Variation in Nymphon brevirostre (Hodge), and the status of N. rubrum (Hodge) (Arthropoda: Pycnogonida)

A series of morphometric analyses conducted on specimens of Nymphon brevirostre Hodge and Nymphon rubrum Hodge from British waters has shown that the characteristics variously used in the past to distinguish these two species are invalid. In this material these characteristics range from the extreme of the typical N. brevirostre form to that of the typical N. rubrum form, with most of the specimens exhibiting an intermediate morphology. Multivariate analyses showed no significant splitting of the material into two groups which might correlate to Hodge's two species. Protonymphon larvae from males of both typical extreme forms were morphologically identical. It is concluded that N. rubrum is a junior synonym of N. brevirostre , a variable species whose morphology tends towards the rubrum form as the animal grows.     

Larvae of three sea spider species of the genus <Emphasis Type="Italic">Nymphon</Emphasis> (Arthropoda: Pycnogonida) from the White Sea

Larvae of three Nymphon species are described. For Nymphon brevirostre Hodge, 1863 and Nymphon micronyx Sars, 1888 protonymphon larvae are examined, larvae of N. micronyx have been studied for the first time. The fine external morphology of larvae is studied in detail using SEM methods; the internal structure of N. brevirostre protonymphon is additionally described. SEM data on external morphology and fine surface structures are given for several postembryonic stages of Nymphon grossipes Fabricius, 1780. Different types of postembryonic development are revealed in examined Nymphon species. N. brevirostre and N. micronyx have a “typical protonymphon” developmental pathway. A special developmental pathway named “lecithotrophic protonymphon” is suggested for N. grossipes and other Nymphonidae having large yolky eggs.     

Salinity and temperature tolerance by Nymphon gracile (Leach) and Achelia echinata (Hodge) (Pycnogonida)

The tolerance of two species of pycnogonid, Achelia echinata (Hodge) and Nymphon gracile (Leach) to a range of salinities, temperatures and relative humidities has been examined. In most instances N. gracile is more resistant than A. echinata and indeed is found higher in the littoral zone during the warmer months and thus likely to be subjected to greater environmental fluctuation.     

Structure of the body cavity of the sea spider <Emphasis Type="Italic">Nymphon brevirostre</Emphasis> Hodge, 1863 (Arthropoda: Pycnogonida)

The microscopic anatomy and ultrastructure of the body cavity and adjacent organs in the sea spider Nymphon brevirostre Hodge, 1863 (Pycnogonida, Nymphonidae) were examined by transmission electron microscopy. The longitudinal septa subdividing the body cavity are described: (1) Dohrn’s horizontal septum, (2) lateral heart walls, and (3) paired ventral septa consisting of separate cellular bands. The body cavity is a hemocoel, it has no epithelial lining and is only bordered by a basal lamina. The epidermis, heart, and Dohrn’s septum are not separated from each other by basal laminae and may have a common origin. The cellular bands forming the longitudinal ventral septa are not covered with the basal lamina and presumably derive from cells belonging to the hemocoel. The roles of the morphological structures studied for the circulation of hemolymph are discussed. The gonad lies inside Dohrn’s septum, it is covered with its own basal lamina and surrounded by numerous lacunae of the hemocoel entering the septum. The gonad wall is formed with a single layer of epithelium. The same epithelial cells form the gonad stroma. The gonad cavity is not lined with the basal lamina; muscle cells are present in the gonad wall epithelium, thus rendering the lumen similar to a coelomic cavity. Freely circulating cells of two types are found in the hemocoel: small amebocytes containing electronic-dense granules that are similar to granulocytes of other arthropods, as well as hemocytes with large vacuoles of varying structure that are comparable with plasmatocytes; however some of these may be activated granulocytes.     

