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Serphitoidea (disused)
Taxonomy
Cydnides was named by Billberg (1820). It is extant. Its type is Cydnus.
It was reranked as the subfamily Cydninae by Scudder (1878), Scudder (1890); it was corrected as Cydnidae by Archibald and Mathewes (2000), Wappler (2003), Lis et al. (2015), Lis et al. (2018), Szwedo (2018), Wu et al. (2018), Popov (2019).
It was assigned to Pentatomidae by Scudder (1878), Scudder (1890); to Heteroptera by Archibald and Mathewes (2000), Wappler (2003); and to Pentatomoidea by Lis et al. (2015), Lis et al. (2018), Szwedo (2018), Wu et al. (2018), Popov (2019).
It was reranked as the subfamily Cydninae by Scudder (1878), Scudder (1890); it was corrected as Cydnidae by Archibald and Mathewes (2000), Wappler (2003), Lis et al. (2015), Lis et al. (2018), Szwedo (2018), Wu et al. (2018), Popov (2019).
It was assigned to Pentatomidae by Scudder (1878), Scudder (1890); to Heteroptera by Archibald and Mathewes (2000), Wappler (2003); and to Pentatomoidea by Lis et al. (2015), Lis et al. (2018), Szwedo (2018), Wu et al. (2018), Popov (2019).
Synonymy list
| Year | Name and author |
|---|---|
| 1878 | Cydninae Scudder p. 769 |
| 1890 | Cydninae Scudder p. 436 |
| 2000 | Cydnidae Archibald and Mathewes p. 1445 |
| 2003 | Cydnidae Wappler |
| 2015 | Cydnidae Lis et al. p. 846 |
| 2018 | Cydnidae Lis et al. p. 258 |
| 2018 | Cydnidae Szwedo p. 113 |
| 2018 | Cydnidae Wu et al. |
| 2019 | Cydnidae Popov p. 385 |
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If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.
Superfm. †Serphitoidea Brues 1937
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Diagnosis
| Reference | Diagnosis | |
|---|---|---|
| C. Callaway 1877 | Oval, moderately convex; axial furrows tapering gradually backwards from where they meet the facial suture in front of the eye to two thirds the length of the tail; head and thorax about equal in length, tail shorter. | |
| C. D. Walcott 1910 |
Dorsal shield broad, ovate. Cephalon large, semicircular in out- line, about one-third the length of the dorsal shield genal angles ; extended into spines ; facial sutures rudimentary or in a condition of symphysis ; eyes crescentic, with ridges uniting them with the anterior lobe oi the glabella ; glabella elongate, with a relatively small anterior lobe and three posterior transverse lobes ; strong occipital ring. Thorax with twent}'-eight segments ; body of pleurcie nearly straight ; pleural furrow broad and parallel to the transverse axis of the jilcune ; pleura; terminating in long, curved spines that are much shorter on the posterior eleven segments in the type species which are without a distinct, furrowed pleural lobe. Pygidium small, without pleural lobes and transverse furrows. Surface minutely granular and with irregular network of fine, irregular, anastomosing ridges. | |
| C. D. Walcott 1910 | Cephalon very large, wider than long, genal angles with spines ; intergenal spines developed in young and may be pres- ent in adult. Facial suture rudimentary, .or in a condition of syn- thesis. Eyes crescentic or semicircular and attached more or less closely to the anterior lobe of the glabella by a rounded ridge ; visual surfaceofeyeswithfacetsarrangedinquincunxorder. Hypostoma usually with more or less spinose posterior margin. Thorax long, composed of from 13 to 27 free segments. Pygidium siuall, margin usually entire but may have from one to three spines. Surface of test in adult specimens granular and usually with network of very fine thread-like raised inosculating ridges. | |
| J. H. Bradley 1930 | Cephalon subelliptical in outline, weakly and evenly convex, about four-fifths as long as wide. The anterior and lateral slopes of the glabella are somewhat steeper than the posterior slope. Median lobe slightly more elevated than the lateral lobes, evenly convex from front to back. The first glabellar furrows are narrow but sharp, converging very little posteriorly, thus making the median lobe almost parallel sided, becoming obsolete at the base of the lateral lob,ls. The lateral lo·bes· are well circumscribed on the inside and in front and slope laterally to the margin of the shield. Second glabellar furrows obsolete; third glabellar furrows shallow, poorly defining the fourth glabellar lobes. Occipital lobes very small, showing as a slight intumescence on the bases of the fourth glabellar lobes. Occipital furrow straight, deep, wider than the glabellar furrows. Anterior margin of the shield is flattened and extends well around the lateral lobes before being intercepted by the facial suture. Fixed cheeks poorly preserved, but probably small and non-protruding. Surface pustulose. Known only from the cranidium. | |
| E. S. Cobbold 1931 | Dorsal shield convex, with relatively strong axial lobe. Test granular, the granules tending towards linear arrangement. Cephalon elongate, semi-elliptical to semicircular. Cranidium trapezoidal, features in strong relief; axial lobe narrowing towards the occipital ring; glabella extending to the anterior border, with four pairs of side furrows, more or less obsolete anteriorly, and tending to sink into pits within, and independent of, the dorsal furrow; dorsal furrow clearly marked, sinking to a pit at a point in advance of the palpebral ridge independently of the glabellar furrows, coalescing with the anterior intra-marginal furrow; front border consisting of a narrow, convex, marginal rim limited by a sharply marked furrow; occipital ring well defined ; fixed cheeks sub-triangular, laterally steeply inclined; palpebral ridge short, not connected to the glabella; palpebral lobe rather long (one-third, more or less, of the length of the cranidium), depressed; facial suture, posterior branch a slightly sigmoidal curve extending outwards and downwards to the rounded lateral angles, anterior branch unknown. :Free cheek triangular with convex border and genal spine. Thorax unknown. Pygidium (?). In the pygidium associated with the genotype the anterior axial lobe is partially divided by a transverse groove, and the bases of the pleural spines are seen to originate from the anterior half, while the corresponding axial spine rises from the
posterior portion. | |
| C. E. Resser 1936 | Small trilobites with large glabella extending to the anterior edge. Glabella expanded forward so that its width at the front end is about twice that dimension at the neck ring; short, faint gla- bellar furrows present. Eyes small, situated forward of the midpoint. Fixed cheeks wide in rear, tapering rapidly to extinction at the an- terior end of the eyes, except for tiny anterior lobes.
Thorax in C. minor has seven segments. Pygidium small, flat, with segments fused. Axial and pleural fur- rows present. | |
| C. E. Resser 1936 | Cranidium alone known. Glabella large, convex, rising above the fixed cheeks and extending forward to the rim. A strong occipital furrow separates a neck ring which is extended into a stout spine. Three sets of glabellar furrows extend across the glabella, slop- ing backward in the usual manner. Eyes fairly large, situated well back. Eye lines heavy. Brim consists of a thickened rim only. Facial sutures diverge slightly anterior to the eyes. | |
| C. E. Resser 1936 | All features normal for the family. Cephalon and pygidium like Bailiella except for the rim of cephalon. Rim thickened and extended backward in the middle, sufficiently in some species to connect the anterior furrow with the dorsal furrow.
Some species were referred to Ctenocephalus because the thickened rim simulates a median boss, but examination of the specimens shows at once that the structure is quite different. | |
| C. E. Resser 1936 | Eyeless trilobite, oval in outline. Glabella tapered, with several sets of short, recurved furrows ; extends nearly to the anterior furrow. Deep marginal furrow extends all around the cephalon, being interrupted only at the genal angles. Two short diverging furrows cross the preglabellar area, joining the dorsal with the marginal furrow, thus making a distinct lobe anterior to the glabella. Eye ridges faint.
Thorax of 15 segments, with blunt tips. Pygidium fused, transverse | |
| C. E. Resser 1936 | Cephalon alone known. Cephalon semicircular, eye- less. Glabella well defined, tapered, a little more than half the length of the head. Glabellar furrows present, the rear pair recurved, sepa- rating marginal lobes. Cheeks marked by irregular lines radiating from the dorsal furrow. Upturned rim with coarse granules ; other scattered granules on cheeks. A depression extends across the pre- glabellar area. | |
| C. E. Resser 1936 | A rather small trilobite with convex cephalon. Gla- bella distinctly marked by shallow dorsal furrow. Marginal furrow alsoshallow; thoraxandpygidiumtypicalofthefamily. | |
| C. E. Resser 1936 | Small trilobites of average form. Cranidium rather wide, convex in both directions ; facial sutures diverging slightly an- terior to the eyes, but owing to the infolding of the anterior angles appear to converge. Glabella well defined, tapering, truncated or rounded in front ; two thirds or more of the length of the cranidium. Glabellar furrows two or three short pairs ; neck furrow and ring well defined. Eyes small, situated in middle or somewhat back of the middle of the head. Eye lines present, brim consisting of a more or less thickened rim and a preglabellar area of varying relative width.
Judging from somewhat uncertain material, the thorax consists of about 15 segments. Pygidium small, well fused, with axis well defined, and with pleural grooves. | |
| C. E. Resser 1936 | Small trilobites of which the cranidium alone is known. Glabella narrow, tapering slightly ; truncated, about two-thirds as long as cranidium ; in some species ill defined on upper surface of test. Glabellar furrows showing on inner side of test. Brim convex with faint frontal furrows extending inward from the anterior angles for varying distances. Facial suture converging anterior to eyes. Eyes small. Occipital furrow developed on the cheeks, but sometimes al- most obsolete on the cranidium. | |
| C. E. Resser 1936 | Cranidium only part known. Broad, with wide gla- bella and fixed cheeks. Glabella wide, rounded in front, apparently with a faint rear pair of joined glabellar furrows. Eyes slightly be- hind middle of head ; eye lines developed. Brim consists of narrow rim only. | |
| C. E. Resser 1936 | Cephalon semicircular ; glabella well defined, tapered, with glabellar furrows. Eyes lacking. Fixed cheeks large, apparently extending to the lateral margins. Marginal furrow extends entirely around the cephalon. Rim generally of even width but sometimes slightly expanded in the middle. Preglabellar area always present and exceeds that of Conocoryphe.
Thorax and pygidium typical of the family. Surface rarely smooth but usually pustulose or lined or both. Usu- ally a ridge or vein extends outward across the cheeks from the dorsal furrow, arising near the front of the glabella. This ridge occupies the position of, and resembles, an eye line, but in some species it is more like a vein branching as it advances. Anterior to this ridge the system of surface ornamentation characterizing the species gives way frequently to lines or veins. | |
| C. Lochman 1947 | Cranidium subquadrate, wider than long; glabella conical with slightly rounded front, convexity moderate, regular; three pairs of distinct glabellar furrows, anterior pair short, ·nearly straight, second and third pairs longer and arcuate; dorsal furrow of medium width, clear along sides, shallower across front; occipital furrow narrow, ·distinct, deeper at sides; occipital ring of ated on midline through glabella; palpebral furrow narrow, shallow; ocular ridge narrow, distinct, curved; posterolateral limb one and one-third times as long as occipital ring, of medium width, crossed by . a broad, deep intramarginal furrow. Free cheek not known. Facial suture cutting anterior margin far out at sides, curving out to marginal furrow, then running straight back and in to and curving around palpebral lobes; thence running straight out and then curving back · to cut posterior margin. Outer surface of test finely granulated. Thorax and pygidium not known | |
| V. J. Okulitch 1951 | The species differs from Nevadia weeksi Walcott by having a longer and narrower glabella with a more pronounced forward taper. The preglabellar area is much nar- rower than in N. weeksi. The palpebral lobes of N. addyensis are narrower and closer to the glabella and at least partially separated from it by dorsal furrows. Nevadia cartlandi (Walcott) differs from the new species in the different position of its eyes. | |
| C. Lochman 1952 | The species is most closely related to the other two Cordilleran Wanneria species, W. rowe i(Walcott) and W. occidens Walcott, but in so far as comparison can be made it differs from them in (i) the sudden bulge of the anterior lobe just infront of the palpebral lobes, (2) the very short occipital spine, and (3) the somewhat shorter eyes. The author recognizes this form as a species differing from described ones as noted above; and this particular form as a variety of that species in which the intergenal and genal spines are placed as noted. | |
| G. A. Cooper et al. 1952 | The species is very close to P.transitans Walcott. It appears to differ from that species only in the more rounded anterior lobe of the glabella and the straight posterior margin of the cephalon. As the type of P.transitans is a flattened, somewha tcrushed shale specimen, the author is unable to determine how much the difference in matrix has affected these details of the cephalon. She considers it feasible for the present to describe the Mexican material as a new species but to call attention to its close relation to P.transitans. In studying the species of Paedeumias the author concluded that nearly all the same features listed for Olenelliis constitute features of specific value in Paedeumias, i.e. (i) length of eye, (2) shape of anterior glabellar lobe, (3) width of brim and structure of median anterior ridge. In Paedeumias, however, there appears to be no variation in the width of the border, the narrow, rim like border being apparently a generic character. | |
| C. Lochman 1952 | The holotype of this variety is a cephalon 24mm. in length, and though not complete, does permit specific determination. It is most interesting to find the occurrence of this variety of the common Appalachian species in the Cordillerantrough. It agrees with Wanneria walcottana (Wanner) and differs from all other described species in the following features: (i) A regular and smooth expansion of the anterior lobe of the glabella; (2) the presence of a row of pits along the inner edge of the border; (3) the eyes and palpebral lobes reaching just to the center of the fourth glabellar lobe; (4) an occipital node rather than spine; (5) a very small intergenal spine, apparently aborted in large adults; (6) a medium-sized genal spine. | |
| C. Lochman 1952 | This species is described from a single small cranidium and seven pygidia ranging from 4 mm. by 2 mm. to 8 mm. by 4 mm. All the pygidia are poorly preserved; only one specimen shows the very small marginal spine. The species is close to three described species but differs from them as indicated. Bonnia tensa Resser and Bonnia fieldensis (Walcott) have all proportions and convexity practically the same, but both differ in that the glabella merges into the border as the marginal furrow is almost obsolete; and B. tensa Resser also has a steeper frontal slope of the glabella. A small cranidium in the B. fieldensis lot, which is the same size as the Mexican specimen, differs from it in exactly the same features as the larger B. fieldens is cranidia. This indicates that the specific features are already established in cranidia of this small size. Pygidia of the same size as B. fieldens is and B. sonorado not show any differences.The Mexican species is closest to fall to B. columbensis Resser from the Mount Whyte formation. The cranidium differs in (i) the steep slope of the front part of the fixed cheeks, and (2) the nearly flat profile of the glabella. In the pygidium of B. columbens is the marginal furrow is more distinct, especially around the back, and the posterior border tends to flatten out. With the present limited material representing both species, it is possible that, when more material is obtained, the Mexican species can be shown to be the same as B. columbensis Resser. | |
| C. Lochman 1952 | The species is most similar to the genotype, A. subcoronatum (Hall and Whitfield) differing from it mainly in the weaker development of the median bulge and the steeper descent of the border. In the genotype the border is essentially horizontal in position. | |
| C. Lochman 1952 | It may be distinguished from A. typicalis Resser by the descending nature of the narrow border as contrasted to the distinct upcurve in the latter species. The Mexican species also has a less convex glabella and a shallower dorsal furrow. It is doubtful how much importance should be put on these features, because all the types and duplicates of A. typicalis are internal molds in a punky yellow sandy leached limestone. Such preservation tends to emphasize the depth of furrows and convexity of glabella. A. mexicana dififers from A. brighamensis Resser in the flat horizontal border of the latter species, but as the types are crushed and flattened in shale this may be only a feature of preservation. The preservation of A. typicalis and A. brighamensis fails to show the low median bulge although some cranidia of the former species have a suggestion of it. | |
| C. Lochman 1952 | They definitely belong to Kochaspis but are not like any previously described species. It is most like the pygidia assigned to K. dispar Resser {K. maladensis Resser) but differs from them in the slight convexity of the pleural platforms and the direction of the marginal spines. In the holotype pygidium, chosen because of its completeness, the convexity is probably somewhat lower than that attained in the larger adult specimens. | |
| C. Lochman 1952 | The Mexican species is very similar to P. rossensis (Walcott) in the structure of the cranidium, showing only an occipital node rather than the usual strong spine. In the pygidium it appears that the four pairs of marginal spines were of medium length, and the species differs from all other described forms in the possession of two axial spines and the axial node on the third segment of the pygidium. | |
| C. Lochman 1952 | This species appears to differ from all described ones in (i) the gentle expansion and low convexity of the front of the glabella, and (2) the narrow, descending border of the pygidium. In fact, only two other species show a similar narrow border - G. helesis (Walcott), preserved in shale, and with the border apparently horizontal in position, and the paratype U.S.N.M.No.62695 of G.mckeei Resser. On this specimenal so the narrow border apparently lies in horizontal position and so is not comparable to the Mexican species. Attention should be called to the fact that the holotype of G.mckeei has a pygidial border of medium width and so could not represent the same species as the above-mentioned paratype. | |
| C. Lochman 1952 | The species appears quite close to Albertella bosworthi Walcott both in the forward expansion of the glabella and the five segments of the axis of the pygidium. The pygidium of A. proveedora Lochman differs, however, in greater width of the pleural lobes in proportion to the axis, and the apparent absence of the median nodes on the axis. The cranidium differs in the medium length of the palpebral lobes. Those of A. bosworthi Walcott appear to have been somewhat longer even though the lengthening due to crushing of the shale matrix is discounted. This situation presents a problem which should receive further study as it appears that in all limestone specimens of the cranidium the palpebral lobes are of medium length, where as in all shale specimens they appear to be larger. | |
| C. Lochman 1952 | The species is readily distinguished from Mexicaspis stenopyge Lochman, by (i) posterior part of glabella wider so that forward expansion appears less, (2) glabella moderately convex, (3) dorsal furrow in front of glabella and marginal furrow distincton outer surface, (4) pygidium with broader proportions, and (5) the lateral direction of the posterior pair of marginal spines. | |
| C. Lochman 1952 | This species differs noticeably from A. caborcana Lochman in the descending position of the border and the different proportions of the brim and border. Otherwise the two species are very close. | |
| C. Lochman 1952 | Of the described species of Inglefieldia it appears to be closest to I. discreta Poulsen, as in that species the brim and border are practically the same width, but the brim is descending, where as in I. imperfecta Lochman, new species, it appears to have been normally horizontal in position. The poor preservation of all the specimens makes it difficult to be sure about certain features of the carapace as indicated in the description. | |
| C. Lochman 1952 | The species is close to P. typicalis Resser, differing from it mainly in the narrower and less steeply sloping brim and the narrower marginal furrow. The small Mexican cranidia also appear to have stronger glabellar furrows and stronger convexity of the glabella when compared with the larger holotype cranidium of P. typicalis Resser, but as certain of the smaller cranidia of the latter species show a more convex glabella and deeper furrows than the holotype, the author considers these differences to be caused by size and not to be specific differences. | |
| C. Lochman 1952 | The species differs from P. deborra Lochman, new species, in the narrower and more steeply sloping brim and the somewhat wider border—the brim in front of the glabella being actually slightly narrower than the border on the midline. The border is also slightly sloping as contrasted to the horizontal position in P. deborra. | |
| A. R. Palmer 1954 | Cephalon subsemicircular in outline; posterior margin straight; distinct genal spines extend backward from posterolateral corners. Cranidium with glabella extending to or nearly to anterior margin, straight sided behind next to last glabellar furrow, distinctly expanded anterior to this furrow; anterolat- eral corners strongly rounded; anterior margin broadly rounded. Three pairs of straight glabellar furrows visible; posterior pair extended backward at a sharp angle with dorsal furrow; middle pair approximately perpen- dicular to dorsal furrow; anterior pair extended forward from dorsal furrow. Occipital furrow distinct, deepest at sides. Occipital ring flat, unspined. Width of fixed cheeks about one-third width of glabella. Palpebral lobes long, distinct; anterior ends nearly touching dorsal furrow in front of first distinct glabellar furrow. Pos- terior limbs longer than occipital ring, narrow.
Thorax of 7 or 8 segments. Axial lobe narrower than pleural lobes. Pleural lobes of each segment subequally divided by broad pleural furrow and extended distally into a short pointed spine. Pygidium subsemicircular in outline, nearly as large as cephalon. Axial lobe prominent, furrowed, about as wide as pleural platforms, not extended onto concave border. Pleural lobes with pleural furrows and inter- pleural grooves about equally spaced, usually extended onto border. | |
| A. R. Palmer 1954 | Cephalon subsemicircular in outline; posterior margin straight; distinct genal spines extend backward from posterolateral corners. Cranidium with glabella moderately well to poorly defined by dorsal furrow, extended to anterior margin, slightly expanded forward or of nearly constant width. Glabellar furrows shallow or absent; when visible, four pairs usually present. Occipital furrow shallow. Occipital ring flat, unspined. Border absent. Brini, when present, nar- row, only at sides of glabella. Fixed cheeks about one- half width of glabella; palpebral lobes long, poorly defined; anterior ends nearly touching dorsal furrow, posterior eads about opposite occipital furrow. Poste- rior limbs narrow, about as long as occipital ring; marginal furrow shallow.
Free cheek with evenly rounded margin and prominent genal spine. Border differentiated from ocular platform, variable in width. Thorax of 7 or 8 segments. Axial lobe prominent, width equal to or less than that of pleural lobes. Pleural lobes of each segment subequally divided by pleural furrow. Segments terminate distally in short, sharp spines. Pygidium subsemicircular in outline about as large as cephalon. Axial lobe prominent, tapered slightly posteriorly, reaching to or onto broad, flat or concave border. Furrows on axial and pleural lobes shallow or absent. | |
| A. R. Palmer 1954 |
Diagnosis.—Cranidium like Olenoides Meek. It dif- fers from typical forms of that genus by having the anterior end of the glabella overhanging the anterior margin, a shallow occipital furrow, but no occipital spine. Pygidium with axial lobe moderately well defined, narrow, three-fourths or more the length of pygidium, consisting of four segments and a rounded terminal portion. Pleural lobes consisting of four segments; distinct pleural furrows and interpleural grooves pres- ent; pleural furrows parallel and adjacent to groove marking posterior margin of each pleural segment. First and second pleural segments rapidly expanding distally, second segment largest, and extended into moderately long, straight, posterolaterally directed marginal spines; second pair of marginal spines origi- nates posterior to end of axial lobe. Third pleural seg- ment narrower than second segment; pleural furrow and interpleural groove nearly parallel to dorsal furrow. Fourth pleural segment, when discernable, less than one-third width of third segment, poorly defined, approximately parallel to dorsal furrow. Third and fourth segments not extended into marginal spines. Posterior margin of pygidium between spines of second pleural segment, curved gently forward. | |
| A. R. Palmer 1954 | Cephalon subsemicircular in outline; posterior margin nearly straight; genal spines originating from posterolateral corners. Cranidium subtrapezoidal in outline. Glabella parallel sided or slightly expanded forward, prominently elevated above cheeks, reaching to border, smooth or faintly furrowed, well defined by dorsal furrow. Occipital furrow distinct. Occipital ring usually bears median spine. Fixed cheeks narrow, less than one half width of glabella. Ocular ridges distinct, strongly backswept from anterolateral margin of glabella. Distinct pits are present in dorsal furrow at junction with ocular ridge. Palpebral lobes small, situated opposite middle third of glabella. Posterior limbs long, wide; marginal furrow distinct.
Thorax consists of about seven segments. Axial portion of segments often with median node or spine. Pleural portions with distinct, posterolaterally directed furrow originating near inner anterior corner of lobe. Marginal spines sharp. Axial lobe gently to moderately tapered posteriorly. Pygidium subsemicircular in outline. Axial lobe well defined by dorsal furrow, tapered posteriorly, reaching to or nearly to the inner margin of the border. Pleural lobes with distinct interpleural grooves and diagonal pleural furrows. Pleural furrows usually deepest. Margin with four or more pairs of posteriorly or posterolaterally directed, nearly horizontal spines. | |
| A. R. Palmer 1954 | Cranidium with glabella prominent, nearly parallel sided, extended to border. Glabellar furrows marked by three pairs of pits often connected by shallow furrows across top of glabella but not con- nected to dorsal furrow. Occipital furrow distinct,
.with deep pits near dorsal furrow. Brim not present in front of glabella. Border flat, short. Width of fixed cheek one-half or more width of glabella. Palpe- bral lobe moderately large, opposite middle third of glabella, connected to dorsal furrow by distinctly to poorly defined ocular ridge. Facial suture straight for- ward in front of palpebral lobe, divergent behind. Free cheek narrow, margin evenly rounded. Border well defined by marginal furrow, extended posteriorly into genal spine. Thora,x of 6 or 7 segments. Axial lobe prominent, width greater than one-half width of pleural lobes. Pleural lobe of each segment with deep diagonal pleural furrow; distal spines of variable length. Pygidium subsemicircular in outline, consisting of 5 segments and a small terminal portion. All segments visible on axial and pleural lobes; pleural furrows deeper than interpleural grooves. Each segment extended distally into a marginal spine. | |
| A. R. Palmer 1954 | Cephalon subsemicircular in outline; posterior margin straight; long genal spines extend out- ward and backward from lateral margins; short inter- genal spines may be present at distal ends of posterior limbs on cranidium.
Cranidium with glabella moderately well defined by dorsal furrow, parallel sided, slightly expanded forward or slightly tapered forward, not reaching to anterior margin. Glabellar furrows variable in depth, 3 or 4 pairs usually visible. Occipital furrow distinct, straight. Occipital ring flat, with or without occipital spine. Frontal area short, broad, with depressed brim and raised border. Fixed cheeks about one-half width of glabella, semicircular hi outline. Palpebral lobes narrow, well defined by palpebral furrow, long, curved; anterior ends adjacent to dorsal furrow opposite third pair of glabellar furrows anterior to occipital furrow; posterior ends opposite occipital furrow, about one-third width of glabella from dorsal furrow. Posterior limbs narrow, about as long as occipital ring, often with distinct, posteriorly directed spines at distal extremities. Facial sutures widely divergent anterior and posterior to palpebral lobe. Free cheek with evenly rounded margin and distinct, narrow border. Long, slender genal spine directed outward and backward from lateral margin. Thorax of 8 or 9 segments. Axial lobe as wide or wider than pleural lobes exclusive of spines. Pleural portion of each segment with two laterally converging marginal furrows that leave a triangular raised median portion near the dorsal furrow. Each segment extended distally into a long slender spine. Pygidium smaller than cephalon with prominent furrowed axial lobe occupying one-third or more width of pygidium, extended nearly to posterior margin. Anterolateral corners of pygidium strongly rounded. Posterior margin with three or more pairs of posteriorly directed spines of varying lengths. | |
| A. R. Palmer 1954 | Micropygous ptychoparioidea. Frontal area long. Brim and border present but poorly differentiated. Median swelling extends from brim onto border. Width of fixed cheeks greater than one-half width of glabella. Thorax of 17 to 23 segments. | |
| A. R. Palmer 1954 | Cephalon subsemicircular in outline, posterior margin nearly straight, genal angles rounded. Cranidium with glabella tapered forward, strongly rounded anteriorly, poorly defined by shallow dorsal furrow. Glabellar furrows not visible. Occipital fur- row shallow. Occipital ring gently rounded. Frontal area with deep, curved marginal furrow. Border gently arched upward, width constant. Fixed cheeks down-sloping from glabella; palpebral lobes not defined by palpebral furrow, situated opposite middle third of glabella. Length of posterior limbs about equal to width of glabella at occipital furrow; marginal furrow broad, shallow. Facial sutures divergent anterior and posterior to palpebral lobes. Anterior course extended onto border and then turned abruptly inward to cut anterior margin intermediate between anterolateral corners of cranidium and longitudinal axis. Posterior course extended backward and outward in even curve to cut margin within genal angle.
Free cheek with evenly rounded lateral margin. Marginal furrow moderately deep, parallels margin. Genal angle rounded. Thorax consists of nine segments. Axial lobe prominent, narrower than pleural lobes. Pleural lobes of each segment subequally divided by deep pleural furrows near axial lobe, flattened and rounded distally. Pygidium about as large as cephalon, subsemicircular in outline, width less than twice length. Axial lobe prominent, tapered posteriorly, extended to or onto border. Border of nearly constant width, narrower than widest portion of pleural platform. Axial and pleural furrows shallow. | |
| A. R. Palmer 1954 | Cranidium with anterior margin gently and evenly rounded, glabella prominent, tapered for- ward, bluntly rounded anteriorly; dorsal furrow deep at sides, shallow across front. Four pairs of glabellar furrows visible on well-preserved specimens. Occipital furrow deep at sides, shallow on midline. Occipital ring with or without spine. Frontal area with distinct brim and border separated by shallow marginal furrow or by sudden change in slope; border maintains con- stant width; axial length of frontal area less than half that of glabella. Width of fixed cheeks about one-half that of glabella. Prominent ocular ridges present. These touch dorsal furrow opposite anterior pair of glabellar furrows. Palpebral lobes poorly defined by shallow palpebral furrow, depressed below general level of cheeks, opposite middle third of glabella. Length of posterior limbs about equal to width of glabella at occipital furrow. Anterior and posterior courses of facial sutures divergent from palpebral lobes; anterior course intramarginal on border nearly to axial line.
Free cheek with margin evenly rounded. Border narrower than ocular platform, well defined by shallow marginal furrow, extended into short, pointed genal spine. Thorax of 13 segments. Axial lobe prominent, narrower than pleural lobes. Distal ends of pleural lobes of each segment rounded or extended into short sharp spines. Pygidium with axial length one-half or more width. Axial lobe tapered posteriorly, not extended to posterior margin, composed of 3 to 5 segments. Pleural lobes with 2 or more pairs of pleural furrows and interpleural grooves of about equal strength; each pair of furrows directed more posteriorly than preceding pair. Border not present, but distal margins of downsloping pleural lobes flexed outward. Surface of carapace smooth. | |
| A. R. Palmer 1954 | Cranidium subquadrate in outline; gla- bella prominent, relatively small, tapered forward, truncate anteriorly. Glabellar furrows shallow or ab- sent. Occipital furrow shallow. Frontal area nearly as long as glabella; border not distinctly differentiated from brim; brim with large median boss extending from front of glabella to border. Fixed cheeks as wide or wider than glabella. Palpebral lobes small, opposite anterior third of glabella; palpebral furrow absent. Facial sutures straight forward in front of palpebral lobes, widely divergent behind.
Free cheek, thorax, and pygidium unknown. | |
| A. R. Palmer 1954 | Carapace elongate subovate in outline, widest at posterior margin of cephalon.
Cephalon subsemicircular in outline, genal angles extended into genal spines. Cranidium with anterior margin gently and evenly rounded. Glabella well defined by shallow dorsal furrow, straight sided, tapered forward, strongly and evenly rounded anteriorly. Glabellar furrows shallow or absent. Occipital furrow deepest at sides, shallow on midline. Occipital ring flat or gently arched, un- spined. Frontal area with distinct brim and border; border flat or gently arched upward; marginal furrow shallow. Width of fixed cheeks one-half or more width of glabella. Paipebral lobes opposite middle third of glabella, well defined by shallow palpebral furrow. Ocular ridges indistinct. Length of posterior limbs about equal to width of glabella at occipital furrow. Marginal furrow broad, shallow. Facial sutures diver- gent anterior and posterior to paipebral lobs; anterior course turns inward after reaching border and cuts margin about midway between anterolateral corners of cranidium and axial line; posterior course cuts margin before reaching genal spine. Free cheek with evenly curved lateral margin. Border well defined by shallow marginal furrow; ex- tended into short genal spine. Thorax of 13 segments. Axial lobe prominent; width less than half that of pleural lobes. Pleural lobe of each segment subequally divided by deep pleural fur- row; distal extremity extended into short, sharp point. Pygidium subsemicircular in outline, smaller than cephalon; axial length less than one-half width. Axial lobe prominent, not reaching posterior margin; width about one-half that of pleural lobes. Narrow border present, poorly defined by shallow marginal furrow. Furrows on axial and pleural lobes shallow. Surface of carapace smooth. | |
| A. R. Palmer 1954 | Cranidium subquadrate in outline, gently to moderately rounded anteriorly. Glabella well de- fined by dorsal furrow, straight sided or with sides slightly bowed outward, tapered forward, bluntly rounded anteriorly. Three pairs of deep glabellar fur- rows present. Occipital furrow deep. Occipital ring gently to moderately arched upward. Frontal area with distinct brim and border; border gently arched upward; marginal furrow shallow on midline, broad, deep distally. Width of fixed cheeks more than one- half width of glabella; prominent ocular ridges present. Palpebral lobes situated opposite middle third of gla- bella, poorly defined by shallow palpebral furrow. Length of posterior limbs about equal to width of glabella at occipital furrow; marginal furrow deep. Anterior course of facial suture nearly straight forward from palpebral lobe; posterior course divergent from longitudinal cranidial axis.
Free cheek and thorax unknown. Pygidium, excluding spines, subquadrate in outline. Axial lobe prominent, distinctly furrowed, variable in length; width slightly more than one-half width of pleural lobes. Pleural segments coalesced distally to form a pair of posteriorly directed marginal spines. Outer surface of carapace covered with scattered large granules; inner surface smooth | |
| J.L. Wilson 1954 | Cranidium of moderate size (less than 1 cm. long), longer than wide, moderate relief to individual parts. Glabella prominent but not highly arched, rather depressed posteriorly but well defined by dorsal furrow which shallows anteriorlv,., well taoered and rounded anteriorly, with occipital ring occupying about two-thirds of axial length disregarding anterior spine; three pairs of very faint glabellar furrows, posterior two curved sharply backwards proximally; occipital ring projected backward axially, occipital spine may be present. Frontal area highly variable, brim very short but border projected anteriorly a t axis. Fixed cheeks narrow (one-fourth to one-third basal glabellar width), elevated and convex, bearing wellmarked obliquely backward trending ocular ridges and small semicircular palDebral lobes elevated above them and locatedAabout on the midwidth line through glabella; posterior limbs narrow and extended straight out, deeply furrowed.
Facial suture cuts anterior margin a t long axis on non-spinose species, proceeds straight back to eye from anterolateral corners of cranidium, indents slightly behind same and trends straight out along posterior limbs. A pygidium assigned questionably to one of the species of this genus is figured. It is convex, a little wider than long, with a large elevated axis bearing three segments and a terminal portion; pleural lobes well marked by two or three interpleural grooves which fade out posterolaterally; border lacking. | |
| J.L. Wilson 1954 | The size range of the specimens (all from a single boulder) is from 5 to 10 mm. long. Almost half the cranidial length is formed by the border which is prolonged into a convex blunt spine. This is counter-balanced by a large thick occipital spine of about equal size although this is broken off all but one specimen. The test is finely tuberculate. The free cheek possesses a triangular ocular platform, steeply depressed laterally, a wellmarked marginal furrow and a relatively wide, moderately convex border. Presumably the anterior facial suture cuts the margin of the cranidium a t some place along the side of the cranidial spine. | |
| J.L. Wilson 1954 | Coosia with glabella and occipital ring about three-fourths of total axial length of cranidium; glabella elevated and convex but frontal area flat, rather short compared to that of other species, except C. robusta. Border about four times axial length of brim, separated from it by a well-defined marginal furrow; border bears no secondary (false) marginal furrow. Fixed cheeks about one-fifth basal width of glabella, palpebral lobes on or anterior to center of glabella, relatively small for the genus. Most distinctive specific feature is broad backcurvin- g .po sterior limbs.
Facial sutures cut anterior margin of cephalon away from axis, converge slightly anterior to eye and behind same trend outward obliquely a t about 60" to longitudinal axis. Pygidium almost quadrate, pleural platforms elevated and convex, axis half or less than half total axial length, bears three or four segments plus rounded terminal. Posterior border flat and unsegmented, marginal inbend present a t axis. | |
| A. R. Palmer 1954 | Like Olenoides but with only pleural furrows visible on the pleural platforms of the pygidium. | |
| A. R. Palmer 1954 | Pygidium with long, slender marginal spines. | |
| A. R. Palmer 1954 | As this is the only described species of the genus, its characters are those of the genus. | |
| J. Kukalová 1955 | Genus of the family Archimylacridae, characterised by the following features of the anterior wing: the ratio of the width to the length is 2.5. The costal area, expanded at the base, extends to 2/3 of the length of the wing. R is directed moderately obliquely below the apex. The first branch of R starts near the base. M strongly reduced, narrow and spar- sely ramified. Cu well developed. Anal area exceeding only a little 1/3 of the total length of the wing. The vena dividens forms a quite gentle arc. The regular transversal veins pass proximally into irregular ones. | |
| J. Kukalová 1955 | Genus of the family Arclzimylacridae, which is characterised by the following features: anterior wing suboval, ratio of width to length about 2.5. Costal area expanded at the base, extending beyond the middle ofthe total length of the wing. R is divided into two branches, and direct- ed above the apex. The media divides dichotomically before the middle. In comparison with R it is less rich in branches. The branches R and Cu + vena dividens connect at the base. The anal area ends in the + M
same plane as the costal area, and extends to about the middle of the length of the wing. The vena dividens runs in a moderate arc. The regular transverse weins pass into a dense reticulation. The posterior wing is characterised by a very short Sc running near the anterior margin. R divides near the base into two branches of different structure. Of these especially Rs is richly ramified and occupies the whole apical part of the wing. The legs of the third pair are provided with several rows of broad spines. Oculi compositi composed of ommatidia of circular shape, 0.032 mm. in size. | |
| A. B. Shaw 1955 | Olenellus with ten thoracic segments and a pygidium behind the spined fifteenth segment. The eyes are short with the posterior extremities lying opposite the pre-occipital lobe of the glabella, unlike O. thompsoni. Glabella long, with second pair of furrows behind anterior lobe short and incomplete. | |
| A. A. Stoyanow and T. Susuki 1955 | Diagnostic differences: from Anoria and Glossopleura, by eight thoracic segments; from S. torresi, by the thoracic tuberculation; from S. gomezi, by the course of facial sutures. | |
| H. J. Harrington 1956 | Olenellinae with well advanced genal spines. Metagenal spines absent in adult. Metagenal angle narrowly obtuse to acute. Glabella with three pairs of lateral glabellar lobes, strongly constricted at the level of the preoccipital furrow. Second (3p) furrow transglabellar, gently curved backward, not reaching the axial furrows. Preoccipital (2p) furrow curved, chevron shaped, disconnected at middle. Frontal glabellar lobe pyriform, expanded in front and encroaching over anterior border of cephalon. Occipital ring broad (sag.) and as wide (tr.) as frontal glabellar lobe. Eye lobes short, submedian (slightly anterior), reaching level of preoccipital furrow. Cephalic border raised, narrowing anteriorly. | |
| H. J. Harrington 1956 | Olenellinae with well advanced genal spines. Metagenal spines ab- sent in the adult. Metagenal angle sub-rectangular. Glabella parallel-sided with three pairs of lateral glabellar lobes, the an- terior (4p) narrower than the remainder and ill-defined. Frontal glabellar lobe not expanded, evenly rounded anteriorly, slightly encroaching over anterior border of cephalon. Eye lobes short, posterior, reaching level of mid-length of preoccipital glabellar lobe. Cephalic border wide, slightly raised andlflat. | |
| H. J. Harrington 1956 | Wannneriinae with well advanced genal spines. Metagenal spines well developed in adult. Metagenal angle slightly obtuse to slightly acute. Glabella with three pairs of lateral glabellar lobes and a pair of small depressions in front of the first lateral glabellar furrow. Glabella tapering backward. Frontal glabellar lobe ex- panded anteriorly, reaching the anterior border of cephalon. Eye lobes short, sub-median (slightly posterior) reaching level of anterior third of second (3p) lateral glabellar lobe. Posterior genal ridge conspicuous. Cephalic border wide, slightly raised and flat. | |
| H. J. Harrington 1956 | Olenellidae with moderately advanced genal spines. Metagenal spines absent or minute in the adult. Metagenal angle broadly obtuse. Glabella parallel-sided, with three pairs of lateral glabellar lobes. Anterior (4p) lobes slightly narrower than the remainder, well defined by the second (3p) transglabellar furrow. Frontal lobe of glabella moderately expanded, reaching anterior border of cephalon. Eye lobes short, submedian (slightly posterior), reaching level of anterior third of preoccipital glabellar lobe. Cephalic border raised, narrowing anteriorly. | |
| T. E. Tjernvik 1956 | Distance between eyes in front of glabella about half the width of the glabella at rear end. Pygidium subtriangular in outline, with a wide, flat tened border. | |
| G. Henningsmoen 1957 | A Ctenopyge species with : almost parallcl-siclecl glabella; Sl strong, usually united ae1·oss glabella, other glabellar furrows very short ; oblique eyc ridges ; centres of palpebral lo bes opposite Sl or L2 ; interocular cheeks somewhat narrower than glabella at eye line ; postocular cheeks slightly less to slightly more than 1 . 5 times as wicle as occipital ring ; frec cheeks with long flattened spine. obtuse genal angle, and straight posterior lateral margin slightly shorter than com-ex anterior l ateral margin ; thorax with pleural regwns ( excluding spines) somewhat w ider than axi:; and lang flat pleural spines ( at !east posteriori y) ; hypo::toma with relatively short brim. Pygidium unknown. | |
| G. Henningsmoen 1957 | A Ctenopyge species with : almost parallel-sided glaodla ; S l united across axis, other glabellar furrows Yery short ; straight transverse eye ridges ; centres of palpebral lobes opposite S2 ; intcrocular -cheeks as wide as glabella at eye line ; postocular cheeks almost l . S times as wide as .occipital ring ; postocular sutures oblique ; f ree chceks with long curved and flattened spine, slightly convex posterior lateral margin slightly shorter than almost straight anterior lateral margin. Other parts of shield unknown. | |
| J. A. Weir 1959 | Isocolidae with a single pair of pit-like lateral glabellar furrows confluent with occipital furrow; lateral border furrows conf1uent with preglabellar furrow; facial sutures situated on lateral border. | |
| J. A. Weir 1959 | Shumardia with isolated basal lateral glabellar lobes; glabella overhanging anterior margin. Posterior cephalic border furrows broad (sag.). | |
| J. A. Weir 1959 | Isocolidae with a single pair of pit-like lateral glabellar furrows confluent with occipital furrow; lateral border furrows conf1uent with preglabellar furrow; facial sutures situated on lateral border. | |
| J. A. Weir 1959 | Phylacops wilh oblique laleral glabellar furrows placed far back; one pair of tubercles situated between lateral glabellar and occipital furrows. | |
| Z. Kielan 1960 | Preglabellar field long, equal to one-fourth of cephalic length. Length
of the eye equal to one -sixth that of cephalon . Glabellar tubercle present. Hypostome unknown, thorax imp erfectly known, thoracic pleurae straight (transverse), not produced into spines. Pygidium sub-semicircular. Rhachis equa l to almos t six-eighths of pygidial length . 10-12 rhachial rings, 7-8 pleurae . Almost the whole bod y ornamented with anas tomosing lines. | |
| C. Lochman and C. Hu 1962 | Cranidium equiconvex (sag and tr); glabella convex, tapered forward to a narrowly to broadly rounded front, no lateral furrows, occipital furrow shallow, occipital ring low, frontal area somewhat more than 4 glabellar length, concave, anterior border furrow nearly obsolete; fixigenae downsloping, palpebral area flat, about one-half glabellar width at midline, palpebral lobe small, opposite anterior ? of glabella, furrow obsolete, eye ridge very faint, posterior area triangular, 3 length of occipital ring with bluntly rounded end, broad shallow posterior furrow. Librigena not known.
