The status of Bogidiella Balearica Dancau, 1973, a stygobiont amphipod from Mallorca
ENDINS, n . O 13. 1987. Ciutat de Mallorca.
by Jan H. STOCK (*) & Thornas M. ILIFFE (**)
Se presenta la redescripción de Bogidiella balearica Dancau, 1973, un anfípodo es-
tigobionte de las cuevas anquihalinas de Mallorca. Queda demcstrado que se trata de
una buena especie, clasificable dentro del subgénero Bogidiella s. str., y que se puede
diferenciar fácilmente de Bogidiella chappuisi Ruffo, 1952 (con la cual se intentó sino-
nimizar con anterioridad), perteneciente a otro subgénero, Medigidiella.
Redescription of Bogidiella balearica Dancau, 1973, a stygobiont amphipod from
anchihaline caves i n Mallorca. It proves t o be a good species, t o be classified with the
subgenus Bogidiella s. str., and it is well-distinguished from Bogidiella chappuisi
Ruffo, 1952 (with which it was tentatively synonyrnized i n the past), belonging to a
different suhgenus, Medigidiella.
Bogidiella balearica Dancau, 1973
Bogidiella balearica was described by D. Dan-
Dancau, 1973: 114-1 19, figs. 14; Kararnan, 1979:
cau (1973) after specirnens frorn two caves in Ma-
24-25 (synonyrny discussed); Stock, 1981: 354 (cited
llorca, Coves del Drac and Cova des Pont.
only); Kararnan, in Ruffo, 1982: 253 (cited only).
The status of B. balearica was discussed by
1 8, Mallorca 86-002, Coves del
Kararnan, 1979: 25, who regarded it as a apossible
Drac (Manacor), in upper layers of 2nd lake of corn-
synonyrn)) of B. chappuisi Ruffo, 1952. This state-
rnercial cave; surface salinity 3 ppt, surface ternpe-
rnent was repeated by Kararnan (in Ruffo, 1982: 253).
rature 18.8' C; 10 Jan. 1986.
In a series of Arnphipoda collected early 1986
1 d , Mallorca 86-004, Coves dels Harns (Mana-
in Mallorcan caves by the junior author, two rnale
cor), in upper layers; surface salinity 14 ppt, sur-
specirnens of a Bogidiella were encountered,
face ternperature 19.6O C; 11 Jan. 1986.
which were rnorphologically in good agreernent
The following notes rnay serve to supplernent
with Dancau's description of B. balearica. Moreo-
Dancau's description, which is in general quite
ver, one of the specirnens carne frorn the sarne
cave (Coves del Drac) as Dancau's type-material.
Body length 2.2 and 2.4 rnrn. Head (fig. 1):
On the basis of the new material, the taxonornic
Ocular lobe narrow, rounded; antennal sinus shal-
status of B. balearica is clarified; it appears to be a
l o w but distinct.
good species.
First antenna (fig. 2): First peduncle segrnent
with medioventral spine. Aethetasks on al1 8 flagel-
lurn segrnents, each as long as corresponding seg-
lnstitute of Taxonomic Zoology, University of Amsterdam,
P. O. Box 20125, 1000 HC Amsterdam, The Netherlands.
rnent. Accessory flagellurn rather long, 3-segment-
(**) Bermuda Biological Station for Research, Ferry Reach 1-15,
ed; short aesthetask on tip of segrnent 3.
Second antenna (fig. 3): Gland cone elongate-

