Frontal Filaments Research Articles

From swimming towards sessility in two metamorphoses – the drastic changes in structure and function of the nervous system of the bay barnacle Amphibalanus improvisus (Crustacea, Thecostraca, Cirripedia) during development

Knowledge about the development of the nervous system in cirripeds is limited, particularly with regard to the changes that take place during the two metamorphoses their larvae undergo. This study delivers the first detailed description of the development of the nervous system in a cirriped species, Amphibalanus improvisus by using immunohistochemical labeling against acetylated alpha-tubulin, and confocal laser scanning microscopy. The development of the nervous system in the naupliar stages corresponds largely to that in other crustaceans. As development progresses, the protocerebral sensory organs differentiate and the intersegmental nerves forming the complex peripheral nervous system appear, innervating the sensory structures of the cephalic shield. During metamorphosis into a cypris the lateral sides of the cephalic shield fold down into a bilateral carapace, which leads to a reorganization of the peripheral nervous system. The syncerebrum of the cypris exhibits the highest degree of complexity of all developmental stages, innervating the frontal filaments, nauplius eye, compound eyes and the antennules. During settlement, when the second metamorphosis occur, the closely associated frontal filaments and compound eyes are shed together with the cuticle of the carapace and the antennules. In adults, the syncerebral structures are reduced while the ventral nerve cord and the peripheral nervous system increase in complexity. The peripheral nervous system plays an important role in processing sensory input and also in settlement. In summary, through the larval development we observed a structural and thus also functional increase of complexity in favor of the peripheral nervous system and the ventral nerve cord.

Phylogenetic, ecological and biomechanical constraints on larval form: A comparative morphological analysis of barnacle nauplii.

Barnacle naupliar larvae are differentiated from other zooplankton by their unique pair of frontal lateral horns, frontal filaments, and a pear-shaped cephalic shield. Their morphology impose constraints on their ecological functions and reflect their evolutionary history. To explore the potential functional basis underlying the similarities and differences in barnacle larval form, we conducted a meta-analysis on the shape of the barnacle nauplii’s cephalic shield and examined its relation to larval size, trophic mode, pelagic larval duration and habitat. Nauplii cephalic shield morphology of 102 species were quantified with normalized elliptic Fourier analysis. Most of the species were distributed around the center of the morphospace but a few extreme groups occupied the periphery: nauplii that were large and lecithotrophic. Subsequent principal component regression analyses showed that larval size was a good predictor of the first shape variations axis (aspect ratio). After allometry adjustment, nauplii from different trophic modes differentiated along the second axis of the major shape variations (relative frontal horn length). Habitat was a poor predictor of variations in naupliar body form, but it could be used to differentiate extreme morphology groups from other nauplii. Our result suggests that size-related biomechanical or developmental constraints and feeding requirements are important in shaping the evolution of the naupliar body form. Within the limitations of these functional constraints, habitat drives the divergence of extreme morphology groups from the majority of species. Our comparative morphometrics analysis demonstrated how variations in larval body form can be quantitatively linked to the functional needs that constrain or drive their diversity, and inform further empirical experiments on larval functional morphology.

Fine morphology of frontal filaments in nauplii of cirriped crustaceans.

Fine morphology of the frontal filaments (FFs) at all nauplius stages of two barnacle species (Verruca stroemia and Hesperibalanus hesperius) has been investigated by scanning electron microscopy. FFs have been detected at the second nauplius stage and persist during all stages. FFs contain a wide proximal and a fine distal parts, but they are not actually separated as segments of the limbs, and the area between them looks like a single cuticular crease. Apical and subapical pores have been found at the top of each FF in the larvae of both species, which may indicate the chemoreceptor function of these organs.

The nature of non-appendicular anterior paired projections in Palaeozoic total-group Euarthropoda

