PHYLOGENETIC REVISION OF THE MICRO-ECHINOID GENUS, ECHINOCYAMUS
As A Thesis submitted to the faculty of % San Francisco State University In partial fulfillment of 2-OlS the requirements for bloL the Degree ■nyi Master of Science
In
Biology: Ecology and Systematic Biology
by
Kelly McNeal Markello
San Francisco, California
August 2015 Copyright by Kelly McNeal Markello 2015 CERTIFICATION OF APPROVAL
I certify that I have read Phylogenetic Revision o f the Micro-Echinoid Genus,
Echinocyamus by Kelly MeNeal Markello, and that in my opinion this work meets the criteria for approving a thesis submitted in partial fulfillment of the requirement for the degree Master of Science in Biology: Ecology and Systematic Biology at San Francisco
State University.
Rich Mooi, Ph.D. Curator, California Academy of Sciences
Peter Roopnarine, Ph.D. Curator, California Academy of Sciences
Karen Crow, Ph.D. Associate Professor of Biology PHYLOGENETIC REVISION OF THE MICRO-ECHINOID GENUS, ECHINOCYAMUS
Kelly McNeal Markello San Francisco, California 2015
Sea urchins (Echinoidea) comprise a group exhibiting enormous diversity in shape and size. At the lowest end of the size scale is Echinocyamus, a poorly known genus of highly miniaturized relatives of the sand dollars sometimes called “sea peas.” There are sixteen known extant species of Echinocyamus and two species in the sister genus,
Mortonia. Using elliptical Fourier analysis to compare these species, 13 of them with sufficiently large sample sizes (> ten specimens) were significantly distinct based on test shape alone. A morphological phylogeny indicates that two species, E. planissimus and
E. platytatus, should be removed not only from the genus, but from the family to which they belonged, Fibulariidae. With the removal of these two species, Echinocyamus and
Mortonia are monophyletic and sister to Fibularia. Mortonia was nested well within
Echinocyamus and should be sunk into that genus.
I certify that the Abstract is a correct representation of the content of this thesis.
Chair, Thesis Committee Date ACKNOWLEDGEMENTS
I would like to thank my thesis advisor, Rich Mooi, and my committee, Peter Roopnarine and Karen Crow, for their feedback on my thesis project. The California Academy of
Sciences provided many of the resources necessary for my project, including imaging equipment in the Project Lab and specimens for my work. I would also like to thank the
Museum of Comparative Zoology at Harvard University, the Smithsonian Institution’s
National Museum of Natural History, the University of California’s Museum of
Paleontology, and the Zoological Museum of the University of Copenhagen for loaning specimens for this project and allowing me to spend time visiting their collections. This project was supported by an Ernst Mayr Travel Grant in Animal Systematics from the
MCZ. Jon Fong’s assistance with X-raying specimens provided key evidence. Chrissy
Piotrowski, Liz Kools, and Bob Van Syoc also assisted me with lab support and specimen curation. Brian Simison and the Phylocluster provided computing resources to facilitate my tree searches. My labmate Kristen Vollrath inspired me to keep going. Finally, I would like to thank my husband Jay for his endless patience and invaluable support throughout my master’s program.
v TABLE OF CONTENTS
List of Tables...... vii
List of Figures...... viii
List of Appendices...... xiii
Introduction...... 1
Methods and Materials......
Morphometric Analysis...... 8
Phylogenetic Analysis...... 9
Character Descriptions...... 10
Results......
Morphometric Analysis...... 17
Phylogenetic Analysis...... 18
Taxonomy...... 22
Discussion...... 54
Works Cited...... 59
Appendices...... 65
Tables...... 66
Figures...... 72 LIST OF APPENDICES
Appendix Pi
1. Character matrix...... 6 LIST OF TABLES
Table Page
1. List of extant species with taxonomic authority...... 66 2. List of specimens photographed for morphometric analysis...... 67 3. Principal components that show a significant correlation with length...... 71
viii LIST OF FIGURES
Figure Page
1. Diagram of the basic anatomy of Echinocyamus...... 72 2. Left view of E. m egapetalus...... 73 3. Aboral, oral, and left view of Echinocyamus pusillus...... 74 4. X-ray of Echinocyamus pusillus...... 75 5. Aboral, oral, and left view of the holotype of Echinocyamus apicatus...... 76 6. X-ray of Echinocyamus apicatus...... 77 7. Aboral and oral view of Echinocyamus convergens...... 78 8. Aboral, oral, and left view of the holotype of Echinocyamus convergens...... 79 9. X-ray of Echinocyamus convergens...... 80 10. Aboral, oral, and left view of Echinocyamus crispus...... 81
11. X-ray of Echinocyamus crispus...... 82 12. Aboral, oral, and left view of Echinocyamus elegans...... 83 13. X-ray of Echinocyamus elegans...... 84 14. Aboral, oral, and left view of Echinocyamus grandiporus...... 85 15. Aboral, oral, and left view of a syntype of Echinocyamus grandiporus...... 86
16. X-ray of Echinocyamus grandiporus...... 87 17. Aboral, oral, and left view of the holotype of Echinocyamus incertus...... 88 18. X-ray of the holotype of Echinocyamus incertus...... 89 19. Aboral, oral, and left view of Echinocyamus macrostomus...... 90 20. Aboral, oral, and left view of a syntype of Echinocyamus macrostomus...... 91 21. X-ray of Echinocyamus macrostomus...... 92 22. Aboral, oral, and left view of Echinocyamus megapetalus...... 93 23. X-ray of Echinocyamus megapetalus...... 94 24. Aboral, oral, and left view of a syntype of Echinocyamus platytatus...... 95 25. X-ray of a syntype of Echinocyamus platytatus...... 96 ix LIST OF FIGURES (CONTINUED)
Figure Page
26. Aboral, oral, and left view of Echinocyamus provectus...... 97 27. X-ray of Echinocyamus provectus...... 98 28. Aboral, oral, and left view of Echinocyamus scaber...... 99 29. X-ray of Echinocyamus scaber...... 100 30. Aboral, oral, and left view of Echinocyamus sollers...... 101 31. Aboral, oral, and left view of a cotype of Echinocyamus sollers...... 102 32. X-ray of Echinocyamus sollers...... 103 33. Aboral, oral, and left view of Echinocyamus australis...... 104 34. X-ray of Echinocyamus australis...... 105 35. Aboral, oral, and left view of Echinocyamus polyporus...... 106 36. Aboral, oral, and left view of a type specimen of Echinocyamus polyporus...... 107 37. X-ray of Echinocyamus polyporus...... 108 38. Aboral, oral, and left view of the holotype of Echinocyamus planissimus...... 109 39. Aboral, oral, and left view of a paratype of Echinocyamus planissimus...... 110 40. X-ray of the holotype of Echinocyamus planissimus...... 111 41. Canonical Variate Analysis on combined dataset of lateral and aboral outlines - variate 1 vs. variate 2 ...... 112 42.. Canonical Variate Analysis on combined dataset of lateral and aboral outlines - variate 1 vs. variate 3 ...... 113 43. Principal component analysis of aboral view...... 114 44. Principal component analysis of lateral view...... 115 45. Linear regression of length versus first principal component from aboral view of E. australis...... 116 46. Linear regression of length versus second principal component from aboral view of E. crispus...... 117 x LIST OF FIGURES (CONTINUED)
Figure Page
47. Linear regression of length versus third principal component from aboral view of E. polyporus...... 118 48. Linear regression of length versus third principal component from aboral view of E. provectus...... 119 49. Linear regression of length versus third principal component from aboral view of E. pusillus...... 120 50. Linear regression of length versus second principal component from aboral view of E. scaber...... 121 51. Linear regression of length versus third principal component from aboral view of E. scaber...... 122 52. Linear regression of length versus first principal component from lateral view of E. crispus...... 123 53. Linear regression of length versus first principal component from lateral view of E. platytatus...... 123 54. Linear regression of length versus second principal component from lateral view of E. provectus...... 124 55. Linear regression of length versus third principal component from lateral view of E. pusillus...... 124 56. Linear regression of length versus second principal component from lateral view of E. scaber...... 125 57. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. australis...... 125 58. Linear regression of length versus first principal component from combined analysis of lateral and aboral views of E. crispus...... 126
xi LIST OF FIGURES (CONTINUED)
Figure Page
59. Linear regression of length versus third principal component from combined analysis of lateral and aboral views of E. elegans...... 126 60. Linear regression of length versus first principal component from combined analysis of lateral and aboral views of E. incertus...... 127 61. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. incertus...... 127 62. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. polyporus...... 128 63. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. provectus...... 128 64. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. p u sillu s...... 129 65. Strict consensus tree...... 130 66. Majority rule consensus tree...... 131 67. Character 1: location of maximum test heigh...... 132 68. Character 2: internal radial partitions...... 133 69. Character 3: circumferential partitions...... 134 70. Character 4: periproctal spines...... 135 71. Character 5: number of plates in periproctal membrane...... 136 72. Character 6: accessory podial pores along sutures...... 137 73. Character 7: ratio of test height versus length...... 138 74. Character 8: number of petaloid pore pairs...... 139 75. Character 9: coverage of test by petals...... 140 76. Character 10: ratio of test length to width...... 141 xii LIST OF FIGURES (CONTINUED)
Figure Page
77. Character 11: shape of petaloid pore pair series...... 142 78. Character 12: depth of infundibulum...... 143 79. Character 13: shape of peristome...... 144 80. Character 14: peristome size...... 145 81. Character 15: number of hydropores...... 146 82. Character 16: hydropore groove...... 147 83. Character 17: width of interporiferous zone (IPZ) relative to poriferous zone (PZ)...... 148 84. Character 18: glassy tubercles...... 149 85. Character 19: periproct shape...... 150 86. Character 20: relative size of ocular and genital pores...... 151 87. Character 21: inflation of ambital region...... 152 88. Biogeography of species in Clade A ...... 153 89. Biogeography of species in Clade B ...... 153 90. Biogeography of species in Clade C ...... 154 91. Biogeography of species in Clade D ...... 154
xiiii 1
Introduction
The “sea peas” constitute the genus Echinocyamus, a widespread group of diminutive
urchins, about which little is known. Sea peas can be found in large aggregations, as evidenced
by large specimen lots from single collecting events, but patchiness and small size make them
difficult to find in the field. Their ecological importance is not known, but likely to be
significant in sandy habitats as they chum up the substrate, consume organic material, and alter
the microbial biome. They provide a valuable trophic link between organic debris in the
substrate and their predators, such as fish, starfish, snails, and other urchins. Sea peas may also
serve as an important reservoir of calcium carbonate, as that substance is the primary
component of the tightly sutured plates that make up their internal test. Despite their ecological
potential, little is known about these tiny urchins.
Sea peas are echinoderms, along with about 7000 extant species of starfish, sea lilies,
sea cucumbers, brittle stars, and other sea urchins (Brusca and Brusca, 2003). As their name
suggests, many echinoderms are covered in spines and have benthic lifestyles in their
exclusively marine habitats. All echinoderms have an internal water vascular system that helps
maintain the tube feet (sometimes called podia) that are the main way by which echinoderms
interact with their environment. Tube feet are not only used for locomotion and feeding, but
also in respiratory gas exchange.
Most echinoderms show some degree of pentaradial symmetry in their adult form, while their larvae are bilateral. However, irregular urchins (formally known as the
“Irregularia”) are an exception to this rule. The Irregularia is a monophyletic group of urchins, which includes sand dollars, sea biscuits, lamp urchins, and heart urchins, that have evolved 2
bilateral symmetry in their adult form by moving their periproct (or anal region) relative to their
peristome (or mouth) (Smith and Kroh, 2011). “Regular” urchins have their periproct located at the center of the on top (apex) of the test, surrounded by gonopores (reproductive pores which are connected to their gonads and allow gametes to be released during spawning). The peristome of both irregular and “regular” urchins is located on the bottom of the organism, where it is closest to the substrate and probable food sources. The periproct of irregular urchins has shifted from the apex to the margin, or edge of the test. Sometimes the periproct has moved so far as to be only slightly posterior of the peristome (Mooi, 1990).
The Clypeasteroida is a clade of irregular urchins including sand dollars and sea biscuits. Their spines are extremely miniaturized and used primarily for locomotion (Durham,
1966). Most clypeasteroids have a fairly flattened, disk-like test, but the family Fibulariidae is an exception to this. Fibularia, Echinocyamus, and Mortonia are micro-echinoids (urchins that grow to less than 2 cm long) that range from an elongate bean shape to almost spherical. The genus Echinocyamus comprises approximately 15 extant species of micro-echinoids. Among the smallest of all sea urchins, they range in length from 2 to 10 mm and closely resemble juveniles of other, more disk-shaped clypeasteroids. However, micro-echinoids have long been known to represent fully mature but miniaturized taxa, because of the presence of gonopores in adult specimens. Sea peas are generally off-white when live, turning green when distressed.
They have a rounded, slightly flattened bean shape, hence the name “sea pea.” The presumed sister genus of Echinocyamus, Mortonia, is also a group of micro-echinoids containing two species with lengths from 6 to 18 mm. 3
Little is known of the natural history of sea peas, as only the type species,
Echinocyamus pusillus, has been studied in detail (Ghiold, 1982, Telford and Mooi, 1983).
Telford and Mooi (1983) found that E. pusillus prefers exposed areas with strong currents in waters generally 10-20 m deep, with shelly sediment. It is likely that they prefer this habitat because of higher aeration through the substrate (Telford and Mooi, 1983). Sea peas could be largely nocturnal, as they have been observed to be more active at night (Telford and Mooi,
1983). E. pusillus does not use food grooves (long, shallow indentations in the surface of the body that lead to the mouth) for feeding like other clypeasteroids, but rather moves individual particles with specialized, sucker-tipped tube feet towards the peristome. Once food particles reach the vicinity of the mouth, E. pusillus uses specialized, circum-oral spines and the lip-like membrane in which the mouth is situated to manipulate sediment into the esophagus. Sea urchins and sand dollars possess a five-part jaw apparatus known as the “Aristotle’s lantern.”
This apparatus contains five sharp teeth used to scrape organic material from ingested particles
(Telford and Mooi, 1983). This feeding method is unique to micro-echinoids such as
Echinocyamus, as most other echinoids use their lantern to grind sand particles and ingest both organic and inorganic debris. Predators of sea peas are likely to be fish, starfish, snails, and large urchins, as some specimens of Echinocyamus incertus have been found in the gut of a heart urchin, Scrippsechinus fisheri (Allison, Durham, and Mintz, 1967). Their planktonic larvae may serve as an important food source, but little is known about the volume or timing of spawning. There are no reports of parasites or symbionts of sea peas.
Sea peas have a global distribution, with species found off every continent except
Antarctica (Mortensen, 1948). The majority of species are found in the tropical Indo-Pacific, 4
with a few exceptions. E. pusillus is found from the North Sea to the Mediterranean, while E. grandiporus and E. macrostomus are found in the Caribbean. E. insularis is found in the
southeast Pacific, near Easter Island and the Nazca Ridge (Mironov and Sagaidachny, 1984). E. provectus and E. platytatus are known from southeast Australia and Tasmania, while the related form, Mortoniapolyporus is found only in New Zealand and the Kermadec Islands (see table 1 for more detailed distributions).
