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Lattice organs of cypris larvae (A, B, E, F–I Lithoglyptidae; C, D Trypetesidae); note the distinct median crest in the organs in C–E; dotted arrows indicate anterior direction: (A) Weltneria spinosa , anterior lattice organs, dorsal view; (B) Armatoglyptes mitis , anterior lattice organs, lateral view of one side; (C) Trypetesa lateralis , anterior lattice organs, lateral view of one side; (D) T. lateralis , lattice organ 2, lateral view; (E) A. mitis , posterior lattice organs, lateral view; (F) Weltneria spinosa , posterior lattice organs, lateral view; (G–I) W. spinosa , lattice organs 3 (G), 4 (H) and 5 (I) lateral views. k 1⁄4 median keel; lcp 1⁄4 large, central, lo1–5 1⁄4 lattice organs 1–5; pf 1⁄4 pore field of lattice organ; tp 1⁄4 terminal pore of lattice organ. Scale bars in m m. 

Lattice organs of cypris larvae (A, B, E, F–I Lithoglyptidae; C, D Trypetesidae); note the distinct median crest in the organs in C–E; dotted arrows indicate anterior direction: (A) Weltneria spinosa , anterior lattice organs, dorsal view; (B) Armatoglyptes mitis , anterior lattice organs, lateral view of one side; (C) Trypetesa lateralis , anterior lattice organs, lateral view of one side; (D) T. lateralis , lattice organ 2, lateral view; (E) A. mitis , posterior lattice organs, lateral view; (F) Weltneria spinosa , posterior lattice organs, lateral view; (G–I) W. spinosa , lattice organs 3 (G), 4 (H) and 5 (I) lateral views. k 1⁄4 median keel; lcp 1⁄4 large, central, lo1–5 1⁄4 lattice organs 1–5; pf 1⁄4 pore field of lattice organ; tp 1⁄4 terminal pore of lattice organ. Scale bars in m m. 

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Article
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Scanning electron microscopy was used to provide a full morphological description of cypris morphology in the acrothoracican species Lithoglyptes mitis and L. habei (Lithoglyptidae). Special attention was given to lattice organs, antennules, thorax, thoracopods, abdomen, and furcal rami. Cypris larvae of the Acrothoracica share some putative plesio...

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Context 1
... the Lithoglyptidae and the Trypetesidae the lattice organs are elongate and narrow depressions (7-18 mm by 0.8-1 mm) perforated by numerous very small rounded pores (Fig. 5D, G, I, and H). The depression is normally traversed by a median crest, which can be very distinct (Fig. 5C, D, and I). All lattice organs have a distinct, large terminal pore, situated anteriorly in LO2 but posteriorly in LO1 and LO3-5 ( Fig. 5A-D, G, H, and I). LO1-2 are arranged around a large, central ''pore'' in the dorsal midline of the ...
Context 2
... the Lithoglyptidae and the Trypetesidae the lattice organs are elongate and narrow depressions (7-18 mm by 0.8-1 mm) perforated by numerous very small rounded pores (Fig. 5D, G, I, and H). The depression is normally traversed by a median crest, which can be very distinct (Fig. 5C, D, and I). All lattice organs have a distinct, large terminal pore, situated anteriorly in LO2 but posteriorly in LO1 and LO3-5 ( Fig. 5A-D, G, H, and I). LO1-2 are arranged around a large, central ''pore'' in the dorsal midline of the carapace, whilst LO3-5 lie around a similar, posteriorly situated ''pore'' ( Fig. 5A and ...
Context 3
... elongate and narrow depressions (7-18 mm by 0.8-1 mm) perforated by numerous very small rounded pores (Fig. 5D, G, I, and H). The depression is normally traversed by a median crest, which can be very distinct (Fig. 5C, D, and I). All lattice organs have a distinct, large terminal pore, situated anteriorly in LO2 but posteriorly in LO1 and LO3-5 ( Fig. 5A-D, G, H, and I). LO1-2 are arranged around a large, central ''pore'' in the dorsal midline of the carapace, whilst LO3-5 lie around a similar, posteriorly situated ''pore'' ( Fig. 5A and ...
Context 4
... can be very distinct (Fig. 5C, D, and I). All lattice organs have a distinct, large terminal pore, situated anteriorly in LO2 but posteriorly in LO1 and LO3-5 ( Fig. 5A-D, G, H, and I). LO1-2 are arranged around a large, central ''pore'' in the dorsal midline of the carapace, whilst LO3-5 lie around a similar, posteriorly situated ''pore'' ( Fig. 5A and ...