Recent studies have clarified the segmental organization of appendicular and exoskeletal structures in the anterior region of Cambrian stem-group Euarthropoda, and thus led to better understanding of the deep evolutionary origins of the head region in this successful animal group. However, there are aspects of the anterior organization of Palaeozoic euarthropods that remain problematic, such as the morphological identity and significance of minute limb-like projections on the anterior region in stem and crown-group representatives. Here, we draw attention to topological and morphological similarities between the frontal filaments of extant Crustacea and the embryonic frontal processes of Onychophora, and distinctive anterior paired projections observed in several extinct total-group Euarthropoda. Anterior paired projections are redescribed in temporally and phylogenetically distant fossil taxa, including the gilled lobopodians Kerygmachela kierkegaardi and Pambdelurion whittingtoni, the bivalved stem-euarthropod Canadaspis perfecta, the larval pycnogonid Cambropycnogon klausmuelleri, and the mandibulate Tanazios dokeron. Developmental data supporting the homology of the ‘primary antennae’ of Onychophora, the ‘frontal appendages’ of lower-stem Euarthropoda, and the hypostome/labrum complex of Deuteropoda, argue against the morphological identity of the anterior paired projections of extant and extinct panarthropods as a pair of pre-ocular appendages. Instead, we regard the paired projections of fossil total-group euarthropods as non-appendicular evaginations with a likely protocerebral segmental association, and a possible sensorial function. The widespread occurrence of pre-ocular paired projections among extant and extinct taxa suggests their potential homology as fundamentally ancestral features of the anterior body organization in Panarthropoda. Non-appendicular paired projections with a sensorial function may reflect a critical – yet previously overlooked – component of the panarthropod ground pattern.

Naupliar and Metanaupliar Development of Thysanoessa raschii (Malacostraca, Euphausiacea) from Godthåbsfjord, Greenland, with a Reinstatement of the Ancestral Status of the Free-Living Nauplius in Malacostracan Evolution.

The presence of a characteristic crustacean larval type, the nauplius, in many crustacean taxa has often been considered one of the few uniting characters of the Crustacea. Within Malacostraca, the largest crustacean group, nauplii are only present in two taxa, Euphauciacea (krill) and Decapoda Dendrobranchiata. The presence of nauplii in these two taxa has traditionally been considered a retained primitive characteristic, but free-living nauplii have also been suggested to have reappeared a couple of times from direct developing ancestors during malacostracan evolution. Based on a re-study of Thysanoessa raschii (Euphausiacea) using preserved material collected in Greenland, we readdress this important controversy in crustacean evolution, and, in the process, redescribe the naupliar and metanaupliar development of T. raschii. In contrast to most previous studies of euphausiid development, we recognize three (not two) naupliar (= ortho-naupliar) stages (N1-N3) followed by a metanauplius (MN). While there are many morphological changes between nauplius 1 and 2 (e.g., appearance of long caudal setae), the changes between nauplius 2 and 3 are few but distinct. They involve the size of some caudal spines (largest in N3) and the setation of the antennal endopod (an extra seta in N3). A wider comparison between free-living nauplii of both Malacostraca and non-Malacostraca revealed similarities between nauplii in many taxa both at the general level (e.g., the gradual development and number of appendages) and at the more detailed level (e.g., unclear segmentation of naupliar appendages, caudal setation, presence of frontal filaments). We recognize these similarities as homologies and therefore suggest that free-living nauplii were part of the ancestral malacostracan type of development. The derived morphology (e.g., lack of feeding structures, no fully formed gut, high content of yolk) of both euphausiid and dendrobranchiate nauplii is evidently related to their non-feeding (lecithotrophic) status.

The morphology of the frontal filaments in the nauplii of Verruca stroemia (Müller, 1776) and Hesperibalanus hesperius (Pilsbry, 1916) (Cirripedia: Thoracica)

Scanning electron microscopy was used to investigate the fine morphology of the frontal filaments (FFs) in nauplial stages of the barnacles Verruca stroemia (Muller, 1776) and Hesperibalanus hesperius (Pilsbry, 1916). Changes in the FF structure in the course of development were examined. The FFs that were revealed in nauplial stage II were retained at all the subsequent nauplial stages. Small openings, which were located either apically or subapically, were found at the top of each FF in both species. In addition, the surface of an FF bears longitudinal grooves. The FFs are not separated from the body; however, each of them has a constriction at the base. In the nauplii of both species, the FFs have a wide proximal and a thin distal part. The area between the proximal and distal parts in V. stroemia has the form of a single crease; in H. hesperius, it resembles an accordion-shaped series of creases. The length ratio of the proximal/distal parts remains constant during all nauplial stages, but it varies between species. The proximal part constitutes 20–25% of the FF length in V. stroemia, while in H. hesperius the proximal and distal parts are approximately equal. The frontal filament length increases proportionally to the length of the larval body in the course of development. The possible functions of FFs as sense organs, their occurrence among the crustaceans, and possible homology with the preantennal limbs are discussed.

