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Tropical Zoology
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Interstitial fauna of th e
Galapagos: Duplacrorhynchinae,
M acrorhynchinae, Polycystidinae,
Gyratricinae (Platyhelm inthes
Polycystididae)
T. J. Artois a & E. R. S ch o ck a erta
a Research Group Zoology, D epartm ent of Chemistry, Biology and
Geology, Limburgs Universitair Centrum , B-3590, Diepenbeek,
Belgium E-mail:
Version of record first published: 30 Jul 2012.
To cite this article: T. J. Artois & E. R. Schockaert (2001): Interstitial fauna of th e Galapagos:
Duplacrorhynchinae, M acrorhynchinae, Polycystidinae, G yratricinae (Platyhelm inthes
Polycystididae), Tropical Zoology, 14:1, 63-85
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Tropical Zoology 14 : 63 - 85 , 2001
In terstitia l fa u n a o f th e G a la p a g o s:
D u p la cro rh y n ch in a e, M acrorh yn ch in ae,
P o ly c y stid in a e , G yratricinae
(P la ty h elm in th es P o ly cy stid id a e)
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T.J.
A r t o is a n d
E.R.
S
chockaert
Research G roup Zoology, D epartm ent o f Chemistry, Biology and. Geology, L im burgs
Universitair Centrum, B-3590 Diepenbeek, B elgium (E-m ail: tom .artois@ luc.ac.be)
Received 17 January 2000, accepted 30 August 2000
Seven species from th e G alapagos of d ifferen t su b fam ilies of th e Polycysti­
d id a e a re d e sc rib e d a n d d isc u ssed . F o u r o f th e m a re new to science, th re e of
w h ich belong to th e su b fam ily D u p lac ro rh y n c h in a e S c h o c k a e rt & K arling 1970:
Djeziraia incana n. sp. d iffers fro m th e o th e r sp ecies o f th e g e n u s b y its long,
th in stylet; P honorhynchoides haegheni n. sp. is u n iq u e w ith in th e genus, as th e
p ro s ta te sty let is m u c h s h o rte r th a n th e a ccesso ry stylet; P arachrorhynchus jo n ­
delii n. sp. is c h a ra c te rise d b y th e p re se n c e of a b u n d le of a tr ia l g lan d s, p a ire d
sem in al vesicles, a very larg e s p h in c te r a t th e d ista l en d o f th e fem ale d u ct, an d
a rin g of h a rd p la te s a t th e d is ta l e n d o f e a ch ovary. O f th e M a c ro rh y n c h in a e
E v d o n in 1977, M acrorhynchus m anusferrea n. sp. d iffers fro m th e o th e r species
of th e g en u s b y th e sh a p e o f th e stylet a n d th e p re se n c e o f a fem ale b u rsa . The
subfam ily P o lycystidinae S c h o ck aert & K arling 1970 is re p re se n te d by Polycystis
ali S c h o c k a e rt 1982 a n d Paulodora felis (M arcu s 1954). T h e G alap ago s p o p u la ­
tio n of th e la tte r species is co n sid ered a se p a ra te subspecies, P. felis asym m etrica
n. subsp., b a se d o n differences in th e fem ale system . Finally, Gyratrix herm aphro­
ditus E h ren b erg 1831 is discussed. T h ree co lo u r v a ria n ts o f th is species com plex
a re fo u n d in th e G alapagos, w h ich re p re se n t th re e m o re species in th e com plex.
F o r all new taxa, a d iag n o sis is given. At th e end, w e pro v id e a species list of th e
p olycystidid fau n a o f th e G alapagos Isles.
key w o rd s:
D u p lacro rh y n ch in ae, M a cro rh y n ch in ae, Polycystidinae, G y ratricinae,
Polycystididae, G alapagos, new taxa.
I n t r o d u c t i o n ..................................................................................................................
M aterial and m ethods
............................................................................................
Abbreviations
Results
.........................................................................................................................
Subfamily D uplacrorhynchinae Schockaert & Karling 1970
. . . .
Djeziraia incana n. sp.............................................................................................
Phonorhynchoides haegheni n. sp..........................................................................
Parachrorhynchus jondelii n. sp.
..................................................................
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T.J. A rtois a n d E.R. S chockaert
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Subfamily M acrorhynchinae Evdonin 1977
Macrorhynchus manusferrea n. sp.......................................................................
Subfamily Polycystidinae Schockaert &Karling 1970
Polycystis ali Schockaert 1982
Paulodora felis (M arcus 1954) Artois & Schockaert 1998 asymmetrica n.
subsp....................................................................................................................
Subfamily Gyratricinae Graff 1905
..............................................................
Gyratrix hermaphroditus Ehrenberg 1831 species complex
. . .
Diagnoses of the new taxa
.....................................................................................
Polycystididae of the Galapagos: species l i s t .........................................................
Acknowledgements
....................................................................................................
References
..................................................................................................................
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INTRODUCTION
F o rm e r c o n trib u tio n s o n th e p o ly cy stid id fa u n a fro m th e G alapagos d e a lt
w ith th e su b fa m ilie s P o rro c y stid in a e E v d o n in 1 9 7 7 ( A r t o i s & S c h o c k a e r t 1 9 9 9 )
a n d T yphlopolycystidinae E v d o n in 1 9 7 7 ( A r t o i s & S c h o c k a e r t 2 0 0 0 ) . In th is th ird
c o n trib u tio n , m e m b e rs o f th e re m a in in g su b fam ilie s are d e scrib ed a n d d iscussed,
m ak in g th e survey of th e p o ly cy stid id fa u n a o f th e a rc h ip e la g o com p lete. The
re m a in in g p a rt of th e E u k a ly p to rh y n c h ia (six re p re se n ta tiv es o f th e K o in o cy stid i­
d ae a n d one o f th e G n ath o rh y n ch id ae) w ill be d iscussed in a la ter paper, w h ich will
c o n clu d e th e series o n th e k a ly p to rh y n ch fa u n a o f th e G alapagos. The
S ch izo rh y n ch ia w ere tre a te d by N o l d t & H o x h o l d ( 1 9 8 4 ) .
T his c o n trib u tio n is p a rt of a series o f p ap e rs o n th e m eio fa u n a o f th e G ala­
pagos, a survey o f w h ich can be fo u n d in M i e l k e ( 1 9 9 7 ) .
MATERIAL AND M ETHODS
T he m a te ria l fro m th e G alap ag o s w as co llec ted b y S. H o xhold a n d P. S c h m id t, w ho
d rew a n d p h o to g ra p h e d th e live a n im a ls (fo r d e ta ils see Ax & S c h m i d t 1973). S. H o x hold
m ad e th e m ea su re m e n ts o n live a n im als. W h en su fficien t an im als w ere available, so m e w ere
fixed w ith B o u in ’s fixative, e m b e d d e d in p a ra ffin a n d serially se ctio n ed (4 p m sectio n s). The
sections w ere sta in e d w ith H eid en h a in ’s iro n h em ato x y lin w ith eo sin as c o u n tersta in .
A d d itio n al m a te ria l of so m e sp ecies w as co llected in o th e r lo c a litie s b y th e a u th o rs.
S om e of th e sp ecim en s w ere w hole m o u n te d using lac to p h en o l as fixative. S pecim ens in te n d ­
ed for sectioning w ere tre a te d as d escrib ed above.
B rief d e sc rip tio n s o f th e lo calities in th e G alap ag o s a re follow ed b y a n u m b er. T his
n u m b e r refers to th e d etailed d escrip tio n s of th e lo calities b y Ax & S c h m id t (1973: 21-26).
M ea su re m e n ts o f h a rd p a rts a re axial. F ig u re s w ith o u t a scale a re free h an d . P h o ­
to g rap h s w ith o u t scale a re from live m aterial. The p o sitio n of th e g o n o p o re is expressed as a
p ercen tag e of b o d y len g th (d istan ce from th e a n te rio r tip of th e body).
T he species e p ith e t of M a crorhynchus m anusferrea n. sp. is c o p ie d fro m I. H e n z e ’s
D ip lo m arb eit (U niversity o f G ö ttin g en 1987).
Type m a te ria l w ill b e d e p o site d in th e Z oological M u seu m of th e U niversity of G ö ttin ­
gen (G erm any), except fo r th a t o f P honorhynchoides haegheni, w h ich w ill b e d ep o sited in th e
c o llectio n of th e R e se a rc h G ro u p Z oology o f th e L im b u rg s U n iv e rsitair C e n tru m (LUC),
D iepenbeek, B elgium .
In te rstitia l fa u n a of th e G alapagos
65
ABBREVIATIONS
acg: accessory glands; ast: a ccesso ry stylet; b : b u rsa ; b s : b u rs a l stalk; d b : d is ta l p a r t of
b u rsa; de: ejacu lato ry d uct; ds\ in se m in a tio n duct; e\ eyes; fd: fem ale d uct; ga: c o m m o n g en ital
atriu m ; gd: g enital duct; gg: glands; gp: gonopore; m : m o u th ; ma: m ale a triu m ; od: oviduct; ov:
ovarium ; p: p roboscis; pb: p ro x im a l p a r t of b u rsa; pc: p re p h a ry n g e a l cavity; ph: p h ary n x ; sd:
sem in al duct; sph: sp h in c ter; sr: se m in a l recep tacle; sí: stylet; t: testis; ut: u te ru s; va: vagina;
vd: vitelloduct; vg: p ro s ta te vesicle; vi: vitellariu m ; vs: se m in al vesicle; v, w, x, y, i;. see in text.
RESULTS
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Subfam ily D u p lacro rh y n ch in ae S cho ck aert & K arling 1970
D je zira ia in c a n a n. sp. (Figs 1, 4A-B)
Localities 1. S a n ta Cruz: B ah ia A cadem y (IX. 6b). S a n ta Cruz: B ahia B o rre ro
(IX. 3). S an ta Cruz: B ah ia T ortuga (IX. 7b). B a rrin g to n (XI. 2).
Material. D raw ings a n d p h o to g ra p h s of live a n im a ls b y S. H oxhold. Several serially sec­
tio n e d specim ens, o n e sagittally sectio n ed sp ecim en d e sig n ated as holotype.
