Corresponding author: Karla D. A. Soares (
Academic editor: Peter Bartsch
The genus
Contrasting hypotheses on the classification of catsharks are widespread in literature and divide opinions of many authors (e.g.
Although the paraphyly of
Doubts concerning the monophyly of the genus
Detailed descriptions of all
Thirty-five taxa were included as terminals in the phylogenetic analysis. Species representing the three genera assigned to the
List of species examined (except
Species examined | Data available | Origin of material |
---|---|---|
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||
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External morphology, dermal denticles, musculature, skeleton | AMNH, USNM |
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External morphology, dermal denticles, musculature, skeleton, clasper | SAIB |
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External morphology, dermal denticles, musculature, skeleton | USP |
|
External morphology, dermal denticles, musculature, skeleton | AMS |
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External morphology, dermal denticles, musculature, skeleton, clasper | SAIAB |
|
External morphology, dermal denticles, musculature, skeleton, clasper | SAIAB |
|
||
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External morphology, dermal denticles, musculature, skeleton, clasper | HUMZ |
|
External morphology, dermal denticles, musculature, skeleton, clasper | AMS |
|
External morphology, dermal denticles, musculature, skeleton, clasper | CSIRO, MZUSP |
|
External morphology, neurocranium, clasper | ZMH |
|
External morphology, dermal denticles, musculature, skeleton, clasper | USNM |
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External morphology, dermal denticles, musculature, skeleton, clasper | CSIRO, MZUSP |
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External morphology, dermal denticles, musculature, skeleton, clasper | UF |
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External morphology, dermal denticles, musculature, skeleton, clasper | SAIAB |
|
External morphology, dermal denticles, musculature, skeleton, clasper | AMNH, BMNH |
|
External morphology, dermal denticles, musculature, skeleton, clasper | SAIAB |
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External morphology, dermal denticles, musculature, skeleton, clasper | CAS |
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External morphology, musculature, skeleton, clasper | CAS |
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External morphology, dermal denticles, musculature, skeleton, clasper | UERJ, ZMH |
This study was based on the examination of 84 morphological characters (79 qualitative and five quantitative) that included external morphology, branchiomeric and hypobranchial cranial muscles, clasper morphology, dermal denticles and skeleton. External morphological characters were observed directly or with the aid of a stereomicroscope. Anatomical preparation was performed through manual dissections. For the examination of clasper anatomy, the left clasper was chosen to study the external morphology and the right clasper for the internal anatomy. Neurocrania and musculature of adult specimens were examined through dissection. Skin samples were taken for examination of dermal denticles from the right side of the body above the pectoral fin, below the origin of the first dorsal fin and below the insertion of the second dorsal fin. Dermal denticles were photographed using scanning electron microscopes (DSM 940 and ZEISS SIGMA VP), housed in the Departamento de Zoologia of the Universidade de São Paulo. Data of intestinal valves, tooth and vertebral counts were obtained directly from the examined specimens or taken from
Radiographs were taken in the
Terminology for neurocranium and jaws follows
Character descriptions, related to meristic data, are presented first, followed by characters of external morphology, myology, skeleton and clasper. Skeletal characters are grouped into character complexes, such as neurocranium, jaws, hyoid and gill arches and pectoral girdle. The number preceding each character in the description corresponds to its number presented in the character matrix. A brief summary of each character and its states is followed by its recovered consistency and retention indices (
Characters were illustrated with photographs and schematic drawings made from digital photographs. Photographs were taken with a digital camera (Canon Power Shot SX610 HS). Characters and their states are indicated by arrows and numbers in the figures. Figures were digitised and edited with the aid of Adobe Photoshop CS6. Whenever a character is described in the text for a genus without a species citation, that citation refers only to the species examined in the present study and does not imply that the character is present in all congeners.
Hypotheses of phylogenetic relationships were proposed using the cladistic method formalised by
The section ‘Description and character analysis’ presents the description of each character, its variation within the subfamily
1 Counts of monospondylous vertebrae: minimum = 28; maximum = 54. (
2 Counts of diplospondylous vertebrae: minimum = 67; maximum = 131. (
3 Upper tooth row counts: minimum = 33; maximum = 110. (
4 Lower tooth row counts: minimum = 29; maximum = 102. (
Tooth row counts are presented for many species in descriptions or taxonomic reviews, but never used as a phylogenetic character. Regarding the differences between upper and lower jaws, we considered upper and lower tooth row counts as distinct characters. In
5 Counts of intestinal valves: minimum = 5; maximum = 17. (
6 Extension of anterior nasal flap: (0) entirely covering excurrent nasal aperture and posterior nasal flap, but not covering the upper lip; (1) partially covering excurrent nasal aperture and not covering posterior nasal flap nor upper lip; (2) entirely covering excurrent aperture, posterior nasal flap and upper lip. (ordered;
The anterior nasal flap is a triangular or subrectangular structure and is situated medial to the incurrent aperture and lateral to the excurrent one. This flap can cover partially or entirely the excurrent aperture and posterior nasal flap, which is situated on the posterior border of the excurrent aperture (
Ventral view of the head. A,
7 Distance between anterior nasal flaps: (0) distant by one-half or more of the width of the flap; (1) distant by less than one-half. (
In relation to the distance between anterior nasal flaps, these are separated by one-half or more of the width of the flaps in
8 Configuration of anterior nasal flap: (0) flap consisting of a single structure; (1) flap separated into lateral and medial two portions. (
Most scyliorhinids present a single anterior nasal flap. In
9 Mesonarial crest: (0) inconspicuous; (1) prominent. (
The presence of a mesonarial crest was observed and described by
Nasoral region with lifted anterior nasal flap and exposed posterior nasal flap. A,
10 Muscular nasal barbel on anterior nasal flap: (0) absent; (1) present. (
The presence of a muscular nasal barbel is observed in
11 Posterior nasal flap: (0) present; (1) absent. (
A posterior nasal flap, associated with the excurrent nasal aperture, is present in all scyliorhinines, most scyliorhinids and
