#NEXUS [ File saved by NDE version 0.5.0, Tue Nov 19 12:18:52 2013 ] BEGIN TAXA; DIMENSIONS NTAX=56; TAXLABELS Gastrotricha Kinorhyncha Loricifera Nematomorpha Nematoda Aysheaia Peripatus Kerygmachela Microdictyon Tardigrada 'Acanthopriapulus horridus' 'Ancalagon minor' 'Anningvermis multispinosus' 'Chalazoscolex pharkus' 'Corynetis brevis' 'Cricocosmia jinningensis' 'Eximipriapulus globocaudata ' 'Fieldia lanceolata' 'Guanduscolex minor' 'Halicryptus higginsi' 'Halicryptus spinulosus' 'Laojieella thecata' 'Louisella pedunculata' 'Maccabeus cirratus' 'Maccabeus tentaculatus' 'Maotianshania cylindrica' Markuelia 'Meiopriapulus fijiensis' 'Ottoia prolifica' 'Palaeopriapulites parvus' 'Palaeoscolex piscatorum' 'Paraselkirkia jinningensis' 'Paratubiluchus bicaudatus' 'Priapulites konecniorum' 'Priapulopsis australis' 'Priapulopsis bicaudatus' 'Priapulopsis cnidephorus' 'Priapulus abyssorum' 'Priapulus caudatus' 'Priapulus tuberculatospinosus' 'Scolecofurca rara' 'Selkirkia columbia' 'Sicyophorus rarus' 'Tabelliscolex hexagonus' 'Tabelliscolex maanshanensis' 'Tubiluchus arcticus' 'Tubiluchus australensis' 'Tubiluchus corallicola' 'Tubiluchus philippinensis' 'Tubiluchus remanei' 'Tubiluchus troglodytes' 'Tubiluchus vanuatensis' 'Tylotites petiolaris' 'Xiaoheiqingella peculiaris' 'Xystoscolex boreogyrus' 'Yunnanpriapulus halteroformis' ; ENDBLOCK; BEGIN CHARACTERS; DIMENSIONS NCHAR=88; FORMAT DATATYPE=STANDARD MISSING=? GAP=- SYMBOLS="0123456789"; CHARLABELS [1] 'Degree to which the introvert can be invaginated' [2] 'Anterior branched frontal appendages' [3] 'Scalids (nonspecific and sensu lato, including both scalids and presumed scalid derivatives)' [4] 'Mouth cone' [5] 'Zone I' [6] 'Arrangement of zone I armature' [7] 'Radial arrangement of armature' [8] 'Morphology of zone I armature' [9] 'Number of elements comprising the first three circlets and, hence, defining the number of longitudinal rows of elements on the introvert' [10] 'Sequence of zone I elements' [11] 'Basal circlet of zone I armature separated from more anterior by a constriction (as in loriciferans) or by insertion of longitudinal or circular muscles (kinorhynchs)' [12] 'Number of elements in the proximal circlet of zone II' [13] 'Elongate spines at the base of Zone II' [14] 'Number of circlets of zone III armature' [15] 'Morphology of proximal circlets of zone III armature ("teeth")' [16] 'Morphology of middle circlets of zone III armature (teeth)' [17] 'Morphology of the distal circlets of zone III armature (teeth)' [18] 'Number of elements in first circlet of pharyngeal armature (base of zone III)' [19] 'Number of proximal, pentagonal circlets in zone III of the proboscis' [20] 'length of zone III relative to zone II' [21] 'Width of the distal portion of zone Ill' [22] 'Eversibility of zone Ill' [23] 'All zone III elements size ' [24] 'Armament of cervical region' [25] 'Aspect ratio of body length to width in adult' [26] 'Paired, lateral, locomotory appendages' [27] Annulation [28] 'Number of trunk annuli' [29] 'Trunk spines, fine spines, or setae in adult' [30] 'Trunk papillae in adult' [31] 'Trunk sclerites in adult' [32] 'Trunk sclerites' [33] 'Trunk tumuli' [34] 'Trunk tubuli' [35] 'Trunk tubercles of soft, coriaceous consistency' [36] 'Flosculi, N-flosculi or sensory spots' [37] Clavulae [38] Tube [39] 'Arc or ring of posterior spines or hooks (distinct from the bilaterally symmetrical structures present in Markuelia and many palaeoscolecids)' [40] 'Posterior ring papillae' [41] 'Eversible bursa' [42] 'Posterior abdomen greatly extensible' [43] 'Position of the anus' [44] 'Posterior tubulae or setae' [45] 'Caudal appendage(s)' [46] 'Division of caudal appendage(s) or tail' [47] 'Caudal appendage(s)' [48] 'Surface