Copulatory Structures Research Articles

Atlas of Australasian hormurid scorpions. I. The genus Hormurus Thorell, 1876 in Papua New Guinea. Exceptional morphological diversity in male and female copulatory structures suggests genital coevolution

Atlas of Australasian hormurid scorpions. I. The genus Hormurus Thorell, 1876 in Papua New Guinea. Exceptional morphological diversity in male and female copulatory structures suggests genital coevolution

Taxonomy and phylogeny of Dalytyphloplanida Willems et al., 2006 (Platyhelminthes: Rhabdocoela), with the description of a new family, a new genus, and sixteen new species from Cuba and Panama

Twenty-two marine species of Dalytyphloplanida from Cuba, Panama, and Lanzarote are presented, sixteen of which are new to science. Five known species are recorded from Cuba: Kytorhynchus microstylus, Ceratopera paragracilis, Trigonostomum armatum, T. franki, and T. vanmecheleni. Neokytorhynchus pacificus is recorded for the first time in the Atlantic (Lanzarote). For one species, a new genus is erected to be included in a new family. A new genus and thirteen new species from Cuba and three new species from Panama are described. The new species are mainly distinguished from their congeners by the specific morphology of the copulatory structures. The new taxa are discussed in the context of a new phylogenetic analysis of Dalytyphloplanida. This analysis includes sequences of 238 species, 14 of which were sequenced for the first time. The phylogenetic analysis and the detailed morphology suggest that K. microstylus may constitute a complex of cryptic species. The internal phylogenetic relationships of Trigonostomum and Promesostoma were separately analysed to include new sequence data of the mitochondrial cox1 gene. These genera are shown to consist of several diagnosable clades, concurring with the results of early morphological studies of these taxa. Morphological and molecular data support the notion that Trigonostomum sinensis is a junior synonym of T. vanmecheleni, and we therefore propose their synonymisation. A new family is erected, Schockaertiidae fam. n., to include a monophyletic clade of neodalyellids with a pharynx rosulatus and a copulatory bulb devoid of hard structures (Schockaertia aprostatica gen. n. sp. n. and Einarella argillophyla). We propose a new classification into families of Thalassotyphloplanida, which envolves elevating the status of Adenorhynchinae to family level (Adenorhynchidae status novus). All previously recognised thalassotyphloplanid families are recovered as monophyletic (Kytorhynchidae, Byrsophlebidae, Promesostomidae, and Trigonostomidae). No support was found for the previously described subfamilies, except Paramesostominae, and we, therefore, propose to suppress these taxa.

Anatomy of male and female genitalia of Acanthoscelidesobtectus (Say, 1831) (Coleoptera, Chrysomelidae, Bruchinae) in interaction.

Armatures of the male intromittent copulatory structures have been surmised to increase male fitness by imposing physiological costs on female re-mating. Female kicking could, consequently, be a counterstrategy to avoid wounding or to prevent males from mating. The membranous endophallus of male Acanthoscelidesobtectus (Say, 1831) is armed with denticles. Checking if these denticles penetrate the wall of the female genital tract during copulation revealed that only the tip of the median lobe of the aedeagus is intromitted into the female genital opening during copulation. The everted endophallus extends over the full length of the ovipositor, and the spermatophore is placed in the bursa. Identification by means of light microscopy and Micro-CT of the exact relative position of male and female copulatory organs while mated confirmed that the denticles do not cause wounds in the vagina wall. Parts of the inner wall of the bursa copulatrix are covered with inward pointing denticles. Already mated females kick mounting males by vehement movements of their hind legs, thereby preventing mating. In contrast, virgin females usually accept the first male they encounter and terminate copulation by slower movements of their hind legs. The same applied to females who accepted re-mating the second day after the first copulation. Acanthoscelidesobtectus females kick males off to prevent rather than to terminate copulation. Copulatory structures as well as behaviour may have different functional roles in different beetle species, even within the Bruchinae.

Two new species of Cyatholaimidae (Nematoda: Chromadorida) from the Southeastern Brazilian coast with emphasis on the pore complex and lateral pore-like structures.

