M - Paleopolis

Zoologica Scripta. Vol. 4, 37-40, 1975

Terminology in Phoronida Christian C. Emig Station Marine d'Endoume, Marseille, France Receiced 1974-12-/6

Abstract Emig, C. C. (Station Marine d'Endoumc, F· 13007 Marseille. France.) Termina/ogy in Phoronida. Zoo!. Ser. 4 (1): 37--40. 1975.-A terminalogy (in three languages) of the main descriptive terms of the anatorny of the phylum Phoronida is estabLished and proposed to facilitate com-

munication in future anatomieal studics and taxonomieal works. A bricf discussion is given of the names (Latin and comman) of Ibis phylum. The present note must be considered as a preliminary work on nomenclature problems.

(racha (Latin name of the larva): an actinotroch. une actinotroque, eine Actinotroche. As general common Dame of the phylum (phoronid and actinotroch together), the following terms should he applied: the Phoronida, les phoronidiens, die Phoronideen.

3. Terminology 3.1. Pharonid termina/ogy

1. Introduction

The terminology used at the moment is frequcntly obscure and Dot entirely satisfactory; in sorne cases it is even ambiguous, the same ward being useful for several different senses or organs. Aiso the comparison of the terminology in English, French or German studies often leads to a very confusing situation. ln the present work, in order to facilitate communication and easy exchange between research workers, 1 propose a compiled list of main descriptive terms applied to the anatomy and morphology of Phoronida: this list is given in Englisb, French and German for each word. With the intention of establishing a c1ear and precise terminology in these languages, it was found necessary to create new terms in sorne of the languages. An exhaustive inventory through the bibliography of the phylum Phoronida has been compiled, especially in Cori (1889, 1939), Selys-Longchamps (1907), Hyman (1959), Zimmer (1964), Emig (1971, 1973, 1974). Ali (erms are presented in diagram form and indicated without aoy definition (these definitions are given by.the cited authors). Schemes and lerms have also been established to make easier the taxonomy and systematics of the species of Phoronida. 2. Names of the phylum Phoronida

Müller (1846) named Actinatracha the larva of the phoronids, which was considered by him to be an adult animal. The adult was described by Wright (1856) under the name Phoronis, which corresponds, according to this author, (0 a sumame of the Egyptian goddess Isis.' Gilchrist (1907) created a second genus, called Phoronopsis. In order to clarify our understanding, the present author proposes as common name for the adult: a phoronid, une phoronide, eine Phoronide (Phoronis and Phoronopsis are strictly used as genus names), and as common na me of ActinoZimmer (1964) cODsiders that this na me corresponds to Phoroneus. one of the two sons of the chief river-god Inachos, from the people of Argolis.

1

Three diagrams iIlustrate the termioology and their captioos group the main technical terms of the phoronid morphology and anatomy (Figs. 1-3). Yet, sorne remarks should follow

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Fig. 1. Diagram of a mature PllOronis hippocrepia vicwcd From the distal end (species characters, sec Emig, 1974). a, anus - anus - After. ep, epistome - épistome - Epistom. m, mouth - bouche - Mund. ne, nephridiopore - néphridiopore - Nephroporus. np, nephridial papi lia - papille néphridiale - Nephridienpapille. arg, accessory reproductive glands - glandes sexuelles annexes - Fortpflanzungsnebendrüsen. 01, lophophoral organ (accessory spermatophoral organ) - organe lophophoral - Lophophororgan. bng, basal nidamenlal gland - glande nidamenlaire basale - Basalbrutdrüse. Ing, tenlacular nidamental gland - glande nidamentaire tentaculaireTentakelbrutdrüse. 10, lophophore - lophophore - Lophophor. of, outer tentac1e row - tentacules externes (rangée de) - âussere Tentakelreihe. il, ioner tenlac1c row - tentacules internes (rangée de) - innere Tentakelreihe. /cc, lophophoral eoncavily - coneavilé lophophorale - LophophorkonkaviUl.l. /C, lophophoral cavily - cavité lophophorale - Lophophorkavitât. e, end of the lophophore (horseshoe-shaped, spiral-coiled or helicoidal) - pointe du lophophore (forme de fer à cheval, enroulée en spirale ou hélicoidale) - Lophophorarm (hufeisenformig, spiralig eingeroUt oder helikoidal). Zoologica Scripfa 4

