taxonID	type	description	language	source
03F487A0FFAFFFA7FFDCFF48FBAE973E.taxon	description	1. Homology of apophysis on the three-segmented P 3 endopod in the male. Although the derivation of this apophysis in the Cylindropsyllidae and at least the freshwater component of the Canthocamptidae is now fully understood and proven homologous (Martínez Arbizu & Moura, 1994; R. Huys & S. Conroy- Dalton, unpubl. data), it needs to be stressed that this modification is not a synapomorphy for the Canthocamptidae, as it is found in a wider group of families, including the Louriniidae, Huntemanniidae, Rhizothrichidae, and in all probability also the Cletodidae. Therefore, it cannot be used as evidence for the allocation of the Cylindropsyllinae to the Canthocamptidae. 2. Similarity in structure of the female genital field with a posteriorly displaced copulatory pore. The authors claim that this posterior position is displayed by all genera of the Canthocamptidae, and that they also observed this character in a number of cylindropsyllid genera such as Evansula, Cylindropsyllus Brady and Stenocaris Sars. In reality, the copulatory pore is not posteriorly displaced in the primitive genus Evansula (see below) or the more derived Stenocaris (R. Huys, pers. observ.), and does not represent the ancestral state for the family, but has evolved secondarily only in the Cylindropsyllus – Cylinula Coull lineage. Therefore, the similarity found in both families is a result of convergence. Moura & Pottek (1998) regarded the pore patterns on the female genital somite (in particular the pore clusters or ‘ sieves’) as an additional synapomorphy supporting a relationship between the Canthocamptinae and Cylindropsyllinae, but this evidence is not convincing. Pore triplets flanking the copulatory pore have been recorded in other families of the canthocamptoid complex, such as the Ancorabolidae (e. g. Conroy-Dalton, 2001), Cletodidae (e. g. Gee, 1994, 1999; Gee & Huys, 1996) and Huntemanniidae (Dahms & Pottek, 1992). In addition to their evolutionary labile nature within families, the phylogenetic significance of genital pore clusters is further limited by their presence in taxa that are not related to the Canthocamptidae and its allies, such as the Miraciidae (Huys & Böttger-Schnack, 1994) and Clytemnestridae (Huys & Conroy-Dalton, 2000). There is as yet no evidence questioning the monophyly of the Cylindropsyllidae. The unique sexual dimorphism on the male P 2 exopod, the structure of the P 4, and the composite seta V of the caudal ramus are distinctive synapomorphies for the family. Conversely, it is at present not clear at all what characters define the Canthocamptidae. Martínez Arbizu & Moura’s (1994) proposal to amalgamate the Cylindropsyllinae within the Canthocamptidae is also an unwise one for pragmatic reasons. Currently, three subfamilies exclusively containing freshwater genera are recognized in the Canthocamptidae: Canthocamptinae, Morariinae and Epactophaninae. Pesta (1932) lumped all the marine and brackish-water canthocamptids in the catch-all taxon ‘ Halocanthocamptinae’, a family group name that, for nomenclatural reasons, is unavailable as it was not based on the stem of an available genus (in fact the genus name Halocanthocamptus has never been proposed). Lang (1948) argued strongly against the recognition of this subfamily and finally abandoned it. The marine genera, however, have never been placed satisfactorily and are still floating in the system of the Canthocamptidae. This state of affairs has inspired other people to include even more marine genera in this family and, in one instance, an entire subfamily was transferred (Por, 1986). The Canthocamptidae, once a morphologically homogeneous, primarily freshwater-inhabiting family, currently contains 55 genera, 18 of which are found in fully marine or estuarine habitats (Boxshall & Halsey, 2004; Karanovic, 2004; Karaytu ğ & Huys, 2004). Paradoxically, subsuming the Cylindropsyllidae within this family would increase this number to 27, half of the total number of genera. Transferring a monophyletic group to a larger polyphyletic group without considering the possible implications this may have is generally regarded as bad practice. The systematic arrangement of the Canthocamptidae has not reached any stability and cannot be expected to do so within the foreseeable future. The Cylindropsyllinae, on the other hand, is a sharply delimited group with definite characteristics that deserves separate family rank. It is not even at all established that the Canthocamptidae, or any subgroup currently contained in it, is the sister taxon of the Cylindropsyllidae, as the characters that would support such a relationship are found in other taxa as well. It is the misapplication of the shared presence of very few characters that has caused the Canthocamptidae to become a repository for anything that could not fit the diagnoses of the other families contained in the second volume of Lang’s (1948) monograph. A lack of decisive facts permits every taxonomist to have their own opinion about the rank of groups. However, any objective observer cannot fail to notice the unfortunate consequences of amalgamating various cletodid genera with doubtful affiliations in a loosely defined group, assigning this group the rank of subfamily and accommodating it in the Canthocamptidae (Por, 1986). The concept of the ‘ Halocanthocamptinae’ introduced by Pesta (1932) deserves to be abolished, as Lang (1948) suggested. Yet from the recent addition of the marine Hemimesochrinae (Por, 1964), Cylindropsyllinae (Martínez Arbizu & Moura, 1994), and a group of genera formally designated ‘ Canthocamptidae incertae sedis ’ (Por, 1986), it seems that this unsatisfactory taxonomic practice is being kept alive by certain taxonomists. Pending a full-blown phylogenetic analysis of the canthocamptoid complex and the arrival of molecular sequence data, we retain the Cylindropsyllidae as a valid family comprising the following genera: Cylindropsyllus Brady, 1880; Evansula T. Scott, 1906 b; Stenocaris Sars, 1909; Cylinula Coull, 1971; Boreopontia Willems, 1981; Stenocaropsis Apostolov, 1982; Willemsia Huys & Conroy-Dalton, 1993; Navalonia Huys & Conroy-Dalton, 1993; and Selenopsyllus Moura & Pottek, 1998. Of these, the genus Cylindropsyllus is possibly paraphyletic and Stenocaris undoubtedly polyphyletic (R. Huys & S. Conroy- Dalton, unpubl. data).	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
03F487A0FFACFFA6FFD7F97FFC1E926A.taxon	diagnosis	Diagnosis: Cylindropsyllidae. Body slender, cylindrical. Rostrum elongate, defined at base. Genital doublesomite female without chitinous traces marking original segmentation. Copulatory pore large, semicircular, positioned slightly posterior to gonopores. Copulatory duct and paired anterior extensions very well chitinized. Caudal ramus of moderate length, conical; with seven setae; seta I vestigial; seta V with defined flexure zone between proximal styliform and distal flagellate parts, fused to seta IV; seta VI vestigial. Sexual dimorphism in antennule, P 2 – P 3 (exopod and endopod), P 5, P 6, ventral ornamentation of anal somite, and in genital segmentation. Occasionally, also in P 2 basis (slight), P 4 endopod and caudal rami. Antennule with segment 1 approximately as long as segment 2; segment 2 with nine setae; segment 6 with three setae in female; seven-segmented and with aesthetasc on segments 4 and 7 in female; indistinctly nine-segmented, haplocer and with aesthetasc on segments 5 and 9 in male; apical segment in both sexes with terminal acrothek consisting of two long setae fused basally to slender aesthetasc. Antenna with incompletely fused allobasis and bisetose, onesegmented exopod. Mandible with two-segmented, uniramous palp consisting of unisetose basis and endopod bearing one lateral and four distal setae. Maxillule with one claw and one seta on coxa, five setae plus one claw on basis; endopod and exopod represented by a small, trisetose segment and two setae, respectively. Maxillary endopod discrete, one-segmented, with four setae; proximal endite of syncoxa with three setae / spines. Maxillipeds well developed, subchelate, syncoxa with zero to one seta, endopod represented by strong claw. P 1 – P 4 with three-segmented exopods and one- (P 2 – P 4) or two-segmented (P 1) endopods. P 1 with outer and inner seta on basis; exopod with outer spine on middle segment and two geniculate setae plus two spines on distal segment; endopod prehensile, proximal segment longer than exopod, with inner seta being plumose proximally and pinnate distally (except E. arenicola), distal segment with two geniculate setae (or spines) and one setule. P 2 – P 4 bases with outer seta. P 3 endopod of female with two distal spines. Inner distal spine of P 3 – P 4 exp- 3 distinctly shorter than outer distal one. Inner setae of P 4 exp- 3 and endopod serrate. Armature formula of swimming legs: Exopod Endopod P 1 0.0.112 1.111 P 2 0.0.022 110 P 3 0.0.022 020 P 4 0.0.122 211 P 2 male with inner seta of exp- 3 transformed into claw; endopod with slight setal modifications. P 3 exopod male usually without or with reduced hyaline frill on exp- 1; exp- 3 with secretory pore and elongate outer distal seta. P 3 endopod male two- or three-segmented; enp- 1 minute or incorporated in middle segment; enp- 2 (enp- 1 if ramus two-segmented) with large apophysis and pinnate seta; enp- 3 (enp- 2 if two-segmented) with one short seta. P 5 baseoendopod and exopod completely fused, forming elongate plate in both sexes; with eight setae / spines in female; drawn out into spinous pinnate process and with six setae / spines in male. P 6 asymmetrical in male, with three setae each; represented by small operculum with three setules in female. Type species: Tetragoniceps incertus T. Scott, 1892 = Evansula incerta (Scott, 1892) [by original designation: Scott (1906 a)]. Other species: E. pygmaea (T. Scott, 1903 a); E. arenicola Nicholls, 1939; E. cumbraensis sp. nov.; E. spinosa sp. nov.; E. polaris sp. nov. Species incertae: Evansula (?) spec. sensu Noodt (1955 c).	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
