In defence of the calcichordates

The cornutes and mitrates, collectively known as stylophorans or calcichordates, are the most controversial members of the traditional group of the 'carpoid echinoderms'. In a recent review, Philip has rejected Ubaghs' interpretation of them as aberrant echinoderms with a single feeding arm, has also rejected my own interpretation of diem as primitive chordates, and has made some tentative proposals of his own. The present paper answers Philip's criticisms of the calcichordate interpretation and discusses his own proposals and his comments on Ubaghs' views.     

Life on the seafloor: adaptations and strategies in Stylophora (Echinodermata)

Stylophorans are a Palaeozoic group of non‐pentamerous echinoderms, morphologically well‐adapted to a benthic mode of life on soft sediment seafloors. By developing a thin and wide theca, they successively increased the surface area in contact with the substrate resulting in an even distribution of the body mass resting on the ground, and efficiently preventing the body from sinking into non‐indurated sediments (snowshoe strategy). In stylophorans, the body surface is dramatically increased in result of the expansion of two integumentary areas on the lower thecal surface. While infracentral areas are reduced in primitive forms to the detriment of massive marginals; in boot‐shape cornutes, infracentral areas are larger, polyplated and framed by very delicate marginals. In heart‐shaped cornutes, the reduction of both infracentral areas is compensated by the development of spiny elements on the theca outline. In forms where the degree of bilateral symmetry is high, the left infracentral area is larger than the right area, resulting in an elongated thecal shape. In structural geology, changes of shape of a rock submitted to a strain can be categorized by translating the obtained deformation into a strain ellipsoid plotted on a Flinn diagram, while in biology changes of shapes of organisms are traditionally measured through morphometric analysis. By applying Flinn's principle of strain ellipsoid to biological objects, the present study aims to characterize different life adaptations across stylophorans, observing changes of shape of both infracentral areas interpreted as two ellipsoids. Once plotted on a Flinn diagram, three significantly separated clusters are observed when focusing on the left area. This concerns forms with reduced infracentral areas, highly and weakly asymmetrical forms. According to these results, three new morphological adaptations are described (water strider, flat fish and stream‐lined body). These newly described adaptations enabled snowshoe strategist stylophorans to remain stable on top of the seafloor.     

Brief review of the stylophoran debate

The "calcichordate" theory interprets an extinct group of calcite-plated invertebrates, the stylophorans, as chordates. In this theory, cornute stylophorans are interpreted as stem chordates, whereas mitrate stylophorans are primitive members of the acraniates, tunicates, and craniates. However, this theory discounts major synapomorphies between cornutes and mitrates. These groups constitute a natural, monophyletic group which is here argued to lie within the echinoderm radiation. The "calcichordate" theory is, therefore, rejected because it relies on assumption-driven hypotheses of character transformation which are supported by ambiguous, poor, or missing fossil evidence. Stylophorans may lie at the base of the echinoderm clade and primitively lack pentameral symmetry, therefore casting light on the near-ancestral body organization of the phylum.     

A new calcichordate from the Ordovician of Bohemia and its anatomy, adaptations and relationships

The paper describes a new member of a group of Lower Palaeozoic marine fossils which partly bridge the gap between echinoderms and chordates. Evidence suggests that this group included the ancestors of the vertebrates. Its members are traditionally regarded as primitive echinoderms, but are better seen as primitive chordates with echinoderm affinities. They form a basal subphylum of chordates-the Calcichordata Jefferies 1967. The Calcichordata, in accordance with an early suggestion by Gislén, are probably ancestral to all living chordates. The new calcichordate is named Reticulocarpos hanusi gen et sp. nov. It comes from the Lower Ordovician ?árka Formation (Llanvirn) of ?árka near Prague, Czechoslovakia and is placed in the family Amygdalothecidae Ubaghs 1970. It is important because of its position in the Calcichordata. This group is divided into two very different orders–the Cornuta and the Mitrata. The Cornuta are the more primitive order and gave rise to the Mitrata, which had the structure of giant, calcite-plated tunicate tadpoles. Many features show that the new species is a very advanced cornute, closely related to the stock that gave rise to the mitrates. For this reason it is important in the general history of the chordates, since some primitive mitrate was probably the latest common ancestor of the living chordate subphyla i.e. of tunicates, of amphioxus and its allies and of the vertebrates. Being a mitrate-like cornute, the new species allows the cornutes and mitrates to be compared more confidently than before. Four results are especially important. Firstly it is likely that the stem (=tail) of mitrates is equivalent only to the anterior part of the stem of cornutes. This is significant, because traditional views as to which was the upper surface in mitrates have been based on stem homologies now seen as false. Secondly Reticulocarpos hanusi is adapted to stay up on very soft mud, using only the strength of the mud for support. The mitrates, on the other hand, supported themselves on soft mud by a much more reliable method resembling buoyancy. Thirdly, the new form had paired transpharyngeal eyes which are otherwise known only in mitrates, and which are the earliest type of paired eyes in chordates. Fourthly, it becomes possible to homologize the thecal plates of cornutes with those of mitrates. Reticulocarpos hanusi represents an important phase in chordate evolution dominated by the necessity of staying up on mud by a very precarious method. During this phase many pre-adaptations for swimming were acquired. Primitive mitrates, descended from a very similar form, were probably the first chordates that could swim.     