D erivation o f the nam e. T he species n a m e refers to th e grey co lo u r of th e liv­
ing an im al. In c a n u s (Lat.): to tally grey.
D escription. A nim als a re 1-1.2 m m in length, w ith tw o eyes. They a p p e a r grey
in tra n s m itte d light. T he g en eral o rg a n is a tio n c o rre sp o n d s w ith th a t of D. pardii
S ch o ck aert 1971 in m o st resp ects, su c h as in th e d im en sio n o f th e pro b o scis (± 7%
o f th e body len g th ) w ith th re e p ro b o scis re tra c to rs a n d tw o in te g u m en t retra c to rs,
in th e p o sitio n o f th e p a ire d g o nads, in th e g en ital a triu m bein g c o n n ec ted to th e
su b te rm in a l gen ital p o re (at ± 90%) by a long genital canal, etc. Fem ale duct, m ale
d u c t a n d u te ru s e n te r th e genital a triu m sep arately a t th e a n te rio r side, the fem ale
d u c t being m o st d orsal.
The sle n d e r a n d slightly curved single-w alled stylet is 162-198 p m long (m ean
= 179, n = 3) even u p to 295 p m in one specim en. The opening o f the stylet is te r­
m in a l. T he se m in a l vesicle, a p p a re n tly w ith o u t m uscle sh e a th , is relatively large,
w ith a very th in e p ith eliu m w ith fla tte n e d nuclei. The in terp o sed p ro sta te vesicle is
sm all, w eakly m uscular, a n d c o n ta in s m ain ly a c o arse-g rain ed baso p h ilic secretio n
th a t is so m etim es su p p le m e n te d w ith very sm all eo sin o p h ilic glands. The nuclei of
th e p ro sta te g lands a re b o th in tra- a n d extracapsular.
The long, n a rro w b u rsa l stalk ru n s anteriorly, th e n bends back w ard to en d in
th e te rm in a l b u rsa . Its e p ith e liu m is r a th e r low a n d a n u c le a te d , m o stly ru ffled
("pseu d o ciliatio n "), a n d it is s u rro u n d e d by lo n g itu d in al m uscles a ro u n d th e distal
p a rt, a n d stro n g c irc u la r m u scles a ro u n d th e p roxim al half. The sh o rt fem ale d u c t
e n te rs th e sta lk a t a b o u t its m id d le. T his fem ale d u c t (w ith o u t an y m u scle coat)
w id en s to a larg e sem in al re c e p ta cle in to w h ich th e o v iducts enter. T he in se m in a ­
tio n d u c t lin k s th e se m in a l re c e p ta cle w ith th e p ro x im al en d of th e b u rsa l stalk.
1 The type locality is S anta Cruz. The exact location rem ains uncertain. The holotype is
labelled 31 a/b, 3 1 a was collected at the first locality (IX. 6b), 31 b at the second (IX. 3).
T.J. A rtois a n d E.R. S chockaert
66
A
ov
,vs
m
-v i
vs
.ov
ut
ov
ds
gP
Fig. 1. — Djeziraia incana n. sp. A, general organisation; B, atrial organs; C, reconstruction of the
female atrial organs from the left h and side (A-B from live animals).
L o n g itu d in al m u scles s u rro u n d th is d u ct, w hile a t b o th en d s th ere is also a stro n g
c irc u la r m u scle layer. T he in s e m in a tio n d u c t is lin ed w ith a very low ep ith eliu m ,
w h ich is m ostly re d u c e d to th e b a se m e n t m em b ran e, form ing a so-called pseudocuticula, especially n e a r th e sem in al receptacle.
D iscu ssio n . Djeziraia incana is th e th ird species d escrib ed in th is very h o m o ­
geneo u s genus. D. euxinica (M ack-Fira 1 9 7 1 ) S ch o ck aert 1 9 8 2 o ccu rs in th e B lack
Sea ( M a c k - F i r a 1 9 7 1 ) a n d in th e M e d ite rra n e a n (S a rd in ia, T.J. A r t o i s pers. obs.),
w h ile D. pardii S c h o c k a e rt 1971 is fro m th e E a st A frican co asts. The new species
ca n be in itially d istin g u ish e d by its lo n g ( 1 6 2 - 1 9 8 p m a n d u p to 2 9 5 p m in one
specim en) a n d very slen d er stylet, w ith a term in al opening. The stylet o f D. eu xin i­
ca also h a s a te rm in a l opening, b u t is s h o rte r [ 1 0 5 p m in th e M e d iterra n ean speci­
m e n (T.J. A r t o i s pers. obs.), 1 2 0 p m a c c o rd in g to M a c k - F ir a ( 1 9 7 1 ) ] . T he stylet of
D. pardii is 8 4 p m long w ith a su b te rm in a l opening.
The very thin-w alled fem ale d uct, com ing from th e sem inal receptacle, e n ters
th e b u rsa l stalk a t a b o u t its m id d le in D. euxinica a n d in D. incana; it is very sh o rt
in th e la tte r species. In D. pardii it e n te rs th e g en ital a triu m . In all th re e species,
th e in se m in a tio n d u c t co n n ects to th e sem in al receptacle. T his w as in d o u b t in the
orig in al d e sc rip tio n o f D. pardii. C areful o b serv atio n of th e type m aterial, however,
revealed th a t a c o n n e c tio n o f th e "d iv erticulum " ( S c h o c k a e r t 1 9 7 1 ) a n d th e sem i­
nal recep tacle does exist. A strik in g fe a tu re only fo u n d in D. pardii, is the presence
o f tw o h o rn s a t th e c o n n e c tio n . T he sp e rm d u c t o f D. pardii is highly m uscular,
being su rro u n d e d by a very th ick c irc u la r m uscle sh eath , w hile in D. incana circ u ­
la r m uscles a re only p re se n t a t th e tw o ends of th e d uct. The situ a tio n in D. eu xin i­
ca is n o t yet k n ow n 2.
2 Data on the internal organisation of D. euxinica (not given in the original description) are
from an unpublished reco n stru ction by M a c k - F ir a and K a r l in g , kept at th e Sw edish M useum of
N atural History in Stockholm.
In te rstitia l fa u n a of th e G alapagos
67
P h o n o rh yn ch o id es haegh en i n. sp. (Figs 2-3, 4C-D)
Localities in the Galapagos. S a n ta Cruz: B ah ia B o rre ro (IX. 3). S a n ta Cruz:
B ah ia Academ y: n o rth e rn side (IX. 5c a n d IX. 5d). B artholom é: n o rth e rn side (VII.
1). Tower: B ahia D arw in (V. la ).
O ther localities. S. H u tc h in so n Isla n d (F lo rid a, USA), a t a brid g e n e a r th e
park; m ed iu m fine sa n d ta k e n a t ca 0.5 m deep a t high tide, 16.XI.1994 (type locali­
ty). S. H u tc h in so n Isla n d (F lo rid a, USA), Je n se n B each, a little b it n o rth o f "B ath
Tub" a t th e ICH -side; m ed iu m fine sa n d w ith som e d e tritu s from ca 30 cm deep, a t
th e foot of som e rocks, n e a r a m angrove area, 24.IX.1994.
Material. Several a n im als stu d ie d alive. T h ree w hole m o u n ts (F lorida), o n e of th e m d es­
ig n a te d h o lo ty p e (LUC no. 213), a n o th e r o n e p a ra ty p e (LUC no. 214). T h re e a n im a ls from
F lo rid a serially sectioned, 22 sectio n ed sp ecim en s from th e G alapagos.
D erivation o f the nam e. This species is d ed icated to David Van D er H aeghen, a
frien d o f T.J. Artois.
vs
ae g
vi
ast
ov
50 |tm
B-C
Fig. 2. — Phonorhynchoides haegheni n. sp. A, atrial organs; B, accessory stylet; C, stylet (A from a
specimen from the Galapagos, B-C from the holotype).
T.J. A rtois a n d E.R. S chockaert
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68
D escription. Live an im als a re long a n d slender, 1-1.4 m m , w ith p aire d eyes and
a d ra b grey colour. T he e x creto ry system is w ell developed. The e p id e rm is is 3 pm
h ig h w ith cilia 2 p m long, a n d w ith fla tte n ed , lo b ate nuclei, n u m e ro u s opticallyem pty vacuoles a n d elo n g ated rh a b d ite s o f length a b o u t 1/3 to 1/2 of th e e p ith eliu m
h eig h t. In so m e sp ecim en s th e vacu o les a re filled w ith a c o a rse-g rain e d b a so p h ilic
secretion. C audal glan d s are well developed.
The p ro b o scis is very sm all, ± 5% o f th e body length. The apex is b arely visi­
ble in th e live an im al. The b asal an d apical ep ith elia of the eone are relatively high.
The ex tern al m u sc u la tu re o f th e p ro b o scis co n sists of eight p ro tra c to rs w ith seven
to eig h t m u scle fibres each, six fix ato rs a n d a ven tral a n d do rsal p a ir of re tra cto rs.
The p resen ce o f th e u su a l th ird , d o rso la te ral p a ir of re tra c to rs could n o t be esta b ­
lish ed w ith c e rta in ty o n su c h a sm all p ro b o scis. All re tra c to rs in se rt o n th e body
w all in th e p h a ry n g e a l reg io n . A dditionally, a d o rsa l a n d a v e n tra l p a ir o f in te g u ­
m e n t re tra c to rs are presen t.
The p h ary n x is s itu a te d in th e first body h a lf a n d is ca 70 p m in diam eter. It
h a s th e sam e s tru c tu re as in P honorhynchoides som aliensis S ch o ck ae rt 1971 (see
S c h o c k a e r t 1 9 7 1 ).
The g en eral o rg a n is a tio n o f th e g en ital system is th e sam e as in P.
so m a lien sis. T he g en ital p o re is s itu a te d ventrally, su b term in ally , a t ca 95%. The
co m m o n genital a triu m h a s a ruffled, d eg en eratin g ep ith eliu m (pseudociliation); in
so m e a re a s th e e p ith e liu m h a s d isa p p e a re d , leaving only a th ic k b a se m e n t m em ­
b ra n e (p se u d o c u tic u la ). The g en ital a triu m is s u rro u n d e d by lo n g itu d in a l m uscles
only. The lo n g itu d in al sclero tized ridges o bserved in P. som aliensis are lacking.
