Search

Lucena, Rudá Amorim (2021): Discovery and re-description of the holotype of Pycnogonum gibberum (Pycnogonida: Pycnogonidae). Journal of Natural History 55 (1-2): 61-66, DOI: 10.1080/00222933.2021.1893402, URL: http://dx.doi.org/10.1080/00222933.2021.1893402

Sea spiders (Pycnogonida) are a small group of arthropods, sister to other chelicerates. They have an unusual adult bauplan, oligosegmented larvae, and a protracted postembryonic development. Pycnogonum litorale (Strøm, 1762) is an uncommonly long-lived sea spider with a distinctive protonymphon and adult anatomy. Although it was described ~250 years ago, little is known about its internal organization and development. We examined the anamorphic and early epimorphic development of this species using histology, light microscopy, and SEM, and provide the first comprehensive anatomical study of its many instars. Postembryonic development of P. litorale includes transformations typical of pycnogonids: reorganization of the larval organs (digestive, nervous, secretory), formation of the abdomen, trunk segments (+ appendages), primary body cavity and reproductive system. Specific traits include the accelerated articulation of the walking legs, formation of the subesophageal and posterior synganglia, and the system of twin midgut diverticula. In addition, P. litorale simultaneously lose the spinning apparatus and all larval appendages. We found that developmental changes occur in synchrony with changes in ecology and food sources. The transition from the anamorphic to the epimorphic period in particular is marked by considerable anatomical and lifestyle shifts. HIGHLIGHTS: Postembryonic development of P. litorale includes numerous anamorphic and epimorphic stages. The instars acquire abdomen, trunk segments, body cavity, and gonads, while losing all larval appendages. Developmental changes are synchronized with changes in lifestyle and food sources.

We describe a new pycnogonid species, Pantopipetta lenis sp. nov., collected from 141–152 m depth in southern Japan, North Pacific Ocean. Pantopipetta lenis closely resembles Pantopipetta auxiliata...

Developmental modes in pycnogonids are variable, even between congeneric species. Most of our previous knowledge of pycnogonid life history stages is based on morphological descriptions of the larval and those postlarval instars still retained on male ovigers, whereas the free-living instars are largely unknown. In this paper, a combination of morphological and molecular tools is used in order to link those postlarval instars retained on male ovigers to free-living specimens representing putatively later stages in the postembryonic development of a sea spider species. Male specimens of Nymphon australe carrying postembryonic instars were collected from localities off the South Shetland Islands, the Weddell Sea, the Drake Passage, the Bransfield Strait, and the Ross Sea. Three subsequent free-living instars were collected from Drake Passage. Nymphon australe hatch as a postlarval instar with two-articled palps and limb buds of the two first-walking legs. Postlarval instars 1 and 2 (both carried on the ovigers of males), postlarval instars 3 and 4, and a juvenile (all three free living) are all described and illustrated. A molecular phylogenetic based on the cox1 gene links all free-living stages to N. australe populations from the Antarctic Peninsula. The postlarval development of N. australe is discussed here and compared with those known from other pycnogonid species with similar development pathways.

Higher-level phylogenetics of Pycnogonida has been discussed for many decades but scarcely studied from a cladistic perspective. Traditional taxonomic classifications are yet to be tested and affinities among families and genera are not well understood. Pycnogonida includes more than 1300 species described, but no systematic revisions at any level are available. Previous attempts to propose a phylogeny of the sea spiders were limited in characters and taxon sampling, therefore not allowing a robust test of relationships among lineages. Herein, we present the first comprehensive phylogenetic analysis of the Pycnogonida based on a total evidence approach and Direct Optimization. Sixty-three pycnogonid species representing all families including fossil taxa were included. For most of the extant taxa more than 6 kb of nuclear and mitochondrial DNA and 78 morphological characters were scored. The most parsimonious hypotheses obtained in equally weighted total evidence analyses show the two most diverse families Ammotheidae and Callipallenidae to be non-monophyletic. Austrodecidae + Colossendeidae + Pycnogonidae are in the basal most clade, these are morphologically diverse groups of species mostly found in cold waters. The raising of the family Pallenopsidae is supported, while Eurycyde and Ascorhynchus are definitely separated from Ammotheidae. The four fossil taxa are grouped within living Pycnogonida, instead of being an early derived clade. This phylogeny represents a solid framework to work towards the understanding of pycnogonid systematics, providing a data set and a testable hypothesis that indicate those clades that need severe testing, especially some of the deep nodes of the pycnogonid tree and the relationships of ammotheid and callipallenid forms. The inclusion of more rare taxa and additional sources of evidence are necessary for a phylogenetic classification of the Pycnogonida. © The Willi Hennig Society 2006.