The (not very) typical protonymphons of Pycnogonum litorale

Sea spiders are unique and poorly known marine chelicerates. Their larvae are even less studied, especially at the ultrastructural level. Here, we examined the hatchlings of Pycnogonum litorale (Strøm, 1,762) using histology, SEM and TEM. Existing classifications place these larvae among “typical” protonymphons, together with Nymphon brevirostre. Our results, however, revealed major differences between the two species. Hatchlings of P. litorale are endotrophic for 1–2 weeks, with yolk deposits in the body wall and a reduced secretory apparatus. They lack a body cavity, demonstrate an unusual modification of the midgut sheath cells and a complex subesophageal ganglion, which includes neuromeres of the prospective walking legs 1. These larvae also possess well-developed glia and complex sensory structures: eyes, V-shaped mechanoreceptive bristles, integrated chemo- and mechanoreceptors, and three types of concealed mechanoreceptors embedded into the body wall and only seen on the sections. In this paper we also propose a new interpretation of the pycnogonid larval types: we present a set of traits useful for diagnosis and a preliminary classification. Finally, we discuss the complexity of glial types in sea spiders and other arthropods.     

Synopsis of a new collection of sea spiders (Arthropoda: Pycnogonida) from the Ross Sea,Antarctica

The biodiversity research expedition TAN0204 with RS Tangaroa to the Ross Sea in 2004 yielded a new collection of 2,687 specimens of pycnogonids. As much as 25 different species encompassing 14 genera and eight families were identified and their records are discussed herein. The collection is archived in the Marine Invertebrate Collection of the New Zealand National Institute of Water and Atmospheric Research (NIWA). The majority (69%) of specimens are from the Nymphon australe group (Nymphonidae), although species richness and abundance varied among the stations sampled. The collection includes several specimens from polymerous taxa; Pentanymphon antarcticum (Nymphonidae), Decolopoda australis (Colossendeidae) and Pentapycnon bouvieri (Pycnogonidae). All species were classified based on morphological characters, and DNA sequences (from the 18S, 12S, 16S and COI regions) for 21 of the representative morphotypes are given. The DNA sequences confirmed the species-level distinctiveness of these morphotypes. No species new to science were identified, although further detailed morphometric and/or molecular analyses may reveal cryptic or sibling species, especially in species such as the highly abundant Nymphon australe group. An erratum to this article can be found at     

Postembryonic development of Nymphon unguiculatum Hodgson 1915 (Pycnogonida, Nymphonidae) from the South Shetland Islands (Antarctica)

Male specimens of the sea spider species Nymphon unguiculatum, carrying eggs, larvae and postlarvae in various stages of development, were collected off the South Shetland Island at water depths between 112 and 472 m in austral summer 2006/2007. Here, we describe the external morphology of four postembryonic stages (protonymphon, instar 1, instar 2, and instar 3) carried by these specimens. We found that (1) protonymphon larvae hatch from the eggs; (2) larvae and postlarval stages have yolk reserves and are characterized by a relatively large size (average body lengths of 0.46, 0.55, 0.65 and 0.73 mm in the successive stages); (3) postlarvae remain on the ovigerous legs of males during several moults; (4) a spinning apparatus is present; (5) the development of walking legs is sequential. The larval and postlarval development of N. unguiculatum is compared with that known from other pycnogonid species.     

Pycnogonid sperm. An example of inter- and intraspecific axonemal variation

Summary The flagella of the motile sperm cells of Nymphon leptocheles and N. rubrum (Pyonogonida, Arthropoda) exhibit a 12+0 and a 9+0 axoneme pattern, respectively. Central tubules, central sheath, spokes and arms are absent. The doublets are connected by a circular nexus. The functional significance of this axonemal composition is discussed. Aberrant axonemes occurring in high frequencies both within the species and within single specimens are probably explained by the loose axonemal connection, due to the absence of a central complex. This absence is further suggested to have facilitated the evolution from the 9+0 type to the 12+0 type.     