The fate of the onychophoran antenna

Recent gene expression data suggest that the region on which the onychophoran antenna is situated corresponds to the anteriormost, apparently appendage-less region of the arthropod head. The fate of the onychophoran antenna (or any appendage-like precursor), also called the primary antenna, has been discussed intensively, and there are conflicting suggestions that this anteriormost non-segmental appendage gave rise either to the arthropod labrum or, alternatively, to the so-called frontal filaments found in certain crustaceans. Our data on early axogenesis in anostracan crustaceans show that even in the earliest embryos, before the antennula and antennal nerves are developed, the circumoral anlagen of the brain display very prominent nerves which run into the frontal filament organ (also known as the cavity receptor organ). This situation resembles the development of the antennal nerves in onychophorans, which leads us to conclude that the frontal filaments are indeed homologous to the primary antenna. Frontal filaments also appear to be more common in crustaceans than previously thought, removing the need for a complicated scenario of transformation from a primary antenna into the labrum.

The development of the nervous system in Laevicaudata (Crustacea, Branchiopoda): insights into the evolution and homologies of branchiopod limbs and ‘frontal organs’

We investigated the development of the external morphology and of the nervous system in Lynceus biformis and Lynceus brachyurus (Laevicaudata, Branchiopoda), by using immunohistochemical methods in combination with a confocal laser scanning analysis. In both Lynceus species, a free-swimming nauplius larva, equipped with three appendages, hatches from resting eggs. Despite their close phylogenetic relationship to each other, considerable differences are present in their external morphology. Hatching L. brachyurus larvae are equipped with a large and flattened labrum, where in contrast, the L. biformis larvae possess a smaller labrum with four conspicuous posteriorly directed spines at its margin. Despite these differences, the development of the nervous system is quite similar in both species. The hatching larvae are equipped with a naupliar nervous system, and only in the more advanced stages, the development of the ventral nerve cord starts. Furthermore, our investigation into the nervous system provided insights into architecture and evolution of protocerebral sensory organs, the dorsal setae field and the dorsal frontal organ, only present in Laevicaudata. The identification of frontal filaments with an associated frontal filament organ in Lynceus revealed—after a comprehensive comparison with other branchiopods—that these organs exist throughout Branchiopoda and are comparable to those in other crustaceans. Additionally, our results of the peripheral nervous system analysis showed that the innervation pattern of the naupliar appendages (antenna and mandible) and the trunk appendages could be serially homologized, despite much difference in gross morphology of these. Based on the innervation pattern of limbs, we suggest that the larval uniramous mandibular palp, found in the larvae of all ‘large’ branchiopods, is largely exopodal of nature (contrary to most earlier statements) and that the endopodite of the trunk limbs consists of only one distal endite-like segment (confirming some earlier statements) and not of three as proposed by others.

Surface morphology of Diplodon expansus (Küster, 1856; Mollusca, Bivalvia, Hyriidae) gill filaments after exposure to environmentally relevant concentrations of atrazine herbicide

Brazilian endemic species Diplodon expansus (Küster, 1856) is found in freshwater bodies in the country's southeast, in large anthropogenic influence regions especially with an extensive agriculture emphasis. One of the main pesticides used in the species occurrence region is the atrazine herbicide, which has a great contamination potential in the aquatic environment. Therefore, several studies into its toxicity in aquatic systems have been developed. However, the tested concentrations are usually very high and rarely found in the environment and the short-term exposure responses in other aquatic organisms such as native bivalves are still scarce. Thus, this study sought to consider the potential effects of environmentally realistic concentrations of atrazine herbicide on the surface morphology of gill filaments of the bivalve D. expansus under laboratory-controlled conditions after short-term exposure. None of the animals died before the end of the experiment. The main alterations were observed on the frontal surface of filaments, which include mucus accumulation, cilia loss, and disruption. Mucus increased secretion and accumulation in the frontal filaments region preceded as a protective mechanism. Cilia loss and disruption on the frontal surface of the gill filament indicated that ciliated frontal cells were more sensitive to atrazine exposure and these alterations may cause gills functional damages, compromising the uptake of food particles and respiration. Therefore, higher sublethal concentrations of atrazine may compromise the survival and consequently the population of D. expansus in agriculture areas after a longer period of continuous exposure.

