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Phuke Ma ine Biological Cen e Special Publica ion 23(1): 273–280 (2002)
PARAPROTELLA SALTATRIX, A NEW SPECIES OF THE CAPRELLIDEA (CRUSTACEA:
AMPHIPODA) FROM PHUKET ISLAND, THAILAND
Ichi o Takeuchi1 and José Manuel Gue a-Ga cîa2
1Depa men o Li e Conse a ion Ecology, Facul y o Ag icul u e, Ehime Uni e si y, 3-5-7 Ta umi,
Ma suyama, Ehime 790-8566, JAPAN
2Labo a o io de Biologîa Ma ina, Depa amen o de Fisiologîa y Biologîa Animal, Facul ad de Biologîa,
Uni e sidad de Se illa, Apdo. 1095, 41080 Se illa, SPAIN
ABSTRACT
Pa ap o ella sal a ix sp. no . is desc ibed om specimens om Phuke Island, he Andaman Sea
coas o Thailand. Pa ap o ella sal a ix di e s om P. p ima Maye , 1903 by he lack o la e al p ojec ions
on head and pe eoni e II and by he u opod being educed o a single se a. No single male specimen o P.
sal a ix could be ound among he mo e han 70 specimens o P. sal a ix collec ed, indica ing he
possibili y ha P. sal a ix is pa henogene ic.
INTRODUCTION
Al hough he opical a eas o he Indo-Paci ic
a e well ecognised as ha ing a high species
di e si y o ma ine in e eb a es, cap ellidean
amphipods ha e a ely been eco ded om hese
a eas wi h he excep ion o Maye (1903) and
Laubi z (1991). Recen ly, Takeuchi and Sawamo o
(1998) eco ded i e species o he Cap ellidea
om he Sou h China Sea and Gul o Thailand.
Howe e , mos o hese specimens had been
damaged and accu a e desc ip ions o es ablish new
axa o o edesc ibe specimens could no be
achie ed. In Thailand, hi een species o
cap ellidean a e known om he coas o he Gul
o Thailand (Maye , 1903; McCain and S einbe g,
1970) bu , un il now, none ha e been eco ded
om he Andaman Sea coas .
Du ing Ma ch 1999, he senio au ho , had
an oppo uni y o conduc a p elimina y ield
su ey on he sou he n coas o Phuke Island while
isi ing he Phuke Ma ine Biological Cen e , and
collec ed se e al species o cap ellidean. Among
he collec ed ma e ial we e 71 specimens o an
undesc ibed species o which males a e appa en ly
lacking. This species is he e desc ibed.
MATERIALS AND METHODS
Ma e ial om he BIOSHELF p og am was
sen o he senio au ho o inclusion in his
olume. Un o una ely his ma e ial was in a
condi ion ha p ecluded accu a e desc ip ion o
axonomic pu poses. Howe e , he senio au ho
had made collec ions om his egion, and ha
ma e ial is he e desc ibed.
Samples o a ious ypes o subs a a on
which cap ellideans we e expec ed o inhabi we e
ca e ully de ached om ocks and opes. Samples
we e collec ed om se e al locali ies a ound sou h-
wes a ea o Phuke Island and placed immedia ely
in indi idual plas ic bags. Samples we e b ough
o he labo a o y and opened in a shallow ay illed
wi h seawa e . The cap ellideans we e hen picked
ou using o ceps and p ese ed in 5 % neu alised
seawa e – o malin.
O he specimens examined, wo indi iduals,
a ma u e emale (holo ype) and a p ema u e emale
(pa a ype ‘sma ’), we e d awn using a came a
lucida: he body leng h o all specimens was
measu ed using a p o ile p ojec o . Holo ype and
se e al pa a ype specimens a e deposi ed in he
Re e ence Collec ion o Phuke Ma ine Biological
Cen e (PMBC), Thailand.
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In e na ional Wo kshop on he C us acea o he Andaman Sea
TAXONOMY
Family Cap ellidae
Genus Pa ap o ella
Pa ap o ella sal a ix sp. no .