of caudal appendage' [49] 'Terminally posterior spines, hooks or cones of basal diameter >20% trunk diameter' [50] 'Posterior warts' [51] Polythyridium [52] 'Ventral nerve cord unpaired throughout its length' [53] 'Ventral nerve cords merge caudally' [54] 'Dorsal nerve cord unpaired' [55] 'Circumpharyngeal brain' [56] 'Brain with anterior posterior sequence of perikarya-neuropil-perikarya' [57] 'Apical part of the brain composed of perikarya' [58] 'Pharyngeal nervous system composed of numerous tooth ganglia connected by a diagonal nerve net' [59] 'Two rings of introvert retractors attached through the collar-shaped brain' [60] 'Developmental mode' [61] 'Loricate stage' [62] 'Cuticle of the larva/juvenile dorso-ventrally flattened (at least in older stages), with six lateral plates in-folded and accordion like. Cuticle of the lorica thickened in dorsal and ventral plates (at least) with sculpture of four to six longitudinal row' [63] 'Larvae/juveniles with six long pharynx retractor muscles' [64] 'Teeth of second circle of the larvae/juveniles with very small median denticle' [65] 'Division of the body into a distinct proboscis and abdomen in larva/juvenile' [66] 'Cloaca in both sexes' [67] Protonephridia [68] 'Urogenital system attached to the body wall by a ligament' [69] 'Spermatozoa with a flagellum' [70] 'Genital field extremely simplified' [71] 'Copulatory organs' [72] 'Perigenital setae' [73] 'Cuticle predominantly containing collagen' [74] 'Distribution of chitin in the cuticle' [75] 'Cuticle with homogeneous layer at surface/beneath epicuticle' [76] 'Cuticle with middle layer of distinct composition' [77] 'Cuticle with radially striated/vertical canal middle layer' [78] 'Cross-wise fibres in cuticle' [79] ' Large helical fibres in the cuticle' [80] 'Cuticle surface with ornament of tessellating polygons' [81] 'Division of the body into a distinct proboscis and abdomen in adult' [82] 'Terminal mouth' [83] 'Percentage of animal length taken up by the introvert' [84] 'Voluminous primary body cavity' [85] 'Movement by peristaltic movement of the pharynx/introvert' [86] 'Pharyngeal lumina' [87] 'Circular body musculature' [88] 'Nucleation of "peritoneal" membrane' ; STATELABELS 1 'no introvert, or introvert not invaginable at all ' 'partially invaginable (i.e., part of zone I) ' 'completely invaginable into the trunk (i.e., to the base of zone I)', 2 absent present, 3 'absent ' 'present and composed exclusively of cuticle ' 'present and with cuticle limited to a thin outer covering ', 4 'absent ' 'present but non-inversible ' 'present and inversible ', 5 'unarmed ' 'armed ', 6 'not in rows ' 'discrete parallel longitudinal rows ' 'rows aligned diagonal to the anterior-posterior axis of the animal ', 7 'pentaradial ' 'hexaradial ', 8 'papillae ' 'simple spines ' ' hooks or spinose hooks ' 'conical scalids ' 'telescopiform scalids ' 'curved scalids and dentoscalids ' ' complex scalids ' 'glandular scalids, trifid spines, sensory spines, and double, tentaculite scalids ' 'scalids with pectinate hood ' 'spinoscalids and clavoscalids (kinorhynchs and lorificerans) ', 9 'none ' '<20 ' '25 ' '>25 ', 10 'elements as a single series (all elements identical or with differing morphologies) ' ' elements organized into two or more transverse bands or series, possibly with different element morphologies within each series, but the sequence of morphologies being comparable between subsequent series ', 11 'absent ' 'present ', 12 'none or zone unarmed ' 'eight ' 'more than eight ', 13 ' absent ' 'present ', 14 'none or unarmed ' 'one to four ' 'six to fourteen ' 'sixteen or more ', 15 'spines or papillae ' ' multispinose' ' multispinose but massively reduced ' 'hooks ' 'conical with a fringe of