Rostrate pedicellariae: A morphologically distinct form of echinoid test appendage

The term rostrate was introduced by Mortensen ('07) to describe a type of pedicellaria he found in spatangoids. These pedicellariae resemble tridentate ones but have arching valves. Unlike the main categories of echinoid pedicellariae, no clear diagnosis of the rostrate form exists. This work examines the detailed morphology of the valves of rostrate pedicellariae observed by light and scanning electron microscopy and compares the shapes and dimensions of their component parts with tridentate pedicellariae. The data reveal considerable differences between the two, which warrant the recognition of rostrate pedicellariae as a distinct form. A diagnosis is given. © 1993 Wiley-Liss, Inc.     

Taxonomic significance of spine morphology in the echinoid genera Diadema and Echinothrix

Abstract. The spine morphology of all established species of Diadema and Echinothrix, including 2 color morphs of E. calamaris, were examined externally and internally via transverse sectioning to identify diagnostic species features and to assess the morphological relationship between species. Forty‐nine different morphological characters were measured and analysed using ordination by multi‐dimensional scaling (MDS) and cluster analysis. Specimens of Diadema paucispinum and D. setosum had very distinct spine structures. In D. paucispinum, the spines were more robust than those of other species of Diadema. This was evident in the spine's internal structure, with large, closely packed solid wedges, a small axial cavity, and rings of trabeculae throughout the spine's length. The spines in D. setosum were distinctive because of their length in relation to test size and the reduced flaring of their verticillations. The spines of other members of this genus were very similar to each other. Without careful sectioning, the spines from specimens of D. antillarum, D. ascensionis, D. mexicanum and D. savignyi were difficult to differentiate. The internal structures of spines for each species did, however, possess a combination of features that differentiated the species. Such features included the shape, orientation, and number of solid wedges, the presence or absence of spokes and rings of trabeculae between the solid wedges, and the presence or absence of tissue within the axial cavity. Individuals of Diadema palmeri also had spines morphologically similar to other species, however, the red pigmentation of these spines (in life and when preserved) made them easily distinguishable. The spine structures of the 2 species of Echinothrix were starkly different, while the white and brown color morphs of E. calamaris had morphologically distinctive ambulacral and interambulacral spines. The blunt, open‐tipped interambulacral spines, with reticular tissue present in the axial cavity of the white color morph, were easily distinguished from the pointed, closed‐tipped spines, with a hollow axial cavity found in the brown color morph. Such differences indicate that the brown color morph is either a subspecies or a separate species. Taken together the data show that each species has significant morphological differences in the structure of the spines. It is evident from our data that spine morphology is a useful tool to differentiate these commonly confused species.     

Lunar periodicities of diadematid echinoids breeding in Fiji

Gonad indices for two species of Diadema and two species of Echinothrix, including two color morphs of Echinothrix calamaris, are described for a 12-month period on Sosoikula Reef and Nukubuco Reef, Viti Levu, Fiji. Seasonal fluctuations in salinity (36.11±0.88 ppt) and water temperature (26.35±0.91°C) occurred. Gonad index data showed monthly reproductive rhythms, closely attuned to the lunar cycle. Diadema savignyi and Echinothrix diadema spawned on the full moon, and Diadema setosum and E. calamaris (white and brown color morphs) spawned on the new moon. Breeding periodicities coincided with the spring tides, thus maximizing chances of fertilization and dispersal. Such breeding cycles indicate how closely related species can co-exist with minimum risk of hybridization. Unusual sex ratios were found for all species, with an exceptionally low incidence of males. Reasons for this deviation from the typically reported 1:1 sex ratio may relate to the exceptionally high levels of tributyltin (TBT) recorded in Suva Harbor.     