The marine biodiversity of the tropical northwestern Atlantic (TNWA) has been explored by many great naturalist expeditions. After more than one century of marine exploration, how well do we know its biodiversity? As a poorly studied taxon, sea spiders (Arthropoda: Pycnogonida Latreille, 1810) are excellent candidates to address this issue. Here, we report the results from the Madibenthos Survey conducted by the Museum national d’Histoire naturelle (Paris) on the coasts of Martinique during Fall 2016, where sea spiders were collected during 138 sampling events by different methods. A total of 67 pycnogonid species, including 13 species probably new to science, were distinguished during this survey, whereas only 20 species were previously known from Martinique. Relying on an extensive and intensive sampling, the collection of pycnogonids from the Madibenthos Survey is one of the richest for TNWA. DNA barcoding based on 172 new CO1 sequences helped to discriminate species with inconspicuous diagnostic characters and pointed out 11 additional possible cryptic species. Richness estimators and species rarity indicate that the diversity of Martinique still remains underestimated. Results suggest that Martinique is not a hot spot for sea spiders and that similar levels of biodiversity might be expected in most other islands of the TNWA if similar sampling methods were applied. Finally, we show that the Caribbean and Atlantic coasts of Martinique harbor different sea spider faunas.

We studied Pycnogonida, sea spiders, collected from 54 samples of Corallina officinalis belts in the vicinity of Pula and the Brijuni National Park representing both exposed and sheltered localities as well as different levels of human impact. Seven species were identified, namely Achelia echinata , A. langi , Tanystylum conirostre , Anoplodactylus angulatus , A. pygmaeus, Trygaeus communis , and Callipallene tiberi . As we used a quantifiable standard sample size of 5 cm 2 , we could perform a statistical analysis of species richness and abundances. The exposed low human impact sites showed a significantly higher amount of both, specimens and species than the sheltered high impact sites. C. tiberi and A. echinata showed a significant preference for exposed low impact sites while T. conirostre was equally distributed among the habitat subtypes.

Individuals of the marine chelicerate lineage Pycnogonida (sea spiders) show considerable regenerative capabilities after appendage injury or loss. In their natural habitats, especially the long legs of sea spiders are commonly lost and regenerated, as is evidenced by the frequent encounter of specimens with missing or miniature legs. In contrast to this, the collection of individuals with abnormally developed appendages or trunk regions is comparably rare. Here, we studied a remarkable malformation in a postlarval instar of the species Phoxichilidium femoratum (Rathke, 1799) and describe the external morphology and internal organization of the specimen using a combination of fluorescent histochemistry and scanning electron microscopy. The individual completely lacks the last trunk segment with leg pair 4 and the normally penultimate trunk segment bears only a single aberrant appendage resembling an extension of the anteroposterior body axis. Externally, the proximal units of the articulated appendage are unpaired, but further distally a bifurcation into two equally developed leg‐like branches is found. Three‐dimensional reconstruction of the musculature reveals components of two regular leg muscle sets in several of the proximal articles. This confirms interpretation of the entire appendage as a malformed leg and reveals an externally hidden paired organization along its entire proximodistal axis. To explain the origin of this unique malformation, early pioneering studies on the regenerative potential of pycnogonids are evaluated and (a) an injury‐induced partial fusion of the developing limb buds of leg pair 3, as well as (b) irregular leg regeneration following near complete loss of trunk segments 3 and 4 are discussed. Which of the two hypotheses is more realistic remains to be tested by dedicated experimental approaches. These will have to rely on pycnogonid species with established laboratory husbandry in order to overcome the limitations of the few short‐term regeneration studies performed to date. Deutsche Forschungsgemeinschaft http://dx.doi.org/10.13039/501100001659 National Science Foundation http://dx.doi.org/10.13039/100000001