The sem in al vesicle is sp in d le-sh ap ed . It h a s a th in ep ith eliu m w ith som e flat­
te n e d n u c le i a n d a very th in , sp ira l m u scle sh e a th . T he sem in al d u c t ru n s axially
th ro u g h th e ovoid p ro sta te vesicle, w h ich is su rro u n d e d by a n a lm o st c ircu lar m u s­
cle c o a t a n d c o n ta in s a lig h t e o sin o p h ilic se cre tio n . N uclei o f th e p ro sta te glan d s
are b o th in tra- a n d ex tracap su lar. T he p ro sta te stylet co n d u cts sp erm a n d p ro sta te
se c re tio n , a n d is a s tra ig h t single-w alled tu b e, 87-98 p m long (m ea n = 91, n = 3),
ta p e rin g from 12 pm to 5 pm in th e specim ens from F lorida. The an im als from the
G alapagos have stylets 47-58 p m long (n = 2). The stylet lies in a n a rro w canal sur-
VI
acg
vs
ov
sr d s
a st
Fig. 3. — Phonorhynchoides haegheni n. sp. R econstruction of the atrial system from the left hand
side.
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In te rstitia l fa u n a of th e G alapagos
69
ro u n d e d by a lo n g itu d in a l m u scle s h e a th only (a t least, no ep ith e liu m c o u ld be
seen). This m uscle s h e a th is c o n tin u o u s w ith the m uscles of the genital atriu m .
The accesso ry se c re tio n reservoir, to w h ic h th e accesso ry stylet is con n ected , is
m u c h larg er th a n th e p ro sta te vesicle. It is elongated, ovoid, su rro u n d e d by a spirallyru n n in g , a lm o st c irc u la r m u scle c o a t a n d filled w ith a b a so p h ilic se cretio n . The
(five?) g lands th a t p ro d u c e th is se c re tio n ea c h have a very large e x tra c a p su lar n u cle­
ate d p a rt. T ow ards th e accessory stylet th e reservoir n arro w s to form a b ro a d duct. In
sectio n ed m a te ria l, th e accesso ry reserv o ir is invariably o rie n ta te d w ith its d istal end
in th e cephalic d irectio n , w ith th e d u c t tu rn in g over 180° to w ard s th e accessory stylet.
The accesso ry sty let is slender, slig h tly b en t, a n d en d s in a s h a rp p o in t. In th e sp eci­
m e n s fro m F lo rid a it is 264-325 p m lo n g (m e an = 294, n = 3) a n d 3 pm b ro a d in its
m iddle. It ten d s to be s h o rte r in th e G alapagos individuals: 209-279 pm (n = 2).
The fem ale b u rs a is larg e a n d d iv id ed in to a m u sc u la r p a rt a n d a re so rb e n t
p art, c o n n ected to one a n o th e r via a m u sc u lar pore. T he m u sc u la r p a rt is globular,
su rro u n d e d by a n o u te r c irc u la r a n d a n in n e r lo n g itu d in al m uscle coat, a n d is filled
w ith a n eo sin o p h ilic secretio n . In som e sp ecim ens it also co n ta in s som e sperm . Its
ep ith e liu m is re d u c e d to a p se u d o c u tic u la , w h ich is folded in th e d ista l p a rt. The
b u rsa l stalk, co m in g from th e gen ital a triu m , en ters th is p a rt of th e b u rsa ventrally.
The d istal p a rt o f th e b u rsa l w all folds b ack giving th e im p ressio n o f a th im b le su r­
ro u n d in g th e p ro x im al p a rt o f th e b u rsa l stalk. The b u rsa l stalk is covered w ith a
low ep ith eliu m , in som e specim en s red u ced to a pseu d o cu ticu la, a n d is m ainly su r­
ro u n d e d by c irc u la r m u scles a n d so m e lo n g itu d in a l in n e r m u scles n e a r th e co m ­
m o n genital a triu m . A sp h in c te r g u a rd s th e m o st proxim al p a rt o f th e stalk. D istally
fro m th is s p h in c te r lo n g itu d in a l m u scles a tta c h to th e stalk a n d ru n to w a rd s the
d ista l w all of th e m u s c u la r p a rt of th e b u rsa . The in se m in a tio n d u c t leaves the
m u s c u la r p a rt o f th e b u rsa d o rsally a n d jo in s th e sem in al recep tacle to w h ich th e
s h o rt o v id u cts a re co n n e c te d . T his d u c t is lin ed w ith a p seu d o c u tic u la a n d is s u r­
ro u n d e d by a c irc u la r m u scle coat. In th e sp ecim ens from F lo rid a it is ra th e r short,
b u t it is so m ew h at longer in th e G alapagos specim ens. The fem ale duct, co n necting
th e sem in al recep tacle w ith th e u teru s, is long a n d narrow , w ith a very th in e p ith e ­
liu m a n d m ain ly th in lo n g itu d in al m uscles. O u ter c irc u la r m uscles, however, occur
a t b o th ends. The u te ru s jo in s th e fem ale d u c t in th e p roxim ity o f th e a triu m .
D iscu ssio n . The F lo rid a a n d G alap ag o s p o p u la tio n s of th is species exh ib it
som e m in o r differen ces: in th e G alap ag o s sp ec im en s th e in se m in a tio n d u c t is
slig h tly lo n g e r a n d th e size of th e accesso ry g la n d u la r o rg a n is so m e w h a t larger.
B o th p o p u la tio n s also d iffer in th e le n g th o f th e h a rd p a rts, b o th sty lets being
lo n g e r in th e sp e c im e n s fro m F lo rid a. H ow ever, th e p ro p o rtio n s o f th e le n g th s of
th e p ro s ta te sty let a n d th e accesso ry sty let c an be c o n sid e red th e sam e. In the
F lo rid a p o p u la tio n th e m a in sty let is 31% o f th e a ccesso ry sty let o n average (2833%), w hile it is 22% (21-22% ) in th e p o p u la tio n from the G alapagos. S im ilar v ari­
a tio n in le n g th c a n be fo u n d in other, co n g e n eric sp ecies (T.J. A r t o i s pers. obs.).
However, th e sh ap e o f b o th th e accessory a n d m a in stylet is the sam e in b o th p o p u ­
latio n s o f P. haegheni, a n d clearly se p a ra te s th em from th e o th e r Phonorhynchoides
species. In o th e r w ords, th e d ifferences b etw een the tw o p o p u latio n s o f P. haegheni
a re m u c h sm a lle r th a n th e d ifferen ces b e tw ee n th e m a n d th e o th e r con g en eric
species. T herefore, w e do n o t co n sid er th e d ifferences betw een th e p o p u latio n s o f P.
haegheni sufficient to a ttrib u te th em to tw o d ifferen t species.
The differences w ith th e o th e r species of th e genus are m ainly to be fo u n d in
th e sh a p e a n d le n g th o f th e h a rd p a rts. In P. som aliensis, P. carinostylis Ax &
T.J. A rtois a n d E.R . S cho ck aert
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70
Fig. 4. — Djeziraia incana n. sp. A, stylet; B, exceptionally long stylet; Phonorhynchoides haegheni n.
sp. C, atrial organs; D, stylet an d distal tip o f accessory stylet (all from live animals).
In te rstitia l fa u n a of th e G alapagos
71
A rm onies 1987 a n d P. flagellatus B eklem ischev 1927 th e m a in stylet is m u ch longer
th a n th e accessory stylet. The m a in stylet is also m o re o r less corkscrew -shaped in
th ese th re e species, w hile it is stra ig h t in P. haegheni. The division of th e b u rsa into
a m u sc u la r a n d a n o n -m u sc u la r p a rt is only fo u n d in P. haegheni.
P a r a c h ro r h y n c h u s jo n d e lii n. sp. (Figs 5-6)
Locations. S an ta Cruz: B ahia Academ y: S o u th e rn B each (IX. 6c) (type locality).
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Material. Drawings and photographs by S. Hoxhold of one live animal that was later
sagittally sectioned (holotype).
D erivation o f the n a m e. N am ed a fte r D r Ulf Jo n d e liu s (U ppsala U niversity,
U ppsala, Sw eden).
D escription. T he a n im a ls a re 0.7-0.8 m m long, colourless, w ith tw o eyes. The
ep ith eliu m is 3 pm h ig h w ith cilia 4 p m long. R h ab d ites a re ro d -shaped, a b o u t 3/4
o f th e h e ig h t o f th e e p ith e liu m in len g th ; th ey a re lacking a ro u n d th e p ro b o scis
p o re a n d a re especially n u m e ro u s a ro u n d th e caudal end, w here a m o re ovoid kind
o f rh a b d ite s is also p resen t. The c au d al glands a re well developed.
The p ro b o scis is a b o u t 20% o f th e body length. An in n e r c ircu lar a n d a n o u ter
lo n g itu d in al m uscle sh e a th su rro u n d th e p ro boscis cavity. N eith er th e eone n o r the
p ro b o sc is cavity h as in tra -e p ith e lia l nuclei; nu clei are obviously in su n k in th e cell
m a ss a ro u n d th e p ro b o sc is b u lb (see S c h o c k a e r t & B edini 1977 re g ard in g Polycy­
stis naegelii). T he ex act o rg a n is a tio n of th e m o tio n a l m u scles o f th e p ro b o sc is is
d ifficu lt to d e te rm in e , b u t th e n u m b e r o f re tra c to rs a n d in te g u m e n t re tra c to rs
a p p e a rs to be th re e a n d tw o p airs, respectively.
ut
Fig. 5. — Parachrorhynchus
jondelii n. sp. General organ­
isation (from a live animal).
gp
ga
Fig. 6. — Parachrorhynchus jondelii n. sp. Reconstruction of the
atrial organs from the left hand side.
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72
T.J. A rtois a n d E.R. S chockaert
The p h ary n x is situ a te d in th e first body h alf a n d is inclined slightly forw ards.