triangular, rather short. Flagellum 5-segmented,
First uropod (fig. 24): Peduncle with strong
aesthetasks on segments 2 and 5, subequal to
proximoventral spine. Margins of rami unarmed.
lenght of corresponding segment.
Exopodite slightly shorter than endopodite, each
Upper lip as illustrated (fig. 4).
ramus with 4 distal spines.
Mandibles (figs. 5, 6): Molar seta present both
Second uropod (fig. 25): Endopodite longer
lefi and right. Left lacinia mobilis with 5 coarse
than peduncle and longer than exopodite. Exopo-
teeth, right lacinia finely toothed (9 teeth).
dite with 4 distal spines (3 short, 1 long). Endopo-
Lower lip (fig. 7), contrary to Dancau's descrip-
dite with 4 shorter and 1 longer distal spines, non
tion with well-developed inner lobes.
of them modified in male.
First maxilla (fig. 8): Outer lobe with 7 spines
Third uropod (fig. 26): Rami slender, at least
(2 pluridentate, 1 with 3 denticles, 2 with 2 denti-
2.5 times as long as peduncle. One of the distal
cles, 2 with 1 denticle).
spines of each ramus very long (>33 % of length
Second maxilla as illustrated (fig. 9).
of ramus).
Maxilliped: lnner lobe (fig. 11) with bicuspidate
Telson (fig. 27): Rather deep, widely V-shaped
spines on distal margin. Outer lobe (fig. 10) with 3
distal cleft. Two plumose sensorial setae on' either
finely denticulated, simple spines.
side. Each telson lobe with 2 long distal spines
First gnathopod (fig. 12): Coxal plate trapezoi-
(longest spine longer than telson, shortest spine
dal, wider than long. Posterior margin of basis with
about 5/6 of length of longest spine).
1 short and 2 long setae. Palmar index (sensu
Ruffo, 1973) 0.46. Palmar margin with 5 bifid spi-
nes, some setae, and 2 rows of fine denticles: an
Angle row (A in fig. 13) and a row at the Base of
the claw (B in fig. 13); the B-row is short and the
A-row is implanted in a very shallow palmar angle
The absence of modified elements (spines, se-
sinus. Three setule-tipped palmar angle spines.
tae) on the endopodite of the second uropod and
Second gnathopod (fig. 14): Coxal plate wider
the exopodite of the second pleopod in the male of
than long. Posterior margin of basis with 1 short
the Mallorcan taxon, show that it belongs to the
and 1 long seta. Palmar index 0.46. Palmar margin
subgenus Bogidiella s. str. (see Stock, 1981, and
fig. 15) with 6 bifid spines, some setae, and short
Karaman, 1982). Its alledged senior synonym, B. chap-
rows of A- and B-spinules. Two setule-tipped pal-
puisi Ruffo, 1952 (see Karaman, 1979) belongs to
mar angle spines.
the subgenus Medigidiella, since it possesses mo-
Third pereiopod (fig. 16): Basis with 4 spines
dified spines, presumably serving for sperm trans-
o n anterior margin, 3 on posterior margin. Merus
fer, on the second male uropod.
3.5 times as long as wide. Propodus with 3 setae
We have compared the Mallorcan material with
o n posterior margin.
a sample of B. (M.) chappuisifrom the type area (lit-
Fourth pereiopod (fig. 17): Almost identical to
toral interstitial waters, Roussillon coast, France),
P3. Coxal plate very short. Coxal gills on P4 - P6,
and have observed several additional characters,
ovate, with short peduncle.
allowing separation of both sexes of balearica and
Fifth pereiopod (fig. 18): Coxal plate vaguely
chappuisi. Some of these are shown in figs. 28-34
equilobate. Merus 4.2 times as long as wide.
of the present paper; moreover the correct illustra-
Sixth pereiopod (fig. 19) much longer than fifth.
tions in Karaman, 1979, figs. 1-IV, and in Karaman
Coxal plate slightly anterolobate. Four spines on
(in Ruffo), 1982, in particular fig. 171, based on to-
posterior margin of basis. Merus 5.1 times as long
potypes, may serve very well for comparison.
as wide. Anterior margin of propodus with 2 setules.
The discriminating characters are: (1) The
Seventh pereiopod (fig. 20) longer than sixth.
greater elongation of several appendages (pedun-
Coxal plate hardly lobate. Posterior margin of basis
cle of first antenna, mandible palp, pereiopods 3
with 5 spines. Merus 4 times as long as wide.
through 7, third uropod) in balearica. For instance,
Some setae on anterodistal end of carpus. Very
the merus of P3, P5, and P6 is 2.5, 2.15, and 2.8
long setae on propodus.
times as long as wide in chappuisi, against 3.5,4.2,
Lentiform organs small, rounded, slightly
and 5.1 times, respectively, in balearica. (2) The
elleptical, smooth-edged, in basal part of basis of
presence of 3 bicuspidate spines on the inner lobe of
P3 - P7.
the maxilliped (2 bicuspidate spines and an unarm-
Epimeral plates (fig. 21) unarmed. Posteroven-
ed swelling in chappuisi, fig. 28). (3) The presence
tral corner produced into small tooth.
of 3 setae (2 long, 1 short) on the posterior margin
Pleopods 1 to 3 similar, without endopodite.
of the basis of gnathopod 1 (1 long and 1 short in
Second pleopod (fig. 22) not modified in male. Two
chappuisi, fig. 29). (4) Palmar margin of gnathopod
retinacula on inner side of peduncle of each pleo-
1 with ca. 6 bifid spines (0-3 in chappuisi, fig. 30).
pod, anchor-shaped, with 3 pairs of hooks (fig. 23).
(5) The palmar angle sinus of gnathopod 1 is shal-

Figs. 1-8. Bogidiella (B.) balearica Dancau, 1973, 6 (Manacor,

1, cephalosome, from the left (scale WX); 2, first antenna (WX);
3, second antenna (WX); 4, upper lip (M);
5, right mandible
(WZ); 6, left mandible, palp omitted (WZ); 7, lower lip W ) ;
8, first maxilla (WZ). Scales below fig. 19.