A cladistic analysis of the cornutes (stem chordates)

At the root of the calcichordate controversy is the problem of recognizing homologous similarity. Only when a criterion or set of criteria for hypothesizing homologies is established can the calcichordate theory be tested against its main rival—the Stylophoran theory—by applying the principle of parsimony. The criterion of topological similarity is applied to a three-laxon problem involving a mitrate, the crown-group Echinodermata and the crown-group Chordata. The mitrate shares more features with extant chordates than with extant echinoderms and the calcichordate theory is supported by a simple parsimony analysis. In a cladistic analysis of the cornutes, four monophyletic families are recognized (Cothurnocystidae, Scotiaecystidae, Phyllocystidae and Hanusiidae) and their interrelationships resolved using Hennig86 and the successive-weighting procedure of Farris. Because the known fossil record of cornutes and mitrates is very poor, the correlation between the cladogram produced for cornutes and their order of appearance in the geological record is weak. All of the cornute families must have originated in the Lower Cambrian at the latest.     

A new stem-group chordate (Cornuta) from the Llandeilo of Czechoslovakia and the cornutemitrate transition

A new stem chordate, Prokopiytis mergli, from the Middle Ordovician (Llandeilo) of Bohemia, Czechoslovakia is reconstructed and described. It proves to be the most crownward cornute yet known, sharing with the mitrates (primitive crown chordates) a number of derived features unknown in other cornutes. Such features include: the position of the gonopore-anus, the absence of major dorsal plates from the fore tail and the expansion of the marginal plates of the head onto the dorsal surface. Some other characteristics of P. mergli are a fore tail composed of only three segments and an elongate head. Only three hind-tail ossicles are known. The elongate head and small number of fore-tail segments are autapomorphies of P. mergli. The locomotory cycle is reconstructed and it is concluded that P. mergli moved rearwards, pulled largely by its mid and hind tail. The calcichordate theory is defended and the orientation of the mitrates considered. P. mergli reinforces the view that the convex surface of the mitrates was ventral in life. The origin of the atrial chambers of mitrates is discussed and a new homology proposed between the anterior ventral fore-tail plates of cornutes and what were formerly called plates g and j of mitrates. Nový chordát z kmenové skupiny/stem-group/Cornuta zeského Ilandeila, pechod mezi skupinami Cornuta a Mitrata. V práci je popsán a rekonstruován nový zástupce kmene chordátu Prokopicystis mergli gen. et sp. n., z eského llandeila/eskoslovensko/. Jde o dosud známého nejvýe organizovaného kornuta, vykazukícíhoadu znaku shodných s chordáty mitrátníimi a zcela neznámýc u ostatních pedstavitelå skupiny Cornuta. Tyto znaky zahrnují: pozici gonopor-análníiho otvoru, nepítomnost hlavních dorzálních desek v proximálníásti ocasu a expanzi marginálníc desek hlav na dorzální stranu. K dalím charakteristickým znakåum druhu P. mergli patí protáhlá hlava a proximálníást ocasu tvoená jen temi segmenty. V distálníásti ocasu jsou rovn známy jen 3 destiky. Protáhá halva a maly pocet segmentåu proximálníásti ocasu jsou u. P. mergli autapomorfiemi. V práci je také rekonstruován pohybový cyklus ivoicha se závrem, e P. mergli se pohyboval dozadu, pozvolným tahem stední a zadníáSTI OCASU> Je zde hájena tzv. kalcichordátová teorie a diskutována orientace mitrátåu. VyAsledky studia podporují domnnku, e skupiny Mitrata byla za ivota konvexní strana jejich theky stranou ventrální. Je diskutován påuvod atriálnich koma?rek skupiny Mitrata. Souasn jsou pova ovány pední ventrálni desek pedníásti za homologicke s desek díve oznaovanými jako g a j u mitr     