It is o f th e n o rm a l p o ly cy stid id type w ith th e u su a l tw o k inds of g lands, a n d th e
fo u r sclero tized te e th a t th e p ro x im al o p en in g o f th e p h ary n g eal lum en. T here are
24 in te rn a l lo n g itu d in al m uscles.
The g o n ad s a re p a ire d . T he r a th e r sm all testes lie dorsolaterally, ju s t b e h in d
th e phary n x , th e o varies ju s t b e h in d th e gonopore. B o th v itellaria extend d o rso la t­
erally, fro m b e h in d th e p ro b o sc is to th e c a u d a l en d o f th e an im al. H ere th ey are
very close a n d m ay even be c o n n ected to each other. The genital pore is a t 75% and
su rro u n d e d by a sp h in c te r o f six to seven m uscle fibres. The co m m o n genital a tri­
u m is lined by a low e p ith eliu m w ith very few nuclei, a n d su rro u n d e d by lo n g itu d i­
n a l m u scles. A larg e b u n d le o f e o sin o p h ilic g lands e n te rs th e a triu m above th e
u te ru s, w h ic h jo in s th e a triu m th ro u g h th e a n te rio r w all. It is o f th e n o rm a l poly­
cystidid type.
The sem in al vesicles a re lin ed w ith a low n u c le ate d e p ith eliu m a n d have one
layer of m ain ly lo n g itu d in al m uscles. They jo in to fo rm a long sem inal duct, w hich
h a s a n a n u c le a ted ep ith eliu m an d is s u rro u n d e d by a th in lo n g itu d in al a n d a th ick ­
er o u te r c irc u la r m uscle sh eath . It e n ters a vesicle in w h ich also the p ro sta te glands
em p ty th e ir c o n te n ts. T his vesicle c a n be c o n sid ere d th e p ro sta te vesicle (in c o m ­
p a riso n w ith th e o th e r sp ecies of D u p la c ro rh y n c h in ae). It is lin e d w ith a h ig h
n u c le a te d e p ith e liu m a n d su rro u n d e d by sp ira l m u scles. T here a re tw o k in d s of
p ro s ta te glands, e o sin o p h ilic a n d b a so p h ilic ones, b o th w ith co a rse g ran u les. The
se m in a l d u c t ru n s to w a rd s th e m ale a triu m , e c c en trically th ro u g h th e p ro sta te
se c re tio n w ith in th e p ro s ta te vesicle. T he m ale a triu m leaves th e p ro sta te vesicle
(th ro u g h a n o p en in g w ith a sp h in c te r) a n d e n te rs th e co m m o n a triu m do rsally on
th e rig h t side. T he m ale a triu m (o r m ale d u c t) is lin ed w ith a low a n u c le a ted
ep ith e liu m th a t tu rn s in to a p se u d o c u tic u la in th e n a rro w p ro x im al p a rt o f th e
duct. T ow ards th e co m m o n a triu m th e m ale d u c t enlarges. In th is part, th e e p ith eli­
u m is m u c h higher. The m ale a triu m is su rro u n d e d by a n in n e r lo n g itu d in al a n d an
o u te r c irc u la r m u scle sh e a th ; th e la tte r is, how ever, a b se n t a ro u n d th e pro x im al
p a rt a n d fo rm s a sp h in c te r a t th e e n tra n c e to th e co m m o n a triu m .
The fem ale d u c t e n te rs th e c o m m o n g en ital a triu m caudally. H ere a large
g lo b u lar sp h in c te r is fou n d . The d u c t is covered w ith a p se u d o c u tic u la a n d is su r­
ro u n d e d by a th in lo n g itu d in a l a n d a th ic k c irc u la r m u scle co at. Proxim ally, it
receives th e o v id u cts a n d th e v itello d u cts fro m b o th sides a n d c o n tin u e s fo r som e
distan ce to en d in th e te rm in a l b u rsa. T here is a rin g of (10?) knobs a t the e n tra n ce
of th e b u rsa. T he o viducts a re very sh o rt, lined by a pseu d o cu ticu la, a n d s u rro u n d ­
ed by c irc u la r m u scles a n d very w eak in te rn al lo n g itu d in a l m uscles, w h ich c o n tin ­
ue a ro u n d th e ovaries over a sh o rt d istan ce. At th e last oocyte a rin g o f tria n g u la r
p la te s (p ro b a b ly five: x in Fig. 6) is fo u n d . The lo n g itu d in a l m u scles a ro u n d th e
oviducts a n d ovaries seem to converge to w ard s th ese plates.
D iscu ssio n . T h ree re p re se n ta tiv e s of th e g en u s P arachrorhynchus a re now
know n: P. axi a n d P. bergensis, b o th d e sc rib e d by K a r l i n g (1956) fro m th e N o rth
E u ro p e a n A tlantic coast, a n d P. jondelii n. sp. The m a in co m m o n fe atu res of these
species are th e long m ale a triu m (so m etim es referred to as th e cirrus), the p ro state
vesicle th ro u g h w h ich th e sem in al d u c t ru n s eccen trically (i.e. a n asy m m etric co n ­
ju n c ta type: te rm in o lo g y o f K a r l i n g 1956), th e te rm in a l b u rsa , a n d th e o v iducts
th a t jo in th e b u rsa l stalk o n e ith e r side.
O nly P. bergensis h a s sp in es in th e m ale d u ct. T his p a rt c a n be ev erted as a
p ap illa (as o rig in ally d escrib ed ). H ow ever th is "papilla" is n o t a p e rm a n e n t struc-
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In te rstitia l fa u n a of th e G alapagos
73
tu re , as w as su b seq u en tly o bserved in a live a n im a l by th e late T.G. K a r l i n g (pers.
com m .). In P. bergensis th e re is a clearly d e lim ite d m ale b u rsa (c o n n e cted to the
a triu m by a n a rro w canal), in P. axi th e re is a caudal bulge a t th e d istal end o f the
m ale d u c t (w ith a d ifferen t e p ith eliu m as well), w hile in P. jondelii th ere is a sim ple
e n la rg e m e n t a t th e e n d of th e m ale a triu m . S perm can be sto re d h ere in th e th ree
species (n o t su re for P. bergensis).
As in P. axi, th e re is a rin g o f little k n o b s a t th e e n tra n c e o f th e b u rsa in P.
jondelii. S u ch k nobs w ere n o t observed in P. bergensis, b u t th e individual on w hich
th e d e sc rip tio n w as b ased clearly h a d n o t reach ed full fem ale m aturity.
The p ro b o sc is of P. jondelii is a b o u t 1 /5 o f th e to ta l body len g th , th e m o st
c o m m o n p ro p o rtio n fo u n d in P olycystididae. In th e tw o o th e r species it is only 7 13% o f th e body length, a p ro p o rtio n fo u n d in m o st o th e r D u p lacro rh y n ch in ae (see
A r t o i s & S c h o c k a e r t 1 9 9 9 ) . O th er strik in g differences a re th e p a ired sem in al vesi­
cles, th e h uge sp h in c te r a t th e fem ale d u c t a n d th e large glands en terin g th e co m ­
m o n a triu m . A u n iq u e fe a tu re is th e rin g o f se p a ra te p late s a t th e ovaries. S im ilar
s tru c tu re s a re fo u n d in th e g en u s Paulodora M arcus 1 9 4 8 (th e "c u tic u la r nozzles"
o f S c h o c k a e r t & K a r l i n g 1 9 7 5 ) . How ever, th e sim ila ritie s m ay be su p e rficia l, as
th e a p p e a ra n ce s in b o th lateral a n d fro n ta l view s are clearly different. In Paulodora
th e "nozzle" a t each ovary looks like a sm all u m b rella w ith a little stalk a n d it is a
sim ple circle in fro n ta l view w ith no se p a ra te plates to be found. T hus th e h o m o lo ­
gy of th is s tru c tu re in P. jondelii a n d in th e Paulodora species is n o t a t all certain.
M ost o f th e c h a ra c te rs sh a re d by th e th re e P arachrorhynchus species m ay be
c o n sid e re d sy m p le sio m o rp h ie s. W ith in th e D u p la c ro rh y n c h in ae , to w h ich Para­
chro rh yn ch u s b elo n g s (see A r t o i s & S c h o c k a e r t 1 9 9 9 ) , th ey are th e only species
w ith a co p u la to ry o rg a n w ith a n asy m m etrical, a lth o u g h in terp o sed , p ro sta te vesi­
cle. This c h a ra c te r c a n be p u t fo rw ard as a likely synapom orphy.
S ubfam ily M acro rh y n ch inae E v donin 1977
M a c r o r h y n c h u s m a n u s fe r r e a n. sp. (Figs 7-8, 11A-B)
Localities. S an ta Cruz: B ah ia B o rrero (IX. 3) (type locality). S an ta Cruz: B ahia
T ortuga (IX. 7b).
M aterial. D raw in g s a n d p h o to g ra p h s o f live a n im a ls b y S. H oxhold. Two serially sec­
tio n e d specim ens, o n e of th e m d esig n ated as h olotype (sagittally sectioned).
D erivation o f the nam e. T he sty let is g ra p n e l-sh ap e d . M anus fe rre a (Lat.):
grapnel.
D escription. A ccording to S. H o x h o l d 's notes, th e live an im als are u n pigm ented, th o u g h o ra n g e u n d e r in c id e n t light, a n d filled w ith b lack -b ro w n g lan d s u n d e r
th e ep id erm is w h en seen in tra n s m itte d light. H ow ever no su ch glands w ere found
in se c tio n e d m a te ria l. T he a n im a ls a re 0.8-1.2 m m long a n d have a p a ir o f eyes.
The e p id e rm is is sy n cy tial w ith n u m e ro u s vacuoles, m o st o f th e m em pty, som e
filled w ith a n eo sin o p h ilic secretion. The rh a b d ite s are sm all, 1/3 of th e ep ith eliu m
h e ig h t long, s le n d e r a n d sp in d le -sh a p e d . T he c o a rse-g rain e d , b a so p h ilic cau d al
glan d s a re w ell developed.