The bathymetric range of sea peas is quite large, but it can be difficult to discern if some localities are viable habitat for sea peas or simply a death assemblage, in which their tests accumulate at greater offshore depth because of prevailing currents. Specimens with spines still attached can be assumed to be either collected live or recently dead, but localities from the taxonomic literature or in museum databases do not always describe the condition of specimens collected, or if spines were removed after collection. With these limitations in mind, it can be reported that E. macrostomus was collected live at 2500 m, with dead specimens found at 3100 m (Mironov and Sagaidachny, 1984). E. crispus has been collected live from 1800 m (Mironov and Sagaidachny, 1984). E. grandiporus has been collected at 2500 m, but might not have been alive (MCZ collection). Knowledge of the depth ranges of most sea peas is imprecise due to a lack of collection data, so it is quite likely that other species have deep ranges exceeding the presently known limits. Several species are also found in quite shallow depths, such as E. pusillus mentioned earlier. E. crispus has been collected at 3 m, but it is not clear if the specimen was live (USNM collections).
The name Echinocyamus had a taxonomically challenging origin. Van Phelsum (1774) first described Echinocyamus from 14 forms from the Adriatic Sea and America, but he did not 5
assign binomial names to any of these “species.” Leske (1778) later assigned names to these
forms, but O.F. Muller (1776) was the first to use the name Spataguspusillus for the common
sea pea from the northeastern Atlantic now known as Echinocyamuspusillus. Most of Leske’s
species were reassigned to other genera, and the only valid Echinocyamus species were found to
be synonymous with Spatagus pusillus, which was transferred to Echinocyamus by Agassiz
(1841). Agassiz established the use of Echinocyamus for flat, bean-shaped forms and Fibularia
for globose forms, but Lambert challenged this in 1891, when he claimed that van Phelsum’s
figure depicts Echinocyamus as globular forms (Lambert, 1891). However, the only extant
species present in Europe, where van Phelsum collected his specimens, is Echinocyamus pusillus, a flattened, bean-shaped form, so it was deemed likely that van Phelsum’s figures were
simply inaccurate (Mortensen, 1948). This became a central point behind arguments to the
International Commission on Zoological Nomenclature (ICZN) in an attempt to stabilize the
nomenclature. Subsequently, in 1948 the ICZN ruled that Echinocyamus pusillus is the type species of Echinocyamus. Fibularia ovulum, an almost spherical micro-echinoid, is the type
species of Fibularia.
Mortensen (1948:171) reported on the “nomenclatural crime” by Lambert (1891), by which the fossil species of Echinocyamus and Fibularia remained in taxonomic confusion.
Lambert, based on figures in van Phelsum (1774), argued that the commonly accepted usage of
Echinocyamus for low, bean-shaped tests and Fibularia for high, globular tests should be reversed. Several other paleontologists followed suit, and with the range of intermediate and variable shapes, it remains quite difficult to determine which species are true Echinocyamus without close examination of internal structures or suture lines (Mortensen, 1948). The ICZN 6
(1948) ruled in favor of keeping the original usage, in which Echinocyamus consists of flattened
forms with well-developed internal buttresses (sometimes known as “partitions”). In contract,
Fibularia contains forms that are generally more high-bodied and completely lack internal
buttresses. However, most species described after Lambert’s (1891) reversal of these concepts
but before 1948 are still in need of revision. With this uncertainty in mind, the fossil record
suggests that Echinocyamus first appeared in the Eocene, as several species, reliably ascribed to
Echinocyamus, have been described from this time period (Mortensen, 1948). Due to the added
taxonomic difficulty of determining to which genus the fossil species belong, fossil specimens
were largely excluded from the morphometric analysis performed herein. A few well-described
fossil species were included in the parsimony analysis using characters from the taxonomic
literature.
As a result of this complicated taxonomic history, the validity of several extant
Echinocyamus and Mortonia species is still unknown. This paper seeks to resolve these issues
and to explore the phylogeny of this understudied group of urchins characterized by an unusual
morphology. Sea peas can provide a vital glimpse into how evolution produces miniaturized
forms and the possible role of heterochrony, changes in the timing of development that lead to
morphological differences. The vast majority of Echinocyamus specimens are denuded,
bleached skeletons with little tissue remaining, preventing employment of molecular methods to establish species boundaries or phylogenetic relationships within Echinocyamus. Despite this
limitation, this investigation sought to meet the following goals using data collected through
morphological and morphometric methods: 7
1. Determine validity of Echinocyamus and Mortonia species using elliptical Fourier
analysis.
2. Explore phylogenetic relationships within Echinocyamus and with the genera Mortonia
and Fibularia.
3. Determine any morphological synapomorphies that support monophyly of
Echinocyamus.
4. Determine the necessity of the clade Echinocyamidae to contain Echinocyamus and
related forms versus retaining Echinocyamus and Mortonia together in the Fibulariidae.
5. Explore biogeographic patterns of speciation in Echinocyamus and Mortonia using
phylogenetic relationships. 8
Methods and Materials
Morphometric Analyses
Specimens examined during this study were from the collections of the Department of
Invertebrate Zoology & Geology at the California Academy of Sciences (CASIZ), the Museum
of Comparative Zoology at Harvard (MCZ), the Invertebrate Zoology department of the
National Museum of Natural History (USNM), University of California Museum of
Paleontology (UCMP), and Zoological Museum - University of Copenhagen (ZMUC). Over
400 specimens were photographed using either the automontage or the “Big Kahuna” imaging
systems, depending on the size of the specimen. Specimens larger than 1 cm were
photographed on the “Big Kahuna” which was a Canon E05D with 100 mm lens and 1,4x
extender, mounted on a motor-driven stand. Lightroom and ZereneStacker v. 1.04 were used to
import the images and make Z-stacked images, respectively. Specimens were photographed in
aboral, oral, and left side views. Some specimens were too broken or fragile to use for all three
views (see Table 2 for which specimens were included in each analysis). Fewer than 10
specimens were used for E. apicatus, E. sollers, E. convergens, E. planissimus, and M polyporus, due to the scarcity of material. X-rays were also taken of each species at CAS.
Instead of the traditional landmark analysis, elliptical Fourier analysis was chosen because it
doesn’t require homologous landmarks, which are scarce on sea peas. For the Fourier analysis,
specimens were outlined in Photoshop, then converted to chain coordinates using the program
SHAPE ver. 1.3 (Iwata and Ukai, 2002). Elliptical Fourier analyses were run in PAST ver.
2.17c (Hammer, Harper, and Ryan, 2001). Canonical variates analysis (CVA) was used to
detect significant variances between species, and principal components analysis (PCA) was 9
used to visualize the shape changes accounting for the largest variance. Both CVA and PCA
were run in PAST. Allometric relationships in the PCA results were assessed using linear
regression of principal component scores versus length.
Phylogenetic Analysis
For the phylogenetic analysis, 18 ingroup taxa were included. All of the species of
Echinocyamus in Table 1 were included, with the exception of E. grandis. No specimens of this
species were examined, and the taxonomic descriptions given in the literature were not detailed
enough to derive the necessary characters. Mortensen (1948) suggested that this species is
probably a large E. crispus, but this cannot be confirmed without examining the type material.
Two fossil species, E. parviporus Kier, 1964 and E. petalus Kier, 1964, were included in the
ingroup as they were described after Lambert’s taxonomic swap was resolved. Only one
species of Mortonia, M. australis, was included because morphometric analyses could not differentiate M. australis from M. polyporus. Ten outgroup taxa were chosen based on a recent phylogeny of the putative sister genus Fibularia and related forms (Mooi et al., 2014).
The character matrix (Appendix 1) included 21 characters, 12 of which are binary and nine are multistate, for a total of 30 different character transitions (theoretical minimum tree length). Descriptions of each character are provided below. Phylogenetically uninformative characters were excluded, and question marks represent missing data. Characters were left unordered for the analysis. The matrix was compiled in Mesquite ver. 3.02 (Maddison and
Maddison, 2015). Parsimony analysis was run in PAUP* ver. 4 (Swofford, 2003) using the heuristic search algorithm. One thousand replicates were run with random tree addition and tree bisection and reconnection to rearrange trees. Bootstrap analyses were run using PAUP* on the 10
Phylocluster at CAS. One thousand bootstrap replicates were run, with 10 random additions per replicate. To reduce computing time, the bootstrap analysis was restricted to 1 million rearrangements per addition-sequence replicate. TreeRot ver. 3 (Sorensen and Franzosa, 2007) and FigTree ver. 1.4.2 (Rambaut, 2014) were used to calculate Bremer support values.
Character descriptions and character state codings
Character 1 - Location of maximum test height
Looking at the test laterally, most clypeasteroids reach their maximum height at the apex, which is usually located at the midpoint along the anterior-posterior axis of the test. However, the test of some Echinocyamus is inflated so that the point of maximum test height is shifted either posteriorly (towards the periproct) or anteriorly. In the outgroups, the position of greatest test height is variable, but tends to be central.
0 = point of maximum height central; 1 = point of maximum height anterior; 2 = point of maximum height posterior
Character 2 - Internal radial partitions
Most clypeasteroids have a complex system of internal supports arranged in pillars and buttresses. The latter are sometimes called “partitions,” or walls. Radial partitions extend interiorly from the margin of the test and end before reaching the peristome. Fibulariids have very minimal internal structures, entirely lacking pillars (Mooi, 1990). Fibularia lacks both pillars and partitions. Echinocyamus and Mortonia have 5 interambulacral pairs of partitions and a pair of partitions in the posterior interambulacrum only, respectively.
0 = ten radial walls; 1 = two radial walls; 2 = no radial walls 11
Character 3 - Circumferential partitions
In addition to five radial partitions, clypeasteroids can have internal “circumferential partitions” that branch orthogonally from the radiating walls, parallel to the ambitus. Fibulariids lack this elaboration of the internal supports.
0 = circumferential partitions present; 1 = circumferential partitions absent
Character 4 - Periproctal spines
Spines are sometimes present on the plates embedded in the periproctal membrane surrounding the anal opening. The presence or absence of periproctal spines can be difficult to determine in fossil specimens or denuded tests, as periproctal plates are quite fragile and quickly dissociate from the test after death. Most fibulariids lack these spines.
0 = periproctal spines present; 1 = periproctal spines absent
Character 5 - Number of plates in periproctal membrane
Most clypeasteroids have many (>20) plates in their periproctal membrane. However, fibulariids generally only have 5-20 plates. This character cannot be determined for many fossil taxa due to the usual loss of the periproctal membrane in fossil material.
0 = more than twenty plates in periproctal membrane; 1 = less than twenty plates in the periproctal membrane
Character 6 - Accessory podial pores along plate sutures
Podial pores connect the tube feet to the internal water vascular system. Echinocyamus tends to have these pores along the horizontal border between plates, forming distinctive lines of pores 12
along the ambitus (see figure 2). Fibularia and other clypeasteroids have podial pores scattered
throughout the plates.
0 = podial pores not concentrated along plate sutures; 1 = podial pores concentrated along plate
sutures
Character 7 - Ratio of test height versus length
The ratio of test height to length expresses how flat the test of any given species is, determining
the range of shapes within each species. “Flat” is defined as the height being less than 35% of the length of the specimen. “High” is defined as 35-50%, while “globular” is defined as over
50%.
0 = flat; 1 = high; 2 = globular
Character 8 - Number of petaloid pore pairs
Petaloid pores connect the respiratory tube feet to the internal water vascular system. These
pores form the five-petaled “flower” shape with two columns of paired pores in each of the 5
petaloids. Most clypeasteroids have hundreds of pore pairs in their petaloids, but fibulariids tend to have much fewer, with some species having only a few dozen. “Few” pore pairs is defined as < 50 total pore pairs in mature specimens. “Moderate” pore pairs is defined as 50 to
100 total pore pairs, while “many” pore pairs is defined as over 100 total pore pairs.
0 = many pore pairs; 1 = moderate pore pairs; 2 = few pore pairs 13
Character 9 - Coverage of test by petals
This character describes what percentage of the aboral side of the test is covered by the petaloid
pore pair regions. This character was estimated by measuring the length of the petals to
determine how much of the test, including the area between petaloids, was covered and
comparing it to the total area of the test. Some fibulariids have relatively small petals compared
to their test, while others have petals that reach all the way to the margin. Low coverage is
defined as less than 40%, while moderate coverage is defined as 40-70% of the test. High
coverage is over 70% of the test is covered by the petals.
0 = high coverage of test by petaloids; 1 = moderate coverage of test by petaloids; 2 = low
coverage of test by petaloids
Character 10 - Ratio of test length to width
This character describes the elongation of the test by dividing the width of the specimen by its
length and expressing that as a percentage. “Subcircular” is defined as the width being greater
than 85% of the length, while “elongate” is defined as the width being less than 85% of the
length.
0 = subcircular; 1 = elongate
Character 11 - Shape of petaloid pore pair series
Each petal begins at the apex of the test, with an ocular pore at the adapical end of each
petaloid. A petal typically consists of two series of pore pairs that are fairly close together, adapically. As the pores spread out from the apex, the series tend to diverge slightly. As the 14
pore series continue, they may either converge, stay parallel to one another, or continue
diverging from each other to form a “V” shape.
0 = convergent; 1 = parallel; 2 = divergent
Character 12 - Depth of infundibulum
The infundibulum is a concavity on the oral side of the test, centered around the peristome but
sometimes including the periproct. Fibulariids have infundibula of varying depths, and some
clypeasteroids have no infundibulum at all.
0 = slight infundibulum; 1 = no infundibulum; 2 = deep infundibulum
Character 13 - Shape of peristome
Most clypeasteroids have an almost round peristome, with the length roughly equal to the width and no angles to the curvature of the peristome’s perimeter. A few fibulariids and certain other clypeasteroids have a rounded, pentagonal peristome, with five “corners” at the end of each
interambulacral.
0 = round peristome; 1 = rounded pentagonal peristome
Character 14 - Peristome size
Most clypeasteroids have relatively small peristomes compared to the area of the oral surface.
However, some species of fibulariids have comparatively large peristomes. Small peristomes are defined as less than 15% of test length, medium peristomes are 15-25% of test length, and
large peristomes are greater than 25%.
0 = small peristome; 1 = medium peristome; 2 = large peristome 15
Character 15 - Number of hydropores
Most clypeasteroids have dozens or even hundreds of hydropores in the madreporic plate.
Some fibulariids, however, tend to have only one pore, while Fibularia itself seems to have more than one hydropore in most cases.
0 = more than 10 hydropores; 1 =2-10 hydropores; 2 = one hydropore
Character 16 - Hydropore groove
Some clypeasteroids such as Laganum retrict their hydropores to the bottom of a groove.
Echinocyamus lacks this groove.
0 = groove present; 1 = groove absent
Character 17 - Width of interporiferous zone relative to poriferous zone
The poriferous zone refers to the space occupied by the pore pair columns in a single petaloid, while the interporiferous zone is the region between the two pore pair columns (see fig.l).
Some fibulariids have interporiferous zones that are significantly wider than the poriferous zone, while others have interporiferous zones that are equal in width or slightly narrower than the poriferous zones.
