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29
April
1988
PROC.
BIOL.
SOC.
WASH.
101(1), 1988,
pp.
221-226
TULUMELLA UNIDENS,
A NEW GENUS AND
SPECIES OF THERMOSBAENACEAN
CRUSTACEAN FROM THE YUCATAN
PENINSULA, MEXICO
Thomas
E.
Bowman and Thomas M. Iliffe
Abstract.
-
Tulumella,
new genus, the sixth genus of Thermosbaenacea, with
type species
T. unidens,
new species, is reported from Najaron Cave, near
TulGm, Quintana Roo, Mexico. It is characterized by having small
non-func-
tional eyestalks, a scale on antenna 2, mandibles that may lack a lacinia mobilis
and have reduced molars, a biramous pereopod 1, and a reduced pleopod 2.
The family Monodellidae is recognized as valid, and a key is given to the
families and genera of Thermosbaenacea.
In November 1986 the second author
made collections in nine caves and cenotes
on the Yucatan Peninsula, Mexico, mostly
in the vicinity of the ancient Mayan city of
TulGm. From one of these caves he ob-
tained three specimens of the new
ther-
mosbaenacean described below.
Order Thermosbaenacea Monod, 1927
Family Monodellidae Taramelli, 1954
Telson separate from pleonite 6. Seven
pairs of pereopods present. Maxilliped of
6
with or without endopod.
Tulumella,
new genus
Diagnosis.
-Small eyestalks present,
without visual elements. Antenna 1 long,
with up to
18
flagellar segments. Antenna 2
with scale (exopod). Right or both mandi-
bles without lacinia mobilis; molar slender,
cylindrical; chewing surface divided into a
few slender spiniform teeth. Maxilla 2 with
2
broad basal endites armed with spoon-
shaped setae; exopod setose, inserted well
lateral to
endopod. Maxilliped with broad
endopod and exopod, both with several
marginal setae. Pereopod 1 biramous,
en-
dopod 5-segmented (including basis); pe-
reopods 2-7 with 2-segmented exopod and
6-segmented
endopod. Pleopod 1 a small
pyriform unarticulated lobe; pleopod 2 ar-
ticulated, elongate. Uropod exopod slightly
longer than
endopod; 1st segment slightly
longer than 2nd.
Type species.
-
Tulumella unidens,
new
species.
Etymology.-From
the ancient Mayan
city of Tulum, with the diminutive Latin
suffix
"ella."
Tulumella unidens,
new species
Figs. 1-2
Material.
-Mexico: Quintana Roo, near
ruins of Mayan city of TulGm:
Najaron (Na-
haron) Cave, leg. T. M. Iliffe 11 Nov 1986
(collection no.
86-106), 3 specimens: ho-
lotype, 2.9 mm, USNM 233394; paratypes,
3.0 and 1.8 mm, USNM 233395.
Etymology.
-From the Latin "uni-"
(one)
+
"dens" (tooth), referring to the lack
of a lacinia mobilis in both mandibles.
Diagnosis.
-As for the genus.
Description.
-Length up to about 3 mm
(the 3 mm
paratype is in 2 pieces; hence its
measurement is approximate). Carapace
covering pereonites 1-6. Eyestalks oval,
close together, partly covered by carapace.
222
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig.
1.
Tulumella unidens:
A, Habitus; B, Antenna
1;
C, Antenna
2;
D, Right mandible; E, Left mandible;
F,
Right mandible; G, Maxilla 1;
H,
Maxilla
2;
I, Maxilliped;
J,
Pereopod
1.
VOLUME
101,
NUMBER
1
223
Telson linguiform,
l/3
longer than wide; pos-
terior margin with medial pair of short spines
flanked by 2 pairs ofdistinctly longer spines.