spines ' 'sclerotized trabeculae ' 'pectinate ' 'conical papillae terminating in a long spine (prickle) ' 'oral stylets ', 16 'absent ' 'spines (of any length) or papillae ' 'multispinose (reduced or otherwise) ' 'pectinate ', 17 ' absent ' ' spines (of any length)' ' multispinose (reduced or otherwise) ' ' pectinate ', 18 'elements absent ' 'first circlet of 5 elements ' ' first circlet of 8-10 elements ' 'first circlet of numerous elements (more than 10) ', 19 'none ' 'five ' 'six ' 'seven ', 20 'zone III less than twice the width of zone II ' 'zone III equal to or greater than twice the width of zone II ', 21 'distal zone III parallel to proximal zone III or tapering gradually ' 'distal zone III expanded into a bulb ', 22 'zone III completely eversible ' 'zone III incompletely eversible, but eversible beyond the proximal teeth ' 'zone III normally eversible only as far as the proximal teeth ', 23 ' approximately equal size ' 'decreasing regularly in size from the posterior to the anterior (anteriormost elements less than half the size of the posteriormost) ', 24 'none ' 'cuticular plates (as in larvae/juveniles) ', 25 '<10 ' '10-20 ' '>20 ', 26 'absent ' 'present ', 27 'absent ' 'homonomous ' 'heteronomous ', 28 '<7 ' '7-11 ' '30-60 ' '61-120 ' '140 or more ', 29 'absent ' 'present ', 30 'absent ' 'present and simple ' 'extremely long and arranged into two discrete, longitudinal rows ', 31 'absent ' 'present and small to medium in size ' 'present and greatly enlarged ', 32 'absent ' 'in two longitudinal rows ' 'in annular rings ', 33 'absent ' 'present ', 34 'absent ' 'present ', 35 'absent ' 'present ', 36 'absent ' 'flosculi present with between 7 and 11 petals ' 'flosculi present and invariably with 8 petals ', 37 'absent ' 'present ', 38 'absent ' 'tube composed of plant debris: absent ' 'tube flattened, annulated, tapering and chitinous ', 39 'absent ' 'present ', 40 'absent ' 'present ', 41 'absent ' 'present ', 42 'no ' 'yes ', 43 'anus terminal, whether within bursa or otherwise ' 'anus in posterolateral or posteroventral surface of the abdomen ', 44 'absent ' 'present ', 45 absent 'present but less than the length of the body ' 'present and up to three times the lenth of the body ', 46 'undivided ' 'pseudosegmented ', 47 'single and positioned terminally ' 'single and positioned dorso-medially ' 'bicaudal as in some species of Priapulopsis and Paratubiluchus ', 48 'smooth ' 'vesiculae ' 'large warts ', 49 'absent ' 'present and arranged in pairs about the sagittal plane ', 50 'absent ' 'small (as in Halicryptus) ' 'large (as in Priapulus) ', 51 'absent ' 'present ', 52 'absent ' 'present ', 53 'absent ' 'present ', 54 'absent ' ' present', 55 'absent ' 'present ', 56 'absent ' 'present ', 57 'absent ' 'present ', 58 'absent ' 'present ', 59 'absent ' 'present ', 60 'direct ' 'biphasic ', 61 'absent ' 'present ', 62 'absent ' 'present ', 63 'absent ' 'present ', 64 'absent ' 'present ', 65 'absent ' 'present ', 66 'absent ' 'present ', 67 'absent ' 'present but do not flow into the gonoduct and are not integrated into the gonad ' 'present and flow into the gonoduct and/or are integrated into the gonad ', 68 'absent ' 'present ', 69 'absent ' 'present ', 70 'no ' 'yes ', 71 'absent ' 'present ', 72 'absent or reduced ' 'prominent ', 73 'absent ' 'present ', 74 'none ' ' predominantly within the middle cuticle layer (exocuticle)' 'predominantly within the lowermost cuticle layer (endocuticle) ', 75 'absent ' 'present ', 76 'absent ' 'present ', 77 'absent ' 'present ', 78 'absent ' 'present ', 79 'absent ' 'present and composed of platy subunits ' 'present and composed of unpaired fibrils ' 'present and composed of paired, wound fibrils ', 80 'absent ' 'present ', 81 'absent ' 