0 = interporiferous zone wider than poriferous zone; 1 = interporiferous zone equal in width or narrower than poriferous zone
Character 18 - Glassy tubercles
Glassy tubercles are extremely glossy bumps scattered over the test that do not support spines or pediceltariae. Some Echinocyamus species, such as E. scaber, have higher, denser glassy 16
tubercles than other species. Fibularia kieri is not known to have glassy tubercles, but nearly all clypeasteroids have at least a few.
0 = smaller glassy tubercles; 1 = larger glassy tubercles; 2 = no glassy tubercles
Character 19 - Periproct shape
Most clypeasteroids have a circular or slightly transverse periproct. However, a few species have a periproct that is elongated along the anterior-posterior axis.
0 = circular periproct; 1 = anterior-posteriorly elongate periproct
Character 20 - Relative size of ocular and genital pores
In most clypeasteroids, ocular pores are so small as to be invisible except under high magnification, while genital pores (gonopores) are much larger. However, in a few species of
Echinocyamus the ocular pores are the same size as the gonopores.
0 = ocular pores small; 1 = ocular pores as large as genital pores
Character 21 - Inflation of ambital region
Most clypeasteroids are thickest at the apex and gradually become thinner towards the margin of the test. However, a few species have an inflated ambital region in which the test increases in thickness.
0 = inflation present; 1 = inflation absent 17
Results
Morphometric Analysis
A pairwise comparison of MANOVA results (using both lateral and aboral outlines) showed significant differences between most species tested. Three species could not be distinguished from any other species - E. planissimus, E. sollers, and M. polyporus. All three species had 10 or fewer representative specimens, so it is more likely that the test for significance failed because of small sample size rather than morphological similarity. E. scaber could not be differentiated from E. incertus and E. grandiporus. E. incertus is easily differentiated from E. scaber based on the relative size of gonopores. E. scaber and E. grandiporus are morphologically similar to one another, but E. scaber is found in the
IndoPacific while E. grandiporus is a Caribbean species.
Results from CVA on the combined dataset of both aboral and lateral outlines can be seen in figures 41-42. Some species, such as E. megapetalus and M. australis, are easily distinguished based on the first two variates, while other species, such as E. crispus and E. macrostomus, show less overlap in figure 42 than figure 41. The classifier function, which removed a priori assumptions about species identity, was able to identify each specimen’s species designation with 80% accuracy.
The first two components of the PCA of the aboral view (figure 43) account for 95% of the variance, with the first component accounting for 56% and the second component 39%. The first two components of the PCA of the lateral view (figure 44) account for 94% of the variance, with the first component accounting for 82% and the second component 12%. Species with significant allometric correlations in their PCA scores are listed in table 3 and figures 44-64. 18
Phylogenetic Analysis
A heuristic search in PAUP* resulted in 1,379 equally parsimonious trees with a minimum length of 85 steps. The consistency index was 0.352, the retention index was 0.565, and the rescaled consistency index was 0.199. The strict consensus tree (fig. 65) shows little resolution, with Laganum laganum, Sismondia occitana, Peronella peronii, and Echinocyamus planissimus falling outside a large, mostly unresolved polytomy containing the rest of the taxa.
Three small clades composed of Lenicyamidia compta + Cyamidia nummulitica, E. grandiporus
+ E. macrostomus, and E. crispus + E. megapetalus + M. australis were recovered in all of the equally parsimonious trees.
The majority rule consensus tree (figure 66) shows Fibularia and Fibulariidae
(excluding Echinocyamus, sensu Kroh and Smith, 2010) as monophyletic clades.
Echinocyamus can only be considered monophyletic if E. planissimus and E. platytatus are excluded from the genus, and if Mortonia australis is moved back within Echinocyamus.
Fibulariidae, including Fibularia and Echinocyamus, can be considered monophyletic if E. planissimus and E. platytatus are excluded. Four clades within Echinocyamus were recovered in greater than 50% of the most parsimonious trees. Clade A was recovered in 58% of trees with
E. pusillus and E. apicatus. Clade B, E. crispus, E. megapetalus, and M. australis, were recovered in 100% of the trees. Clade C, E. insularis, E. incertus, E. convergens, and E. provectus, were recovered in 65% of the trees, and 100% of trees recovered Clade D, E. sollers,
E. scaber, E. grandiporus, and E. macrostomus.
Each character was traced on the majority consensus tree in figures 67-87. For cases in which the ancestral state was ambiguous, slight preference was given for ACCTRAN over 19
DELTRAN, but it depended on the individual character state. The ancestral character states for the genus Echinocyamus are based on the most likely state at the node between E. parviporus and the rest of the ingroup Echinocyamus and Mortonia (excluding E. planissimus and E. platytatus). Based on the majority rule consensus tree and character matrix, the maximum test height of the ancestor of Echinocyamus was central, with several later clades developing either an anterior or posterior position. Ten radial internal partitions, with no circumferential partitions, were present. The circular periproct supported no spines and had few plates in the periproctal membrane. Podial pores were primarily along plate sutures along the ambitus, with no ambital inflation, and the test was high but not globular as in most Fibularia. The petals were moderately long with 50-100 pore pairs in two parallel series and covered 40-70% of the test. The interporiferous zone was wider than the poriferous zone of the petals. The test was likely subcircular, with length almost equal to width. The small, round peristome was probably in a slight infundibulum. The apical system contained ocular pores smaller than the genital pores, with one hydropore and no hydropore groove. Small glassy tubercles were present.
These characters are consistent with what is known of the fossil species E. parviporus. No synapomorphy for Echinocyamus was found.
Clade A, which consists of E. pusillus and E. apicatus, is fairly smilar to the ancestral
Echinocyamus character states. Instead of a central position, the maximum height of the test has shifted anteriorly. Also, E. pusillus has petals with divergent pore series rather than parallel pore series.
Clade B, which consists of E. crispus, E. megapetalus, and M. australis, has a few interesting divergences from the ancestral state. This is the only ingroup clade with a unique 20
synapomorphy - only species in Clade B have a deep infundibulum. Species in Clade B are
also elongate, not subcircular, and have a rounded pentagonal peristome instead of a round
peristome. E. megapetalus and Mortonia australis differ from E. crispus in that they both have
larger petals with more pore pairs. E. megapetalus is unique in the clade for having divergent,
instead of parallel, petaloids. Mortonia australis has several unique traits within the clade: anterior maximum height, only two radial partitions, periproctal spines, larger numbers of periproctal plates, and a small peristome relative to the total test size.
Clade C consists of E. insularis, E. incertus, E. convergens, and E. provectus. It is not clear what the ancestral position of the maximum test height is for this clade, as the ancestral
Echinocyamus likely had a central maximum height, but E. insularis (which branches off first within this clade) has an anterior maximum height while the others have a posterior maximum height. All of the species within Clade C are elongate as in Clade B. E. insularis has several other unique traits within the clade, such as periproctal spines and a small peristome. All of the species in Clade C except E. provectus fewer than 50 pore pairs, the interporiferous zone is equal to or narrower than the poriferous zone, and the peristome is round. E. convergens shows reduced coverage of its test by the pore pairs to less than 40%. E. convergens and E. provectus developed converging pore series in their petals, and their glassy tubercles are larger than their primary tubercles.
Clade D contains Echinocyamus sollers, Echinocyamus scaber, Echinocyamus grandiporus, and E. macrostomus. At the base of this clade, ocular pores evolved to be similar in size or larger than genital pores. However, this character was lost in E. macrostomus, which has the same state as the ancestral Echinocyamus with small ocular pores. With the exception 21
of E. sollers, Clade D species have large glassy tubercles and less than 50 pore pairs in each petal. The interporiferous zone is narrower or similar in size to the poriferous zone in E. sollers and E. scaber, but it reversed to the ancestral state in E. macrostomus and E. grandiporus. E. macrostomus has a larger peristome relative to its total test size.
E. planissimus and E. platytatus were both placed outside the clade containing the rest of Echinocyamus. These species have a flattened test, rather than the bean-shaped test of other
Echinocyamus, and they have more than one hydropore. E. platytatus also possesses a reduced number of petal pore pairs with fewer than 50. E. planissimus differs from the ancestral
Echinocyamus in several other characters. Circumferential partitions are present, which places
E. planissimus closer to the out-outgroup laganids. The podial pore pairs of E. planissimus are not found along the sutures between plates, its petaloids are convergent, the test is elongate, its peristome is pentagonal, and the ambitus is inflated. Considering all these differences, it is not surprising that E. planissimus was found not to be a true Echinocyamus. However, E. platytatus is less obviously outside the genus Echinocyamus in terms of the character distributions used in the analysis, but it is much flatter than most other Echinocyamus, and does show some vestiges of circumferential internal buttressing. 22
Taxonomy
Class Echinoidea Leske, 1778
Subclass Euechinoidea Bronn, 1860
Infraclass Acroechinoidea Smith, 1981
Irregularia Latreille, 1825
Microstomata Smith, 1984
Neognathostomata Smith, 1981
Order Clypeasteroida L. Agassiz, 1835
Infraorder Laganiformes Desor, 1857
Family Fibulariidae Gray, 1855
Small echinoids with reduced, simple, internal supports with no branching or circumferential partitions. Internal supports entirely lacking in some taxa.
Genus Echinocyamus van Phelsum, 1774
Type-species. Spataguspusillus O.F. Muller, 1776, by ICZN designation (Opinion 207).
Diagnosis: Accessory podial pores in lines along horizontal sutures of plates on aboral side of test. Usually only one hydropore, with one exception (E. platytatus). Internal supports consist of radial walls, either five partitions or two posterior partitions. 23
Echinocyamus pusillus (O.F. Muller, 1776)
Figures 3, 4.
1776. Spatagus pusillus O.F. Muller, p. 236.
1791. Echinus pusillus (O.F. Muller). Gmelin, p. 3198.
1791. Echinus minutus Gmelin, p. 3194.
1812. Echinuspulvinulus Pennant, p. 140.
1816. Fibularia tarentina Lamarck, p. 17.
1828. Clypeasterpulvinulus (Pennant). Van Den Ende, p. 301
1830. Echinocyamus minutus (Gmelin). Blainville, p. 195.
1847. Echinocyamus tarentinus (Lamarck). L. Agassiz & Desor, p. 140.
1850. Fibularia equina Aradas, p. 15.
1850. Echinocyamus minimus Girard, p. 367.
1856. Echinocyamus angulosus Llitken, p. 5.
1867. Echinocyamus speciosus Costa, p. 4.
1912. Fibulariapusilla (O.F. Muller). Lambert, p. 55, PI. IV.
Type material. None known.
Material studied. CASIZ 111220, Scotland, CASIZ 104183, Azores, CASIZ 111440
Bay of Biscay, France, CASIZ 198063, France. 24
Type locality. Adriatic Sea.
Description.
Size and shape - High intraspecific variation in shape. Mostly round (width >85% of total length) but some elongate with width only 30% of length. Test high, 30-47% of length.
Maximum height towards anterior. Some specimens with slight depression on posterior aboral side behind apex. Size range 2.7-15.0 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores.
Ambulacra - Petals with 8-18 pore pairs each; pores diverging slightly; petaloid region covering approximately 50% of test. Interporiferous zones wider than poriferous zones.
Glassy tubercles - Moderate size and scattered among the spine and pedicellarial tubercles, more strongly expressed in size and number on aboral side.
Peristome - Moderate size, about 20% test length; circular to subcircular. Infundibulum shallow.
Periproct - A few (but always more than 4) large plates on periproctal membrane; no spines on plates.
Spines - Primary spines about 0.5-0.7 mm, tapering, serrate. Miliary spines 0.2-0.3 mm long, with crown-shaped tip.
Pedicellaria - Ophicephalous pedicellariae with narrow, elongate blade densely serrate along outer edge, stalk narrower in middle than distally or proximally at hinge. Valves held 25
together with small stereom loops that prevent disarticulation. Tridentate pedicellariae about
0.08-0.09 mm, blade elongate and serrate, with larger serrations at tip. Triphyllous pedicellariae about 0.04 mm, stalk narrower in middle than at hinge or distal ends, blade with narrow, pin like teeth, proximal part of blade involute to form short, narrow tube.
Geographic range - From Norwegian Sea in the north, south to Northeast Atlantic
Ocean and Azores, east to the Marmara region in the Mediterranean Sea.
Bathymetric range - Littoral to 1250 m.
Remarks. E. pusillus is a highly variable species and might represent several cryptic taxa. However, with its complex taxonomic history, assigning valid names based on prior names that have been ascribed to this species will pose challenges. Future molecular studies are needed to determine if the intraspecific variability warrants splitting E. pusillus into several different taxa
Echinocyamus apicatus Mortensen, 1948
Fig. 5, 6.
1948b. Echinocyamus apicatus Mortensen, p. 5.
Type material. Holotype: MCZ 8051, paratype: ZMUC-ECH-452
Material studied. MCZ 8051 (holotype), MCZ 8095, ZMUC-ECH-452 (paratype), all from off Port Hacking, New South Wales 26
Type locality. Off Port Hacking, 2 miles from Jibbon Strait, New South Wales, 55 fathoms.
Description.
Size and shape - Subcircular test with width 87-94% of length. Apex conspicuous, slightly anterior. Anterior side arched, gradually sloping up toward apex; posterior end flattened, dropping off more sharply posterior to apex. Test high, 40-46% test length. Size range
3.9-6.2 mm.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores.
Ambulacra-Petals with 10-12 pore pairs each; pore series parallel; petaloid region covering approximately 60% of test. Interporiferous zones wider than poriferous zones.
Glassy tubercles - Moderate size and scattered among the spine and pedicellarial tubercles, more strongly expressed in size and number on aboral side.
Peristome - Moderate size, about 20% test length; subcircular. Infundibulum very shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Pedicellaria- Ophicephalous pedicellariae present.
Geographic range - New South Wales, off of Wattamolla Beach north to Broken Bay. 27
Bathymetric range - 70-100 m.
Remarks. E. apicatus is easily recognized by its steep-sided vertex and subcircular to circular outline.
Echinocyamus convergens Mortensen, 1948
Fig. 7-9.
1948b. Echinocyamus convergens Mortensen, p. 6.
Type material. Holotype: ZMUC-ECH-110.
Material studied. ZMUC-ECH-110 (holotype), Seychelles, USNM E36101,
Mozambique Channel
Type locality. Providence Island of the Seychelles, 128 m.
Description.
Size and shape - Elongate test with width 70-80% test length. Test high, 35-45% test length. Maximum height towards posterior. Size range 2.9-9.5 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores.
Ambulacra-Petals with 6-10 pore pairs each; pore series converging; petaloid region covering approximately 35% of test. Interporiferous zones width equal to that of poriferous zones. 28
Glassy tubercles - Larger than spine-bearing primary tubercles.
Peristome - Moderate size, about 20% test length; circular to subcircular. Infundibulum shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Geographic range - Western Indian Ocean, from the Seychelles down to Durban Bay in
South Africa.
Bathymetric range - 130-350 m.
Remarks. Petals are fairly short, and pore series converge more distinctly in this species than in other Echinocyamus. Fairly shallow infundibulum and converging pore series distinguish this species from E. grandis.
Echinocyamus crispus Mazzetti, 1893
Fig. 10, 11.
