taxonID	type	description	language	source
951887EAFFC7FF9BFF51D69AE724FC22.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 2 – P 3 endopods, P 5 – P 6 and urosomal segmentation. Body fusiform; posterior margin of cephalothorax with plain hyaline frill; remaining somites (except P 5 - bearing somite and anal somite) with deeply divided, denticulodigitate, hyaline frills. Rostrum defined at base, large, reaching to halfway second segment of antennule; with parallel sides tapering sharply into pointed apex. Anal operculum semicircular, naked. Caudal ramus wider than long, without conspicuous spinular ornamentation; with seven setae, setae IV – V with fracture planes, not inflated at base in ♀. Antennule slender, segment 1 elongate in both sexes; 9 - segmented in ♀, segments 7 – 9 shortest, with aesthetascs on segments 4 and 9; haplocer and indistinctly 11 - segmented in ♂ (segment 4 largely incorporated in segment 5), with geniculation between segments 7 and 8, and aesthetascs on segments 5 and 11; segmental homologies in ♂: I, II – VIII, IX – XII, XIII, XIV – XVII, XVIII, XIX – XX, XXI – XXII, XXIII, XXIV – XXV, XXVI – XXVIII. Antenna not sexually dimorphic; with completely fused allobasis lacking endopodal pinnate seta on abexopodal margin; exopod 2 - segmented, proximal segment with two setae, distal segment with one lateral and three apical elements; free endopod without penicillate elements. Mandible with three elements on basis; endopod with two lateral and four apical setae; exopod rudimentary, represented by single seta on minute protuberance. Maxillulary coxal epipodite represented by one seta. Maxilla with three endites on syncoxa; endopod not completely discrete at base, with two setae. Maxilliped with two setae on syncoxa; basis with two setae, without longitudinal spinular row on palmar margin; endopod represented by claw with three accessory setae. P 1 inner basal spine not sexually dimorphic. P 1 exopod 3 - segmented, segments subequal in length; exp- 2 with inner seta; exp- 3 with two pinnate spines, one geniculate seta and one non-geniculate bipinnate seta. P 1 endopod 2 - segmented; enp- 1 longer than exopod, with long, proximally inserted, pinnate inner seta, not extending to distal margin of segment, segment margins without area of reduced chitinization; enp- 2 with one naked minute seta and two naked claws of different lengths. P 2 – P 4 rami 3 - segmented; inner seta of P 2 – P 4 exp- 1 of moderate size; distal inner seta of P 3 – P 4 exp- 3 reduced. P 2 endopod ♂ 2 - segmented; compound enp- 2 with one inner seta (inner seta of ancestral enp- 3 lost). P 3 endopod ♂ 3 - segmented; without inner seta (vs two in ♀) on enp- 3, resulting in 1.1.021 pattern; outer spine not transformed into spinous apophysis. Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe with five setae, outermost minute; inner margin without transverse striae; exopod moderately elongate, articulation with baseoendopod very narrow, with six elements. P 5 ♂ endopodal lobe with two small elements, outer one shortest; exopod 2 - segmented; exp- 1 with short outer seta, exp- 2 with three elements. Vestigial P 6 ♀ represented by three setae. P 6 ♂ with three setae.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFC7FF9BFF51D69AE724FC22.taxon	type_taxon	Type and only species. Johnwellsia bipartita sp. nov. (by original designation herein).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFC7FF9BFF51D69AE724FC22.taxon	etymology	Etymology. The genus is named in dedication to the late Prof. John J. B. Wells, in honour of his outstanding career in harpacticoid taxonomy, systematics and biogeography.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFC6FF93FF51D758E7A3FBA5.taxon	description	(Figs. 1 – 5) urn: lsid: zoobank. org: act: 142 A 465 D- 1 E 30 - 404 B- 9962 - CCC 7 A 10 D 3 FE 4	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFC6FF93FF51D758E7A3FBA5.taxon	materials_examined	Type locality. China, Taiwan Strait, Xiamen, Dadeji Beach (24 ° 26 ’ 30 ” N, 118 ° 4 ’ 12 ” E); intertidal zone, gravelly sand. Type material. Holotype adult female dissected on fifteen slides (OUCBL reg. no 2019.001). Paratypes are one adult male dissected on twelve slides (OUCBL reg. no 2019.002), ten female and ten male specimens preserved in 4 % formalin (OUCBL reg. nos 2019.003 – 022), one female dissected on one slide (OUCBL reg. no 2019.023) and one male dissected on one slide (OUCBL reg. no 2019.024). All type specimens collected from the type locality by Dr Er Hua on 24 July 2010.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFC6FF93FF51D758E7A3FBA5.taxon	etymology	Etymology. The specific epithet is derived from the Latin bi, meaning two, and the past participle of partire, meaning to part, and refers to the 2 - segmented condition of the male P 5 exopod.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFC6FF93FF51D758E7A3FBA5.taxon	description	Description of female. Body length 505 – 530 µm (mean = 515 µm, n = 10), semicylindrical, widest at posterior margin of cephalothorax, tapering posteriorly without clear distinction between prosome and urosome (Fig. 1 A – B). All somites with sensillae as illustrated, except for penultimate one. Conspicuous pores present on all somites. Rostrum (Fig. 1 A – B) defined at base, elongated, triangular and with pointed tip; reaching to middle of antennulary segment 2, with pair of subapical sensilla. Cephalothorax with plain hyaline frill; pedigerous somites bearing P 2 – P 4 with distinctly incised hyaline frills dorsally; genital double-somite and abdominal somites 2 – 3 also with incised frills all around posterior margin; incised frills consisting of dentate lappets (Figs 1 A – B, 2 A). P 4 - bearing somite with transverse rows of minute spinules dorsally. P 5 - bearing somite with spinular row around dorsal and dorsolateral posterior margin. Genital and first abdominal somite forming elongate double-somite; original segmentation marked by dorsolateral and lateral, internal, transverse chitinous ribs; ornamentation consisting of two pairs of dorsolateral spinule rows in anterior half, and almost continuous spinular row posteriorly (interrupted dorsally, ventral spinules markedly shorter than lateral ones); with additional rows of minute spnules laterally and dorsally (Figs 1 A – B, 2 A). Second abdominal somite with ventral spinular row in anterior half and almost continuous row of spinules near posterior margin (interrupted middorsally) (Figs 1 A – B, 2 A); with additional transverse rows of minute spinules ventrally and dorsally. Third abdominal somite with ventral row of spinules and additional transverse rows of minute spinules laterally and dorsally (Figs 1 A – B, 2 A). Anal somite with spinules either side of anal operculum and around ventral and lateral posterior margin (Figs 1 A – B, 2 A). Anal operculum (Fig. 1 B) semi-circular, withour ornamentation; anal frill with long hair-like extensions. Caudal rami (Fig. 1 C – E) slightly broader than long, with slightly oblique, transverse row of spinules on dorsal surface and wrapped around inner margin; with several spinules, laterally and ventrally, around bases of setae II – V; with seven setae; lateral and ventral surface with pore (Figs 1 D – E, 2 A). Seta I minute, positioned ventrally near outer margin (Fig. 1 E); seta II naked, originating from outer margin; seta III long and naked, positioned at outer subdistal corner, slender; terminal setae IV – V long and bipinnate (Fig. 1 A), with weakly developed fracture planes, seta IV about half as long as seta V; seta VI long and naked, originating from inner subdistal corner; seta VII triarticulated at base, arising from dorsal surface (Fig. 1 C). Antennule (Fig. 2 B – C) 9 - segmented; segments 1 – 6 slender with segment 1 more elongate than in other parastenheliid genera; segments 7 – 9 short, in particular segment 7; all segments without spinular ornamentation; all setae naked; with aesthetasc on segments 4 and 9; apical acrothek consisting of aesthetasc and two setae. Armature formula as follows: 1 - [1], 2 - [10], 3 - [6], 4 – [3 + (1 + ae)], 5 – [2], 6 - [3], 7 - [3], 8 - [0], 9 - [5 + acrothek]. Antenna (Fig. 2 D). Basis and proximal endopodal segment completely fused, forming allobasis; abexopodal margin with two spinular tufts but no armature elements. Exopod 2 - segmented; proximal segment with two setae, lateral one slender and naked, distal one long and unipinnate; distal segment with four setae: proximal one tiny and naked, apical margin with one unipinnate, spiniform element and two small naked setae. Endopod with sparse spinules along both lateral margins; lateral armature consisting of small naked element proximally and large unipinnate spine distally; distal armature consisting of one strong, unipinnate spine, four geniculate setae (longest one with large spinules around geniculation and fused at base to unipinnate seta) and one naked seta. Mandible (Fig. 3 A – B). Coxal gnathobase well developed, with complex cutting edge bearing several multicuspidate teeth, with strong tooth at ventral corner and short, unipinnate seta at dorsal corner. Basis elongate, with row of spinules on both anterior and posterior surfaces, with three setae near inner distal corner: one spiniform and unipinnate and two setiform and naked. Endopod elongate, 1 - segmented; with two lateral and four terminal setae (two of which fused at base); exopod minute, represented by one tiny seta arising from small protuberance. Maxillule (Fig. 3 C). Praecoxal arthrite with seven spiniform elements around distal margin, with two long setae on anterior surface, and two short elements near inner margin. Coxal endite cylindrical, with four naked setae; coxal epipodite represented by one seta. Basis with three setae on single endite; rami fused to basis but clearly discernible. Endopod 1 - segmented, with three naked, apical setae. Exopod 1 - segmented, with one short, naked and one long, unipinnate seta apically. Maxilla (Fig. 3 D). Coxa with three endites, proximal endite bilobate with one naked seta on inner cusp and two naked setae on outer cusp; middle and outer endite each with two naked setae and one basally fused pinnate spine; allobasal endite drawn out into strong, unipinnate spine and three setae (one of which minute); endopod welldeveloped, 1 - segmented, not completely discrete at base, with two naked setae. Maxilliped (Fig. 3 E). Armature of syncoxa as in male (see below; Fig. 5 H). Basis with two short, naked setae along palmar margin and few spinules around outer margin. Endopod represented by well-developed, unipinnate claw and three accessory, naked setae near its base. P 1 (Fig. 3 F). Intercoxal sclerite small, unadorned. Praecoxa (not illustrated) small, triangular, with spinule row on distal margin. Coxa with row of large spinules along outer margin and medial part of distal margin, and row of small spinules on anterior surface; with pore on anterior surface. Basis with unipinnate inner spine and bipinnate outer spine; with row of spinules near base of outer spine, along inner margin and around base of endopod. Exopod 3 - segmented, segments subequal in length, proximal segment with row of spinules along outer margin and unipinnate spine at outer distal corner; middle segment with row of spinules along outer margin and two setules along inner margin, with unipinnate spine at outer distal corner and short, plumose seta at inner distal corner; distal segment with plumose, inner distal seta, unipinnate, geniculate outer distal seta, and two unipinnate outer spines. Endopod 2 - segmented, prehensile; proximal segment much longer than exopod, with long, uniplumose seta arising from proximal fourth of segment; distal segment small, with few spinules, one minute inner seta and two slightly curved, naked spines apically. P 2 – P 4 (Fig. 3 G, 4 A – B). Praecoxa triangular, with spinule row along distal margin; coxa with 1 – 2 row (s) of spinules along outer margin and row of tiny spinules on anterior surface; basis with row of spinules near proximal inner corner, row of fine setules distally between insertions of rami, and few spinules around base of outer seta, inner distal corner forming small, pointed process. Rami 3 - segmented, slender; endopods about as long as (P 2) or slightly shorter than (P 3 – P 4) exopods; P 2 – P 4 enp- 3 longest, enp- 1 widest and with convex inner margin. All segments with rows of large spinules along outer margin; with few spinules / setules on inner margin of exp- 2, and enp- 2 and - 3; with anterior pore on all segments except for enp- 1; distal inner seta on P 3 – P 4 exp- 3 very small. Armature formulae of P 1 – P 4 as follows: P 5 (Fig. 4 C). Rami discrete and baseoendopods not fused medially. Endopodal lobe small, trapezoid; with five setae (four pinnate, one short and naked) and few spinules around base of outer basal seta; anterior surface with two pores. Exopod about two times longer than maximum width; basal portion articulating with baseoendopod very narrow; with row of setules along inner and outer margins; with six setae: one long, bipinnate, inner seta, two naked setae apically (one very long, other one short), and one short naked and two bipinnate setae along outer margin. Genital field (Fig. 2 A) with large midventral copulatory pore; paired genital apertures positioned anteriorly, each covered by vestigial P 6 bearing three setae, inner two short, outer one long and plumose. Description of male. Sexual dimorphism expressed in urosomal segmentation and ornamentation, antennule, endopods of P 2 – P 4, P 5 and P 6. Body slightly smaller than female, length 415 – 460 µm (mean = 440 µm, n = 10). Genital and first abdominal somites not fused. Urosomal ornamention very similar to female, except for absence of transverse spinule row on ventral and lateral surfaces of penultimate somite. Antennule (Fig. 5 A – B) indistinctly 11 - segmented, haplocer; segment 1 elongate; segment 4 largely incorporated into moderately swollen segment 5; geniculation located between segments 7 – 8 with four segments distal to it; segment 11 with surface suture along dorsal surface; aesthetasc on segments 5 and 11; all setae naked except for bipinnate seta on segment 1; armature formula as follows: 1 - [1], 2 - [10], 3 - [8], 4 - [2], 5 - [4 + (1 + ae)], 6 - [2], 7 - [4 + 3 modified spines], 8 - [1 + 2 modified spines], 9 - [1], 10 - [2], 11 - [6 + (1 + ae)]. Maxilliped (Fig. 5 H). As in female, syncoxa with two setae near inner distal corner. P 2 (Fig. 5 C). Exopod as in female. Endopod 2 - segmented; compound enp- 2 with only one plumose inner seta, arising from proximal third of segment (homologue of inner seta of ♀ enp- 2). P 3 (Fig. 5 D). Exopod as in female. Enp- 3 shorter than in female, without inner setae. P 4 (Fig. 5 E). As in female except enp- 3 slightly shorter. Armature formulae of P 1 – P 4 as follows: P 5 (Fig. 5 F). Baseoendopods fused medially, forming deeply incised transverse plate. Endopodal lobe with row of spinules along outer margin; with two short elements (inner one unipennate and spiniform, outer one naked and shorter than inner one); anterior surface with two pores; outer basal seta bipinnate and arising from short setophore. Exopod 2 - segmented; exp- 1 with row of spinules along outer margin and naked seta at outer distal corner; exp- 2 with short naked seta at outer subdistal corner, long bipinnate seta apically and unipinnate spine at inner subdistal corner; anterior surface of exp- 2 with pore. P 6 (Fig. 5 G) represented on each side by small plate closing off genital aperture; with three setae, middle seta about twice longer than other two, innermost one unipinnate. Variability. In one dissected male the P 2 enp- 2 displayed two inner setae on one side.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFC6FF93FF51D758E7A3FBA5.taxon	discussion	Remarks. The new species cannot be placed in Parastenhelia, Microthalestris, Karaytugia gen. nov. or Penicillicaris gen. nov. since it does not display the sexual dimorphism on the P 3 endopod (enp- 3 with spinous apophysis; see Fig. 7 A – F) that characterizes these four genera. The absence of sexual dimorphism in the male antenna also excludes the species from assignment to Thalestrella and the lack of a transformed inner base spine in the male P 1 prevents it from being accommodated in Foweya, Paraleptomesochra or Psammoleptomesochra. Johnwellsia bipartita sp. nov. appears to be morphologically closest to two species that were previously assigned to Parastenhelia, P. megarostrum and P. pyriformis, but which will be transferred to a new genus Porirualia gen. nov. below. Both species and J. bipartita sp. nov. share the same sexual dimorphism on P 3, involving the loss of setation elements on male enp- 3. In members of Porirualia gen. nov. the proximal inner seta of enp- 3 is lost in the male while the further derived state (involving the additional loss of the distal inner seta) is observed in Johnwellsia gen. nov. (Fig. 7 G – I). Two additional synapomorphies supporting a sistergroup relationships between these genera are the modification of the distal seta on the mandibular basis into a short spine, and the reduction of the outermost endopodal element of the P 5 in both sexes, being represented by a minute seta. Johnwellsia gen. nov. differs from Porirualia gen. nov. in the following characters: (a) rostrum reaching to middle of antennulary segment 2 (vs very large, reaching to at least halfway the fourth antennulary segment in the female); (b) antennule ♀ very slender with elongate segment 1 (vs more compact and segment 1 short); (c) antennary allobasis without abexopodal seta; (d) mandibular exopod rudimentary, represented by single seta on minute protuberance (vs 1 - segmented with three setae); (e) maxillipedal basis with two setae (vs one seta); (f) P 1 enp- 1 inner seta long but not extending beyond distal margin of segment; (g) P 2 endopod ♂ sexually dimorphic (2 - segmented and distal inner seta of ♀ enp- 3 lost in ♂; Fig. 6 C); (h) P 5 ♀ without series of transverse striae along inner margin of endopodal lobe; and (i) P 5 exopod ♂ 2 - segmented, with four elements in total (vs 1 - segmented, with 5 – 6 elements). Species of both genera also differ in the shape of the female P 5 exopod which is moderately elongate and shares a very narrow articulation with the baseoendopod in Johnwellsia gen. nov. (Fig. 4 C). The 2 - segmented P 5 exopod in the male is an unusual condition in the family, being found only in Parastenhelia willemvervoorti sp. nov. and Karaytugia aydini (Kuru & Karaytuğ, 2015) comb. nov. (Vervoort 1964; Kuru & Karaytuğ 2015; Table 1 herein).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFCEFF8DFF51D61EE7BCFCDD.taxon	discussion	The description of two Sri Lankan species, P. hornelli and P. similis, by Thompson & Scott (1903) marked the start of the historical division of the genus into two groups, the spinosa - group, characterized by a distinctly elongate P 1 exp- 2, and the hornelli - group, which unites species with subequal segments in the P 1 exopod. Various species have been added since then to either group, almost solely on the basis of P 1 morphology, without considering other characters that could be of taxonomic or phylogenetic significance. Attempts to delimit species boundaries within Parastenhelia have historically been made by two opposing factions, i. e. lumpers (Lang 1948; Pesta 1959; Vervoort 1964; Wells & Rao 1987; Mielke 1974, 1990) versus splitters (Sewell 1940; Gee 2006; Kuru & Karaytuğ 2015). Proponents of the first school of thought were primarily influenced by Lang (1948) who basically considered P. spinosa an amalgam of forms in which the middle segment of the exopod of P 1 is at least four times as long as broad. Wells & Rao (1987) pointed out that P. spinosa, as revised by Lang, is one of the most variable species of harpacticoids, and that no clearcut distinctions can be made between the various forms that he included. Unfortunately, the adoption of his system as the best approach to the complex taxonomy of the genus has generated tremendous confusion. An extreme example is presented by Vervoort’s (1964) unwarranted recognition of extreme variability in specimens attributed to P. spinosa from the Caroline Islands. His grouping of two different forms – one with subequal exopodal segments in the P 1 (hornelli type), the other with a distinctly elongate P 1 exp- 2 (spinosa type; note that his Fig. 70 b depicts the exopod not the exopod as stated in the figure legend) – into P. spinosa defies any serious attempts to delimit species boundaries in the genus. As pointed out by Wells et al. (1982) the relative stability displayed by the proportions of the P 1 exopodal segments in P. spinosa must invalidate Vervoort’s inclusion of a form which by his own admission approaches P. hornelli in several respects. In his search of apomorphies that could define the genus Parastenhelia as a monophyletic lineage, Mielke (1990) highlighted the significance of the sexual dimorphism of the P 3 endopod, involving the transformation of the outer spine on the distal segment into a fused apophysis and a reduction in the number of inner setae (to one in P. bulbosa and none in all other species) (Fig. 7). The absence of this character in P. reducta and P. megarostrum led him to claim that they do not belong to Parastenhelia. This view was corroborated by Gee (2006), who moved P. reducta as species incertae sedis to Karllangia (= Thalestrella) and suggested that both P. megarostrum and P. pyriformis should be assigned to one or more new genera, a course of action that is formalized here (see below). Various authors have pointed out that a revision of the genus is hampered by the taxonomic confusion surrounding its alleged type species P. spinosa (Vervoort 1964; Wells & Rao 1987; Gee 2006; Kuru & Karaytuğ 2015). However, it has recently come to light that Lang’s (1948) fixation of P. spinosa as the type was invalid since it was not originally included (ICZN Art. 69.2) and that P. hornelli had already been fixed as the type species by subsequent designation (Lang 1934: 24; Huys 2009: 96). This implies that any assessment of the monophyly of Parastenhelia must first centre on a review of P. hornelli (and allied species – hornelli group) since it provides the objective standard of reference for the application of the generic name it bears. Consequently, the genus is here restricted to a core group of four valid species which share a P 1 exopod with three subequal segments in both sexes. Males of these species share a haplocer antennule with three segments distal to the geniculation, a 2 - segmented P 2 endopod (enp- 2 and - 3 forming a compound segment but total armature as in ♀) and a modified P 3 endopod with spinous apophysis on enp- 3 (Table 1; Figs 6 – 7). Thompson & Scott (1903) described two Parastenhelia species from pearl oyster washings in the Gulf of Mannar off northwestern Sri Lanka: P. hornelli, which was represented by several females and males and was also found in general washings of dredged material in deep water off Galle (southwestern Sri Lanka), and P. similis of which only two females were recorded. The two species were considered morphologically similar with females being distinguished on the basis of proportional length differences in the antennulary segments (the whole appendage being distinctly longer and more slender in P. hornelli), the morphology of the maxilliped, and the shape of the P 5 exopod. Additional discrepancies are found in the P 1 with the endopod being more slender, the inner seta of enp- 1 distinctly longer, and the distal claws on enp- 2 markedly longer (relative to the segment) in P. hornelli. Lang (1934) dismissed all these points and sank P. similis as a junior subjective synonym of P. hornelli. Since no variability in the antennule, P 1 and P 5 has so far been documented we regard these differences as sufficient justification to reinstate P. similis as a valid species. Wells & Rao (1987) assessed the “ hornelli group ” based on material from Mozambique, the Andaman Islands and New Zealand. They redescribed both sexes of P. hornelli from specimens of Middle and South Andaman, recognized two distinct size morphs among the female specimens (small form about 630 μm; large form about 885 μm; all ♂♂ 650 μm) and opted for the interim solution to place specimens previously identified as this species by Noodt (1955 a) and Apostolov (1973), as well as the material identified as P. spinosa by Vervoort (1964), in P. hornelli. Careful inspection of Wells & Rao’s (1987) illustrations, however, shows that they were not dealing with P. hornelli but with P. similis. Evidence in support of this claim is found in the female morphology of the antennule (general facies and relative lengths of segments), maxilliped (palmar margin with conspicuous row of long spinules), P 1 (length of endopod, inner seta of enp- 1 and claws on enp- 2), P 5 exopod (exact shape and strong spinular ornamentation along inner and outer margins) and caudal rami (setae IV – V slightly swollen and irregularly shaped at base). The three differences between Thompson & Scott’s (1903) type material of P. similis and Wells & Rao’s (1987) specimens are (a) segmentation of female antennule (the minute segments 7 – 8 in Wells & Rao’s Fig 52 e are shown as complelety fused in the original description), (b) absence of the small distal inner seta on P 4 exp- 3, and (c) absence of short inner seta on the endopodal lobe of P 5, both of which are conceivable the result of imperfect observation. The records of P. hornelli from Odisha (formerly Orissa) in the Bay of Bengal (Rao 1989), Lakshadweep (Rao 1991) and Little Andaman (Rao 1993) are accompanied by very brief, mostly uninformative descriptions and must be considered as unconfirmed. Other Indian records from Bhitarkanika in Odisha (Chadha & Kar 1999), Kundugal channel (Krishnaswamy 1953) and Porites Bay (Krishnaswamy 1957) in Tamil Nadu, and the Andaman Islands (Rao 1993; Jayabarathi 2016; Jayabarathi et al. 2012, 2015; Pongener et al. 2018) are equally indeterminable. Noodt (1955 a) provides a brief redescription, completely lacking in illustrations, of two females from the Sea of Marmara in northwestern Turkey which he attributes to P. hornelli. He states that both P 1 and P 5 agree exactly in form and armature with the illustrations of Thompson & Scott (1903) but it is not clear which of the Sri Lankan species he refers to. Since the shape of the exopod is different between P. hornelli and P. similis, Noodt’s (1955 a) statement that it is widening proximally towards the baseoendopod (“ Insbesondere ist der Exp. P 5 durch seine Erweiterung zur Basis hin typisch ”) may carry some significance since this feature is more expressed in P. hornelli (Thompson & Scott 1903: Plate VII, Fig. 8). Noodt’s (1955 a) report that the antennary exopod is 3 - segmented must be an observational error since it is 2 - segmented in the groundpattern in the Parastenheliidae (Willen 2000); he also states that the proximal segment bears one seta but does not disclose how many elements are present on the remaining segments. Noodt (1955 a) presents the armature pattern of P 2 – P 4, noting variability in the presence / absence of the inner seta on P 2 exp- 3. Although Wells & Rao (1987) pointed out that the pattern of P 4 (exp: 1.1.223; enp: 1.1.221) is congruent with that given by Thompson & Scott (1903) (for both P. hornelli and P. similis) it remains to be confirmed whether the presence of only two inner setae on P 3 – P 4 exp- 3 in the Turkish specimens is genuine or reflects an oversight of the fine inner distal seta. Based on Noodt’s description no positive statement can be made as to the validity of his identification and pending re-examination of the material (and discovery of the male) his record must be considered suspect. The unillustrated records of P. hornelli from two beaches (Cunda Beach and Altınova Beach) in the Aegean Sea (Karaytuğ & Sak 2006), in close proximity to the Sea of Marmara, possibly refer to the same species. Although Vervoort (1964) voiced criticism of Lang’s (1948) broad taxonomic concept of P. spinosa, he indirectly expressed the opinion that both P. hornelli and P. ornatissima may also fall within the range of this species. Among