The probo scis, 1 /5 of th e body length, is o f th e n o rm a l polycystidid c o n stru c ­
tio n a n d h as th re e p a irs of re tra c to rs a n d tw o p airs of in te g u m e n t re tra cto rs.
74
T.J. A rtois a n d E.R. S chockaert
The p h ary n x is s itu a te d in th e first body h a lf a n d is in clin ed fo rw ards. M ore
o r less in th e m id d le o f th e p re p h a ry n g e a l cavity th e re is a rin g o f a ruffled
(p se u d o c ilia te d ) e p ith e liu m . T he re s t of th e p re p h a ry n g e a l cavity is lin ed w ith a
low, a n u c le a te d ep ith e liu m . An o u te r lo n g itu d in a l a n d a n in n e r c irc u la r m u scle
lay er s u rro u n d th e p re p h a ry n g e al cavity. The la tte r fo rm s a sp h in c te r betw een the
m iddle a n d th e pro x im al th ird of th e cavity, b u t is lacking proxim al to th is sp h in c­
ter. T he p h a ry n x c o n ta in s th re e ty p es o f glands: tw o e o sin o p h ilic a n d one
b a so p h ilic. T he b a so p h ilic o n e o p en s in to th e p h ary n g ea l lu m e n b etw e en th e tw o
eo sin o p h ilic ones, th e d a rk e st of w h ich o p ens m o st distally.
The to p o g ra p h y o f th e g o n ad s is sh o w n in Fig. 7A. The g o n o p o re is situ a te d
a t a b o u t 70%. The co m m o n genital a triu m is lined w ith a p seu d o ciliated e p ith eliu m
a n d su rro u n d e d by lo n g itu d in al m uscles.
The long p y rifo rm p ro s ta te vesicle c o n ta in s tw o types o f secretio n : a c o arse­
g rain ed b aso p h ilic one a n d a fin e-g rain ed eosinophilic one. The m u sc u latu re o f the
p ro sta te vesicle a n d o f th e pro x im al p a rt of th e m ale a triu m is ra th e r com plex and
co m p arab le to th a t o f Aí. groenlandicus (L evinsen 1879) G raff 1882 as describ ed by
S c h o c k a e r t & K a r l in g (1975). T h ree m u scle layers su rro u n d th e p ro sta te vesicle;
e a c h o f th e m situ a te d o u tsid e th e m e m b ra n e th a t b o rd e rs th e vesicle. T he in n e r
m uscle layer is m o re o r less circular, a n d only covers th e p ro state vesicle. The o u te r
tw o lay ers ex ten d fu rth e r alo n g th e p ro x im al p a rt of th e m ale a triu m . The m iddle
m
50 |xm
vs
b
— ov
in
Fig. 7. — Macrorhynchus manusferrea n. sp. A, general organisation; B, stylet; C, section through
the prepharyngeal cavity; (A-B from live animals).
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In te rstitia l fa u n a of th e G alapagos
75
lay er is m o re o r less lo n g itu d in a l b u t b eco m es m o re sp ira l a ro u n d th e pro x im al
p a r t o f th e m ale a triu m . T he o u te r lay er is sp ira l a ro u n d th e p ro sta te vesicle a n d
th e m a jo r p a rt o f th e m ale a triu m , b u t b ecom es lo n g itu d in a l to w ard s th e place
w h ere it a tta c h e s to th e m ale a triu m . The re tra c tio n of th ese fibres causes th e wall
o f th e m ale a triu m to fold. In th is fold, a stro n g sp h in c te r o ccurs a ro u n d th e distal
p a rt o f th e stylet.
The stylet is double-w alled w ith a 175 p m long o u te r stylet a n d a n 83 p m long
in n e r stylet, lying in th e d ista l h a lf o f th e o u te r stylet. The o u te r sty let is 31 pm
b ro a d a t th e pro x im al opening, n a rro w in g to 23 p m in a b o u t the m iddle, w here the
in n e r sty let begins, a n d n a rro w in g fu rth e r d istally to 7 pm . T he d istal p a rt o f the
sty let is r a th e r com plex, b u t h a s th e g en eral a p p e a ra n c e o f a hook w ith a g u tte r­
sh a p e d extension. D istally from th e stylet th e m ale a triu m b en d s to w ard s th e co m ­
m o n a triu m , w h ic h it e n te rs dorsally. T he d istal p a rt o f th e m ale a triu m is lined
w ith a relatively h ig h e p ith eliu m a n d is s u rro u n d e d by a w eak c irc u lar a n d a longi­
tu d in a l m u scle layer. The re st of th e m ale a triu m is covered w ith a pseu d o cu ticu la.
A b u n d le of co arse-g rain ed b aso p h ilic glan d s e n te rs th e m ale a triu m dorsally a t the
level of th e d istal tip of th e stylet.
The p a ire d se m in a l vesicles a re lin ed w ith a very low, n u c le a te d e p ith eliu m
a n d have a sp irally -ru n n in g , alm o st lo n g itu d in al m uscle coat. The sh o rt vasa defere n tia jo in e a c h o th e r u n d e r th e p ro s ta te vesicle. The eja c u la to ry d u c t is relatively
b ro a d a n d lin ed w ith a low e p ith eliu m w ith flatten ed nuclei; it is su rro u n d e d by a
circ u la r m uscle layer. It e n te rs th e m ale a triu m ventrally th ro u g h a m u sc u la r p a p il­
la close to th e d istal tip of th e stylet. This p apilla seem s to en d in th e g u tte r form ed
by th e stylet tip.
B oth o viducts a re very sh o rt a n d su rro u n d e d by a very w eak c ircu lar m uscle
layer. W here th ey jo in , a sm all b u rsa is p resen t, in w h ich sp erm a n d som e n e c ro t­
ic n u clei w ere observed. The fem ale d u c t is lined w ith a high, n u c lea te d e p ith eli­
um . It e n te rs th e c o m m o n g en ital a triu m a t th e sam e place as do es th e u te ru s,
vg
sph
ma
vs
m
ov
gp
Fig. 8. — Macrorhynchus manusferrea n. sp. Reconstruction of the atrial organs from the left hand
side.
76
T.J. A rtois a n d E.R. S chockaert
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giving th e im p ressio n o f a sh o rt d u c tu s u tero -co m m u n is, a t least in one specim en.
In o n e live in d iv id u al th e u te ru s c o n ta in e d a n egg cap su le w ith fo u r fully-grow n
young (Fig. 11 A).
D iscu ssio n . M. m anusferrea sh o w s th e a u ta p o m o rp h ie s o f th e genus as given
by A r t o i s & S c h o c k a e r t (1998): th re e p a irs o f p ro b o sc is re tra c to rs su p p le m e n te d
w ith tw o p a irs o f in te g u m e n t re tra c to rs, th e long stylet w ith a sh o rt in tern al stylet
a n d com plex ending. W ithin th e genus, it m o st resem bles M. groenlandicus (Levinsen 1879) G raff 1882. In b o th sp ecies th e e ja cu la to ry d u c t e n te rs th e m ale a triu m
th ro u g h a m u sc u la r papilla. F u rth e rm o re , in b o th species a bu n d le of glands opens
in th e m ale a triu m a t th e d ista l tip o f th e stylet. In M. groenlandicus th ese glands
a re e o sin o p h ilic a n d r a th e r d iffuse ( S c h o c k a e r t & K a r l i n g 1975), w hile th ey are
b aso p h ilic w ith co a rse g ra n u le s in M. m anusferrea. N evertheless th ey m ay be co n ­
sid ered h o m o lo g o u s in th e tw o species, a n d th e d iffe re n t a p p e a ra n c e of th e se cre­
tio n d u e e ith e r to th e co n d itio n o f th e an im als o r a fixation o r stain in g a rtefact.
M. m anusferrea clearly h a s a fem ale b u rsa , resem b lin g th a t o f th e Polycystis
species (see A r t o i s & S c h o c k a e r t 1998). A fem ale b u rsa h a s n o t b een observed in
M. groenlandicus (see S c h o c k a e r t & K a r l i n g 1975). The p resence of su ch a n o rg an
in M. croceus (F ab riciu s 1823) G raff 1882 is m e n tio n e d by G r a f f (1905), a lth o u g h
n o t in h is m o n o g ra p h of 1882. M e i x n e r (1925) explicitly den ies th e p resen c e of
su c h a b u rsa , a n d o u r ow n o b serv atio n s o n m a teria l o f M. croceus co nfirm M e i x n e r 's sta te m e n t. It could be a ssu m ed th a t th e fem ale b u rsa is a tem p o ra ry form ation,
fo rm ed ju s t b efore o r a fte r co p u latio n . B u t th is is ag ain c o n trad icte d by th e o b ser­
v a tio n s o f sp e rm in th e o v id u cts a n d th e stro m a a ro u n d th e ov aries in M. groen­
landicus w ith o u t an y tra c e o f a b u rsa l o rg a n ( S c h o c k a e r t & K a r l i n g 1975).
As to biogeography, M. m anusferrea is th e first M acrorhynchus species fo u n d
o u tsid e th e A tlantic, except fo r a d o u b tfu l rec o rd of M. croceus in th e A driatic Sea
by G r a f f (1913) (see S c h o c k a e r t & K a r l i n g 1975). M oreover it is by fa r th e so u th ­
e rn m o st re c o rd o f th e gen u s, M a ssa c h u se tts (USA) being u n til now th e so u th e rn
m o st b o u n d a ry ( S c h o c k a e r t & K a r l i n g 1975). T here is one re co rd of M. croceus in
th e C an ary Isla n d s by G r a f f (1913) b u t th is w as also c o n sid e re d d o u b tfu l by
S c h o c k a e r t & K a r l i n g (1975).
S ubfam ily P olycystidinae S cho ck aert & K arling 1970
P o ly c y stis a li S ch o ck aert 1982
Locality. S an ta Cruz: B ah ia Academ y: n o rth e rn side (IX. 5b).
D istrib u tio n . E a s t A frican C oast: fro m S o m alia ( S c h o c k a e r t 1982) to K enya
(E.R. S c h o c k a e r t pers. o bs.). C alifo rn ia (USA): several p laces a t P acific Grove, in
tid al pools a n d shallow w a te r ( K a r l i n g 1986).