1893. Echinocyamus crispus Mazzetti, p. 239.
1914. Echinocyamus elongatus H.L. Clark, p. 62.
1914. Fibularia crispa (Mazzetti); Lambert & Thiery, p. 292.
Type material. Unknown.
Material studied. CASIZ 173810, Hawaii. 29
Type locality. Red Sea.
Description.
Size and shape -Elongate test, with width 67-81% test length. Test high, 33-47% test length. Maximum height central. Size range 1.8-10.5 mm long.
Internal buttressing-Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores.
Ambulacra-Petals with 6-20 pore pairs each; pore series parallel; petaloid region covering approximately 70% of test. Interporiferous zones wider than poriferous zones.
Glassy tubercles - Moderate size.
Peristome - Moderate size, about 20% test length; rounded pentagonal shape.
Infundibulum quite deep.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Pedicellaria - Ophicephalous pedicellariae present.
Geographic range - Widespread in Indo-Pacific, from Red Sea and Natal Coast to
Japan, Hawaii, and Guam.
Bathymetric range - Littoral to 1880 m.
Remarks. E. crispus is by far the most widespread and commonly found Indo-Pacific species of Echinocyamus. It’s easily distinguished from other Indo-Pacific species by its deep 30
infundibulum and comparatively short petals with respect to E. megapetalus, which has a deep infundibulum and extensive petals. Despite having many juvenile specimens, it is difficult to determine a specific size at which this species matures (as defined by the appearance of gonopores), as some are mature at 2.4 mm, while juveniles as large as 5.1 mm have been found.
E. crispus may be highly variable in when it matures, or this may indicate a cryptic species complex.
Echinocyamus elegans Mazzetti, 1893
Fig. 12, 13.
1893. Echinocyamus elegans Mazzetti, p. 240.
1925. Fibularia elegans (Mazzetti); Lambert & Thiery, p. 577.
Type material. Unknown.
Material studied. MCZ 8724, USNM E l0681, both from Arabian Sea.
Type locality. Red Sea
Description.
Size and shape - Subcircular test, sometimes rounded pentangular, with width 81-94% test length. Test high, bordering on globular, 37-56% test length. Maximum height towards anterior. Size range 2.1-6.0 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum. 31
Apical system - Single hydropore. Ocular pores smaller than gonopores.
Ambulacra - Petals with 6-20 pore pairs each; pore series parallel; petaloid region covering approximately 90% of test. Interporiferous zones width equal to width of poriferous zones.
Glassy tubercles - Moderate size.
Peristome - Moderate size, about 20% test length; round shape. Infundibulum shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Geographic range - Red Sea east to India and Arabian Sea. Possibly as far south as
Natal Coast, but identification is questionable.
Bathymetric range - 50-275 m.
Remarks. E. elegans is recognizable by its extensive petals which cover 90% of the test, and small, subcircular shape. E. megapetalus also has petals which cover most of the test, but it is generally larger and more elongate. 32
Echinocyamus grandiporus Mortensen, 1907
Fig. 14-16.
1907. Echinocyamus grandiporus Mortensen, p 33, PI. XII.
1914. Fibularia grandipora (Mortensen); Lambert & Thiery, p. 292.
Type material. Syntypes ZMUC-ECH-143, 447, 450, and 451, possible syntype ZMUC-ECH-
452
Material studied. ZMUC-ECH-143, Florida, ZMUC-ECH-447, Azores, ZMUC-ECH-
450, Florida, ZMUC-ECH-451, St. Croix, ZMUC-ECH-452, Cuba, USNM 10623, Mexico,
USNM 1014103, Brazil, USNM E l4552, St. Lucia
Type locality. Caribbean Sea and Azores.
Description.
Size and shape -Subcircular test, with width 81-94% test length. Test high, 38-48% test length. Maximum height central. Size range 2.1-9.5 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores same size as gonopores.
Ambulacra - Petals with 4-8 pore pairs each; pore series parallel; petaloid region covering approximately 30% of test. Interporiferous zones width greater than width of poriferous zones.
Glassy tubercles - Larger than spine-bearing primary tubercles. 33
Peristome - Moderate size, about 20% test length; round shape. Infundibulum shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Spines - Largest primary spines about 1 mm, slender, few serrations. Miliary spines about 0.3 mm, slender, and slightly serrated distally.
Pedicellaria - Ophicephalous pedicellariae with few serrations. Tridentate pedicellariae narrow gradually. Triphyllous pedicellariae with broad blade, finely serrated, head 0.04 mm.
Geographic range - In the West Atlantic and Caribbean from off of South Carolina south to Bahia. Also found in Azores and the Josephine Seamount in the East Atlantic.
Bathymetric range - 30-2500 m.
Remarks. E. grandiporus is easily distinguished from E. macrostomus, the only other
Caribbean species, by its ocular pores which are as large as its gonopores. E. grandiporus also lacks the large peristome of E. macrostomus.
Echinocyamus grandis H.L. Clark, 1925
1925. Echinocyamus grandis H.L. Clark, p. 165, PI. IX.
Type material. Location of holotype not known.
Material studied. None.
Type locality. Seychelles. 34
Description.
Size and shape - Elongate test, with width 79% test length. Test high, 36% test length.
Maximum height towards anterior. Size of single specimen reported as 14 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores.
Ambulacra - Petals with 22-28 pore pairs each; pore series parallel; petaloid region covering approximately 60% of test.
Glassy tubercles - Moderate size.
Peristome - Moderate size, about 20% test length; round shape. Infundibulum very deep.
Periproct - Unknown, periproctal membrane not present on specimen.
Geographic range - Seychelle Islands.
Bathymetric range - 62 m.
Remarks. Mortensen (1948a) believed this specimen was likely a large E. crispus, due to its deep infundibulum. The largest E. crispus found in the present study was 10.5 mm, so 14 mm would be quite a size extension, but not impossible. However, with the poor quality of photos in Clark (1925) and without seeing the holotype, it is difficult to determine if this species is actually a synonym of E. crispus. 35
Echinocyamus incertus H.L. Clark, 1914.
Fig. 17, 18.
1914. Echinocyamus incertus H.L. Clark, p. 64, PI. 128.
1925. Fibularia incerta (H.L. Clark), Lambert & Thiery, p. 577.
Type material. Holotype: USNM 34208.
Material studied. USNM 34208 (holotype), Hawaii, CASIZ 198061, Hawaii, UCMP
36579, 36580, 36581, 36582, Nazca Ridge.
Type locality. Hawaii.
Description.
Size and shape -Elongate test, with width 70-80% test length. Test high, 30-50% test length. Maximum height toward posterior. Size range 1.9-9.0 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores.
Ambulacra - Petals with 6-10 pore pairs each; pore series parallel; petaloid region covering approximately 45% of test. Interporiferous zones width equal to width of poriferous zones.
Glassy tubercles - Moderate size.
Peristome - Moderate size, about 20% test length; round shape. Infundibulum shallow. 36
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Geographic range - Hawaii. Potentially Nazca Ridge off west coast of South America.
Bathymetric range - 20-360 m.
Remarks. E. incertus is the most difficult Echinocyamus to describe, as it is a most unremarkable species. It has a shallow infundibulum, differentiating it from E. crispus. Fairly short petals and an elongate test demarcate it from E. megapetalus, E. (Mortonia) australis, E.
(Mortonia) polyporus, E. apicatus, and E. elegans. Its ocular pores are much smaller than its gonopores, and its glassy tubercles are not prominent, which eliminates E. scaber, E. sollers and
E. provectus. It is not flat enough to be confused for E. platytatus or E. planissimus. Thus, despite having no unique traits, this species is distinct enough from other Echinocyamus to be considered a valid species. However, this can make identification difficult to confirm. Several specimens of E. incertus were collected along the Nazca Ridge (UCMP specimens, Allison et al., 1967), but the specimens are too small to be confident about their identification.
Echinocyamus insularis Mironov & Sagaidachny, 1984
1984. Echinocyamus insularis Mironov & Sagaidachny, p. 192-193.
Type material. Holotype: IOANSSR N XV-65-19 at the Academy of Sciences of the USSR,
Institute of Oceanography, Moscow, Russia.
Material studied. None. 37
Type locality. Easter Island.
Description.
Size and shape - Elongate test, with width 60-80% test length. Test high, 30-35% test length. Maximum height towards anterior. Size range 1.0-4.8 mm long
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores.
Ambulacra - Petals with 8-10 pore pairs each; pore series parallel; petaloid region covering approximately 50% of test. Interporiferous zones width equal to width of poriferous zones.
Glassy tubercles - Moderate size.
Peristome - Small, about 15% test length; round shape. Infundibulum shallow.
Periproct - Spines present, few plates on periproctal membrane.
Pedicellaria - Ophicephalous and triphyllous present.
Geographic range - Southeast Pacific, near Easter Island and Isla Sala y Gomez
Bathymetric range - 50-80 m.
Remarks. Without having specimens at hand, it is hard to differentiate this species from other Echinocyamus. Mironov and Sagaidachny (1984) differentiated this species based on the distance between its posterior gonopores and its small size at maturity, which can have 38
high intraspecific variability in Echinocyamus. However, it is the only species known to be found in the eastern Pacific, except possibly for E. incertus.
Echinocyamus macrostomus Mortensen, 1907.
Fig. 19-21.
1907. Echinocyamus macrostomus, Mortensen, p.36, PI. XII.
1914. Fibularia macrostoma (Mortensen), Lambert & Thiery, p. 292.
1984. Echinocyamus scaber macrostomus Mortensen. Mironov & Sagaidachny, p. ###.
Type material. Syntypes: ZMUC-ECH-214 and 453.
Material studied. ZMUC-ECH-214 and 453, probably Azores?, MCZ 4589, Madeira
Islands, MCZ 7802, Cuba, USNM E14554, Bahamas, USNM E14557, Cuba
Type locality. Azores and Cape Verde Islands
Description.
Size and shape - Subcircular test, with width 80-92% test length. Test high, 37-44% test length. Maximum height central. Size range 4.0-8.8 mm long.
Internal buttressing-Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores. 39
Ambulacra - Petals with 2-6 pore pairs each; pore series parallel; petaloid region covering approximately 30% of test. Interporiferous zones width greater than width of poriferous zones.
Glassy tubercles - Larger than spine-bearing primary tubercles.
Peristome - Large, about 30% test length; round shape. Infundibulum shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Pedicellaria - Ophicephalous, tridentate, and triphyllous pedicellaria similar to those of
E. grandiporus.
Geographic range - Caribbean Sea, from Florida south to Bahia. Eastern Atest lengthantic near the Azores and Jospehina seamount.
Bathymetric range - 180-3000 m.
Remarks. E. macrostomus is easily differentiated from E. grandiporus by its large ocular pores and large peristome. Mortensen (1948a) reports this species as being more abyssal than E. grandiporus, but their bathymetric ranges overlap a fair bit, with E. grandiporus being found as deep as 2500 m. 40
Echinocyamus megapetalus H.L. Clark, 1914
Fig. 2, 22, 23.
1914. Echinocyamus megapetalus, H.L. Clark, p. 60, PI. 126.
1925. Fibularia megapetala (H.L. Clark), Lambert & Thiery, p. 577.
Type material. Holotype: MCZ 4747, paratypes: MCZ 2294.
Material studied. MCZ 2294 (paratypes), Mauritius, USNM E08491, Marshall Islands
Type locality. Mauritius, potentially, but might be misreported.
Description.
Size and shape -Elongate test, with width 69-81% test length. Test high, 38-46% test length. Maximum height central. Size range 3.9-7.9 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore, sometimes distorted to resemble groove with two hydropores. Ocular pores smaller than gonopores.
Ambulacra - Petals with 12-30 pore pairs each; pore series diverging; petaloid region covering approximately 95% of test. Interporiferous zones width greater than width of poriferous zones.
Glassy tubercles - Moderate size.
Peristome - Moderate size, about 25% test length; rounded pentagonal shape.
Infundibulum deep. 41
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines
on plates.
Geographic range - Initially collected at Mauritius, but may not be valid. From Hawaii
south to Marshall Islands and Tahiti.
Bathymetric range - 20-75 m.
Remarks. Clark reports the type locality as Mauritius, but this species has not been
found there by later investigations (Mortensen 1948a). It is possible that the locality was
reported incorrectly, as E. megapetalus has only been found in Hawaii, Micronesia, and
Polynesia since the original description. E. megapetalus is easily distinguished from other
Pacific species by its well-developed petals which cover the majority of its test and its fairly
deep infundibulum.
Echinocyamus platytatus H.L. Clark, 1914
Fig. 24, 25.
1914. Echinocyamus platytatus H.L. Clark, p. 63, PI. 127.
1925. Fibulariaplatytata (H.L. Clark), Lambert & Thiery, p. 577.
Type material. Syntypes: MCZ 2306.
Material studied. MCZ 2306 (syntypes), Victoria, MCZ 2307, Victoria, MCZ 5020,
South Australia, MCZ 5021, South Australia 42
Type locality. Port Phillip, Victoria.
Description.
Size and shape -Subcircular test, with width 75-99% test length. Test flat, 20-35% test
length. Maximum height central. Size range 2.6-8.7 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - One to seven hydropores. Ocular pores smaller than gonopores.
Ambulacra - Petals with 4-16 pore pairs each; pore series parallel; petaloid region
covering approximately 40% of test. Interporiferous zones width greater than width of
poriferous zones.
Glassy tubercles - Moderate size.
Peristome - Small, about 15% test length; round shape. Infundibulum shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines
on plates.
Pedicellaria - Ophicephalous only.
Geographic range - Southeastern Australia, from St. Vincent’s Gulf, S.A., to Broughton
Island, N.S.W.
Bathymetric range - 10-400 m.
Remarks. This species is superficially similar to E. planissimus, as they are both fairly flat, Australian species. However, the petals of E. platytatus are much narrower at their 43
proximal end. It is interesting to note that this is the only valid species of Echinocyamus to have more than one hydropore; however, this character alone is not enough to remove it from the genus.
Echinocyamus provectus de Meijere, 1903
Fig. 26, 27.
1903. Echinocyamus provectus de Meijere, p. 6.
1914. Fibulariaprovecta (de Meijere), Lambery and Thiery, p. 292.
Type material. Syntype: ZMUC-ECH-260. There may be other type specimens not found during search.
Material studied. ZMUC-ECH-260 (syntype), Timor, USNM E09328, China Sea,
USNM E35874, Philippines, MCZ 2308, New South Wales, MCZ 7237, Tasmania, MCZ 8031,
New South Wales
Type locality. Timor, Indonesia.
Description.
Size and shape -Elongate test, with width 69-94% test length, with the majority <85% test length. Test high, 29-50% test length. Maximum height toward posterior. Size range 1.8-8.5 mm long. 44
Internal buttressing - Ten radial walls, one pair in each interambulacrum, slightly thickened on medial end.
Apical system - Single hydropore. Ocular pores smaller than gonopores.
Ambulacra - Petals with 6-18 pore pairs each; pore series converging; petaloid region covering approximately 50% of test. Interporiferous zones width greater than width of poriferous zones.
Glassy tubercles - Larger than spine-bearing primary tubercles, particularly large on posterior.