Nasoral region with lifted anterior nasal flap and exposed posterior nasal flap. A,
12 Degree of development of posterior nasal flap: (0) corresponding to one-half of the area of the anterior nasal flap; (1) reduced and only bordering the posterior tip of the excurrent aperture. (
Scyliorhinines present a well-developed posterior nasal flap, corresponding to one-half of the area of anterior nasal flap, as do the genera
13 Position of the posterior nasal flap: (0) situated on the posterior border of the excurrent aperture; (1) laterally situated to the excurrent aperture. (
In relation to the position of the posterior nasal flap,
14 Nasoral grooves: (0) absent; (1) present. (
A nasoral groove, which links the excurrent aperture and the mouth, is observed only in
15 Upper labial furrow: (0) present; (1) absent. (
An upper labial furrow is absent in
16 Lower labial furrow: (0) present; (1) absent. (
A lower labial furrow is present in
17 Projected flap on the upper lip margin: (0) absent; (1) present. (
In
18 Configuration of labial furrows: (0) continuous and fused laterally; (1) discontinuous and upper furrow ventral to the lower one. (
In taxa, where both labial furrows are found, there is a difference concerning their configuration. In
19 Number of upper labial cartilages: (0) two; (1) one. (
Labial cartilages. A, detail of labial cartilages in
20 Postoral groove: (0) absent; (1) present. (
In species of
21 Pelvic apron: (0) absent; (1) present. (
The fusion of the pelvic inner margins, known as the pelvic apron and defined by
22 Extension of pelvic apron: (0) fusion extending up to one-half the length of pelvic inner margins; (1) fusion extending up to two thirds of the length of pelvic inner margins; (2) pelvic inner margins almost entirely fused. (ordered;
Amongst the taxa presenting the pelvic apron, there is some variation in its extension. In
Pelvic apron. A, detail of pelvic apron in
23 Origin of the first dorsal fin: (0) closer to the vertical line that passes through the insertion of pelvic fins; (1) closer to the vertical line that passes through the origin of pelvic fins. (
The posteriormost origin of the first dorsal fin is the main character used to diagnose the family
24 Origin of second dorsal fin: (0) posterior to the vertical line that passes through half-length of anal fin base; (1) anterior to the vertical line that passes through half-length of anal fin base. (
Two conditions were observed concerning the origin of the second dorsal fin: posterior (most scyliorhinids; state 0, Fig.
Caudal region. A, detail of dorsal fins in
25 Cusplets of dermal denticles on dorsolateral body surface: (0) present; (1) absent. (
The crown of the dermal denticles on the dorsolateral surface of the body varies from ‘teardrop’ to ‘trident’ shape due to the presence or absence of cusplets lateral to the principal cusp of the crown. Cusplets are present in most scyliorhinids (state 0; Fig.
Dermal denticles above the origin of the first dorsal fin. A,
26 Extension of ectodermal pits in dorsal surface of the crown denticles: (0) extending through more than half the length of the crown; (1) restricted to anterior portion of the crown (
Ectodermal pits were observed and illustrated by
27 Median ridges on dermal denticles: (0) two ridges; (1) one ridge. (
28 Caudal crest of enlarged dermal denticles: (0) absent; (1) present. (
The presence of a caudal crest of dermal denticles distinct from the denticles on dorsolateral surfaces and situated on the upper lobe of caudal fin, is found in
Detail of the caudal crest in
29 Muscle
The postorbital musculature is composed of three muscles: m.
Detail of the postorbital musculature. A,
30 Insertion of the muscle
In most taxa examined, the m.
Detail of the insertion region of the muscle
31 Insertion of the muscle
In most taxa examined, the muscle
Hypobranchial musculature. A,
32 Configuration of muscles bundles of the m.
The presence of m.
33 Origin of the muscles
In
Detail of muscle
The insertion of the m.
34 Rostral cartilages: (0) fused; (1) united only by connective tissue. (
In scyliorhinines,
Neurocranium; dorsal view. A,
35 Relation between lateral rostral cartilages and anterior fontanelle: (0) rostral cartilages distant from anterior fontanelle; (1) rostral cartilages confluent with lateral borders of anterior fontanelle. (
The distance between lateral rostral cartilages may vary, positioned medially or laterally to the lateral borders of the anterior fontanelle. In some cases, the lateral rostral cartilages are confluent with the lateral borders of the anterior fontanelle, connected to it through ridges that extend from the base of the rostral cartilages to the border of the fontanelle; this condition was observed in
36 Relationship between median rostral cartilage and anterior fontanelle: (0) median rostral cartilage and anterior fontanelle separated by internasal space; (1) median rostral cartilage confluent with anterior fontanelle. (
The distance between the median rostral cartilage and anterior fontanelle varies amongst taxa examined. In scyliorhinines and other taxa examined, the median rostral cartilage and the anterior fontanelle are separated by the internasal space, distant by at least two thirds of the length of the median rostral cartilage (state 0; Fig.
Detail of rostral region and anterior fontanelle of
37 Orientation of nasal capsules: (0) nasal capsules perpendicular to the anteroposterior axis of the neurocranium; (1) nasal capsules oblique. (
In
Neurocranium; ventral view. A,
38 Relative position between nasal apertures: (0) incurrent aperture anterior to excurrent one; (1) nasal apertures at the same level. (
Nasal apertures may be positioned at the same level (
39 Fusion of the external nasal cartilage to the dorsal position of the nasal capsule: (0) present; (1) absent. (
The external nasal cartilage, situated anteriorly to the nasal apertures and ventrally to the nasal capsules, may or may not be fused to the anterodorsal portion of the nasal capsules (
40 Degree of development of the subnasal plate: (0) restricted to the medial portion of the nasal capsules and ventral to the internasal septum; (1) laterally expanded and united to the lateral border of the nasal capsule. (
The subnasal plate is the ventral floor of the nasal capsules, generally associated with a cavity posteromedial to the incurrent aperture and covered by a layer of connective tissue (nasal fontanelle of
41 Epiphyseal notch: (0) absent; (1) present. (
The anterior fontanelle, the anterodorsal aperture of the neurocranium covered by a layer of connective tissue, presents different shapes amongst species and also varies between sexes (
42 Supraorbital crest: (0) present; (1) absent. (
The occurrence of a supraorbital crest on the neurocranium is widely used for identification and separation of shark genera and families. The presence of this crest is considered primitive for elasmobranchs and its absence secondary in some sharks and rays (
43 Distance between internal carotid foramina: (0) greater than the distance between internal carotid and stapedial foramina; (1) smaller than the distance between internal carotid and stapedial foramina; (2) equal to the distance between internal carotid and stapedial foramina. (ordered;
Four foramina are present on the posterior portion of the basal plate: two for the medial internal carotid arteries and two for the lateral stapedial arteries.