Speleonectes williamsi, a new species of Remipedia (Crustacea) from the Bahamas

We describe a new species of the genus Speleonectes (Crustacea, Remipedia, Nectiopoda) from an anchialine cave on Grand Bahama Island in the northern Bahamas. Speleonectes williamsi n. sp. is morphologically highly similar to Speleonectes emersoni from the Dominican Republic. However, morphological differences between the two species were detected in dissected body parts, such as the setal patterns of the antennae and trunk limbs, the terminal claws of maxillae and maxillipeds, and the frontal filaments.

The formation of the nervous system during larval development in Triops cancriformis (Bosc) (crustacea, Branchiopoda): An immunohistochemical survey

We provide data of the development of thenervous system during the first five larval stages of Triops cancriformis. We use immunohistochemical labeling (against acetylated α-tubulin, serotonin, histamine, and FMRFamide), confocal laser scanning microscopy analysis, and 3D-reconstruction. The development of the nervous system corresponds with the general anamorphic development in T. cancriformis. In larval stage I (L I), all brain parts (proto-, deuto-, and tritocerebrum), the circumoral connectives, and the mandibular neuromere are already present. Also, the frontal filaments and the developing nauplius eye are already present. However, until stage L III, the nauplius eye only consists of three cups. Throughout larval development, the protocerebral network differentiates into distinct subdivisions. In the postnaupliar region, additional neuromeres and their commissures emerge in an anteroposterior gradient. The larval nervous system in L V consists of a differentiated protocerebrum including a central body, a nauplius eye comprising four cups, a circumoral nerve ring, mandibular- and postnaupliar neuromeres up to the seventh thoracic segment, each featuring an anterior and a posterior commissure, and two parallel connectives. The presence of a protocerebral bridge is questionable. The distribution of neurotransmitters in L I is restricted to the naupliar nervous system. Over the course of the five stages of development, neurotransmitter distribution also follows an anteroposterior gradient. Each neuromere is equipped with two ganglia innervating the locomotional appendages and possesses a specific neurotransmitter distribution pattern. We suggest a correlation between neurotransmitter expression and locomotion.

Godzilliognomus schrami, a new species of Remipedia (Crustacea) from Eleuthera, Bahamas

Godzilliognomus schrami, a new species of the crustacean class Remipedia, Yager, 1981 is the second species assigned to the genus. The new species, with an average body length of 6.8 mm, was collected from an anchialine cave on the Bahamian island of Eleuthera. Godzilliognomus schrami can be distinguished from the other species in the genus, Godzilliognomus frondosus Yager, 1989 by narrower and less trapezoidal tergites, frontal filaments that differ regarding the shape and insertion of the medial process, and dorsal antennular rami composed of only ten segments.

Antennular specialization in cyprids of coral-associated barnacles

We used video, light and scanning electron microscopy to study the cypris larvae in species of coral inhabiting barnacles (Pyrgomatidae) in search of adaptations to settlement on their highly unusual substratum. Species studied were Savignium crenatum, Trevathana jensi, Trevathana margaretae, Trevathana mizrachae and Trevathana sarae. In all five species the third antennular segment was shaped like a spearhead with only an extremely narrow attachment disc. This morphology represents the most extreme antennular specialization known from cirripedes, and it is not even matched by those parasitic barnacles that use the antennule for active penetration of cuticles. Compared to the third segment, the first, second and fourth segments exhibit no obvious specializations, and the armament of sensory setae is also as very comparable to that seen in balanomorphan cirripedes. The coral barnacles also have the nauplius eye, compound eyes, frontal filaments, lattice organs and cement glands known from other barnacles. Only T. sarae differed by having two unusually shaped setae terminally on the fourth segment. Video observations showed that the coral barnacle cyprids display the exploratory walking behaviour known from cyprids settling on hard bottom substrata. We interpret the pointed structure of the third antennular segment as an adaptation to penetrate through live coral tissue. Projecting this character onto a recent phylogenetic tree of balanomorphan species indicates that it is an apomorphy for a large clade of coral inhabiting barnacles.