Pygidium broad-triangular, axis narrow, tapered 6 length to narrow rounded end, dorsal furrow shallow, all others obsolete on outer surface, present on inner, seven axial rings and terminal, pleural platform same width anteriorly as axis, narrowing posteriorly, slightly convex, horizontal, 5-6 narrow pleural segments crossed by narrow pleural furrows, and separated by broader interpleural grooves, pleural furrows end at border furrow but grooves may run onto border, narrow curved border furrow, border very narrow anteriorly, steep, widening posteriorly with moderate slope, outer edge becomes horizontal in sag. line. Outer surface smooth, inner surface punctate. | |
| A.R. Palmer 1962 | Members of Olenaspella with pygidium having 3 or 4 ring furrows behind articulating furrow on axis. Pleural regions with 2 or 3 shallow pleural furrows curved abruptly backward near inner edge of poorly defined narrow border, and extended onto bor- der. Shallow interpleural grooves apparent between first and second pleural segments near outer edge of pleural field. Margin with 2 or 3 pairs of spines. Most specimens with two pairs of spines, outer pair longest; each pair connected to posterior band of first or second pleural segment by low narrow ridge. Third pair of spines, if present, short, adjacent to inner edge of second pair of spines (pi. 5, fig. 3). | |
| A. R. Palmer 1965 | External surface of cranidium, exclusive of furrows and palpebral lobes, covered with coarse granules. Intergranular distance about equal to or greater than granule diameter.
Ocular platform on free cheek has granular ornamentation as on cranidium. Lateral border, exclusive of genal spine, has more concentrated granules. Genal spine has scattered coarse pits. Pygidium has granules on tops of axial segments, pleural ribs, and border. Granules closely-packed on border and at end of axis. Surfaces of molds of most parts have mixture of scattered coarse granules and pits. | |
| A. R. Palmer 1965 | Cranidium having glabella well defined by axial furrows; preglabellar furrow and prominent fossulae generally best defined in members of younger populations. Frontal area has well-defined bri~ and border; brim moderately to strongly convex sagittally, makes distinct angle with border; border flat or slightly convex; sagittal length slightly less ~han length of brim. Fixed cheeks slightly upslop1ng; palpebral lobes moderately defined by shallow palpebral furrow.
Free cheek has moderately broad border and slender genal spine extending posteriorly to about fifth thoracic segment. Lateral border furrow moderately to poorly formed. Thorax composed of 12 or 13 segments. Pleural tips short, sharp, directed posterolaterally. Pygidium has well-defined axis bearing two or three ring furrows behind articulating furrow. Pleural fields moderately convex and have one or two moderately well-defined pleural furrows. Border separated from pleural field principally by change in slope; border narrow behind axis, abruptly expanded laterally, flat or slightly concave, downsloping. External surfaces of cranidium and free cheek, except for furrows and genal spines, generally have distinct shallow pits visible after whitening. External surfaces of pygidium and tips of genal spines have fine granular ornamentation apparent only after whitening. Surfaces of molds of all parts pitted. | |
| P. D. Pack and H. B. Gayle 1971 | A relatively smooth cephalon with well defined border of medium width, rounded genal angles and no genal spines. Glabella characterized by indistinct lat eral and axial furrows. Palpebral lobes short,directed upward and posteriorly, close along the sides of the glabella extending from the postero-lateral parts of the anterior glabellar lobe. Small eyes located laterally below the palpebral lobes slightly behind center of the glabella. Thorax consists of 15 prothoracic segments and 10 or more opisthothoracic segments. Third prothoracic segment is strongly macropleural and the fifteenth segment bears a long axial spine. | |
| P. D. Pack and H. B. Gayle 1971 | Exoskeleton subovate. Glabella subcylindrical, contracted at middle with axial furrows indistinct and lateral furrows absent or indistinct. Well defined border. Genal spines absent; genal angles rounded Thorax consists of 15 prothoracic segments 10 or more opisthothoracic segments. The third prothoracic segment is very strongly macropleural. | |
| E. F. Riek 1973 | Similar to Austroprosbole from which it differs mainly in the more basad origin
of Rs and the more arched basal half of Rs. The fusion of the upper branch of M wi th Rs for a short distance is probably only an abberation. Whole of wing membrane sculptured. Costal space very broad. A 'stridulation' ridge, ventrally, in the costal space towards base, and fore margin decidedly thickened towards base. Node on fore margin very clearly defined. | |
| R. P. Tripp 1976 | Glabella weakly convex; anterior tongue large, protuberant; axial furrows shallow. Free cheek moderately broad; librigenal angle produced backwards in broad-based spine. Surface closely covered with fine striae arranged in Bertillon pattern. | |
| R. A. Henderson 1976 | Aphelaspis characterized by a short frontal area measuring about 40% of the glabellar length, elongate palpebral lobes, and a pygidium which is twice as long as wide. | |
| K. Sdzuy 1978 | Characters of genus. Glabella conical, but diminishing anteriorly only moderately for the genus. Extra-ocular cheeks of moderate width. Ornamentation distinct. Tiny metagenal spines present. | |
| R. A. Fortey and R. M. Owens 1978 | Cephalic doublure and eye position like M. major; glabella apparently lacking terrace lines on frontal lobe, which is more expanded than in other species; cranidial section of lateral border furrow very short; palpebral lobes shortest in genus; hypostoma with fine terrace lines, sparser and less backwardly-bowed than on other species, posterior margin non-acuminate; pygidial doublure broad, with inner margin parallel to posterior pygidial margin. | |
| K. Sdzuy 1978 | Characters of genus. Glabella strongly tapered anteriorly. Extraocular cheeks narrow. Ornamentation fine. Tiny metagenal spines present. | |
| K. Sdzuy 1978 | Glabella parallel sided or slightly narrowing towards front, frontal lobe usually distinctly less wide than lobe behind it. Three pairs of glabellar furrows, the anterior ones very short and directed anteriorly, the posterior ones frequently vaguely confluent. Border very narrow in front of glabella and distinctly angular. Eye lobes thick, widening near glabella, and rather evenly arcuate. Genal spine well advanced and very short. | |
| K. Sdzuy 1978 | Glabella distinctly tapered towards front. Three pairs of moderately deep glabellar furrows, indistinctly bifurcated and the posterior ones nearly confluent. Width between anterior branches of suture slightly less than that between eye lobes. The latter thick and raised above cheeks, with well-defined ocular furrow and parafrontal band. Preglabellar field slightly wider (sag.) than border. Genal spine with broad base, of medium length. Surface with granules tending to a netlike arrangement, sharp subparallel lines on border. | |
| R. A. Fortey and R. M. Owens 1978 | Porterfieldia species with narrow (sag., exsag.) preglabellar field and pitted anterior border furrow. Glabellar and axial furrows subdued. Hypostoma as wide as long, with prominent maculae, posterior margin truncate. Pygidium with three axial rings, carrying sculpture of fine terrace lines. | |
| R. A. Fortey and R. M. Owens 1978 | Microparia species with six thoracic segments, eyes widely separated at anterior
margin. Glabella expan9ing in width to about one-third cephalic length, tapering anteriorly; narrow (trans.) eyes, with correspondingly curved inner margins. Palpebral rims narrow, effaced posteriorly. Semicircular pygidium with short, narrow axis, with single axial ring clearly defined; border faint, especially postaxially. | |
| R. A. Fortey and R. M. Owens 1978 | Closely similar to M. selwynii except that cephalic doublure is broader, so that front
end of eye is closer to paradoublural line; pygidial doublure wider, inner margin not parallel to pygidial margin, so that doublure has its greatest width exsag. | |
| R. Ludvigsen 1979 | A species of Pseudogygites with a long preglabellar field (= 18— 23% of sagittal
cephalic length); a subtriangular pygidium with 12 to 14 distinct pleurae, short and faint pleural furrows evident only on anterior portion of pygidium, well-defined border furrow, and relatively narrow border. Median connective suture is fused. | |
| A. Pribyl and J. Vanek 1980 | Exoskeleton small, slightly convex. Cephalon semielliptical in outline,
more convex in sagittal axis than transversely. Glabella strongly convex, globose, well-defined by deep axial furrows, ovoid in outline. A smooth minute mesial node lies slightly behind the centre of glabella. Anterior pits shallow. Preoccipital part of glabella not continuing posteriorly into a spine. Preglabellar field narrow, weakly convex, sloping forward; anterior border very narrow. Fixigenae narrow (tr.), convex. Alae inconspicuous. Anterior branches of facial sutures almost subparallel, slightly diverging in front of eyes, gently curved outward; posterior branches of facial sutures sigmoidally bent, directed outward and slightly forward behind eyes. Thorax consisting of 6 segments. Axis relatively narrow, tapering backward; it is devoid of spines on the third and fourth segments. Pleurae straight, with very distal fulcrum, and wide submeidan furrow parallel to margin. Pleural extremities bend backwards, ending as short spines. Pygidium semielliptical to subtrangular in outline, elongated transversely with almost unsegmented short axis and smooth lateral regions on which only articulating facet is perceptible. Border broad sloping steeply downward. | |
| R. P. Tripp 1980 | Anterior tongue at base less than 60% width of glabella; length of tongue more than 65% length of palpebral lobe. Field of free cheek faintly striate. Subgenal notch large. | |
| R. A. Fortey 1981 | The diagnosis given in the Treatise (Richter & Richter in Moore, 1959) is extended to include cyclopygids with seven thoracic segments, and relatively large fixigenal areas, with concomitantly smaller eyes. | |
| R. A. Fortey 1981 | Cyclopygidae retaining cephalic median suture. Remnant fixigenal areas larger
than on any later cyclopygid, and eyes not extending as far posteriorly. Well denned glabella which expands forwards; three pairs shallow glabellar furrows. Thorax of seven segments. Pygidium with two axial rings, anterior pleural furrows deep. Surface sculpture of fine lines. | |
| F. Nikolaisen 1983 | Median area rather convex, almost two-thirds as long as wide, with sculptural lines in cordate pattern mesially. Glabellar tongue less than one-fifth as wide as median area. Preglabellar field at axial line two-fifths as long as glabellar tongue. Occipital furrow less than half as wide as median area. | |
| Y. C. Hong 1984 | incertae sedis | |
| P. A. Jell 1985 | Cephalon sub-semicircular; lp furrow well impressed; alae distinct but low; occipital tubercle prominent; radial anastamosing caecal network on cheeks and brim with moderately large pits between caeca on brim gradually decreasing towards glabella; brim prolonged only a very short distance behind posterior of occipital ring, markedly concave posterolateral^ long genal spine at rear of prolongation; girder prominent, extending to tip of prolongation; check roll widened at level of posterior border furrow, extending adaxially almost as far as exsagitlal line of outer edge of eye tubercle. | |
| P. A. Jell 1985 | Alae absent; eye tubercles small, siluated level with anterior of glabella; preglabellar boss present; girder fading out just before tip of prolongation; brim wide anteriorly, tapering rapidly along relatively short prolongation; spine present on tip of prolongation. | |
| P. A. Jell 1985 | Member of Australoharpes without alae; eye tubercles situated behind glabellar anterior; preglabellar boss extremely low; girder fading out just before lip of prolongation; spine present on tip of brim prolongation; cheek roll on prolongation relatively wide; thorax of at least 18 segments. | |
| P. A. Jell and B. Stait 1985 | Member of Scotoharpes with only very faint lp furrow evident on glabella, an occipital node, very faint alae evident as extremely weak depressions or no! evident at all, with short posteriorly directed spine on posterior of prolongation. | |
| P. A. Jell and B. Stait 1985 | Member of Asaphopsoides with parallel-sided to anteriorly tapering glabella broadly rounded anleriorly; palpebral lobes of moderate size, highly arcuate laterally, situated at or behind glabellar midlcngth; pygidium with narrow tapering axis of at least seven rings and a long terminal piece which may represent several more rings; marginal spines small, inconspicuous, widely separated; posterior margin broadly rounded. | |
| P. A. Jell and B. Stait 1985 | Ekeraspid with very low convexity, axial furrow extremely poorly impressed; glabella with rounded anterior, vvaisted at level of palpebral lobes; palpebral lobes small, wide, semicircular, situated behind midlength of cranidium; facial suture diverging forward of palpebral lobes, eoneavc forward of palpebral lobe to widest point, sigmoidal behind palpebral lobe, meeting posterior margin at large angle, as it runs posteroaxially. Free cheek with long genal spine. Hypostome highly convex, with complete rounded posterior margin, with widest point near midlength. Pygidium subtriangular, with long terminal spine decreasing in length with growth; pleural and interpleural furrows weakly impressed on anterior segments. | |
| B. R. Pratt 1988 | A species of Kainella with moderately divergent facial suture. Lateral glabellar furrows absent. Row of pits located forward of anterior border furrow. Pygidium wide, bearing a single pair of backward-curving furrows and two pairs of spines. | |
| R. Ludvigsen et al. 1989 | A genus of Loganellidae with palpebral lobes located opposite Is furrow and some distance from glabella. Anteriorly-rounded glabella is parallel-sided or gently tapered and has firmly
impressed lateral glabellar furrows on testate surfaces. Anterior border is short band. Pygidium is broad and semielliptical in outline with length less than half of width. | |
| R. Ludvigsen et al. 1989 | A genus of Loganellidae with a relatively long anterior border. Palpebral lobe is long and located close to glabella. Anteriorly truncate glabella is parallel-sided or gently tapered; glabellar furrows are shallow on testate surfaces. Broad pygidium is subelliptical in outline with length less than half of width. | |
| R. Ludvigsen et al. 1989 | A genus of Loganellidae with pygidium of subrectangular outline and bilobate posterior margin. Frontal area of cranidium is divided into short preglabeller field and longer, gently convex anterior border. | |
| G. A. Young and R. Ludvigsen 1989 | A species of Zacanthoides with a moderately long, rectangular glabella. Occipital ring extended into a very small spine. Palpebral lobe long, semilunate. Cranidial prosopon finely granulate. Pygidium subovate, wider than long. Axis of three rings and a terminal piece. Pleural furrows very long, shallow. lnterpleural furrows very shallow or absent. Four pairs of short, flattened marginal spines decrease in length posteriorly. Pygidial prosopon of coarse, sparse granules. | |
| G. A. Young and R. Ludvigsen 1989 | A species of Onchocephalites with a strongly effaced cranidium. Lateral glabellar furrows very shallow. Occipital ring semielliptical; median tubercle very small. Axial and preglabellar furrows very shallow, anterior border furrow absent. Palpebral lobe moderately long, narrow. Eye ridge very weak. Cranidial prosopon of dense pits. | |
| G. A. Young and R. Ludvigsen 1989 | A species of Elrathina with a narrow glabella. Is to 3s furrows long and shallow. Occipital ring semielliptical, bearing large median tubercle. Anterior border long, convex, strongly curved. Palpebral lobe very short, narrow. Eye ridge weak or absent. Anterior branches of facial suture convergent, convex. Cranidium smooth; smallest specimens bear granulate prosopon. | |
| R. Ludvigsen et al. 1989 | A species of Memscocoryphe with a semielliptical subtriangular cephalon with blunt genal comers. | |
| R. A. Fortey 1990 | Libristoma having ventral median suture. Only primitive forms retain natant hypostomal condition, the majority of the group being conterminant or impendent. Higher Asaphida also have a distinctive inflated protaspis - the asaphoid protaspis. | |
| R. A. Fortey 1990 | Libristoma having fusiform glabella in the protaspis stage, and natant hypostomal condition attained early in ontogeny. Larval hypostome margin lacking spines. Most species natant also in adult; some secondarily conterminant or impendent. Median occipital tubercle usually present. | |
| R. A. Fortey 1990 | Trilobites having natant hypostomal condition, or with secondarily conterminant or impendent hypostomal condition | |
| P. D. Kruse 1990 | Cranidium subquadrate. Glabella rectangular or truncated conical, with three pairs of glabellar furrows. Occipital ring crescentic. Fixed cheeks approximately as wide as glabella at midlenght. Palpebral lobes of small to moderate size; eye ridges variably developed. Preglabellar field flat to gently convex. Anterior border and preglabellar field subequal in lenght (sag.). Anterior body flat. Facial suture divergent anteriorly from palprebal lobes. Posterolateral limb wider (sag.) than anterior boder. Posterior border furrow distinct.
Thorax with at least 14 segments. Pygidium small, broad; pygidial axis with up to three rings; up to four pleural and interpleural furrows. Pygidial border flat, narrow, poorly defined. | |
| P. D. Kruse 1990 | Cranidium subquadrate. Glabella truncated-conical, with three to four pairs of glabellar furrows; the posterior two pairs may be bifurcate adaxially. Occipital ring crescentic. Fixed cheeks about half or less of glabellar width. Palpebral lobes of small to moderate size. Preglabellar field subequal in lenght or slightly longer (sag.) than anterior boder. Anterior border flat to gently convex, longer (sag.) medially than laterally. Anterior border furrow may include a pair of elongate lateral pits. Facial suture divergent anteriorly from palpebral lobes. Posterolateral limbs wider than anterior border. Posterior border furrow distinct.
Thorax with 12 or more segments. Pygidium small, broad: pygidial axis with about three rings and as many pleural furrows. Axis short, not reaching posterior margin. Posterior margin with shallow median reentrant. | |
| R. A. Fortey 1990 | A paraphyletic libristomate group retaining primitive characters; a combination of: relatively narrow rostral plate present, glabella barrel-shaped to tapering; natant hypostomal condition (possibly some secondarily conterminant); hypostome unspecialized; protaspis with forward-expanding axis reaching far forwards; with few exceptions micro- to heteropygous. | |
| R. A. Fortey 1990 | Ptychopariida with thin dorsal exoskeleton; free cheeks yoked together as single unit connected by narrow (sag.) doublure (rostral plate present only in the most primitive members of Olenus). | |
| R. A. Fortey 1990 | Libristoma having small eyes placed anteriorly relative to glabella; cephalon broadly convex (sag. tr.) with long preglabellar area, and often a brim, with stout genal spines or genal prolongations of the brim; free cheeks narrow (tr.) or lost altogether, fused together; thoracic segments numerous (12 or more) and characteristically narrow (sag.) with long pleurae in contact along their length; pygidium relatively small and often transverse. | |
| S. R. Westrop 1992 | A species of Coosina with firmly impressed axial and preglabellar furrows. Frontal area equally divided into downsloping preglabellar field and convex border by well-defined border furrow. Axis occupies about four-fifths of pygidial length and consists of three axial rings plus terminal piece. Pygidial border gently concave.
Description. Tapered, anteriorly rounded glabella weakly convex and raised only slightly above adjacent areas of cheeks; outlined by firmly impressed axial and preglabellar furrows and occupies about three-quarters of cranidial length. Three shallow lateral glabellar furrows evident on parietal surfaces: 1s and 2s geniculate, whereas 3s convex. Occipital furrow wide and bowed gently forward medially but narrows and curves forward near axial furrows. Palpebral lobe located opposite 2p glabellar lobe and separated from wide, weakly arched cheek by broad, shallow palpebral furrow. Frontal area equally divided into downsloping preglabellar field and convex border by well-defined border furrow. Narrow posterior border separated from cheek by broad, firmly impressed posterior border furrow. Anterior branches of facial suture subparallel before swinging abruptly inward along anterior cranidial margin; posterior branches diverge backwards in a weakly sigmoid curve. Parietal surfaces finely pitted, with the addition of fine, transverse ridges on anterior border. Free cheek lacks genal spine and possesses convex lateral border separated from convex cheek by deep border furrow. External surface cames prosopon of fine granules, with faint caecal markings on cheek; border cames fine transverse ridges. Pygidium sub-semielliptical in outline, with length about threefifths of width, and posesses a very weak median embayment along posterior margin. Axis occupies about four-fifths of pygidial length and consists of three axial rings plus terminal piece. Convex pleural fields crossed by four pairs of broad, well-defined pleural furrows and slope down to a relatively narrow, concave border. External surface cames prosopon of fine granules; internal molds finely punctate. | |
| S. Y. Storozhenko 1992 | Forewing fairly flat, bristles not preserved, venation distinct. Apex of wing rounded. Sc ending on C. Costal area narrow, cut by regular branches of Sc. Subcostal area very wide, divided by regular crossveins. RS diverging from R in anterior third ot wing; R in distal half of wing curved, with comb of branches directed toward frontal margin of wing; apical third of wing containing anastomoses between Rand RS. M dividing into MA and MP just proximally of division of RS into branches. CuA! strongly curved into undulations; CuA2 almost straight. CuP straight and slightly sclerotized; area between CuA and CuP with regular straight crossveins. Anal area narrow and long, and evidently with two anal veins. | |
| S. Y. Storozhenko 1992 | Head large, wider than pronotum; compound eyes and ocelli well developed; mandibles large and pointed at vertices. Pronotum transverse, with full ring of wide rims; its width 1.3-1.9 times exceeding its length. Femora and tibiae lacking spines and spurs. Ovipositor slightly curved upward. Eggs elongated-oval; chorion strongly sclerotized, with sharp longitudinal costulae. Forewing lacking bristles and with almost straight frontal margin; Sc abutting on C in distal third of wing; R without branches; RS diverging from R in anterior third of wing and having five to eight branches. Base of M entirely merged with CuA; MA and MP diverging from CuA separately from each other, simple or branching CuAl simple or with two or three branches; Cu~ and CuP simple; area between CuA and CuP with nonbranching regularly S-curved crossveins. Ai having three and A2 two to four branches. Hindwing like forewing, but always showing distinct anastomosis between RS and MA; CuAl simple. | |
| S. Y. Storozhenko 1992 | Forewing lacking bristles and with almost straight frontal margin. Sc joining C in apical third of wing. RS extending from R in anterior third of wing and having three branches, each of which, or at least one of them, giving rise to short fork near wing apex. Base of M fully merged with CuA; MA and MP diverging independently from CuA; MA with three to five branches, MP bifurcate and desclerotized in middle. CuA! with two branches, CuA2simple. CuP simple and straight. At with three or four branches, Az bifurcate. Crossveins in area between CuA and CuP simple and slanting. Hindwing like forewing, but for considerable extent showing anastomosis between RS and MA. | |
| S. Y. Storozhenko 1992 | Forewing having almost straight frontal margin, with widely rounded posterior margin. Sc abutting on C in distal third of wing; costal area 15 times narrower than greatest width of wing and 4.2 times narrower than subcostal area. Radial area clearly narrower than subcostal, and intersected by simple crossveins. RS with five branches. MA bifurcate, MP simple. CuA! bifurcate; area between CuA and M divided by simple straight or slightly curved crossveins, and short false vein near posterior margin of wing. Area between CuAl and CuA2 having thick, short crossvein at point of bending of CuAl and faint crossveins at posterior margin of wing, forming double row of cells. | |
| A. R. Palmer and L. N. Repina 1993 | Holmiinae with intergenal spines not ap-
parent. Glabella expanded anteriorly; L4 with distinct lateral projections situated in front of ocular lobe and segregated by longitudinal furrow. Posterior tips of ocular lobes opposite Ll. Thorax narrow, with pleural spines short but not constricted at base; each segment with axial nodes; number of segments not known. | |
| A. R. Palmer and L. N. Repina 1993 | Archaeaspididae with glabella slightly ta- pered forward, L4 slightly conical. Interocular area wider than extraocular area opposite midlength of ocular lobe. Posterior tips of ocular lobes approximately opposite oc- cipital furrow. Genal spines short, broad-based. Border broad, convex, well defined; sagittal length greater than length of preglabellar field. Exoskeletal surface strongly reticulate. | |
| A. R. Palmer and L. N. Repina 1993 | Bristohinae with posterior margin of cephalon strongly and evenly curved back distally; genal spines short, at posterolateral corners, situated well behind level of occipital ring. Intergenal spine absent. Preglabellar field short. Width of interocular area about half width of glabella at L2. Posterior tips of ocular lobes opposite Ll. | |
| A. R. Palmer and L. N. Repina 1993 | Biceratopsinae with posterior margin of cephalon nearly straight. Genal angles broadly rounded; genalandintergenalspinesabsent.Preglabellarfield present; sagittal length equal to or less than that of border. Ocular lobes prominent, close to glabella; posterior tips opposite SI, elevated above top of glabella. Opisthothorax of at least 11 segments. | |
| A. R. Palmer and L. N. Repina 1993 | Bristoliinae with posterior margin of cephalon nearly straight or deflected anterolaterally distal to intergenal spine or intergenal angle. Genal spine origi- nates opposite or posterior to Ll. In tergenal spine present on some species. L4 prominent, subglobular, may overlap border in dorsal view. Preglabellar field absent. Posterior tips of ocular lobes approximately opposite occipital furrow. | |
| A. R. Palmer and L. N. Repina 1993 | Neltneriidae with preglabellar field present, its sagittal length about equal to length of border. Interocular area poorly defined, width more than half width of extraocular area. Posterior tips of ocular lobes opposite occipital furrow. Thorax with 18 segments: pleurai regions of each segment narrower than axis; no macropleural segments. | |
| A. R. Palmer and L. N. Repina 1993 | Archaeaspididae with glabellar outline subcylindrical; anterior end of glabella strongly rounded. Sagittal length of preglabellar field about equal to that of border. Width of interocular area less than width of extraocular area opposite midlength of ocular lobe. Ocular lobes raised above level of glabella. | |
| A. R. Palmer and L. N. Repina 1993 | Bristollinae with cephalon subpentagonal to subquadrate in outline. Genal spine originating opposite or anterior to L2. Intergenal spine absent. Preglabellar field absent. Posterior tips of ocular lobes opposite or anterior to Ll. Fifteenth thoracic segment with long axial spine. Opisthothorax of at least 17 segments.
| |
| A. R. Palmer and L. N. Repina 1993 | Nevadiidae with glabella slightly tapered forward. Ocular lobes small, posterior tips about opposite SI or anterior part of Li. Width of interocular area equal to or greater than that of extraocular area. Thorax of 17 or 18 segments, maintaining width or widening slightly backward to eighth segment, then tapering posteriorly; posterior segment may be fused with anterior part of simple pygidium. Pleural spines short; pleural regions only slightly wider than axis. | |
| A. R. Palmer and L. N. Repina 1993 | Callaviinae with intergenal spines well developed, adjacent to genal spines. Intergenal ridge usually present. | |
| A. R. Palmer and L. N. Repina 1993 | Daguinaspididae with anterior margin of cephalon gently curved. Glabella moderately to strongly tapered forward. Width of interocular area opposite tip of ocular lobe nearly equal to basal glabellar width; width of extraocular area about equal to width of ocular lobe. Sag- ittal length of frontal area greater than that of occipital ring. Intergenal ridge well developed. | |
| A. R. Palmer and L. N. Repina 1993 | Daguinaspidinae with anterior margin of cephalon bluntly pointed. Width of interocular area less than one-half basal glabell-ar width; width of extraocular area about equal to width of ocular lobe. Sagittal length of frontal area greater than that of occipital ring. Intergenal ridge well developed. Thorax of 16 (17?) segments; pleura about equal in width to axial lobe, unmodified. Pygidium small, subquadrate. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidinae with posterior margin of
cephalon nearly straight; small intergenal spine on poste- rior border approximately midway between axial furrow and genal spine. Glabella slightly tapered forward. Ocular lobes directed posterolaterally from junction with L4. Width of interocular area equal to or greater than width of ex- traocular area opposite midlength of ocular lobe. Occipi- tal ring simple. Thorax narrow; axis wider than pleural regions; third segment macropleural. External surface strongly pitted. | |
| A. R. Palmer and L. N. Repina 1993 | -Archaeaspididae with glabella slightly ta- pered forward; anterior end bluntly rounded. Preglabellar field short, crossed by plectrum. Posterior tips of ocular lobes opposite Ll. Width of interocular area about half or less width of extraocular area opposite midlength of ocu- lar lobe. Si and S2 usually continuous across glabella. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidinae with posterior margin of cephalon straight or gently curved forward distally. Intergenal spine not developed, but intergenal ridge inter- secting posterior border closer to glabella than to genal spine. Glabella slightly tapered forward. Ocular lobes directed posterolaterally from junction with L4. Width of interocular area half or less width of extraocular area. Thorax of 21 segments; 17 in prothorax, 4 in opisthothorax. Opisthothorax not strongly differentiated. Pleural regions, excluding spines, about same width as axis. Third thoracic segment macropleural. Pygidium small, subquadrate. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellinae with external surface smooth. Genal spines strongly advanced; genal spine angle anterior to S2. Intergenal angle nearly a right angle. Preglabellar field absent. Posterior tips of ocular lobes opposite ante- rior part of LI. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellidae with posterior margin of cephalon curved forward. Intergenal angle variably developed, situated slightly distal to midlength of posterior margin. Genal spine originating opposite or anterior to Si. Glabella bullet-shaped. Ocular lobes close to glabella, tips opposite or posterior to occipital furrow. Thorax with at least 17 segments; width of thoracic axis greater than width of inner parts of pleural region; third thoracic segment not modified; fifteenth segment with long axial spine. Opisthothorax not clearly differentiated. Pygidum elongate, sides convergent posteriorly, end bluntly pointed. | |
| A. R. Palmer and L. N. Repina 1993 | Holmiinae with intergenal spine generally well developed at or slightly proximal to midlength of posterior cephalic margin. Glabella expanded anteriorly. Posterior tips of ocular lobes opposite or posterior to Li. Thorax narrow, with 16 or 17 segments; pleural spines thornlike; each segment with axial spines. Pygidium small, subquadrate; posterior margin nearly straight transversely. | |
| A. R. Palmer and L. N. Repina 1993 | Holmiinae with outline of cephalon subpentagonal; genal spine prominent, originating oppo- site or anterior to S3; short intergenal spine or node lo- cated at or on axial side of intergenal angle. Glabella expanded anteriorly. Posterior tips of ocular lobes opposite or posterior to Li. Pygidium large, wider than long, of at least 3 segments; anterior two segments extended into short border spines. | |
| A. R. Palmer and L. N. Repina 1993 | Judomiidae with glabella barely touching border or with preglabellar field present. Border well defined with sagittal length greater than that of preglabellar field (when present) and about equal to or slightly greater than sagittal length of occipital ring. Posterior tips of ocular lobes opposite occipital ring. Thorax with 15 to 17 thoracic segments; pleural furrows weak or absent. Opisthotliorax not developed. Pleural spines long; trans- verse length of thoracic pleurae more than twice width of axis. Pygidium elongate, with posterior median notch; last two thoracic segments may be fused with pygidium in axial region only. | |
| A. R. Palmer and L. N. Repina 1993 | Judomiidae with cephalon strongly arched transversely. Glabella reaching nearly to anterior margin. Border poorly differentiated. Ocular lobes adjacent to glabella. Interocular area poorly differentiated from ocular lobe. Posterior tips of ocular lobes reaching to posterior margin of cephalon. | |
| A. R. Palmer and L. N. Repina 1993 | Callaviinae with posterior margin of cephalon nearly straight or deflected slightly forward distal to intergenal swellings, which are located closer to genal angle than to axial furrows | |
| A. R. Palmer and L. N. Repina 1993 | Laudoniinae with cephalon subquadrate to subpentagonal in outline. Genal spine originating opposite or anterior to L3. Procranidial spines not developed in adult. Intergenal ridge distinct. Preglabellar field absent or shorter than border. Posterior tips of ocular lobes opposite or anterior to Li. Thorax of about 20 segments. External sculpture reticulate.
| |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidinae with posterior margin of cephalon directed anterolaterally distal to intergenal angle; intergenal angle closer to axial furrow than to genal spine. Glabella slightly tapered forward. Interocular area narrow; width less than half width of extraocular area. Poste- rior tips of ocular lobes about opposite occipital furrow. Externalsurface reticulate. | |
| A. R. Palmer and L. N. Repina 1993 | Species of Olenellus with posterior margin of
cephalon angled forward distal to intergenal angle. Preglabellar field absent or very short, with sagittal length generally equal to or less than that of anterior border. Posterior tips of ocular lobes directed nearly straight pos- teriorly, situated opposite or anterior to medial part of LI. Interocular area may be extended posteriorly as low, broad ridge. Most species with transverse width of inner part of macropleural third segment, exclusive of spine, less than 1.5 times width of axis. Pygidium, known only for type species, has two pairs of short marginal spines. | |
| A. R. Palmer and L. N. Repina 1993 | Olencilinae with posterior margin of
cephalon nearly straight. Parts of external surface may be reticulate. Intergenal swelling distal to midlength of pos- terior margin. Preglabellar field absent or sagittal length less than that of border. S3 deep, continuous across glabel- la. Occipital spine may be present. Ocular furrow deep; oilier band of ocular lobe narrower than inner band. Third thoracic segment generally only weakly macropleural. | |
| A. R. Palmer and L. N. Repina 1993 | Neltneriidae with preglabellar field absent. Interocular area poorly defined, width less than half that of extraocular area. Posterior tips of ocular lobes opposite L l. Thorax with 17 segments. Pleural spines short; width of pleural regions about same as width of axis; eleventh segment macropleural; twelfth to seventeenth segments decreasing in width rapidly and progressively backward. Pygidium small, subquadrate, possibly of 2 or 3 segments. | |
| A. R. Palmer and L. N. Repina 1993 | Bristohinae w ith posterior m argin of cephalon directed slightly posterolaterally to intergenal spine or intergenal swelling that is situated near slightly advanced genal spines. Preglabellar field short. Width of interocular area approximately half or more width of gla- bella at L2. Posterior tips of ocular lobes opposite Li. Third thoracic segment macropleural, with extremely long pleural spines; tips posterior to end of thorax. Prothorax of 13 segments; opisthothorax of at least 17 segments. Fifteenth segment lacks strong axial spine. | |
| A. R. Palmer and L. N. Repina 1993 | Nevadiidae with glabella distinctly tapered forward, sides slightly concave so that width is narrowest opposite L2 or S2. Preglabellar field short (sag.) or absent. Basal glabellar width about equal to width of extraocular area opposite midlength of ocular lobe. Posterior tips of ocular lobes opposite Ll. Width of interocular area more than one-fourth width of extraocular area opposite midlength of ocular lobe. Thorax of 17 to 23 segments. Pleural spines long, falcate. Opisthothorax not clearly differentiated. Pygidium small, subquad rate. | |
| A. R. Palmer and L. N. Repina 1993 | Nevadiidae with width of interocular area less than one-fourth width of extraoctihu- area. Preglabellar- field present. Thorax of 27 segments. Pleural spines long. Transverse length of pleurae at least twice width of axis. Transition to opistluoliorax between segments 15 and 18. Pygidium small, subquadrate. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellinae with external surface generally smooth or with faint Bertillion markings, rarely granular or reticulate. Occipital ring smooth or with small node near posterior margin; occipital spine rare. Genal spine angle opposite or posterior to Ll. Genal spine slender. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidinae with posterior margin of cephalon nearly straight or slightly backswept. Glabella moderately tapered forward. Plectrum strongly developed. Ocular lobes directed posterolaterally from junction with
L4. Width of interocular area slightly more than half width of extraocular area opposite midlength of ocular lobe. Posterior tips of ocular lobes opposite occipital ring. | |
| A. R. Palmer and L. N. Repina 1993 | Judomiidae with glabella weakly tapered forward to nearly parallel-sided. Preglabellar field present. Posterior tips of ocular lobes opposite occipital ring. Interocular area distinctly developed. | |
| A. R. Palmer and L. N. Repina 1993 | Biceratopsinae with posterior margin of cephalon nearly straight. Glabella extended to or nearly to border furrow. Posterior tips of ocular lobes approximately opposite position of Sl. Genal spines at posterolateral corners or slightly advanced, strongly inflated. Opisthothorax of at least 10 segments. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidinae with posterior margin of cephalon nearly straight. Glabella gently tapered forward. Occipital ring with prominent axial spine. Sagittal length of preglabellar field greater than length of border. Width of interocular area slightly more than half width of ex- traocular area. Ocular lobes directed posterolaterally from junction with L4. Posterior tips of ocular lobes about opposite occipital furrow. External surface coarsely pitted, grading to reticulate. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellinid(?) with posterior margin of cephalon nearly straight. Glabella parallel-sided, bluntly rounded anteriorly; occipital ring unusually short. Pal- pebral lobes short, prominent, located about midway be- tween axial and lateral border furrows, and connected to posterior part of L4 by narrow ocular ridge; posterior tips of ocular lobes opposite L2. Preglabellar field longer
(sag.) than anterior border. Genal spines well developed; intergenal spines not indicated. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidinae with posterior margin of cephalon straight or slightly backswept. Ocular lobe directed only slightly posterolaterally at junction with L4. Glabella tapered forward, but area opposite L3 and L4 merging laterally with ocular lobes. Interocular area wider than extraocular area opposite midlength of ocular lobe. Posterior tips of ocular lobes opposite Ll . Cephalic bor- der concave in sagittal profile. Lateral and posterior border furrows not connected across base of genal spine. External surface coarsely pitted, grading to reticulate. | |
| A. R. Palmer and L. N. Repina 1993 | Nevadiidae with border furrows weakly developed. Cephalon strongly convex transversely; posterior margin strongly curved backward. Glabella slightly tapered forward. Basal glabellar width about equal to wi(ltll of extraocular area. Lateral parts of poorly differentiated border nearly vertical. Ocular lobes poorly differentiated from interocular area; posterior tips about opposite occipital furrow. | |
| A. R. Palmer and L. N. Repina 1993 | Holmiinae with posterior margin of cephalon nearly straight; intergenal spines not developed. Glabella with prominent axial spine on anterior margin of occipital ring and encroaching on occipital furrow. Posterior tips of ocular lobes opposite or posterior to LI. Thorax narrow; pleural spines short, not constricted at base. Pygidium small, subquadrate. | |
| A. R. Palmer and L. N. Repina 1993 | Archaeaspididae with glabella slightly ta- pered forward, sides slightly concave; anterior end strongly rounded. Posterior tips of ocular lobes about opposite inidlength of 1,1. Width of interocular area about equal to width of extraocular area. Sagittal length of border about equal to that of preglabellar field. | |
| A. R. Palmer and L. N. Repina 1993 | Judomiidae with L4 slightly expanded in front of ocular lobes. Border narrow, its sagittal length less than sagittal length of occipital ring. Sagittal length of preglabellar field approximately equal to that of border. Posterior tip of ocular lobe opposite posterior part of occipital ring. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellidae with L4 enlarged. Posterior margin of cephalon straight or curved backward towards base of genal spine. Intergenal spine absent. Preglabellar field absent. Posterior tips of ocular lobes opposite or anterior to occipital furrow. Thorax of 17 segments, not divided into prothorax and opisthothorax; third segment unmodified; fifteenth segment bearing long axial spine. Pygidium small, subquadrate, with prominent median notch. External sculpture reticulate; polygons may have central granule. On well-preserved specimens, boundaries of polygons marked by rows of perforations on underside of exoskeleton. | |
| A. R. Palmer and L. N. Repina 1993 | Daghinaspidinae with anterior margin of cephalon gently curved. Glabella gently tapered forward. Extraocular area steeply downsloping. Width of interocular area more than half basal glabellar width. Frontal area short; sagittal length about equal to that of occipital ring. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellidae with glabellar furrows weakly to moderately defined. Transverse width of anterior part of LI equal to or only slightly less than transverse width of occipital ring. L4 slightly to moderately expanded anteriorly. Posterior tips of ocular lobes ranging from opposite L2 to opposite posterior part of occipital ring. Posterior margin of cephalon nearly straight or slightly deflected forward distal to intergenal spine or swelling. Intergenal spines or distinct intergenal angle usually present. Prothorax, where known, with axis narrower than inner part of pleurae. Third thoracic segment weakly to moderately macropleural, with pleural spine not greatly elongated. Fifteenth thoracic segment with long axial spine. Opisthothorax with variable number of uniformly small segments, each bearing narrow pleurae. Pygidium small, subquadrate, with 1 or 2 pairs of short marginal spines. | |
| A. R. Palmer and L. N. Repina 1993 | Olenelloidea with width of interocular area generally half or less that of extraocular area. Third thoracic segment slightly to strongly macropleural. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellina with L4 usually enlarged and glabella narrowest at L2 or SI. L3 usually modified distally, posterolateral part bending backward at the expense of L2. Ocular lobe connected only to posterolateral part of L4. | |
| A. R. Palmer and L. N. Repina 1993 | Redlichiida lacking dorsal cephalic sutures. | |
| A. R. Palmer and L. N. Repina 1993 | Fallota.spidoidea with glabella weakly to strongly tapered forward in dorsal view. L4 elongate. Ocular lobe connected only to posterior part of L4. Interocular area narrow. Posterior tips of ocular lobes op- posite or anterior to occipital furrow. Width of interocular area variable. Thorax of 15 to 27 segments, without development of macropleurae. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidoidea with projection of anterior margin of ocular lobe more or less tangent to anterior end of IA or continuous as parafrontal band in front of L4. Glabella slightly to moderately tapered forward. Length of L4 less than one-third length of glabella, exclusive of occipital ring. Interocular area ranging from significantly wider to slightly narrower than extraocular area.
