eunice siciliences

7/22/2019 Eunice Siciliences

1/23

Makalah Bahan Alam Kelautan

Eunice siciliences(Annelida : Polychaeta, cacing laut/nyale)

sebagai bahan Antibakteri


2/23

BAB I

Pendahuluan

Annelida berasal dari bahasa latin, Annulus yang berarti cincin dan iodos yang berarti

bentuk. Annelida berarti cincin kecil dan tubuh bersegmen yang mirip dengan serangkaian

cincin yang menyatu merupakan cirri khas cacing filum Annelida. Terdapat sekitar 1.!!!

spesies filum Annelida, yang pan"angnya berkisar antara kurang dari 1 mm sampai # m pada

cacing tanah Australia. Anggota filum Annelida hidup di laut, sebagian ada "uga di air ta$ar,

dan tanah lembab.

%alam klasifikasi modern, Annelida dibedakan men"adi beberapa kelas, yakni

Polychaeta, &lygochaeta, dan 'irudinae. %asar pengklasifikasian ini mengacu pada ciri

morfologi, fisiologi, maupun anatomi dari he$anhe$an ini. %engan demikian, kita akan

menemukan adanya perbedaan ciri diantara tiap kelas dalam filum ini.

Annelida memegang peranan yang sangat penting bagi ekosistem, terutama dalam

kehidupan manusia. eberapa spesies dalam filum Annelida memegang peranan dalam upaya

penyuburan tanah dan ada pula yang men"adi bahan makanan dan obatobatan yang telah

dikembangkan se"ak dulu. *elain itu, beberapa spesies dari filum ini dapat men"adi ekoparasit

seperti halnya pacet yang mengisap darah. erbagai peranan tersebut men"adikan pengetahuan

mengenai filum ini men"adi sangat penting.

erdasarkan uraian pada pendahuluan latar belakang tersebut, maka saya menganggap

sangat perlu untuk menyusun makalah mengenai Annelida. 'al ini saya harapkan dapat

menambah pengetahuan bagi saya khususnya dan bagi kita semua mengenai filum ini

terutama dari spesies Eunice Siciliences. +elihat dari beberapa penelitian sebelumnya

mengenai spesies ini, saya merasa sangat men"an"ikan "ika Eunice Siciliencesini di"adikan

sebagai bahan makalah dan sebagai bahan penelitian. arena prospek kedepannya sangat

bagus dan men"an"ikan sebagai sumber obat baru.


3/23

BAB II

Pembahasan

A. Klasifikasi

ingdom : Animalia

-ilum :Annelida

elas :Polychaeta

*pesies :Eunice Siciliences

B. Anatomi dan Fisiologi

Anantomi filum Annelida secara umum berbentuk gelang yang memiliki tubuh

meman"ang, simetri bilateral, bersegmen, dan permukaannya dilapisi kutikula. %inding

tubuhnya dilengkapi otot. +emiliki prostomium dan system sirkulasi, system pencernaan

lengkap, system ekskresi sepasang nephrida di setiap segmen. *ystem syaraf tangga tali.


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*ystem respirasi terdapat pada epidermis. ebanyakan cacing Annelida hidup akuatik di laut

dan terrestrial

C. System Gerak

Polychaeta merupakan Annelida berambut banyak. Tubuhnya dibedakan men"adi

daerah kepala (prostomium), antenna, dan sensor palpus. Pada setiap segmen tubuhnya

terdapat sepasang struktur seperti dayung (parapodia) yang berfungsi sebagai alat gerak dan

"uga mengandung pembuluh darah sehingga "uga berfungsi sebagai insang untuk bernapas.

Pergerakan disebabkan oleh perpaduan gerak antara parapodia, otot dinding tubuh, dan cairan

rongga tubuh.

D. System es!irasi

Parapodia selain digunakan sebagai alat gerak "uga digunakan sebagai alat pernapasan.

Polychaeta bernapas dengan insang ketika berada di perairan, namun pertukaran gas ia

permukaan tubuh ter"adi secara difusi. Tiap ruas terdapat insang, kecuali u"ung anterior dan

posterior.

". System Pen#ernaan

Terdapat ruas pada anterior yang mengandung mulut disebut peristomium. uas terakhir

atau pigidium mengandung anus. Tipetipe system pencernaan :

a. aptorial feeder : aertebrata kecil ditangkap dengan faring atau proboscis yang

di"ulurkan, terdapat rahang kitin.

b. %eposit feeder : menelan pasir dan lumpur dalam lorong 0 bahan organic dicerna dan

partikel mineral dikeluarkan le$at anus, atau melalui tentakel silia yang berlendir.

c. -ilter feeder : tidak punya proboscis tutup kepala dilengkapi radiola untuk menyaring

detritus dan plankton.


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F. System "kskresi

*ystem ekskresinya belum sempurna dan masih menggunakan organorgan khusus

sebagai alat ekskresi. Tidak mempunyai pembuluh darah berupa protonefridia solenosit,

namun mempunyai pembuluh darah berupa metanefridia. Alat ekskresi terdiri dari nefrostom

yaitu corong bersilia, nefridal kanal yaitu pembuluh ekskresi, bermuara pada neuropodium.

efridia "uga berfungsi sebagai alat osmoregulasi.

G. System Syaraf

*ystem syarafnya berupa system syaraf tangga tali. Alat indra utama dari mata,

2nuchal organ3, dan statocyst. +ata berkembang baik (errantia), bintik mata/tidak ada

(sedentaria) dan berfungsi sebagai fotoreseptor. uchal organ berfungsi sebagai kemoreseptor

untuk mendeteksi makanan. *el peraba terdapat diseluruh tubuh,terutama parapodia dan

kepala.