(Figs 1–4)
Ma e ial examined
Holo ype: PMBC 14936, Female, ma u e, o
no h-eas co ne o Aeow Island, 07°45.92´N,
098°24.06´E, 28.5°C , SCUBA, 12 m, coll. I.
Takeuchi, 23.03.1999.
Pa a ypes: PMBC 14937, Female ‘sma ’,
p ema u e, PMBC 14938, Lo A, 2 ma u e, 10
p ema u e and 3 imma u e emales, same da a as
holo ype, and Lo B, 4 ma u e, 31 p ema u e and
17 imma u e emales, PMBC pie , 07°48.10´N,
098°24.35´E, 29.0°C, by hand, 1 m, coll. I.
Takeuchi, 25.03.1999.
Desc ip ion
Ma u e emale, holo ype. Body leng h, 8.08
mm; head and pe eoni es slende . Pe eoni e V
longes , ollowed by pe eoni e III. Leng h o head
0.57 mm; leng h o pe eoni es I–VII 0.76, 1.29,
1.45, 1.22, 1.67, 0.63 and 0.49 mm, espec i ely.
Head wi h elonga e iangula p ojec ion; eyes la ge
and dis inc i e. Pe eoni e I wi h o wa dly ben
p ojec ion nea pos e io end o do sal su ace.
Pe eoni e II wi h pai o small mid-la e al
p ojec ions and a pos e io p ojec ion. Pe eoni e
III and IV wi h a shallow mid-do sal p ojec ions,
espec i ely.
An enna I elonga e, nea ly 0.8 as long as body;
p opodus a icle 2 longes . Flagellum sligh ly longe
han p opodus, 19-segmen ed. An enna II a li le
mo e han 0.3 imes as long as an enna I; p opodus
a icles 2 o lagellum a icle 1 sca cely se ose.
Gna hopod I, me us o p opodus se ose.
P opodus 1.5 imes as long as wide.
Gna hopod II posi ioned abou 0.3 om he
an e io end. Basis 1.5 imes as long as pe eoni e
II, dis ally wi h iangula p ojec ion. P opodus
elonga e, 2.5 imes as long as wide, wi h 3
p ojec ions on palm; hooked spine on small
p ojec ion si ua ed 0.20 om p oximal end; second
p ojec ion si ua ed 0.40 om dis al end; hi d
p ojec ion sepa a ed by “U” no ch om second
p ojec ion 0.25 om dis al end o palm; palm
be ween g asping p oximal p ojec ion and second
p ojec ion wi h se a i o m ee h, sca cely se ose.
Dac ylus alca e.
Pe eopod III minu e; a icle 1 ec angula ,
a icle 2 o a e, sca cely se ose; a icle 3 es igial
wi h 3 se ae. Pe eopod IV simila o pe eopod III.
Pe eopods V, VI and VII becoming
p og essi ely mo e obus . Pe eopod V elonga e;
ca pus longes ollowed by basis, me us and
p opodus. P opodus cylind ical and con ex wi h
a pai o long se ae nea p oximal end o palm.
Pe eopod VI wi h p opodus longe han ca pus.
P opodus o pe eopod VII de eloping pai o
g asping spines nea p oximal end o palm ins ead
o long se ae on pe eopods V and VI.
Mou hpa s. Maxilliped inne pla e o
ec angula , wi h 2 pai s o se ae nea dis al ma gin;
ou e pla e o al, 2.5 imes as long as inne pla e,
wi h 2 se ae on dis al ma gin and 5 se ae on la e al
ma gin. Palp a icle 2 wi h 5 se ae on medial
ma gin; a icle 3 se ose on dis al hal o la e al pa ;
a icle 4 alca e weakly in e io ly cu ed and inely
se ose nea dis al end; maxilla 1 ou e pla e o oblong
wi h 5 s ou apical ee h, dis al pa o palm wi h 4
apical ee h ollowed by 2 se ae. Ou e pla e o
maxilla II apezoid, wi h apical se a; palm wi h
ow o apical se ae. Le mandible wi h a icle 3 o
palp wi h 2-7-1 se ae o mula (see ema ks); a icle
2 wi h 3 se ae on la e al pa ; inciso di ided in o 5
ee h, and lacinia mobilis di ided in o 5 ee h
ollowed by 3 inely se ose se ae. Righ mandible
simila o le mandible excep o lacinia mobilis
wi h 7 small ee h ollowed by 2 inely se ulose
se ae. Uppe lip weakly no ched. Lowe lip wi h
inne lobe sepa a ed in o 2 ound lobe and ou e
lobe.