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| A. R. Palmer and L. N. Repina 1993 | Olenelloidea with width of interocular area more than half width of extraocular area. Third thoracic segment generally unmodified; prothorax and opisthothorax not differentiated. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidoidea with anterior part of L4 anterior to line tangent to anterolateral margin of ocular lobe. Parafrontal band usually present, continuing to or nearly to anterior end of L4. Width of interocular area approximately same as extraocular area. Genal spines usually present. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidoidea with posterior margin of cephalon straight or curved backward distally. Glabella typically parallel-sided; glabellar furrows weakly to moder- ately developed. L4 elongate. Ocular lobe connected only to posterior part of L4. Interocular area very narrow or absent; width equal to or less than width of ocular lobe. Inner margin of ocular lobe typically undifferentiated or only weakly differentiated from interocular area. Posterior tips of ocular lobes posterior to occipital furrow. Intergenal spines not apparent. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspidoidea with posterior margin of cephalon deflected slightly forward distally. Glabella parallel-sided, bluntly rounded anteriorly. L4 elongate. Ocular lobe connected only to posterior part of L4. Ocu- lar lobes close to glabella; interocular area poorly differen- tiated. Thorax has 17 segments. | |
| A. R. Palmer and L. N. Repina 1993 | Species of Olenellus with posterior margin of cephalon nearly straight or only slightly angled forward distal to position of intergenal spines. Intergenal spine or swelling generally closer to genal spine than to axial fur- row. Sagittal length of preglabellar field characteristically greater than twice sagittal length of anterior border. Posterior tips of palpebral lobes convergent toward glabella, opposite or posterior to posterior part of Li. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellina with glabellar outline typically cylindrical or slightly conical in dorsal view. L3 simple, form similar to L2. | |
| A. R. Palmer and L. N. Repina 1993 | lohniidae with glabella expanded forward. Cephalic border generally convex in cross-section. Inner part of pleural region of thorax narrower than axis. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellidae with L4 enlarged. Posterior margin of cephalon straight or curved backward towards base of genal spine. Intergenal spine absent. Preglabellar field absent. Posterior tips of ocular lobes opposite or anterior to occipital furrow. Thorax of 17 segments, not divided into prothorax and opisthothorax; third segment unmodified; fifteenth segment bearing long axial spine. Pygidium small, subquadrate, with prominent median notch. External sculpture reticulate; polygons may have central granule. On well-preserved specimens, boundaries of polygons marked by rows of perforations on underside of exoskeleton. | |
| A. R. Palmer and L. N. Repina 1993 | Holmiinae with posterior margin of cephalon straight or slightly angled forward distal to intergenal swelling. Intergenal spine absent. Posterior tips of ocular lobes opposite or slightly anterior to occipital furrow. Sagittal length of preglabellar field equal to or slightly greater than that of anterior border. Thorax mod- erately wide, consisting of 18 segments, pleural regions of adults lacking macropleurae. Fifteenth through eighteenth segments with strong axial spines. Pygidium small, subquadrate, with single pair of short an terolateral spines. External surface with some reticulate areas. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellidae with glabella usually strongly constricted at SI or L2; transverse width of anterior part of Li usually distinctly narrower than occipital ring; glabellar furrows generally well developed. Preglabellar field shorter than border or absent. Posterior tips of ocular lobes oppo- site or anterior to Ll. Third thoracic segment with inner part of pleural region strongly expanded distally; pleural spine may be extended posterior to end of thorax. Opisthothorax well developed. Pygidium not known. | |
| A. R. Palmer and L. N. Repina 1993 | Latidoniinae with ccphalon subliexagonal in outline; prominent procranidial, genal, or intergenal spines at each angle of the hexagon. Intergenal spines close to glabella. L4 subglobular. Preglabellar field ab- sent. Width of narrow interocular area about equal to width of equally narrow ex traocular area opposite midlength of ocular lobe. Posterior tips of ocular lobes opposite 1.2. Thorax of nine segments; axis wider than inner part of pleural region exclusive of spines; third and sixth thoracic segments macropleural. | |
| A. R. Palmer and L. N. Repina 1993 | Species of Olenellus with genal spines at pos-
terolateral cephalic corners or slightly advanced to posi- tion opposite Ll. Intergenal spines, if present, small, close to genal spines, and directed slightly posterolater-ally. Preglabellar field absent or very short, sagittal length usu- ally less than twice sagittal length of anterior border. Ocu- lar lobes curved; posterior tips convergent toward glabella and situated opposite or posterior to posterior part of Ll. S2 isolated from axial furrow, usually present as distinct transverse slits. | |
| A. R. Palmer and L. N. Repina 1993 | Species of Olerzelluswith posterior margin of cephalon directed slightly anterolaterally distal to position of intergenal spine. Intergenal spines intermediate between genal spine and axial furrow or closer to axial furrow than to genal spine. Sagittal length of preglabellar field equal to or slightly greater than that of anterior border. Posterior tips of ocular lobes directed nearly straight backward, situated opposite or anterior to midlength of Li. | |
| A. R. Palmer and L. N. Repina 1993 | Species of Olenellus with posterior margin of cephalon directed slightly forward distal to position of intergenal spine or intergenal angle. Sagittal length of preglabellar field equal to or slightly greater than sagittal length of border. Intergenal spine or angle about midway between genal spine and axial furrow or closer to axial furrow. Posterior tips of ocular lobes convergent towards glabella, situated opposite or posterior to occipital furrow. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellidae with glabella narrowest at midlength; glabellar furrows very poorly developed; trans- verse width of LI less than transverse width of occipital ring. Ocular lobes close to glabella; posterior tips opposite or anterior to Li. Genalspines absent or opposite midlength of ocular lobes. Intergenal spines absent; intergenal angle may be developed. Thorax divided into prothorax and opistbothorax; third segment strongly expanded distally and pleural spine extremely long, tips posterior to body; 15th segment with long axial spine. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellidae with posterior margin of cephalon curved forward. Intergenal angle variably developed, situated slightly distal to midlength of posterior margin. Genal spine originating opposite or anterior to Si. Glabella bullet-shaped. Ocular lobes close to glabella, tips opposite or posterior to occipital furrow. Thorax with at least 17 segments; width of thoracic axis greater than width of inner parts of pleural region; third thoracic segment not modified; fifteenth segment with long axial spine. Opisthothorax not clearly differentiated. Pygidum elongate, sides convergent posteriorly, end bluntly pointed. | |
| A. R. Palmer and L. N. Repina 1993 | Olenellidae with cephalon subquadrate to subhexagonal in outline. Genal spine strongly advanced, originating anterior to Si. Intergenal spine strongly devel- oped in adult at posterolateral corner of cephalon. Trans- verse width of anterior part of Li nearly equal to occipital ring. Thorax has third segment weakly macropleural; fifteenth segment lacks axial spine. Opisthothorax not clearly differentiated. Pygidium small, bilobate. | |
| A. R. Palmer and L. N. Repina 1993 | Holmiidae with glabella subcylindrical in outline or slightly expanded anteriorly; L4 extended onto inner part of border. Preglabellar field absent. Border broad. Posterior tips of ocular lobes opposite or posterior to midlength of Li. Thorax broad, with 16 to 18 segments; not clearly differentiated into prothorax and opisthothorax. Pleural regions lacking macropleurae. Pleural spines elongate. Pygidium small, subquadrate. | |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspididae with genal spines present. Interocular area slightly wider to slightly narrower than extraocular area.
| |
| A. R. Palmer and L. N. Repina 1993 | Fallotaspididae with interocular area significantly wider than extraocular area. Genal spines absent. | |
| F. A. Sundberg 1994 | Dorypygidae with cranidium with medially inflated to subquadrate glabellae, effaced to nearly effaced lateral glabellar furrows; fixigenae convex, slightly downsloping. Pygidium with well-defined axis generally extend- ing to border, 3-5 ring furrows; pleural fields crossed by 3-5 generally deep pleural furrows; interpleural furrows shallow, exsagittal width of anterior pleu- ral bands increases laterally; width of posterior pleural bands uniform; border moderately well defined, generally 3-4 pairs of spines of equal length, a posterior pair of greater length, and most pos- terior pair very short, unevenly spaced along mar- gin. | |
| F. A. Sundberg 1994 | Dorypygidae with cranidium with subquadrate to slightly expanded glabella wich shallow lateral glabellar furrows; fixi- genae convex, slightly downsloping. Pygidium with well-defined axis general ly extending to border, 5- 7 ring furrows; pleural fields crossed by 5 - 6 gen- erally deep pleural and interpleural furrows, exsag- mal width of anterior pleural bands increases laterally; width of posterior pleural bands uniform; border poorly defined, generally 4 or more pairs of spines of equal length, evenly spaced along margin. | |
| F. A. Sundberg 1994 | Oryctocephalinae with subtrape- zoidal cranidium with glabella with medial sulcus, furrow pits connected with axial furrows and not connected with transglabellar furrow, and undu- lating axial furrows; posterior border furrow connecting with Fl furrows, occipital furrow con- necting with posterior border. Pygidium with moderately deep pleural and interpleural furrows, 5 pairs of spines. | |
| F. A. Sundberg 1994 | Zacanthoididae with cranidia with expanding glabella, preglabellar field absent, anterior border present, long, narrow, mod- erately curved palpebral lobes terminating opposite preoccipital lobe, facial sutures widely divergent anterior of palpebral lobes. Thoracic axial lobe not as wide (trans.) as pleural lobes exclusive of spines. Small,transverselyelongatepygidiumwithaxiallobe 40% or greater of pygidial width, pleural furrows moderate depth, border furrow indistinct, 4-6 pairs of posteriorly directed, rounded spines on posterior margin. | |
| F. A. Sundberg 1994 | Tonkinellinae with lateral glabellar furrows connecting with the axial furrows at the F2 and F3 positions. | |
| F. A. Sundberg 1994 | DIAGNOSIS.Micropygous Doli- chometopidae with cranidia with anteriorly ex- panding glabella, shallow axial furrows, tapering posterior area of fixigenae, and relatively shorter palpebral lobes. Posterior thoracic segment not projecting adjacent to pygidium. Small, transversely elongate pygidium with poorly defined border furrow, less distinct interpleural furrows, and either complete margin or incipient spines. | |
| F. A. Sundberg 1994 | \Zacanthoididae with cranidia with slightly expanding glabella, preglabellar field and anterior border present, long, narrow, strongly curved palpebral lobes terminating opposite preoccipital lobes, facial sutures widely divergent anterior of palpebral lobes. Thoracic axial lobe as wide or wider than pleural lobes exclusive of spines. Pygid- ium with axial lobe 33% or greater of pygidial width, pleural furrows shallow, border furrow indistinct, 3 or more pairs of posteriorly directed, flat spines on posterior margin. | |
| F. A. Sundberg 1994 | Large, micropvgous Altiocculinae with cranidium with typically moderate length (20- 30% glabellar length), posteriorly set, elevated pal- pebral lobes; strongly downsloping anterior area of fixigena; very strongly tapered glabella; median swelling posterior of anterior border furrow; no deep pits in anterior border furrow; secondary preoccipital lobes; narrow posterior area of fixi- genae (15-30% glabellar length); coarse granular
ornamentation. Pygidium subelliptical; transversely elongate; narrow, level border; moderately to moderately weak convex anterior pleural bands; strongly convex posterior pleural bands. | |
| F. A. Sundberg 1994 | Small Ptychopariida with subtrap- ezoid cranidium with glabella elevated well above fixigena, moderately convex transversely; deep, short lateral glabellar furrows; rounded frontal lobe; con- vergent, deep axial furrows; downsloping, moder- ately wide fixigenae (60-65% glabellar width); nar- row preglabellar field; convex, slightly curved, sagittally narrow, tapering anterior border; short palpebral lobes; weak, thin ocular ridges; short pos- terior area of fixigenae. | |
| F. A. Sundberg 1994 | Large, micropygous Ptychopariida with subrectangular to subpentagonal cranidium with strongly to very strongly tapered glabella; secondary basal glabellar lobe weakly developed; gla bella width approximately 25% cranidial width; short to moderate length, medially set, non-elevat- ed palpebral lobes; slightly to very strongly down- sloping anterior area of fixigena and preglabellar field; median swelling, when present, posterior of anterior border furrow; moderately wide posterior area of fixigenae, rounded terminations; smooth or fine granules. | |
| F. A. Sundberg 1994 | Small Ptychopariida with subrec- tangular cranidium with a relatively large glabella elevated well above fixigena; moderately convex transversely; parallel to slightly convergent, mod- erate depth axial furrows; shallow to moderately shallow lateral glabellar furrows; rounded frontal lobe; downsloping, moderately narrow fixigenae; narrow or absent preglabellar field; convex, mod- erately to moderately strong curved, transversely wide, sagittally narrow, slightly tapering anterior border; moderate to short length palpebral lobes; moderate length posterior area of fixigenae. | |
| F. A. Sundberg 1994 | Ehmaniellinae with subrectangular cranidium with divergent facial su- tures anterior of palpebral lobes; moderate to long palpebral lobes (35-50% glabellar length); well-de- fined anterior border, narrow to moderate width (10-20% cranidial length), moderately to slightly tapered; preglabellar field sagittally narrow to moderately wide (30-60% frontal area length); moder- ately tapered glabella; bluntly rounded frontal lobe; exsagittally narrow posterior area of fixigenae (20- 35% glabellar length). Pygidium suboval to subrect- angular to alate with blunt axis; 3-4 axial rings; moderate depth pleural furrows; moderately con- vex posterior pleural bands; moderately convex to weak anterior pleural bands. | |
| F. A. Sundberg 1994 | Ehmaniellinae with subrectangular cranidium with divergent facial su- tures anterior of palpebral lobes, moderate to long palpebral lobes (30-45% glabellar length); well-de- fined anterior border, narrow to moderate width (10-20% cranidial length), moderately to strongly tapered; preglabellar field sagittally moderately wide (25-45% frontal area length); moderately to strongly tapered glabella; moderately to strongly rounded frontal lobe; exsagittally narrow posterior area of fixigenae (20-35% glabellar length). Pygidium sub- rectangular to subelliptical to subtriangular with elongate axis; 4-6 axial rings; moderate depth pleu- ral furrows; moderately to strongly convex poste- rior pleural bands; moderate to absent anterior pleural bands. | |
| F. A. Sundberg 1994 | Small, micropygous Ptychopariida with subrectangular cranidium with convergent fa- cial sutures anterior of the palpebral lobes, glabella elevated above fixigena; parallel to slightly conver- gent, moderate depth axial furrows; shallow to ef- faced lateral glabellar furrows; strongly rounded frontal lobe; slightly downsloping, moderately wide fixigenae; narrow preglabellar field; convex, slightly to moderately curved, sagittally narrow, tapering anterior border; short palpebral lobes anterior of glabellar midlength. Free checks lack significant development of genal spines. Triangular pygidium. | |
| F. A. Sundberg 1994 | Asaphiscidae with cranidia with short, tapered glabella, lateral glabellar furrows shallow to absent, frontal glabellar lobe rounded to bluntly rounded, shallow to moderate depth preglabellar and axial furrows, straight to slightly concave axial furrows, moderately strong ocular ridges, long palpebral lobes, posterior area of fixigena exsagittally narrower than occipital ring, fixigena moderately narrow, slightly convex preglabellar field and anterior area of fixigena, wide, moderately to strongly concave anterior border, anterior border furrow typically very faint. Pygidia large, subcircular, rounded anterolateral corners; multisegmented, well-defined, narrow axis reaching to border furrow or extending into border; pleural field well defined, pleural furrows shallow, extend- ing onto border, interpleural furrows very shallow typically not extending onto border, low convexity; wide, slightly to moderately concave border; smooth, evenly rounded lateral and posterior margin. | |
| F. A. Sundberg 1994 | Ehmaniellinae with subrectangu- lar cranidium with divergent facial suture anterior of palpebral lobes; short to moderate length pal- pebral lobes (25-40% glabellar length); well-defined anterior border, narrow (5-10% cranidial length), slightly to very slightly tapered; preglabellar field sagittally wide (55-75% frontal area length); mod- erately to strongly tapered glabella, rounded frontal lobe; exsagittally wide posterior area of fixigenae (35-50% glabellar length). Pygidium alate with blunt axis, 2-3 axial rings; shallow depth pleural furrows; weakly to moderately convex posterior pleural bands; weak anterior pleural bands. | |
| F. A. Sundberg 1994 | Small to moderate size Altioccu- linae with cranidium with large (40-50% glabellar length), medially set, elevated palpebral lobes; mod- erately strong downsloping anterior area of fixigena and preglabellar field; strongly tapered glabella; elongated, narrow, median swelling either within or anterior of anterior border furrow; no deep pits in anterior border furrow; moderate width poste- rior area of fixigenae (35% glabellar length); fine granular ornamentation. Pygidium subelliptical, transversely elongate; tapered, narrow, level border; anterior pleural bands weakly convex; poste- rior pleural bands moderately to weakly convex. | |
| F. A. Sundberg 1994 | Ehmaniellinae with subpentagonal cranidium with divergent facial sutures anterior of palpebral lobes; moderately to long palpebral lobes (35-40% glabellar length); poorly defined anterior border, gently and uniformly convex preglabellar field and anterior border, shallow anterior border furrow, moderate width (15-20% cranidial length), very strongly tapered; preglabellar field sagittally moderately wide (40-60% frontal area length); moderately to strongly tapered glabella; bluntly rounded frontal lobe; exsagittally wide posterior area of fixigenae (25-30% glabellar length). Pygid- ium subelliptical with elongate axis; 4 axial rings; moderate depth pleural furrows; strongly convex posterior pleural bands; without anterior pleural bands. | |
| F. A. Sundberg 1994 | Small, micropygous Ptychopariida with subrectangular cranidium with glabella ele- vated well above fixigena; slightly convergent, mod- erate depth axial furrows; shallow to effaced lateral glabellar furrows; bluntly rounded from lateral lobe; downsloping, moderately narrow to fixigenae; nar- row or absent preglabellar field; convex, moder- ately curved, sagittally narrow, strongly tapering, medially swollen anterior border; short length pal- pebral lobes; short (trans.) posterior area of fixigenae. Librigenae narrow, rounded at genal angle. Pygidium small, alate, elevated axis, 1-2 shallow pleural furrow pairs, poorly defined border. | |
| F. A. Sundberg 1994 | Ehmaniellinae with subpentagonal to trapezoidal cranidium with con- vergent facial sutures anterior of palpebral lobes; short to moderate palpebral lobes (20-35% gla- bellar length); well-defined anterior border, narrow to moderate width (5-20% cranidial length), modrate to strongly tapered; preglabellar field sagittally narrow to moderately wide (15-45% frontal area length); strongly to very strongly tapered glabella; moderately to bluntly rounded frontal lobe; exsag- ittally wide posterior area of fixigenae (35-55% gla- bellar length). Pygidium subcircular to subelliptical with elongate axis; 5 axial rings; moderate depth pleural furrows; moderate to strong posterior pleural bands; weak to absent anterior pleural bands. | |
| F. A. Sundberg 1994 | Ehmaniellinae with subrectangu- lar cranidium with divergent facial sutures anterior of palpebral lobes; moderate to long palpebral lobes (35-45% glabellar length); well-defined anterior border, moderately narrow (10-15% cranidial length), slightly to moderately tapered; preglabellar field sagittally moderately wide (50-55% frontal area length); slightly tapered glabella; moderately round- ed frontal lobe; exsagittally narrow posterior area of fixigenae (20-35% glabellar length). Pygidium subtriangular with elongate axis, 4 axial rings; mod- erate depth pleural furrows; moderately convex anterior and posterior pleural bands. | |
| F. A. Sundberg 1994 | Multisegmented, rni- cropygous Ptychopariida with cranidium with mod- erate length glabella, lateral glabellar furrows shal- low to absent, frontal glabellar lobe truncated to slightly rounded, deep preglabellar furrow, anterior portion of axial furrows very deep, straight to con- cave axial furrows, ocular ridges typically strong, palpebral lobe moderate length, posterior area of fixigenae wader than occipital ring, posterior border furrow distally curved anteriorly, fixigenae mod- erately wade, convex preglabellar field and anterior area of fixigena, anterior border slightly to strongly convex. | |
| R. Wolfart 1994 | Cranidium trapezoidal (sometimes subtriangular) in outline, lenght (sag.) 2/3 of basal width; glabella long (without occipital ring), not quite twice as long as basal glabellar width and narrow lateral glabellar furrows obscured or faintly developed, preglabellar field short, slightly convex, frontal border furrow and border distinct, slightly arched (transv.)
Posterior facial suture diverges rapidly behind palpebral lobes, forming very wide postero-latral limbs of fixigenae; anterior part of facial suture subparallel to convergent, short. Position of palpebral lobes in anterior half of cranidium, unusally far from glabellar midlenght. Eye-ridges obscure to faintly developed. Lenght of palpebral lobes up to 1/20 cephalic lenght. Librigenae with short, stout genal spine. General curvate moderate( to rather strong). Test finely granulose. | |
| R. Wolfart 1994 | The new genus Centonella is a member of the FAmily Dolichometopidae, becaus it has a strongly clavate glabella with the frontal cranidial
limb lacking and the clavation beginning in front of F1, comperatively wide(transv.) fixigenae between rather long palpebral lobes, ocular ridges well developed, three paris of short lateral glabellar furrows and a fourth preoccipital pair crossing the glabellar crest as a rather distinct, though wide and shallow furrow. Position of palpebral lobes about opposite midlenght of glabella, which is extremly inflated. Occipital ring medium in lenght (sag.) bearing a median node or small spine. Free cheeks with short general spines. In well preserved specimens, the back line of the occipital ring is triangular. Pygidium with well defiend border which is not crossed by ribs or furrows. Rachis with three or four rings, pleura with three widely rounded ribs. Posterior margin of pygidium not intended. | |
| F. A. Sundberg 1994 | Dorypygidae with cranidium with medially expanded to anteriorly ex- panded glabellae, nearly effaced lateral glabellar furrows; fixigenae convex, slighrly downsloping. Pygidium wirh well-defined axis generally extend- ingroborder, 3-5 ringfurrows; pleural fieldscrossed by 3-5 generally deep pleural furrows; inrerpleural furrows very shallow ro effaced, borh anterior and posterior pleural bands uniform in widrh; border moderately defined, generally 5-7 pairs of spines of equal length, evenly spaced along margin. | |
| F. A. Sundberg 1994 | Kootenia with pygidium having 6 pairs of moderately thin, short to moderate length spines; 4-5 axial rings; weak to strong axial nodes; moderately shallow pleural fur- rows; and narrow border (5-15% pygidial length). Cranidium with slightly divergent, convex axial furrows, short palpebral lobes (20-25% glabellar length). | |
| J. M. Adrain 1994 | Richterarges ethnikos has a plesiomorphic pattern of nine border spines similar to that of Borealarges, although other aspects of its morphology, particularly cranidial features, make assignment to Richterarges unambiguous. | |
| F. A. Sundberg 1994 | Kootenia with pygidium having 6 pairs of moderately thick, short to moderate length spines; 4 axial rings; no axial nodes; moderately shallow pleural furrows; and wider border (10-15% pygidial length). Cranidium with parallel, concave axial furrows; short palpebral lobes (20-25% glabellar length). | |
| F. A. Sundberg 1994 | Elrathiella with cranidium with elongate glabella (70-80% cranidial length), mod- erately to strongly tapered, strongly rounded fron- tal lobe; anterior border strongly convex, moder- ately wide (55-65% frontal area length) unevenly curved, nearly straight medially, moderately ta- pered; deep anterior border furrow; moderately wide fixigenae (55-60% glabellar width); moderately coarse granular ornamentation. Pygidium subelliptical, moderately coarse granular ornamentation; anterior pleural bands moderately to weakly developed. | |
| F. A. Sundberg 1994 | Pseudoalokistocare with cranidium with median swelling in anterior border furrow; moderately curved, strongly tapered, flat to slightly convex, level anterior border. | |
| F. A. Sundberg 1994 | Dorypyge with pygidium having 5 pairs of spines, anterior 4 pairs of lateral spines increasing in length from anterior to posterior, pos- terior pair of spines very short, 4 axial rings, borderincreasing in width posteriorly. Cranidium smooth, glabella with slightly divergent, slightly concave to straight axial furrows; frontal lobe slightly smaller than greatest glabellar width. | |
| F. A. Sundberg 1994 | Kootenia with pygid-ium having 6 pairs of moderately thin, long spines; 4 axial rings; no axial nodes; moderately deep pleural furrows; and narrow border (5-15% pygidial length). Cranidium with slightly divergenr, concave axial furrows; moderate length palpcbral lobes (30- 35% glabcllar lengrh). | |
| F. A. Sundberg 1994 | Kootenia with pygidium having 5 pairs of rhin, moderate length spines; 4 axial rings; no axial nodes; moderately shallow pleural furrows; and border very poorly defined. Cranidium with parallel, concave axial furrows; shorr co moderare length pa lpebral lobes (15- 30% glabellar length). | |
| F. A. Sundberg 1994 | Same as genus. [Oryctocephalinae with subtrape- zoidal cranidium with glabella with medial sulcus, furrow pits connected with axial furrows and not connected with transglabellar furrow, and undu- lating axial furrows; posterior border furrow con- necting with Fl furrows, occipital furrow connecting with posterior border. Pygidium with moderately deep pleural and interpleural furrows, 5 pairs of spines.] | |
| F. A. Sundberg 1994 | onkinella with up- turned anterior border on cranidium. Pygidium with smooth margin, moderate ro moderately shallow pleural furrows of unifom depth not reaching border, narrow posterior pleural segment, segment width 7-11.5% pygidial width, 3-5 axial rings. | |
| F. A. Sundberg 1994 | .Tonkinella with cra- nidium with strongly posterolateral directed ocular ridges, slightly upturned anterior border, low con- vex frontal lobe, plectrum, and medial sulcus in frontal lobe. Pygidium with smooth margin, mod- erate to moderately shallow pleural furrows of uni- form depth not reaching border, wide posterior pleural segment (13-19% pygidial width), and 5-6 axial rings. | |
| F. A. Sundberg 1994 | Wenkchemnia with cranidium with exsagittally narrow posterior area of fixigena (20- 30% glabellar length), slightly expanding glabella (105-110%), occipital spine or node. Pygidium with converging pleural and interpleural furrows, pygid- ial axial nodes, and pygidial margin with 3 incipient spines. | |
| F. A. Sundberg 1994 | enkchemnia with cranidium with exsagittally narrow posterior area of fixigenae (20- 30% glabellar length), greatly expanded frontal lobe (140-155%), short occipital spine or node. Pygidium with upturned border, posteriorly directed pygidial field furrows, short axis, short spine on L1 axial ring. | |
| F. A. Sundberg 1994 | Parkaspis with cranidium with slightly expanding frontal lobe (110-120%), non- recurved posterior area of fixigenae, convex ante- rior border. Pygidium with short sagittal length (50- 60% width), moderately tapering axis, no axial nodes, short pygidial spines. Smooth exoskeleton. | |
| F. A. Sundberg 1994 | Zacanthoides with cranidium with strongly divergent anterior facial sutures, wide frontal area (10-25% cranidial length) with moderately well-defined plectrum. Pygidium subrectangular with 4 short, flat spines of unequal length, and granules on pleural lobes | |
| F. A. Sundberg 1994 | Zacanthoides divergensis with cranidium with anterior border moderately well defined and not upturned, shallow anterior border furrow, frontal area 18-23% cranidial length. | |
| F. A. Sundberg 1994 | Zacanthoides with cranidium with strongly upturned preglabellar field, anterior border not defined; nearly effaced lateral glabellar furrows; short ocular ridges. Pygidium subrectangular, 4 pairs of short, thin, rounded, unequal length spines pointing toward midline of pygidium. | |
| F. A. Sundberg 1994 | Zacanthoides with cranidium with moderately divergent anterior facial sutures, frontal area 5-10% cranidial length, very shallow anterior border furrow. Pygidium subovoid with 1 pair of moderate length, lateral spines; 1-3 pairs of short to very short, flat, posterior spines | |
| F. A. Sundberg 1994 | Ehmaniella with cranidium with bluntly rounded frontal lobe; fine granular orna- mentation; flat to slightly concave, moderately strongly curved, sightly downsloping, slightly ta- pered, wide anterior border, 55-70% of frontal area length; narrow fixigenae (30-40% glabellar width). Pygidium alate, elongate axis, moderately convex anterior pleural bands. | |
| F. A. Sundberg 1994 | Ehmaniella with cranidium with very bluntly rounded frontal lobe; smooth to very fine granular ornamentation; slightly convex to con- cave, moderately curved, downsloping, moderately tapered, wide anterior border (55-65% of frontal area length); wide fixigenae (50-60% glabellar width). Pygidium alate, elongate axis, moderately convex anterior pleural bands. | |
| F. A. Sundberg 1994 | Ehmaniella fronsplanata with cra- nidium with flat to very slightly convex anterior border, no medial swelling; smooth exoskeleton. | |
| F. A. Sundberg 1994 | Ehmaniella fronsplanata with cranidium with moderately convex anterior border, fine granular ornamentation, no medial swelling; scattered coarse granules on fixigena. | |
| F. A. Sundberg 1994 | Ehmaniella fronsplanata with cranidium with moderately concave anterior border, elongate medial swelling on anterior border, fine granular ornamentation | |
| F. A. Sundberg 1994 | Ehmaniella with cra- nidium with bluntly rounded frontal lobe; fine gran- ular ornamentation; strongly convex, moderately curved, upturned, slightly tapered, narrow anterior border (40-55% of frontal area length); moderately wide fixigenae (40-50% glabellar width). Pygidium subrectangular, blunt axis, weakly to moderately convex anterior pleural bands. | |
| F. A. Sundberg 1994 | Elrathiella with cra- nidium with elongated glabella (70-80% cranidial length), strongly tapered, strongly rounded frontal lobe; anterior border convex, wide (65-75% frontal area length), evenly curved, moderately to strongly tapered; shallow anterior border furrow; moder- ately wide fixigenae (50-60% glabellar width); coarse granular ornamentation. Pygidium subelliptical to subrectangular, smooth; anterior pleural bands moderately to weakly convex. | |
| F. A. Sundberg 1994 | Elrathiella with cranidium with moderately elongate glabella (60-70% cranidial length), strongly tapered, strongly rounded frontal lobe; anterior border flat, moderately wide (55- 65% frontal area length), evenly curved, slightly tapered; shallow anterior border furrow; moderately narrow fixigenae (40-55% glabellar width); very fine granular ornamentation. Pygidium subel- liptical, smooth; anterior pleural bands weakly to moderately convex. | |
| F. A. Sundberg 1994 | Elrathiella with cranidium with elongated glabella (75-80% cranidial length), mod- erately to strongly tapered, sharply rounded frontal lobe; anterior border strongly convex, moderately wide (55-65% frontal area length) unevenly curved, straight medially, moderately tapered; medially shallow to laterally moderate depth anterior border furrow; moderately narrow fixigenae (45-55% glabellar width). Pygidium subrectangular; anterior pleural bands weakly convex to absent. | |
| F. A. Sundberg 1994 | Elrathiella? with cranidium with elongated glabella (70-75% cranidial length), mod- erately tapered, moderately rounded frontal lobe; anterior border flat, moderately wide (55-65% frontal area length) evenly curved, very slightly ta- pered; shallow anterior border furrow; moderately wide fixigenae (50-55% glabellar width); fine granular ornamentation. Pygidium subelliptical, smooth; weakly developed anterior pleural bands. | |
| F. A. Sundberg 1994 | Elrathiella with cranidium with moderately elongate to elongated glabella (60-75% cranidial length), moderately tapered, moderately rounded frontal lobe; anterior border slightly con- vex, wide (65-75% frontal area length) evenly curved, slightly tapered; deep anterior border furrow; narrow fixigenae (30-40% glabellar width); non-granular ornamentation. Pygidium subrectangular, smooth; with anterior pleural bands absent or very weakly developed. | |
| F. A. Sundberg 1994 | Proebmaniella with cranidium with moderately rounded frontal lobe; frontal area moderately narrow (25-30% cranidial length); anterior border narrow (25-35% of frontal area), slightly curved, moderately convex; slightly posterolaterally directed ocular ridges (80-85° to axial trend); moderately wide fixigenae (50-60% glabellar width). Pygidium alate, blunt axis, anterior pleural bands very weak. | |
| F. A. Sundberg 1994 | Proehmaniella with cranidium with anterior border moderately wide (40-45% frontal area length), moderately curved, strongly convex; moderately posterolaterally directed ocular ridges (65-70° to axial trend); very fine granular ornamentation. Pygidium alate, blunt axis, with 2 axial rings; anterior pleural bands moderately developed; scat- tered, relatively large granules. | |
| F. A. Sundberg 1994 | Proehmaniella with cranidium with blunt to moderately rounded frontal lobe; frontal area moderately narrow (25-35% cranidial length); anterior border moderately wide (35-50% frontal area length), moderately curved, slightly convex to flat; moderately posterolateral directed ocular ridges (65-70° from axial trend); wide fixi- genae (60-65% glabellar width); moderately fine granular ornamentation. Pygidium subcircular, elongated axis, anterior pleural bands moderately well developed; smooth exoskeleton. | |
| F. A. Sundberg 1994 | Same as genus. [Ehmaniellinae with subpentagonal cranidium with divergent facial sutures anterior of palpebral lobes; moderately to long palpebral lobes (35-40% glabellar length); poorly defined anterior border, gently and uniformly convex preglabellar field and anterior border, shallow anterior border furrow, moderate width (15-20% cranidial length), very strongly tapered; preglabellar field sagittally moderately wide (40-60% frontal area length); moderately to strongly tapered glabella; bluntly rounded frontal lobe; exsagittally wide posterior area of fixigenae (25-30% glabellar length). Pygidium subelliptical with elongate axis; 4 axial rings; moderate depth pleural furrows; strongly convex posterior pleural bands; without anterior pleural bands.] | |
| F. A. Sundberg 1994 | Trachycheilus with moderate size, subpentagonal cranidium with narrow to moderate width anterior border (55-70% frontal area), no median inbend; glabella with deep axial and lateral glabellar furrows, strongly convex, rounded frontal lobe; scattered coarse granular ornamentation; no nodes on posterior border. Pygidium subtriangular with weakly convex or absent anterior pleural bands, strongly developed posterior pleural bands, mod- erately dense medium size granules. | |
| F. A. Sundberg 1994 | Trachycheilus with small subpen- tagonal cranidium with sagittally narrow to wide (65-85% frontal area) and transversely wide (55- 75% cranidial width) anterior border, no or very slight median inbend; glabella with moderate depth axial and lateral glabellar furrows, slightly flattened to flattened frontal lobe; smooth exoskeleton, nodes present or absent on posterior border. Pygidium subelliptical with moderately to weakly convex anterior and posterior bands, smooth surface. | |
| F. A. Sundberg 1994 | Trachycheilus whirlwindensis with cranidium with anterior border with very slight me- dian inbend; wide anterior border furrow, palpebral lobes 28-37% glabellar length, posterior area of fixigenae 22-35% glabellar length with nodes on posterior border; frontal area 20-24% cranidial length. | |
| F. A. Sundberg 1994 | Trachycheilus whirlwindensis with cranidium with anterior border without median in- bend; narrow anterior border furrow, palpebral lobes 22-29% glabellar length, posterior area of fixigenae 39-48% glabellar length without nodes on posterior border; frontal area 11-18% cranidial length. | |
| F. A. Sundberg 1994 | Same as genus. [Ehmaniellinae with subrectangu- lar cranidium with divergent facial sutures anterior of palpebral lobes; moderate to long palpebral lobes (35-45% glabellar length); well-defined anterior border, moderately narrow (10-15% cranidial length), slightly to moderately tapered; preglabellar field sagittally moderately wide (50-55% frontal area length); slightly tapered glabella; moderately round- ed frontal lobe; exsagittally narrow posterior area of fixigenae (20-35% glabellar length). Pygidium subtriangular with elongate axis, 4 axial rings; moderate depth pleural furrows; moderately convex anterior and posterior pleural bands.] | |
| F. A. Sundberg 1994 | Altiocculus with subpentagonal cranidium with median swelling extending from preglabellar furrow to anterior furrow; unevenly, strongly curved, strongly tapered, flat, level anterior border; shallow, relatively narrow, anterior border furrow, slightly convex axial furrows in larger spec- imens; posterior area of fixigenae with rounded terminations. Pygidium subelliptical with moderately convex anterior pleural bands, strongly convex pos- terior pleural bands. | |
| F. A. Sundberg 1994 | Altiocculus with hemicircular cra- nidium with low relief, median swelling extending from preglabellar furrow to anterior furrow; even- ly, strongly curved, slightly tapered, strongly con- vex, level anterior border; deep, relatively wide, anterior border furrow; posterior area of fixigenae with sharp terminations. Pygidium subelliptical with strongly recurved pleural and interpleural furrows near margin, furrows reach margin; moderately weak convex anterior pleural bands, moderately convex posterior pleural bands. | |
| F. A. Sundberg 1994 | Glyphaspis with cranidium with short, strongly tapered glabella from lobe L2 to anterior end, frontal lobe bluntly rounded; trans- versely wide anterior margin (80-95% cranidial width); sagittally long frontal area (45-50% cranidi- al length); concave, wide anterior border; very shallow anterior border furrow. Pygidium relatively transversely elongate, axis reaches margin of pleural field or extends onto border, faint postaxial ridge. | |
| F. A. Sundberg 1994 | Same as genus. [Small Ptychopariida with subtrap- ezoid cranidium with glabella elevated well above fixigena, moderately convex transversely; deep, short lateral glabellar furrows; rounded frontal lobe; con- vergent, deep axial furrows; downsloping, moder- ately wide fixigenae (60-65% glabellar width); nar- row preglabellar field; convex, slightly curved, sagittally narrow, tapering anterior border; short palpebral lobes; weak, thin ocular ridges; short posterior area of fixigenae.] | |
| F. A. Sundberg 1994 | Chanciaopsis with two cranidial types. Specimens without prominent medial swell- ing with convergent, concave facial suture anterior of palpebral lobes; flat, level anterior border, very slight median swelling in anterior border furrow; moderate to large palpebral lobes, moderately pos- terior directed ocular ridges; anterior area of fixi- gena and preglabellar field slightly downsloping to very strongly downsloping; and moderately narrow posterior area of fixigenae (30-40% glabellar length). Specimens with prominent medial swelling with di- vergent convex facial sutures anterior of palpebral lobes; flat, level anterior border relatively wide; moderate to large palpebral lobes, moderately posterior directed ocular ridges; anterior area of fixigena and preglabellar field moderately downsloping to very strongly downsloping; and moderately nar- row posterior area of fixigenae. | |
| F. A. Sundberg 1994 | Deltina with cranidium with relatively wide preglabellar field (30-50% frontal area), moderate length palpebral lobes (30% glabellar length), anterior border curved moderately strong, granular ornamentation. | |
| F. A. Sundberg 1994 | Deltina with cranidium with very narrow to narrow preglabellar field, moderate length to long palpebral lobes (30-45% glabellar length), moderately curved anterior border, smooth exo- skeleton. | |
| F. A. Sundberg 1994 | Elrathina with cranidium with transversely wide anterior border, transversely wide fixigenae (70-80% glabellar width), moderate size recurved occipital spine. | |
| F. A. Sundberg 1994 | Spencella? with cranidium with bluntly rounded glabella; convex axial furrows; moderately dorsally arched anterior border, 65- 70% of frontal area, medial swelling absent or poorly developed; moderately high glabellar convexity (sag. and trans.); and exoskeleton smooth. | |
| F. A. Sundberg 1994 | Spencella? with cranidium with bluntly rounded to rounded frontal lobe; straight to convex axial furrows; moderately to strongly, dorsally arched anterior border, 60-95% of frontal area, medial swelling strong; moderately high glabellar convexity (sag. and trans.); and fine granular ornamentation | |
| F. A. Sundberg 1994 | Yuknessaspis with cranidium with medial swelling, downsloping preglabellar field and anterior area of fixigena. | |
| F. A. Sundberg 1994 | Yuknessaspis benningtonis with cranidium with sagittally elongate, low, medial swelling; very strong to moderately strong down- sloping preglabellar field and anterior area of fixi- gena; slightly tapered glabella; posterior portion of axial furrows shallow; slightly convex anterior bor- der; transversely long (115-155% glabellar length), anteriorly bent posterior area of fixigenae. | |
| F. A. Sundberg 1994 | Yuknessaspis benningtonis with cranidium with rounded very weak to moderately strong medial swelling; very strong downsloping preglabellar field and anterior area of fixigena form- ing sharp flexure in front of glabella; slightly tapered glabella; posterior portion of axial furrows deep; moderately to strongly convex anterior border; transversely shorter (100-110% glabellar length) slightly backswept posterior area of fixigenae. | |
| W. H. Fritz 1995 | Holmiid with cephalon approximately twiceas wide as long. Maximum width of extraocular area less than twice that o f interocular area. Anterior and lateral borders well de fined. Glabella wide, exhibiting only slight restriction at level of posterior glabellar furrow (SI); anterior lobe terminating at or near anterior border. Third pair of glabellar lobes (L3) not appreciably different from second pair (L2). Ocular lobes nar row, strap-like, lacking ocular furrow. Intergenal spines located at or proximal to midlength on posterior border. Genal angles usually moderately advanced. Occipital ring bearing spine with narrow base. Inner pleurae o f thorax narrower (tr.) than axis, transverse, bearing short thorn-like spines that rapidly narrow from the base. Pygidium small, axis comprised of one or two indistinct rings and terminal piece, margin smooth(?) or bearing up to three(?) pairs of small spines. | |
| A. T. Nielsen 1995 | Large species. GlabeIla of low relief, pestleshaped; usually no anterior cranidial border; anterior margin of cranidium rounded (dorsal view) ; genal angle rounded; cephalic terrace lines sparse. Pygidium quite convex, short; no concave border. Axis long, slender, inconsiderably raised above pleural fields, evenly tapering backwards. Surface sculpture of pygidium consists of a few vaguely impressed terrace lines on anterior portion of pleural fields dose to axis. Doublure narrow (cephalon and pygidium). | |
| A. T. Nielsen 1995 | Cranidium of low convexity (both ways), anterior margin evenly rounded and with tlattened rim; anterolateral expansion of glabella strong (tr. ) , but short (exsag. ) ; axial furrows well-impressed; mesial glabellar tuberde situated relatively far back. Pygidium covered, axis indusive, with extremely densely spaeed, very fine terrace lines; axis slightly raised, moderately tapering; border very narrow and steep, ill-defined. Pygidial doublure show more than 25 terrace lines. | |
| A. T. Nielsen 1995 | Cranidium distinetly triangular in outline with only a short ( tr.) anterior border; glabella narrow ( tr.) , moderately inflated, with weakly impressed musde scars; convexity (exsag.) of fixigenae low; pygidium with slender (tr. ) axis, narrow, steep border and a restricted median arch; convexity overall low. | |
| A. T. Nielsen 1995 | Cranidium strongly convex (exsag.) , glabeIla distinetly vaulted, pointed, overhanging anterior margin; glabellar musde scars almost effaced; posterior border narrow, delimited by nearly straight border furrow; cephalic test surface smooth. Pygidium short with high border; axis continues onto border, although with a very low relief. [Emended from V. Poulsen 1965.] | |
| F. C. Shaw 1995 | Cephalic outline semicircular, maximum width (tr.) nearly twice length (sag.), genal prolongations long, about one-third length of sagittal cephalic length. Glabella high, ex- panding anteriorly; posterior end one-eighth total cephalic width, maximum width at inner fringe margin one third total cephalic width, anterior one-fourth overhangs fringe. Only S1 present as small lateral notch at about one-eighth total glabellar length. Occipital furrow present, ring shows occipital spine base and/ or spine in only a very few specimens (e.g., MCZ 2244, not figured). Genal field smoothly inflated, surface detail absent on most molds, but rarely showing deep pits and three transverse distally radiating ridges. | |
| W. H. Fritz 1995 | Palmettaspis species with slightly expanded an terior glabellar lobe, ocular lobes terminating opposite anterior third of posterior glabellar lobes (LI), strongly advanced genal spines, and short, thorn-like pleural spines that narrow rapidly from base. | |
| W. H. Fritz 1995 | Palmettaspis species with nearly parallel-sided glabella, anterior pair (L3) of glabellar lobes markedly shorter (exsag.) than each of two preceding pairs, ocular lobes terminate opposite occipital furrow, and genal spines only slightly ad vanced. | |
| W. H. Fritz 1995 | Esmeraldina species with short (sag.) anterior glabellar lobe that does not reach anterior border furrow. Pos terior margin of occipital lobe inclined back to broad apex from which narrow spine protrudes. Ocular lobe terminates opposite intersection of axial and occipital furrow. Surface of cephalon smooth. | |
| W. H. Fritz 1995 | Esmeraldina*? with a relatively narrow, parallel- sided glabella, transverse posterior margin, and long genal spines. | |
| G. Geyer 1996 | Daguinaspidinae with interocular area sub- horizontally or gently sloping laterally, subequal in width to occipital lobe; glabella clearly tapers forward, with narrow frontal lobe; anterior margin of cephalon gently curved; genal spine angle with rudimentary genal spine in !arge individuals. | |
| G. Geyer 1996 | Daguinaspidinae with interocular area that slopes gently laterally and width less than two-thirds the width of the occipital lobe of glabella; glabella distinctly tapering forward, with narrow frontal lobe; an- terior margin of cephalon bluntly pointed . | |
| G. Geyer 1996 | Fallotaspidinae with glabella slightly ta- pered forward ; ocular Iobes directed posterolaterally from L4, eye ridges distinctly narrewer than posterior ocular Iobes; width of interocular area equal, slightly smaller or greater than width of extraocular area; small intergenal spine present on posterior border; thorax with macropleural third segment; opisthothorax barely differentiated. | |
| G. Geyer 1996 | Fallotaspidinae with glabella tapered for- ward; ocular Iobes directed posterolaterally, eye ridges faintly narrewer than posterior ocular Iobes ; width of interocular area about half or less width of extraocular area; intergenal spine only visible in immature indivi- duals; thorax with macropleural third segment; opis- thothorax weakly differentiated. | |
| G. Geyer 1996 | A species of Fa//otaspis with the following characteristics: Glabella weakly tapers for- ward; occipital ring with weil defined anterolateral nodes and extended into a moderately strong terminal spine; ocular Iobes short, posterior tips opposite LI in adult specimens; extraocular area wide ; thoracic tergites 4 to 13 with conspicuous median Spines on axial rings. | |
| G. Geyer 1996 | Species of Daguinaspi with adult cephalon heart-shaped in out! ine width about equal to length, anterior margin with geniculation; glabella moderately Iong, distinctly tapering forward, with narrow frontal lobe; preglabellar field sagittally wide. | |
| G. Geyer 1996 | Figs . 19 , 32 , 35 , I, 37, 38 , 42 , 65 , 1-65 ,8 | |
| G. Geyer 1996 | A species of Fallotaspis with the glabella weakly tapering forward· occipital ring with weil defined anterolateral nodes, without terminal spine; ocular Iobes moderately long, posterior tips op- posite anterior margin af occipital ring in adult speci- mens; extraacular areas comparatively narrow · anterior and lateral border furrows form shallow, poorly de- fined furrows ; proximal parts af anterior and lateral border with shallow depression; posterolateral cephalic margins almost parallel ; thoracic tergites with rather small axial nades. | |
| G. Geyer 1996 | A species of Fallotaspis with a relatively hort , moderately tapering glabella with slightly truncate frontal lobe; occipital ring with weil defined anterolateral nodes, extended into a terminal spine· ocular Iobes moderately long, posterior tips opposite anterior part of occipital lobe in adult specimens; extraocular area moderately wide. | |
| G. Geyer 1996 | A specie s of Fallotaspis with ce- phalon semiparabolic or , rarely, semicircular in out- line; glabella relatively strongly tapering forward · oc- cipital ring with weil defined anterolateral nodes, Iacks terminal spine; ocular Iobes moderately long, posterior tips opposite anterior margin of occipital ring in adult specimens ; extraocular areas relatively wide . | |
| G. Geyer 1996 | A species of Fallotaspis with cephalon semiparabolic to semicircular in outline , gla- bella slightly tapering , cephalic front separated from anterior border furrow occipital ring with weil deve- l oped anterolateral nodes and mode r ately long, slender occipital spine; posterior cephalic ma r gin with strong forwa rd bend an d resultant anterior position of genal spine angle; anterior and lateral border relatively wide; pleural spines of the third thoracic tergite extremely long. | |
| G. Geyer 1996 | Species of Fallotaspis with a weakly tape- ring glabella, cephalic front relatively close to anterior border furrow, occipital ring with broad-based, stro.ng terminal occipital spine, posterior cephalic margin with moderate forward bend, preglabellar field sagittally narrow , extraocular areas relatively wide, anterior border moderately wide. | |
| G. Geyer 1996 | Species of Choubertel/a with moderately curved anterior margin of cephalon, strongly tapering glabella, relatively lang and curved ocular Iobes, thick eye ridges, and relatively thin ante- rior and lateral borders. | |
| G. Geyer 1996 | Species of Daguinaspis with ce- phalon wider than long, broadly rounded anteriorly but margin with faint medial geniculation; glabella moder- ately long and strongly tapered forward; frontal lobe narrow; preglabellar field moderately wide sagittally. | |