Cyatholaimidae is a common and diverse family of mainly marine nematodes, potentially, with a large number of species to be discovered. The taxonomy of the group is marked by a lack of information about the evolutionary history of the characters and of detailed descriptions of morphological structures that may be taxonomically relevant. Two new species of the family are described from a sublittoral region in Southeastern Brazil, emphasizing the importance of the distribution and morphology of pore complex and pore-like structures present on the cuticle. The taxonomic importance of the cuticle ornamentation and spicule shape for the Biarmifer species, as well as the precloacal supplements structures of Pomponema species, are discussed. Biarmifer nesiotes sp. nov. differs from other species of the genus by the presence of eight longitudinal rows of pore complex on the cuticle and by the shape of the copulatory structure. Pomponema longispiculum sp. nov. differs from the most similar species, P. stomachor Wieser, 1954, by the smaller number of turns of the amphidial fovea, the shorter tail and the beginning of the cuticle lateral differentiation (3/4 of the pharynx length vs. end of the pharynx, respectively). We also obtained the SSU rDNA sequence from Pomponema longispiculum sp. nov., which is closely related to Pomponema sp. (MN250093) by about 91%. Updated tabular keys to species identification of each genus (Biarmifer and Pomponema) are included, containing morphometric data, characters related to cuticle ornamentation, and copulatory structures.

Testing Morphological Relationships Between Female and Male Copulatory Structures in Bats.

The lower reproductive tract of female mammals has several competing functions including mating, tract health maintenance, and parturition. Diverse vaginal anatomy suggests interactions between natural and sexual selection, yet despite its importance, female copulatory morphology remains under-studied. We undertook a comparative study across the species-rich mammalian order Chiroptera (bats) with a focus on the suborder Yangochiroptera (Vespertilioniformes) to examine how female vaginal features may have coevolved with male penis morphology to minimize mechanical damage to their tissues during copulation. The penis morphology is diverse, presenting great potential for post-copulatory sexual selection and coevolution with the female morphology, but vaginas have not been carefully examined. Here we test the hypotheses that vaginal thickness and collagen density have coevolved with features of the male penis including the presence of spines and a baculum. We present histological data from females of 24 species from 7 families of bats, and corresponding data on male penis anatomy. We also examine the role of phylogenetic history in the morphological patterns we observe. We found evidence that female vaginal thickness has coevolved with the presence of penile spines, but not with baculum presence or width. Collagen density did not appear to covary with male penile features. Our findings highlight the importance of considering interactions between the sexes in influencing functional reproductive structures and examine how these structures have been under selection in bats.

Limitations of sperm transfer in the complex reproductive system of spiders

Abstract In spiders, sperm transfer from the male to the female is indirect via secondary copulatory structures, the pedipalps. At the time of transfer the sperm are not mobile and the ejaculate needs to move through narrow male and female ducts to the female sperm storage organ. In addition, copulation duration can be very short, often limited to just a few seconds. Finally, sexual cannibalism and genital damage limits male life-time mating opportunities. These features of the reproductive biology in spiders are likely to result in sperm transfer constraints. Here we review the intrinsic and extrinsic sperm transfer limitations and conduct a meta-analysis on sperm transfer data from published data. Most of the information available relates to orb-web spiders, but our meta-analysis also includes non-orb-web spiders. Our review identifies some of the behavioural factors that have been shown to influence sperm transfer, and lists several morphological and physiological traits where we do not yet know how they might affect sperm transfer.

Studies on raspy crickets: Caudafistulus rubrinervosus (Serville, 1838) n. comb. (Orthoptera: Gryllacrididae).

Caudafistulus n. gen. is described to accommodate Brachybaenus rubrinervosus, a species of Raspy Cricket from the Colombian Andes, endemic to the highlands of the Aburra Valley and the eastern subregion of the department of Antioquia. Caudafistulus rubrinervosus n. comb. differs from Brachybaenus species, due to its conspicuous frontal ocellus, male terminalia with processes on the tenth tergite, subgenital plate modified, and genitalia with sclerotized areas, being one of the few genera of gryllacridids known with this peculiarity in the world. Finally, the similarity of copulatory structures with old-world taxa and their possible function for mating is discussed.

