Size and shape in the phylum Phoronida

The lophophorate phylum Phoronida consists of about 13 species, which differ in body length and width, number of longitudinal muscles, lophophore geometry and number of lophophore tentacles. In absolute terms large species have a larger body width, more tentacles, more longitudinal muscles and greater coiling of the lophophore than small species. However, size and shape analyses suggest that with increasing size: (I) the body surface area to volume ratio increases because body length increases faster than body width; (2) the relative number longitudinal muscles decreases, and (3) the relative feeding surface area of the lophophore decreases because tentacle diameter is constant while tentacle number increases at the same rate as body length and tentacle length increases more slowly than tentacle number. Coiling and spiraling of the lophophore in large species may be an attempt to compensate for this last relationship. We suggest that the habits, mode of growth and feeding mechanism of phoronids constrain size-related changes in shape.     

Architecture and function of the lophophore in the problematic brachiopod Heliomedusa orienta (Early Cambrian, South China)

The detailed structure of the lophophore is a key diagnostic character in the definition of higher brachiopod taxa. The problematic Heliomedusa orienta Sun and Hou, from the Lower Cambrian Chengjiang Lagerstätte of Yunnan, southwestern China, has a well-preserved lophophore, which is unlike that of any known extant or extinct brachiopods. Based on a comparative study of lophophore disposition in H. orienta and the extant discinid Pelagodiscus atlanticus, the in- and excurrent pattern and shell orientation of H. orienta are described and discussed. Reconstructions of lophophore shape and function are based on numerous specimens and comparison with P. atlanticus. The lophophore is composed of a pair of lophophoral arms that freely arch posteriorly rather than coiling anteriorly as commonly seen in fossil and recent lingulids. The lophophore is attached to the dorsal lobe of the mantle; it has neither calcareous nor chitinous supporting structures, and is disposed symmetrically on either side of the valve midline. The mouth can be inferred to be located at the base of the two brachial tubes, slightly posterior to the anterodorsal projection of the body wall. The lophophoral arms bear laterofrontal tentacles with a double row of cilia along their lateral edge, as in extant lingulid brachiopods. The main brachial axes are also ciliated, which presumably facilitated transport of mucous-bound nutrient particles to the mouth. The unique organization of the lophophore in Heliomedusa is not like any known fossil and living brachiopods. This clearly demonstrates that H. orienta is not a member of any crown group. It is here considered as a member of the brachiopod stem group, which challenges recent interpretations of a close discinid affinity.     

Modern Data on the Innervation of the Lophophore in Lingula anatina (Brachiopoda) Support the Monophyly of the Lophophorates

Evolutionary relationships among members of the Lophophorata remain unclear. Traditionally, the Lophophorata included three phyla: Brachiopoda, Bryozoa or Ectoprocta, and Phoronida. All species in these phyla have a lophophore, which is regarded as a homologous structure of the lophophorates. Because the organization of the nervous system has been traditionally used to establish relationships among groups of animals, information on the organization of the nervous system in the lophophore of phoronids, brachiopods, and bryozoans may help clarify relationships among the lophophorates. In the current study, the innervation of the lophophore of the inarticulate brachiopod Lingula anatina is investigated by modern methods. The lophophore of L. anatina contains three brachial nerves: the main, accessory, and lower brachial nerves. The main brachial nerve is located at the base of the dorsal side of the brachial fold and gives rise to the cross neurite bundles, which pass through the connective tissue and connect the main and accessory brachial nerves. Nerves emanating from the accessory brachial nerve account for most of the tentacle innervation and comprise the frontal, latero-frontal, and latero-abfrontal neurite bundles. The lower brachial nerve gives rise to the abfrontal neurite bundles of the outer tentacles. Comparative analysis revealed the presence of many similar features in the organization of the lophophore nervous system in phoronids, brachiopods, and bryozoans. The main brachial nerve of L. anatina is similar to the dorsal ganglion of phoronids and the cerebral ganglion of bryozoans. The accessory brachial nerve of L. anatina is similar to the minor nerve ring of phoronids and the circumoral nerve ring of bryozoans. All lophophorates have intertentacular neurite bundles, which innervate adjacent tentacles. The presence of similar nerve elements in the lophophore of phoronids, brachiopods, and bryozoans supports the homology of the lophophore and the monophyly of the lophophorates.     