| G. Geyer 1996 | Species of Daguinaspis with cephalon heart-shaped in outline; glabella long and faintly tapering forward ; preglabellar field narrow sagittally. | |
| F. A. Sundberg and L. B. McCollum 1997 | Oryctocephalinae with cranidium with medially to anteriorly expanded glabella; lateral glabellar fur rows with pits at SI-S3, not extending to axial furrows, typically SI pits connected with transglabellar furrow, S4 pits small and faint; occipital ring typically without longitudinal furrows pro jecting from So depression to the posterior margin; anterior mar gin of rostral-hypostoma plate slightly wider than maximum gla bellar width. Pygidium with typically 5 or 6 pairs of cylindrical spines and moderately developed interpleural furrows. | |
| F. A. Sundberg and L. B. McCollum 1997 | Isopygous with cranidium subpentagonal, gla bella medially expanded without medial anterior sulcus, only S1 lateral glabellar furrow extending to axial furrow, SI and S2 transglabellar furrows, sinuous axial furrows; basal lobes on LI; fixigena narrow, tapering, posterior area sharply terminated; pal pebral lobes long; surface granular; opisthoparian sutures. Five thoracic segments with pleural bands of uniform length (exsag.), bluntly terminated. Pygidium semicircular, without marginal spines; axis narrow, 4 axial rings; 5 pleural and interpleural fur rows, each pleura equally divided in anterior and posterior bands; pygidial border moderately well defined. | |
| F. A. Sundberg and L. B. McCollum 1997 | Same as genus. [Isopygous with cranidium subpentagonal, gla bella medially expanded without medial anterior sulcus, only S1 lateral glabellar furrow extending to axial furrow, SI and S2 transglabellar furrows, sinuous axial furrows; basal lobes on LI; fixigena narrow, tapering, posterior area sharply terminated; pal pebral lobes long; surface granular; opisthoparian sutures. Five thoracic segments with pleural bands of uniform length (exsag.), bluntly terminated. Pygidium semicircular, without marginal spines; axis narrow, 4 axial rings; 5 pleural and interpleural fur rows, each pleura equally divided in anterior and posterior bands; pygidial border moderately well defined.] | |
| F. A. Sundberg and L. B. McCollum 1997 | Cranidium with a conical glabella, SI to S 3 transglabellar furrows, e y e ridges directed slightly poster- olaterally, wide fixigena (92 ± 1 1 percent glabellar width), near ly straight anterior margin, and anterior border without curvature towards frontal lobe, and no intergenal angle. Twelve thoracic segments each with cylindrical pleural spines. Pygidium with 3 axial rings; anterior pleural bands long (exsag.), posterolaterally to posteriorly directed with cylindrical spines; and V I to V 3 spines decreasing in size and projecting posteriorly. | |
| H. B. Whittington et al. 1997 | Glabella broadly conical; preglabellar field short (sag.) or absent. Thorax with 9 to 13 segments. Pygidium large and spinose (except
in Parapaokannia), usually with 3 to 6 pairs of spines | |
| F. A. Sundberg and L. B. McCollum 1997 | Cranidium with glabella parallel sided, SI to S3 transglabellar furrows, eye ridges directed strongly pos- terolaterally (50-65 degrees), fixigena narrow (55-70 percent glabellar width), anterior margin moderately curved, and ante rior border without curvature towards frontal lobe. Pygidium with 5 axial rings; anterior pleural bands short (exsag.) with cylindrical spines; VI and V2 spines projecting laterally; V3 spine projects more posteriorly; V4 and V5 spines projecting posteriorly; spines increasing in length posteriorly except V5 which is small. | |
| F. A. Sundberg and L. B. McCollum 1997 | Cranidium with a concave-sided gla bella and expanded frontal lobe, SI to S2 or S3 transglabellar furrows, eye ridges directed moderately posterolaterally (68 ± 3 degrees to axis), wide fixigena (92 ± 4 percent glabellar width), curved anterior margin (159 ± 3 degrees) with curvature towards frontal lobe, and intergenal angle. Fifteen thoracic seg ments each with flat pleural spines. Pygidium with 3 axial rings; anterior pleural bands short (exsag.), posteriorly directed with flat spines; and VI to V3 spines decreasing in size and projecting posteriorly. | |
| F. A. Sundberg and L. B. McCollum 1997 | Cranidium with medially expanded glabella, cir cular glabellar pits at the SO to S3 positions, SI transglabellar furrow, longitudinal furrows connecting SI to S3 pits, lateral glabellar furrows not extending to axial furrows, and occipital ring without longitudinal furrows. Seven thoracic segments each with long, cylindrical spine. Pygidium with 5 pairs of cylindrical spines. | |
| F. A. Sundberg and L. B. McCollum 1997 | Cranidium with medially expanded glabella, cir cular glabellar pits at the SI to S2 positions, slit-shaped pits at So and S3, SI transglabellar furrow present or absent, longitu dinal furrows absent, S3 extending to axial furrows, and occip ital ring without longitudinal furrows. Nine thoracic segments each with long, cylindrical spine. Pygidium with 3 pairs of cy lindrical spines. | |
| G. E. Budd and J. S. Peel 1998 | Large, Xenusion-like lobopod, but differing in possessing rows of four trunk nodes instead of two, in the probable presence of annular nodes, and in the lack of fleshy limb outgrowths. Hardranax bears a pair of long, branched, probably frontal appendages that are not known from Xenusion. | |
| D. J. Holloway and P. D. Lane 1998 | Cephalon strongly convex (sag.), curvature in sagittal plane subtending more than 90°, height in lateral profile greater than or equal to sagittal length. Omphalus and anterolateral internal pit present. Axial furrow diverging moderately behind and immediately in front of lunette, dying out anteriorly behind omphalus. Eye large, situated less than its own length from posterior cephalic margin; socle not strongly convex (tr.). Posterior branch of facial suture strongly diverging backwards; anterior branch diverging moderately forwards. Genal angle broadly rounded. Rostral plate sub-triangular, gently convex (sag., exsag.) over anterior 70 per cent. and gently concave in posterior part, without upturned posterior flange; connective suture meeting hypostomal suture close to sagittal line; vincular furrow present across posterior edge of doublure. Thorax with very wide, gently arched axis comprising 60--70 per cent. segmental width (tr.); axial furrow weak; fulcrum situated very close to axial furrow; pleurae abaxial to fulcrum almost continuous in slope with lateral part of axial rings. Pygidium moderately convex (sag., exsag.), lenticular in dorsal view, maximum width just in front of midlength; anteriorrnost pleural furrow and holcos very weak or not defined. Terrace ridges present over most of dorsal surface of cephalon and pygidium. | |
| J. D. Eddy and L. B. McCollum 1998 | Cranidia with anteriorly expanded glabella; pal- pebral lobe situated opposite glabellar midlength; median node on occipital ring; broadly expanded fixigena behind palpebral lobe; fine granular ornamentation on anterior third of glabella only. Pygidia with four axial ring segments; poorly developed, thin, linear transverse nodes on anterior two axial rings, four pleural furrows; shallow border furrow; marginal spines directed posteriorly to slightly convex inward; fine granular ornamentation on border and spines only. | |
| J. H. Stitt 1998 | A species of Menomonia with conical glabella, strongly tapering to rounded point. Palpebral lobes elevated, located opposite anterior end of glabella. Palpebral areas very narrow. Anterior border broadly triangular, more convex at anterior margin than at posterior end. Internal mold of cranidium and librigena covered by fine granules. | |
| B. S. Lieberman 1999 | Anterior cephalic border prominently separated from extraocular area by fur- row, developed as raised, rounded ridge; plectrum absent; L4 contacts anterior border furrow; anterior margins of L4 at each side of midline deflected posteriorly at roughly 40 degree angle relative to transverse line; length (sag.) of L4 equal to 1.5 times length of L0 and L1 (sag.); L4 expands prominently dorsally; lateral margins of L4 distal to lateral margins of L0; region of anterior part of ocular lobe between visual surfaces and L4 nar- row (tr.); ocular lobe with ocular furrow; line from posterior tip of ocular lobe to junc- tion of posterior margin of lobe with glabella forms roughly 10 degree angle relative to sagittal line; transverse profile of ocular lobes convex dorsally; interocular area arched; posterior tips of ocular lobes developed opposite S0 or medial part of distal margin of L1; width (tr.) of interocular area roughly equal to width of ocular lobe; S3 jaggedly convex, conjoined medially; entire margin of S3 arched furthest anteriorly at point three-quarters of way between midline of glabella and axial furrows; L1, L2, and L3 of strong transverse convexity; L2 and L3 do not merge; S2 conjoined medially; distal margins of L2 when proceeding anteriorly subparallel; S1 conjoined medially; distal sector of S0 straight, con- joined medially, proximal end well posterior of distal end; axial part of L0 with axial node; posterior margin of L0 convex posteriorly; lateral lobes on L0 absent; posterior cephalic border between L0 and intergenal angle deflected posterolaterally; prominent anasto- mosing ridges present on extraocular area; width (tr.) of extraocular region opposite L1 equal to roughly 75% of width of glabella at L1; genal spine near cephalic border directed posterolaterally at angle greater than or equal to 25 degrees relative to sagittal line; genal spine length (exsag.) greater than or equal to length of first 4 to 5 thoracic segments; intergenal angle developed two-thirds of way between distal tips of ocular lobes and genal spine angle, or directly behind genal spine angle. | |
| B. S. Lieberman 1999 | Anterior cephalic border between L4 and genal spine angle relatively short, length (exsag.) less than or equal to length (sag.) of L1; anterior cephalic border developed as elevated, flattened ridge, prominently separated from extraocular area; frontal lobe con- tacts anterior border furrow; anterolateral margins of frontal lobe deflected posteriorly at roughly 10 degree angle relative to transverse line; frontal lobe moderately long (sag.), length equal to length of L0 and L1 medially; glabellar furrows prominently incised; S3 same depth laterally as medially; lateral margins of L2 when proceeding anteriorly bulging laterally relative to L0; lateral margins of glabella between posterior margin of L0 and L1 constrict; distal tips of S0 straight; extraocular region flattened; intergenal ridge visible as faint trace; intergenal spine absent in adult; genal spine length (exsag.) of approximately first eight thoracic segments; thorax divided into pro- and opisthothorax; nodes present on medial part of thoracic axial rings; T3 macropleural, spine length (exsag.) greater than length of thoracopygidium; anterior margin of T3 deflects anteriorly before flexing poste- riorly; anterior margin of pleural furrow on T3 parallels transverse line before flexing pos- teriorly; anterior margin of thoracic pleural furrow separated from body of segment by prominent ridge; thoracic pleural spines, behind T4, developed as short sweeping projec- tions extending two to three thoracic segments back, lying in roughly same dorso-ventral plane as pleural segments; prominent spine on axial ring of T15 (see Harrington 1956 and Lieberman 1998 for additional characters). | |
| B. S. Lieberman 1999 | Anterior cephalic border developed as rounded ridge, prominently separated by furrow from extraocular area; frontal lobe contacts anterior border furrow, lateral margins distal of lateral margins of L0, expands prominently dorsally; anterolateral margins of frontal lobe prominently separated from extraocular area by furrow (not anterior border furrow); length (sag.) of L4 equal to 1 to 1.1 times length of L0 and L1; dorsal notch in ocular lobe where it contacts lateral margins of frontal lobe; S3 evenly convex, medial and distal tips as far forward; anterodistal margins of L3 formed by axial furrows; distal margin of L3 straight; lateral margins of glabella opposite L2 diverging anteriorly; medial and distal tips of S2 as far forward; S1, S2 and S3 conjoined medially; posterior edge of ocular lobe opposite distal tip of S0; line from posterior margin of ocular lobe to junction of lobe with glabella forms roughly 5 to 15 degree angle relative to sagittal line; L0 with spine posterior of midline; lat- eral lobes absent from L0; intergenal angle with small spine directly behind distal tip of oc- ular lobe; intergenal angle forms roughly 10 to 15 degree angle with transverse line; extraoc- ular area prominently vaulted, width (tr.) at L1 45% to 55% width of glabella at L1; genal spine angle opposite distal margin of L0; lateral margin of genal spine near spine angle forms roughly 30 to 40 degree angle with sagittal line; genal spine extends back roughly 8 thoracic segments; thoracic pleural spines on first four segments extend back only one segment; lat- eral margins of spines inclined at 40 to 50 degree angle relative to sagittal line; posterior mar- gins of first four thoracic segments near spines flex slightly anteriorly; length (sag.) and width (tr.) of pygidium roughly equal; posterior margin of pygidium roughly transverse. | |
| B. S. Lieberman 1999 | Length (exsag.) anterior cephalic border near but not directly anterior to frontal lobe very long, equal to length (sag.) of L1 and L2; length (sag.) L4 equal to 1.5 times length (sag.) of L0 and L1; distal margins of L2 subparallel; S0, S1 not conjoined medially; L0 smooth, convex posteriorly; genal spine near cephalic border directed posterolaterally at 35 to 45 degree angle relative to sagittal line, length (exsag.) equal to four times length (sag.) L0; genal spine angle opposite medial part of distal margin of L3; cephalic posterior border transverse between intergenal angle and L0. | |
| B. S. Lieberman 1999 | Anterior cephalic border not prominently separated from extraocular area by furrow; frontal lobe of glabella contacts anterior border furrow; anterolateral parts of glabella not prominently separated from extraocular area by furrow; prominent parafrontal band not visible in dorsal view; anterior margins of frontal lobe at each side of midline deflected posteriorly at roughly 40 degree angle relative to transverse line; length (sag.) of frontal lobe equal to 1.5 times length of L0 and L1 medially; L4 expands dorsally; lateral margins of L4 distal to lateral margins of L0; abrupt dorsal notch-like truncation of ocular lobes at margin of L4 absent; ocular lobe smoothly merges into extraocular area; oc- ular lobe with prominent furrow; transverse profile of ocular lobes convex dorsally; sur- face of interocular area developed as flattened shelf; anterodistal margins of L3 formed by ocular lobes; distal margins of L3 convex outward; S3 conjoined medially, jaggedly convex; medial and distal tips of S3 as far forward; lateral margins of glabella at L2 when proceed- ing anteriorly diverging; L1 to L3 strongly transversely convex; line between ends of S2 transverse; S0 not conjoined medially; distal sector of S0 convex anteriorly; posterior mar- gin of ocular lobe opposite distal tip of S0; L0 with axial node posterior of midline; lateral lobes present on L0; extraocular region prominently vaulted; genal spine near genal spine angle directed posterolaterally at roughly 10 to 20 degree angle relative to sagittal line; genal spine extends back four to five thoracic segments; genal spine angle opposite medial part of distal margin of L0; intergenal angle posterior of lateral margins of ocular lobes; posterior margin of thoracic pleural furrow on T3 directed evenly posterolaterally; lateral margins of prothoracic pleural spines inclined at roughly 45 degree angle relative to sagit- tal line; posterior margins of first four thoracic segments near spine bases flex slightly an- teriorly before flexing posteriorly; boundary between thoracic pleural furrow and anterior band gradational; length (exsag.) of thoracic pleural furrows at medial part of thoracic segment 1 to 1.3 times length of pleural furrows on distal part of segment; length (exsag.) of thoracic pleural furrows at medial part of segment equal to length (exsag.) of posterior band; spine on 15th thoracic segment short, length (sag.) of two thoracic segments; opisthothorax with prominent pleurae which continue orientation of prothoracic seg- ments; pygidium with length (sag.) equal to width (tr.); posterior margin of pygidium weakly convex. | |
| B. S. Lieberman 1999 | Anterior cephalic border developed as flattened plane, prominently separated from extraocular area by furrow; frontal lobe contacts anterior border furrow, lateral mar- gins distal to lateral margins of L0, expands prominently dorsally; anterolateral margins of glabella prominently separated from extraocular area by furrow; length (sag.) of frontal lobe 1 to 1.1 times length (sag.) of L0 and L1; dorsal notch-like truncation in ocular lobes where they contact frontal lobe of the glabella; anterodistal margins of L3 formed by axial furrows; distal margins of L3 straight; S3 evenly convex, medial and distal tips as far for- ward; lateral margins of glabella opposite L2 subparallel; S1, S2 and S3 not conjoined me- dially; medial and distal tips of S2 as far forward; line from posterior edge of ocular lobe to junction of lobe with glabella forms roughly 10 to 15 degree angle with sagittal line; poste- rior margin of ocular lobe opposite distal tip of S0; lateral lobes present on L0; node or spine on L0 posterior of midline; intergenal angle with small spine directly behind distal tip of ocular lobe; intergenal angle forms 80 to 90 degree angle with transverse line; extraocu- lar region prominently vaulted, very narrow, width (tr.) at L1 10% to 15% width of glabella at L1; genal spine angle opposite distal tip of L3 or L4; lateral margin of genal spine near spine angle forms roughly 60 to 70 degree angle relative to sagittal line; pygidium twice as wide (tr.) as long (sag.), posterior margin notched. | |
| B. S. Lieberman 1999 | Anterior cephalic border not prominently separated from extraocular area by deep furrow, developed as flattened, angled plane; frontal lobe contacts anterior border fur- row, does not expand prominently dorsally; lateral margins of L4 directly anterior to lateral margins of L0; anterolateral margins of glabella not prominently separated from extraocu- lar area by furrow; length (sag.) of frontal lobe equal to 1.5 times length (sag.) of L0 and L1; pre-ocular furrow on frontal lobe directed inward and forward from glabellar margin; an- terodistal margins of L3 formed by ocular lobes; postero- and anterodistal margins of L3 with same orientation; lateral margins of glabella opposite L2 diverging when proceeding anteriorly; posterior margins of ocular lobes opposite distal tip of S0; line from posterior margin of ocular lobe to junction of posterior margin of lobe with glabella forms 20 to 25 degree angle relative to sagittal line; S3 evenly convex, distal tip further forward than medial tip; distal tip of S2 further forward than medial tip; S0, S1, S2 and S3 conjoined medially; S0 sinuous; L0 with spine posterior of midline of L0; lateral lobes on L0 present; intergenal angle developed midway between ocular lobe and genal spine, with small spine; intergenal angle forms 10 to 15 degree angle with transverse line; extraocular area gently sloping from eye to lateral border furrow, width (tr.) opposite L1 equal to 45% to 55% width of glabella at L1; genal spine angle opposite medial part of distal margin of L0; genal spine close to genal spine angle forms roughly 10 to 15 degree angle relative to sagittal line; genal spine extends back roughly 4 to 5 thoracic segments. | |
| B. S. Lieberman 1999 | Anterolateral margins of frontal lobe of glabella not prominently separated from extraocular area by furrow; posterior margins of frontal lobe expanding where ocular lobes intersect with it; anterior portions of posterior margins of ocular lobes opposite medial part of distal margins of L1; posterodistal margins of L3 formed by ocular lobes; length (exsag.) of genal spine less than or equal to length (sag.) of first four thoracic segments; lateral lobes absent from L0; intergenal angle forms 30 to 50 degree angle with transverse line; lateral margins of each prosomal thoracic axial ring converging when proceeding from anterior to posterior edges; field of thoracic pleural furrows at medial portion long (exsag.), equal to approximately 1.3 times length of posterior pleural band; anterior margin of T3 medially flexes anteriorly; posterior margin of thoracic pleural furrow on T3 directed evenly poste- riorly laterally; width (tr.) of thoracic axis 100% width of pleural field excluding spines. | |
| B. S. Lieberman 1999 | Anterior cephalic border between frontal lobe and genal spine angle with length (exsag.) equal to length of L0; anterior border prominently separated from extraocular area by furrow; anterolateral margins of frontal lobe separated from extraocular area by furrow (not anterior border furrow); anterior margins of frontal lobe at each side of midline de- flected posteriorly at roughly 10 degree angle relative to transverse line; prominent parafrontal band not visible in dorsal view; L4 expands dorsally; lateral margins of L4 dis- tal to lateral margins of L0; pre-ocular furrow on frontal lobe directed inward and forward from glabellar margin; transverse profile of ocular lobes convex dorsally; posterolateral tip of ocular lobe directly behind anterolateral tip or rotated slightly laterally; vertical shelf of ocular lobe separated from extraocular area by furrow (except in M. truncatooculatus); S3 jaggedly convex or carat shape, conjoined medially; line between ends of S2 transverse; S2 convex anteriorly; S1 conjoined medially; distal sector of S0 straight; lateral lobes on L0 pre- sent; posterior margin of L0 convex; extraocular area gently flattened or convex; intergenal angle developed posterior of point halfway between ocular lobes and genal spine; extraoc- ular region opposite L1 with width (tr.) equal to roughly 75% of width of glabella at L1; in- tergenal ridge visible as trace; intergenal area with small pointed spine (could not be deter- mined in M. macer); intergenal angle roughly parallels a transverse line; genal spine angle developed opposite medial part of distal margin of L0; T3 macropleural; boundary between thoracic pleural furrow and anterior band sharp; thoracic pleural furrows extend onto spines; length (exsag.) of thoracic pleural furrows (excluding those on T3) at medial part of segment equal to half length (exsag.) of posterior band at similar point; prominent antero- lateral lobes variably present or absent on thoracic axial rings; thoracic pleural spines on segments T5 to T8 extend back approximately four segments; spine on T15 short, length (sag.) of five thoracic segments (could not be determined in M. macer or M. muralensis); and base of spine on T15 nearly as wide (tr.) as axis of segment (could not be determined in M. macer); pleurae present on opisthothorax (could not be determined in M. macer or M. muralensis). | |
| B. S. Lieberman 1999 | Anterior cephalic border prominently separated from extraocular area by fur- row; plectrum absent; L4 contacts anterior border furrow; anterior margins of L4 at each side of midline deflected posteriorly at roughly 10 to 20 degree angle relative to transverse line; length (sag.) of L4 equal to 1.5 times length of L0 (sag.); L4 expands prominently dor- sally; lateral margins of L4 distal to lateral margins of L0; region of anterior part of ocular lobe between visual surfaces and L4 broad (tr.), one-third width of glabella at L1; ocular lobe without prominent ocular furrow; line from posterior tip of ocular lobe to junction of posterior margin of lobe with glabella forms roughly 45 degree angle with sagittal line; transverse profile of ocular lobes convex dorsally; posterior tips of ocular lobes developed opposite medial part of distal margin of L1; width (tr.) of interocular area greater than width of ocular lobe; S3 jaggedly convex, not conjoined medially; L2 and L3 do not merge; S2 convex anteriorly, conjoined medially; distal margins of L2 when proceeding anteriorly diverging; S1 conjoined medially; S0 convex anteriorly, not conjoined medially, proximal and distal ends on transverse line; L0, L1, L2 and L3 with axial spine; posterior margin of L0 convex posteriorly; prominent anastomosing ridges absent on extraocular area; width (tr.) of extraocular region opposite L1 equal to 75% of width of glabella at L1; genal spine near cephalic border directed posterolaterally at roughly 35 to 45 degree angle relative to sagittal line; genal spine short (exsag.), length approximately equal to length of first 2 to 3 thoracic segments; genal spine angle developed opposite medial part of distal margin of L0; intergenal angle deflected anteriorly at least 30 degrees relative to transverse line; medial part of posterior border between intergenal angle and L0 flexes posteriorly. | |
| B. S. Lieberman 1999 | Anterior cephalic border prominently separated from extraocular area by fur- row, border developed as rounded ridge; frontal lobe does not contact anterior cephalic border furrow (variably developed in “P.” cometes); frontal lobe expands prominently dor- sally; anterolateral margins of glabella prominently separated from anterior extraocular area; length (sag.) of frontal lobe equal to 1 to 1.1 times length (sag.) of L0 and L1; S3 evenly convex; notch in ocular lobes where they contact frontal lobe (not in “P.” cometes); S1, S2 and S3 conjoined medially; posterior margin of ocular lobe opposite distal tip of S0 or me- dial part of distal margin of L0; line from posterior margin of ocular lobe to junction of posterior margin of lobe with glabella forms roughly 0 to 15 degree angle with sagittal line; L0 with spine, posterior of midline; intergenal angle with small spine directly behind distal tip of ocular lobe; extraocular region prominently vaulted, width (tr.) at L1 45% to 55% width of glabella at L1; genal spine extends back roughly eight thoracic segments; thoracic pleural spines very short (exsag.), extend back only one segment, lateral margins inclined at 10 to 20 degree angle relative to sagittal line (see Fritz 1995 for additional characters). | |
| B. S. Lieberman 1999 | Length (exsag.) anterior cephalic border near but not directly anterior to frontal lobe equal to length (sag.) of L0; cephalic anterior border not prominently separated from extraocular area by furrow; frontal lobe of glabella contacts anterior border furrow; length of L4 (sag.) equal to length (sag.) of L0 and L1; posterolateral margins of L4 divergent anteri- orly, distal to lateral margins of L0; anterodistal margins of L3 formed by ocular lobes; distal margin of L3 convex outward; S3 carat shaped, not conjoined medially; distal margins of L2 diverge anteriorly; posterior tips of ocular lobes developed opposite medial part of distal margin of L1; L0 with lateral lobes absent, node present; line between ends of S2 transverse; S0, S2 convex anteriorly; length (exsag.) of genal spine equal to length (sag.) of first four to five thoracic segments; intergenal spine faintly developed; intergenal angle developed poste- rior to point half way between ocular lobes and genal spine; anterior margin of thoracic pleural furrow on third segment parallels transverse line axially, abaxially flexes strongly pos- teriorly; thoracic pleural furrows extend only half width of inner pleural region; prominent spine of length (sag.) five thoracic segments present on 15th axial ring, width (tr.) of base of spine nearly as wide as axis of segment; length (sag.) of pygidium equal to width (tr.). | |
| B. S. Lieberman 1999 | The Olenellinae is characterized by the following shared characters when present in combination: 1. posterolateral margins of L4 divergent anteriorly; 2. transverse profile of ocular lobes flattened dorsally; 3. posterior tips of ocular lobes developed opposite medial part of distal margin of L0 or distal tip of S0; 4. S3 not conjoined medially; 5. line between ends of S2 directed inward and posteriorly at 45 degree angle relative to transverse line; 6. S0 not conjoined medially; 7. lateral lobes on L0 absent; 8. glabellar furrows moderately to strongly incised; 9. genal ridge prominently developed; 10. extraocular region opposite L1 broad, width (tr.) greater than or equal to 75% of width of glabella at L1; 11. genal spine near where it hits cephalic border directed posterolaterally at roughly 10 to 20 degree angle relative to sagittal line; 12. length (exsag.) of genal spine roughly equal to length (sag.) of first four to six thoracic segments; 13. genal spine angle developed opposite medial part of distal margin of L0; 14. intergenal angle developed adjacent to or directly behind genal spine angle; 15. intergenal angle with small, pointed spine; 16. intergenal angle deflected at roughly 10 degree angle relative to transverse line; 17. medial part of cephalic posterior border between intergenal angle and L0 weakly flexed anteriorly; 18. anterior margin of third thoracic pleural segment, before flexing strongly posteriorly, parallel to transverse line or weakly deflected posteriorly; 19. posterior margin of thoracic pleural furrow on third thoracic segment directed evenly posterolaterally; 20. thoracic pleural spines on seg- ments T5 to T8 developed as broad sweeping projections extending roughly four segments back; 21. length (exsag.) of thoracic pleural furrows (excluding those of T3) at medial part of segment equal to roughly 1.3 times length (exsag.) of posterior band of pleural segment; 22. width of thoracic pleural spines T5 to T8 at spine midlength more than two-thirds length (exsag.) of medial part of inner pleural region; 23. length (sag.) of spine on 15th thoracic segment roughly equal to length of whole prothorax; 24. base of spine on 15th thoracic segment nearly as wide (tr.) as axis of segment; 25. segments of opisthothorax with reduced pleurae; 26. pygidium length (sag.) 1.5 times width (tr.); 27. posterior mar- gin of pygidium weakly convex. Of these characters, only character 25 is unique to the Olenellidae. | |
| B. S. Lieberman 1999 | Plectrum present; length (sag.) of preglabellar field approximately 15% to 25% length (sag.) of L4; lateral margins of L4 directly anterior of L0 or medial of L0; S2 not con- joined medially; lateral margins of L2 bulging laterally relative to L1; S1 conjoined medially; ocular lobe with prominent furrow across entire margin; posterior margin of ocular lobe extends back to medial part of distal margin of L0; extraocular area gently convex; anterior margins of T3 deflected anteriorly relative to transverse line at 5 to 10 degree angle; T3 ex- tends back length of entire thorax; ratio of width (tr.) T7, at articulating tips, relative to width of T3 at same position, 0.70 to 0.75. | |
| G. E. Budd 1999 | Nektaspids with more than one axial articulation. | |
| G. E. Budd 1999 | Nektaspids with more than one axial articulation. | |
| H. H. Basibuyuk et al. 1999 | "...fossils appear to belong to a single species which displays a combination of characters not known in any extant subfamily and we therefore erect a new subfamily here to receive them. Further, the
presence in these of both cyclostome mouthparts and hindwing vein 2-CU has led us to reappraise the homologies and significance of various features of hindwing venation within the family." | |
| B. S. Lieberman 1999 | Anterior cephalic border not prominently separated from extraocular area by furrow; frontal lobe contacts anterior border furrow, not expanding prominently dorsally; distal margins of L4 deflected as far or farther laterally than distal margins of L0; length (sag.) of frontal lobe 1.5 times length of L0 and L1; anterodistal margins of L3 formed by ocular lobes; lateral margins of L3 convex outward; S3 and S2 with medial and distal tips as far forward; lateral margins of glabella opposite L2 diverging when proceeding anteriorly; S1, S2 and S3 conjoined medially; posterior margin of ocular lobe opposite medial part of distal margin of L0; lateral lobes on L0 present; L0 with node or spine posterior of midline of L0; intergenal angle directly behind distal tip of ocular lobe; intergenal angle forms 0 to 15 degree angle with transverse line; intergenal angle with small spine; extraocular region gently sloping from eye to lateral border furrow; genal spine angle opposite medial part of distal margin of L0. | |
| B. S. Lieberman 1999 | The Mesonacinae can be defined by the possession of the following diagnostic char- acters in combination (thoracic and pygidial characters are not preserved in all species as- signed to the Mesonacinae): 1. anterior part of ocular lobe close to L4; 2. ocular lobe with prominent furrow; 3. width (tr.) of interocular area approximately equal to width of ocular lobe; 4. axial part of L0 with node; 5. extraocular area flattened, lacking prominent anasto- mosing ridges (except in M. svalbardensis); 6. extraocular region opposite L1 40% to 50% width (tr.) of glabella at L1 (except in M. fremonti); 7. intergenal angle directed anteriorly at angle greater than or equal to 30 degrees relative to transverse line; 8. anterior margin of thoracic pleural furrow on third segment parallels a transverse line before flexing strongly posteriorly when proceeding from proximal to distal edge; 9. width (tr.) of thoracic pleural spines T5 to T8 at spine midlength less than or equal to half length (exsag.) of correspond- ing segment between spine and axis; 10. segments of opisthothorax with prominent pleu- rae (except in M. fremonti); 11. pygidium length (sag.) 1.5 times width (tr.). None of these characters are unique to the Mesonacinae, meaning they all show some homoplasy or are symplesiomorphic within the Olenelloidea. | |
| B. S. Lieberman 1999 | Anterior cephalic border developed as raised ledge; anterior border promi- nently separated from extraocular area by furrow; plectrum absent; anterior part of glabella prominently separated from anterior extraocular area by furrow (not anterior bor- der furrow); prominent parafrontal band not visible in dorsal view; anterior margins of frontal lobe at each side of midline deflected posteriorly at roughly 40 degree angle relative to transverse line; length (sag.) of L4 equal to roughly 1.5 times length (sag.) of L0 and L1; L4 expands prominently dorsally; lateral margins of L4 distal to lateral margins of L0; pre-ocular furrow on frontal lobe directed inward and forward from glabellar margin; an- terolateral margins of ocular lobe separated from extraocular area by prominent furrow; ocular lobe with prominent ocular furrow; line from posterior tip of ocular lobe to junc- tion of posterior margin of lobe with glabella forms roughly 10 to 15 degree angle with sagittal line; transverse profile of ocular lobes convex dorsally; surface of interocular area arched; width (tr.) of interocular area approximately equal to maximum width of ocular lobe; anterodistal margins of L3 formed by ocular lobe; distal margins of L3 convex out- ward; S3 jaggedly convex, conjoined medially; line between ends of S2 transverse; L2 and L3 do not merge; S2 convex anteriorly; distal margins of L2 when proceeding anteriorly di- verging; distal sector of S0 convex anteriorly, well anterior of proximal end; posterior mar- gin of L0 roughly transverse; lateral lobes on L0 present; extraocular area flattened, width (tr.) roughly equal to 75% width of glabella at L1; length (exsag.) of genal spine roughly equal to length (sag.) of first four to five thoracic segments; genal spine angle developed opposite medial part of distal margin of L0; intergenal angle developed posterior of point half way between ocular lobes and genal spine; intergenal angle roughly parallel to trans- verse line; medial part of posterior border between intergenal angle and L0 roughly trans- verse; thorax broken up into pro- and opisthothorax; anterior margin of T3 before flexing strongly posteriorly roughly parallel to transverse line; T3 macropleural, projects posteri- orly 6 to 8 thoracic segments; anterior margin of thoracic pleural furrow on T3 when pro- ceeding from proximal to distal edge directed weakly posteriorly, before flexing strongly posteriorly; posterior margin of thoracic pleural furrow on T3 directed evenly posterolat- erally; thoracic pleural spines on T5 to T8 developed as broad sweeping projections ex- tending roughly four segments back; prothoracic axial rings without prominent lateral fur- rows; single nodes very faint, but present on median portion of some thoracic axial rings; thoracic pleural furrows extend width of inner pleural region; boundary between thoracic pleural furrow and anterior band sharp; thoracic pleural furrows short (exsag.), length equal to roughly half length (exsag.) of posterior band; thoracic pleural spines T5 to T8 at spine midlength with width (tr.) one half to two-thirds length (exsag.) of medial part of inner pleural region; prominent spine on 15th thoracic axial ring of length (sag.) of entire prothorax, base of spine nearly as wide (tr.) as axis of segment; segments of opisthothorax with prominent pleurae; pygidium with length (sag.) equal to width (tr.); posterior mar- gin of pygidium weakly convex (only some of these thoracic characters, and none of the pygidial characters, could be determined in F. lapworthi; none of the pygidial characters could be determined in F. reticulatus). | |
| B. S. Lieberman 1999 | Length (sag.) of L4 equal to roughly 1.5 times length of L0 and L1 (sag.); anterior margins of frontal lobe at each side of midline deflected posteriorly at roughly 40 de- gree angle relative to transverse line; lateral margins of L4 distal to lateral margins of L0; preocular furrow on L4 directed inward and forward from glabellar margin; posterior margins of L4 divergent anteriorly; distal margins of L3 convex outward; middle sector of S3 convex anteriorly; L2 and L3 do not merge; distal margins of L2 diverging anteriorly; line between ends of S2 transverse; S2 convex anteriorly; ocular lobe with prominent ocu- lar furrow; line from posterior tip of ocular lobe to junction of posterior margin of lobe with glabella forms 10 to 20 degree angle with sagittal line; anterodistal margins of L3 formed by ocular lobes; intergenal angle developed posterior of point halfway between oc- ular lobes and genal spine angle; genal and intergenal ridges prominently developed; dis- tal sector of S0 with proximal end well posterior of distal end; axial part of L0 with node; posterior margin of L0 convex posteriorly; lateral lobes on L0 absent; thoracic pleural fur- rows extend onto spines; length (exsag.) of furrows equal to length of posterior band; an- terior margin of thoracic pleural furrow on T3 parallels a transverse line before flexing strongly posteriorly; long spine on 15th thoracic segment, spine broad (tr.) at its base; py- gidium twice as broad as long, posterior margin weakly convex. | |
| B. S. Lieberman 1999 | Bristoliinae is redefined as a
116 Peabody Museum Bulletin 45 monophyletic subfamily that includes only the genera Fremontella, Lochmanolenellus and Bristolia. This subfamily can be defined by the possession of the following diagnostic char- acters (thoracic characters only found in a few species of Bristolia, the only members of the subfamily for which thoracic remains exist): 1. anterior cephalic border developed as an el- evated ridge, and prominently separated from extraocular area by furrow; 2. anterolateral margins of frontal lobe not prominently separated from anterior extraocular area by furrow (not anterior border furrow); 3. prominent parafrontal band not visible in dorsal view; 4. anterolateral margins of frontal lobe at each side of midline deflected posteriorly at roughly 10 degree angle relative to transverse line; 5. S3 same depth laterally as medially; 6. an- terodistal margins of L3 formed by ocular lobes; 7. surface of interocular area slopes evenly from tip of ocular lobe to glabella; 8. genal spine angle developed opposite S0 or position anterior of S0; 9. posterior edge of ocular lobe developed anterior of or opposite medial part of distal margin of L1; 10. lateral lobes absent from L0; 11. distal sector of S0 transverse; 12. intergenal angle relative to transverse line directed anteriorly at least 35 degrees; 13. T3 macropleural, pleural spine of T3 projects further posteriorly than length of entire protho- rax; 14. anterior margin of thoracic pleural furrow on T3, when proceeding from proximal to distal edge, parallels a transverse line before flexing strongly posteriorly; 15. posterior margin of thoracic pleural furrow on T3 with medial part directed strongly posteriorly, dis- tal part parallel to a transverse line or weakly flexing anterolaterally; 16. thoracic pleural spines on segments 5 to 8 developed as short projections extending two thoracic segments back; 17. sharp boundary between thoracic pleural furrow and anterior band; 18. prominent spine present on 15th thoracic axial ring. | |
| B. S. Lieberman 1999 | Length (exsag.) of anterior cephalic border between lateral margins of L4 and genal spine angle equal to 50% to 60% length (sag.) of L0; L4 does not contact anterior border furrow and does not expand prominently dorsally; plectrum not visible in dorsal view; prominent parafrontal band visible, width (tr.) near ocular lobes equal to length (sag.) medially; ocular lobe only subtly separated from extraocular area by shelf; posterior end of ocular lobes extend back to S0; S3 jaggedly convex; S3, S2, S1 and S0 conjoined medially; S2 and S3 prominently incised; S1 transverse; intergenal angle represented by distinct dorsal swelling; posterior cephalic border between intergenal angle and genal spine transverse. | |
| B. S. Lieberman 1999 | Anterior cephalic border anterolateral of frontal lobe length (exsag.) 0.75 times length (exsag.) of L2 at distal tip; anterior cephalic border elevates from anterior border furrow at approximately 45 degree angle relative to dorsal flat plane, then intersects raised lineament, anterior of this point, border lies in flattened dorsal plane before deflecting ventrally; anterior part of frontal lobe does not intersect anterior border furrow; lateral margins of frontal lobe deflected as far laterally as lateral margins of L0; distal margins of glabella opposite L2 subparallel; S2 straight, not conjoined medially; lateral margins of glabella between posterior margin of L0 and point midway forward on distal tip of L1 subparallel; line from posterior tip of ocular lobe to junction of ocular lobes with glabella forms roughly 5 degree angle with sagittal line; S0 conjoined medially; lateral lobes pre- sent on L0; posterior margin of ocular lobes opposite medial part of distal margin of L0; glabellar furrows faintly incised; intergenal angle forms 30 to 35 degree angle with trans- verse line; genal spine length (exsag.) approximately length (sag.) of 8 to 10 thoracic seg- ments; pleural spine of T3 extends back 6 to 8 thoracic segments; anterior margin of T3 roughly parallels a transverse line medially; posterior margin of pleural furrow of T3 par- allels transverse line before flexing posterolaterally; thoracic pleural spines behind T3 ex- tend back 4 to 7 segments posteriorly; spines wide (tr.), one-half length (exsag.) of corre- sponding pleural segment at its medial part; width (tr.) of thoracic axis 100% of width of pleural field excluding spines; base of large axial spine on T15 less than half width (tr.) of axis of segment. | |
| B. S. Lieberman 1999 | Posteromedial part of L4 at same elevation as glabella at L3; entire margin of S3 arched furthest anteriorly at point midway between midline of glabella and axial furrows; medial and distal tips of S2 deflected as far forward anteriorly; S2 arching first anteriorly then posteriorly when going from medial to lateral tip; posterior margin of ocular lobes de- veloped opposite medial part of distal margin of L1; medial part of intergenal angle devel- oped at point about two-thirds of way between distal tip of ocular lobes and genal spine angle; intergenal angle forms 30 to 35 degree angle with transverse line; genal spine angle opposite distal edge of medial margin of L0. | |
| B. S. Lieberman 1999 | Plectrum present; medial part of intergenal angle arches anteriorly at point mid- way between distal tip of ocular lobe and genal spine angle; S2 does not contact axial fur- rows; medial part of cephalic posterior border flexes posteriorly; genal spine angle devel- oped opposite medial part of distal margin of L0; length (exsag.) of medial part of thoracic pleural furrows approximately 1.5 times length (exsag.) posterior pleural band. | |
| B. S. Lieberman 1999 | Anterior cephalic border between L4 and genal spine angle very short (exsag.), length equal one-half length (sag.) L0; genal spine angle opposite medial part of distal mar- gin of L1 or S2; intergenal angle forms roughly 50 to 60 degree angle with transverse line; slight ventral depression across entire margin of frontal lobe at point where ocular lobe in- tersects frontal lobe; lateral and distal tips of S3 as far forward; S2 convex or transverse, me- dial edge declined posteriorly relative to lateral edge, lateral edge generally not contacting axial furrows; line from posterior tip of ocular lobe to junction of posterior margin of lobe with glabella forms 10 degree angle relative to sagittal line; posterior edge of ocular lobe op- posite medial part of distal margin of L1; width (tr.) of extraocular area roughly equal to two-thirds width between distal most tips of ocular lobes; spine on T15 developed as long needle shaped projection. | |
| F. A. Sundberg 1999 | Glabella with a bluntly rounded to rounded frontal lobe. Anterior border concave, long (27 ± 3 percent cranidial length), wide (86 ± 7 percent cranidial width). Fixigena moderately wide (62 ± 4 percent glabellar width), posterior area of fixigena broad (51 ± 6 percent glabellar width). Circular medial swelling on the preglabellar area and crossing onto the anterior border. | |
| O. A. Hoel 1999 | The species differs from P. (B.) stenorhachis in the following features: Cranidium narrower in front of eyes. Glabella reaching to anterior border. Palpebral lobes smaller. Pygidium shorter, with dorsal furrows curving slightly outwards. Pleural platforms with half-rib and about six pleural ribs. | |
| F. A. Sundberg and L. B. McCollum 2000 | Cranidia with an anterior border slight- ly tapered distally, no medial inbend, poorly defined anterior border furrow. Glabella moderately elongated, strongly tapered from S0 to S2 then slightly tapered, S1 and S2 bifurcated; axial furrows deepest posteriorly. S0 moderately shallow. Posterior area of fixigena strap-like with sharp termination, border widens distally; border furrow extends to suture. Anterior branches of facial sutures slightly to strongly divergent to midlength (exsag.) of anterior border; posterior branches strongly divergent to pos- terior border furrow, then nearly parallel. Librigena spine flat- tened, broad-based. Pygidium suboval, micropygous; axis with one or two axial rings; poorly developed anterior pleural band; anterolateral corners rounded, opposite anterior half of the axis; posterior margin with median notch arched in posterior view; doublure absent behind axis. | |
| F. A. Sundberg and L. B. McCollum 2000 | Cranidium moderately convex; anterior margin narrow (width 59 | |
| F. A. Sundberg and L. B. McCollum 2000 | Cranidium with frontal area subequally divided (anterior border 60 | |
| F. A. Sundberg and L. B. McCollum 2000 | Cranidium slightly convex; anterior margin mod- erately narrow (width 71 | |
| F. A. Sundberg and L. B. McCollum 2000 | Cranidium moderately convex; anterior margin moderately wide (width 84 | |
| F. A. Sundberg and L. B. McCollum 2000 | Cranidium with glabella moderately inflated; gla- bellar furrows very shallow to absent. Axial furrows moderately shallow; convex and convergent from the S0 to S2 and then near- ly parallel to frontal lobe. Ocular ridges very faint to absent. Li- brigena with relatively long spine (approximately 40 percent li- brigenal length). | |
| F. A. Sundberg and L. B. McCollum 2000 | Cranidium with a preglabellar area 47 | |
| S. R. Westrop and E. Landing 2000 | A species of Protolenus in which strongly convexglabella lacks occipital spine. Anterior border occupies less than half of frontal area length. Fixigenae are wide, so that glabellar width is equal to no more than 40 percent of cranidial width between palpebral lobes. | |
| Geyer and Landing 2001 | The Agraulidae are difficult to characterize as a natural group, and the family requires a careful revision. General characters in this group include a subtrapezoidal outline and a low to moderate convexity of the glabella. The glabella has three pairs of short and relatively faint lateral glabellar furrows with S1 directed obliquely rearward and S2 and S3 roughly normal to the axis. Internal glabellar molds show a tendency to development of a crestline with nearly flat, lateral abaxial slopes. The rest of the cranidium and cephalon show a moderate convexity, with the cephalic regions defined by shallow and rather wide, often poorly defined furrows. Consequently, the glabella, although not very convex, is somewhat elevated above this low domal platform, and is laterally defined by shallow and moderately wide axial furrows. The palpebral lobes are quite small and located opposite the anterior half of the glabella; this morphology creates gently curved and distinctly extended wing-like posterior sectors of the fixigenae. The fixigenae are characteristically slightly inclined to the basal plane of the cephalon so that the posterior ends of the palpebral lobes are located slightly above the anterior tips. In addition, the facial suture curves abaxially before it reaches the posterior tips of the palpebral lobes so that a tiny subtriangular fixigenal area is abaxial to the posteriormost section of the palpebral lobes. | |
| M. A. S. McMenamin 2001 | As for genus. [A fallotaspidid trilobite with a wide cephalon, wide extraocular region, a plectrum, very long and slenderly tapering genal spines, distinct border furrow, tapered glabella, and relatively long frontal lobe (IA).] | |
| B. S. Lieberman 2001 | Anterior cephalic border developed as narrow rounded ridge, prominently separated from extraocular area by furrow; plectrum present; LA (L4) with length (sag.) roughly equal to length (sag.) of L0 and L1; ocular lobes gradually in- crease dorsoventral elevation between axial furrows and mid- point of ocular lobes, contact anterior and posterior parts of LA; width (tr.) of interocular area approximately equal to width of ocular lobe; anterior of eyes extraocular area greatly reduced (exsag.); anterodistal margins of L3 formed by axial furrows; S2 not conjoined medially; glabellar furrows weakly incised; width (tr.) extraocular region opposite L1 equal to roughly 35 percent width of glabella at L1; genal spine angle developed opposite medial part of distal margin of L0; intergenal angle developed posterior of lateral margins of ocular lobes. | |
| B. S. Lieberman 2001 | Anterior cephalic border prominently separated from extraocular area, very short (exsag.), length about one-half length (sag.) of L0; plectrum present; LA (L4) length about 1.1 times length (sag.) of L0; ocular lobes gradually increase dorso- ventral elevation between axial furrows and mid-point of ocular lobes; posterior tips of ocular lobes developed opposite medial part of distal margin of L1; extraocular region broad, width (tr.) opposite L1 more than 150 percent width of glabella at L1; genal spine angle developed opposite medial part of first thoracic seg- ment; medial part of cephalic posterior border between intergenal angle and L0 flexes posteriorly. | |
| B. S. Lieberman 2001 | Parafrontal band long (exsag.) anterior of anterolateral margins of LA (L4), 50 percent length (sag.) of L0; length (sag.) of LA (L4) equal to length of L0 and L1; outer band of ocular lobe near lateral margin of LA (L4) expands prominently exsagittally; ocular lobes of constant dorso-ventral elevation between axial furrows and mid-point of ocular lobes, contact anterior and posterior parts of LA; ocular lobe between visual surfaces and LA equal to approximately 33 percent width (tr.) of glabella at L1; distal margins of L3 convex outward; S3 jaggedly convex; distal margins of L2 diverge anteriorly; width (tr.) of extraocular region opposite L1 equal to approximately 125 percent width of glabella at L1; genal spine angle developed opposite medial part of distal margin of L0; intergenal angle directed anteriorly at roughly 30 degree angle; anteromedial margin of third thoracic pleural segment parallels transverse line; third thoracic segment macropleural; anteromedial margin of third thoracic pleural furrow directed weakly posteriorly; thoracic pleural spines on segments 5–8 developed as short projections extending three segments back; thoracic pleural furrows extend roughly width of inner pleural region.