Antenna 1 nearly
314
length of body. Pe-
duncle segments progressively shorter,
armed medially with long setae; segment 1
with lateral flange produced distally into
blunt process bearing 2 long setae, segment
3 with median distal process bearing 3 api-
cal setae. Outer flagellum 18-segmented,
segments 4-1 5 each with 1 or 2 long esthetes
and
1
or 2 shorter curved setae. Inner fla-
gellum about 0.7 length of outer flagellum,
14-segmented, each segment with several
distal setae of varying lengths. Antenna 2
scale about 3.5
x
as long as wide, with 10
marginal setae; flagellum about half as long
as inner flagellum of antennae 1, 10-seg-
mented.
Incisor of mandible slender with long
neck, left 6-cuspate, right 4-cuspate. Left
spine-row with
8
spines, right with 6, gaps
I
between spines decreasing toward molar.
i
Segment
2
of mandibular palp with single
row of
4
pectinate spines on distal half, seg-
1
ment
3
with double row of
3
pectinate spines
(6 in all) and pair of longer naked apical
spines.
Maxilla 1, coxal endite with 15 plumose
setae; basal endite with
9
apical spines with
denticulate medial margins; endopod (palp)
2-segmented, distal margin of 2nd segment
armed with 2 tricuspid spines, 2 spatulate
apically ciliate spines, and 1 naked seta.
Maxilla
2
coxa with row of about 22 long
setae on medial margin; coxal endite with
about
9
marginal setae; basal endites with
11 and 6 spoon-shaped setae respectively;
endopod subequal in length to basal endites
but much narrower, armed with
1
seta on
medial margin, 1 subapical seta, and 3 api-
cal setae; exopod oval, with 4 marginal
se-
tae.
Coxa of maxilliped not produced into
en-
dite; distal margin with 2 long setae reaching
distal margin of basal endite and lateral to
them a seta about
l/3
as long. Basal endite
with 13 setae on apical margin as shown in
Fig.
11 and
1
surface seta near medial mar-
gin.
Endopod a broad shallow lobe with
5
marginal setae. Exopod oval, with narrow
base and 3 setae on apical margin.
Pereopod 1 basis expanded anteriorly,
with row of long setae on anterior margin.
Ischium
3/4
length of basis, expanded ante-
riorly with
1
long seta on anterior margin.
Merus and carpus with long setae on pos-
terior (flexor) margin.
Propus broadening
distally; distal margin with 3 spiniform and
1 slender setae, largest (anterior) spiniform
seta interpreted as dactyl.
Pereopods 2-7 of uniform structure (Fig.
2A). Flexor margin of dactyl with row of
delicate peg-shaped spines on proximal half
and minutely serrate apex.
Pleopod 1 a short pyriform lobe with very
long apical seta. Right and left pleopods sep-
arated by distance equal to
2/3
length of api-
cal seta. Near lateral margin of pleonite 1,
a second pyriform lobe with
2
setae at apex,
1 on lateral margin, and 1 at base of medial
margin. Posterior margin of pleonite 1 be-
tween 2 lobes armed with 4 short setae with
swollen bases (Fig. 2C). It is not known
whether pleopod 1 is represented by the me-
dial lobe only or by both lobes plus the in-
tervening setae.
Pleopod
2
a pair of elongate rami nearly
8
x
as long as wide inserted close together
in an emarginate medial part of pleonite 2.
Apex of
ramus with apical seta more than
1.5
x
length of ramus; lateral margin with
5 setae, penultimate of which nearly as long
as
ramus.
Exopod of uropod about
l/4
longer than
endopod, 1 st segment slightly longer than
2nd. Medial margin of 1 st segment and both
margins of 2nd and of
endopod armed with
plumose setae. Distolateral corner of 1 st
segment of exopod with 3 spines increasing
in size distally and several setae as shown
in detail of Fig. 2E. Telson about 1.4 as long
as wide; posterior margin armed with 3 pairs
of spines with lengths (anterior to posterior)
2>1>3.
Comparisons.
-The presence of a scale
224
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig.
2.