Across the benthic and pelagic realms: a species‐level phylogeny of Benthesicymidae (Crustacea : Decapoda)

Benthesicymidae is a monophyletic group of Decapoda adapted to a life on the sea-floor, in the near-bottom layer, in the bathy- and in the mesopelagic, within an impressive depth range from a few hundred metres (Gennadas) to several thousand metres (Benthesicymus). Higher taxa are known to conquer all main oceanic biotopes such as the benthic, benthopelagic, and pelagic and a wide depth range but few family-level groups have clades evolved within all these oceanic realms. Therefore, the global fauna of Benthesicymidae provides a rare opportunity for an insight into phylogenetic processes favouring colonisation of all principal oceanic biotopes. The first comprehensive phylogenetic study of Benthesicymidae (all 37 valid species) is based on six molecular markers and 105 morphological characters (including 72 female and male copulatory characters). Analyses resulted in trees with similar topology and the same set of robust clades. Molecular methods based on 167 sequences (84 new) provided better resolution of deeper nodes and generally higher support of the clades, while morphological methods allowed analyses of all valid species of the global fauna. Phylogenetic analyses support the monophyly and robustness of all currently known genera except Gennadas, which was split into Gennadas Bate, 1881, Amalopenaeus Smith, 1882, and Notogennema gen. nov. We also retrieved two major clades for which we erected two new subfamilies: Benthesicyminae subfam. nov. (presumably benthic, genera Altelatipes, Bathicaris, Benthesicymus, and Benthonectes) and Gennadinae subfam. nov. (presumably pelagic, genera Amalopenaeus, Bentheogennema, Benthoecetes, Boreogennema, Gennadas, Maorrancaris, and Notogennema gen. nov.). We revealed two groups of morphological characters, that are interlinked evolutionarily: (1) petasma and thelycum; (2) body, mouthparts, and pereopods. Morphological traits within benthic and pelagic clades are different, a model explaining the differences is proposed. Along with previous studies, our results confirm the idea that the elaboration of the copulatory structures is a key to successful colonisation of the pelagic realm. These results extend our knowledge about evolution in the largest habitual biotope of our planet and phylogenetic processes favouring colonisation of all principal oceanic biotopes.

A phylogeny-based revision of the shrimp genera Altelatipes, Benthonectes and Benthesicymus (Crustacea: Decapoda: Benthesicymidae)

AbstractA phylogenetic study of deep-sea dendrobranchiate genera Altelatipes, Benthesicymus and Benthonectes based on four molecular markers and 91 morphological characters is presented. All currently recognized species of these genera, representatives of all other genera and species groups of Benthesicymidae, and three outgroups were included in the analyses. The molecular and morphological methods retrieved similar results, the molecular methods provided better resolution of deeper nodes and higher clade support. Both types of analyses showed paraphyly of Benthesicymus, which encompass five robust clades, four of which are diagnosed as new genera (type species in parentheses): Benthesicymus s.s. (B. crenatus), Bathicaris gen. nov. (Benthesicymus brasiliensis), Dalicaris gen. nov. (Benthesicymus altus), Trichocaris gen. nov. (Benthesicymus bartletti) and Maorrancaris gen. nov. (Benthesicymus investigatoris). Altelatipes was found to be monophyletic. The evolution of the major clades of Benthesicymidae is shown to be linked to trophic specialization, while further divergence at the genus level is mainly related to sexual evolution seen in the elaboration of the copulatory structures. We provide amended diagnoses of the previously recognized and new genera, key to species of each of these genera and include an updated key to genera of Benthesicymidae.

Nerves innervating copulatory organs show common FMRFamide, FVRIamide, MIP and serotonin immunoreactivity patterns across Dinophilidae (Annelida) indicating their conserved role in copulatory behaviour