03F487A0FFADFFA8FFC7FCCDFA1194A4.taxon	description	Original description: Scott (1892: 254 – 255, plate XII, figs 1 – 17). Additional descriptions: All subsequent redescriptions are unreliable, as they were based on another species (Sars, 1911; Wilson, 1932) or on an amalgam of several species of which E. incerta may or may not have been part of (Klie, 1929; Kunz, 1935; Scheibel, 1972). Type locality: Scotland, Firth of Forth, not specified by Scott (1892), but according to a later paper (Scott, 1906 b) the types were dredged off St. Monans. Material examined: (1) NHM, reg. nos. 44505 – 507: Firth of Forth, Scotland; 2 ♀♀, 1 ♂ in alcohol (labelled as ‘ cotypes’); coll. T. Scott, 9 September 1894; as part of Cannon A. M. Norman collection (1911.11.8); 1 ♀ (on 11 slides) and 1 ♂ (on seven slides) dissected. This vial also contained 1 ♂ of E. spinosa sp. nov. (see below); (2) Royal Museum of Scotland, Edinburgh, reg. no. 1955.63.115: Firth of Forth, coll. T. Scott, June 1901 (labelled Tetragoniceps incertus): vial empty. Redescription Female: Total body length: 695 – 710 µm (N = 2; mean = 702.5 µm). Body slender, cylindrical (Fig. 1 A, B), semitransparent, light brown; no distinct separation between prosome and urosome. Genital double-somite completely fused (Figs 1 B, C, 2 D, 6 A), original segmentation marked dorsally by paired anterior and posterior sensillae (Fig. 1 A). Anal somite only slightly longer than wide (57 × 60 µm), with three pairs of secretory pores laterally (Fig. 6 E); posterior margin with two short spinular rows on either side of ventral midline (Fig. 6 A, C). Anal operculum weakly developed, unarmed (Fig. 6 D). Caudal ramus conical (Fig. 6 A, C), length (measured along the outer margin) approximately 1.9 times the proximal width; dorsal surface without chitinous spur; with seven setae, setae I – VI in distal and seta VII in proximal half (Fig. 1 A); seta I diminutive; setae II – III long and bare; seta IV short and bare, approximately half length of styliform part of V (Fig. 6 C); seta V long, with distinct flexure zone between styliform part and long distal flagellate part, fused at base with vestigial seta IV; seta VI vestigial; seta VII tri-articulate at base and located along proximal inner margin; ventral surface with one simple and two tube-pores (Fig. 6 C), dorsal surface with one tube-pore. Rostrum elongate (Fig. 2 A, C), with parallel margins in proximal half, tapering distally; distinctly shorter than first antennulary segment; demarcated at base; base surrounded by area of flexible integument; with two long sensillae; subapical pore positioned midventrally. Antennule seven-segmented (Fig. 3 A). Segment 1 only slightly shorter than segment 2, with small sclerite around proximal posterior margin, with pattern of spinular rows on anterior surface, as illustrated in Figure 3 B; segment 2 longest, without secretory pore; segment 4 with distal cylindrical process bearing large aesthetasc (90 µm). Armature formula: 1 - [1], 2 - [8 + 1 pinnate], 3 - [5], 4 - [1 + (1 + ae)], 5 - [1], 6 - [3], 7 - [7 + acrothek]. Apical acrothek consisting of two long setae and one slender aesthetasc (35 µm). Antenna (Figs 2 A, 3 C, D) with small unarmed coxa; basis and first endopod segment incompletely fused forming allobasis, abexopodal margin without ornamentation; exopod a narrow segment, with one long, pinnate and one shorter, naked seta apically; endopod with two lateral spines and distal armature consisting of two pinnate spines, two geniculate setae and one large geniculate spine bearing spinules at approximately mid-margin and fused at base with a short, pinnate seta. Labrum (Fig. 2 A, B) a well-developed, ventrally produced extension; distal margin with short, blunt spinules; lateral margins with finer setules. Mandible (Fig. 4 A). Gnathobase well developed; uniramous palp, consisting of basis and onesegmented endopod; basis elongate, with one lateral, pinnate seta; endopod with one outer and four apical setae fused in two clusters. Maxillule (Fig. 3 E). Praecoxal arthrite with ten spines / setae around distal margin and two tubular setae (one long, one rudimentary) on anterior surface. Coxal endite with one geniculate claw and one seta. Basis and rami largely fused into elongate palp; basal armature represented by two lateral setae, and three setae plus a geniculate claw apically. Endopod represented by a small semidiscrete segment with three setae, exopod by two small setae. Maxilla (Fig. 4 B, C). Syncoxa with two endites, proximal endite with three articulating setae, distal endite with one pinnate claw and two setae, all articulating. Allobasis drawn out into a claw-like pinnate endite armed with two additional setae; with a distinct tube-pore. Endopod a discrete segment with four long setae and with a small sclerite around its base (Fig. 4 C). Maxilliped (Figs 2 A, B, 3 F) well developed, subchelate, directed inwards. Syncoxa well developed, with one pinnate seta and two spinular rows on anterior surface. Basis elongate, with two to three long spinules anteriorly and a spinular row along inner margin posteriorly. Endopod represented by a strong, curved, bare claw. P 1 (Fig. 4 D, E). Praecoxa strongly developed. Coxa with three spinular rows. Basis with long, naked, inner seta and short, pinnate, outer seta. Exopod three-segmented; with outer spine on exp- 2, with two spines and two geniculate setae on exp- 3. P 1 endopod prehensile, distinctly longer than exopod; proximal segment approximately ten times as long as average width, with pinnate inner seta being plumose in proximal third; distal segment short, with subdistal setule, and two geniculate spines distally. Swimming legs P 2 – P 4 (Fig. 5 A – F). P 4 distinctly longer than P 2 – P 3. Width of intercoxal sclerites decreasing in antero-posterior direction (Fig. 5 B, D, E). Praecoxae well developed, with spinular row on anterior surface in P 2 and P 4. Coxae with pattern of spinules, as in Figure 5 A, B, C, D, E, F, with a large tubepore on anterior surface of P 3 and P 4. Bases with outer seta (short and pinnate in P 2, long and plumose in P 3, long and bare in P 4); with spinular rows on anterior surface only (Fig. 5 A, C, E). Exopods threesegmented, endopods one-segmented. Exopodal spines of P 3 minutely serrate; inner distal spine shorter than outer distal one (Fig. 5 C). Inner setae of P 4 endopod and P 4 exp- 3 serrate. Inner element of P 2 endopod setiform, bare, approximately 0.3 times the length of distal spine. Armature elements of P 3 endopod spiniform; inner spine less than half length of outer spine. Seta and spine formulae as for genus. Fifth pair of legs (Figs 5 G, 6 A) not fused medially, no distinct intercoxal sclerite. Baseoendopod and exopod fused into a common, elongate plate, tapering distally; apex with strong, articulating spine, distinctly longer than plate and with flagellate tip; outer margin with three sparsely plumose setae (including seta derived from baseoendopod); inner margin with two serrate spines, one pinnate seta and one long, pinnate seta fused to plate; anterior surface with three large tube-pores. Sixth legs (Fig. 2 D) each represented by small operculum closing off gonopore; armature consisting of spiniform outer element and two accessory setules. Genital apertures not fused medially; copulatory pore large, located slightly posterior to gonopores; leading via a short chitinized copulatory duct to paired anterior extensions positioned anterior to genital apertures; copulatory pore flanked by two small secretory pores. Single egg-sac (Fig. 1 B, C) containing approximately seven eggs arranged in a biserial way, enclosed in a common egg-sac membrane; egg-sac connected with each genital aperture via transparent string. Male: Body length: 705 µm (Fig. 7 A). Spermatophore 110 µm. Anal somite without spinules near ventral anterior (Fig. 6 B) and hind margins (Fig. 6 