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Zoologica Scripta. Vol. 4, 37-40, 1975
Terminology in PhoronidaChristian 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 anatornyof the phylum Phoronida is estabLished and proposed to facilitate communication in future anatomieal studics and taxonomieal works. Abricf discussion is given of the names (Latin and comman) of Ibisphylum. The present note must be considered as a preliminary work onnomenclature problems.
(racha (Latin name of the larva): an actinotroch. une actino
troque, eine Actinotroche. As general common Dame of thephylum (phoronid and actinotroch together), the followingterms should he applied: the Phoronida, les phoronidiens, die
Phoronideen.
3. Terminology
ne np
3.1. Pharonid termina/ogy
Three diagrams iIlustrate the termioology and their captioosgroup the main technical terms of the phoronid morphology
and anatomy (Figs. 1-3). Yet, sorne remarks should follow
Fig. 1. Diagram of a mature PllOronis hippocrepia vicwcd From the distalend (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 - Fort
pflanzungsnebendrüsen.01, lophophoral organ (accessory spermatophoral organ) - organe
lophophoral - Lophophororgan.bng, basal nidamenlal gland - glande nidamenlaire basale - Basal
brutdrüse.Ing, tenlacular nidamental gland - glande nidamentaire tentaculaire
Tentakelbrutdrüse.10, lophophore - lophophore - Lophophor.
of, outer tentac1e row - tentacules externes (rangée de) - âussereTentakelreihe.
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 heli
coidal) - pointe du lophophore (forme de fer à cheval, enroulée enspirale ou hélicoidale) - Lophophorarm (hufeisenformig, spiraligeingeroUt oder helikoidal).
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1. Introduction
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 secondgenus, called Phoronopsis.
In order to clarify our understanding, the present authorproposes as common name for the adult: a phoronid, unephoronide, eine Phoronide (Phoronis and Phoronopsis are
strictly used as genus names), and as common name of Actino-
The terminology used at the moment is frequcntly obscure
and Dot entirely satisfactory; in sorne cases it is even ambi
guous, the same ward being useful for several different sensesor organs.
Aiso the comparison of the terminology in English, Frenchor 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 compiledlist of main descriptive terms applied to the anatomy andmorphology of Phoronida: this list is given in Englisb, French
and German for each word. With the intention of establishinga c1ear and precise terminology in these languages, it wasfound necessary to create new terms in sorne of the languages.
An exhaustive inventory through the bibliography of thephylum 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 taxonomyand systematics of the species of Phoronida.
1 Zimmer (1964) cODsiders that this na me corresponds to Phoroneus.one of the two sons of the chief river-god Inachos, from the people ofArgolis.
Zoologica Scripfa 4
38 Co Co Emig
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aFig. 2. (a) Diagram of a phoronid, Phoronis psammophila (speciescharaeters. see Emig, ]974); (b) cross-section of the met8some; cavitiesand mesenteries are used to establish the conventional muscuJarformula (Selys~Longchamps. 1907) as follows:
om
IIm loc 1roc rlmlac rae
am
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 - Muskelrumpf
lei!.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)
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 OralhOhleTOC, right oral cavity - cavité oraIe droite - rechte Oralhohleloc,kft anal cavity - cavité anale gauche -linke AnalhohleTOC, 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 Lateral
mesenterium.dm, rigbt lateral mesentery - mésentère latéral droit - recbtes Lateral
mesenterium.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.
Zoologico Scr;pto 4
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·?fipcr
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Fig. 4. Diagram of an actinotroch (Aclino1rol'hu brancltiata) near ilsmctamorphosis. Sorne species show only one lenladc type: larvallentac!es become adult tcntades (Emig, 1973). Il stems also that thepiriform 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 onCori, 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 - Rumpf
ringmuskeln.dr, dorsal retractor muscle - musele rétracteur dorsal - Dorsalretraktor
muskcl.co, eXlensor muscle of œsophagus - muscle extenseur de rœsophage
Exlensormuskel des Œsophagus.Ims, levator muscle of melasomal sac - muscle élévateur du sac méla
somique - Levalormuskel des Metasomdivenikel.Irm, lobe retractor muscle - muscle rétracteur du lobe - Retraktormuskcl
des Lappens:ra, retraetor muscle of anus - muscle rétracteur de l'anus - Afterrelrak
tormuskeLrami, 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 - Rumprre
traktormuskel.
N, nephridium - ncphridie - Nephridium.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 NephridienkanalNS, nervous system - système nerveux - Nervensystem.
gn, nervous ganglion - ganglion nerveux - Nervcngangliollm, 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 - epithe
liales Nervennetz.Fig. 3. Diagrams of the circulatory system in Phorotlis Ol>alis (b) and inthe other species (a).
0100, afferenl lophophoral ring vessel - arc lophophoral afférentausteigendes 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 - Neben
lateralgefâss.tu, tentaeular vessel - capillaire tentaculaire - Tentakelgefass.
Zoologico Sl'ripta 4
40 C. C. Emig
to define the morphological orientation that leads to an casiernomenclature 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 correspondsal 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, hinterw
and proximal towards the ampuJIa (these definitions do nottake the ontogenetic disposition ioto account). Body. corps,
Korper shows the whole animal; trunk. tronc, Rumpf. dividedioto 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 thephoronid.
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 pre
Iiminary 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 esta
blished 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 Histologieder 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 régé
nération 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 PhOTO
nida. - Univ. Microfilm, Ann Arbor.
Zoologica Scripta 4
Dr Christian C. EmigStation Marine d'EndoumeRue de la Batterie·des-LionsF-J3007 MarseilleFrance
Printed 1975-06-20