistencies such as his illustration of the P5 exopod which shows only 4 setae and his statement that the P2—P4 enp-3 setal pattern is 6-5-5, indicating that he has confounded P2 and P3. The author also claims that the male P5 is modified and the female antennule 7-segmented. The small size (0.65 mm) seems to tule out conspecificity with G. clausi. Chen et al.’s (1974) record of G. rostratus from the East China Sea and Mori’s (1937) from Japanese waters are indeterminable on the basis of the few illustrations provided. The short female P5 suggests a species different from G. rostratus. Similarly, Marques (1958) did not give convincing evidence for her record from Angola since only the habitus of the male and body length measurements (2 : 0.4- 0.94 mm; d: 1 mm) were provided. DISCUSSION Generic concepts and species discrimination The generic concepts of Goniopsyllus and Clytemnestra (as R. HUYS AND S. CONROY-DALTON Goniopelte) introduced by Claus (1891b), but dismissed by subse- quent authors, are reinstated here. Claus based the distinction on differences in antennule segmentation and setation of the antennary exopod, and on the presence or absence of sexual dimorphism in the caudal rami. Goniopsyllus is clearly more advanced than Clytemnestra, being illustrated by several reductions in the cephalic appendages, P1 and male P6 which provide additional discrepancies between both genera. In Goniopsyllus the number of distal setae on the antennary endopod is reduced (the missing elements being marked by rudiments; arrowed in Fig. 21B), the armature of the maxillule is represented by a single apical element, the distal syncoxal endite of the maxilla bears only 2 elements and the long syncoxal seta representing the proximal endite is lost. The latter character should be used with caution in generic discrimination since convergent loss of the proximal endite has happened in at least one representative of Clytemnestra (Fig. 16E). All species of Goniopsyllus lack the outer basal seta of P1 and have lost the inner seta of its exopod. The male sixth legs are weakly developed bearing only 1 seta in Goniopsyllus (Fig. 11C) but are produced into con- spicuous, elongate, trisetose processes in Clytemnestra (Fig. 11A—B), resembling the condition found in the Aegisthidae and Cerviniidae. Although Clytemnestra is the more primitive genus, it can be readily identified by the absence of the outer spine on P2 exp-1. As far as we could ascertain this is a unique character in harpacticoids with a 3-segmented P2 exopod. The caudal ramus sexual dimor- phism displayed only by Clytemnestra requires further ontogenetic study before it can be considered a potential autapomorphy for the genus. The typical caudal ramus condition found in the majority of the Harpacticoida shows normally developed terminal setae IV and V. In the Clytemnestridae this condition is exhibited only by the males of Clytemnestra (e.g. Fig. 5B), the atypical female state (Fig. 5A) showing reduced setae. In contrast to swimming leg sexual dimorphism which is nearly always the result of deviations in male ontogeny, secondary sexual characters in the caudal rami are exclu- sively expressed by the female, and as a rule are not expressed until the final moult. This timing of expression has been demonstrated in various families displaying caudal ramus sexual dimorphism, in- cluding the Canuellidae, Cylindropsyllidae and Canthocamptidae. In these families it is intrinsically linked with precopulatory mate guarding where female caudal ramus modification shows substan- tial congruence with male antennule morphology. Since the atypical condition in female Clytemnestra is also found in both sexes of Goniopsyllus — and thus unlikely to be the result of transformation at the final moult — a different ontogenetic explanation must apply. This is further corroborated by examination of early copepodids (including Cop V 3d) of C. asetosa and G. clausi which revealed similarly reduced caudal setae in both species. The male caudal setae in Clytemnestra must therefore undergo transformation at the final moult. Hence, it is assumed here that reduction of setae [V—V represents the ancestral state in the family and that elongation evolved only secondarily in male Clytemnestra, not being linked to mate guarding but possibly enhancing its capacity during mate location. Examination of the genital field has revealed significant differ- ences between both genera. In Goniopsyllus the copulatory pore is located halfway down the genital double-somite in a large circular depression (Fig. 27A) and connects via a strongly chitinized duct with the anteriorly positioned seminal receptacles (Fig. 23C—D). In Clytemnestra the copulatory pore is represented by a posteriorly directed minute slit (arrowed in Fig. 27B), located between the genital apertures far anteriorly on the genital double-somite, and a copulatory duct is hardly differentiated (Fig. SA). The polarity of copulatory pore displacementis difficult to assess, however, outgroup GENERIC CONCEPTS IN CLYTEMNESTRIDAE 45 Table 1 Diagnostic characters of Clytemnestra species [Al = antennule; GDS = genital double-somite; AS = first adominal somite]. Length measure- ments are based on material examined in this paper. scutellata gracilis farrani longipes asetosa size ? (in um) 1121 1309-1562 927-947 2 758-830 size 3 (in pm) 1064 1420-1531 939-945 1211 920 cephalothoracic processes present present absent obsolete absent setal number segment 2 Al 12 12 12 u 10 proximal endite maxilla present present present present absent P2 exp-3 formula 223 223 222 222 222 P3 exp-3 formula 323 323 323 322 322 P4 exp-3 formula 323 323 323 322 322 setal number P5 exopod $/d 6 6 5 5 5 P5 apex 2 vs GDS posterior margin coinciding distad coinciding ? proximad P5 apex d vs AS posterior margin proximad coinciding proximad coinciding proximad - spinules 2nd abdominal somite 2 absent present present ? absent _ spinules 1st abdominal somite 3 absent absent absent present absent comparison with the Tegastidae, Peltidiidae and Tisbidae suggests that migration happened anteriorly and the condition in Clytemnestra is apomorphic. Species discrimination in Clytemnestra is most easily achieved by comparing primarily cephalothorax shape, swimming leg spine pattern, urosomal ornamentation and setation of the maxillae and antennules (Table I). Conversely, identification of Goniopsyllus species is strenuous and largely based on size, maxillipedal orna- mentation and proportional lengths of caudal ramus setae. The reported variability in body size and/or PS setation for both C. scutellata and G. rostratus (e.g. Boxshall, 1979; Huys et al., 1996) is based on erroneous identifications and observational errors. Relationships _ Prior to Claus’ (1891a) study the relationships of the Clytemnestridae | were believed to lie with the planktonic poecilostomatoid families, in particular the Sapphirinidae (Car, 1890). This concept was partly _ based on the superficial similarity in dorsoventrally depressed body shape, laterally displaced fifth legs and the failure to recognise the geniculate antennules in the male (Car, 1890). More significantly, this assignment was based also on the strongly reduced mouthparts and the sexual dimorphism displayed by the maxillipeds, two characters regarded as highly diagnostic for the Poecilostomatoida (Huys & Boxshall, 1991). Sexual dimorphism in the maxillipeds is uncommon in the Harpacticoida. Huys (1988) reviewed the topic, showing that there is clear dimorphism only in the Aegisthidae (as a result of male atrophy), some Tisbidae (e.g. Boxshall, 1979) and deepwater Huntemanniidae (Metahuntemannia, Talpina). Dahms & Schminke (1993) demonstrated that in Tisbe bulbisetosa the male maxilliped is involved in precopulatory mate guarding by holding the female’s caudal setae IV and V prior to spermatophore transfer, the antennules playing only an auxiliary role during this process. We speculate that the modified male maxillipeds in clytemnestrids perform a similar function, the elongate endopodal claw probably being involved in holding the female’s caudal rami or swimming legs. Boxshall & Huys (1998) pointed out that the antennulary chemo- sensory system of C. rostratus (= G. clausi sp. nov.) is secondarily enhanced in both sexes by transformation of three setae into aesthetasc-like elements. The middle and distal of these elements are fused basally to an aesthetasc. This study has revealed this pattern to be diagnostic for all Clytemnestridae and can be consid- ered an apomorphy for the family. Examination of copepodid stages showed these transformed setae to be present from at least copepodid III onwards. Modification of antennulary elements into putative chemosensors is rare in harpacticoid copepods and has thus far only been recorded in some deep-sea species. Gee & Huys (1991) described a densely opaque, bulbous element on the distal antennulary segment in both sexes of the paranannopid Leptotachidia iberica Becker, 1974. The only report of a similar structure is that by Por (1969) who figured a modified bulbiform element on the antennule of Cerviniopsis obtusirostris Brotskaya, 1963 (Cerviniidae) which he called the ‘Brodskaya organ’. The complete lack of swimming leg sexual dimorphism impedes an assessment of the relationships of the Clytemnestridae. The 1- segmented P1 exopod is found in several interstitial Paramesochridae, Leptastacidae and Laophontidae, yet it is diagnostic at the family level only in the Rotundiclipeidae and Tegastidae. Lang (1948) recognised a close relationship between the latter, the Peltididiidae and the Clytemnestridae. He based this affinity solely on P1 mor- phology, including the non-prehensile nature of the endopod and the presence of maximum 5 elements on the distal exopod segment. Within this group of tisbidimorph families he placed the Peltidiidae as the sistergroup of the Clytemnestridae on account of the dorso- ventrally flattened body and the reduction of the PS baseoendopod in the female. The usefulness of Lang’s (1948) characters is limited due to their homoplastic nature, however, there are at least two other features which appear to substantiate a close relationship between these three families. First, the aesthetasc pattern on the male antennule (with an additional aesthetasc on ancestral segment XI) is displayed by all three families. Secondly, the modification of the distal palmar element on the maxillipedal basis into a pad-like sensory element (Fig. 10B) is a unique synapomorphy (see Huys et al. (1996) for examples in Peltidiidae and Tegastidae). A detailed phylogenetic analysis of the Peltidiidae is nevertheless required before its sistergroup relationship with the Clytemnestridae can be substanti- ated. Indeed, an alternative evolutionary scenario could be that the latter represent only a specialized terminal branch of the former. Most species of the peltidiid genus Alteutha Baird are common members of the coastal plankton, performing pronounced diurnal vertical migrations in the water column. This may well be viewed, either ecologically or evolutionary, as a transitionary step towards the holoplanktonic lifestyle exhibited by the Clytemnestridae. ‘Taxonomic Impediment’ and Marine Plankton The present revision has quadrupled the number of species in the family solely by examination of the relatively limited material deposited in the NHM. There is no doubt that this number would 46 have been significantly higher had the geographic coverage been wider. Indicative of this is the discovery of three species of Clytemnestra in a small sample from the Great Barrier Reef. Pre- liminary examination of material from Brazilian waters (Rio Grande do Sul) revealed a similar sympatry for both Clytemnestra and Goniopsyllus. Although the discovery of several closely related species in both genera is noteworthy, it is not unexpected nor exceptional for a marine planktonic taxon. For example, recent taxonomic studies have uncovered several important species com- plexes in the Oncaeidae (Heron, 1977; Heron & Bradford-Grieve, 1995; Bottger-Schnack, 1999). Although this family is morphologi- cally distinctive and arguably the most speciose in the marine plankton, the continuing discovery of pseudo-sibling species and frequent confusion about the validity of rank of its species and morphs tarnish its literature, both taxonomic and ecological. Current research on another planktonic poecilostomatoid genus, Pachos Stebbing, resulted in the recognition of several new but previously misidentified species (Huys & KrSinié, in prep.). The taxonomy of pelagic harpacticoids is plagued by consider- able conservatism and inadequate study of morphological features. With the exception of the mesopelagic tisbid genera (Boxshall, 1979) all planktonic harpacticoids were known well before the turn of the century (Kr@yer, 1846; Dana, 1847, 1849; Boeck, 1865; Brady, 1883; Giesbrecht, 1891; T. Scott, 1894), yet, their morpho- logical definition and supposedly cosmopolitan breadth of their distribution have hitherto remained unchallenged. The genus Microsetella Brady & Robertson currently encompasses only two species, however, one can expect its number of species to increase by an order of magnitude if the many undescribed sibling species are considered (unpubl. data). Similarly, Euterpina acutifrons (Dana, 1847) is commonly regarded as a cosmopolitan species but compari- son of distant ‘populations’ suggests that there is no factual justification for this universally accepted view. In Fleminger & Hulsemann’s (1977) scholarly study demonstrat- ing the taxonomic divergence in three sympatrically occurring sibling species of Calanus in the North Atlantic, one sentence deserves wide currency: *. . ., the quality of knowledge about circulating oceanic habitats and their entrained ecosystems rests upon the reliability of three interrelated sets of information: system- atics of the biota, routine identifications of species, and assessments of their ranges, horizontally and vertically’. Unfortunately, routine identifications in ecological investigations are generally not condu- cive to the recognition of sibling species and all too often wide geographical distributions have been uncritically accepted as the natural consequence of potentially broad oceanic dispersal. The latter perception is often coloured by underlying assumptions of the lack of isolating physical barriers and global uniformity in the open pelagic environment. Pseudo-sibling species can only be readily distinguished once the appropriate characters are considered. Our study demonstrated that for the last 110 years species discrimination in the Clytemnestridae was based exclusively on generic characters, the current recognition of cryptic species being only an artifact of previous ignorance. Hence, there is considerable doubt involved in collating records of the occurrence of these species from the litera- ture to produce distribution maps. Though C. scutellata and G. rostratus have universally been regarded as cosmopolitan, this distributional concept is now no longer tenable and the compilation of distribution records must start from scratch. It would be best to consider earlier records primarily as evidence of the occurrence of the respective genera, a useful attribute considering their virtual absence at latitudes above 60° N and 45° S. Although the geographic location of the collection and/or body size can occasionally be used as indicators of species identity, these R. HUYS AND S. CONROY-DALTON approaches are limited in areas of sympatry where often more sophisticated techniques are required. Like Clytemnestra in the harpacticoids, Calanus is an unusual calanoid genus in that the morphology of the female P5 does not discriminate all of the species (Frost, 1971, 1974). Bucklin et al. (1995) showed however, that despite their exceptional morphological similarity, species of Calanus are quite distinct genetically. They obtained similar results for the genus Metridia, confirming the distinctiveness of M. lucens (Boeck, 1865) and M. pacifica (Brodsky, 1948). Frost (1989) concluded, based on morphological characters other than size, that there are seven species within Pseudocalanus. For some, no absolute mor- phological criterion could be found to distinguish females, however, their validity was inferred from trends in several morphological characters. Sévigny ef al. (1989) used patterns of allozyme variation at the GPI (glucose phosphate isomerase) locus to show that Frost’s (1989) sibling species were genetically isolated from each other. Their results agreed with McLaren etal.’s (1989a—c) studies demon- strating differences in genome size and life cycle characteristics among Pseudocalanus species. Bucklin et al. (1998) showed by DNA sequencing of two mitochondrial genes that the sibling species P. moultoni and P. newmani can be reliably discriminated. Bucklin et al.’s (1996) genetic analysis of DNA sequence variation separated the widespread Nannocalanus minor into two genetically distinct types that may represent the previously described N. m. forma major and N. m. forma minor which differ primarily in size range and geographic distribution. Finally, McKinnon ef al. (1992) demon- strated the presence of three sympatric sibling species of Acartia using allozyme electrophoresis. Molecular analysis of marine planktonic copepods is likely to continue to reveal taxonomically-significant genetic partitioning of species populations, including cryptic species. The application of molecular techniques should not however, be an end in itself. Methods used to discriminate sibling species such as protein electro- phoresis or discriminant function analysis profit significantly from or even require a priori morphological recognition of groups or morphotypes whose distinctiveness can be subsequently tested. In fact, how can one demonstrate the accuracy and resolving power of morphological analysis better than to refer to the thorough revisions by Fleminger (1973) and Fleminger & Hulsemann (1974) who presented most compelling evidence for sibling speciation in marine calanoid copepods long before the deluge of molecular data. 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Boletim da Faculdade de Filosofia e Ciéncias, Universidade de Sao Paulo, Zoologia e Biologia Marinha, N.S. 30: 419-447. Sars, G.O. 1921. Copepoda Supplement. Parts [IX & X. Harpacticoida (concluded), Cyclopoida. An Account of the Crustacea of Norway, with short descriptions and figures of all the species 7: title-page, preface, 93-121, pls. 65-76. Scott, A. 1909. The Copepoda of the Siboga Expedition. Part I. Free-swimming, littoral and semi-parasitic Copepoda. Siboga Expedition Monographs 29a: \-323. Scott, T. 1894. Report on Entomostraca from the Gulf of Guinea, collected by John Rattray, B.Sc. Transactions of the Linnean Society of London, Zoology (2)6: 1-161. Sévigny, J.-M., McLaren, I.A. & Frost, B.W. 1989. Discrimination among and variation within species of Pseudocalanus based on the GPI locus. Marine Biology 102(3): 321-327. Sewell, R.B.S. 1940. Copepoda, Harpacticoida. Scientific Reports of the John Murray Expedition 7(2): 117-382. Suarez Morales, E. & Gasca, R. 1998. Updated checklist of the free-living marine Copepoda (Crustacea) of Mexico. Anales del Instituto de Biologia. Universidad Nacional de México, Serie Zoologia 69(1): 101-119. Thompson, I.C. 1888. Report on the Copepoda collected in Maltese seas by David Bruce, M.B., during 1886—7-8. Proceedings of the Liverpool Biological Society 2: 137-151. Vega-Perez, L.A. 1993. Estudo do zooplancton da regiao de Ubatuba, Estado de Sao Paulo. Publicagao Especial do Instituto Oceanografico 10: 65-84. Vilela, M.H. 1965. Copépodes da Ria de Faro-Olhao. Notas e estudos do Instituto de Biologia Maritima, Lisboa 31: 1-38. . 1968. Copépodes da campanha do N.R.P. «Faial», 1958-1959. Notas e estudos do Instituto de Biologia Maritima, Lisboa 35: 1-55. Wells, J.B.J. 1970. Copepoda. I. Sub-order Harpacticoida. Fiches d’Identification du Zooplancton 133: 1-7. . 1976. Keys to aid in the identification of marine harpacticoid copepods. 215p. Department of Zoology, University of Aberdeen, Aberdeen. Wilson, C.B. 1932. 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(Zool.) 66(1):49—107 Issued 29 June 2000 Basal resolution of laophontid phylogeny and the paraphyly of Esola Edwards RONY HUYS AND WONCHOEL LEE Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD CONTENTS EEA BERRI RLCEL CNM Menece aaa eager «semen e ated aedctan Meee ec rtcece ee dsaa seven sa todat Onis asce~ Avec cuneate stacdatets ca tte cena vacerd suet Cededcact deca dnatasaadsecetpdcuredeccdiotss 50 Materials and Methods ...... epee srcccuans cer eenarere teuessacedacan enone toa te wastes satan eben ust vive terete nts wuts vstucedenusatueve cusecvastercnccvesrseaceussdetsstetes 50 MEMEMC ACM OSES AMAISPECIES CESCHIPIOMS ycsvccov. scxs-soeectnceessecsoctaeciesceusercattest cuasscoversarosetl«uacestvseatchae sceorsussuaviestsivedssatvicectt arses 50 PAMMVAeAO POM tt dae wr Ss COC ODS ewe Aten ccs cco tot ss sachs cc tecacaste se tece auscuaeconesceacteeecedcnsctaavaasdes sassdcenccdstuesstuccecesenenhaeseetetees 50 (Cie JEG af end Ea 6 To ee rece na Bec PPE OE ESSER O SEE De CEE CHT POP REE ECR ECC CEC OSES BEE? SRDELEC EEL EEE CER EL CRO CECE PEEL EEE CeCe 50 BO laMloniercaHdaieGwatdss ISOM ei aitecceceteccescte once cvotstconduececeaaese cesuecuavesuenestetseaacs seadesdsdveatucver edeceasdveadecdiecnesestestenevacs 51 Lat Da CaM GN (oh rears ta gS 1 Be rete eee ee SRS ee eee aR a oe ea ee eee 52 Esolarcalapazoensis Niele: iQ BilMorad Sao viii. cstaes eve eae sececee as ca ere veoe van teeaccwacualiccs Di na dutavaesettees soubeecusestacdsadstsnesucineas 59 TECGUG? CA GPANIRS RYO). SON Cec cep cer ec Be Dao Cae ccc RE cca oe PERERA EEC ere CeCe Ra ace ER EE EERE SO ee eee ae 59 JEONG ALATA VANES OF, 110) Pepe OPEC ROO RPE CR OT PERC PPCORER EE OSPPCR OPER POC ERE LEDC CERES > AEE AE oe SB RARE RES a ne aS 62 ESC) | ND EO WPI CUSTOM pret See gaara See ce te ae acc ts tao en dost vcs avasutcagaeeme caer a caeomen nae i annas sndas aoesacecvescesuens dectadap stuck ster saceecds 63 PASCO) VE TROD TETAS PD ML ON nies Sas ayaa dencasenks sxostescuneneawcdsacuasavedadensalatdassatedcstasacceascarsnsvasdovasesaseanadvacsocdtedvestassissdensecéstevatsciviesnteas 63 Eola longicavdd Bawatds, Soi cers NOOGE (LOSS) .:..c 87 PA CHESOL GE LAINOPIEL SP ILO W:5 needy seretepee setateCa tae rac eCer cscs caus’oteco see scetantetvrneconadeseeshecnsveteedesucesaeecadtsavectevisteasdueensracthontetese ve 87 PESOLGUS MSH ISHACS MIS LET ON QUI GI erseeaeeer cents cebeedere cu sceohsscaasainitasecdaes j) HIS an) = = Se LLL we IS a =: —<—s SS SSS La. Wy oe oF bo SS ————— : ig IG A Ml ges OSs SS = Z Zz =< LEE ———— LE c rior; D, P3 endopod (¢), anterior. rior; C, P4 enp-1 (9), ante rior; B, P4 (9), ante Fig. 4 sola bulbifera (Norman, 1911). A, P3 (9), ante BASAL LAOPHONTID EVOLUTION Sy) SSS WD es IPE ZZ tan Se ry We X SN Wy Li 0\\\ Y Way LL. MULAN \a\5 \ HAN AAR LOS) 41S “4 4M, 4 4 NZ Wa Onis BONES 4 Ve All We: \ ATR a Wes A WN it Ny WM YA hn ih wl THAR AUT a ey, WN Sidi Lhe pe ea ity Asif! wale yy RNS 4 i] \ Wiig PW / LUN. EDN, 7] AYE IE Win, , B ¥ NN \ AIMS TINK DWN SQN ae in i Ly Wi Fig.5 sola bulbifera (Norman, 1911) (3). A, Habitus, dorsal; B, antennule, dorsal (armature of segments 3-6 omitted); C, antennulary segments 3-4, ventral; D, antennulary segment 5, ventral; E, antennulary segments 6-7, ventral; F, left caudal ramus, ventral; G, right PS, anterior; H, left P6, anterior. 