the material from Ifalik Atoll, Vervoort describes a male and a female which he assigns to P. spinosa while admitting that they approach P. hornelli in the structure of the P 1 exopod, where the three segments have about the same length. Vervoort (1964) dismisses the significance of this difference by accepting Lang’s (1948) view that it falls within the variability of P. spinosa – a concept that is mistakenly based on comparison of geographically separated and, most likely, non-conspecific populations. Since intrapopulation variability in the length of P 1 exp- 2 has never been recorded in any species so far, Vervoort’s (1964) interpretation is rejected here. His material displays a number of distinct differences with both P. hornelli and P. similis: (a) P 3 – P 4 exp- 3 without inner seta; (b) in addition to the common sexual dimorphism expressed in the endopods of P 2 – P 3, the male P 4 endopod also differs from that of the female in having only one inner seta on enp- 3; this type of sexual dimorphism has previously been observed in Microthalestris littoralis f. penicillata sensu Willey (1935) and Parastenhelia bulbosa (Gee 2006), and as a further derived state in P. minuta and Penicillicaris penicillata sp. nov. where both inner setae are lost in the male (Willey 1935; Pallares 1982; Fig. 6 F – G herein); and (c) male P 5 exopod 2 - segmented; this condition was recently described in P. aydini (Kuru & Karaytuğ 2015) and in Johnwellsia bipartita gen. et sp. nov. (present account). Based on these differences it is obvious that Vervoort’s (1964) material from stations 589 and 590 cannot be attributed to P. spinosa or to either of Thompson & Scott’s (1903) species and must be assigned distinct specific rank (see below). Wells et al. ’ s (1982) claim that Vervoort’s description and illustrations of the antenna are contradictory is irrelevant. His lateral habitus view (Fig. 68 a) shows indeed the antennary exopod as 3 - segmented under low magnification but the detailed drawing of the antenna (Fig. 68 d) clearly reveals it as 2 - segmented (as reiterated in the text: p. 186). Apostolov (1973: 273, Fig. 9) reported two females and one male of P. hornelli on brown algae (Cystoseira barbata) near Tsarevo (formerly Michurin) along the Black Sea coast of Bulgaria. No text description was given, except for his statement that the male fits Thompson & Scott’s (1903) original description perfectly. Apostolov (1973) provides illustrations of the male P 2 endopod (similar to that illustrated by Thompson & Scott), P 3 endopod (previously unknown) and P 5. The latter differs significantly in that it is only 1 - segmented and the armature elements all are very short, consequently ruling out the possibility that the Bulgarian material belongs to either P. hornelli or P. similis. Similar to Noodt’s (1955 a) specimens the antennary exopod is erroneously illustrated as 3 - segmented causing Wells & Rao (1987) to suggest that both sets of specimens are conspecific. The female antennule and P 5 exopod resemble their respective conditions as illustrated by Thompson & Scott (1903) for P. hornelli, the P 4 also displays the same armature pattern (although confirmation of the exact number of inner setae on exp- 3 is required in both), but the P 1 endopod is comparatively longer and more slender. The only significant difference with Noodt’s (1955 a) material is found in the absence of the inner seta on P 2 enp- 1 (a condition that is mirrored in the male). In summary, Apostolov’s (1973) Bulgarian specimens share several characters with P. hornelli from Sri Lanka but the 1 - segmented condition of the male P 5 exopod and deficiencies in the description of the antenna (and possibly P 3 – P 4 exopods) prevent them from being assigned to the type species. Pending a re-examination of the Black Sea material, P. hornelli sensu Apostolov (1973) must be classified as indeterminable. Apostolov & Pandourski (1999) reported one female and one male of P. hornelli from a littoral sandbank in Livingston Island (South Shetlands archipelago), north of the Antarctic Peninsula (see also Pesce & Pandourski 2002: 134). The authors do not provide a text description but state that their material confirms the variability previously observed in this species and cite the different number of setae and spines on P 3 exp- 3 and enp- 3 as an example. Although they do not cite the work it is likely that Apostolov & Pandourski (1999) used Wells & Rao’s (1987) redescription of P. hornelli (= P. similis) as a reference for comparision. Their illustration of the female P 3 endopod (they do not figure the exopod) shows three well developed inner setae on the distal segment which contradicts the condition in P. similis (Table 1). The female P 5 exopod approaches the shape (although more truncated) of Thompson & Scott’s (1903) P. hornelli while the length of the endopodal setae is different. The morphology of the male is said to be identical with P. hornelli but only the P 2 endopod is illustrated; the proximal segment of this limb lacks the inner seta found in the type species. No information was given on the armature pattern of P 2 – P 4 (except P 3 endopod ♀) or the segmentation and setation of the male P 6. Apostolov & Pandourski’s (1999) description is too fragmentary to allow any conclusions to be made about its identity and consequently it must be classified as an indeterminable record of P. hornelli. Based on Wells & Rao’s (1987) report the only confirmed records of P. similis are from Sri Lanka (Thompson & Scott 1903), Inhaca Island in Mozambique (Wells 1967), northeastern New Zealand (Wells et al. 1982) and the Andaman Islands (Wells & Rao 1987). Authenticated records of P. hornelli remain restricted to the type populations from Galle and the Gulf of Mannar in Sri Lanka (Thompson & Scott 1903). This is in stark contrast with the previously assumed circum-subtropical distribution (Wells et al. 1982; Wells & Rao 1987) which included unconfirmed and (possibly) false records from the Aegean / Black Sea region (Noodt 1955 a; Apostolov 1973), Barbados (Coull 1970 a) and the U. S. Virgin Islands (Coull 1971 a; Hartzband & Hummon 1974) in the Atlantic, and the east coast of Peninsular Malaysia (Zaleha et al. 2006) and the Caroline Islands (Vervoort 1964) in the Pacific. The armature pattern of P 2 – P 4 given by Zaleha et al. (2006) shows that they were dealing with a different species. In addition to the unconfirmed Indian records mentioned above, P. hornelli has also been reported from Gazi Bay in Kenya (De Troch 2001 – as P. cfr. hornelli) and along the Pacific seaboard of North America including the Nanaimo River delta (Sibert 1981; Kask et al. 1982) and Fraser River estuary (Sibert et al. 1982; Bravender et al. 1993; Johansen & Reis 1994) in British Columbia, Padilla Bay (Simenstad et al. 1988) and Puget Sound (Simenstad et al. 1991) in Washington State and the Santa Barbara Channel in southern California (Spies et al. 1988). All these records are unillustrated and must be considered as unconfirmed and probably false. Parastenhelia oligochaeta has so far been recorded from the Andaman and Nicobar Islands (Chidyatapu, Mayabandar, Car Nicobar, Katchal and Great Nicobar Islands – Wells & Rao 1987) and from Bhitarkanika in Odisha, mainland India (Chadha & Kar 1999).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFCEFF8DFF51D61EE7BCFCDD.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 2 – P 3 endopods, P 5 – P 6 and urosomal segmentation; occasionally P 4 endopod and caudal ramus setae IV – V. Body subcylindrical, short; posterior margin of cephalothorax, somites bearing P 2 – P 5 and remaining urosomites (except anal somite) with denticulodigitate hyaline frills. Rostrum defined at base, of moderate size, reaching at most to distal margin of second antennulary segment; linguiform. Anal operculum semicircular, naked or bordered with fine spinules. Caudal ramus wider than long, without conspicuous spinular ornamentation on dorsal surface; with at least six setae (seta I possibly absent), setae IV – V slightly swollen at base in ♀ in some species (P. similis). Antennule elongate and 8 - or 9 - segmented in ♀, segment 1 not elongate, segments 7 – 8 (when 9 - segmented) shortest, with aesthetascs on segments 4 and 8 or 9; haplocer in ♂ but segmentation and segmental homologies unconfirmed, with three segments distal to geniculation, segment 5 swollen and with aesthetasc. Antenna not sexually dimorphic; proximal endopodal segment largely separated from basis or forming allobasis, with pinnate seta on abexopodal margin; exopod 2 - segmented, proximal segment with two setae, distal segment with 1 – 2 lateral and three apical elements; distal endopodal segment without penicillate elements. Mandible with 2 – 4 elements on basis; endopod with eight setae; exopod 1 - segmented with three setae or represented by single seta arising from minute knob. Maxillulary coxal epipodite represented by one seta. Maxilla with three endites on syncoxa; endopod discrete, with 2 – 3 setae. Maxilliped with 2 – 3 setae on syncoxa; basis with 1 – 2 seta (e) on palmar margin; endopod represented by curved claw, accompanied by 1 – 2 accessory seta (e). P 1 inner basal spine not sexually dimorphic. P 1 exopod 3 - segmented; exp- 2 about as long as others, typically with short inner seta; exp- 3 small, with two pinnate spines, one geniculate seta and one naked seta. P 1 endopod 2 - segmented; enp- 1 elongate, about 1.3 – 1.5 times length of exopod, with long, pinnate, inner seta inserted in proximal third, segment margins without area of reduced chitinization; enp- 2 very small, with one naked minute seta and two dentate claws. P 2 – P 4 rami 3 - segmented. P 2 endopod ♂ 2 - segmented; total number of spines / setae as in ♀. P 3 endopod ♂ 3 - segmented; without inner setae on enp- 3 and outer spine modified into spinous apophysis. P 4 endopod ♂ occasionally with one less inner seta on enp- 3 (P. willemvervoorti sp. nov.). Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe with five setae, outermost shortest; inner margin without transverse striae. P 5 ♀ exopod elongate, with six elements. P 5 ♂ endopodal lobe with two elements, outer one shortest; exopod 1 -, 2 - or 3 - segmented, with 6 – 7 elements in total. Vestigial P 6 ♀ represented by two minute setae. P 6 ♂ with three setae.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFCEFF8DFF51D61EE7BCFCDD.taxon	type_taxon	Type species. Parastenhelia hornelli Thompson & Scott, 1903 (by subsequent designation; Lang 1934: 24).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFCEFF8DFF51D61EE7BCFCDD.taxon	discussion	Other species. P. similis Thompson & Scott, 1903; P. oligochaeta Wells & Rao, 1987; P. willemvervoorti sp. nov. Species inquirendae. Parastenhelia hornelli Thompson & Scott, 1903 sensu Noodt (1955 a), sensu Apostolov (1973), sensu Apostolov & Pandourski (1999) and sensu Zaleha et al. (2006).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFD0FF8DFF51D054E6B1F93E.taxon	description	Original description. Vervoort (1964): 181, 184 – 188; Figs. 66 a – b (♂ – from locality 590), 68 a – c (♀ – from locality 589), 69 a – f (♂ – from locality 590) [as Parastenhelia spinosa].	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFD0FF8DFF51D054E6B1F93E.taxon	materials_examined	Type material. The dissected male specimen illustrated by Vervoort (1964: 181, 187) in his figures 66 (a – b) and 69 (a – f) and collected on 16 October 1953 is here designated as the holotype of P. willemvervoorti sp. nov. (ICZN Arts 16.4 and 72.5.6). The species can be differentiated by the characters listed in the diagnosis below and those mentioned and illustrated in Vervoort (1964) (ICZN Art. 13.1). Type locality. Federated States of Micronesia, Caroline Islands, Ifalik (Ifaluk) Atoll; seaward reef about middle of Falarik. Vervoort (1964) collected males from two different stations (589 and 590). The male holotype came from station 590 (sand sample taken about 24 m from reef margin).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFD0FF8DFF51D054E6B1F93E.taxon	diagnosis	Differential diagnosis. Parastenhelia. Body length 290 – 470 μm in ♀, 270 – 650 μm in ♂. Rostrum as long as antennulary segments 1 – 2 combined. Antennule 8 - segmented in ♀; haplocer in ♂ but segmentation unconfirmed, with three segments distal to geniculation. Antenna with 2 - segmented exopod bearing two setae on exp- 1 and one lateral and three apical elements on exp- 2. P 1 exopod about 0.6 times length of endopod; exp- 2 not elongate, slightly longer than exp- 3. P 1 enp- 1 about 1.5 times length of exopod. P 3 – P 4 exp- 1 without inner seta, exp- 3 with three inner setae. P 2 endopod ♂ 2 - segmented; enp- 2 elongate, reaching apex of exopod, with additional inner seta [1.221 vs 1.1. 121 in ♀]. P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]; P 4 endopod ♂ 3 - segmented, with only one inner seta on enp- 3 [1.1.121 vs 1.1. 221 in ♀]. Armature of P 1 – P 4 – see Table 1. P 5 ♀ with elongate exopod (about 2.5 times as long as wide), with six elements; armature of endopodal lobe unknown. P 5 ♂ 2 - segmented, with one outer seta on exp- 1 and six elements on exp- 2; endopodal lobe with two elements. P 6 ♂ with three setae. Anal operculum smooth. Caudal ramus seta IV – V ♀ not swollen at base.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFD0FF8DFF51D054E6B1F93E.taxon	etymology	Etymology. The species is named after the late Prof. Willem Vervoort (12 June 1917 – 18 August 2010), eminent copepodologist, who discovered this species.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFD7FFBCFF51D13AE09BFBFD.taxon	discussion	Various authors have reported on what they individually regard as Parastenhelia spinosa or varieties / species sharing some of its morphological characteristics, but recognise a wide range of variation in features such as segmentation of the female antennule, modification of the caudal ramus setae (particularly V), setation of the antennary exopod and endopod, relative proportions of P 1 rami and form of their terminal elements, armature formulae of P 2 – P 4, shape and armature of ♀ P 5 exopod, and segmentation of ♂ P 5 exopod. The magnitude of this variability, or the identity of differences which might represent true genetic discontinuity (and thus distinct species), is in many cases impossible to determine due to the inaccurate and / or incomplete descriptions and illustrations [especially the original description by Fischer (1860)] which have accompanied many published reports. The currently accepted concept of P. spinosa being a highly variable, cosmopolitan taxon has grown over time since 1860 by the gradual accumulation of differences observed in what were wrongly assumed to be geographically isolated populations of the same species. The historical review presented below aims at (a) pointing out some important misconceptions in the taxonomy of the species, (b) assigning distinct specific status to some of its radically divergent “ populations ”, and (c) formulating a working hypothesis and baseline for future comparisons and revisionary work. Parastenhelia spinosa was first described as Harpacticus spinosus by Fischer [1860: 665 – 666, Table XXI (figures 51 – 53), XXII (figure 66)] from the island of Madeira [Lang (1948: 589) stated incorrectly that Fischer did not disclose the type locality] but the description is severely lacking in any real detail (8 - segmented antennule, ovate female P 5 with four spines on exopod and baseoendopod, caudal ramus seta V swollen at base) and only his illustration of the P 1 provides the barest minimum to identify it as a member of the Parastenheliidae. The inaccuracies expressed in the few poorly rendered drawings (maxilliped, P 1, female P 5, caudal rami) and the lack of the male make the description virtually useless for comparative purposes. Harpacticus spinosus is placed in Microthalestris solely on account of its elongate P 1 exp- 2 but cannot be treated as anything more than a species inquirenda in this genus. Had Lang’s (1948) fixation of H. spinosus as the type of Parastenhelia been valid, this would have threatened stability and universality in the application of the name of the type species since its taxonomic identity cannot be determined from its name-bearing type. Although Lang (1936 a: 21) considered the possibility that Fischer’s (1860) species was conspecific with Parastenhelia forficula var. littoralis (sensu Sars 1911) he recommended to set it aside on the grounds that it cannot be identified with any degree of confidence. Unfortunately, he changed his opinion by not only reinstating it as a valid species but also by illegitimately fixing it as the type of the genus (Lang 1944: 13; 1948: 586 – 588). Claus (1863: 131 – 132, Plate XVII, figs 7 – 11) described Thalestris forficula Claus, 1863 as one of ten species assigned to his new genus Thalestris Claus, 1862 (type species by subsequent designation: Thalestris longimana Claus, 1863). The author reluctantly included H. spinosus in the genus as well as two other Harpacticus species previously described by Fischer (1860). His description of T. forficula, based on material from Messina (Sicily), included illustrations of the antennule of both sexes, P 1, caudal rami and the female genital field. The most significant information that can be extracted from Claus’s (1863) concise description is the 8 - segmented condition of the female antennule, the presence of an apophysis on the male P 3 endopod, the modified caudal seta (V) displaying a basal swelling, and the length of the relatively short inner seta on P 1 enp- 1 which extends only to about 40 % of the segment length. No information was given about the armature of P 2 – P 5 in both sexes, or the segmentation of the male P 5 exopod. ...... continued on the next page 17 Based on Lang’s (1936 a: Fig. 37) supplementary description (as Pa. gracilis). 18 Pallares (1982: 10) erroneously stated that P 3 – P 4 exp- 3 display a [333] pattern in the male; probably not having consulted Pallares’s drawings of the male, Mielke (1990: 166) accepted this as an additional difference between the type population and his material collected from Bahía Lapataia near Ushuaia (Argentina). 19 Armature pattern of enp- 3 021 in ♂. 20 0.1.12 + apo in ♂. 21 The single ♀ observed by Mielke (1974: 20) showed an aberrant setal pattern on one side of P 2 enp- 2 (with two inner setae) and P 4 exp- 3 (with inner seta). 22 There is confusion over the exact number of elements on the exopod. Pallares (1963) states that there are three outer, one inner and two apical elements but in a subsequent account (Pallares 1968: 65) explicitly states that there are seven without going into detail about their position. We have adopted Pallares’s original observation pending reexamination of new material. 23 Lang’s (1936 b: 24) only female displayed the atypical 121 pattern on enp- 3. 24 Mielke (1990) recorded a high degree of variability in the armature of P 2 – P 4. Some specimens displayed (a) an inner seta on P 2 – P 3 exp- 2, (b) only one (instead of two) inner seta on P 3 exp- 3, (c) two (instead of one) inner setae on P 4 exp- 3, (d) two (instead of one) inner setae on P 3 – P 4 enp- 3, and / or (e) only two (instead of three) outer spines on P 4 exp- 3. One male lacked the inner seta on P 4 enp- 2. 25 According to Wells (2007: 596) the original report of three inner setae on P 2 exp- 3 is wrong. 26 The endopodal armature pattern differs from Mielke’s (1994 a: 257) in two aspects; the apical spiniform element on enp- 3 is here considered as the homologue of the outer spine in other genera, and the two minute and fine setae originating from the inner corner of enp- 3 are included in the formulae. 27 Inner seta minuscule (Mielke 1994 a: Fig. 4 C). 28 2 - segmented in ♂ (1.321). 29 1.1. 321 in ♂; Kunz (1975) did not illustrate the ♀ condition but listed the armature formula of P 3 endopod as 1.1. 221 in his Table 5; Gee (2006) re-examined the type material and noted that the armature formula is as in F. anglica. 30 2 - segmented in ♂ (1.121). 31 1.1.021 in ♂. 32 Willey (1935) reported some specimens with a 3 - segmented endopod in the ♀ (1.1.221). 33 P 3 endopod ♂ and P 5 ♂ based on Vervoort’s (1964) description of Pa. spinosa [partim]. According to Sewell (1940: Text-fig. 27 H) P 2 exp- 1 lacks an inner seta but Vervoort shows a small inner element which is included in the armature formula here. 34 Scott (1894 a: Plate 12, Fig. 39) most likely overlooked the short inner seta on exp- 3. 35 Sewell (1940: 196) claims the proximal inner seta was overlooked in Scott’s (1894 a) description, bringing the total to five. 36 1.221 in ♂. In a paper written in old Danish and completely lacking in illustrations, Boeck (1865: 266 – 267) described a new species, Thalestris karmensis Boeck, 1865, based on specimens collected from Karmøy in Norway; this species fell into oblivion until it was discussed by Sars (1905 – see below). In a similar contribution, Boeck (1873: 56) provided a short unillustrated text description of Dactylopus longipes Boeck, 1873, obtained from 29 m depth in the Oslofjord in Norway. His female specimens were characterized by a 9 - segmented antennule, a long and slender P 1 exopod which is shorter than enp- 1, a 2 - segmented P 1 endopod with a very long enp- 1 which bears a seta in the proximal quarter of the inner margin, and a P 5 exopod with eight elements. Based on this short description Lang (1936 a: 31) would later synonymize D. longipes with Thalestris forficula which he had previously placed in Parastenhelia (Lang 1934: 24). Thomson’s (1883: 104 – 105; Plate X, Figs 16 – 21) material of Thalestris forficula from Otago Harbour in New Zealand agrees with the description of Claus (1863) in the 8 - segmented condition of the female antennule and the basally inflated setae V on the caudal ramus but differs in size (1,000 vs 800 μm) and the morphology of P 1. The female P 5 bears six and five setae on the exopod and endopodal lobe, respectively. Lang (1934: 25) lists Thomson’s record but does not appear to question its identity. In later accounts he accepts it as a record of either Parastenhelia forficula var. littoralis (Lang 1936 a: 52) or P. spinosa (Lang 1948: 588). Car (1884: 248 – 249, Plate XVIII, Figs 1 – 8) provided an illustrated description of a new species, Thalestris pectinimana Car, 1884, based on a single ovigerous female collected in the vicinity of Trieste, northeastern Italy. The author did not compare T. pectinimana with other congeners nor did he present a justification for its recognition as a distinct species. Illustrations were provided for the female in lateral aspect, antennule, antenna, maxilliped, distal segments of the P 1 rami and P 5. The species is characterized by the 8 - segmented antennule, the large P 5 with seven setae on the foliaceous exopod and five elements on the endopodal lobe, and the presence of distinct pectinate spines on the terminal segments of P 1 exopod and endopod. No information was given on the armature of P 2 – P 4. The species has not been recorded again since its original description. Pesta (1920: 590) listed it as a junior subjective synonym of Phyllothalestris mysis (Claus, 1863) but this was rejected by Lang (1936 a: 49) who considered it a valid species of Thalestris. Lang (1948: 497, 588) revised his previous opinion and considered T. pectinimana conspecific with Parastenhelia spinosa, more specifically (based on P 5 morphology) with the form described as Microthalestris littoralis f. penicillata by Willey (1935: 82). This claim is dubious since Willey did not describe the female P 5 but only stated that it displays “ … the fifth foot of littoralis ”. According to Sars’s (1911: Suppl. Plate 11) description of M. littoralis the female P 5 has only six setae on the exopod while T. pectinimana exhibits seven.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFD7FFBCFF51D13AE09BFBFD.taxon	discussion	Scott & Scott (1894: 142 – 144, Plate IX, Figs 4 – 9) described Thalestris forficuloides from mud near the lowwater mark at Seafield in the vicinity of Leith, Firth of Forth, Scotland. An expanded description of the female and additional illustrations of the male were provided by Scott (1894 b: 255 – 256, Plate X, Figures 13 – 25) based on specimens from the type locality (note that the mandible and maxillule were transposed in his Plate X). As the name suggests this species was considered close to T. forficula, displaying differences in the segmentation of the female antennule (9 - segmented vs 8 - segmented in T. forficula), the length of the P 1 endopod, and some unspecified discrepancies in the ” … proportional lengths of the other thoracic feet ” (Scott & Scott 1894: 144). Sars (1905: 122) proposed a new genus Microthalestris in the Thalestridae to accommodate T. forficula as its type and only species and stated that the forms recorded by Boeck (1865) as Thalestris karmensis and by Scott & Scott (1894) as T. forficuloides, both belong to M. forficula (Claus, 1863). His redescription of the latter (Sars 1905: 123 – 124, Plate LXXVI) corresponds exactly to the illustrations of Scott (1894 b) in the segmentation of the antennule, morphology of P 1, and the basally inflated caudal ramus seta (V), however, also shows a number of important differences with T. forficuloides including (a) P 3 exp- 1 with inner seta instead of without, (b) P 3 exp- 3 with two inner setae instead of three, (c) P 5 exopod ♀ comparatively longer, and (d) P 5 exopod ♂ 3 - segmented instead of 1 - segmented. Although Sars was not specific about the total number of setae on the male P 5 exopod it appears from his illustration that there are only six as opposed to the seven recorded in later descriptions. There is a hint of a short setal element among the spinular cluster on the outer margin of exp- 1, however, the seta usually found in that position is typically very long (e. g. Chislenko 1967: Fig. 45; Mielke 1974: Fig. 9 D). Lang’s (1936 b: 23, Fig. 52) observation of a similar condition in a male from the Øresund indicates that it is genuinely lost (or extremely reduced) rather than broken off during dissection. Although Lang (1934: 24) expressed doubts, primarily based on differences in antennulary segmentation, about the conspecificity of Sars’s (1905) Norwegian specimens of M. forficula and Claus’s (1863) type material of T. forficula, he cited the former as a synonym of the new combination Parastenhelia forficula (Claus, 1863). Sars (1911: 369 – 370, Supplement plate 11 - 1) added the new species M. littoralis Sars, 1911 to the genus which showed more or less the same distribution in Norway as M. forficula, including various localities along the south and west coasts and further northwards to the Trondhjem Fjord (Bejan). Sars (1911) only reported the female which differs from that of the type species in the more compact antennule (the number of segments was not disclosed), the longer and more slender P 1, the number of exopodal setae on the P 5 (six vs eight) and the normally developed seta V on the caudal ramus. Unfortunately, no information on the armature formula of P 2 – P 4 was given which inevitably led to the incorrect assignment of various non-conspecific populations to this species. Farran (1913) observed considerable variation in body size (500 – 850 μm) in his material of M. littoralis from Clare Island and Blacksod Bay, Co. Mayo (Ireland) (see also Farran et al. 1915; Southern 1915) which he viewed as a possible indication of the presence of a second species. All specimens agreed with Sars’s (1905) description in the morphology of the female P 5. Pesta (1920: 591 – 593) summarized records of Microthalestris in the Adriatic, including M. forficula in the Venice Lagoon (Grandori 1912, 1914) and M. littoralis from the Palagruža (Pelagosa) archipelago (Steuer 1912). Brian (1921: 77 – 80) reported M. littoralis from the Gulf of Genoa, illustrated naupliar stages I and III – V (Text figures 23 – 25; Plate VI, Fig. 3) and described copepodids III – V (Plate VIII, Figs 10 – 22). Dahms & Hicks (1996) pointed out that Brian’s naupliar illustrations in reality refer to NIV – NVI. Brian’s (1921) description reveals little information about the female except for the lateral aspect view of an ovigerous