Grazing preferences of diadematid echinoids in Fiji

Grazing preferences of two species of Diadema and two species of Echinothrix, including two colour morphs of Echinothrix calamaris, were investigated by gut contents analysis, in situ feeding observations, and grazing preferences trials in aquaria. Grazing preferences were compared to the distribution and abundance of diadematid sea urchins and the percentage cover of algal and seagrass species throughout Sosoikula Reef and Nukubuco Reef, Fiji. Results showed that grazing was selective, with distinct preferences between sea urchin genera, species, and colour morphs of E. calamaris. Preferred species of algae were non-calcareous, with reportedly low concentrations of tannins, phenols and bioactive compounds. Both Diadema savignyi and D. setosum selected Codium geppiorum as their most preferred species, followed by Hydroclathrus clathratus. All Echinothrix favoured Hy. clathratus, with both colour morphs of E. calamaris next selecting Padina pavonica and E. diadema selecting green filamentous algae. The seagrass species Syringodium isoetifolium was only grazed in significant quantities by E. calamaris (b) and E. diadema. This reflected species distributions in the seagrass bed. Peak abundances of diadematid sea urchins coincided with many of their grazing preferences at their maximum percentage cover. However, only E. diadema and D. savignyi had significant correlations with preferred algal/seagrass species (E. diadema with Co. geppiorum, and D. savignyi with G. marginata) throughout the subhabitats identified on the reefs.     

Structural characteristics of marsupial brood pouches of the antarctic sea urchins Abatus nimrodi and Abatus shackletoni (Echinoidea: Spatangoida)

Marsupial spines, tubercles, and pedicellariae of the antarctic brooding spatangoids Abatus nimrodi and Abatus shackletoni have been examined by scanning electron microscopy. Individual brood pouches of A. nimrodi may hold up to 28 embryos and juveniles and those of A. shackletoni may hold up to 38 lecithotrophic embryos and juveniles. Juveniles can be divided into (a) those with early development of external elements and a mean size of 2.0 and 1.6 mm, respectively, and (b) those equipped with fully developed external elements and a mean length of 4.3 and 2.8 mm, respectively. Mean diameters of aboral brood pouch openings of A. nimrodi and A. shackletoni were 6.5 and 4.1 mm, respectively. Brood pouches contain tall, distally enlarged spines, and smaller, layered cover-spines, which form a protective arch over the marsupia. There are also slender brood-pouch-bottom spines, which have an extremely thickened spinal epidermis. A. nimrodi has mainly bidentate, but also triand quadrodentate pedicellariae. A. shackletoni has two forms of tridentate, rostrate, and globiferous pedicellariae. In A. shackletoni, marsupial spine density is significantly lower than in A. nimrodi. These differences may be related to distinct sediment characteristics in their respective habitats. © 1993 Wiley-Liss, Inc.     

Fine structure and presumed functions of the pedicellariae of Echinocardium cordatum (Echinodermata,Echinoida)

Summary Tridactylous, trifoliate, and globiferous pedicellariae occur on the body surface of Echinocardium cordatum. Tridactyles have three forms: the typical, the rostrate, and the large forms. Both typical and rostrate tridactyles and trifoliates occur all around the echinoid body (trifoliates are, however, 4 times more numerous than tridactyles). Large tridactylous and globiferous pedicellariae are restricted to the peribuccal area.As a general rule tridactyles and trifoliates are similar in morphology. The distal part of the valves forms an open blade and bears lateral teeth and/or denticles (single or in combs). The stalk consists of a rigid proximal part supported by an axial rod and a flexible distal part which includes an axial fluid-filled cavity. The cavity is surrounded by muscle fibers and acts as an hydroskeleton, allowing the undulating-coiling movements of the flexible part of the stalk. Trifoliates are always active while tridactyles react only to direct or indirect mechanical stimulation.The valves of the globiferous pedicellariae have a tubular distal part whose upper opening is surrounded by teeth. There is no differentiated venom gland but a cluster of epithelial glandular cells located at the level of the valve upper opening. A small ciliary pad occurs just below the glandular cluster. Globiferous stalks are not flexible, being supported for their full length by an axial rod. Globiferous pedicellariae appear to be sensitive only to chemical stimulation.The presumed functions of E. cordatum pedicellariae are (1) cleaning of the body surface and ciliary structures (trifoliates), (2) protection against sedimenting particles (tridactyles), and (3) defense of the peribuccal area against potential small predators (globiferous pedicellariae).     

