taxonID	type	description	language	source
E35187DFFFC80073EDA6FC176A89FBC8.taxon	materials_examined	Material examined (Table 2): Type specimens deposit- ed in the collection of the National Institute of Water and Atmospheric Research, New Zealand. Holotype: female, Stn 59, top net, NIWA 86594 (three slides). Paratypes: one male, Stn 59, top net, NIWA 86595 (three slides). Paratype lots in 4 % formalin: one female, Stn 59, top net, NIWA 86596 (one vial, one slide); one female, Stn 59, bottom net, NIWA 86597 (one vial, one slide); two females, Stn 121, top net, NIWA 86600 (one vial); one female, one male, Stn 121, bottom net, NIWA 86599 (one vial); one female, one male, Stn 59, bottom net, NIWA 86598 in 95 % ethanol (one vial); one female, Stn 59, top net, NIWA 85993, NHMUK 2013.25. Etymology: The generic name Pinkertonius (gender masculine) is derived from the name of Dr Matt Pinkerton who is project leader for the NIWA research programme ‘ Marine Food Web Dynamics’ on the Chatham Rise, where these specimens were found. The specific name ambiguus derives from the Latin, referring to the ambiguous nature of the morphology of this species in not exactly fitting into any of the previously described families. Genetic description The genetic description, from one specimen, is based on the nuclear large (28 S) and small (18 S) subunits ribosomal RNA and the mitochondrial genes for the proteins cytochrome c oxidase subunit I (COI) and cytochrome b (cyt b). These sequences were deposited in GenBank accession nos: KF 753813 – KF 753816, respectively, which correspond to the catalogued specimen, Co. 449.1.1, held in the Marine Science Department, University of Connecticut. Morphological description Female: Total length 1.8 – 2.0 mm, urosome 28 % of total length (Fig. 2 A, B). Prosome ovoid with head (cephalosome) and pedigerous somite 1 separate; pedigerous somites 4 and 5 separate, posterolateral corners of pedigerous somites 1 – 4 extend into small triangular projections, pedigerous somite 5 produced into pointed lappets extending more than halfway along genital double somite. Rostrum in form of ventrally directed rounded plate with pair of distal filaments (Fig. 2 D). Urosome of four free somites, first three bordered posteriorly by unevenly serrated hyaline fringe, –, samples not examined. Measurements taken along posterior border of each segment but two (posterior (shortest) and anterior) measurements taken of ancestral segment I. largely intact ventrally. Genital double somite with slight anterior swelling in dorsal view; in lateral view swollen ventroanteriorly; in ventral view genital field asymmetrical, gonopores slightly unequally developed, being larger on left, genital operculum skewed to left with hinge aligned at about 45 ° to anterior – posterior axis of somite, so that right gonopore not completely covered, right side of genital field bordered by ridge aligned anterior – posteriorly, no such flange on left, copulatory pore and seminal receptacle not obvious, although sac on left apparently linked to left gonopore (Fig. 2 E). Caudal rami slightly asymmetrical, longer on right, with seven setae each (Fig. 2 F, G) more or less symmetrically arranged on each side. Seta I vestigial, seta II spiniform, seta V longest, about 1 mm long, seta IV next longest followed by setae VI and III, seta VII small and spiniform and inserted on dorsoinner distal corner, left caudal ramus inner border lined with fine setules; on right ramus row of long setules arranged obliquely on anterior part of ventral surface. Antennule (Table 3): Twenty-seven segmented, extending to posterior border of pedigerous somite 5 (Fig. 2 C; Fig. 3 A, B, C). One aesthetasc present on each of segments I, III, VII, XI, XIV, XVI, XVIII, XXI, XXV, and XXVIII; small cuticular thickenings found on segments II – X, XII, XIII, XV, and XVII; one seta on eight proximal segments, wider than other setae, and attenuated distally into curved narrow tip. Setal formula: I, 1 s, 1 a; II, 2 s; III, 2 s, 1 a; IV, 2 s; V, 2 s; VI, 2 s; VII, 2 s, 1 a; VIII – X, 2 s; XI, 2 s, 1 a; XII to XIII, 2 s; XIV, 2 s, 1 a; XV, 2 s; XVI, 2 s, 1 a; XVII, 2 s; XVIII, 2 s, 1 a; XIX to XX, 2 s; XXI, 2 s, 1 a; XXII to XXIII, 1 s; XXIV, 1 + 1 s; XXV, 1 + 1 s, 1 a; XXVI, 1 + 1 s; XXVII – XXVIII, 5 s, 1 a. Antenna: Coxa and basis separate, coxa with one seta, basis with two setae (Fig. 4 A). Endopod two-segmented, with traces of fusion between segments 2 and 3, and between