Abdomen. U opod degene a ed in o 1 se a.
Abdominal lobe wi h 2 apical se ae.
Female, pa a ype ‘b’. Body leng h, 5.81 mm.
Pe eoni e V longes . Pe eoni es III and IV wi h
s aigh do sal line. An enna I wi h lagellum 11-
segmen ed.
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Phuke Ma ine Biological Cen e Special Publica ion 23(1): 273–280 (2002)
Figu e 1 Pa ap o ella sal a ix sp. no . A, ma u e emale, holo ype, PMBC 14936; B, p ema u e emale, pa a ype ‘b’, PMBC 14937.
Ba indica es 1.0 mm.
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In e na ional Wo kshop on he C us acea o he Andaman Sea
Figu e 2 Pa ap o ella sal a ix sp. no . Ma u e emale, holo ype, PMBC 14936. A, an enna I; B, an enna
II; C, gna hopod I; D, gna hopod II. Ba s indica e 0.2 mm.
Type locali y
Aeo Island, o he sou h-eas e n coas o
Phuke , Thailand.
E ymology
The epi he sal a ix is de i ed om ‘dancing
lady’ alluding o he shaking and clinging beha iou
o Pa ap o ella sal a ix sp. no .
Rema ks
Gene ic and species diagnoses o he
Cap ellidea ha e been usually based on ma u e
males. Maye (1903) es ablished he genus
Pa ap o ella Maye , 1903 based on wo species:
P. p ima Maye , 1903 ( om Singapo e, Thailand
and Japan) and P. secunda Maye , 1903 ( om
Japan) (Maye 1903, A imo o 1976). McCain and
S einbe g (1970) designa ed P. p ima, as he ype
species o Pa ap o ella.
Al hough no adul male specimen was
a ailable, compa ison be ween he p esen ma e ial
o Pa ap o ella sal a ix and he diagnosis and
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Phuke Ma ine Biological Cen e Special Publica ion 23(1): 273–280 (2002)
Figu e 3 Pa ap o ella sal a ix sp. no . Ma u e emale, holo ype, PMBC 14936. A, pe eopod III; B,
pe eopod IV; C, pe eopod V; D, pe eopod VI; E, pe eopod VII. Ba s indica e 0.5 mm.
desc ip ion o he ma u e emale o P. p ima
(Maye , 1903; A imo o, 1976) p o ided a clea
di e ence in he species diagnosis. 1) In P. sal a ix
head and pe eoni e II lack la e al p ojec ions, while
in P. p ima hese wo somi es ca y dis inc
p ojec ions. 2) In P. sal a ix, pe eoni es III and
IV lack he la e al p ojec ions, while in P. p ima
hese pe eoni es III and IV ca y dis inc
p ojec ions. 3) In P. sal a ix he u opod o he
ma u e emale is es igial and has degene a ed in o
one se a, while in P. p ima new species i is
uni amous. O he abo e h ee diagnos ic poin s,
he u opod cha ac e has usually been ega ded as
o gene ic signi icance. Recen ly, he u opod on
he abdomen o Cap ella has been obse ed o
de elop om single se a o be p obably uni amous
du ing pos ma supial g ow h (Takeuchi, 1989;
Mo i, 1999). This, oge he wi h he lack o male
specimens, sugges s ha he di e ence be ween
he u opods o P. sal a ix and P. p ima is no

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In e na ional Wo kshop on he C us acea o he Andaman Sea
Figu e 4 Pa ap o ella sal a ix sp. no . Ma u e emale, holo ype, PMBC 14936. A; maxilliped; B, maxilla
I; C, maxilla II; D; le mandible; E, igh mandible; F, lowe lip; G, uppe lip; H, en al iew o abdomen.
Ba s indica e 0.05 mm.
necessa ily su icien o jus i y a new genus.