BackgroundMales of the microscopic annelid family Dinophilidae use their prominent glandomuscular copulatory organ (penis) to enzymatically dissolve the female’s epidermis and thereafter inject sperm. In order to test for putative conserved copulatory structures and neural orchestration across three dinophilid species, we reconstructed the reproductive myo- and neuroanatomy and mapped immunoreactivity patterns against two specific neurotransmitter markers with reported roles in invertebrate male mating behaviour (FVRIamide, MIP) and three general neural markers (acetylated α-tubulin, serotonin, FMRFamide).ResultsSeminal vesicles (one or two pairs), surrounded by a thin layer of longitudinal and circular muscles and innervated by neurites, are found between testes and copulatory organ in the larger males of Dinophilus vorticoides and Trilobodrilus axi, but are missing in the only 0.05 mm long D. gyrociliatus dwarf males. The midventral copulatory organ is in all species composed of an outer muscular penis sheath and an inner penis cone. Neurites encircle the organ equatorially, either as a ring-shaped circumpenial fibre mass or as dorsal and ventral commissures, which are connected to the ventrolateral nerve cords. All three examined dinophilids show similar immunoreactivity patterns against serotonin, FMRFamide, and FVRIamide in the neurons surrounding the penis, supporting the hypotheses about the general involvement of these neurotransmitters in copulatory behaviour in dinophilids. Immunoreactivity against MIP is restricted to the circumpenial fibre mass in D. gyrociliatus and commissures around the penis in T. axi (but not found in D. vorticoides), indicating its role in controlling the copulatory organ.ConclusionsThe overall myo- and neuroanatomy of the reproductive organs is rather similar in the three studied species, suggesting a common ancestry of the unpaired glandomuscular copulatory organ and its innervation in Dinophilidae. This is furthermore supported by the similar immunoreactivity patterns against the tested neurotransmitters around the penis. Smaller differences in the immunoreactivity patterns around the seminal vesicles and spermioducts might account for additional, individual traits. We thus show morphological support for the putatively conserved role of FMRFamide, FVRIamide, MIP and serotonin in dinophilid copulatory behaviour.

A hard-earned draw: phylogeny-based revision of the deep-sea shrimp Bentheogennema (Decapoda: Benthesicymidae) transfers two species to other genera and reveals two new species

AbstractThe phylogenetic study of the deep-sea genus Bentheogennema is based on four molecular markers and 79 morphological characters. All four previously recognized species and two new species of Bentheogennema, representatives of all other genera and species groups of Benthesicymidae, and three outgroups were included in the analyses. We have examined and coded six major groups of morphological characters related to the carapace (three characters), the pleon and the telson (14), the mouthparts (nine), the armature of the pereopods (five), the thelycum (27) and the petasma (21). Results of morphological and molecular analyses were similar. Two species were transferred from Bentheogennema to other genera (for one of them a new genus was erected) and two new species of Bentheogennema were described. Three pelagic genera (Gennadas, Bentheogennema and a new genus) created a robust clade. The divergence of this clade is linked to ‘smoothening’ of the body (reduction of the branchiostegal spine on the carapace, reduction and loss of the dorsolateral spines and the end-piece on the telson) and elaboration of the copulatory structures. We provide amended diagnoses of these three pelagic genera and key to species of Bentheogennema.

First record of Polycystididae (Platyhelminthes, Kalyptorhynchia) from Cuba, with the description of a new genus and five new species, and remarks and the description of one new species from Panama.

The first records of Polycystididae from Cuba are provided and discussed. In total nine species have been collected, five of which are new to science, one representing a new genus. Polycydora intermedia gen. n. sp. n. shows intermediate features between Polycystis Kölliker, 1845 and Paulodora Marcus, 1948. The ovaries are kidney shaped, with the oocytes arranged in a row, and lack the hard "nozzles" of Paulodora. A female bursa as in Polycystis is present. In P. intermedia gen. n. sp. n. lacks the strong muscle bulb at the male bursa and the accessory glands type I in the male atrium, which are present in Polycystis. The male atrial organs include a prostate stylet type II connected to a free prostate vesicle type II. The four new species (Phonorhynchoides minor sp. n., Phonorhynchopsis capillaris sp. n., Phonorhynchopsis sublinguatus sp. n., Myobulla armenterosi sp. n.) are distinguished from their congeners by the specific shape and length of the male hard copulatory structures. Phonorhynchoides minor sp. n. has the smallest stylets within the genus, and these stylets are more straight than those of the other species of the genus. In Phonorhynchopsis capillaris sp. n. the prostate stylet type IV is only 20% of the length of the accessory stylet type IV, the lowest relative length within the genus. The accessory stylet in this species is extremely thin; it is only 2µm wide. Phonorhynchopsis sublinguatus sp. n. has a prostate stylet widened in the distal half, ending in a rounded tip, not twisted in the middle. Because of its strong resemblance to M. armenterosi sp. n., M. berti sp. n., a new species of Myobulla Artois Schockaert, 2000 collected at the Atlantic coast of Panama is described here too. Both species of Myobulla have a prostate stylet type III that shows a 90° turn at some point, as is the case in some other species of Myobulla. In M. berti sp. n. the stylet is smaller overall and is bent in the middle; in M. armenterosi sp. n. the stylet is larger and it is bent more distally. All of the four known species have a very wide geographical distribution: Phonorhynchopsis haegheni, Alcha evelinae, Paraustrorhynchus elixus, and Gyratrix hermaphroditus.