58 seta and 2 small processes; with 4 medially directed, spinous processes (Fig. 2H). First postgenital somite with backwardly produced lateral angles, bearing spinular tuft (Fig. 1C); without ventral ornamentation. Penultimate and anal somites distinctly narrower; ventral posterior border with spinules. Anal somite (Figs. 1D; 31A) with spinulose anal operculum. Caudal rami (Figs. 1D; 31A) widely separated; slightly longer than widest portion; proximal half distinctly bulbous with major swelling medially, dorsally and ventrally (Fig. 1C); ventral surface with very large semi-circular concavity (Fig. 21) leading to small tube-pore; dorsolateral surface with minute spinules; seta I small, setae II-III well developed, naked and closely set; setae [V and V pinnate and with fracture planes, seta V 2.5 times as long as seta IV; setae VI-VII naked. Rostrum (Figs 1A; 2A) large, rounded anteriorly; delimited at base by transverse surface suture; with paired sensillae anteriorly and median tube-pore dorsally. Antennule (Fig. 2C) slender, incompletely 7-segmented, with 1 minute (obscured by large distal one) and 2 well developed spinous processes on posterior margin of segment 1, no processes on long segment 2. Segment 1 with short spinules posteriorly between processes and large spinular patch around anterior margin. Armature formula: 1-[1], 2-[4 + 4 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[2], 7- [6 + 1 pinnate + acrothek]. Aesthetasc on segment 4 fused basally to 2 setae. Acrothek consisting of aesthetasc and 2 naked setae; set on apical pedestal. Boundary between segments 6 and 7 only expressed posteriorly. Antenna (Fig. 2D) with elongate exopod bearing 2 lateral and 2 apical pinnate elements, and a longitudinal row of fine spinules. Allobasis with pinnate abexopodal seta and spinular patch opposite exopod. Endopod with lateral armature consisting of | pinnate spine and 2 setae; distal armature consisting of 2 unipinnate spines and 3 geniculate setae (outermost fused basally to small tube-seta). Labrum (Fig. 2B) with spinules around distal margin; anterior face with dense pattern of fine spinules and distal patch of overlap- ping scales. Mandible (Fig. 2E) with short gnathobase and small 1-segmented palp probably representing fused basis and endopod; with 2 lateral (basal) pinnate setae and 3 distal (endopodal) setae (1 pinnate, 2 bare). Paragnaths highly ornate lobes as in Fig. 3F. Maxillule (Fig. 2F) with well developed praecoxal arthrite bear- ing 1 seta on anterior surface and 9 elements around distal margin. Coxal endite with 1 spine and 1 seta, basal endite with 1 spine and 2 setae. Exopod a short segment with 2 distal setae; endopod incorpo- rated into basis, represented by 2 setae. Maxilla (Fig. 2G). Syncoxa with very long spinules around outer margin and dense surface spinulation as figured; with 3 endites; praecoxal endite small, with 1 plumose seta; middle endite drawn out into pinnate claw, with 2 tube-setae; distal endite with 3 ele- ments. Allobasis produced into strong curved claw; accessory armature consisting of 1 spine and | seta; with spinular patch proximal to endopodal setae. Endopod incorporated into allobasis, represented by 2 bare and 2 pinnate setae. Maxilliped (Fig.3E) slender, with elongate basis and endopodal claw. Syncoxa with 2 plumose setae. Basis with spinular ornamen- tation as figured; spinules present along entire palmar margin. Endopod represented by very long, minutely pinnate claw bearing 1 accessory seta and tube-pore at base. P| (Fig. 3A) with dense ornamentation on praecoxa, coxa and basis. Basis with pinnate seta on anterior surface and along outer margin. Exopod 2-segmented, small compared to endopod; exp-1 R. HUYS AND W. LEE } not extending to distal margin of basal pedestal, with pinnate outer spine; exp-2 with 3 pinnate outer setae and 2 geniculate setae apically. Endopod slender; enp-1 about 2.5 times as long as basis, with long setules along inner margin and fine spinules along outer margin; enp-2 about 3 times as long as wide, with slender minutely pinnate claw and small accessory seta (Fig. 3B). P2—P4 (Figs 3C; 4A—B) with 3-segmented exopods and 2-seg- mented endopods. P2 basis with long, bipinnate outer spine; P3—P4 bases with bare outer seta. P2—P3 enp-1 with multipinnate inner seta; P4-enp-1 (Fig. 4C) with basally swollen, minute seta. Outer spine of P2—P4 enp-2 very long and setiform. Tube-pore present near distal outer corner of P3—P4 enp-2. Armature formula as follows: Exopod Endopod P2 0.1.123 1.221 iP) 0.1.223 iL2y2Il [d: 1.1.220] P4 0.1.223 1.221 P5 (Fig. 3D). Endopodal lobe small, extending just beyond insertion sites of proximal outer setae of exopod; with 1 short and 1 long pinnate seta apically, and 2 long widely separated setae along inner margin; tube-pores present near articulation with exopod, between apical setae and proximal to innermost seta. Exopod elon- gate, produced apically into tubular extension bearing 1 bare seta; inner margin with 1, outer margin with 4 pinnate setae; inner seta distinctly longer than apical one. Both baseoendopod and exopod with elaborate ornamentation pattern as figured. MALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 512 um (n=2, range 500-524 um). Maximum width (168 tm) measured at posterior margin of cephalothorax. Body (Fig. 5A) more compact and abbreviated than in 2; covered with similar dense pattern of minute spinules. Pattern of cup-shaped pores as in @ except for paired lateral pores present on genital somite. Cephalothorax wider than free somites; body constricted at level of genital somite. None of urosomites with backwardly pro- duced posterolateral corners. Genital somite with large cup-shaped pores laterally, each partly closed off by fringe of setular extensions (Fig. 5H). Sixth legs represented by well developed opercula, one articulating and clos- ing off left or right genital aperture; each produced into cylindrical process bearing | lateral and 1 apical seta. Antennule (Fig. 5B—E) 7-segmented, subchirocer, with geniculation between segments 5 and 6. Segment 1 with spinules/ setules around anterior margin and 2 spinous processes along poste- rior margin. Segment 2 longest; segment 4 minute, represented by incomplete sclerite. Segment 5 with large proximal process anteriorly, bearing modified bifid spine (Fig. 5D); forming cylindrical process bearing long aesthetasc. Segment 6 with 3 spinous processes along anterior margin. Distal portion of segment 7 elongated, displacing acrothek to position isolated from other armature. Armature for- mula: 1-[1], 2-[4 + 5 pinnate], 3-[6 + 1 pinnate], 4-[2], 5-[7 + 2 pinnate + | bifid spine + (2 + ae)], 6-[1 + 3 processes], 7-[7 + acrothek]. Apical acrothek consisting of aesthetasc and 2 bare setae. P3 endopod (Fig. 4D) 3-segmented; enp-1 as in 9, enp-2 with inner seta and short outer apophysis; enp-3 small, with tube-pore, 2 lateral and 2 apical setae. P5 (Fig. 5G) medially fused, positioned ventrolaterally. Baseoendopod without endopodal lobe; medial margin with 2 spinules and 2 tube-pores; outer basal seta arising from short spinulose pedestal. Exopod free; with 1 apical and 1 inner and 3 outer pinnate setae. BASAL LAOPHONTID EVOLUTION Caudal ramus (Fig. 5F) rectangular, without bulbiform expan- sions; about 1.6 times as long as wide; with medioventral cup-shaped concavity as in 9; ventral ornamentation more elaborate than in 9. REMARKS. Lang (1948) synonymized L. bulbifera with E. longicauda, alluding to the congruence in the female P5 baseoendopod between Gurney’s (1927) description of L. bulbifera and Edwards’ (1891) original description of E. longicauda (i.e. with 4 setae; Norman (1911) figured only 3), and in the number of processes on the first antennulary segment between the males of L. rhodiaca (cf. Brian, 1928a) and E. longicauda (cf. Edwards, 1891) and the female of L. bulbifera (cf. Norman, 1911). He also referred to Willey’s (1935) discovery of L. bulbifera in Bermuda as addi- tional zoogeographical evidence for this course of action. It is clear however that (1) the morphological grounds for this synonymy only prove generic identity and not conspecificity, (2) Willey’s (1935) record is both unreliable and unconfirmed, and (3) Gurney’s (1927) records from the Suez Canal in reality refer to another species E. canalis sp. nov. (see below). Our redescription diverges from Norman’s illustrations in only two aspects: (1) the presence of 4 setae on the baseoendopod of the 2 PS, the innermost being overlooked by Norman as already sus- pected by Lang (1948), and (2) the inner seta of P4 enp-1 which is minute (checked against the holotype) instead of very well devel- oped as figured by Norman. E. bulbifera can be differentiated from its congeners on the basis of the following combination of characters: antennule indistinctly 7-segmented, P1 enp-1 2.5 times as long as basis, P1 enp-2 3 times as long as wide, P2-P4 enp-1 with inner seta (that of P4 minute), outermost seta of 2P5 baseoendopod extending to distal margin of exopod, caudal rami @ distinctly bulbous. E. bulbifera is widely distributed around the British Isles with reliable records from Ireland (Farran, 1913, 1915; Holmes & O’Connor, 1990), the west coast of Scotland (Norman, 1911) and Norfolk (Hamond, 1969). Moore’s (1973) record of E. longicauda from St. Abb’s probably also refers to this species. It has not been reported anywhere else in northwest Europe, however, its syn- onymy with L. rhodiaca Brian, first suspected by Nicholls (1941b) and later confirmed by Lang (1948), has considerably extended its distribution, including the Mediterranean, Gulf of Suez and Western Australia. Nicholls based his conviction on similarities in the antennule, antennary exopod, P1 and P4 and the modified caudal rami although he admitted that the latter were not bulbous in L. rhodiaca. Brian’s (1928a) original description, based on a single male specimen from Rhodes in the Aegean Sea (Brian, 1928a—b), shows very few discrepancies with our material from Ireland and Norfolk. The caudal rami are somewhat longer in the Mediterranean specimen, the inner seta on P4 enp-! is more developed, the antennule shows an additional segment distal to the geniculation and small proportional length differences can be noted in the antennulary segments and P4 endopod. Lang (1948) had already pointed out that Brian had overlooked one of the outer spines on the P2 exopod. We regard these differences insufficient to warrant the reinstatement of L. rhodiaca and tentatively regard it as a junior subjective synonym of E. bulbifera. Nicholls’ (1945) few illustrations of a male from Port Denison in Western Australia which he attributed to L. rhodiaca do not contradict Brian’s description. In the absence of information on the swimming legs (except P3 endopod) and the female this geographically widely separated record cannot be verified abso- lutely. Monard’s (1928) brief description of L. bulbifera from the Banyuls area does not contain the level of detail to either confirm or deny his identification. The setae on the P5 baseoendopod were probably not drawn at their full length even though the outermost one appears to 59 be exceptionally short, his spine formula would infer a 123 pattern on P4 exp-3 and the size of his female specimens (0.8 mm) falls outside our recorded range. Monard (1937) recorded the species a second time from Algers but the specimens were apparently dis- tinctly smaller (0.64 mm). The Croatian records of L. bulbifera from Rovinj and Split in the northern Adriatic (Douwe, 1929; Klie, 1941) could not be con- firmed. It is conceivable that VriSer’s (1984, 1986) records of E. longicauda from the Gulf of Trieste and Petkovski’s (1955) record from Montenegro refer to the same species. Esola galapagoensis Mielke, 1981 grad. nov. Esola longicauda galapagoensis Mielke, 1981 TYPE LOCALITY. Cabo Douglas, Fernandina (Galapagos). Wells & Rao (1987) expressed reluctance about the subspecific rank attributed to the Galapagos population of E. longicauda. Although Mielke (1981) acknowledged the reported variability and cosmopolitanism of the latter to some extent, he considered the differences exhibited by his material sufficient to warrant the recog- nition of a distinct subspecies. Mielke diagnosed E. longicauda galapagoensis on the basis of the following characters: (1) P1 exp- 2 with 4 setae/spines, (2) Pl enp-2 with remarkably short claw, (3) P4 enp-1 without inner seta, and (4) P5 baseoendopod @ with strongly reduced outer apical seta. Additional diagnostic features not mentioned by the author include (1) inner seta of P6 d extremely reduced, (2) outer setae of PS exopod ¢ naked, (3) outer spine of P2— P4 enp-2 remarkably short, and (4) caudal rami very elongate with conspicuous medial swelling in °. Based on this suite of characters we feel it justified to upgrade Mielke’s form to full species rank as E. galapagoensis. The species has thus far been recorded from two localities in the Galapagos archipelago (Mielke, 1981). Esola canalis sp. nov. Laophonte bulbifera Norman, 1911 sensu Gurney (1927) TYPE LOCALITY. Suez Canal, Port Taufiq (Egypt). TYPE MATERIAL. Holotype @ dissected on 10 slides (BMNH 1999.993): paratype 2 in alcohol (BMNH 1999.994); from material originally registered as Laophonte bulbifera (BMNH 1928.4.2.116) collected during the Cambridge Expedition to the Suez Canal in 1924; det. R. Gurney. ETYMOLOGY. The species name refers to the type locality. DESCRIPTION. FEMALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 621 um (n=2; range: 585-658 um). Maxi- mum width (137 pm) measured at posterior margin of cephalothorax. Body as in E. bulbifera; cephalothorax with paired cup-shaped pores both anterodorsally and anteroventrally on either side of rostrum, and with distinct transverse spinule row dorsally about halfway down the cephalothorax length. Genital double-somite (Fig. 6A) wide and dorsoventrally flat- tened; original segmentation marked by bilateral constriction and spinule row arising from transverse surface ridge dorsally and laterally; anterior (= genital) half with large cup-shaped pores laterally; ventral surface without spinular ornamentation. First postgenital somite with backwardly produced lateral angles, bearing spinular tuft (Fig. 6A); without ventral ornamentation. Penultimate and anal somites distinctly narrower; ventral posterior border with spinules (Fig. 6C). Anal somite (Fig. 6B) with spinulose anal operculum; spinules coarser than in E. bulbifera. 60 R. HUYS AND W. LEE \ avin wi iL Vays ut HF ayy ‘wt SNS ANTIK Mop Tree tp wat " S \ \ NN Soin rae fi nh /) \ | yh AN My Wit Up Wily iyi! \ \\ wih ht y Hi Nt NUMAN HINA WA UMN NAT NS NY \\ it ni iH i Whiyat NHK \ MM iW SHAAN Wy, S A N NOVAY Erg age AUWUUNARATUTTANAT LANNY 25 EF Fig. 6 Esola canalis sp. nov. (2). A, Urosome (excluding P5-bearing somite), lateral; B, anal somite and left caudal ramus, dorsal; C, anal somite and caudal rami, ventral; D, antennule, dorsal; E, mandibular palp; F, antennary exopod. LLL L—L£L_S BASAL LAOPHONTID EVOLUTION NINN ~ = =e SS = : SSS Z = Ss Sex Ww = —— » Vv i / tAZAl XZ ‘ : <— = GA Ss Wil le Ss a. Zi LSS = SSE oe ter EY Se SSS SST WS Sa E rior; C, P3 endopod, anterior; D, P4 endopod, anterior; E, P5, anterior v. (9). A, P1, anterior; B, P2 endopod, ante sp. no lis Fig.7 Esola cana maxilliped. 62 R. HUYS AND W. LEE Table 1. Diagnostic characters of Esola species [species A = Esola spec. sensu Mielke (1997)]. CR = caudal rami, SD = sexual dimorphism. longicauda _ bulbifera galapagoensis canalis profunda vervoorti lobata species A 2 size (um) ? 643-714 330-460 585-658 500-529 510 490-530 470 3 size (um) 550 500-524 300-360 2 2 389-415 380-450 ? cephalothorax dorsal spinule row ? present u present present absent ? ? mandible — endopod ? fused, 3 setae fused,2setae free,3setae free,3setae free,3setae free, 3 setae free, 2 setae? — exopod ? absent absent 1 seta absent absent absent absent — basis ? 2 setae 2 setae 1 seta 2 setae 1 seta 2 setae 2 setae? P1 exp-2 setal number 4 3) 4 5) 5) 3) 3) 4 P1 exp-2 outer apical seta SD eo = = ? ? + ~ y ratio Pl enp-1 : enp-2 claw 0.83 0.50 0.37 0.50 0.55 0.55 0.55 y P2—P3 enp-1| inner seta present present present present present absent present present P3 enp-2 apophysis d ? smooth smooth ? ? dentate smooth ? P4 enp-1 inner seta normal vestigial absent vestigial absent absent absent vestigial P5 benp 2 outer apical seta plumose plumose naked plumose plumose plumose plumose uy — length* short long vestigial very short short short ~short ? CR — pore position ventral medioventral _ ventral mediodorsal ventral medial ventral 2 — medial swelling weak strong strong moderate moderate moderate slight moderate — @length : distal width 3.0 28} 3.4 3.0 DS Del 3.8 ? — 6 length : distal width 2 PhS) 3.4 i ? i157) 37) ? *: very short = not extending to insertion level of middle outer exopodal seta; short = extending to about insertion level of middle outer exopodal seta; long = extending to about apex of exopod [Note that in E. lobata sp. nov. the endopodal lobe is secondarily elongated so that its outer apical seta extends beyond the insertion level of the middle outer exopodal seta despite being short]. Caudal rami (Fig. 6A—C) widely separated; gradually tapering posteriorly and about as long as anal somite; proximal half expanded with major swelling medially and dorsally, and to a lesser extent ventrally (Fig. 6A); large cup-shaped pore located mediodorsally (Fig. 6A) leading to small tube-pore. Armature as in E. bulbifera. Antennule (Fig. 6D) slender, 6-segmented, with 1 large (proxi- mal) and 2 small spinous processes along posterior margin of segment 1. Segment 1 with long spinules around anterior margin. Segments 2 and 3 equally long. Armature formula: 1-[1 pinnate], 2- [7 + 1 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[9 + acrothek]. Aesthetasc on segment 4 fused basally to 2 setae. Acrothek consisting of aesthetasc and 2 naked setae; set on apical pedestal. Antennary exopod (Fig. 6F) elongate exopod bearing 2 lateral and 2 apical pinnate elements, and a longitudinal row of coarse spinules proximally. Labrum with ornamentation as in E. bulbifera. Mandible (Fig. 6E) with small 2-segmented palp; proximal seg- ment with 1 inner (basal) seta and 1 small outer seta representing exopod; endopod a free segment with 3 setae. Maxilliped (Fig.7F). Basis more slender than in E. bulbifera and spinules along outer margin coarser. P1 (Fig. 7A) similar to that of E. bulbifera but basis forming shorter pedestal for endopod, and both exopod (but exp-1 extending to distal margin of basal pedestal) and enp-2 somewhat shorter; exp- 2 with 3 outer setae and 2 geniculate setae apically. P2—P4 (Fig. 7B—D). P2—P3 enp-1 with multipinnate inner seta; P4 enp-1 with vestigial inner seta. Outer spine of P2—P4 enp-2 shorter than in E. bulbifera. Armature formula as follows: Exopod Endopod P2 0.1.123 122 P3 0.1.223 1.321 [d: probably 1.1.220] P4 0.1.223 122A P5 (Fig. 7E). Endopodal lobe small, not extending beyond inser- tion sites of proximal outer setae of exopod; with 2 apical and 2 widely separated inner setae; outer apical seta very short. Exopod more slender than in E. bulbifera; with 1 apical, | inner and 4 outer setae; length of inner (ratio to exopod length 1.15 vs 1.5 in E. bulbifera) and apical seta (ratio to exopod length 1.25 vs 2.2 in E. bulbifera) distinctly shorter. Unknown. REMARKS. Gurney (1927) collected this species from the plankton at Port Taufiq and Le Cap, and in sediment samples from El Ferdane. He attributed his material to L. bulbifera but remarked on some differences with Norman’s (1911) holotype, such as the discrepancy in body size (0.68 mm instead of 0.80 mm), the P1 endopod which is more slender in the Scottish specimen and the presence of an additional seta (the innermost) on the PS baseoendopod. We have found these differences to be of no value in discriminating both species. Norman (1911) clearly overlooked the innermost seta (as indicated by the gap along the medial margin in his figure of the baseoendopod). Also, based on a larger sample of E. bulbifera we found this species on average to be significantly smaller than Nor- man’s observed size of 0.8 mm, approximating the mean length of E. canalis (681 xm vs 621 um; see also Table I). There is no significant difference in the P1 endopod of both species although the proximal segment appears to be longer in E. canalis and the distal segment to be longer in E. bulbifera. Gurney (1927) illustrated the P5, caudal rami and the female habitus in lateral view. His illustration of the caudal rami gives a slightly distorted view in that the rami appear to be much longer than in reality. Por & Marcus (1972) recorded E. longicauda from four localities in the Suez Canal; itis likely that these records and Por’s (1967) previous record from the Gulf of Elat refer to E. canalis. E. canalis is most closely related to E. bulbifera. Females of the former can be differentiated by the conical caudal rami, the medi- odorsal position of the cup-shaped pores on these rami, and the P5 endopodal lobe which is significantly shorter and has a much smaller outer apical seta. Additional differences can be found in the proportional lengths of the proximal antennulary segments, the slenderness of the maxilliped and the size of particular setae on the P2—P4 endopods and PS exopod. E. canalis is the only species of the genus which has retained a vestige of the mandibular exopod. MALE. Esola lobata sp. nov. Esola longicauda (Edwards, 1891) sensu Mielke (1997) TYPE LOCALITY. Bunaken Island near Manado, North Sulawesi (Indonesia); sublittoral sand between seagrass and corals. BASAL LAOPHONTID EVOLUTION ETYMOLOGY. The species name refers to the well developed endopodal lobe of the P5 in both sexes. P2—P4 setal formula: Exopod Endopod P2 0.1.123 1.221 P3 0.1.223 1.321 [d: 1.1.220] P4 0.1.223 0.221 Mielke (1997) provided an excellent description of Sulawesi females and males which he attributed to E. Jongicauda. His illustra- tions show sufficient differences to warrant separate species status. E. lobata is similar to E. profunda from the Mediterranean and both E. vervoorti and E. galapagoensis from the Pacific in the loss of the inner seta on P4 enp-1. The species can, however, be readily distinguished by the long endopodal lobe in the @ PS, a well developed bulbous extension on the baseoendopod of the d PS, the elongate caudal rami which are relatively little modified in the female, and the short P1 endopod. Discrepancies are also noted in the female antennule, particularly in the relative lengths of the proximal segments, and the size and precise position of the spinous processes on segment |. The species is thus far known only from the type locality. Esola profunda sp. nov. TYPELOCALITY. Ligurian Sea (Western Mediterranean; 42°39’ 12" N, 08°39°30" E), northwest of the Bay of Calvi (Corsica); depth 760 m. TYPE MATERIAL. 