$. e!roduksi

Perkembangbiakan secara seksual. Pembuahan dilakukan diluar tubuh terutama

didalam air. etina dan "antan ditandai dengan perbedaan $arna dimana betina ber$arna hi"au

dan "antan ber$arna coklat. Pada saat ka$in tubuh "antan dan betina bergerakgerak menu"upermukaan air dengan gerakan spiral. 4erakan spiral ini kemudian menyebabkan sperma dan

sel telur keluar. Pertemuan sel telur dan sperma menghasilkan 5igot yang kemudian menu"u

dasar laut. Telur yang dibuahi ini (5igot) akan men"adi lara yang disebut trakofora.

I. $abitat

'abitatnya di lautan. iasanya dalam pasir atau menggali batubatuan pasang surut air

laut ataupun membentuk tabung. 'idupnya bersembunyi sehingga sulit untuk ditemukan.

%. "kologi

*ecara ekologi berperan penting sebagai makanan he$an dasar seperti ikan dan udang.

Pada ekosistem terumbu karang turut mrnyumbang kalsium karbonat.


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K. Penyebaran

%i 6ndonesia cacing "enis ini umumnya muncul di daerah Pantai *ager, 7ombok

*elatan. +erupakan he$an tyahunan yang muncul hanya pada $aktu$aktu tertentu, sekitar

pertengahan -ebruari sampai a$al +aret. 8acing yang ber$arna$arni ini hanya muncul pada

musim ka$in. 9ang umum ditemukan di 7ombok adalah "enisEunice Siciliences.

&. Senya'a Kimia (ang Dihasilkan

a. romopyrrole.

b. romophenol : mempunyai sifat anti mikroba dalam hal ini bromophenol mampu

menghambat respirasi mikroba.

c. ahan aktif (natural product) lainnya yang dapat mematikan atau menghambat

pertumbuhan makhluk hidup lainnya (seperti bakteri), khususnya pada "urnal ini

adalah bakteri benthos.


7/23

. lack *ea/+editerranean ;nironment= (?!1?)

")I"* A+IC&"

A #he#k,list of !oly#haete s!e#ies -Annelida Poly#haeta/ from the

Bla#k Sea

G0ley Kurt 1ahin23 Melih "rtan 45nar6

1%epartment of iology, -aculty of Arts and *ciences, *inop @niersity, !!!, *inop,T@;9?%epartment of 'ydrobiology, -aculty of -isheries, ;ge @niersity, #1!!, ornoa65mir,T@;9BCorres!onding author gkurtsahinCsinop.edu.tr

Abstra#t

Polychaetes from the lack *ea $ere reie$ed based on the aailable literature. 7ists of thealid species, reassigned species and dubious species (nomen dubium or indeterminablespecies) reported from the region are proided. The checklist comprises ?#= alid speciesbelonging to > families. T$enty three species belonging to = families $ere considered asnomen dubium or indeterminable. -iftyone species reported from the lack *ea bet$een1=D= and ?!11 hae been synonymised $ith other species. Among the lack *ea countries,the coast of ulgaria is represented by the highest number of polychaete species (1E? species),follo$ed by the coasts of @krania (1D1 species) and omania (11 species). The highestnumber of species (11E species) $ere found on sandy substratum, the lo$est number of

species ( species) on sponges. ine alien polychaete species (Hesionides arenaria,Streptosyllis varians, Glycera capitata, Nephtys ciliata, Polydora cornuta, Prionospiopulchra, Streblospio gynobranchiata, Capitellethus dispar and Ficopomatus enigmaticus)$ere reported from the lack *ea up to no$. The most speciose families in the area $ere*yllidae (#? species), *pionidae (#1 species), Phyllodocidae (1D species) and ereididae (1species). y presenting a complete oerie$ of the species reported from the area, this $orksummarises our current kno$ledge about the diersity of the lack *ea polychaetes andproides a database for future studies.

Key'ords Polychaeta, annelida, checklist, lack *ea

Introdu#tionThe lack *ea is one of the largest enclosed seas in the $orld, coering an area of about >.? F1! km?0 the maFimum depth of the sea is ??1? m and the total olume of $ater of #>.!!!km#. +ost of this $ater (>?#.!!! km# that lies belo$ a depth of 1!G?!! m) is anoFic andcontaminated $ith hydrogen sulphide. The lack *ea drainage basin coers more than ?million km? and more than 1D! million people lie in this area. The total length of the lack*ea coastline is oer >>!! km. *iF countries (Turkey, ulgaria, omania, @kraine, ussian-ederation and 4eorgia) border the lack *ea. The large ariety of geomorphologic types of


8/23

these coasts corresponds to different geological enironments surrounding the lack *ea(Panin ?!!).The scientific researches concerning polychaetes in the lack *ea $ere started during the late1=!!s. The first detailed studies on polychaetes in the lack *ea $ere made by obret5ky

(1=D=, 1=!, 1=1, 1==1) and 85erniasky (1==!, 1==1, 1==?). obret5ky performed hisstudies in the 4ulf of *eastopol and 85erniasky focused on polychaetes distributed alongthe coasts of 9alta and *ukhumi.The polychaete species liing along the Turkish coast of the lack *ea $ere reported in a fe$papers (Pinar 1E>0 at et al. ?!!!0 Hinar and 4InlJgJr%emirci ?!!0 AKLrbaM et al. ?!!=04Ikkurt et al. ?!!=0 4I5ler et al. ?!!E, ?!1!0 %agli and 8inar ?!11). Polychaetes inhabitingthe Prebosphoric region of the lack *ea $ere studied by akuboa (1E>=), +arino (1EEb,1ED>), 8aspers (1ED=), %umitrescu (1ED!, 1ED?), ullier (1ED#), 4illet and Nnsal (?!!!) and@ysal et al. (?!!?).

esult and Dis#ussion

Polychaete iversityAs a result of the compilations of papers on polychaetes reported from the lack *ea bet$een1=D= and ?!11, a total of ?#= alid species belonging to > families $ere determinedThe aims of this study is to summarise the aailable information about the polychaetediersity in the lack *ea and to prepare a database for the lack *ea polychaetes.