Functional disposition of the lophophore in living Brachiopoda

Emig, C. C. 1992 07 15: Functional disposition of the lophophore in living Brachiopoda.
The shape and disposition of adult brachiopod lophophores relate to in- and excurrent apertures. to the internal water irrigation system, to shell orientation at substratum and to near-bottom currents. The arrangement of the mantle canals and gonads of different lophophores are induccd by water circulation. The trocholophe (2% of living species) is considered as a plesiomorphic character which represents the basic plan of the lophophore, shared by all Lophophorata. Three different types of schizolophe (10%) are represented in terebratuloids, thecidioids and discinids. The spirolophe (19%), characteristic of rhynchonellides and most inarticulate brachiopods, except the schizolophe Pelagodiscus , has evolved divergently into specific arrangements of the mantle canals and gonads. The zygo-plectolophe (67%) is characteristic of most Terebratulida. The ptycholophe (2%) probably evolved independently in Megathlris and the thecidioids. The mesolophe, known in the fossil chonetdceans, is considered to be a primitive zygo-plectolophe. The median brachiopod sulcus increases the efficiency of the excurrent system and is considered as an evolved character but a homoplasy within the brachiopods. The characteristics of Recent lophophore types have to be taken into account when reconstructing the lophophore in fossil forms. Brachiopoda, Lophophorata, lophophore, water system, orientation, evolution .     

Survival, development and life tables of two congeneric ladybirds in aphidophagous guilds

Two congeneric aphidophagous ladybirds, Coccinella septempunctata and Coccinella transversalis, were reared on three aphid species, Lipaphis erysimi, Myzus persicae and Aphis nerii, to estimate the effect of prey quality and intra- and interspecific interactions on their survival and development of life stages. Mortality of first instar ladybirds of both species was highest feeding on A. nerii. Preimaginal mortafity was lowest when feeding on L. erysimi （C. septempunctata, 1.6% and C. transversalis, 3.2%）, and highest when feeding on A. nerii （ C. septempunctata, 6.2% and C. transversalis, 8.2%）. Comparatively higher weight and larger size of C. septempunctata along with the lower levels of mortality recorded suggested that it is more likely to have acted as an intraguild predator than C. transversalis. High recorded mortality of C. transversalis is attributed to probable intraguild predation on account of its smaller size. The major sources of mortality were probably cannibalism, intraguild predation and other unknown factors. Lower prey quality increased the incidence of cannibalism and intraguild predation, especially in C. transversalis. The investigation suggests an intrinsic competitive advantage for C. septempunctata over C. transversalis in guilds of three aphid species.     

Ultrastructure and immunocytochemistry of the nervous system of the larvae ofLingula anatina andGlottidia sp. (Brachiopoda)

Summary Planktotrophic brachiopod larvae ofGlottidia sp. have been investigated for the occurrence of glyoxylic acid induced fluorescence in catecholamines (CA), and serotonin-like (5-HT) and neuropeptide FMRFamidelike (FMRFamide) immunoreactivity (ir). The location of CA, 5-HT-ir and FMRFamide-ir cells and processes were compared with the location of neurons and nerve processes found by transmission electron microscopy. The apical ganglion contains 5-HT-ir and FMRFamideir cells and processes and CA processes. From the dorsal part of the apical ganglion extend dorsal 5-HT-ir and FMRFamide-ir processes; from the nine pairs of tentacles stage (9. pt) they project to the ventral ganglion. These dorsal lophophore processes follow themusculus lophophoralis and them. brachialis. The 5-HT-ir and some of the FMRFamide-ir processes project along the muscles to the tentacles. From the ventral part of the apical ganglion extend CA, 5-HT-ir and FMRFamide-ir processes which follow the ciliary band of the lophophore and project to the tentacles. An intense band of CA processes was also observed in the lophophore, but the dorsal/ventral location could not be ascertained. The ventral ganglion contains 5-HT-ir and FMRFamide-ir cells which project either caudally on the metasome or rostrally as part of the dorsal lophophore processes. The neuropil of the ventral ganglion contains CA, 5-HT-ir and FMRFamide-ir processes. The nervous system of the planktotrophic brachiopod larvae seems to consist of a ventral lophophore system innervating the ciliary bands and a dorsal lophophore system including the ventral ganglion innervating the body musculature. The latter system develops later in ontogeny and is regarded as a specialization due to the presence of shells and associated musculature. The former system is regarded as homologous with the nervous system of actinotroch larvae (Phoronida) and planktotrophic larvae of the echinoderms.     