Despite significant advances in invertebrate phylogenomics over the past decade, the higher-level phylogeny of Pycnogonida (sea spiders) remains elusive. Due to the inaccessibility of some small-bodied lineages, few phylogenetic studies have sampled all sea spider families. Previous efforts based on a handful of genes have yielded unstable tree topologies. Here, we inferred the relationships of 89 sea spider species using targeted capture of the mitochondrial genome, 56 conserved exons, 101 ultraconserved elements, and 3 nuclear ribosomal genes. We inferred molecular divergence times by integrating morphological data for fossil species to calibrate 15 nodes in the arthropod tree of life. This integration of data classes resolved the basal topology of sea spiders with high support. The enigmatic family Austrodecidae was resolved as the sister group to the remaining Pycnogonida and the small-bodied family Rhynchothoracidae as the sister group of the robust-bodied family Pycnogonidae. Molecular divergence time estimation recovered a basal divergence of crown group sea spiders in the Ordovician. Comparison of diversification dynamics with other marine invertebrate taxa that originated in the Paleozoic suggests that sea spiders and some crustacean groups exhibit resilience to mass extinction episodes, relative to mollusk and echinoderm lineages. Deutsche Forschungsgemeinschaft BR5039/3-1 National Science Foundation ANT-0551969, ANT-0440577 Ministerio de Economía y Competitividad POL2006-06399/CGL, CTM2012-39350- C02-01 Antarctic Science AA3010 National Science Foundation IOS-1552610

Ten intertidal sea spider (Pycnogonida) species from five families - Pycnogonidae (1 species), Ammotheidae (3 species), Callipallenidae (3 species), Phoxichilidiidae (2 species) and Endeidae (1 species) - are reported in the present study. Out of these ten species, four are first records from Pakistan, including a new species, Achelia karachiensis sp. nov. Intertidal zoanthid colonies have been found to provide shelter and food to a large and diverse Pycnogonid population at Sandspit and the Buleji, Karachi coast. This study highlights the significance of the intertidal fauna of pycnogonids of these areas and contributes to our understanding of the species diversity from the Karachi coast.

Despite significant advances in invertebrate phylogenomics over the past decade, the higher-level phylogeny of Pycnogonida (sea spiders) remains elusive. Due to the inaccessibility of some small-bodied lineages, few phylogenetic studies have sampled all sea spider families. Previous efforts based on a handful of genes have yielded unstable tree topologies. Here, we inferred the relationships of 89 sea spider species using targeted capture of the mitochondrial genome, 56 conserved exons, 101 ultraconserved elements, and three nuclear ribosomal genes. We inferred molecular divergence times by integrating morphological data for fossil species to calibrate 15 nodes in the arthropod tree of life. This integration of data classes resolved the basal topology of sea spiders with high support. The enigmatic family Austrodecidae was resolved as the sister group to the remaining Pycnogonida and the small-bodied family Rhynchothoracidae as the sister group of the robust-bodied family Pycnogonidae. Molecular divergence time estimation recovered a basal divergence of crown group sea spiders in the Ordovician. Comparison of diversification dynamics with other marine invertebrate taxa that originated in the Paleozoic suggests that sea spiders and some crustacean groups exhibit resilience to mass extinction episodes, relative to mollusk and echinoderm lineages.