Materials and Methods

The checklist for the polychaete species reported from the lack *ea $as made follo$ing aneFhaustie bibliographic reie$ and analysis of the published articles. -or the checklist,species names recorded in publications concerning the lack *ea $ere compiled, checked fortheir alidity and synonyms made eident. The alid species, dubious species (considered asnomen dubium or indeterminable) and synonymised species are gien in the different tables(Tables 1, ?, #). The polychaeta diersity in the lack *ea $ere eFamined according to thecountries (including Prebosphoric region): Turkey, ulgaria, omania, @kraine, ussia and4eorgia. 6n each country, all reports of the species are gien as numbers in Table 1 that areindicated in the reference section. Ohen aailable, ecological data in the papers are alsoealuated. All calculations are based on the alid species.

Polychaete !amilies

The most speciose families in the lack *ea are sho$n in -igure 1. *yllidae and *pionidae arethe most dominant families in terms of the number of species (respectiely, #? and #1species), follo$ed by Phyllodocidae (, 1D species), ereididae (D, 1 species),

ephtyidae (>, E species) and *abellidae (>, E species). Thirteen families (*igalionidae,8hrysopetalidae, Pisionidae, Amphinomidae, Pilargidae, 4oniadidae, Arabellidae,8tenodrilidae, -labelligeridae, *ternaspidae, Trichobranchidae, *accocirridae andParergodrilidae) are represented by only one species.


9/23

eferen#es

Agirbas, ;., 4o5ler, A. +., *ahin, 8., 'acimurta5aoglu, . (?!!=) %oKu aradeni53dedaKLlLm gIsteren "lva fasiesinin poliket faunasL.#ournal o! Fisheries Science.com ?: >?>#1.

Alemo, *.


10/23

7at. Am. . ARuat. es., #D(1): #D1, ?!!= Polychaeta from the +eFican Pacific%&6: 1!.#=D/ol#Dissue1fullteFt>

Poly#haetes -Annelida Poly#haeta/ des#ribed for the Me7i#an

Pa#ifi# an histori#al re8ie' and an u!dated #he#klist Pablo$ern9nde:,Al#9ntara23 Mar;a Ana +o8ar,$ern9nde:6 ). The species descriptions for the +eFicanPacific can be diided into four main periods: the first, during the 1E1!s, includes mainly8hamberlin3s studies of deep sea fauna. The second, in the 1E>!s, comprises studies carried

out by io"a (intertidal 5one) and 'artman (including specimens from the Allan 'ancock-oundation collection). The third period began around the 1E!s $hen -auchald3s studies$ere published and currently alid species $ere described for $estern +eFico. The fourthperiod began in the 1E=!s and continues to date, being characteri5ed by descriptions of speciesdone mainly by +eFican scientists. The differences in the number of species described duringeach period and for the different regions of the +eFican Pacific are directly related to thesampling effort carried out along these coasts.

Key'ords Polychaeta, benthos, biodiersity, geographic distribution, +eFican Pacific.

&os !oliuetos -Annelida Poly#haeta/ des#ritos en el Pa#;fi#o me7i#ano

re8isi?n hist?ri#a y lista faun;sti#a a#tuali:ada

"S=M">. *e presenta una lista taFonUmica de las especies de poliRuetos descritos en elPacVfico meFicano y una reisiUn histUrica de su estudio. ;l listado incluye referenciasnomenclaturales, informaciUn sobre la localidad tipo y sinonimias basadas en reisionessistemWticas. @n total de #1# especies y ?1 gXneros han sido descritos en el Wrea de estudio, deellas, ?= especies son actualmente Wlidas. ;l ?= de las descripciones de las especiesWlidas no incluyeron el hWbitat de la localidad tipo. 7as 1EE especies Wlidas restantes fueron


11/23

descritas en una amplia ariedad de hWbitats: algas (11 especies), manglares (?), fondos duros(??), fondos blandos (plataforma continental D especies0 mar profundo = especies),entilas hidrotermales (1) y otros (parWsitos, formas larales planctUnicas, epitocas) (>). 7adescripciUn de especies en el PacVfico +eFicano puede ser diidida en cuatro periodos

principales: el primero en la dXcada de 1E1! incluye principalmente los traba"os de8hamberlin sobre la fauna de aguas profundas. ;l segundo, en la dXcada de 1E>!, comprendelos estudios reali5ados por io"a en la 5ona intermareal y por 'artman Rue incluyUespecVmenes de la colecciUn de la Allan 'ancock -oundation. ;l tercer periodo se iniciUalrededor de 1E! en Rue se publicaron los estudios de-auchald Rue describen especies actualmente Wlidas del oeste de +XFico. ;l cuartoperiodo se iniciU en la dXcada de 1E=! y continYa hasta la fecha, caracteri5ado por ladescripciUn de especies reali5ada por cientVficos meFicanos. 7as diferencias obseradas en elnYmero de especies descritas en cada periodo y en cada regiUn del PacVfico meFicano estWnasociadas con el esfuer5o de muestreo lleado a cabo a lo largo de estas costas.

Palabras #la8e Polychaeta, bentos, biodiersidad, distribuciUn geogrWfica, PacVficomeFicano.7at. Am. . ARuat. es., #D(1), ?!!=

I>+JD=C+IJ>

The study of polychaetes in the +eFican seas started during the second half of the nineteenthcentury, $hen inberg (1=) recorded the amphinomid Hermodice carunculata (Pallas,1D). @nfortunately, he did not record the sampled site precisely, only noting Z+eFico[ asthesampling location. 'o$eer, $e consider it plausible that the amphinomid $as collected inthe


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nomenclature and the criteria for taFa definitions hae al$ays been common. This is in partdue to the relatiely freRuent reisions of genus and synonym procedures, but also because thelack of integratie taFonomic monographs and updated checklists for this region has causedconfusion in nomenclature and in the real distribution ranges of many species. This study aims

to present an updated list $ith the original and currently alid names for the polychaetespecies $hose locus typicus is in the +eFican Pacific, to carry out an historical reie$, and toanaly5e the recorded species distributions and habitats. Oith this, $e hope to contribute to the$orld$ide effort to standardi5e polychaete nomenclature, in this case, in the ;astern Pacific.