Postembryonic development of pycnogonids: A deeper look inside

Sea spiders form a small, enigmatic group of recent chelicerates, with an unusual bodyplan, oligosegmented larvae and a postembryonic development that is punctuated by many moults. To date, only a few papers examined the anatomical and ultrastructural modifications of the larvae and various instars. Here we traced both internal and external events of the whole postembryonic development in Nymphon brevirostre HODGE 1863 using histology, SEM, TEM and confocal microscopy. During postembryonic development, larvae of this species undergo massive reorganization: spinning apparatus and chelar glands disappear; larval legs redifferentiate; three new segments and the abdomen are formed with their corresponding internal organs and appendages; circulatory and reproductive systems develop anew and the digestive and the nervous systems change dramatically. The body cavity remains schizocoelic throughout development, and no traces of even transitory coeloms were found in any instar. In Nymphon brevirostre, just like in Artemia salina LINNAEUS 1758 the heart arises through differentiation of the already existing schizocoel, and thus the circulatory systems of arthropods and annelids are not homologous. We found that classical chelicerate tagmata, prosoma and opisthosoma, are inapplicable to adult pycnogonids, with the most striking difference being the fate and structure of the seventh appendage-bearing segment.     

Hox genes in sea spiders (Pycnogonida) and the homology of arthropod head segments

The pycnogonids (or sea spiders) are an enigmatic group of arthropods, classified in recent phylogenies as a sister-group of either euchelicerates (horseshoe crabs and arachnids), or all other extant arthropods. Because of their bizarre morpho-anatomy, homologies with other arthropod taxa have been difficult to assess. We review the main morphology-based hypotheses of correspondence between anterior segments of pycnogonids, arachnids and mandibulates. In an attempt to provide new relevant data to these controversial issues, we performed a PCR survey of Hox genes in two pycnogonid species, Endeis spinosa and Nymphon gracile, from which we could recover nine and six Hox genes, respectively. Phylogenetic analyses allowed to identify their orthology relationships. The Deformed gene from E. spinosa and the abdominal-A gene from N. gracile exhibit unusual sequence divergence in their homeodomains, which, in the latter case, may be correlated with the extreme reduction of the posterior region in pycnogonids. Expression patterns of two Hox genes (labial and Deformed) in the E. spinosa protonymphon larva are discussed. The anterior boundaries of their expression domains favour homology between sea spider chelifores, euchelicerates chelicerae and mandibulate (first) antennae, in contradistinction with previously proposed alternative schemes such as the protocerebral identity of sea spider chelifores or the absence of a deutocerebrum in chelicerates. In addition, while anatomical and embryological evidences suggest the possibility that the ovigers of sea spiders could be a duplicated pair of pedipalps, the Hox data support them as modified anterior walking legs, consistent with the classical views.Supplementary material is available for this article at and is accessible for authorized users.Guest editors Jean Deutsch and Gerhard Scholtz     

Molecules,morphology, missing data and the phylogenetic position of a recently extinct madtom catfish (Actinopterygii: Ictaluridae)

Taxa missing large amounts of data pose challenges that may hinder the recovery of a well‐resolved, accurate phylogeny and leave questions surrounding their phylogenetic position. Systematists commonly have to contend with one or two species in a group for which there is little or no material available suitable for recovering molecular data. It is unclear whether these taxa can be better placed using analyses based on morphological data only, or should be included in broader analyses based on both morphological and molecular data. The extinct madtom catfish Noturus trautmani is known from few specimens for which molecular data are unavailable. We included this taxon in parsimony and Bayesian analyses of relationships of madtom catfishes based on a combination of morphological and molecular data. Results indicate that using a combination of morphological and molecular data does a better job at providing a phylogenetic placement for N. trautmani than morphology alone, even though it is missing all of its molecular characters. We provide a novel hypothesis of relationships among Noturus species and recommendations for classification within the group. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 60–75.     

A new species of Armodoris (Mollusca, Gastropoda, Nudibranchia, Akiodorididae) from McMurdo Sound, Antarctica

Recently collected specimens of Armodoris from McMurdo Sound, Antarctica, were morphologically examined and sequenced. Comparison between this new material and literature sources revealed that it belongs to an undescribed species, Armodoris anudeorum. Although this new species is externally very similar to Armodoris antarctica (the only previously known species of Armodoris), these two species differ in several details of their external morphology, and particularly in their reproductive anatomy and radular morphology. This is the second known species of Armodoris; thus, this paper doubles the known diversity of this exclusively Antarctic group.     