Morphometric analysis of Tremadocian (earliest Ordovician) kirkocystid mitrates (Echinodermata, Stylophora) from the Taebaeksan Basin, Korea

Abundant isolated remains of stylophoran echinoderms (cornutes and mitrates) are reported for the first time in the late Tremadocian (Asaphellus Zone) Tumugol Formation of Korea. Mitrate remains include numerous adorals of Kirkocystidae. Several new important anatomical features have been observed on these adorals, as an internal calcitic layer that is associated to s₂ and possibly also to the palmar complex. This observation suggests that the palmar complex would be present not only in mitrocystitid mitrates, but also in peltocystitids. For the first time, several morphometric analyses have been undertaken based on isolated kirkocystid adorals, so as to explore the morphological diversity displayed by Korean adorals, but also in order to compare their morphology with that of other Gondwanan kirkocystids. Morphometric analyses indicate the occurrence of two contrasted morphologies within Korean adorals (morphotypes A and B), and of three distinct morphologies within European and North African forms (“Anatifopsis”, “Balanocystites”, and “escandei” morphotypes). Comparison of Korean adorals with those from Europe and North Africa shows that: (1) morphotypes B and “Anatifopsis” are equivalent; consequently, the two Korean specimens referred to morphotype B are assigned to the genus Anatifopsis; (2) morphology of most Korean adorals, which belong to morphotype A, is clearly distinct from that of all other described kirkocystids from Europe and North Africa. The small size, juvenile morphology, and great morphological variability observed in the morphotype A of the Korean adorals are suggestive of possible heterochronic processes (peramorphosis).     

Epizoan encrustation on Marsupites testudinarius – additional argument favoring an epifaunal mode of life of uintacrinoids?

Uintacrinoids (Uintacrinoidea Zittel) are among the best-known Late Cretaceous crinoids, but owing to their unusual morphology and widespread distribution their mode of life has become a subject of much discussion. Of several competing hypotheses, nektonic, pseudoplanktonic, hemipelagic, semi-infaunal and epibenthic lifestyles have been suggested. Recent study synthesizing and extending previous data has shown that these crinoids were epibenthic, holding their arms vertically for feeding. However, evidence supporting a rheophilic epibenthic model over an alternative rheophobic semi-infaunal model is still limited. Here we report epizoans, mostly represented by serpulids and bryozoans, encrusting thecal plates of Marsupites testudinarius from the Lägerdorf in Germany. Although a definitive interpretation whether recorded infestations occurred syn vivo or post mortem is not certain, it is remarkable that all epizoans (or their traces) are attached to the convex side (latera) of well-preserved isolated plates displaying no signs of reworking. Furthermore, a bryozoan colony crossing plate boundaries has been also found on the surface of a sub-articulated theca suggesting that it may have been colonised syn vivo. Recorded epibiotic associations, whether syn vivo or post-mortem, must have developed prior to burial of the specimens, above the surface of sea floor, and thus provide another argument against rheophobic semi-infaunal mode of life of uintacrinoids.     

Evolution of bilaterally asymmetrical larvae in freshwater mussels (Bivalvia: Unionoida: Unionidae)

Bilaterally asymmetrical glochidia (i.e. bivalved parasitic larvae bearing a large marginal appendage on a single valve) have been reported from five Asian freshwater mussel genera belonging to two separate subfamilies, the Gonideinae (i.e. Pseudodon, Solenaia, and Physunio) and Rectidentinae (i.e. Contradens and Trapezoideus). This classification requires that the bilaterally asymmetrical glochidium‐bearing mussels are not monophyletic, and suggests that this atypical larval morphology evolved twice in the same geographic region. Although homoplastic glochidium characters are known (e.g. marginal appendages and size), we hypothesized that bilaterally asymmetrical glochidia represent a novel morphological synapomorphy. We tested the monophyly of the mussels bearing bilaterally asymmetrical glochidia using a molecular matrix consisting of representatives from all six freshwater mussel families and three molecular markers (28S, 16S, and COI). Bayesian inference, maximum likelihood, and ancestral state reconstruction were employed to estimate the phylogeny and larval trait transformations. The reconstructed phylogeny rejects the monophyly of the asymmetrical glochidium‐bearing mussels and resolves two putative origins of asymmetrical glochidia; however, ancestral state reconstruction supports asymmetrical glochidia as a synapomorphy of only one supraspecific taxon of the Rectidentinae. In the Gonideinae, asymmetrical glochidia were autapomorphic of Pseudodon cambodjensis (Petit, 1865). That is, no other taxa resolved among the Gonideinae had bilaterally asymmetrical glochidia, including other Pseudodon species. We describe how the alleged intrageneric glochidial variation in Pseudodon, and in the other genera of the Gonideinae reported to have asymmetrical glochidia (i.e. Solenaia and Physunio), challenge the resolved convergence of asymmetrical glochidia. Our results are discussed in the context of freshwater mussel larval evolution, patterns in life‐history traits, and the classification of freshwater mussels generally. © 2015 The Linnean Society of London     