MA+"IA&S A>D M"+$JDS

The checklist for the polychaete species described from the +eFican Pacific $as madefollo$ing an eFhaustie bibliographic reie$ and analysis of the original descriptions. The listincludes the nomenclature reference for each polychaete species (original name, year ofpublication, pages, figures), data for the type locality, depth and habitat (as complete aspossible, based on the information in the original description), and the current alid name

(citing the author $ho introduced the taFonomic change). 6n addition, the number of speciesfor each biogeographic proince is indicated. The state to $hich the type locality belongs isnoted only in order to facilitate the search $ithin the country since the political diisions haeno biogeographic meaning. At the end of this paper, the complete references of the consultedpublications are proided. 6n this $ay, $e hope to reduce the time inested $hen furtherinformation is needed for other specific ob"ecties.

Me7i#an Pa#ifi# #oasts and biogeogra!hi# !ro8in#es

The +eFican Pacific littorals coer >,!> km and are located bet$een 1_ and #?_, alongthe Tropic of 8ancer, crossing the southern a"a 8alifornia region (-ig. 1). 6ts $ater masses ofdifferent origins (arctic, subarctic, tropical, subtropical) (Oirtky, 1ED) combine $ith thediersified physiography to create many habitats: muddy and sandy beaches, rocky shores,bays, coastal lagoons and estuaries, about ?>! islands and islets of different origins andgeological ages, mangroes and seagrass beds, and een actie eFpansion centers in thebottoms of the central 5ones $ith their corresponding hydrothermal ent areas (4uaymasasin and ?1_). The different marine regions in the +eFican Pacific are separated by steepthermal gradients to the north and south and by an area of open ocean (;ast Pacific arrier) tothe $est. The boundaries of the biogeographic proinces used herein are taken from seeralauthors: riggs (1EE), 'endrickF (1EE?), and 'astings (?!!!), $ho separated them on thebasis of the distribution patterns of fishes and marine inertebrates. The north$estern littoralsof +eFico are enironmentally classified as $armtemperate and belong to the 8alifornianbiogeographic proince $hich goes from south of Point 8oncepcion (near *anta arbara,8alifornia) do$n to northern +agdalena ay, south$est of the a"a 8alifornia Peninsula(rusca, 1E=!0 'astings, ?!!!) (-ig. 1). The topographic characteristics of the a"a 8aliforniaPeninsula shoreline protect it from the cold southbound 8alifornia 8urrent and help to create a$armtemperate area along the southern 8alifornia coasts (riggs, 1EE). The Tropical;astern egion eFtends from the 4ulf of 8alifornia south$ard to northern Peru, $here a fauna$ith tropical affinities dominates (rusca, 1E=!0 'astings, ?!!!). 6n the +eFican Pacificarea, the tropical region encloses the 8ortes, the +eFican, and the Panamanian biogeographicproinces. The 8ortes Proince corresponds to the 4ulf of 8alifornia, including the entireeastern and most of the south$estern coasts of the a"a 8alifornia peninsula. The 4ulf of


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8alifornia is primarily tropical in origin but could $ell be considered subtropical: itsnorthernmost areas are primarily populated by eurythermal tropical species, $hereas thesouthern fauna gradually eoles spatially to a more typical tropical biota (rusca, 1E=!). The+eFican Proince includes the coasts from +a5atlWn (*inaloa) do$n to the 6sthmus of

Tehuantepec in southern +eFico. At about 1D_, in the northern part of the 4ulf ofTehuantepec, another faunal change can be obsered0 the area from that point south$ard tothe 4ulf of 4uayaRuil (#_*) is kno$n as the Panamic Proince (riggs, 1EE).

"S=&+S

+a7onomi# ri#hness of !oly#haetes in Me7i#an Pa#ifi# 'aters

6n the +eFican Pacific, the number of polychaete species recorded to date is approFimately1,1!! ('ernWnde5AlcWntara, ?!!?)0 of these, #1# species (?= currently alid) and ?1 generahae their locus typicus in the +eFican Pacific (Table 1). The serpulid Eupomatus humilis(currently alid asHydroides humilis) (ush, 1E!>) $as the first species found $hose locustypicus is in the +eFican Pacific. %uring the t$entieth century, significant contributions $ere