13 pages, 2 figures, 2 tables, We aim to study the sea spider specimens collected from a deep cruise in the economic exploitation zone of Uruguay from 250 to 3800 m with the Marianne fishing vessel. We gather the information from deep-sea Uruguayan pycnogonids from literature together with the ones reported in a recent checklist of sea spiders present in Uruguayan waters published by Scarabino et al. (2019) and including some biogeographic considerations. This new cruise includes two new species for Uruguayan waters and a great enlarging new bathymetric distribution of an already identified species in the area

Specimens of Pycogonida belonging to the genus Ammothea collected during the Polarstern ANT-XXIX/3 cruise (January 22–March 18, 2013), to Bransfield Strait and Eastern Weddell Sea, were studied. Eight species are recognized in this collection: Ammothea bicorniculata, A. calmani, A. carolinensis, A. clausi, A. minor, A. nadiae n. sp., A. spinosa, and A. striata. Morphological variability is discussed. The bathymetric range of A. bicorniculata and A. minor is enlarged and A. bicorniculata is reported for only the second time. Ammothea nadiae n. sp. is described and compared with its closest congeners. An updated key to species is presented.

The Pycnogonida or sea spiders are exclusively marine invertebrates, numbering about 1,300 described species worldwide. Given their remarkable position in phylogeny as basal chelicerates or even basal euarthropods, the structure of their sense organs can reveal important characters, which—in a comparative framework—provide arguments to phylogenetic discussions and help to develop scenarios of evolutionary transformations. This review summarizes current knowledge and presents new original data on the sense organs in pycnogonids, that is, the eyes, the lateral sense organs and the ciliary or sensillar sense organs. Except for the eyes, there are not many detailed studies available. The ultrastructure of the R-cells of the four eyes located on the ocular tubercle is described as “pseudoinverted”. The eyes are innervated to two visual neuropils located in the protocerebrum. The features of the lateral sense organ, also located on the ocular tubercle, are hitherto not conclusively resolved, a chemo- or thermoreceptive function is suggested. Finally, an overview of the various ciliary or sensillar sense organs distributed all over the body is given and the fine structure of branched setae is shown for the first time. The morphology of the sense organs of pycnogonids is compared with that of other arthropod taxa and assessed against the background of current theories of arthropod evolution.

Nymphonella tapetis (Pycnogonida, Ascorhynchidae) is an endoparasitic sea spider affecting bivalves. Recently, sea spiders have been found on a massive scale in the commercially important Manila clams (Veneridae, Ruditapes philippinarum) in Japan (Tokyo Bay). Simultaneously, mass mortality has occurred in this area. Local fishers assumed that this mass mortality was caused by the parasitic sea spider, despite the effect of the parasite and parasite intensity on the host being unknown. To evaluate the susceptibility of the Manila clam to sea spider infestation and the impact on mortality levels, we established six treatments at different infection intensities (density of newly hatched larvae of sea spiders) over a 6-month long laboratory experiment. We monitored mortality and three susceptibility indices (clearance rate, sand-burrowing speed and adductor muscle strength) under sufficient food conditions. Parasitization by sea spider affected clearance rate and sand-burrowing speed. The pattern of parasitic intensity effects on survival of Manila clam hosts was shown to be dependent on the levels of parasite numbers, i.e. clams with lower parasitic levels (total of

Postembryonic developmental stages of an endoparasitic pycnogonid, Ammothella biunguiculata in Izu Peninsula, Japan are described. Eleven stages were identified beginning with a protonymphon larva attached to the male oviger. We found endoparasitic individuals in the host actinian from the second to tenth instar, and forms in the ninth stage to adult were found free-living. This indicates a transition from being endoparasitic to free-living during the ninth to tenth instar stages. The first instar protonymphon attached to the adult male oviger has a gland duct on the anterior margin of each chelifore scape which completely disappears with the second instar. The disappearance of the chelifore gland duct coincides with the beginning of an endoparasitic stage in the development of this species. Although the larval morphology and the postembryonic development of pycnogonids have been summarized by several authors, the present study concludes that much remains to be learnt.