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a Fig. 2. (a) Diagram of a phoronid, Phoronis psammophila (species charaeters. see Emig, ]974); (b) cross-section of the met8some; cavities and mesenteries are used to establish the conventional muscuJar formula (Selys~Longchamps. 1907) as follows:

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a. anus (sec Fig. 1). amp. ampuUa - ampoule - Ampulla. op, anal papiUa - papille anale - Analpapilla oder Afterpapille. cf. collaT raid - repli ~pidermique - Ringfalte. ~P. epistome (sec Fig. 1). ino, invagination - invagination - Einfaltung. m, moutb (sec Fig. 1). mtp, muscular trunlc part - r~gion musculaire du tronc - Muskelrumpflei!. Cal. coetam - cœlome - Cœlom oder Côlom. p, prosome, protocœlom (or epistomecœlom) prosome, protocœ!ome (ou cœlome de l'épistome) Prosome, Protocœlom (oder Epistomcœlom) m, mesosome, mesocœlom (or lophophorecœlom) Zoologico Scr;pto 4

mésosome, mésocœlome (ou cœlome du lophophore) Mesosoma, Mesosomcœlom (oder Lophophorcœlom) M, metasome, metacœlom (or trunkcœlom) métasome, métacœlome (ou coelome du tronc) Metasoma, Metasomcœlom (oder Rumpfcœlom) foc. left oral cavity - cavité orale gauche - linke OralhOhle TOC, right oral cavity - cavité oraIe droite - rechte Oralhohle loc,kft anal cavity - cavité anale gauche -linke Analhohle TOC, rigbt anal cavity - cavité anale droite - recbte AnalbObie d, diaphragm - diaphragme - Diaphragma. am, anal mesentery - mésent~re anal - Analmesenterium. mm, Median mesentery - mésentère médian - Mediaomesenterium. om, oral mesentery - mésentère oral - Oralmesenterium /lm, left lateraI mesentery - rMsentère lateral gauche - tinkes Lateralmesenterium. dm, rigbt lateral mesentery - mésentère latéral droit - recbtes Lateralmesenterium. DS, digestive system - tractus digestif - Darmtractus as, oesophagus - œsophage - CEsophagus. ps, prestomach - préestomac - Vannagen cs, ciliated strip - gouttière cilée - Wimpcrrinne. i, intestine - intestin - Dünndarm oder Intestinum. s, stomach - estomac - Magen. py, pylorus - pylore - Ringklappe.

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Fig. 4. Diagram of an actinotroch (Aclino1rol'hu brancltiata) near ils mctamorphosis. Sorne species show only one lenladc type: larval lentac!es become adult tcntades (Emig, 1973). Il stems also that the piriform organ docs not appear in ail Jarva. For a, i. m, 11(:', tu, PY• .l': sec Fig. 2, and gn: see Fig. S. ap. apical plate - plaque upicale - Scheitclplattc. tif, adull tentacle - tentacule adulte - Adulttentakel. It, larval tentade - tentacule larvaire - Larvaltentakcl. m. mesocœl - mésocœlc - Mcsocôl. M, metacœl - métacœle - Mctaci)1. M.f. mctasomal sac - sac mélasomique - Metasomdivcrtikcl. P, protocoel (or preoral lobe cœlom) - protocœlc (ou cœlome du lobe préoral) - Prolacôl (oder Pracorallappencœlom). per, perÎanal cilialcd ring - couronne ciliée pèrianale - .mal Wimperring. pn, proloncphridium - protonephridie - Protonephridium. o,piriform-organ (or sensory papilla) -organe piriforme - Piriformorgan • (oder Sinnesorgan). pol, preoral lobe (or prcoral hood) - lobe préoral - Praeorallappen (odcr Episphaere). psc, preseplal cavity - cavilé préseptale - Vorseptalhôhle. sd. stomacal divertÎculuOl - diverlicule stomacal - Darmdiverlikel. .ft', septum - seplum - Septum. v, vestibule - vestibule - Yeslibulum. vm, ventral mesenlery - mésentére ventral - Yentralmesenterium. Fig. 5. Schcme of Ihe lurval nervous syslem in an aClinotroch (AclinoIrocha A, after Zimmcr, 1964). cnr, collar nerve ring - nerf circulaire collaire - Kragenringnerv. gn, nervous ganglion - ganglion nerveux - Nervenganglion. Ine, laierai nerves of collar - nerfs latéraux collaires - Kragenlateral· nerven.