Peristome - Moderate size, about 20% test length; rounded pentagonal shape.
Infundibulum shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Spines - Primary spines short, 0.26 mm long. Miliary spines 0.2 mm long.
Geographic range - From southwest Indian Ocean, off Mozambique, to Western Pacific
Ocean with Hong Kong in the north and Tasmania in the south.
Bathymetric range - 10-390 m.
Remarks. E. provectus is identifiable by the converging pore series in its petals and prominent glassy tubercles. It is unusual in that its glassy tubercles are often particularly prominent in the posterior portion of the test, while most other species with prominent glass tubercles have them evenly scattered on the test. 45
Echinocyamus scaber de Meijere, 1903
Fig. 28, 29.
1903. Echinocyamus scaber de Meijere, p. 5.
1914. Fibularia scabra (de Meijere). Lambert & Thiery, p. 292.
Type material. None found.
Material studied. ZMUC-ECH-541 (syntypes for E.s. subconicus), Kai Islands, CASIZ
198062, Hawaii
Type locality. Indonesia.
Description.
Size and shape -Subcircular test, with width 85-96% test length. Test high, 32-48% test length. Maximum height central. Size range 2.7-8.0 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores similar size to gonopores.
Ambulacra - Petals with 4-12 pore pairs each; pore series parallel; petaloid region covering approximately 40% of test. Interporiferous zones width equal to width of poriferous zones.
Glassy tubercles - Larger than spine-bearing primary tubercles. 46
Peristome - Moderate size, about 20% test length; round shape. Infundibulum shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Spines - Primary spines about 0.5 mm, slightly tapered, strong longitudinal ribs.
Miliary spines 0.22-0.30 mm.
Geographic range - Indo-Pacific, from South Africa to Hawaii and New South Wales.
Bathymetric range - 250-1950 m.
Remarks. E. scaber is recognizable by its prominent glassy tubercles, short petals, and subcircular test.
Echinocyamus sollers Koehler, 1922
Fig. 30-32.
1922. Echinocyamus sollers Koehler, p. 132. PI. XII, XV.
1925. Fibularia sollers (Koehler). Lambert & Thiery, p. 577.
Type material. A cotype is at ZMUC, but did not have a unique identifier assigned to it yet.
There are likely to be more types at other museums.
Material studied. ZMUC cotype, MCZ 6023, Andaman Sea
Type locality. Laccadive Sea. 47
Description.
Size and shape -Subcircular test, with width 91-95% test length. Test high, 43-47% test length. Maximum height central. Size range 5.4-7.0 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum.
Apical system - Single hydropore. Ocular pores similar in size to gonopores.
Ambulacra - Petals with 8-14 pore pairs each; pore series parallel; petaloid region covering approximately 60% of test. Interporiferous zones width equal to width of poriferous zones.
Glassy tubercles - Moderate size.
Peristome - Moderate size, about 20% test length; round shape. Infundibulum shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Spines - Primary spines up to 1 mm long, gradually widen from base then taper quickly near tip, strong denticulation. Miliary spines on average 0.035 mm long.
Pedicellaria - Ophicephalous pedicellariae with head about 0.1 mm, strong denticulation along blade. Triphyllous pedicellariae with valves about 0.08 mm, teeth become finer towards tip.
Geographic range - From Laccadive Sea off southern India to Andaman Sea.
Bathymetric range - 250-900 m. 48
Remarks. E. sollers is most similar to E. scaber, but lacks the prominent glassy tubercles found in the latter.
Echinocyamus (Mortonia) australis (des Moulins, 1835)
Fig. 33, 34.
1835. Fibularia australis des Moulins, I p. 187.
1847. Echinocyamus australis (des Moulins). L. Agassiz & Desor, p. 140.
1851. Mortonia australis (des Moulins). Gray, p. 38.
1948a. Echinocyamus (Mortonia) australis (des Moulins). Mortensen, p. 197.
Type material. None found.
Material studied. MCZ 4639, Hawaii.
Type locality. South Sea Islands.
Description.
Size and shape -Elongate, bordering on subcircular test, with width 69-91% test length, with the majority <85% test length. Oral side rises gently anterior to peristome. Test high, 31 -
53% test length. Maximum height towards anterior. Size range 1.8-18.1 mm long.
Internal buttressing-Two radial walls in the posterior interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores. 49
Ambulacra - Petals with 20-52 pore pairs each; pore series parallel; petaloid region covering approximately 75% of test. Interporiferous zones width greater than width of poriferous zones.
Glassy tubercles - Moderate size.
Peristome - Small, about 10% test length; rounded pentagonal shape. Infundibulum deep.
Periproct - Spines present, many irregular plates on periproctal membrane.
Spines - Primary spines 1 mm long, simple, and club-shaped. Miliary spines slightly widened distally.
Pedicellaria- Ophicephalous pedicellaria present. Tridentate pedicellariae small, head about 0.1 mm long.
Geographic range -South Pacific, from Hawaii, Midway Islands, Palmyra, and Loyalty
Islands.
Bathymetric range - 10-75 m.
Remarks. Based on the phylogenetic analysis, this species can no longer be considered a separate genus without causing Echinocyamus to be polyphyletic. Therefore, this study proposes returning Mortonia to subgenus rank, so that E. (M.) australis and E. (M.) polyporus can be considered distinct for their unusual periproct characters and reduced internal walls, but still part of the Echinocyamus clade. Echinocyamus (Mortonia) is also notable for having the two largest species of Echinocyamus. 50
Echinocyamus (Mortonia) polyporus Mortensen, 1921
Fig. 35-37.
1921. Echinocyamus polyporus Mortensen, p. 176, PI VI.
1925. Fibulariapolypora (Mortensen). Lambert & Thiery, p. 577.
1948a. Echinocyamus (Mortonia) polyporus Mortensen, p. 200.
1955. Mortonia polyporus (Mortensen). Durham, p. 134.
Type material. Holotype: ZMUC-ECH-263. Cotypes: ZMUC-ECH-263 (same # as holotype, but with two specimens). Paratype: MCZ 5016.
Material studied. ZMUC-ECH-263 (holotype and cotypes), Cook Strait, MCZ 5016
(paratype), New Zealand, MCZ 3305, Kermadecs, USNM El 6321, Norfolk Island.
Type locality. Cook Strait.
Description.
Size and shape -Elongate test, with width 70-87% test length. Test high, 31 -39% test length. Maximum height towards anterior. Size range 6.1-13.2 mm long.
Internal buttressing - Two radial walls in the posterior interambulacrum.
Apical system - Single hydropore. Ocular pores smaller than gonopores. 51
Ambulacra - Petals with 12-32 pore pairs each; pore series parallel; petaloid region
covering approximately 75% of test. Interporiferous zones width greater than width of
poriferous zones.
Glassy tubercles - Moderate size.
Peristome - Small, about 10% test length; rounded pentagonal shape. Infundibulum
deep.
Periproct - Spines present, many plates on periproctal membrane.
Geographic range - Southwest Pacific, from Norfolk Island, Kermadec Islands, New
Zealand, and Lord Howe Island.
Bathymetric range - 10-100 m.
Remarks. This species can be quite difficult to differentiate from E. (M.) australis. In general, specimens of the same size will have fewer pore pairs in E. (M.) polyporus, but this character is not always reliable. It is likely that this species could be synonymized with E. (M.) australis. 52
Incertae sedis
“Echinocyamus”planissimus H.L. Clark, 1938
Fig. 38-40.
1938. Echinocyamus planissimus H.L. Clark, p.422, PI.27.
Type material. Holotype: MCZ 7234. Paratypes: MCZ 7235, ZMUC-ECH-258, J.6160 at the
Australian Museum.
Material studied. MCZ 7234 (holotype), MCZ 7235 (paratypes), ZMUC-ECH-258
(paratypes), MCZ 7236, all from Western Australia
Type locality. Pearl Shoal, Broome, Western Australia, 5-7 frns.
Description.
Size and shape -Elongate, bordering on subcircular test, with width 75-89% test length.
Test flat, 23-26% test length. Maximum height central. Size range 5.3-8.1 mm long.
Internal buttressing - Ten radial walls, one pair in each interambulacrum, with
orthogonal circumferential walls (see fig. 40).
Apical system - Three to seven hydropores. Ocular pores smaller than gonopores.
Ambulacra - Petals with 8-16 pore pairs each; pore series converging; petaloid region covering approximately 60% of test. Interporiferous zones width greater than width of poriferous zones.
Glassy tubercles - Moderate size. 53
Peristome - Small, about 10% test length; rounded pentagonal shape. Infundibulum shallow.
Periproct - A few (but always more than 4) plates on periproctal membrane; no spines on plates.
Pedicellaria - Ophicephalous and tridentate only.
Geographic range - Reported from Broome, Western Australia and off of Twofold Bay,
New South Wales.
Bathymetric range - 10-110 m.
Remarks. This species is certainly not an Echinocyamus or even a fibulariid due to the presence of small “endcaps” on each of its internal radial walls. E. planissimus is also a bit odd for an Echinocyamus due to its multiple hydropores and particularly wide petals that converge distally to be almost closed. E. planissimus should be moved to Laganidae, most likely as a species of Peronella which has a similar petal shape and 4 gonopores. 54
Discussion
Based on the results of the elliptical Fourier analysis, almost all species of
Echinocyamus were found to be morphologically distinct from one another, supporting the validity of these species. Of the species that did not show a significant difference from other
Echinocyamus, all but one were represented in the analysis by ten or fewer specimens.
Therefore, there was not enough material for the analysis to differentiate these taxa: E. planissimus, E. sollers, and E. polyporus. E. scaber, on the other hand, was represented by a large sample size but still could not be differentiated from E. incertus and E. grandiporus on the basis of shape alone. However, E. incertus and E. scaber are easily differentiated based on the greater size of the ocular pores in the latter. E. scaber and E. grandiporus are somewhat more similar to each other morphologically, but they have very different, non-overlapping distributions, with the former in the Indo-Pacific and the latter in the Caribbean. These two species were also found to be in the same clade within Echinocyamus, as will be discussed later.
The allometric patterns found in the PCA results indicate previously undetected growth patterns. In analyses of the ambital shape, the relative width of the test increased faster than test length in E. australis, E. crispus, E. polyporus, and E. pusillus showing how these species became rounder as they matured (fig.45-47, 49). E. provectus and E. scaber also showed statistically significant changes in their ambital shape, but no clear patterns emerged when the shapes were compared (fig. 48, 50, 51). In the lateral view, E. crispus and E. platytatus appear to shift their apex from an anterior position to a more central position as they mature (fig. 52,
53). The aboral surface of E. provectus and E. pusillus tends to flatten as they mature (fig. 54,
55), while E. scaber develops a higher test and more prominent apex (fig.56). Conclusions 55
about these patterns are preliminary and need to be tested with additional specimens from a greater size range to further hone descriptions of these ontogenetic changes.
The topology inferred by the majority rule consensus tree lacked statistical support, with a rescaled Cl of only 0.199 among the characters (RI=0.565), but there were several informative and consistent trends. To resolve the tree further, it is clear that additional characters that can be analyzed throughout ontogeny need to be developed. Such character analysis might be helpful towards increasing confidence in hypotheses of homology. However, some aspects of the analysis allow for several important inferences regarding the evolution and taxonomy of the genus. Among extant taxa, and in accord with most prior interpretations (Kroh and Smith, 2010, Mooi et al., 2014), the sister genus of Echinocyamus was found to be
Fibularia. However, Mortonia was clearly nested within Echinocyamus, with E. megapetalus strongly supported as a sister taxon.
There is also strong evidence from the morphological characters alone that both E. planissimus and E. platytatus are outside Echinocyamus and might not be part of the fibulariid clade at all. Based on the presence of internal walls with circumferential branches in E. planissimus (see fig. 40), this species belongs in the family Laganidae, not Fibulariidae. E. planissimus could actually be a Peronella based on the number of pore pairs and petals that are relatively wide proximal to the apex, but more study of the plate architecture of E. planissimus as well as of all laganids, including Peronella, is needed before E. planissimus can be placed unequivocally in any genus within Laganidae. It is even possible that E. planissimus represents a new genus of miniaturized laganiform. If E. planissimus is a true laganid, then miniaturization may have occurred more than once in the Laganiformes, once in the basal 56
fibulariid lineage and later separately in E. planissimus alone. Alternatively, if E. planissimus is basal within the laganids, miniaturization could have occurred only once at the beginning of the
Laganiformes lineage, and was subsequently reversed within Laganidae. This is unlikely, however, given that previous analyses indicate multiple derivations of miniaturization in the other clypeasteroid clades, and that relationships among them support the idea that for each of these clypeasteroid clades, the ancestral morphology was a relatively large, flattened form that followed the “sand dollar paradigm” (Seilacher, 1979, Mooi, 1990).
Unambiguous synapomorphies are rare on the consensus tree (fig. 67-87). Almost all characters display at least some homoplasy. The most promising synapomorphy for
Echinocyamus, the presence of podial pores primarily in sutures between plates, was also found in the laganid Sismondia occitana and in E. platytatus, which is not a true Echinocyamus according to the current consensus tree (fig. 66). No completely unequivocal synapomorphy differentiating all Echinocyamus species from the other fibulariids was found. Within
Echinocyamus, the posterior position of the maximum test height was a synapomorphy within
Clade C, with the exclusion of E. insularis. Another synapomorphy, the presence of a deep infundibulum, was found in all members of Clade B. It is likely that future analyses using molecular data will resolve some of the nodes more clearly, but at present, material to do this work is lacking.
Kroh and Smith (2011) resurrected the family Echinocyamidae to contain
Echinocyamus and Mortonia, rather than retaining them in Fibulariidae, as had been commonly accepted since before Mortensen (1948). The latter clarified this relationship in a monograph on all the clypeasteroid species. However, without an unequivocal synapomorphy to support 57
Echinocyamidae, and the current sister relationship of Echinocyamus and Fibularia, it seems unnecessary to retain Echinocyamus in its own family. Additionally, Leniechinus and E. platytatus currently split off individually at the base of the clade containing Echinocyamus and
Fibularia. If this clade was split into two separate families, these two taxa would need two new monotypic families to prevent Fibulariidae from being paraphyletic. The current data supports retention of all these taxa in the family Fibulariidae.
The four clades suggested to exist within Echinocyamus do not shed light on specific biogeographic patterns within the genus. However, considering that the sister taxa of
Echinocyamus (i.e. Fibularia and fossil taxa) are Indo-Pacific, it is most parsimonious to assume that Echinocyamus also originated in this region. Clade A has a highly disjunct distribution, with E. pusillus in the Mediterranean and Northeast Atlantic, and E. apicatus off southeast Australia. This suggests that E. pusillus moved into the European region later (but still very early in the radiation of the genus) through the Tethyan seaway. Clades B and C are widely distributed throughout the Indo-Pacific. Clade D is also found throughout the Indo-
Pacific, with two species, E. macrostomus and E. grandiporus, found only in the Caribbean. E. macrostomus and E. grandiporus are sister taxa, so it is likely that Echinocyamus colonized the
Caribbean once and speciated later.