segments 3 and 4; segment 1 with two inner setae, segment 2 with nine plus seven terminal setae and outer transverse row of spinules marking boundary between putative endopod segments 3 and 4. Exopod shorter than endopod, eight-segmented, ancestral segments VIII and IX fused, segments I – VII each with one well-developed seta, compound distal segment VIII – IX with three terminal setae and one inner subterminal seta, outermost seta on terminal endopod segment lined proximally with small spinules. Mandible: Gnathobase with seven marginal teeth, ventralmost largest, five dorsal teeth bicuspid, small spinulated seta inserted dorsally (Fig. 4 B). Basis with four apparently naked setae; endopod two-segmented, segment 1 with inner lobe and four setae, segment 2 with ten terminal setae, distoinner border with short row of spinules, transverse row of spinules at about midlength. Exopod five-segmented, with 1, 1, 1, 1, 2 setae. Maxillule: Praecoxal arthrite with 14 spines and setae, including four on posterior surface and one on dorsal surface (Fig. 4 C); coxal endite with four setae; basal endites 1 and 2 with four and five setae, respectively; endopod segments 1 and 2 fused, segments 2 and 3 separate, with four, three, and seven setae, respectively; exopod with 11 setae, of which three terminal setae short and bordered by fine setules along inner border; basal exite without seta; coxal epipodite with nine setae, of which three proximal setae short. Maxilla: Praecoxa, coxa, and basis clearly separated, endites 1 – 4 with seven (one very short), three, three, and three setae, respectively (Fig. 3 D, E); basal endite with three setae, one of them stout and spiniform; inner setae on endites 2 – 5 lined with long spinules; endopod segment 1 endite with three setae, segments 2 – 4 with two, two, and three setae, respectively. Maxilliped: First syncoxal endite with one seta (Fig. 4 D); endites 2 and 3 with two and four spinulose setae, respectively, crescent-shaped row of fine spinules at base of endite 3 on inner surface; endite 4 with three setae * Illustrated specimen with extra seta, absent in another specimen. and small peg-like structure, and a few small spinules. Basis with two setulose setae and proximal border lined by spinules. Endopod well-developed, longer than basis, endopod segment 1 apparently separate from basis, endopod segments 1 – 6 with two, four, four, three, three plus one, and four spinulose setae, respectively, outer seta of segment 6 wider and longer than adjacent seta, and terminally inserted. Swimming legs (Table 4): Legs 1 – 5 biramous, all rami three-segmented (Fig. 5). Leg 1 basis with distal lobe on posterior surface situated between exopod and endopod (this lobe is visible in lateral view in the whole animal, as it projects posteriorly); ornamented with transverse row of inner setules; inner distal seta slightly curved. Coxae of legs 2 – 5 decorated with patches of long spinules on posterior surface; basis of legs 2 – 4 with blunt tooth on anterior surface between endopod and exopod; outer seta on basis of leg 3 spiniform; outer proximal border of exopod segments 2 and 3 of legs 2 – 4 each with small knob-like projection; exopods of legs 2 – 5 with heavily built outer spines bordered by very small blunt denticles, terminal exopod spines also heavily built, and with outer edge lined by tiny small blunt teeth. Surfaces of both rami of legs 1 – 5 ornamented with patches of very small spinules, most dense on posterior surfaces. Leg 5 articulation of exopod segment 3 with segment 2, oblique and reminiscent of species of Ridgewayia (Fig. 1 I, H); proximal end of segment 3 narrowing, distance between two pivot points (Fig. 2 H) ensures region of articulation is as wide as exopod segment 3 at level of proximal inner seta of exopod segment 3. Outer distal extension of exopod segment 2 not reaching origin of proximal outer spine of exopod segment 3. Male: Total length 1.80 – 1.88 mm, urosome 28 % of total length (Fig. 6 A, B). Prosome ovoid with head and pedigerous somite 1 separate; pedigerous somites 4 and 5 separate, posterolateral corners pedigerous somites 1 – 4 extended into small triangular projections, pedigerous somite 5 extending into pointed lappets reaching beyond posterior border of genital somite. Rostrum in form of ventrally-directed rounded plate, with pair of distal filaments. Urosome of five free somites, anterior four somites bordered posteriorly by unevenly serrated hyaline