2 22from type locality. The bottom sample was taken on 10 June 1986 with a small, modified Reineck box corer (170 cm?) by K. Soetaert. The median grain size of the sediment is 4 um and the silt-clay amount averages 78.5%. The CPE value is about 0.59 g/cm? of which 12.6% is represented by chl a. Holotype dissected on 11 slides (BMNH 1999.991), paratype @ preserved in alcohol (BMNH 1999.992). ETYMOLOGY. The species name is derived from the Latin profun- dus (meaning deep) and refers to the bathyal distribution of this species. DESCRIPTION. FEMALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 515 um (n=2; range: 500-529 um). Maxi- mum width (129 um) measured at posterior margin of cephalothorax. Body (Fig. 8A) as in E. bulbifera but constrictions between pedigerous somites less defined; cephalothorax with paired cup- shaped pores both anterodorsally and anteroventrally on either side of rostrum, and with distinct transverse spinule row dorsally about halfway down the cephalothorax length. Urosomites with dense spinulation and irregular pattern of sur- face ridges laterally and dorsally (Fig. 8B). Genital double-somite (Fig. 8A—B) with large cup-shaped pores laterally in anterior half; ventral surface without spinular ornamentation; posterolateral angles slightly produced. First postgenital somite with backwardly pro- duced lateral angles, bearing spinular tuft; without ventral Ormamentation. Penultimate and anal somites distinctly narrower (Fig. 8A); ventral posterior border with spinules (Fig. 8D). Anal somite (Fig. 8C) with spinulose anal operculum. Caudal rami (Fig. 8C—D) widely separated; with slight swelling medially and virtually no expansion ventrally (Fig. 8B); dorsal surface with 2 chitinous processes in posterior half; large cup- 63 shaped pore located ventrally (Fig. 8D) leading to small tube-pore. Armature as in E. bulbifera. Antennule (Fig. 9A) slender, 6-segmented, with 1 large (proxi- mal) and 2 small spinous processes along posterior margin of segment 1.Segment | with long spinules around anterior margin. Segment 2 distinctly longer than segment 3. Armature formula: 1-[1 pinnate], 2-[7 + 1 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[9 + acrothek]. Aesthetasc on segment 4 fused basally to 2 setae (Fig. 9B). Acrothek consisting of aesthetasc and 2 naked setae; set on apical pedestal. Antennary exopod (Fig. 9D) elongate exopod bearing 2 lateral and 2 apical pinnate elements; no ornamentation discernible. Labrum with ornamentation as in E. bulbifera. Mandible (Fig. 9E) with small 2-segmented palp; proximal seg- ment with 2 inner, (basal) setae; endopod a free segment with 3 setae. Maxillule (Fig.10D) as in E. bulbifera but outer apical seta of exopod naked and shorter and distal spine on basis stouter. P1 (Fig. 8E) similar to that of E. bulbifera but both endopodal segments and terminal claw shorter; exp-1 extending to distal margin of basal pedestal; exp-2 with 3 outer setae and 2 geniculate setae apically. P2—P4 (Figs 9C; 10A-—B). Outer basal spine of P2 distinctly shorter and more setiform. P2—P3 enp-1 with multipinnate inner seta; P4 enp-1 inner seta absent. Outer spine of P2—P4 enp-2 shorter than in E. bulbifera. Armature formula: Exopod Endopod P2 0.1.123 1.221 P3 0.1.223 1.321 [d: probably 1.1.220] P4 0.1.223 0.221 P5 (Fig. 10C). Endopodal lobe elongate, clearly extending bey- ond insertion sites of proximal outer setae of exopod; with 2 apical and 2 widely separated inner setae; outer apical seta distinctly shorter. Exopod more slender than in E. bulbifera; with 1 apical, 1 inner and 4 outer setae; anterior proximal seta and distalmost outer seta much shorter. MALE. Unknown. REMARKS. E. profunda is known only from the type locality and represents the deepest record for the genus. It is similar to E. lobata in the elongate endopodal lobe of the 2 P5, the mandibular palp setation, the ventral position of the caudal ramus pores and the absence of the inner seta on P4 enp-1. It differs from this species in the elongate 2 PS exopod, caudal ramus shape (presence of dorsal chitinous processes) and the longer P1 enp-2. Esola vervoorti sp. nov. Esola longicauda (Edwards, 1891) sensu Vervoort (1964) TYPELOCALITY. Ifaluk Atoll, Caroline Islands, North Pacific; stn 592 (Vervoort, 1964). TYPE MATERIAL. National Museum of Natural History, Washing- ton, D.C.: holotype 2 dissected on 12 slides (NMNH 109702); paratypes are 2 dd in alcohol (NMNH 288048). Originally labelled E. longi-cauda; det. W. Vervoort; 16 October 1953. Two other vials with identical labels contained different species: (a) NMNH 109789: cope-podid V 2 of E. longicauda var. sensu Vervoort (1964), from stn 591; (b) NMNH 109790: 1 2and 1 3 of Paralaophonte sp., from stn 590. ETYMOLOGY. The species is named in honour of Dr Willem Vervoort (Rijksmuseum van Natuurlijke Historie, Leiden) who first illustrated this species. 64 R. HUYS AND W. LEE Dl \\ Ki ay PH i) PAN “Py, 7 a bOP~ Gee Fe Wy) \ AIAN hl WANN WN RNY WAN ASA ip ATA RN \y Y “yy Al Yu TANI \ HAIN Ws ine i fy Wi PONTOON i Uj fi) < Ye ra Wh , be Mini iil fit ¥ ey adil Way Ni & PPI my Wi i : Yj / “ Fe natn RGN Wi Miypiy i ; i mV ADM hi ins NN Wi Ny Nh i ipa parry ral ANY KA NN Wi Hh fc NN \ ANN AY we Hi i ue i: ‘ Dea Hy PN ARR Mh iy) Wy Wi) Uy) Ti) 1 Wp svi ANNE i oe nah ANNU I : iumaaieras iN VAN deen i RUAN , HN oe ‘ oo TN i Ne ; wiht nil \WN SANA hla NN AA i Ai ; Mh eh HWA AANA \y AY AY ; yey ti Ss Wily) (A \ = i Wi HY, fi ATH a EN \ SS Wy LRN Nx! * a ae ae Mii in \ ae ie ‘ (a At Ans mya AK é Upp ' t AVL YAY we AS EN se A b \ AIAN NK ne fh NY \ v4 2 iN ANAS X ARN ; AN SY : wd ‘XS Ns I\N f I) i \Y iy} My 1) AN NYS a ih Ny : « ¢ ZY ES a a oo GD bal nN HANA \ ; oy ah \\ Ny \ ; — ~ NN \ — x a es LiL dyed Ul . Wy v. (9). A, P2, anterior; B, P3, anterior; C, P5, anterior; D, maxillule, anterior. Fig. 10 Esola profunda sp. no BASAL LAOPHONTID EVOLUTION DESCRIPTION. FEMALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 510 um; maximum width 120 um (Vervoort, 1964). MALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 401 um (n=3; range: 389-415 um). Maxi- mum width (121 um) measured at posterior margin of cephalothorax. Body (Fig. 11A) more compact and abbreviated than in 9; covered with similar dense pattern of minute spinules. Cephalothorax with small cup-shaped pores anterodorsally and anteroventrally on either side of rostrum; wider than free somites; without transverse spinule row dorsally. Urosome distinctly narrower than prosome; none of urosomites with backwardly produced posterolateral corners. Genital somite with ventrolateral cup-shaped pores (Fig. 11B-—C). Sixth legs (Fig. 11B—C) represented by well developed opercula, one articulating and closing off left or right genital aperture; each produced into cylindrical process bearing | lateral and | apical seta. Antennule (Fig. 12A—D) 7-segmented, subchirocer, with geniculation between segments 5 and 6. Segment 1 with spinules/ setules around anterior margin and 2 spinous processes along poste- rior margin. Segment 4 minute, represented by incomplete sclerite. Segment 5 longest, with large proximal process anteriorly, bearing modified spine; forming cylindrical process bearing long aesthetasc fused basally to 2 setae (Fig. 12C). Segment 6 with 3 spinous processes along anterior margin. Segment 7 triangular. Armature formula: 1-[1 pinnate], 2-[7 + 2 pinnate], 3-[6], 4-[2], 5-[7 + 2 pinnate + | spine + (2 + ae)], 6-[1 + 3 processes], 7-[7 + acrothek]. Apical acrothek consisting of aesthetasc and 2 bare setae. Mandibular palp (Fig. 11F) small, comprising elongate basis with 1 pinnate seta and free endopod bearing 3 apical setae. Pl (Fig. 12E) with broader basal pedestal and more robust endopod than in E. bulbifera; enp-1 stouter and enp-2 slightly shorter. Exopod small; exp-1 not extending to distal margin of basal pedestal, with stout outer spine; exp-2 with 3 outer setae and 2 apical setae, outer apical seta much shorter than in 2 and not geniculate. P2—P4 without inner seta on enp-1 (Fig. 12F—H). P3 endopod (Fig. 12G) 3-segmented; enp-1 as in 2; enp-2 with inner seta and dentate outer apophysis; enp-3 small, with tube-pore, 2 lateral and 2 apical setae. Armature formula: Exopod Endopod P2 0.1.123 0.221 P3 0.1.223 0.1.220 [ 2: 0.321] P4 0.1.223 0.221 P5 (Fig. 11E) medially fused, positioned ventrolaterally. Baseoendopod without endopodal lobe; medial margin with 2 tube- pores; outer basal seta arising from short spinulose pedestal. Exopod free; with 1 inner seta and 1 apical plus 3 outer pinnate spines; spines markedly shorter than in E. bulbifera. Caudal ramus (Fig. 11B—D) rectangular, without bulbiform ex- pansions; about 1.7 times as long as wide; with medial cup-shaped concavity as in 2. REMARKS. Vervoort (1964) inclined to assign specific status to his material from the Ifaluk Atoll, however, refrained from doing so due _ to the uncertainty about the widely recorded variability for E. longicauda. E. vervoorti occupies an isolated position in the genus for a number of reasons: (1) the absence of the inner seta on P2—P4 enp-1, (2) the dentate type of apophysis on the male P3 endopod, (3) absence of transverse spinular row on cephalothorax, (4) reduced 67 mandibular palp, (5) very short 2 caudal rami, and (6) the sexual dimorphism of the outer apical seta on P1 exp-2. The latter character is unique within the Laophontidae; Vervoort (1964) also illustrated this sexual dimorphism but did not mention it as a feature of high significance. Esola longicauda Edwards, 1891 sensu Noodt (1955) Noodt (1955) illustrated a single ovigerous female of E. Jongicauda recorded from the Sea of Marmara. His specimen is much larger (0.79 mm) than any other species in the genus (Table I) and like Edwards’ (1891) types shows a strongly developed seta on P4 enp- 1. It resembles E. bulbifera in the bulbiform caudal rami and the incompletely 7-segmented antennule which according to Noodt (1955) displays a partly subdivided apical segment. His statement that the endopodal lobe has only 3 setae is clearly based on an error. Without further information the identity of this specimen cannot be determined. Esola longicauda Edwards, 1891 var. sensu Vervoort (1964) This variety, known from a single male, differs from Vervoort’s (1964) typical specimens of E. longicauda (here designated as E. vervoorti sp. nov.) in the slender and almost haplocer antennule, the presence of an inner seta on P2—P4 enp-1 and the shorter P4 endopod and P5 exopod. This combination of characters rules out conspecificity with both E. vervoorti and E. lobata, the only estab- lished species from the Western Pacific. It also differs from Mielke’s (1997) Esola spec. from Sulawesi by the presence of 5 setae on the distal exopod segment of P1. It is conceivable that this variety represents yet another species, however, the discovery of the female is crucial before it can be attributed such status. Esola longicauda Edwards, 1891 sensu Wells & Rao (1986) Wells & Rao’s (1986) record of a single female from Havelock Island (South Andaman) is virtually indeterminable. It is probably conspecific with Sewell’s (1940) specimens of Laophonte bulbifera recorded from Nankauri Harbour in the Nicobar Islands and Addu Atoll in the Maldive Archipelago. Both share the absence of the inner seta on P4 enp-1 and their antennulary segments have similar proportional lengths. Esola spec. sensu Mielke (1997) Mielke (1997) provided figures and additional information of a single female which is potentially sympatric with E. lobata in North Sulawesi. This form differs from the latter in the size of the proc- esses on the first antennule segment, the shape and setal length of the antennary exopod, mandibular armature, P1 exp-2 setation, pres- ence of a vestigial seta on P4 enp-1 and caudal ramus shape. The presence of only 4 setae on the distal exopod segment of P1 relates it to E. galapagoensis and E. longicauda, however, differences in the antennules and P4 endopod make conspecificity unlikely. Genus Mourephonte Jakobi, 1953 Moerephonte Jakobi, 1953: lapsus calami by Vervoort (1964). Jakobi (1953) established this genus to accommodate a new species M. catharinensis described from the coast of Santa Catarina, Brazil. Vervoort (1964) expressed severe doubts as to the validity of this genus, assuming that the completely reduced P2 endopod and the 68 R. HUYS AND W. LEE L atonunuineiinnmprini NU, we { / @ Se SS LALA ASS Mii i Ani AW Joop rag tient | my Wy ont WN | WS Mil ‘ i f Winall \\ Why tyy ALA hy Me Ahh hit et SN 1 1 Ny Uy —— SSS SS - 7 Chia ie) beth OREN RH yh prey ay SY ww HH HEHEHE RRR = = aS. == =~ FS SS BCD aS | 4 | i rh hint i Minh i\ AM i Arlt HAA betes = SS D t ,, N 4 " , Fig. 11 Esola vervoorti sp. noy. (3). A, Habitus, dorsal; B, urosome (excluding P5-bearing somite), ventral; C, same, lateral; D, anal somite and caudal rami, dorsal; E, left PS, anterior; F, mandibular palp. 69 BASAL LAOPHONTID EVOLUTION Fig.12 Esola vervoorti sp. nov. (3). A, Antennule, ventral (armature of segments 2—5 and 7 omitted); B, antennulary segments 2-4; C, antennulary segment 5, ventral; D, antennulary segment 7; E, P1, anterior; F, P2 endopod, anterior; G, P3, exopod, anterior; H, P4, endopod, anterior. 70 aberrant setal formula most likely resulted from imperfect dissec- tion. In addition, he suspected that Mourephonte was a junior subjective synonym of Esola and claimed that M. catharinensis was probably nothing more than an inadequately illustrated specimen of Esola longicauda. Lang (1965) pointed out that Jakobi’s species had already been described as Laophonte longiseta by Nicholls (1941a) and regarded the absence of the P2 endopod and the reduced armature of P2—P4 as sufficient grounds to maintain Mourephonte as a distinct genus. Jakobi’s material, consisting of an unspecified number of males collected from the tidal zone at Itapocoroy and Porto Belo, is no longer extant, the only specimen available being Nicholls’ holotype male of L. longiseta deposited in the South Australian Museum, Adelaide. This specimen forms the basis of the redescription given below. The female is as yet unknown. DIAGNOSIS (based on 6 only). Laophontidae. Body cylindrical. Integument of cephalothorax and body somites with dense pattern of spinules and setules. Rostrum large, partly delimited at base. Cup- shaped pores present both anterodorsally and anteroventrally on cephalic shield, laterodorsally on caudal rami; absent on genital somite. Anal operculum dentate. Caudal rami rectangular, short. Sexual dimorphism in antennule, P3 endopod, P5, P6, genital segmentation and caudal rami. Antennules slender; haplocer and 7-segmented in d; segment 1 with 2 small processes along posterior margin; swollen segment 5 with very long aesthetasc (fused basally to 2 setae) but without distinct anterior outgrowth. Antenna with 4 setae on exopod; allobasis with abexopodal seta. Labrum with marginal spinules distally. Man- dible with small 1-segmented palp bearing 1 lateral and 3 apical setae. Maxillule without defined exopod, represented by | setae. Maxilla with 3 endites on syncoxa; endopod represented by 4 setae. Maxilliped slender; syncoxa with 2 setae; entire palmar margin with spinules; endopodal claw elongate. P1 very large compared to other legs; with 2-segmented exopod bearing 4—5 setae on exp-2 and elongate endopod; enp-1 without inner seta, enp-2 with minute seta and long, slender claw. P2—P4 with 3-segmented exopods; endopods entirely absent (P2) or 2- segmented (P3—P4). Bases with plumose (P2) or naked (P3—P4) short outer seta. P2—P4 without inner setae on exp-2 and -3. P4 enp- 2 with widely separated apical setae. P3 endopod ¢ indistinctly 3-segmented with incomplete surface suture between enp-2 and -3; enp-2 with inner seta and short outer, spinous apophysis. Armature formula as follows: Exopod Endopod P2 0.0.022 — P3 0.0.022 1.1.110 [in 2 presumably 1.211] P4 0.0.022 0.111 P5 6 without endopodal lobe; exopod short, with 1 inner, 2 apical and 2 outer setae/spines. P6 asymmetrical; membranous flaps with 2 setae arising from cylindrical process. TYPE AND ONLY SPECIES. Laophonte longiseta Nicholls, 1941a = Mourephonte longiseta (Nicholls, 1941a) Mourephonte longiseta (Nicholls, 1941a) Laophonte longiseta Nicholls, 1941a Mourephonte catharinensis Jakobi, 1953 TYPE LOCALITY. ‘Tidal zone at Itapocoroy and Pérto Belo, Santa R. HUYS AND W. LEE Catarina State, Brazil; holdfasts of Endocladia and Codium. MATERIAL EXAMINED. South Australian Museum, Adelaide: Holotype 3 of Laophonte longiseta, dissected on slide Tc 13437 (SAM C5550); Sellick Beach, south of Port Willunga, South Aus- tralia; coll. H.M. Hale, 31 January 1937, froma stone in 1.5 mat low tide on south edge of reef. Jakobi’s (1953) type material of M. catharinensis is lost. REDESCRIPTION. Unknown. MALE. Body length 0.25 (Jakobi, 1953) to 0.30 mm (Nicholls, 1941a). Cephalic shield with paired cup-shaped pores both anterodorsally and anteroventrally on either side of rostrum. Antennule (Fig. 14A—F) 7-segmented, haplocer; geniculation between segments 5 and 6; proximal segments without conspicuous spinous processes but segment | with 2 small protuberances; seg- ment | with spinular row distally and tiny spinules along anterior margin; segment 2 longest; segment 5 with very long aesthetasc (150 um). Armature formula: 1-[1], 2-[8 + 1 pinnate], 3-[6], 4-[2], 5- [8 + 1 pinnate + 1 spine + (2 + ae)], 6-[1 + modified seta], 7-[7 + acrothek]. Acrothek consisting of 2 basally fused setae. Antennary exopod (Fig. 13D) with 2 pinnate setae laterally and 2 pinnate spines distally. Labrum with marginal spinules distally; without overlapping scales. Mandibular palp (Fig. 13C)1-segmented, bilobate, rami com- pletely incorporated; with 1 pinnate seta laterally (probably basal in origin) and 3 bare setae distally (representing incorporated endopod). Maxillule and maxilla as in the genus Esola. Maxilliped (Fig. 13B) slender; syncoxa with 3 spinular rows and 2 pinnate setae; basis elongate, with long spinular row on palmar margin and spinular patch on outer margin; endopod represented by tiny setule and very long claw, exceeding length of basis. Pl (Fig. 13A) large compared to P2—P4; protopodal segments with rows and patches of fine spinules as illustrated; basis with outer spine near joint with coxa and inner pinnate spine on anterior surface. Exopod 2-segmented, exp-1 with pinnate outer spine; exp-2 with 2 spines and 3 geniculate setae. Endopod very long; enp-1 without inner seta; enp-2 with 3 spinular rows, | setule and long, denticulate claw. P2—P4 (Fig. 14G-I) with 3-segmented exopods; endopod 2- segmented (P3—P4) or entirely absent (P2). P3 enp-2 partly subdivided along anterior surface by short transverse suture; apo- physis on outer margin short and slightly sigmoid, bare. P4 enp-2 with apical setae widely separated and flanking secretory tube-pore. Armature formula of P2—P4 as for genus. P5 (Fig. 14J) with baseoendopod fused to somite, endopodal lobe not developed. Exopod rectangular; with 2 outer, 1 apical and 2 inner setae. P6 asymmetrical, produced into cylindrical process at outer corner, bearing long apical and shorter inner seta. Pleural areas of genital somite without modified pores. Posterior margins of abdominal somites with row of long spinules (Fig. 13F). Anal operculum dentate (Fig. 13E). Caudal rami (Fig. 13E—F) short, about 1.3 times as long as wide; with 6 setae (seta I absent), seta VII tri-articulate at base and plumose, setae [TV and V well developed and fused at base. Inner proximal margin with cup-shaped depression (specialized pore) dorsally, marked by row of tiny spinules set on strongly chitinized margin; cup filled with secretory substance. REMARKS. Neither Jakobi (1953) nor Nicholls (1941a) illustrated cup-shaped pores on the cephalic shield. Vervoort (1964) pointed out that Jakobi had shown an anteriorly directed middorsal spinous FEMALE. BASAL LAOPHONTID EVOLUTION a2 AS | i , as i} /} Ny i Ip y bin h EN NANNY Hypa ram r Dili ya PPD Nyy p yqyy yyy AAR Ic i i _"_ SLLSLA TTI F 959 EE ~ | f, wi a & Be oe < ZEAL) LLL | anal somite and right caudal ramus, dorsal; F, posterior urosomites and right caudal ramus, ventral. Hm me ey Ge uy BZ Ne Vy | yy cs 7: = SE S — oS = \\\ =, ae ME, —— 5 a Vai | oA ees Fig. 13 Mourephonte longiseta (Nicholls, 1941a) (3). A, P1, anterior; B, maxilliped; C, mandibular palp; D, coxa, allobasis and exopod of antenna; E, 72 S555 = Se Ze Lkllezs a eee SS PVS=s SS SSS LES ee 2 SoS Le ) SS SSS Ss aS = | = Co —7 largely omitted); B—F, antennulary segments 3-7; Fig. 14 Mourephonte longiseta (Nicholls, 1941a) (3). A, Antennule, ventral (armature of segments 3 G, P2, anterior; H, P3, posterior; I, P4, anterior (apical tube-pore arrowed); J, right P5, anterior. BASAL LAOPHONTID EVOLUTION process which was not illustrated in the lateral habitus view, possi- bly indicating that the author had indeed observed but incorrectly figured the cephalic pores. We have re-examined Nicholls’ slide material and found remnants of the cephalic shield, confirming the presence of both anterodorsal and anteroventral pores as in Esola. Scrutinous observation failed to reveal any such structures on the genital somite. With only two records known, M. longiseta appears to display a remarkably disjunct distribution. Topotype material from Brazil is required to confirm whether the morphometric discrepancies in Jakobi’s description result from imperfect observation or reflect species level differences between the Brazilian and Australian popu- lations. } Genus Archilaophonte Willen, 1995 DIAGNOSIS. Laophontidae. Body elongate and slender; cephalo- thorax slightly wider than rest of body; posterolateral corners of 2 genital double-somite and second abdominal somites not laterally or backwardly produced. Integument of cephalothorax and body somites with dense pattern of spinules and setules; cup-shaped pores on cephalothorax, genital (double-)somite and caudal rami absent. Rostrum very large, partly delimited at base. Integumental cup- shaped pores absent. Anal operculum spinulose. Caudal rami very long, cylindrical with posterior halves diverging. Sexual dimorphism in body shape, antennule, P3 endopod, P5, P6 and in genital segmentation. Antennules short; 6-segmented in 2, subchirocer and 7-seg- mented in d;posterior margin of segment 1 with small blunt process, that of segment 2 with distinct spinous process; with aesthetasc on segment 4 () or 5 (6) and as part of apical acrothek on distal segment; segment 6 of ¢ not particularly modified; proximal aesthetasc fused to 1 seta. Antenna with 4 setae on exopod; allobasis with abexopodal seta. Mandible with biramous palp bearing discrete 1-segmented rami; basis with | lateral seta, exopod with 1, endopod with 3 apical setae. Maxillule with seta at base of exopod. Maxilla with 3 endites on syncoxa; endopod represented by 4 setae. Maxilliped moderately slender; with 3 setae on syncoxa; endopodal claw long and slender. Pl with 2-segmented exopod bearing 5 setae on exp-2 and elongate endopod; enp-1! with inner seta, enp-2 with minute seta and long, slender claw. P2—P4 with 3-segmented exopods and 2-seg- mented endopods. P2 basis with normally developed outer spine. Outer spine of P4 enp-2 not very long. P3 endopod d 3-segmented; enp-2 with inner seta and very long, slender, sigmoid apophysis. P3 exopod d weakly modified with exp-3 being shorter than in 9 Armature formula as follows: Exopod Endopod P2 0.1.123 Pl P3 0.1.223 1.321 [d: 1.1.220] P4 0.1.223 1.221 P5 2with separate rami; exopod large and elongate, with 6 setae/ spines; baseoendopod well developed, with 5 setae/spines. PS 3 with trapezoid endopodal lobe bearing 2 long setae; exopod rectan- gular, with 1 inner, 1 outer and 2 apical setae/spines. P6 2forming opercula closing off paired genital apertures; with 2 ‘long setae. P6 d asymmetrical; membranous flaps with | tiny seta. TYPE AND ONLY SPECIES. [by monotypy]. Archilaophonte maxima Willen, 1995 73 TYPE LOCALITY. 