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... a highly reduced abdomen with short telson (Kolbasov et al., 1999), the ascothoracid larvae possess developed 4-5-segmented abdomen with long telson (Kolbasov et al., 2008; own data), although their telson lacks cuticular ridges forming plates, is flattened laterally and bears only a pair of telsonic spines (in contrast to normally 4-6 telsonic spines in y-cyprids, see Table 1). All six thoracomeres in cypridiform larvae of Ascothoracida and Cirripedia are separated but the tergite of first thoracomere in y-cyprids is fused with the cephalon or with the second thoracomere in Grygier's (1987) inter- The cirripede cyprids also possess four-segmented antennules, but they lack a curved hook comparable to that in y-cyprids, and their third segment bears an attachment disc covered with cuticilar villi and having exit pores for both the multicellular cement gland and unicellular glands (Bielecki et al., 2009;Høeg et al., 2003). ...
... a highly reduced abdomen with short telson (Kolbasov et al., 1999), the ascothoracid larvae possess developed 4-5-segmented abdomen with long telson (Kolbasov et al., 2008; own data), although their telson lacks cuticular ridges forming plates, is flattened laterally and bears only a pair of telsonic spines (in contrast to normally 4-6 telsonic spines in y-cyprids, see Table 1). All six thoracomeres in cypridiform larvae of Ascothoracida and Cirripedia are separated but the tergite of first thoracomere in y-cyprids is fused with the cephalon or with the second thoracomere in Grygier's (1987) inter- The cirripede cyprids also possess four-segmented antennules, but they lack a curved hook comparable to that in y-cyprids, and their third segment bears an attachment disc covered with cuticilar villi and having exit pores for both the multicellular cement gland and unicellular glands (Bielecki et al., 2009;Høeg et al., 2003). ...
Article
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Although the naupliar and cypridiform stages of the enigmatic y-larvae of Facetotecta have been found in the marine plankton worldwide, they still represent the last significant group of crustaceans for which the adult forms are still unknown. From a number of y-cyprids representing different taxa from different locations, we employ scanning electron microscopy to describe fine morphological details of all external structures of this unique larval form. We document different segmentation patterns of the abdomen and presence/absence of the labrum and structural differences in the antennules, labrum, paraocular process, thoracopods, and telson lend support for the erection of several new genera as opposed to the single Hansenocaris. The data presented here emphasize the morphological limits of the genus Hansenocaris and the "bauplan" of cyprydiform larvae of Facetotecta. Although the optimum pathway is a joint analysis of both molecular and morphological characters, we use the morphological characters of y-cyprids to align them cladistically and determine the limits of the genus Hansenocaris s.s. and describe common characters for all y-cyprids including six pairs of the lattice organs instead five pairs considered as a ground pattern for all Thecostraca. We also determine plesiomorphic and apomorphic characters of all known y-cyprids and separate them from other thecostracan cypridiform larvae.
... and Synagoga arabesque; furthermore, contrary to other members of both genera, Synagoga arabesque has an anterior pore in lo2 . Such findings tend to refute the idea of cladespecific positioning of the pores of the first two pairs of lattice organs, but species of both Sessilogoga and Synagoga do share one additional potential synapomorphy: the anterior position of the terminal pores in the third pair of lattice organs is contrary to most thecostracans, in which lo3 has a posterior terminal pore (e.g., Jensen et al. 1994;Kolbasov et al. 1999;Høeg et al. 2004;Celis et al. 2008). Similar remarks by were based on then-unpublished observations of the present new species. ...
Article
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More than 40 specimens of a new endoparasitic ascothoracidan species, described herein as Sessilogoga captiva Kolbasov & Grygier sp. nov. in the family Synagogidae Gruvel, 1905, were found at 35 m depth near Green Island off southeastern Taiwan infecting a colony of the antipatharian Antipathes sp. aff. A. atlantica Gray, 1857. Its sole known congener is S. elongata Grygier, 1990a, an endoparasite of an unidentified antipatharian from Guam. We photographed some specimens in life and used scanning electron microscopy (SEM) and light microscopy to extensively document the fine-scale external morphology of both sexes, including carapace ornamentation (with special attention to lattice organs), the frontal filament complex, the antennular armature, and details of the mouthparts. We also examined the first two naupliar larval stages by SEM. Among the three most plesiomorphic genera of ascothoracidans, the two species of Sessilogoga Grygier, 1990a exhibit several synapomorphies, probably connected with their endoparasitic way of life, that distinguish them from the vagile, possibly micro-predatory species of Synagoga Norman, 1888 and the ectoparasitic, possibly vagile species of Waginella Grygier, 1983a: (i) a reduced frontal filament complex, (ii) absence of epaulets on the sixth thoracomere, (iii) reduction of the setation of the proximal antennular segments, and (iv) more seminal receptacles in any given pair of thoracopods. The new species appears to be gonochoric with females generally larger than males; our find of a juvenile female smaller than any adult male tends to refute the likelihood of protandry.