44 Relative size of postorbital groove: (0) groove corresponds to more than one-half the height of the hyomandibular facet; (1) groove corresponds to less than one-half the height of the hyomandibular facet. (
The postorbital groove is situated posteriorly to the orbits and ventral to the postorbital processes, limited dorsally by the opisthotic process and ventrally by the hyomandibular facet; the lateral vein of the head passes along this groove (
Neurocranium; lateral view. A,
45 Fenestra for the infraorbital canal of the lateral line: (0) present; (1) absent. (
The pre- and postorbital processes are laterally expanded from the neurocranial roof, as wide as or wider than the nasal capsules. In most scyliorhinids, the distal tip of the postorbital process has a large fenestra through which passes the infraorbital canal of the lateral line (
46 Labial ridge of the quadrate process: (0) present; (1) absent. (
Detail of jaws; lighter portion of Meckel’s cartilage is less calcified portion. A,
47 Position of the orbital processes of the palatoquadrate: (0) at the anterior one-fourth of each antimere; (1) closer to the half-length of each antimere. (
The palatoquadrate articulates to the neurocranium by ethmopalatine ligaments, which are inserted on the postorbital processes of the palatoquadrates and originate from the orbital notches; these notches are situated between the posteroventral region of the nasal capsules and the preorbital wall. Orbital processes are situated in variable positions in the dorsal border of each antimere of the palatoquadrate, delimitating the extension of palatine and quadrate processes. In most scyliorhinids, orbital processes are situated at the anterior one-fourth (state 0; Fig.
48 Degree of calcification of the medial portion of Meckel’s cartilage: (0) similar calcification throughout; (1) medial portion less calcified than the rest of Meckel’s cartilage. (
Meckel’s cartilages present distinct degrees of calcification in some scyliorhinids. In
49 Articular region of the quadratomandibular joint of Meckel’s cartilage: (0) posterior lingual condyle situated between anterior labial condyle and facet; (1) anterior and posterior condyles forming a unit and distant from the facet; (2) posterior lingual condyle opposite to the facet. (ordered;
Detail of the articular region of the quadratomandibular joint of Meckel’s cartilage. A,
50 Thyroid foramen: (0) present; (1) absent. (
The basihyal cartilage, situated ventromedially to other components of the hyoid arch, is a structure that presents variable dimensions amongst taxa. This cartilage may or may not present an opening in its anterior portion, the thyroid foramen (
Detail of the basihyal cartilage. A,
51 Internal surface of the hyomandibular cartilage: (0) smooth; (1) concave. (
In
Detail of the hyomandibular cartilage; internal surface. A,
52 Anterior border of the basihyal cartilage: (0) not bifurcated; (1) bifurcated. (
The occurrence of a bifurcation on the anterior border of the basihyal cartilage, anterior to the thyroid foramen and not confluent with it, was observed in
53 Lateral processi rastriformis: (0) present; (1) absent. (
Processi rastriformis were observed and illustrated in
Detail of the lateral processi rastriformis (
54 Oropharyngeal denticles: (0) small and not forming rows on internal face of gill components; (1) large and forming rows on internal face of gill components. (
Detail of the oropharyngeal denticles in
55 Shape of the gill pickax: (0) elongated and sling-like; (1) short and triangular. (
The fusion between the dorsal tips of gill arches IV and V, forming a unique plate known as the gill pickax (
Detail of the gill pickax. A,
56 Ventral extrabranchial cartilages: (0) four; (1) three. (
Ventral extrabranchial cartilages are present amongst muscle bundles of the
57 Medial projection of the coracoid bar: (0) present; (1) absent. (
The presence of a medial projection on the coracoid bar was observed in many orders of elasmobranchs by
Coracoid bar; ventral view. A,
58 Degree of development of the medial projection of the coracoid bar: (0) reduced to less than twice the size of the lateral portion of the coracoid bar; (1) well developed, more than twice the size of the lateral portion of the coracoid bar. (
In taxa in which a medial projection of the coracoid bar is present, differences concerning its shape and degree of development were observed. In
59 Lateral processes on pectoral girdle: (0) present; (1) absent. (
Lateral processes on the coracoid bar, medial to the articular region between pectoral girdle and fins, were observed in
60 Dermal denticles on the dorsal surface of clasper glans: (0) present; (1) absent. (
Dermal denticles on the dorsal surface of clasper glans were observed in most taxa examined. In
61 Distribution of dermal denticles on dorsal surface of clasper glans: (0) denticles present only on the exorhipidion; (1) denticles over all of the dorsal surface except on rhipidion and terminal dermal cover. (
Clasper; external morphology. A,
62 Dermal denticles on the medial border of the exorhipidion: (0) absent; (1) present. (
63 Terminal dermal cover: (0) present; (1) absent. (
64 Extension of the terminal dermal cover: (0) restricted to the distal tip of the clasper glans; (1) extending up to one-third of clasper glans. (
Regarding its extension, the terminal dermal cover may be restricted to the distal clasper tip or it extends up to one-third of the clasper glans, covering the posterior borders of the cover rhipidion and exorhipidion. The former condition was observed in
65 Configuration of the terminal dermal cover: (0) smooth; (1) rough. (
66 Degree of development of the rhipidion: (0) well developed and presenting a prominent posterior margin; (1) reduced and consisting in a narrow strip. (
Some differences regarding the degree of development of the rhipidion were observed in taxa that have this structure (
Detail of the rhipidion. A,
67 Extension of the rhipidion: (0) extending throughout the clasper glans; (1) extending up to one-third of the clasper glans. (
The extension of the rhipidion varies, depending on the species examined. In
68 Cover rhipidion: (0) poorly developed; (1) well developed and medially expanded. (
69 Exorhipidion: (0) medially expanded; (1) poorly developed. (
The exorhipidion is a ventromedially situated flap, covering totally or partially the ventral terminal cartilage. In all species of
70 Envelope: (0) present; (1) absent. (
The envelope is a distinct projection anterior to the exorhipidion, which covers the accessory terminal cartilage, posterior border of the ventral marginal cartilage and anterior border of the ventral terminal cartilage. According to our observations, this structure is present in
71 Degree of development of the envelope: (0) poorly developed; (1) medially expanded. (
A well-developed envelope, projecting medially and covering the anterior border of the cover rhipidion is observed in
72 Accessory terminal cartilage: (0) present; (1) absent. (
73 Accessory dorsal marginal cartilage: (0) present; (1) absent. (
Clasper; skeleton. A,
74 Dorsal marginal 3 cartilage: (0) absent; (1) present. (
The dorsal marginal 3 cartilage is situated dorsally and external to the accessory dorsal marginal cartilage or posterior to it. A dorsal marginal 3 cartilage is absent in scyliorhinines and most of the scyliorhinids examined (state 0). In
75 Ventral marginal 2 cartilage: (0) present; (1) absent. (
The presence of a ventral marginal 2 cartilage was reported for
76 Position of the ventral marginal 2 cartilage: (0) continuous to the posterior border of the ventral marginal cartilage; (1) lateral to the posterior border of the ventral marginal cartilage. (
77 Dorsal terminal 2 cartilage: (0) present; (1) absent. (
The dorsal terminal 2 cartilage was described by
78 Shape of the dorsal terminal 2 cartilage: (0) elongated and rod-like; (1) rhomboidal. (
Variations in the shape of the dorsal terminal 2 cartilage were observed amongst the taxa examined. In
79 Ventral terminal 2 cartilage: (0) present; (1) absent. (
80 Position of the ventral terminal 2 cartilage: (0) anteriorly situated and sometimes attached to the anterior tip of the ventral terminal cartilage; (1) posteriorly situated, posterior to the half-length of the ventral terminal cartilage. (
A different condition from the one described by
81 Extension of the clasper siphon: (0) extending beyond the half distance between the coracoid and cloaca; (1) shorter than the coracoid-cloaca half distance. (
Detail of the clasper siphon. A,
82 Colour pattern composed of saddles: (0) present; (1) absent. (
The presence of transverse bands darker than the background colour over most of the body, known as ‘saddles’, is widespread amongst catsharks.