Effects of shell shape,size, and sculpture in burrowing and anchoring abilities in the freshwater mussel Potamilus alatus (Unionidae)

Because of the sedentary lifestyle of freshwater mussels, studies examining their movement capabilities are scarce. However, the ability to burrow into the substrate and the ability to remain stationary are likely crucial components of their behavioural repertoire. The performance of these different tasks is likely to be affected by the presence of the shell ornamentation characteristic of many mussel species. Previous studies have suggested that shell ornamentation results in a trade‐off between burrowing ability and remaining stationary when an extrinsic force attempts to dislodge it from the substrate once buried. We examined the effect of morphology and shell ornamentation on burrowing performance and anchoring ability by artificially creating shell ornamentation on a relatively smooth‐shelled species (Potamilus alatus). Burrowing behaviours and performance and the force required to dislodge mussels (anchoring ability) were quantified with and without ornamentation. Interestingly, we found that the artificial shell ornamentation had no significant effect on burrowing behaviours and performance or dislodgement force. Burrowing and dislodgement, however, were both highly influenced by shell size and shape. All of the available information suggests that shell size, shape, and sculpture influence burrowing and anchoring in complex ways that needs further examination. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 136–144.     

Elucidating Ernietta: new insights from exceptional specimens in the Ediacaran of Namibia

Exceptionally preserved specimens of Ernietta in a shallow‐marine gutter cast from southern Namibia reveal that all previously figured specimens of this iconic Ediacaran megafossil are incomplete, representing only the base of a larger and more complex organism. The complete organism is interpreted as comprising a buried, sand‐filled anchor exhibiting the classical Ernietta morphology that passes distally into a trunk that is crowned by two facing fans that extended into the overlying water column. All parts of Ernietta, from the base of its buried anchor to the tip of its fans, appear to have been composed of a palisade of tubular elements that have been variably preserved. Similarity of tubule morphology despite the inherent difference in function between these constructions supports the view that these tubes were integral to all anatomical parts and functions of Ernietta. This style of architecture, construction and function is unique to the Erniettomorpha, supporting the view that it represents an extinct Ediacaran clade in the early evolution of multi‐cellular life.     

Taxonomic significance of asymmetrical helmet and lance bristles in the genus Mallomonas (Synurophyceae) and their discovery in Eocene lake sediments

Complex bristle types formed by species in the genus Mallomonas include those with helmet or lance-shaped apices. The ornamentation on each side of the helmet has been thought to be equivalent or symmetrical, whereas on a lance-shaped bristle an expanded portion folds over one side of the shaft to form an asymmetrical structure. We describe, for the first time, helmet bristles with a distinctly asymmetrical design, also formed by the folding of a siliceous membrane over one side of the helmet. We postulate that the asymmetrical helmet represents a structure that combines the formation of a symmetrical helmet and a lance-shaped design on the same bristle. Further, we report structurally similar asymmetrical helmet bristles, lance-shaped bristles and scales that are unambiguously assigned to Mallomonas asmundiae in Middle Eocene sediments from a maar lake in northern Canada, supporting the hypothesis that scale and bristle morphology in the Synurophyceae has undergone extensive prolonged evolutionary stasis. Given differences in scale morphology and the presence of asymmetrical helmet bristles, we transfer the North American endemic Mallomonas acaroides var. muskokana to the rank of species. Further, we formally describe Mallomonas dispar and M. lancea, fossil species with asymmetrical helmet bristles and lance-shaped bristles, respectively. The taxonomic and biogeographic significance of asymmetrical and lance-bearing bristles is discussed.     