fringe. Caudal rami symmetrical, with seven setae each: seta I vestigial, seta II spiniform, seta V longest, seta IV next longest followed by setae VI and III, seta VII small and spiniform and inserted on dorsoinner distal corner, inner borders of caudal rami lined with fine setules (Fig. 6 D). Antennule: Twenty-six segmented on left, 25 - segmented on right, extending just beyond posterior border of fourth pedigerous somite; right antennule geniculate between ancestral segments XX and XXI, segments XV – XIX enlarged (Fig. 7). One seta on proximal segment on each side wider than other setae and attenuated distally into curved narrow tip. Left antennule with following setation: I, 1 s, 1 a; II – III, 3 s, 5 a (two on segment II, three on segment III); IV, 2 s, 1 a; V, 2 s, 2 a; VI, 2 s, 1 a; VII, 2 s, 2 a; VIII to XVIII, 2 s, 1 a; XIX to XX, 2 s; XXI, 2 s, 1 a; XXII to XXIII, 1 s; XXIV, 1 + 1 s; XXV, 1 + 1 s, 1 a; XXVI, 1 + 1 s; XXVII – XXVIII, 5 s, 1 a. Right antennule (Fig. 8) with following setation: I, 1 s, 1 a; II – IV, 6 s, 5 a (one on segment II, three on segment III, one on segment IV); V, 2 s, 2 a; VI, 2 s, 1 a; VII, 2 s, 2 a; VIII to XVIII, 2 s, 1 a; XIX to XX, 2 s (on segment XIX distal seta fused spatulate distal extension, segment XX with anterior proximal ridge, proximal seta modified as fused spine-like element and one seta); XXI – XXIII, two fused spine-like elements, 1 a, 2 s; XXIV – XXV, 2 + 2 s (without fused distal process; see Appendix S 5), 1 a; XXVI, 1 + 1 s; XXVII – XXVIII, 5 s, 1 a. Antenna, mandible, maxillule, maxilla, maxilliped, and swimming legs 1 – 4 identical to those of female. Leg 5: Leg 5 biramous on both sides, with both rami three-segmented, asymmetrical (Fig. 6 E); exopod slightly longer on right. Right exopod segment 2 with triangular inner attenuation, segment 3 in form of claw with terminal spine, rounded distally, fused to segment, with two articulated spines (one medioproximal, other on outer border more distally inserted), with additional outer fused spinule and pore opening. Left leg exopod segment 2 with inner border swollen, bearing seta modified into scalpel-shaped element thickened along its outer border; exopod segment 3 simple, about twice as long as wide, short terminal spine fused to segment at level adjacent to lateral pore opening, one articu- lated inner spine and outer articulated spine more proximally inserted on posterior surface. Posterior surfaces of exopods and endopods ornamented with scattered patches of very small spinules. Ecological notes: The samples that contained P. ambiguus sp. nov. came from the hyperbenthic zone above sediments with high pelagic calcium carbonate content on the flanks of the Chatham Rise, but were absent from shallower regions on the Mernoo Saddle or on the scoured slopes of the rise (Carter, Neil & McCave, 2000). The Chatham Rise traps the Subtropical Front east of New Zealand, and is the location of year-round elevated primary productivity (Bradford-Grieve et al., 1997). Remarks: Pinkertonius ambiguus gen. et sp. nov. retains a number of plesiomorphic features: the female genital double somite has a genital operculum; the caudal rami have a vestigial seta I; the female antennule has all of ancestral segments I – XXVII separated; all swimming legs have both rami on each side three-segmented, male leg 5 of relatively simple construction; the proximal seven segments of the antenna exopod are separate; the endopod of the mandibular palp is two-segmented, welldeveloped, and bears ten terminal setae on segment 2; the maxillule has nine setae on the coxal epipodite, four setae on the posterior surface of the praecoxal arthrite, and 11 exopod setae; on the maxilla, endopod segment 1 and its endite are separated from the basis; the maxilliped endopod segment 1 is separate from the basis, and endopod segment 5 has an outer seta. Superficially, P. ambiguus sp. nov. resembles Miheptneria abyssalis, especially in general body shape, but differs from this species and all other genera currently in the Epacteriscidae, Pseudocyclopidae, Ridgewayiidae, and Boholinidae in having several autapomorphies: in the female the longer caudal ramus is on the right and there is an oblique row of long setules on the ventral surface, which we consider to be homologous with the inner row of setules on the left ramus; the segmental distribution of aesthetascs differs greatly between the sexes (female ancestral segments II, IV – VI, VIII – X, XII – XIII, XV, and XVII are without aesthetascs, whereas the male has aesthetascs on all of segments I to XVIII). Pinkertonius ambiguus gen. et sp. nov. is unique among the above group of families in having the apomorphic condition of multiple aesthetascs on some segments in the male (three on ancestral segment III and two each on segments V and VII on both sides) and one aesthetasc on right segment II and two aesthetascs on left segment II. The proximal outer borders of exopod segments 2 and 3 of legs 2 – 4 each have a knob-like projection. The following analyses allow us to place P. ambiguus sp. nov. in a systematic hierarchy.	