Circumpolar structure and distribution of the Antarctic Circumpolar Current fronts: 1. Mean circumpolar paths

High resolution hydrographic sections and maps of the gradient of sea surface height (SSH) reveal that the Antarctic Circumpolar Current (ACC) consists of multiple jets or frontal filaments. Here we use a 15 year time series of SSH observations to determine the circumpolar structure and distribution of the ACC fronts. The jets are consistently aligned with particular streamlines along the entire circumpolar path, confirming and extending the results of an earlier study restricted to the region south of Australia. The intensity of the fronts (as measured by the cross‐front gradient of SSH) varies along the fronts and the individual branches merge and diverge, often in response to interactions with bathymetry. Maps of absolute velocity at 1000 m depth derived from Argo trajectories confirm the existence of multiple current cores throughout the Southern Ocean. High resolution hydrographic sections and profiles of temperature and salinity from Argo floats are used to show that the front locations derived from fitting SSH contours to maps of SSH gradient are consistent with locations inferred from the traditional criteria based on water mass properties, suitably modified to account for multiple frontal branches. Three regions are examined in detail: the Crozet Plateau, the Kerguelen Plateau and the Scotia Sea. These examples show how recognition of the multiple jets of the ACC can help resolve discrepancies between previous studies of ACC fronts.

The chelifores of sea spiders (Arthropoda, Pycnogonida) are the appendages of the deutocerebral segment

Within the last decade, gene expression patterns and neuro-anatomical data have led to a new consensus concerning the long-debated association between anterior limbs and neuromeres in the arthropod head. According to this new view, the first appendage in all extant euarthropods is innervated by the second neuromere, the deutocerebrum, whereas the anterior-most head region bearing the protocerebrum lacks an appendage. This stands in contrast to the clearly protocerebrally targeted "antennae" of Onychophora and to some evidence for protocerebral limbs in fossil euarthropod representatives. Yet, the latter "frontal appendages" or "primary antennae" have most likely been reduced or lost in the lineage, leading to extant taxa. Surprisingly, a recent neuro-anatomical study on a pycnogonid challenged this evolutionary scenario, reporting a protocerebral innervation of the first appendages, the chelifores. However, this interpretation was soon after questioned by Hox gene expression data. To re-evaluate the unresolved controversy, we analyzed neuro-anatomy and neurogenesis in four pycnogonid species using immunohistochemical techniques. We clearly show the postprotocerebral innervation of the chelifores, which is resolved as the plesiomorphic condition in pycnogonids when evaluated against a recently published comprehensive phylogeny. By providing direct morphological support for the deutocerebral status of the cheliforal ganglia, we reconcile morphological and gene expression data and argue for a corresponding position between the anterior-most appendages in all extant euarthropods. Consequently, other structures have to be scrutinized to illuminate the fate of a presumptive protocerebral appendage in recent euarthropods. The labrum and the "frontal filaments" of some crustaceans are possible candidates for this approach.

Immunocytochemical studies on the naupliar nervous system of Balanus improvisus (Crustacea, Cirripedia, Thecostraca)

The nervous system of nauplii of the crustacean taxon Cirripedia was analysed in the species Balanus improvisus Darwin, 1854 using for the first time immunocytochemical staining against serotonin, RFamide and α-tubulin in combination with confocal laser scanning microscopy. This approach revealed a circumoesophageal neuropil ring with nerves extending to the first and second antennae and to the mandibles, all features typical for Crustacea. In addition, RFamidergic structures are present in the region of the thoraco-abdomen. A pair of posterior nerves and a pair of lateral nerves run in anterior-posterior direction and are connected by a thoracic nerve ring and a more posteriorly situated commissure. A median nerve is situated along the ventral side of the thoraco-abdomen. The innervation of frontolateral horns and the frontal filaments are α-tubulin-positive. Several pairs of large neurons in the protocerebrum, along the circumoesophageal connectives and in the mandibular ganglion stain only for serotonin. Due to the almost complete absence of comparable data on the neuroanatomy of early (naupliar) stages in other Crustacea, we include immunocytochemical data on the larvae of the branchiopod, Artemia franciscana Kellogg, 1906 in our analysis. We describe several characteristic neurons in the brains of the nauplius larvae of both species which are also found in decapod larvae and in adult brains of other crustaceans. Furthermore, our data reveal that the naupliar brain of cirripedes is more complex than the adult brain. It is concluded that this ontogenetic brain reduction is related to the sessile life style of adult Cirripedia.

Speleonectes emersoni, a new species of Remipedia (Crustacea) from the Dominican Republic

A new species of Remipedia (Crustacea), Speleonectes emersoni, is described from two anchialine caves in the Dominican Republic. It represents the first record of remipedes for the island of Hispaniola. Speleonectes emersoni is a comparatively small-sized remipede. It can be distinguished from other species in the genus Speleonectes by a combination of features that include: 1) a relatively large antennal exopod, 2) frontal filaments with subapically inserted medial processes extending over the tip of the main filament, 3) an arc-shaped horseshoe-type claw with 7–10 small denticles, and 4) an anal segment that is longer than wide with caudal rami nearly as long as the anal segment.