The present contribution describes and discusses the observed morphological variability of pycnogonids in the genus Pallenopsis collected during the Italica XIX cruise to Victoria Land, Ross Sea, Antarctica (3 Feb-4 Mar, 2004). Four species are recognized in this collection and two of them are proposed as new species Pallenopsis gracilis n. sp. and Pallenopsis rotunda n. sp. The new species are described, illustrated, and compared morphologically with their closest congeners. The morphological diagnostic characters of the subgenus Bathypallenopsis are discussed and considered insufficient to justify a separation into subgenera. The observed morphological variability is instead proposed to be accommodated in one inclusive genus Pallenopsis. Finally, we propose an updated dichotomous key to all currently recognized Antarctic and Sub-Antarctic Pallenopsis species (adult form), and discuss some of the more disputable taxa, highlighting the lack of knowledge of the morphological and molecular characters of this genus.

Background Sea spiders (Pycnogonida) are an abundant faunal element of the Southern Ocean (SO). Several recent phylogeographical studies focused on the remarkably diverse SO pycnogonid fauna, resulting in the identification of new species in previously ill-defined species complexes, insights into their genetic population substructures, and hypotheses on glacial refugia and recolonization events after the last ice age. However, knowledge on the life history of many SO pycnogonids is fragmentary, and early ontogenetic stages often remain poorly documented. This impedes assessing the impact of different developmental pathways on pycnogonid dispersal and distributions and also hinders pycnogonid-wide comparison of developmental features from a phylogenetic-evolutionary angle. Results Using scanning electron microscopy (SEM) and fluorescent nuclear staining, we studied embryonic stages and postembryonic instars of three SO representatives of the taxon Pallenopsidae (Pallenopsis villosa, P. hodgsoni, P. vanhoeffeni), the development of which being largely unknown. The eggs are large and yolk-rich, and the hatching stage is an advanced lecithotrophic instar that stays attached to the father for additional molts. The first free-living instar is deduced to possess at least three functional walking leg pairs. Despite gross morphological similarities between the congeners, each instar can be reliably assigned to a species based on body size, shape of ocular tubercle and proboscis, structure of the attachment gland processes, and seta patterns on cheliphore and walking legs. Conclusions We encourage combination of SEM with fluorescent markers in developmental studies on ethanol-preserved and/or long term-stored pycnogonid material, as this reveals internal differentiation processes in addition to external morphology. Using this approach, we describe the first known cases of pallenopsid development with epimorphic tendencies, which stand in contrast to the small hatching larvae in other Pallenopsidae. Evaluation against current phylogenetic hypotheses indicates multiple gains of epimorphic development within Pycnogonida. Further, we suggest that the type of development may impact pycnogonid distribution ranges, since free-living larvae potentially have a better dispersal capability than lecithotrophic attaching instars. Finally, we discuss the bearing of pycnogonid cheliphore development on the evolution of the raptorial first limb pair in Chelicerata and support a multi-articled adult limb as the plesiomorphic state of the chelicerate crown group, arising ontogenetically via postembryonic segmentation of a three-articled embryonic limb. Electronic supplementary material The online version of this article (10.1186/s40851-018-0118-7) contains supplementary material, which is available to authorized users.