B – F). Antennule (Fig. 7 B, C) indistinctly nine-segmented; geniculation between segments 7 and 8; segment 1 with one minute seta and spinular pattern as in female; segment 2 longest, with one plumose and eight naked setae; segment 3 with six setae; segment 4 minute, forming an incomplete ring, with two short setae; segment 5 distinctly swollen, with six elements along the anterior margin and with a distal cylindrical process bearing basally fused seta and large, constricted aesthetasc (53 µm); segment 6 with one long seta and one short pinnate spine; segment 7 with three spiniform elements and one seta; segment 8 with one seta and four modified elements (modified as in Figure 24 C for E. cumbraensis); segment 9 with five posterior setae, one anterior seta and one seta plus an acrothek apically. Apical acrothek consisting of two naked setae and one slender aesthetasc (15 µm). P 2 (Fig. 8 A) with inner distal corner of basis not modified into spinous process (as in other cylindropsyllid genera) but lateral margin more chitinized than in female. Endopod slightly larger than in female; outer margin with more spinules; apical seta distinctly longer, extending beyond distal margin of exp- 2, with sparser ornamentation than in female; inner seta larger than in female. Exp- 3 modified; outer distal element setiform and distinctly longer than in female; inner distal element transformed into strong claw, directed medially and posteriorly, and with distal half pinnate and tapering to a fine tip. Spines of P 3 exopodal segments with pinnate ornamentation (Fig. 8 B); exp- 1 without hyaline frill; exp- 3 with secretory pore on anterior surface near articulation with exp- 2; outer distal element of exp- 3 setiform and distinctly longer than in female. P 3 endopod (Fig. 8 B – D) distinctly two-segmented; enp- 1 small, with short, serrate, posterior seta and long, rigid apophysis arising from the anterior surface; distal half of apophysis bilaterally compressed (cf. Fig. 8 C, D); enp- 2 tapering distally, with one short, pinnate seta apically. Fifth legs (Figs 6 B, 8 E) not fused medially, no distinct intercoxal sclerite. Baseoendopod and exopod fused into a common elongate plate, tapering distally towards long, pinnate, spinous process, which is longer than the plate; inner margin with serrate spine; outer margin with small serrate spine, three naked setae and a sparsely plumose seta derived from baseoendopod; anterior surface with three large tube-pores. Sixth pair of legs (Figs 6 B, 8 F) asymmetrical, with three sparsely pinnate setae each, decreasing in length medially. Left or right leg articulating according to sinistral or dextral development of testis and vas deferens. First postgenital somite with transverse spinular row near ventral anterior margin (Fig. 8 F). Caudal ramus conical (Fig. 6 D – F), slightly longer than in female, length (measured along the outer margin) approximately 2.2 times the proximal width; seta IV long and uniplumose, extending to flexure zone of seta V. Differential diagnosis: Evansula incerta and E. arenicola are the only species in the genus that have retained the syncoxal seta on the maxilliped and that lack the presence of a raised spinular row or spinous process on the dorsal surface of the caudal rami. In both species seta c of the female P 5 (cf. Figure 36 for reference position) is fused to the segment. However, this character is also shared by E. spinosa sp. nov. Females of E. incerta and E. arenicola can be differentiated by the general facies of leg 1, the shape of the caudal rami and seta V, the length of the inner distal seta of the P 2 endopod, and the presence / absence of ventral anterior spinule rows on the anal somite. Males can be distinguished by the endopodal segmentation of P 3 and P 4. Distribution: Scotland: St. Monans in Firth of Forth (Scott, 1892, 1906 b). Although many authors have recorded specimens they attribute to E. incerta, there is good reason to believe that in fact they have often mistaken other undescribed species for it. The reasons for this conclusion lie in the frequently repeated statements about the difficulty in differentiating E. incerta, E. pygmaea, and E. arenicola. With the discovery of several new species in the North Sea, the reliability of previously published records from north-western Europe becomes uncertain. The true state of confusion reigning in the genus is illustrated by the situation in the Firth of Forth – the type locality of both E. incerta (St. Monans) and E. pygmaea (Musselburgh). Examination of a single, intertidal sandflat sample taken at Elie (near St. Monans) revealed the presence of E. cumbraensis sp. nov. (type locality Isle of Cumbrae, and widely distributed in the North Sea) and resulted in the discovery of two other new species, which are currently under study. Surprisingly, the sample did not contain any E. incerta or E. pygmaea. A re-examination of T. Scott’s material of the Forth River produced the second record of E. spinosa sp. nov. (type locality Korshavn, Norway), raising the number of Evansula species in the Firth of Forth to six.	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
03F487A0FFADFFA8FFC7FCCDFA1194A4.taxon	description	Sweden: Hållö (Por, 1964), Isle of Bonden (Por, 1964; Swedmark & Teissier, 1967). Scotland: River Ythan (Hockin, 1981, 1982 a, b, c, 1983, 1984; Hockin & Ollason, 1981). Wales: Porth-y-Post and Port Swtan (Church Bay), Anglesey (Geddes, 1972). Germany: Kieler Bucht (Klie, 1929, 1950; Remane, 1933; Kunz, 1935), off Bottsand and Weisenhaus in Kieler Bucht (Noodt, 1956, 1957), Boknis Eck in Kieler Bucht (Scheibel, 1976), Helgoland (Kunz, 1938; Klie, 1950), Isle of Sylt (Noodt, 1952, 1957), Amrum (Noodt, 1957). Belgium: North Sea coastal zone (Govaere et al., 1980). France: Kersaint, Finistère (Bodin & Boucher, 1981; Bodin, 1988), Charente-Maritime (Bodin, 1976, 1977), Bassin d’Arcachon, Gironde (Renaud-Debyser, 1963 a, b), Contis-Plage, Landes (Noodt, 1955 a, b; Delamare Deboutteville, Gerlach & Siewing, 1955; Delamare Deboutteville, 1960). Hockin (1984) recorded the presumed ectocommensal suctorians Thecacineta inclusa Meunier, 1903 and Thecacineta cothurnoides Collin, 1909 from E. incerta in the River Ythan estuary, Aberdeenshire, Scotland.	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
03F487A0FFA3FFB6FFD3F9A7FDC99052.taxon	description	Original description: Scott (1903 a: 117 – 118, plate IV, figs 11 – 19). Additional descriptions: None. The additional illustrations given by Mielke (1975) and Martínez Arbizu & Moura (1994) are based on specimens of E. cumbraensis sp. nov. Type locality: Scotland, Firth of Forth, near Musselburgh. Material examined: NHM, reg. nos 45033 – 035: Firth of Forth, Scotland; 1 ♀ in alcohol (labelled ‘ cotype’); coll. T. Scott, 1894; as part of Cannon A. M. Norman collection (1911.11.8); dissected on eight slides. Redescription Female: Total body length: 475 µm. Body slender, cylindrical (Fig. 9 A, B), semitransparent, yellowish; no distinct separation between prosome and urosome. Genital double-somite completely fused (Figs 9 A, B, 12 C); original segmentation marked dorsally by paired anterior and posterior sensillae (Fig. 9 A, B). Anal somite only slightly longer than wide (50 × 45 µm), with three pairs of secretory pores laterally (Fig. 12 D, E); ventral surface with four rows of tiny spinules near anterior border (Fig. 12 D); posterior margin with two short spinular rows on either side of ventral midline (Fig. 12 D). Anal operculum weakly developed, unarmed (Figs 10 E, 12 E). Caudal rami slightly convergent (Figs 9 A, 10 E, 12 E); distinctly constricted around anterior rim resulting in strongly convex contour of proximal inner margin; length (measured along outer margin) approximately 2.8 times proximal width; dorsal surface without chitinous spur but with raised spinular row anterior to seta VII (Fig. 10 E) and with blunt dorsal process (discernible in lateral aspect: Fig. 12 E); with seven setae, setae I – VI in distal and seta VII in proximal half (Fig. 10 D, E); seta I diminutive; setae II – III long and bare; seta IV long and naked, more than twice the length of styliform part of V; seta V long, with distinct flexure zone between styliform part and long distal flagellate part, fused at base with seta IV; seta VI vestigial and spiniform; seta VII triarticulate at base, somewhat flattened near tip (Fig. 10 E) and located along proximal inner margin; ventral and dorsal surfaces with one tube-pore each (Fig. 12 D, E). Rostrum elongate (Fig. 12 A), with slightly convex margins in proximal half, tapering distally; distinctly shorter than first antennulary segment (Fig. 9 A); with two long sensillae; median pore positioned dorsally near apex of rostrum. Antennule seven-segmented (Fig. 10 A). Segment 1 only slightly shorter than segment 2, with small sclerite around proximal posterior margin, anterior surface with pattern of spinular rows, as illustrated in Figure 10 B; segment 2 longest, without secretory pore; segment 4 with