A phylogenetic study of krill (Crustacea: Euphausiacea) reveals new taxa and co-evolution of morphological characters.

The first comprehensive phylogenetic study of Euphausiacea (all 86 valid species) is presented. It is based on four molecular markers and 168 morphological characters (including 58 characters of the petasma). Phylogenetic analyses support the monophyly and robustness of the families Bentheuphausidae and Euphausiidae and reveal three major clades for which we erect three new subfamilies: Thysanopodinae, Euphausiinae and Nematoscelinae. All genus-level clades are statistically supported (except Thysanopoda in molecular analyses), deeply nested within the subfamily-level clades, and encompass 14 new species groups. Copulatory structures have a major impact on tree topology in the morphological analysis, the removal of which resulted in only half the number of supported clades and genera. We revealed three groups of morphological characters, which are probably coupled with the same biological role and thus interlinked evolutionarily: (i) antennular peduncle and petasma (copulation); (ii) eyes and anterior thoracopods (feeding); and (iii) shape of carapace and pleon (defence). We analysed the evolutionary pathways of the clades into main oceanic biotopes and compared them with morphological adaptations most likely to be coupled with this process.

The morphology of mouthparts, wings and genitalia of Paleozoic insect families Protohymenidae and Scytohymenidae reveals new details and supposed function

Megasecoptera is an extinct group of insects with specialized rostrum-like mouthparts, which is a synapomorphy shared with all members of the Late Paleozoic Palaeodictyopterida, and markedly slender wings that are unable to flex backwards. Here we describe the close up morphology of Protohymenidae and Scytohymenidae and uncover new aspects of the endoskeleton (tentorium) of the head, structure of the mouthparts with discernible proximal part of stylets controlled by muscles, surface of compound eyes that consist of a hexagonal pattern of large facets, structure and microstructures on the wings and reconstruct male and female external genitalia using ESEM and light stereomicroscopy. Furthermore, we describe Protohymen novokshonovi sp. n. based on an exceptionally well preserved fossil from the early Permian at Tshekarda in Russia, which shows crucial details, and the earliest species of Protohymenidae, Carbohymen testai gen. et sp. n. from a late Carboniferous siderite nodule at Mazon Creek in Illinois, USA. Our comparative study confirmed a set of structural and microstructural details on their wings, such as the composite anterior wing margin, development of an apical cell and the previously unknown external genitalia. Based on the results and comparison of homologous structures known primarily for extant relatives, such as mayflies and dragonflies, we outline for the first time the function of the mouthparts, in particular, the stylets, structure of the tentorium, vision provided by large hexagonal ommatidia and male copulatory structures bearing curved claspers for holding a female during copulation and penial lobes with seminal grooves.

The male sexual apparatus in the order Scorpiones (Arachnida): a comparative study of functional morphology as a tool to define hypotheses of homology

BackgroundInsemination in scorpions is carried out by means of a partly sclerotized structure, the spermatophore, which is composed of two separate halves, the hemispermatophores. In most genera these reproductive structures can be used to differentiate species. However, many taxa such as the genus Euscorpius and the family Diplocentridae lack the morphological diversity observed in the copulatory organs of many other arthropods, rendering them useless for species level taxonomy. Such structural stasis, however, suggests that hemispermatophores have evolved relatively slowly and may thus provide a stronger phylogenetic signal for recognizing supra-generic ranks than previously thought. Based on the postulate that the phenotypic stability observed in some groups is the consequence of functional constraint, the most comprehensive comparative study of the male sexual apparatus to date was conducted for a complete reassessment of the morphology, phylogenetic value and hypotheses of homology of these structures.ResultsHemispermatophores, pre- and post-insemination spermatophores, as well as the inherent mechanisms of insemination, were studied across the whole order, allowing the recognition and description of a series of five basic bauplans for the capsular region. For the most part, these patterns appear to be consistent within each major taxonomic group, but several cases of incongruence between spermatophore morphology and taxonomy raises questions about the monophyly of some clades. The Bothriuridae are traditionally regarded as a basal scorpionoid family. However, except for the genus Lisposoma, bothriurid hemispermatophores and spermatophores are morphologically more similar to those of the Chactoidea than to those of scorpionoids. On the other hand, the male copulatory structures of the hormurid clade (Hormiops (Hormurus + Liocheles)) are more akin to those of Diplocentridae and Heteroscorpionidae than to those of other hormurids.ConclusionsSpermatophore capsular patterns appears to be congruent with a recent phylogeny of the order Scorpiones based on phylogenomic data that placed Bothriuridae outside of Scorpionoidea and Liocheles outside of Hormuridae, in contradicton with earlier phylogenetic reconstructions based on morphology. This raises questions about the potential use of functionally constrained traits to assess the reliability of contradicting phylogenetic hypotheses and emphasizes the need for a thorough reassessment of the scorpion phylogenetic relationships.