| |
| B. S. Lieberman 2001 | Same as for genus. [Parafrontal band long (exsag.) anterior of anterolateral margins of LA (L4), 50 percent length (sag.) of L0; length (sag.) of LA (L4) equal to length of L0 and L1; outer band of ocular lobe near lateral margin of LA (L4) expands prominently exsagittally; ocular lobes of constant dorso-ventral elevation between axial furrows and mid-point of ocular lobes, contact anterior and posterior parts of LA; ocular lobe between visual surfaces and LA equal to approximately 33 percent width (tr.) of glabella at L1; distal margins of L3 convex outward; S3 jaggedly convex; distal margins of L2 diverge anteriorly; width (tr.) of extraocular region opposite L1 equal to approximately 125 percent width of glabella at L1; genal spine angle developed opposite medial part of distal margin of L0; intergenal angle directed anteriorly at roughly 30 degree angle; anteromedial margin of third thoracic pleural segment parallels transverse line; third thoracic segment macropleural; anteromedial margin of third thoracic pleural furrow directed weakly posteriorly; thoracic pleural spines on segments 5–8 developed as short projections extending three segments back; thoracic pleural furrows extend roughly width of in- ner pleural region.] | |
| B. S. Lieberman 2001 | Frontal lobe of glabella (LA) does not contact anterior border furrow; L4 (LA) moderately long (sag.), length equal to length (sag.) of L0 and L1; lateral margins of L4 (LA) directly anterior to lateral margins of L0; ocular lobes contact anterior and posterior parts of frontal lobe (LA); ocular lobes of constant dorso-ventral elevation between axial furrows and mid-point of ocular lobes; anterodistal margins of L3 formed by axial furrows; distal margins of L3 straight; S0 not conjoined medially; glabellar furrows weakly incised; intergenal spine not developed; thoracic pleural furrows extend only half width of inner pleural region, boundary between thoracic pleural furrow and anterior pleural band sharp; posteriormost segments of thorax merge with pygidium. | |
| M. A. S. McMenamin 2001 | A fallotaspidid trilobite with a wide cephalon, wide extraocular region, a plectrum, very long and slenderly tapering genal spines, distinct border furrow, tapered glabella, and relatively long frontal lobe (IA). | |
| M. A. S. McMenamin 2001 | A nevadioid trilobite with relatively narrow cephalon, narrow extraocular region, an advanced genal spine, prominent occipital spine, straight-sided to slightly concave glabella, and thin ocular lobes that reach back posteriorly to LO [occiptal ring]. | |
| M. A. S. McMenamin 2001 | As for genus. [A nevadioid trilobite with relatively narrow cephalon, narrow extraocular region, an advanced genal spine, prominent occipital spine, straight-sided to slightly concave glabella, and thin ocular lobes that reach back posteriorly to LO [occiptal ring].] | |
| A. Zylinska 2001 | A species of Leptoplastides with palpebral lobes situated opposite L2, two sinuous lateral glabellar furrows, anterior part of facial suture sub-parallel to slightly divergent in front of palpebral lobes, librigenae with medium sized slender spine, thorax with at least 11 segments, pygidium with two axial rings and two pairs of short, broadly based spines. | |
| A. Zylinska 2001 | A species of Leptoplastides with transglabellar lateral glabellar furrows, small palpebral lobes situated opposite L3, no eye-ridges, narrow fixigenae and librigenae with short spine. | |
| A. Zylinska 2001 | A species of Leptoplastides with small palpebral lobes situated opposite L3, three sinuous lateral glabellar furrows, sub-parallel anterior parts of facial suture, librigenae with short, slender spines, pygidium with two axial rings and at least two short, broadly-based pygidial spines. | |
| B. S. Lieberman 2002 | Frontal lobe of glabella does not contact anterior border furrow; plectrum present; anterior margins of frontal lobe at each side of midline deflected posteriorly at roughly 40 degree angle relative to transverse line; length (sag.) of preglabellar field equal to 0.25–0.7 times length (sag.) of L0; posterior tips of ocular lobes developed opposite medial part of distal margin of L0; ori- entation of S2 straight, abaxially directed inward and posteriorly at roughly 35–45 degree angle to transverse line; midinterocular ridge prominently developed, inclined at roughly 10 degree angle relative to sagittal line. | |
| F. A. Sundberg and L. B. McCollum 2002 | Cranidium with glabella elongated, strongly con vex, and deep axial furrows; preglabellar field short or absent; and anterior border straight or slightly curved. Pygidium trans versely elongate, subrectangular, posterior border nearly straight, wide; long, relatively narrow-based pygidial spines; wide posterior border; strong pleural bands, postaxial notch absent. | |
| D. S. Aristov 2002 | Head and eyes small. Pronotum large, longitudional, trapezoidal, widening distally. Length of ones in 1,3 times more than width. Paranota wide, narrowing toward to head, with cut in anterior margin. Anterior margin of forewing nearly straight, abruptly rounded near base of wing. SC in distal part of wing parallel to anterior margin. Basal part of CuA convex. | |
| D. S. Aristov 2002 | Head diametral, moderately large; eyes small, round; antennae thin. Pronotum smaller than head, rounded; paranota broad with cut in anterior margin. Mesonotum large, its width more than length; prescutum small. Metanotum similar to mesonotum, but smaller.
Forewing: Anterior margin convex, apex rounded, posterior margin convex. Costal area wide (three to four times wider than subcostal) with dense ridge of fore branches of SC. SC extend to distal thirds of wing. R nearly straight, RS originates near middle of wing, parallel to R, radial area very narrow. M dividing early, basal parts of M free. MA connected with origin of RS by strong crossvein, MAarea wide with slightly curved crossveins, MP closed together with CuA. CuA S-shaped, with hind branches in basal part of CuA-area. Cup weak, AI simple, almost straight. Hindwing: Anterior margin straight, apex rounded. Costal area two times wider than subcostal; SC ending in distal quarter of wing; CuA straight. Abdomen does not reach apex of wings, narrowing to apecal; ovipositor probably short. | |
| O. Béthoux and A. Nel 2002 | Forewing distinctly narrower than T. noinskii (width not exceeding 11 mm instead of 15 mm in T. noinskii); also branching of CuPa into CuPaα and CuPaβ opposite distal end of ScA on anterior margin. | |
| O. Béthoux and A. Nel 2002 | Forewing width exceeding 15 mm; CuPa branching into CuPaα and CuPaβ, basal of end of ScA on anterior wing margin | |
| O. Béthoux et al. 2002 | only the forewing characters are known, other parts of the body being unknown. Forewing narrow; network of irregular cross-veins covering the distal half of the wing, but variable in extension; variable irregular cross-veins between anterior margin and ScP after ScA reaches anterior margin; anterior branches of ScP long, some ramified, those of the distal half of ScP being curved towards the apex; RA posteriorly curved opposite the separation of MA and MP; the most basal anterior branchesof RAare basal to the fusion of RP with MA1; proximal part of RP very long; fusion of RP with MA1 long but variable; basal free part of MA1, basal of fusion with RP, short, shorter or equal to fusion; first branch of RP in a very distal position relative to the separation of MA1 and RP; MA1 distally ramified; MA2 posteriorly pectinate; area between MA/MA2 and MP wide; MP simple; base of MP close to the origin of CuA; area between MP and CuA + CuP1a narrow; basal free part of CuP1a twice the length of basal free part of MP before their fusion; branches of CuA + CuP1a reaching posterior margin obliquely; presence of subhorizontal cross-veins between branches of CuA + CuP1a, CuP2 and 1A. | |
| F. A. Sundberg and L. B. McCollum 2002 | Cranidia with moderately short glabella (66 ± 2 percent cranidial length), moderately deep lateral gla bellar furrows, moderately posteriorly directed ocular ridges, an terior border furrow evenly curved, concave to flat anterior bor der, frontal area long (34 ± 2 percent cranidial length) and mod erately wide (approximately 60 percent cranidial width), and mod erate size granules. Pygidium subrectangular; relatively wide; nearly straight sides; nearly level pleural field; scattered granules; sharp, moderately long lobes (147 ± 19 percent axial length); and 2 to 3 axial rings. | |
| B. S. Lieberman 2002 | Length (sag.) of preglabellar field roughly 0.9–1.2 times length (sag.) of L0; anterior tip of visual surfaces between S2 and S3; axial part of L0 with spine; length (exsag.) of pleural spine on T3 roughly equal to length (exsag.) of pleural spines on T2, T4–T6; prominent anterolateral nodes on thoracic axis. | |
| B. S. Lieberman 2002 | Preglabellar field short (sag.); anterior part of oc- ular lobe deflects strongly anteriorly near L3; anterior tips of vi- sual surfaces developed opposite adaxial part of S3; axial part of L0 with faint node; length (exsag.) of pleural spine on T3 roughly equal to length (sag.) of entire thorax; prominent anterolateral nodes on thoracic axis absent. | |
| F. A. Sundberg and L. B. McCollum 2002 | Cranidia with rectangular outline (approximately 60 percent width), preglabellar field present (25 to 30 percent cranidial length), straight anterior border in larger specimens, plectrum present, wide fixigena (45 to 60 percent glabellar width), strongly posterolaterally directed ocular ridges, coarse and fine granules. Pygidium subrectangular with 3 to 4 axial rings, large terminal piece not overhanging pygidial border, and longer (exsag.) pleural furrows that extends only a short distance on to relatively narrow-based spines | |
| F. A. Sundberg and L. B. McCollum 2002 | Cranidia with elongated glabella (73 ± 2 percent cranidial length), moderately deep lateral glabellar furrows, strongly to moderately posteriorly directed ocular ridges, anterior border furrow evenly curved, slightly convex anterior border, frontal area short (27 ± 2 percent cranidial length) and moder ately wide (60 to 65 percent cranidial width), and coarse granules. | |
| F. A. Sundberg and L. B. McCollum 2002 | Cranidia with moderately short glabella (64 ± 2 percent cranidial length), shallow to moderately deep lateral gla bellar furrows, moderately posteriorly directed ocular ridges, an terior border furrow evenly curved, concave anterior border, fron tal area long (36 ± 1 percent cranidial length) and moderately wide (64 ± 2 percent cranidial width), and moderate size gran ules. Pygidium subrectangular; relatively wide; nearly straight sides; nearly level pleural field; scattered granules; rounded, short lobes (141 ± 23 percent axial length); and 2 to 3 axial rings. | |
| F. A. Sundberg and L. B. McCollum 2002 | Cranidia with rectangular outline (length 50 to 60 percent width), no preglabellar field, straight anterior border in larger specimens, plectrum absent, wide fixigena (47 ± 4 percent glabellar width), moderately posterolaterally directed ocular ridg es, coarse and fine granules. Pygidium subrectangular with 3 axial rings, large terminal piece that overhanging the pygidial border, and longer (exsag.) pleural furrows that extend to nearly the tip of relatively broad-based spines. | |
| D. H. Kim et al. 2002 | A species of Eccaparadoxides with axis that occupies about 75 percent of pygidial length in large holaspids. S3 and S4 furrows shallow in late meraspids and holaspids. | |
| D. H. Kim et al. 2002 | A species of Eccaparadoxides with deeply incised S3 and S4 glabellar furrows and large median node on occipital ring. Anterior cranidial border narrows medially. Cranidium has coarsely granulose sculpture. | |
| D. H. Kim et al. 2002 | A species of Eccaparadoxides having raised ridges and irregular tubercles on L4 and frontal glabellar lobes; remainder of glabella with coarsely granulose sculpture. | |
| J. O. R. Ebbestad and G. E. Budd 2002 | A species of Burlingia with a broad (tr.) glabella having a rounded frontal glabellar lobe with laterally bulbous L4, maximum width at posterior part of cephalon with a posteriorly tapering body and a narrowly rectangular pygidium with a short distance between end of axis and posterior border. | |
| F. A. Sundberg and L.B. McCollum 2003 | "Cranidia with glabella with deep axial furrows that bow inwards in outline, and S1 and S2 lateral glabellar furrows moderately deep. Palpebral lobes slightly upturned and modrately long (averages 40-45 per cent glabellar lenght). Ocular ridge directed slightly posterolaterally (averages 75-80 degrees). Anterior border level; border furrow shallow to moderately shallow; weakly curved, and interborder furrow absent. Anterior branches of facial suture slightly bowed. Fixigena with anterior area slightly downsloping; posterior area of moderate width (with averages 75-90 per cent glabellar lenght),moderately short (averages 35-40 of basal glabellar lenght); lateral extension of posterior branches not parallel. Librigenae with border area having triangular cross-section, genal spine moderately long, straight, and with moderately wide base. Thoracic segments with well-developed fulcrum, short pleural spines. Pygidia with pleural furrows weakly developed; anterolateral corners rounded, no pygidial spines or lobes. Pustules absent." | |
| F. A. Sundberg and L.B. McCollum 2003 | Cranidium with slightly expanding glabella, pal pebral lobes moderately short, and posterior area of the fixigena tapered. Pygidium subrectangular with maximum width near an terior end of axis; pleural and axial ring furrows moderately strong; anterior border poorly defined, anterior border furrow ab sent; margin upturned; each anterior pleural band with long, up turned spines. | |
| O. Béthoux and A. Nel 2003 | ScA long parallel to anterior margin, distally vanishing in a pigmented (sclerotized?) apical area; area between anterior margin and ScA pigmented (sclerotized?); apex slightly pointed and slightly rejected posteriorly; ScP ending on RA just distal of midway of wing; few cross-veins between ScA and ScP, irregularly spaced, with one cross-vein just opposite end of ScP on RA; ScP and R basally posteriorly bowed; all cross-veins between RA and RP close to or distal of end of ScP on RA, few in apical area; RP with numerous branches, first posterior branch simple; MP with at least two branches; M and MP aligned and MA looking like an oblique secondary anterior branch; MA shortly connected with RP; origin of CuA from Cu distinct and long; Cu and CuP aligned and CuA looking like an oblique secondary anterior branch; CuA shortly connected with M; short basal oblique cross-vein opposite to a narrowing of area between CuA and CuP, just distal of connection of CuA with M; three more or less parallel series of gradate cross-veins between branches of RP, MA, branches of MP and CuA; less than ten pigmented spots, all roughly located basal of a cross-vein, in areas between main veins; apical and posterior wing margin more or less pigmented; cerci twice body length (only known in specimen MNHN-LPR. 52937, here assigned to Diaphanoptera incertae sedis). | |
| O. Béthoux and A. Nel 2003 | RP with 6–7 branches; MP with at least 4–5 branches. | |
| O. Béthoux and A. Nel 2003 | RP with five branches; MP with two branches | |
| P. Vršanský 2003 | Head more or less globular, antenna with wide segments and very long setae in transversal rows; pronotum mostly short and narrow, with reduced paranota; forewing heavily elytrised, with cup–like cells called "bunky" and reduced venation; hindwing venation generally of early polyphagoid groundplan (simplified venation with CuA without tertiary branches, CuP simple, A1 curved, with short branches), but with reduced venation of R (with few branches – possibly with pterostigma), M branched. Cerci with numerous long hair; females with short external ovipositor. | |
| P. Vršanský 2003 | Elytra not colored, radius poorly branched, fused with simple M; Cu strongly curved, S–shaped; anal veins multiple. All veins distinct. Pronotum roughly punctuate. | |
| P. Vršanský 2003 | Forewing only partially elytrised. Hindwing with numerous cross–veins, R reduced. Hind leg comparatively long, of the average roach appearance (not as short and thick as in the rest Umenocoleidae except Jantaropterix). | |
| P. Vršanský 2003 | Head not wider than pronotum (both with numerous short chaetae). Forewing well sclerotised (sometimes except apex), dark coloured with pale maculas, and pubescent throughout, with most veins distinct, Sc long and branched, R comparatively richly branched, M simplified up to 3 branches, Cu expanded, A multiple). Hindwing lacking pterostigma, with R1 apparently two–branched, and with cross–veins. | |
| P. Vršanský 2003 | Individuals very large, robust (unlike Petropterix), with very thick and heavily sclerotised elytra with apparent, reduced venation. Coloration with many (over 6) pale maculas. | |
| P. Vršanský 2003 | Very small species with surface of elytra, head, pronotum and body densely pubescent. Head possibly hypognathous, antenna thin. Legs thin and long, of the average roach appearance (not short and strong). Forewing with monochromatic coloration and venation indistinct (CuP may be exception). Hindwing with rich R1 (sharply curved) and RS, with pterostigma. Cross–veins present, but without numerous reticulations. | |
| F. A. Sundberg and L.B. McCollum 2003 | Antecior border moderate length (54 ± 5 percent of frontal area) and fixigena wide (52 ± 4 percent of glabellar width). | |
| F. A. Sundberg and L.B. McCollum 2003 | Same as for genus. [Cranidium with slightly expanding glabella, pal pebral lobes moderately short, and posterior area of the fixigena tapered. Pygidium subrectangular with maximum width near an terior end of axis; pleural and axial ring furrows moderately strong; anterior border poorly defined, anterior border furrow ab sent; margin upturned; each anterior pleural band with long, up turned spines.] | |
| F. A. Sundberg and L.B. McCollum 2003 | Cranidium with glabella that has poorly defined lateral glabellar furrows and small occipital spine; anterior border with flat lateral margins, narrows in front of glabella; posterior portions of fixigena parallel-sided. Surface punctate. Pygidium el- liptical shape with maximum width near posterior end of axis, pleural and axial ring furrows weak to effaced, posteromedial notch weak to absent, marginal spines absent. | |
| F. A. Sundberg and L.B. McCollum 2003 | Cranidium possibly with a small occipital node. Pygidial spines at posterolateral comers, straight or slightly curved posterior margin, posteromedial notch absent; axis wide; pleural bands distinct. longer laterally, level. Anterior border furrow absent at corners. Fixigena flat to slightly concave, level, width 35 to 50 percent glabellar width (K2). Palpebral lobes crescent shape, level, narrow and long, length 40 to 50 percent glabellar length; anterior margin located opposite S3 or L3. Ocular ridge short, weak, straight, directed very strongly posterolaterally from glabella at 35 to 40 degrees to axis. Posterior area of fixigena unknown.
Librigena moderate to large sized, length 6.1 to 9.7 mm (n = 2); wide, width 50 to 65 percent length without spine; lateral margin slightly curved. Genal field moderately convex. Border moderately convex, width approximately 25 percent librigenal width; border furrows moderately deep. Genal spine narrow, slightly curved, long, approximately 90 percent librigenal length. Rostral plate, hypostoma, and thorax unknown. Pygidium moderate size, length 4.3 to 6.0 mm (n = 3); sub- rectangular, length 75 to 85 percent width; moderately convex (sag. and trans.); posteromedial notch absent. Spines developed from lateral margin, narrow, sharp, directed posteriorly, upturned. Axis slightly tapered, anterior width approximately 45 percent pygidial width; length 75 to 85 percent pygidial length, postaxial ridge absent; three axial rings, strongly convex, axial nodes ab- sent; terminal axial piece small size, rounded; axial ring furrows moderately deep. Pleural regions with three anterior pleural fur- rows moderately wide, deep to shallow, extending to near pos- terior margin, interpleural furrows moderately strong to faint; first anterior pleural band moderately well developed, others moder- ately convex, posterior pleural bands weakly developed. Border defined as smooth area. Margin-parallel ridges on the occipital ring and librigenal bor- der. Genal caeca on genal area of the librigena and pleural area of pygidium. | |
| F. A. Sundberg and L.B. McCollum 2003 | ranidium with anterior border moderately wide (width 63 ::!::4 percent cranidial width), moderately long, mod- erate curved, not arched dorsally, and moderately convex; ocular ridges slightly posterolaterally directed; fixigena moderately wide (width 66 ::!::6 percent glabellar width); palpebral lobes moder- ately long (length 40 ::!::3 percent glabellar length). Pygidium wide (length 29 ::!::3 percent width). | |
| F. A. Sundberg and L.B. McCollum 2003 | Cranidium slightly convex; anterior margin mod- erately narrow (67 ± 12 percent cranidial width); preglabellar area short (30 ± 5 percent frontal area length); anterior border slightly convex, furrow without inbending, medial swelling ab- sent; anterior branches of facial suture relatively short; glabellar length 70 ± 2 percent cranidial length, convexity low, lateral glabellar furrows moderately defined; axial furrows straight; oc- cipital node absent; ocular ridge moderately defined; posterior area of fixigena sharply terminated, directed slightly posteriorly; posterior border convex. Librigena with moderately narrow genal area, moderately defined border furrows, interborder furrow pre- sent, librigenal spine long (55 to 60 percent librigenal length), and recurved. Pygidium suboval, moderately arched, axis has two axial rings, plural bands poorly defined, anterolateral comers moderately rounded, and posteromedial notch well developed. Fine granular ornamentation on glabella, fixigena, and pygidium. | |
| F. A. Sundberg and L.B. McCollum 2003 | Cranidium pentagonal in outline with moderately low relief; anterior branches of facial suture parallel to mod~rately convergent; posterior branch slightly divergent, nearly parallel; glabella moderately to strongly tapered, frontal lobe bluntly rounded, straight or slightly convex glabellar outline, lateral gla- bellar furrows and axial furrows nearly effaced; anterior border narrow, upturned; border furrow wide; palpebral lobes moderately long, posteriorly placed; fixigena slightly convex, level; anterior area moderately downsloping, posterior area extending slightly past palpebral lobes. Exoskeleton smooth. | |
| F. A. Sundberg and L.B. McCollum 2003 | Same as for genus. [Cranidium pentagonal in outline with moderately low relief; anterior branches of facial suture parallel to mod~rately convergent; posterior branch slightly divergent, nearly parallel; glabella moderately to strongly tapered, frontal lobe bluntly rounded, straight or slightly convex glabellar outline, lateral gla- bellar furrows and axial furrows nearly effaced; anterior border narrow, upturned; border furrow wide; palpebral lobes moderately long, posteriorly placed; fixigena slightly convex, level; anterior area moderately downsloping, posterior area extending slightly past palpebral lobes. Exoskeleton smooth.] | |
| B. Waggoner 2003 | As for type species. [Relatively small aglaspidids with highly arched axial region, no genal spines, very short pleural spines, large smooth eyes relatively far apart.] | |
| B. Waggoner 2003 | Relatively small aglaspidids with highly arched axial region, no genal spines, very short pleural spines, large smooth eyes relatively far apart. | |
| F. A. Sundberg and L.B. McCollum 2003 | Cranidium with a parallel-sided glabella, S1 transglabellar furrow, eye ridges directed moderately posterolaterally, moderately wide ®xigena (71 6 8 per cent glabellar width), intergenal spines, slightly curved anterior margin, and anterior border without curvature towards frontal lobe. Twelve thoracic segments, each with flat pleural spines. Pygidium with two axial rings; anterior pleural bands long (exsag.), posterolaterally to posteriorly directed with flat spines; and V1 to V2 spines decreasing in size and projecting posteriorly. | |
| F. A. Sundberg and L.B. McCollum 2003 | Cranidium has shallow lateral glabellar furrows, axial furrows, anterior border furrow, median inbend of the anterior border, and wide palpebral lobes. | |
| F. A. Sundberg and L.B. McCollum 2003 | Cranidium with glabella narrow and slightly tapered; anterior margin of palpebral lobes adjacent to L3 or S3; fixigena moderately wide; anterior of ®xigena strongly downsloping; granular surface. Librigena with genal spine. | |
| F. A. Sundberg and L.B. McCollum 2003 | Cranidium with a nearly straight, slightly convex anterior border with shallow anterior border furrow. | |
| P. Vršanský 2004 | Medium-sized species. Antenna with numerous strong and long chaeta on each segment. Pronotum much wider than long, with pale margins. Cercus thin and long, with very strong and long sensilla chaetica (even in initial immature stages). Legs with strong spines. | |
| P. Vršanský 2004 | Large species with rich venation, distinct intercalaries and cross-veins. Sc branched in the forewing, RS differentiated, cerci strong and short. Pronotum very wide. | |
| P. Vršanský 2004 | Small species. Wing rather wide (apomorphy), with venation reduced compared with other Cretaceous blattellids. Rs differentiated. Cu reduced to several veins, branched at the terminal fourth of their length. Clavus is extremely long, reaching almost the half of the wing length, A simple. | |
| P. Vršanský 2004 | Large cockroach (known species with wing about 22.5 mm long). Wing rather narrow, with Sc simple, R and A systems rich, R with several terminal veinlets (plesiomorphy), M reduced to several veins, Cu expanded, clavus of ordinary length, some anal veins branched. | |
| G. Geyer and E. Landing 2004 | Species of Hamatolenus (Hamatolenus) with glabella faintly tapering forward in the posterior third, progressively tapering anteriorly; occipital ring with median node; parafrontal band relatively narrow; eye ridges almost straight, slightly thinner and more elevated than palpebral lobes; palpebral lobes clearly longer (>1.2 times) than anterior branches of facial suture, but posterior tips clearly distant from posterior border furrow; preglabellar field moderately narrow; anterior border barely wider than preglabellar field, modestly elevated. | |
| L. Amati and S. R. Westrop 2004 | A species of Thaleops with strong longitudinal convexity. Muscle scar attachment sites well defined on internal moulds as two longitudinal ridges along the lateral margins of the glabella with a shallow depression between. Relict segmentation present on ventral surface of pygidial axis. Anterior margin of pygidial doublure with second, smaller pair of forward projections between larger pair of forward projections on either side of axis. | |
| T. P. Fletcher et al. 2005 | Glabella widening markedly to L4, with anterior margin less than semicircular; S0-S2 transglabellar, S3 and S4 short, shallow; anterior cranidial border flat, not well defined by impressed border furrows, widening (sag.) markedly abaxially; in more mature stages, border furrows barely discernible. | |
| J. R. Pollitt et al. 2005 | Moderately long preglabellar field (character 1). Longitudinal furrow can be effaced but, if not, is not continuous with S0 (character 2). S1 conjoins to S2 to circumscribe the lobe L2, may be effaced (character 3). Median glabella does not overhang the anterior border (character 7). Glabella lobe is L2 (character 10). Sagittal convexity of cranidium is weak (character 20). Path of anterior section of facial suture is strongly divergent (except in Acidaspidella, where it is subparallel: character 21). No interpleural furrows are present (character 35). All taxa have more than one complete pygidial inter-ring furrow (character 39). | |
| R. G. Martins-Neto et al. 2005 | Apart from the characteristics Triassic blattopteran insects from Argentina 719 discussed by Vischnyakova (1982), tegmen with RP and MA secondary branches restricted to the apical area, above the apex. RA and CuP distal extremity perpendicularly aligned to both costal and anal margin and MA origin after this alignment (lcm = aml < oMA ). Two-branched MA | |
| R. G. Martins-Neto and O. F. Gallego 2005 | Elongated tegmen, with multibranched ScP , with the last distal branches reaching the costal margin at the same level of CuP distal extremity; unbranched RA ; short RP , sigmoid, reaching the costal margin at around 2/3 of the tegmen length; CuA branches bend strongly backwards, running subparallel to each other and the posterior wing border. | |
| R. G. Martins-Neto et al. 2005 | Tegmen with MA origin and CuP distal extremity perpendicularly aligned to both costal and anal margins and the distal part of the last RA branch oblique to the MA / CuP alignment, at an angle around 45º ( oMA = aml < lcm ). CuA secondary branches well marked. Number of MP secondary branches reduced. | |
| R. G. Martins-Neto and O. F. Gallego 2005 | Elongated tegmen with narrow costal area and multibranched ScP , whose last branch reaching 1/3 of the wing length at the same level of CuP distal extremity; no pronounced RA ; branched MA ; twobranched MP multiple forked; CuA branches arise by forking. CuP origin at R , relatively far from the tegmen base. Anal area occupying around 70% of the wing width. No cross venation present | |
| R. G. Martins-Neto and O. F. Gallego 2005 | RA reaching the costal margin circa 1/3 of apex. MA origin at 1/3 of the tegmen base. Anal area occupying around 40% of the tegmen length and 50% of its width | |
| R. G. Martins-Neto et al. 2005 | Small sized tegmen. Anal area occupying more than a half of the total wing length and anal veins multibranched. | |
| R. G. Martins-Neto et al. 2005 | As for the family. Tegmen three times longer than wide. ScP long and unbranched. RP apparently twobranched. | |
| J. S. Hollingsworth 2005 | An olenellid genus with LA rounded forward, reaching border furrow without preglabellar field. LA approximately equal in width (tr.) to L0; L3 wider than LA. Glabella has distinctly impressed furrows; S2 slightly curved forward. Ocular lobes long, nearly reaching poste- rior border furrow, with distinct ocular furrows; lobes strongly curved with small interocular area. Intergenal spine small, posterlaterally directed, located one-half to two- thirds distance from axial furrow to genal spine angle; inter- genal ridge lacking. Hypostome has five pairs of short pos- terolateral spines. Thoracic axis narrow, less than twice the width of adjacent inner pleural region. T15 has a large axial spine; two thoracic segments posterior to T15. Pygidium elongate. | |
| J. S. Hollingsworth 2005 | Same as the generic diagnosis [An olenellid genus with LA rounded forward, reaching border furrow without preglabellar field. LA approximately equal in width (tr.) to L0; L3 wider than LA. Glabella has distinctly impressed furrows; S2 slightly curved forward. Ocular lobes long, nearly reaching poste- rior border furrow, with distinct ocular furrows; lobes strongly curved with small interocular area. Intergenal spine small, posterlaterally directed, located one-half to two- thirds distance from axial furrow to genal spine angle; inter- genal ridge lacking. Hypostome has five pairs of short pos- terolateral spines. Thoracic axis narrow, less than twice the width of adjacent inner pleural region. T15 has a large axial spine; two thoracic segments posterior to T15. Pygidium elongate.] | |
| J. S. Hollingsworth 2005 | species of Polliaxis with glabella strongly inflated transversely and longitudinally, widest at L3. Eye ridges distinct, nearly as high as palpebral lobe. Thorax long, slightly tapered, 22 segments, axial rings have medial spine. Pleural spines flattened, triangular, slightly curved posteriorly, and directed posterolaterally. Pygidium with wide strongly tapered axis and notably elevated rounded trian- gular terminal piece. | |
| J. S. Hollingsworth 2005 | A species of Keeleaspis? with tapered gla- bella rounded anteriorly, which extends into border furrow. Glabellar furrows deeply incised, not parallel, S3 nearly transverse, S1 curved posteriorly nearly intersecting S0. Palpebral lobe short, one-third of glabellar length and mid- line of lobes is at posterior third of glabella. Thorax at least 27 segments, fewer on small specimens. Pygidium tiny with
two axial segments and narrow pleural areas. Exoskeleton covered with spinules. | |
| T. P. Fletcher et al. 2005 | Exoskeleton narrow (Fig. 6.1) or broad (Fig. 6.4), up to 27 cm long; glabella with an evenly rounded anterior margin; palpebral lobes medium-sized; genal spines long, extending posteriorly as far as the tenth thoracic segment; thorax of 17 or 18 segments of which the last four pleural spines commonly extend beyond the posterior pygidial margin and gradually decrease in length; pygidium subcircular, subquadrate, with one well-defined ring on a relatively long, bluntly pointed, triangulate axis | |
| J. S. Hollingsworth 2006 | Holmiid with genal angle slightly to
strongly advanced, genal spine wide at base. Intergenal spine well developed, located proximal to midlength of posterior margin. Anterior and lateral borders well defined. Glabella slightly ex- panded anteriorly, slightly narrowed at S1, reaching or nearly reaching anterior border, L1 to L3 similar in shape, glabellar fur- rows not joined at axis. L0 with spine or node. Posterior tips of ocular lobes align with L1 or anterior of L0. Extraocular area between one and two times width of interocular area. Exoskeleton smooth to strongly granular. Thorax narrow, with 17 segments, inner pleural region narrower (tr.) than axis, pleural spines short, thornlike, each segment with an axial spine. Pygidium small, ex- panded posteriorly. | |
| J. S. Hollingsworth 2006 | Holmiid with advanced genal spines ini- tially directed outward to slightly forward from anterolateral mar- gin of cephalon, posterolateral margin rounded, intergenal spines or node a short distance in from posterolateral corner, or may be absent. Glabellar sides concave, LA expanded and front broadly curved against anterior border furrow. Ocular lobes long. Extra- ocular area ranges from wider than interocular area to much less than interocular area width. Small occipital spine present on some species. Thorax of 17 segments, pleural regions slightly narrower than axis. Pygidium large for a holmiid, wide, one axial ring clearly defined and a second defined by pits and a shallow furrow. Posterior margin transverse, bearing short spines. | |
| V. Kordule 2006 | Glabella tapering forward, with weakly rounded anterior; four pairs of glabellar furrows; cephalic border prominent, rather long (sag.); anterior border furrow deep; genal spine prominent, short, rapidly tapering, extending backward toward the third segment; genal caeca prominent on external surface of preglabellar field, preocular field of fixigenae and librigenal field of librigenae; pygidium with 5 to 6 axial rings and terminal piece; pleural lobes with distinct pleural and interpleural furrows; exoskeleton surface with finely tuberculate sculpture. | |
| Z. Zhu and S. Peng 2006 | Koldinioidia with stout occipital spine, occipital furrow faint and straight; thorax of five segments with axial spine on the third thoracic segment. | |
| D. S. Aristov et al. 2006 | Pronotum trapezoid, slightly wider than long, without paranota, as broad as head and mesonotum. Wing sheaths long, with poorly visible traces of venation. Middle and hind legs equally long, fore legs slightly shorter. Middle and hind tibiae directed backward. Abdominal tergites with median crest, broad pointed paranota and rather long unsegmented cerci. | |
| D. S. Aristov et al. 2006 | Pronotum with paranota twice as wide as head and as wide as mesonotum. Pronotal paranota without incisions on their anterior margins, their lateral margins broadly rounded. Middle tibiae directed backward, tibiae armed and not broadened apically, tarsus short with two to five tarsomeres. Abdominal tergites with paranota, body colored. Length of nymph body, 4.5–43 mm; length of subimago, 41 mm; and length of imago, 47.1–49.8 mm | |
| D. S. Aristov et al. 2006 | Pronotum with paranota twice as broad as head and as broad as mesonotum. Each pronotal paranotum with incision on its fore margin. Fore femora thickened; tibiae unarmed, fore and middle tibiae directed forward, hind tibiae directed backward. Fore and middle tarsi unsegmented, hind tarsus 2-segmented. Body length, 13–14 mm. | |
| D. S. Aristov et al. 2006 | Pronotum with paranota twice as broad as head and as broad as mesonotum. Each pronotal paranotum with incision on its fore margin. Tibiae unarmed, all three pairs directed forward; tibiae and femora with hairs; tarsus shortened, most probably 3-segmented. Coxal gills visible near fore and hind coxae. Abdomen somewhat narrowed near its base, abdominal paranota narrow. Body length, 14.5 mm. | |
| D. S. Aristov et al. 2006 | Pronotum with paranota as long as wide, less than twice as broad as head and as broad as mesonotum. Paranota narrow. Hind legs longer than middle legs. Middle and hind tibiae armed and directed backwards; middle tibia longer than 3-segmented tarsus; first tarsomere longer than third. Abdominal tergites narrow, with small paranota. Body length, 20 mm. | |
| R. G. Martins-Neto et al. 2006 | Fore wing with costal area proximally broad; forking of RA and RP relatively close to base of wing, within the proximal third (plesiomorphy); MP anteriorly close to apex, CuA more [Saaloscytina] or less [Chanarescytina] resolved into reticulation over the whole length (apomorphy); sculpture coarsely reticulate. | |
| R. G. Martins-Neto et al. 2006 | Costal area relatively narrow (slightly narrower than in Saaloscytina). RP bipartite. Supporting veins r-m and m-cu well defined (apomorphy). | |
| R. G. Martins-Neto et al. 2006 | Polycytellid fore wing small-sized (length about 3 mm), venation reduced: RA hidden
in a slightly zigzagged line between the two rows of cells close to the anterior margin. RP and MA unbranched, MA originating from RP approximately in the mid-length of the wing | |
| D. S. Aristov et al. 2006 | Nymphs with slender body. Head large with long antennae. Pronotum as wide as head, transverse, with narrow paranota. Mesonotum somewhat wider than pronotum, trapezoid; metanotum equal to mesonotum. Legs elongated and lender, middle andhind tibiae directed backward, tarsus long, 5-segmented. Abdominal tergites without paranota, cerci very long. | |
| D. S. Aristov et al. 2006 | Pronotum with paranota as broad as
mesonotum and less than twice as broad as head. Pronotum elongate, slightly broadening to base. All three pairs of legs approximately equally long. Middle and hind tibiae directed backward; tarsus shortened, 5-segmented. Abdominal tergites with narrow paranota | |
| D. S. Aristov et al. 2006 | Head large with small eyes. Pronotum with paranota as broad as mesonotum and less than twice as broad as head. Pronotum quadrate, paranota narrow. Fore and middle legs of approximately equal length, hind legs noticeably longer. Middle and hind tibiae directed backward, tarsus 5-segmented. Abdominal tergites with narrow paranota. | |
| D. S. Aristov et al. 2006 | Head large, with thick antennae. Pronotum with paranota narrowing backward, less than twice as broad as head and as broad as mesonotum. Middle and hind tibiae unarmed, directed backward; middle tarsus as long as tibia; tarsus 3-segmented, first tarsomere as long as third. Abdominal tergites without paranota. | |
| D. S. Aristov et al. 2006 | Pronotum broadening forward, pronotum with paranota broader than mesonotum. Hind tibiae directed backwards, unarmed. Tarsus shortened, 3-segmented; first two tarsomeres short and equal to each other. Abdominal tergites with paranota. Body length, about 18 mm. | |
| D. S. Aristov et al. 2006 | Pronotum without incision on its fore margin, pronotum with paranota less than twice as broad as head and as broad as mesonotum. All three pairs of tibiae directed forward; tarsi shortened, with three subequal segments. Abdominal tergites with paranota. Body length, 18.5 mm | |
| R. G. Martins-Neto and O. F. Gallego 2006 | Fore wing with RA and RP distally fused; ra-rp absent; unbranched RA; RP with two or
three secondary branches. Genera included: Dysmorphoptilioides EVANS, 1956 (Triassic Australia), Gallegomorphoptila MARTINS-NETO in MARTINS-NETO, GALLEGO & MELCHOR 2003), (Triassic, Argentina) and Bandelnielsenia MARTINS-NETO et GALLEGO in MARTINSNETO, GALLEGO &MELCHOR 2003 (Triassic, Chile). | |
| J. S. Hollingsworth 2006 | Holmiid with preglabellar field. Glabella narrow,
LA equal to or slightly wider than L0, S3 faint, L3 shorter (exsag.) than L2, L1, or L0 which are about equal, L0 with pos- terior axial spine, axial furrow uniform. Ocular lobes long. Ex- traocular area wide. Reticulate sculpture prominent on larger ce- phala. Thorax of 17 segments, T3 mildly amplipleural, pleural spines weakly thornlike to sentate. T13 has long, posteriorly di- rected axial spine; posterior to T13, thorax strongly tapered. Py- gidium expands backward, posterior margin straight, posterolateral corners rounded. Axis triangular to subtriangular, not reaching posterior margin. | |
| J. S. Hollingsworth 2006 | Holmiid with LA impinging on the anterior bor- der. Lateral cephalic border wide, one-third to over two-thirds width (tr.) of L0. S3 furrow faint or obsolete, and L3 incorporated in dorsally inflated LA. Glabella wider at LA than at L0 and constricted at S1. L0 expands posteriorly into stout spine. Ocular lobes short with posterior tips opposite anterior of L1 or posterior of L2. Extraocular area one to one and one-half times width (tr.) of interocular area. Intergenal spine located closer to axial furrow than to genal angle. Dorsal surface granular. Thorax of 17 seg- ments, pygidium unknown. | |
| J. S. Hollingsworth 2006 | Cephalon with strongly advanced genal spines, and a small node on L0. Thorax narrow, widest at T5. | |
| J. S. Hollingsworth 2006 | Glabella parallel-sided from L0 through L2. Pos- terior margin angled back from axial furrow to intergenal spine, then distinctly curved forward to genal angle. Genal spine mod- erately advanced, aligning with mid-L0 to S0, and initially slight- ly outward-directed curving to straight back. | |
| J. S. Hollingsworth 2006 | Cephalon slightly subtrapezoidal, narrowing for- ward, with rounded posterolateral margins, intergenal spine or node absent. Genal spine, long, advanced to align with anterior quarter of LA, slightly forward-directed at base, then curved backward. Ocular lobes long. Extraocular area narrower than in- terocular area. | |
| J. S. Hollingsworth 2006 | Cephalon subtrapezoidal with rounded posterolat- eral margins. Genal spine advanced to anterior portion of ce- phalon, initially forward-directed then curving outward to straight backward to a length greater than length of entire carapace. Gla- bella slightly expanded anteriorly with shallow S3 furrow. Ocular lobes very close to glabella; extraocular area is about 2.5 times width (tr.) of interocular area. Pygidium wide (tr.), more than twice pygidial length, pygidial axis narrow, about one-fourth pygidial width. | |
| V. Kordule 2006 | Ptychoparia with S1, S2, S3 and S4 deeply incised and clearly defined; adaxial portion of S1 forked; anterior border prominent, with average LCr/LAB cca 6.0; genal caeca with anastomosing canals very prominent in external surface of the cephalon; paradoublural furrow distinctly separated from border furrow; distal ends of pleurae extended into short spines in all segments; pygidial axis with six axial rings and terminal piece; pygidial pleural fields with five pairs of ribs and rudimentary sixth rib; exoskeleton generally larger than in P. striata. | |
| V. Kordule 2006 | Glabella tapering forward, with weakly rounded anterior; four pairs of glabellar furrows; cephalic border short (sag.); anterior border furrow deep; genal spine conspicuous, long, weakly tapering; genal caeca weakly defined on external surface of preglabelar field, preocular field of fixigenae and librigenal field of librigenae; pygidium with six to eight rings and terminal piece, pleural field with distinct pleural furrows; inter pleural furrows obscure or weakly defined; surface of cephalon with minute tubercles. | |
| V. Kordule 2006 | Mikaparia with subtrapezoidal, forward tapering and anteriorly rounded glabella; S1 long, strongly curved backward, S2, and S3 shorter and shallower, S3 almost normal to sagittal axis; S4 weekly impressed; anterior border conspicuous, short (sag.), gently convex; average LCr/LAB cca 10.0; pygidial axis with six rings and sub triangular terminal piece; five ribs on pleural field. | |
| V. Kordule 2006 | Glabella highly convex, bluntly conical, anteriorly rounded, extending nearly toward border furrow; axial furrows deep; occipital ring vaulted, with small median node; S1, S2 and S3 shallow; preglabellar field flat; anterior border short (sag.), high, convex, separated from glabella by deeply concave border furrow; ocular ridges prominent; palpebral lobes small; genal caeca obscure; genal spine short; thorax of 14 segments; pygidium transverse, with axis with three to five rings and large terminal piece; pleural furrows present, inter pleural furrows absent; surface of exoskeleton finely tuberculate. | |