specimen (Plate III, Fig. 7), showing the presence of a single egg sac, and the ventral view of the abdomen (Plate V, Fig. 15) which shows that the caudal ramus seta V is not swollen at the base. His illustrations of the male (Plate IV, Fig. 9; Plate V, Fig. 5; Plate IX, Figs 9 – 14) document three characters of particular significance, (a) the presence of two penicillate spines on the antenna (already expressed in copepodid IV: his Plate VIII, Fig. 17), (b) the 3 - segmented P 3 endopod in the male bearing an inner seta on enp- 1 and - 2, in addition to the apophysis and two apical setae on enp- 3, and (c) the 1 - segmented condition of the male P 5 exopod bearing seven setae. No information was provided on the segmentation of the female antennule or armature of the swimming legs and P 5. Monard (1927) retained Microthalestris in the Thalestridae as an “ insufficiently described genus ” (p. 157) in his Synopsis universalis generum harpacticoidarum but placed the closely related Parastenhelia in the Diosaccidae (= Miraciidae). He admitted that the latter genus may belong to the same family as Microthalestris on the basis of P 1 morphology but was unable to make a firm recommendation for its exclusion from the Miraciidae because the number of egg sacs was apparently unknown [this being untrue since both Car (1884) and Brian (1921) had made cursory observations and illustrated a single egg sac]. Gurney (1927: 545) confirmed the presence of a single egg sac in a damaged female of an undescribed Parastenhelia species from Port Taufiq (Suez Port), located at the southern boundary of the Suez Canal in Egypt. He classified the genus in the Canthocamptidae. Monard (1928: 348 – 349, Fig. XVIII – 2) reported numerous specimens of M. forficula from Port-Vendres and Banyuls-sur-Mer on the French mediterranean coast and considered the species as extremely variable with M. littoralis definitely falling within its limits. Contrary to Sars (1905), Monard also appeared to support the concept that M. forficula assumes a cosmopolitan distribution by accepting doubtful records from New Zealand (Thomson 1883) and the Gulf of Guinea (Scott 1894 a) as valid. The French material differs from the Norwegian specimens used in Sars’s (1905) description in a number of significant aspects such as (a) the segmentation of the female antennule (eight vs nine), (b) antennary endopod with two penicillate elements, (c) caudal ramus setae IV – V with an inflated base (this also occurs in Claus’s description of T. forficula), (d) claws on both rami of P 1 distinctly shorter, (e) P 3 exp- 3 with three inner setae vs two, (f) P 4 enp- 3 with two inner setae vs one, (g) P 5 exopod ♀ with six elements vs eight, and (h) P 5 exopod ♂ 1 - segmented vs 3 - segmented. Monard (1928) also reported variability in the armature of the female P 5 exopod with some (rare) specimens displaying seven elements, and variation in body length (650 – 800 μm). Sewell (1940: 191) stated that Monard misidentified his specimens and was in reality dealing with M. littoralis sensu Sars (1911) but there is no justification for this claim. Stephensen (1929) recorded M. littoralis from two localities in the Faroe Islands, i. e. off Trangisvaag (Suðuroy = Suderø), and between Streymoy (Strømø) and Eysturoy (Østerø). Lang (1948) re-examined Stephensen’s material and confirmed the identity of the specimens from the latter site but confirmed that the male from Trangisvaag belongs to Amonardia normani (Brady, 1872) (Thalestridae). Wilson (1932 a) added the first North American record of M. littoralis from the plankton in Chesapeake Bay, Maryland, stating that he found both sexes. His record was not accompanied by any illustrations but the author seems to indicate that he accepts Brian’s (1921) identification of the male of this species as correct. Wilson (1932 b) subsequently recorded both M. forficula (Penzance Pond) and M. littoralis (Cuttyhunk Island between Buzzards Bay and Vineyard Sound) from the Woods Hole region, Massachusetts. The report of the first species included illustrations of P 1, P 5 of both sexes and male P 3 endopod, all of which agree well with Sars’s (1905) description except for the latter which shows an inner seta on the distal segment (only P. bulbosa shares this character; Table 1), and the male P 5 which displays a long outer seta on the proximal segment. No information was given about the segmentation of the female antennule, swimming leg armature or the form of caudal ramus seta V. The North American specimens are distinctly smaller (♀♀: 400 – 500 μm; ♂♂: 350 μm) than their European counterparts. The authenticity of Wilson’s (1932 b) material of M. littoralis is indeterminable since the few illustrations provided are reproduced from Sars (1911). His text reveals that females (no males were found) correspond in the setation of the female and measure 500 – 600 μm. Interestingly, Wilson (1932 b: 572) maintained Parastenhelia and Microthalestris as separate genera. Jakubisiak (1933: 115) recorded M. littoralis from the Isle of Šolta along the Dalmatian coast of Croatia and opposed Monard’s (1928) view that this species must be subsumed under M. forficula, stating that it not only differs in the morphology of the female P 5 but also in body size and the segmentation of the antennule. Lang (1934) formally synonymized Microthalestris with Parastenhelia and transferred the latter to the Thalestridae based primarily on the similar position of the inner seta on P 1 enp- 2 and the reported presence of a single egg sac in members of the genus (cfr. Gurney 1927: 545). Morphological comparisons between single species of these genera were also used as additional evidence in support of their synonymy, e. g. similarities were pointed out in the morphology of ♀ P 5 (P. hornelli and M. littoralis), ♂ P 5 (P. hornelli and M. forficula), P 1 (P. antarctica and M. forficula / littoralis) and caudal ramus seta V (P. similis and M. forficula). Lang (1934: 25) relegated M. littoralis to a mere variety of Parastenhelia forficula (Claus, 1863), dismissing the most clearcut character that consistently separates these species (number of exopodal setae on ♀ P 5) along most of the Norwegian coastline (Sars 1911) as of no significance. This course of action was based on the incongruous argument that differences had been reported between the widely disjunct “ populations ” of T. forficula (and M. littoralis) from New Zealand (Thomson 1883), Gulf of Guinea (Scott 1894 a) and Ireland (Farran 1913). The first two sources contain incomplete and partly inaccurate descriptions (based on a single specimen in Scott’s case) while the third is completely lacking in illustrations and by Farran’s own account is probably based on an amalgam of two species. Lang’s (1934) paper set the scene for his authoritarian stance in shaping a very broad, encompassing taxonomic concept of Parastenhelia spinosa, with the result that it has unjustly come to be regarded as a highly variable, cosmopolitan species. He further fuelled the controversy by assigning a pair in amplexus from Perseverance Harbour on Campbell Island in the southern Pacific to P. forficula var. littoralis. Differences in P 1, P 5 and caudal rami indicate that his subantarctic specimens belong to a different, as yet unnamed species (see below). Lang (1934) listed T. karmensis and T. forficuloides as synonyms of the typical P. forficula. Monard (1935 a: 43) recorded both M. forficula and M. littoralis from the Roscoff region (Brittany). The former displayed a 3 - segmented P 5 exopod in the male while females of the latter were excessively large (900 μm) compared to the Norwegian populations (590 μm; cfr. Sars 1905). He continued to place Parastenhelia in the Miraciidae while maintaining Microthalestris and his newly established genus Thalestrella in the Thalestridae. During the same year, Monard (1935 b: 47), adopting a different nomenclature, stated that he had found both the “ typical ” form and the littoralis variety of Parastenhelia forficula in Cartaghe (Salammbô), Tunisia and admitted having difficulties separating them. He reported variability in the number of segments in the female antennule (one specimen having seven!) and occasional left-right asymmetry in the setation of the female P 5 exopod in some individuals. Without illustrations it is impossible to decide on the identity of his material. Willey (1935: 82 – 83) distinguished two forms in his material of M. littoralis from Harrington Sound in Bermuda. One ovigerous female (500 μm) conformed to Sars’s (1911) description in displaying 9 - segmented antennules and unmodified caudal ramus setae. The endopods of P 2 – P 4 of this littoralis - form are distinctly 3 - segmented, have a constricted mobile articulation between enp- 2 and enp- 3 and the inner distal corner of enp- 2 produced into a sharp spur beyond the insertion of the inner seta (his Fig. 121); the formula of P 4 enp- 3 is 121. Furthermore, the proximal inner seta of P 4 exp- 3 is longer than the second inner seta, which is comparatively short and only slightly thickened. All other specimens (♀♀: 500 – 700 μm; ♂♂: 400 – 500 μm) had 8 - segmented antennules and forcipately curved caudal setae IV – V with swollen bases. Willey (1935: Figs 119 – 120, 122 – 126) assigned them to a separate penicillata - form, based on the presence of two penicillate spines amongst the apical armature elements of the antenna similar to those figured by Brian (1921) for M. littoralis from the Gulf of Genoa. In addition, both claws at the distal margin of P 1 enp- 2 are coarsely penicillate, and P 1 exp- 3 bears a slender inner seta, a geniculate seta, and two penicillate spines. In the penicillata - form P 2 – P 3 enp- 2 and - 3 are more often indistinctly separated, without a functional articulation between them. Specimens with distinctly 3 - segmented endopods of P 2 – P 3 lack a spur at the inner distal corner of enp- 2. The armature formula of P 4 enp- 3 is sexually dimorphic, being 221 in the female, and 121 in the male. On P 3 – P 4 exp- 3, the distal inner seta is excessively fine and curving outwards, and the proximal inner seta of P 4 exp- 3 is distinctly shorter than the middle one. Females of both forms share the typical P 5 setation pattern of M. littoralis (six elements on the exopod, five on the endopodal lobe). Males of the penicillata - form have a 3 - segmented P 3 endopod with two apical setae and a spinous apophysis on enp- 3, and a 1 - segmented P 5 exopod with seven elements. Lang (1936 b) recorded both P. forficula (one ♀, one ♂) and P. forficula var. littoralis (one ♀) from the Øresund between Zealand (Denmark) and Scania (Sweden) with all three specimens having been collected from different localities. His P. forficula specimens agree with Sars’s (1911) description in the general facies of the P 1 and P 5 of both sexes but differs in the presence of only one inner seta on P 3 enp- 3 (instead of two) in the female; however, it must be noted that the latter observation was based on a single female and not accompanied by an illustration. Lang (1936 b) assigned, except for his own record from Campbell Island (Lang 1934), several older records to P. forficula var. littoralis without giving any explanation. These include the reports of Microthalestris littoralis from Norway (Sars 1911), western Ireland (Farran 1913), the Gulf of Genoa (Brian 1917, 1921), Palagruža in Croatia (Steuer 1912), Chesapeake Bay (Wilson 1932 a) and Cuttyhunk Island, Massachusetts (Wilson 1932 b), of Microthalestris forficula from Banyuls-sur-Mer (Monard 1928), and of Thalestris forficula from New Zealand (Thomson 1883) and Ghana (Scott 1894 a). Lang (1936 a) proposed a new subfamily Parastenheliinae within the Thalestridae and fixed Parastenhelia as its type and only genus. Contrary to his earlier opinion (Lang 1934: 24) he accepted the conspecificity of Claus’s (1863) T. forficula and Sars’s (1905) M. forficula, and assigned the previous records of T. forficula from Lille-Karajak fjord, western Greenland (Vanhöffen 1897) and of M. forficula from Woods Hole (Wilson 1932 b) to Parastenhelia forficula. He maintained Sars’s (1911) M. littoralis as a variety of P. forficula, made Dactylopus longipes a synonym of this species, but surprisingly made no mention of Fischer’s (1860) Harpacticus spinosus. The two females of P. gracilis he reported from the Falkand Islands are here assigned to P. antarctica (see below). Monard (1937: 59, 62, Fig. 3 – E) collected material from Algiers and Bou Ismaïl (formerly Castiglione) in Algeria and assigned it to P. forficula littoralis. He figured the male P 5 exopod as 1 - segmented bearing a total of seven elements and based on his previous observations (Monard 1928, 1935 a) prudently postulated that this character may be dimorphic among males. In his review of the genus Parastenhelia Sewell (1940: 189 – 196 )) reassessed the great degree of reported variability in antennule segmentation, armature of P 2 – P 4, segmentation of male P 5 exopod, and shape of terminal caudal ramus setae. He forcefully opposed the views of Lang (1934, 1936 a, 1936 b) and Monard (1935 b, 1937) for uncritically accepting the synonymy of T. forficula, T. forficuloides and M. littoralis, claiming that these authors and others before them had been dealing with a mixture of species. Sewell (1940) reinstated the latter two as valid species of Parastenhelia and maintained Willey’s (1935) penicillate variety as a distinct form of P. littoralis. His material from seaweed washings in Addu Atoll, the southernmost atoll of the Maldives, revealed three different forms which he attributed to P. littoralis, P. littoralis f. penicillata, and a new form, P. littoralis f. scotti. Sewell’s P. littoralis material (his Text-Fig. 26) is characterized by the following features: (a) body length 440 – 560 μm (♀), unknown (♂); (b) antennule ♀ 9 - segmented; (c) antennary endopod with two penicillate spines; (d) P 1 enp- 1 very long, about 1.45 times length of exopod; enp- 2 with two penicillate claws; exp- 3 with three penicillate elements and one long seta; (e) P 3 endopod ♂ 3 - segmented, with armature formula [1.1.02 + apo]; (f) P 4 exp- 1 without inner seta and exp- 3 with seven elements; endopod with inner seta on exp- 1 and - 2, and four elements on enp- 3; (g) P 5 exopod ♀ elongate, about 3.7 times as long as maximum width, with six setae; endopodal lobe with five setae; (h) P 5 exopod ♂ 1 - segmented, with seven setae; and (i) caudal ramus seta V unmodified or only slightly swollen at the base. Despite the absence of information about female antennulary segmentation, antennary morphology and swimming leg armature in M. littoralis, it is still obvious that Sewell’s (1940) specimens do not belong to this species; this is best illustrated by the morphology of the P 1 and the length: width ratio of the ♀ P 5 exopod. The absence of the inner seta on P 1 exp- 2 is most likely based on an observational error. It is puzzling why Sewell listed T. forficula sensu Thomson (1883) and M. forficula sensu Monard (1928) as misidentified records of P. littoralis since both sets of specimens are significantly divergent from Sars’s (1905) description. Sewell’s (1940: Text-Fig. 27) second form, based on females only and assigned to P. littoralis f. penicillata, is characterized by the following features: (a) body length 600 μm (♀), unknown (♂); (b) antennule ♀ 8 - segmented; (c) antennary endopod with two penicillate spines; (d) P 1 enp- 1 long, about 1.15 times length of exopod; enp- 2 with two penicillate claws; exp- 3 with two pinnate claws and two (?) setae; (e) P 2 – P 3 exp- 3 with six and seven setae / spines, respectively, P 2 exp- 1 without inner seta; (f) P 2 – P 4 enp- 3 with four, five and five elements, respectively; (g) P 5 exopod ♀ elongate, about three times as long as maximum width, with six setae; endopodal lobe with five setae; and (h) caudal ramus setae IV – V markedly swollen at the base (occasionally less pronounced in seta IV). According to Sewell (1940) there is no doubt that his material is identical to Willey’s (1935) specimens from Bermuda but a comprehensive comparison is hampered by the lack of information on (a) relative segment lengths of P 1, armature of P 2, and setation and length: width ratio of the female P 5 exopod in the Bermudian material, and (b) armature of P 4 exopod, sexual dimorphism in P 4 enp- 3 setal pattern, and both segmentation and armature of the male P 5 exopod in the Maldivian specimens. It is unclear whether the absence of the thin, outwardly curved, distal inner seta on P 3 (and possibly P 4) exp- 3 in Sewell’s (1940: Text-Fig. 27 – I) specimens is genuine or due to an observational error; a minute notch along the distal protion of the inner margin appears to hint at its presence. The final form, based on a single female (Text-Fig. 28), was described as a new variety, forma scotti, of P. littoralis. It is characterized by the following features: (a) body length 560 μm; (b) antennule 8 - segmented; (c) antennary endopod without penicillate spines; (d) P 1 enp- 1 very long, about 1.33 times length of exopod and twice as long as exp- 2; enp- 2 with two pinnate claws; exp- 3 with three claws and one seta; (e) P 5 exopod large (reaching to middle of third abdominal somite), elongate-oval, about three times as long as maximum width (measured in proximal 1 / 3), with six setae all inserting in distal one-third; endopodal lobe with five setae; and (f) caudal ramus very broad (about 4.5 times as wide as long), seta V forming pyriform swelling at the base. Sewell (1940) considered this form identical with Scott’s (1894 a) ovigerous female of T. forficula from Ghana, primarily on account of the general shape of the P 5 exopod and the modification of caudal seta V. Although both females are strikingly similar in these two characters and Sewell dismissed the absence of the proximal inner seta on the P 5 endopodal lobe in Scott’s specimen as an observational error, others demonstrate that they are clearly different. Such features are particularly related to the P 1 (relative lengths of rami, armature of exp- 3) and P 5 (length: width ratio, size relative to urosome, position and length of setae). A comparison of the swimming legs is impossible since only the P 4 was figured by Scott (1894 a: Plate XII, Fig. 39) and no information was given by Sewell (1940) other than that they “ … appear to resemble those of P. littoralis ”. Lang (1944: 13) invalidly fixed Harpacticus spinosus as the type species of Parastenhelia. Nicholls (1945: 3 – 4, Text fig. 2 A – B) illustrated two forms of P. forficula from the reef fringing Leander Point at Port Denison near Dongarra in Western Australia. Both forms differ considerably in the shape of caudal ramus setae IV – V, P 1 and female P 5. Adopting the view of both Monard (1928) and Lang (1934, 1936 a, 1936 b) he considered M. littoralis a variety of P. forficula. Three females (720 μm), which were assigned to the “ typical ” form of P. forficula, display (a) distincty basally swollen caudal setae IV – V, (b) a P 1 in which exp- 2 and enp- 1 are extremely elongate, being much longer than in Sars’s (1905) material from Norway, and (c) a P 5 which is ovate in shape and shows the 6 - element arrangement typical of P. littoralis instead of the eight setae / spines in typical P. forficula. Nicholls (1945: 4) himself regarded the identification of his material as provisional, suggesting it may represent a new species. The second form, described as P. forficula var. littoralis (♀♀: 630 – 670 μm.; ♂: 540 μm), exhibits unmodified caudal setae IV – V. Females differ slightly from Sars’s (1911) description of M. littoralis in (a) the proportionally longer P 1 exp- 2 and enp- 1, and (b) the P 5 endopodal lobe which is narrower and less truncate. Nicholls (1945) claimed that the male P 5 (1 - segmented with seven elements) did not conform with the descriptions of Sars and Lang but this statement cannot be substantiated since Sars (1911) did not describe the male and Lang’s (1934: Fig. 56) illustration of the P 5 refers to a different species (see below). Having previously claimed that Harpacticus spinosus was indeterminable (Lang 1936 a: 21), Lang (1948: 311) stated that “ Fischer’s illustrations of this species clearly show that it is the same species that Claus described as Thalestris forficula and which Lang (1934) subsequently placed in the genus Parastenhelia. That I make this identification without any reservation, is based on the fact that there is no other harpacticoid whose P 1 and P 5 are built in the way Fischer depicts ” [our translation]. As a result of Lang’s groundless reinstatement of H. spinosus as a valid species it not only became the senior synonym of a whole string of species and forms previously considered as conspecific with M. forficula and M. littoralis but was also unjustly designated as the type species (P. spinosa) of Parastenhelia. Lang (1948) assembled arguments in favour of his very broad concept of P. spinosa, including all the species, forms and varieties that in the course of time had been described and he considered conspecific – mostly on dubious grounds. These include Thalestris forficula, T. forficuloides, T. karmensis, T. pectinimana, Dactylopus longipes, Microthalestris littoralis and M. littoralis f. penicillata. With regard to the latter Lang (1948: 586) stated emphatically that comparison with Fischer’s (1860) description of Harpacticus spinosus shows beyond doubt that both forms are identical. Since Fischer only illustrated, and quite inadequately, the maxilliped, P 1, female P 5 and the caudal ramus, it is inexplicable how Lang could have arrived at this conclusion because none of these appendages (except for the caudal ramus) was figured or described by Willey (1935). Given that Fischer’s (1860) illustrations do not inspire confidence in their accuracy it would not be extraordinary to state that there is no tangible evidence for synonymizing any of the other species / forms mentioned above with H. spinosus. Lang (1948: 586) suggests in his review that M. forficula, M. littoralis and M. littoralis f. penicillata should be regarded as three forms (forma typica, forma littoralis and forma penicillata, respectively) of P. spinosa, which however are not geographically or ecologically distinct. Separation of these forms was based on the shape and / or modification of caudal ramus setae IV – V, number of segments in the female antennule, presence / absence of penicillate spines on the antennary endopod, shape and ornamention of the distal claws on P 1 enp- 2, number of setae on P 3 – P 4 exp- 3, P 4 enp- 3 (in both sexes where known) and P 5 exopod ♀, and segmentation of P 5 exopod ♂. Rather than exploring the possibility that P. spinosa may represent a species complex Lang claimed that it is impossible to keep the three forms distinct because a number of intermediate forms are known to exist and the combination of characters varies in every conceivable manner. Lang (1948: 587) underpinned his claim by shoehorning into P. spinosa every form and variety from geographically disparate locations such as Campbell Island (Lang 1934), the Woods Hole region (Wilson 1932 b), Gulf of Guinea (Scott 1894 a), Brittany (Monard 1935 a), the Banyuls area (Monard 1928) and the Gullmar Fjord in Sweden (Lang 1948), irrespective of the morphological variation reported. Since Lang failed to recognize sibling species boundaries, which negatively affects studies on biodiversity, speciation and ecology, his attempt to present a unifying, deliberately broad, taxonomic concept of P. spinosa should be seen as an exercise in futility. Unfortunately, most subsequent workers have uncritically adopted this concept and even attempted to expand it (e. g. Vervoort 1964) rather than subjecting it to scrutiny [Gee (2006) and Kuru & Karaytuğ (2014) are notable exceptions]. The species is now widely viewed as assuming an allegedly cosmopolitan distribution, covering all continents and including records from the Arctic (Vanhöffen 1897; Scott 1899; Scott & Scott 1901; Sars 1909; Brotskaya 1962; Mielke 1974; Chislenko 1967, 1977; Chertoprud et al. 2005; Kornev & Chertoprud 2008; Dvoretsky & Dvoretsky 2010) and remote islands such as Tristan da Cunha (Wiborg 1964) and Moce Island, Fiji (Wells 1978). Lang’s (1948) view was not generally accepted by some Irish workers who continued to separate “ populations ” of P. spinosa into forms or even species. Williams (1954) recorded both Microthalestris forficula and M. littoralis from pools at Ardglass (the latter also from similar habitats at Kilclief) in Co. Down, Northen Ireland. Roe (1958) reported both P. spinosa var. littoralis and P. spinosa var. penicillata in intertidal pools around Dalkey Island and The Muglins (lighthouse), Co. Dublin, Ireland. Both forms were subsequently obtained from littoral stone and seaweed washings and net sweepings in Lough Hyne (Ine), West Cork, Ireland (Roe 1960). Noodt (1955 b: 204) suggested that Sewell’s (1940) Parastenhelia littoralis f. scotti probably falls within the boundaries of Lang’s concept of P. spinosa. Božić (1955: 5, 11, Plate 4, Figs 1 – 3) described P. spinosa from Kernic, west of Roscoff, Brittany and illustrated the female caudal ramus, antennary exopod and mandibular palp. Females display a 9 - segmented antennule and a P 5 with six setae on the exopod and five on the endopodal lobe. Males have a 3 - segmented P 5 exopod bearing seven elements. Božić (1955) reported sexual dimorphism in the shape of caudal ramus seta IV, being distinctly swollen and bulb-shaped in the female and unmodified in the male. In both sexes the inner terminal seta V was more or less normal at the base. The material was accorded distinct specific rank by Gee (2006 – see below). Şerban (1959) reported P. spinosa from Agigea along the Romanian Black Sea coast, stating that his specimens approached Willey’s (1935) penicillata form because of the 1 - segmented exopod in the male P 5, a condition he attributed to neotenic development rather than variability. It is not clear whether his material displayed the penicillate spines on the antenna and P 1 endopod. Pór (1960) subsequently recorded P. littoralis from Eforie and Agigea and advocated that P. spinosa (which he regarded as conspecific with Microthalestris forficula) and P. littoralis are separate species. Pesta (1959: 128 – 129; Figs 66 – 69) collected both sexes of P. spinosa from algal washings at Sorrento in the Bay of Naples, Italy. Illustrations were provided for the P 1, the female P 5 and abdomen, and the male P 3. Pesta accepted Lang’s (1948) concept of P. spinosa, being a variable species containing three ill-defined and overlapping forms, and stated that the females in his material displayed the P 5 armature of f. littoralis and the basally swollen caudal ramus seta V of f. typica, and that one of the males resembled f. penicillata in the structure of the P 3 endopod. The body size of the male was erroneously given as 4.4 mm (it is more likely 440 μm). Among all the records that have been attributed to P. spinosa, Pesta’s is probably the one that is most similar to Claus’s (1863) original from Messina. Apart from being the one that is geographically closest to the type locality in Sicily, the Napolitanean female agrees with the original description in the relative proportions of the P 1 rami, the degree of modification of caudal ramus seta V and the ovate shape of the P 5. Except for the male P 3 Pesta (1959) does not provide any information about the armature of the remaining swimming legs or the segmentation and setation of the male P 5. In his first paper on the harpacticoids from the Isles of Scilly Wells (1961) reported finding populations of P. spinosa containing a mixture of f. typica, f. littoralis and many females which were identical with Božić’s (1955) unnamed form characterized by the bulbous base of caudal ramus seta IV. The species was found on all four principal islands (St Mary’s, Tresco, St Martin’s and St Agnes). Wells (1963 a: 14) subsequently found the bulbous form in clean sandy substrates at two sites in the estuary of the River Exe and formally proposed to name it P. spinosa forma bulbosa. A second form, P. spinosa forma littoralis was also recorded from one of the stations