Al hough Pa ap o ella secunda has no been
as well-desc ibed as P. p ima (Maye , 1903),
Maye (1903) desc ibed he p esence o la ge
la e al p ojec ion on pe eoni e VI o male, 7 mm in
body leng h, and ea u e o emale, 8 mm in body
leng h. Pa ap o ella sal a ix which o a simila
size o P. secunda, bu he lack o la e al p ojec ion
in P. sal a ix allows he clea sepa a ion om P.
secunda.
In gene al, se a o mula o he hi d a icle o
mandibula palp in Pa ap o ella and ela ed gene a
was exp essed by o mula ‘1-x-y-1’ (Maye , 1903;
McCain, 1968; A imo o, 1976). This o mula
indica es he p esence o 1 long se a a ei he end
o ow o a iable numbe o sho se ae (x) and
in e media e one (y) (McCain, 1968). The se al
ow o he hi d a icle o mandibula palp in P.
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Phuke Ma ine Biological Cen e Special Publica ion 23(1): 273–280 (2002)
sal a ix has 2 long se ae nea p oximal end
ollowed by 6 sho se ae and 1 long se ae. Thus,
he se al o mula o he p esen species is newly
assigned as 2 (long se ae nea p oximal end)-x
(numbe o sho se ae)-1(long se ae nea apical
end).
Pa ap o ella sal a ix may be i s eco ded
case o pa henogenesis in he Cap ellidea. No
males we e ound among he 71 specimens o P.
sal a ix collec ed om Aeo Island and he Cape
Panwa (Fig. 5). The imma u e s age o P. sal a ix
anged 2.95–5.25 mm in body leng h, he
p ema u e s age om 4.00–6.40 mm, and ma u e
s age eached a maximum o 8.20 mm. This biased
sex a io is no expec ed om he ma ing sys em
o he Amphipoda and om he p e ious s udies
on he sex a io o he Cap ellidea. Cap ella, he
mos well-s udied genus o he Cap ellidea, and
Deu ella a e epo ed no o be ex emely biased
om 1:1 (males: emales) (e.g., Lewbel, 1978;
Bynum, 1978; Caine, 1979; Takeuchi and Hi ano,
1991, 1992). Gene ally, emales o Amphipoda do
no s o e spe m, and emales a e able o
successi ely p oduce emb yos se e al imes a e
sexual ma u i y (e.g., Bo owsky, 1991). Thus, once
a emale a ains he ma u e s age, a male mus
accompany he emale o each ime o e ilisa ion
which occu s a e e e y moul ing. Pa heno-
genesis in he Amphipoda has been suspec ed only
in a ew species o he subo de s Gamma idea and
Hype idea; e.g., Co ophium bonnelli Milne
Edwa ds, 1830 (C aw o d,1937; Moo e, 1981;
Mye s e al., 1989), and Rhabdosoma, a pelagic
genus o he Hype idea (Fage, 1954; Cha niaux-
Co on, 1960). In he nea u u e, labo a o y
ea ing expe imen s on P. sal a ix om newly
ha ched ju eniles un il hei ma u a ion in isola ion
a e expec ed o elucida e hei li e his o y, and allow
hei ep oduc i e beha iou o be de ailed,
especially whe he his species is pa henogenesis
o no .
ACKNOWLEDGEMENTS
The au ho s exp ess hei since e hanks o
M . Somchai Bussa awi and s a o he Phuke
Ma ine Biological Cen e o hei kind help o he
sampling nea he Cen e , and D . Niel L. B uce
o kindly a anging he submission o he p esen
pape o he P oceedings o he Wo kshop:
Biodi e si y o C us acea in he Andaman Sea. The
p esen s udy was pa ially suppo ed by he
Coope a i e In e na ional Resea ch on Ma ine and
Coas al En i onmen .
Figu e 5 Size dis ibu ion o body leng h (mm) in Pa ap o ella sal a ix sp. no . collec ed om wo
s a ions, Aeo Island and Cape Panwa, bo h o he sou h-eas coas o Phuke Island.
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In e na ional Wo kshop on he C us acea o he Andaman Sea
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