... Species of both Synagoga and Sessilogoga share anterior terminal pores in lo3 and posterior terminal pores in lo4 and lo5. This is opposite to the condition in most thecostracans, which have a posterior terminal pore in lo3 (e.g., Jensen et al. 1994;Kolbasov et al. 1999;Høeg et al. 2004;Celis et al. 2008), and thus represents a potential synapomorphy of these two genera (unpublished data). The different position of terminal pores of the lattice organs even within congeners (terminal pores of anterior lattice organs in Synagoga) shown here for the first time might be evidence that the configuration of lattice organs in ascothoracidans is not constant, at least in adult stages. ...
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A new ascothoracidan species has been discovered off Taiwan in the north part of the west Pacific at SCUBA depths. Twelve specimens including both sexes of the new species, described herein as Synagoga arabesquesp. nov. , were collected from colonies of the antipatharian Myriopathes cf. japonica Brook, 1889. Three previously described species of Synagoga , morphologically the least specialized ascothoracidan genus, have been found as ectoparasites of antipatharians and an alcyonarian, whereas all other records of this genus have been based on specimens collected from the marine plankton. This is the second study of a new form of Synagoga to be based on more than a few mature specimens of a single sex or on a single juvenile. Furthermore, it is the second in which SEM has been used to document the fine-scale external morphology. The position of terminal pores in the anterior pairs of the lattice organs is different in Synagoga arabesquesp. nov. than those in S. grygieri Kolbasov & Newman, 2018 and S. millipalus Grygier & Ohtsuka, 1995. Species of Synagoga are small, host-specific predators or ectoparasites of antipatharians. This genus exhibits a major Tethyan reliction pattern.
... Together with the Cirripedia and Facetotecta, they form the class Thecostraca (Grygier 1987b), which apparently may also encompass the Tantulocarida (Newman 1987;Petrunina et al. 2014). A number of morphological features including a well-developed and segmented abdomen, multisegmented prehensile antennules, and carapace lattice organs of a particular kind place the ascothoracidans as the most generalized of the thecostracans ( Newman et al. 1969;Grygier 1987a, b;Kolbasov et al. 1999;Høeg and Kolbasov 2002;Pérez-Losada et al. 2009). Their body is covered by a bivalved carapace. ...
... Therefore, it is not surprising that they concentrate, along with cuticular pores, at the anterior end of the carapace where prehensile antennules and mouth parts are situated. Similar mantle structures are found in ascothoracid larvae and males of other Ascothoracida ( Grygier 1982Grygier , 1988Grygier , 1991Itô and Grygier 1990;Grygier and Ohtsuka 1995;Kolbasov et al. 2008), and also in many cyprid larvae of the Cirripedia ( Kolbasov et al. 1999). The cuticular ornamentation of the mantle in adults of S. grygieri sp. ...
... So it seems reasonable to assume that the function of these pits is to aid in the search of a partner, especially since most ascothoracidans lack such sensory structures like eyes. These structures are likely homologous with frontolateral pores of the cyprids of the Cirripedia, the assumed function of which is chemosensory ( Kolbasov et al. 1999;Kolbasov and Høeg 2007), although TEM and electrophysiological studies are needed to be certain. ...
... Together with the Cirripedia and Facetotecta, they form the class Thecostraca (Grygier 1987b), which apparently may also encompass the Tantulocarida (Newman 1987;Petrunina et al. 2014). A number of morphological features including a well-developed and segmented abdomen, multisegmented prehensile antennules, and carapace lattice organs of a particular kind place the ascothoracidans as the most generalized of the thecostracans (Newman et al. 1969;Grygier 1987a, b;Kolbasov et al. 1999;Høeg and Kolbasov 2002;Pérez-Losada et al. 2009). Their body is covered by a bivalved carapace. ...
... Therefore, it is not surprising that they concentrate, along with cuticular pores, at the anterior end of the carapace where prehensile antennules and mouth parts are situated. Similar mantle structures are found in ascothoracid larvae and males of other Ascothoracida (Grygier 1982(Grygier , 1991Itô and Grygier 1990;Grygier and Ohtsuka 1995;Kolbasov et al. 2008), and also in many cyprid larvae of the Cirripedia (Kolbasov et al. 1999). The cuticular ornamentation of the mantle in adults of S. grygieri sp. ...
... So it seems reasonable to assume that the function of these pits is to aid in the search of a partner, especially since most ascothoracidans lack such sensory structures like eyes. These structures are likely homologous with frontolateral pores of the cyprids of the Cirripedia, the assumed function of which is chemosensory (Kolbasov et al. 1999;Kolbasov and Høeg 2007), although TEM and electrophysiological studies are needed to be certain. ...