72°52.3'S, 19°34.7' W, Weddell Sea, Antarctic; 495 m depth. REMARKS. Willen (1995) described A. maxima in great detail; the slight sexual dimorphism illustrated for the P3 exopod was not mentioned in the text. The species is known from two localities in the Weddell Sea. Genus Applanola gen. nov. DIAGNOSIS. Laophontidae. Body strongly depressed and com- paratively short; cephalothorax much wider than rest of body; posterolateral corners of ¢ genital double-somite and second ab- dominal somites laterally and backwardly produced. Integument of cephalothorax and body somites with dense pattern of spinules and setules. Rostrum very large, partly delimited at base. Four pairs of integumental cup-shaped pores present: anterodorsally on cephalothorax, near ventrolateral margins of cephalic shield, later- ally on genital (3) or genital double-somite (9) and ventrally on caudal rami. Anal operculum spinulose. Caudal rami short, squar- ish. Sexual dimorphism in body shape, antennule, P2—P4 exopods, P3 endopod, P5, P6 and in genital segmentation. Antennules short; 6-segmented in 9, subchirocer and 7-seg- mented in 3; segments 1—2 without distinct processes; with aesthetasc on segment 4 () or 5 (d) and as part of apical acrothek on distal segment; segment 6 of d with large bilobate outgrowth dorsally; proximal aesthetasc fused basally to 2 setae. Antenna with 4 setae on exopod; allobasis with abexopodal seta. Labrum with distal patch of long spinules. Mandible with elongate 1-segmented palp with 1 lateral and 3 apical setae. Maxillule with elongate defined exopod. Maxilla with 3 endites on syncoxa; endopod represented by 4 setae. Maxilliped large and robust; with 2 setae on syncoxa; endopodal claw relatively short. P1 with 2-segmented exopod bearing 5 setae on exp-2 and robust endopod; enp-1 without inner seta, enp-2 with minute seta and short, strongly curved claw. P2—P4 with 3-segmented exopods and 2- segmented endopods. P2 basis with very long outer spine. Outer spine of P4 enp-2 very long. P3 endopod d 3-segmented; enp-2 with inner seta and outer dentate apophysis. P3 exopod ¢ strongly developed with modified outer and distal spines on exp-3; exopods of P2 and P4 similar in size to 2 but with stronger ornamentation on outer spines. Armature formula as follows: Exopod Endopod P2 0.1.123 1.220 PS 0.1.223 1.321 [d: 1.1.220] P4 0.1.223 1.221 P5 ¢ with separate rami; exopod elongate, with 6 setae/spines; baseoendopod slightly developed, with 4 setae/spines. P5 d without endopodal lobe; exopod short, with 1 inner, 2 apical and 2 outer setae/spines. P6 ¢ forming opercula closing off paired genital apertures; with one seta and 2 small processes at outer corner. P6 ¢ asymmetrical; membranous flaps without armature. TYPE AND ONLY SPECIES. Laophonte hirsuta Thompson & A. Scott, 1903 = Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. ETYMOLOGY. The generic name is derived from the Latin ad (to) and planatus (flattened), and alludes to the dorsoventrally depressed body. Gender: feminine. 74 Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. Laophonte hirsuta Thompson & A. Scott, 1903 Esola hirsuta (Thompson & A. Scott, 1903): Lang (1948) Thompson & A. Scott (1903) described Laophonte hirsuta from washings of pearl oysters and other unidentified invertebrates dredged in the Gulf of Manaar, Sri Lanka. A. Scott (1909) reported the species from 1595 m in the Banda Sea (Indonesia) but this record is almost certainly the result of contamination by ashallow water sample (Lang, 1948; Lee & Huys, 1999). The unknown male was described by Gurney (1927) from Port Taufiq in the Suez Canal. Por’s (19645) records from Haifa Bay and off the coast of Caesarea are likely the result of Lessepsian migration. Both Krishnaswamy (1957) and Krishna Murty (1983) reported the species from the Bay of Bengal. Krishnaswamy collected adults and developmental stages from sponges taken off the Krusadai Islands. Krishna Murty reported some occasional specimens in algal washings from the Visakhapatnam coast. The only other record outside the Indo-Pacific is that by Pesta (1916) from Sao Tomé in the Gulf of Guinea. Lang (1944, 1948) placed the species in the Jongicauda-group of Esola. TYPE LOCALITY. Muttuvaratu, Sri Lanka; washings of pearl oys- ters and other dredged invertebrates. MATERIAL EXAMINED. Cambridge Suez Canal Expedition 1924; Port Taufiq (Egypt): 1 2 dissected on 14 slides (BMNH 1999.982), 1 d dissected on 11 slides (BMNH 1999.983); 3 29(1 damaged), 2 36 and | copepodid V 6 in alcohol (BMNH 1928.4.2.111). REDESCRIPTION. FEMALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 664 um (n=3; range: 650-690 um). Maxi- mum width (281 wm) measured at posterior margin of cephalothorax. Body very dorsoventrally depressed, covered with dense pattern of minute spinules dorsally (Fig. 30A). Cephalothorax much wider than free somites, posterolateral angles backwardly produced; with paired cup-shaped pores both anterodorsally and anteroventrally on either side of rostrum (arrowed in Fig. 15A—B and 30A-B), anterodorsal set partly closed off by fringe of setular extensions. Posterior margin of cephalothorax and all body somites with row of long spinules dorsally and laterally. Ventrolateral areas of cephalic shield and pleurotergites of first two pedigerous somites with long spinules and setules (Fig. 31B; ventral surface with distinct vent- pore at level of mandibles (Fig. 31B). Pleurotergite of P5-bearing somite wide. Genital double-somite (Fig. 17A—B) only slightly narrower than pedigerous somites (Fig. 15A); original segmentation marked by bilateral constriction and dorsal transverse spinule row; anterior (= genital) half with large cup-shaped pores laterally (Fig. 29A), each partly closed off by fringe of setular extensions (Fig. 29B—C); posterior half with backwardly directed lobate extensions bearing spinular tuft (Fig. 29A); ventral surface without spinular ornamen- tation except for spinule row around posterior margin; genital field located near anterior margin. Sixth legs (Fig. 17C) forming well developed opercula closing off paired genital apertures; each with naked seta and 2 small processes at outer corner; inner corner produced into paired, medially directed, spinous processes. Postgenital somites with spinules around ventral hind margin; second abdominal somite with posteriorly directed lateral angles, bearing spinular tuft; penultimate and anal somites distinctly nar- rower. Anal somite with paired oblique spinule rows on ventral surface; anal operculum spinulose. Caudal rami (Fig. 15A, C) widely separated; shorter than wide; R. HUYS AND W. LEE inner margin with medial protrusion; ventral surface with 2 spinule rows and large slit-like pore (arrowed in Figs. 15C; 30C) connected with spacious subsurface duct, extending into anal somite; entrance to pore with fine setules (Fig. 30D); dorsal surface with minute spinules; setae I-III all well developed, naked and closely set; setae IV and V pinnate and with fracture planes, seta V twice as long as seta IV; setae VI-VII naked. Rostrum (Fig. 15A) large, rounded anteriorly; partly delimited at base by transverse surface suture (Fig. 30A); with paired sensillae anteriorly. Antennule (Figs 15A; 16A) short, 6-segmented, without proc- esses on segments 1—2. Segment 1 with dorsal spinular patch. Armature formula: 1-[1 pinnate], 2-[4 + 4 pinnate], 3-[2 + 2 pin- nate], 4-[(2 + ae)], 5-[1], 6-[6 + 3 pinnate + acrothek]. Acrothek consisting of aesthetasc and 2 naked setae; set on apical pedestal. Antenna (Fig. 16B) with well developed exopod bearing 2 lateral and 2 apical pinnate elements. Allobasis with pinnate abexopodal seta accompanied by setular patch. Endopod with lateral armature consisting of 2 spines and | seta; distal armature consisting of 2 unipinnate spines and 3 geniculate setae (outermost shortest and fused basally to setule). Labrum with elaborate ornamentation around distal margin (Fig. 20E) but without spinules or scales on anterior face (Fig. 29D). Mandible (Fig. 16C) with elongate gnathobase and long 1-seg- mented palp (Fig. 31B) probably representing fused basis and endopod; with | lateral and 3 distal pinnate setae. Paragnaths densely hirsute lobes as in Fig. 20D. Maxillule (Fig. 16D) with well developed praecoxa bearing | seta on anterior surface and 8 elements around distal margin. Coxal endite with 2 setae, basal endite with 1 spine and 2 setae. Exopod an elongate segment with 2 distal setae; endopod incorporated into basis, represented by 2 setae. Maxilla (Fig. 20F). Syncoxa with long coarse spinules around outer margin; with 3 endites; praecoxal endite small and unisetose; middle endite drawn out into pinnate claw, with 2 setae; distal endite with 3 elements. Allobasis produced into strong curved claw; acces- sory armature consisting of | spine and 1 seta. Endopod a minute segment with 4 setae of different lengths. Maxilliped (Fig. 17D) compact, with relatively short basis and endopodal claw. Syncoxa with 2 pinnate setae. Basis with spinular ornamentation as figured. Endopod represented by unipinnate claw bearing | accessory seta and tube-pore at base. P1 (Fig. 19E) with narrow coxa and basis. Basis with pinnate seta on anterior surface and along outer margin. Exopod 2-segmented, small compared to endopod; exp-1 with pinnate outer seta; exp-2 with 3 distinctly pinnate outer setae and 2 geniculate setae apically. Endopod robust; enp-1 with long setules along inner margin; enp-2 with short, hook-like, naked claw and small accessory seta. P2-P4 (Figs 17F; 18A, C) with 3-segmented exopods and 2- segmented endopods. P2 basis with very long, multipinnate outer spine; P3—P4 bases with bare outer seta. P2—P4 exp-2 with well developed inner seta. P2—P4 enp-1 small, with inner seta. P2 enp-2 without outer spine; outer spine of P3—P4 enp-2 very long. Tube- pore present near distal outer corner of P3—P4 enp-2. Armature formula as for genus. P5 (Fig. 17E). Endopodal lobe reduced, not extending beyond proximal outer setae of exopod; with 1 short and 1 long pinnate seta apically, and 2 long widely separated setae along inner margin; anterior face with 2 tube-pores. Exopod elongate, produced apically into tubular extension bearing | bare seta; inner margin with 1, outer margin with 4 pinnate setae; inner seta much shorter than apical one. Both baseoendopod and exopod with elaborate ornamentation pat- tern as figured. BASAL LAOPHONTID EVOLUTION 15 OL Eee eee TA see 44444 ss 444A ttd RRR RRR REE SESSA SEARS ASSES ys a 4 Vy ak Y us Fig! \ieayy Winy yy, { bin, PUTT RH wy i, f Wi ee NN tt ¥ Y y wry th ay Lay Ly MM) iat Lise, ay all ha i Wj "4, ae 4 ——- i ip Why vial! ANON e Ma han ii MAR RMR Tce yy We HI vy by \ Vy WYN yypPo any Lane CC EES “Aa ATTA Hh PS Hh a HN . Hike mh [161d Dey pba CT ENS Vihar ys AT naa 7 tay ayt, daly My C LHF: f Wining h AMR : TMA Hy Lippy Ni 7 iE NA\\\ ML PL NUN My , ON WEN 1 TIN ia a) TIWIN Ab 4) h \ N Fae ray a Pe I AA Cros OTT iL Te Uh La ea Kh VA Mitt 1 < ca GALA \ AB /200p FT a, ro) we Ue ere | Fig. 15 Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. (9). A, Habitus, dorsal; B, habitus, lateral; C, left caudal ramus, ventral. [Arrows indicating anteroventral cup-shaped pores in A-B, ventral one in C]. R. HUYS AND W. LEE le Sa ae 7 v. (9). A, Antennule, dorsal; B, antenna; C, mandible; D, maxillule. Fig. 16 Applanola hirsuta (Thompson & A. Scott, 1903) comb. no BASAL LAOPHONTID EVOLUTION 77 Fig.17 Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. A, Urosome 2 (excluding P5-bearing somite), ventral; B, same, lateral; C, left genital aperture 9, ventral; D, maxilliped; E, PS Q anterior; F P2 2 anterior; G, P2 exopod d, anterior. : R. HUYS AND W. LEE |B 5B, rior; D, P4 exopod 4, anterior ; C, P4 9 ante rior; B, P3 d, anterior Fig. 18 Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. A, P3 &, ante rticulated enp-2 and -3, anterior. P3 6, disa 79 BASAL LAOPHONTID EVOLUTION 7 it 7 141 Fig. 19 Applanola hirsuta 1 Mh pth CAN byt ie feisty LAAN Le AN Ae yt me Np 4 ar it f N Wah Wh AG nN AULA RN ! LA TT UPL eae yy tt 1 / if Mate hdin i, ai wnt tt . § A Jy LMU GAVIINS «Ayes Leen G0) PD ADORE RNS Ugh Wes! LAUT NRCS IR RN rae VN LN wily ~ UT ERA Vi Mi H ‘ We Ns, Wt i 7 MEL ad a iit nt yt rail Y PP HSER Ss) iecere! PAU ‘ ANA Ay ayy whiny NN Wye + SNS yay NS Sos * oS S Wiss YES) TiN UT) 7 oli YE: Why TA Ay e DD MAT HEAT oe SRR uly Fete ea nny Yi Hp hn\ re SUN AN SAY ints PAYS VOW OE das PAAR ALOTEH NLRC OR SSS ar TTT TA i Whiley 0 Wy un’ dit i I ( Is Wi YG) 3 TUSTIN MITA NNO CARTS if i "\) 1 (Thompson & A. Scott, 1903) comb. nov. A, Habitus 3, dorsal; B, urosome ¢, ventral; C, PS 6, anterior; D, genital apertures 6, ventral [arrows indicating absence of armature]; E, P1, anterior. 80 R. HUYS AND W. LEE ; Kl cece Fig. 20 Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. A, Antennule 6, dorsal [armature of segments 4—5 omitted]; B, antennulary segments 3-4 of d, dorsal; C, antennulary segment 5 of d, anterior; D, left paragnath; E, labrum, anterior; F, maxilla. BASAL LAOPHONTID EVOLUTION MALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 581 um (n=3, range 570-595 um). Maximum width (248 um) measured at posterior margin of cephalothorax. Length of d copepodid V: 571 um. Body (Fig. 19A) very dorsoventrally depressed, covered with dense pattern of minute spinules as in 2. Pattern of cup-shaped pores as in 2 except for paired lateral pores present on genital somite. Cephalothorax much wider than free somites, posterolateral angles backwardly produced. Posterior margin of cephalothorax and all body somites with row of long spinules dorsally and laterally. Pedigerous somites decreasing in width posteriorly. Urosome (Fig. 19B) slender and narrow; pleurotergite of P5-bearing somite nar- row; posterolateral corners of all urosomites with spinular tuft and posterior margin with spinules all around. Genital somite with large cup-shaped pores laterally, each partly closed off by fringe of setular extensions (Fig. 19D); ventral surface without spinular ornamentation except for spinule row around pos- terior margin. Sixth legs represented by membranous flaps, one articulating and closing off left or right genital aperture; without armature at outer corner. Antennule (Fig. 20A—C) 7-segmented, subchirocer, with geniculation between segments 5 and 6. Segment | with spinules/ setules around anterior margin. Segment 2 with minute knob near dorsal posterior margin. Segment 4 minute, represented by incom- plete sclerite. Segment 5 with spinous outgrowth on anterior margin, probably interlocking with similar processes on segment 6 (Fig. 20C); forming cylindrical process bearing long aesthetasc. Segment 6 with bilobed outgrowth on ventral surface near posterior margin. Distal portion of segment 7 elongate, displacing acrothek to position isolated from other armature. Armature formula: 1-[1 pinnate], 2-[4 +5 pinnate], 3-[7 + 1 pinnate], 4-[2], 5-[7 + 1 pinnate + 1 spine + (2 + ae)], 6-[1 + 2 processes], 7-[7 + acrothek]. Apical acrothek consisting of aesthetasc and 2 bare setae. P2 exopod (Fig. 17G). Outer spines of all segments with much longer pinnules than in &. P3 (Fig. 18B, E). Exopod more robust than in &, slightly bent medially; outer spine of exp-1 with longer pinnules than in 2; middle and distal outer spines and apical spine of exp-3 enlarged, with minute spinules; inner and inner apical setae reduced in length. Endopod 3-segmented; enp-1 larger than in 9, densely setulose along outer margin; enp-2 with inner seta and short outer apophysis bearing small spinous processes along both inner and outer margin (Fig. 18E); enp-3 small, with long tube-pore and 4 setae. P4 exopod (Fig. 18D). Proximal segment slightly more robust than in 2 Outer spines of exp-2 and -3 stubby and somewhat enlarged; spinules typically longer than in &. P5 (Fig. 19C) medially fused (Fig. 19B) positioned ventrolater- ally. Baseoendopod without endopodal lobe; medial margin with setules and tube-pore; outer basal seta arising from short spinulose pedestal. Exopod free; with 3 multipinnate (1 apical, 2 outer) and 2 bipinnate (inner) setae, all well developed. REMARKS. Thompson & A. Scott (1903) illustrated the female P5 with only 3 setae on the baseoendopod, a character included with hesitation by Lang (1948) in the diagnosis of the species. Re- examination revealed that the innermost seta on the endopodal lobe was overlooked. This seta is implanted medially at considerable distance from the others and was also missed by Norman (1911) in his description of Laophonte bulbifera. According to Lang’s (1948) table XXIV the swimming leg armature formula is constant within the Jongicauda-group, including amongst other patterns the pres- ence of the outer spine on P2 enp-2. One cannot but conclude that Lang (1948) must have overlooked Gurney’s (1927) statement that 81 this segment has 2 inner and 2 apical setae. The present redescription has revealed the sexual dimorphism of the exopods of P2 and P4, the presence and pattern of integumental cup-shaped pores, and the detailed morphology of the genital area in both sexes. Krishnaswamy’s (1957) redescription is grossly inadequate and potentially misleading. The ramus labelled *P2 end 2” is the male P3 endopod, his illustration of the female P2 is in fact based on the P3 and the real P2 is figured as the P7(!). In view of these inaccuracies the tabulated setal formula and the author’s remarks on the generic placement of the species are best ignored. Krishnaswamy’s descrip- tion of the first copepodid is of similarly abominable quality. Genus Archesola gen. nov. This genus is proposed to include Esola typhlops and a number of closely related species. It is difficult to understand why Lang (1965) regarded Laophonte lamellipes Nicholls as most closely related to E. typhlops. This doubtful statement was based on the similarity in the long caudal rami and the erroneous fact that males of both species show no modifications on the P3 endopod. Noodt (1955) suggested arelationship with the Laophonte setosa-group but did not elaborate on this view. Re-examination of Nicholls’ (1944) type material (BMNH 1947.10.6.23—27) revealed the true nature of the modified male P2 endopod, confirming close affinity with the genus Paralaophonte Lang. DIAGNOSIS. Laophontidae. Body cylindrical or dorsoventrally depressed; posterolateral corners of 9 genital double-somite and second abdominal somite laterally but not backwardly produced. Integument of cephalothorax and body somites with irregular pat- tern of minute surface lamellae. Rostrum large, partly delimited at base by surface furrow. Integumental cup-shaped pores absent on cephalothorax, genital (double-)somite and caudal rami. Anal oper- culum smooth or bordered with spinules. Caudal rami cylindrical and elongate; not sexually dimorphic. Sexual dimorphism in antennule, P3 endopod, P5, P6 and in genital segmentation. Antennules slender; 7-segmented in 9, haplocer and 7-segmented in 6;segments 1—2 without spinous processes along posterior margin; with aesthetasc on segment 4 ( 9) or 5 (3) andas part of apical acrothek on distal segment; segment 5 d not swollen, without anterior out- growthbut with very long cylindrical pedestal for aesthetasc; proximal aesthetasc fused to 2 setae. Antenna with 4 setae on exopod; allobasis with abexopodal seta. Labrum with distal spinular ornamentation. Mandible with discrete 1-segmented exopod bearing | seta; endopod (3 setae) and basis (2 setae) incompletely fused. Maxillule with minute, defined exopod. Maxilla with 3 endites on syncoxa; endopod represented by 4 setae. Maxilliped slender; syncoxa with 2 setae; palmar margin naked; endopodal claw elongate. Pl with 3-segmented exopod bearing 4 setae on exp-3 and elongate endopod; enp-1 with inner seta, enp-2 with