Sea spiders (Pycnogonida) constitute a group of marine benthic arthropods that has a particularly high species diversity in the Southern Ocean. The “longitarsal” group of the sea spider family Colossendeidae is especially abundant in this region. However, this group also includes some representatives from other oceans, which raises the question where the group originates from. Therefore, we here investigated the phylogeny of the group with a hybrid enrichment approach that yielded a dataset of 1607 genes and over one million base pairs. We obtained a well-resolved phylogeny of the group, which is mostly consistent with morphological data. The data support an Antarctic origin of the longitarsal Colossendeidae and multiple dispersal events to other regions, which occurred at different timescales. This scenario is consistent with evidence found in other groups of marine invertebrates and highlights the role of the Southern Ocean as a source for non-Antarctic biota, especially of the deep sea. Our results suggest an initially slow rate of diversification followed by a more rapid radiation possibly correlated with the mid-Miocene cooling of Antarctica, similar to what is found in other taxa.

Asari clam is one of the most important fishery resources in Japan. The clam has been utilized as not only a food but also a recreational harvestable species. Main habitat of the asari extends from tidal area to shallow subtidal area of sandy soft bottom. Asari is harvested by hand raking or short-handled basket raking in tidal areas and long-handled basket raking, dredging, helmet diving in subtidal areas. Major areas of asari harvest are Tokyo Bay, Mikawa Bay, Ise Bay, Suo Nada and Ariake Bay. Annual harvest of the asari in the major areas, however, decreased largely after late 1980s except Mikawa Bay. Stock decline of natural asari is remarkable and seriously problematic for coastal fisheries in Japan. Along with application of the various countermeasures intended to restore the natural stocks of asari, a number of factors have been discussed to be responsible for the asari decline. Some of these are decrease of suitable habitat areas, deterioration of water and substrate quality, erosion of the clams by strong wave disturbance, new predators and parasites, oligotrophication, and excess harvesting. We still have not determined significant factors responsible for the asari decline in Japan. In the discussed factors, we compared the effect of five factors; blue tide, river flood, strong wave disturbance, infection of sea spider, and hypoxia on clam larvae, based on the studies in Tokyo Bay. The impacts (mortalities) of the factors for the total asari stock by the blue tide in Sanbanze in 2008, the river flood in Sanbanze in 2007, winter wave mortality in the three areas in 2009–2010, and infection of sea spider in Kisarazu in 2009 were calculated to be 0.16，0.07，0.95，0.30, respectively. Among these factors, winter wave mortality showed the most significant effect on the asari stock in Tokyo Bay. Comparing the properties of the estimated factors, the factors that affected wider areas and longer period with higher mortality were supposed to give severer impact for the asari stock. In order to take effective countermeasures for the decline of asari stock in Japan, further assessment of the impact of the suspected factors is required to be progressed.

Pycnogonids (sea spiders) are commonly collected at bathyal and abyssal depths all around the world; however, little is known about species from deep-water habitats. The present study explores the life history of Nymphon hirtipes collected in northeastern Newfoundland between 700 and 1450 m depth and monitored in mesocosms for over 2 years. The pycnogonids were found in association with octocorals, hydrozoans and sea anemones. Adult females developed mature oocytes between June and August. Paired mating followed by oviposition occurred between early July and mid-October. Up to three egg masses were brooded by each male. It took a maximum of 4 months for the propagules to hatch from the egg mass. Offspring developed from walking leg-bearing larvae to early juveniles over the next ~5 months before leaving the male’s protection. Mating and oviposition coincided with the highest water temperatures of the annual cycle and dispersal of juveniles occurred in spring, when phytodetritus deposition was high and as ocean temperature rose markedly. All the females died after oviposition and the males ~9 months later, after juvenile dispersal. The sex ratio of mature individuals (~55–60 mm leg span) was 2 males for 3 females. Fecundity was estimated to be 184–288 eggs per adult female. Adults of N. hirtipes were seen feeding on hydrozoan polyps, small sea anemones (Stephanauge nexilis) and nudibranchs. Larvae and early juveniles did not feed while brooded by the male. Upon dispersal, their feeding apparatus became functional and they began feeding on hydrozoan polyps. After 13 months of growth post hatching, the juveniles reached ~21 mm leg span. Curve fitting estimated that ~7 years are required to reach the adult size of 55 mm leg span.