Article
A new ascothoracidan species has been discovered in the Macaronesia region of the eastern Atlantic Ocean at SCUBA depths. Ten specimens including both sexes of the new species, described herein as Synagoga grygieri sp. nov., were collected from colonies of the antipatharian Antipathella wollastoni (Gray, 1857). Two previously described species of Synagoga, morphologically the most generalized ascothoracidan genus, were found as ectoparasites of an antipatharian and an alcyonacean, respectively, whereas all other records of this genus have been from marine plankton. Synagoga grygieri sp. nov. currently appears to be endemic to Macaronesia, but its true distribution may be wider as its host is known to range from the Mediterranean to the west coast of Africa, and a potentially synonymous congener of the host ranges somewhat northward. The new species is described as having separate males and females, although protandry cannot be fully excluded. This is the first study of a new form of Synagoga to be based on more than a few mature specimens of a single sex or on a single juvenile. Furthermore, it is the first in which SEM has been used to document features other than carapace ornamentation, in particular the distal antennular armature, frontal filament complex, and details of the mouth parts. On the carapace of adult stages, the first and third pairs of lattice organs are oriented with the terminal pore anterior, as in S. millipalus Grygier & Ohtsuka, 1995.
... The early-stage larvae (i.e. nauplii) are brooded as ''nauplii-in-eggs" within the mantle of females and later released as cypris larvae for settlement (Batham and Tomlinson, 1965;Kolbasov et al., 1999;Kolbasov and Høeg, 2007;Tomlinson, 1969;Turquier, 1985). Unlike other acrothoracicans, cryptophialid cyprids lack swimming thoracopods; thus, they can only engage in an exploratory ''walk" with antennules on the surface close to their parents and settle afterward (Darwin, 1854;Kolbasov and Høeg, 2007) or be dispersed passively with water currents. ...
Article
The barnacles of the superorder Acrothoracica are small, burrowing, epibiotic, and dioecious (large female with dwarf male) crustaceans largely found in the carbonate sediments and skeletons of marine invertebrates. The acrothoracicans represent the Cirripedia with the most plesiomorphic characters and have prominently featured in phylogenetic speculations concerning these crustaceans. Traditionally, Acrothoracica was divided into two main orders, Pygophora and Apygophora. The Apygophora had uniramus cirri and no anus. The Pygophora had biramus terminal cirri and an anus and was further divided into two families, Lithoglyptidae and Cryptophialidae. Kolbasov (2009) revised the superorder Acrothoracica on the basis of morphological examinations of females, dwarf males, and cyprids and rearranged the acrothoracican species into two new orders, Lithoglyptida and Cryptophialida. The present study is the first attempt to reconstruct the phylogenetic relationships of acrothoracican barnacles by sequencing two mitochondrial (cytochrome C oxidase I and 16S ribosomal DNA) and two nuclear (18S ribosomal DNA and histone H3) markers of 8 of the 11 genera comprising 23 acrothoracican species. All monophylies of the eight acrothoracican genera sampled in this study were strongly supported. The deep interfamilial relationship constructed is consistent with the recent morphological phylogenetic relationship proposed by Kolbasov, Newman, and Høeg (Kolbasov, 2009) that Cryptophialidae (order Cryptophialida) is the sister group to all other acrothoracicans (order Lithoglyptida). According to an ancestral character state reconstruction analysis, the posterior lobes of females; armament of opercular bars, attachment stalk, lateral projections of the body, and aperture slits in dwarf males; and habitat use appear to have phylogenetic importance.
... Recently, two studies on cirripedes from different orders addressed this problem. In the acrothoracicans Lithoglyptes mitis and Lithoglyptes habei, a scanning electron microscopy analysis reported the presence of four reduced segments located between the thorax and telson in cypris larvae (Kolbasov et al. 1999). In the rhizocephalan Sacculina carcini, we analysed the expression pattern of the segmentation gene engrailed during larval development: this study revealed five small engrailed stripes in a postero-dorsal region behind the sixth thoracic segment, that we interpreted as a vestigial abdomen (Gibert et al. 2000). ...
... It is worth mentioning that among represen tatives of the superorder Rhizocephala FFs are present in all species with free swimming larvae [4,27,33,61], but are absent from species whose larvae develop in a brood pouch [7,27]. Among members of the superorder Acrothoracica, FFs have also been found in free living larvae during the nauplial (when these stages are present) and cyprid stages of development [44,46,47]. ...
... It is worth mentioning that among represen tatives of the superorder Rhizocephala FFs are present in all species with free swimming larvae [4,27,33,61], but are absent from species whose larvae develop in a brood pouch [7,27]. Among members of the superorder Acrothoracica, FFs have also been found in free living larvae during the nauplial (when these stages are present) and cyprid stages of development [44,46,47]. ...
Article
Full-text available
Scanning electron microscopy was used to investigate the fine morphology of the frontal filaments (FFs) in nauplial stages of the barnacles Verruca stroemia (Müller, 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.