Two closely related species of Nipponaphis (Hemiptera: Aphididae) that migrate between Distylium racemosum and Machilus trees in Japan

Nipponaphis loochooensis Sorin, 1996 and N. machilicola (Shinji, 1941) form fig‐shaped or globular galls on Distylium racemosum in southern Japan. Nipponaphis machilicola migrates to the lauraceous evergreen Machilus thunbergii. Nipponaphis loochooensis has also been supposed to migrate to M. thunbergii, but its secondary‐host generation has not been found in the field to date. Through sampling Nipponaphis aphids from trees of M. japonica in addition to M. thunbergii, and sequencing their mitochondrial DNA, we found that the two species form colonies on twigs of both Machilus species, and that the two at times colonize on the same trees and even form mixed colonies. Only N. loochooensis forms colonies on leaves of M. japonica, but neither species colonizes on leaves of M. thunbergii. The secondary‐host generations of the two species could be clearly discriminated from each other, based on morphology. It was confirmed by examining the type specimens of Nipponaphis amamiana Takahashi, 1962 that the name is a junior synonym of N. machilicola.     

The synonymy of Haplochromis pharyngalis and Haplochromis petronius (Cichlidae)

Haplochromis pharyngalis and Haplochromis petronius, two endemic cichlids from the Lake Edward system (Uganda, Democratic Republic of the Congo), are very similar in general morphology but have been reported to differ in pharyngeal jaw morphology and distribution. This study analysed 51 morphometrics and various qualitative characteristics of 48 specimens from different localities. The morphological traits of both species strongly overlap, and differences in the pharyngeal jaw morphology correspond to a geographic morphocline. We conclude that all specimens belong to one valid species, H. pharyngalis, and consider H. petronius to be a synonym.     

Morphological variation in British Atherinids, and the status of Atherina presbyter Guvier (Pisces: Atherinidae)

A series of morphometric and meristic analyses conducted on specimens of sand smelt, Atherina , from populations around the British Isles, and including classic A. boyeri and A. presbyter forms, has shown that the characteristics variously used in the past to distinguish these two species are invalid. Multivariate analyses showed no significant splitting of the material into two groups which might correlate to these two species; classic A. boyeri and the very large Atlantic coast A. presbyter individuals represent the tails of a continuum of form. From these and published data, it is concluded that the A. boyeri morphology varies under the influence of conditions of temperature and salinity during embryo development, and the isolation of populations maintains comparative morphological distinctions resulting from local selection and random genetic drift.     

Phylogenetics and seed morphology of African Nymphoides (Menyanthaceae)

We evaluated the boundaries among Nymphoides (Menyanthaceae) species in Africa, a region where the genus has received relatively little attention. We gathered morphological data from seeds using light and scanning electron microscopy, and we conducted molecular phylogenetic analyses using nuclear sequence data from the internal transcribed spacer (ITS) region. Morphological and molecular features distinguished six species and affirmed their respective geographic ranges. No African specimens were attributable to Nymphoides indica, even though this paleotropical species previously had been understood to grow in Africa. We establish the new combination Nymphoides senegalensis (G. Don) Tippery, based on an established African basionym, to accommodate the specimens formerly identified as N. indica. Phylogenetic analyses resolved two distinct clades containing African species. The majority of African species are closely related to neotropical species, with which they share similar petal ornamentation. The morphologically distinct N. ezannoi shares floral and phylogenetic similarity with species from North America and Asia. Results presented here support prior hypotheses that allopolyploid species in the Americas may have originated from one or more parental lineages in Africa. Seed morphological characters remain some of the most reliable features for identifying species, particularly for herbarium specimens lacking observable floral characters.     