Rejection Mechanism of Plectolophous Lophophore of Brachiopod <Emphasis Type="Italic">Coptothyris grayi</Emphasis> (Terebratulida,Rhynchonelliformea)

Brachiopoda is a relict group of invertebrate filter feeders that used a tentacle organ, lophophore, for capturing food particles from the water column. Brachiopod extinction apparently occurred due to low productivity of their filtering organ in comparison with more advanced filter-feeders. Investigation of the filtering mechanism of modern brachiopods is essential to understanding their evolutionary fate. This study is devoted to the rejection mechanism of large waste particles from the plectolophous lophophore of brachiopod Coptothyris grayi. The waste particles gather inside of the lophophore on the outer side of the brachial fold. The particles form rows along frontal grooves of outer tentacles and are carried successively to the tentacle tips and move along them, slimed by mucus. One portion of the particles comes off the lophophore and falls down the mantle, while another part is carried to the abfrontal surface of the tentacles. Due to repeated reversals of abfrontal cilia, the particles wavily move along the abfrontal surface of tentacles. Such movement contributes to the secretion of mucus and the formation of particle clots. The clots come off the lophophore and fall down the mantle. The particles are transported along the mantle by cilia to the anterior part of the mantle margin. Here the ciliary reversals that facilitate secretion of mucus and formation of pseudofeces also take place. The latter takes away from the mantle cavity. Thus, only outer tentacles participate in the rejection of large waste particles from the lophophore. Ciliary reversals of the abfrontal surface of tentacles and the mantle are discovered in brachiopods for the first time. This facilitates the additional secretion of mucus and formation of pseudofeces, easing their exit from the mantle cavity. The results contribute to the knowledge of lophophore function and evolution of tentacle organs in Bilateria.     

On the organization of the lophophore in phoronids (Lophophorata: Phoronida)

The organization of the lophophore is the main feature used for the identification of phoronid species. The structure of the lophophore and tentacles in seven phoronid species (Phoronis ovalis, P. ijimai, P. hippocrepia, P. svetlanae, P. australis, Phoronopsis harmeri, and Ph. malakhovi) collected in different areas of the World Ocean was studied. Two new patterns of the phoronid lophophore structure were found: “transition to horseshoe-shaped” (as in P. ovalis from Aniva Bay and in P. ijimai from the coast of Iturup Island, Sea of Okhotsk) and “transition to spiral” (in burrowing specimens P. hippocrepia from Aniva Bay, P. svetlanae and Ph. harmeri from Vostok Bay, Sea of Japan). For the first time it was shown that phoronid species with different types of the lophophore structure possess different kinds of tentacles. Thus, five types of phoronid tentacles were identified that vary in the shape of their cross section: rounded, oval, ellipsoid, rectangular, and skittle-shaped. A correlation was found between lophophore organization and the type of tentacles in phoronids. A table of the correlation between body size, lophophore organization, tentacle structure, and mode of life in different phoronid species is proposed.     