Material. Five serially sectio n ed sp ecim en s a n d d raw in g s of live a n im als b y S. H oxhold.
A d d itio n a l in fo rm a tio n . T he in te rn a l o rg a n isa tio n o f th e sp ecim e n s fro m th e
G alap ag o s d o es n o t d iffer fro m e a rlie r d e sc rip tio n s ( S c h o c k a e r t 1982), n o r fro m
th a t of P. naegelii K ölliker 1845 (see th e d e scrip tio n by K a r l i n g 1956).
P. ali h a s b een fo u n d in th re e d ista n t localities (see above), a n d K a r l i n g ( 1 9 8 6 )
gave th e th re e p o p u la tio n s d ifferen t n a m e s as "form as". The S om ali a n d G alapagos
In te rstitia l fa u n a of th e G alapagos
77
p o p u la tio n s a re b o th u n ifo rm ly blue-black an d the stylet is alm o st th e sam e size an d
sh a p e . H ow ever, n o w h o le m o u n ts a re av ailab le o f th e G alapagos m a te ria l a n d a
close c o m p a riso n is difficu lt. T he C a lifo rn ian sp e cim en s a re co lo u rless a n d the
sty let is c o n sid e ra b ly larger. T he la tte r p o p u la tio n m ay th u s re p re se n t a se p a ra te
species, w hile th e S om ali an d G alapagos rep resen tativ es m ig h t be conspecific.
P a u lo d o ra fe lis (M arcus 1954) A rtois & S ch o ck aert 1998 a s y m m e tr ic a n. subsp.
(Figs 9-10, 11C-D)
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Locality in the Galapagos (P. felis asym m etrica). S a n ta Cruz: B ah ia A cadem y:
so u th e rn b e a c h (IX. 6b).
D istrib u tio n o f P. felis felis. B razil (M a rc u s 1954), Ja m a ic a ( T h e r r ia u lt &
K o la s a 1999), M exico: M azatlan , algae in a tid a l pool o n ro ck s in fro n t of H otel
V alentino (4.X II.1996) (ow n d a ta ). C uraçao: D am di C abicuchi (S p aan se W ater) at
th e side o f th e S p aan se W ater; m ixed algae (m ainly Caulerpa a n d H alim eda gantia)
w ith a lot o f d e tritu s o n th e th a llu s (14.X II.1998 an d 30.X II.1998) (ow n data).
Material. Galapagos: drawings and photographs of live animals by S. Hoxhold. Several
serially sectioned animals. Brazil: two whole mounts and three serially sectioned specimens
A
B
C
ov
vg-
y
ds
ov
od
od
b
Fig. 9. — Paulodora felis asymmetrica n. subsp. A, general organisation; B-D, stylets of different
specimens; E, reconstruction of the female atrial organs from above (bursal stalk and female duct
not indicated) (A-D from live animals).
78
T.J. A rtois a n d E.R. S chockaert
(m a te ria l from E. M arcu s in th e S w ed ish M useu m of N a tu ra l H istory, S tockholm ). C uraçao:
six w hole m o u n ts. Jam aica: one w hole m o u n t (leg. J. K olasa). M exico: th re e w hole m o u n ts.
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D erivation o f the nam e. The n am e of th e subspecies refers to th e asym m etricallyb u ilt fem ale system .
D escription. T he stylet is a lm o st id e n tic al in all p o p u la tio n s exam ined. It is
double-w alled, w ith a n in n e r stylet th a t c a n be observed over m o st o f th e length of
th e o u te r stylet. In its course, th e stylet m akes a tu rn o f ab o u t 270°. As a result, the
d istal p a rt ru n s p e rp e n d ic u la r to th e p ro x im al p a rt. W hen heavily squeezed, th e d is­
tal p a rt can fold d ow nw ards, giving th e stylet th e shape of a sickle. The o u te r stylet
h as a flap-like p ro je c tio n in its proxim al p a rt (som etim es difficult to observe) along
w h ich th e sp erm are d rain ed . The len g th of th e stylet is co m p arab le in th e different
p o p u latio n s: 77-88 p m (m ean = 84, n = 6) (G alapagos), 77-84 p m (m ean = 81, n =
2) (B razil), 76-82 p m (m ean = 79, n = 2) (M exico), 73 p m (Jam aica) a n d 81-92 pm
(m ean = 90, n = 6) (C uraçao). The large m ale b u rsa is co n n ected to th e m ale a tr i­
u m by a sh o rt b u rsa l stalk, leaving th e m ale a triu m th ro u g h its ca u d a l w all, close
to th e c o m m o n atriu m .
T he fem ale d u c t is r a th e r long a n d n arro w , lin e d w ith a high, n u c le a te d
ep ith eliu m a n d w ith a th in lo n g itu d in al m uscle coat. It leaves th e c o m m o n a triu m
a t th e ca u d a l side, ru n s fo rw ard s a n d b ifu rca tes in to th e oviducts above th e stylet.
The o v id u cts a re e n la rg e d a n d filled w ith sp erm , clearly fu n c tio n in g as sem in al
rec e p ta cle s, a n d a re s u rro u n d e d by a n in n e r lo n g itu d in a l a n d a n o u te r c irc u la r
m uscle sh eath . N ear th e ovaries th e lo n g itu d in al m uscles becom e thinner, w hereas
th e c irc u la r m uscles a re lacking com pletely. The fem ale system is co n n ec ted to the
m ale b u rsa by a m u sc u la r p o re a t th e b ifu rc a tio n o f th e fem ale d u ct. In th e speci­
m en s fro m th e G alap ag o s th e fem ale system is asy m m etric ally b u ilt. O n th e rig h t
side, th e o v id u ct is c o a te d by a relativ ely h ig h e p ith eliu m , w ith som e n u clei in
som e in d iv id u als. A larg e sp erm -receiv in g vesicle (y in Figs 9-10) is c o n n e c te d to
th e ovid u ct by a b ro a d p o re n e a r th e ovary. In som e specim ens th e vesicle is com ­
pletely filled w ith a diffu se m a te ria l, th e e p ith e liu m is unclear, a n d no sp erm are
a p p a re n t. In o th e r in d iv id u a ls it is co m p letely filled w ith sp erm an d lin ed by a
ov
vs
ma
Fig. 10. — Paulodora felis asymmetrica n. subsp. R econstruction of the atrial organs from th e left
hand side w ith om ission of the left h an d side of the female system.
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In te rstitia l fa u n a of th e G alapagos
79
C
Fig. 11. — Macrorhynchus manusferrea n. sp. A, uterus w ith an egg capsule containing four young
specimens; B, stylet; Paulodora felis asymmetrica n. subsp. C, stylet; D, stylet of an o th er specim en
(all from live animals).
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T.J. A rtois a n d E.R. S chockaert
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high, n u c le a te d ep ith e liu m . T he left ovary alw ays h as a slightly m o re c a u d a l p o si­
tio n th a n th e rig h t ovary. H ere a s e p a ra te sp erm -receiv in g vesicle is lacking. The
left o v id u ct d iffers fro m th e rig h t o n e in th a t it is lin e d by a n ep ith e liu m w hose
nuclei a re u n d e rn e a th th e m uscles, a n d w h ich pro b ab ly h as a secreto ry fu n ction. In
th e sp ecim en s fro m th e G alapagos, a "bridge" (z in Figs 9-10) c o n n ects th e p ro x i­
m al en d s o f th e ovid u cts to ea c h other. It is b ro a d o n th e left side, n a rro w o n the
rig h t side. It is s u rro u n d e d by very th in lo n g itu d in al m uscles, lined by a very low,
n u c le a te d e p ith e liu m a n d filled w ith fine g ran u lar, eo sin o p h ilic m a te ria l, w h ich is
co n d en sed in som e places.
D iscu ssio n . T he n ew su b sp e c ie s d iffers fro m th e n o m in a l su b sp ecie s P. felis
felis (M arcus 1954) in th e c o n stru c tio n o f th e fem ale system . A ccording to M a r c u s
(1954), th e b u rsa e n te rs th e genital a triu m from b ehind, w ith th e exit of th e fem ale
d u c t above it. R e -e x a m in a tio n o f M a r c u s 's m a te ria l revealed th a t th e s itu a tio n is,
in stead , th e sa m e as d e sc rib e d h e re fo r P. felis asym m etrica. T here are, however,
so m e m a jo r differen ces: th e rig h t a n d left o v id u cts a re th e sa m e in P. felis felis,
w ith o u t th e g la n d u la r e p ith e liu m o n th e left oviduct, n o r th e larg e vesicle o n th e
rig h t one. A "bridge" b etw een th e ovaries does n o t exist. These differences m ay ju s ­
tify giving th e n ew ta x o n th e s ta tu s o f species. H ow ever, w ith th e m a te ria l av ail­
able, n o d ifferences c a n be fo u n d in th e c o n stru c tio n o f th e stylet. As th e sh ape of
th e stylet is very typical fo r species w ith in the genus Paulodora M arcus 1948 a n d to
avoid la te r confu sio n , w e p refer to keep th e tw o taxa w ith in th e sam e species, even
th o u g h th e d ifferen ces obviously can clearly be seen in living sp ecim en s th a t have
re ach ed full fem ale m aturity.
The tw o subsp ecies are clearly g eographically sep arated : all individuals found
on th e A tlantic side of th e A m ericas belong to P. felis felis. So far, P. felis asym m etri­
ca h a s only b een fo u n d in th e G alapagos.
S ubfam ily G y ratricinae G raff 1905
G y ra trix h e r m a p h r o d itu s E h re n b e rg 1831 species com plex (Fig. 12)
Localities, (a) C olourless v a ria n t (G alapagos species I). Jervis N o rth (VIII. 1).
S a n ta Cruz: Playa B o rrero : co a rse sa n d in ro ck po o ls (IX. 2). S a n ta Cruz: B ah ia
B o rrero : co a rse sa n d in ro ck p o o ls (IX. 3). S a n ta Cruz: B ah ia A cadem y: n o rth e rn
side (IX. 5b). S a n ta Cruz: B ah ia Tortuga: co arse sa n d in rock pools a t th e edge of
m angroves (IX. 7). B arrin g to n : co arse sa n d in rockpools (XI. 1). (b) Yellowish vari­
a n t (G alap ag o s sp ecies II). S a n ta C ruz: B ah ia A cadem y: s o u th e rn b e a c h (IX. 6b).