Systematics of Cladophora spp. (Chlorophyta) from North Carolina,USA, based upon morphology and DNA sequence data with a description of Cladophora subtilissima sp. nov.

Identification of Cladophora species is challenging due to conservation of gross morphology, few discrete autapomorphies, and environmental influences on morphology. Twelve species of marine Cladophora were reported from North Carolina waters. Cladophora specimens were collected from inshore and offshore marine waters for DNA sequence and morphological analyses. The nuclear‐encoded rRNA internal transcribed spacer regions (ITS) were sequenced for 105 specimens and used in molecular assisted identification. The ITS1 and ITS2 region was highly variable, and sequences were sorted into ITS Sets of Alignable Sequences (SASs). Sequencing of short hyper‐variable ITS1 sections from Cladophora type specimens was used to positively identify species represented by SASs when the types were made available. Secondary structures for the ITS1 locus were also predicted for each specimen and compared to predicted structures from Cladophora sequences available in GenBank. Nine ITS SASs were identified and representative specimens chosen for phylogenetic analyses of 18S and 28S rRNA gene sequences to reveal relationships with other Cladophora species. Phylogenetic analyses indicated that marine Cladophorales were polyphyletic and separated into two clades, the Cladophora clade and the “Siphonocladales” clade. Morphological analyses were performed to assess the consistency of character states within species, and complement the DNA sequence analyses. These analyses revealed intra‐ and interspecific character state variation, and that combined molecular and morphological analyses were required for the identification of species. One new report, Cladophora dotyana, and one new species Cladophora subtilissima sp. nov., were revealed, and increased the biodiversity of North Carolina marine Cladophora to 14 species.     

Facilitation of Nothofagus antarctica (Fagaceae) seedlings by the prostrate shrub Empetrum rubrum (Empetraceae) on glacial moraines in Patagonia

Abstract Although positive interactions among plants are believed to be common in primary succession, they have rarely been demonstrated on glacial moraines. In Patagonia, the prostrate shrub Empetrum rubrum is dominant in early succession in recently deglaciated valleys, eventually being replaced by the small tree Nothofagus antarctica. This study experimentally evaluates the effects of E. rubrum shrubs on density, biomass, growth and survival of N. antarctica seedlings. The initial density of the seedlings was significantly higher under the canopy of Empetrum than in open areas. A removal experiment was performed to evaluate the influence of Empetrum on survival and growth of seedlings that germinated naturally beneath the canopy of this species. The shrub cover was removed from half of a sample of randomly selected seedlings, and left intact above the remainder. One year later, seedlings with an intact cover of E. rubrum showed significantly higher survival, growth and leaf number than seedlings which had been exposed. The results suggest that the presence of E. rubrum facilitates the establishment of N. antarctica in post‐glacial succession, mainly as a result of more favourable microclimatic conditions under the shrub canopy.     

Analysis of the clinal variation pattern inSilene latifolia (Caryophyllaceae) pollen morphology

Reticulate and microechinate pollen forms withinSilene latifolia (S. alba, S. pratensis) were first described from N. America, where the species is an introduced weed. A previous study showed that the two forms also exist in Europe and intergrade along a zone roughly congruent with intergradation zones in seed morphology and flavone glycosylation genotypes. The present survey of pollen from herbarium specimens is more extensive and covers localities from nearly the whole of the species' native Eurasian and North African range. The first axis of a detrended correspondence analysis (DCA) of qualitative characterstate data for 11 pollen morphological characters shows a gradient between the two extreme exine types, reticulate and microechinate. When mapped, the co-ordinates of pollen samples on this axis indicate a broad zone of intergradation which coincides approximately with the somewhat sharper transition between low- and high-tubercle seeds in C. and N. Europe, but becomes diffuse in Italy and the Mediterranean region and diverges from the seed transition zone in SW. Asia and the Middle East. The biological significance of the pollen morphs is unknown; the variation pattern in pollen morphology, unlike that in seeds, is not consistently correlated with macroclimate.