distal cylindrical process bearing large aesthetasc (55 µm). Armature formula: 1 - [1], 2 - [8 + 1 pinnate], 3 - [5], 4 - [1 + (1 + ae)], 5 - [1], 6 - [3], 7 - [7 + acrothek]. Apical acrothek consisting of two long setae and one slender aesthetasc (25 µm). Antenna, mandible, maxillule, and maxilla as in E. incerta. Maxilliped (Fig. 12 B) well developed, subchelate, directed inwards. Syncoxa well developed, with two spinular rows, without seta. Bases elongate, with two to three long spinules on anterior surface and a spinular row along posterior inner margin. Endopod represented by strong, curved, bare claw. P 1 (Fig. 10 C, D). Praecoxa strongly developed, unarmed. Coxa with three spinular rows. Basis with long, semiplumose, inner seta and short, pinnate, outer seta. Exopod three-segmented; with two spines and two geniculate setae on exp- 3. P 1 endopod prehensile, distinctly longer than exopod; proximal segment approximately nine times as long as average width, with pinnate inner seta being plumose in proximal third; distal segment short, with subdistal setule, and two geniculate spines distally. Swimming legs P 2 – P 4 (Fig. 11 A – F) somewhat more robust than in E. incerta. Praecoxae well developed, with spinular row on anterior surface (Fig. 11 A, C, E). Coxae with pattern of spinules as in Figure 5 A – F, with large tube-pore on anterior surface of P 3 (but not P 4). Bases with outer seta (short and pinnate in P 2, long and plumose in P 3, long and bare in P 4); with spinular rows on anterior surface only (Fig. 11 A, C, E). Exopods three-segmented, endopods one-segmented. Exopodal spines of P 3 pinnate (Fig. 11 C). P 2 endopod approximately 0.8 times as long as exp- 1; with two spinular rows on anterior and one spinular row on posterior surface; inner element setiform, pinnate, only slightly shorter than distal spine. Armature elements of P 3 endopod spiniform; inner spine approximately half length of outer spine. Seta and spine formulae as for genus. Fifth pair of legs (Fig. 11 G) with baseoendopod and exopod fused into a common elongate plate, tapering distally; apex with strong, articulating spine, slightly longer than plate and with flagellate tip; outer margin with three sparsely plumose setae (including seta derived from baseoendopod), middle and distal ones distinctly shorter than in E. incerta; inner margin with three pinnate setae (middle one longer than in E. incerta) and one short serrate spine (much shorter than in E. incerta); none of armature elements fused to plate; anterior surface with three large tube-pores. Inner margin setae sometimes with foreign tubular filaments attached to individual pinnules. Sixth legs (Fig. 12 C) each represented by a small operculum closing off gonopore; armature consisting of setiform outer element and two accessory setules. Genital apertures not fused medially; copulatory pore large, located slightly posterior to gonopores; leading via short chitinized copulatory duct to paired anterior extensions, positioned anterior to genital apertures; copulatory pore flanked by two small secretory pores. Male: Based on Scott’s (1903 a) original description. P 3 endopod distinctly three-segmented; enp- 1 small and unarmed; enp- 2 drawn out into long, bent apophysis (the accessory seta on this segment was probably overlooked by T. Scott); enp- 3 with one short apical seta. P 5 baseoendopod and exopod fused into common triangular plate, tapering distally towards curved, naked, spinous process, which is slightly longer than the plate; inner margin with one seta; outer margin with five armature elements, the distalmost one short- est and setiform; no details given about the ornamentation of these elements. Remark: The plumosity of the long inner setae on the female P 5 in Scott’s (1903 a) original description is an artefact, being caused by attached filamentous bacterial epibionts. Differential diagnosis: Perhaps the most characteristic feature of E. pygmaea is the peculiar shape of the slightly convergent caudal rami with their strongly convex proximal inner margins. The species is most closely related to E. cumbraensis sp. nov. and E. polaris sp. nov. with which it shares the raised spinular row on the caudal ramus. E. pygmaea differs from E. cumbraensis sp. nov. in the length of the outer spines on P 2 exp- 2 and - 3, the size of the copulatory pore, the presence of an anteroventral spinular row on the female anal somite and in both size and ornamentation of the setae on the female P 5 (in particular the short inner distal spine). The major discrepancy with the Arctic species is found in the shape and length of the caudal ramus. Provided Scott’s (1903 a) illustration (plate IV, Fig. 17) is accurate, additional differences can be found in the male P 5. Distribution: Scotland: Musselburgh in Firth of Forth (Scott, 1903 a, 1906 b). The following list contains all records attributed to E. pygmaea that do not provide enough information for a positive identification to be made. Scotland: Firemore, Loch Ewe (McIntyre & Murison, 1973), River Ythan (Hockin, 1982 c). Wales: Trearddur Bay, Traeth Lligwy, Traeth Bychan, and Tal-y-Foel, Anglesey (Geddes, 1972). England: Whitsand Bay, Cornwall (Harris, 1972 a, b, c), St. Andrews and St. Agnes, Isles of Scilly (Wells, 1961, 1970), River Exe (Wells, 1963). Ireland: Silver Strand, Galway (Bodin & Jackson, 1989; Holmes & O’Connor, 1990). Germany: Isle of Sylt (Noodt, 1952, 1957), Kieler Bucht (Klie, 1929, 1950; Remane, 1933; Kunz, 1935; Scheibel, 1972, 1973; Scheibel & Noodt, 1975; Anger & Scheibel, 1976), Bottsand and Weissenhaus in Kieler Bucht (Noodt, 1956, 1957), Bocknis Eck in Kieler Bucht (Scheibel, 1976), Helgoland (Kunz, 1938; Klie, 1950), Elbe estuary (Riemann, 1966). The Netherlands: Oosterschelde (Smol et al., 1994), Westerschelde estuary (Van Damme, Heip & Willems, 1984). Belgium: Sluice Dock, Ostend (Thielemans & Heip, 1984), Kwinte Bank (Willems et al., 1982 a, b), coastal zone (Heip, Herman & Vincx, 1984).	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
03F487A0FFBDFFBCFCABFEEAFAE4931D.taxon	description	Original description: Nicholls (1939: 299 – 302, figs 23, 24). Additional description: Wilson (1932) (as E. incerta). Type locality: Washings from coarse sand, taken by grab at 8 m depth, at Baie de Mille Vaches on the north shore of the St. Lawrence River, Quebec (Canada). Material examined: NHM, reg. nos 1940.5.1.73 – 78: syntypes, 8 ♀♀ and 2 ♂♂ in alcohol; 1 ♀ (on nine slides) and 1 ♂ (on six slides) dissected; leg. A. G. Nicholls, 19 August 1937. Redescription Female: Total body length: 710 – 750 µm (N = 5; mean = 730 µm). Body slender, cylindrical (Fig. 13 A), semitransparent, yellowish; no distinct separation between prosome and urosome. Genital double-somite completely fused (Figs 13 A, 14 A); original segmentation marked dorsally by paired anterior and posterior sensillae (Fig. 13 A) and ventral chitinous patches (Fig. 14 A). Anal somite only slightly longer than wide (84 × 76 µm), with two pairs of secretory pores laterally (Fig. 16 D); ventral surface with four rows of tiny spinules near anterior margin (Fig. 14 A); posterior margin with two short spinular rows on either side of ventral midline (Fig. 13 D). Anal operculum weakly developed, unarmed (Fig. 17 E). Caudal rami slightly divergent, cylindrical except for proximal third, which is swollen both dorsally and medially (Figs 13 D, 16 D, 17 E), length (measured along outer margin) approximately 2.8 times the proximal width; dorsal surface without chitinous spur or raised spinular row; with seven setae, seta VII in proximal third, setae I – II at approximately half ramus length and setae III – VI in distal third (Fig. 17 E); seta I diminutive; seta II long and pinnate; seta III long and bare; seta IV relatively long, unipinnate at approximately halfway its length, longer than swollen part of V; seta V long, with distinct flexure zone between short, proximal bulbous part and long, distal flagellate part, fused at base with seta IV (Figs 13 D, 17 E); seta VI vestigial; seta VII tri-articulate at base and located along proximal inner margin; with three tube-pores, one dorsally, one ventrally, and one laterally (Figs 13 D, 16 D, 17 E). Rostrum elongate (Fig. 13 B), only slightly shorter than first antennulary segment; proximal third with inflated lateral margins, distinctly tapering distally; with two long sensillae; median pore positioned dorsally near apex of rostrum. Antennule seven-segmented (Fig. 13 B). Segment 1 approximately as long as segment 2, with small sclerite around proximal posterior margin, anterior surface with pattern of spinular rows, as illustrated in Figure 13 C; segment 2 without secretory pore; segment 4 with distal cylindrical process bearing large aesthetasc (115 µm). Armature formula: 1 - [1], 2 - [8 + 1 pinnate], 3 - [5], 4 - [1 + (1 + ae)], 5 - [1], 6 - [3], 7 - [7 + acrothek]. Apical acrothek consisting of two long setae and one slender aesthetasc (40 µm). Antenna, mandible, maxillule, and maxilla as in E. incerta. Maxilliped (Fig. 14 E) well developed, subchelate, directed inwards. Syncoxa well developed, with one pinnate seta and two spinular rows. Basis elongate, with three to four long spinules on anterior surface and a spinular row along posterior inner margin. Endopod represented by strong, curved, sparsely pinnate claw. P 1 (Fig. 15 A). Praecoxa strongly developed, with a row of tiny spinules. Coxa with two spinular rows on posterior surface and eight spinular rows on anterior surface. Basis with one posterior and two anterior spinular rows; with long, plumose, inner seta and short, bare, outer seta; anterior surface with secretory pore. Exopod three-segmented; with two spines and two geniculate setae on exp- 3. P 1 endopod prehensile, distinctly longer than exopod; proximal segment approximately nine times as long as average width, with pinnate inner seta not plumose in proximal third; distal segment short, with three spinular rows, a subdistal pinnate setule, and two geniculate setae distally (both being distinctly longer than in other species). Swimming legs P 2 – P 4 (Figs 15 B, 16 A, 17 A). P 4 distinctly longer than P 2 – P 3. Praecoxae well developed, with spinular row on anterior surface in P 2 – P 4. Coxae with pattern of spinules as in Figures 15 B, 16 A, 17 A; with large tube-pore on anterior surface of P 3 and P 4. Bases with outer seta (pinnate and spiniform in P 2, long and plumose in P 3, long and bare in P 4); with spinular rows on anterior surface only (Figs 15 B, 16 A, 17 A). Exopods three-segmented, endopods onesegmented. Inner distal spine of P 3 – P 4 exp- 3 shorter than outer distal one (Figs 15 B, 17 A). Inner setae of P 4 endopod and exp- 3 serrate. Inner element of P 2 endopod spiniform, pinnate, less than half length of distal spine. Armature elements of P 3 endopod spiniform; of approximately equal length with inner spine, slightly longer than outer. Seta and spine formulae as for genus. Fifth pair of legs (Fig. 16 C) with baseoendopod and exopod fused into a common elongate plate, tapering distally; apex with strong, articulating spine, approximately as long as the plate and with a flagellate tip; outer margin with three plumose setae (including seta derived from baseoendopod); inner margin with two serrate spines, one plumose seta, and one long, pinnate seta fused to plate and distinctly swollen in proximal half; anterior surface with three large tube-pores. Sixth legs (Fig. 14 A, D) each represented by small operculum closing off gonopore; armature consisting of three setiform elements with outer slightly longer than others. Genital apertures not fused medially; copulatory pore of moderate size, located slightly anterior to gonopores; leading via short chitinized copulatory duct to paired anterior extensions positioned anterior to genital apertures; copulatory pore flanked by two small secretory pores. Single egg-sac (Fig. 13 A) containing approximately 12 – 13 eggs arranged biserially, enclosed in a common egg-sac membrane; egg-sac connected with each genital aperture via transparent string. Male. Body length: 690 µm. Spermatophore 100 µm. Anal somite without spinules on ventral anterior or hind margins (Fig. 14 B). Antennule as in E. incerta. P 2 (Fig. 16 B) with inner distal corner of basis not modified into spinous process. Exp- 3 modified; outer distal element more setiform than in female; inner distal element transformed into strong claw, directed medially and posteriorly, pinnate along middle third. Endopod slightly shorter than in female; anterior surface with only one spinular row (four in female); apical spine distinctly longer and more slender than in female; inner spine bare. P 3 (Fig. 15 C) exp- 1 with reduced hyaline frill; exp- 3 with secretory pore on anterior surface near joint with exp- 2; outer distal spine of exp- 3 distinctly shorter than in female. P 3 endopod (Fig. 15 C, D) distinctly threesegmented, longer than exp- 1; enp- 1 small, without armature; enp- 2 with serrate, posterior seta and long, rigid apophysis arising from anterior surface; distal half of apophysis with flimsy barb; enp- 3 tapering distally, with one strong, bare spine apically (bifid at tip). P 4 endopod two-segmented (Fig. 17 C); enp- 1 with spinular row, without armature; enp- 2 with two serrate setae along inner margin and two pinnate spines apically, inner distal spine distinctly shorter than in female. Fifth legs (Fig. 17 D) with baseoendopod and exopod fused into a common, elongate plate, tapering distally towards long, spinous process, which is longer than the plate, slightly curved and bare; inner margin with serrate spine; outer margin with small serrate spine (partly fused to the plate), three naked setae and a sparsely plumose seta derived from baseoendopod; anterior surface with three large tube-pores. Sixth pair of legs (Fig. 14 B, C) asymmetrical, with three setae each, decreasing in length medially; outer seta sparsely plumose. Left or right leg articulating according to sinistral or dextral development of testis and vas deferens. First postgenital somite with transverse spinular row near ventral anterior margin (Fig. 14 B). Caudal ramus (Fig. 17 F) as in female, except for length and shape of proximal styliform part of seta V, which is distinctly longer than caudal ramus length. Variability: Nicholls (1939) described the female P 4 endopod as being partly divided into two segments. This partial subdivision, marked by a suture line halfway along the outer margin (Fig. 17 B), was found in most female specimens examined. However, in some individuals and sometimes in the same specimen, no such subdivision could be observed (Fig. 17 A). Surprisingly, the separation in two distinct segments in the male was overlooked by Nicholls (1939). Remarks: Nicholls (1939) figures two geniculate setae and only one outer spine on P 1 exp- 3 of the female, whereas for the male he states ‘ ... with 3 terminal setae on exopod’. Re-examination of all syntypes revealed that Nicholls’ illustration was based on a damaged or aberrant female. There is little doubt that Wilson’s (1932) illustrations of E. incerta were based on E. arenicola. Evidence for this is found in his illustrations of the male, which show a two-segmented P 4 endopod and agree in most other aspects, and in his statement that the female seta V of the caudal ramus ‘ ... is stout and spiniform proximally and passes abruptly at an angle into a filiform distal portion’. For the male he characterized the latter seta as being enlarged at the base, which conforms to the sexual dimorphism found in E. arenicola. Differential diagnosis: Evansula arenicola clearly occupies an isolated position in the genus by virtue of its two-segmented P 4 endopod in the male. It is also the only species that displays sexual dimorphism in the caudal seta V (bulbiform in the female) and the inner distal spine of the P 4 endopod (reduced in the male). The male P 2 endopod is significantly smaller in comparison with other species. The male P 3 endopod, on the contrary, is much larger than in its congeners and differs in the presence of a strong, naked apical spine. The general shape of the caudal rami and the P 1 (enp- 1 inner seta not plumose in proximal part; enp- 2 with much longer geniculate elements) provide additional differentiating characters. Distribution: Canada: St. Lawrence River, Quebec (Nicholls, 1939).	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
03F487A0FFBDFFBCFCABFEEAFAE4931D.taxon	materials_examined	USA: Katama Bay, Marthas Vineyard, Massachusetts (Wilson, 1932); it is conceivable that Coull’s (1971, 1977) records of E. incerta from the North Carolina continental shelf also belong to E. arenicola.	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