| V. Kordule 2006 | Ptychoparioides with highly convex, forward rapidly tapering glabella, which is distinctly higher than fixigenae; S1 and S2 are moderately deep and inclined backward, S3 and S4 are short, shallow, transverse; preglabelar field flat, very short; anterior border short, evenly broad (sag.), weakly convex, with edge directed anterodorsally, separated from preglabelar field by broadly concave border furrow; ocular ridges conspicuous, almost straight, strongly inclined backward towards small palpebral lobes; pygidium transverse, with axis 30–35% of pygidial width; five axial rings and terminal piece; two distinct and third weakly defined pleural furrows in lateral pygidial lobe; inter pleural furrow weakly defined between the first and second pleura; third and fourth ribs poorly defined. | |
| V. Kordule 2006 | Ptychoparioides with slightly convex glabella, weakly tapering forward as high as fixigena, with evenly rounded anterior lobe; S1 and S2 are shallow, inclined backward, S3 and S4 are short, shallow, transverse; preglabelar field flat, very short; anterior border short, evenly broad (sag.), weakly convex anteriorly and dorsally, with edge directed dorsally, separated from preglabelar field by broadly concave border furrow; ocular ridges conspicuous, weakly curved and inclined backward towards small palpebral lobes; pygidium short (sag.) transverse, with axis cca 25% of pygidial width; three axial rings and terminal piece; two distinct and third weakly defined pleural furrows in lateral pygidial lobe; interpleural furrow weakly defined between the first and second pleura; third and fourth ribs poorly defined. | |
| J. S. Hollingsworth 2006 | Sagittal length of preglabellar field equal to or slightly less than sagittal length of anterior border. Glabella typically parallel-sided with LA equal in width (tr.) to L0, anterior of LA rounded to somewhat pointed. L3 wider (tr.) than L2, S2 short and faint or absent at axial furrow. Ocular lobes gently curved, close to axial furrow. Narrow occipital spine arises as ridge on posterior half of L0. Axial rings of thorax have small longitudinal ridge on posterior half. Lateral margins of pleural spines backward-directed. | |
| J. S. Hollingsworth 2006 | Preglabellar field equal to sagittal length of anterior border on small specimens ranging to virtually absent on large specimens. Glabella slightly narrowed at S2. LA broadly rounded anteriorly and slightly wider (tr.) than L0. Ocular lobes semicircular. Occipital spine arises at posterior margin of L0. Ax- ial rings of thorax have nodes or small spines. Lateral margins of pleural spines outward-directed, about 45 | |
| N. E. Vaccari et al. 2006 | Endymioniinae with cephalic doublure extended forward and slightly downward into long, sharp, pointed spine; glabella strongly inflated; preglabellar field pointed forward and arched medially. Pygidium with axis extended on to posterior border; border wide, steeply down-turned. | |
| K. J. McNamara and R. Feist 2006 | Small scutelluine in which anterior margin of frontal lobe in contact with convex anterior border; glabellar fur- rows, especially S2 and S3, isolated from each other and very weakly developed; occipital ring very long, two-fifths glabellar length, with short occipital spine in small holaspids; eye lobe small to very small; no genal spine. Pygidium extremely vaulted, height up to 60% of length; bell-shaped in posterior view; short, up to 1.4–1.9 times wider than long; axis prominent, with small anterolateral lobes, inconspicuous medial lobe; well-incised, broad axial furrow; fulcrum approximately midway between axial furrow and lateral margin; median rib not divided, broad adaxi- ally; interrib furrows narrow and deep; smooth, flat border well developed with entire margin, making angular junction with very convex pleural field. | |
| R. A. Fortey 2006 | Trinucleus species with 50 or more radial sulci on the relatively narrow (sag.) fringe; dorsal surface sculpture comprising fine reticulum; pygidium with up to ten pleural ribs. | |
| F. A. Sundberg 2006 | Glabella expanding forward (120- 135% basal glabellar width), frontal lobe moderately convex with no medial sulcus, S1 transglabellar furrow effaced to moderately developed, S2 and S3 very shallow, connected to axial furrows. Ocular ridges effaced, moderately posterolaterally directed (75°), palpebral lobe length 25-40% glabellar length. Plectrum absent, anterior border horizontal. Pygidium with spines on the anterior three or four pleural segments, no curvature of anterior border of pygidium, pleural furrows shallow to moderately deep, reaching margin; with 13-15 pleural ribs, anterior pleural ribs with slight backward curvature of the distal portion, posterior pleural rib moderately narrow with rib width 6 ± 1% pygidial width. Pygidial axis moderately long (75 ± 4% pygidial length), with 5-6 axial rings. | |
| A. I. Varlamov et al. 2006 | Dorsum (D) elliptical, 5–6 cm long.
C broadly semicircular, approximately 0.33 times as long as D; length of cranidium (Cr) equal or less than its width at palpebra (Pal). Arculum (Ar) convex, rarely, weakly convex and flattened. Anterior branches of facial sutures sharply diverging. Librigenae broad, with equal posterior and lateral borders; long genal spines developed at posterolateral corners. Thorax (T) consisting of 16 segments, with distinct fulcral lines gently arcuate outward. Axis of Pyg having 4 to 7 rings, posterolateral parts of borders of Pyg broadened into backward oriented triangles, serving as bases for a pair of rather large flattened spines. | |
| M. Mergl 2006 | Orometopid with high, anteriorly widening, subglobose glabella; large palpebral lobes highly raised above fi xigena; librigenae with very high, subvertical fringe with prominent crests on inner and outer margins; long posterodorsally directed occipital spine; pygidium without border, border furrow, or pleural furrows; pygidial axis smooth, with rings distinct only on inner surface; weak posterior arch present. | |
| V. Kordule 2006 | Mikaparia with subtrapezoidal, forward tapering and anteriorly rounded glabella; S1 long, strongly curved backward, S2 and S3 shorter and shallower, S3 almost normal to sagittal axis; S4 weekly impressed; anterior border narrow (sag.), gently convex; average LCr/LAB cca 9.0; pygidial axis with seven to eight rings and subtriangular terminal piece; six ribs on pleural field; interpleural furrows absent; genal caeca obscure on external surface of cephalon. | |
| L. E. Babcock and J. S. Peel 2007 | Glabella slightly tapering forward; ocular lobe relatively small, posterior opposite L1; interocular area narrower than extraocular area; genal spine short, rarely extending posteriorly beyond the first thoracic segment. Thorax with 17-18 segments; posterior segment may be fused with anterior part of simple pygidium; pleural spines short (exsag.), slender. | |
| L. E. Babcock and J. S. Peel 2007 | Cephalon semicircular, width about 1.5 to 2.25 length; ocular lobe smoothly semicircular to slightly abaxially geniculated in posterior one-third; width (tr.) of interocular area slightly narrower than extraocular area; genal area of variable width. Thorax variable but moderately wide, about 0.5 to 0.6 of maximum sagittal length, widening slightly backward to about 6th segment, then tapering posteriorly; inner pleural regions slightly wider (tr.) than axis. | |
| J. S. Hollingsworth 2007 | An archaeaspidid with cephalon rounded posterolaterally, genal spine small or absent. Cephalon and glabella only gently arched dorsally. Preglabellar field wide (sag.). Interocular area variably sloped dorsally, width (tr.) about equal to extraocular area width. | |
| J. S. Hollingsworth 2007 | Fritzaspis with wide, slightly convex anterior and lateral border. Preglabellar field wider than anterior border. Glabella moderately tapered. SO distinct at axial furrow, crosses axis as broad shallow depression. LO only slightly expanded posteriorly with small node. Interocular area rises gently from axial furrow to ocular lobe. | |
| J. S. Hollingsworth 2007 | A fallotaspidid trilobite with a wide cephalon and outwardly directed genal spines. Genal spine angle is advanced to align (tr.) with L1. Anterior border is wide, about 25% of cephalic length and strongly convex dorsally. Glabella narrow, parallel-sided to slightly tapered.
Ocular lobes connected to LA in posterior twothirds by eye ridges as wide as, but less elevated than the ocular lobes. At LA, eye ridges form an angle of 65º with the sagittal axis. Ocular lobes wide and elevated posteriorly. Interocular area about as wide (tr.) as extraocular area. | |
| D. Y. Huang and A. Nel 2007 | Legs: three (or four?) segmented-tarsi. Fore wing: a long slightly curved vein between Sc and C (unique apomorphy in Grylloblattodea); M divided into simple MA and simple MP at arculus level; RS simple; CuA2 simple; CuA1 pectinate; a long and broad cell in anal area; A1 simple, A2 pectinate with three posterior branches | |
| G. D. Edgecombe et al. 2007 | Eokosovopeltis with axial furrow
relatively strongly divergent anterior to S2. Glabellar furrows faintly impressed on external and internal surfaces of exoskeleton, none pit-like; S1 and S2 confluent with axial furrow; L2 gently swollen as round lateral lobe. Cranidium wider across β than γ , with relatively broad anterior margin and wide preocular fixigenal field. | |
| M. Webster 2007 | Biceratopsids with short, strongly divergent ocular lobes (posterior tip opposite or anterior to midlength of L1; line drawn from posterior tip of ocular lobe to contact between adaxial margin of ocular lobe and axial furrow forms angle of 25°-50° with sagittal axis). Glabella hourglass-shaped, constricted at S1 or L2; axial furrow shallow or absent at lateral margins of L1; S2 gently convex anteriorly, contacts axial furrow; S3 elongate furrow, oriented anterolaterally away from axis until contact with ocular lobes, contacts axial furrow; LA hemispherical to transversely oblate, summit slightly higher than more posterior glabellar lobes. Genal spine longer than length of LO (exsag.), base located at point of maximal cephalic width (tr., excluding genal spines). Fourteen prothoracic segments; pleural spine of T3 stout, macrospinous, blunt-tipped; first segment of opisthothorax with long axial spine. Summit of interocular area of higher dorsal elevation than summit of ocular lobes at small cephalic size (approximately 1.5 mm sagittal length). | |
| M. Webster 2007 | Species of Paranephrolenellus with very strong adgenal angle (deflecting posterior margin of cephalon anteriorly by 60°-80°), base of genal spine opposite L2, S2, or L3; S2 gently convex anteriorly either side of sagittal axis, abaxial and adaxial ends on roughly transverse line; L3 transverse to weakly ‘V’-shaped. | |
| M. Webster 2007 | Species of Paranephrolenellus with strong adgenal angle (deflecting posterior margin of cephalon anteriorly by 40°-70°), base of genal spine opposite LO, SO, or L1; S2 gently convex anteriorly either side of sagittal axis, abaxial and adaxial ends on roughly transverse line; L3 roughly transverse to weakly ‘M’-shaped. Fourteen prothoracic segments; T3 weakly hyperpleural, macrospinous, pleural spine stout, broad (tr.), blunt-tipped; long axial spine on T15. | |
| M. Ghobadi Pour et al. 2007 | raphiophorine with an almost circular exoskeleton; short, divergent genal spines; large cranidium ornamented with fine, concentric, raised ridges; ovoid, strongly convex glabella, not overhanging the cranidial margin with a median tubercle located near cranidial mid-length; wide posterior border furrow; fixigenae with a pair of deep, subtriangular pits on both sides of the glabella bordered anterolaterally by rudimentary persistent eye ridges in adults. Thorax with five thoracic segments, almost as wide as the cranidium. Pygidium strongly transverse, semioval to sub- triangular, with up to five axial rings and four well defined pleural ribs distally raised and undercut near the narrow, nearly vertical border; axis convex, reaching the posterior border, with deep axial furrows. | |
| A.P. Van Viersen 2007 | Pygidial median rib gradually narrowing towards anterior; proximally slightly widened and connected to axis. Pygidial pleurae distally slightly wider (tr.) than adjacent interpleural furrows. | |
| T.-Y. Park et al. 2008 | Cranidium wider than long; frontal area short, 20–25% of cranidial length, with a distinctive preglabellar boss outlined by parallel-sided furrows emerging from antero-lateral corners of glabella, not differentiated into preglabellar field and anterior border; glabella slender, 30–40% of palpebral cranidial width, with obsolete glabellar furrows; occipital ring with a long and stout median spine; palpebral lobes located at glabellar mid-point. | |
| M. Stein and J. S. Peel 2008 | One diagnostic pygidial character of Perissopyge
not mentioned in the original description (Blaker et al. 1997) is that the two anterior pleural bands of the pygidium are laterally drawn out into small, posteriorly directed spines, best seen in well-preserved material of the type species, Perissopyge phenax (Fig. 4C). It can be confirmed even for P. triangulata from Nevada (Fig. 2A). | |
| A. L. Stigall and J. H. Hartman 2008 | Carapace moderate in size, elliptical in outline, valves moderately convex, length approximately 1.5× greater than height; dorsal growth bands broad and flat transitioning to narrow bands between the tenth and fifteen bands, with large, sievelike, cavernous ornamentation formed by aggregation of numerous punctae; growth lines smooth; cavernous ornamentation throughout growth band except distal margin; distal margin of growth bands ornamented by linear bands of punctae. Soft part and egg morphology unknown. | |
| A. L. Stigall and J. H. Hartman 2008 | As for the genus. | |
| H.-L. Luo et al. 2008 | Body large, cranidium subtrapeziform in outline; glabella wide and cone-shaped, with 3 paries of shallow glabellar furrows; occipital furrows shallow and wide; neck ring shows uniform width. Dorsal furrows narrow and shallow. Eye ridges shorts and unclear; eye lobes relatively large and convex outward. Fixigena relatively narrow. Posterolateral limb wide and large; posterior border furrow shallow and wide. Anterior limb narrow; anterior border wide and convex slightly. Anterior branch of the facial suture extends outward; posterior branch of the facial suture relatively long, extending backward and outward. Thorax with at least 16 thoracicsegments; axial lobe wide, columnar in outline, with a node on each ring. Pleurae relatively narrow; pleural spines become longer backward. Pygidium probably small. | |
| H.-L. Luo et al. 2008 | Same as for the genus | |
| M. Stein and J. S. Peel 2008 | Examination of the type suite illustrated by Blaker et al. (1997, fig. 7) indicates shortening, stretching and oblique
distortion of individual specimens, presumably as a result of tectonic deformation. Thus, the holotype is antero-posteriorly shortened relative to the paratype illustrated here (Fig. 2A). The presence of pleural spines on the lateral border of the pygidium (Fig. 2A) sheds doubt on the reported presence of seven thoracic segments in the holotype (LACMIP 7716) of P. triangulata. The tagma boundary between thorax and pygidium is not readily identified in that specimen (Fig. 2B). At least one of the supposed thoracic segments may belong to the pygidium. For other specimens (e.g. LACMIP 7417, 7418), seven thoracic segments can be confirmed. As the holotype specimen is larger than the remainder, it seems unlikely that it represents a late meraspis. It should be noted that the cephalothoracic boundary is poorly preserved, the posterior cephalic border being somewhat deformed. It is possible that the anteriormost thoracic tergite partly underlies the posterior margin of the cephalon and hence is not recognized. | |
| R. G. Martins-Neto et al. 2008 | Forewing tegminous, costal margin thickened basally, with ScR and M origin close to the base; both veins having a short common stem, r-m close to the wing apex; RP origin located little before the wing mid length, CuA leaves basal cell separately, m-cu veinlet (arculus) strongly oblique next to the anal margin, narrow medial area. | |
| T.-Y. Park et al. 2008 | Cranidium as long as wide; glabella generally 45–50% of cranidial length; frontal area moderately long (30– 35% of cranidial length); occipital ring long (20–25% of cranidial length); palpebral lobes small (ca 20% of cranidial length); a pair of short furrows on frontal area strongly divergent; anterior branches of facial sutures convergent convex; posterior branches of facial sutures less divergent, straight. | |
| T.-Y. Park et al. 2008 | Cranidium with weakly convex preglabellar field defined by a pair of weakly divergent short furrows on frontal area; long glabella (greater than one-half of cranidial length); short occipital ring (15–20% of cranidial length); and large palpebral lobes. | |
| B. Wang et al. 2008 | Hindwing oval, anterior margin curved in distal part; costal area wide, maximal width in basal 1/3rd; vein R fused with vein Sc basally, and branching into veins Sc+RA and RP; vein Sc terminating in costal margin just basad of wing indentation; branch RP fused with vein M1 for a long distance basad of level of indentation; veins M1, M2 and M3+4 arising from stem M at wing base; vein M3+4 unbranched and almost straight; veins CuP and Pcu simple; vein A1 absent; membrane with geometric colour pattern. | |
| T.-Y. Park et al. 2008 | Cranidium ca 1.1–1.3 times wider than long; palpebral lobes located at level corresponding to anterior half of glabella; short occipital ring with a large median node; eye ridges nearly transverse; narrow librigenae without genal spines; and surface ornamented with more-or-less evenly spaced granules. | |
| T.-Y. Park et al. 2008 | Cranidium generally as long as wide; glabella truncato-conical with faintly impressed glabellar furrows; occipital ring semi-circular, with a small median node; palpebral lobes crescentic, small to intermedi- ate in size, located at a level corresponding to anterior half of glabella; eye-ridges weakly to clearly developed, transverse to directed ob- liquely rearwards; preglabellar area convex, ca one-third of cranidial length; a pair of incomplete furrows emerge from antero- lateral corners of glabella and diverge ob- liquely forwards on frontal area; anterior border absent or very narrow. Librigena long and narrow, with a postero-laterally directed long genal spine. Pygidium small, micropy- gous, transversely elliptical in outline; axis one-quarter to one-third of pygidial width, nearly reaching to posterior margin, with four to five axial rings and a terminal piece; border narrow or undeveloped. | |
| T.-Y. Park et al. 2008 | A species of Cyclolorenzella with pustulate prosopon. | |
| T.-Y. Park et al. 2008 | Cranidium with a convex pregla- bellar boss defined clearly by a pair of deep, broad and complete divergent furrows on frontal area. | |
| T.-Y. Park et al. 2008 | Cranidium longer than wide; frontal area long (35–40% of cranidial length); palpebral lobes small (ca 20% of cranidial length); a pair of diverging furrows on frontal area insignificantly impressed. | |
| E. Liñán et al. 2008 | Bigotinid trilobites with arched eyeridge and palpebral lobe; librigenae with obtuse genal angle and asmall sublateral spine. Thoracic axial ring (sag.) as wide as the pleural field, which is arched and presents an elliptical pleural furrow. Semielliptical micropygidium smaller than cranidium whit well developed axis and pleurae. | |
| K. J. McNamara et al. 2009 | Cephalon very vaulted, with strongly tapering prolongations; brim and genal roll both strongly convex and steeply inclined sagittally, less so laterally. Girder very distinct and smooth. Prominent large preglabellar boss, as wide as posterior of glabellar, on genal roll in some morphotypes. Genal area of low convexity. Eye lobes anteriorly positioned. Eye ridges broad, short and weak. Glabella small, strongly vaulted. Ala prominent; alar furrows and posterior alar depression deep. Posterior border furrow broad and shallow. Brim, genal roll and prolongations with irregular but prominent pitting, except on lateral genal roll, where more radially aligned. Genal area, occipital ring and glabella ornamented with fine granules. Hypostome, thorax and pygidium unknown. | |
| Żylińska and Szczepanik 2009 | Ornamentaspis with narrow glabella c. 82% of cranidial length and c. 50% of cranidial width across centre of palpebral lobes; glabella with three pairs of rather shallow lateral glabellar furrows; occipital ring narrow, without spine; fixigenae c. 70% of maximum glabellar width; palpebral lobes less than 30%of cranidial length; preglabellar field and anterior border poorly differentiated. | |
| B. Wang et al. 2010 | Body slender, length about 25 mm. Mesonotum almost as wide as abdomen. Forewing triangular, length about 33–36 mm, width about 16 mm; vein Sc unbranched; veins RA, RP and M separating at one point; branch CuA bifurcating basal of the level of nodal indentation. | |
| B. Wang et al. 2010 | Body robust. Mesonotum wider than abdomen. Forewing triangular, length 65–75 mm, maximum width (at the middle of wing) about 30 mm; vein Sc unbranched; veins RA, RP and M separating at one point; branch RP subparallel to branch RA; antenodal region trapezoid; postnodal region reduced; clavus narrow, less than one-third of wing length. Hindwing small, about half as long as forewing. | |
| B. Wang et al. 2010 | Body robust. Mesonotum width about 20 mm, clearly wider than abdomen. Forewing triangular, length 43–46 mm; vein Sc unbranched; veins RA, RP and M separating at one point. Hindwing small; vein M4-branched. | |
| R. M. Owens et al. 2010 | Cranidium with transverse anterior margin with border; frontal glabellar lobe short; glabellar furrows pit-like; L1 expanding laterally, merging adaxially with middle lobe; three low tubercular swellings on middle glabellar lobe and occipital ring; pygidium with 15 broad, slightly convex pleural spines of length equal to or less than width of pleural ield adaxial to spines; spines without median keels but with marginal grooves deined by very narrow, elevated border ridges. Dorsal surface smooth or with sculpture of inconspicuous granules. | |
| A. C. Daley and G. E. Budd 2010 | An anomalocaridid with a segmented frontal appendage that is elongated and tapering in outline, with each podomere bearing one thick spine on its inner margin and a single row of numerous small spines along its outer margin. Appendages have a convex, bell-shaped
proximal margin and a pointed distal end bearing a curved terminal spine. | |
| T. A. Hegna and D. Ren 2010 | Anteriorly-rounded dorsal carapace fused with the head; labrum fused ventrally to carapace; reduved antennae; paired sessile eyes; 11 pairs of ostia in the heart; female genital opening at the base of thoracic limb series; male genital opening a simple pore at the base of the 11th pair of thoracic limbs. (Partially modified from characters used by Richter et al. 2007) | |
| J. R. Paterson et al. 2010 | Nektaspida with tagmosis composed of cephalic
shield with posterior margin nearly straight and transverse, short thorax of three or four segments, and elongate pygidium, much longer than cephalic shield. First thoracic segment as wide as cephalic shield, without contraction in width behind cephalon. Hypostome elongate, with natant attachment, placed well behind anterior margin of cephalic shield, divided by a transverse suture into a subtrapezoidal anterior half with an anterior wing and shield-shaped posterior half with a rounded posterior margin. Antennae relatively short, laterally deflected from their attachment behind anterior wing of hypostome; three pairs of postantennal cephalic limbs. Pygidium with distinct border furrow. | |
| J. R. Paterson et al. 2010 | Emucaridid with four thoracic segments and
relatively narrow, elongate, subtriangular pygidium about twice the length of cephalic shield. Axis lacking significant relief or impression of axial furrows on cephalic shield and thorax but markedly raised on pygidium; postaxial ridge continued onto doublure. Pygidial axis with polygonal ornament on dorsal surface; c. 44 axial rings defined by closely spaced ring furrows on ventral surface; intersegmental furrows shallow or indistinct on pygidial pleurae; pygidial border relatively wide; doublure of identical dimensions to dorsal pygidial border | |
| J. R. Paterson et al. 2010 | . Emucaridid with very narrow, rim-like doublure
on cephalic shield and pygidium. Cephalic shield c. 66 per cent length (sag.) of pygidium. Three thoracic segments. Pygidial axis with strong ring furrows that define c. 12–13 segments; strongly impressed intersegmental furrows on pygidial pleurae; narrow, segmented pygidial border | |
| A. C. Daley and G. E. Budd 2010 | A radiodontan with 14 podomeres per frontal appendage, which are elongated and tapering in outline. A pair of ventral spines extends from the ventral surface of each podomere. Appendages have a convex bell-shaped proximal margin, a terminal spine protrudes from the distal end in some species (emended from Daley & Budd 2010, p. 727). | |
| S. R. Westrop et al. 2010 | Very short frontal area that occupies less than 20% of cranidial length. Librigena with long, stout genal spine directed backward steeply upward. Pygidium relatively narrow, with length equal to half of width; anterior margin curved sharply backward abaxially; articulating facet long. | |
| S. R. Westrop et al. 2010 | Long, wide, gently tapered, tongue-shaped anterior border that occupies 33% (31–34) of cranidial length in larger holaspids; border length of larger holaspids is 48% (46–51) of maximum border width. Preglabellar field long in larger holaspids (e.g. Fig. 6A–C, G–I), equal to 14% (13–16) of preoccipital glabellar length but somewhat shorter in smallest cranidia (e.g. Fig. 7H–J), equal to 12% of preoccipital glabellar length. Occipital ring with stout spine that curves upward and backward. | |
| A.P. Van Viersen and B. De Wilde 2010 | Lateral glabellar furrows S3 are firmly impressed. Anterolateral pygidial border is wholly smoothly rounded. Pygidial median rib is broad (tr.) distally, exceedingly narrow proximally, and rudimentary close to the axis. Pygidial pleural ribs are dorsally flattened. | |
| M. J. Hopkins 2011 | Amended from Palmer (1965b). Anterior border longer (tr.) abaxially, tapers strongly laterally. Glabella tapers strongly anteriorly; axial furrows gently concave between palpebral lobes in dorsal view. Node on occipital ring | |
| M. J. Hopkins 2011 | Amended from Palmer (1965b) and Stitt et al. (1994). Anterior border concave. Preglabellar field down- sloping to noticeably convex. Coarse pits or granules in row along anterior border furrow of internal mold (character 6). | |
| M. J. Hopkins 2011 | Amended from Palmer (1960). Pleural ridges extend postero-laterally, curve posteriorly at border furrow, extend to the posterior margin in some species. | |
| M. J. Hopkins 2011 | Palmer (1965b) is sufficient. | |
| G. O. Poinar and J. T. Huber 2011 | Fore wing narrow, venation extending about two-thirds wing
length, marginal vein probably longer than submarginal vein (wing base not clearly visible), and postmarginal vein apparently absent (Fig. 1); antenna with 8-segmented funicle and 2-segmented clava (Fig. 2); palpi (probably maxillary) distinctly 3-segmented (Fig. 2); tarsi 5-segmented and long; metanotum with anterior and posterior margins parallel; petiole ring-like, apparently shorter than wide; and ovipositor sheaths with several setae near apex. | |
| M. J. Hopkins 2011 | Palmer (1965b) is sufficient. | |
| M. J. Hopkins 2011 | Amended from Palmer (1965b). Anterior bor- der narrower (sag.) than preglabellar area, defined by narrow, deep furrow. Glabella tapering anteriorly, glabellar furrows poorly defined. Axial furrows strongly defined, straight. Anterior border furrow divergent. Interocular areas upslop- ing. Palpebral lobe one third length of glabella (sag.), prominent, well defined by palpebral furrow. Node on occipital ring. Transversely subovate pygidium, three or four axial furrows behind articulating furrow, no border furrow. Smooth to roughened exoskeletal surface. Faint caecal markings on frontal area | |
| M. J. Hopkins 2011 | Palmer (1965b) is sufficient. | |
| M. J. Hopkins 2011 | Palmer (1965b) is sufficient. | |
| M. J. Hopkins 2011 | Amended from Palmer (1965b). Posterior margin smoothly curved to gently inbent. | |
| N. C. Hughes et al. 2011 | Effaced cranidium with long (sag.), concave frontal area with anterior branch of facial suture running abaxially anterior of the palpebral lobe, then sweeping adaxially in open curve to intersect anterior margin in front of fixigena. Glabellar conical with weakly incised anterior margin. Posterior border furrow and SO firmly incised. Posterior lateral margin of LO pit-like. Isopygous with convex, semicircular pygidium with distinct pleural facet, axis tapering posteriorly, border broad, flattened with broad doublure. | |
| N. C. Hughes et al. 2011 | Effaced cranidium with long (sag.), concave frontal area with anterior branch of facial suture running abaxially anterior of the palpebral lobe, then sweeping adaxially in open curve to intersect anterior margin in front of fixigena. Glabellar conical with weakly incised anterior margin. Posterior border furrow and SO firmly incised. Posterior lateral margin of LO pit-like. Isopygous with convex, semicircular pygidium with distinct pleural facet, axis tapering posteriorly, border broad, flattened with broad doublure. | |
| A. P. Van Viersen and H. Prescher 2011 | A species of Scutellum with a combination of the following features: anterior border of cranidium protruding slightly (in smaller specimens) to distinctly (in large specimens) from cephalic outline along with anterior margin of glabella. Frontal glabellar lobe laterally expanded; having indistinct, rounded anterolateral corners. S3 weakly impressed, especially abaxially. Small eyes. Abaxial half of lateral portion of pygidium distinctly protrudes laterally from parabolic outline of axis. Of all ribs, pair adjacent to median rib has narrowest proximal half. Interpleural furrows widening quickly between anterior pleural ribs but widening from nearby margin of elevated central area of pygidium between remaining ribs. | |
| A. P. Van Viersen and H. Prescher 2011 | Large Scutellum species with the following combination of features: pygidia easily attaining 30 mm in sagittal length. Cephalon wholly moderately weakly vaultred (tr.) and of widely and smoothly rounded outline. Axial furrows making an angle of 60° from L1. Border in front of glabella. Librigenal field slopes smoothly abaxially. Sharp, posteriorly directed genal angle. Median rib proximally as wide as, and less vaulted (tr.) than, adjacent pleural rib pair. | |
| M. Ghobadi Pour et al. 2011 | Fringe with pits on upper lamella dominantly radially arranged in relatively narrow, deep sulci, containing pits E1–2 and I1–2. I pits interior of I2 in radial alignment with sulci frontally, randomly distributed and out of radial alignment laterally and posterolaterally. Fringe with E1 complete; E2 developed abaxially very close to E1, diverging from it posterolaterally until maximum curve of fringe, then converging to it posteriorly. I1 and I3 complete; I2 very close to I3 frontally, with median two I2/I3 pit pairs merging; I2 increasing in distance from I3 laterally, but cannot be traced anterolaterally due to irregular arrangement of I pits between I1 and I3. Apart from I1 and I3, I pits irregularly arranged. Around four irregular arcs between I1 and I3 at anterolateral corners, up to eight posterolaterally. Lateral eye ridges and tubercles absent. Pygidium with five axial rings plus terminal piece. | |
| M. Ghobadi Pour et al. 2011 | Cranidium of about equal length and width, with broadly and evenly rounded anterior margin; preglabellar field smooth, with very weakly expressed longitudinal median ridge; pygidium about 70% as long as wide, with axis occupying 20% of pygidial width and 75% of maximum pygidial length, and terminating with a short postaxial ridge; axial furrows are shallow and wide, almost straight; 12 well-defined axial rings; pleural field flattened, with six pairs of pleural ribs which do not reach border. | |
| M. Ghobadi Pour et al. 2011 | Glabella slightly elongate, with maximum width at anterior margin; cranidial axial furrows strongly divergent anteriorly; very narrow cephalic border; large semicircular eye; very weakly defined posterior cephalic border; pygidium transverse, semioval, with axis completely lacking segmentation; long, well-defined postaxial ridge. | |
| M. Ghobadi Pour et al. 2011 | Cephalon semioval, about 80% as long as wide; elongate, parallel axial furrows with evenly rounded anterior margin confined to the anterior border furrow; three pairs of very weakly defined glabellar furrows; occipital ring transverse with a small median tubercle; convex cephalic border of equal width with short median spine; medially (exsag.) situated palpebral lobes about 25% of cephalic length; eight thoracic segments, and pygidium with conical axis about 35% of maximum pygidial width, four axial rings and very short terminal piece; pleural fields with three to four pairs of ribs; border narrow, fading posteriorly abaxially. | |
| M. Høyberget and D. L. Bruton 2012 | Glabella convex, slightly tapering, longer than wide. Distinct occipital furrow, S1 pronounced abaxially. S2 and S3 short, shallow to obsolete. Anterior border upturned, moderately to strongly convex in frontal view. Centres of palpebral lobes opposite S1 to L2. Eye ridges oblique. Palpebral area markedly narrower than glabella at eye line. Posterior area as wide as to slightly wider than occipital ring (tr.). Posterolateral projection ventrally downflexed. Librigena with long and curved spine rounded in cross- section. Thorax consists of ten segments, pleural fields about as wide (tr.) as axis. Tenth segment with long axial spine. Pleural spines short, seventh pair elongated. Pygidium minute and triangular, one to two axial rings. Very long, backwardly curved caudal spine. One pair of short marginal spines. Pleural field less than half the width of axis. | |
| G. Yang et al. 2012 | Head width longer than length; eyes large and ovoid; antennae filiform, ten-segmented, scapus shortest and thickest, pedicelli enlarged, slightly longer than scapus, all flagellomeres slender except for apical segment swollen; rostrum straight, long, three-segmented. Femora stout, distinctly shorter than tibiae; tibiae slender, and hind tibiae longer than fore and mid tibiae; tarsi two-segmented, first segment longer than second. Forewings broadened in subapical part; vein R2 long, R1 reduced; Rs reaching to costal margin; stem of R slightly shorter than vein M+CuA; M with two long branches; cell m1+2 variously elongated, always longer than cell cua1; CuA forked into long CuA1 and short CuA2; anal field long, almost half length of forewing. | |
| J. R. Carlucci et al. 2012 | Predominant shape of lunette is elliptical (character 4, state 1), path of the cranidial axial furrow is straight (character 5, state 0), 10 thoracic segments (character 8, state 2), large paired muscle scars present on pygidium (character 11, state 1), ratio of pygidial length to width is high (76–83%) (character 16, state 3), axial furrow expressed as notches flanking centre of the pygidial anterior margin (character 17, state 0). | |
| J. Peng et al. 2012 | A dorsal exoskeleton divided by a single transverse articulation into two shields, a cephalic shield and a posterior shield. The cephalic shield is round to half-round in dorsal view, devolving spinose or nonspinose in genal angles, both lateral margins and rear margin of po- sterior shield; one pair of lateral project antennas in ante- rior cephalic area. Axial lobe weak, both lateral axial lobe with alimentary system, including alimentary diverticulae and alimentary canal, with appendages in abdominal re- gion. | |
| J. R. Paterson et al. 2012 | As for type species by monotypy | |
| J. R. Paterson et al. 2012 | —Exoskeleton an elongate oval in outline; maximum
exoskeletal width (tr.) 40–50 percent sagittal length (excluding pygidial spines). Cephalic shield approximately 10 percent length of complete exoskeleton (sag.), its anterior margin rounded and lacking a median notch; eyes lacking dorsal expression in cephalic shield. Trunk consists of 23 thoracic tergites with blunt, spatulate tips and a small pygidium with inflated axis and pair of long, flat, divergent pleural spines. Endopods short, terminating around mid-width of pleurae. | |
| X.-L. Zhang et al. 2012 | Arthropod with well-developed pleural fold, subelliptical
shape and weakly trilobite. Head shield crescentic in outline, one-third of the total length; anterolateral margin notched. Posterior section composed of 27 distinct tergites withwell-developed pleural spines and a terminal piece; anteriormost tergite reduced, tergites 2–7 almost straight, and tergites 8–27 curved in dorsal aspect compared to anterior ones, the curvature of these tergites increasing posteriorly so that the posterior 16 tergites are highly curved to semicircular in dorsal aspect | |
| X.-L. Zhang et al. 2012 | Arthropod with well-developed pleural fold, subelliptical
shape and weakly trilobite. Head shield crescentic in outline, one-third of the total length; anterolateral margin notched. Posterior section composed of 27 distinct tergites with well-developed pleural spines and a terminal piece; anteriormost tergite reduced, tergites 2–7 almost straight, and tergites 8–27 curved in dorsal aspect compared to anterior ones, the curvature of these tergites increasing posteriorly so that the posterior 16 tergites are highly curved to semicircular in dorsal aspect. | |
| M. Høyberget and D. L. Bruton 2012 | A Sphaerophthalmus species with centres of palpebral lobes opposite S1. Glabellar furrows shallow. Palpebral area convex, elevated above palpebral lobes. Librigenal posterior lateral margin nearly straight and shorter than convex anterior lateral margin. | |
| W. Zhang et al. 2013 | Body length medium (c. 5–6 mm), broad. Compound eyes large, reniform, protrusive. Antennae relatively long, first antennomere stout, nearly twice diameter of remaining antennomeres (Fig. 7C). Pronotum divided into anterior and posterior lobes, with explanate
and semitranslucent margins present; lateral margins of anterior lobe nearly straight, anterior angles rounded, posterior margin convex. Scutellum large, triangular. Forewing costal fracture very long; membrane with cells. Tarsal formula 2-2-3; metatarsi relatively long, about one-half length of corresponding tibiae. Connexiva absent. The synapomorphy for this group is the pronotum with explanate and semitranslucent margins. | |
| W. Zhang et al. 2013 | Body length medium (c. 6–7 mm). Compound eyes small, reniform, protrusive; ocelli present, close to each other.Anteclypeus and paraclypeus elongate (Fig. 7D). Posterior lobe of pronotum much broader than anterior lobe, lateral margins of anterior lobe slightly converging posteriorly; anterior angles acute; posterior margin of pronotum distinctly concave. Scutellum large, triangular. Forewing separated into corium and membrane although boundary between corium and membrane not distinct;membrane with
four cells. Connexiva present. The synapomorphies for this group are the elongate anteclypeus and paraclypeus, and the characteristic pronotum. | |
| L. Xing et al. 2013 | Epichnia, convex burrows (Fig. 7A) with dilated parts considered to be chambers (Fig. 7B). Traces columnar, branching, convex, with Y-shaped branches (Fig. 7C). Traces wind around the surfaces of the bones. At junctions other than Y-shaped branches, traces overlap rather than intersect directly (Fig. 7D). Surface texture rough. Traces hollow with thin mud constructed walls and cement infilled. | |
| D.-C. Lee 2013 | Species of Remopleurides with non−spinose, entire pygidial margin and longitudinal ridges on occipital ring and thoracic axial rings | |
| A. C. Daley et al. 2013 | Hurdia with an elongated H-element that has a maximum length twice as long as the width. | |
| A. C. Daley et al. 2013 | Hurdia with a short, wide H-element that has a maximum length 1.5 times longer than the width. | |
| M. F. Tortello and S. B. Esteban 2014 | A species of Asaphellus with exoskeleton of large size, flat or very slightly convex; posterior facial suture transverse, strongly directed outward; genal spines small; pygidial axis long and narrow, with traces of five rings on its anterior part; pleural fields smooth or with very faint indications of five or six pleural furrows; pygidial border well-developed, depressed (emended from Harrington and Leanza, 1957). | |
| J. Ortega-Hernandez et al. 2014 | Large and weakly biomineralized arthropod exhibiting the following unique combination of charac- ters: cephalon wide (tr.), with weakly defined glabellar region, genal spines inserting laterally, abaxially procurved posterior margin, and apparently no dorsal eyes or ecdy- sial sutures; 11 trunk tergites, T1–8 with edge-to-edge articulations and sickle-shaped pleural spines inserting an- terolaterally, T9–11 characterized by an abrupt change in tapering and reduction of the pleural regions into long, backward-facing pleural spines. | |
| J. Ortega-Hernandez et al. 2014 | arge and weakly biomineralized arthropod exhibiting the following unique combination of charac- ters: cephalon wide (tr.), with weakly defined glabellar region, genal spines inserting laterally, abaxially procurved posterior margin, and apparently no dorsal eyes or ecdy- sial sutures; 11 trunk tergites, T1–8 with edge-to-edge articulations and sickle-shaped pleural spines inserting an- terolaterally, T9–11 characterized by an abrupt change in tapering and reduction of the pleural regions into long, backward-facing pleural spines. | |
| L. Amati 2014 | A species of Isotelus with very short (sag.) cranidium, only slightly longer than maximum width (tr.) in front of palpebral lobes. Facial sutures with only slight inflection in front of palpebral lobes then nearly parallel. Pygidium with long border, longest directly behind axis. | |
| L. Amati 2014 | A species of Isotelus with facial sutures that diverge only slightly and for short distance in front of palpebral lobes then run nearly parallel until converge to mid-line. Frontal area very long (sag.), length about 20% total length of cranidium. Eye-stalks tall, extend well above maximum height of glabella. Lateral border of pygidium strongest over posterior 1/3 of pygidium, becomes much wider directly behind axis. | |
| L. Amati 2014 | A species of Isotelus with anterior branches of facial sutures directed inward in front of palpebral lobes then curving outward so that lateral margins of cranidium in front of palpebral lobes strongly sinuous. Anterior branches of facial sutures curve abruptly forward just before intersecting making anterior margin of cranidium sharply pointed. Ornament of coarse pits. Pygidium subtriangular but short (sag.) for genus and without border furrow. | |
| L. Amati 2014 | A species of Ectenaspis with border furrow of pygidium becoming obsolete through holaspid ontogeny. Weak axial rings and pleural furrows present. Cranidium wide anterior to palpebral lobes; facial sutures converge at base of proboscis. | |
| L. Amati 2014 | A species of Stegnopsis with facial sutures converging at front of cranidium at low angle; maximum width in front of palpebral lobes close to anterior margin. Convexity of cranidium very high. | |
| L. Amati 2014 | A species of Stegnopsis with very wide, evenly rounded cranidial border; anterior facial sutures directed abruptly forward just before intersecting to form sharp point. Palpebral lobes large. Librigenae without genal spines in large holaspids. Pygidium with relatively well-defined axial furrows. | |
| L. Amati 2014 | A species of Anataphrus with long (exsag.), wide (tr.) palpebral lobes. Pygidium long (sag.) relative to width (tr.); axial lobe weakly defined on internal molds. | |
| E. Bushuev et al. 2014 | Cephalon moderately extended in width by 1.6 times its length, with gently curved anterior margin. Gla- bella moderately tapered forward, convex, with three pairs of glabellar furrows oriented obliquely backwards when traced adaxially (S1, S2, S3). Anterior border narrow (less than 0.1 of glabella length), flattened and tilted back. Occipi- tal ring of rounded triangle outline, with moderately long, robust axial spine, rising upwards and backwards. Occipital furrow strongly curved back, deep, reaches axis. Distinct thin, paired, well-impressed “facial lines”, which onset at the merging of the ocular lobe with the eye ridge and run ob- liquely forward on the extraocular area. Small intergenal spines located along the posterior margin at distinct interge- nal angle. Medium length genal spines. Cephalic surface co- vered a system of raised ridges, intersecting to form a net- work of fine polygons with diameters up to 0.05 mm. | |
| E. Bushuev et al. 2014 | Cephalon semicircular in outline, relatively wide. Glabella moderately tapering forward, conical, with slightly acuminate LA. Poorly expressed, slightly convex parafrontal band in front of glabella; plectrum sometimes present on preglabellar field. Ocular lobes long, curved; posterior tips opposite or slightly behind the occipital fur- row. Interocular area narrow, less than half the width of glabella, opposite S1. Extraocular area moderately wide, slightly wider than L2, slightly convex, strongly tilted to- wards outer cephalic margins.