in the estuary. Gee (2006: 2634 – 2643, Figs 14 – 19) re-examined Wells’s (1963 a) forma bulbosa based on new material from the Isles of Scilly and accorded it full specific rank (as P. bulbosa). Both Bodin & Boucher (1983) and Bodin (1988) had previously recorded the taxon from Nord-Finistère (Brittany) as P. spinosa bulbosa. Since neither satisfied the criteria of availability (ICZN Arts 11 – 18) they cannot claim authorship for the subspecific name (ICZN Art. 45.5.1). Wells’s (1970: 261) record of P. spinosa from Port Hellick on St Mary’s was also confirmed to be conspecific with P. bulbosa (Gee 2006). Vervoort (1964: 182) found that most of his Ifaluk material, although showing great variability in body length, agreed with Lang’s (1948) forma typica of P. spinosa (formerly M. forficula). No illustrations were given but Vervoort stated that the females were characterized by a 9 - segmented antennule, normally developed spines on the antennary endopod, P 1 exp- 2 (Vervoort erroneously referred to as the “ 2 nd endopodal segment ”) 4 – 5 times as long as wide with an inner seta near the apex, P 4 enp- 3 with one inner seta, P 5 exopod with eight elements, and a slightly basally swollen caudal ramus seta V. Males displayed a 3 - segmented P 5 exopod bearing six setae. One female and one male of a different form (from his locality 592), which he claimed would formerly have been assigned to Willey’s (1935) forma penicillata, were described in considerable detail (Vervoort 1964: 182 – 184, Figs 64 c, 66 c, 67, 68 d, 70 a – g). The female (520 μm) exhibits a fairly long 8 - segmented antennule, two penicillate spines on the antennary endopod, P 1 exopod with two penicillate spines on exp- 3 (note that Vervoort had misinterpreted this ramus as the endopod), basally swollen caudal ramus setae IV – V, and a P 5 exopod with six elements. No information was provided about the female armature of P 2 – P 4 or P 5 endopodal lobe but Vervoort did illustrate the exopods of P 2 – P 3 and the endopod of P 3 of the male (420 μm). His material is assigned distinct species status below. Bodin (1964: 128, Plate III, Fig. 15) recorded specimens of P. spinosa f. penicillata in high abundance in the Marseille region, southern France. Females display eight segments in the antennule but the proportional segment lengths are different from those reported by Monard (1928) and Sewell (1940). The only illustration given is that of the male P 3 endopod which differs from Willey’s (1935) account in the absence of the inner seta on enp- 1. Chislenko (1967: 140 – 144, Figs 45 – 46) provided illustrations of both sexes and an aberrant specimen of P. spinosa from the White Sea. His material (♀: 550 – 800 μm, ♂: 400 – 420 μm) shows some similarities with Sars’s (1905) illustrations of M. forficula (e. g. 9 - segmented antennule and eight elements on P 5 exopod ♀) but differs in the length: width ratio of the female P 5. The P 1 has a similar exopod: endopod length ratio but the rami are significantly longer in the White Sea specimens. The males among Chislenko’s material display the full complement of seven elements on the P 5 exopod while the outer seta is absent on the proximal exopodal segment in the Scandinavian populations of M. forficula (Sars 1905; Lang 1936 b). Considerable variability was observed in the degree of the basal swelling of caudal ramus seta V, including left-right asymmetry in the same specimen. In one female individual the P 5 exopod on one side only had seven instead of the typical eight setae but the taxonomic significance of this aberration was dismissed. Chislenko (1967) drew particular attention to the structure of the female genital field, pointing out that the condition in the White Sea material differed significantly from that of Lang’s (1934: Fig. 51) illustration of the structure in his forma littoralis but was more similar to Claus’s (1863: Plate XVII, Fig. 11) approximate representation of that in the typica form. He expressed a sceptical attitude towards the recognition of forms or varieties in P. spinosa, claiming instead that a proper comparative analysis of the female genital field of these infrasubspecific taxa would confirm their specific status. Apostolov (1968: 398, Plate 2, Figs 1 – 10) reported both sexes of P. spinosa from a sandy beach in Tsarevo (formerly Michurin), Bulgaria, provided a partial description, and assigned them to a new form, f. bulgarica. The Black Sea specimens are characterized by unmodified caudal ramus setae IV – V, the absence of the inner seta on P 2 exp- 1 and enp- 1, and the presence of seven setae on the ♀ P 5 exopod. The male displays a 1 - segmented P 5 exopod with seven elements and a bisetose endopodal lobe which is distinctly larger than in any other reported population. Apostolov (1968) illustrated sexual dimorphism in the setation of the antennary exopod but did not comment on it in the text. In a later paper Apostolov (1973: 270, 272 – 273, Fig. 8) recorded additional specimens from Tsarevo and supplemented his original descriptions of the female with illustrations of the genital field, P 1, P 2, P 4, P 5 and caudal rami. The latter two differ in some aspects (e. g. P 5 exopod with only six setae) from his 1968 drawings, rendering future morphological comparison difficult. Mielke (1974: 20 – 22, Fig. 9) reported on Parastenhelia specimens he collected in the intertidal zone at Longyearbyen in Svalbard. He assigned the material to P. spinosa, stating that it could not be attributed to any of the forms recognized so far. Illustrations were presented of the antennary exopod and P 1 of the female, and the P 3 endopod and P 5 of the male. The female (590 μm) is characterized by a 9 - segmented antennule, normally developed caudal ramus setae, and eight and five setae on the exopod and endopodal lobe of the P 5, respectively. Males (420 – 510 μm) possess a characteristic, acutely recurved, spinous apophysis on the P 3 endopod. Some aberrations were recorded in the armature of P 2 enp- 2 (one or two inner setae) and P 4 exp- 1 (with or without inner seta). Marinov (1974: 80 – 82, Figs 1 – 2) provided a description of P. spinosa specimens collected from algal washings at Sozopol on the southern Bulgarian Black Sea coast. Females display an 8 - segmented antennule, six and five elements on the P 5 exopod and endopodal lobe, respectively, and what appear to be unmodified caudal ramus setae IV – V. The distal endopod segment of P 1 bears two pinnate (or penicillate) claws while enp- 1 is excessively long, measuring 2.2 times the length of exp- 2. The inner seta on P 4 enp- 1 is exceptionally long, extending beyond the distal tip of the endopod. The male shows the typical modification on the P 3 endopod [1.1.02 + apo] and a 1 - segmented P 5 exopod with seven elements. Marinov (1974) reported some aberrant specimens that had a 1 - segmented exopod in P 3 – P 4, each carrying two apical elements, and observed a malformation in one male exopod which carried only six setae. Due to its dubious taxonomic status Apostolov’s (1968) forma bulgarica was dismissed from further consideration. Chislenko (1977: 246, Fig. 6 – 1,2) collected two females (675 – 700 μm) of P. spinosa from washings of the demosponge, Semisuberites cribrosa (Miklucho-Maclay, 1870), in Franz Josef Land. He maintained that his specimens were morphologically similar to the material he described from the White Sea (Chislenko 1967) which is confirmed by the two illustrations presenting the genital field and the P 5 (with eight elements on the exopod). According to Wells & Rao (1987) specimens (two ♀♀, one ♂) of P. spinosa from the Andaman Islands appear to be morphologically most similar to Sewell’s (1940) forma scotti, with eight segments in the female antennule, non-pectinate spines on the antennary endopod and a relatively short P 1 exopod. The exopod of the male P 5 is 3 - segmented with six setae. Gee (2006) stated that the Langian concept of P. spinosa as a very variable, cosmopolitan species is probably erroneous and a thorough reappraisal of this species is required. He convincingly demonstrated that P. spinosa var. bulbosa must be accorded distinct specific status as P. bulbosa and argued that Willey’s (1935) forma penicillata displays several characteristics warranting its elevation to species level. Kornev & Chertoprud (2008: 196 – 197) provided an illustrated description of a White Sea population which they identified as P. spinosa. It appears that most of the text and at least some of the illustrations are a verbatim account of Chislenko’s (1967) description, including the substantial variation recorded in female body length. The authors do not specify whether the observed variability in P 3 – P 4 armature is based on their White Sea material or on previous literature reports. Kuru & Karaytuğ (2015) added a new species, P. aydini, from the eastern Mediterranean and Aegean coasts of Turkey and considered it morphologically closest to P. spinosa. The species is unique in the family by displaying a 2 - segmented P 5 exopod with a total of five setae in the male (a 2 - segmented condition with different armature has been observed in Johnwellsia bipartita gen. et sp. nov. and Parastenhelia willemvervoorti sp. nov.). According to Kuru & Karaytuğ (2015), females of P. aydini differ from those of P. spinosa by the 9 - segmented antennule, the reduction of the spinular rows around the outer margins of the bases of P 1 – P 4, the presence of only four elements on P 4 enp- 3, ornamentation of the P 5 baseoendopod and exopodal setae different, and the normally developed caudal ramus seta V. Males differ in the absence of a modified element on the seventh segment of the antennule (apparently present in P. spinosa), the notched apophysis on P 3 enp- 3 and the presence of only four setae on P 4 enp- 3. It is not clear which material of P. spinosa the authors used as their reference for comparison. Zaleha et al. (2018) provided a short text description of a single ovigerous female of Parastenhelia sp. from sediment in seagrass bed in Sungai Pulai, Johor, Malaysia. The antennule was described as 8 - segmented and the 2 - segmented P 1 endopod as being much longer than the exopod. However, the armature of P 2 – P 4 and the endopodal lobe of P 5 is radically divergent from any other species in the genus, casting serious doubts on the validity of their identification. Zaleha et al. (2010) had previously identified Parastenhelia littoralis from rockpool seaweeds and sediment in Pulau Besar, Melaka. Cordell (2018: 117 – 118, Fig. 5) found several Parastenhelia specimens on debris, generated from the earthquake and tsunami that struck Japan on 11 March 2011, that landed in North America. The species was recorded on floating structures (docks, vessels, buoys) in several localities along the coasts of Washington and Oregon. While Cordell assigned the material to P. spinosa he acknowledged that they may be a different species within a larger species complex. Photographs were provided of the P 1 and female urosome, showing that the caudal ramus seta V is somewhat swollen near the base. Although this character was observed in every female, Cordell (2018) mentioned that specimens he had previously examined from the northeast Pacific had almost no swelling at the base of this seta.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFD7FFBCFF51D13AE09BFBFD.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 3 endopod, P 5 – P 6 and urosomal segmentation; occasionally endopods of P 2 and P 4, and caudal ramus setae IV – V. Body subcylindrical, short; posterior margin of urosomites (except anal somite) typically with denticulodigitate hyaline frills. Rostrum defined at base, of moderate size, reaching at most to distal margin of second antennulary segment; linguiform. Anal operculum semicircular, typically bordered with fine spinules. Caudal ramus wider than long, without spinular row on dorsal surface; with at least seven setae (seta I possibly absent); setae IV and / or V slightly swollen at base in ♀; seta IV ♀ with bulbous base in some species (M. bulbosa comb. nov.). Antennule ♀ elongate and 9 - segmented in ♀, segment 1 not elongate, segments 7 – 8 (when 9 - segmented) shortest, with aesthetascs on segments 4 and 8 or 9. Antennule ♂ haplocer with four segments distal to geniculation but segmentation and segmental homologies unconfirmed for most species; in M. bulbosa comb. nov. segment 5 swollen and with aesthetasc, segmental homologies as follows: I, II – VIII, IX – XII, XIII, XIV – XVII, XVIII, XIX – XX, XXI – XXII, XXIII, XXIV – XXV, XXVI – XXVIII. Antenna not sexually dimorphic; proximal endopodal segment largely separated from basis or forming allobasis, with pinnate seta on abexopodal margin; exopod 2 - segmented, proximal segment with 1 – 2 seta (e), distal segment with 1 – 2 lateral and three apical elements; distal endopodal segment without penicillate elements. Mandible with 3 – 4 elements on basis; endopod with eight setae; exopod 1 - segmented with 2 – 5 setae. Maxillulary coxal epipodite represented by one seta. Maxilla with three endites on syncoxa; endopod discrete, with 2 – 3 setae. Maxilliped with 2 – 3 setae on syncoxa; basis with two setae on palmar margin; endopod represented by curved claw, accompanied by up to three accessory seta (e). P 1 inner basal spine not sexually dimorphic. P 1 exopod 3 - segmented; exp- 2 elongate, about 2 – 2.5 times length of exp- 1 and between two-thirds and 80 % length of endopod, with inner seta; exp- 3 very small, typically with two pinnate spines, one geniculate seta or claw, and one non-geniculate seta. P 1 endopod 2 - segmented; enp- 1 elongate, longer than exopod, with well developed inner seta originating either from near end of proximal quarter of segment or from middle third of segment, segment margins without area of reduced chitinization; enp- 2 very small, with one naked minute seta and two dentate claws (or one of claws replaced by geniculate seta / spine). P 2 – P 4 rami 3 - segmented. P 2 endopod ♂ typically 3 - segmented and not modified; occasionally 2 - segmented but with same total number of spines / setae as in ♀ (M. costata comb. nov., M. variabilis sp. nov.). P 3 endopod ♂ 3 - segmented; without inner setae on enp- 3 and outer spine modified into spinous apophysis. P 4 endopod ♂ occasionally with one (M. bulbosa comb. nov.) or two (M. minuta comb. nov.) less (er) inner seta (e) on enp- 3. Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe with five setae; inner margin without transverse striae. P 5 ♀ exopod elongate, with six or eight elements. P 5 ♂ endopodal lobe with two elements, outer one shortest; exopod 1 - or 3 - segmented, with 6 – 7 elements in total. Vestigial P 6 ♀ represented by three minute setae. P 6 ♂ with three setae.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFD7FFBCFF51D13AE09BFBFD.taxon	type_taxon	Type species. Thalestris forficula Claus, 1863 = Microthalestris forficula (Claus, 1863) comb. nov. (by monotypy; Sars 1905: 123) [species inquirenda].	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFD7FFBCFF51D13AE09BFBFD.taxon	discussion	Other species. Thalestris forficuloides Scott & Scott, 1894 = M. forficuloides (Scott & Scott, 1894) comb. nov.; Parastenhelia gracilis Brady, 1910 = M. gracilis (Brady, 1910) comb. nov.; M. littoralis Sars, 1911; Parastenhelia antarctica Scott, 1912 = M. antarctica (Scott, 1912) comb. nov.; Parastenhelia costata Pallares, 1982 = M. costata (Pallares, 1982) comb. nov.; Parastenhelia minuta Pallares, 1982 = M. minuta (Pallares, 1982) comb. nov.; Parastenhelia bulbosa Gee, 2006 = M. bulbosa (Gee, 2006) comb. nov.; M. campbelliensis sp. nov.; M. polaris sp. nov.; M. santacruzensis sp. nov.; M. sarsi sp. nov.; M. variabilis sp. nov. Additional species inquirendae. Harpacticus spinosus Fischer, 1860; Parastenhelia spinosa (Fischer, 1860) sensu Pesta (1959), sensu Marinov (1974), sensu Wells & Rao (1987) and sensu Kornev & Chertoprud (2008); Parastenhelia spinosa f. bulgarica Apostolov, 1968; Microthalestris forficula (Claus, 1863) sensu Wilson (1932 b); Microthalestris littoralis Sars, 1911 sensu Willey (1935) and sensu Nicholls (1945). Species incertae sedis. Thalestris karmensis Boeck, 1865; Dactylopus longipes Boeck, 1873.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE1FFBAFF51D568E78BFD72.taxon	discussion	The original descriptions of two species that had previously been designated, validly or invalidly, as a type species (Sars 1905: 123; Lang 1944: 13), cannot be used as the basis for their identification. Fischer’s (1860) description of Harpacticus spinosus is grossly inadequate and renders the identification of the species indeterminable. Pending the collection of material from Madeira it must be set aside as a species inquirenda in Microthalestris since it does not qualify as a reference for morphological comparison with populations of Microthalestris (or Parastenhelia) spp. that have subsequently been assigned to it. The various morphs of P. spinosa described by Pesta (1959), Wells & Rao (1987) and Kornev & Chertoprud (2008) as well as Apostolov’s (1968) forma bulgarica are indeterminable (see Table 1 for armature formulae of P 1 – P 5).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE1FFBAFF51D568E78BFD72.taxon	description	The allegedly cosmopolitan distribution of Microthalestris spinosa comb. nov. is a direct result of the inadequacy of the original description and the adoption of Lang’s (1948) excessively broad concept of the species. The great majority of the records below are probably false or indeterminable: Iceland: Bay of Faxaflói (Ólafsson et al. 2001); Seltjarnarnes (Steinarsdóttir et al. 2003); Hvassahraun (Steinarsdóttir & Ingólfsson 2004). Norway: Fanafjorden, south of Bergen (Dommasnes 1969). Sweden: Øresund (Dahl 1948); Gullmarfjord (Lang, 1948); northern Øresund (Hagerman 1966). Germany: Kiel Bay (Klie 1949; Anger & Scheibel 1976). Scotland: St. Abbs (Moore 1973, 1974; Hicks 1980); Firth of Forth (Shiells 1983); Great Cumbrae Island (Preston & Moore 1988, 1990). England: Wembury (Colman 1940) and Plymouth Sound (Wieser 1952) in Devon; Isles of Scilly (Wells 1961, 1970); Northumberland (Moore 1973, 1974). Wales: Pembrokeshire (Crothers 1966); Anglesey and Menai Strait (Geddes 1972); north Wales (Lintas & Seed 1994); Holyhead Island (Jarvis & Seed 1996). Isle of Man (Moore 1932; Bruce et al. 1963). France: Bassin d’Arcachon (Renaud-Debyser 1963 a, 1963 b); Nord-Finistère (Brittany) (Le Guellec 1988). Spain: Canary Islands, Tenerife (Noodt 1955 b); River Ebro estuary (Sabater 1986). Portugal: Azores (Chapman 1955; Chapman & Santler 1955). Italy: Sorrento (Pesta 1959); Laguna di Lesina (Ceccherelli & Mistri 1990); Valle Smarlacca, northern Adriatic (Cristoni et al. 2004). Montenegro: Budva (Petkovski 1955). Romania: Agigea (Pintea & Klemens 1957; Şerban 1959). Bulgaria: Gulf of Varna and Sozopol (Caspers 1951); Sozopol (Marinov 1974). Black Sea (Sergeeva 2003). Turkey: Datça-Bozburun Peninsulas, Muğla (Alper 2009; Alper et al. 2010); Dilek Peninsula, Aydın (Alper et al. 2015); Çatal and Kara Islands of Bodrum (Yıldız & Karaytuğ 2018). Israel: Achziv, Caesarea (= Keisarya or Qaysaria) and Palamachim (Por 1964). Russia: Kandalaksha Bay, White Sea (Kornev & Chertoprud 2008); Barents Sea (Dvoretsky & Dvoretsky 2010); Sea of Okhotsk (Chertoprud et al. 2015). Mozambique: Inhaca Island (Wells 1967). Andaman and Nicobar Islands: Aberdeen (Port Blair) and Mayabandar (Wells & Rao 1987); South Andaman (Jayabarathi 2016 – as P. spinosa spinosa). China: South China Sea (Chertoprud et al. 2009). Australia: Port Phillip Bay, Victoria (Walker-Smith 2003). New Zealand: Island Bay, Wellington (Coull & Wells 1983); undisclosed locality (Webber et al. 2010). Fiji: Moce Island (Wells 1978). Bermuda: Baileys Bay and Trunk Island (Coull 1970 b; Coull & Herman 1970). Tristan da Cunha (Wiborg 1964). Canada: Nanaimo Estuary (Kask et al. 1982); Lower Prospect, Nova Scotia (Johnson & Scheibling 1986, 1987 a, 1987 b). U. S. A.: North Carolina continental shelf (Coull 1971 b); central and northern California (Watkins 1983); Port Valdez, Prince William Sound (Cordell 2000; Hines & Ruiz 2001) and Kachemak Bay, Cook Inlet (Ruiz et al. 2006) in Alaska; Puget Sound (Haas et al. 2002), Elliot Bay (Toft & Cordell 2006; Toft et al. 2010), Crescent Bay and Freshwater Bay (Shaffer et al. 2020) in Washington State; recorded on Japanese tsunami-driven marine debris along west coast of North America from Alaska to California (Carlton et al. 2013, 2017; Cordell 2018; Gillman 2018; Miller et al. 2018). Brazil: Porto de Galinhas (Sarmento et al. 2009; Sarmento & Santos 2012 a, 2012 b; Barroso et al. 2018) and Serrambi, Tamandaré, Rio Doce, Piedade and Paiva (Barroso et al. 2018) in Pernambuco State; Arraial do Cabo, Rio de Janeiro State (Sarmento et al. 2012). Similarly, Claus’s (1863) description of Thalestris forficula is so inadequate that it is only recognizable as a Microthalestris by a figure of the P 1. Although it should be classified as a species of doubtful identity needing further investigation (species inquirenda) and – as a designated type species – will not best serve stability and universality, this does not of itself invalidate Sars’s (1905) type fixation. The following records of Microthalestris forficula are considered indeterminable: Iceland: Seydisfjord (Jespersen 1940; Klie 1941 – both as Parastenhelia forficula). Faroe Islands: off Suðuroy and between Streymoy and Eysturoy (Stephensen 1929). Northern Ireland: Ardglass, Co. Down (Williams 1954). Germany: Kiel Bay (Klie 1929; Kunz 1935). Scotland: Fairlie and Hunterston in the Firth of Clyde (Scott 1900 – as Thalestris forficulus); between Leith and Portobello, Firth of Forth (Scott 1906). England: Plymouth region, Devon (Norman & Scott 1906; Marine Biological Association 1931, 1957). Isle of Man: Port Erin and Port St Mary (Fraser (1936). France: Roscoff, Brittany (Monard 1935 a; Jakubisiak 1936). Italy: Venice Lagoon (Grandori 1912, 1914; Pesta (1920). Tunisia: Cartaghe (Salammbô) (Monard (1935 b – as Parastenhelia forficula). U. S. A.: Gulf of Maine (north of Georges Bank) (Bigelow 1926); Woods Hole (Wilson 1932 b). Australia: Port Denison, Western Australia (Nicholls 1945 – as Parastenhelia forficula). New Zealand: Otago Harbour (Brady 1899 – as Thalestris forficula).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE7FFBBFF51D1E2E7DAFDC2.taxon	description	Original description. Scott & Scott (1894): 142 – 144, 149; Plate IX, figs 4 – 9. Additional description. Scott (1894 b): 255 – 256; Plate X, figs 13 – 25.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE7FFBBFF51D1E2E7DAFDC2.taxon	materials_examined	Type locality. Scotland, Firth of Forth; mud near low-water mark at Seafield, in the vicinity of Leith.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE7FFBBFF51D1E2E7DAFDC2.taxon	diagnosis	Differential diagnosis. Microthalestris. Body length 730 μm in ♀, smaller in ♂. Antenna with 2 - segmented exopod bearing one seta on exp- 1 and two lateral and three apical elements on exp- 2. P 1 exopod about two-thirds length of endopod; exp- 2 elongate, about twice as long as exp- 1, and about 45 % length of enp- 1; insertion point of inner seta of enp- 1 at 25 % of inner margin length; exp- 3 with two unipinnate spines and two geniculate setae; enp- 2 with one minute seta, one geniculate seta and one geniculate claw. Armature pattern of ♀ P 2 – P 4: P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ with elongate exopod (about 2.4 times as long as maximum width), inner margin and proximal half of outer margin straight, with eight elements, proximal outer one long, outer apical one short; endopodal lobe with five elements, innermost one short. P 5 ♂ exopod 1 - segmented, with seven elements; endopodal lobe with two elements, outer one shortest. Armature of P 6 ♂ unconfirmed. Caudal ramus seta V with slightly swollen proximal part. Notes. Unconfirmed records of M. forficuloides include Inveraray and near Largabruach in Loch Fyne, Scotland by Scott (1897), Holy Island in Northumberland, England by Brady (1904) and Norman & Brady (1909 – as M. forficula) and the Isle of Man by Herdman (1897 – as Thalestris forficuloides). Scott (1899) recorded T. forficula (= probably Microthalestris polaris sp. nov.; see below) dredged at 55 m depth off Cape Gertrude in Franz Josef Land and added, with a slight inclination, that T. forficuloides is probably to be regarded as merely a local form of the latter. This view was reiterated in a later paper reporting on the presence of the species (as “ T. forficulus ”) in Fairlie and Hunterston in the Firth of Clyde (Scott 1900). Scott (1906), accepting Sars’s (1905) synonymy, reported the species (as M. forficula) from pools near low-water between Leith and Portobello in the Firth of Forth but it is likely that his record referred to T. forficuloides. Norman & Scott (1906) recorded M. forficula from rock-pools at and above the high-water mark in Plymouth, Devon, listing T. forficuloides as a synonym, but the identity of this record remains indeterminable.