minute seta and strong claw. P2—P4 with 3-segmented exopods and 2-segmented endopods. P2 basis with long outer spine. Outer spine of P2—P4 enp- 2 setiform and very long in P3—P4. P3 endopod ¢ 2-segmented; enp-2 with 3 inner setae and short outer basally fused spine. Arma- ture formula as follows: Exopod Endopod P2 0.1.123 1.221 P3 0.1.223 1.321 [2and 3] P4 0.1.223 1.221 82 P5 ¢ with separate rami; exopod elongate, with 6 setae/spines; baseoendopod slightly developed, with 5 setae/spines. P5 d without endopodal lobe; exopod short, with 2 outer, 1 apical and 2 inner elements (distal inner spiniform). Outer basal seta arising from long, articulating, cylindrical setophore in both sexes. P6 ¢forming opercula closing off paired genital apertures; with 2 small setae at outer corner. P6 ¢ asymmetrical; membranous flaps with 1 apical and 1 lateral seta. TYPESPECIES. Laophonte typhlops Sars, 1908 =Archesola typhlops (Sars, 1908) comb. nov. OTHER SPECIES. Laophonte longiremis TY. Scott, 1905 = A. longiremis (T. Scott, 1905); A. hamondi sp. nov. SPECIES INQUIRENDAE. Esola sp. sensu Chislenko (1967); Esola typhlops pontoica Por, 1959 = A. typhlops pontoica (Por, 1959) comb. nov. ETYMOLOGY. The Greek prefix arche alludes to the primitive position of the genus. Archesola typhlops (Sars, 1908) comb. nov. Laophonte typhlops Sars, 1908 Esola typhlops (Sars, 1908) Lang (1948) TYPE LOCALITY. Flekker6, south coast of Norway, 36 m depth. MATERIAL EXAMINED. (1) West Runton, Norfolk, England: 1 6d dissected on 8 slides (BMNH 1999.1079); collected among Polyclinum and Morchellium under rocks; leg. R. Hamond, September 1971; (2) Frierfjord/Langesundfjord, Norway: 4 damaged 2° (3 in alco- hol: BMNH_ 1999.1081—1083; 1 dissected on 5 slides: BMNH 1999.1080); 99 m, mud, leg. R. Huys, 1985; (3) Gullmar Fjord, Sweden: 1 2 in alcohol (NMNH 90955); 30 m, sand; leg. K. Lang, 08 July 1942. REDESCRIPTION. FEMALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 585 um (n=4; range: 575-592 um). Body cylindrical, not dorsoventrally depressed, covered with dense pattern of minute surface ridges dorsally and laterally. Cephalothorax with almost parallel lateral margins in posterior two- thirds, without paired cup-shaped pores. Posterior margin of cephalothorax and all body somites with row of long setules dorsally and laterally. Posterior margin of urosomites with spinules all around (Fig. 23B); ventrolateral areas of cephalic shield and pleurotergites of pedigerous somites with longer setules. Pleurotergite of P5-bearing somite narrowest. Genital double-somite (Fig. 23A) wide and dorsoventrally flat- tened; original segmentation marked by bilateral constriction and transverse surface ridge dorsally; without cup-shaped pores in ante- rior half; lateral lobes in both anterior and posterior halves with backwardly directed strong spinules; ventral surface without orna- mentation except for spinules around hind margin and 2 pairs of medial tube-pores. Genital field located near anterior margin (Fig. 23A); copulatory pore minute. Sixth legs forming well developed opercula closing off paired genital apertures; each with 2 naked setae. Anal somite (Fig. 23B) with coarse spinules on anal operculum. Caudal rami (Figs. 23B; 24F; 31C—D) widely separated, cylindri- cal and slightly tapering posteriorly; without cup-shaped pores; about 4 times as long as wide; setae I-III closely set, I minute, I-III very long and thin; setae IV and V pinnate and with fracture planes, R. HUYS AND W. LEE seta IV distinctly longer than caudal ramus; setae VI-VII naked. Vent-pore and small tube-pore present ventrally near insertion sites of setae I-III (Fig. 31C—D). Rostrum as in d (Fig. 22B); large, trapezoid with straight anterior margin; delimited at base by transverse surface suture; with paired sensillae anteriorly and median tube-pore ventrally. Antennule (Fig. 22A) slender, 7-segmented; segments 1—2 with- out processes. Segment | with spinules around anterior margin; segment 4 forming large cylindrical pedestal ventrally. Armature formula: 1-[1], 2-[8 + 1 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[2], 7- [7 + acrothek]. Aesthetasc on segment 4 fused basally to 2 setae. Acrothek consisting of aesthetasc and 2 naked setae; set on small tubercle. Antenna (Fig. 23C) with elongate exopod bearing 2 lateral and 2 apical pinnate setae, and a longitudinal row of coarse spinules. Coxa with few large spinules, allobasis with pinnate abexopodal seta. Endopod with lateral armature consisting of | seta, 1 large and 1 small spine; distal armature consisting of 2 unipinnate spines and 3 geniculate setae (outermost fused basally to small seta). Labrum as in A. hirsuta. Mandible (Fig. 25A) with short gnathobase and small bilobed palp representing partially fused basis and endopod; with 2 lateral (basal) pinnate setae and 3 distal (endopodal) setae; exopod repres- ented by minute segment bearing 1 apical seta. Maxillule (Fig. 25B) and maxilla as in E. bulligera. Maxilliped (Fig.23D) slender, with elongate basis and endopodal claw. Syncoxa with 2 pinnate setae. Basis with naked palmar margin and setules around outer margin. Endopod represented by very long, naked claw bearing 1 accessory seta at base. P1 (Fig. 22F) with sparse ornamentation on coxa and basis. Basis with pinnate seta on anterior surface and along outer margin. Exopod 3-segmented, well developed; exp-1 with long pinnate outer spine; exp-2 with | naked outer spine; exp-3 with 2 unipinnate lateral setae and 2 geniculate setae apically. Endopod long and slender; enp-1 with long setules along inner margin and shorter spinules along outer margin, with thin inner seta in distal quarter (arrowed in Fig. 22F); enp-2 about twice as long as wide, with strong minutely pinnate claw and small accessory seta. P2—P4 as in Sars (1908). P3 enp-2 (Fig. 24B) with setiform outer spine (arrowed). Armature formula typical for genus. P5 (Fig. 23E). Endopodal lobe well developed, not extending beyond insertion sites of proximal outer setae of exopod; with distinctly stepped inner margin bearing 2 strong spines and | long distal seta (extending beyond apex of exopod); apex with 2 setae, outer one about twice length of inner one; tube-pores present near apical setae and proximal to innermost spine; outer basal seta inserting on cylindrical articulating setophore. Exopod narrow and elongate, produced apically into long tubular extension bearing | bare seta; inner margin with 1, outer margin with 1 naked and 3 pinnate setae. Both baseoendopod and exopod with elaborate or- namentation pattern as figured. MALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 475 um. Sexual dimorphism in antennule, P3 endopod, P5, P6 and genital segmentation. Antennule (Fig. 22B—E) 7-segmented, haplocer, with geniculation between segments 5 and 6. Segment 1 with spinules/setules around anterior margin; segment 2 longest; segment 4 minute, represented by incomplete sclerite (Fig. 22C). Segment 5 with large process proximally but forming long cylindrical pedestal distally (Fig. 22D). Segment 6 with 3 spinous processes along anterior margin (Fig. 22E). Distal portion of segment 7 elongated, displacing acrothek to position isolated from other armature (Fig. 22E). Armature formula: BASAL LAOPHONTID EVOLUTION prey ya on A eee ; $ UO RS TAC vin aS AX ESE d ‘ =o : TS ah / te rie A Rat Pee Mach pity ytd A gare itp i, \ Hig tae 7 rs! t 1 Si AML ris | = ¥ BI AFRAIC RS ES ST RE OSFK OEM KA, YY, LES ae ST t\. ‘4 Bes SOYA J A+100 yd 7 ea B} 100, i ; 1 ip ton fe \ Mi PEPPPPER TODD. \ 5 eo sp AN RU | ca rr “nm " 8 < LI ~s a4 eo : 1 \ X \y R&S ‘a \ XN Vy\y Vas pep ayy \ wi Tifity VASSA ASSES iN ' S pa yaa AS 1 ! \ MK 1 ee s Say \ ys AMANITA it ww oD is Y/ Mh \ \ % an \ al he . Fig. 21 Archesola hamondi gen. et sp. nov. A, Habitus 9, dorsal. Corbulaseta bulligera (Farran, 1913) comb. nov. B, habitus 9, dorsal. 83 84 R. HUYS AND W. LEE SEE ADR Es Sy pS —— GRY, mae Nttisres Fig. 22 Archesola typhlops (Sars, 1908) comb. noy. A, Antennule °, ventral; B, rostrum and antennule 4, dorsal [armature of segments 3—7 omitted]; C antennulary segments 3-4 6; D, antennulary segment 5 d; E, antennulary segments 6-7 6; F, P1 Q anterior [inner seta on enp-2 arrowed]. a; D, ( tral; B, anal somite and right caudal ramus, dorsal; C, antenn v. F, PS , anterior. v. (2). A, Genital double-somite, ven Fig. 23. Archesola typhlops (Sars, 1908) comb. no maxilliped; E, P5, anterior. Archesola hamondi gen. et sp. no BASAL LAOPHONTID EVOLUTION R. HUYS AND W. LEE WE SSS — ’ Sif iil = sa! ; eS N udal ramus, dorsal. Archesola v. B, P3 endopod 9 anterior [elongate outer spine arrowed]; C, P3 protopod and endopod 4, anterior [apophysis arrowed]; D, right P5 d, anterior [endopodal tube-pore arrowed]; F, left ca np-2 arrowed]; E, anal somite and left ca udal ramus, lateral; G, P3 enp-2 d. ta on e v. (@). A, P1, anterior [inner se Fig. 24 Archesola hamondi gen. et sp. no typhlops (Sars, 1908) comb. no BASAL LAOPHONTID EVOLUTION 1-[1], 2-[8 + 1 pinnate], 3-[7], 4-[2], 5-[8 + 1 pinnate spine + 1 spinous process + (2 + ae)], 6-[1 + 3 processes], 7-[8 + acrothek]. Apical acrothek consisting of aesthetasc and 2 bare setae. P3 endopod (Fig. 24C, G) 2-segmented; enp-1 as in 2; enp-2 with 3 inner setae (proximal one being distinctly shorter than in 9), 2 long apical setae and short pinnate outer spine (fused basally to segment); tube-pore present near outer apical seta. PS (Fig. 24D) medially fused, positioned ventrolaterally. Baseoendopod without endopodal lobe or armature; medial margin with few setules and 2 tube-pores (longest arrowed); outer basal seta arising from long, articulating, cylindrical setophore. Exopod free, rectangular; with | long pinnate seta apically; inner margin proximal seta and distal bipinnate spine; outer margin with 2 bare setae. Sixth legs represented by well developed opercula, one articulat- ing and closing off left or right genital aperture; each produced into cylindrical process bearing 1 lateral and 1 apical seta. REMARKS. Drzycimski (1969) corrected two major errors in Sars’ (1908) description. First, he pointed out the presence of the thin inner seta on the proximal endopod segment of P1. Within the Laophontidae this element is further only found in Archilaophonte maxima. Secondly, Drzycimski remarked that the inner seta on the baseoendopod of the male P5 is not well developed as in Sars’ illustration but greatly reduced. In reality, Drzycimski referred to the short hyaline tube-pore located closely to the exopod whereas in Sars’ (1908) illustration it was the longer medial tube-pore (arrowed in Fig. 24D) which was misinterpreted as a genuine seta. One character that has traditionally been used to differentiate A. typhlops from A. longiremis is the setation of the female P5 exopod. This distinction is invalid since it is based on the erroneously reported absence of the proximal surface seta in Sars’ description of A. typhlops. The same error also served to distinguish E. typhlops pontoica from the type population (Por, 1959, 1964a). Reliable records of A. typhlops include Flekkeré (Sars, 1908), Bergen ( Drzycimski, 1969) and Frierfjord/Langesundfjord (this account) in Norway, Gullmar Fjord (Lang, 1948) and the Isle of Bonden (Por, 1964a) in Sweden, and Norfolk in England (this account). The Scottish records from the River Ythan (Aberdeen- shire) by Hockin & Ollason (1981) and Hockin (1982a—b, 1984) and that from Newbiggin (Northumberland) by Moore (1973) may be based on A. longiremis. Archesola longiremis (T. Scott, 1905) comb. nov. Laophonte longiremis T. Scott, 1905 Esola longiremis (T. Scott, 1905) Lang (1948) TYPE LOCALITY. Granton, Firth of Forth, Scotland; old quarry opening to the sea (T. Scott, 1905, 1906). TYPE MATERIAL. _T. Scott (1905) recorded an unspecified number of females; this material has not been deposited in any of the British museums (London, Newcastle-upon-Tyne, Edinburgh) and is there- fore almost certainly lost. REMARKS. This species is very close to A. typhlops and can be differentiated primarily by the shorter caudal rami (only twice as long as wide) and the smaller body size (0.6 mm). Lang (1948) pointed out that T. Scott’s (1905) drawing of the P5 showed an aberrant setation on the endopodal lobe (total of 7 setae: 3 inner, 3 apical, 1 outer). The short apical seta is almost certainly the equiva- lent of the long tube-pore found in this position in A. typhlops, however, the presence of the supernumerary outer seta is more difficult to explain since no laophontoidean is known to display more than 5 elements on the endopodal lobe of the female P5 (Huys, 87 1990a; Huys & Lee, 1999). We suspect that this seta is the result of an observational error. The species has never been figured again since T. Scott (1905) nor has the male been discovered. Wells (1961) illustrated some features of a male specimen from St. Martin’s (Isles of Scilly) which he attributed to E. longiremis. The P5 shows only 3 setae on the exopod and the endopodal armature is represented by 2 fine setae (one of which likely to be a tube-pore). The P6 bears 2 strong setae but is not drawn out into a cylindrical process as in other species of the genus. These characters in conjunction with his statement that the male antennule is subchirocerate and the endopod 3-segmented clearly exclude the possibility that Wells was dealing with a species of Archesola or any other esolinid genus. Wells (1963) also recorded the species from Exmouth (Devon) but this record remains uncon- firmed. The genus Archesola consists of a complex of closely related species which can be differentiated primarily by morphometric characters, such as caudal ramus length and Pl exopod: endopod ratio, and various setal length differences on the PS. Coull’s (1971) identification of E. longiremis from North Carolina suggests an amphi-Atlantic distribution for the genus Archesola, however, in view of the relatively subtle differences between congeners, the specific identity of his record remains to be confirmed. Archesola hamondi sp. nov. TYPE LOCALITY. 53°10.34'N 00°56.34'E; depth 12-13 m; fine sand with high silt and shell gravel content. TYPE MATERIAL. This species is only known from the holotype 2 (leg. R. Hamond; 06 May 1992) which unfortunately was acciden- tally destroyed before the description could be completed. The brief description below provides sufficient information to warrant the proposal of a new species. ETYMOLOGY. This patronym is dedicated to Dr Richard Hamond who collected the holotype, in recognition of his significant contri- butions to laophontid systematics. DESCRIPTION. FEMALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 600 pm. Body (Fig. 21A) dorsoventrally depressed and much wider than in A. typhlops; covered with dense pattern of minute surface lamel- lae dorsally and laterally. Cephalothorax bell-shaped, distinctly widening towards posterior margin; lateral and hind margins fringed with long setules; without paired cup-shaped pores. Setular fringes also present laterally on pedigerous somites and urosomites, some- times forming tufts locally. Posterior margin of urosomites without distinct ornamentation dorsally except for penultimate somite bear- ing transverse row of fine spinules. Genital double-somite (Fig. 21A) wide and dorsoventrally flat- tened; original segmentation marked by bilateral constriction only; without cup-shaped pores in anterior half; lateral lobes without backwardly directed strong spinules. Genital field as in A. typhlops. Anal somite (Fig. 24E) with distinct setular fringe around anal opening; anal operculum completely bare; posterolateral margins with fine spinules. Caudal rami (Fig. 24E) cylindrical and slightly swollen in anterior half; distinctly wider than inA. typhlops; about 3 times as long as wide. Seta II shorter and seta II posteriorly displaced compared to A. typhlops; setalV reduced, lacking fracture planes, shorter than caudal ramus; seta V well developed, pinnate, without fracture planes; setae VI-VII naked. Large vent-pore present at outer subdistal corner. 88 Rostrum (Fig. 21A) longer than in A. typhlops; trapezoid with straight anterior and concave lateral margins; delimited at base by transverse surface suture; with paired sensillae anteriorly and me- dian tube-pore ventrally. Antennules to maxillipeds as in A. typhlops. P1 (Fig. 24A) as in A. typhlops except for (a) basal pedestal bearing endopod wider, (b) inner seta on enp-1 inserting more distally, and (c) enp-1 about twice the length of exopod (distinctly shorter in type species). P2—P4 as in A. typhlops. P5 (Fig. 23F). Endopodal lobe well developed, well extending beyond insertion sites of proximal outer setae of exopod; with distinctly stepped inner margin bearing 2 strong spines (more closely set than in A. typhlops) and 1 bare distal seta (not extending beyond apex of exopod); with 2 apical setae, outer one about 1.5 times length of inner one; tube-pores present near apical setae and proxi- mal to innermost spine; outer basal seta inserting on cylindrical articulating setophore. Exopod elongate but distinctly shorter than in A. typhlops, produced apically into short tubular extension bear- ing 1 bare seta; inner margin with 1, outer margin with 4 pinnate setae. Both baseoendopod and exopod with elaborate ornamentation pattern as figured. MALE. Unknown. REMARKS. Differentiation of A. typhlops and A. hamondi is best achieved by comparison of the general body shape, caudal ramus outline and armature pattern, and 2 PS morphology and morpho- metry Esola sp. sensu Chislenko (1967) Chislenko illustrated a male which he obtained in Laminaria saccharina washings from the White Sea and identified as Esola sp. His drawings of the caudal ramus, P3 endopod and P1 leave little doubt that this species belongs to Archesola and is obviously close to A. typhlops and A. longiremis. The caudal ramus L:W ratio appears to be intermediate between the latter two species and the P1 endopod and exopod have slightly different proportions. Chislenko’s male (0.35 mm) is smaller than those of A. typhlops recorded by Drzycimski (1969) from the Bergen area (0.45 mm) and our single male from West Runton (0.475 mm). The antennary exopod bearing the atypical number of 5 setae is obviously based on an aberrant specimen. His illustration of the P3 endopod lacks the proximal inner seta on the distal segment, its location being indicated by the distinct step in the inner margin. Finally, the small inner seta illustrated on the P5 baseoendopod is probably a tube-pore. The White Sea material identified by Brotskaya (1961) as E. longiremis is likely to be conspecific with this species, the true identity of which is as yet uncertain. Consequently, Chislenko’s species is tentatively ranked specues inquirenda in Archesola. Archesola typhlops pontoica (Por, 1959) comb. nov. Esola typhlops pontoica Por, 1959 TYPE LOCALITY. Black Sea coast, Rumania. TYPE MATERIAL. Dr Ileana Negoescu (Museum ‘Grigore Antipa’, Bucharest) informed us that the syntypes no longer exist. REMARKS. Por (1959, 19645) established this subspecies for 3 22 found at 61-69 m depth off the Rumanian coast, however it is doubtful whether his material deserves such status. The author discriminated the Black Sea population on the basis of the presence of 6 setae on the 2P5 exopod (Sars (1908) erroneously figured only 5), the slightly shorter caudal rami and the incompletely 3-seg- mented P1 exopod (a feature displayed in only 1 specimen!). The R. HUYS AND W. LEE most significant difference, not mentioned by Por, is found in the proportional lengths of the distal antennulary segments (segments 6 and 7 being of equal length). Examination of new material is necessary to resolve the identity of the Rumanian population; E. typhlops pontoica is considered here as subspecies inquirenda. Genus Corbulaseta gen. nov. The diagnosis below is based on Vervoort’s (1964) redescription of E. bulligera and personal observations of Wells’ (1970) material from Great Britain Rock, Isles of Scilly, and additional specimens collected from the Belgian North Sea coast by the senior author. DIAGNOSIS. Laophontidae. Body cylindrical; posterolateral cor- ners of 2 genital double-somite and second abdominal somite laterally and backwardly produced. Integument of cephalothorax and body somites with dense pattern of spinules and setules. Ros- trum large, partly delimited at base by incomplete surface furrow. Cephalothorax with one pair of large, anterodorsal cup-shaped pores; such pores absent on genital (double-)somite and caudal rami. Anal operculum spinulose. Caudal rami rectangular, short; not sexually dimorphic. Sexual dimorphism in antennule, P3 endopod, P5, P6 and in genital segmentation. Antennules slender; 6-segmented in 9, subchirocer and 7-seg- mented in 6; segment 1 with 1—2 minute processes along posterior margin; with aesthetasc on segment 4 () or 5 (d) and as part of apical acrothek on distal segment; segment 5 ¢ swollen, without anterior outgrowth; proximal aesthetasc fused basally to 2 setae. Antenna with 4 setae on exopod; allobasis with abexopodal seta. Labrum with distal spinular ornamentation. Mandible with 1-seg- mented palp; exopod and endopod represented by small tubercles bearing 1 and 3 setae, respectively; basis represented by 2 apical setae. Maxillule with minute, defined exopod. Maxilla with 3 endites on syncoxa; endopod represented by 3 setae. Maxilliped slender; syncoxa with 2 setae; entire palmar margin with long setules; endopodal claw elongate. P1 with 2-segmented exopod bearing 5 setae on exp-2 and elongate endopod; enp-1 without inner seta, enp-2 with minute seta and long, slender claw. P2—P4 with 3-segmented exopods and 2- segmented endopods. P2 basis with short outer spine. Outer spine of P2—P4 enp-2 setiform and very long in P3-P4. P4 endopod modified in both sexes; distal inner seta proximally dilated, bearing enlarged spinules which enclose long secretory tube-pore arising from seg- ment. P3 endopod d 3-segmented; enp-2 with inner seta and short outer spinous apophysis. Armature formula as follows: Exopod Endopod P2 0.1.123 Ail P3 0.1.223 e372 [d: 1.1.220] P4 0.1.223 0.221 *: or 1.220 in Vervoort’s (1962) 2 specimen of E. bulligera from New Caledonia. P5 2 with separate rami; exopod elongate, with 6 setae/spines; baseoendopod slightly developed, with 4 setae/spines. PS d without endopodal lobe; exopod short, with 1 inner, 2 apical and 2 outer setae/spines. P6 forming opercula closing off paired genital apertures; with 2 small setae at outer corner. P6 d asymmetrical; membranous flaps