Color patterns. A,
83 Dark spots: (0) present; (1) absent. (
84 Dark stripes: (0) absent; (1) present. (
A colour pattern, composed of dark stripes running in different directions, was observed in
Anterior nasal flaps in
Muscle
The muscle
Muscle
Origin of the muscle
The m.
Clasper hooks
In
The phylogenetic analysis of the data matrix (Appendix
Character listings for clades numbered in Figure
Strict consensus cladogram of the three equally most-parsimonious trees (L = 233,
Monophyly of clade 1
The hypothesis of the monophyly of the
Lower diplospondylous vertebral count [char. 2, 0.520–0.660 > 0.280–0.310].
Muscle
Nasal apertures at the same level in nasal capsules [char. 38, 0 > 1].
Articular region of the quadratomandibular joint of Meckel’s cartilage composed by a posterior lingual condyle opposite to the facet [char. 49, 0 > 1].
Three ventral extrabranchial cartilages [char. 56, 0 > 1].
Terminal dermal cover extending up to one-third of the clasper glans [char. 64, 0 > 1].
Accessory dorsal marginal cartilage absent [char. 73, 0 > 1].
Clasper siphon short and restricted to the pelvic region [char. 81, 0 > 1].
Monophyly of clade 2
The monophyly of
Anterior nasal flap divided into two portions, medial and lateral [char. 8, 0 > 1].
Muscular nasal barbel present [char. 10, 0 > 1].
Accessory terminal cartilage absent [char. 72, 0 > 1].
Ventral terminal 2 cartilage posteriorly situated, posterior to the half-length of the ventral terminal cartilage [char. 80, 0 > 1].
Colour pattern not composed of transverse saddles [char. 82, 0 > 1].
Projected flap present on the upper lip margin [char. 17, 0 > 1]. Independently acquired in
Muscle
Colour pattern composed of stripes [char. 84, 0 > 1].
Dermal denticles present on the medial border of the exorhipidion [char. 62, 0 > 1].
Monophyly of clade 3
The monophyly of the clade formed by
Mesonarial crest prominent [char. 9, 0 > 1].
One upper labial cartilage [char. 19, 0 > 1].
Lateral processi rastriformis similar to dermal papillae in length [char. 53, 0 > 1].
Coracoid bar with a well-developed medial projection, more than twice the size of its lateral portion [char. 58, 0 > 1].
Rhipidion well-developed and presenting a prominent posterior margin [char. 66, 1 > 0].
Rhipidion extending throughout the clasper glans [char. 67, 1 > 0].
Ventral marginal 2 cartilage absent [char. 75, 0 > 1].
Monophyly of clade 4
The monophyly of
Higher monospondylous vertebral count [char. 1, 0.540–0.650 > 0.730].
Higher upper tooth row count [char. 3, 0.230–0.350 > 0.450–0.480].
Higher lower tooth row count [char. 4, 0.230–0.370 > 0.490–0.530].
Lower labial furrow absent [char. 16, 0 > 1].
Postoral grooves present [char. 20, 0 > 1].
Origin of second dorsal fin anterior to half-length of the anal-fin base [char. 24, 0 > 1].
Muscle bundles of muscle
Higher diplospondylous vertebral count [char. 2, 0.280–0.310 > 0.670–1.00].
Higher upper tooth row count [char. 3, 0.450–0.480 > 0.570–1.00].
Higher lower tooth row count [char. 4, 0.490–0.530 > 0.580–1.00].
Monophyly of clade 5
The clade, formed by species
Lower diplospondylous vertebral count [char. 2, 0.280–0.310 > 0.130–0.170].
Lower diplospondylous vertebral count [char. 2, 0.130–0.170 > 0.060–0.080].
Monophyly of clade 6
This clade is formed by
Oropharyngeal denticles large and forming rows on internal face of gill components [char. 54, 0 > 1].
No unique autapomorphies were found for
Higher monospondylous vertebral count [char. 1, 0.730 > 0.810].
Monophyly of clade 7
The monophyly of
Lower values for upper tooth row count [char. 3, [0.230–0.350 > 0.210–0.220].
Lower values for lower tooth row count [char. 4, 0.230–0.370 > 0.220].
Projected flap present on the upper lip margin [char. 17, 0 > 1].
Pelvic apron present [char. 21, 1 > 0].
Ephyseal notch present [char. 41, 0 > 1].
Monophyly of clade 8
This clade is formed by the species
Envelope present on clasper [char. 70, 1 > 0].
Envelope medially expanded [char. 71, 0 > 1].
No unique autapomorphies were found in the present analysis for
Colour pattern without dark spots [char. 83, 0 > 1].
is characterised by the following autapomorphy:
Colour pattern composed of dark stripes [char. 84, 0 > 1].