Is geography an accurate predictor of evolutionary history in the millipede family Xystodesmidae?

For the past several centuries, millipede taxonomists have used the morphology of male copulatory structures (modified legs called gonopods), which are strongly variable and suggestive of species-level differences, as a source to understand taxon relationships. Millipedes in the family Xystodesmidae are blind, dispersal-limited and have narrow habitat requirements. Therefore, geographical proximity may instead be a better predictor of evolutionary relationship than morphology, especially since gonopodal anatomy is extremely divergent and similarities may be masked by evolutionary convergence. Here we provide a phylogenetics-based test of the power of morphological versus geographical character sets for resolving phylogenetic relationships in xystodesmid millipedes. Molecular data from 90 species-group taxa in the family were included in a six-gene phylogenetic analysis to provide the basis for comparing trees generated from these alternative character sets. The molecular phylogeny was compared to topologies representing three hypotheses: (1) a prior classification formulated using morphological and geographical data, (2) hierarchical groupings derived from Euclidean geographical distance, and (3) one based solely on morphological data. Euclidean geographical distance was not found to be a better predictor of evolutionary relationship than the prior classification, the latter of which was the most similar to the molecular topology. However, all three of the alternative topologies were highly divergent (Bayes factor >10) from the molecular topology, with the tree inferred exclusively from morphology being the most divergent. The results of this analysis show that a high degree of morphological convergence from substantial gonopod shape divergence generated spurious phylogenetic relationships. These results indicate the impact that a high degree of morphological homoplasy may have had on prior treatments of the family. Using the results of our phylogenetic analysis, we make several changes to the classification of the family, including transferring the rare state-threatened species Sigmoria whiteheadi Shelley, 1986 to the genus Apheloria Chamberlin, 1921—a relationship not readily apparent based on morphology alone. We show that while gonopod differences are a premier source of taxonomic characters to diagnose species pairwise, the traits should be viewed critically as taxonomic features uniting higher levels.

Male genital titillators and the intensity of post-copulatory sexual selection across bushcrickets

Animal genitalia are diverse and a growing body of evidence suggests that they evolve rapidly under post-copulatory sexual selection. This process is predicted to be more intense in polyandrous species, although there have been very few comparative studies of the relationship between the complexity of genital structures in males and measures of the degree of polyandry. In some bushcricket families, males possess sclerotised copulatory structures known as titillators, which are inserted into the female’s genital chamber and moved rhythmically. Like other genital structures, bushcricket titillators are widely used as important taxonomic characters and show considerable variation across species in structure, shape and the extent to which they are spined. Here, we examine relationships between the presence/absence of titillators, titillator complexity and both mating frequency and the degree of polyandry in bushcrickets, using phylogenetic comparative analyses. Using published sources combined with original observations, data were obtained for the mean level of polyandry, the duration of the male and female sexual refractory periods and the level of complexity of titillators. To analyse data, we fitted phylogenetic generalised least squares models. No significant relationships were found between titillator presence or complexity and either the level of polyandry, duration of the male’s sexual refractory period or the ratio of the female and male sexual refractory periods. The duration of the female’s refractory period, however, was positively associated with titillator presence and negatively associated with titillator complexity. The data therefore partially support the hypothesis that post-copulatory sexual selection drives genital evolution in this taxon.

Phylogeny and biogeography of the scaleless scale worm Pisione (Sigalionidae, Annelida).