03F487A0FFB7FF87FFFEFD2EFBB29635.taxon	description	Original description: Mielke (1975: 98 – 100, abb. 66, as E. pygmaea: figs 2 b, 3 a – c). Additional description: Martínez Arbizu & Moura (1994) (as E. pygmaea). Type locality: Scotland, Isle of Cumbrae, Fintray Bay; exposed sandy beach with a steep profile; coarse sand. Material examined: (1) From type locality: holotype ♀ (NHM reg. no. 2005.2025) and paratype ♂ (NHM reg. no. 2005.2026) dissected on six and five slides, respectively; other paratypes in alcohol 15 ♀♀ and 7 ♂♂ (NHM reg. nos 2005.2027 − 48); coll. R. Huys & S. Conroy-Dalton; 27 August 1993; Karaman- Chappuis method (Delamare Deboutteville, 1954); (2) Firth of Forth, Elie Bay (Scotland): 15 ♀♀ and 10 ♂♂ in alcohol; coll. R. Huys & S. Conroy-Dalton; 26 August 1993; sandbank sampled at low tide; clean medium sand; deposited in NHM (reg. nos 2005.2049 − 58); (3) From Dr W. Mielke: Isle of Sylt; 5 ♀♀ and 5 ♂♂ in alcohol (NHM reg. nos 2005.2059 − 68). Etymology: The species is named after the type locality, the Isle of Cumbrae in Scotland. Description Female: Total body length: 500 – 560 µm (N = 10; mean = 535 µm). Body slender, cylindrical, semitransparent, colourless; no distinct separation between prosome and urosome. Genital double-somite completely fused (Fig. 18 B, C); original segmentation marked dorsally by paired anterior and posterior sensillae. Anal somite longer than wide (60 × 44 µm), with three pairs of secretory pores laterally (Fig. 21 C); posterior margin with two short spinular rows on either side of ventral midline (Figs 18 B, 21 E). Anal operculum weakly developed, unarmed (Fig. 21 B). Caudal ramus slightly conical (Figs 18 B, 21 B – E), length (measured along outer margin) approximately 2.5 times the proximal width; dorsal surface without chitinous spur but with raised spinular row forming crest covering base of seta VII (Figs 21 B, 27 C, D); with seven setae, setae I – VI in distal and seta VII in proximal half (Fig. 21 B); seta I diminutive, tubular (Fig. 27 A, B); setae II – III long and bare; seta IV long, longer than styliform part of V, plumose half its length; seta V long, with distinct flexure zone between styliform basal part and long distal flagellate part, fused at base with seta IV (Fig. 27 A); seta VI vestigial; seta VII tri-articulate at base (Fig. 27 D) and located along proximal inner margin; ventral surface with one tube-pore near anterior margin (Fig. 21 E), lateral surface with two tube-pores (Figs 21 C, 27 A, B). Rostrum elongate (Figs 18 F, 23 A, B), with parallel margins in proximal half, tapering distally; distinctly shorter than first antennulary segment; base surrounded by area of flexible integument (Fig. 23 A), with two long sensillae; median pore raised and positioned dorsally near apex of rostrum (Fig. 23 A, B). Antennule seven-segmented; armature formula as in E. incerta; segment 1 only slightly shorter than segment 2, with small sclerite around proximal posterior margin (Fig. 23 A), with pattern of spinular rows as illustrated in Figure 21 A. Antenna, mandible, maxillule, and maxilla as in E. incerta. Labrum, a massive lobe (Fig. 22 A) overlying mouth and mandibular gnathobases; with series of overlapping spinular rows around midventral margin (Figs 22 C, 23 D). Paragnaths, well-developed lobes, closely adpressed to gnathobases of mandibles (Fig. 22 C); with two rows of spinules each. Maxilliped (Fig. 22 A) as in E. pygmaea. Syncoxa without seta. Palmar margin of basis with spinular row posteriorly and patch of longer setules anteriorly. Endopodal claw distinctly curved, bare. Area between maxillipedal syncoxae and intercoxal sclerite of P 1 provided with midventral, trifid, spinous process (Figs 22 A, 23 C). P 1 (Fig. 19 A, B). Praecoxa strongly developed. Coxa with three distinct spinular rows. However, scanning electron microscopy (Fig. 23 C) revealed additional rows of tiny spinules, which cannot be discerned using differential interference contrast microscopes. This additional ornamentation was observed in E. arenicola and it is probable that the same spinular pattern is present in all species. Basis with long, naked, inner seta and short, pinnate, outer seta. Exopod three-segmented; with two spines and two geniculate setae on exp- 3. P 1 endopod prehensile, distinctly longer than exopod; proximal segment approximately ten times as long as average width, with pinnate inner seta being plumose in proximal third; distal segment short, with two spinular rows, a subdistal setule, and two short, geniculate spines distally. Swimming legs P 2 – P 4 (Fig. 20 A – F). P 4 distinctly longer than P 2 – P 3. Width of intercoxal sclerites decreasing in antero-posterior direction (Fig. 20 A, D – F). Praecoxae well developed, with spinular row on anterior surface. Coxae with pattern of spinules as in Figures 20 A, B, D – F, with a large tube-pore on anterior surface of P 3 and P 4. Bases with outer seta (short and bare in P 2, long and plumose in P 3, long and bare in P 4); with spinular rows on anterior surface only (Fig. 20 A, B, D – F). Exopods three-segmented, endopods one-segmented. Exopodal spines of P 2 exp- 2 and exp- 3 elongate. Inner distal spine of P 3 – P 4 exp- 3 shorter than outer distal one. Inner setae of P 4 endopod and exp- 3 serrate. Inner element of P 2 endopod spiniform, pinnate, less than half length of distal spine. Armature elements of P 3 endopod spiniform; inner spine longer than half length of outer spine. Seta and spine formulae as for genus. Fifth pair of legs (Figs 18 B, 20 G) with baseoendopod and exopod fused into a common elongate plate, tapering distally, with stepped inner and outer margins; apex with strong, articulating spine, distinctly longer than plate and with flagellate tip; outer margin with one bare (seta derived from baseoendopod) and two pinnate setae; inner margin with two serrate spines and two long, pinnate setae (all free at base); anterior surface with three large tube-pores. Sixth legs (Figs 18 C, 26 A) each represented by small operculum closing off gonopore; armature consisting of spiniform outer element and two accessory setules. Genital apertures not fused medially; copulatory pore very wide, located at level of gonopores (Fig. 26 A); leading via short chitinized copulatory duct to paired anterior extensions positioned anterior to genital apertures; copulatory pore flanked by two complex secretory pores, each consisting of four vents (Fig. 26 B). Male: Body length: 480 – 525 µm (N = 5; mean = 505 µm) (Fig. 18 A). Spermatophore 65 µm. Anal somite without spinules on ventral anterior and hind margins (Figs 21 F, 25 D). Antennule indistinctly nine-segmented; geniculation between segments 7 and 8 (Fig. 24 A); segment 1 with one minute seta and spinular pattern as in female; segment 2 longest, with one plumose and eight naked setae; segment 3 with six setae; segment 4 minute, forming an incomplete ring, with two short setae; segment 5 distinctly swollen, with six elements along the anterior margin and with a distal cylindrical process bearing a basally fused seta and large, constricted aesthetasc; segment 6 with one long seta and one short pinnate spine; segment 7 with three spiniform elements and one seta (Fig. 24 D); segment 8 with one seta and three interdigitating modified setae (Fig. 24 B, C); segment 9 with five posterior setae, one anterior seta and one seta plus an acrothek apically. Apical acrothek consisting of two naked setae and one slender aesthetasc. P 2 (Fig. 19 D, E) with inner distal corner of basis not modified into spinous process. Exp- 3 modified; inner distal element transformed into slender claw, directed medially and posteriorly, and with pinnules in distal half (Fig. 19 E). Endopod of approximately the same size as in female; outer margin with two spinular rows; apical seta longer and more slender, extending beyond distal margin of exp- 2; inner seta minute, bare. P 3 (Fig. 19 F) with reduced hyaline frill on exp- 1 (Fig. 25 B); exp- 3 with secretory pore on anterior surface near joint with exp- 2 (Fig. 25 A). P 3 endopod (Fig. 19 F, G) distinctly three-segmented; enp- 1 small, without ornamentation or armature; enp- 2 short, with serrate, posterior seta and rigid, sigmoid apophysis arising from anterior surface; distal third of apophysis with denticulate margin; enp- 3 a short bulbous segment, with one short, pinnate seta apically. Fifth legs (Fig. 18 D) with baseoendopod and exopod fused into a common elongate plate, tapering distally towards a long, bare, spinous process, which is longer than the plate; inner margin with serrate spine; outer margin with small serrate spine, three naked setae and sparsely plumose seta derived from baseoendopod; anterior surface with three large tube-pores. Sixth pair of legs (Figs 18 E, 25 C) asymmetrical, with two long, sparsely pinnate setae and one short, bare seta each. Left or right leg articulating according to sinistral or dextral development of testis and vas deferens. First postgenital somite with pattern of transverse spinular rows near ventral anterior margin (Figs 18 E, 25 C). Caudal ramus (Fig. 21 F) as in female. Remarks: Mielke (1975) expressed some reservations about the identity of his Evansula material from the Isle of Sylt. He decided to attribute his specimens to E. pygmaea on the following grounds: (1) body length shorter than in E. incerta, (2) distal spines of P 1 enp- 2 claw-like, and (3) similarity in female P 5 with Scott’s (1903 a) original description. A comparison based on one female and one male from Sylt revealed no differences with the type material of E. cumbraensis except for (1) the P 2 endopod, which, as well as its distal spine, appears to be slightly longer in the German specimens (Fig. 20 C), (2) the shape of the geniculate spines on P 1 enp- 2, which appear to be slightly more stout (Fig. 19 C). However, the precise length and shape of these elements are dependent on their orientation on the slide (cf. Fig. 19 A, B), and (3) small differences in caudal ramus contours and length of styliform part of seta V (but see under variability). Mielke seems to have found only E. cumbraensis. However, other studies reporting both E. pygmaea and E. incerta (Noodt, 1952, 1957) potentially indicate the occurrence of other sympatric species on the Isle of Sylt. Martínez Arbizu & Moura’s (1994) drawings of the genital field (cf. position and size of copulatory pore; however, the innermost setule on P 6 was overlooked) and the P 2 of both sexes (length of exopodal spines, endopod in male) of E. pygmaea from Sylt leave no doubt that they were also dealing with E. cumbraensis. Mielke (1975) commented on the structure of the ‘ plumose’ setae found on the female P 5. He described the pinnules as composed of ‘ ... einem kurzen, verstärkten Basalteil und einem langen Endteil, der leicht abbrechen kan. Bei Verlust einer grösseren Anzahl von Fiederendstücken werde kurz behaarte Borsten vorgetäuscht’. Scanning electron microscopy revealed that these plumose setae are clearly pinnate (Fig. 26 D) and the plumosity is an artefact created by epibiotic filamentous bacteria attached at and around the short pinnules (Fig. 26 C: distal part of second innermost seta). McAllen & Hannah (1999) observed similar biofouling by filamentous bacteria on the exoskeleton of the high-shore rockpool harpacticoid Tigriopus brevicornis (O. F. Müller). In this species, the major biofoulant is Leucothrix mucor OErsted, which tends to colonize between the body tergites where the exoskeleton is at its thinnest for the action of chitinolytic enzymes. The filamentous bacteria found in E. pygmaea, E. cumbraensis, and E. arenicola appear to show a high site specificity, attaching themselves only to the long setae of the P 5 (usually the female). Variability: Found only in the proximal styliform part of seta V, the length and shape of which can vary considerably within a population. Figure 21 G illustrates the intraspecific variation found in the Elie population. The Sylt population clearly fits within this range (Fig. 21 E; Mielke, 1975: abb. 66 B). Differential diagnosis: The most important diagnostic features of female E. cumbraensis are the elongated exopodal spines on the middle and distal segments of P 2 and the very large copulatory pore. It shares with E. incerta the absence of ventral spinular rows near the anterior margin of the anal somite in the female. Males of the closely related E. cumbraensis and E. polaris can be distinguished by the detailed morphology of the P 3 endopod (segmentation, shape of enp- 3) and P 5 (spinous process smooth vs. pinnate). Distribution: Scotland: Fintray Bay, Isle of Cumbrae (present account); Elie in Firth of Forth (present account). Germany: Isle of Sylt (Mielke, 1975, 1976; Martínez Arbizu & Moura, 1994).	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
03F487A0FF8DFF8CFD75F9BEFA859635.taxon	description	Original description: Sars (1911: 415 – 416, suppl., plate 39). Additional description: Scheibel (1972). Type locality: Norway, Korshavn, near Lindesnes; at 30 – 50 fathoms. Material examined: (1) From type locality: holotype ♀ dissected on eight slides; found among spirit- preserved specimens of Neobradya pectinifera T. Scott, 1892, deposited in Zoologisk Museum, Oslo; coll. G. O. Sars; deposited in NHM (reg. no. 1995.428); (2) NHM, reg. no. 1995.429: Firth of Forth (no further details specified), Scotland; 1 ♂ paratype [found among spirit-preserved ‘ cotypes’ of E. incerta: reg. nos 44505 – 507; as part of Cannon A. M. Norman collection (1911.11.8)]; coll. T. Scott, 9 September 1894; dissected on eight slides; (3) NHM, reg. no. 2005.2069: Southern Bight of North Sea, off Suffolk (UK), 51 ° 57 ′ 24 ′ N 2 ° 10 ′ 57 ′ E; 42.7 m depth; 1 ♀ paratype in alcohol; coll. R. Huys, 30 March 1992. Etymology: The species name is derived from the Latin spina, meaning spine, and refers to the dorsal spinous process on the caudal ramus. Description Female: Total body length: 700 µm. Body slender, cylindrical (Fig. 28 A, B), semitransparent, light brown; no distinct separation between prosome and urosome. Genital double-somite completely fused (Figs 28 A, B, 29 D); original segmentation marked dorsally by paired anterior and posterior sensillae (Fig. 28 A). Anal somite only slightly longer than wide (72 × 66 µm), with three pairs of secretory pores laterally (Fig. 32 D); anterior margin with ventral transverse row of tiny spinules (Fig. 29 D); posterior margin with two short spinular rows on either side of ventral midline (Figs 29 D, 32 E). Anal operculum weakly developed, unarmed (Fig. 32 C). Caudal ramus cylindrical (Figs 29 D, 32 C – E) with slightly concave inner margin, length (measured along the outer margin) approximately 3.5 times the proximal width; dorsal surface with chitinous spur (Fig. 32 C, D); with seven setae, setae I – VI in distal and seta VII in proximal half (Fig. 32 C); seta I small, larger than in other species; setae II – III long and bare; seta IV bare, longer than styliform part of V; seta V long, with distinct flexure zone between styliform part and long distal flagellate part, fused at base with seta IV; seta VI vestigial; seta VII tri-articulate at base and located near proximal inner margin; ventral surface with one tube-pore (Fig. 32 E), lateral surface with one tubular and one simple pore (Fig. 32 D). Rostrum elongate (Fig. 28 A), with parallel margins in proximal half, tapering distally; distinctly shorter than first antennulary segment; with two long sensillae. Antennule seven-segmented (Fig. 28 A, B); armature formula as in E. incerta; segment 1 slightly longer than segment 2. Antenna with spinular row on coxa (Fig. 31 B); basis and first endopod segment incompletely fused to form allobasis, abexopodal margin without ornamentation; exopod small, with one long, pinnate and one shorter, naked seta. Antennary endopod, mandible, maxillule, and maxilla as in E. incerta. Maxilliped as in E. pygmaea. Syncoxa without seta. P 1 (Fig. 29 A). Praecoxa strongly developed, with three spinular rows. Coxa with six spinular rows on anterior and two on posterior surface. Basis with long, sparsely plumose inner seta and shorter, pinnate outer seta. Exopod three-segmented; with two spines and two geniculate setae on exp- 3. P 1 endopod prehensile, distinctly longer than exopod; proximal segment approximately ten times as long as average width, with pinnate inner seta being plumose in proximal third; distal segment short, with three spinular rows, a subdistal setule, and two geniculate spines distally. Swimming legs P 2 – P 4 (Fig. 30 A – C). P 4 distinctly longer than P 2 – P 3. Width of intercoxal sclerites decreasing in antero-posterior direction. Praecoxae well developed, with spinular row on anterior surface in P 2 – P 4. Coxae with pattern of spinules as in Figure 30 A, B, C, with large tube-pore on anterior surface of P 3 and P 4. Bases with outer seta (short and naked in P 2 and P 4, long and plumose in P 3); with spinular rows on anterior surface only. Exopods three-segmented, endopods one-segmented. Exopodal spines of P 3 – P 4 minutely serrate. Inner distal spine of P 3 – P 4 exp- 3 shorter than outer distal one. Inner setae of P 4 endopod and exp- 3 serrate. Inner element of P 2 endopod setiform, pinnate, approximately half length of distal spine. Armature elements of P 3 endopod spiniform; inner spine longer than half length of outer spine. Seta and spine formulae as for genus. Fifth pair of legs (Figs 29 D, 30 D) with baseoendopod and exopod fused into a common elongate plate, tapering distally; apex with strong, articulating spine (lost on both sides during dissection), approximately as long as the plate and with a flagellate tip; outer margin with three sparsely plumose setae (including seta derived from baseoendopod); inner margin with two serrate spines, one pinnate seta and one long pinnate seta fused to plate; anterior surface with three large tube-pores. Sixth legs (Fig. 29 D, E) each represented by small operculum closing off gonopore; armature consisting of spiniform outer element and two accessory setules. Genital apertures not fused medially; copulatory pore large, located slightly posterior to gonopores; leading via short chitinized copulatory duct to paired anterior extensions positioned anterior to genital apertures; copulatory pore flanked by two small secretory pores. Egg-sac not confirmed. Male: Body length: 670 µm. Spermatophore 75 µm. Anal somite without spinules near ventral anterior and hind margins (Fig. 31 C). Antennule (Fig. 31 A) distinctly nine-segmented; geniculation between segments 7 and 8; segment 1 with one minute seta and spinular pattern as in female; slightly longer than segment 2; segment 5 distinctly swollen, with distal cylindrical process bearing basally fused seta and large aesthetasc (100 µm); acrothek on segment 9 with slender aesthetasc (27 µm). P 2 (Fig. 32 A) with inner distal corner of basis not modified into spinous process but lateral margin slightly more chitinized. Exp- 3 modified; inner distal element transformed into strong claw, directed medially and posteriorly, and with distal half pinnate. Endopod slightly shorter than in female; outer margin with two spinular rows; apical seta distinctly longer, clearly extending beyond distal margin of exp- 2, with sparser ornamentation than in female; inner