ICHNOLOGICAL INSIGHTS INTO MITRATE PALAEOBIOLOGY

Abstract:  Mitrates are a controversial group of extinct deuterostomes; there is little agreement over their affinities, functional morphology or even the orientation of their upper and lower surfaces. Four slabs of slate from the Lower Devonian Hunsrück Slate (Bundenbach, Germany) are here described, showing trace fossils ( Vadichnites transversus igen. et isp. nov.) associated with the mitrate Rhenocystis latipedunculata . These new findings clearly demonstrate that the mitrate appendage was used in locomotion and that this movement took place appendage-first. Such a functional interpretation suggests that mitrates were oriented with the flat body surface upwards in life and argues against a phylogenetic position in the echinoderm crown-group.     

Ontogenetic changes in habitat use by whitefish,Coregonus lavaretus

Synopsis The whitefish, Coregonus lavaretus, in the lake Mj?sa exhibited two niche changes during their life cycle. Juveniles (< 25 cm body length) were confined to the shallow (0–30 m) epibenthic zone. Medium sized whitefish (25–35 cm body length) expanded their habitat use to include the deep (30–90 m) epibenthic zone as well as the pelagic zone. From a body length of 35 cm, habitat use was restricted to the deep epibenthic zone. Small fish in the shallow epibenthic zone ate small and medium-sized prey (zooplankton, insect larvae and surface insects). Medium-sized fish in this zone were in addition feeding on the larger amphipod, Pallasea quadrispinosa. In the pelagic zone, the diet of medium-sized whitefish was dominated by zooplankton, although some larger prey like surface insects and age-0 smelt, Osmerus eperlanus, were also eaten. In the deep epibenthic zone, the diet of both medium-sized and large (< 35 cm) whitefish consisted mainly of the large prey P. quadrispinosa.     

Exceptionally preserved soft parts in fossils from the Lower Ordovician of Morocco clarify stylophoran affinities within basal deuterostomes

The extinct echinoderm clade Stylophora consists of some of the strangest known deuterostomes. Stylophorans are known from complete, fully articulated skeletal remains from the middle Cambrian to the Pennsylvanian, but remain difficult to interpret. Their bizarre morphology, with a single appendage extending from a main body, has spawned vigorous debate over the phylogenetic significance of stylophorans, which were long considered modified but bona fide echinoderms with a feeding appendage. More recent interpretation of this appendage as a posterior “tail-like” structure has literally turned the animal back to front, leading to consideration of stylophorans as ancestral chordates, or as hemichordate-like, early echinoderms. Until now, the data feeding the debate have been restricted to evaluations of skeletal anatomy. Here, we apply novel elemental mapping technologies to describe, for the first time, soft tissue traces in stylophorans in conjunction with skeletal molds. The single stylophoran appendage contains a longitudinal canal with perpendicular, elongate extensions projecting beyond hinged biserial plates. This pattern of soft tissues compares most favorably with the hydrocoel, including a water vascular canal and tube feet found in all typical echinoderms. Presence of both calcite stereom and now, an apparent water vascular system, supports echinoderm and not hemichordate-like affinities.     

DIRECT AND INDIRECT SEXUAL SELECTION AND QUANTITATIVE GENETICS OF MALE TRAITS IN GUPPIES (POECILIA RETICULATA)

Abstract.— The ornamentation and displays on which sexual attractiveness and thus mating success are based may be complex and comprise several traits. Predicting the outcome of sexual selection on such complex phenotypes requires an understanding of both the direct operation of selection on each trait and the indirect consequences of selection operating directly on genetically correlated traits. Here we report the results of a quantitative genetic analysis of the ornamentation, sexual attractiveness, and mating success of male guppies (Poecilia reticulata). We analyze male ornamentation both from the point of view of single ornamental traits (e.g., the area of each color) and of composite measures of the way the entire pattern is likely to be perceived by females (e.g., the mean and contrast in chroma). We demonstrate that there is substantial additive genetic variation in almost all measures of male ornamentation and that much of this variation may be Y linked. Attractiveness and mating success are positively correlated at the phenotypic and genetic level. Orange area and chroma, the area of a male's tail, and the color contrast of his pattern overall are positively correlated with attractiveness and/or mating success at the phenotypic and genetic levels. Using attractiveness and mating success as measures of fitness, we estimate gradients of linear directional sexual selection operating on each male trait and use equations of multivariate evolutionary change to predict the response of male ornamentation to this sexual selection. From these analyses, we predict that indirect selection may have important effects on the evolution of male guppy color patterns.     