en	Bradford-Grieve, Janet M., Boxshall, Geoffrey A., Blanco-Bercial, Leocadio (2014): Revision of basal calanoid copepod families, with a description of a new species and genus of Pseudocyclopidae. Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 171 (3): 507-533, DOI: 10.1111/zoj.12141, URL: http://dx.doi.org/10.1111/zoj.12141
E35187DFFFD1006FEEC3FEC86D2AFC53.taxon	diagnosis	Diagnosis: Plesiomorphic calanoid copepods with underlying pattern of full development of arthrodial membranes between body somites and limb segments, with some exceptions: antennule of female up to 27 - segmented, ancestral segments XIX, XX, and XXIII usually without asthetascs; male antennule always geniculate on right and tendency for ancestral segment XXV to have distoanterior process. Male and female mouthparts identical. Antenna exopod nine-segmented, segments I – VIII with one seta each. Maxilliped endopod segment V nearly always with outer border seta (except Edaxiella); swimming legs 1 – 5 with both rami usually three-segmented; legs 1 and 2 exopod segment 3 with two or three outer border spines, leg 3 exopod segment 3 with two or three outer border spines, legs 4 and 5 (female) with two or three outer border spines.	en	Bradford-Grieve, Janet M., Boxshall, Geoffrey A., Blanco-Bercial, Leocadio (2014): Revision of basal calanoid copepod families, with a description of a new species and genus of Pseudocyclopidae. Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 171 (3): 507-533, DOI: 10.1111/zoj.12141, URL: http://dx.doi.org/10.1111/zoj.12141
E35187DFFFD10068EE00FC5A6CD7FA5C.taxon	diagnosis	Diagnosis: Female caudal rami of equal lengths (except Pinkertonius gen. nov., longer on right); male caudal rami usually symmetrical (except Stargatia); seta II spiniform. Female antennule ancestral segment IV without aesthetasc. Tendency for aesthetascs to be absent from additional segments. Mandible endopod well-developed, two-segmented, ranging from greater than half length of exopod to extending well beyond exopod (except Exumella), segment 1 usually with four setae (exceptions Robpalmeria, Normancavia, Exumella, and Stargatia), segment 2 usually with between eight and 11 setae, except for Normancavia, Exumella, and Exumellina, which have six, six, and seven setae, respectively. Maxillule coxal epipodite always with nine setae. Maxilla basis usually less than twice estimat- ed length of coxa (except Exumella), basal endite usually elongate (except in Placocalanus), and endopod setae normal (exception in Boholina, which has spine-like setae). Maxilliped endopod segments 2 – 6 usually longer than length of coxa (not including praecoxa) (exception Placocalanus and Pseudocyclops) and endopod setae normal. Swimming legs 1 – 5 with both rami threesegmented. Leg 1 basis posterior surface with tendency to have posterior surface process; exopod segment 2 with tendency to have outer distal corner produced into spinous lobe inner to articulated spine; exopod segment 3 with variable number of inner setae. Female leg 5 has tendency towards specialization of joint between exopod segments 2 and 3 composed of lengthening of outer distal part of segment 2, forming an oblique distal margin, and narrowing of proximal articulating region of segment 3; endopod reduced to two segments in Boholina and reduced or absent in Ridgewayia. Male left leg 5 with inner process present on exopod segment 2; tendency towards reduction in arthrodial membrane formation in endopods and increasing complexity in form of exopods (Table 6). Type genus: Pseudocyclops Brady, 1872. Remarks: The genera included in this family are: Badijella, Boholina, Brattstromia, Exumella, Exumellina, Hondurella, Normancavia, Pinkertonius gen. nov., Placocalanus, Pseudocyclops, Ridgewayia, Robpalmeria, Stargatia, and Stygoridgewayia.	en	Bradford-Grieve, Janet M., Boxshall, Geoffrey A., Blanco-Bercial, Leocadio (2014): Revision of basal calanoid copepod families, with a description of a new species and genus of Pseudocyclopidae. Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 171 (3): 507-533, DOI: 10.1111/zoj.12141, URL: http://dx.doi.org/10.1111/zoj.12141