Das filter-feeding-System bei Spiriferida

SEM studies of the brachidial microstructure of brachiopods of the Spirifer group, Atrypa group, and A thyris suggest that the lophophore of each group was fixed to the basal side of the lamella. On this basis, there are four possible constructions of the spiriferoid lophophore, but the most probable one is a doubled lophophore acting as a filter tube which draws water into the mantle cavity laterally and ejects it through a median exhalant aperture.
Rasterelektronenmikroskopische Untersuchungen der Brachidien von Spiriferen, Atrypen und Athyris erlauben die Schlußfolgerung, daß bei allen drei Brachiopodengruppen der Lophophor an der proximalen bzw. basalen Seite der kalkigen Spirallamelle gehangen hat. Von den dann möglichen vier Modellen des Lophophorensystems erweist sich das eines Filterdoppelkorbs am wahrscheinlichsten, der das Frischwasser seitlich in das Gehäuse einströmen und in der Mitte ausströmen ließ.     

Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study

Riisgård, H.U., Okamura, B. and Funch, P. 2009. Particle capture in ciliary filter‐feeding gymnolaemate and phylactolaemate bryozoans – a comparative study. —Acta Zoologica (Stockholm) 91 : 416–425. We studied particle capture using video‐microscopy in two gymnolaemates, the marine cheilostome Electra pilosa and the freshwater ctenostome Paludicella articulata, and three phylactolaemates, Fredericella sultana with a circular funnel‐shaped lophophore, and Cristatella mucedo and Lophophus crystallinus, both with a horseshoe‐shaped lophophore. The video‐microscope observations along with studies of lophophore morphology and ultrastructure indicated that phylactolaemate and gymnolaemate bryozoans with a diversity of lophophore shapes rely on the same basic structures and mechanisms for particle capture. Our study also demonstrates that essential features of the particle capture process resemble one another in bryozoans, brachiopods and phoronids.     

The mesolophe, a new lophophore type for chonetacean brachiopods

Rachebaeuf, P. R. & Copper, P. 1990 10 15: The mesolophe, a new lophophore type for chonetacean brachiopods. Lethaia , Val. 23, pp. 341–346. Oslo. ISSN 0024–1164.
Following a summary of previous lophophore reconstructions for the chonetaceans, we describe an unusual pyritized structure within the calcite infill of an exceptionally preserved shell of Archeochonetes primigenius (Twenhofel) from the Late Ordovician (Ashgill) of Anticosti Island, Quebec, Canada. The brachial valve interior of most Lower Devonian to Permian chonetaceans shows the development of three depressed deepened areas (gutters) in the valve floor. The disposition of these gutters coincides remarkably with the shape of the pyritized structure, which we postulate as a new type of lophophore, the mesolophe. ▭ Brachiopoda, Chonetacea, functional morphology, lophophore .     

Body cavities in bryozoans: Functional and phylogenetic implications

Based on morphological evidence, Bryozoa together with Phoronida and Brachiopoda are traditionally combined in the group Lophophorata, although this view has been recently challenged by molecular studies. The core of the concept lies in the presence of the lophophore as well as the nature and arrangement of the body cavities. Bryozoa are the least known in this respect. Here, we focused on the fine structure of the body cavity in 12 bryozoan species: 6 gymnolaemates, 3 stenolaemates and 3 phylactolaemates. In gymnolaemates, the complete epithelial lining of the body cavity is restricted to the lophophore, gut walls, and tentacle sheath. By contrast, the cystid walls are composed only of the ectocyst-producing epidermis without a coelothelium, or an underlying extracellular matrix; only the storage cells and cells of the funicular system contact the epidermis. The nature of the main body cavity in gymnolaemates is unique and may be considered as a secondarily modified coelom. In cyclostomes, both the lophophoral and endosaccal cavities are completely lined with coelothelium, while the exosaccal cavity only has the epidermis along the cystid wall. In gymnolaemates, the lophophore and trunk cavities are divided by an incomplete septum and communicate through two pores. In cyclostomes, the septum has a similar location, but no openings. In Phylactolaemata, the body cavity is undivided: the lophophore and trunk coeloms merge at the bases of the lophophore arms, the epistome cavity joins the trunk, and the forked canal opens into the arm coelom. The coelomic lining of the body is complete except for the epistome, lophophoral arms, and the basal portions of the tentacles, where the cells do not interlock perfectly (this design probably facilitates the ammonia excretion). The observed partitioning of the body cavity in bryozoans differs from that in phoronids and brachiopods, and contradicts the Lophophorata concept.     