Monophyly of clade 9
The monophyly of the clade, formed by
Accessory terminal cartilage absent [char. 72, 0 > 1].
is hypothesised as the sister group of all remaining species of
Higher intestinal valve count [char. 5, 0.420 > 0.670–0.750].
In some trees:
Higher monospondylous vertebral count [char. 1, 0.460–0.540 > 0.580–0.690].
Monophyly of clade 10
This clade is formed by
Terminal dorsal 2 cartilage rhomboidal [char. 78, 0 > 1].
In some trees:
Lower monospondylous vertebral count [char. 1, 0.540–0.580 > 0.420–0.460].
Higher intestinal valve count [char. 5, 0.420 > 0.250].
Monophyly of clade 11
The monophyly of the clade formed by
Envelope present on clasper [char. 70, 1 > 0].
No unique autapomorphies were found in the present analysis for
Dermal denticles restricted to the exorhipidion on dorsal surface of clasper glans [char. 61, 1 > 0].
Monophyly of clade 12
The monophyly of the clade, formed by
Colour pattern without dark spots [char. 83, 0 > 1].
Higher upper tooth row count [char. 3, 0.280–0.310 > 0.470].
Ventral terminal 2 cartilage absent [char. 79, 0 > 1].
Higher monospondylous vertebral count [char. 1, 0.460 > 0.540–0.770].
Monophyly of clade 13
The monophyly of the clade, formed by
Pelvic inner margins almost entirely fused. [char. 22, 1 > 2].
Lower monospondylous vertebral count [char. 1, 0.350–0.540 > 0.080–0.270].
Lower upper tooth row count [char. 3, 0.170–0.220 > 0.00–0.120].
Lower values for lower tooth row count [char. 4, 0.190–0.220 > 0.010–0.180].
Envelope present on clasper [char. 70, 1 > 0].
Monophyly of clade 14
This clade is formed by
Accessory terminal cartilage present [char. 72, 1 > 0].
Higher monospondylous vertebral count [char. 1, 0.350–0.540 > 0.620–0.690].
Lower values for lower tooth row count [char. 4, 0.210–0.220 > 0.260–0.770].
Terminal dermal cover rough [char. 65, 0 > 1].
Dorsal terminal 2 cartilage rhomboidal [char. 78, 0 > 1].
Monophyly of clade 15
The monophyly of the clade, formed by
Rhipidion reduced and consisting of a narrow strip [char. 66, 0 > 1].
Rhipidion extending up to 1/3 of the clasper glans length [char. 67, 0 > 1].
Monophyly of clade 16
The monophyly of the clade, formed by
Anterior nasal flap entirely covering excurrent aperture, posterior nasal flap and upper lip [char. 6, 0 > 2].
Colour pattern composed of dark spots [char. 83, 1 > 0].
Higher monospondylous vertebral count [char. 1, 0.350–0.540 > 0.770].
Monophyly of clade 17
Monophyly of the clade, formed by
Anterior nasal flaps distant from each other by less than half of their length [char. 7, 0 > 1].
Posterior nasal flap laterally situated to the excurrent aperture [char. 13, 0 > 1].
Nasoral groove present [char. 14, 0 > 1].
Terminal dermal cover rough [char. 65, 0 > 1].
Lower intestinal valve counts [char. 5, 0.420 > 0.250].
Accessory terminal cartilage absent [char. 72, 0 > 1].
The phylogenetic relationships of species of the subfamily
The monophyly of
Species of
Clasper morphology contributed important characters that helped elucidate the phylogenetic relationships among species of
According to
Characters from the nasoral region, dermal denticles, claspers, vertebrae and intestinal counts were revealed to be extremely important to shed light on the phylogenetic relationships amongst scyliorhinines and may contribute to future phylogenetic analyses concerning scyliorhinids. A more detailed examination of the nasal flaps and labial furrows allows for the identification of differences amongst the genera
Data from tooth morphology of catsharks are scarce and the only study that reported tooth characters for
Despite the relevance of characters associated to claspers in species identification and phylogenetic analyses, information on the internal anatomy of these organs are found only for some species and mainly in classical works about clasper morphology (
The monophyly of the subfamily
Phylogenetic hypotheses of the subfamily
The monophyly of the genus
In the present study, we contribute to the understanding of the phylogenetic relationships amongst
Despite the contributions presented here for the phylogeny of
The monophyly of Scyliorhininae is supported by four characters proposed by
The monophyly of
Characters from the nasoral region, dermal denticles, claspers, vertebrae and intestinal counts were revealed to be extremely important to shed light on the phylogenetic relationships amongst scyliorhinines and may contribute to future phylogenetic analyses concerning scyliorhinids and carcharhiniforms.
Results presented here mostly agree with those obtained in recent phylogenetic analyses, but further work integrating molecular and morphological data is still needed.
The authors wish to acknowledge Barbara Brown (AMNH), David Catania (CAS), Rob Robbins (FLMNH), Toshio Kawai (HUMZ), Karsten Hartel and Andrew Williston (MCZ), Patrice Pruvost (MNHN), Marcelo Brito (MN/UFRJ), Alessio Datovo (MZUSP), Oliver Crimmen and Ralf Britz (BMNH), Sven Kullander (NRM), Masanori Nakae and Gento Shinohara (NSMT), Ofer Gon and Nkosinathi Mazungula (SAIAB), Albe Bosman (SAM), Ulisses Gomes and Hugo Santos (UERJ), Ricardo Rosa (UFPB), Otto Gadig (UNESP, São Vicente), José Ortiz (USAC), Lynne Parenti, Jeffrey Williams, Kris Murphy and Sandra Raredon (USNM), Peter Bartsch (ZMB), Ralf Thiel (ZMH), and Marcus Krag (ZMUC) for permission to visit the collections and examine the specimens under their care. José Ortiz (USAC) for sending photos of specimens of
List of examined material
Matrix summarising quantitative characters used in the phylogenetic study. 1, monospondylous vertebral counts; 2, diplospondylous vertebral counts; 3, upper tooth row counts; 4, lower tooth row counts; 5, intestinal valve counts. Absolute values and normalised ones (in parentheses) are given.