'present ', 82 'absent ' 'present ', 83 'None (no introvert) ' '2% (as in Halicryptus higginsi) ' 'between 2% and 30% ' '30 to 50% ', 84 'absent ' 'present ', 85 'absent ' 'present ', 86 'round ' ' triradiate ', 87 'absent ' 'present ', 88 'membrane without nuclei or simply with amoebocytes in association with the surface ' 'membrane containing scattered nuclei ', ; MATRIX Gastrotricha -000----0- -0-----0-- ---0000010 0000000000 00000---00 00000000-0 0-0-001010 000000-000 0100011? Kinorhyncha 2022120910 10-1811200 0201000010 0000010000 00000---00 00?0111010 0-0-101010 0001110000 11200(01)10 Loricifera 2022120910 1101801201 0101000000 0000010000 00000---00 0000111011 1000102110 0002110000 11200110 Nematomorpha 0010101110 0????????? ???020(01)411 0000000010 00100---10 0011111001 0-00110000 001210-1(123)1 0120000? Nematoda 0010101110 0????????? ???020(01)410 0000000000 (01)0000---00 0011111000 0-0-010000 0012(01)1(01)(01)20 01200(01)0? Aysheaia -100----0- -0-----0-- ---0112211 0000000000 000?0---00 0????????? ???-0????? ?????????? 010????? Peripatus -000----0- -0-----0-- ---0111?11 0000000000 00100---00 000?000000 0-0-000010 0002110000 0000011? Kerygmachela -100----0- -0-----0-- ---0112411 0000000000 00?00--?00 0????????? ???-0????? ?????????? 010???1? Microdictyon -100----0- -0-----0-- ---0111300 2100000000 00000---00 0????????? ???-0????? ?????????? 010????? Tardigrada -100----0- -0-----0-- ---0011100 (01)100000000 00100---00 000?1000?0 0-0-0(01)0010 0002(01)1(01)00? 0(01)00010? 'Acanthopriapulus horridus' 2021110320 0????????? 02?0001201 0000000001 0010111100 0100111111 1111102110 000?????00 11311011 'Ancalagon minor' 00(12)?120210 00-1300200 0200101410 0000000000 00000---00 0????????? ?????????? ?????????? 112????? 'Anningvermis multispinosus' ??(12)?02--1- -2131111?1 0000101211 0000000000 ?0??100000 0????????? ?????????? ?????????? 112????? 'Chalazoscolex pharkus' ?0(12)?1??1?? 0?-??????? 1??0101400 2200000000 10000---00 ?????????? ?????????? ?????????? 112???1? 'Corynetis brevis' ??(12)?02--1- -2131111?1 0000101211 0000000001 ?00?100000 0????????? ?????????? ?????????? 112????? 'Cricocosmia jinningensis' ?0(12)?12?210 0003011101 1000201300 2101000000 00000---10 0????????? ?????????? ?????????? 112????? 'Eximipriapulus globocaudata ' 10(12)?110(23)30 00010????0 02?0001311 0000000001 00000---00 0????????? ????1????? ?????????0 11211??? 'Fieldia lanceolata' ?0(12)?12?210 00-1000300 0?00100010 0000000000 00000---00 0????????? ?????????? ?????????? 112????? 'Guanduscolex minor' ?0(12)?120?10 0003011300 0000101300 1100000000 00000---00 0????????? ?????????? ?????????? 112????? 'Halicryptus higginsi' 2021110520 0203121121 0210101410 0000000001 00110---01 0100111111 1110102110 000211(01)000 11111011 'Halicryptus spinulosus' 2021110520 0203121121 0210001310 0000000001 00110---01 0100111111 1110102110 000211(01)000 11211011 'Laojieella thecata' ?0(12)?11?110 0003011200 0000100000 0000000000 00001020?0 0????????? ?????????? ?????????? 112????? 'Louisella pedunculata' 10(12)?120010 0213011301 1000101?12 ??00000000 10000---00 0????????? ?????????? ?????????? 112????? 'Maccabeus cirratus' 2021110(27)20 01025211(23)0 0200001300 0000110110 01110---00 0100111111 1110102110 000?????00 11211010 'Maccabeus tentaculatus' 2021110(27)20 01025211(23)0 0200001300 0000110110 01110---00 0100111111 1110102110 000?????00 11211010 'Maotianshania cylindrica' 20(12)?120??0 0??3???3?0 0010201300 0000000000 00000---10 0????????? ?????????? ?????????? 112????? Markuelia 202?10?120 00-0---0-- ---01013(01)0 0000000000 00000---10 0????????0 0-?01????? ?????????0 112????? 'Meiopriapulus fijiensis' 2021110830 0102633300 0100000010 0011020010 00000---00 1100110110 0---?02110 000????100 11211010 'Ottoia prolifica' 20(12)?120230 02034(12)2301 1010001300 0000000010 10000---00 010??????? ?????????? ?????????? 