| |
| I. Rabano et al. 2014 | Cephalon parabolic in outline, with modera tely large eyes situated about 1.5 times their own length from posterior margin; glabella wide, about half the width of cephalon posteriorly, axial furrows subparallel to slightly curved inwards in vicinity of the lunette (ca opposite posterior end of the palpebral lobes), genal angles rounded. Librigenae with weak subocular furrow. Rostral plate broad (tr.), weakly convex (sag.), posterolaterally defined by almost transverse connective sutures and with a short upwardly and forwardly turned flange on the posterior edge. Hypostome with a narrow and globose middle body. Thorax of ten segments with well-defined axis that is broader than pleurae in dorsal view; axial furrows subparallel; pleurae adaxial to fulcrum successively increasing in width (tr.) posteriorly. Pygidium subtriangular in outline, larger than cephalon; axis weakly defined, subconical; pleural region concave posteromedially. Pygidial doublure broad, divided into convex outer half and flattened or weakly concave inner half by a shallow vincular furrow running approximately parallel to the inner margin. Dorsal surface of carapace almost smooth, with sculpture of fine terrace lines and small pits only developed on the anteriormost part of cephalon and the lateral borders of pygidium. | |
| M. F. Tortello and S. B. Esteban 2014 | An Asaphellus species with perceptible axial furrows, a moderately wide (tr.) palpebral area of the fixigena, and a narrow (sag.) pygidial border. | |
| M. F. Tortello and S. B. Esteban 2014 | A Nileus species with a subrectangular, anteriorly truncate glabella; palpebral lobes small for the genus; test surface of pygidium with weak indications of a few terrace lines on anterolateral part of pleural fields; pygidial axis vaguely perceptible on internal molds, with undifferentiated axial rings; pygidial border very narrow; doublure covered with 14 terrace lines. | |
| P. Cong et al. 2014 | Radiodontan with a pronounced, four-segmented neck; the head bears an oval to subcircular dorsal plate; mouth sub-circular to quadratic in outline, characterized by concentric wrinkled furrows; frontal appendage short, with a proximal podomere bearing a stout, blade-shaped endite with pectinate, sclerotized spines that gradually become larger distally; body flap of first trunk segment larger than succeeding body flaps. | |
| G. Geyer and J. Corbacho 2015 | Genus of the Burlingiidae with glabella slender, tapering forward; eye lobes short (exsag.), its posterior tips opposite S2, anterior tips opposite frontal lobe; with low tubercles on genal field adjacent to L3; preocular area with slightly raised portion; thorax composed of 11 or 12 segments; pygidium fan-shaped, posterior margin forming the rear end of the shield | |
| G. Geyer and J. Corbacho 2015 | Genus of the Burlingiidae with a glabella with subparallel sides or slightly tapering forward, with well-defined margins; frontal lobe subacute, with faint anterolateral corners; eye lobes long (exsag.), located close to the glabella, its posterior tips opposite S1, anterior tips adjacent to slight anterolateral corners of the glabella; preocular area a flat platform slightly sloping towards anterior margin; thorax composed of 11, 12 or 13 segments; pygidium slender, minute. | |
| D. J. Holloway and P. D. Lane 2016 | Cephalon and pygidium of low to moderate convexity. Omphalus and/or anterolateral internal pit present; axial furrow commonly extending forward to omphalus on internal molds. Shallow border furrow may be present on librigena; eye socle absent; genal angle broadly rounded to acute. Rostral plate lenticular to rhombic in outline; rostral portion of hypostomal suture transverse or gently convex forward. Thorax with very broad axis comprising 65%–75% segmental width and with ‘nonfulcrate’ pleurae. Pygidium with first pleural furrow impressed, meeting distinct holcos distally. | |
| M. F. Tortello and S. B. Esteban 2016 | Exoskeleton elongated and elliptical, about 70% longer than wide. Cranidium smooth, glabella almost indistinct laterally, preglabellar furrow ill-defined, convex forward. Genal spines widely based, and short (about 20% of free cheek maximum length). Pygidial border flat to slightly concave, wider anteriorly than posteromedially, barely defined axis in the anterior third of the pygidium. | |
| G.-Y. Ren et al. 2017 | Axial furrow wide and distinct. Cranid-ium subrhomboidal in outline exclusive of librigenae, and withfrontal projection as a long spine, a short club. Glabella trun-cated conical with obscure lateral furrows. Fixigenae moderatelywide, strongly convex. Palpebral lobe from moderately longto long, located posteriorly; anterior border furrow narrow andshallow, straight or slightly bending forward. Preglabellar fieldnarrow to absent. Librigenae narrow, with a very long genal spine. Thorax of 12 segments. Pygidium transversal semicircu-lar in outline, with 6 pairs of marginal spines, of which the lastpair is broader and triangular. Axis narrow and short, with 4–5axial rings and a short postaxial ridge. Pygidial border wider,without distinct border furrow. | |
| S. Pates and A. C. Daley 2017 | aryosyntrips with one pair of ventral spines on each podomere, and no dorsal spines. Distal most three podomeres are less tall than the others. Distal most podomere ends in a recurved terminal spine. | |
| S. Pates and A. C. Daley 2017 | Caryosyntrips with dorsal and ventral surfaces straight, a rounded distal end and no terminal spine. Paired ventral spines on each podomere are simple triangles that end in a sharp point. A single row of small (1 mm tall), curved, distally pointing, dorsal spines spaced 1 mm apart runs along the length of the appendage. A single large dorsal spine, the same size and shape as the ventral spines, is present on each of the distal most seven podomeres. | |
| L. M. E. McCobb and L. E. Popov 2017 | Fringe with E1-2, I n-1 complete; no F pits present.
E1 and E2 pits generally radially aligned with pit pairs set in sulci, but with occasional aberrant pits unpaired and out of alignment. In-1 pits radially aligned with those of In anteriorly, but posteriorly In-1 irregularly distributed and sometimes very close to In ; In pits not cut off by In-1 pits; I1 apparently complete; 2 to 3 I arcs between I1 and In-1 frontally, up to approximately 10 I arcs postero-laterally, randomly distributed. Glabella anterior greatly expanded and gently convex. Occipital ring lacks occipital spine. No lateral eye tubercles or eye ridges present. Pygidium short and wide, with width approximately three-anda-third times length; axis with five axial rings. | |
| R. Lerosey-Aubril et al. 2017 | Species of Glypharthrus characterized by the following unique combination of characters: body (telson excluded) as wide (tr.) as long (sag.); cephalon broadly semielliptical in outline, with particularly narrow (exs.) preocular field and exceptionally large eyes; trunk anterior to telson subtriangular in outline, bearing deep pleural furrows. | |
| A. L. Stigall et al. 2017 | Thin carapace valves of medium size, elliptical to ovate in outline, umbo centrally located, extending slightly above hinge line; height to length ratio from 0.7 to 0.9, valve convexity moderate; growth bands of two orders: flat growth bands ~0.4mm wide in umbonal and central carapace, and rounded growth bands ~0.04 mm (40 μm) wide at distal margin. Weak radial lirae and wrinkles present on dorsal growth bands. | |
| G. Geyer and J. S. Peel 2017 | Species of Elrathina with subparallel to slightly tapering glabella with a shallow rounded front; palpebral lobes small, but not minute, distinctly anterior to the cephalon’s half; eye ridges with only gentle curvature; anterior border narrow to moderately broad (sag.), occasionally with a minute to considerable medial swelling at its rear side; preglabellar field of variable sagittal width; librigenae with rounded posterolateral corner; geniculation of thoracic pleurae slightly abaxially to pleural mid-length; pygidium sublenticular to subelliptical outline; pygidial rhachis broad, with one to three rings, its posterior end with pair of slightly inflated bulbs separated by a longitudinal furrow, reaching to or almost to the posterior margin; pygidial pleurae with two or three furrows. | |
| T. P. Fletcher 2017 | A species of Agraulos with a pronounced truncate front to the isosceles-trapezoidal glabella (on internal moulds, the front is slightly kinked backwards over the axis); an ocular ridge commonly stemming from the middle inside part of the palpebral lobe (only well defined on the best preserved specimens); an occipital ring medially swollen backwards to a rounded/blunt point; a very short, outwardly directed, genal spine; a thorax with the first three axial rings with terrace lines and incipient median nodes on the succeeding three rings. | |
| S. Pereira et al. 2017 | Cephalic axial furrows gently curved; palpebral lobes corresponding to 18–19% of cranidial length; librigena bearing a short genal spine; pygidial length/width ratio from 58% to 65%; maximum pygidial width across posterior limit of axis; axial furrows indistinct; axis about 40% of pygidial length. | |
| T. P. Fletcher 2017 | Agraulinae with a relatively large subquadrate glabella, narrow (sag.), slightly convex, preglabellar field, conspicuously narrow (sag.) lunate cranidial anterior | |
| B. W. Rasmussen et al. 2017 | Parallel-sided glabella, S1 united across glabella and connected with axial furrow. Centre of palpebral lobes situated opposite L2. Interocular cheeks ¾ as wide as glabella at eye line. Postocular cheeks about as wide as occipital ring. Eye ridges oblique. Free cheeks with very wide flattened spine with centred rib or pair of ribs (on both sides). Genal angle acute. | |
| S. R. Westrop and A. A. Dengler 2017 | Palpebral lobe elongate, oblique but nearly straight, length equal to half preoccipital glabellar length on largest specimens (Figs. 2c, 2n), but shorter (equal to about 40% of preoccipital glabellar length) on smaller specimens (Figs. 2d, 2j); finely etched palpebral furrow; long, convex, arcuate, anterior border occupies about half of frontal area length. Punctate sculpture well defined over entire cranidial surface aside from anterior and posterior borders, palpebral lobes, and various furrows; smallest specimen (Fig. 2j) with weak scattered tubercles and faint punctae. | |
| B. W. Rasmussen et al. 2017 | Cranidium with slightly tapering glabella rounded in front, 3 pairs of glabellar furrows. Medium-sized palpebral lobes situated opposite of S2–L3. Free cheek with spine that follows the course of lateral margin. Subtriangular pygidium without marginal spines and 5–6 axial rings. | |
| R. Lerosey-Aubril and S. Pates 2018 | Hurdiid radiodont exhibiting the following unique combination of characters: central element lanceolate (length/width ratio > 2), with a tiny antero-sagittal spine, well-developed lateral lips, a strong postero-lateral constriction, and an indented posterior margin; paired lateral elements tall, with a straight to slightly concave postero- dorsal margin; frontal appendage equipped with at least seven plate-like endites, including two proximal ones bearing at least seven anterior auxiliary spines organised in two rows, and five longer distal ones fringed anteriorly by two rows of 50–60 narrowly-spaced setae. | |
| S. Pates et al. 2018 | Frontal appendage with 11 distally tapering podomeres; one robust double-pointed or single-pointed spine projects from the dorsal surface of P2–11 either vertically or at an oblique angle; P2–6 bear a spinous ventral blade slightly curved distally 2–3 times the length (sag.) of the height (sag.) of the dorsal spines (adapted from Caron et al. 2010). | |
| F. A. Sundberg 2018 | Cranidium with glabella slightly rounded in outline, wider (36 ± 2% glabellar width vs. cranidial width), slightly tapered (84 ± 5%); shallow glabellar furrows. Fixigena narrower (49 ± 6% anterior vs. posterior cranidial width), posterior portion moderately narrow (63 ± 3% posterior limb length to glabellar length), eye ridge nearly straight. Anterior border moderately tapered, curved and wide (56 ± 4% anterior vs. posterior cranidial width). Librigenal with rounded genal corner. Exoskeleton with fine granular surface. | |
| I. H. Escapa et al. 2018 | Leaves subopposite, narrow-elliptical to narrow-lanceolate, up to 12 cm long, length to width ratio 7–9:1. Leaves with margin entire, apex obtuse to acute, and basally reduced in width. Leaves amphistomatic, although scarce and isolated stomata are seen on the adaxial surface, with majority of stomata oriented parallel to venation. Pollen cones axillary, cylindrical at maturity, ca. 40 × 5 mm, subtended by adpressed basal bracts (at least two pairs); micro-sporophylls peltate, imbricate. Pollen monad, apolar; inaperturate, densely ornamented with irregularly spaced gemmae. Ovuliferous complexes broadly triangular, broader than long, apical margin thickened and upturned or recurved, a single seed scar centrally disposed, lacking basal embayments. Seeds composed by a central body and two lateral, slightly asymmetrical wings. | |
| C. Cronier et al. 2018 | Cephalon roughly semicircular. Brim gently inclined anteriorly and laterally, about one-third of cephalic length. External rim rather wide. Subrectangular elongated glabella with no tuberculation. Genal roll gently convex, sloping steeply, and short. Alae as wide as L1. No distinct eye ridges. No granulation or pits on genal area. | |
| B. Mayers et al. 2019 | Naraoiid nektaspids with the following characters: trunk length/total length ratio: <0.65; cephalic and trunk margins smooth or spinose. (Emended from Zhang et al. 2007.) | |
| B. Mayers et al. 2019 | Trilobitomorph artiopodans with the following characters as their ground pattern: lateral eyes reduced or absent; hypostome with natant attachement; extensive articulation overlap between unfused trunk tergites; fused pleurae not forming lateral spines. (Emended from Raymond 1920.) | |
| M. J. Hopkins 2019 | Cloacaspis ceryx species with relatively narrow preglabellar field (0.06–0.10 length of glabella) and smooth exoskeletal surface. | |
| B. Mayers et al. 2019 | Naraoiid nektaspids with the following characters: trunk length/total length ratio: ≥ 0.65; cephalic and trunk margins smooth. (Emended from Zhang et al. 2007.) | |
| B. Mayers et al. 2019 | Nektaspid trilobitomorphs with the following characters: cephalon with ovoid truncated shape; convex posterior cephalic margin; presence of ramifications on secondary digestive structures; anterior tapering of trunk relative to cephalic shield; complete effacement of tergite boundaries within fused section of the trunk. (Emended from Zhang et al. 2007.) | |
| G. Geyer 2020 | Cranidium relatively narrow; glabella constricted at level of L2, with three pairs of short lateral glabellar furrows separated from axial furrow; occipital ring wide (sag.); preglabellar field as long (sag.) as anterior border; palpebro-ocular ridge arcuate, slightly expanded in front; anterior sections of facial suture moderately divergent; preglabellar field (sag. and exsag.) absent. Thorax narrow, composed of 14 segments; pleurae narrower (tr.) than axis, spinose; first six thoracic pleurae ending in short, subequal spines; spines increasing rapidly in size between 7th and 11th pleurae with segment 11 macrospinose and fulcrum distal; pleurae of posterior segments (= opisthothorax) with short spines. Pygidium small, with ca. four pairs of tiny marginal spines; axis narrow, moderately well segmented; pleural regions with indistinct furrows only. | |
| G. Geyer 2020 | Lateral glabellar furrows slightly obliquely backward directed from axial furrow; posterior tips of palpebral lobes reaching to level of midlength of occipital ring; preglabellar field (sag. and exsag.) very short or absent; anterior border narrow to moderately wide; anterior branches of facial suture in a comparatively adaxial position. Thorax with 12–14 segments, with segments 9, 10 or 11 being macrospinose; tr. length of normalsized pleurae smaller than width axis in the anterior and middle part of the thorax, slightly growing in length. Pygidium subparabolic in outline; axis with up to 12 axial rings and with moderately large to large terminal axial piece; up to 10 pleural ribs developed; lateral border moderately wide. | |
| G. Geyer 2020 | Genus of the Despujolsiidae with glabella tapering forward, anterior margin of frontal lobe curved; lateral glabellar furrows slightly obliquely backward directed from axial furrow, less well developed than occipital furrow; palpebral lobes slightly oblique to axis, its posterior tips reaching to mid-level of L1; eye ridges, moderately long, weakly curved, distinctly oblique to axis; fixigenae moderately wide (tr.), subtriangular in outline; preglabellar field (sag. and exsag.) very short, without plectrum or swelling; anterior border moderately broad (sag., exsag.); anterior branches of facial suture in a comparatively abaxial position. Pygidium longitudinally subelliptical in outline; axis with well defined rings developed only in the anterior half; pleural fields narrow, with furrows well developed only in the anterior part; lateral border moderately wide, low. | |
| G. Geyer 2020 | Lateral glabellar simple, slightly obliquely rearward directed from axial furrow; posterior tips of palpebral lobes moderately long; eye ridges tapering forward, curved; preglabellar field (sag. and exsag.) absent; anterior border narrow to moderately wide. Thorax composed of 12, 13 or predominantly 14 segments; tr. length of pleurae smaller than width axis in the anterior and middle part of the thorax, slightly growing in length rearward but without distinctly extended pleura or macrospinose segment. Pygidium subsemicircular to spathulate in outline, posterior margin regularly curved or with slight indentation on sagittal line; with three or four axial rings and equal number of pleural rings; axis with small terminal axial piece; lateral border relatively narrow, tapering rearward. | |
| A. Bignon et al. 2020 | Usually small ovoid-globular (commutavi) protaspides. Short eyes, when present, con- nected to glabella by eyeridge without palpebral lobe. A long (sag.) glabella compared to its maximal width (tr.). Three or fewer short glabellar furrows. Tubercle on gla- bella or occipital ring. Free cheeks anteriorly fused. Facial sutures are marginal or dorsal. Perforated fringe present in more derived forms. Genal spines long and narrow at base. Distal extremities of thoracic segments pointed. Pygidium triangular or semi-circular, smaller than cephalon, wider (tr.) than long (sag.), with a curved anterior border. Pygidial axis narrow, may extend onto posterior border. | |
| F. A. Sundberg 2020 | Cranidium subtrapezoidal, moderate length frontal area (36 ± 3% cranidial length), nearly parallel anterior facial sutures, anterior border moderately wide (width 61 ± 7% cranidial width) and narrow (length 16 ± 2% cranidial length), false anterior furrow absent; genal caeca well developed on preglabellar field; palpebral lobes dorsally arched; fixigena slightly convex, gently sloping upwards; bacculae absent. Librigenae with genal caeca; genal spine relatively short (length ∼60% librigenal length), broad based with shallow lateral border furrow extending into spine. Pygidia nearly effaced, faint border furrow and anterior pleural furrow, anterior lateral corners located at the level of the posterior axial lobe, anterior margin strongly curved posterolaterally approximately 1/3 distance from axial lobe. Cranidium and librigena covered in fine granular ornamentation. | |
| M.B. Lara and A. Bashkuev 2020 | RA2 simple. Medial cell trapezoid. M4, CuA1 and CuA2 much closer to each other than in T. rhaetica. | |
| M.B. Lara and A. Bashkuev 2020 | Cuyanoscytinum gen nov. is referred to Scytinopteridae Handlirsch, 1906 based on the following characters: forewing tegminous, costal margin thickened basally and costal area wide, Scb reduced,
stem of R, M and Cu come off separately from a point close to the base, branches of MP and CuA short, and crossveins reduced (Tillyard 1919). Cuyanoscytinum gen nov. compared to Tipuloidea Wieland, 1925 differs in position of medial cell and crossvein m-cua. | |
| M.B. Lara and A. Bashkuev 2020 | Only the forewing characters are known. RA2 simple. Medial cell trapezoid, between stem M1 and M2. M3 + 4. Crossvein m-cua connects stem of Mand CuA1. | |
| G. Geyer 2020 | Glabella tapering slightly forward; preglabellar field (sag. and exsag.) short or absent, with three simple lateral glabellar furrows; eye ridge and palpebral lobe united with a palpebro-ocular ridge; palpebral lobe long (exsag.), its posterior extremity distant from axial furrow. Librigena with narrow ocular platform, oblique posterior margin, and long genal spine. Thorax with 12 to 15 segments, pleurae short anteriorly and growing in tr. width posteriorly, or 9th or 11th segment macrospinose. Pygidium subparabolic or cordate in outline, posterior margin occasionally with slight indentation on sagittal line; axis broad, conical, with up to 10 rings; pleural regions with more or less clearly developed subequal pleurae. | |
| M. Armstrong et al. 2020 | Frontal area 25% (23–26) of cranidial length, with sharp change in slope between preglabellar field and anterior border, and well-defined anterior border furrow. Seven thoracic segments. Pygidium semielliptical in outline with inflated pleural field, well-defined border furrow, and down-sloping border of even width occupying 10% (8–12) of pygidial length (sag). | |
| M. Armstrong et al. 2020 | Gently rounded anterior cranidial margin. Frontal area occupies 28% of cranidial length; down-sloping preglabellar field and slightly shorter, nearly flat anterior border differentiated largely by change in slope. Anterior border furrow shallow but clearly defined. Facial sutures diverge conspicuously anteriorly from palpebral lobes. Subtrapezoidal glabella with well-rounded front. Pygidium with short border of even width equal to about one-fifth (19%; 17–21) pygidial length (sag); separated from pleural field by strongly incised border furrow. Internal mould expresses at least nine axial rings and terminal piece of two segments, and faint, straight interpleural furrows. | |
| M. Armstrong et al. 2020 | Anterior border increases in length medially, producing nasute outline; maximum length roughly equal to length (sag.) of preglabellar field. Abrupt slope change between down-sloping preglabellar field and nearly flat anterior border; shallow anterior border furrow. Palpebral lobes situated posteriorly, near mid-length of glabella. Triangular pygidium with border occupying 25% of length (sag.) but narrowing abaxially. Border furrow faint; moderately incised axial furrows that weaken at axis tip. Pleural field gently inflated above border. At least nine axial rings and short terminal piece expressed on internal moulds, as are pits corresponding to small projections on
underside of exoskeleton. | |
| M. Armstrong et al. 2020 | Anterior border narrows abaxially with well-incised border furrow; steeply sloping preglabellar field. Moderately convex glabella that narrows (tr.) anteriorly to a broadly rounded front, and very shallow axial furrows. Posterolateral projections long and triangular, with posteriorly deflected margins. Pygidium subtriangular in outline with faint border, axial, and pleural furrows that all weaken posteriorly. Pitting (corresponding to small projections on underside of exoskeleton) preserved on internal mould. | |
| M. Armstrong et al. 2020 | Frontal area 24% (23–24) of cranidial length (sag.) with steeply sloping preglabellar field. Cranidium is slightly wider than long with evenly rounded anterior margin. Strongly convex glabella with nearly parallel sides, well-rounded front, and shallow occipital furrow expressed on the exoskeleton. Posterolateral projections long, extend to a narrow, tapered point. Semielliptical pygidium has distinct axial furrows that shallow at the terminal piece; pleural and interpleural furrows expressed only on internal mould. Border furrow nearly effaced on external surface, but inner edge of border marked on mould by termination of pleural and interpleural furrows (Pl. 27, fig. 4); border flexed steeply downward. Internal mould with pitted surface. | |
| J. D. Beech and J. C. Lamsdell 2021 | Eye tubercles each with two lenses; semicircular alae adjacent to posterior glabellar lobes; bilamellar fringe with opposed pits in outer surfaces, genal rolls steeply sloping, brim gently sloping, with stout girder on lower lamella separating these two parts, flattened prolongations of fringe varying in length; cephalic suture skirts marginal band of fringe. Hypostome pear-shaped in outline, with ovate middle body, large anterior, small posterior, and wings. Thorax with 12–29 segments, pleurae bent down ventrally at tips. Pygidium small, short (sag.), triangular, with few segments. External surface of glabella and genae with raised ridges in reticulate pattern, tuberculate, or smooth; minute tubercles on fringe between pits and marginal band, on internal rim, and elsewhere (modified from Fortey & Owens 1997). | |
| Y. Wu et al. 2021 | Houcaris with elongate distally tapering frontal appendage consisting of 16 podomeres, including three base podomeres and 13 claw podomeres; Cp2–Cp8 nearly square, Cp9–Cp12 rectangular and much longer than height; En2–En8 at least one and a half times as long as the height of the associated podomeres, and bear 5 distal auxiliary spines and 2 proximal auxiliary spines; auxiliary spines near the base part of endite are shorter; En1 stout and ventro-distally curved; two smaller setules projected from distal ventral surface of each claw podomere; Cp10–Cp13 project paired elongate dorsal spines arching forward; Cp13 also bearing one terminal spine and a pair of secondary dorsal spines (Modified from Hou et al. 1995). | |
| J. D. Beech and J. C. Lamsdell 2021 | Cephalic border not sharply set off from convex genae and preglabellar field; alae small, semicircular; facial sutures marginal, or with parallel anterior and posterior sections running close to each other and directed anterolaterally from eye tubercles to margin; genal caeca radiating over cheek lobes and in some cases extending onto cephalic border. Hypostome subrectangular, length (sag.) equal to that of glabella. Thorax with 20 or more segments; axis narrow; long (tr.) pleurae curving back at outer part may be extended into spines, with deep pleural furrows and convex posterior bands (Fortey & Owens 1997). | |
| J. D. Beech and J. C. Lamsdell 2021 | Exoskeleton small. Cephalon semicircular, characterized by anterior and posterior sections of facial sutures close to each other, both directed outward and backward; librigenae fused together through doublure, consisting of narrow dorsal strips connecting eyes to margin and to genal spines (modified from Fortey & Owens 1997). | |
| J. D. Beech and J. C. Lamsdell 2021 | Angle of cephalic curvature greater than 90°. Yoked librigenae, but lacking true bilamellar fringe. Facial sutures directed outward and forward. Small eye lobes, diverging posteriorly, with eye ridges anterolaterally directed. High cephalon convexity and equilateral glabellar lateral margins. Highly convex genae. Lacking alae. Narrow, rounded genal prolongations (i.e. spines). Pygidium of four to eight segments. | |
| D. J. Holloway 2021 | Anterior cephalic border less than twice as long sagittally as exsagittally, with broadly rounded margin; L1a small for genus; front of palpebral lobe lying well behind widest part of bullar lobe; anterior branch of facial suture converging gently in front of β–β, width α–α being much greater than width of fron- tal lobe. Hypostome 1.3 times as wide across shoulder as long (sag.); lateral margin strongly flexed outwards behind anterior wing and lateral border expanding gently; anterior lobe of middle body only slightly wid- er anteriorly than at outer end of middle furrow. Py- gidium with lateral margins subparallel, weakly con- vex in outline; maximum width approximately level with posterior end of axis; axis 0.3 times maximum pygidial width and approximately 0.4 times sagittal length; postaxial rib gently tapering backwards for less than half its length and subsequently expanding so as to exceed its anterior width distally. | |
| D. J. Holloway 2021 | Anterior cephalic border approximately 4 times as long sagittally as exsagittally, anterior margin well rounded; palpebral lobe extending forwards almost as far as maximum glabellar width across bullar lobe; anterior branch of facial suture converging strongly in front of β, width α–α being equal to width of frontal lobe. Hypostome 1.1 times as wide across shoulder as long (sag.); anterior lobe of middle body narrowing moderately strongly backwards to outer end of middle furrow; lateral border narrowing slightly posteriorly. Pygidium with maximum width situated well behind axis; lateral margins moderately convex in outline; axis about 0.25 times maximum pygidial width, with only one, poorly defined axial ring; postaxial rib subparallel-sided. | |
| D. J. Holloway 2021 | Occipital ring sloping steeply forwards medially; median glabellar lobe moderately convex sagittally, posterior half not strongly elevated above
bullar lobe; bullar lobe ovate, wider than median lobe measured across distal end of S1, expanding forwards. Three pairs of prominent, paired tubercles present on median lobe, another three on bullar lobe arranged in a forwardly-diverging row, and a central one on L1b together with a slightly smaller one situated more anterolaterally. Pygidium 1.6 –1.8 times as wide as long (sag.), maximum width level with posterior end of axis; second axial ring very poorly defined. Anteriormost marginal spine with distal end level with 0.75 sagittal length of pygidium; second spine terminating level with apex of notch between third spines; third spine acutely pointed. | |
| D. J. Holloway 2021 | Cranidium moderately convex; anterior margin strongly curved in outline, roughly semicircular;
anterior border long (sag., exsag.), only slightly shorter medially than laterally; longitudinal glabellar furrow extending backwards to occipital furrow but shallow and poorly defined posterior to bullar lobe which is confluent with L1b; L1a present; median part of L1 gently convex (sag., exsag.), narrowest part of median glabellar lobe immediately in front weakly depressed; bullar lobe subparallel-sided, oriented with long axis about 45° to sagittal line; frontal lobe extending in front of bullar lobe by about 40 % glabellar length (sag., excluding occipital ring); palpebral lobe situated with midlength opposite medial part of S1. Pygidium subpentagonal in outline, with three pairs of broad, flattened and rather long marginal spines separated by broad notches; pleura on first segment curves gradually backwards abaxially, with tip of spine lying more or less opposite midlength (sag.) of postaxial region; axis comprising 25 % maximum pygidial width anteriorly or less, and about 33 % sagittal length. | |
| D. J. Holloway 2021 | Cranidium curving through about 110°–120° in lateral profile, vertical or only slightly overhanging anteriorly in dorsal view; axial furrow converging strongly forwards immediately in front of posterior cephalic margin, thereafter subparallel as far as lunette, very short and weak in front of lunette; glabella with slight independent convexity (tr.) posterior to lunette, lacking independent convexity farther forward; anterior branch of facial suture converging weakly forwards. | |
| Y. Wu et al. 2021 | Tamisiocaridids with paired elongate, distally tapered frontal appendages consisting three base podomeres and 13 claw podomeres; claw podomeres in proximal and intermediate region bear paired elongate endite altering long/short on odd/even numbered claw podomeres, each of which processes multiple distal auxiliary spines; En1 stout and slightly curved; claw podomeres are separated by triangular flexible arthrodial membrane. | |
| Y. Wu et al. 2021 | Radiodonts with frontal appendages bearing paired thin endites much longer than the height of the associated podomeres; endites on proximal and intermediate claw podomeres bearing multiple slender auxiliary spines (modified from Pates and Daley 2019). | |
| D.-G. Jiao et al. 2021 | Radiodont with slender frontal appendages, which comprise at least 17 podomeres; proximal podomeres have a tall rectangular lateral outline, which changes to approximately square distally; endites alternate long/short on subsequent podomeres, and reduce in length along the appendage; small triangular terminal spine on distalmost podomere. | |
| M. F. Tortello 2022 | A species of Athabaskia having short genal spines; pygidial axis subparallel sided to slightly tapered backwards, containing four rings and an extended terminal piece, length ∼75–85% of pygidium; pleural fields and pygidial border of moderate proportions, showing five apparent pleural and interpleural furrows of equal length that almost reach the pygidial margin; rear pygidial margin broadly rounded or slightly indented; well-preserved pygidia show a pair of delicate fossulae on the fifth pleural furrow, close to the posterolateral corners of the axis. | |
| D. J. Holloway et al. 2022 | Cephalon of low convexity (sag., exsag., tr.). Glabella very wide posteriorly, there comprising about 75% cranidial width, narrowing forwards abruptly for short distance immediately in front of posterior margin, thereafter narrowing less strongly as far as lunette, expanding at similar rate in front of lunette as far as axial furrow is distinguishable. Palpebral lobe of moderate size, situated its own length or less from posterior cephalic margin; anterior branch of facial suture diverging moderately strongly forwards from γ to β. Librigena with posterior margin curving gently backwards to strongly rounded genal angle, and with very weak vincular furrow running close to and subparallel with inner edge of doublure adaxial to genal angle. Rostral plate lenticular in ventral view, gently convex (sag., exsag.). Hypostome parabolic in outline behind large anterior wings, without shoulders; anterior lobe of middle body elliptical in outline, posterior lobe with prominent maculae. Thorax nonfulcrate, with very wide axis comprising most of segmental width (tr.); axial furrow indistinct. Pygidium more convex than cranidium, subelliptical in outline, as wide or a little wider than long, maximum width well in front of midlength (sag.); axis very wide anteriorly, there comprising about 60% maximum pygidial width, otherwise undefined; anteriormost pleural furrow impressed; doublure increasing in width posteromedially, its inner margin with a bifid forward projection medially. | |
| D. J. Holloway et al. 2022 | Cephalon excluding genal spines subpentagonal in dorsal outline; fringe short (sag., exsag.) frontally, markedly expanded in width laterally where it is strongly rounded to subangular in outline; two pit arcs present in front of glabella, progressively increasing to five regularly arranged arcs laterally where they are aligned diagonally to curvature of outer margin of fringe at its widest part, less complete arcs lying external to more complete ones; pits in two arcs frontally and inner three arcs anterolaterally and laterally arranged in about 15–17 radial rows (half fringe, excluding posterior row); pits in outer two regular arcs on lateral part of fringe out of phase with those in inner three arcs and with each other; pits external to these arcs on lateral extremity of fringe, including marginal pits, mostly small and mostly lacking regular arrangement; no F pits apart from posterior fossula. Glabella with convex (sag., exsag.) preoccipital ring; occipital ring without medial spine. No preglabellar field. Lateral eye tubercles situated relatively close to glabella. Reticulate sculpture on glabella and pitted sculpture on genal field. Pygidium very short, only first segment clearly defined on axis and pleural region. | |
| N. M. Handkamer et al. 2022 | Zacanthoidid with axial furrow moderately defined, two pairs of lateral glabellar furrows poorly defined, palpebral lobe short, posterior border long and narrow; genal spine oriented obliquely backwards; four thoracic segments; macropygous, ring furrows effaced, pleural field narrow, pleural furrows poorly defined or effaced, one pair of border spines oriented obliquely backward or subparallel. | |
| N. M. Handkamer et al. 2022 | Zacanthoidid with a glabella subrectangular, palpebral lobe long; advanced genal spines long; two or four thoracic segments, axis and pleura equal in width; macropygous, six or eight axial rings, three to five pleural furrows well defined, one pair of posteriorly located border spines. | |
| N. M. Handkamer et al. 2022 | Zacanthoidid with glabella anteriorly wide, palpebral lobe long; genal spine non-advanced; nine thoracic segments, pleural spines long, macropleural spines on the fifth and ninth segments; oval-shaped pygidium subisopygous, four axial rings, four pleural furrows, border furrow well defined, border spines absent. | |
| P. Cederström et al. 2022 | Species of Ellipsostrenua with glabella tapering forward to 70–76% maximum width across L1, maximum width 39–44% maximum cranidial width across centre of palpebral lobes; frontal lobe with relatively distinct median kink at anterior margin; faint lateral extension of frontal lobe present on internal moulds only; occipital ring of 16–20% cephalic length; palpebral furrows shallow in its anterior part, better developed posteriorly, palpebral lobes slightly tilted against slope of fixigenae; anterior cephalic margin with comparatively strong curvature and with slight median angulation. Cuticle granulose. (Data for adult individuals only). | |
| P. Cederström et al. 2022 | Species of Ellipsostrenua with glabella tapering forward to 69–75% maximum width across L1, maximum width 38–42% maximum cranidial width across centre of palpebral lobes; frontal lobe with slight median arcuation at its anterior margin; occipital ring of 16–19% of cephalic length exclusive occipital spine, extended into a long occipital spine; palpebral furrows well‐developed in the median and posterior sectors, well recognizable on the exterior of the cuticle; preglabellar field slightly broader (sag.) than anterior border, well elevated; anterior cephalic margin with moderately strong curvature, without distinct median angulation. External surface of cuticle granulose except for principal furrows. (Data for adult individuals only). | |
| T. Unger et al. 2022 | S1 shallowing and curving backwards axially, S2 and S3 curving forward, S3 and S4 short, all well-defined; occipital ring broadens axially; pygidium nearly circular in outline with wide axis ending in a blunt point at about half pygidial length (based on Lake 1934, 1935; Fletcher 1972b, with modifications). | |
| D. E. G. Briggs et al. 2023 | Head bearing five paired appendages, the first bearing three flagella, the second and third uniramous with a substantial gnathobasic basipod; the posterior two head appendages and all trunk appendages bear a robust spinous endopod, a flap-like exopod with a strengthening rod, and an exite proximally with an array of long filaments. The trunk consists of 10 appendage-bearing segments, an apodous abdomen of four segments and a tail spine. | |
| M. F. Tortello 2023 | A species of Cedaria with a large exoskeleton; noticeable lateral glabellar furrows; strongly divergent anterior branches of the facial suture; a well-developed preglabellar field that is much longer (sag.) than cranidial border; elevated palpebral lobes lacking palpebral furrows; and a semicircular pygidium with a narrow (tr.), long pygidial axis composed of 7–8 rings, five pairs of distinct pleural furrows, and a slender border. | |
| R. G. Johnson 2024 | Genicranium narrow, length (sag.) between 75% and 85% of width (tr.). Glabella inset and preglabellar furrow bowed upwards in anterior view. No tubercles on occipital ring. Posterior border moderately wide (tr.), between 41% and 50% of width (tr.) of occipital ring. Alar depression present and course of inner margin of fringe across anterior of genal area straight. Genal roll sloping moderately anteriorly (sag.) at between 420 and 500 to horizontal. Brim wide (sag.) >35% of cephalic length (sag.) and steepening to less than vertical on prolongations. Brim width ratio between 0.86 and 0.94. Spaces between brim perforations inflated. Prolongations moderately short (exsag.), between 76% and 85% of cephalic length (sag.). | |
| R. G. Johnson 2024 | Cephalon strongly vaulted and ovoid. Preglabellar field narrow (sag.) (20% to 30% of
width (tr.)) and posterior border short (tr.) (20% to 30% of width (tr.) of occipital ring). Genal area narrow, width (tr.) under 54% of length (exsag.). Alae large, extending over 60% of way across (tr.) genal area. Eye lobe forms prominent inflated cone or oval extending from edge of axial furrow to near internal margin of fringe. Anterior boss absent or barely inflated and girder without kink. Genal roll and brim free of caeca. Brim slightly concave (sag.), straight laterally, gently to moderately sloping and steepening quickly to near vertical on prolongations. Brim narrowing (tr.) moderately strongly to very strongly posteriorly (brim width ratio <0.68). Extension of girder meeting internal rim one third of the way down prolongation. Thorax up to 31 segments | |
| R. G. Johnson 2024 | Cephalon not strongly vaulted and genicranium widest at alae. Glabella wholly or partly carinate and narrow (tr.), at S1 between 54% and 66% of length (sag.) of glabella anterior to S0. Eye lobes weakly inflated, genal area moderately wide, width (tr.) between 54% and 66% of length (exsag.). Brim sloping moderately steeply at between 210 and 300 and strong network of caeca across whole brim. | |
| R. G. Johnson 2024 | Cephalon strongly vaulted. Occipital furrow medially straight (tr.). Alae depression
absent. Eye lobe short (exsag.), between 15% and 31% width (tr.) of glabella at S0. Course of inner margin of fringe across anterior of genal area convex. Girder moderately wide (sag. and exsag.), between 2.2% and 3% of width (tr.) of genicranium. Brim barely convex to moderately convex and sloping steeply at between 250 and 450. Standardised brim perforations moderately fine, less than 150 μm. Exterior rim fine and marginal band sloping down under cephalon. Thorax of up to 16 segments. | |
| R. G. Johnson 2024 | Cephalon ovoid. Occipital furrow broad (sag.), equal to or wider (exsag.) than posterior
border. Width (tr.) of posterior border between 31% to 40% of occipital ring width (tr.). Genal area moderately broad, inner margin of fringe across anterior of genal area straight and Eye lobes with three lenses each. Tubercles on occipital ring, palpebral lobes and dorsal surface of internal rim. Girder narrow (sag. And exsag.), 1.5% to 2.2% of genicranium width (tr.). Brim width ratio between 0.70 and 0.85. Extension of girder meeting internal rim halfway down prolongation. Thorax with 20 to 24 thoracic segments. Pleural furrows shallow and narrow, and distal ends of pleurae rounded. | |
| R. G. Johnson 2024 | Glabella inset and narrow, width (tr.) being less than 67% of length (sag.). Preglabellar
furrow straight in anterior view. Alae small, posterior border wide (tr.). Fine granulation on glabella, occipital ring and genal area. Inner margin of fringe marked by narrow, faint furrow. Genal roll sloping gently anteriorly, at under 500, more steeply laterally at over 550. Perforations on genal roll smaller than brim perforations. On brim, spaces between perforations strongly inflated. In lateral view, prolongations are strongly tapered, with internal and external rims joining to form long genal spines. | |
| R. O'Flynn et al. 2024 | Emended after Hou and Bergström (1997). Six cephalic segments covered by head shield; from anterior to
posterior, these are: protocerebral/ocular segment that carries paired optic lobes/eyes; post-ocular segment with paired raptorial deutocerebral frontal appendages (A1); and four subsequent segments each bearing a biramous appendage pair. A1 anterolaterally orientated, consisting of ≥ 16 podomeres that bear paired inward-facing spines that decrease in length gradually towards tip. Thorax comprises seven tergites with short pleural spines, each thoracic segment with one pair of biramous appendages. Large pygidium with a terminal tail spine and two pairs of marginal spines. Axis terminates in posterior half of pygidium. | |
| R. G. Johnson 2024 | Cephalon ovoid with widest point across or just anterior to alae. Brim strongly convex, less steeplysloping than genal roll; S1 furrow narrow and deeply incised. Alae anterolaterally directed, eyes lobes small (exsag.) and reniform. Course of inner margin of fringe across anterior of genal area concave. Brim not steepening on prolongations and inner rims subparallel. Prolongations long, pointed, dagger-like in shape, with long terminal spine carried unusually on upper lamella. | |
| R. G. Johnson 2024 | Furrow at S1 curves around top of muscle insertion area on glabella. Relatively large and wide (tr.) ala, anterolaterally directed; course of inner margin of fringe across anterior of genal area concave, anterior boss inflated and preaxial furrows converging towards girder. Brim width ratio between 0.66 and 0.75, standardized brim perforations fine (<100μm) and exterior rim with fine granulation. | |
| J. Drohojowska et al. 2024 | The new genus is similar to modern day species in the genus Tetraleurodes based on its oval body shape; dark colouration; striate submargin apparently differentiated from the dorsal disk by a suture/furrow and encompassing the body; its toothed lateral margin undifferentiated at the tracheal and caudal opening; and its cordate shaped vasiform orifice. It is also similar to Aleurolobus species which have a trilobed figure over the vasiform orifice; however several Tetraleurodes species also have a similar trilobed area as in the new genus and the other characteristics such as the shape of the vasiform orifice, apparent lack of dorsal setae and undifferentiated lateral margin indicate that it is more similar to Tetraleurodes species.