Passive feeding in spiriferide brachiopods: an experimental approach using models of Devonian Paraspirifer and Cyrtospirifer

Shiino, Y. 2009: Passive feeding in spiriferide brachiopods: an experimental approach using models of Devonian Paraspirifer and Cyrtospirifer. Lethaia, Vol. 43, pp. 223–231. Passive feeding flows of two Devonian spiriferide brachiopods, Paraspirifer bownockeri and Cyrtospirifer sp., were examined experimentally using transparent hollow models and a flowing water tank. The models were constructed with polycarbonate plates using a vacuum heat press method. Another model of Paraspirifer with a spiral brachidium was constructed for the visualization of passive flow. The results of the ventral and dorsal directions of the hollow models suggest that outflow was generated through lateral gapes in the shell, while intake may have occurred through a sulcus gape. The passive internal flow inside the models invariably exhibited gyrating behaviour, and the axis of the rotation was similar to that of the spiral lophophore in spiriferides. Comparing the results using the hollow and backbone models indicates that the internal structures, which consisted mostly of the spiral brachidium, aided in adjusting the gyrating flows around the brachidium, even when the lateral region of the shell faced upstream. Extant terebratulides are known to generate gyrating flows around the median coils of the major feeding area, and this model best fits the relationship between the passive internal flows and the form of the lophophore. As a consequence, the shell forms of spiriferides could generate passive feeding flows from the gyrating movement surrounding the lophophore that are effective for feeding by the spiral lophophore. □Brachiopoda, flume experiment, functional morphology, impingement feeder, physiology, suspension feeder.     

A reconstruction of the lophophore of Devonian rhynchonellids (Brachiopoda) by using X-Ray micro-CT

The X-ray microtomographic study has revealed contrast inclusions (possibly iron compounds) in several shells of Devonian (Emsian–Famennian) rhynchonellids (Brachiopoda) from Transcaucasia. Judging by the location of inclusions, they may correspond to soft tissues of lophophores. The spire-shaped inclusion in the shell of the holotype of the Late Devonian Sharovaella mirabilis Pakhnevich, 2012 has typical features of spirolophe and is interpreted as a part of one of lophophore spires. This find suggests that in the Late Devonian the spirolophe already existed in rhynchonellids. The dorsoventrally directed spirolophous lophophore is an ancient conservative feature of the order Rhynchonellida.     

Yellow Traps Can Be Used to Monitor Populations of Coccinella transversalis (F.) and Adalia bipunctata (L.) (Coleoptera: Coccinellidae) in Cotton Crops

During 1992–4, square (30 times 30 cm) sticky traps of various colours were used on a commercial cotton farm to trap adults of Coccinella transversalis and Adalia bipunctata which are both major predators of Helicoverpa spp. Both insects were attracted most to yellow traps which also reflected the most visible light between 500 nmand 600 nm(where green foliage reflects most light). When yellow was diluted with white to produce yellow-white hues, the light reflected between 500 nm and 600 nm was reduced and the numbers of C. transversalis and A. bipunctata adults caught on these traps were also significantly reduced. This suggests that C. transversalis and A. bipunctata adults can discriminate foliage-like hues (500–580 nm) from non-foliage-like hues (<500 nm and >580 nm) and are attracted to colours that suggest the foliage of host plants that may harbour their prey. Yellow sticky traps placed 25–50 cm above ground caught significantly more C. transversalis and A. bipunctata adults than those placed at 75–150 cm and are the most appropriate traps to monitor populations of C. transversalis and A. bipunctata adults in cotton farms.     