E35187DFFFD60069EEFFFA406A44F941.taxon	diagnosis	Diagnosis: As for Pseudocyclopidae, except female caudal ramus longer on right. Mandible endopod longer than exopod. Leg 1 basis without outer edge seta, mediodistal seta present and posterior surface process present; exopod segment 2 without spinous lobe; endopod segment 3 with three inner setae. Leg 3 outer distal corner of basis with one spine-like seta; exopod segment 1 with one inner seta; segment 3 with three outer spines. Leg 4 exopod segment 1 with one inner seta; segment 3 with four inner setae. Female leg 5 endopod formula: 0 – 1; 0 – 1; 2, 2, 2. Exopod segment 2 extended distolaterally. Male leg 5 formula similar to that of female, except for exopods: I- 0; I- 1; I, 0, I (left) and I- 0; I- 0; I, 0, I (right); left and right exopod segment 2 with inner processes. Type species: Pinkertonius ambiguus gen. et sp. nov. by original designation. Remarks: The most distinctive shared characteristics that link this genus to the family Pseudocyclopidae are: the absence of an aesthetasc on ancestral antennular segment IV; the presence of a well-developed, elongate two-segmented mandibular endopod with ten terminal setae; the presence of nine setae on the coxal epipodite of the maxillule; the presence of a posterior surface process on the basis of leg 1; exopod segment 2 of female leg 5 is distally extended, and the articulation between segments 2 and 3 is at an oblique angle to the main axis of the limb; and the right exopod segment 2 of male leg 5 has a triangular inner process and left exopod segment 2 has a scalpel-like inner projection that is directed distally.	en	Bradford-Grieve, Janet M., Boxshall, Geoffrey A., Blanco-Bercial, Leocadio (2014): Revision of basal calanoid copepod families, with a description of a new species and genus of Pseudocyclopidae. Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 171 (3): 507-533, DOI: 10.1111/zoj.12141, URL: http://dx.doi.org/10.1111/zoj.12141
E35187DFFFD7006AED81F9576A88FE6F.taxon	diagnosis	Diagnosis: Female caudal rami usually of equal lengths (exceptions: Balinella longer on right; Gloinella longer on right); male caudal rami symmetrical or asymmet- rical, seta II spiniform, setiform, or apparently absent in Miheptneria, female seta VI tending to be asymmetrical, with one seta having one side bearing short setules, the other with long setules (exceptions Balinella, Cryptonectes, Oinella, Bunderia, Edaxiella, and Enantronoides). Female antennule ancestral segment IV with aesthetasc (except for Oinella). Mandible with exopod and basis forming major axis of palp; endopod poorly developed, two- or one-segmented, or absent, being at most half length of exopod segment 1, which when present, with at most two setae, segment 2 with four or fewer setae, except for Erebonectes and Miheptneria, which have seven and nine setae, respectively. Maxillule coxal epipodite with seta 9 absent (exception Miheptneria), and tendency for further setae to be lost up to seta 6. Maxilla basis usually more than twice estimated length of coxa, basal endite with low profile, and endopod setae spine-like (exception Miheptneria, which has elongate basal endite and normal setae). Maxilliped endopod segments 2 – 6 shorter than length of coxa (not including praecoxa) and endopod setae spine-like (exceptions Miheptneria, Bomburiella, and Edaxiella, which appear to have normal setae on endopod). Leg 3 endopod segment 1 with outer distal corner bifid or trifid in clade 20. Male left leg 5 without inner process present on exopod segment 2 (only exception Balinella) (Table 7). Type genus: Epacteriscus Fosshagen, 1973. Remarks: The genera included in this family are: Balinella, Bofuriella, Bomburiella, Bunderia, Cryptonectes, Edaxiella, Enantiosis, Enantronia, Enantronoides, Epacteriscus, Erebonectes, Gloinella, Iboyella, Miheptneria, Minnonectes, and Oinella.	en	Bradford-Grieve, Janet M., Boxshall, Geoffrey A., Blanco-Bercial, Leocadio (2014): Revision of basal calanoid copepod families, with a description of a new species and genus of Pseudocyclopidae. Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 171 (3): 507-533, DOI: 10.1111/zoj.12141, URL: http://dx.doi.org/10.1111/zoj.12141