Pisione is a scaleless group of small scale worms inhabiting sandy bottoms in shallow marine waters. This group was once considered rare, but now 45 described species can be characterized, among others, by their paired, segmental copulatory organs (one to multiple external pairs), which display a complexity of various accessory structures. The evolutionary significance of these unique organs was suggested in the late 1960s, but has been heavily debated since the late 1990s and remains controversial. In the present paper, we study the internal relationships within Pisione, employing combined phylogenetic analyses of both molecular and morphological data from 16 terminals of Pisione, as well as two terminals of Pisionidens, and eight additional scale worms as outgroups. Our taxon sampling covers all geographical areas where the genus has been reported, as well as most of their morphological and copulatory variability, including representatives of the “africana,” “remota,” “crassa,” and “papuensis” groups, established previously by Yamanishi. We hereby provide a first insight into the relationships of the genus, testing previously proposed hypotheses on the evolutionary significance of male copulatory structures within Pisione, while attempting to understand patterns of distribution. The phylogenetic analyses using maximum likelihood and Bayesian methods consistently recovered two large clades spanning the East Atlantic (including the Mediterranean) and the Indo‐Pacific–West Atlantic, respectively. Character optimization on our trees revealed a high degree of homoplasy in both non‐reproductive and sexual characters of Pisione, with buccal acicula found to be the sole apomorphy among the morphological features assessed herein, with none defining the biogeographical subclades within. Overall, our comparative analyses highlight the high degree of morphological variation in this widely distributed genus, rejecting previous assertions of an increasing number and complexity of copulatory structures across the genus.

The role of the male copulatory organs in the colonization of the pelagic by shrimp-like eucarids

Five major taxa of shrimp-like eucarid are recognized: the order Euphausiacea, the suborder Dendrobranchiata, and three infraorders of the suborder Pleocyemata (Caridea, Procarididea, and Stenopodidea). These animals are very successful in the colonization of the pelagic but the role of the sexual structures (the appendix masculina and the petasma) in this process remains uncertain. We revise the vertical distribution of pelagic species and genera within the shrimp-like eucarids and the presence and types of the male copulatory organs. We further test the hypothesis that the pelagic habitat is associated with the elaborated male copulatory organs and discuss possible mechanisms of association. We also consider possible effects of the depth and coastal factors. In the turbulent and fluid pelagic, successful copulation depends on the perfect fixation and possible stimulation of mates during spermatophore transfer and thus on the development and elaboration of the copulatory structures. The presence of the appendix masculina is necessary for copulation in the pelagic and thus for colonization of the pelagic. The ‘family success’ in colonizing the pelagic depends on the presence of the elaborate petasma, whilst the ‘species success’ is not so obviously related to the presence of this organ. The colonization of the water column by the eucarids may basically occur at the family level and be associated with the elaboration of the petasma. Within monophyletic pelagic groups, a greater development of the petasma is observed in the epi- and mesopelagic; some reduction of this organ in the bathypelagic may be related to the reduction in current velocities and turbulence. Simplification of the petasma is also observed in coastal waters. The proposed hypothesis is invariant to phylogenetic models and suggests an explanation to why (but not in which consequence) the elaborated copulative organs have been developed/retained in the pelagic or reduced/lost in the benthic.

Meiofaunal cryptic species challenge species delimitation: the case of the Monocelis lineata (Platyhelminthes: Proseriata) species complex

Given the pending biodiversity crisis, species delimitation is a critically important task in conservation biology, but its efficacy based on single lines of evidence has been questioned as it may not accurately reflect species limits and relationships. Hence, the use of multiple lines of evidence has been portrayed as a means to overcome identification issues arising from gene/species tree discordance, morphological convergence or recent adaptive radiations. Here, the integrative taxonomic approach has been used to address the study of the Monocelis lineataspecies complex. The taxonomic resolution of the complex is challenging, as the species lacks sclerotised copulatory structures, which as a rule of thumb aid identification in Proseriata. Eighteen populations, which encompass most of the geographic range of the complex, were studied using morphology, karyology, crossbreeding experiments and molecular analysis. These different markers provided evidence of four (karyology) to eight (morphology) discrete entities, whereas crossings showed various degrees of intersterility among the tested populations. Molecular species delimitation revealed a different number of candidate species, spanning from five (ABGD and K/θ) to 11 (GMYC). Such incongruences reflect the multifaceted evolutionary history of M. lineata s.l.and hamper the full taxonomic resolution of the complex. However, two candidate species were consistently validated by all of the markers and are described as new species: Monocelis algicolanov. sp. and M. exquisitanov. sp. The latter species appear to have a restricted distribution, and the possibility that meiofaunal taxa may be of conservation concern is discussed.