with 1 long and 1 minute seta. BASAL LAOPHONTID EVOLUTION TYPE AND ONLY SPECIES. Laophonte bulligera Farran, 1913 = Corbulaseta bulligera (Farran, 1913) comb. nov. ETYMOLOGY. The generic name is derived from the Latin corbula (little basket) and seta (bristle) and refers to the modified distal inner seta of P4 enp-2, the proximal setules of which form a trapping basket typically enclosing a secrete bolus. Corbulaseta bulligera (Farran, 1913) comb. nov. Laophonte bulligera Farran, 1913 Esola bulligera (Farran, 1913) Lang (1948) Laophonte rosei Monard, 1926 Laophonte Rosei Monard, 1926: Monard (1928) Esola rosei (Monard, 1928) Lang (1948) TYPE LOCALITY. Blacksod Bay, Co. Mayo (Ireland); 1.8—5.4 m depth MATERIAL EXAMINED. Farran’s (1913) type material is lost (J.M.C. Holmes, pers. comm). (a) Isles of Scilly, Great Britain Rock: 1 @ in alcohol (BMNH 1967.10.31.76); coll. University of London Sub-Aqua Expedition 1966; det. J.B.J. Wells; (b) Belgium, North Sea coast, 51°30"N 2°00"E: 19, 1 3; 08 April 1986, depth 14.1 m, sandy substrate; leg. R. Huys. ADDITIONAL OBSERVATIONS. FEMALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 570-590 um. Body (Fig. 21B) cylindrical, slightly depressed; covered by irregular pattern of minute surface spinules. Cephalothorax widest, subrectangular; with pair of large cup-shaped pores anterodorsally (Fig. 25C); ventral pores absent; posterior margin and anterior half of ventral margin with setular fringe; posterior half of ventral margin bordered by tiny spinules; posterolateral corner produced forming distinctive lobate extension (Fig. 25C). Prosome gradually tapering posteriorly; all somites with dorsal transverse spinular row and setular fringe around hind mar- gin. Genital double-somite dorsoventrally depressed; original seg- mentation marked by bilateral constriction and dorsal transverse spinular row set on surface ridge; ventral surface without conspicu- ous ornamentation; cup-shaped pores absent. Genital aperture closed off by sixth legs bearing | naked seta. Posterolateral corners of second abdominal somite backwardly produced; remaining urosomites distinctly narrower. Ventral posterior margin of penulti- mate somite with medial fringe of fine setules flanked by strong spinules (decreasing in length ventrolaterally). All urosomites with spinules around dorsal posterior margin. Anal operculum spinulose. Caudal rami short, slightly longer than wide; all setae arranged in posterior quarter; setae IV and V well developed, pinnate, with fracture planes; no conspicuous pores present. Rostrum (Fig. 21B) trapezoid, delimited at base by incomplete surface suture; with 2 long sensilla apically and tube-pore ventrally. Antennule 6-segmented; posterior margin of segment | with slight bulbous swelling but no real spinous processes; aesthetasc on segment 4 fused basally to 2 long setae; armature formula: 1-[1], 2- [7 + 1 pinnate], 3-[6], 4-[(2 + ae)], 5-[1], 6-[9 + acrothek]; acrothek consisting of aesthetasc and 2 naked setae. Antennary exopod with strong pinnate outer apical spine and 3 pinnate setae. Labrum with Sparse ornamentation resembling condition in A. hirsuta. Mandibu- lar palp 1-segmented, with ancestral setation, i.e. 2 basal, 1 exopodal and 3 endopodal setae. Maxillule as in E. bulbifera, with endopod represented by 2 setae. Maxilla as in E. bulbifera. Maxilliped with 2 setae on syncoxa; palmar margin with long fine spinules; endopodal 89 claw slender and longer than basis, with 1 accessory seta. P1 as in Farran’s (1913) description except for outer spine of exp- 1 being longer and pinnate and proximal and middle outer spines of exp-2 distinctly shorter. P2 basis with bipinnate outer spine, P3—P4 bases with smooth outer seta. Outer spine of P3—P4 enp-2 very long and setiform (Fig. 25D). P4 (Fig. 25D) with 2-segmented endopod; enp-1 short, without inner seta; enp-2 (Fig. 25E-F) highly distinctive: distal inner seta with dilated base bearing comb of long curved setules on both anterior and posterior outer margins; this ornamentation forming trapping basket enclosing large secrete bolus produced by long anterior surface tube-pore located near distal margin of enp-2. PS as in original description. MALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 530 um. Body more slender than in °; none of urosomites with backwardly produced posterolateral corners. Ven- tral posterior margin of postgenital (except anal) somites with median fringe of fine setules flanked by strong spinules. Antennule subchirocerate; 7-segmented with geniculation between segments 5 and 6. Segment | without distinct processes, segment 5 without anterior outgrowth. P3 endopod 3-segmented; very similar to that of EF. bulbifera (Fig. 4D). PS without endopodal lobe; medial margin frimged with long spinules and | tube-pore; outer basal seta arising from short setophore. Exopod elongate, about 3.5 times as long as wide; with | seta and 1 spine along inner margin, apex with | long bipinnate seta, outer margin with 2 bipinnate spines. P6 asymmetrical; each opercular flap with cylindrical extension at outer corner bearing long outer seta and minute inner seta. REMARKS. Nicholls (19416) pointed out that Laophonte rosei, described from Banyuls (Monard, 1926) may well be a junior synonym of L. bulligera since the difference between them appears to be based on two doubtful characters. The ‘sensory organ’ illustrated on the P4 endopod of L. bulligera by Farran (1913) was not described for L. rosei by Monard (1926) although the latter did illustrate the adjoining modified seta (see also Monard (1928)). Secondly, the different number of setae on the P5 endopodal lobe is based on Monard’s failure to observe the seta near the base of the baseoendopod, a portion of which appears to have been lost in L. rosei. Lang (1948) also expressed strong reservations about the distinctiveness of L. rosei but like Nicholls (1941b) and Vervoort (1967) nevertheless maintained it as a valid species. We can see no justification for this distinction and formally relegate E. rosei to a junior subjective synonym of E. bulligera. Pesta (1959) published an incomplete description of the male (as E. rosei) but did not mention the transformed P4 endopod. The discrepancy found in the length of the P1 endopod casts some doubt on his identification. Pending the re-examination of mediterranean material, the known records suggest an almost continuous boreo-mediterranean distri- bution pattern with records from Ireland (Farran, 1913, 1915), Isles of Scilly (Wells, 1970), Belgian coast (unpubl.), Banyuls-sur-Mer (Monard, 1926, 1928) and possibly Naples (Pesta, 1959). Por & Marcus (1972) recorded the species also in the Great Bitter Lake and off Port Taufig in the southern part of the Suez Canal and considered the species an Atlantic (anti-Lessepsian) immigrant. There is no morphological evidence supporting Alheit & Scheibel’s (1982) record from Harrington Sound in Bermuda. The isolated record from New Caledonia by Vervoort (1962) is difficult to interpret, particularly because his single female speci- men deviates from European E. bulligera in the absence of the outer spine on P2 enp-2. Vervoort (1962) did not remark on this character ee R. HUYS AND W.LEE — A Vimas ; 3 oo l \) h 4 : \ . 5 tS ; Da TIN = 7 co = *. es x > ‘ ce . i > ESA : : a SS SS So. Nya: SO SN, SESS | Wes, yi Nea | | KAN fi Yh | Nia DNA] SRR / Ad i YONT WY i Hiei D\ f \SSTSNE IN Uys / {AN \SUIRSRN } \ (DANS Ys bE A WSS PANTING > LS ae TE Af V dee —— ee s Z — A < NAG VE: ieee — Jf 7+ LES Fig. 25 Archesola typhlops (Sars, 1908) comb. nov. A, Mandibular palp; B, maxillule, anterior. Corbulaseta bulligera (Farran, 1913) comb. nov. C, cephalothorax, lateral; D, P4 2, anterior; E, P4 endopod 2, anterior; F, P4 enp-2 °, medial [contours of secrete bolus stippled in E-F]. Laophonte parvula Sars, 1908. G, P5 2, anterior [anteriorly displaced outer seta arrowed]. BASAL LAOPHONTID EVOLUTION presumably because of the lack of a base for comparison in Farran’s (1913) description and illustrations which omitted P2 and P3. The problem is exacerbated by the aberrant left-right asymmetry (1.220 vs 1.320) displayed on the P2 endopods. It is unclear whether the reduced setal formula is real and therefore indicative for the pres- ence of a second species in the western Pacific. There is very little additional evidence pointing in this direction except for the different cephalothorax shape (in lateral aspect: compare Fig. 25C) and some morphometric discrepancies in the caudal rami, which appear to be longer, and in the exopods of P2—P4, which are more abbreviated. E. bulligera cannot be retained in the genus Esola because of the absence of (1) distinct spinous processes on the first antennulary segment, (2) cup-shaped integumental pores on the genital (double-) somite and caudal rami, (3) characteristic labral ornamentation and (4) caudal ramus sexual dimorphism. It is reminiscent of Bathyesola compacta (see below) in the presence of only one pair of cup-shapes pores on the cephalothorax but differs from it in the reduced armature on the 2 P5 exopod and the transformed P4 endopod which is the most significant autapomorphy of E. bulligera, justifying its placement in a new genus Corbulaseta. Genus Bathyesola gen. nov. DIAGNOSIS (based on 2 only). Laophontidae. Body cylindrical; posterolateral corners of 2 genital double-somite and second ab- dominal somite laterally and backwardly produced. Integument of cephalothorax and body somites with dense pattern of spinules and setules. Rostrum large, partly delimited at base. Anterolateral pair of small integumental cup-shaped pores present on cephalothorax. Caudal rami not modified in 9, cylindrical and elongate. Sexual dimorphism presumably in antennule, P3 endopod, P5, P6, and genital segmentation. Antennules slender; 7-segmented in 2; segment | without spinous processes along posterior margin; with aesthetasc on segment 4 (fused basally to 2 setae) and as part of apical acrothek on segment 7. Antenna with 4 setae on exopod; allobasis with abexopodal seta. Labrum without overlapping scales distally but with pattern of spinules anteriorly. Mandible with 2-segmented palp; endopod free, with 3 setae; exopod represented by single seta; basis represented by 2 setae. Maxillule with defined exopod. Maxilla with 3 endites on syncoxa; endopod represented by 3 setae. Maxilliped robust; syncoxa with 2 setae; entire palmar margin with spinules; endopodal claw relatively stout. P1 with large 3-segmented exopod bearing 4 setae on exp-3 and relatively short endopod; enp-1 without inner seta, enp-2 with minute seta and short, curved claw. P2—P4 with 3-segmented exopods and 2-segmented endopods. P2 basis with moderately long outer spine. Inner seta of P2—P4 exp-2 reduced. Outer spine of P2—P4 enp- 2 setiform, short in P2—P3, long in P4. Armature formula as follows: Exopod Endopod P2 0.1.123 e222 P3 0.1.123 0.321 [d presumably 0.1.220] P4 0.1.123 0.221 P5 2 with separate rami; exopod relatively short, with 6 setae/ spines; baseoendopod well developed, with 5 setae/spines, apical setae reduced; outer basal seta on short setophore. P6 Yforming opercula closing off paired genital apertures; with 2 small setae. TYPE AND ONLY SPECIES. Bathyesola compacta gen. et sp. nov. 91 ETYMOLOGY. The generic name refers to the bathyal distribution of the type species. Bathyesola compacta gen. et sp. nov. TYPE LOCALITY. 18°50'S, 173°29'W, ‘White Lady’ site on North Fiji Ridge, west of Fiji; 2765 m depth. Accompanying harpacticoid fauna: several 92 and dod of Xylora bathyalis Hicks, 1988 (Thalestridae: Donsiellinae). TYPE MATERIAL. Holotype 2 dissected on 6 slides, deposited in Muséum National d’ Histoire Naturelle, Paris under MNHNP Cop- 1869; collected during STARMER II expedition, station 14 (Kaiyo 87), dive 19; 14 July 1989; leg. L. Laubier. ETYMOLOGY. The species name alludes to the compact P1, dis- playing a short and robust endopod. DESCRIPTION. FEMALE. Body length from anterior margin of rostrum to posterior margin of caudal rami 360 um. Maximum width (105 um) measured at posterior margin of cephalothorax. Body (Fig. 26A—B) cylindrical, slightly dorsoventrally depressed, covered with dense pattern of minute spinules dorsally and laterally. Cephalothorax slightly wider than free somites, posterolateral angles backwardly produced forming small lobate extension (Fig. 26B); with pair of small lateral cup-shaped pores. Posterior margin of cephalothorax and all body somites with row of long setules dorsally and laterally. Pleurotergite of P5-bearing somite almost as wide as anterior somites. Genital double-somite wide and dorsoventrally flattened; with lateral, backwardly produced extensions in posterior (=abdominal) half; original segmentation marked by bilateral constriction and spinule row arising from transverse surface ridge dorsally and laterally; posterior half with backwardly directed lobate extensions bearing spinular tuft; ventral surface without spinular ornamenta- tion; genital field located near anterior margin. Sixth legs forming well developed opercula closing off paired genital apertures; each with 2 small setae. First postgenital somite with backwardly produced lateral angles, bearing spinular tuft; without ventral ornamentation. Penultimate and anal somites distinctly narrower; ventral posterior border with long spinules. Anal somite with spinulose anal operculum. Caudal rami (Fig. 26A—B) widely separated; about 4 times as long as average width; maximum width measured at base; dorsal surface with minute spinules; seta I small, setae II-III well developed, naked and closely set; setae IV (naked) and V (pinnate) with fracture planes, seta V 2.8 times as long as seta IV; setae VI-VII naked. Rostrum (Figs 26A) large, blunt anteriorly; delimited at base by transverse surface suture; with paired sensillae anteriorly and me- dian tube-pore dorsally. Antennule (Fig. 26A—B) relatively short, distinctly 7-segmented, without spinous processes on segments 1—2. Segment 1| with large spinular patch around anterior margin. Armature formula: 1-[1], 2- [4 + 4 pinnate], 3-[6], 4-[1 + (1 + ae)], 5-[1], 6-[2], 7-[5 + 1 pinnate + acrothek]. Acrothek consisting of aesthetasc and 2 naked setae; set on apical pedestal. Antenna (Fig. 27A). Coxa with spinules on both inner and outer margins. Exopod short, bearing 2 lateral and 2 apical pinnate elements, and a longitudinal row of fine spinules along outer margin. Allobasis with pinnate abexopodal seta. Endopod with lateral arma- ture consisting of 2 spines and 1 minute seta; distal armature consisting of 2 naked spines and 3 geniculate setae (outermost fused basally to minute seta). 92 Fig. 26 R. HUYS AND W. LEE im im Tan Ne i Tit 1 ni ty Tree fn \f stu eA ail i nmin i mtn i Me = Gee ecr , — oi TTA ee Hh) AHN) ( a op Tan RTT aes i rari om Gh oe ene itl ar ea Fj aN iN ee uy, NN ON pe = — WAY, Lt +) 100 yu 1 sr Bathyesola compacta gen. et sp. nov. (@). A, Habitus, dorsal; B, habitus, lateral. - BASAL LAOPHONTID EVOLUTION ity, _ Fig.27 Bathyesola compacta gen. et sp. nov. (2). A, Antenna; B, mandible; C, maxillule, anterior; D, maxilla; E, maxilliped; F, P1, anterior. ‘d > r) t 93 ? R. HUYS AND W. LEE 94 pee phe hb = heeds. pit WA wth pb eee —~ IF bf Y o li A Se Na Gc / ac eX = es, Se ee Se aes C= e Yipee UTE oe Srna aaa aS SS Se = NG 2 : Lh SES en ie 8 RNS 50u Fig. 28 Bathyesola compacta gen. et sp. nov. (@). A, P2, anterior; B, P3, anterior; C, P4, anterior; D, P5, anterior. Paralaophonte pilosoma Vervoott, 1964. E, P3 endopod 4, anterior. BASAL LAOPHONTID EVOLUTION Labrum with spinular patches on anterior face but no overlapping scales. Mandible (Fig. 27B) with short gnathobase and small 2-seg- mented palp representing free endopod and fused basis and exopod; endopod a minute segment with 3 pinnate setae; basal armature represented by 2 lateral pinnate setae, exopod represented by single seta. Paragnaths highly ornate lobes as in E. bulbifera. Maxillule (Fig. 27C) with elongate arthrite bearing | seta on anterior surface and 9 elements around distal margin. Coxal endite with | spine and | seta, basal endite with 1 spine and 2 setae. Exopod a short segment with 2 distal setae; endopod incorporated into basis, represented by 2 setae. Maxilla (Fig. 27D). Syncoxa with very long setules around outer margin and few additional spinule rows as figured; with 3 endites; praecoxal endite small, with | naked seta; middle endite drawn out into spine, with 2 setae; distal endite with 3 elements. Allobasis produced into strong curved claw; accessory armature consisting of 2 setae. Endopod incorporated into allobasis, represented by 3 bare setae. Maxilliped (Fig.27E) compact, basis and endopodal claw not particularly elongate. Syncoxa with 2 pinnate setae. Basis with spinular ornamentation as figured; spinules present along entire palmar margin. Endopod represented by stout, minutely pinnate claw bearing | accessory seta and tube-pore at base. P1 (Fig. 27F) with dense ornamentation on praecoxa, coxa and basis. Basis with pinnate seta on anterior surface and along outer margin. Exopod large, 3-segmented; exp-1 with pinnate outer seta; exp-3 with 2 unipinnate outer spines and 2 geniculate setae apically. Endopod robust and relatively short; enp-1 about 2.2 times as long as basis, with long setules along inner margin and fine spinules along outer margin; enp-2 about as long as wide, with short unipinnate claw and small accessory seta. P2—P4 (Figs 28A—C) with 3-segmented exopods and 2-segmented endopods. P2 basis with long, bipinnate outer spine; P3—P4 bases with bare outer seta. P2 enp-1 with pinnate inner seta, P3—P4 enp-1 unarmed. Inner seta of P2 exp-2 reduced. Outer spine of P2—P4 enp- 2 setiform, very long in P4. Pore present near distal outer corner of P3—P4 enp-2. Armature formula as for genus. P5 (Fig. 28D). Endopodal lobe well developed, extending to halfway down the exopod; with 2 reduced bare setae apically, and 2 long widely separated setae along inner margin; pores present near articulation with exopod, at base of apical setae and proximal to innermost seta. Exopod relatively short, produced apically into short tubular extension bearing | bare seta; inner margin with 1, outer margin with 4 pinnate setae; inner seta slightly longer than apical one. Both baseoendopod and exopod with spinulation as figured. MALE. Unknown. REMARKS. The discovery of B. compacta at 2765 m depth at the North Fiji Ridge represents the deepest record thus far for the family Laophontidae (Lee & Huys, 1999). It displays a mozaic of primitive (7-segmented 2 antennule; 3-segmented P1 exopod; 2 P5 endopodal lobe with 5 setae/spines) and advanced characters (P3—P4 enp-1 without inner seta; P3—P4 exp-3 with | inner seta) which serves to distinguish the species from other esolinids. Status of Esola Spelaea (Chappuis, 1938) Lang (1944, 1948) placed Laophonte spelaea in the genus Esola without giving any explicit reasons. From his generic diagnosis and the phylogenetic scheme presented on p. 1450 (Lang, 1948), one can 95 infer that his course of action was based solely on the presence of an outer spine on the distal endopod segment of P2. Although this character was diagnostic for Esola in Lang’s sense it is clearly a symplesiomorphy shared by all genera in the Archilaophonte-Esola lineage (with the exception of Mourephonte) and consequently of no value in inferring relationships. Lang (1944, 1948) subdivided Esola into two species groups, diagnosed by the number of setae on the male P5 endopodal lobe and the armature of the P3 in both sexes. His spelaea-group included only E. spelaea and has until now remained monotypic. It differed from the Jongicauda-group in the presence of 2 setae (rather than | or 0) on the d P5 endopodal lobe and a reduced armature on the P3 exopod (exp-3 with only 2 outer spines) and endopod (enp-2 @ with only 2 inner setae; endopod 3 without inner seta on enp-2 and with only 3 setae on enp-3). Chappuis’ (1938) description is very brief and provides illustra- tions of the male P2—P5 only. Unfortunately the author did not give any information about the position of the setae on the female P5 which could have provided the justification for including L. spelaea in the Archilaophonte-Esola lineage since in all of its members (1) the proximal seta of the endopodal lobe is medially displaced and (2) the insertion sites of the 2 proximal setae of the exopod are superim- posed. Chappuis’ statement that there are 4 setae on the baseoendopod and 5 or 6 setae on the exopod can be interpreted in the light of this generalized pattern. His reservation about the correct number of exopodal setae might indicate the close or overlapping position of some of these elements. Secondly, due to its strong medial displace- ment the proximal endopodal seta has frequently been overlooked or lost during dissection (Thompson & A. Scott, 1903; Norman, 1911; Monard, 1926, 1928; Noodt, 1955), leaving open the possibility of a similar observational error made in Chappuis’ (1928) description. The actual number of endopodal setae on the female P5 of L. spelaea could therefore be five rather than four. Chappuis’(1928) armature formula of P2 exp-3 tabulated as 222 (i.e. with 2 outer spines) is unlikely to be correct when both P2 and P4 reportedly have 3 outer spines on exp-3. No laophontid described thus far displays a [3-2-3] outer spine pattern for P2—P4 and hence we suspect 123 (as in B. compacta) to be the correct formula for P2 exp-3. Chappuis (1938) described L. spelaea from three caves in Apulia, southern Italy (Abisso and La Zinzulusa near Castro, Grotta dei Diavoli near Badisco) and regarded it as a marine relict. The caves exhibit a tidal regime but the salinity approaches that of freshwater (‘. . . das Wasser schmeckt aber fast stiss’) which appears to be confirmed by the presence of the stygobiont mysids Spelaeomysis bottazzii Caroli and Stygiomysis hydruntina Caroli and the palaemonid Typhlocaris salentina Caroli, all of which are endemic to coastal caves and phreatic waters in the Apulia region. Both Pesce (1985) and Rouch (1986) consider the species as a descendant from a marine ancestral stock which successfully colonized subterranean freshwater habitats via littoral karstic systems, possibly during regression periods in the Tertiary (“Regression Model Evolution’ ). Laophonte spelaea cannot be accommodated in any of the exist- ing laophontid genera. It appears to be related to Bathyesola in certain aspects (see above) but differs from it in the presence of an inner seta on P3—P4 enp-1, only 2 inner setae on P3 enp-2 and more primitive setal formula on the P4 exopod. In view of the strong ecological divergence between B. compacta and L. spelaea we prefer to establish a new genus for the latter. The male P3 endopod in Troglophonte gen. nov. does not accord with the pattern found in the other esolinid genera. The absence of an inner seta on the middle segment could be related to the reduced 1.221 pattern in the female but might also indicate a relationship with a large group of other laophontid genera which typically lose the proximal inner seta during male P3 ontogeny. 