.,,",trl, major nerve ring of lobe - nerf circulaire majeur du lobe - Grossringnerv des Praeorallappens. 11/11'1, minor nerve ring of lobe - nerf circulaire mineur du lobe - Minderringnerv des Praeorallappens. tn, tenlucular nerve - nerf tentaculaire - Tentakelnerv. ("l, Iripartite median ncrve of lobe - triple nerf médian du lobe - drei Mcdiannervenfiiserzüge. SS, ser.sory papi lia - plaque sensorielle - Sinnespapille (sec also: po, Fig. 4). Fig. 6. Diagrams of Ihe musculalure of various actinotrochs (based on Cori, 1939, and Zimmer, 1964); ra: is no! rcprescnted in Ihe rigurc. clml', circular and longitudinal muscles of collar - muscles circulaires et longitudinaux colla ires - Ring- u. Langmuskeln des Kragens. l'mt, circular muscles of Irunk - muscles circulaires du tronc - Rumpfringmuskeln. dr, dorsal retractor muscle - musele rétracteur dorsal - Dorsalretraktormuskcl. co, eXlensor muscle of œsophagus - muscle extenseur de rœsophage Exlensormuskel des Œsophagus. Ims, levator muscle of melasomal sac - muscle élévateur du sac mélasomique - Levalormuskel des Metasomdivenikel. Irm, lobe retractor muscle - muscle rétracteur du lobe - Retraktormuskcl des Lappens: ra, retraetor muscle of anus - muscle rétracteur de l'anus - AfterrelraktormuskeL rami, radial and annular muscles of lobe - muscles radiaires et annu· laires du lobe - Radial und Ringmuskeln des Praeorallappens. Irm. trunk retraclor muscle - muscle rétracteur du tronc - Rumprretraktormuskel.

N, nephridium - ncphridie - Nephridium.

afferenl lophophoral ring vessel - arc lophophoral afférent austeigendes Lophophorringgefass 00, "accessory" vessel - vaisseau "accessoire" - "Nebengefass". cc, capîllary ccteum - cœcum sanguin - Cœcalgefiiss. eloo, ereerent lophophoral ring vessel - arc lophophoral effcrent abstcigendcs lophophorringgefâss. lib, left lateral branch - branche latéral gauche - linker lateralast. I/u, !cft lateral vessel - vaisseau latéral gauche - Iinkes Lateralgefass. lou, Lophophoral vessel - vaisseau lophophoral - Lophophoralgefâss. lu, laierai yesscl - vaisseau latéral - Lateralgefâss. mu, median vessel - vaisseau médian - Mediangefüss. rlb, right lateral branch - branche latéral droite - reehter Lateralast. rio, righl lateral vessel - vaisseau latéral droit - rechtes Lateralgefass. sbp, stomacal bloodplexus - sinus péristomacal- Magcnblutsinus. slu, secondary lateral vessel - vaisseau latéral secondaire - Nebenlateralgefâss. tu, tentaeular vessel - capillaire tentaculaire - Tentakelgefass.

ne, nephridiopore (see FiC. 1). ob. ascending nephridiaJ branch - branche néphridiale ascendante aufsteigender Nephridienkanal. /, funnel (if two: oral and anal funne!) - entonnoir (si deux: entonnoir oral et entonnoir anal) - Trichter (wenn zwei: Oral- u. Analtrichter). db, descending nephridial branch - branche néphridiale descendante absteigender Nephridienkanal NS, nervous system - système nerveux - Nervensystem. gn, nervous ganglion - ganglion nerveux - Nervcnganglioll m, ring nerve - nerf circulaire - Ringncrv. Ig/, Idt giant fibre - fibre nerveuse géante gauche - linker Lateralnerv oder Langnerv. rg/. right giant fibre - fibre nerveuse géante droite - rechter Lateralnerv oder Langnerv. enp, epilhelial nervous plexus - plexus nerveux épithéliale - epitheliales Nervennetz. Fig. 3. Diagrams of the circulatory system in Phorotlis Ol>alis (b) and in the other species (a).