E35187DFFFD40064EDCCFE526948FA1C.taxon	description	Genetic data and a revised phylogeny confirm the basal position of a newly defined superfamily Pseudocyclopoidea (which now includes families previously assigned to the synonymized Epacteriscoidea) within the Calanoida, as well as improving the organization and resolution of the relationships among superfamilies. The revised molecular phylogeny (Fig. 12) reinforces the results obtained in the previous genebased study (Blanco-Bercial et al., 2011), although the lack of members of the basal families, the epibenthic Ridgewayiidae, Boholinidae, Pseudocyclopidae, and Epacteriscidae in the previous analysis was problematic. With the inclusion of sequences from the superfamily Pseudocyclopoidea, the reconstructed phylogeny agrees partially with the topology of early morphologybased phylogenies (Andronov, 1974; Park, 1986). Here superfamilies Pseudocyclopoidea, Augaptiloidea, and Centropagoidea sequentially split off from a main stem. The remaining superfamilies form a single clade, the topology of which is similar to the topology described in Bradford-Grieve et al. (2010) based on morphological data. The lower support than that found by Blanco-Bercial et al. (2011), evident in many parts of the reconstruct- ed phylogeny, could result from incomplete gene coverage of the new taxa added (Appendix S 1). It is likely that the addition of COI and cyt b to the clades where they are missing would add stronger support to the analyses. For example, the inclusion of mitochondrial genes can improve resolution at deeper nodes (Fisher-Reid & Wiens, 2011; Cornils & Blanco-Bercial, 2013). We found that without the addition of mitochondrial gene sequences from P. ambiguus sp. nov., not only was the phylogeny not recovered, as it is here, but the resulting superfamilies were also polyphyletic or paraphyletic in some cases (data not shown). All superfamilies were recovered as monophyletic, even in cases where very divergent groups were included, supporting the conclusions drawn from morphological characters (Ho, 1990; Huys & Boxshall, 1991). This fact is very significant in Centropagoidea, where the two divergent families Diaptomidae (the only entirely freshwater family) and Pseudodiaptomidae clustered with the other Centropagoidea in a single clade. The resolution of monophyletic clades, representing the superfamilies Eucalanoidea, Megacalanoidea, and Bathypontioidea, sister to the Clausocalanoidea and Spinocalanoidea, does not agree with morphological studies (Andronov, 1974; Park, 1986; Bradford-Grieve et al., 2010). Although interesting, this result should be considered with caution, because the family identified as intermediate between these two clades (Ryocalanidae; superfamily Ryocalanoidea) is missing from the molecular analysis, and its addition might result in changes to the present topology of the phylogeny. Thus, the presented revised molecular phylogeny provides testable hypotheses for future work. PHYLOGENY OF PSEUDOCYCLOPOIDEA Among the pseudocylopoidean genera, the morphologybased phylogenetic signal was possibly obscured by the high degree of homoplasy and may have interfered with accurate tree inference. Some of the homoplasy in our data may be the result of the possibility that we are not always dealing with homologous characters or that the character states are not accurately recorded in the literature. It is also possible that some characters reflect lifestyle rather than preserving a phylogenetic signal. In this analysis, it was not possible to determine homologies in some characters relating to the mandible upon which feeding niche strongly impacts (Itoh, 1970). We note, however, that certain types of modification of the mandible, maxilla, and maxilliped are strongly linked to the two major clades (Pseudocyclopidae and Epacteriscidae), and so may contain a robust phylogenetic signal. In the Pseudocyclopidae, the mandible generally has small teeth (although Exumellina and Stargatia have two elongate ventral teeth) and the endopod is well developed, often with four setae on segment 1 and more than nine setae on segment 2. The maxilla has a normally developed basis with an elongate endite and normal endopod setae. The maxilliped usually has an elongate endopod that is furnished with normal setae. We note that this family contains genera that live in open water habitats and the only freshwater