B arrin g to n : fine sa n d in ro ck pools (XI. 1). (c) R eddish v a ria n t (G alapagos species
III). S an ta Cruz: B ah ia Academ y: so u th e rn b each (IX. 6b).
Other localities. R eco rd ed fro m all over th e w orld in fresh w ater, b rac k ish and
m a rin e localities.
Material. S everal live a n im a ls stu d ie d b y S. H oxhold. S ectio n ed sp ecim en s: 20 c o lo u r­
less specim ens, 5 yellow ish sp ecim en s a n d 2 re d d ish specim ens.
A dditional data and discussion. This eu ry h alin e co sm o p o litan w as fo u n d a b u n ­
d an tly in several lo calities in th e G alapagos a n d in th re e co lo u r variants: colourless,
yellow ish an d reddish.
In te rstitia l fa u n a of th e G alapagos
81
The c o lo u rle ss a n d yellow ish v a ria n ts have eyes a n d th e ir in te rn a l o rg a n isa ­
tio n co rre sp o n d s w ith th e d escrip tio n s fo u n d in th e lite ra tu re (e.g. K a r l i n g 1 9 6 3 ) .
The colourless v a ria n t can re a c h a len g th of 1.5 m m . The m e asu rem en ts (n =
7) of th e h a rd p a rts are: stylet 112-151 p m (m ean = 121), sh e a th 25-35 p m (m ean =
31), stalk of sh e a th 47-78 pm (m ean = 60).
The yellow ish v a ria n t ten d s to be sm aller, a b o u t 1 m m long. M easu rem en ts (n
= 5) of h a rd p arts: stylet 115-122 p m (m ean = 118), sh e a th 27-29 pm (m ean = 28),
stalk 77-86 p m (m ean = 82).
The re d d ish v a ria n t is 0.8-0.9 m m long a n d h a s no eyes. The in te rn a l o rg an i­
sa tio n o f these specim en s deviates from all fo rm er d escrip tio n s a n d from th e o th er
tw o v a ria n ts fro m th e G alap ag o s in th e o rg a n isa tio n o f th e fem ale system . The
extern al vagina is situ a te d ventrally b etw een th e m ale an d th e fem ale pore. It opens
alm o st im m e d ia te ly in to th e b u rsa . O p p o site th e vagina, th e b u rsa show s a fla t­
ten ed sp ace w ith a th ick sclero tized ro o f (w in Fig. 12). F rom th is "roof", lo n g itu d i­
nal m u scles ru n in all d irectio n s to w ard s th e v en tral side of the b u rsa a n d are co n ­
tin u o u s w ith th e lo n g itu d in a l m u scles th a t s u rro u n d th e b u rsa. In th e fro n tal p a rt
A
h
V
OV
r\A
W -
va
ogp
ds
20 (im
Fig. 12. — Gyratrix herm aphroditus. A, reco n stru ction of the fem ale system from the right h and
side; B, stylet of the reddish form; C, stylet of the yellowish form; D, stylet of the colourless form
(B-D from live animals).
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T.J. A rtois a n d E.R. S chockaert
of th e b u rsa a re tw o se p a ra te d c o m p a rtm e n ts filled w ith sp erm (v in Fig. 12). They
n a rro w to w ard s th e fro n ta l rim of th e b u rsa a n d a re co n n ected w ith th e oviduct by
tw o sh o rt in se m in a tio n d u cts (only o n e in o th e r Gyratrix species). A high nucleated
e p ith e liu m a n d lo n g itu d in a l m u scles s u rro u n d th e m . The ep ith e liu m o f th e d istal,
n a rro w e r p a rt o f th e c o m p a rtm e n ts is c o m p a ra b le w ith th a t o f th e oviduct, w hile
th e e p ith eliu m of th e pro x im al p a rt sta in s less deeply. The m ea su rem en ts (n = 2 ) of
th e h a rd p a rts are: stylet 1 7 1 - 1 7 5 pm , sh e a th 3 4 - 3 6 pm , stalk 5 0 - 5 9 pm .
The s tro n g p o ly m o rp h ism o f G . h erm aphroditus h a s b e en k n o w n fo r a long
tim e; G r a f f n a m e d a first sub sp ecies in 1 9 0 5 (G . herm aphroditus coeca) a n d a sec­
o n d o n e in 1 9 1 1 (G . h erm a p h ro d itu s m a culata). K aryological stu d ie s have also
revealed w ide v a ria tio n w ith in th e species ( R e u t e r 1 9 6 1 , L 'H a r d y 1 9 8 6 , several co n ­
trib u tio n s by C u r i n i - G a l l e t t i & P u c c i n e l l i : see refe ren ce s in C u r i n i - G a l l e t t i &
P u c c i n e l l i 1 9 9 8 ) . T hese la st a u th o rs co n sid er th e ta x o n as a species com plex w ith
m an y sibling species, som e even o c c u rrin g sym patrically. K aryotypes, size, a n d p ro ­
p o rtio n s o f th e h a rd p a rts m ay vary considerably. It is clea r th a t in th e G alapagos
a t least th re e "species" a re p re se n t (G alapagos species I-III).
The G . herm a p h ro d itu s m aculata o f G r a f f ( 1 9 1 1 ) is th e only yellow v a ria n t
know n. However, h e re yellow co rp u scles w ith in th e an im al cause th e yellow colour.
In th e G alap ag o s p o p u la tio n th e a n im a ls a re u n ifo rm ly yellow ish. The only red
v a ria n t kn o w n so fa r w as fo u n d by d e B e a u c h a m p ( 1 9 4 2 ) in the in te stin e (tak en as
food?) o f som e O ligochaeta fro m a n a q u a riu m in R ennes. Eyeless v arian ts, G . her­
m aphroditus coeca G raff ( 1 9 0 5 ) , w ere previously only reco rd ed from fresh w a ter in
p laces w ith little o r n o light: th e d eep w a te rs o f Lake G eneva, w ells in B ohem ia
(C zech R ep u b lic), b u t also in so m e floods in M oscow (for referen ces: see G r a f f
1 9 1 3 ) . The re d G . h erm a p h ro d itu s o f th e G alapagos is th u s th e first free-living
m a rin e v a ria n t th a t is red a n d w ith o u t ocelli. M oreover, it also deviates in its a n a to ­
m y fro m th e o th e r re p re se n ta tiv e s o f th e species com plex. We re fra in , how ever,
fro m n a m in g it form ally, avoid in g fu rth e r co n fu sio n , a n d co n sid er it a m e m b e r of
th e species com plex th a t can easily be reco gnised o n m o rphological grounds.
DIAGNOSES OF T H E N EW TAXA
Djeziraia incana n. sp. Djeziraia sp ecies w ith a very long, sle n d er stylet ( 1 6 2 pm ; 2 9 5 p m ) w ith a te rm in a l o p en in g . S h o rt fem ale d u c t e n te rin g th e b u rsa l
s ta lk in a b o u t th e m id d le. S p erm d u c t w ith c irc u la r m u scle s h e a th only a ro u n d
b o th ends, a n d w ith o u t sclero tized "horns".
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P honorhynchoides haegheni n. sp. P honorhynchoides species w ith a single­
w alled p ro s ta te stylet; 5 3 (G alapagos), 9 1 (F lo rid a) p m long. A ccessory se c re tio n
reserv o ir very large, w ith a large e x tra c a p su lar p art. A ccessory stylet single-w alled:
2 4 3 (G alapagos), 2 9 4 (F lorida) p m long. B u rsa w ith a stro n g m u sc u la r d istal p art,
co n n e c te d to th e p ro x im al p a rt by a m u sc u la r pore. In se m in a tio n d u c t a n d b u rsa l
stalk leave th e m u sc u la r p a rt of th e b u rsa a t d ifferen t places.
P arachrorhynchus jondelii n. sp. P arachrorhynchus species w ith a p ro b o scis ±
o f th e b o d y len g th . C o p u lato ry o rg a n w ith a n u n a rm e d m ale a triu m (cirrus),
th e te rm in a l p a rt en larg ed as m ale b u rsa. S em inal vesicles p aired. B undle of a tria l
g lan d s p re se n t. F em ale d u c t d ista lly w ith a larg e sp h in cter. B u rsal e n tra n c e s u r­
20%
In te rstitia l fa u n a of th e G alapagos
83
ro u n d e d by sm all teeth . O viducts proxim ally provided w ith a kind of closing a p p a ­
ra tu s co n sistin g o f tria n g u la r plates.
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M acrorhynchus m anusferrea n. sp. M acrorhynchus species in w h ich th e ejaculato ry d u c t e n te rs th e m ale a triu m th ro u g h a v en tral m u scu lar p apilla a t the distal
tip o f th e stylet. A ccessory glan d s in th e m ale system p resent. D ouble-w alled stylet
g ra p n e l-sh a p e d , 175 p m long. S m all fem ale b u rsa p re se n t a t th e p ro x im al e n d of
th e fem ale duct.
Paulodora felis (M arcu s 1954) A rtois & S ch o c k a ert 1998. Paulodora species
w ith a 73-92 p m long stylet. The d istal p a rt o f th e stylet ru n s p e rp e n d ic u la r to the
pro x im al p a rt in live an im als; w h en sq u eezed it can have the sh ap e of a sickle. The
fem ale d u c t is ra th e r long, a n d co m m u n icates w ith the m ale b u rsa th ro u g h a m u s­
c u la r p o re a t th e level o f its b ifu rc a tio n in to b o th oviducts. The o v iducts fu n c tio n
as sem in al receptacles.
P. f. felis (M arcu s 1954). F em ale sy stem sy m m etrica l, w ith o u t a d d itio n a l
sperm -receiving vesicle. W ith o u t a co n n e c tio n ("bridge") betw een th e proxim al ends
o f b o th oviducts.