made to the taFonomy of +eFican polychaetes and #1# polychaete species, grouped in 1D?alid genera and >1 families, $ere described. As our kno$ledge on the systematics of thegroup hae increased, the taFonomicstatus has changed freRuently and so, of the original #1#species described, only ?= polychaete species are alid no$adays. *pecies belonging to thefamilies Acoetidae (Polyodontes cali!ornicus) and 7opadorrhynchidae (*opadorrhynchusparvus) hae been synonymi5ed0 therefore, these families hae no ne$ species in the +eFicanPacific region. 6n 1EDE, 'artman established the family *abellongidae for the genusSabellonga (as monotype) and the species Sabella dis+uncta, collected at 11 m in 8edros6sland, a"a 8alifornia. 'o$eer, according to -it5hugh (1E=E), 'artman interpreted theentral lips or maybe the entral sacs, as palps, and the species type (S. dis+uncta) lacks atentacular cro$n but uncini and chaetae resemble those found in *abellidae. 'e thenconcluded that the genus Sabellonga is a member of the family *abellidae. The taFonomicstatus of D species has changed, but they are still considered to be alid. -or three of thesespecies, taFonomic ad"ustments hae been made at the family leel: according to lake(1EE#),Pilargis mirasetis, initially included in the family Pilargidae, belongs to the genusSantelma, in the family autiliniellidae established in 1EE# by lake0 Flabelligellamacrochaeta(-labelligeridae) is a member of the family Acrocirridae by 7ight (1E=)0 andthe serpulid Hydroides glandi!erum is a type of the genus lgaharmania by io"a (1E>1b)$ithin the *erpulidae. 8onsidering the number of currently alid species, the ereididae and&nuphidae are the richest families (?> species each), follo$ed by the Polynoidae (1E species),the *erpulidae (1 species), and the Ampharetidae and 7umbrineridae (1D and 1 species,respectiely). -or 1!! species, of $hich E are alid, the habitat of their type localities $asnot indicated in the original description. 'o$eer, the distribution of the remaining alidspecies on the soft bottoms of the study area sho$s that D alid species (?#) $ere collectedon the continental shelf and = alid species (?=) in the deepsea. The remaining habitatsrecorded include less species: ?? species on hard bottoms from the littoral and sublittoral5ones, 1 species from the hydrothermal ents both at 4uaymas and ?1_, and 1 species inother habitats. According to their locus typicus, the number of polychaete species aries ineach geographic 5one. 6n the $armtemperate 8alifornian Proince, D alid species haebeen identifiedin the 8ortes Proince, $ith subtropical conditions, the number of species is thehighest (1 alid species)0 finally, the +eFican Proince in the southern +eFican Pacific


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(tropical characteristics) had ># alid species (Table #). *tudies of the polychaete faunadistributed along the southern border of the +eFican Pacific (Panamic Proince) are scarceand only four ne$ alid species from this area hae been identified.

DISC=SSIJ>The description of #1# species (?= currently alid) from the +eFican Pacific oer the lasthundred years at a rate of ?.E species per year is largely due to the monumental $ork done by-auchald in the 1E!s. 'e alone described E# of the currently alid species (#>), $hich ismore than a third of the total. These contributions made +eFico3s Pacific 5one the region $iththe largest number of described taFa in the country. -rom 1E=! on, the rate of ne$ speciesdescribed decreased to t$o species per year due to a reduction in sampling and in taFonomicstudies carried out after -auchald3s $ork and not because fe$er ne$ species remain to bedescribed in the +eFican littorals. 6n fact, in ?!!D, four ne$ species $ere described('ernWnde5AlcWntaraet al., ?!!D0 *ala5arC"S

anse, . 1E!. The small species ofEuchone +almgren (*abellidae, Polychaeta). Proc. iol.*oc. Oash., #: #=>!=.


15/23

astidaQaala, .. 1EE!.*ycastopsis rio+ai, a ne$ species of polychaete (Polychaeta:ereidae) from the 4ulf of 8alifornia. e. iol. Trop., #=(?): >1>?!.

astidaQaala, .. \ .A. de 7eUn4on5Wle5. ??!!?!!?. A ne$ species of Hydroides

(Polychaeta: *erpulidae) from $estern +XFico. . +ar. iol. Ass. @., ?: 1!#1=.

erkeley, ;. \ 8. erkeley. 1E#=.-amphobrachium longisetosum, sp. n., $ith someobserations on the regeneration of the speciali5ed anterior setae. Ann. +ag. atur. 'ist., 1:>?=>#.

erkeley, ;. \ 8. erkeley. 1E#E. &n a collection of Polychaeta, chiefly from the $est coastof +eFico. Ann. +ag. atur. 'ist., series 11, #(#=): #?1>D.

erkeley, ;. \ 8. erkeley. 1E>1. &n a collection of Polychaeta from southern 8alifornia.ull. *outh. 8alif. Acad. *ci, >!: 1DD!.

erkeley, ;. \ 8. erkeley. 1ED!. otes on some Polychaeta from the $est coast of +eFico,Panama, and 8alifornia. 8an. . Qool., #=: #D?.lake, .A. 1E=1.Polydora and%occardia species (Polychaeta: *pionidae) from $estern+eFico, chiefly from calcareous habitats. Proc. iol. *oc. Oash., E#(>): E>ED?.

lake, .A. 1E=. Polychaeta from the icinity of deepsea geothermal ents in the easternPacific, 1. ;uphrosinidae, Phyllodocidae, 'esionidae, ereididae, 4lyceridae, %orilleidae,&rbiniidae and +aldanidae. Proc. iol. *oc. Oash., D: D1!1.


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+a7onomy -Annelida3 Poly#haeta/ +he state of affairs

$A( A. +"> $J)" < "&">A K. K=PI(A>J)A

Zoological Museum, University of Amsterdam POB 94766, 19 !" Amsterdam, "#e

$et#erlands

%&mail' ()A)ten(ove*uva)nl

+%art# and %nvironmental ciences, University of Adelaide A -- Adelaide Australia1

%&mail' lena).u/riyanova*gmail)com,elena.*ynu)ac)0/

Introdu#tion

The family *erpulidae is a discrete group of sedentary calcareous tube$orms $ithin the largeclade *abellida, $hich shares a presence of radiolar cro$n and separation of the body into

thoracic and abdominal regions, as diergence from the usually rather uniformly segmentedmotile polychaete form. The ie$s on the relationships $ithin *erpulidae hae undergonemany changes oer the years (see upriyanoa et al. ?!!D for the details). The *erpulidaeafinesRue, 1=1 had been traditionally diided into subfamilies *erpulinae afinesRue, 1=1(although probably in most papers attributed to +ac7eay, 1=>!) and *pirorbinae 8hamberlin,1E1E until io"a (1E?#) established the subfamily -ilograninae. Pillai (1ED!) included brackish$ater serpulid genera in the subfamily -icopomatinae but ten 'oe \ Oeerdenburg(1E=) reised the group and placed all its genera in the genus Ficopomatus. @chida (1E=)created 11 subfamilies and numerous ne$ genera, but his scheme, strongly critici5ed by ten'oe (1E=>) has not been accepted $idely. Pillai (1E!) eleated *pirorbinae to the family*pirorbidae, but later a number of authors suggested that *pirorbidae are more closely related