96 R. HUYS AND W. LEE Fig. 29 SEM micrographs. Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. (2). A, Lateral margin of genital double-somite, ventrolateral; B, lateral cup-shaped pore on genital double-somite [pore exit arrowed]; C, setular extensions bordering dorsal margin of cup-shaped pore; D, labrum and mandibular gnathobases. [Scale bars: 2 um (C), 10 um (B, D), 20 um (A)]. BASAL LAOPHONTID EVOLUTION 97 Fig. 30 SEM micrographs. Applanola hirsuta (Thompson & A. Scott, 1903) comb. nov. (2). A, Cephalothorax and rostrum, frontal [anterodorsal pore arrowed]; B, cephalothorax, ventral [anteroventral pore arrowed]; C, anal opening and caudal ramus, ventral [ventral pore arrowed]; D, right caudal ramus, ventral, showing cup-shaped pore. [Scale bars: 6 um (D), 15 um (C), 20 um (B), 60 um (A)]. Genus Troglophonte gen. nov. DIAGNOSIS. Laophontidae. Body shape unknown but somites not well demarcated. Rostrum short, presumably fused at base. Integumental cup-shaped pores unconfirmed. Anal operculum spinulose. Caudal rami short, squarish. Sexual dimorphism in antennule, P3 endopod, P5, P6 and in genital segmentation. Antennules 7-segmented in 9, segmentation unknown in d. An- tenna with 4 setae on exopod; allobasis with abexopodal seta. Mouthparts unknown. Maxilliped very slender. P1 with 3-segmented exopod bearing 4 setae on exp-3 and slender endopod; enp-1 without inner seta, enp-2 with minute seta and slender, strong claw. P2-P4 with 3-segmented exopods and 2-seg R. HUYS AND W. LEE x009Z dx3 emubey 4 i 4016 ey ubew a o =) < © ~ ole os dx3 Fig. 31 SEM micrographs. Esola bulbifera (Norman, 1911). A, Anal somite and caudal rami, dorsal. Applanola hirsuta (Thompson & A. Scott, 1903) comb. noy. B, mandibular palp and inner face of cephalothorax showing vent-pore [arrowed]. Archesola typhlops (Sars, 1908) comb. nov. C, caudal ramus, ventral; D, caudal ramus, area around setae I-III showing pores [arrowed]. [Scale bars: 5 um (D), 10 um (B), 20 um (A, C)]. BASAL LAOPHONTID EVOLUTION mented endopods. P3 endopod d 3-segmented; enp-2 with outer pinnate apophysis but without inner seta. Armature formula as follows: Exopod Endopod Pp 0.1.123 1.221 PS 0.1.123 (or 0.1.2226) 1.221 [d: 1.0.120] P4 0.1.223 22 P5 2 with separate rami; exopod elongate, with 5 or 6 setae/ spines; baseoendopod slightly developed, with 4 setae/spines. PS 3 with trapezoid endopodal lobe; exopod short, with 1 inner, 2 apical and 2 outer setae/spines. P6 unknown in both sexes. TYPE AND ONLY SPECIES. Laophonte spelaea Chappuis, 1938 = Troglophonte spelaea (Chappuis, 1938) comb. nov. ETYMOLOGY. The generic name is derived from the Greek trogle, meaning hole, and refers to the stygobiont life style of the type species. Gender: feminine. 99 MATERIALEXAMINED. None. Chappuis’ (1938) material no longer exists and the species has not been recorded again since its original description. PHYLOGENETIC ANALYSIS Taxa and characters The analysis was executed at species level in order to test the monophyly of the genus Esola and its relationships to both Mourephonte and Archilaophonte. Onychocamptus and the cornuta- group of the genus Laophonte were also included as separate taxa in the analysis on the basis of their ancestral P3 endopod sexual dimorphism. The highly advanced genus Folioquinpes Fiers & Rutledge, although having been positively identified as the sistergroup of Onychocamptus (Lee & Huys, 1999), was excluded from the analysis. The residual Laophontidae were replaced by their hypothetical ancestor (Table 4: Other Laophontidae) which was constructed by combining the most plesiomorphic state encountered for each character. Table 2. Laophontidae with 3 inner setae [3(1-2)(0-1) setation pattern] on P3 enp-2 9. P2 exp enp Laophonte Group I OM23 1.220 Laophonte adduensis 0.1.122 1.220 Laophonte ciliata 0.1.122 1.220 Onychocamptus Group I 0.1.123 0.220 Onychocamptus besnardi O23 0.220 Onychocamptus anomalus 0.1.123 0.220 Onychocamptus taifensis 0.1.123 0.120 Onychocamptus krusensterni 0.1.123 0.220 Laophonte galapagoensis 0.1.123 0.220 Laophonte confusa 0.1.123 0.220 Laophonte Group II 0.1.123 0.220 Laophonte lignosa 0.1.123 0.220 Laophonte setosa 0.1.123 0.220 Laophonte elongata Oaln23 0.220 Laophonte Group III 0.1.123 0.220 Laophonte nordgaardi O23 0.120 Bathylaophonte spp. 0.1.123 0.220 Microlaophonte trisetosa 0.1.122 0.220 Pseudonychocamptus carthyi 0.1.123 0.220 Paralaophonte Group I 0.1.123 0.220 Paralaophonte panamensis 0.1.123 0.220 Paralaophonte Group II 0.1.123 0.220 Paralaophonte tenera 0.1.123 0.220 Paralaophonte innae 0.1.123 0.220 Paralaophonte aenigmaticum 0.1.123 0.220 Heterolaophonte campbelliensis 0.1.123 0.220 Heterolaophonte Group I 0.1.123 0.220 Heterolaophonte Group II 0.1.123 0.220 Heterolaophonte manifera 0.1.123 0.220 Heterolaophonte hamata ONE123 0.220 Heterolaophonte minuta 0.1.123 0.220 Paronychocamptus spp. 0.1.123 0.1—220 Asellopsis hispida 0.1.123 0.220 Asellopsis duboscqui 0.1.122 0.120 Folioquinpes chathamensis 0.1.123 0.220 P3 P4 exp enp 2 enp 3 exp enp 0.1.223 1.321 1.1.220 0.1.223 1.221 0.1.223 S21 1.1.220 0.1.223 1.221 0.1.222 1.321 1.1.220 0.1.222 1.221 0.1.123 0.321 0.1.220 0.1.123 0.111 0.1.123 0.321 0.1.220 0.1.022 0.111 0.1.123 0.321 0.1.220 0.1.122 0.111 0.1.123 0.321 0.1.220 0.1.123 0.111 0.1.123 0.321 0.1.220 0.1.122 0.111 0.1.223 0.321 0.0.220 0.1.223 1.121 0.1.223 0.321 0.0.220 0.1.223 1.120 0.1.223 0.321 0.0.220 0.1.223 0.221 0.1.223 0.321 0.0.220 0.1.223 0.121 0.1.223 0.321 0.0.220 0.1.223 0.111 0.1.223 0.321 0.0.220 0.1.123 0.111 0.1.123 0.321 0.0.220 0.1.123 0.111 0.1.123 0.311 0.0.210 0.0.023 0.111 0.1.223 0.321 0.0.220 0.1.223 0.221 0.1.222 0.321 0.220 0.1.222 0.221 0.1.223 1.321 0.220 0.1.223 ii 0.1.223 0.321 0.0.220 0.1.223 0.121 0.1.223 0.321 0.0.220 0.1.222 0.121 0.1.123 0.321 0.0.220 0.1.123 0.121 0.1.123 0.321 0.0.120 0.1.123 0.121 0.1.223 0.320 0.320 0.1.223 0.121 0.1.123 0.320 0.320 0.1.022 0.120 0.1.223 0.321 0.0.220 0.1.223 0.121 0.1.123 0.321 0.0.220 0.1.022 0.121 0.1.123 0.321 0.220 0.1.123 0.121 0.1.123 0.321 0.220 ONeI22 0.121 0.1.123 0.321 0.220 0.1.022 0.121 0.1.123 0.321 0.220 0.0.022 0.121 0.1.223 0.321 0.0.220 0.1.122 0.111 0.1.223 0.321 0.0.220 0.1.223 0.111 0.1.222 0.321 0.0.220 0.1.222 0.111 0.1.123 0.321 0.321 0.1.123 0.120 Laophonte: Group I = cornuta, expansa, plana; Group II = inornata, parvula, serrata; Group III = adamsiae, thoracica. Onychocamptus: Group | = mohammed, bengalensis, vitiospinulosa Paralaophonte: Group I = asellopsiformis, brevirostris, congenera, dieuzeidei, gurneyi, hyperborea, lacerdai, majae, meinerti, ormieresi, pacifica, pilosoma, royi; Group II = karmensis, lunata, spitzbergensis, zimmeri. Heterolaophonte: Group I = discophora, variabilis,; Group Il = murmanica, stromi, uncinata. 100 Characters used in the analysis are listed in Table 3. Apomorphic character states are explained inside square brackets using the multistate system. The scores for each character and taxon are compiled in matrix format in Table 4. A question mark indicates missing data, either because the appendage or structure is unknown in that species (certain sexually dimorphic characters could not be scored because only one sex is known) or because it was impossible to score the character accurately due to incompleteness or the lack of detail in the original descriptions. Esola typhlops pontoica, E. longicauda vat. sensu Vervoort (1964) and the unnamed forms of E. longicauda identified by Noodt (1955) and Wells & Rao (1987) were excluded from the analysis because of their questionable status. Huys & Boxshall’s (1991) study of ordinal copepod phylogeny demonstrated that oligomerization was the dominant trend of evolu- tionary transformation within the Copepoda. Armature counts used in this analysis were scored according to this overall polarisation mode. Most characters in Table 3 are self-explanatory but additional notes are provided for the following: Integumental pores (characters I-4) The conspicuous cup-shaped integumental pores on the cephalothorax and genital (double-)somite have remained unnoticed Table 3. Characters used in phylogenetic analysis. Apomorphic character states are referred to in square brackets. 1 Paired anterodorsal cup-shaped pores on cephalothorax absent [present] 2 Paired anteroventral cup-shaped pores on cephalothorax absent [present] 3 Paired cup-shaped pores on genital double-somite of 9 and genital somite of ¢ absent [present] 4 Caudal rami without large pore medially or ventrally [present] 5 Cephalothorax without transverse spinular row dorsally [present] 6 Caudal rami not sexually dimorphic [modified in 9] 7 Antennule ?7-segmented [6-segmented; failure in separation of segments 6 and 7] 8 Antennule d with 3 segments distal to geniculation [with 2 segments: segments 7 and 8 fused] 9 Aesthetasc of segment 4 in 2(and segment 5 in d) fused basally to seta [fused to two setae forming trifid compound element] 10 Antennule segment 1 without processes in 9/d [with 3 spinous processes along posterior margin] 11 Antennule segment 2 with large spinous process arising from posterior margin in 9/d [absent] 12 Antennule segment 5 of 3d without anterior cylindrical process (bearing large spine) [present] 13 Labrum without conspicuous ornamentation on anterior surface [with overlapping scales distally and dense pattern of fine spinules proximally] 14 Maxillulary endopod represented by 3 setae [2 setae, outermost seta lost] 15 Maxillipedal syncoxa with 3 setae [state 1: 2 setae, proximal seta lost; state 2: 1 seta] 16 Pl exopod 3-segmented [2-segmented; exp-2 and -3 fused] 17 Pl exopod 2-segmented, exp-2 with 3 outer spines and 2 apical geniculate setae [exp-2 with 2 outer spines and 2 apical geniculate setae] 18 P1 enp-1 with inner seta [absent] 19 P2 enp-2 with outer spine/seta [absent] 20 P3 endopod d 3-segmented [2-segmented; neotenic development] il P3 enp-2 6 with inner seta [absent] 22 P5 baseoendopod @ with 5 setae [state 1: with 4 setae, middle inner seta lost; state 2: with 3 setae] 23) P5 baseoendopod ¢ with 2 setae [setae absent] 24 P5 basoendopod 9/¢ without distinct setophore for outer basal seta [basal seta positioned on long cylindrical setophore] 25 P5 exopod 2 with all outer setae arranged around margin [proximal 2 outer setae displaced with overlapping insertion sites] R. HUYS AND W. LEE in previous descriptions except for Vervoort (1962, 1964) who briefly described the anterodorsal pores in C. bulligera and E. vervoorti and suspected them to be eyes. Various authors (e.g. Jakobi, 1953; Hamond, 1969; Mielke, 1981, 1997) have uninten- tionally figured the modified pores on the caudal rami, however, incorrect interpretation of the internal chitinized walls of the ducts as external ridges (“Chitinleiste’) made them fail to recognize these structures as true pores. Huys (1990b) pointed out that the trans- formed cup-shaped pores in Esola are not serially homologous with the pleural glands of the Adenopleurellidae and consequently can- not serve as a basis for phylogenetic affinity. With the exception of Archilaophonte and the typhlops-group of Esola all other esolinids appear to exhibit a propensity for developing modified secretory pores. The functional correlation between pores of different body- regions is unknown and in view of their positional disparity and structural differences it is unlikely that their expression is controlled by a single gene. We postulate that the cup-shaped pore type evolved from a surface precursor pore by major integumental invagination and secondary development of setular extensions. These marginal extensions either protect the depression or (more likely) maintain the secrete bolus in close contact to the body wall. The degree of invagination is obviously morphologically constrained and this is particularly the case in swimming leg segments which are typically depressed along the antero-posterior body axis. Although the ‘trap- ping basket’ seta on the P4 endopod of C. bulligera represents a radically divergent modification, it can be viewed as an external analogue of the internal cup-shaped pore which developed in response to this constraint. The tube-pore, which is also found in most other esolinids, is enclosed by the long setules arising from the proximally dilated distal inner seta (Fig. 25E-F) which hold the secrete bolus in position. Since there are no differences in pore pattern between the sexes a possible role in mate recognition is considered unlikely. Huys (1992) demonstrated that in the interstitial Leptastacidae the mucopolysaccharid strands produced by the caudal ramus glands are intimately involved in mucus-trap feeding. We suggest that in esolinids the secretory products discharged by the cup-shaped pores perform a similar role in trophic gardening. It should be noted that the caudal ramus pores located near the insertion sites of setae I-III in E. typhlops (Fig. 31C—D) are not homologous to the large slit-like pores found in Esola and Mourephonte. Caudal ramus sexual dimorphism (character 6) Females of Esola typically have bulbous caudal rami, displaying a variety of swelling medially, ventrally and/or dorsally. Although the secondary expansion appears to be correlated with the size of the transformed pores, it is decoupled here from character 4 (presence of caudal ramus pores) and scored separately. This is justified by the absence of caudal ramus sexual dimorphism in A. hirsuta despite the presence of modified pores in both sexes. Setal fusion on antennules (character 9) In most esolinids (except Archilaophonte) the proximal aesthetasc (on segment 4 in 9, segment 5 in <) is fused at the base to 2 setae. This trifid compound element is a unique character in the Harpacticoida. Antennulary processes (characters 10-12) Within the esolinid grouping a spinous process along the posterior margin of the second antennulary segment (character 11) is present only in Archilaophonte. This is not an autapomorphy for the genus but considered a retention of the ancestral state, based on outgroup comparison with the remaining families of the Laophontoidea (Huys, 1990a; Huys & Lee, 1999). The presence of auxiliary processes BASAL LAOPHONTID EVOLUTION along the posterior margin of the first segment (character 10) is a unique feature displayed by the species related to E. longicauda. There are no equivalent structures known from other Laophontidae and consequently this feature should be regarded an evolutionary novelty for this species-group. In males of the same group the enlarged fifth segment has produced an anterior sub-cylindrical outgrowth bearing a stout modified spine (character 12). Minute outgrowths are found on the first segment of C. bulligera but these are not considered important enough to warrant a separate score. Maxillulary endopod armature (character 14) The maxillulary endopod typically bears 2 setae along the outer margin of the basis, representing the incorporated endopod. This condition is found in all esolinids while in several other laophontid genera the endopod is represented by a cluster of 3 setae (e.g. Langia, Quinquelaophonte: Mielke (1997)). A notable exception is Archilaophonte in which the outermost third seta is secondarily displaced to a more proximal position, i.e. at the base of the exopod. Consequently, character 14 is scored 0 for A. maxima despite the clearly derived positional pattern. Male P3 endopod segmentation (character 20) The P3 endopod in the males of A. typhlops and Esola sp. sensu Chislenko (1967) is 2-segmented as in the female. The outer spine forming the apophysis in the males of other esolinids has remained largely unmodified except for reduction in size and basal fusion. This virtual absence of sexual dimorphism is considered the apomorphic state on the basis of ontogenetic evidence. Huys (1990a) demonstrated that the typical 3-segmented condition is accom- plished at the final moult by secondary subdivision of the distal segment and allometric growth of the spinous apophysis. The atypical pattern in A. typhlops, resembling the condition of a copepodid V stage, is interpreted here as the result of neoteny, i.e. the decrease in developmental rate has delayed the segmentation beyond the final moult. Male P3 endopod armature (character 21) The modification of the male P3 endopod in esolinids has no effect on the number of armature elements. In particular, the homologue of the outer spine in the female is transformed into a spinous process or apophysis arising from the middle segment in the male (but see character 20), and the proximal inner seta on enp-2 of the 2-segmented endopod in the female is retained on enp-2 of the 3-segmented endopod in the male [typically 1.1.220 pattern]. The presence of the latter seta in males is a particularly conservative character in primitive laophontids, however, outside the esolinid grouping it is found only in Onychocamptus and one species group of the genus Laophonte. The fate of this seta during male development can only be traced in Laophontidae displaying the full complement of 3 inner setae in the female enp-2. In these species (Table 2) the endopodal armature pattern is most commonly [0-1.321] but can also be [0.311] in Laophonte nordgaardi Sars or [0.320] in some species of Paralaophonte Lang. Except for 6 species of Laophonte and all species of Onychocamptus, the proximal inner seta is consistently lost in the male, resulting in a 0.0.220 pattern. The only exceptions with 3 inner setae in the male are those that have lost sexual dimorphism altogether (Folioquinpes, Paralaophonte innae Chislenko, P. aenigmaticum Wells, Hicks & Coull). Vervoort (1964) reported a very long inner seta on the middle segment of Paralaophonte pilosoma but re-examination of the holotype (USNM reg. no. 109763) has proven this to be erroneous (Fig. 28E). The loss of the proximal inner seta in the male is an apomorphy of pivotal importance in laophontid evolution since it unifies nearly 101 95% of all species. Since many genera have only 0, 1 or 2 inner setae in the female we have assumed that they are descendants from an ancestral stock which displayed the 3-setae condition in the female but lost the proximal one in the male. Female P5 exopod armature (character 25) Female esolinids can be readily identified by the setal arrangement around the outer margin of the PS exopod. The two proximal setae are displaced so that their respective insertion sites have become superimposed on one another. Lang (1948) and Willen (1995) pointed out that a similar displacement also occurs in Laophonte parvula Sars (arrowed in Fig. 25G), however, we concur with the latter author that this is the product of convergence. Results Analysis was performed with PAUP 3.1.1 (Swofford, 1993) using the exact Branch and Bound algorithm (Hendy & Penny, 1982) that is guaranteed to find all most parsimonious trees (MPTs), with all characters set irreversible up and arbitrary solutions (zero- length branches) suppressed. Analysis of the complete data (Table 4) produced 84 MPTs with tree length 40 and consistency index 0.675. The strict component consensus tree is illustrated in Fig. 32 and has a slightly longer length (42) and lower consistency index (0.643). Relationships within the crown-group Esola are poorly resolved, however construction of the majority-rule com- ponent consensus tree revealed an additional group (bulbifera-canalis-profunda). This boreo-mediterranean majority component appears in 48 (57%) of the trees. A. longiremis, A. hamondi and Esola sp. sensu Chislenko (1967) all have different combinations of missing entries, however each is also a potential taxonomic equivalent of A. typhlops (Table 4) and can therefore be safely deleted (Wilkinson, 1995). Safe taxonomic reduction of these taxa reduces the number of MPTs to 14 but does not alter tree length or consistency index. The strict component consensus (Fig. 32) reveals a strongly supported basal dichotomy which divides the Laophontidae into two major clades. In order to reflect the robustness of this dichotomy, subfamilial rank is attributed to the two corresponding lineages. The Esolinae subfam. nov. includes Archilaophonte, Mourephonte and all species previously assigned to Esola. It is supported by male antennulary segmentation (character 8) and the female P5 exopodal setation pattern (character 25). The primitive position of Archilaophonte conjectured by Willen (1995) is confirmed. The genus represents the first offshoot in the evolution of the Esolinae and is tentatively defined by the following suite of autapomorphies: (a) 6-segmented @ antennule (segment 6 compound), (b) 2-segmented P1 exopod (fusion exp-2 and -3), (c) Pl enp-2 secondarily elongated P2, (d) P2 enp-2 with only 1| inner seta, (e) P3 enp-2 d with very long sigmoid apophysis, (f) P5 exopod 6 with 4 setae (loss of proximal inner seta), and (g) extremely elongation of caudal rami. In addition, the maxillulary palp shows a peculiar setal arrangement along the outer margin with 1 seta positioned at the base of the bisetose exopod. Outgroup comparison with the Normanellidae indicates that this seta is of endopodal origin and must therefore have been secondarily displaced to a more proximal position. The basal position of Archilaophonte is sup- ported by the presence of (a) a spinous process on the posterior margin of the 2nd antennulary segment, (b) maxillulary endopod represented by 3 setae, (c) 3 setae on the maxillipedal syncoxa, and (d) the well developed ¢ P5 endopodal lobe bearing 2 long setae. The apomorphic alternatives of these characters (Table 3) in con- junction with the formation of a trifid compound element on 102 ESOLINAE subfam. nov. 3 2 Se a ee ee a eee mS Xx 2 2 Sy Re: ie) — x Sg = S & = = se U4y 46S Mimo Se RS Sess g ss 8 wk eon ee ee a TOSS eS S Kei = 8) SI Ss Noy ral wt Be S e : nm N N wm N So | Applanola gen. nov. hirsuta 19 R. HUYS AND W. LEE LAOPHONTINAE ade 3 = =; : es °: > : 9 -#& (=) Sw nc bake = vo i—| ° ee : Ss : 8D go “ss Ss A oe 6s S jaa) = 8 8 > ) oo 0 YY OeH yy Oa YO yt OF OF