P. f. a sym m etrica n. su b sp . F em ale sy stem asy m m etrica l. A d ditional sperm receiv in g vesicle c o n n e c te d to th e p ro x im al p a rt o f th e rig h t oviduct. L eft oviduct
w ith o u t su c h a vesicle. N uclei of th e e p ith eliu m of the left oviduct in su n k u n d e r the
m u scle sh eath s. P roxim al ends of b o th o viducts co n n ected by a "bridge".
POLYCYSTIDIDAE OF T H E GALAPAGOS: SPEC IES LIST
D u p lacro rh y n ch in ae S ch o ck aert & K arling 1970
Djeziraia incana n. sp.
P honorhynchoides haegheni n. sp.
Parachrorhynchus jondelii n. sp.
M acro rh y n ch in ae E v d o n in 1977
M acrorhynchus m anusferrea n. sp.
P olycystidinae S ch o ck aert & K arling 1970
Polycystis ali S ch o ck aert 1981
Paulodora felis (M arcus 1954) asym m etrica n. subsp.
P o rro cy stid in ae E vd o n in 1977
A ustrorhynchus galapagoensis A rtois & S cho ck aert 1999
G alapagorhynchus hoxholdi A rtois & S cho ck aert 1999
Pygm orhynchus pygm aeus A rtois & S ch ockaert 1999
G y ratricin ae G raff 1905
Gyratrix herm aphroditus E h re n b e rg 1831
C olourless v a ria n t (G alapagos species I)
Y ellowish v a ria n t (G alapagos species II)
R ed d ish v a ria n t (G alapagos species III)
T yphlopolycystidinae E v d o n in 1977
M yobulla m yobulla A rtois & S ch o ck aert 2000
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T.J. A rtois a n d E.R. S chockaert
M yobulla dunata A rtois & S ch o ck aert 2000
Sabulirhynchus axi A rtois & S ch o ck aert 2000
L im ipolycystis sp.
ACKNOW LEDGEM ENTS
We th a n k P rof. D r P e te r Ax fo r k in d ly se n d in g us all th e m a te ria l, in clu d in g
se c tio n e d m a te ria l as w ell as p h o to g ra p h s a n d draw in g s. D r Ulf Jo n d e liu s is
ackno w led g ed fo r sen d in g u s th e m a te ria l p re se n t in th e co llection of th e M useum
o f N a tu ra l H isto ry o f S to ck h o lm (S w eden) (d raw in g s o f Djeziraia euxinica an d
w hole m o u n ts a n d se c tio n s o f Paulodora felis a n d th e M acrorhynchus species). We
also th a n k D r N ikki W atson for critical read in g of the m a n u sc rip t a n d valuable co m ­
m en ts, a n d M r F ra n k V an B elleghem fo r h is h elp w ith th e figures. The G alapagosp ro ject w as fin an ced by th e S tiftu n g V olksw agenw erk. T his p a p e r is a c o n trib u tio n
to p ro je c t G .0086.96, fin a n c e d by th e F u n d o f S cien tific R ese arc h - F la n d e rs (Bel­
gium ).
R EFE R E N C E S
T.J. & S c h o c k a e r t E.R. 1998. A c lad istic re -asse ssm e n t of th e Polycystis species co m ­
plex (Polycystididae, E u k aly p to rh y n ch ia). Hydrobiologia 383: 97-102.
A r t o i s T.J. & S c h o c k a e r t E.R. 1 9 9 9 . In te rs titia l fa u n a o f th e G alapagos: P o rro c y stid in a e
(P laty h elm in th es P olycystididae). Tropical Zoology 1 2 : 3 0 9 - 3 2 4 .
A r t o is T.J. & S c h o c k a e r t E.R. 2000. In te rstitia l fa u n a of th e G alapagos: T yphlopolycystidinae
(P laty h elm in th es P olycystididae). Tropical Zoology 13: 141-158.
Ax P. & S c h m i d t P. 1973. In te rstitie lle F a u n a v o n G alap ag o s I. E in fü h ru n g . M ikro fa u n a
Meeresboden 20: 1-38.
B e a u c h a m p P. d e 1942. P se u d o p a ra sitism e d es G yratrix (R h ab d o co eles) d a n s u n O ligochète.
B ulletin de la Société Zoologique de France 67: 59-60.
C u r i n i G a l l e t t i M. & P u c c in e l l i I. 1998. T he G yratrix h erm a p h ro d itu s sp ecies com plex
(K alyptorh y n ch ia: P o lycystididae) in m a rin e h a b ita ts o f E a ste rn A ustralia. H ydrobiolo­
gia 383: 287-298.
G r a f f L. v o n 1 8 8 2 . M o n o g ra p h ie d e r T u rb ellarien . I. R h ab d o co elid a. Leipzig, Verlag vo n Wil­
helm E n g elm a n : I, IX + 4 4 2 pp.
G r a f f L. v o n 1905. M a rin e T u rb e lla rie n O ro tav as u n d d e r K ü ste n E u ro p a s. E rg e b n isse
einiger, m it U n te rstü tz u n g d e r k ais. A kadem ie d e r W isse n sch a fte n in W ien (au s d em
L egate W edl) in d e n J a h re n 1902 u n d 1903 u n te rn o m m e n S tu d ie n re ise . II. R h a b d o ­
coela. Zeitschrift fü r W issenschaftlichen Zoologie 83: 68-148.
G r a f f L. v o n 1 9 1 1 . Acoela, R h ab d o c o ela u n d A lloeocoela des O stens d e r V ereinigten S ta a te n
von A m erika. E rg e b n isse einiger. M it N a c h trä g e n zu d e n “M a rin e n T u rb e llarie n O ro­
tavas u n d d e r K ü sten E u ro p a s”. Zeitschrift fü r W issenschaftlichen Zoologie 99: 1-108.
G r a f f L. v o n 1913. P la ty h e lm in th e s. T u rb e lla ria II. R h a b d o c o elid a . Das Tierreich 35: XX +
A r t o is
484 pp.
I. 1985. In te rstitielle E u k aly p to rh y n ch ia (P lath elm in th es) v o n G alapagos. D iplom arbeit
z u Erlangung des Grades eines D iplom-Biologen, G öttingen, 83 pp.
K a r l i n g T.G. 1956. M o rp h o lo g isch -h isto lo g isch e U n te rsu c h u n g en a n d e n m ä n lic h e n A trialo r­
g an en d e r K aly p to rh y n ch ia (T urbellaria). A rkiv fö r Zoologi 2 (9): 187-279.
K a r l i n g T.G. 1963. D ie T u rb ellarie n O stfe n n o sk an d ie n s V. N e o rh a b d o c o e la 3. K aly p to ­
rh y n ch ia. Fauna Fennica 17: 1-59.
H enze
In te rstitia l fa u n a of th e G alapagos
T.G. 1986. F ree-living m a rin e R h a b d o c o e la (P la ty h e lm in th e s) fro m th e N. A m erican
Pacific coast. W ith re m a rk s o n species fro m o th e r areas. Zoologica Scripta 15 (3): 201-219.
L’H a r d y J.P. 1986. K aryology of m a rin e p o p u la tio n s o f G yratrix h erm aphroditus (T urbellaria:
R h ab d o co ela) a n d c h ro m o so m a l ev o lu tio n in th is sp ecies com plex. H ydrobiologia 132:
233-238.
M ack -F ira V. 1971. Deux T urbellariés nou v eau x de la M er N oire. R evue R o u m a in e de Biologie
(Zoologie) 16 (4): 233-240.
M a r c u s E. 1954. T u rb e llaria B rasileiro s-X I. Papéis a vulsos do D epartem ento de ZoologiaSecrataria da Agricultura-S. Paulo-Brasil 11 (24): 419-489.
M e i x n e r J. 1925. B eitrag z u r M o rphologie u n d zu m System d e r T u rb ellaria - R h ab docoela: I.
Die K alyptorh y n ch ia. Zeitschrift fü r M orphologie u n d Ökologie der Tiere 3: 255-343.
M i e l k e W. 1997. In te rs titia l fa u n a o f G alap ag o s. XL. C opepoda, p a r t 8. M icrofauna M arina
11: 153-192.
N o l d t U. & H o x h o l d S. 1984. In te rstitie lle F a u n a v o n G alap ag o s. XXXIV. S c h iz o rh y n c h ia
(P lath elm in th es, K aly p to rh y n ch ia). M icrofauna M arina 1: 199-256.
R e u t e r M. 1961. U n tersu c h u n g e n ü b e r R assen b ild u n g b ei Gyratrix herm aphroditus (T urbellar­
ia N eorhabdo co ela). Acta Zoologica Fennica 100: 1-32.
S c h o c k a e r t E.R. 1971. T u rb e lla ria fro m S o m alia I. K a ly p to rh y n c h ia (P a rt 1). M onitore Z oo­
logica Italiano (N uova Serie) S u pplem enta 4 (5): 101-122.
S c h o c k a e r t E.R. 1982. T u rb ellaria fro m S o m alia II. K a ly p to rh y n ch ia (P a rt 2). M onitore Zoo­
logica Italiano (N uova Serie) S u pplem enta 7 (2): 81-96.
S c h o c k a e r t E.R. & B e d i n i C. 1977. O b serv atio n s o n th e u ltra s tru c tu re of th e p ro b o sc is
ep ith elia in Polycystis naegelii K ölliker (T urbellaria E u k aly p to rh y n ch ia) a n d so m e asso ­
c iated stru ctu res. Acta Zoologica Fennica 154: 175-191.
S c h o c k a e r t E.R. & K a r l i n g T.G. 1975. A natom y a n d tax o n o m y o f som e species o f P olycysti­
did ae (T urbellaria, K aly p to rh y n ch ia) from N. A tlantic a n d M e d ite rran e an co asta l areas.
Zoologica Scripta 4: 133-143.
T h e r r i a u l t T.W. & K o l a s a J. 1999. N ew sp ecies a n d re c o rd s of m ic ro tu rb e lla ria n s from
co astal ro ck p o o ls o f Jam a ica , W est Indies. A rchiv fü r Hydrobiologie 144 (3): 371-381.
K a r l in g
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