to *erpulinae than to -ilograninae (ten 'oe 1E=>, -it5hugh 1E=E, *mith 1EE1, ouse \-it5hugh 1EE>) and that the maintenance of the family *pirorbidae is not "ustified. ecentphylogenetic analyses confirmed the position of *pirorbinae as a subfamily of *erpulidae(upriyanoa ?!!#, upriyanoa et al. ?!!D, 7ehrke et al. ?!!, upriyanoa \ ouse?!!=). Ten 'oe (1E=>) regarded the -ilograninae as paraphyletic and a morphologybasedcladistic analysis of some *erpulidae (upriyanoa ?!!#) supported his conclusions.+oreoer, the most recent phylogenetic analyses of *erpulidae using 1=* ribosomal %A(7ehrke et a l. ?!!) and another using combined molecular and morphological data(upriyanoa et al. ?!!D) suggested that both traditionally formulated subfamilies *erpulinaeand -ilograninae are not monophyletic. upriyanoa et al. (?!!D) refrained from reising theserpulid classification and suggested that a ma"or reision of serpulid taFonomy is needed

based on more genera than used in their study. The ma"or obstacle to a comprehensiephylogenetic analysis of the *erpulidae remains the state of its alpha taFonomy. 6t is almostproerbial to say that serpulid taFonomy is ery confused and most currently recogni5edserpulid genera hae long and conoluted taFonomic histories. Oithin *erpulidae, specificidentification has traditionally been based on a combination of characters such as morphologyof the operculum and opercular peduncle (if present), degree of deelopment of the collar andthoracic membranes, structure of collar chaetae and tube and, to a lesser degree, structure ofchaetae and uncini. *erpulid genera hae been described on the basis of uniRue characters or
mailto:[email protected]:[email protected]:[email protected]


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on uniRue combinations of characters (een on absence of characters) rather than on presenceof shared deried characters. Although traditionally only fe$ characters hae been used inserpulid taFonomy, ariability of these characters remains largely unstudied. There hae beenery fe$ reie$s of serpulid taFonomy. The ery first reision (+Irch 1=D#) $as follo$ed by

early reie$s by *aintoseph (1=E>), ush (1E!), and PiFell (1E1?, 1E1#). 8hamberlin(1E1E) gae a key to the serpulid genera $ithout attempting to reise the family, and so did*outh$ard (1ED#), half a century later. -auchald (1E) compiled a list of generic diagnosesand a key to genera for all polychaetes, including serpulids and spirorbids. 6n addition to the*pirorbidae, he ackno$ledged ##1 species of serpulids, diided into # subfamilies0 the*erpulinae $ith >> genera, the -ilograninae $ith genera, and the -icopomatinae $ith genera. &f these > genera, ?? $ere monotypic and another 1# had only ? species. @chida(1E=) proided a systematic reie$ of the group $ith a description of ne$ species and ne$genera, but gae no key. 'e mentioned only ?## species, as compared to the ##1 of -auchald(1E). &f the D1 genera distinguished by @chida (1E=), ?D $ere monotypic, and 1 had onlyt$o species. o attempts to reie$ *erpulidae hae been made eer since and no$, thirty

years later, -auchald (1E) still remains the most commonly used source of information onthe generic composition of serpulids. %uring the last three decades serpulid taFonomyunder$ent significant changes, $ith numerous taFa being synonymi5ed, older diagnosesemended and eFtended, ne$ species described and about 1! genera added.

Material and methods

The aim of this reie$ is to proide uptodate information on the current state of taFonomyof *erpulidae sensu lato. Although the position of *pirorbinae $ithin *erpulidae has beendetermined (upriyanoa ?!!#, upriyanoa et al. ?!!D, 7ehrke et al. ?!!), spirorbins arenot included in the present paper because composition and phylogenetic relationships $ithinthis monophyletic group recently hae been treated else$here (+acdonald ?!!#). Themorphology of serpulids (and ariability of morphological characters) is reie$ed $ithrespect to features that can be used as characters in forthcoming cladistic analyses. Zotobsered[ in the diagnoses belo$ indicates that no data hae been gien in the literature andmaterial either could not be (re) eFamined by us, or $as not presered $ell enough. *ince amere literature compilation $ould not be sufficient $hen dealing $ith a group $ith such acompleF taFonomic history, $e eFamined $ith use of light microscopy representaties(mostly preiously unpublished material) of all genera currently considered alid in*erpulidae sensu lato. 6t should be noted that some of the characters are sub"ect tointerpretation, changing gradually rather than in distinct steps. +oreoer, $hile structure ofchaetae and uncini do proide important characters for serpulid taFonomy, many eFistingdescriptions, especially the early ones, $ere published $ith ery sketchy linedra$ings ofchaetal structures made under a compound light microscope. These illustrations often do notproide adeRuate details of chaetal ultrastructure, and een can gie a $rong impression $hencompared $ith images done $ith scanning electron microscopy (*;+) (ten 'oe \ ansenacobs 1E=>: 1>0 compare for instance -auel 1E? fig. 1?1R $ith reton \


18/23

$ere that incomplete that full redescriptions of their typespecies hae been included.Authors3 names and year of publication for alid serpulid taFa can be found in the Table of8ontents and in the releant sections and lists of species. -or the remaining taFa, thisinformation is gien $ith their first occurrence in the teFt.