or S'S SOO srs OO) Ot (1 SS) a SS oot ee ee a a a a ia) SBNN VW Ve Se Vee ee yee eee eR oO oF Ee SH OF KS KP OOK Coo r YE HS EHP - SS So Sore Ss OO SS SS SS SS SOOO Py py) Pe uy fer) Pet ee) eed ey SS Sis) Sem oQooc”*ooddqdcoocroooraqooqcond cooo7yrRrOowovrvvVFROoOwooooocno°o eoovoovonvrvvornrvnvyooocnoeoocoeo ON OF NER KH OOr OF OF eRe Ere yo COrOowr Re NR NIN NOR Re DOR RR Re RRO qooocorococorocorcrdqocoqocooo°qc”oo a oe el el further elaborated in both Mourephonte and the residual species of Esola by the development of an accessory pair of anteroventral pores on the cephalothorax (character 2) and of ventral or medial pores on the caudal rami (character 4). Finally, the lateral pores on the genital (double-)somite (character 3) evolved not until after the divergence of Mourephonte. The genus Esola is redefined here to encompass the terminal polychotomy containing the type species E. longicauda and 7 other species (Fig. 32). This strongly supported, cosmopolitan crown- group is characterized by distinctive labral ornamentation, caudal ramus sexual dimorphism, formation of 3 spinous processes on the first antennulary segment and modification of segment 5 in the male antennule E. hirsuta is the only species that shares genital cup- shapes pores with this clade, however it is excluded from Esola and placed in a monotypic genus Applanola on account of the following autapomorphies: (1) dorsoventrally depressed body morphology, (2) elongation of mandibular palp, (3) modification of P1 endopod, (4) exopodal sexual dimorphism of P2—P3, (5) loss of outer spine on P2 enp-2, and (6) strong reduction of the male sixth legs. The sexual dimorphism on the P2—P3 exopod is unique in the Esolinae. Al- though this character is globally homoplastic within the Laophontidae it can be informative locally (see Lee & Huys, 1999) and should not therefore be routinely ignored in phylogenetic analyses. The three remaining species, compacta, spelaea and bulligera, are identified as independent lineages splitting off successively between the basal Archesola clade and the terminal ((Mourephonte + Esola) + Applanola) clade. Corbulaseta gen. nov., accommodat- ing E. bulligera, is most closely related to the latter clade because of shared fusions in the female antennule (segments 6-7) and Pl exopod (exp-2 and -3). The modified distal inner seta forming a trapping-basket is a unique autapomorphy for this genus. The position of Troglophonte is tentative pending the confirmation of cup-shaped pores on the cephalothorax and of the armature patterns of P2 exopod and P5 in both sexes. The basal position of the genus Bathyesola is caused by its retention of the maximum number of setae on the female PS baseoendopod. The genus Mourephonte is radically divergent from other esolinids. The extreme development of the P1, the complete absence of the P2 endopod, the loss of the inner seta on the P2—P4 exopods and the wide separation of the apical setae on P4 enp-2 form a remarkable combination of autapomorphies which places it on a distinct evolu- tionary lineage, ruling out possible inclusion in the genus Esola under a broader concept. The residual laophontids, comprising 95% of the known species, are grouped in the subfamily Laophontinae. All 54 genera have lost the inner seta on P| enp-1 and the outer spine on P2 enp-2, and bear a maximum of 2 setae on the maxillipedal syncoxa (absence of proximal seta). With the exception of the genus Onychocamptus and the Laophonte cornuta-group all Laophontinae are characterized by the P3 endopod sexual dimorphism involving the loss of the proxi- mal inner seta of enp-2 (character 21). The isolated position of the cornuta-group (= Laophonte Group I + adduensis + ciliata: Table 2) testifies to the widely accepted polyphyletic status of the genus Laophonte and has major nomenclatural consequences because of its inclusion of the type species L. cornuta Philippi. Restriction of the generic concept to the cornuta-group will require the other 37 species of Laophonte to be re-allocated to other existing or new genera. This is a major task which can only be accomplished by sound phylogenetic analysis involving the remaining laophontinid genera. The sistergroup relationship between the cornuta-group and Onychocamptus depicted in Fig. 32 is not be taken as absolute since other advanced but closely related genera such as Folioquinpes have deliberately been omitted from the outgroup to the Esolinae. Al- though inclusion of these genera in future analyses may introduce additional basal nodes changing the relative position of Laophonte and Onychocamptus, we envisage that the latter will consistently show up as an early speciation event predating the evolution of the other Laophontinae. Subfamilial division ESOLINAE subfam. nov. Rostrum delimited at base by surface suture; antennule 2 6- or 7- segmented, usually without spinous process on segment 2 but frequently with processes on segment 1; 7-segmented and haplocerate or subchirocerate in 6, with only 2 segments distal to geniculation; 104 proximal aesthetasc typically fused to 2 setae (except Archilao- phonte). Antennary exopod with 4 well developed setae. Mandible typically biramous (except Applanola and Mourephonte). Maxilla with 3 endites on syncoxa. Maxilliped with 2—3 setae on syncoxa. P1 with 2- or 3-segmented exopod, retaining full complement of setae (0.0.022 or 0.023); enp-1 occasionally with inner seta. P2 enp- 2 with outer spine (except Applanola) or entire P2 endopod absent (Mourephonte). P3 endopod ¢ retaining proximal inner seta of 2 enp-2 (except for Troglophonte where it is lost in both sexes). Armature formula as follows: Exopod Endopod p2 0.1.123 0-1.(1—2)2(0-1) or absent P3 0.1.(1—2)23 0-1.321 P4 0.1.(1=2)23 0-1.221 P5 @ with separate rami; exopod elongate, with 6 setae/spines; proximal two setae along outer margin with superimposed insertion sites; baseoendopod trapezoid, slightly developed, with 4—5 setae/ spines. P5 d without endopodal lobe (except for Archilaophonte, bearing 2 long setae), no endopodal armature; exopod 5 setae/ spines. Typically with cup-shaped transformed pores on cephalothorax, genital (double-)somite, and/or caudal rami. TYPE GENUS. Esola Edwards, 1891 OTHER GENERA. Mourephonte Jakobi, 1953; Archilaophonte Willen, 1995; Applanola gen. nov.; Archesola gen. noy.; Bathyesola gen. nov.; Corbulaseta gen. nov.; Troglophonte gen. nov. Laophontinae T. Scott, 1905 Antennule d with up to 3 segments distal to geniculation; proximal aesthetasc fused to 1 seta. Mandible typically uniramous. Maxilliped with maximum 2 setae on syncoxa. P| enp-1 without inner seta. P2 enp-2 without outer spine. P3 endopod ¢ typically not retaining proximal inner seta of 2enp-2 (except for Laophonte cornuta-group and Onychocamptus). Proximal outer setae of 2 P5 exopod with distinctly separated insertion sites. Cup-shaped transformed pores on cephalothorax, genital (double-) somite, and/or caudal rami never present. TYPE GENUS. Laophonte Philippi, 1840 OTHER GENERA. Fifty-five; see Lang (1948), Bodin (1997), George (1997) and Lee & Huys (1999) for complete list. KEY TO GENERA OF ESOLINAE [PR 2endopodkabsemtgeceer secre ee Mourephonte Jakobi, 1953. PAendopodspresenty2-Se SMe Med meecsee esas eee eee De 2. Antennulary segment 2 with large spinous process along anterior margin; P2 enp-2 with 1 inner seta; P5 baseoendopod d with 2 long ISVS Spo acen aa ee ses ssoe soe eSoeeeceeuassendebocoocececaee Archilaophonte Willen, 1995. Antennulary segment 2 without spinous process along anterior margin; P2 enp-2 with 2 inner setae; P5 baseoendopod ¢ without setae ....... 3: 3. Antennule 2 6-segmented; P1 exopod 2-segmented; caudal rami with Mecialormven thal anno chiki dap OLesmeeewes ee see eee eee ene eee eee 4. Antennule 2°7-segmented; P1 exopod 3-segmented; caudal rami with- QULBISUCIY POLES acess vest es eo eonicees cae sms asctecepae cere conn aAcattucs seeeeeeaees heen 6. 4. Body short, dorsoventrally flattened; P2 enp-2 outer spine absent; P3 R. HUYS AND W. LEE exopod ¢ strongly modified 00.0.0... Applanola gen. nov. Body elongate, sub-cylindrical; P2 enp-2 outer spine present; P3 exopod G MOt tmOdified ez. c, Be ececkeeacecee eo ose scene seed se a: 5. Antennulary segment | with 3 spinous processes along posterior mar- gin; distal inner seta of P4 endopod not transformed; caudal rami 2 modified, with bulbous swelling dorsally, ventrally and medially ....... ss hevanstehs Seth ts «ons dane dR ences aicasessaaiitenessseneae ani Esola Edwards, 1891. Antennulary segment | without distinct spinous processes; distal inner seta of P4 endopod transformed; caudal rami not sexually dimorphic, Gylim det Calls. 5 lake cs eae Mees seete casks een eee Corbulaseta gen. nov. 6. P3—P4 exp-3 with 1 inner seta; P3—P4 enp-1 without inner seta ......... SEE Er ee ee 2 ae eee Bathyesola gen. nov. P3—P4 exp-3 with 2 inner setae *; P3—P4 enp-1 with inner seta....... 7. 7. P3 enp-2 with 3 inner setae; P3 endopod d 2-segmented; P5 baseoendopod with long articulating setophore in both sexes ............. seus nstnn jedan ato sugbhes bevdteocaeettneds voscunecentettte stot rence weer Archesola gen. nov. P3 enp-2 with 2 inner setae; P3 endopod ¢ 3-segmented; P5 baseoendopod of both sexes without articulating setophore................ sdaacdibiaigia snasat Biaesov satel niveasees tosvOueciede eect Troglophonte gen. nov. * Note that Chappuis’ (1938) setal formula of P3 exp-3 can also be interpreted as 123, implying the presence of only 1 inner seta. ECOLOGICAL RADIATION OF ESOLINAE Although none of the 18 species can be considered as truly cosmo- politan, the subfamily as a whole occurs in all oceanic basins, including the Antarctic Ocean. Superimposing habitat utilization upon the phylogeny presented in Fig. 32 reveals an interesting but complex ecological radiation pattern. Esolinae are essentially shal- low water inhabitants, however, the variety of additional habitats exploited by this lineage is startling for its small number of known species. Considered against the background of the overwhelming evolutionary success of their sister-lineage Laophontinae, esolinids can be viewed as relicts of a formerly diverse group. Lee & Huys (1999) reviewed published deepwater records of Laophontidae and regarded the colonization of the deep sea by this family as remarkably unsuccessful. There is no single lineage containing all deepwater forms, and the three exclusively bathyal genera in the Laophontinae, Cornylaophonte Willen, Weddellao- phonte Willen and Bathylaophonte Lee & Huys can be considered as independent colonists of this habitat. Colonization of the deep sea by the Esolinae follows a similarly erratic trend with early attempts by the monotypic genera Archilaophonte in the Antarctic and Bathyesola in the western Pacific. Within the genus Esola, E. profunda repres- ents a third, secondary deepwater invasion derived from a shallow water inhabiting ancestral stock (Fig. 32). According to Pesce (1985) and Rouch (1986) the genus Troglophonte is likely to be derived from a marine ancestor stranded during the lowering of sea level during the Tertiary. It is highly endemic to freshwater lenses in several Apulian caves in southern Italy (Chappuis, 1938). These caves are separated from the littoral zone by macroporous karstic rock and exhibit a detectable tidal current which appears insufficient to ensure substantial mixing of the water inside the caves. The strong stratification with freshwater lenses overlying the poorly oxygenated deeper layers has clearly prevented the establishment of a diverse marine benthic fauna. Rather than considering Troglophonte a Tethyan relict, its present restricted distribution can also be regarded as a relatively recent landward habitat range extension from a primarily shallow-subtidally residing ancestral stock. Although some Laophontidae are regularly BASAL LAOPHONTID EVOLUTION found in salt-marsh and mudflat habitats within river estuaries (Noodt, 1957; Barnett, 1968; Bodin, 1976) or in brackish lagoons (Heip, 1969; Hamond, 1972), tolerance to oligohalinity may have appeared convergently only twice in the family. Both colonization events presumably occurred early in the evolution of the family (Fig. 32), however their nature is fundamentally different. The evolution- ary success of the Troglophonte lineage has clearly remained limited, both in dispersal and speciation. It can be considered as a freshwater incursion without further radiation or diversification. The second invasion of low salinity environments is cosmopolitan in scope and probably of Tethyan origin, containing the genera Onychocamptus Daday and Folioquinpes Fiers & Rutledge (Lee & Huys, 1999). Little is known about the possible dispersal of Laophontidae in marine caves. Pesta (1959) reported E. rosei from a submarine cave near Naples and several unidentified Laophontidae were recorded by Huys (1996) from the anchialine Walsingham Cave on Bermuda. Examination of samples from Caye Chapel Cave in Belize, the type locality of the recently discovered family Novocriniidae (Huys & Iliffe, 1998), resulted in the discovery of a single male belonging to a new genus of Esolinae. The new genus has several characters in common with Archilaophonte such as the presence of a spinous process on the second antennulary segment, the displacement of the outermost endopodal seta on the maxillule, the presence of 3 setae on the maxillipedal syncoxa, Pl with 2-segmented exopod and elongate enp-2, P2 enp-2 with only | inner seta and presence of a very long apophysis on P3 endopod. Phylogenetic analysis identi- fied the Belize genus unambiguously as the sistergroup of Archilaophonte, suggesting the evolution of an independent cavernicolous lineage in the western Atlantic. The genus Archesola is exclusively boreo-arctic in distribution and restricted to the Atlantic basin, with a single known outlier from the Black Sea (Por, 1959). Its southernmost limit based on reliable records is Norfolk (England), however, confirmation of the doubtful records of A. longiremis from the south coast of England (Wells, 1961, 1963, 1970) and North Carolina (Coull, 1971) may extend this limit further southward. The genus occurs primarily at higher lati- tudes, showing limited dispersal in Arctic waters such as the White Sea (Brotskaya, 1961; Chislenko, 1967). It is suggested that the strongly discontinuous, bipolar distribution of the two basal clades, with Archesola restricted to northern Europe and Archilaophonte to the Antarctic, indicates a wider, perhaps continuous, horizontal zonation of primitive stenothermal esolinids at greater depths. This trend of ‘Equatorial Submergence’ appears to be supported by the discovery of Bathyesola in the deep tropical western Pacific, the first lineage to diverge after Archesola (Fig. 32). A major event in the evolution of the Esolinae was the episode of rapid speciation within the genus Esola. This event is revealed as a polychotomy in the cladogram (Fig. 32) although it is clear that the low resolution is partly attributable to the abundance of missing entries for several taxa which are known from one sex only (Table 3). Many of these species are small-sized (Table 1) and adapted to a mesopsammic life-style in shallow subtidal localities and sandy beaches, while others are frequently found associated with algal substrates. Results show that only a fraction of the species is known. Although the genus assumes a cosmopolitan distribution it is pre- dominantly restricted to the circum-tropical belt. This zone coincides with the former Tethyan seaway separating the northern and south- ern continents, which continued into Palaeogene times with free marine continuity along its length not being interrupted until the beginning of the Neogene. One Pacific-Caribbean subgroup, com- prising E. longicauda, E. galapagoensis and Esola sp. (Fig. 32), probably originated from an ancestral stock in the western Pacific. From there, eastward dispersal was greatly influenced by tectonic 105 plate movement, particularly the formation of the Caribbean plate at the beginning of the Oligocene. This was established by decoupling of the eastward protruding tongue of the East Pacific plate, causing the formation of a subduction zone along what is now the western coast of southern Central America, and the subsequent westward motion of North and South America past a nearly stationary Carib- bean plate (Malfait & Dinkelman, 1972; Coney, 1982). The entry of the ancestor of E. longicauda into the Caribbean must have preceded the closing of the Panama land bridge approximately 3.1—3.5 Ma (Keigwin, 1978). Applanola displays amore disjunct distribution than its sistergroup Esola, provided that Pesta’s (1916) record from the Gulf of Guinea is correct. The dorsoventrally depressed body, robust maxillipeds and powerful Pl endopod indicate that A. hirsuta may be loosely associated with invertebrate hosts. Thompson & A. Scott (1903) obtained the species from washings of pearl oysters and other dredged invertebrates but did not present any firm evidence for a clear association. ACKNOWLEDGEMENTS. Dr Danielle Defaye (Muséum National d’ Histoire Naturelle, Paris), Dr Chad T. Walter (National Museum of Natural History, Washington D.C.), Karen Gowlett-Holmes (South Australian Museum) and Dr Mark Holmes (National Museum of Ireland) kindly made type and other material available. Dr Richard Hamond (Norfolk, U.K.) provided us with specimens of E. bulbifera and A. typhlops. The material of Bathyesola compacta was collected during the STARMER II expedition to the Fiji Basin (chief scientist: L. Laubier). One of us (W.L.) acknowledges financial support from the Korea Research Foundation provided for the programme year 1997. REFERENCES Alheit, J. & Scheibel, W. 1982. Benthic harpacticoids as a food source for fish. Marine Biology, Berlin 70(2): 141-147. Barnett, P.R.O. 1968. Distribution and ecology of harpacticoid copepods of an intertidal mudflat. /nternationale Revue der gesammten Hydrobiologie 53: 177-209. Bodin, P. 1976. Les Copépodes Harpacticoides (Crustacea) des cétes Charentaises (Atlantique). Données écologiques et biologiques sur les espéces principales. Bulle- tin du Muséum national d’ Histoire naturelle, Paris (3)353 (= Ecologie générale 29): 1-45. . 1997. Catalogue of the new marine harpacticoid copepods. Studiedocumenten van het K.B.I.N. 89: 1-304. Brian, A. 1928a. Descrizione di specie nuove 0 poco conosciute di Copepodi bentonici del mare Egeo. (Nota preliminare). Bollettino dei Musei di Zoologia e Anatomia comparata della R. Universita di Genova (2)7(18): 1-37. . 1928b. I Copepodi bentonici marini. Archivio zoologico italiano 12: 293-343. Brotskaya, V.A. 1961. 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Archilaophonte maxima gen. n., spec. n., a new taxon of the Laophontidae (Copepoda, Harpacticoida) from the high Antarctic (Weddell Sea). Hydrobiologia 302: 241-255. ——. 1996. Two new genera of Laophontidae (Copepoda: Hapacticoida) from the high Antarctic Weddell Sea. Journal of natural History 30: 1297-1327. Willey, A. 1935. Harpacticoid Copepoda from Bermuda. — Part II. Annals and Maga- zine of natural History (10)15: 50-100. Seren d ai iene on ty ae a" —— ow Mes ty ) GA ew xe? .oxrd ik Try 0) vase bere - f ly A alate ll Nandi IE SS Fee ee een ee ey is. eee ep mat its eae im ad etapa Yeo tor AT) walle? aay meete —— wel ene Gm PS tapi agree Rom tod A=eerige fae 1 tee - toll alae OD rm si citi ~ & Ut cee ahem, A a: ne AR ae ee pedbewttieahinemaceg tape “nae ah cape ag ee - 4 ae aren > cham etyt aps ee oath i re , en ee Se ee ha late a oo 0 Lore erage] iets Re! VG) ae 7 Tn &.4e8 a et atin > * tytn ite te eae) £0 Nd ae -_ - ie et ee ee Se [Nha Owes S Oy ee OS te efoto aDse.§ wee A IEOD Erart 6 OT lenient oacep) teen ‘ j : . 2 TE eget cme a, iinet au Aiea ear aie ye! omy tm a me| yas LD & Gaead ae @ sei e oO. ee oh ie Goat! /© AP ehh TS awl sO 5 eae “hos ee oe e _ é ‘ > 24 ni hg { P - « i » = és ~ — ~ = ¥ . - = te 0 «- es ' 4 pa a ee \ i rn ’ Se = = 7 - « ig i] = ve 7 : = » = se = — » — ts e . 2 o- ~ e ‘ _*. - = ~~ a. “i 4 — > - S ? 2 ‘ ‘ a —s GUIDE FOR AUTHORS Aims and scope. The Bulletin of the British Museum (Natural History) Zoology, was established specifically to accommodate manuscripts relevant to the Collections in the Department of Zool- ogy. It provides an outlet for the publication of taxonomic papers which, because of their length, prove difficult to publish elsewhere. 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Drawings should be in black on white stiff card with a line weight and lettering suitable for the same reduction throughout, ideally not more than 40%. After reduction the smallest lettering should be not less than 10 pt (3 mm). Tracing paper should ideally be avoided because of the possibility of shadows when scanned. All artwork must have bulletin, author and figure number included, outside of the image area, and must be free of pencil, glue or tape marks. PHOTOGRAPHS — All photographs should be prepared to the final size of reproduction, mounted upon stiff card and labelled with press-on lettering (eg Letraset). They can be mounted on white or black background; a black background must be evenly black all over; any background must be free of all pencil and glue marks within the image area. All figures should be numbered consecutively as a single series. Legends, brief and precise, must indicate scale and explain symbols and letters. Photos, when components of figure- plates should be abutted, trimmed as regular rectangles or close trimmed up to edge of specimen. Joins etc. can be removed at the scanning stage but at extra cost. Cropping instructions, if any, should be indicated on an overlay or marked on a photocopy of the figure. SIZE — Maximum size of artwork for use of flatbed scanners is A3. Larger artwork has to be reduced photographically prior to scan- ning, therefore adding to expense. Symbols in text. Male and female symbols within the text should be flagged within curly brackets to enable setter to do a swift global search. Reprints. 25 reprints will be provided free of charge per paper. Orders for additional reprints can be submitted to the publisher on the form provided with the proofs. Later orders cannot be accepted. CONTENTS 1 Generic concepts in the Clymnestridae (Copepoda, Harpacticoida): revision and revival R. Huys and S. Conroy-Dalton 49 Basal resolution of laophontid phylogeny and the paraphyly of Esola Edwards R. Huys and W. Lee Sulletin of The Natural History Museum ZOOLOGY SERIES Vol. 66, No. 1, June 2000