groundwater genus, as well as marine cave-dwellers. It is deduced from the form of the mouthparts that many of these genera are fine-particle feeders. Those that deviate in having reduced mandibular endopod setation appear to have other modes of feeding. For example, Exumella seems to be a benthic scavenger (Jaume & Boxshall, 1995), and Exumellina and Stargatia, which also have paddle-like modification to the mandible and maxillule endopods, may be raptorial feeders feeding on delicate prey (Fosshagen & Iliffe, 1998). A large subset of these genera also have female leg 5 modified so that the mode of articulation of exopod segment 3 directs this segment into the midline, or ensures that it has an even greater arc of movement (Badijella, Pseudocyclops, Ridgewayia, Robpalmeria, Normancavia, Brattstromia, Exumella, Exuminella, Pinkertonius gen. nov., and possibly Hondurella, Placocalanus, and Stargatia). It is tempting to hypothesize that these genera are adapted to digging in sediment. In the Epacteriscidae, the mandible generally has the ventral tooth enlarged and separated by a much larger gap than exists between the remaining teeth; an even more specialized gnathobase is found in Epacteriscus, where the cutting blade has a prominent extension bearing sharp teeth that extends well out from the body (Fosshagen, 1973). Most species have a reduced endopod that is either one-segmented with only one seta or is absent (except for Balinella, Bofuriella, Erebonectes, and Miheptneria, which have two-segmented endopods). The endopods of the maxillae and maxillipeds are very condensed and nearly always have spine-like setae, with a few exceptions (Miheptneria, Bomburiella, and Edaxiella), and the maxilla basis is enlarged and has a very low-profile endite. Taken together, these character states indicate that most genera are carnivores. The basal taxa in each family (Pinkertonius gen. nov. and Miheptneria) appear to be adapted to fineparticle feeding. Their mandibles have short teeth, and the endopods of the maxilla and maxilliped bear normal setae. The remaining genera in each clade have rather uniform modifications to the mandible, maxilla, and maxilliped. A similar situation exists in the family Heterorhabdidae, within which distinct transformations occur from small-particle feeders such as Disseta, to highly specialized carnivores, such as Heterorhabdus and Neorhabdus. This evolutionary shift in feeding type is accompanied by a strong reduction or loss of dorsal teeth on the mandible and an increase in size of the ventralmost tooth (Nishida & Ohtsuka, 1996). The phylogenetic analysis of the Heterorhabdidae by Ohtsuka, Soh & Nishida (1997) recovered the small particle-feeding genera as basal branches and the specialized carnivorous genera as terminal branches. We infer that carnivory has evolved independently within all of these families. Our phylogenetic analysis and conclusions concerning the taxonomic hierarchy at the base of the calanoid phylogeny are similar to the suggestions of Andronov (2007). Nevertheless, we have raised the rank of his family and subfamily names. Our analysis places Miheptneria in the same taxon as that of Andronov (2007), but separates out Azygonectes, the affinities of which he was not sure about, along with Erebonectoides and Caiconectes. We consider these genera currently to be incertae sedis (of uncertain placement). There is reasonably strong evidence for a differential diagnosis of a new family based on Caiconectes. It has a uniquely primitive setation pattern on the endopod of leg 1. It is the only calanoid with seven setae on the third endopodal segment. It also is the only calanoid we are aware of with five setae on the basis of the maxilla (most others have a maximum of four setae). In addition to these unique plesiomorphies, there is a cluster of other shared maximum plesiomorphic states relating to the distribution of aesthetascs on the antennules in both sexes, especially on segments XIX and XX (Appendix S 5 a, b). In addition, there are some apomorphies (e. g. reduced setation on the mandibular endopod and maxillule, and the coarse outgrowths of the long feeding setae on the maxilliped). At least on the basis of morphology, we would expect Caiconectes to be robustly recovered as the basal offshoot of the Calanoida in future analyses. On the other hand, Azygonectes and Erebonectoides is a potentially unstable group. These two genera share many key characters with other