The known fossil record of Echinocyamus begins with a rapid diversification in the
Eocene, with few transitional forms (Kier, 1982). It was once generally assumed that fibulariids represented the basal clypeasteroid form (Durham, 1955, Kier, 1982). This was partly supported by the inclusion of Togocyamus and other diminutive fossil echinoids in the
Fibulariidae. Togocyamus displays many ancestral characteristics, but might not even be a true 58
clypeasteroid. It is certainly not a fibulariid due to its lantern support structure, periproct position, and podial arrangement (Mooi, 1990, p.26). The removal of E. planissimus from
Echinocyamus further supports the hypothesis that miniaturization is a derived state that evolved independently in several different clypeasteroid lineages, rather than representing the basal state of Clypeasteroida. Miniaturization in clypeasteroids is often a result of paedomorphosis, as the adult microechinoids retain many juvenile characteristics of their larger sister taxa (Mooi, 1990). Further study of the developmental pathways in clypeasteroids could shed light on why miniaturization occurs so frequently.
Further work is needed to refine the phylogeny of Echinocyamus. Molecular analysis is a promising direction to obtain more phylogenetic data, but whole specimens with viable DNA are difficult to acquire. Resolving the status of Fibularia and Echinocyamus in the fossil record will be necessary to uncover the evolutionary history of microechinoids. MicroCT scanning will undoubtedly reveal more informative morphological traits, such as plate maps and a more detailed view of the internal buttresses. Additional specimens of some of the rarer species, such as E. sollers, and the more unusual species, such as E. planissimus, are needed for more complete morphometric analyses, and to determine placement and taxonomic status of such forms. Nevertheless, this study provides some supportable refinements concerning monophyly not only of Echinocyamus, but of Fibulariidae, as well as glimpses into the difficulties surrounding studies of this challenging taxon. 59
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Appendices.
Appendix 1. Character matrix. Numbers in bold at top indicate character number represented in each column.
1 1 1 1 1 1 1 1 1 1 2 2 Taxa 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 Echinocyamus pusillus 1 0111111102001210 0 0 0 0 E. apicatus 1 0 1 1 1 1 1 1 1 010012100000 E. convergens 2 0 1 1 1 1 1 2 21000121110 0 0 E. crispus 001111111112112100000 E. elegans 1 011111100100121 1 0 0 0 0 E. grandiporus 0 0 1 1 1 1 1 2 2 0 1 0 0 1 2 1 0 1 0 1 0 E. incertus 2 01111121110012110000 E. macrostomus 001111122010022101000 E. megapetalus 0 0 1 1 1 1 1 0 0 1 2 2 1 1 2 1 0 0 0 0 0 E. planissimus 00 11 0111001 0 1 1 0 0 0 0 1 E. platytatus 0 0 1 1 1 1 0 2 2 010001100000 E. provectus 201111111100112101000 E. scaber 0 0 1 1 1 1 1 2 1 0 1 0 0 1 2 1 1 1 0 1 0 E. sollers 001111111010012110010 E. australis 1 1 1 0 1 1 0 0 1 1 2 1 0 2 1 0 0 0 0 0 E. insularis 1 0 1 0 1 ? 121110002110000 Fibularia ovulum 1 21110221011111010000 F. dubarensis ? 2 1 7 1 021101000101 7 0 0 0 F. cribellum 0 211101201 10 0 1 1 0 1 0 0 0 0 Laganum laganum 0 0000000000 0 0 0 0o I 0 0 1 0 1 Sismondia occitana 0 0 0 7 7 1 1 0 0 0 0 0 1 0 0 0 0 7 0 0 0 Leniechinus herricki 2 0 1 7 7 0 0 0 0 1 1 0 0 0 2 1 0 7 0 0 0 Lenicyamidia compta 1 2 1 7 7 0 1 1 1 0 0 0 0 2 2 1 1 7 1 0 0 Cyamidia nummulitica 1 2 1 7 7 0 1 1 0 0 1 0 1 0 2 1 1 7 1 0 0 7 ? F. kieri 0 1 1 0 2 2 1 0 1 1 0 1 2 1 *1 1 2 0 0 0 E. petalus 0 0 1 7 7 1 1 1 1 0 1 0 0 1 2 1 0 7 0 0 0 E. parviporus 0 0 1 7 7 1 1 1 1 0 1 0 0 0 2 1 0 7 0 0 0 Peronella peronii 0 0 0 1 0 0 0 0 1 0 0 0 1 0 0 1 0 0 , 1 0 1 66
Tables.
Table 1. List of extant species with taxonomic authority. Asterisk marks type species of genus.
Species Taxonomic authority Terra typica Depth range Echinocyamus apicatus Mortensen, 1948b Tasman Sea 70-100 m Echinocyamus convergens Mortensen, 1948b West Indian Ocean 130-350 m Echinocyamus crispus Mazzetti, 1893 Indo-Pacific, 0-1880 m Hawaii Echinocyamus elegans Mazzetti, 1893 Arabian Sea 50-275 m Echinocyamus Mortensen, 1907 Caribbean Sea 30-2500 m grandiporus Echinocyamus grandis H.L. Clark, 1925 Seychelles 60 m Echinocyamus incertus H.L. Clark, 1914 Hawaii 20-360 m Echinocyamus insularis Mironov & Sagaidachny, Southeast Pacific 50-80 m 1984 Echinocyamus Mortensen, 1907 Caribbean Sea 180-3000 m macrostomus Echinocyamus H.L. Clark, 1914 South Pacific 20-75 m megapetalus Echinocyamus H.L. Clark, 1938 Western Australia 10-110 m planissimus Echinocyamus platytatus H.L. Clark, 1914 Tasman Sea 10-400 m Echinocyamus provectus de Meijere, 1903 Indo-Pacific 10-390 m Echinocyamus pusillus * (O.F. Muller, 1776) Mediterranean Sea, 0-1250 m North Atlantic Echinocyamus scaber de Meijere, 1903 Indo-Pacific 250-1950 m Echinocyamus sollers Koehler, 1922 North Indian 250-900 m Ocean Mortonia australis* (des Moulins, 1837) Hawaii 10-75 m Mortonia polyporus (Mortensen, 1921) Tasman Sea 10-100 m 67
Table 2. List of specimens photographed for morphometric analysis, a = anterior, 1 = left, o = oral. ZMUC* specimens do not have a catalog number.
Catalog number Species # specimens type status views locality MCZ 8051 E. apicatus 1 holotype a,l,o New South Wales MCZ 8095 E. apicatus 1 a,o New South Wales ZMUC-ECH- 542 E. apicatus 1 syntype a,o New South Wales USNM E36086 E. convergens 1 a,o Mozambique USNM E36094 E. convergens 1 a,o Mozambique Indian Ocean, USNM E36101 E. convergens 7 a,o off South Africa Providence Island, ZMUC-ECH- West Indian 110 E. convergens 1 holotype a,l,o Ocean Arabian Sea, off CASIZ 101538 E. crispus 3 a,l,o India CASIZ 104508 E. crispus 1 a,l,o Ryukyu Islands CASIZ 115632 E. crispus 12 a,l,o Hawaii CASIZ 173810 E. crispus 17 a,l,o Hawaii Arabian Sea, off MCZ 8724 E. elegans 28 a,l,o Pakistan Arabian Sea, off USNM El0681 E. elegans 3 a,l,o India Arabian Sea, off USNM E036138 E. elegans 1 a,o India CASIZ 90537 E. elongatus 5 a,l,o Hawaii MCZ 3812 E. elongatus 1 a,l,o Guam MCZ 4212 E. elongatus 1 a,l,o Hawaii USNM 34235 E. elongatus 1 holotype a,l,o Hawaii USNM 34236 E. elongatus 1 paratype a,l,o Hawaii USNM 34237 E. elongatus 1 paratype 0 Hawaii USNM 38204 E. elongatus 1 a,1,0 Guam USNM E07502 E. elongatus 6 a,l,o Marshall Islands USNM 10623 E. grandiporus 2 a,l,o Yucatan Channel USNM 1014103 E. grandiporus 12 a,l,o Brazil Saint Lucia, USNM E14552 E. grandiporus 12 a,l,o Lesser Antilles USNM E36170 E. grandiporus 2 a,l,o Florida 68
Table 2 (continued).
Catalog # type number Species specimens status views locality ZMUC- ECH-143 E. grandiporus 1 syntype a, l,o off Key West, Florida ZMUC- ECH-447 E. grandiporus 1 syntype a,l,o Azores ZMUC- ECH-450 E. grandiporus 1 syntype a,l,o Florida ZMUC- ECH-451 E. grandiporus 1 syntype a,l,o St. Croix ZMUC- possible ECH-452 E. grandiporus 1 syntype a,l,o Havana, Cuba UCMP 36579 E. incertus 1 hypotype a,l,o Nazca Ridge UCMP 36580 E. incertus 1 hypotype a,l,o Nazca Ridge UCMP 36581 E. incertus 1 hypotype a,l,o Nazca Ridge UCMP 36582 E. incertus 1 hypotype a,l,o Nazca Ridge CASIZ 198061 E. incertus 17 a,l,o Hawaii USNM 034208 E. incertus 1 holotype a,l,o Hawaii USNM E07374 E. incertus 1 a,l,o Marshall Islands ZMUC* E. incertus 1 o Honolulu, Hawaii E. MCZ 4589 macrostomus 2 a,l,o Madeira Islands E. MCZ 7802 macrostomus 2 a,l,o Cuba USNM E. E14554 macrostomus 17 a,l,o Bahamas USNM E. E14557 macrostomus 15 a,l,o Cuba ZMUC- E. ECH-214 macrostomus 1 cotype a,l,o Unknown ZMUC- E. ECH-453 macrostomus 2 syntypes a,l,o St. 2044? CASIZ 206938 E. megapetalus 1 a,l,o Hawaii 69
Table 2 (continued).
Catalog # type number Species specimens status views locality CASIZ 187427 E. megapetalus 1 a,l,o Philippines MCZ 2294 E. megapetalus 3 paratypes a,l,o Mauritius USNM E08491 E. megapetalus 27 a,l,o Marshall Islands ZMUC* E. megapetalus 1 a,l,o Honolulu, Hawaii MCZ 7234 E. planissimus 1 holotype a,l,o Western Australia MCZ 7235 E. planissimus 2 paratypes a,l,o Western Australia MCZ 7236 E. planissimus 1 a,l,o Western Australia ZMUC* E. planissimus 3 a,l,o New South Wales ZMUC- ECH-258A E. planissimus 1 paratype a,l,o Western Australia MCZ 2306 E. platytatus 11 syntypes a,l,o Victoria MCZ 2307 E. platytatus 2 a,l,o Victoria MCZ 5020 E. platytatus 7 a,l,o South Australia MCZ 5021 E. platytatus 14 a,l,o South Australia St. Vincent Gulf, South ZMUC* E. platytatus 1 a,o Australia Backstairs Passage, ZMUC* E. platytatus 3 a,l,o South Australia ZMUC* E. platytatus 2 a,l,o New South Wales MCZ 2308 E. provectus 2 a,l,o New South Wales MCZ 7237 E. provectus 1 a,l,o Tasmania MCZ 8031 E. provectus 2 a,l,o New South Wales USNM E09328 E. provectus 1 a,l,o China Sea USNM E35874 E. provectus 31 a,l,o Philippines USNM E36024 E. provectus 16 a,l,o Mozambique ZMUC* E. provectus 2 a,l,o Banda Sea ZMUC* E. provectus 2 a,l,o Kei Islands ZMUC- ECH-260 E. provectus 1 syntype a,o Timor? CASIZ 104183 E. pusillus 2 a,l,o Azores 70
Table 2 (continued).
Catalog # type number Species specimens status views locality CASIZ 111220 E. pusillus 11 a,l,o Scotland CASIZ Bay of Biscayne, 111440 E. pusillus 9 a,l,o France CASIZ 198063 E. pusillus 13 a,l,o Northern France CASIZ 198062 E.scaber 29 a,l,o Hawaii USNM 34253 E.scaber 1 a,l,o Hawaii ZMUC-ECH- E.scaber 541A var.subconicus 1 syntype a,l,o Kei Islands ZMUC-ECH- E.scaber 541B var.subconicus 3 syntype a,l,o Kei Islands MCZ 6023 E. sollers 3 a,l,o Andaman Sea ZMUC* E. sollers 2 cotype a,l,o Investigator St. 250 CASIZ 90527 M. australis 2 a,l,o Midway Island CASIZ 90535 M. australis 2 a,l,o Hawaii CASIZ 90549 M. australis 2 a,l,o New Guinea CASIZ 90550 M. australis 3 a,l,o Midway Atoll CASIZ 90551 M. australis 2 a,l,o Midway Atoll CASIZ 96780 M. australis 3 a,l,o Pitcairn Islands CASIZ 112807 M. australis 4 a,l,o Marshall Islands MCZ 4639 M. australis 11 a,l,o Hawaii ZMUC* M. australis 2 a,l,o Honolulu, Hawaii CAS 173811 M. australis 15 a,l,o Hawaii MCZ 3305 M. polyporus 3 a,l,o Kermadecs USNM E16321 M. polyporus 1 a,l,o Norfolk Island ZMUC-ECH- 263A M. polyporus 1 type a,l,o Cook Strait ZMUC-ECH- 263B M. polyporus 2 cotypes a,l,o Cook Strait ZMUC-ECH- 462 M. polyporus 4 syntype a,l,o Kermadecs Table 3. Principal components that show a significant correlation with length.
Significant PC - Significant PC - Significant PC - Species Aboral Lateral Combined M. australis PCI None PC2 E. crispus PC2 PCI PCI E. elegans None None PC3 E. grandiporus None None PC2 E. incertus None None PCI, PC2 E. macrostomus None None None E. megapetalus None None None E. planissimus None None None E. platytatus None PCI None M. polyporus PC3 None PC2 E. provectus PC3 PC2 PC2 E. pusillus PC3 PC3 PC2 E. scaber PC2, PC3 PC2 None E. sollers None None None 72
Figures.