POPULATION STRUCTURE AND SPECIATION IN TROPICAL SEAS: GLOBAL PHYLOGEOGRAPHY OF THE SEA URCHIN DIADEMA

Abstract.— The causes of speciation in the sea are rarely obvious, because geographical barriers are not conspicuous and dispersal abilities or marine organisms, particularly those of species with planktonic larvae, are hard to determine. The phylogenetic relations of species in cosmopolitan genera can provide information on the likely mode of their formation. We reconstructed the phylogeny of the pantropical and subtropical sea urchin genus Diadema, using sequences of mitochondrial DNA from 482 individuals collected around the world, to determine the efficacy of barriers to gene flow and to ascertain the history of possible dispersal and vicariance events that led to speciation. We also compared 22 isozyme loci between all described species except D. palmeri. The mitochondrial DNA data show that the two deepest lineages are found in the Indian and West Pacific Oceans. (Indo‐Pacific) Diadema setosum diverged first from all other extant Diadema, probably during the initiation of wide fluctuations in global sea levels in the Miocene. The D. setosum clade then split 3‐5 million years ago into two clades, one found around the Arabian Peninsula and the other in the Indo‐West Pacific. On the lineage leading to the other species of Diadema, the deepest branch is composed of D. palmeri, apparently separated when the climate of New Zealand became colder and other tropical echinoids at these islands went extinct. The next lineage to separate is composed of a currently unrecognized species of Diadema that is found at Japan and the Marshall Islands. Diadema mexicanum in the eastern Pacific separated next, whereas D. paucispinum, D. savignyi, and D. antillarum from the western and central Atlantic, and (as a separate clade) D. antillarum from the eastern Atlantic form a shallow polytomy. Apparently, Indo‐Pacific populations of Diadema maintained genetic contact with Atlantic ones around the southern tip of Africa for some time after the Isthmus of Panama was complete. Diadema paucispinum contains two lineages: D. paucispinum sensu stricto is not limited to Hawaii as previously thought, but extends to Easter Island, Pitcairn, and Okinawa; A second mitochondrial clade of D. paucispinum extends from East Africa and Arabia to the Philippines and New Guinea. A more recent separation between West Indian Ocean and West Pacific populations was detected in D. setosum. Presumably, these genetic discontinuities are the result of water flow restrictions in the straits between northern Australia and Southeast Asia during Pleistocene episodes of low sea level. Diadema savignyi is characterized by high rates of gene flow from Kiribati in the central Pacific all the way to the East African Coast. In the Atlantic, there is a biogeographic barrier between the Caribbean and Brazil, possibly caused by fresh water outflow from the Amazon and the Orinoco Rivers. Diadema antillarum populations of the central Atlantic islands of Ascension and St. Helena are genetically isolated and phylogenetically derived from Brazil. Except for its genetic separation by the mid‐Atlantic barrier, Diadema seems to have maintained connections through potential barriers to dispersal (including the Isthmus of Panama) more recently than did Eucidaris or Echinometra, two other genera of sea urchins in which phylogeography has been studied. Nevertheless, the mtDNA phylogeography of Diadema includes all stages expected from models of allopatric differentiation. There are anciently separated clades that now overlap in their geographic distribution, clades isolated in the periphery of the genus range that have remained in the periphery, clades that may have been isolated in the periphery but have since spread towards the center, closely related clades on either side of an existing barrier, and closely related monophyletic entities on either side of an historical barrier that have crossed the former barrier line, but have not attained genetic equilibrium. Except for D. paucispinum and D. savignyi, in which known hybridization may have lodged mtDNA from one species into the genome of the other, closely related clades are always allopatric, and only distantly related ones overlap geographically. Thus, the phylogenetic history and distribution of extant species of Diadema is by and large consistent with allopatric speciation.     

Coexistence in a sea urchin guild and its implications to coral reef diversity and degradation

Summary Coexistence between the coral reef inhabiting sea urchins Echinometra mathaei, Diadema savignyi and D. setosum was studied by comparing differences in body morphology, distribution, diet, susceptibility to predators, intra- and interspecific competition and settlement. The three species share similar diets and broad within-habitat distributions but differ in their microspatial preferences. E. mathaei is the smallest species, has the highest settlement rates and lives territorially within small burrows or crevices. D. savignyi is intermediate in size and lives frequently in intermediate size crevices or occassionally in social groups. D. setosum is the largest species and occassionally lives in large crevices or more frequently in social groups. Both Diadema have similarily low settlement rates. Competition experiments showed that E. mathaei was consistently the top competitor for crevice space. Diadema species shared larger crevices but competition occured within smaller crevices and was frequently won by the largest individual, regardless of species. D. savignyi may be the top competitor for crevice space between the Diadema species due to a reduced spine length/test size ratio which gives it a larger test for the same crevice size requirement. Predation rates were high for E. mathaei and low for both Diadema species. Coexistence is mediated by predation on the competitive-dominant while predation coupled with different body morphologies and behavior allows spatial resource partitioning of the reef's variable topography. Consequently, the three variables of predation, topographic complexity and differing body shapes create the observed species diversity. A reduction in predators due to stochastic fluctuations or from fishing pressure can lead to E. mathaei population increases and competitive exclusion of Diadema.     