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Mor!hology

+he tube

Ohereas tubes of the closely related sabellid family are constructed of mucus and muddy orsandy sediments (e.g., onar 1E?0 $ith the eFception of the calcareous tube in the sabellid

Glomerulaielsen, 1E#1 (including Calcisabella Perkins, 1EE1), e.g.,


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radioles each bearing a double ro$ of ciliated pinnules. &ne of the radioles is usuallytransformed into the opercular peduncle (-ig. 8, %, pd) and distally bears the operculum(-ig. G8, op). The base of the branchial cro$n is surrounded by the collar, $hich continuesas the thoracic membranes, a structure found only in serpulids. The border bet$een thoraF and

abdomen is marked by chaetal inersion, $ith the dorsal notochaetae and entral combshapedneurochaetae (uncini) of the thoraF changing places such that abdominal uncini become dorsal(notopodial) and abdominal chaetae become entral (neuropodial) in the abdomen. Althoughserpulids are often ery brightly coloured and the colour of the animals indeed may be usefulin the field (-ig. 1), as a taFonomic character the type of colouration is of little use as colour israpidly lost in preseraties, particularly in alcohol. The colouration may also be a sub"ect tosignificant interspecific ariability (e.g., in the Spirobranchus corniculatus compleF: -oss` \ilsen ?!!!: 1>!, 1>0 *ong ?!!D, as S. giganteus). 6t may een ary $ithin a singlespecimen, for instance, in Spirobranchus the colour of both lobesof a branchial cro$n may beso different as blue and red (ten 'oe unpubl.). -]yn \ 4"]en (1E#) describe a +endelianpattern found in the colouration of branchial cro$ns of Pomatoceros tri&ueter, $ith ?D?=

bro$n,?1= blue, and only ? orange branchial cro$ns.+he bran#hial #ro'n

The branchial cro$n, used for feeding and respiration, $ith each radiole bearing ro$s ofpaired ciliated pinnules is a distinct feature of sabellids and serpulids. The cro$n is consideredto be prostomial (cf. *egroe 1E>1). The radioles of the branchial cro$n are attached to pairedlobes (-ig. D-, bl) located laterally on both sides of the mouth. The branchial lobes arecompletely separate from one another in serpulids, but are fused together in some sabellids(e.g., Chone ush, 1E! and Sabella 7innaeus, 1D, see -it5hugh (1E=E)). The number ofradioles used to delineate some taFa (e.g., Salmacina tribranchiata) is an unreliable character.6n indiidual species, the lo$er limit of the number of radioles has no alue, since all"ueniles hae fe$er radioles than adults. The upper limit is possibly an eFponent of si5e,possibly genetically determined.'o$eer, the ariation $ithin indiiduals of larger species (e.g., Spirobranchus andProtula)is enormous. upriyanoa (1EEE) sho$ed that $ithin some Serpula species number ofradioles (as $ell as number of opercular radii) is directly correlated $ith animal si5e. Thebases of the radioles in some sabellids and serpulids may be "oined $ith an interradiolarmembrane (-ig. #A, ;, mb). 6n serpulids, the interradiolar membrane is ery high inPomatoleios and it unites radioles for up to half of their length in Pyrgopolon. The membraneis also $ell deeloped in Spirobranchus (and may bear processes in some Spirobranchusspp.), Pomatoceros, Pomatostegus, Galeolaria, asynema, and Neovermilia. 6t is alsocommonly found in species of Serpula, Spiraserpula, and Crucigera, but is ery rare inHydroides (see astidaQaala \ ten 'oe ?!!?). The membrane is absent in the serpulidgenera Ficopomatus, Filograna, Pseudochitinopoma, Pseudovermilia, Salmacina, andermiliopsis.

"yes. Photoreceptors may be found not only in the anterior region but almost any$here inannelids, including *abellida, from ephemeral eyespots on epitokous segments (notably in;unicidae) to those on pygidia (e.g., Fabricia lainille, 1=?=0 Augeneriella anse, 1E(both in -it5hugh 1E=E)). The serpulids are no eFception to this plasticity, as e.g., sho$n bythe girdle of thoracic (peristomial) redpigmented ocelli in Placostegus (-ig. 1-, te). &ne


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difficulty is that the eyespots may disappear in preseratie in a comparatiely short time. 6nthe diagnoses gien belo$, Zpresence[ or Zabsence[ has been obsered in fresh material0 Znotobsered[ indicates that no data hae been gien in the literature and material could not be(re)eFamined by us. Another difficulty is that there has been no consistent terminology, Zeye[

or Zeyespot[ can hae any of the meanings gien belo$.

Prostomial o#ellar #lusters. +any serpulids possess a pair of brainassociated clusters ofocelli in the prostomium, apparently the continuation of the laral Zeyes[ (*mith 1E=>a, b) .-or instance, more than ?! presered specimens of Filograna imple(a sho$ed ? ro$s of >GDpigmented cells in the prostomial area, presumed to be prostomial Zeyespots[, ho$eer$ithout lenses0 on the other hand, more than ?! nonoperculate specimens of Salmacina spec.from +arseille lacked pigmented spots in the prostomial area (ten 'oe \ Pantus 1E=).)etavermilia multicristata has prostomial Zeyes[ (Qibro$ius 1ED=a: =D, 1?=, as ermiliopsis),presumably simple ocelli. &n the other hand, in fresh ). multicristata specimens from the*eychelles Zeyespots[ $ere inisible (ten 'oe unpublished). %ranchial eyes. 6n serpulids,