ABORAL SURFACE (TOP) ORAL SURFACE (BOTTOM) fplate columns labeled according to Loven’s system! ANTERIOR a interambulacrum
ambulacrum POSTERIOR b 5 a
B perradialt suture
Figure 1. Diagram of the basic anatomy of Echinocyamus. A. Aboral (left) and oral (right) view of denuded test. B. Close-up view of ambulacrum, illustrating structures within a petal, abc = ambulacral basicoronal plate, b = buccal pore, g = gonopore, h = hydropore, ibc = interambulacral basicoronal plate, o = ocular pore, p = pore pair, ppr = periproct, pst = peristome. Illustrated by R. Mooi. 73
Figure 2. Left view of E. megapetalus (USNM E08491, specimen 11). Arrows show location of podial pores in lines along plate sutures. Scale bar = 1 mm. Figure 3. Aboral, oral, and left view of Echinocyamus pusillus (CASIZ 111220, spec. 10). Figure 4. X-ray of Echinocyamus pusillus (CASIZ 111220, spec. 10). 76
Figure 5. Aboral, oral, and left view of the holotype of Echinocyamus apicatus (MCZ 8051). 77
Figure 6. X-ray of Echinocyamus apicatus (MCZ 8095, spec. 2). 78
Figure 7. Aboral and oral view of Echinocyamus convergens (NMNH E36101, spec. 1). 79
Figure 8. Aboral, oral, and left view of the holotype of Echinocyamus convergens (ZMUC- ECH-110). Figure 9. X-ray of Echinocyamus convergens (NMNH E36101, spec. 8). 81
Figure 10. Aboral, oral, and left view of Echinocyamus crispus (CASIZ 173810, spec. 7). Figure 11. X-ray of Echinocyamus crispus (CASIZ 173810, spec. 2). 83
Figure 12. Aboral, oral, and left view of Echinocyamus elegans (MCZ 8724, spec. 34). Figure 13. X-ray of Echinocyamus elegans (MCZ 8724, spec. 25). 85
Figure 14. Aboral, oral, and left view of Echinocyamus grandiporus (NMNH E14552, spec. 6). 86
Figure 15. Aboral, oral, and left view of a syntype of Echinocyamus grandiporus (ZMUC-ECH- 447, spec. 1). Figure 16. X-ray of Echinocyamus grandiporus (NMNH El 4552, spec. 9). 88
Figure 17. Aboral, oral, and left view of the holotype of Echinocyamus incertus (NMNH 034208). 89
Figure 18. X-ray of the holotype of Echinocyamus incertus (NMNH 34208). 90
Figure 19. Aboral, oral, and left view of Echinocyamus macrostomus (MCZ 7802, spec. 2). 91
Figure 20. Aboral, oral, and left view of a syntype of Echinocyamus macrostomus (ZMUC- ECH-453, spec. 2). Figure 21. X-ray of Echinocyamus macrostomus (NMNH E14557, spec. 10). 93
Figure 22. Aboral, oral, and left view of Echinocyamus megapetalus (NMNH E8491, spec. 11). 94
Figure 23. X-ray of Echinocyamus megapetalus (NMNH E8491, spec. 1). 95
Figure 24. Aboral, oral, and left view of a syntype of Echinocyamus platytatus (MCZ 2306, spec. 6). 96
Figure 25. X-ray of a syntype of Echinocyamus platytatus (MCZ 2306, spec. 6). 97
Figure 26. Aboral, oral, and left view of Echinocyamus provectus (NMNH E3874, spec. 5). 98
Figure 27. X-ray of Echinocyamus provectus (NMNH E35874, spec. 13). 99
Figure 28. Aboral, oral, and left view of Echinocyamus scaber (CASIZ 198062, spec. 8). 100
Figure 29. X-ray of Echinocyamus scaber (CASIZ 198062, spec. 4). 101
Figure 30. Aboral, oral, and left view of Echinocyamus sollers (MCZ 6023, spec. 1). 102
Figure 31. Aboral, oral, and left view of a cotype of Echinocyamus sollers (ZMUC). 103
Figure 32. X-ray of Echinocyamus sollers (MCZ 6023, spec. 1). 104
Figure 33. Aboral, oral, and left view of Echinocyamus australis (CASIZ 90550, spec. 3). Figure 34. X-ray of Echinocyamus australis (MCZ 4639, spec. 8). 106
Figure 35. Aboral, oral, and left view of Echinocyamus polyporus (MCZ 3305, spec. 1). 107
Figure 36. Aboral, oral, and left view of a type specimen of Echinocyamus polyporus (ZMUC- ECH-263, spec. 1). Figure 37. X-ray of Echinocyamus polyporus (MCZ 3305, spec. 1). 109
Figure 38. Aboral, oral, and left view of the holotype of Echinocyamus planissimus (MCZ 7234). 110
Figure 39. Aboral, oral, and left view of a paratype of Echinocyamus planissimus (MCZ 7235, spec. 1). Ill
Figure 40. X-ray of the holotype of Echinocyamus planissimus (MCZ 7234). variate 12. vs.variatevariate Figure 41. Canonical Variate Analysis on combined dataset of outlineslateral aboraland- dataset of VariateonAnalysiscombined Canonical 41.Figure
CVA2 8
♦ -6 V1 S CVA2 VS. CVA1 -8 -6 8 CVA1 AA m E. provectus - x E.x scaber ♦ x x + E.megapetalus ▲ + E. platytatus E incertus E.elegans M. australis M.
++ ■ ■ • x x ▲ ♦ x - E. sollers E. pusillus M.polyporus E. planissimus E.macrostomus E.grandiporus E.crispus 8
112 ait 13.variate vs.variate Figure 42. Canonical Variate Analysis on combined dataset of outlineslateralaboraland - onAnalysisFigurecombined dataset of VariateCanonical 42.
-8 CVA 3
♦ -6 V1 S CVA3 VS CVA1 6 V _ , _ 1 CVA * X -r H ^ + + + + ? . x x ?. .0 ♦ ♦ ME. provectus £ - 4 4 + £ +£ x x *++ + V - E.elegans australis M. £ £ f. " t “ ' incertus platytatus megapetalus scaber
■ £ ■ £ m A ♦ X x £ £ x - £ £ - £ £ £ £ £ M.polyporus pusillus planissimus sollers crispus macrostomus grandiporus 8
113
Figure 43. Principal component analysis of aboral view. Uppermost side of outlines is anterior. is outlines of side Uppermost view. aboral of analysis component Principal 43. Figure Principal Component 2 - 39% 0.3 Principal Component Analysis Principal Component of Aboral rnia opnn 56% 1 - Component Principal 0 . 1 • % View . • • 0.3 • • • • • • • • • • m E. sollers • m E.mega petal us % E.confer gens • E.scaber E.australis E.macrostomus E.incertus E.grandiporus E.crispus E. pusillus E.provectus E. polyporus E. platytatus E. planissimus E.apicatus E.elegans
114
Figure 44. Principal component analysis of lateral view. Right side of outlines is anterior. is outlines of side Right view. lateral of analysis component Principal 44. Figure PRIncipal component 2 - 12% . -0 1.8 C ____ m ) < 6 -0 .6 • • Principal Component AnalysisLateralPrincipal of Component View V | , ■4 ..
w rn m • • . ...»
rnia opnn 82% 1 - component Principal 0.15 0.25 0.2 . • • • • • • m E. platytatus • • • m E.incertus • m E.crispus •
E. sollers E.scaber E. pusillus E. provectus E.macrostomus E.elegans E. polyporus E.grandiporus E. planissimus E.megapetalus E.australis
115 116
Figure 45. Linear regression of length versus first principal component from aboral view of E. australis. rpearson= -0.50, p < 0.05. Uppermost side of outlines is anterior. 117
Figure 46. Linear regression of length versus second principal component from aboral view of E. crispus. rpearson= 0.35, p < 0.05. Uppermost side of outlines is anterior. 118
Figure 47. Linear regression of length versus third principal component from aboral view of E. polyporus. rpearson= -0.73, p < 0.05. Uppermost side of outlines is anterior. 119
9.0
8.0 • • • m m • • 7.0 % • • • 6.0
5.0 • ■ m 4.0 • • m % m
3.0
2.0
1.0
0.0 -6
Figure 48. Linear regression of length versus third principal component from aboral view of E. provectus. rpearson= -0.34, p < 0.05. Uppermost side of outlines is anterior. 120
Figure 49. Linear regression of length versus third principal component from aboral view of E. pusillus. rpearson= 0.44, p < 0.05. Uppermost side of outlines is anterior. 121
8.0 • 7.0 • • 1 6.0 ‘ L A . m .... • • / • • 5.0 V • 4.0 • • ••r— ” ■ ~~j • • 1 . • 3.0 • : . j v • • <1 . 2.0 '
1.0
0.0 4 ' —— 1 — ij------j------1L.------■-----—- ■ f...... j ■ ------1 -6
Figure 50. Linear regression of length versus second principal component from aboral view of E. scaber. rpearson= -0.54, p < 0.05. Uppermost side of outlines is anterior. 122
8.0 • 7.0 rt'i^ ' “T '? '' ...... |.... 1 ^| ...... 1• • 6.0 9 m # m •# • I • • * * • 5.0 • ; W •\W~~ m 4.0 # i • • • m 3.0 * # • • 2.0
1.0 L . 0.0 r_ -3 _ -2 -1 0
Figure 51. Linear regression of length versus third principal component from aboral view of E. scaber. vpeaTSOn= -0.52, p < 0.05. Uppermost side of outlines is anterior. 123
12.0 [— ■■ —! ' 10.0 • • «) 8.0 m
t • • 6.0 U— — #J % • » • • • • 4.0 ______$ • __ m j • i> m § • • • m j• 2.0 #
0.0 -2 10
Figure 52. Linear regression of length versus first principal component from lateral view of E. crispus. rpearson= -0.45, p < 0.05. Right side of outlines is anterior.
10.0 • 9.0 L___
2.0
1.0
0.0 -6 -4 -2 8 10
Figure 53. Linear regression of length versus first principal component from lateral view of E. platytatus. rpearson= -0.61 ,P<0 .05. Right side of outlines is anterior. 124
9.0
8.0 ___ i I • i §
7.0 1 • L...... w ^ | • 6.0 • # w • • <1 • 5.0 __ ___ ft. j • • • • • ...jtu .... _ • ..3 ,.. • » : 4.0 • • \ 3.0 ..m ...... • 2.0
1.0
0.0 -6 -2
Figure 54. Linear regression of length versus second principal component from lateral view of E. provectus. rpearson= -0.42, p < 0.05. Right side of outlines is anterior.
12.0
10.0
8.0
•% • 6.0
4.0
2.0
0.0 -4
Figure 55. Linear regression of length versus third principal component from lateral view of E. pusillus. rpearson= 0.59, p < 0.05. Right side of outlines is anterior. 125
8.0 • 7.0 1 • • • 6.0 • i • • • •• .JL__^• • • • 5.0 • W • :j .■ ■1 r • • 4.0 • # 1 • 3.0 • t vw 1 • 2.0
1.0
0.0 -6 -4
Figure 56. Linear regression of length versus second principal component from lateral view of E. scaber. rpearson= 0.43, p < 0.05. Right side of outlines is anterior.
Figure 57. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. australis. rpearson= 0.49, p < 0.05. 126
12.0
10.0 • • • 8.0 L______...... #
• • 9 _ m 6.0 • r • • • • • • • 4.0 A • j...... , j ______...... ■ ^ .....______i • • i • • * « * 2.0 * *
0.0 T ~ " - - “ I — ' ' ~ — r — ------— 1— — - — ...... — ------— T f ------f — ------1 -8 -6 -4 -2
Figure 58. Linear regression of length versus first principal component from combined analysis of lateral and aboral views of E. crispus. rpearson= 0.52, p < 0.05.
6.0 • 5.0 ~~~~9 .. > .* • i m ^...... ; 4.0 9 W • : • • •i * • • • • j ^ ; ...... j 3.0 • ■ v * • • • • • 2.0
1.0
0.0 -4 -3 -2 -1
Figure 59. Linear regression of length versus third principal component from combined analysis of lateral and aboral views of E. elegans. rpearson= 0.53, p < 0.05. 127
10.0 9.0
8.0
7.0 • • i 6.0 5.0 4.0 3.0 •• « # 2.0
1.0
0.0 -6 -4 -2 0 2 4 6 8 10 12
Figure 60. Linear regression of length versus first principal component from combined analysis of lateral and aboral views of E. incertus. rpearson= 0.48, p < 0.05.
10.0 9.0 • 8.0 •
7.0 • 6.0 m •• 5.0 • 4.0 •• 3.0 z.u1 o 1.0 0.0 -4 -2 0 2 4 6 8 10 12
Figure 61. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. incertus. rpearson= -0.55, p < 0.05. 128
• • » • • • • • •
% ..
-4 -2 0 2 4 6 8 10
Figure 62. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. polyporus. rpearson= -0.65, p < 0.05.