The origin and paleoecologic significance of the trace fossil Asteriacites in the Pennsylvanian of Kansas and Missouri

The trace fossil Asteriacites, recorded in Cambrian to Recent shallow- and deep-marine facies, is traditionally interpreted as the resting trace of asterozoans. Well-preserved specimens of A. lumbricalis are abundant in Pennsylvanian (Upper Carboniferous) shallow- and marginal-marine siliciclastic deposits of eastern Kansas and western Missouri. Detailed morphologic analysis of these specimens suggests that they record the activities of mobile epifaunal ophiuroids. Evidence of a brittle star (ophiuroid) producer rather than sea star (asteroid) is provided by (1) trace-fossil morphologic features reflecting the anatomy of the producer (e.g., well-differentiated central structure, slender vermiform arms) and ophiuroid burrowing technique (e.g., proximal arm expansion, arm branching), and (2) mode of occurrence (e.g., gregarious behavior, horizontal and vertical repetition). Vertical and horizontal repetition produces complex aggregates of A. lumbricalis that are interpreted either as escape structures (fugichnia) or as feeding structures, respectively. Ophiura texturata is proposed as a modern analogue for the A. lumbricalis producer, based on inferred life habit and feeding behavior. Asteriacites lumbricalis is present in two different intertidal trace-fossil assemblages. The first assemblage is characterized by high diversity and records tidal flats developed outside of embayments under normal marine conditions. The second assemblage consists of A. lumbricalis together with a few other ichnotaxa and represents a depauperate association that developed in restricted tidal flats within an embayment or estuarine setting. This challenges the conventional view of Asteriacites as a normal-marine salinity indicator. Some echinoderms, and particularly asterozoans, penetrate and inhabit modern environments of depressed salinity. The presence of Asteriacites in Pennsylvanian marginal-marine facies of Kansas and Missouri provides evidence that ophiuroids had adapted to brackish-water conditions by the late Paleozoic.     

Pollen morphology of Trigonostemon and its relatives (Euphorbiaceae)

The pollen of Trigonostemon and the related genera Dimorphocalyx, Ostodes, Tritaxis and Jatropha (outgroup) has been studied with light microscopy, and scanning and transmission electron microscopy. The two major pollen types within Trigonostemon correlate well with macromorphological characters. Species belonging to the Trigonostemon reidioides type have pollen with ‘croton pattern’ ornamentation, a pistil with deeply divided stigmas (to at least half the length of the stigma arm) and stamens with a protruding appendage on the connective, while species of the Trigonostemon verrucosus type have verrucate (to almost gemmate) pollen, stigmas that are shortly cleft and stamens without an appendage on the connective. Dimorphocalyx, Ostodes, Tritaxis and Jatropha (outgroup) have similar pollen morphology, while Trigonostemon deviates from these genera in the absence of the ‘vertically’ striate ornamentation on the subunits. Therefore, when compared with an existing phylogeny of the Euphorbiaceae, the pollen characters of Trigonostemon appear to be derived. Moreover, because the ‘croton pattern’ ornamentation itself is widely shared by the ‘inaperturate crotonoids’, the loss of that structure in the Trigonostemon verrucosus type pollen is considered a further apomorphy.     

Testing the success of palaeontological methods in the delimitation of clam shrimp (Crustacea,Branchiopoda) on extant species

Fossil spinicaudatan taxonomy heavily relies on carapace features (size, shape, ornamentation) and palaeontologists have greatly refined methods to study and describe carapace variability. Whether carapace features alone are sufficient for distinguishing between species of a single genus has remained untested. In our study, we tested common palaeontological methods on 481 individuals of the extant Australian genus Ozestheria that have been previously assigned to ten species based on genetic analysis. All species are morphologically distinct based on geometric morphometrics (p ≤ 0.001), but they occupy overlapping regions in Ozestheria morphospace. Linear discriminant analysis of Fourier shape coefficients reaches a mean model performance of 93.8% correctly classified individuals over all possible 45 pairwise species comparisons. This can be further increased by combining the size and shape datasets. Nine of the ten examined species are clearly sexually dimorphic but male and female morphologies strongly overlap within species with little influence on model performance. Ornamentation is commonly species-diagnostic; seven ornamentation types are distinguished of which six are species-specific while one is shared by four species. A transformation of main ornamental features (e.g. from punctate to smooth) can occur among closely related species suggesting short evolutionary timescales. Our overall results support the taxonomic value of carapace features, which should also receive greater attention in the taxonomy of extant species. The extensive variation in carapace shape and ornamentation is noteworthy and several species would probably have been assigned to different genera or families if these had been fossils, bearing implications for the systematics of fossil Spinicaudata.