Functional morphology of the pedicellariae of the asteroid Marthasterias glacialis (Echinodermata)

Summary Marthasterias glacialis bears two kinds of pedicellariae. The straight pedicellariae are single and occur everywhere on the asteroid body surface except in the ambulacral groove. The crossed pedicellariae are clustered on mobile structures (the rosettes) build around marginal and abactinal spines.Basically, each pedicellaria has a head and a stalk. A skeleton occurs only in the pedicellarial head. It consists of two valves and a basal piece. Muscular bundles are anchored on these skeletal ossicles. The straight pedicellariae have two pairs of adductor muscles (the inner and the outer adductors) and one pair of abductor muscles, these latter being weakly developed. Longitudinal muscle fibers occur all along the stalk of straight pedicellariae. The crossed pedicellariae have two pairs of adductor muscles (the distal and the proximal adductors) and two pairs of abductor muscles (the distal and the proximal abductors). The proximal adductors of crossed pedicellariae are homologous to the stalk muscles of straight pedicellariae.The pedicellariae are able to react to direct and indirect tactile stimuli. There is a great deal of individual variation among pedicellarial responses. Moreover, the reactions occur at random and lack coordination. The seemingly aberrant behavior of the pedicellariae is interpreted as a preventive activity that protects the asteroid body surface against unwanted materials and organisms.     

Effects of a nonnative, invasive lovegrass on Agave palmeri distribution, abundance, and insect pollinator communities

Nonnative Lehmann lovegrass (Eragrostis lehmanniana) has invaded large areas of the Southwestern United States, and its impact on native plants is not fully understood. Palmer’s agave (Agave palmeri), an important resource for many pollinators, is a key native plant potentially threatened by E. lehmanniana. Understanding potential impacts of E. lehmanniana on A. palmeri is critical for anticipating the future of the desert community where they coexist and for addressing management concerns about associated threatened and endangered species. Our study provides strong indications that E. lehmanniana negatively impacts A. palmeri in several ways. Areas of high E. lehmanniana abundance were associated with significantly lower densities and greater relative frequencies of small A. palmeri, suggesting that E. lehmanniana may exclude A. palmeri. There were no significant differences in species richness, abundance, or community composition when comparing flower associates associated with A. palmeri in areas of high and low E. lehmanniana abundance. However, we did find significantly lower connectedness within the pollination network associated with A. palmeri in areas with high E. lehmanniana abundance. Although E. lehmanniana forms thick stands that would presumably increase fire frequency, there was no significant association between E. lehmanniana and fire frequency. Interestingly, medium to high densities of A. palmeri were associated with areas of greater fire frequency. The complex ramifications of E. lehmanniana invasion for the long-lived A. palmeri and interlinked desert community warrant continued study, as these species are likely to continue to be found in close association due to their similar soil preferences.     

The evolution of pedicellariae in echinoids: an arms race against pests and parasites

Coppard, S.E., Kroh, A. and Smith, A.B. (2010). The evolution of pedicellariae in echinoids: an arms race against pests and parasites. —Acta Zoologica (Stockholm) 00 :1–24. Sea urchins (Echinoidea) have evolved a diverse array of jawed appendages termed pedicellariae to deter pests and predators. Pedicellarial structure and function are reviewed and their distribution mapped in 75 extant genera. Using a phylogeny of echinoids at family level constructed from 353 skeletal characters scored across 162 extant and fossil taxa, the evolution of pedicellarial form and function is reconstructed. For much of the Palaeozoic echinoids possessed a very restricted pedicellarial armament. By the early Mesozoic a diverse array of pedicellarial types had become established, implying that the threat from predators and pests markedly increased at this time. Since the Triassic, echinoids have continued to improve their defensive capability by evolving more effective venom delivery in globiferous pedicellariae, developing spatulate‐tips and curved blades for a more efficient grab in tridentate pedicellariae, and stouter, more robust valves with a stronger bite in ophicephalous pedicellariae to disable and remove ectoparasites. However, pedicellarial types are shown to be particularly prone to subsequent secondary loss, especially among infaunal echinoids, and thus have higher homoplasy levels than other phylogenetically useful skeletal structures.     