Terminal | 1 | 2 | 3 | 4 | 5 |
---|---|---|---|---|---|
|
38 | 118 | 47–62 | 49–59 | 10 |
(0.38) | (0.8) | (0.18–0.38) | (0.27–0.41) | (0.42) | |
|
30–33(0.08–0.19) | ? | 35–45(0.03–0.16) | 29–40(0–0.15) | 15–17(0.83–1) |
|
? | ? | 61–62(0.35–0.37) | 56–62(0.37–0.45) | ? |
|
39–46(0.42–0.69) | 110–115(0.67–0.75) | 56–73(0.30–0.52) | 50–59(0.29–0.41) | 11–13(0.5–0.67) |
|
45–46(0.65–0.69) | 103–109(0.56–0.65) | 50–70(0.22–0.48) | 45–59(0.22–0.41) | 16(0.92) |
|
45–48(0.65–0.77) | 71–72(0.06–0.08) | 50–70(0.22–0.48) | 45–65(0.22–0.49) | ? |
|
49(0.77) | 75–91(0.12–0.37) | 67–84(0.44–0.66) | 67–87(0.52–0.79) | 10(0.42) |
|
47–54(0.73–1) | 110–131(0.67–1) | 77–110(0.57–1.00) | 71–102(0.57–1.00) | ? |
|
44–47(0.61–0.73) | 72–77(0.08–0.16) | 68–82(0.45–0.63) | 68–82(0.53–0.73) | ? |
|
28–35(0–0.27) | 67–71(0–0.06) | 54–66(0.27–0.43) | 54–68(0.34–0.53) | 5–6(0–0.08) |
|
35–38(0.27–0.38) | 105–111(0.59–0.68) | 54–57(0.27–0.31) | 54–62(0.34–0.45) | ? |
|
33–39(0.19–0.42) | ? | 56(0.30) | 52(0.31) | 6–8(0.08–0.25) |
|
31–33(0.11–0.19) | 92–100(0.39–0.51) | 56(0.30) | 47(0.25) | ? |
|
33–40(0.19–0.46) | 88–100(0.32–0.51) | 66–75(0.43–0.54) | 58–72(0.40–0.59) | 6–7(0.08–0.17) |
|
28–33(0–0.19) | 78–103(0.17–0.56) | 60–70(0.35–0.48) | 62–67(0.45–0.52) | 7(0.17) |
|
38–39(0.38–0.42) | 71–108(0.06–0.64) | 67–71(0.44–0.49) | 78–82(0.67–0.73) | 7–8(0.17–0.25) |
|
42–45(0.54–0.65) | 73–92(0.07–0.39) | 45–55(0.16–0.29) | 42–49(0.18–0.27) | 9–13(0.33–0.67) |
|
32–46(0.15–0.69) | 70–78(0.05–0.17) | 45–51(0.16–0.23) | 40–46(0.15–0.23) | 8–13(0.25–0.67) |
|
35–40(0.27–0.46) | 80–129(0.20–0.96) | 53–65(0.26–0.42) | 41–56(0.16–0.37) | ? |
|
39–42(0.42–0.54) | 82–95(0.23–0.43) | 39–49(0.08–0.21) | 40–45(0.15–0.22) | ? |
|
37–39(0.35–0.42) | 83–85(0.25–0.28) | 45–58(0.16–0.32) | 44–50(0.20–0.29) | 6–8(0.08–0.25) |
|
35–40(0.27–0.46) | 83–95(0.25–0.43) | 40–61(0.09–0.36) | 36–50(0.09–0.29) | 10–11(0.42–0.5) |
|
44–46(0.61–0.69) | 78–88(0.17–0.32) | 46–76(0.17–0.56) | 48–85(0.26–0.77) | 10–11(0.42–0.5) |
|
40–45(0.46–0.65) | 80–91(0.20–0.37) | 44–58(0.14–0.32) | 42–52(0.18–0.31) | ? |
|
40(0.46) | 97(0.46) | 50(0.22) | 43(0.19) | ? |
|
35–37(0.27–0.35) | 83–88(0.25–0.32) | 42–48(0.04–0.19) | 36–44(0.09–0.20) | 8(0.25) |
|
48(0.77) | 83(0.25) | 46(0.17) | 44(0.20) | ? |
|
36–40(0.31–0.46) | 81–87(0.21–0.31) | 48–54(0.19–0.27) | 42–53(0.18–0.33) | 6–8(0.08–0.25) |
|
39–42(0.42–0.54) | 85–96(0.28–0.45) | 39–49(0.08–0.21) | 38–46(0.12–0.23) | ? |
|
46–48(0.69–0.77) | 84–90(0.26–0.35) | 46–52(0.17–0.25) | 43–50(0.19–0.29) | ? |
|
38–42(0.38–0.54) | 84–93(0.26–0.40) | 36–55(0.04–0.29) | 34–50(0.07–0.29) | 10–11(0.42–0.5) |
|
43–46(0.58–0.69) | 87–89(0.31–0.34) | 40–52(0.09–0.25) | 33–50(0.05–0.29) | 13–14(0.67–0.75) |
|
32–37(0.15–0.35) | 73–89(0.09–0.34) | 50–76(0.22–0.56) | 45–81(0.22–0.71) | 10–11(0.42 –0.5) |
|
30–35(0.08–0.27) | 81–83(0.21–0.25) | 33–42(0–0.04) | 31–42(0.03–0.18) | 6–8(0.08–0.25) |
|
38–39(0.38–0.42) | 81–96(0.21–0.45) | 47–56(0.18–0.30) | 45–53(0.22–0.33) | 6–8(0.08–0.25) |
Discrete morphological characters utilised in the phylogenetic analysis. Characters numbers correspond with those in the character descriptions (see text).