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE6FFB9FF51D147E149FE76.taxon	discussion	Comparison of the various reports dealing with M. littoralis reveals the true state of confusion reigning in the genus. In Norway the species shows virtually the same distribution as M. sarsi sp. nov., extending from the south coast northwards to the Trondhjem Fjord (Sars 1911). Several authors have reported on the sympatric occurrence of M. littoralis with other species but in the great majority of the cases the authenticity of the former is rendered doubtful by the absence of an adequate description or a definite statement of authority for the identification. For example, according to Wells (1961) three forms of P. spinosa (typica, littoralis, bulbosa) occur sympatrically in the Isles of Scilly while the latter two also coexist in the River Exe estuary (Wells 1963 a). Both Monard (1935 a) and Jakubisiak (1936) found that M. littoralis and M. forficula are sympatric in the Roscoff region in Brittany while Wilson (1932 b) recorded both species from the Woods Hole region. Farran (1913) documented significant variation in female body size (500 – 850 μm) in his material of M. littoralis from Clare Island and Blacksod Bay in Ireland which he perceived as a possible indication of the presence of a second species. Lang’s (1934, 1936 b, 1948) failure to recognize the sympatric occurrence of Microthalestris species and his subsequent proposal of an all-encompassing, highly variable species concept of P. spinosa are the primary reasons why most published records of both M. littoralis and M. sarsi sp. nov. cannot be accepted as accurate. Although most workers accepted Lang’s views and perpetuated the myth of a cosmopolitan morphologically plastic species, others advocated that M. littoralis deserved distinct species status (e. g. Willey 1935; Jakubisiak 1936; Sewell 1940; Nicholls 1945; Roe 1958, 1960; Pór 1960). Sars (1911) claimed that M. littoralis differed from his M. forficula (= M. sarsi sp. nov.) in the more compact antennule, the longer and more slender P 1, the number of exopodal setae on the P 5 (six vs eight) and the normally developed seta V on the caudal rami. Although the species was recorded from virtually all around the Norwegian coastline, no males were discovered by Sars (1911). Despite the absence of information on the armature formula of P 2 – P 4, M. littoralis is reinstated here based on the differences listed above. The only reliable records of the species are those by Sars (1911). Several authors have provided concise illustrated descriptions of what they believed to be M. littoralis. Brian’s (1921: 77 – 80) description of M. littoralis from the Gulf of Genoa included detailed illustrations of various developmental instars (naupliar stages I, III – V and copepodids III – V) while his descriptions of the adults are unfortunately not of a comparable standard. Little useful information is revealed about the morphology of the female except for the presence of a single egg sac and the normally developed caudal ramus seta V. The armature formula of P 1 – P 5 is completely unknown although Brian (1921: 78) does mention that the distal endopodal segment of P 3 has one spine and three plumose setae (“ … una gracile spina e tre setole piumate … ”). His description of the male shows the typical sexual dimorphism on the P 3 endopod (bearing an inner seta on enp- 1 and - 2, and an apophysis and two apical setae on enp- 3), and the 1 - segmented condition of the male P 5 exopod (bearing seven setae). The presence of two penicillate spines on the antenna (already expressed in copepodid IV: his Plate VIII, Fig. 17) provides evidence that Brian’s (1921) material does not belong to Microthalestris, representing instead a member of Penicillicaris gen. nov. (see below). Given its geographical proximity it may be conspecific with Car’s (1884) Thalestris pectinimana. Wilson (1932 b: 205) recorded ten females from Cuttyhunk Harbor, Massahusetts which he identified as M. littoralis. Unfortunately, the only drawings of the habitus in dorsal aspect and the P 5 are adapted from Sars (1911), making it impossible to decide on the identity of his material. Lang’s (1934) illustrated record of P. forficula var. littoralis from Campbell Island differs from M. littoralis in the morphology of the P 1, female P 5 and caudal ramus seta V and is here renamed as M. campbelliensis sp. nov. (see below). Willey (1935: 82 – 83) recorded one ovigerous female from Harrington Sound in Bermuda which according to the author agreed with Sars’s (1911) description of M. littoralis in displaying 9 - segmented antennules (although the segmentation was not disclosed by Sars) and unmodified caudal ramus setae. Much of Willey’s discussion centres around morphological differences in the antenna and swimming legs encountered between this “ typical ” specimen and his more abundant, sympatric, forma penicillata. The latter is here upgraded to species level and fixed as the type of Penicillicaris gen. nov. (see below) while the former is considered indeterminable based on the limited information presented by Willey. Monard’s (1937: 59, 62, Fig. 3 – E) description of P. forficula littoralis from Algeria, presenting a single illustration of the male P 5 exopod, does not contain the bare minimum required for any meaningful statement on its identity. Sewell (1940: 191 – 196) distinguished three different forms in his material of Parastenhelia littoralis obtained from seaweed washings in Addu Atoll, Maldives. The typical form, which he considered identical with Sars’s (1911) M. littoralis, shows distinct differences with the original description in the female antennule, P 1 and P 5, unambiguously ruling out such conspecificity. The presence of penicillate spines on the antennary endopod and on exp- 3 and enp- 2 of P 1 indicates that Sewell’s (1940) material represents a member of Penicillicaris gen. nov. (see below). Given their morphological disparity, we cannot concur with the author that both Thalestris forficula sensu Thomson (1883) and M. forficula sensu Monard (1928) are misidentified records of M. littoralis. Nicholls (1945: 4) provided illustrations of both sexes of Parastenhelia forficula var. littoralis from Port Denison in Western Australia. Females differ slightly from Sars’s (1911) description of M. littoralis in (a) the proportionally longer P 1 exp- 2 and enp- 1, and (b) the P 5 endopodal lobe which is narrower and less truncate. Pending information on the antennule, antenna and swimming legs in both forms and on the male of M. littoralis, the Australian population is provisionally regarded as distinct from the Norwegian one.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE6FFB9FF51D147E149FE76.taxon	description	Original description. Sars (1911): 369 – 370, Supplement plate 11, Fig. 1.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE6FFB9FF51D147E149FE76.taxon	materials_examined	Type locality. Sars (1905) recorded material from several places on the south and west coasts of Norway, and in the Trondhjem Fjord. Since he did not specify which specimens the illustrations were based on, the type locality encompasses all of their respective places of origin (ICZN Art. 73.2.3).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE6FFB9FF51D147E149FE76.taxon	diagnosis	Differential diagnosis. Microthalestris. Body length 600 μm in ♀, unknown for ♂. Antennule segmentation and antenna unconfirmed. P 1 exopod about 80 % length of endopod; exp- 2 elongate, about twice as long as exp- 1, and about half as long as enp- 1; insertion point of inner seta of enp- 1 at 25 % of inner margin length; exp- 3 with two unipinnate spines and two geniculate setae; enp- 2 with one minute seta, one geniculate seta and one geniculate claw. Armature pattern of ♀ P 2 – P 4 and ♂ P 3 endopod unknown. P 5 ♀ with elongate exopod (about three times as long as maximum width), inner margin and proximal two-thirds of outer margin straight, with six elements; endopodal lobe with five elements, innermost one short. P 5 – P 6 ♂ unconfirmed. Caudal ramus seta V normally developed.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE6FFB9FF51D147E149FE76.taxon	discussion	Notes. Most records of M. littoralis (either as a distinct species or as a form of Parastenhelia forficula or P. spinosa) are not accompanied by a description and must therefore remain indeterminable: Sweden: Øresund (Lang 1936 b – as Parastenhelia forficula var. littoralis). Ireland: Clew Bay and Blacksod Bay, Co. Mayo (Farran 1913; Farran et al. 1915); Clare Island (Southern 1915); Dalkey Island and The Muglins, Co. Dublin (Roe 1958 – as Parastenhelia spinosa var. littoralis); Lough Hyne (Ine), Co. Cork (Roe 1960 – as P. spinosa var. littoralis). Northern Ireland: Ardglass and Kilclief, Co. Down (Williams 1954). England: Isles of Scilly (Wells 1961 – as P. spinosa forma littoralis); River Exe (Wells 1963 a – as P. spinosa forma littoralis). France: Roscoff, Brittany (Monard 1935 a; Jakubisiak 1936). Italy: Gulf of Genoa Brian (1917, 1921); Sardinia (Brian 1923 a). Croatia: Palagruža (Steuer (1912); Rovinj (Brian 1923 b; Vàtova 1928); Island of Šolta (Jakubisiak 1933). Greece: Dodecanese Islands (Brian 1928 a); Rhodes, Astypalaia, Tilos, Symi, Kos, Karpathos (Brian 1928 b). Algeria: Îlot de la Marine (Amirauté) and Bab-el-Oued (Monard 1936 – as Parastenhelia (Microthalestris) forficula littoralis); Algiers and Bou Ismaïl (Castiglione) (Monard 1937 – as P. forficula littoralis). Ukraine: Yalta, Crimean Peninsula (Marcus & Pór 1960 – as Parastenhelia littoralis). Bermuda: Harrington Sound (Willey 1935). U. S. A.: Chesapeake Bay (Wilson (1932 a); Cuttyhunk Island (Wilson (1932 b). India: Porites Bay off Krusadai Island, Tamil Nadu (Krishnaswamy 1957); Visakhapatnam coast (Sarma 1974 a, 1974 b, 1974 c; Krishna Murty 1983) and Visakhapatnam Harbour (Sarma & Ganapati 1975) in Andhra Pradesh (all as Parastenhelia littoralis). Andaman and Nicobar Islands: South Andaman (Jayabarathi 2016 – as Parastenhelia spinosa littoralis). Malaysia: Peninsular Malaysia (Zaleha et al. 2010 – as Parastenhelia littoralis); Straits of Johor (Mahadi et al. 2014). Australia: Port Denison, Western Australia (Nicholls 1945 – as Parastenhelia forficula littoralis).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE4FFB6FF51D2EBE615F8E5.taxon	description	Sars (1905) treated Thalestris karmensis and T. forficuloides as synonyms of M. forficula but did not provide any justification. Although Sars and Boeck were contemporaries in the early stages of their careers it is unlikely that the former had the opportunity to examine Boeck’s material of T. karmensis thirty years after his untimely death in 1873. Boeck (1865) did not present any illustrations and the essence of his concise description translates as follows: “ The body is elongated, but the cephalothorax is somewhat inflated. The abdomen is short and wide, with rows of spinules around the posterior margin of the two middle somites. The caudal rami are wider than long. The antennules are short, with the second segment being about as long as the following two segments combined. The bases of the maxillipeds are oval and their claws very long. The first pair of swimming legs are elongated, with the endopod bearing a short seta at the end of the proximal quarter of the inner margin. The P 1 exopod is shorter than the endopod; the middle segment has spinules along the outer margin, and a longer spine near its distal corner; the claws on exp- 3 are very strong and several times longer than the segment. The oval exopod of P 5 is longer than the endopodal lobe. ” Based on Boeck’s (1865) description no positive statement can be made as to the validity of T. karmensis or its relationship to other parastenheliids. The little information that is given raises grave doubts that it belongs to Microthalestris at all or that it is a member of the Parastenheliidae. The species must be redescribed and pending this it is considered here as species incertae sedis in Microthalestris. Microthalestris forficuloides is similar to Sars’s material in P 1 exopod / endopod length ratio, the position of the inner seta on P 1 enp- 1, the absence of the inner seta on P 4 exp- 1, the presence of only one inner seta on P 4 exp- 3, the maximum number of eight elements on the female P 5 exopod, and the slightly swollen proximal part of caudal ramus seta V in the female, however, differs from it in the following suite of characters: (a) antennary exopod has one lateral seta on exp- 1 and two lateral setae on exp- 2 (vs two and one, respectively), (b) P 3 exp- 1 lacks the inner seta, (c) P 3 exp- 3 has three inner setae (vs two), (d) the female exopod is shorter (2.4 vs 2.8 times as long as maximum width), (d) the male exopod is 1 - segmented (vs 3 - segmented), and (e) the larger body size (♀♀: 730 μm vs 580 μm). On the basis of these morphological discrepancies, the previously suggested conspecificity with M. forficuloides can be ruled out and, consequently, Sars’s (1905) form is here attributed distinct specific rank as M. sarsi sp. nov. Lang (1948: 586) summarized earlier records attributed to Parastenhelia spinosa and stated that three forms of the species had been reported in the literature, forma typica, forma littoralis (for Microthalestris littoralis), and forma penicillata (for M. littoralis var. penicillata Willey, 1935). Although the diagnosis of his forma typica is clearly based on Sars’s material of M. forficula (and not Claus’s original description of the species), it is clear from the contents of his work that he did not intend to propose it as new (in which case “ M. typica ” would have become an available and the valid name for Sars’s material). Lang (1948: 587) unambiguously revealed that the three forms had no taxonomic meaning by stating that “ In Wirklichkeit is es unmöglich, die 3 Formen auseinanderzuhalten, den eine Menge Zwischenformen sind vorhanden, und die Merkmalkombinationen variieren in jeder denkbaren Weise ”. Although the forma typica could be construed as a name proposed for an infrasubspecific entity, it remains unavailable since it was not adopted for a species or subspecies before 1985 (ICZN Art. 45.6). Original description. Sars (1905): 123 – 124, Plate LXXVI. Additional description. Lang (1936 b): 23 – 25; Figs 50 – 52 (as Parastenhelia forficula).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE4FFB6FF51D2EBE615F8E5.taxon	materials_examined	Type material. The female specimen illustrated by Sars (1905: Plate LXXVI) is here designated as the holotype of M. sarsi sp. nov. (ICZN Arts 16.4 and 72.5.6). The species can be differentiated by the characters listed in the diagnosis below and those mentioned and illustrated in Sars (1905) (ICZN Art. 13.1). Type locality. Sars (1905) recorded material from several places on the south and west coasts of Norway, and in the Trondhjem Fjord. Since he did not specify which specimens the illustrations were based on, the type locality encompasses all of their respective places of origin (ICZN Art. 73.2.3).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE4FFB6FF51D2EBE615F8E5.taxon	diagnosis	Differential diagnosis. Microthalestris. Body length 580 μm in ♀, considerably smaller in ♂. Antenna with 2 - segmented exopod bearing two setae on exp- 1 and one lateral and three apical elements on exp- 2. P 1 exopod about two-thirds length of endopod; exp- 2 elongate, about 2.1 times as long as exp- 1, and about 60 % length of enp- 1; insertion point of inner seta of enp- 1 at 25 % of inner margin length; exp- 3 with two unipinnate spines and two geniculate setae; enp- 2 with one minute seta, one geniculate seta and one geniculate claw. Armature pattern of ♀ P 2 – P 4: P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ with elongate exopod (about 2.8 times as long as maximum width), inner margin and proximal half of outer margin straight, with eight elements, proximal outer one long, outer apical one short; endopodal lobe with five elements, innermost one well developed. P 5 ♂ exopod 3 - segmented, with six elements (outer element of exp- 1 absent or extremely reduced); endopodal lobe with two elements. Armature of P 6 ♂ unconfirmed. Caudal ramus seta V with slightly swollen proximal part.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE4FFB6FF51D2EBE615F8E5.taxon	etymology	Etymology. The specific epithet is named after Georg Ossian Sars (20 April 1837 – 9 April 1927), eminent Norwegian copepodologist, who provided the first illustrated account of the species under the name Microthalestris forficula (Claus, 1863). Notes. Microthalestris sarsi sp. nov. can readily be distinguished from other congeners that exhibit a 3 - segmented exopod in the male by the absence (or possibly extreme reduction) of the outer seta on exp- 1. Confirmation of this character in Lang’s (1936 b) male of Parastenhelia forficula from the Øresund corroborates its conspecificity with M. sarsi sp. nov. Sars (1905) himself listed the records from the British Isles (Scott 1894 b, 1897; Scott & Scott 1894 – all as T. forficuloides; Scott 1900 – as T. forficulus), the Bohuslän coast in Sweden (P. T. Cleve, unpubl. data), between Kolguev and Novaya Zemlya (Scott & Scott 1901 – as T. forficulus), Bear Island (Bjørnøya) and Hope Island, Svalbard (Scott & Scott 1901 – as T. forficulus), Franz Josef Land (Scott 1899 – as T. forficula), and the Arctic islands north of Grinnell Land, Canada (subsequently published by Sars 1909) as valid but considered the conspecificity of Scott’s (1894 a – as T. forficula) record from the Gulf of Guinea questionable. It is now clear that at least the illustrated records of T. forficuloides (Scott 1894 b; Scott & Scott 1894) are not conspecific with M. sarsi sp. nov. (see above) and that the Arctic records by Scott (1899), Scott & Scott (1901) and Sars (1909) may in reality refer to M. polaris sp. nov. (see below).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEAFFB4FF51D38FE5E5FB82.taxon	description	Additional descriptions. Chislenko (1967: 140 – 144, Figs 45 – 46); Chislenko (1977: 246, Fig. 6 – 1, 2).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEAFFB4FF51D38FE5E5FB82.taxon	materials_examined	Type material. The female specimen illustrated by Mielke (1974: Abb. 9 A – B) is here designated as the holotype of M. polaris sp. nov. (ICZN Arts 16.4 and 72.5.6). The species can be differentiated by the characters listed in the diagnosis below and those mentioned and illustrated in Mielke (1974) (ICZN Art. 13.1). Type locality. Svalbard archipelago, Spitsbergen, Longyearbyen; littoral zone.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEAFFB4FF51D38FE5E5FB82.taxon	diagnosis	Differential diagnosis. Microthalestris. Body length 550 – 800 μm in ♀, 400 – 420 μm in ♂. Antenna with 2 - segmented exopod bearing two setae on exp- 1 and two lateral and three apical elements on exp- 2; armature of endopod unconfirmed but likely without penicillate spines. P 1 exopod about 75 % length of endopod; exp- 2 elongate, about 2.5 times as long as exp- 1, and about 55 % length of enp- 1; insertion point of inner seta of enp- 1 at 25 % of inner margin length; exp- 3 with two unipinnate spines and two geniculate setae; enp- 2 with one minute seta and two non-geniculate claws. Armature pattern of ♀ P 2 – P 4: P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ with elongate exopod (about 2.4 times as long as maximum width), inner margin and proximal half of outer margin straight, with eight elements, proximal outer one long, outer apical one short; endopodal lobe with five elements, innermost one well developed. P 5 ♂ exopod 3 - segmented, with seven elements, outer seta of exp- 1 very long, longer than or as long as outer basal seta; endopodal lobe with two elements. Armature of P 6 ♂ unconfirmed. Caudal ramus seta V with slightly swollen proximal part.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEAFFB4FF51D38FE5E5FB82.taxon	etymology	Etymology. The species name is derived from the Latin polaris, meaning polar, and refers to the Arctic distribution of this species. Notes. The species appears so far to be restricted to the Arctic Ocean and its marginal waterbodies such as the White Sea. It has been recorded from Kandalaksha Bay (Chislenko 1967), Svalbard (Mielke 1974) and Franz Josef Land (Chislenko 1977). Kornev & Chertoprud (2008: 196 – 197, Fig. 5.94 – A, Б, B) also reported P. spinosa from the White Sea, including an aberrant specimen, and reproduced Chislenko’s (1967) illustrations of the P 1 and P 5 of both sexes. They mentioned substantial variation in female body length (550 – 800 μm) but this is a verbatim account of Chislenko’s (1967) measurements. It is also not clear whether the variability included in their armature formulae of P 3 – P 4 (number of inner setae on P 3 – P 4 exp- 3 and P 4 enp- 3; Table 1) is based on their own obervations of White Sea material or reflects variability reported in the literature. Since the authors provided no direct evidence that they were dealing with M. polaris sp. nov., their record is at present to be considered as unconfirmed. This uncertainty is exacerbated by the fact that, despite the reported variability on other segments, the White Sea specimens appear to be consistent in the lack of the inner seta on P 3 exp- 1 (which is present in M. polaris sp. nov.). Chislenko (1967) collected two females from washings of the demosponge, Semisuberites cribrosa (Esperiopsidae), in Franz Josef Land but this association is to be regarded as accidental (Huys 2016). Previous records of Thalestris forficula from Arctic localities may refer to M. polaris sp. nov. but in the absence of morphological or other compelling evidence they must remain unconfirmed. These include the records from Lille- Karajak fjord in western Greenland (Vanhöffen 1897), Cape Gertrude in Franz Josef Land (Scott 1899), between Kolguev and Novaya Zemlya (Scott & Scott 1901 – as T. forficulus), Bear Island (Bjørnøya) and Hope Island, Svalbard (Scott & Scott 1901 – as T. forficulus), the Arctic islands north of Grinnell Land, Canada (Sars 1909) and Seydisford in Iceland (Jespersen 1940; Klie 1941 – both as Parastenhelia forficula). Similarly, the Arctic records of Parastenhelia spinosa from Iceland (Ólafsson et al. 2001; Steinarsdóttir et al. 2003; Steinarsdóttir & Ingólfsson 2004), Kandalaksha Bay in the White Sea (Brotskaya 1962; Chertoprud et al. 2005) and from mesozooplankton in the Barents Sea (Dvoretsky & Dvoretsky 2010) require authentication before they can be considered conspecific with M. polaris sp. nov.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE9FFB5FF51D7B2E705FAC1.taxon	discussion	Microthalestris gracilis (Brady, 1910) comb. nov. belongs to a close-knit group of subantarctic species characterized by the position of the inner seta on P 1 enp- 1 which originates from the middle third of the segment. This subantarctic group includes M. gracilis, M. antarctica (Scott, 1912) comb. nov., M. costata (Pallares, 1982) comb. nov., M. santacruzensis sp. nov. and M. variabilis sp. nov. In all other members of the genus the inner seta of P 1 enp- 1 is situated further proximally, at around 25 % the length of the inner margin of the segment. Interestingly, the latter condition is also displayed by M. minuta (Pallares, 1982) comb. nov. from Tierra del Fuego and M. campbelliensis sp. nov. from Campbell Island, suggesting the independent colonization of the subantarctic region by two different lineages. Microthalestris gracilis comb. nov. is known from a single female collected in the Baie de l’Observatoire (Observatory Bay) along the northern shore of the Golfe du Morbihan in the Kerguelen (Brady 1910: 513; Textfig. VIII). A second record by Jayabarathi (2016) from seagrass beds in South Andaman is highly questionable. The original description includes illustrations of the antennule, P 1, P 2, P 5 and caudal rami but is deficient in many aspects. The armature is unknown for P 3 – P 4 except for Brady’s (1910) dubious claim that their distal endopodal segments have only three elements. The P 2 is shown to have two inner setae on exp- 3 which is also questionable. Since within the Parastenheliidae this condition is only found in members of Thalestrella and Foweya (Table 1) it may indicate that Brady’s (1910) illustration is either based on an observational error or in reality refers to P 3 or P 4. Alternatively, the possibility that P. gracilis represents a member of Foweya cannot be excluded but this would require information on the male and the sexual dimorphism in this species. Another unusual feature of P. gracilis is the endopodal lobe of P 5 which is remarkably small and displays only four setae. Lang’s (1948) concept of P. spinosa grouped together all the various forms in which the middle segment of P 1 exopod was at least four times as long as broad, with the exception of P. gracilis which was maintained as a distinct species mainly on the difference in the site of origin of the inner seta of P 1 enp- 2 and the shape of the female P 5. Scott (1912: 561; Plate IV, figs. 25 – 33) described a second subantarctic species, P. antarctica, from Scotia Bay (60 ° 43 ’ 42 ” S, 44 ° 38 ’ 33 ” W) in the South Orkney Islands and compared the female (males are unknown) with P. gracilis. Gurney (1927: 544) suggested that P. antarctica appeared to be identical with P. gracilis and Lang (1934: 23) formally treated them as synonyms but neither author gave a justification for this claim. This course of action appears premature since significant differences can be observed in the morphology of P 1 (length of exopod, in particular exp- 2; length of claws on enp- 2) and P 5 (shape of exopod), both of which were dismissed as irrelevant by Lang (1934, 1936 a, 1948). Although an unusual degree of intra-individual and intra-population variability has been recorded in the armature patterns of P 2 – P 4 in some species of Parastenhelia (e. g. Mielke 1974, 1990) such variability is not expressed in the morphology of either P 1 or P 5. Lang (1936 a: 53; Figs 36 – 37) reported two females from Berkeley Sound, Port Louis in the Falkland Islands which he attributed to P. gracilis. His illustration of the P 1 leaves no doubt that he was dealing with the geographically closer P. antarctica since it is fully compatible in the relative lengths of its rami, exp- 2 and claws on exp- 3 with Scott’s (1912) original description. Thomson (1883, 1913) recorded numerous Thalestris - like specimens from Otago Harbour, Dunedin in the South Island of New Zealand and assigned them to Claus’s (1863) T. forficula [see also Brady (1899) who found it abundantly in the same location]. Lang (1934, 1936 a) initially listed Thomson’s record under the synonymy of Parastenhelia forficula var. littoralis and subsequently, again without giving any specific justification, classified it as conspecific with P. spinosa (Lang 1948). Wells et al. (1982) suggested that the Otago Harbour material may belong to P. megarostrum but were unable to make a firm recommendation for this assignment on morphological grounds (see below). Thomson’s (1883) illustrations (Plate X, figs 16 – 21) show several similarities with the original description of P. antarctica, including (a) the general outline and proportional segmental lengths of the female antennule; (b) P 1 exopod with elongate exp- 2; (c) P 1 endopod showing similar L: W ratio for enp- 1; claws on enp- 2 elongate as in P. antarctica; and (d) female P 5 setation and general shape of exopod. Based on these similarities we consider it likely that Thomson’s (1883) T. forficula belongs to P. antarctica, the only main difference being the position of the inner seta on P 1 enp- 1 which inserts more distally. The female antennule of Thomson’s specimens was described as 8 - segmented but reinterpretation of his illustration (Plate X, fig. 17) suggests it has nine segments (as in P. antarctica). The presence of four setae on the proximal segment indicates that it represents the second segment and that the real first segment was not observed or lost during dissection. The aesthetasc that originates from a ventral pedestal of the fourth segment is shown to insert on the anterodistal corner of this segment (in reality the fifth) in Thomson’s drawing; this can be explained by assuming that the author observed the antennule in dorsal aspect. Based on the arguments above, Microthalestris antarctica (Scott, 1912) comb. nov. is here reinstated as a valid species. It is very close to M. costata comb. nov. in the morphology of the P 1 and the foliaceaous ♀ P 5 exopod but differs from it in the armature of the distal endopodal segment of P 4 (Table 1).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE8FFB2FF51D675E4B1FC8A.taxon	description	Original description. Lang (1934): 25 – 27, Figs 49 – 56.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE8FFB2FF51D675E4B1FC8A.taxon	materials_examined	Type material. The dissected male specimen illustrated by Lang (1934: Figs 52 – 56) is here designated as the holotype of P. campbelliensis sp. nov. (ICZN Arts 16.4 and 72.5.6). The species can be differentiated by the characters mentioned in the diagnosis below and those illustrated in Lang (1934) (ICZN Art. 13.1). Type locality. New Zealand, Campbell Island group, Campbell Island (Motu Ihupuku); Perseverance Harbour (South Harbour); 40 m depth.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE8FFB2FF51D675E4B1FC8A.taxon	diagnosis	Differential diagnosis. Microthalestris. Body length unknown. Antenna with 2 - segmented exopod, armature unconfirmed; bearing two setae on exp- 1 and two lateral and three apical elements on exp- 2; armature of endopod unconfirmed but likely without penicillate spines. P 1 exopod about 70 % length of endopod; exp- 2 elongate, about 2.1 times as long as exp- 1, and about 55 % length of enp- 1; insertion point of inner seta of enp- 1 at 20 % of inner margin length; exp- 3 with four elongate setiform elements; enp- 2 with one minute seta and two non-geniculate claws. Armature pattern of ♀ P 2 – P 4: see top of next page. P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ with elongate exopod (about three times as long as maximum width), inner margin and proximal half of outer margin slightly convex, with six elements, outer ones about equally long, outer apical one shorter than inner apical one; endopodal lobe with five elements, innermost one short. P 5 ♂ exopod 3 - segmented, with seven elements, outer seta of exp- 1 very long, longer than outer basal seta; endopodal lobe with two elements. Armature of P 6 ♂ unconfirmed. Caudal ramus seta V with slightly swollen proximal part.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFE8FFB2FF51D675E4B1FC8A.taxon	etymology	Etymology. The species name alludes to Campbell Island (Motu Ihupuku), an uninhabited subantarctic island of New Zealand, where the type locality is situated. Note. Although Lang (1934: 26) claimed his material consisted solely of a pair in amplexus the condition of the caudal ramus (Fig. 49) was figured in two different females. The species is so far only known from the type locality.