9.0
8.0 • m • •
7.0 WWW • 6.0 v • •
5.0 • •• • 4.0 • m m
3.0 w^ .., .. i • 2.0
1.0 0.0 -8 -6 -4 -2
Figure 63. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. provectus. rpearson= -0.57, p < 0.05. 129
12.0
Figure 64. Linear regression of length versus second principal component from combined analysis of lateral and aboral views of E. pusillus. rpearson= 0.44, p < 0.05. 130
Laganum laganum Echinocyamus planissimus Sismondia occitana t Peronella peronii Echinocyamus pusillus Echinocyamus apicatus Echinocyamus convergens Echinocyamus elegans Echinocyamus incertus Echinocyamus platytatus Echinocyamus provectus Echinocyamus insularis Fibularia ovulum Fibularia dubarensis f Fibularia cribellum Leniechinus herricki f Fibularia kieri\ Echinocyamus petalus t Echinocyamus parviporus f Lenicyamidia compta t Cyamidia nummulitica t Echinocyamus crispus Echinocyamus megapetalus Mortonia australis Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus Echinocyamus macrostomus Figure 65. Strict consensus tree, showing relationships that were supported by all equally parsimonious trees. Daggers mark extinct taxa. 131
Laganum laganum Sismondia occitana t Peronella peronii 83 Echinocyamus planissimus Echinocyamus platytatus 83 Leniechinus herricki t Lenicyamidia compta f 50/100 53/100 Cyamidia nummulitica f 99 83 Fibularia dubarensis t 98 Fibularia cribellum I — 89' L Fibularia ovulum Fibularia kieri f 83 Echinocyamus parviporus t Echinocyamus elegans Echinocyamus petalus f Echinocyamus pusillus 83 58 Echinocyamus apicatus Echinocyamus crispus 100 Echinocyamus megapetalus B 83 Mortonia australis Echinocyamus insularis 65 Echinocyamus incertus Echinocyamus convergens ^ C E Echinocyamus provectus Echinocyamus sollers 100 Echinocyamus scaber i J — T ~ Echinocyamus grandiporus D 1001— 1_ Echinocyamus macrostomus
Figure 66. Majority rule consensus tree. Black numbers next to nodes show the percentage of trees that contained this branching pattern. Red numbers show the bootstrap support. Nodes without bootstrap support in the figure had less than 50% support. Daggers mark extinct taxa. Letters mark named clades within Echinocyamus. 132
- * Laganum laganum •• Sismondia occitana f -+ Peronella peronii “• Echinocyamus planissimus "• Echinocyamus platytatus Leniechinus herricki f Lenicyamidia compta f Cyamidia nummulitica f Fibularia dubarensis t Fibularia cribellum Fibularia ovulum Fibularia kieri f Echinocyamus parviporus f Echinocyamus elegans Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus Max test height Echinocyamus crispus Echinocyamus megapetalus B Central Mortonia australis Anterior Echinocyamus insularis Posterior Echinocyamus incertus Unknown Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocya mus gra ndiporus D Echinocyamus macrostomus
Figure 67. Character 1: location of maximum test height. Red = maximum test height is anterior; blue = maximum test is anterior; green = maximum test height is posterior; grey = location of maximum test height is unknown. 133
“• Laganum lagatium -• Sismondia occitana t -• Peronella peronii “• Echinocyamus planissimus Echinocyamus platytatus “• Leniechinus herricki t Lenicyamidia compta f Cyamidia nummulitica t Fibularia dubarensis t Fibularia cribellum Fibularia ovulum Fibularia kieri t Echinocyamus parviporus t “• Echinocyamus elegans “• Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus A Internal partitions Echinocyamus crispus Echinocyamus megapetalus B Ten • U-CE Mortonia australis Two • Echinocyamus insularis None • Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers • Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 68. Character 2: internal radial partitions. Red = ten internal radial partitions; blue = two internal radial partitions; green = no internal radial partitions. 134
• Laganum laganum • Sismondia occitana t • Peronella peronii • Echinocyamus planissimus • Echinocyamus platytatus • Leniechinus herricki f Lenicyamidia compta t Cyamidia nummulitica t Fibularia dubarensis f • Fibularia cribellum Fibularia ovulum Fibularia kieri f Echinocyamus parviporus f Echinocyamus elegans “• Echinocyamus petalus t Echinocyamus pusillus Echinocyamus apicatus Circumferential Echinocyamus crispus partitions Echinocyamus megapetalus B - p - c Mortonia australis Present • 4 Absent • Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Ech inocya mus macros to m us
Figure 69. Character 3: circumferential partitions. Green = circumferential partitions are present; red = circumferential partitions are absent. 135
* Laganum laganum * Sismondia occitana t * Peronella peronii * Echinocyamus planissimus * Echinocyamus platytatus m Leniechinus herricki f Lenicyamidia compta t Cyamidia nummulitica t I------Fibularia dubarensis f » i * Fibularia cribellum Fibularia ovulum Fibularia kieri f "* Echinocyamus parviporus f “• Echinocyamus elegans “• Echinocyamus petalus t Echinocyamus pusillus Echinocyamus apicatus Periproctal spines Echinocyamus crispus Echinocyamus megapetalus B Present • Mortonia australis Absent • Echinocyamus insularis Unknown • Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus ■ EchinocyamusE sollers Echinocyamus scaber L i E D i i— chinocyamus grandiporus i— Echinocyamus macrostomus
Figure 70. Character 4: periproctal spines. Red = periproctal spines present; green = periproctal spines absent; gray = presence of periproctal spines unknown. •J
136
Laganum laganum ■* Sismondia occitana f -• Peronella peronii "• Echinocyamus planissimus Echinocyamus platytatus “• Leniechinus herricki f Lenicyamidia compta f Cyamidia nummulitica t Fibularia dubarensis t * Fibularia cribellum Fibularia ovulum Fibularia kieri t
" • Echinocyamus parviporus f Echinocyamus elegans Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus Periproctal plates Echinocyamus crispus Echinocyamus megapetalus B >20 HT Mortonia australis <20 Echinocyamus insular is Unknown Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Echinocyam us tnacrostom us
Figure 71. Character 5: number of plates in periproctal membrane. Green = more than twenty plates in the periproctal membrane; red = less than twenty plates in the periproctal membrane; gray = number of perproctal plates unknown. •+ Laganum laganum Sismotidia occitana f •» Peronella peronii “• Echinocyamus planissimus *+ Echinocyamus platytatus “• Leniechinus herricki t Lenicyamidia cotnpta f Cyamidia nummulitica t Fibularia dubarensis f Fibularia cribellum Fibularia ovulum Fibularia kieri f Echinocyamus parviporus f Echinocyamus elegans Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus Podial pores Echinocyamus crispus Echinocyamus megapetalus B Along sutures - K £ Mortonia australis Not along sutures Echinocyamus insularis Unknown Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Echinocyarnus macrostomus
Figure 72. Character 6: accessory podial pores along sutures. Red = podial pores not concentrated along plate sutures; green = podial pores concentrated along plate sutures; gray podial pore location unknown. 138
“• Laganum laganum •* Sismondia occitana f - * Peronella peronii "* Echinocyamus planissimus "• Echinocyamus platytatus ■* Leniechinus herricki t *Lenicyamidia compta f »Cyamidia nummulitica t Fibularia dubarensis f Fibularia cribellum Fibularia ovulum Fibularia kieri f Echinocyamus parviporus f Echinocyamus elegans Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus Test height Echinocyamus crispus Echinocyamus megapetalus B Flat • L - t c Mortonia australis High • Echinocyamus insularis Globular* Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Ech inocya rn us macros to m us
Figure 73. Character 7: ratio of test height versus length. Red = flat; green = high; blue = globular. - • Laganum laganum - • Sismondia occitana f -• Peronella peronii "• Echinocyamus planissimus "• Echinocyamus platytatus “• Leniechinus herricki t Lenicyamidia compta f C Cyamidia nummulitica t Fibularia dubarensis t Fibularia cribellum Fibularia ovulum Fibularia kieri f ■* Echinocyamus parviporus f "• Echinocyamus elegans Echinocyamus petalus t Echinocyamus pusillus Echinocyamus apicatus # of petaloid pores Echinocyamus crispus Echinocyamus megapetalus > 100 - M - c Mortonia australis 50-100 Echinocyamus insularis < 50 Echinocyamus incertus Echinocya m us con vergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus Echinocyamus macrostomus
Figure 74. Character 8: number of petaloid pore pairs. Blue = many pore pairs; green moderate pore pairs; red = few pore pairs. 140
Laganum laganum “• Sismondia occitana t •+ Peronella peronii "• Echinocyamus planissimus “• Echinocyamus platytatus “* Leniechinus herricki f Len icy a m idia compta t Cyamidia nummulitica t Fibularia dubarensis f Fibularia cribellum Fibularia ovulum Fibularia kieri t Ech inocya m us pa rviporus f Echinocyamus elegans Echinocyamus petalus t Echinocyamus pusillus Echinocyamus apicatus Coverage of test Echinocyamus crispus by petals Echinocyamus megapetalus B >70% • LHCE Mortonia australis 40-70% • Echinocyamus insularis <40% • Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D ^ 5 Echi nocya in us macrostomus
Figure 75. Character 9: coverage of test by petals. Blue = high coverage of test by petaloids; green = moderate coverage of test by petaloids; red = low coverage of test by petaloids. 141
— • Laganum laganum — • Sismondia occitana t — —• Peronella peronii Echinocyamus planissimus Echinocyamus platytatus — * Leniechinus herricki t Lenicyamidia compta f C Cyamidia nummulitica t I...... • Fibularia dubarensis t | J * Fibularia cribellum I r * Fibularia ovulum Fibularia kieri t * Echinocyamus parviporus t * Echinocyamus elegans * Echinocyamus petalus f Echinocyamus pusillus C Echinocyamus apicatus »» Length v. width | ...... 5 Echinocyamus crispus I i— Echinocyamus megapetalus g Subcircular* 1— Mortonia australis Elongate # " Echinocyamus insularis - Echinocyamus convergens Echinocyamus provectus . mEchinocyamus sollers —4 Y——•Echinocyamus scaber l—4 — mEchinocyamu$ grandiporus 1—« Echinocyatnus macrostomus
Figure 76. Character 10: ratio of test length to width. Red = subcircular; green = elongate. 142
■ * Laganum laganum - • Sismondia occitana t • » Peronella peronii ’* Echinocyamus planissimus Echinocyamus platytatus wm Leniechinus herricki t Lenicyamidia compta t C Cyamidia nummulitica f Fibularia dubarensis t Fibularia cribellum Fibularia ovulum Fibularia kieri f Echinocyamus parviporus t "• Echinocyamus elegans 'Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus Petal shape Echinocyamus crispus Echinocyamus megapetalus B Convergent* L— Mortonia australis Parallel Echinocyamus insular is Divergent • Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers * Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 77. Character 11: shape of petaloid pore pair series. Blue = convergent; green = parallel; red = divergent. 143
Laganum laganum Sismondia occitana f ■ * Peronella peronii “* Echinocyamus planissimus "• Echinocyamus platytatus Leniechinus herricki f Lenicyamidia compta f Cyamidia nummulitica t Fibularia dubarensis f Fibularia cribellum Fibularia ovulum Fibularia kieri t Echinocyamus parviporus t Echinocyamus elegans Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus Infundibulum Echinocyamus crispus depth Echinocyamus megapetalus B L m = c Mortonia australis Slight • Echinocyamus insular is None • Echinocyamus incertus Deep • Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 78. Character 12: depth of infundibulum. Green = slight infundibulum; red = no infundibulum; blue = deep infundibulum. 144
“• Laganum laganum — Sismondia occitana t - • Peronella peronii Lf ■* Echinocyamus planissimus LI Echinocyamus platytatus Leniechinus herricki t L, r —* Len icy a m idia compta f *—+Cyamidia nummulitica t Fibularia dubarensis t • Fibularia cribellum Fibularia ovulum Fibularia kieri t "• Echinocyamus parviporus f Echinocyamus elegans "*Echinocyamus petalus t Echinocyamus pusillus Echinocyamus apicatus Peristome shape Echinocyamus crispus I— 1- Echinocyamus megapetalus B Round • Mortonia australis Pentagonal • Echinocyamus insularis Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers • Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 79. Character 13: shape of peristome. Red = round peristome; green = rounded pentagonal peristome. 145
“• Laganum laganum Sismondia occitana f •+ Peronella peronii ~* Echinocyamus platiissimus "• Echinocyamus platytatus “• Leniechinus herricki f Lenicyamidia compta t Cyamidia nummulitica f Fibularia dubarensis f Fibularia cribellum Fibularia ovulurn Fibularia kieri f lEchinocyamus parviporus f Echinocyamus elegans Echinocyamus petalus t Echinocyamus pusillus Echinocyamus apicatus A Peristome size Echinocyamus crispus Echinocyamus megapetalus B Small • Mortonia australis Medium Echinocyamus insularis Large • Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 80. Character 14: peristome size. Red = small peristome; green = medium peristome; blue = large peristome 146
* • Laganum laganum Sismondia occitana t Peronella peronii "• Echinocyamus planissimus Echinocyamus platytatus ’* Leniechinus herricki f ~ L Lenicyamidia compta f Cyamidia nummulitica t L Fibularia dubarensis f Fibularia cribellum Fibularia ovulum Fibularia kieri t \Echinocyamus parviporus f \Echinocyamus elegans \Echinocyamus petalus t Echinocyamus pusillus Echinocyamus apicatus # hydro pores Echinocyamus crispus Echinocyamus megapetalus B >10 • Mortonia australis 2-10 • Echinocyamus insularis Echinocyamus incertus One • Echinocya m us con vergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 81. Character 15: number of hydropores. Blue = more than 10 hydropores; Green = 2-10 hydropores; red = one hydropore. 147
- — • * Laganum laganum — —• Sismondia occitana t — * Peronella peronii ~~~* Echinocyamus planissimus Echinocyamus platytatus — * Leniechinus herricki f Lenicyamidia compta f C Cyamidia nummulitica t I 1 Fibularia dubarensis t I \...... * Fibularia cribellum l i~~* Fibularia ovulum * Fibularia kieri t * Echinocyamus parviporus t * Echinocyamus elegans * Echinocyamus petalus f Echinocyamus pusillus C Echinocyamus apicatus Hydropore groove B Present* Absent •
...... • Echinocyamus sollers —4 ■ mEchinocyamus scaber ^ j-—»Echinocyamus grandiporus 1—mEchinocyamus macrostomus
Figure 82. Character 16: hydropore groove. Green = groove present; red = groove absent. 148
■ * Laganum laganum •• Sismondia occitana t -• Peronella peronii "* Echinocyamus planissimus "• Echinocyamus platytatus Leniechinus herricki t 'Lenicyatnidia compta t *Cyamidia nummulitica f Fibularia dubarensis f Fibularia cribellum - L j — Fibularia ovulum M i Fibularia kieri f "• Echinocyamus parviporus t Echinocyamus elegans • Echinocyamus petalus f —* Echinocyamus pusillus Echinocyamus apicatus IPZ width vs. PZ Echinocyamus crispus Echinocyamus megapetalus B IPZ > PZ • - < c Mortonia australis IPZ < PZ • Echinocyamus insularis Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Ech inocya m us macrostomus
Figure 83. Character 17: width of interporiferous zone (IPZ) relative to poriferous zone (PZ). Red = interporiferous zone wider than poriferous zone; green = interporiferous zone equal in width or narrower than poriferous zone. 149
■ * Laganum laganum - • Sismondia occitana t -* Peronella peronii Echinocyamus planissimus "• Echinocyamus platytatus “* Leniechinus herricki t ’Lenicyamidia compta i 1Cyamidia nummulitica t Fibularia dubarensis t Fibularia cribellum Fibularia ovulum Fibularia kieri t Echinocyamus parviporus f Echinocyamus elegans Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus Glassy tubercles Echinocyamus crispus Echinocyamus megapetalus B Small • Mortonia australis Large • Echinocyamus insularis None • Echinocyamus incertus Unknown • Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 84. Character 18: glassy tubercles. Red = smaller glassy tubercles; green = larger glassy tubercles; blue = no glassy tubercles; gray = presence of glassy tubercles unknown. 150
* Laganum laganum * Sismondia occitana t * Peronella peronii L * Echinocyamus planissimus * Echinocyamus platytatus * Leniechinus herricki f mLenicyamidia compta t '•Cyamidia nummulitica t Fibularia dubarensis f * Fibularia cribellutn Fibularia ovulum Fibularia kieri t Echinocyamus parviporus f Echinocyamus elegans Echinocyamus petalus t Echinocyamus pusillus Echinocyamus apicatus A Periproct shape Echinocyamus crispus Echinocyamus megapetalus B Circular • - t c Mortonia australis Elongate * Echinocyamus insularis Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 85. Character 19: periproct shape. Red = circular periproct; green = anterior-posteriorly elongate periproct. 151
“• Laganum laganum - • Sismondia occitana t -• Peronella peronii * Echinocyamus planissitnus * Echinocyamus platytatus * Leniechinus herricki t Lenicyamidia compta t C Cyamidia nummulitica f Fibularia dubarensis t Fibularia cribellum Fibularia ovulum Fibularia kieri t Echinocyamus parviporus t \Echinocyamus elegans \Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus Ocular pore size Echinocyamus crispus Echinocyamus megapetalus B Small • Mortonia australis Large • Echinocyamus insularis Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 86. Character 20: relative size of ocular and genital pores. Red = ocular pores small; green = ocular pores as large as genital pores. 152
- • Laganum laganum " • Sismondia occitana t ■ * Peronella peronii *• Echinocyamus planissimus Echinocyamus platytatus “• Leniechinus herricki f Lenicyamidia compta t Cyamidia nummulitica t Fibularia dubarensis f Fibularia cribellum Fibularia ovulum Fibularia kieri t \Echinocyamus parviporus t lEchinocyamus elegans \Echinocyamus petalus f Echinocyamus pusillus Echinocyamus apicatus Ambital inflation Echinocyamus crispus Echinocyamus megapetalus B Present • h SMortonia australis Absent • Echinocyamus insularis Echinocyamus incertus Echinocyamus convergens Echinocyamus provectus Echinocyamus sollers Echinocyamus scaber Echinocyamus grandiporus D Echinocyamus macrostomus
Figure 87. Character 21: inflation of ambital region. Green = inflation present; red = inflation absent. 153
# E. apicatus • E. pusillus
Figure 88. Biogeography of species in Clade A.
australis % E. crips us # E. megapetalus
Figure 89. Biogeography of species in Clade B. 154
• e provectus # E. incertus • *=• convergens
Figure 90. Biogeography of species in Clade C.
Figure 91. Biogeography of species in Clade D.