Multiple Jets of the Antarctic Circumpolar Current South of Australia*

Abstract Maps of the gradient of sea surface height (SSH) and sea surface temperature (SST) reveal that the Antarctic Circumpolar Current (ACC) consists of multiple jets or frontal filaments. The braided and patchy nature of the gradient fields seems at odds with the traditional view, derived from hydrographic sections, that the ACC is made up of three continuous circumpolar fronts. By applying a nonlinear fitting procedure to 638 weekly maps of SSH gradient (∇SSH), it is shown that the distribution of maxima in ∇SSH (i.e., fronts) is strongly peaked at particular values of absolute SSH (i.e., streamlines). The association between the jets and particular streamlines persists despite strong topographic and eddy–mean flow interactions, which cause the jets to merge, diverge, and fluctuate in intensity along their path. The SSH values corresponding to each frontal branch are nearly constant over the sector of the Southern Ocean between 100°E and 180°. The front positions inferred from SSH agree closely with positions inferred from hydrographic sections using traditional water mass criteria. Recognition of the multiple branches of the Southern Ocean fronts helps to reconcile differences between front locations determined by previous studies. Weekly maps of SSH are used to characterize the structure and variability of the ACC fronts and filaments. The path, width, and intensity of the frontal branches are influenced strongly by the bathymetry. The “meander envelopes” of the fronts are narrow on the northern slope of topographic ridges, where the sloping topography reinforces the β effect, and broader over abyssal plains.

Micropacteridae, a new family of Remipedia (Crustacea) from the Turks and Caicos Islands

Recent diving explorations of anchialine caves on the Turks and Caicos Islands yielded a rather small and slender new species of Remipedia. Micropacter yagerae n. gen., n. sp. is distinguished from all other species of nectiopod remipedes by a number of autapomorphic characters, including an oval body terminus with fused segments, unequal pairs of terminal claws on maxilla and maxilliped, an almost complete reduction of sternal bars and pleurotergites, molar processes with relatively few, but strong spines, and frontal filaments with bifurcate processes. Based on the unique combination of derived and primitive characters, we propose to erect a new family, Micropacteridae, for this new species and genus of Remipedia. Taxonomic diagnoses for the class Remipedia, order Nectiopoda (emended due to discovery that the maxilliped is 9-segmented), and for the families Speleonectidae and Godzilliidae are presented and discussed.

Cypris Larvae inPolysaccus MediterraneusandMycetomorpha Vancouverensis: Their Importance in Analyzing the Phylogeny and Sexual Evolution of Parasitic Barnacles (Crustacea: Cirripedia: Rhizocephala)

We used SEM to investigate the cypris larvae in two species of parasitic barnacles, Polysaccus mediterraneus (Polysaccidae) from the eastern Mediterranean and Mycetomorpha vancouverensis (Mycetomorphidae) from the northern Pacific. Both species belong to the rhizocephalan suborder Akentrogonida and have morphologies of the adult parasite that greatly differ from that seen in the "classical" suborder Kentrogonida. In contrast, both P. mediterraneus and M. vancouverensis have cypris larvae that are remarkably similar to those of the Kentrogonida. For both species, similarities include the presence of aesthetascs on the third and fourth antennular segments and a spinous process distally on the attachment disc. For P. mediterraneus additional similarities to the kentrogonid cyprids are terminal pores in the lattice organs, frontal horn gland pores, frontal filaments, and four terminal setae on the fourth antennular segment. Cyprids of Mycetomorpha have, like most other Akentrogonida, either lost or specialized (terminal setae) these structures. Compared to the stereotyped life cycle found in the Kentrogonida, species of the Akentrogonida have a diversity of reproductive systems and advanced modes of metamorphosis, and this is reflected in specializations of their cyprids. The larval morphologies of P. mediterraneus and M. vancouverensis suggest that these two species have a sexual system and life cycle much closer to that seen in the Kentrogonida. Our results show that adult and larval morphologies are not closely correlated in the Rhizocephala, and provide important information for tracing how the Akentrogonida originated and radiated from within a paraphyletic Kentrogonida.