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEFFFB3FF51D0BBE585FDBA.taxon	description	urn: lsid: zoobank. org: act: 6 A 60 E 30 B- 3 CF 6 - 4 E 72 - 915 D- 0 EF 22 C 6 BD 66 B	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEFFFB3FF51D0BBE585FDBA.taxon	discussion	Pallares (1963, 1968) recorded both sexes of a Parastenhelia species from the Río Deseado, Santa Cruz in Argentina. Although the material was attributed to P. gracilis it is radically divergent from both this species and P. antarctica in the shape of the female P 5 exopod. In the Argentine specimens the inner margin and the proximal two-thirds of the outer margin are virtually straight while in both P. gracilis and P. antarctica they are distinctly convex. Additional differences can be observed in the length of some of its elements, in particular the proximal outer seta of the exopod which is clearly longer and the inner element on the endopodal lobe which is minute. Pallares (1968) states that the male P 5 exopod has seven elements but does not figure the limb nor discloses the number of segments; however, in her Ph. D. dissertation (Pallares 1963) she stated “ Exopodo con tres sedas externas, una interna y dos largas apicales ” suggesting that all elements originate from a single segment and the total is six rather than seven. The Argentine material differs from all species currently assigned to Parastenhelia in the reduced armature formula of the swimming legs. Pallares (1968) only illustrates the male P 3 and gives no information in the text about P 2 and P 4; however, the absence of inner setae on P 2 exp- 1 and - 2 in conjunction with the presence of only one inner seta on exp- 3 differentiates the Río Deseado specimens from all known members of the genus. They further differ from P. gracilis in the length of the P 1 exopod (exp- 2 being distinctly longer) and the shape of caudal ramus seta V (swollen at the base). Although generally accepted as the first and only illustrated report of the male of P. gracilis (Mielke 1990; Bodin 1997; Wells 2007) there is no doubt that Pallares’s (1968) description deals with a different, as yet undescribed, species for which we coin the name P. santacruzensis sp. nov. Pallares (1975), in a subsequent report on littoral harpacticoids from Tierra de Fuego, recorded P. gracilis from Bahía Cook on Isla de Los Estados (Tierra del Fuego) but the authenticity of this record remains to be confirmed.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEFFFB3FF51D0BBE585FDBA.taxon	description	Original description. Pallares (1968): 63 – 65, Plate XIX.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEFFFB3FF51D0BBE585FDBA.taxon	materials_examined	Type material. Pallares’s (1968) original material no longer exists (M. Sciberras, pers. commn). The dissected male specimen illustrated by Pallares (1968: 64) in her Plate XIX is here designated as the holotype of P. santacruzensis sp. nov. (ICZN Arts 16.4 and 72.5.6). The species can be differentiated by the characters listed in the diagnosis below and those mentioned and illustrated in Pallares (1968) (ICZN Art. 13.1). Type locality. Pallares (1968) collected material from a number of localities in the Deseado River, Santa Cruz Province, Argentina: Punta Cavendish, Dos Hermanas, Punta Guanaco, Restinga Chaffers, Isla Quiroga and Bahía Uruguay. Since Pallares (1968) did not specify which specimens she based her illustrations on, the type locality encompasses all of their respective places of origin (ICZN Art. 73.2.3).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEFFFB3FF51D0BBE585FDBA.taxon	diagnosis	Differential diagnosis. Microthalestris. Body length 890 μm in ♀, 700 μm in ♂. Antennule 9 - segmented and haplocer in ♂. Antenna with 2 - segmented exopod bearing two setae on exp- 1 and one lateral and three apical elements on exp- 2; endopod without penicillate spines. P 1 exopod about 0.8 times length of endopod; exp- 2 elongate, about 2.5 times as long as exp- 1. P 3 exopod armature pattern [0.0.123]; endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ with elongate exopod (about 3.2 times as long as wide), inner margin and proximal two-thirds of outer margin straight, with six elements, proximal outer one long, outer apical one minute; endopodal lobe with five elements, innermost one minute. P 5 ♂ with six elements on 1 - segmented exopod; endopodal lobe with two elements. P 6 ♂ with three setae. Anal operculum smooth. Caudal ramus seta V with swollen base.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEFFFB3FF51D0BBE585FDBA.taxon	etymology	Etymology. The species is named after the Santa Cruz Province, located in the southern part of Argentina where the type locality is located.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEEFFB0FF51D1AAE1D1FD8C.taxon	description	Original description. Mielke (1990): 158 – 164, Figs 1 – 5.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEEFFB0FF51D1AAE1D1FD8C.taxon	materials_examined	Type material. The dissected female specimen illustrated by Mielke (1990: Figs 1 – 3, 4 A, 5) is here designated as the holotype of P. variabilis sp. nov. (ICZN Arts 16.4 and 72.5.6). The species can be differentiated by the characters mentioned in the diagnosis below and those illustrated by Mielke (1990) (ICZN Art. 13.1). Type locality. Argentina, Province of Tierra del Fuego, Antarctica and South Atlantic Islands; Isla Grande Tierra del Fuego, Bahía Lapataia near Ushuaia.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEEFFB0FF51D1AAE1D1FD8C.taxon	diagnosis	Differential diagnosis. Microthalestris. Body length 580 – 780 μm in ♀, 420 – 560 μm in ♂. Antenna with 2 - segmented exopod bearing two setae on exp- 1 and two lateral and three apical elements on exp- 2; endopod without penicillate spines. P 1 exopod about 70 % length of endopod; exp- 2 elongate, about 1.8 times as long as exp- 1, and about 40 % length of enp- 1; insertion point of inner seta of enp- 1 at 40 % of inner margin length; exp- 3 with two unipinnate spines and two geniculate setae; enp- 2 with one minute seta, one geniculate and one non-geniculate claw. Armature pattern of ♀ P 2 – P 4: P 2 endopod ♂ 2 - segmented (enp- 2 and - 3 forming compound segment), armature pattern [1.221]. P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ with ovate exopod (about 1.9 times as long as maximum width), with six elements, distal outer one longest, outer apical one short and naked; endopodal lobe with five elements, innermost one well developed. P 5 ♂ exopod 3 - segmented, with seven elements, outer seta of exp- 1 long and naked; endopodal lobe with two short spiniform elements. Armature of P 6 ♂ unconfirmed. Caudal ramus seta V normally developed, seta IV with bulbous swelling near base.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEEFFB0FF51D1AAE1D1FD8C.taxon	etymology	Etymology. The species name alludes to the high level of variability observed in the swimming leg armature.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFEDFFB0FF51D1B1E1DCF9E7.taxon	discussion	As pointed out above, Brady’s (1910) description of M. gracilis is rudimentary, his illustrations are far from precise and the armature formula of P 2 – P 5 is misleading. The species is consequently omitted from the key and the reader is advised to consult the original description. The same applies to M. littoralis which is only known from the female and for which no information on swimming leg armature exists.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFECFFAEFF51D276E0FFFB01.taxon	description	Mielke (1994 b: 152) recognized two evolutionary lineages within Karllangia, the more primitive branch comprising K. arenicola arenicola and K. tertia, and the more advanced one represented by K. psammophila, K. arenicola bengalensis and both Costa Rican species (K. obscura, K. pulchra). As pointed out by Gee (2006) the outstanding problem within the genus Karllangia is the fact that its type species, K. arenicola, apparently does not display the most important synapomorphy that supports the common ancestry of the remaining species (including the type species of Thalestrella). This character state which distinguishes Karllangia from other members of the Parastenheliidae is the modification of the male antenna (allobasal seta much more plumose, exp- 1 smaller and with only one strong, plumose seta, exp- 2 broader with more strongly plumose setae than in ♀). The original description by Noodt (1964) of K. arenicola from coralline sands on the Egyptian coast of the Red Sea does not mention any sexual dimorphism in the male antenna which could be attributed to either an observational error or to genuine absence. This ambiguity surrounding the type species has now been removed by synonymizing Karllangia with its senior subjective synonym Thalestrella. The presence of antennary sexual dimorphism in its type species, T. ornatissima, has been demonstrated by Gee (2006) who confirmed a number of additional genus-level apomorphies in the species, including the compressed appearance of the distal antennulary segments in the female, the presence of only three setae on exp- 2 of the antenna, two setae on the mandibular exopod, and one seta on the maxillipedal syncoxa; the absence of the inner seta on P 2 – P 4 exp- 1; and, a male P 5 exopod with only four setae. Additional apomorphies which were not observed by Gee (2006) include (a) male antennule subchirocer, compound segment 5 (XIV – XX) swollen; (b) the male antennule with only three segments distal to the geniculation (XXI – XXIII, XXIV – XXV, XXVI – XXVIII); (c) ventral surface of segment 2 of male antennule with characteristic, backwardly directed, spiniform element, either displaying a serrate or flagellate tip (Wells & Rao 1987: Fig. 110 b; Mielke 1994 b: Fig. 6 A); and (d) outer spine on P 3 enp- 3 reduced in size in ♂ (Mielke 1994 b: Fig. 6 C; Gee 2006: Fig. 5 C; Fig. 7 herein).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFECFFAEFF51D276E0FFFB01.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 3 enp- 3 outer spine, P 5 – P 6 and urosomal segmentation. Body subcylindrical; posterior margin of cephalothorax (typically) and all somites (except anal somite) typically with deeply divided, denticulodigitate or lobate, hyaline frills (plain in T. obscura comb. nov. and possibly T. arenicola comb. nov.). Rostrum defined at base, of moderate size, reaching at most to end of second segment of antennule; linguiform. Anal operculum semicircular, variously ornamented; usually bordered with fine spinules or denticles. Caudal ramus at most as wide as long, with dorsal spinular row; with seven setae, setae IV – V not basally inflated in ♀. Antennule ♀ compact, 8 - or indistinctly 9 - segmented; segment 1 not elongate, distal four or five segments small with combined length only slightly longer than segment 4; with aesthetascs on segments 4 and 8 (or 9). Antennule ♂ subchirocer, 7 -, 8 - or indistinctly 9 - segmented; ventral surface of segment 2 with characteristic, backwardly directed, spiniform element; with geniculation between segments 5 and 6 (or 6 and 7 when indistinctly 9 - segmented) and 2 – 3 segments distal to geniculation; compound segment 5 (XIV – XX) swollen, occasionally subdivided dorsally by transverse surface suture; with aesthetasc on segments 5 and 7 or 8; segmental homologies of 8 - segmented antennule: I, II – VIII, IX – XII, XIII, XIV – XX, XXI – XXIII, XXIV – XXV, XXVI – XXVIII. Antenna sexually dimorphic. Female antenna with proximal endopodal segment largely fused to basis forming allobasis, with short pinnate seta on abexopodal margin; exopod 2 - segmented (or segments partially fused), proximal segment with 1 – 2 setae, distal segment with one lateral and two apical elements (surrounded at base by serrate hyaline frill); distal endopodal segment without penicillate elements. Male antenna with enlarged, highly plumose seta on allobasis; exopodal segments of different size, with only one seta on proximal segment and all lateral setae enlarged and highly plumose. Mandible with three elements on basis; endopod with eight setae; exopod 1 - segmented, with 2 – 4 setae. Maxillulary coxal epipodite represented by one seta. Maxilla with three endites on syncoxa; endopod discrete, with 4 – 5 setae. Maxilliped with one seta on syncoxa; basis with 0 – 2 setae on palmar margin; endopod represented by pinnate claw, accompanied by one accessory seta. P 1 exopod 3 - segmented; segments subequal; exp- 2 with or without inner seta; exp- 3 with two pinnate spines and one geniculate spine and one (non-) geniculate seta. P 1 endopod 2 - segmented; enp- 1 elongate, about twice as long as exopod, with well developed, pinnate inner seta originating from proximal half in a region of reduced chitinization of segment wall; enp- 2 very small, with one naked minute seta and two unipinnate claws, longest of which geniculate. P 2 – P 4 rami 3 - segmented; outer spine of P 3 enp- 3 ♂ reduced in size. Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe triangular, with five setae; inner margin without transverse striae; exopod elongate (2 – 5 times longer than wide), with 5 – 6 elements. P 5 ♂ endopodal lobe with two elements; exopod 1 - segmented, with 4 – 5 elements. Vestigial P 6 ♀ represented by 2 – 3 short setae. P 6 ♂ with three setae / spines.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFECFFAEFF51D276E0FFFB01.taxon	type_taxon	Type species. Thalestrella ornatissima Monard, 1935 a (by monotypy). Other species. Karllangia arenicola Noodt, 1964 = T. arenicola (Noodt, 1964) comb. nov.; K. psammophila Wells, 1967 = T. psammophila (Wells, 1967) comb. nov.; Parastenhelia reducta Apostolov, 1975 = T. reducta (Apostolov, 1975) comb. nov.; K. arenicola bengalensis Wells & Rao, 1987 = T. bengalensis (Wells & Rao, 1987) comb. nov.; K. obscura Mielke, 1994 b = T. obscura (Mielke, 1994 b) comb. nov.; K. pulchra Mielke, 1994 b = T. pulchra (Mielke, 1994 b) comb. nov. Species inquirenda. Parastenhelia ornatissima (Norman & Scott, 1905) sensu Por (1964).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF3FFAFFF51D632E6D4FCCE.taxon	description	Additional descriptions. Wells (1961): 267 – 268; Fig. 2 B. Gee (2006): 2613 – 2621; Figs 1 – 6.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF3FFAFFF51D632E6D4FCCE.taxon	materials_examined	Type locality. France, Brittany, Roscoff area; near Roc’h Iliévec; dredged from sand. Notes. Por (1964) recorded seven females (400 μm) of Parastenhelia ornatissima from detritus-laden littoral sand in Achziv, northern Israel (Levantine coast). He provided illustrations (Plate 11; figs 95 – 96) of the female P 5 and the posterior half of the abdomen in dorsal aspect. Bodin (1968) reported both sexes in a midlittoral sand sample from Banc du Bûcheron (Ile de Ré) near La Rochelle, but gave no morphological details, while Soyer (1971) subsequently recorded it from Banyuls-sur-Mer. Gee (2006) recorded T. ornatissima comb. nov. as the most abundant species in the harpacticoid community at low tide level on St Martin’s Flat in the Isles of Scilly (only one specimen was found near the high-water mark, cf. Gee 2009). His detailed study of the previously undescribed male confirmed that this species has a sexually dimorphic antenna and belongs in the genus Karllangia (= Thalestrella). Differences with Monard’s (1935 a) description were dismissed as of no significance and attributed to the author’s imprecision in observing the minutiae of the mouthparts and, more importantly, the correct setation of the P 4 exopod (two inner setae on exp- 3 instead of three in the Scillonian material). Conversely, Por’s (1964) specimens, which allegedly share the same armature on P 4 exp- 3 as Monard’s material, were considered a potentially different species. In reality, Por did not illustrate leg 4 nor did he provide a setal formula, stating only that “ Antennae, mouthparts and swimming legs are typical ”. The only two discrepancies observed between Monard’s type material and the specimens from the Israeli coast are related to the anal operculum, which has fewer (7) and larger spinous projections, and the P 5, which has slightly shorter setae (particularly on the endopodal lobe). These characters, together with the length: width ratio of the P 5 exopod (shorter compared to the specimens from the Isles of Scilly), are known to show a certain level of variability and do not warrant the proposal of a new species for the Levantine material, at least not until it has been subjected to thorough re-examination. Gee (2006) showed that the specimens from the sand flats of Tresco and St. Martins (Isles of Scilly), previously identified by Wells (1961) as Parastenhelia anglica, are conspecific with T. ornatissima comb. nov. This claim was primarily based on the abundance and location data of the material, in conjunction with the description of the male P 5 given in Wells (1961). Thalestrella ornatissima comb. nov. shares the presence of the inner seta on P 1 exp- 2 with T. bengalensis comb. nov., T. psammophila comb. nov. and T. pulchra comb. nov. but differs from these species in the presence of only two outer spines on P 3 exp- 3. The only other records of the type species are those by Willems et al. (2008) who obtained it from sandy shell gravel at 17 – 23 m depth in the Koster-area, Swedish west coast, and by Alper & Sak (2018) who collected it in the mediolittoral zone of the Sarımsaklı coast (Balıkesir) in northwestern Turkey. A potentially morphologically close species was recorded under the name P. cf. ornatissima from the northern Gulf of Mexico (Thistle 2003; Thistle et al. 1995 a, 1995 b; Suderman & Thistle 1998).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF2FFAFFF51D07EE509F9B4.taxon	materials_examined	Type locality. Egypt, Red Sea coast near Hurghada; 2 – 4 m depth, coralline sand. Notes. As suggested by Gee (2006: 2645) the sexual dimorphism of the antenna was probably overlooked by Noodt (1964) due to the small size of the species. Other discrepancies which are probably attributable to observational errors include (a) the absence of well developed, deeply incised hyaline frills on the body somites (although plain frills have also been observed in T. obscura comb. nov.); (b) the indistinctly 3 - segmented antennary exopod in the female, displaying an incomplete suture dividing the distal segment, (c) the absence of the minute seta on P 1 enp- 2, (d) the presence of only one seta on P 5 endopodal lobe ♂ (the minute outer seta was probably overlooked), and (e) the absence of a conspicuous dorsal spinular row on the caudal ramus. The inner seta of P 1 enp- 1 is exceptionally long compared to other congeners. Thalestrella arenicola comb. nov. shares the presence of only two segments distal to the geniculation in the male antennule with T. bengalensis comb. nov. and T. reducta comb. nov., the absence of the inner seta on P 1 exp- 2 with T. obscura comb. nov. and T. reducta comb. nov., and the presence of only two inner setae on P 3 exp- 3 with T. ornatissima and T. reducta comb. nov. Thalestrella arenicola has not been recorded again since its original description (Noodt 1964). Rao (1980) listed “ Karllangia arenicala [sic] Noodt? ” from an unspecified locality in the Andaman and Nicobar Islands; in all likelihood this record refers to T. bengalensis. Colangelo et al. (2001) reported on a species, “ Karllangia cfr. arenicola ”, collected from a shallow hydrothermal seepage site off the Island of Panarea (Aeolian Islands) in the Tyrrhenian Sea.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF2FFACFF51D5D9E75EFD5E.taxon	materials_examined	Type locality. Mozambique, Inhaca Island; Ilha dos Portuguesos (Elephant Isle); intertidal zone, clean sand. Notes. Karllangia psammophila was originally described as a distinct species (Wells 1967: 292) but subsequently relegated by Wells & Rao (1987: 132) to a subspecies of K. arenicola along with another form (K. arenicola bengalensis) they described from material collected in the Andaman and Nicobar Islands. Wells & Rao claimed that K. arenicola was identical to K. psammophila in every single aspect except for the lack in both sexes of an attenuated outer distal corner on the posterior margin of the first antennulary segment. However, Gee (2006) reinstated the latter and K. arenicola bengalensis to full species status based on significant differences in the armature of the swimming legs and the ornamentation of the anal operculum. The author also examined paratype specimens of K. psammophila, revealing oversights in Wells’s (1967) description of the antenna (abexopodal margin of allobasis with seta; ♀ exopod with two setae on exp- 1 and one lateral and two distal setae on exp- 2), mandible (basis with three setae; exopod small, with two setae) and maxilliped (syncoxa with one seta; basis with two setae on palmar margin), and suspected that similar corrections apply to K. bengalensis. Thalestrella psammophila shares the presence of only five elements on the female P 5 exopod with T. obscura comb. nov. and T. bengalensis comb. nov., however, it differs from the former in the presence of well developed, denticulodigitate hyaline frills on the body somites and the presence of an inner seta on P 1 exp- 2, and from the latter by the attenuated outer corner of the proximal antennulary segment in both sexes, the second outer seta of the female P 5 exopod being distinctly longer (vs shorter) than the proximalmost seta, and the proximal outer seta of the male P 5 exopod distinctly shorter (vs longer) than the segment.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF1FFACFF51D1CEE5E1F91C.taxon	materials_examined	Type locality. Bulgaria, Burgas Province; Tsarevo (formerly Michurin) and Lozenets; sandy bottom, 5 m depth. Notes. Gee (2006) transferred Parastenhelia reducta as species incertae sedis to Karllangia, based on the female antennule with distinctly compressed distal segments, the maxillipedal syncoxa with only one seta, and the reduced armature and absence of sexual dimorphism of P 2 – P 4. The generic assignment of this species, known only from a single female and male from shallow sandy sediments along the Bulgarian Black Sea coast (Apostolov 1975), was tentative since the male P 5 exopod exhibits five setae (instead of four in the remaining Karllangia species) and, according to the original text description the antenna lacks sexual dimorphism. Armature patterns on the fifth legs are known to vary in many harpacticoid genera and small differences in setal numbers are therefore likely to be of less significance in delimiting their diagnostic boundaries. We concur with Gee (2006) that the hirsute bulbous structure in Apostolov’s Figure 3 h, partly concealed by the second segment of the male antennule, is conceivably the modified antennary exopod. Although a re-examination of Apostolov’s (1975) specimens would be desirable, there appears to be no reason to question its generic assignment. Parastenhelia reducta is here removed from the incertae sedis category of Karllangia and reinstated as a valid species of Thalestrella. Por’s (1964) specimens of T. ornatissima, the geographically closest record of the genus, agree with T. reducta comb. nov. in the length: width proportions of the female P 5 exopod but conspecificity can be ruled out on the basis of differences in the anal operculum and caudal ramus length (as measured in dorsal aspect). Thalestrella reducta comb. nov. can readily be distinguished from its congeners by the reduced armature on P 3 – P 4 (P 4 exp- 3 with two inner setae vs three, P 3 – P 4 enp- 3 with one inner seta vs two) and the presence of six elements on the male P 5 exopod. Apostolov (1975) did not figure the deeply incised hyaline frills on the body somites, however, his illustration (Fig. 3 a) of the urosome showing the long pleural extensions (probably an optical section of the frills) indicates their presence. T. reducta is the only species in the genus that displays four setae on the mandibular exopod.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF0FFADFF51D13AE024FCA0.taxon	materials_examined	Type locality. Costa Rica, beach slope of Manzanillo (Caribbean coast). Notes. Thalestrella obscura comb. nov. occurs sympatrically with T. pulchra comb. nov. at the type locality. It differs from the latter by the absence of well developed denticulodigitate hyaline frills on the abdominal somites and the inner seta on P 1 exp- 2, and by the presence of a tuft of very long setules on the male antennary allobasis near the articulation with the exopod, and only five elements on the female P 5 exopod.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF0FFADFF51D0D5E7C8FB61.taxon	materials_examined	Type locality. Costa Rica, beach slope of Manzanillo (Caribbean coast). Note. See T. obscura comb. nov. for differentiating characters.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF0FFAAFF51D614E734FD40.taxon	discussion	A dichotomous key to the six valid species of Thalestrella was provided by Gee (2006) which is updated below to include T. reducta comb. nov. and to rectify a mistake in his couplet 3 (T. obscura comb. nov. has only five setae on P 5 exopod ♀). Additional unidentified species have been recorded from Papua New Guinea (Willen 2000), the Galápagos Archipelago (Mielke 2003), Zanzibar (Gheerardyn 2007; Gheerardyn et al. 2008; Callens et al. 2012) and possibly Heron Island (Iwasaki 1994), indicating a circum (sub) tropical distribution of the genus.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF7FFABFF51D1F5E6A8FBDA.taxon	discussion	The justification for the proposal of this genus (Wells 1967: 300) was based exclusively on the 2 - segmented condition of the P 1 exopod in the type species P. minima Wells, 1967. Additional apomorphic character states shared with its congener, P. wellsi Rao, 1972, included the 8 - segmented condition of the ♀ antennule, the presence of a modified distal element (“ … expanded, bifid, hyaline structure … ”) on exp- 2 of the antenna, proximal endopodal segment of antenna without allobasis, reduction of the mandibular exopod (absent or represented by short seta), mandibular basis with only two setae; maxilliped with reduced armature (at most one seta on syncoxa, none on basis), inner basal spine of ♂ P 1 modified (with recurved tip), various reductions in the armature of P 2 – P 4 (Tables 1 – 2), P 5 endopodal lobe ♀ with only three elements, and P 5 exopod ♂ with only four elements. The modification of the inner basal spine of P 1 is shared with Psammoleptomesochra and Foweya; however, in members of the latter genus the inner spine is not recurved but typically straight, bifid apically, and carrying a flagellate extension at its tip (cf. Gee 2006: Figs 10 D, 13 A). It is conceivable that, within the Parastenheliidae, Paraleptomesochra is most closely related to Psammoleptomesochra (Mielke 1994 a).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF7FFABFF51D1F5E6A8FBDA.taxon	description	Both species of Paraleptomesochra are small-sized (♀: 274 – 336 μm; ♂: 276 – 315 μm) and exclusively known from sandy beaches. Rao (1980: 166) listed a species under the name “ Paraleptomesochra africana (Kunz) ” but this is a lapsus calami referring to Praeleptomesochra africana (Kunz, 1951) (family Ameiridae) (cf. Dev Roy & Venkataraman 2018). Finally, as yet unidentified species of Paraleptomesochra have been reported from shallow depths off the coast of northwestern Florida (Suderman & Thistle 1998) and from Heron Reef, Great Barrier Reef (Iwasaki 1994).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF7FFABFF51D1F5E6A8FBDA.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 1 inner basal spine, P 5 – P 6 and urosomal segmentation. Body cylindrical, short; posterior margin of prosomites and urosomites (except anal somite) with denticulodigitate hyaline frills. Rostrum fused at base, small, pointed. Anal operculum semicircular, bordered with fine setules. Caudal ramus about as long as wide, with dorsal spinular row; with at least six setae, setae IV – V not basally inflated in ♀. Antennule ♀ elongate and 8 - segmented in ♀, segment 1 not elongate, segments 6 – 7 shortest, with aesthetascs on segments 4 and 8. Antennule ♂ haplocer with three segments distal to geniculation; segmentation partly unconfirmed but probably at least 9 - segmented with segmental homologies as follows: I, II – VIII, IX – XII, XIII, XIV – XVIII, XIX – XX, XXI – XXIII, XXIV – XXV, XXVI – XXVIII. Antenna not sexually dimorphic; proximal endopodal segment completely separated from basis, without seta on abexopodal margin; exopod 2 - segmented, proximal segment with two setae, distal segment with 1 – 2 lateral and two apical elements (one of which enlarged, hyaline and bifid); distal endopodal segment without penicillate elements. Mandible with two elements on basis; endopodal armature unconfirmed; exopod absent or represented by a short seta. Maxillulary coxal epipodite unconfirmed. Maxilla with three endites on syncoxa; endopod unconfirmed. Maxilliped with 0 – 1 seta € on syncoxa; basis without armature on palmar margin; endopod represented by claw without accessory setae. P 1 inner basal spine sexually dimorphic (recurved in ♂). P 1 exopod 2 - segmented; exp- 2 slightly longer than exp- 1, with two distal and two outer pinnate elements. P 1 endopod 2 - segmented; enp- 1 elongate, at most slightly longer than exopod, with short inner seta, segment margins probably without area of reduced chitinization; enp- 2 very small, with one outer distal spine and one inner distal geniculate seta. P 2 – P 4 rami 3 - segmented; enp- 2 and - 3 without inner seta (e); inner seta of P 2 – P 3 exp- 1 reduced in size, absent in P 4. P 2 – P 3 endopods ♂ without sexual dimorphism. Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe with three setae (one inner, two distal); inner margin without transverse striae. P 5 ♀ exopod ovate, with six elements. P 5 ♂ endopodal lobe with 1 – 2 elements; exopod 1 - segmented, with four elements. Vestigial P 6 ♀ represented by one seta. P 6 ♂ with two setae.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF7FFABFF51D1F5E6A8FBDA.taxon	type_taxon	Type species. Paraleptomesochra minima Wells, 1967 (by original designation). Other species. P. wellsi Rao, 1972. Notes. Both Wells (1967) and Rao (1972) state that the rostrum is fused to the cephalothorax which, if correct, would be a unique condition in the family. The segmental homologies of the haplocer male antennule are unconfirmed although Rao (1972) describes it as 7 - segmented with the sixth segment partially divided by an indistinct suture in P. wellsi. The description of the armature on the caudal ramus (with four setae in P. minima, five in P. wellsi) is almost certainly incorrect. The presence / absence of the coxal epipodite of the maxillule is unconfirmed since neither Wells (1967) nor Rao (1972) describe the appendage. Similarly, it is not clear whether the maxillary endopod is expressed or the maxillipedal claw has any accessory elements.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF6FFABFF51D750E60BF88D.taxon	materials_examined	Type locality. Mozambique, Inhaca Island; Ilha dos Portuguesos (Elephant Isle); intertidal zone, clean sand. Notes. The species differs from its congener P. wellsi primarily in the armature of P 4 exp- 3 (121 vs 221), P 2 – P 4 enp- 3 (one apical element vs two) and the endopodal lobe of the male P 5 (two setae vs one). Additional differences include the caudal ramus which is slightly longer than wide in P. minima (vs as long as wide), the length of P 1 enp- 2 which is longer in P. wellsi (and the position of its inner seta being more proximal), the morphology of the setae on P 1 exp- 2 and the inner distal seta of P 2 – P 4 exp- 3 which are characterized by a row of stiff pinnules near the apex in P. wellsi, and the shape of the male P 5 exopod (trapezoid vs oval). Wells (1967: Text-fig. 54 F) shows three setae on the mandibular basis with the two proximal ones at a considerable distance from the distal one. It is conceivable that the short proximal “ setae ” are ornamentation elements similar to those described and figured by Mielke (1994 b: Figs 2 D, 8 A) for Thalestrella pulchra and T. obscura. Rao (1972: 4 F) illustrated similar (but more) setules in his description of P. wellsi in addition to two distal setae. Wells (1967) also described the mandibular endopod as 2 - segmented but this is unlikely; the exopod is apparently lacking but represented by a seta in P. wellsi. The absence of the seta on the maxillipedal syncoxa (present in P. wellsi) requires confirmation. Although Rao (1972) had previously described P. wellsi from mainland India, both Rao (1975) and Wells & Rao (1987) attributed their material from the Andaman and Nicobar Islands (Middle and South Andaman, Car Nicobar Island) to the geographically much more distant P. minima. The latter was subsequently recorded from the littoral zone of Lakshadweep (formerly known as the Laccadive, Minicoy and Aminidivi Islands) (Rao & Misra 1983; Rao 1991) and Little Andaman (Rao 1993).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFBFFA6FF51D38FE624FDC2.taxon	materials_examined	Type locality. India, Andhra Pradesh, Visakhapatnam (formerly Waltair), 17 ° 43 ’ 30 ” N 83 ° 20 ’ 30 ” E; intertidal zone, fine and medium sand, 10 – 30 cm below surface near half-tide level. Notes. Rao (1972) describes the male antennule as 7 - segmented with the sixth segment being partly subdivided. The segmentation pattern is here provisionally reinterpreted as 9 - segmented due to segment 4 (XIII) having been overlooked and the three segments distal to the geniculated accepted here as fully discrete. Rao’s (1972: Fig. 4 D) illustration of two aesthetascs originating from the swollen segment 5 must be attributed to an observational error. The species was subsequently recorded from two localities (Konark = Konarak, Puri) in Odisha (formerly Orissa) in the Bay of Bengal (Rao 1989).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFBFFA7FF51D173E16AFCF4.taxon	discussion	Mielke (1994 a: 258) placed Psammoleptomesochra in the “ Leptomesochra - complex ”, an artificial group of genera in the Ameiridae (Lang 1965; Wells 1967), and recognized a certain closeness with Parapseudoleptomesochra reducta Rao, 1972 (now type species of Raoleptomesochra Karanovic, 2004) and particularly Paraleptomesochra. Psammoleptomesochra displays many autapomorphic character states including, (a) antennary exopod without lateral setae on exp- 2; (b) mandibular basis with two setae; (c) mandibular exopod vestigial, represented by minute seta; (d) coxal epipodite on maxillule absent; (e) maxillipedal basis without setae; (f) P 1 exp- 2 without inner seta; (g) P 2 exp- 3 without inner setae; and (h) P 2 – P 4 enp- 3 with reduced distal setae (represented by two setules).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFBFFA7FF51D173E16AFCF4.taxon	description	Psammoleptomesochra shares the area of reduced chitinization on the elongate P 1 enp- 1 with Paraleptomesochra and Foweya, while the sexual dimorphism of the inner basal spine on P 1 is also displayed in Foweya and Thalestrella. Paraleptomesochra has many reductions in common with Psammoleptomesochra including the loss of setation elements on the mandibular basis and exopod, maxillipedal syncoxa and basis, P 2 – P 4 and endopodal lobe of ♀ P 5 (Table 2). In both genera the armature of P 2 – P 4 is strongly reduced but not all setal losses are evidence for their common ancestry. For example, as a result of convergent evolution, both genera display only four elements on P 3 exp- 3 but in the former the pattern consists of one inner, two distal and one outer element (s) [121] while in the latter there are two distal and two outer elements [022].	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFBFFA7FF51D173E16AFCF4.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 5 – P 6 and urosomal segmentation. Body subcylindrical; posterior margin of cephalothorax and all somites (except anal somite) with deeply divided, denticulodigitate, hyaline frills. Rostrum defined at base, small, reaching to proximal third of second segment of antennule; linguiform with rounded apex. Anal operculum semicircular, bordered with few (around six) strong spinules. Caudal ramus about twice as long as wide, with dorsal oblique row of spinules; with six setae, setae II – III short, pinnate and spiniform, setae IV – V not swollen at base. Antennule short and 9 - segmented in ♀, segment 1 not elongate, segments 7 – 8 shortest, with aesthetascs on segments 4 and 9; haplocer in ♂, segmentation, position of geniculation and segmental homologies unconfirmed. Antenna with basis; proximal endopodal segment with short abexopodal seta; distal endopodal segment without penicillate elements; exopod 2 - segmented, proximal segment with two lateral setae, distal segment with two apical elements. Mandible with two elements on basis; endopod with two lateral and 5 – 6 apical setae; exopod vestigial, represented by short seta. Maxillulary coxal epipodite absent. Maxilla with three endites on syncoxa; endopod discrete, with three setae. Maxilliped with one seta on syncoxa; basis without seta (e) but with short longitudinal spinular row on palmar margin; endopod represented by claw with one accessory seta. P 1 inner basal spine sexually dimorphic, recurved in ♂. P 1 exopod 3 - segmented, segments subequal in length; exp- 2 without inner seta; exp- 3 with two pinnate spines and two geniculate setae. P 1 endopod 2 - segmented; enp- 1 longer than exopod, with well developed, pinnate inner seta arising from area of reduced chitinization in middle third of segment; enp- 2 with one naked minute seta and two claws (one geniculate, one non-geniculate). P 2 – P 4 rami 3 - segmented; endopods short, at most as long as exp- 1 and - 2 combined; inner distal element of P 2 – P 4 exp- 3 setiform and thin; endopodal segments without inner setae except for P 4 enp- 1 bearing long pectinate seta; P 2 – P 4 enp- 3 with two vestigial distal setae and one outer spine. P 2 – P 4 endopods ♂ without sexual dimorphism. Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe with four elements, innermost spiniform; inner margin without transverse striae; exopod elongate, with six naked setae. P 5 ♂ endopodal lobe with two elements, outer one shortest; exopod 1 - segmented, with five elements, inner one vestigial. Vestigial P 6 ♀ represented by two setae. P 6 ♂ with two setae and one spine (innermost element).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFBFFA7FF51D173E16AFCF4.taxon	type_taxon	Type and only species. Psammoleptomesochra australis Mielke, 1994 a (by original designation).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFBFFA7FF51D173E16AFCF4.taxon	materials_examined	Type locality. Chile, Los Lagos Region, Chiloé archipelago, southeastern coast of Isla Chiloé, Quellón Viejo; sandy beach. Original description. Mielke (1994 a): 252 – 258; Figs 1 – 4. Notes. The endopodal armature pattern adopted herein differs from Mielke’s (1994 a: 257) formula in two aspects: the apical spiniform element on enp- 3 is here considered as the homologue of the outer spine in other genera, and the two minute and fine setae originating from the inner corner of enp- 3 are included in the formula as distal elements (Table 1). Mielke (1994 a) observed variability in the armature of the P 5 endopodal lobe (with 2 – 3 apical setae) in the female and the P 5 exopod (distal seta on outer margin variable in length) in the male.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFAFFA4FF51D099E05CFC05.taxon	discussion	Gee (2006) recognized the close resemblance between Parastenhelia anglica and Karllangia tertia and removed both species from their respective genera and placed them in a newly proposed genus Foweya. Both Foweya anglica (Norman & Scott, 1905) and F. tertia (Kunz, 1975) exhibit the narrowing of the outer cuticle of P 1 enp- 1 at around the insertion level of the inner seta (shared with Thalestrella and Psammoleptomesochra), the bifid, sexually dimorphic inner spine on the P 1 basis in the male (shared with Paraleptomesochra and Psammoleptomesochra), the 10 - segmented male antennule with only three segments distal to the articulation, the absence of sexual dimorphism in the endopod of P 3 (shared with Thalestrella, Paraleptomesochra and Psammoleptomesochra), and the most primitive swimming leg armature formula within the family, including two inner setae on P 2 exp- 3 and enp- 3 and three inner setae on P 3 enp- 3. Gee (2006) confirmed that the specimens, previously identified as Parastenhelia anglica from St. Agnes, St. Mary’s and Tresco (Isles of Scilly) by Wells (1961, 1970) and from Namibia by Kunz (1963), are conspecific with Thalestrella ornatissima (type locality: Roscoff, Brittany), and Foweya tertia (type locality: Eastern Cape, South Africa), respectively.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFAFFA4FF51D099E05CFC05.taxon	description	Males of the two known Foweya species can be differentiated by the segmentation of the P 2 endopod (3 - segmented in F. anglica, 2 - segmented in F. tertia) and the morphology of the fifth legs (with five exopodal setae in F. tertia vs six in F. anglica; endopodal lobe relatively weakly developed and both setae equally long in F. tertia vs well developed lobe and outer endopodal seta distinctly longer than inner one in F. anglica). Females (and males) can be distinguished by the ornamentation of the anal operculum (with approximately 30 small, closely set denticles in F. anglica vs numerous very fine spinules in F. tertia).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFAFFA4FF51D099E05CFC05.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 1 inner basal spine, P 5 – P 6 and urosomal segmentation; occasionally in segmentation of P 2 endopod. Body subcylindrical, short; posterior margin of cephalothorax and somites bearing P 2 – P 5 with plain hyaline frills; remaining urosomites (except anal somite) with denticulodigitate hyaline frills. Rostrum defined at base, of moderate size, reaching at most to end of second segment of antennule; linguiform. Anal operculum semicircular, bordered with fine spinules or denticles. Caudal ramus wider than long, with dorsal spinular row; with seven setae, setae IV – V not basally inflated in ♀. Antennule elongate and 9 - segmented in ♀, segment 1 not elongate, segments 5 – 8 shortest, with aesthetascs on segments 4 and 9; haplocer and 10 - segmented with three segments distal to geniculation in ♂, with geniculation between segments 7 and 8, segments 5 – 7 swollen and with aesthetasc on segment 5; segmental homologies in ♂: I, II – VIII, IX – XII, XIII, XIV – XVI, VII – XVIII, XIX – XX, XXI – XXIII, XXIV – XXV, XXVI – XXVIII. Antenna not sexually dimorphic; proximal endopodal segment largely fused to basis forming allobasis, with pinnate seta on abexopodal margin; exopod 2 - segmented, proximal segment with two setae, distal segment with two lateral and three apical elements; distal endopodal segment without penicillate elements. Mandible with three elements on basis; endopod with eight setae; exopod 1 - segmented, with four setae. Maxillulary coxal epipodite represented by one seta. Maxilla with three endites on syncoxa; endopod discrete, with 4 – 5 setae. Maxilliped with three setae on syncoxa; basis with two setae on palmar margin; endopod represented by pinnate claw, accompanied by two accessory setae. P 1 inner basal spine sexually dimorphic (with bifid tip in ♂). P 1 exopod 3 - segmented; exp- 2 only slightly longer than exp- 1, with inner seta; exp- 3 small, with two pinnate spines and two geniculate setae. P 1 endopod 2 - segmented; enp- 1 elongate, almost twice as long as exopod, with moderately long, pinnate inner seta originating from proximal third (at 35 % of ramus length in a region of reduced chitinization of segment wall); enp- 2 very small, with one naked minute seta and two unipinnate claws. P 2 – P 4 rami 3 - segmented (except P 2 endopod ♂ occasionally 2 - segmented); P 3 endopod ♂ not sexually dimorphic. Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe with five setae; inner margin without transverse striae; exopod elongate, with six elements. P 5 ♂ endopodal lobe with two elements; exopod 1 - segmented, with 5 – 6 elements. Vestigial P 6 ♀ represented by three short setae. P 6 ♂ with three setae.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFAFFA4FF51D099E05CFC05.taxon	type_taxon	Type species. Parastenhelia anglica Norman & Scott, 1905 = Foweya anglica (Norman & Scott, 1905) (by original designation). Other species. Karllangia tertia Kunz, 1975 = Foweya tertia (Kunz, 1975).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF9FFA4FF51D737E0BEF9BB.taxon	description	Additional descriptions. Norman & Scott (1906): 148 – 150; Plate X (Figs 10 – 11), Plate XIII (Figs 4, 9), Plate XIV (Fig. 6), Plate XVI (Fig. 4), Plate XVII (Fig. 4), Plate XXI (Fig. 2). Gee (2006): 2623 – 2631; Figs 7 – 11.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF9FFA4FF51D737E0BEF9BB.taxon	materials_examined	Type locality. England, south Cornwall, Fowey; dredging outside of Fowey harbour; 7.3 – 9.1 m.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF9FFA5FF51D5AAE0BBFEC6.taxon	description	Additional descriptions. Kunz (1963 – as P. anglica): 41 – 43; Figs 38 – 48. Gee (2006): 2631 – 2634; Figs 12 – 13.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF9FFA5FF51D5AAE0BBFEC6.taxon	materials_examined	Type locality. South Africa, Eastern Cape, off East London; shell gravel in tidal pools on a reef.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF8FFA2FF51D277E690FD7B.taxon	description	Kuru & Karaytuğ (2015) described a new species, Parastenhelia aydini, from a Turkish beach in the Eastern Mediterranean and considered it morphologically most similar to Pa. spinosa. The authors refer to their unpublished observations but it is not clear which “ population ” they identified as Pa. spinosa. They differentiated Pa. aydini from the latter by the combination of the following characters: (a) ♀ antennule 9 - segmented; (b) spinular rows near outer margin of P 1 – P 4 bases significantly reduced; (c) P 4 enp- 3 with four setae in both sexes; (d) ornamentation of endopodal and exopodal setae of ♀ P 5 different; (e) caudal ramus seta V of ♀ not swollen near the base; (f) modified element on the seventh segment of ♂ antennule not present; (g) apophysis on ♂ P 3 enp- 3 notched; and (h) ♂ P 5 exopod 2 - segmented, five setae in total. Unique characters displayed by Pa. aydini include (a) the modification of the endopodal claw on the maxilliped; the distal half does not taper to a sharp tip as in other members of the family, forming instead a spatulate structure fringed with a series of densely arranged serrations around the inner margin and rounded apex; (b) P 1 exp- 3 with one very long naked seta and three claws of different lengths, the longest one being 0.8 times the length of exp- 2; (c) P 1 enp- 2 claws radically divergent in size, with large one nearly three times longer than short one; (d) inner seta of P 2 – P 3 enp- 1 reduced; (e) spinous apophysis on ♂ P 3 endopod with foot-shaped tip; and (f) ♂ P 5 exopod 2 - segmented, with one very long outer seta on exp- 1 and four elements on exp- 2; the only other species in the Parastenheliidae that display a 2 - segmented exopod are Parastenhelia willemvervoorti sp. nov. and Johnwellsia bipartita gen. et sp. nov. which have six short elements, and one spine and two setae, respectively, on exp- 2. Based on this combination of apomorphic character states, Pa. aydini is her fixed as the type species of a new genus, Karaytugia gen. nov.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF8FFA2FF51D277E690FD7B.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 2 – P 3 endopods, P 5 – P 6 and urosomal segmentation. Body subcylindrical, short; posterior margin of cephalothorax and somites bearing P 2 – P 5 with plain hyaline frills; remaining urosomites (except anal somite) with serrate hyaline frills. Rostrum defined at base, of moderate size, reaching to about halfway second segment of antennule; elongate triangular. Anal operculum semicircular, bordered with fine spinules. Caudal ramus wider than long, with conspicuous spinular ornamentation; with seven setae, setae IV – V not basally inflated in ♀. Antennule elongate and 9 - segmented in ♀, segment 1 not elongate, segments 7 – 8 shortest, with aesthetascs on segments 4 and 9; haplocer and 11 - segmented in ♂, with geniculation between segments 7 and 8, segment 5 swollen and with aesthetasc; segmental homologies in ♂: I, II – VIII, IX – XII, XIII, XIV – XVII, XVIII, XIX – XX, XXI – XXII, XXIII, XXIV – XXV, XXVI – XXVIII. Antenna not sexually dimorphic; proximal endopodal segment largely separated from basis, with pinnate seta on abexopodal margin; exopod 2 - segmented, proximal segment with two setae, distal segment with two lateral and three apical elements; distal endopodal segment without penicillate elements. Mandible with three elements on basis; endopod with eight setae; exopod 1 - segmented, with two setae. Maxillulary coxal epipodite represented by one seta. Maxilla with three endites on syncoxa; endopod discrete, with three setae. Maxilliped with two setae on syncoxa; basis with two setae and longitudinal spinular row on palmar margin; endopod represented by conspicuously serrated claw, accompanied by three accessory setae. P 1 inner basal spine not sexually dimorphic. P 1 exopod 3 - segmented; exp- 2 much longer than others, nearly three times as long as exp- 1 and about half as long as enp- 1, with short inner seta; exp- 3 small, with two pinnate spines, one geniculate seta and one naked seta. P 1 endopod 2 - segmented; enp- 1 elongate, about 1.5 times length of exopod, with short, proximally inserted, pinnate inner seta, segment margins without area of reduced chitinization; enp- 2 very small, with one naked minute seta and two dentate claws. P 2 – P 4 rami 3 - segmented; inner seta of P 2 – P 3 exp- 1 reduced in size, absent in P 4. P 2 – P 3 endopods ♂ 3 - segmented. P 2 slightly more robust in ♂. P 3 endopod ♂ without inner setae (vs two in ♀) on enp- 3 and outer spine modified into apically truncated spinous apophysis. Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe with five setae, outermost well developed, innermost shortest; inner margin without transverse striae. P 5 ♀ exopod elongate, with six elements. P 5 ♂ endopodal lobe with two elements, outer one shortest; exopod 2 - segmented, with one long outer seta on exp- 1 and four elements on exp- 2. Vestigial P 6 ♀ represented by three minute setae. P 6 ♂ with three setae.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF8FFA2FF51D277E690FD7B.taxon	etymology	Etymology. The genus name is dedicated to Prof. Süphan Karaytuğ (Mersin University, Turkey), in recognition of his contributions to the harpacticoid fauna of Turkey, and under whose mentorship the next generation of harpacticoid experts in his home country emerged. Gender: feminine.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFF8FFA2FF51D277E690FD7B.taxon	type_taxon	Type and only species. Parastenhelia aydini Kuru & Karaytuğ, 2015 = Karaytugia aydini (Kuru & Karaytuğ, 2015) comb. nov. (by original designation herein).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFFFFA3FF51D1E9E52CFF0D.taxon	materials_examined	Type locality. Turkey, Kızkalesi, Mersin (36 ° 27.473 ’ N, 34 ° 08.647 ’ E); sand at 0.3 m depth adjacent to rocky shore. Notes. Kuru & Karaytuğ (2015) observed slight sexual dimorphism in the ornamentation of caudal ramus seta III (naked in ♀, minutely bipinnate in ♂). This observation needs confirmation based on a larger sample since only two males were examined. The authors also claim that the male antennule is 11 - segmented and present the armature pattern as follows: 1 - [1], 2 - [10], 3 - [5], 4 - [6], 5 - [3], 6 - [1 + (1 + ae)], 7 - [1 + 6 modified], 8 - [1 + 2 modified], 9 - [1], 10 - [4], 11 - [6 + acrothek]. The presence of four segments (their segments 8 – 11) distal to the geniculation is correct, however, the segmentation pattern proximal to the geniculation is ambiguous even though the authors illustrated it from different angles (their Fig. 7 B – E). Comparison with the ancestral male harpacticoid pattern (as in Hamondia superba Huys, 1990: cf. Huys & Boxshall 1991) clarifies the homologies of the proximal segments in K. aydini comb. nov. and reveals some observational errors in the original description. The homologies between actual segments (indicated in Arabic numerals) and ancestral segments (in Roman numerals) in the male groundpattern of the Harpacticoida are as follows (Huys & Boxshall 1991: 115, Fig. 2.4.4): 1 - [I], 2 - [II], 3 - [III – VIII], 4 - [IX – XII], 5 - [XIII], 6 - [XIV – XVI], 7 - [XVII], 8 - [XVIII], 9 - [XIX – XX], 10 - [XXI – XXII], 11 - [XXIII], 12 - [XXIV], 13 - [XXV], 14 - [XXVI – XXVIII]. In this ancestral 14 - segmented pattern the large aesthetasc is situated on segment 6 (and on segment 4 in the female), however, in all members of the Parastenheliidae segment 2 is not expressed as a discrete segment, forming instead a compound segment [II – VIII] and making the aesthetasc originate from actual segment 5. According to Kuru & Karaytuğ (2015) the aesthetasc arises from segment 6 in K. aydini comb. nov. This condition is extremely unlikely and is further exacerbated by the fact that ancestral segment XIII (= segment 4 in the parastenheliid groundpattern), which is expressed as a U-shaped bisetose sclerite in most male harpacticoids, was overlooked (although there is a hint of it in their Fig. 7 D), thus effectively shifting the position of the aesthetasc even further to segment 7. Their alleged sixth segment (their Fig. 7 C – D) shown to carry this sensory structure is in reality the cylindrical pedestal of segment 5 [= XIV – XVII] which partly conceals the real segment 6 [= XVIII] in ventral aspect. Finally, their segment 7 is a compound double segment homologous to ancestral segments XIX – XX. Kuru & Karaytuğ (2015) claim that the armature for this segment is “ 1 + 6 modified ” but this is almost certainly based on an observational error. Male harpacticoids typically display two spiniform elements on the anterior surface and one seta at the anterodistal corner (cf. Huys & Boxshall 1991: Fig. 115) and the same pattern is expressed in K. aydini comb. nov., the supernumerary elements being spinules. Similar spinular or setular ornamentation elements have been recorded on the same antennulary segment in many other harpacticoids such as in Neobradya pectinifera Scott, 1892 (Huys 1987: Fig. 1 B). The type species was also collected from Denizy-ıldızı beach in Urla (İzmir Province) in the Aegean Sea (Kuru & Karaytuğ 2015).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFEFFA0FF51D23FE6A6FDC2.taxon	description	Ever since the discovery and description of Thalestris pectinimana Car, 1884, the presence of penicillate elements on the antennary endopod and the distal exopodal and endopodal segments of P 1 in various related forms (Scott 1894 a; Brian 1921; Monard 1928; Willey 1935; Sewell 1940; Bodin 1964; Vervoort 1964) has been uncritically accepted as part of the documented variability allegedly displayed by Parastenhelia spinosa (Lang 1948 and others) and unfortunately did not receive the attention it deserved. Sewell (1940) for example, claimed that Monard’s (1928) penicillate form from Banyuls-sur-Mer referred to Parastenhelia littoralis while his own material from Bermuda listed under the latter name cannot possibly be conspecific with the species described by Sars (1911). A new genus, Penicillicaris gen. nov., is proposed here to accommodate four forms that had previously been subsumed in the Parastenhelia spinosa complex and are attributed full species rank. Another three forms are placed with reservations in the genus as species inquirendae: Microthalestris forficula (Claus, 1863) sensu Scott (1894 a), M. littoralis Sars, 1911 sensu Brian (1921) and M. littoralis Sars, 1911 f. scotti sensu Sewell (1940). A number of records refer to the penicillate form but are not accompanied by illustrations. Such indeterminable records include those from Visakhapatnam (formerly Waltair), Andhra Pradesh in India (Krishnaswamy 1957 – as Parastenhelia littoralis f. pencillata [sic]), Dalkey Island and The Muglins, Co. Dublin (Roe 1958) and Lough Hyne (Ine), Co. Cork (Roe 1960; Sloane et al. 1961) in Ireland (all P. spinosa f. penicillata), and Rovinj in Croatia (Zavodnik 1969 – as P. spinosa var. penicillata).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFEFFA0FF51D23FE6A6FDC2.