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Zoologico Sl'ripta 4

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C. C. Emig

to define the morphological orientation that leads to an casier nomenclature of the phoronid body: the terros oral and anal (Fig. 2) take, respectively, the place of ventral and dorsal (ventral and dorsal must no longer he used in phoronids because of their ontogenetic orientation: dorsal corresponds al the median site mouth-anus); the ferros anterior, antérieur, vorder- and distal specify the regions towards the lophophoral extremity (Fig. 2) in contrast to posterior, postérieur, hinter w

and proximal towards the ampuJIa (these definitions do not take the ontogenetic disposition ioto account). Body. corps, Korper shows the whole animal; trunk. tronc, Rumpf. divided ioto two regions (Fig. 2), particularizes the cylindrical body part ( ~ metasoma).

3.2. Actinotroch termin%gy The actinotroch terminology is illustratcd by three schemes and explained by their captions (Figs. 4-6). The larval body is composed of two extemal regions, the preoral lobe and the trunk. It presents a dorsal and ventral side (Fig. 4): the body orientation appears different in the actinotroch and the phoronid. Special terms are used to describe corresponding organs in the adult and the larva in order to avoid all possible future confusing situations (as proposed by Emig, 1973): in the phoronid., the name of the cœlomic cavities ends in -cœlorn (Fig. 2) and in the actinotroch, in -cccl (Fig. 4); also, the term diaphragm is used in the adult and septum in the larva.

4. Conclusion The present short terminology of Phoronida only affects the main organs and specifie terms of this phylum. There are many more. Thus this tenninology represents ooly a preIiminary and basic work on nomenclature problems and the author considers this paper as the beginning of a cJear, precise and unambiguous terminology in Phoronida. 1 al50 suggest that in future ail new terms applied to this phylum are established in the three languages (English. French, German) to facilitate communication and to avoid ail confusion.

References Cori, C. J. 1889. Untersuchungen über die Anatomic und Histologie der Gattung Phoronis. - Z. wiss. Zool. 51: 480-568. - 1939. Phoronidea. - Bronn's KI. Ordn. Tierreîchs 4 (4): 1-183. Emig, C. C. 1971. Taxonomie et systématique des Phoronidiens. Bull. Mus. natn. Hist. nat., Paris (Zool.) 8: 469-568. - 1973. Les processus de l'ontogenèse, comparés à ceux de la régénération des Phoronida. - Z. Morph. Tiere 75: 329-350. - 1974. The systematics and evolution of the phylum Phoronida. - Z. zool. Syst. Evolut.-forsch. 12: 128-151. Gilchrist, J. D. 1907. New fonns of the Hemichordata from South Arrica. - Trans. S. Arr. phil. Soc. 17: 151-l76. Hyman, L. H. 1959. The Invertebrates, Vol. 5. - McGraw-HiII, New York. Müller, J. 1846. Bcricht über einige neue Thierforrnen der Nordsee. Arch. Anal. Physiol. 1846: 101-104. Selys-Longchamps, M. de 1907. Phoronis. - Fauna Flora Golf. Neapel 30, 1-280. Wright, T. S. 1856. Description of two tubicolar animaIs. - Proc. R. Soc. Edinb. 1: 165-167. Zimmer, R. L. 1964. Reproductive biology and development of PhOTOnida. - Univ. Microfilm, Ann Arbor.

Zoologica Scripta 4

Dr Christian C. Emig Station Marine d'Endoume Rue de la Batterie·des-Lions F-J3007 Marseille France

Printed 1975-06-20