most photoreceptors are associated $ith the branchial cro$n (including the operculum), andthese could be termed collectiely Zbranchial eyes[. Apart from ultrastructural differences,and although intermediate types do occur, photoreceptors may roughly be grouped into threegroups for $hich $e propose the follo$ing Zstandard[ terms: celli/ single eyespots $ith (or$ithout a single lens). These may occur on the aFis of radioles (e.g., ermiliopsis spp., sometaFa of the Spirobranchus tetraceroscompleF), but also near the interradiolar membrane(-ig. D%), on the peduncle or the operculum (e.g., a hundred or more ocelli, not rigidlypatterned, on the entral rim of the opercula of Pomatostegus stellatus and Spirobranchuscorrugatus). cellar clusters/ loose, bulging groupings of approFimately ?G?! ocelli,generally $ith as many lenses. &ccurrence on arious radioles (notably Apomatus spp.), orpeduncle (e.g., Semivermilia pomatostegoides, on border bet$een peduncle and opercularampulla). asynema chrysogyrus has GD pairs of ocellar clusters, $ith ?G11 lenses each(6ma"ima \ ten 'oe 1E=>). @chida (1E=) reports)icroprotula ovicellata as haing =G11pairs of Zeyespots[ (ocellar clusters) on each radiole, and a red eye (probably a simpleocellus) in the base of each branchial tuft.Compound eyes/ more or less rigidly patterned groupings of ocelli (-ig. DA). 6n theSpirobranchus giganteus0compleF sensu lato, for instance, there are D!!G1!!! lenses in acompound eye, located at the base of each branchial lobe (-ig. D;), proFimally on the first leftand right radiole. These might ery $ell be capable of receiing isual impressions in asimilar $ay as in crustaceans (*mith 1E=>a). The radii of the opercular funnel in Hydroideslambec1i and of the operculum ofPyrgopolon ctenactis sho$ circulargroupings of ?!G#! redpigmented bulging structures (-ig. D), $hich certainly are ery reminiscent of smallcompound eyes. The knobs at the base of the radioles in Protula balboensis (illustrated by+onro 1E## fig.

+he o!er#ular !edun#le

6n some serpulid taFa, the branchial radiole that bears the operculum is identical to the otherradioles (Filograna, Apomatus, #osephella (-ig. #0 ho$eer, not in#osephella populationsfrom the ;. +editerranean (en;liahu \ Payiatas 1EEE)) and in Protis). 6n most serpulids theoperculum is borne on a distinct peduncle (-ig. #A, 8, %, ;, -). The peduncle may graduallymerge into the basal fleshy part of the operculum (-igs #;, 8), or be separated from it by a


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more or less clear constriction (-ig. #-). +orphologically, an opercular peduncle is usuallyinserted more or less belo$ and bet$een the first and second normal radiole, outside the lineof radioles (-ig. #-). 6n some genera, the peduncle is located at the base of branchial cro$n,coering seeral radioles (Pomatoceros, Spirobranchus, and Galeolaria). 6n other taFa, such

as Semivermilia, )etavermilia and%athyvermilia, the peduncle is clearly positioned as thesecond modified radiole (ten 'oe 1E). 6n small specimens/species the insertion ofpeduncle may be ery difficult to obsere. 6n Semivermilia pomatostegoides the peduncle iseither the second radiole, or inserted belo$ and bet$een first and second.

A#kno'ledgements

This $ork $as supported by the Australian esearch 8ouncil grant %P!=#D. Thanks aredue to ;lly eglinger and %r. %irk Platoet (both Q+A) and the staff of the Adelaide+icroscopy for their help $ith *;+. &er the years $e hae studied material from all themuseums mentioned in the introduction, $e are ery grateful to all staff inoled in the timeconsuming loans, registrations of ne$ material, or help in the collections. This study $ould

not hae been possible $ithout the help of many colleagues $ho oer the years sharedmaterial, literature, and unpublished obserations. Oe both feel priileged to hae beenallo$ed to share ideas and forces in the past and present $ith so many colleagues in andoutside "oint publications, of $hich some can be found in the eferences.

eferen#es

Abildgaard, P.8. (1=E) eschreibung 1. einer grossen *eeblase (Holothuria Priapus 7inn.) ?.5$een Arten des *teinbohrers ($erebella 7inn.) #. einer grossen *andrIhre (Sabella 7inn.).Schri!ter der Gesellscha!t natur!orschenderFreunde 2u %erlin, E, 1##G1>D.

Absolon, . \ 'rab, *. (1E#!) Nber einen neuen *Jss$asserGPolychaeten aus den'Ihlenge$ssern der 'er5ego$ina.3oologischer An2eiger, *eip2ig, ==, E/1!, ?>EG?D>.

Allen, 4.. \ *teene, . (1EE> (reprinted 1EED))'ndo0Paci!ic coral ree! !ield guide. Tropicaleef esearch, *ingapore, #= pp.

Alari^o, A. (1E1) 6ncrustaciones marinas.%oletin del 'nstituto Espa4ol de ceanogra!ia >,1G1?.

AmoureuF, 7. (1E#) uelRues annXlides polychtes de lAfriRue occidentale et XRuatoriale.Cahiers .-.S.$..).,

c5anographie, 11, 1, >1GD.

AmoureuF, 7. (1ED) Serpula 6Paraserpula7 israelitica, nouelle espce de *erpulidae(AnnXlides Polychtes) et une petite collection annXlidienne de la +editerranXe orientale.%ulletin de )us5um National d8Histoire Naturelle, Paris, (#) >!> (Qoologie ?=1), 1!>1!E.

AmoureuF, 7. (1E=) AnnXlides polychtes recoltes par . *tirn en 1EDE, sur les cStesmarocaines du dXtroit de 4ibraltar. Cuademos de Ciencias %iol9gicas, , G##.


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AmoureuF, 7., ullier, -. \ -ishelson, 7. (1E=) *ystXmatiRue et Xcologie dannXlidespolychtes de la presRuil du *inai.'srael #ournal o! 3oology, ?, G1D#.

Augener, '. (1E!D) eports on the results of dredging, under the superision of AleFander

Agassi5, in the 4ulf of +eFico and the 8aribbean *ea, and on the east coast of the @.*. 1=G1==!, by the @.*.*. 8oast *urey *teamerlake. 766, Oestindische Polychaeten.%ulletin o! the )useum o! Comparative 3oology atHarvard College, >#,E1G1ED.

Augener, '. (1E1>) Polychaeta, 66, *edentaria.'n/ +ichaelsen, O. \ 'artmeyer, . (;ds).ie Fauna S:d;estenntnis der )eeres!auna ?esta!ri1as, ol. @. -riedrichsen, 'amburg, pp. DEGD?.

eunice siciliences

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