An integrative approach for species delimitation in the spider genus Grammostola (Theraphosidae,Mygalomorphae)

The mygalomorph genus Grammostola (family Theraphosidae) is endemic to South America. The species Grammostola anthracina is one of the largest spiders in Uruguay and reputed to be the longest lived tarantula in the world. This nominal species has two distinct colour morphs comprising black and reddish‐brown forms with controversial taxonomic status. Here, we present a phylogenetic study based on molecular characters (cytochrome c oxidase subunit I) of haplotypes of G. anthracina and closely related species. Our analysis together with new morphological data and biogeographical information indicates that the two morphs of G. anthracina constitute different species that are not sister to each other. Consequently, a new species, Grammostola quirogai is described, diagnosed and illustrated to encompass the black morph. Phylogenetic relationships and new taxonomic characters for Grammostola species included in this study are discussed.     

On the Fourth Diadema Species (Diadema-sp) from Japan

Four long-spined sea urchin species in the genus Diadema are known to occur around the Japanese Archipelago. Three species (D. savignyi, D. setosum, and D. paucispinum) are widely distributed in the Indo-Pacific Ocean. The fourth species was detected by DNA analysis among samples originally collected as D. savignyi or D. setosum in Japan and the Marshall Islands and tentatively designated as Diadema -sp, remaining an undescribed species. We analyzed nucleotide sequences of the cytochrome oxidase I (COI) gene in the “D. savignyi-like” samples, and found all 17 individuals collected in the mainland of Japan (Sagami Bay and Kyushu) to be Diadema-sp, but all nine in the Ryukyu Archipelago (Okinawa and Ishigaki Islands) to be D. savignyi, with large nucleotide sequence difference between them (11.0%±1.7 SE). Diadema-sp and D. savignyi shared Y-shaped blue lines of iridophores along the interambulacrals, but individuals of Diadema-sp typically exhibited a conspicuous white streak at the fork of the Y-shaped blue iridophore lines, while this feature was absent in D. savignyi. Also, the central axis of the Y-shaped blue lines of iridophores was approximately twice as long as the V-component in D. savignyi whereas it was of similar length in Diadema-sp. Two parallel lines were observed to constitute the central axis of the Y-shaped blue lines in both species, but these were considerably narrower in Diadema-sp. Despite marked morphological and genetic differences, it appears that Diadema-sp has been mis-identified as D. savignyi for more than half a century.     

Sexual dimorphism and food habits of the clingfish,Diademichthys lineatus,and its dependence on host sea urchin

Synopsis Stomach contents of the clingfish Diademichthys lineatus, 10–56 mm standard length, revealed changes in food habits with growth and sexual differences. Soon after settlement, D. lineatus obtained food from their host sea urchin (genus Diadema) and other associated symbionts. They became less dependent on the host with growth. The juveniles ate pedicellariae and sphaeridia of the host and commensal copepods, whereas the adult fish ate burrowing bivalves in corals as well as tube feet of their host and eggs of a commensal shrimp. The young fish were transitional in their food habits. The change in food habits of the fish coincided with behavioral changes; i.e. enlargement of home ranges and less dependency on the host. The adult females, having a longer snout, ate shrimp eggs and bivalves more frequently than the adult males, which ate tube feet of the host more often than the females did. Sexual difference in food habits was apparent after the sex of the fish became identifiable by comparing snout shapes. The polygynous mating system of this species suggests that conspicuous sexual dimorphism might have developed under sexual selection. However, niche partitioning of food is also likely to be related to this sexual dimorphism.     