Figs. 9-17. Bogidiella (B.) balearica Dancau, 1973, d (Manacor,
9, second maxilla (scale WZ); 10, outer lobe of rnaxilliped (WZ);
11, inner lobe of rnaxilliped ( W ) ; 12, first gnathopod (WX); 13, pal-
mar margin of first gnathopod (WY) [A = angle row of spinules,
irnplanted in angle sinus; B = row of spinules at base of claw];
14, second gnathopod (WX); 15, palmar rnargin of second gna-
thopod (WY) [symbols as in fig. 131; 16, third pereiopod (WX);
17, basal part of fourth pereiopod (WX). Scales below fig. 19.

l o w (deeper in chappuisi, fig. 30). (6) The B-row of
Gn.1 and Gn.2 with numerous bifid spines;
spinules on the palma of gnathopod 2 is short
- sernidenticulata: basis of P1 as in aprutina;
(long in chappuisi, fig. 31). (7) Pereiopods 3 to 5
lentiform organs much larger; P3 - P6 ven/ scantily
bear a short seta (P3, P4) ar a spine (P5) in the mid-
armed; proximoventral spine of uropod 1 located
dle of the anterior margin of the merus (absent in
in the middle of the ramus; spines of outer lobe of
chappuisi, fig. 32). (8) The propodus of pereiopods
maxilla 1 with 1 denticle;
3 and 4 bears 3 shorts setae, that of P5 a spine
- vomeroi: basis of Gn.1 as in aprutina; lenti-
(absent in chappuisi, fig. 32). (9) The posterior mar-
form organs located in distal part of basis of pe-
gin of the basis of P7 b e a r ~
several spinules (only 1
reiopods; uropod l
without proximoventral spine;
i n chappuisi). (10) The propodus of P7 bears longer
antennae less slender; spines of outer lobe of max-
setae and the claw is more slender in balearica
illa 1 all pluridentate.
than in chappuisi (fig. 33). (11) The longest telson
Moreover, all 5 species listed above have only
spine is longer than the telson (shorter than the tel-
2 (instead of 3) bifid spines on the inner lobe of the
son in chappuisi).
Study of a large series (>70 specimens) of B. chap-
puisi (from interstitial waters of a gravel bank at
the mouth of La Baillorie, Banyuls, France, chlori-
Other Bogidiella material
nity 24696 mgA), has revealed a broad range of va-
riation in the expression of certain characters
from Mallorca
(number of segments in accessory flagellum of Al,
length of spines on uropod 3, slenderness of pe-
A damaged specimen of Bogidiella, probably a
reiopods, length of telson spines, number of telson
male, was collected from the underflow in the gra-
spines ... ) in what is presumed to be a monospecific
vel bed of the Torrent de Pareis, near La Calobra
population. Similar variations have been noticed
(Mallorca), at ca. 1000 m from the sea, 2 Jan. 1978,
by Karaman (1979) elsewhere in the Mediterranean
chlorinity 800 mgA (ZMA Amph. 108.099). This spec-
belt. It remains to be seen if these populations are
imen, devoid of its P6 and P7, resembles B. (B.)
indeed rnonospecific, or whether they consist of a
balearica in the slenderness of the appendages,
mixture of sibling species. At any rate, the charac-
but has the distal telson armature reduced to I + I
ters 1 t o 11 enumerated above, all fall outside the
spines. Certain other characters of this specimen
range of variation observed in the alledged B. chap-
(armature of basis of Gn.1, armature of P3 - P5) are
puisi, and of course the apomorphic sexual di-
better in agreement with B. (M,) chappuisi than
morphism in the armature of the endopodite of
with B. (B.) balearica. This specimen was briefly
uropod 2 in chappuisi forms already sufficient
mentioned by Stock, 1978: 89. Its taxonomic status
ground for placing B. balearica as a distinct species
must remain uncertain for the moment, awaiting
i n a different subgenus.
more material from this locality. It appears to be
Within the subgenus Bogidiella s. str., with
rare, or at least very localized, since repeated sam-
which balearica is to be classified, the Mallorcan
pling in 1983 and 1985 has failed to produce any
taxon is closely related to aprutina Pesce, 1980,
further specimens.
dalrnatina S. Ka ra man, 1953, niphargoides Ruffo &
Vigna, 1977, semidenticulata Mestrov, 1961, and
vomeroi Ruffo & Vigna, 1977. These species all
share the combination of the following characters
with balearica: (1) telson with II + II apical spines;
The 1985 fieldwork in Mallorca of the senior
(2) presence of lentiform organs on the pereio-
author has been supported financially by EuroUni-
pods; (3) absence of endopodite in the pleopods.
versitas, Munich (FRG).
These species can be distinguished from ba-
The 1986 collections of the junior author in
learica as follows:
Mallorca were supported by grants BSR-8215672
- aprutina: posterior margin of basis of Gn.1
and BSR-8417494 from the National Science Foun-
with 1 long and 1 short seta; lentiform organs cre-
dation (U.S.A.). We extend our appreciation to the
nulated; spines of outer lobe of maxilla 1 with 0-3
following individuals who assisted with logistical
problems and collections and provided data on
- dalmatina: basis of Gn.i as in aprutina; tel-
Mallorcan cave locations: Dr. Guillermo Mateu Ma-
son cleft narrow; telson longer than wide; telson
teu, Ana Maria Abril Duro, Joaquin Gines, and An-
spines very unequal in length; spines on outer lobe
gel Gines. We also thank the owners and manage-
of maxilla 1 with 1-2 denticles;
ment of Coves del Drac and Coves dels Hams for
niphargoides: basis of P I with 1 short and
permitting us to visit and collect from these caves.
3 long setae; telson cleft shallower; flagellum of
This paper is Contribution No. 1095 of the Ber-
first antenna 18-segmented; palmar margin of
muda Biological Station for Research.