The periesophageal celom of the articulate brachiopod Hemithyris psittacea (Rhynchonelliformea,Brachiopoda)

The celomic system of the articulate brachiopod Hemithyris psittacea is composed of the perivisceral cavity, the canal system of the lophophore, and the periesophageal celom. We study the microscopic anatomy and ultrastructure of the periesophageal celom using scanning and transmission electron microscopy. The periesophageal celom surrounds the esophagus, is isolated from the perivisceral cavity, and is divided by septa. The lining of the periesophageal celom includes two types of cells, epithelial cells and myoepithelial cells, both are monociliary. Some epithelial cells have long processes extending along the basal lamina, suggesting that these cells might function as podocytes. The myoepithelial cells have basal myofilaments and may be overlapped by the apical processes of the adjacent epithelial cells. The periesophageal celom forms protrusions that penetrate the extracellular matrix (ECM) of the body wall above the mouth and the ECM that surrounds the esophagus. The canals of the esophageal ECM form a complicated system. The celomic lining of the external circumferential canals consists of the epithelial cells and the podocyte‐like cells. The deepest canals lack a lumen; they are filled with the muscle cells surrounded by basal lamina. These branched canals might perform dual functions. First, they increase the surface area and might therefore facilitate ultrafiltration through the podocyte‐like cells. Second, the deepest canals form the thickened muscle wall of the esophagus and could be necessary for antiperistalsis of the gut. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Aspects of the Functional Morphology of the Lophophore in Articulate Brachiopods

Histologic studies made on three species of articulate brachiopodsprovide information on the structural integrity, epithelialspecialization and other cellular aspects of the lophophore.Similarities in staining properties of granular inclusions foundin the matrix with those found in epithelial modifications ofthe brachial trough and the bases of the filaments suggest movementbetween cells within the matrix and the epithelium. The natureof the granular inclusions raises the possibility of intracellulardigestion along the lophophore of these brachiopods.     

A NEW EARLY CARBONIFEROUS MICRO-PRODUCTID BRACHIOPOD FROM SOUTH CHINA

Abstract:  We describe Muhuarina haeretica gen. et sp. nov. from a recently discovered silicified brachiopod fauna from the Lower Carboniferous of South China. This new microproductid appears to be the oldest representative of the highly differentiated, mainly Permian, aulostegoid family Cooperinidae. The internal structure of the dorsal valve of M. haeretica is dominated by strongly developed, pronounced subperipheral ridges, which suggest support for a simple schizolophe type of lophophore. Brachial ridges, as in other productides, most probably represent an attachment area of the lophophore to the dorsal epithelium. An attempt to homologize subperipheral and brachial ridges among Cooperinidae is presented.     

Prefabricated thin flap using the transversalis fascia as a carrier.

To harvest a thin flap from the groin and hypogastric area, the authors developed a new prefabricated flap using the transversalis fascia as a carrier. The transversalis fascia is a very thin and abundantly vascularized tissue nourished by the deep inferior epigastric vessels. Flap prefabrication was performed by inserting the transversalis fascia between the thinly undermined skin flap and the tissue expander placed beneath the skin flap, followed by a pretransfer delay procedure around the flap. After a 3-week interval, the flap was transplanted with no complications, such as congestion and thrombus of anastomosis. By using this technique, it was possible to elevate an equally thin flap from the groin and hypogastric area while avoiding morbidity of the donor site.     

Body volumes and internal space constraints in articulate brachiopods

Brachiopods were once dominant in all the oceans of the world. but their distributions are non more restricted. There are few species which are found in shallow warm habitats and these are predominantly small. They have exceptionally low metabolic rates and exhibit low energy lifestyles. The majority of living articulate brachiopods are punctate (possessing mantle extensions. or caeca. which traverse the shell). Evidence produced hei-e suggests that the evolution of these phenomena may have been strongly affected by architectural constraints placed on articulate brachiopods by the use of the lophophore for feeding and respiration. They are essentially space limited because of the large volume needed for this organ. In some punctate brachiopods over 75% of their total body volume may be occupied by the lophophore and mantle cavity. This figure is only 60% in an impunctate (no caeca) species and may be only 20% in bivalve molluses. The implications are that caeca evolved to reduce pressure on space requirements, that maximum sizes may be set by the scaling patterns of space allocation and metabolic efficiency is a consequence of space constraints. Current distribution patterns may be strongly affected by the low metabolism and low energy lifestyles. The relative success of small brachiopods in warm shallow seas may have been facilitated by the scaling patterns of space allocations which show small specimens to have similar mantle cavity volumes to bivalve molluscs.