Terminal taxon | 6–10 | 11–20 | 21–30 | 31–40 | 41–50 | 51–60 | 61–70 | 71–80 | 81–84 |
---|---|---|---|---|---|---|---|---|---|
|
00000 | 0000000000 | 0?110??000 | 0000000001 | 0000010001 | 0010000011 | ?100011100 | ?00000001? | 1000 |
|
10000 | 1??0000000 | 0?00001000 | 1000101001 | 0001111121 | 1011000011 | ??1??1?010 | 0001000000 | 1110 |
|
00000 | 0000000000 | 1000000000 | 0000000000 | 0000000000 | 0000000010 | 0000000000 | 0000000000 | 0000 |
|
21000 | 1??1000000 | 0?00010000 | 0001000010 | 1120000000 | 0010000010 | 0000011101 | ?0011?0000 | 0000 |
|
00000 | 1??0000000 | 0?000?00?? | ???1000010 | 01210????? | ?????0???0 | 0000011101 | ?000011?01 | 0000 |
|
00010 | 0000110?11 | 0?01011010 | 0101000110 | 0110010010 | 001011010? | ?????????? | ?????????? | ?000 |
|
00010 | 0000110?11 | 0?01011010 | 0101000110 | 0110010010 | 0011110100 | 1001000101 | ?0101?001? | 1000 |
|
00010 | 0000110?11 | 0?01011010 | 0101000110 | 0110010010 | 001011010? | ?????????? | ????????00 | ?000 |
|
00010 | 0000110?11 | 0?01011010 | 0101000110 | 0110010010 | 001111010? | ?????????? | ?????????? | ?000 |
|
10100 | 0100000000 | 0?11111000 | 0110010000 | 0021001111 | ?010000111 | ??1??1?01? | ?0001?001? | ?110 |
|
10000 | 0100000000 | 0?0000?100 | 0000100000 | 000001?100 | 00?0000010 | 0101011000 | 00001?0000 | 0000 |
|
10000 | 0100000000 | 0?00001100 | 0000101001 | 0000001101 | 00100000?0 | 0101011001 | ?0101?1?00 | 1100 |
|
00000 | 0000000000 | 0?00010000 | 0110000000 | 1020000000 | 0101000001 | ?100000000 | 0001001?00 | 0000 |
|
21000 | 1??1000000 | 0?00010001 | 0010010000 | 0000001011 | 0010001?01 | ??00011011 | ?010000000 | 0000 |
|
10000 | 0100110??0 | 1000001000 | 1010010000 | 1010011010 | 0110000010 | 1001100011 | ?1011?0000 | 1100 |
|
00000 | 0100000100 | 0?1110?100 | 0000100000 | 0020001111 | 1011000011 | ??00011000 | 0000000000 | 0110 |
|
00101 | 0000101?00 | 0?0001?011 | 0001000110 | 0100010010 | 0000110000 | 0001011101 | ?110010001 | 11?1 |
|
00101 | 0000010000 | 0?0001?010 | 0001000110 | 0100010010 | 0000110000 | 0101011101 | ?110010001 | 1100 |
|
10110 | 0100000110 | 0?00111000 | 0010000110 | 1110110010 | 0100001?00 | 0001000100 | 0000000000 | 0000 |
|
00010 | 0000101?10 | 1100011010 | 0001000110 | 1110010010 | 0010010100 | 1001000101 | 10101?0000 | 1000 |
|
00010 | 0000101?10 | 1100011010 | 0001000110 | 1110010010 | 0010010100 | 1001111101 | ?1101?0100 | 1000 |
|
21010 | 0011101?10 | 1200011010 | 0001000110 | 1110010010 | 0010010100 | 1001100101 | ?0101?0000 | 1000 |
|
00010 | 0000101?10 | 1200011010 | 0001000110 | 1110010010 | 0010010100 | 0001000100 | ?0101?0100 | 1010 |
|
00010 | 0000101?10 | 1100011010 | 0001000110 | 1110010010 | 0010010100 | 1001000101 | 01101?0100 | 1000 |
|
00010 | 0000101?10 | 11000110?? | ?????0???? | ?????????? | ?????????0 | 1001000101 | ?1101?0000 | ?010 |
|
21010 | 0011101?10 | 1200011010 | 0001000110 | 1110010010 | 0010010100 | 1001011101 | ?1101?001? | 1000 |
|
20010 | 0000101?10 | ?000?10?? | ?????0???? | ?1???????? | ?????????? | ?????????? | ?????????? | ?000 |
|
00010 | 0000101?10 | 1100011010 | 0001000110 | 1110010010 | 0010010100 | 1001000100 | 01101?0100 | 1000 |
|
00010 | 0000101?10 | 1100011010 | 0001000110 | 1110010010 | 0010010100 | 00???00100 | 1????????? | ?010 |
|
00010 | 0000101?10 | 1100011010 | 0001000110 | 1110010010 | 0010010100 | 1001000101 | ?????????? | ?010 |
|
00010 | 0000101?10 | 1100011010 | 0001000110 | 1110010010 | 0010010100 | 0001000100 | 10101?0000 | 1001 |
|
00010 | 0000101?10 | 1100011010 | 0001000110 | 1110010010 | 0010010100 | 1001000101 | ?1101?0000 | 1000 |
|
00010 | 0000101?10 | 1200011010 | 0001000110 | 1110010010 | 0010010100 | 1001011101 | ?0101?0000 | 1010 |
|
00010 | 0000101?10 | 1200011010 | 0001000110 | 1110010010 | 0010010100 | 1001000100 | 01101?0000 | 1010 |
|
00010 | 0000101?10 | 1100011010 | 0001000110 | 1110010010 | 0010010100 | 1001000100 | 01101?0100 | 1000 |
List of non-ambiguous synapomorphies of clades and terminal taxa based on the three most-parsimonious cladograms obtained. Synapomorphies followed by “!” appear only in some trees.
Clade 1
Char. 2: 0.52–0.66 > 0.28–0.31.
Char. 29: 0 > 1.
Char. 38: 0 > 1.
Char. 56: 0 > 1.
Char. 59: 0 > 1.
Char. 64: 0 > 1.
Char. 73: 0 > 1.
Char. 81: 0 > 1.
Char. 8: 0 > 1.
Char. 10: 0 > 1.
Char. 72: 0 > 1.
Char. 80: 0 > 1.
Char. 82: 0 > 1.
Char. 9: 0 > 1.
Char. 19: 0 > 1.
Char. 53: 0 > 1.
Char. 58: 0 > 1.
Char. 66: 1 > 0.
Char. 67: 1 > 0.
Char. 75: 0 > 1.
Char. 1: 0.54–0.65 > 0.73.
Char. 3: 0.23–0.35 > 0.45–0.48.
Char. 4: 0.23–0.37 > 0.49–0.53.
Char. 16: 0 > 1.
Char. 20: 0 > 1.
Char. 24: 0 > 1.
Char. 32: 0 > 1.
Char. 2: 0.28–0.31 > 0.13–0.17.
Char. 54: 0 > 1.
Char. 3: 0.23–0.35 > 0.21–0.22.
Char. 4: 0.23–0.37 > 0.22.
Char. 17: 0 > 1.
Char. 21: 1 > 0.
Char. 41: 0 > 1.
Char. 70: 1 > 0.
Char. 71: 0 > 1.
Char. 72: 0 > 1.
Char. 1: 0.54–0.58 > 0.42–0.46!
Char. 5: 0.42 > 0.25!