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 3 endopod, P 5 – P 6 and urosomal segmentation; occasionally in P 4 endopod (Pe. penicillata sp. nov.). Body subcylindrical; condition of hyaline frills of cephalothorax and somites unconfirmed. Rostrum defined at base, reaching at most to halfway second segment of antennule; linguiform. Anal operculum semicircular, bordered with fine setules. Caudal ramus wider than long, without conspicuous spinular ornamentation; with at least six setae, setae IV – V typically inflated at base in ♀. Antennule ♀ slender and typically 8 - segmented (occasionally 9 - segmented), with aesthetascs on segments 4 and 8 (or 9). Antennule ♂ haplocer and presumably 10 - segmented; with geniculation between segments 6 and 7, and four segments distal to geniculation; segment 5 swollen and with aesthetasc; segmental homologies in ♂: I, II – VIII, IX – XII, XIII, XIV – XVIII, XIX – XX, XXI – XXII, XXIII, XXIV – XXV, XXVI – XXVIII. Antenna not sexually dimorphic; allobasis completely fused or partially divided, with endopodal pinnate seta on abexopodal margin; exopod 2 - segmented, proximal segment with two setae, distal segment with 1 – 2 lateral and three apical elements; free endopod with two penicillate elements. Mandible with 2 elements on basis; endopod with two lateral and 3 – 4 apical setae; exopod 1 - segmented, with two setae. Maxillulary coxal epipodite and maxilla unconfirmed. Maxilliped with 0 – 1 seta (e) on syncoxa; basis with one seta on palmar margin; endopod represented by claw with unconfirmed number of accessory setae. P 1 inner basal spine not sexually dimorphic. P 1 exopod 3 - segmented; exp- 2 elongate and much longer than others, with inner seta; exp- 3 with two penicillate spines, a geniculate seta or additional penicillate spine, and one non-geniculate seta. P 1 endopod 2 - segmented; enp- 1 longer than exopod, with short, proximally inserted, pinnate inner seta, inner segment margin without area of reduced chitinization; enp- 2 with one naked minute seta and two penicillate claws. P 2 – P 4 rami typically 3 - segmented, P 2 – P 3 endopods ♀ occasionally 2 - segmented (Pe. penicillata sp. nov.); inner seta of P 2 – P 4 exp- 1 reduced in size. P 3 endopod ♂ 3 - segmented; enp- 3 with spinous apophysis and no inner setae. P 4 endopod ♂ occasionally without inner setae on enp- 3 [1.1.021; in Pe. penicillata sp. nov.]. Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe with five setae; inner margin without series of transverse striae. P 5 ♀ exopod elongate, with six elements. P 5 ♂ endopodal lobe with two elements, outer one typically shortest; exopod 1 - segmented, with seven elements. P 6 unconfirmed in both sexes.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFEFFA0FF51D23FE6A6FDC2.taxon	etymology	Etymology. The generic name is derived from the Latin penicillus, meaning paintbrush, and the Greek suffix - caris, meaning shrimp, and refers to the presence of penicillate spines on the antennary endopod and both rami of the first swimming leg. Gender: feminine.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFEFFA0FF51D23FE6A6FDC2.taxon	type_taxon	Type species. Thalestris pectinimana Car, 1884 = Penicillicaris pectinimana (Car, 1884) comb. nov. (by original designation herein). Other species. P. maldivensis sp. nov.; P. penicillata sp. nov.; P. sewelli sp. nov. Species inquirendae. Microthalestris forficula sensu Scott (1894 a); M. littoralis Sars, 1911 sensu Brian (1921); M. littoralis Sars, 1911 f. scotti sensu Sewell (1940).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFDFFA1FF51D196E6ABFF0D.taxon	description	Original description. Car (1884): 248 – 249; Plate XVIII, Figs 1 – 8. Additional descriptions. Monard (1928): 348 – 349; fig. XVIII (as Microthalestris forficula). Bodin (1964): 128 – 129; Plate III, fig. 15 (as Parastenhelia spinosa var. penicillata).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFDFFA1FF51D196E6ABFF0D.taxon	materials_examined	Type locality. Italy, Trieste (Adriatic); plankton.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFDFFA1FF51D196E6ABFF0D.taxon	diagnosis	Differential diagnosis. Penicillicaris. Body length 650 – 800 μm [according to Monard (1928) but sex not specified; sometimes markedly smaller (350 μm!)]. Antennule 8 - segmented in ♀. Antenna with 2 - segmented exopod bearing two setae on exp- 1 and two lateral and three apical elements on exp- 2. Mandibular armature unconfirmed. Maxilliped with one seta on syncoxa and basis. P 1 exp- 3 with two penicillate spines, one geniculate seta and one short non-geniculate seta. P 1 enp- 1 about 1.15 times length of exopod. Armature pattern of ♀ P 2 – P 4: P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ exopod about 2.15 times as long as maximum width; inner and outer margins gradually converging towards apex; with 6 – 7 elements. P 5 endopodal lobe ♀ short, extending to about proximal fifth of exopodal length. P 5 exopod ♂ 1 - segmented, elongate – oval; with seven elements. P 5 endopodal lobe ♂ extending to insertion point of proximal inner seta of exopod; with two elements. Caudal ramus setae IV – V of ♀ inflated near base (forcipate type). Notes. Car (1884) was the first to note the presence of a single egg sac in a member of the Parastenheliidae. Monard (1928) observed variability in the number of setae on the P 5 exopod of the female, with six elements being the typical condition and seven (as in Car’s description) the rare one. The species is so far restricted to the western Mediterranean (Banyuls-sur-Mer, Marseille, Trieste).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFCFFA1FF51D23EE494F978.taxon	description	Original description. Sewell (1940): 191 – 193; Text-fig. 26 A – K.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFCFFA1FF51D23EE494F978.taxon	materials_examined	Type material. The dissected female specimen illustrated by Sewell (1940: Text-fig. 26 A, C, E – G, I – J )) is here designated as the holotype of P. maldivensis sp. nov. (ICZN Arts 16.4 and 72.5.6). The species can be differentiated by the characters mentioned in the diagnosis below and those discussed and illustrated by Sewell (1940) (ICZN Art. 13.1). Type locality. Maldive archipelago, Addu Atoll; seaweed washings.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFCFFA1FF51D23EE494F978.taxon	diagnosis	Differential diagnosis. Penicillicaris. Body length 440 – 560 μm in ♀, unknown for ♂. Antennule 9 - segmented in ♀; with four segments distal to geniculation in ♂. Antenna with 2 - segmented exopod bearing two setae on exp- 1 and one lateral and three apical elements on exp- 2. Mandible with two setae on basis; exopod 1 - segmented, with two setae. Maxillipedal armature unconfirmed. P 1 exp- 3 with three penicillate spines and one long non-geniculate seta. P 1 enp- 1 about 1.5 times length of exopod. Armature pattern of ♀ P 2 – P 4: P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ exopod elongate, about 3.8 times as long as maximum width; proximal half of inner and most of outer margin parallel; with six elements. P 5 endopodal lobe ♀ short, extending to about proximal quarter of exopodal length. P 5 exopod ♂ 1 - segmented, elongate – oval; with seven elements. P 5 endopodal lobe ♂ extending to insertion point of proximal inner seta of exopod; with two elements. Caudal ramus setae IV – V of ♀ not inflated near base.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFCFFA1FF51D23EE494F978.taxon	etymology	Etymology. The species name refers to the Republic of the Maldives where the type locality is located. Notes. Sewell’s (1940) illustration of the P 3 endopod (his Text-fig. 26 H) was inadvertently mislabelled as the P 2 endopod. The absence of the inner seta on P 1 exp- 2 (his Text-fig. 26 G) is probably based on an observational error.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFCFFDEFF51D5ECE699FA56.taxon	description	Original description. Willey (1935): 82 – 83; Figs 119 – 120, 122 – 126.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFCFFDEFF51D5ECE699FA56.taxon	materials_examined	Type material. The dissected female specimen illustrated by Willey (1935: Figs 119 – 120, 122 – 124) is here designated as the holotype of P. penicillata sp. nov. (ICZN Arts 16.4 and 72.5.6). The species can be differentiated by the characters mentioned in the diagnosis below and those discussed and illustrated by Willey (1935) (ICZN Art. 13.1). Type locality. Bermuda, Trunk Island, Harrington Sound; washings of black seaweeds.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFCFFDEFF51D5ECE699FA56.taxon	diagnosis	Differential diagnosis. Penicillicaris. Body length 500 – 700 μm in ♀, 400 – 500 μm in ♂. Antennule 8 - segmented in ♀. Antennary exopod and mouthparts unconfirmed. P 1 exp- 3 with two penicillate spines, one geniculate seta and one well developed non-geniculate seta. Relative length of P 1 enp- 1 unconfirmed. P 2 – P 3 endopods ♀ with enp- 2 and enp- 3 more often very imperfectly separated, without mobile articulation between them (when distinctly 3 - segmented no spur at inner distal corner); P 4 endopod ♂ with armature pattern [1.1.021] (both inner setae of ♀ not expressed in ♂). Armature pattern of ♀ P 2 – P 4: P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ with six elements on exopod; shape and length of exopod and endopodal lobe unconfirmed. P 5 exopod ♂ 1 - segmented, oval; with seven elements. P 5 endopodal lobe ♂ extending to insertion point of proximal inner seta of exopod; with two elements. Caudal ramus setae IV – V of ♀ inflated near base (forcipate type).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFFFCFFDEFF51D5ECE699FA56.taxon	etymology	Etymology. The species named is derived from the Latin penicillus, meaning paintbrush, and refers to the presence of penicillate elements on the antennary endopod and both rami of the first swimming leg. Notes. Willey (1935) inadvertently used two different spellings for his new form which he introduced as pencillata (p. 82) but was cited later in the paper on three occasions as penicillata. Unfortunately, some authors (e. g. Krishnaswamy 1957: 33) adopted the incorrect spelling pencillata. According to Willey (1935) the new species is unique in the morphology of the female P 2 – P 3 endopods which seem to lack a functional articulation between enp- 2 and enp- 3 in most specimens. His discovery of an additional hair-like seta on the distal exopodal segment of P 3 – P 4 means that some caution should be exercised when using previously reported armature patterns to identify other species since some may be incorrect. Penicillicaris penicillata sp. nov. is so far the only member of the genus that displays sexual dimorphism on the P 4 endopod with both inner setae of enp- 3 in the female not being expressed in the male, resulting in an armature pattern [1.1.021].	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF83FFDFFF51D6C6E04AFAE8.taxon	description	Original description. Sewell (1940): 193 – 195; Text-fig. 27 A – J (♀ only). Additional description. Vervoort (1964): 177, 181 – 184, 189; Figs 64 c, 66 c, 67, 68 d, 70 a – g.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF83FFDFFF51D6C6E04AFAE8.taxon	materials_examined	Type material. The female specimen illustrated by Sewell (1940: Text-fig. 27 A – J) is here designated as the holotype of P. sewelli sp. nov. (ICZN Arts 16.4 and 72.5.6). The species can be differentiated by the characters mentioned in the diagnosis below and those discussed and illustrated by Sewell (1940) (ICZN Art. 13.1). Type locality. Maldive archipelago, Addu Atoll; seaweed washings.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF83FFDFFF51D6C6E04AFAE8.taxon	diagnosis	Differential diagnosis. Penicillicaris. Body length 520 – 600 μm in ♀, 420 μm in ♂. Antennule 8 - segmented in ♀. Antenna with 2 - segmented exopod bearing two setae on exp- 1 and two lateral and three apical elements on exp- 2. Mandible with two setae on basis; exopod 1 - segmented, with two setae. Maxillipedal syncoxa unarmed; basis with one seta on palmar margin. P 1 exp- 3 with two penicillate spines, one geniculate seta and one short nongeniculate seta. P 1 enp- 1 about 1.15 times length of exopod. Armature pattern of ♀ P 2 – P 4: P 3 endopod ♂ 3 - segmented, with apophysis on enp- 3, armature pattern [1.1.02 + apo]. P 5 ♀ exopod about three times as long as maximum width; inner margin and proximal two-thirds of outer margin parallel; with six elements. P 5 endopodal lobe ♀ extending to about proximal third of exopodal length. P 5 exopod ♂ 1 - segmented, elongate – oval; with seven elements. P 5 endopodal lobe ♂ well developed, trapezoidal, extending slightly beyond insertion point of proximal inner seta of exopod; with two elements. Caudal ramus setae IV – V of ♀ inflated near base (forcipate type).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF83FFDFFF51D6C6E04AFAE8.taxon	etymology	Etymology. The species is dedicated to Robert Beresford Seymour Sewell, leader of the John Murray Expedition (1933 – 1934) to the Indian Ocean, during which it was collected. Notes. Sewell (1940: Text-fig. 27 G) illustrates an inner seta in the proximal quarter of P 1 exp- 2 but this, in addition to the alleged absence of the short seta on both exp- 3 and enp- 2 of this limb, must be based on observational errors. Vervoort (1964) misinterpreted the exopod as the endopod in both sexes in his text (pp. 182, 194) and illustration (Fig. 70 b); the inner seta on exp- 2 was almost certainly overlooked. According to Sewell (1940: Textfig. 27 H) P 2 exp- 1 lacks an inner seta but Vervoort shows a small inner element which is included in the armature formula here. The only significant difference between the Maldivian and Pacific populations appears to be the armature of the antennary exopod. According to Sewell (1940) the proximal segment has one minute and one long seta, and the distal segment has two lateral and three distal setae. Vervoort’s (1964: 70 g) illustration of the antenna appears to indicate that the proximal segment has two long setae and the distal segment only one lateral in addition to the three distal ones. It is conceivable that this discrepancy is the result of an incorrectly observed segment boundary, causing the proximal inner seta of exp- 2 to erroneously originate from exp- 1, combined with the oversight of the minute proximal element on exp- 1. In addition to the type locality in the Maldives, the species has also been recorded from the Caroline Islands (Federated States of Micronesia).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF82FFDCFF51D69CE1DCFF11.taxon	discussion	The swimming leg armature is incompletely known for all species (Table 1). Willey (1935: Fig. 123) noted that in Pe. penicillata sp. nov. the distal exopodal segment of P 3 – P 4 has three inner setae, the distal one of which is “ … an excessively fine hair curving outwards … ”. It is conceivable that this element has been overlooked in earlier descriptions and thus caution is required when using this key (see couplet 3). Any identification made must be checked against the original description (s).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF81FFDDFF51D217E520FB0F.taxon	description	Wells et al. (1982) placed Parastenhelia megarostrum in close proximity to Pa. anglica, Pa. ornatissima, Pa. reducta and Pa. hornelli, all of which share a 3 - segmented P 1 exopod with segments of subequal proportions. Based on the 9 - segmented antennule in the female, the position and length of the inner seta on P 1 enp- 1, the shape of the female P 5 and armature of the male P 5 exopod, they placed it closest to Pa. hornelli. Although the hornelli - complex was far from resolved at the time, Wells et al. (1982) considered distinct species status for Pa. megarostrum warranted because it displayed characters reflecting considerable conservatism in the genus, such as the massive size of the rostrum, the length / width ratio of P 1 enp- 2, the relative position of the setae on the female P 5 exopod, abdominal ornamentation, and relative proportions of the exopodal setae on the male P 5 exopod. They also attached considerable significance to the distinctive bulbous genital field in the female, which was regarded as an effective isolating factor in the mate recognition of this species. Song et al. (2003) considered Pa. pyriformis most closely related to Pa. hornelli and Pa. megarostrum based on the shared presence of subequal segments in the P 1 exopod, the position and length of the inner seta on P 1 enp- 1 and the 9 - segmented antennule in the female. They also recognized a certain resemblance with Pa. oligochaeta, based on the length and location of the inner seta of P 1 enp- 1, but the latter species differs in the ornamentation of the anal operculum, armature of ♀ P 2 – P 4, segmentation and armature of ♂ P 2 – P 3 endopods and the presence of six elements on the male P 5 exopod. Both Mielke (1990) and Gee (2006) have suggested removing Pa. megarostrum and / or Pa. pyriformis from Parastenhelia because they lack the characteristic sexual dimorphism on the male P 3 endopod that defines this genus. Neither species can be accommodated in Thalestrella because they do not exhibit the sexual dimorphism on the antenna that is typical for this genus nor can they be assigned to Foweya since the males lack the modified inner basal spine of P 1. Both genera are also characterized by an area of reduced cuticle thickness on the P 1 endopod which is not expressed in Pa. megarostrum or Pa. pyriformis. In accordance with earlier views (Mielke 1990; Gee 2006), both species are here transferred to a new genus, Porirualia gen. nov., based on the following suite of apomorphic character states: (a) rostrum very large, reaching to at least halfway the fourth antennulary segment in the female; (b) P 1 enp- 1 inner seta very long, extending beyond distal margin of segment; and (c) P 5 ♀ with series of transverse striae along inner margin of endopodal lobe; these striae are internal chitinous reinforcements and are more strongly developed in Po. megarostrum (Wells, Hicks & Coull, 1982) comb. nov. than in Po. pyriformis (Song, Kim & Chang, 2003) comb. nov. The new genus shares a sistergroup relationship with Johnwellsia gen. nov. based on the following synapomorphies: (a) distal seta on the mandibular basis modified into short spine; (b) P 3 endopod ♂ with only one inner seta on distal segment (homologue of proximal inner seta of ♀ enp- 3 not expressed in ♂; Fig. 7 G – I); and (c) outermost element of P 5 endopodal lobe reduced, represented by minute seta in both sexes. There is controversy over the segmentation of the male antennule. Both Wells et al. (1982) and Song et al. (2003) describe it as 9 - segmented although the latter authors figure ten segments. The antennule is here reinterpreted as 10 - segmented with the geniculation positioned between segments 6 and 7. The small bisetose segment 4 (= ancestral segment XIII; figured but not counted by Song et al. 2003) was overlooked by Wells et al. (1982). The apical segment in Po. megarostrum comb. nov. was described as unarmed, displaying a row of fine marginal hairs; it is unlikely that this observation is correct.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF81FFDDFF51D217E520FB0F.taxon	diagnosis	Diagnosis. Parastenheliidae. Sexual dimorphism in antennule, P 2 – P 3 endopods, P 5 – P 6 and urosomal segmentation. Body fusiform; posterior margin of cephalothorax and all somites (except anal somite) with deeply divided, denticulodigitate, hyaline frills. Rostrum defined at base, very large, reaching to at least halfway fourth segment of antennule; with parallel sides tapering sharply into rounded apex. Anal operculum semicircular, bordered with fine setules. Caudal ramus wider than long, without conspicuous spinular ornamentation; with six setae, seta V slightly inflated at base in ♀. Antennule short and 9 - segmented in ♀, segment 1 not elongate, segments 7 – 8 shortest, with aesthetascs on segments 4 and 9; haplocer and 10 - segmented in ♂, with geniculation between segments 6 and 7, segment 5 swollen and with aesthetasc; segmental homologies in ♂: I, II – VIII, IX – XII, XIII, XIV – XVIII, XIX – XX, XXI – XXII, XXIII, XXIV – XXV, XXVI – XXVIII. Antenna not sexually dimorphic; allobasis completely fused or partially divided, with endopodal pinnate seta on abexopodal margin; exopod 2 - segmented, proximal segment with two setae, distal segment with two lateral and 2 – 3 apical elements; free endopod without penicillate elements. Mandible with 3 – 4 elements on basis, distal one enlarged and spiniform; endopod with two lateral and three apical setae; exopod 1 - segmented, with three setae. Maxillulary coxal epipodite represented by one seta. Maxilla with three endites on syncoxa; endopod discrete, with 2 – 3 setae. Maxilliped with two setae on syncoxa; basis with one seta and longitudinal spinular row on palmar margin; endopod represented by claw with one accessory seta. P 1 inner basal spine not sexually dimorphic. P 1 exopod 3 - segmented, segments subequal in length; exp- 2 with inner seta; exp- 3 with two pinnate spines, one geniculate seta and one naked seta. P 1 endopod 2 - segmented; enp- 1 longer than exopod, with long, proximally inserted, pinnate inner seta extending almost to distal margin of enp- 2, inner segment margin without area of reduced chitinization; enp- 2 with one naked minute seta and two pinnate claws. P 2 – P 4 rami 3 - segmented; inner seta of P 2 – P 4 exp- 1 reduced in size. P 2 – P 3 endopods ♂ 2 - or 3 - segmented. Apical setae of P 2 enp- 3 ♂ occasionally shorter than in ♀. P 3 endopod ♂ with only one inner seta (vs two in ♀) on enp- 3, resulting in 1.1.121 or 1.221 (when ♂ endopod 2 - segmented) pattern; without spinous apophysis. Armature formula of P 2 – P 4 as follows: P 5 ♀ endopodal lobe with five setae, outermost minute; inner margin with series of transverse striae. P 5 ♀ exopod elongate, with six elements. P 5 ♂ endopodal lobe with two elements, outer one minute; exopod 1 - segmented, with 5 – 6 elements. Vestigial P 6 ♀ represented by 1 – 2 minute seta (e). P 6 ♂ with three setae.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF81FFDDFF51D217E520FB0F.taxon	etymology	Etymology. The generic name alludes to the type locality of its type species, Pauatahanui Inlet in Porirua Harbour, Wellington. Gender: feminine.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF81FFDDFF51D217E520FB0F.taxon	type_taxon	Type species. Parastenhelia megarostrum Wells, Hicks & Coull, 1982 (by original designation herein) = Porirualia megarostrum (Wells, Hicks & Coull, 1982) comb. nov. Other species. Parastenhelia pyriformis Song, Kim & Chang, 2003 = Po. pyriformis (Song, Kim & Chang, 2003) comb. nov.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF80FFDAFF51D602E723FD16.taxon	materials_examined	Type locality. New Zealand, North Island, Wellington; Porirua Harbour, Pauatahanui Inlet (since renamed to Te Awarua-o-Porirua Harbour); intertidal beach some 200 m north-east of the State Highway 1 bridge; fine sand with some silt.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF80FFDAFF51D602E723FD16.taxon	diagnosis	Differential diagnosis. This coincides with Wells et al. ’ s (1982) excellent description of Parastenhelia megarostrum. See below for differences with its only congener, Po. pyriformis comb. nov. Notes. Wells et al. (1982) remarked that, despite discrepancies in his illustrations, based on locality and collection details, it appears that Thomson’s (1883) Thalestris forficula is conspecific with Po. megarostrum comb. nov. This claim is considered highly unlikely for a variety of reasons: (a) Thomson’s female specimens are significantly larger (1 mm vs 665 μm); (b) there is no mention of the large rostrum which is one of the most conspicuous characters of Po. megarostrum comb. nov.; (c) the antennule is more slender; (d) P 1 exp- 2 is elongate, about 2.5 times the length of exp- 1 (vs subequal to exp- 1); and (e) P 1 enp- 1 is comparatively longer and its inner seta shorter. Thomson (1883) also stated that the central caudal setae (seta V) are “ … swollen just beyond their basal articulation, and marked along the greater part of their length with annular articulations ” while in Po. megarostrum comb. nov. all caudal ramus setae are unmodified. As stated above it is conceivable that T. forficula sensu Thomson (1883) is conspecific with Microthalestris antarctica comb. nov. Porirualia megarostrum comb. nov. is so far endemic to New Zealand where it is found on fine silty intertidal sandbanks in practically all of the country’s major estuarine and harbour systems (Wells et al. 1982; Wells & Hicks in Dahms & Hicks 1996). Additional records from Porirua Harbour, including Pauatahanui Inlet, have been added by Hicks (1984, 1985, 1986, 1988, 1989, 1991, 1992), Iwasaki et al. (1993, 1999) and Blaschke et al. (2010). On the North Island it has been recorded at localities extending northward from Wellington to Kaipara Harbour on the west coast and to Doubtless Bay on the east, occurring sympatrically with Parastenhelia hornelli in Whangateau Harbour (Wells et al. 1982). The species has also been reported from the South Island, including Portobello Harbour (Otago Harbour) (Stringer et al. 2012 a), Papanui Inlet (Otago Peninsula) (Leduc & Probert 2011; Leduc et al. 2009) and Okains Bay, Canterbury (Stringer et al. 2012 b). Dahms & Hicks (1996) described all six nauplius stages of Po. megarostrum comb. nov. while Dahms (1993) provided details on the copepodid development of this species. Field experiments employing emergence traps indicate that adults do not swim or have only poorly developed swimming ability (Hicks 1986) while nauplii and copepodids are passively mobile, being transported by shifting sandgrains they cling on (Hicks 1988) or resuspended by tidal bottom shear currents (Hicks 1992). Nauplii have never been observed actively swimming (Dahms & Hicks 1996). Porirualia megarostrum comb. nov. is the most abundant epibenthic metazoan on Mana Bank in Pauatahanui Inlet and the dominant food source for young flatfish (Hicks 1984). Study of flat fish predation on copepods at this site indicated that although predation was high, there was little impact on overall abundance of the copepod (Hicks 1985).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF87FFDAFF51D006E0BBFA67.taxon	description	Original description. Song et al. (2003): 221 – 227; Figs 1 – 4; Table 1.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF87FFDAFF51D006E0BBFA67.taxon	materials_examined	Type locality. South Korea, Jeju Island, Sehwa Beach (33 ° 28 ’ 07 ” N, 126 ° 54 ’ 43 ” E); shallow sublittoral zone.	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
951887EAFF87FFDAFF51D006E0BBFA67.taxon	diagnosis	Differential diagnosis. This coincides with Song et al. ’ s (2003) description of Parastenhelia pyriformis. Notes. Song et al. (2003) added a second record from Seungbong Island in the Yellow Sea where it was found to be associated with macroalgae (Zostera japonica) on sandy bottoms (see also Song et al. 2012). The two species can be distinguished, in both sexes, by (a) rostrum size (reaching to distal margin of segment 5 of the antennule in Po. megarostrum vs halfway segment 4 in Po. pyriformis), (b) antennary morphology (allobasis partially divided by surface suture and exp- 2 with four setae / spines vs allobasis without trace of original segmentation and exp- 2 with five elements, i. e. with additional apical seta), (c) armature of mandibular basis (with one spine and three setae vs one spine and two setae), (d) length: width ratio of P 1 enp- 2 (2.2 vs 1.3), and (e) armature of P 3 – P 4 exp- 3 (distal inner seta present and short vs absent). In addition, females can be differentiated by the proportional segment lengths in the antennule (segment 1 longest in Po. megarostrum vs segment 2 longest and entire appendage more slender in Po. pyriformis) and P 5 morphology (L: maximum W ratio 2.2 and distal outer seta longest in Po. megarostrum vs 3.25 and inner distal seta longest in Po. pyriformis). Males differ in the segmentation of P 2 – P 3 endopods (3 - segmented vs 2 - segmented) and the number of elements on the P 5 exopod (six vs five).	en	Huys, Rony, Mu, Fanghong (2021): Johnwellsia, a new intertidal genus of Parastenheliidae (Copepoda, Harpacticoida) from the Taiwan Strait, China, including a review of the family and key to genera. Zootaxa 5051 (1): 236-318, DOI: 10.11646/zootaxa.5051.1.13, URL: https://doi.org/10.11646/zootaxa.5051.1.13