Tulumella unidens:
A,
Pereopod
2;
B, Pereopod
4
dactyl; C, Pleopod
1;
D,
Pleopod
2;
E, Telson and
uropod, dorsal.
on antenna 2 and the absence of a lacinia
mobilis from both mandibles are unique
features for
Tulumella;
the other features
given in the diagnosis are shared with one
or more of the other genera. However,
un-
described species of
Tulumella
from the Ba-
hamas have a lacinia mobilis on the left
mandible
(J.
Yager, pers. comm.).
It is surprising that the family
Monodel-
lidae, proposed by Taramelli (1 954), has not
been recognized in subsequent works except
those of Barker (1959) and
McLaughlin
(1980). Indeed, the family was overlooked
by Bowman and Abele
(1
982) and Bowman
and Iliffe (1 986). In a recent list of all known
Thermosbaenacea, Stock (1 986) recognizes
only one family, Thermosbaenidae Monod,
1927. However, we are convinced that the
differences, given in the following key, are
sufficient to merit the recognition of two
families.
Key to the Families and Genera of
Thermosbaenacea
1. Telson fused with pleonite 6.
Max-
illiped of8 without endopod.
5
pairs
of pereopods
...................
.....
Thermosbaenidae Monod, 1927,
Thermosbaena
Monod, 1927
-
Telson separate from pleonite 6.
Maxilliped of
8 with or without en-
.....
dopod. 7 pairs of pereopods
Monodellidae Taramelli, 1954
... 2
2.
Eyestalks lacking. Exopod of pe-
reopod 6 and 7 l-segmented.
Pleo-
pods 1 and 2 with basal articulation,
nearly as long as their pleonites
. .
..............
Monodella
Ruffo, 1949
-
Eyestalks present. Exopod of per-
eopod 6 1
-
or 2-segmented. Pleopod
1 a small unarticulated lobe or ab-
sent
...........................
3
VOLUME 10 1, NUMBER
1
225
3.
Antenna 2 with scale. Lacinia mo-
bilis present in left mandible or
lacking in both mandibles.
Pereo-
pod 7 exopod 2-segmented
......
...............
Tulurnella,
new genus
-
Antenna 2 without scale. Lacinia
mobilis present in left mandible.
Pereopod
7
exopod 1- or 2-seg-
mented
........................
4
4.
Pereopod
7
exopod 2-segmented
. .
.............
Halosbaena
Stock, 1976
-
Pereopod 7 exopod 1 -segmented
. .
5
5. Pereopod 1
endopod ending in long
acute spine, flanked by 2 short
spines.
hagella of antenna 1 with
3
and 4 segments
.................
...........
Lirnnosbaena
Stock, 1976
Pereopod 1
endopod ending in
3
digitiform spines. Flagella of anten-
na 1 with 14 and 29 segments
....
...
Theosbaena
Cals and Boutin, 1985
Habitat.
-Najaron (Naharon) Cave is a
completely underwater limestone cenote
cave located about 8 km inland from the
Caribbean Sea on the eastern coast of the
Yucatan Peninsula (Coke
&
de Groot 1987).
The spacious underwater entrance to the
upstream, spring cave opens from the far
side of a large open
springlsiphon pool. The
walls of the cave are stained black, as are
the numerous underwater stalactites and
stalagmites. Cave passages are primarily de-
veloped at the depth of the halocline, about
-
15
m. At the halocline, a highly reactive
geochemical zone is produced by the mixing
of fresh ground water with subterranean Ca-
ribbean seawater, thus enhancing carbonate
dissolution and formation of cave passages
(Back et al. 1986). Salinities at the water
surface in the open cenote and at just above
the halocline at
-
14 m were 1.5%, while
those just below the halocline at
-
15 m and
at the bottom at
-
20 m were 32.5 and 3 5Ym,
respectively. Water temperature was 24'C
in November 1986. Water currents are lo-
calized to the upper fresher water layers in
the cave. The spring cave consists of two
main passages, each about 700 m in length.
All biological collections were made from
the Halocline System or East Side of the
cave complex. Most animals were observed
just above the halocline in oligohaline
waters. Collecting was done with a plankton
net and suction bottle from the water col-
umn in
-
10 to
-
18 m depths using scuba.