1121???? 'Palaeopriapulites parvus' ?02?110210 0????????? ???0000000 0000000000 00000---00 0????????? 10???????? ?????????? 1131???? 'Palaeoscolex piscatorum' ?0(12)?10???? 0???011??? 0000201400 2200000000 00000---10 0????????? ?????????? ??1????121 112????? 'Paraselkirkia jinningensis' 10(12)?1001(23)0 0202???300 000010000? 0000000200 00000---?0 0????????? ?????????? ?????????? 112????? 'Paratubiluchus bicaudatus' ?0(12)1110?20 000(123)011??0 0??0000010 001?0??0?? ?0??102000 1????????? ?????????? ?????????? 113????? 'Priapulites konecniorum' 20(12)?110311 0????????? 02?0001200 0000000000 0010101000 0????????? ?????????? ?????????? 1131???? 'Priapulopsis australis' 2021110421 0103021211 0210001201 0010000001 0010100100 0100111111 1111102110 000??????0 11311011 'Priapulopsis bicaudatus' 2021110421 0103221211 0210001201 0010000001 0010102100 0100111111 1111102110 000??????0 11311011 'Priapulopsis cnidephorus' 20(12)111???? 0?03?????? ???0001201 001000000? 00101???00 ?100?????? ????1????? ?????????0 11211?1? 'Priapulus abyssorum' 2021110321 0103121131 0210001201 0000000001 0010111100 0100111111 1111102110 000111(01)000 11311011 'Priapulus caudatus' 2021110321 0103121131 0210001201 0000000001 0010111102 0100111111 1111102110 000111(01)000 11311011 'Priapulus tuberculatospinosus' 2021110321 0103121131 0210001201 0000000001 0010111202 0100111111 1111102110 000111(01)000 11311011 'Scolecofurca rara' ?0??1??0?0 00-3011?01 1?00?01?00 0000000000 ?0000---?0 0????????? ?????????? ?????????? 112????? 'Selkirkia columbia' 10(12)?100130 0202122300 0010100001 0000000200 00000---00 0????????? ?0???????? ?????????? 112????? 'Sicyophorus rarus' ?02?110210 0?0?0112?? 1?00000000 0000000000 00000---00 0????????? 10???????? ?????????? 1131???? 'Tabelliscolex hexagonus' ?0(12)?12?210 0003011101 ???0201300 2101000000 00000---10 0????????? ?????????? ?????????? 112????? 'Tabelliscolex maanshanensis' ?0(12)?12?210 0003011101 ???0201300 2101000000 00000---10 0????????? ?????????? ?????????? 112????? 'Tubiluchus arcticus' 2021110620 0102733300 0101000010 0011011000 0010200000 1100110111 1010102110 000?????00 11211010 'Tubiluchus australensis' 2021110620 0102733300 0100000010 0011011000 0010100000 1100110111 1010102111 000?????00 11211010 'Tubiluchus corallicola' 2021110620 0102733300 0100000010 0011011000 0010100000 1100110111 1010102110 010?????00 11211010 'Tubiluchus philippinensis' 2021110620 0102733300 0100000010 0011011000 0010100000 1100110111 1010102111 110?????00 11211010 'Tubiluchus remanei' 2021110620 0102733300 0100000010 0011011000 0010200000 1100110111 1010102110 100?????00 11211010 'Tubiluchus troglodytes' 2021110620 0102733300 0100000010 0011021000 0010200000 1100110111 1010102110 000?????00 11211010 'Tubiluchus vanuatensis' 2021110620 0102733300 0001000010 0011011000 0010100000 1100110111 1010102110 000?????00 11211010 'Tylotites petiolaris' ?0(12)?12?210 0002011200 1000101210 220?000000 00000---10 0????????? ?????????? ?????????? 112????? 'Xiaoheiqingella peculiaris' ?02?110120 010(123)011??? ???010130? 0000000001 ?00?102000 0????????? ?????????? ?????????? 112????? 'Xystoscolex boreogyrus' ?0(12)110?1?? 0?-?011??? ?000101400 1?00000000 00?00---00 ?????????? ?????????? ?????????? 112???1? 'Yunnanpriapulus halteroformis' ?02?110?20 0??(123)?????? ???0101?01 0000000001 ?00?100000 0????????? ?????????? ?????????? 112????? ; ENDBLOCK; BEGIN ASSUMPTIONS; OPTIONS DEFTYPE=UNORD POLYTCOUNT=MINSTEPS; ENDBLOCK; BEGIN NOTES; [Taxon comments] [Character comments] [Character state comments] [Attribute comments] [Taxon pictures] [Character pictures] [Character state pictures] [Attribute pictures] ENDBLOCK; BEGIN PAUP; ENDBLOCK;