Char. 78: 0 > 1.
Char. 70: 1 > 0.
Char. 83: 0 > 1.
Char. 22: 1 > 2.
Char. 72: 1 > 0.
Char. 66: 0 > 1.
Char. 67: 0 > 1.
Char. 6: 0 > 2.
Char. 83: 1 > 0.
Char. 7: 0 > 1
Char. 13: 0 > 1.
Char. 14: 0 > 1.
Char. 2: 0.13–0.17 > 0.06–0.08.
Char. 1: 0.73 > 0.81.
Char. 2: 0.28–0.31 > 0.67–1.00.
Char. 3: 0.45–0.48 > 0.57–1.00.
Char. 4: 0.49–0.53 > 0.58–1.00.
No autapomorphies.
Char. 17: 0 > 1.
Char. 30: 0 > 1.
Char. 84: 0 > 1.
Char. 62: 0 > 1.
No autapomorphies.
No autapomorphies.
Char. 65: 0 > 1.
Char. 1: 0.35–0.54 > 0.62–0.69.
Char. 4: 0.21–0.22 > 0.26–0.77.
Char. 65: 0 > 1.
Char. 78: 0 > 1.
Char. 61: 1 > 0.
Char. 3: 0.28–0.31 > 0.47.
Char. 79: 0 > 1.
Char. 5: 0.42 > 0.25.
Char. 72: 0 > 1.
Char. 1: 0.35–0.54 > 0.77.
No autapomorphies.
Char. 83: 0 > 1.
Char. 1: 0.46–0.54 > 0.69–0.77.
Char. 84: 0 > 1.
Char. 1: 0.46–0.54 > 0.58–0.69!
Char. 5: 0.42 > 0.67–0.75.
No autapomorphies.
Char. 1: 0.35–0.54 > 0.08–0.27.
Char. 3: 0.17–0.22 > 0.00–0.12.
Char. 4: 0.19–0.22 > 0.01–0.18.
Char. 5: 0.42 > 0.08–0.25.
Char. 70: 1 > 0.
No autapomorphies.
List of character transformation, based on the three most-parsimonious cladograms obtained.
Clade 4: 0.54–0.65 > 0.73.
Clade 7: 0.54–0.65 > 0.46–0.58.
Clade 8: 0.46–0.58 > 0.46–0.54.
Clade 10: 0.46–0.58 > 0.42–0.46.
Clade 13: 0.46–0.58 > 0.35–0.54.
Clade 17: 0.35–0.54 > 0.35–0.46.
Clade 1: 0.52–0.66 > 0.28–0.31.
Clade 2: 0.28–0.31 > 0.17–0.31.
Clade 5: 0.28–0.31 > 0.13–0.17.
Clade 13: 0.28–0.31 > 0.25–0.31.
Clade 1: 0.30–0.35 > 0.23–0.35.
Clade 2: 0.23–0.35 > 0.23–0.29.
Clade 4: 0.23–0.35 > 0.45–0.48.
Clade 7: 0.23–0.35 > 0.21–0.22.
Clade 8: 0.21–0.22 > 0.21.
Clade 13: 0.21–0.22 > 0.17–0.22.
Clade 16: 0.17–0.22 > 0.17–0.19.
Clade 1: 0.29–0.37 > 0.23–0.37.
Clade 2: 0.23–0.37 > 0.23–0.27.
Clade 7: 0.23–0.37 > 0.22.
Clade 4: 0.23–0.37 > 0.49–0.53.
Clade 6: 0.49–0.53 > 0.52–0.53.
Clade 12: 0.22–0.25 > 0.21.
Clade 14: 0.19–0.22 > 0.21–0.22.
Clade 16: 0.20–0.22 > 0.21.
Clade 10: 0.42 > 0.25.
Clade 12: 0.42 > 0.25–0.42.
Clade 14: 0.25–0.42 > 0.42.
Clade 16: 0 > 2.
Clade 17: 0 > 1.
Clade 2: 0 > 1.
Clade 3: 0 > 1.
Clade 2: 0 > 1.
No transformation in
No transformation in
Clade 17: 1 > 0.
Clade 17: 0 > 1.
Clade 1: 0 > 01.
Clade 3: 01 > 1.
Clade 4: 0 > 1.
Clade 7: 0 > 1.
Clade 1: 0 > 01.
Clade 3: 0 > 1.
Clade 4: 0 > 1.
Clade 7: 1 > 0.
Clade 7: 01 > 1.
Clade 13: 1 > 2.
No transformation in
Clade 4: 0 > 1.
No transformation in
No transformation in
Clade 1: 0 > 01.
Clade 3: 01 > 1.
No transformation in
Clade 1: 0 > 1.
No transformation in
Clade 4: 0 > 1.
No transformation in
No transformation in
No transformation in
No transformation in
No transformation in
Clade 1: 01 > 1.
No transformation in
Char. 40 (L = 2)
No transformation in
Clade 7: 01 > 1.
No transformation in
Clade 2: 01 > 0.
Clade 3: 01 > 1.
No transformation in
No transformation in
Clade 1: 01 > 1.
No transformation in
No transformation in
Clade 1: 0 > 1.
No transformation in
No transformation in
No transformation in
Clade 3: 0 > 1.
Clade 6: 0 > 1.
Clade 1: 0 > 01.
Clade 2: 01 > 1.
Clade 4: 01 > 1.
Clade 7: 01 > 0.
Clade 1: 0 > 1.
No transformation in
Clade 3: 0 > 1.
Clade 1: 01 > 1.
No transformation in
Clade 3: 0 > 01.
Clade 8: 01 > 0.
Clade 9: 01 > 1.
No transformation in
Clade 1: 0 > 1.
Clade 2: 01 > 1.
Clade 3: 01 > 0.
Clade 15: 0 > 1.
Clade 3: 01 > 0.
Clade 15: 0 > 1.
No transformation in
No transformation in
Clade 8: 1 > 0.
Clade 11: 1 > 0.
Clade 8: 0 > 1.
Clade 2: 0 > 1.
Clade 9: 0 > 1.
Clade 14: 1 > 0.
Clade 1: 0 > 1.
No transformation in
Clade 3: 0 > 1.
No transformation in
No transformation in
Clade 10: 0 > 1.
Clade 4: 0 > 01.
Clade 2: 0 > 1.
Clade 1: 0 > 1.
Clade 2: 0 > 1.
Clade 12: 0 > 1.
Clade 16: 1 > 0.