The new genus differs from Tetraleurodes and almost all other modern-day whitefly species by having all of the segments of the body distinctly defined by deep sutures; in most extant whiteflies the segmentation of the cephalothorax is not as distinct and the sutures usually do not extend to the submarginal region as in this genus. It is the second fossil genus to be described based on characteristics of the puparium (Drohojowska et al. 2023) and the third fossil species described from the puparium (Rietschel 1983; Drohojowska et al. 2023). Not identified puparia have also been found in upper Eocene deposits of Isle of Wight (Szwedo et al. 2019). | |
| R. G. Johnson 2024 | Narrow cephalon, width (tr.) around 1.2 times sagittal length and widest (tr.) just posterior to eyes. Glabella cone-shaped, anteriorly rounded. Extension of axial furrow across ala distinct. Alae inflated and alar furrows proximally deep, becoming very shallow distally. Alar depression pronounced. Prolongations around same length (exsag.) as cephalon (sag.). In lateral view external rim posteriorly curving upwards and internal rim wavelike in shape and terminated by short genal spine. Over 25 thoracic segments. | |
| R. G. Johnson 2024 | Wide cephalon, width (tr.) across alae about 1.4 times length (sag.). Glabella oval and strongly inflated. Brim barely sloping, slightly concave and narrowing moderately strongly posterolaterally. Alae inflated and axial furrows not extending across alae. Alar furrows become shallower distally. Fine pointed tubercles scattered along top of glabella, on dorsal surface of eye lobes and occipital ring and along dorsal surfaces of external and internal rims. Prolongations up to 15% longer (exsag.) than cephalon (sag.). In lateral view, external rim posteriorly curving upwards and internal rim wave-like in shape. Over 25 thoracic segments. | |
| R. G. Johnson 2024 | Cephalon widest (tr.) across anterior of alae. Large ovoid glabella, anteriorly steeply-sloping with prominent tubercles straddling crest. Tubercles on dorsal surface of eye lobe, occipital ring and internal rim. Alae inflated and moderately large. Alar depression deep and alar and preglabellar furrows well defined. Shallow pitting on post ocular genal area. Narrow (sag.) brim, narrowing moderately posterolaterally (brim width ratio 0.64) with moderately large perforations decreasing in size distally. In lateral view, external rim curves slightly upwards and profile of internal rim weakly wave-like. | |
| R. G. Johnson 2024 | Axial furrows are convex and deep, and preglabellar furrow well defined. Tubercles and faint caeca on genal area. Perforations on genal roll same as on brim. Brim is moderately steeply-sloping and brim width ratio is between 0.68 and 0.75. Dorsal surface of external rim overhangs marginal band and carries tubercles. Prolongations over 90% of cephalic length (sag.), extension of girder meeting internal rim about halfway down prolongation. Internal rim in lateral view strongly wave-like in profile and distal end of prolongation sharply upturned. | |
| R. G. Johnson 2024 | Cephalon widest across eyes and glabella is narrow, cone-shaped, rounded anteriorly with suppressed tubercles straddling its crest. Brim width ratio low at around 0.50, brim slightly concave (sag.), becoming gently convex close to posterior border, alar furrows effaced distally, perforations on brim very fine. Prolongations shorter (exsag.) than cephalon (sag.). Pleurae shortening (tr.) posteriorly over last fifteen thoracic segments to become very short, giving pointed appearance to thorax. | |
| R. G. Johnson 2024 | Cephalon widest across eyes. Glabella narrow anteriorly-rounded cone without tubercles on dorsal surface. Brim broad (sag.), profile straight (neither convex or concave) and moderately steeply-sloping. External border runs straight, slanting adaxially, from point where it passes eyes to two thirds of way down (exsag.) the prolongations. Eyes set close to midway between axial furrow and inner margin of fringe. Perforations on brim moderately large, becoming smaller distally until middle of rim. Perforations on genal roll much finer except proximal to girder, where there are two rows of larger perforations. Terminal spines long, slanting obliquely adaxially. | |
| R. G. Johnson 2024 | Narrow cephalon, widest (tr.) midway between eyes and alae, width (tr.) is just over 1.1 times sagittal length. Glabella anteriorly rounded cone. Eye lobe with short transverse eye ridge on posterior edge. Alar furrows becoming shallower distally. Prolongations are about same length (exsag.) as cephalon (sag.). In lateral view, internal rim has subdued wave-shaped profile and is terminated by short genal spine. Twenty thoracic segments. | |
| R. G. Johnson 2024 | Cephalon ovoid, nearly oval, with external and internal rims curving strongly adaxially. Glabella broad, crest rounded with fine granulation and pitting, and axial furrows convex. Alae anterolaterally directed and eye lobes located away from axial furrow. Brim straight to slightly concave in profile, gently sloping and with large brim perforations (diameter around 220 μm on standardised brim). Well-developed caeca on brim and genal roll. Length (exsag.) of genal prolongations over 90% of cephalic length (sag.). | |
| R. G. Johnson 2024 | Glabella anteriorly tapered with no carination and axial furrows convex. S1 sloping backwards, before curving forward. Occipital furrow medially bent forward, eye ridge poorly defined. Genal roll with caeca and brim perforations moderately large and becoming smaller away from girder. Well-developed caeca crossing brim. Genal prolongations moderately short and internal rims subparallel. | |
| R. G. Johnson 2024 | Genicranium is narrow (tr.), glabella inset and carinate, axial furrows straight, occipital furrow and occipital ring narrower (sag.) than posterior border (exsag.). CMO on occipital node with four pits arranged in a square, alae longer (tr.) than they are wide (exsag.), no alar depression and genal area without caeca, tubercles or granulation. Inner margin of fringe anterior to glabella not marked by perforations larger than those on genal roll. Brim is concave (sag.), straight laterally and quickly steepening to near vertical on prolongations; brim width ratio under 0.60. | |
| R. G. Johnson 2024 | Glabella is conical, carinate and with tectiform flanks and tubercles along crest. S1 short and shallow. Eye lobes inflated, tall, subcircular and with two eye lenses each. Brim is wide (sag.) and steepens strongly on prolongations. Brim width ratio is around 0.70. In lateral view, internal rim nearly horizontal with slightly wave-like profile. Pleural furrows narrow and well defined. | |
| R. G. Johnson 2024 | Cephalon widest at eye lobes. Muscle attachment scars on glabella visible at S2, S3 and S4 barely reaching out of axial furrow. Alae well-defined and with alar furrow and ridge. Eye lobes close to inner margin of fringe; eye ridge and genal ridge present and inner margin of fringe concave across the anterior of genal areas. Genal roll sloping anteriorly moderately steeply (just under 600) down to narrow girder with prominent oval recesses beneath, formed by caecal ridges radiating down from girder. Brim wide (sag.) (between 25% and 38% of cephalic length (sag.)), external rim narrow (less than 10% of width (sag.) of brim). In lateral view, internal rim is straight and slopes down posteriorly at a shallow angle. | |
| R. G. Johnson 2024 | Cephalon sub oval, widest (tr.) at alae. Axial furrows broad and shallow. No muscle insertion furrows at S2, S3 or S4, and S1 only shallow depression in axial furrow. Alar furrows almost effaced. Eye lobes barely inflated and eye ridge indistinct. Genal roll gently sloping. Brim convex and brim width ratio around 0.90. No caeca on brim, and perforations around outside of brim large and widely spaced. External rim tilted outwards with marginal band sloping under cephalon. In lateral view, internal rim only slightly concave and steeply declining to terminal spine. | |
| R. G. Johnson 2024 | Cephalon sub oval, widest (tr.) at alae. Genicranium widest (tr.) at posterior border. Glabella anterior to S0 moderately broad (width (tr.) 0.66 to 0.75 of length (sag.)) and with no furrows at S2, S3 or S4. Alae small (tr.) and well defined, and no alar depression. Eye lobes set close to internal margin of fringe. Inner margin of fringe across anterior of genal area convex. Girder broad (sag.) (>2.3% of width (tr.) of genicranium), meeting internal rim one half or little over one half of way down prolongation. Brim concave, gently sloping with proximal edge steepening slightly on prolongations. Brim width ratio between 0.77 and 0.95. Marginal band concave with ridges top and bottom. 20 to 24 thoracic segments with inner and outer portions of pleurae aligned (tr.). | |
| R. G. Johnson 2024 | Genicranium broad, glabella subcylindrical, rounded anteriorly and preglabellar furrow well defined. Eye lobes with two lenses, and eye and genal ridges well defined. Whole of inner margin of fringe marked by row of bigger perforations. Anterior boss mildly inflated, genal roll steeply inclined (sag. and tr.) and girder kink Σ-shaped. Brim barely concave and width ratio around 0.95. Fine perforations on brim (around 100μm) and finer on genal roll. Internal rims subparallel. Thorax with 20 segments. Blister-like tubercular granulation on dorsal surfaces of external and internal rims, occipital ring and first 5 axial rings of thorax. | |
| R. G. Johnson 2024 | Cephalon nearly ovoid; glabella inset, broad, strongly vaulted and with convex axial furrows, converging anteriorly. Genal roll gently sloping (sag. and exsag.) and without caeca. Axial and alar furrows deep; preaxial furrows forming circular depressions, crossing inner margin of fringe. Eye lobes inflated and with three eye lenses each. Brim width ratio 0.80 to 0.85, and diameter of standardised brim perforations moderate to large (>145 μm). | |
| R. G. Johnson 2024 | Glabella subcylindrical, rounded anteriorly, and S1 curving anteriorly near crest. Alar depression present and ala close to axial furrow is inflated to rise slightly above level of genal area. Eye lobe weakly inflated with four lenses, two very small contiguous central eye lenses (Pl. 26G) dorsolaterally orientated. Anterior boss inflated and girder moderately narrow (just over 1.5% of width of genicranium). Brim barely sloping but steepening slightly posteriorly along inside edge on prolongations. Brim width ratio ≈ 0.95. External rim fine (< 10% of width sag. of brim). Thorax with 20 segments. | |
| R. G. Johnson 2024 | Cephalon widest just anterior to alae, glabella carinate, and no eye ridges. Anterior boss barely inflated and joined to glabella by gentle swelling in preglabellar field. Genal roll sloping anteriorly at under 500. Brim narrow (sag.), at 28% of length of cephalon (sag.); and very fine brim perforations with standardised diameter of ≈70μm. Prolongations short, ≈ 66% of length (sag.) of cephalon. Internal and external rims robust, curving adaxially on prolongations | |
| R. G. Johnson 2024 | Genicranium narrow, glabella very broad without any muscle attachment furrows, and L1 not inflated. Alar furrows and occipital furrows very shallow. Preglabellar furrow barely distinct and anterior boss joined to glabella. Brim barely sloping and with concave profile. Marginal band with three ridges. Internal rims subparallel. | |
| R. G. Johnson 2024 | Genicranium broad and strongly vaulted. Glabella crest carinate, anteriorly rounded and free of tubercles or granulation. Axial furrows converging strongly towards well-defined preglabellar furrow. Occipital node barely inflated; tubercles on genal area. Eye lobes small (exsag.), moderately strongly inflated and with two lenses each, anterior lens being largest. Shape of lenses rectangular with rounded corners to subtriangular. Eye ridge absent. Prominent small tubercles on occipital ring, posterior border and internal rims. No caeca on genal roll. Brim width ratio 0.80, and standardised brim perforations around 170μm. | |
| R. G. Johnson 2024 | Broad (sag.) brim, around 37% of cephalic length (sag.), with moderately high density of fine perforations. Eye tubercles set well away from inner margin of fringe. Transverse eye ridges, which turn anteriorly in axial furrow. Preglabellar boss barely inflated and no median girder kink. Each eye lobe with three contiguous eye lenses. | |
| R. G. Johnson 2024 | Genicranium very narrow. Glabella narrow, with mildly carinate crest and convex flanks. Axial furrows do not cross junction of ala and L1, and converge strongly anteriorly. Preglabellar furrow indistinct and preglabellar field absent. Genal area narrow, width (tr.) around half the length (exsag.). Alae also narrow (tr.) and genal roll moderately sloping (sag.). Brim strongly concave and not steepening on prolongations. Thorax with over 21 segments. | |
| R. G. Johnson 2024 | Cephalon nearly ovoid; genicranium moderately vaulted; genal roll without caeca and sloping gently anteriorly and laterally. Glabella narrow, very strongly vaulted, inset and with fine tubercles and pitting. Axial furrows deep and straight; preglabellar furrows bowed upwards in anterior view; and preaxial furrows forming circular depressions across inner margin of fringe. Occipital ring higher than glabella anterior to S0. Genal area very narrow, and genal ridges present. Eye tubercles tall, with three lenses and fine tubercles and pits on dorsal surface. Brim width ratio high, at 0.87. | |
| R. G. Johnson 2024 | Cephalon ovoid, glabella joined to anterior boss, L1 not inflated, occipital furrow shallow and occipital ring not medially broadened. Alar furrows very shallow or effaced; eye lobes set away from inner margin of fringe and with two lenses each, anterior being largest. Course of inner margin of fringe across anterior of genal area concave. Marginal band profile straight and internal rim meeting external rim before joining at end of prolongation. Genal spines long. | |
| R. G. Johnson 2024 | Genicranium wide, length (sag.) 1/3 width (tr.). Glabella with tectiform flanks, rounded crest and joined to anterior boss. Occipital ring length (sag.) same as posterior border length (exsag.); posterior border wide (tr.), half width of occipital ring (tr.). Alae subtriangular and genal roll sloping gently anteriorly. Brim width ratio just under 0.80; brim gently sloping anteriorly, steepening laterally to over 600 on prolongations. Perforations on brim moderately fine and internal rims are subparallel. | |
| R. G. Johnson 2024 | Glabella strongly vaulted and subcylindrical, tapering anteriorly (axial furrows 150 to axis). Preglabellar furrow very shallow, glabella joining with narrow, moderately inflated anterior boss that does not continue down to girder. Alar furrow very shallow and often effaced. Eye lobes slightly inflated, long (exsag.) and reniform. Brim straight, gently sloping and steepening moderately strongly on prolongations. Brim width ratio between 0.7 and 0.8. Perforations over 150 μm on standardised brim. Marginal band with three lateral ridges. | |
| R. G. Johnson 2024 | Cephalon pyriform, strongly vaulted and widest at eye lobes. Glabella inset and joined to anterior boss. S1 not rising out of axial furrow. Genal areas are strongly convex (sag. and exsag.) and genal roll is steeply sloping, over 600 anteriorly and over 700 laterally. Brim wide (sag.), 35% of cephalic length (sag.), barely sloping and brim width ratio very low, at 0.52. Perforations on brim very fine (< 80 μm) and even finer on genal roll. Brim steepens quickly to near vertical on prolongations. In dorsal view, genal spine not following curve of exterior rim and curving upwards in lateral view. | |
| R. G. Johnson 2024 | Cephalon inverted U in shape, and genicranium strongly vaulted. Occipital ring not medially broadened and in lateral view, nearly upright and lower than glabella anterior to S0. Course of inner fringe across anterior boss not marked by row of large perforations. Anterior boss barely inflated. Brim very narrow, 25% of cephalic length (sag.), and nearly flat. Marginal band with three ridges. Internal and external rims meeting and joining at end of prolongations. Anterior of thorax not broadening posteriorly and pleural furrows effaced. | |
| R. G. Johnson 2024 | Genicranium strongly vaulted and narrow, width (tr.) 1.3 times length (sag.); preglabellar field wide, length (sag.) 16% of width (tr.); and preglabellar furrow bowed up in anterior view. Alae wide (tr.), 45% of glabella width (tr.) at S0, slightly wider than width (exsag.) of ala at axial furrow. Alar depression and faint genal ridge present. Eye lobes oval, inflated and anterior and posterior eye lenses of equal size. Genal roll sloping steeply anteriorly at ≈600 and laterally at ≈700. Brim narrow (sag.), 29% of cephalic length, and brim width ratio low, at 0.54. Marginal band near vertical anteriorly, increasingly sloping down under cephalon posterolaterally. Tubercles on occipital ring, posterior border, internal rims and marginal band, and fine pitting on glabella, genal areas, axial rings and pleurae. | |
| R. G. Johnson 2024 | Cephalon is moderately vaulted, glabella strongly vaulted. Alae are inflated proximally, and alar furrows erased distally. Eye lobes pronounced. Slope of genal roll is moderate, anteriorly continuing anterior slope of glabella. Anterior boss is barely inflated. Perforations on genal roll decrease in size upwards, away from girder. Brim barely sloping; brim perforations widely spaced, decrease rapidly distally to become very fine, before becoming larger again close to rim. | |
| R. G. Johnson 2024 | Cephalon ovoid, glabella with tectiform flanks. S1 short, sloping backwards. L1 weakly inflated and axial furrows effaced at junction of L1 and alae. Occipital furrow shallow, occipital ring pitted and in lateral view, higher than glabella anterior to S0. Genal area very broad (tr.). Anterior boss joined to glabella and preglabellar furrow effaced. Perforations becoming smaller for a short distance away from girder. Girder kink pronounced. Brim narrowing strongly posterolaterally. Marginal band with ridge top and bottom. Tubercles on dorsal surface of internal rims. Pygidium with four pleurae and tubercles on terminal piece. | |
| R. G. Johnson 2024 | Genicranium broad (width (tr.) 1.2 times length (sag.)), and genal roll gently sloping anteriorly. Glabella mildly tectiform, and S1 small depression in axial furrow. Alar furrows effaced. Girder very broad (sag., exsag.) and with central row of large perforations. Prolongation long (exsag,), 1.2 times cephalic length (sag.); external and internal rims flexing abaxially close to distal end of prolongation. | |
| R. G. Johnson 2024 | Cephalon is pyriform, and genicranium narrow to very narrow (width (tr.) over 85% of length (sag.)). Glabella carinate and joined medially to barely-inflated anterior boss. Preglabellar furrow very shallow and bowed upwards in anterior view. Occipital furrow medially bowed posteriorly. Genal roll gently sloping anteriorly (at under 420 with horizontal). Brim convex, sloping steeply at between 310 and 400 and quicky steepening to near vertical on prolongations. External rims tilted outwards and marginal band sloping down under cephalon. Internal rim has straight profile in lateral view. Thorax with over 24 segments. Outer portion of pleurae in line with inner portion and curving anteriorly distally. | |
| R. G. Johnson 2024 | Cephalon mildly pyriform, genal area narrow (width (tr.) 34% of length (exsag.)). Alae are as wide (tr.) as they are long (exsag.). Genal area narrow. Inner margin of fringe across anterior of genal area straight, and anterior boss barely reaching down to girder. Caeca present on genal roll. Brim sloping at 300, narrow (sag.) (27% of cephalic length (sag.) in dorsal view) and has brim width ratio of just over 0.75. Perforations on brim become smaller for short distance away from girder. Exterior rim curves sharply upwards at posterior of prolongation, to join with internal rim at near right angle. Genal spine short, slanting obliquely upwards and adaxially. | |
| R. G. Johnson 2024 | Cephalon mildly pyriform; glabella sloping gently anteriorly and narrow, width (tr.) just over 50% of length (sag.) of glabella anterior to S0. Axial furrows nearly straight and weakly converging anteriorly. Preaxial furrows also converging and initially deep, becoming shallower anteriorly as anterior boss deflates to disappear close to girder. Occipital ring with prominent occipital node. Alar furrow well-defined and eyes elongated (exsag.). Genal roll sloping anteriorly at same angle as anterior of glabella, just under 25º. Laterally, slope steepens strongly to around 55º. Brim strongly convex, moderately broad (sag.) and steeply sloping at around 35º. | |
| R. G. Johnson 2024 | Cephalon strongly pyriform and vaulted. Glabella with blister-like tubercles, slopes gently anteriorly at < 20º. Alae with interalar furrow and alar depression. Pits and faint caeca on genal area. Narrow preglabellar furrow, anterior boss barely inflated and not joined to glabella. Brim very broad and sloping steeply anteriorly at 35º, increasing to over 47º laterally and becoming near vertical on prolongations. In lateral view, in line with eye lobe, brim makes up half the height of cephalon. Brim width ratio low, at around 0.60, no row of larger perforations at proximal edge. | |
| R. G. Johnson 2024 | Cephalon widest at eye lobes, width (tr.) being over 1.46 times length (sag.). Flanks of glabella convex and axial furrows tapered anteriorly. Occipital furrow narrow and occipital ring not medially broadened. Posterior border short (tr.), equal to 25% of occipital ring width (tr.). Perforations on genal roll becoming smaller posterolaterally and no row of larger perforations at the inner margin of fringe across the anterior boss. Brim width ratio high, at around 0.92. Perforations on brim very large and forming net-like web across the dorsal surface. External rim very fine. In dorsal view, internal rim slanting diagonally abaxially, before curving strongly adaxially. No row of larger perforations under internal rim, and brim steepens only moderately on prolongations. | |
| R. G. Johnson 2024 | Cephalon mildly piriform. Genicranium moderately vaulted, glabella more strongly vaulted, tectiform and carinate. Occipital ring in lateral view near vertical. Eye lobes long (exsag.), reniform in shape and eye ridge low and broad. Internal rim slanting abaxially over first 2/3 before curving strongly adaxially. In lateral view, prolongations strongly tapered and external and internal borders converge posteriorly to run adjacent to each other along final third of prolongation. External and internal rim very robust. First five thoracic axial rings with axial tubercle. Pleural furrows narrow (exsag.), deepening posteriorly. | |
| R. G. Johnson 2024 | Cephalon widest at eyes; glabella not inset; L1 not inflated or barely inflated; occipital ring broader (sag.) than posterior border; posterior border moderately short; no preglabellar field; eye lobes large, extending to axial furrow; and no eye ridge. Course of inner margin of fringe across anterior of genal areas slightly convex; anterior boss barely inflated, attached to glabella but not reaching down to girder. No girder kink. Brim wide (sag.; 35% of cephalic length (sag.)). | |
| S. R. Losso et al. 2025 | Concilitergans in which the thorax has tergites with non-effaced boundaries and well-developed pleural spines, and a pygidium with variable number of posterolateral spines and a medial terminal spine
with a broad base. | |
| S. R. Losso et al. 2025 | Artiopodans with broad and nearly flattened dorsal exoskeleton and well-developed rounded anterior sclerite. Axial body region is gently raised but lacking axial furrows. Thorax consists of 6–14 tergites with edge-to-edge articulations and variably fused tergite boundaries. Posterior cephalic margin and anterior tergites gently reflexed anteriorly. | |
| S. R. Losso et al. 2025 | Concilitergans with raised eye bulges on dorsal exoskeleton accommodating ventral stalked eyes. Large anterior sclerite located in central notch of anterior margin of cephalon with notch occupying less than a third of the anterior margin in most taxa (absent only in Tegopelte). Postantennal appendages are homonomous and biramous, including three pairs in the cephalon and one associated with each tergite in the thorax. Endopodites are gracile with slender and rect-
angular distal podomeres. Exopodites bear long, thick lamellae on a single lobe. | |
| S. R. Losso et al. 2025 | Concilitergans in which the thorax has fused tergites and no pleural or pygidial spines. The interpleural furrows nearly completely effaced in some representatives. | |
| S. R. Losso et al. 2025 | A large helmetiid with prominent anterolateral cephalic spines. Six thoracic tergites with progressively larger posterior-facing pleural spines, and two pairs of posterolateral spines on pygidium. Dorsal exoskeleton with serrated margin throughout, except for rounded anterior sclerite. A pair of lateral eyes located underneath slightly domed exoskeletal bulges on cephalon, and two medial eyes underneath the anterior sclerite. 15
pairs of biramous postantennal appendages, three below cephalon, six below the thorax and five below the pygidium. | |
| S. R. Westrop and E. Landing 2025 | Palpebral lobes large but shorten somewhat during holaspid ontogeny from length (exsag.) equal to about 50% of glabellar length (e.g., Figs. 8b and 9a) to 41% of glabellar
length (e.g., Fig. 8a). Pygidium elongate, and lengthens (sag.) by posterior extension of marginal spines during holaspid ontogeny.Median embayment of posteriormarginwell defined in small specimens, becoming longer (sag.) during ontogeny, with posterior corners becoming increasingly spinelike. External surface of cranidium granulose, augmented with terrace ridges on frontal glabellar lobe and margins of L2–L4. External surface of pygidium with scattered coarse granules, most closely spaced on axis and adjacent part of pleural field, but become sparse on pygidial spines; subdued over entire surface on largest specimens. | |
| S. R. Westrop and E. Landing 2025 | Librigena with broad, strongly convex border; firmly impressed border furrow. Genal spine incompletely preserved, but likely long and robust. External surface of cranidium and librigena (aside from furrows) coarsely granulose; granules typically developed over background of minute punctate (usually visible only under high magnification). Paired fixigenal tubercles defined most clearly on smaller sclerites; additional small tubercles present on anterior border. Some granules of librigenal field arrayed along caecal markings. Pygidial outline roughly elliptical; anterior margin curved strongly backward along wide (tr.) articulating facet. Coarse granules of external surface match cephalic sculpture. | |
| S. R. Westrop and E. Landing 2025 | Occipital spine originates near posterior margin of LO. Palpebral ridge well defined on external surface and internal mould. Palpebral area broad, width (tr.) equal to 30% (28–32) of cranidial width across palpebral lobes. Finely granulose external surface of cranidium, with faint relict of fixigenal tubercle array evident on some specimens (e.g., Fig. 20f). Broad pygidium subtriangular in outline; axis with two rings in front of terminal piece composed of at least two segments. |
Measurements
No measurements are available
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| Source: g = genus, f = family, subo = suborder, o = order, c = class | |||||
| References: Kiessling 2004, Jell and Adrain 2002, Tillyard 1921, Aberhan et al. 2004, Hendy 2009, Vršanský 2003, Størmer 1955 | |||||
Collections (15188 total)
Oldest occurrences | |||
| Time interval | Ma | Country or state | Original ID and collection number |
|---|---|---|---|
| Begadean | USA (Nevada) | Fritzaspis generalis (90905) Fritzaspis generalis, Fritzaspis ovalis (90901) Profallotaspis sp., Fritzaspis sp., Fritzaspis generalis, Fritzaspis ovalis, Amplifallotaspis keni (90902) | |
| Placentian | Canada (Nova Scotia) | Trilobita indet. (86747 86748) | |
| Caerfai | Canada (Nunavut) | Trilobita indet., Paedumias sp., Bonniopsis sp. (184) | |
| Caerfai | India (Jammu and Kashmir) | Redlichia noetlingi, Redlichia knjazevi, Tungusella obesa (46443) | |
| Caerfai | USA (New York) | Atops sp., Rimouskia sp., Elliptocephala sp. (181) Olenellus indet. (8236) | |
| Caerfai | Canada (Northwest Territories) | Olenellus sequornalus, Olenellus sp., Wanneria sp. (193) Trilobita indet. (49080) | |
| Caerfai | Canada (New Brunswick) | Ellipsocephalus galeatus (34572 34597) Ellipsocephalus galeatus, Protolenus elegans (34571) Kingaspidoides obliquoculatus (34573) Protolenus articephalus, Protolenus elegans (34567 34751) Protolenus elegans (34569 34570 34595 34596 34752) | |
| Caerfai | USA (Nevada) | Olenellus sp., Bonnia sp. (190) | |
| Early/Lower Cambrian | Spain (Aragon) | Onaraspis altus, Alueva undulata, Hamatolenus ibericus, Kingaspis campbelli, Sdzuyia sanmamesi, Protolenus pisidianus, Protolenus jilocanus (120552) Onaraspis altus, Tonkinella sequei, Alueva undulata, Hamatolenus ibericus, Kingaspis campbelli, Sdzuyia sanmamesi, Protolenus pisidianus, Protolenus jilocanus (120550) | |
| Early/Lower Cambrian | USA (Washington) | Olenellidae indet., Nevadia addyensis (191233) | |
Youngest occurrences |
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| Time interval | Ma | Country or state | PBDB collection number |
| Holocene | USA (Montana) | Melanoplus spretus (185941) | |
| Holocene | Colombia (Santander) | Leptopharsa colombiana (196554) | |
| Holocene | Saint Helena | Labidura loveridgei (155782) | |
| Holocene | USA (Wyoming) | Melanoplus spretus, Spharagemon campestris (185942) | |
| Holocene | Australia (Queensland) | Austrodictya corbouldi (152282) | |
| Holocene | Madagascar | Aradus madagascariensis, Litus beneficus (151372) Evania animensis (224321) Microcaetus formidolosus (152187) Phatnoma madagascariensis, Cysteochila impressa, Cysteochila copal, Hovatlas invaginatus, Gitava fusca (183970) | |
| Holocene | Dominican Republic | Empicoris copal (138880) | |
| Holocene | Benin | Calotermes resinatus, Eutermes handlirschi (150373) Trigonidiinae indet. (123992) | |
| Holocene | USA (California) | Lethocerus americanus (161537) Notonecta badia, Ranatra bessomi, Ranatra asphalti, Belostoma bakeri (161518) | |
| Holocene | Tanzania | Apentacentrus copalicus, Agrabtoblatta symmetrica, Symploce rete (124004) Calotermes batheri, Calotermes erythrops, Termes amicus, Mirotermes resinatus, Microcerotermes latinotus (150372) Ceraphron serrulatus, Ceraphron fasciatus, Alaptus fructuosus (152189) Chaetoessa brevialata, Blatta elliptica, Blatta ruficeps, Poeocera venulosa, Ricania multinervis, Pentatoma schaurothi, Cercopis aurata, Chaetoessa burmeisteri (129861) Cicadidae indet., Cicadellidae indet., Cixiidae indet., Derbidae indet., Termes pusillus, Issidae indet., Typhlocybinae indet., Osaka sp. 2 (123418) Enicocephalus fulvescens, Cercopis strongii (165051) Fulgoroidea indet., Isoptera indet., Braconidae indet., Cixiidae indet., Aleyrodoidea indet., Mantodea indet., Psyllidae indet., Chalcidoidea indet., Issidae indet., Cheloninae indet., Anthocoridae indet. (234540) Oligotoma westwoodi (174113) Palmon bellator, Palmon clavellatus, Palmon capitellatus, Blatta persipicillata, Ricania equestris, Asiraca albipuncta (152160) Periplaneta sp., Mecostibus sp., Termes sp., Amusurgus africanus, Rhicnogryllus zeuneri, Issidae indet., Anaplecta sp., Pochazoides sp. 1, Pochazoides sp. 2, Pochazoides szwedoi, Phyllodromia sp., Acroprivesa msandarusi, Osaka sp. 1, Pocharica sp., Zoraida angolensis, Cicada forsythii, Ricaniidae indet., Phyllodromia woodwardi, Phyllodromia praecursor, Pseudophyllodromia succinica, Ceratinoptera miocenica, Euthyrrhapha pacifica, Ischnoptera provisionalis, Plectoptera antiqua (123991) Plutothrix minutissima (140791) Termes debilis (117251) Termes diaphanus, Termes punctatus (150374) | |