Dimorphisms and fluctuating asymmetry in the forceps of male earwigs

Male dimorphisms are particularly conspicuous examples of the alternative reproductive strategies employed within some species. Such dimorphisms are thought to exist as genetic polymorphisms under ESS conditions, or to be conditional strategies where exogenous conditions determine the adult body plan. Fluctuating asymmetry (FA) is currently considered to be a fitness correlate of significant use in interpreting the functional significance of secondary sexual characteristics. In particular, negative slopes of FA on trait size are thought to arise in traits whose expression is dependent on condition. We measured forceps lengths and asymmetries in 2 island populations of the European earwig Forficula auricularia and Museum specimens of 5 other earwig (Dermaptera, Forficulidae) species from different genera, that appeared to be dimorphic. In a detailed study of Forficula auricularia we found a significant fit to a statistical model for the identification of dimorphisms and, for all species examined, morphs differed in the slope and/or elevation of the allometric relationship between body size and forcep length. Possible determinates of male dimorphisms are suggested from the data. Contrary to expectation, FA was not found to be greater in the minor morphs. Negative relationships between FA and forceps length were absent in both morphs of species examined from museum collections. Of the two island populations of Forficula auricularia, the smaller and more isolated population had higher FA and a negative relationship between FA and forceps length in the major morph. We discuss these patterns in the light of recent theories of FA and honest signalling.     

RELATIONSHIP BETWEEN ISOZYMIC AND MORPHOLOGIC VARIATION IN THE DIPLOIDS CHENOPODIUM FREMONTII,C. NEOMEXICANUM,C. PALMERI,AND C. WATSONII

Levels of variation revealed by starch gel electrophoresis and morphologic character analysis were compared within and among four diploid species of Chenopodium: C. fremontii (sect. Chenopodium subsect. Leiosperma), C. neomexicanum, C. palmeri, and C. watsonii (sect. Chenopodium subsect. Cellulata). The data sets exhibited little concordance. Numerical analysis of morphologic characters demonstrated that C. neomexicanum and C. palmeri are relatively distinct, whereas allozyme analysis indicated a high degree of genetic similarity among populations of the two species. I concluded that C. palmeri should be reduced to a variety of C. neomexicanum. Chenopodium watsonii expressed the highest degree of allozyme heterozygosity and polymorphism and may represent the oldest extant member of subsection Cellulata. Although C. fremontii is classified in another subsection, it exhibited higher genetic identity values with C. neomexicanum and C. palmeri than did C. watsonii. In addition, morphologic and allozymic similarity between C. fremontii and C. neomexicanum suggests the need for further examination and possibly realignment of subsections Cellulata and Leiosperma.     

Morphology of the nervous and muscular systems in the heads of pedicellariae from the sea urchin Echinus esculentus L

Light and electron microscopy reveal that simple receptor cells in the jaw epithelium of sea urchin pedicellariae are connected by nerve tracts to the neuropile that coordinates jaw movements. The muscles responsible for jaw opening and closure and for flexion of the stem are all innervated in this neuropile. At least two types of vesicles occur at the simple synapses between neurone profiles and at neuromuscular junctions. The muscles include both striated and smooth fibres; however, their distribution varies according to pedicellaria type, and an unexpected arrangement exists in trifoliate pedicellariae.     

Five Simuliidae (Diptera) from West Cameroon

The male, female, pupa and larva of a new phoretic species of Simulium are described. A new species-group is established for this species and S.diceros, and its systematic position is discussed. Two other new species of Simulium are described from pupal material, and new forms of S.palmeri and S.berghei are recorded.     

Phenotypic plasticity in the Antarctic fish <Emphasis Type="Italic">Trematomus newnesi</Emphasis> (Nototheniidae) from the South Shetland Islands

The presence of the two morphs, “typical” and “large mouth”, in the Antarctic fish species Trematomus newnesi (Perciformes, Notothenioidei) was recorded for the first time in nearshore waters of the South Shetland Islands (Potter Cove) and western Antarctic Peninsula (Petermann Island). The two morphs were distinguishable in specimens of 60–241 mm total length (TL); about 30% of the specimens constituted intermediate forms. In addition to the previously known characters separating the morphs, we found that the “relative size of the eye” can also be used to identify smaller and larger fish of the typical morph. The ecological significance of the two morphs remains unclear. Ratios of diagnostic characters for identification of the species at two size ranges (60–131 and 132–241 mm TL) are provided.