0.1 mrn
Figs. 18-20. Bogidiella (B.) balearica Dancau, 1973, 8 (Manacor,
18, fifth pereiopod (scale WX); 19, sixth pereiopod (WX); 20, se-
venth pereiopod (WX). All scale elernents (WX, WY, WZ) corres-
pond to 100 pm.

Figs. 21-27. Bogidietla (B.) balearica Dancau, 1973, d (Manacor,

21, epimeral plates 1 to 3, from the right (scale WX); 22, second
pleopod (WX); 23, retinaculum of fint pleopod (free-hand sketch);
24, first uropod (WX); 25, second uropod (WY); 26, third uropod
(WX); 27, telson (WZ). Scales below fig. 19.

Figs. 28-34. Bogidiella (Medigidiella) chappuisi Ruffo, 1952, Q
(mouth of La Baillorie, Banyuls, France).
28, inner lobe of maxilliped (scale WZ); 29, basis of f i n t gnatho-
pod (WX); 30, palmar margin of first gnathopod (WY) [A = angle
r o w of spinules, implanted in deep sinus; B = row of spinules at
base of claw]; 31, palmar margin of second gnathopod (WY) [A
and B as in fig. 301; 32, fifth pereiopod (WX); 33, dista1 part of
seventh pereiopod (m);
34, third uropod W).
Scales below fig. 19.
DANCAU, D. (1973): «Observations sur les Amphipodes souter-
rains de I%le de Majorque. Genre Bogidiellan. Trav. Inst. Spéol.
((Deux nouveaux Amphipodes souterrains de la France .. n
«Emile Racovitza>),
12: 113-119.
C.R. Acad. Sci. Paris, 234: 1636-1638.
KARAMAN. G. S. (1979): ((Bogidiella chappuisi Ruffo 1952 and its
RUFFO, S. (1973): ((Contributo alla revisione del genere Bogidiella
variability with remarks to come other species (fam. Gamma-
Herhog (Crustacea Amphipoda, Gammaridae)>).
Boll. 1st. Ent
r i d a e ) ~ .
Poljoprivreda i Sumarstvo, 25: 17-30.
Univ. Bologna, 31: 49-77.
KARAMAN. G. S. (1982): In S. RUFFO, «The Amphipoda of the Me-
STOCK, J. H. (1978): «A remarkably variable phreatic amphipod
diterranean; Family Gammaridae~. Mém. Inst. océanogr.
from Mallorca, Rhipidogammarus variicauda n. sp.». Bijdr.
Monaco, 13: 245-364.
Dierk., 48 (1): 89-95.
KARAMAN. G. S. (1982): ((Critica1 remarks to the recent revisions
STOCK, J. H. (1981): aThe taxonomy and zoogeography of the fa-
of Bogidiella-group of genera with study of some taxa (farn.
mily Bogidiellidae (Crustacea, Amphipoda). with emphasis
Poljoprivreda i Sumantvo, 28 (3-4): 31-57.
on the West lndian taxan. Bijdr. Dierk., 51 (2): 345-374.