seta slightly larger than in female, bare. Spines of P 3 exopodal segments with pinnate ornamentation (Fig. 29 B); exp- 1 with reduced hyaline frill; exp- 3 with secretory pore on anterior surface near joint with exp- 2. P 3 endopod (Fig. 29 B, C) distinctly three-segmented; enp- 1 small, without armature or ornamentation; enp- 2 with serrate, posterior seta and long, rigid sigmoid apophysis arising from anterior surface; enp- 3 with weakly chitinized inner margin and one pinnate seta apically. Fifth legs (Figs 31 C, D, 32 B) with baseoendopod and exopod fused into a common elongate plate, tapering distally towards long, smooth, spinous process, which is longer than the plate; inner margin with serrate spine; outer margin with small, bare spine completely fused to plate, three naked setae and a sparsely plumose seta derived from baseoendopod; anterior surface with three large tube-pores. Sixth pair of legs (Figs 31 C, E) asymmetrical, with one short, naked and two long, sparsely pinnate setae each. First postgenital somite with transverse spinular row near ventral anterior margin (Fig. 31 E). Caudal ramus cylindrical (Fig. 32 F), shorter than in female; seta IV long and uniplumose; styliform part of seta V comparatively longer than in female. Remarks: Although there are several discrepancies between Scott’s (1892) and Sars’ (1911) descriptions of E. incerta, only the dorsal spinous process has attracted subsequent workers’ attention. The presence or absence of this structure was regarded as part of the intraspecific variability and consequently both the Norwegian and Scottish populations were believed to represent two forms of the same species. Kunz (1938) and Scheibel (1972) claimed that they had found both ‘ forms’ in their samples from Helgoland and the Kieler Bucht, respectively. However, at least in Scheibel’s case, there is evidence that E. spinosa occurred in his material (cf. his illustration of the caudal ramus; Scheibel, 1972: tafel XVII, Fig. 10). It should be noted that Sars (1911) inadvertently reversed the female P 2 and P 3 in his illustrations. Differential diagnosis: Evansula spinosa can be readily distinguished by the presence of a dorsal spur on the caudal ramus in both sexes, and by the fused outer spine on the male P 5. Distribution: Norway: Korshavn, near Lindesnes (Sars, 1911; present account), Troldfjord in Lofoten Islands. Germany: Kieler Bucht (Scheibel, 1972), probably also Helgoland (Kunz, 1938). Scotland: Firth of Forth (present account). England: off Suffolk (present account).	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
03F487A0FF84FF88FD49F94BFD109637.taxon	description	Original description: Scott (1903 b) did not provide any descriptive information. Type locality: Norway, East Finmark, Laksefjord. Material examined: From type locality: holotype ♂ found among spirit-preserved ‘ cotypes’ of E. incerta (reg. nos 44508 – 509) as part of the Cannon A. M. Norman collection (1911.11.8); coll. A. M. Norman, 30 June 1890; det. T. Scott and labelled Tetragoniceps incertus; deposited in NHM (reg. no. 1995.427); dissected on eight slides. Etymology: The species name is derived from the Latin polaris, meaning polar, and refers to the Arctic distribution of this species. Description Female: Unknown. Male: Total body length: 435 µm. Body slender, cylindrical (Fig. 33 A), semitransparent, yellowish; no distinct separation between prosome and urosome. Anal somite only slightly longer than wide (46 × 41 µm), with two pairs of secretory pores laterally (Fig. 33 B); posterior margin with one short spinular row on either side of ventral midline (Fig. 34 D). Anal operculum weakly developed, unarmed (Fig. 34 B). Caudal ramus short and conical (Fig. 34 B – D), proximal inner margin slightly convex, length (measured along outer margin) approximately 2.5 times the proximal width; dorsal surface without chitinous spur but with raised spinular row; with seven setae, setae I – VI in distal and seta VII in proximal half (Fig. 34 B); seta I diminutive; setae II – III long and bare; seta IV uniplumose, slightly longer than styliform part of V; seta V long, with distinct flexure zone between styliform part and long distal flagellate part, fused at base with seta IV; seta VI vestigial; seta VII tri-articulate at base and located along proximal inner margin, with conspicuous swelling in proximal quarter; ventral surface with one tube-pore near anterior margin (Fig. 34 D), lateral surface with two tube-pores (Fig. 34 C). Rostrum elongate (Fig. 33 C), with slightly convex margins in proximal half, tapering distally; slightly shorter than first antennulary segment; with two long sensillae; median pore positioned dorsally near apex of rostrum. Antennule distinctly nine-segmented (Fig. 33 C), geniculation between segments 7 and 8; segment 1 only slightly shorter than segment 2, with small sclerite around proximal posterior margin, with anterior pattern of spinular rows as illustrated for other species; segment 5 distinctly swollen, with a distal cylindrical process bearing a basally fused seta and large aesthetasc (52 µm); acrothek on segment 9 with slen- der aesthetasc (17 µm). Antenna, mandible, maxillule, and maxilla as in E. incerta. Maxilliped well developed, subchelate, directed inwards. Syncoxa well developed, without pinnate seta, with two spinular rows. Basis elongate, with two to three long spinules anteriorly and a posterior spinular row along inner margin. Endopod represented by strong, curved, bare claw. P 1 (Fig. 34 A). Praecoxa strongly developed. Coxa with four spinular rows. Basis with sparsely plumose inner seta and pinnate outer seta. Exopod threesegmented; with two spines and two geniculate setae on exp- 3. P 1 endopod prehensile, distinctly longer than exopod; proximal segment approximately ten times as long as average width, with pinnate inner seta being plumose in proximal third; distal segment short, with three spinular rows, a subdistal setule, and two geniculate spines distally. Swimming legs P 2 – P 4 (Fig. 35 A – D). Praecoxae well developed, with spinular row on anterior surface in P 2 – P 4. Coxae with pattern of spinules as in Figure 35 A, B, D, with large tube-pore on anterior surface of P 3. Bases with outer seta (short and pinnate in P 2, long and plumose in P 3, short and naked in P 4); with spinular rows on anterior surface only. Exopods three-segmented, endopods one-segmented (P 2, P 4) or incompletely three-segmented (P 3). Inner distal spine of P 3 – P 4 shorter than outer distal one. Inner setae of P 4 endopod and exp- 3 serrate. Seta and spine formulae as for genus. P 2 (Fig. 35 A) with inner distal corner of basis not modified into spinous process but with well-chitinized lateral margin. Endopod with three spinular rows; apical seta pinnate and clearly extending beyond distal margin of exp- 2; inner element setiform and naked. Exp- 3 modified; outer distal seta very long; inner distal element transformed into slender claw, directed medially and posteriorly, and with middle third pinnate and distal third denticulate. Spines of P 3 exopodal segments with pinnate ornamentation (Fig. 35 B); exp- 1 with reduced hyaline frill; exp- 3 with secretory pore on anterior surface near joint with exp- 2. P 3 endopod (Fig. 35 B, C) indistinctly three-segmented with middle and distal segments partly fused along posterior surface; enp- 1 small, without armature or ornamentation; enp- 2 with serrate, posterior seta and rigid, slightly sigmoid apophysis arising from anterior surface; enp- 3 with weakly chitinized inner margin and with one short pinnate seta apically. Fifth legs (Fig. 33 B, D) with baseoendopod and exopod fused into a common elongate plate, tapering distally towards long, pinnate, spinous process, which is longer than the plate; inner margin with serrate spine; outer margin with small, pinnate spine, three naked setae and sparsely plumose seta derived from baseoendopod; anterior surface with three large tube-pores. Sixth pair of legs (Figs 33 B, 35 E) asymmetrical, with one short, naked and two long, sparsely pinnate setae each. First postgenital somite with transverse spinular row near ventral anterior margin (Fig. 35 E). Differential diagnosis: Evansula polaris is thus far the only species that possesses spinule rows on the ventral posterior margin of the anal somite in the male (the condition in E. pygmaea is unknown). The characteristic shape of the male P 3 endopod is an additional differentiating feature. E. polaris is the smallest representative of the genus. Distribution: Norway: Laksefjord (Scott, 1903 b). Perhaps also in Troldfjord inside the Lofoten Islands [one male recorded by Sars (1911) as E. incerta].	en	Huys, Rony, Conroy-Dalton, Sophie (2006): Revision of the genus Evansula T. Scott, 1906 (Copepoda, Harpacticoida, Cylindropsyllidae) with a description of three new species. Zoological Journal of the Linnean Society 147 (4): 419-472, DOI: 10.1111/j.1096-3642.2006.00227.x, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2006.00227.x