In addition to
Tulurnella unidens,
speci-
mens of copepods, amphipods, shrimp, and
remipedes
-
all still under study
-
were also
collected from the cave.
Acknowledgments
Cave collections in Yucatan by T. M.
11-
iffe were supported by National Science
Foundation Grants BSR-82 15672 and
BSR-
8417494. We thank James Coke, Dinah
Drago, Juan Jose Fucat, and Michael Mad-
den for assistance with cave diving collec-
tions and Dr. John
Markham for logistical
and collecting aid. We thank Jill Yager for
reviewing the manuscript. This publication
is Contribution No. 1 130 of the Bermuda
Biological Station for Research.
Literature Cited
Back,
W.,
B. B. Hanshaw,
J.
S. Herman,
&
J.
N. Van
Driel. 1986. Differential dissolution of a Pleis-
tocene reef in the ground-water mixing zone of
coastal Yucatan, Mexico.-Geology 14: 137-1 40.
Barker, D. 1959. The distribution and systematic po-
sition of the Thermosbaenacea.
-
Hydrobiolo-
gia 13(1-2):209-235.
Bowman, T. E.,
&
T.
M.
Iliffe. 1986. Halosbaena
fortunata, a new thermosbaenacean crustacean
from the Jameos del Agua marine lava cave,
Lanzarote, Canary
Islands.-Stygologia 2(1/2):
84-89.
,
&
L.
G. Abele. 1982. Classification of the
Recent Crustacea. Pp. 1-27
in
L.
G. Abele, ed.,
The biology of Crustacea, volume I, Academic
Press, New York.
Cals, P.,
&
C. Boutin. 1985. DCcouverte au Cam-
bodge, domaine ancien de la Tethys orientale,
d'un nouveau "fossile vivant" Theosbaena
cambodjiana n.g., asp. (Crustacea, Thermos-
baenacea).-Comptes Rendus Hebdomadaire
des
Seances de 1'AcadCmie des Sciences, Paris,
sene D, 300(8):337-340.
226
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Coke,
J.
G.,
&
J.
de Groot. 1987. Naharon.-Un-
derwater Speleology 14(1): 13-1 7.
McLaughlin,
P.
A.
1980. Comparative morphology
of Recent Crustacea. W.H. Freeman and Com-
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Monod, T. 1927.
Thermosbaena mirabilis
Monod,
remarques sur sa morphologie et sa position
sys-
tkmatique. -Faune des Colonies Frangaises 1
:
29-5
1.
Ruffo, S. 1949.
Monodella stygicola
n.g. n.sp. nuovo
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fau-
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J.
Brill/Dr.
W. Backhuys, Leiden.
Taramelli,
E.
1954.
La
posizione sistematica dei Ter-
mosbenacei quale risulta dallo studio anatomic0
di
Monodella argentarii
Stella.-Monitore
Zoologico Italiano
62(
1 ):9-27.
Crostaceo Termosbenaceo delle acque sottera-
(TEB)
D~~~~~~~~
of
~~~~~~b~~~~ zool-
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nare).-Archivio Zoologico Italiano 34:31-48.
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Stock.
J.
H.
1976.
A
new genus and two new s~ecies
NHB-163, Smithsonian Institution, Wash-
-
of the crustacean order Thermosbaenacea from
ington, D.C. 28560; (TMI) Bermuda Bio-
the West Indies.-Bijdragen tot de Dierkunde
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46(1):47-70.
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1
-
1 5, Bermuda.
.
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in
L.
Note added in proof.-Monod
&
Cals (1988: Comptes Rendus de 11Acad6mie des
Sciences, Paris 306
(SCrie III):99-108) recently rearranged the classification of the Ther-
mosbaenacea, dividing the order into two families: 1. Thermosbaenidea, with the subfam-
ilies Thermosbaeninae and Monodellinae;
2.
Halosbaenidae, new, with the subfamilies
Halosbaeninae, new, and Limnosbaenin, new. Under this scheme
Tulumella
would go
into the Halosbaeninae.