Epacteriscidae: presence of an aesthetasc on segment IV in female; the form of the mandible palp, with a reduced endopod; the asymmetry of caudal seta VI in the female, and of the caudal rami in the male; and endopod segment 1, of at least leg 3, with its distal outer corner bifid or trifid. But these character states alone are obviously not enough (on balance) to cluster them with the Epacteriscidae in this tree. The Pseudocyclopoidea are characterized by numerous homoplasious character states that ensure that the phylogenetic signal in the data is weak; therefore, none of the higher level taxa definitions is based on unique synapomorphies. The wide range of combinations of character states among the taxa in this superfamily hints at these taxa being a sparse sampling of a once much more diverse, ancient taxon, from which the ancestors of the Augaptiloidea, Centropagoidea, and a clade containing the remaining superfamilies evolved (Fig. 12). Part of their evolutionary capacity may have includ- ed the possession of character states that were easily reversed or independently reacquired, hence the currently observed diversity of character state combinations. Although the addition of new taxa or the revision of some character states may change relationships at the base of the tree, in our judgement, the pseudocyclopid and epacteriscid clades are likely to remain intact. The old concepts of the monogeneric Pseudocyclopidae and Boholinidae were based, at least in part, on the obvious external separation of the paired gonopores of the adult female (e. g. Huys & Boxshall, 1991: fig. 2.2.16). A similar arrangement was noted in at least some members of the Arietellidae, but the variation in structure of the female genital system within the family prompted Ohtsuka, Boxshall & Roe (1994) to recognize five major trends involving fusion of copulatory pores to form a single common pore, various migra- tions of gonopores and copulatory pores, and the asymmetrical enlargement of copulatory pores. Despite this variability in structure, Bradford-Grieve et al. (2010) were only able to use three characters based on female genital systems in their phylogenetic analysis of the Calanoida, and two of those were based on seminal receptacles and ducts. Variability in female genital structures can be found even in the more derived calanoid taxa, such as the clausocalanoidean family Stephidae. Unlike all of its congeners, the adult female of Stephos vivesi Jaume, Boxshall, Gràcia, 2008 possesses a pair of separate gonopores. Jaume et al. (2008) interpret- ed this condition as secondary, possibly derived by the loss of the genital operculum concealing the paired gonopores and their subsequent migration and separation. This serves to highlight the scale of intrafamilial variability in certain calanoid families, and the variability within the revised and enlarged concept of the family Pseudocyclopidae should be interpreted from this perspective. It is interesting to note that the two taxa that are basal to the Epacteriscidae and Pseudocyclopidae live in the open ocean at depth, and that the terminal taxa are cave-dwelling in the Epacteriscidae and cavedwelling, shallow-water, or even groundwater-dwelling in the Pseudocyclopidae. Thus, it appears that any hypothesis about the sequence of events surrounding the colonization of anchialine cave environments may be exactly the opposite from that proposed by Boxshall & Jaume (2000) for the Misophrioida. The misophrioid family Speleophriidae currently comprises eight genera and 19 species, almost all of which occur only in coastal anchialine habitats. The exception is Archimisophria Boxshall, 1983, which contains two species, both found in the deep hyperbenthic community of the tropical Atlantic. This genus is not basal within the family, and is recovered as the sister taxon of the cave-dwelling genus Expansophria Boxshall & Iliffe, 1987 (Boxshall & Jaume, 2000). In the case of the speleophriids, Boxshall & Jaume (2000) inferred that the presence of species in the deep sea was secondary, and that they were probably descended from shallow-water ancestors.	en	Bradford-Grieve, Janet M., Boxshall, Geoffrey A., Blanco-Bercial, Leocadio (2014): Revision of basal calanoid copepod families, with a description of a new species and genus of Pseudocyclopidae. Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 171 (3): 507-533, DOI: 10.1111/zoj.12141, URL: http://dx.doi.org/10.1111/zoj.12141
