Troglobitic Ostracoda
(Myodocopa: Cypridinidae,
Thaumatocyprididae) from
Anchialine Pools on Santa
Cruz Island, Galapagos Islands
LOUIS S. KORNICKER
?
and
THOMAS M. ILIFFE
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 483
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 4 8 3
Troglobitic Ostracoda
(Myodocopa: Cypridinidae, Thaumatocyprididae)
from Anchialine Pools on
Santa Cruz Island, Galapagos Islands
Louis S. Kornicker and Thomas M. lliffe
SMITHSONIAN INSTITUTION PRESS
Washington, D.C.
1989
A B S T R A C T
Komicker, Louis S., and Thomas M. Hiffe. Troglobitic Ostracoda (Myodocopa: Cypridinidi-
dae, Thaumatocyprididae) from Anchialine Pools on Santa Cruz Island, Galapagos Islands.
Smithsonian Contributions to Zoology, number 483, 38 pages, 17 figures, 15 tables,
1989.?Two new species, Skogsbergia galapagensis and Danielopolina styx, are described
from 3 anchialine pools (grietas) in fissures on the south coast of Santa Cruz Island, Galapagos
Islands. Skogsbergia galapagensis is the first known troglobitic Cypridinidae. Danielopolina
styx is the first troglobitic Danielopolina from the Pacific, and the fifth troglobite in the genus
to be described. The ontogeny of both species is described, and the ontogeny of S. galapagensis
is compared to that of S. lerneri, an open sea, shallow water Atlantic form, and found to be
similar.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is
recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral
Montastrea cavernosa (Linnaeus).
Library of Congress Cataloging in Publication Data
Komicker, Louis S., 1919?
Troglobitic Ostracoda (Myodocopa: Cypridinidae, Thaumatocyprididae) from anchialine pools on Santa Cruz Island,
Galapagos Islands.
(Smithsonian contributions to zoology ; no. 483)
Bibliography: p.
SupL of Docs, no.: SI 1.27:483
1. Skogsbergia galapagensis?Galapagos Islands?Santa Cruz Island?Classification. 2. Danielopolina styx?
Galapagos Islands?Santa Cruz Island?Classification. 3. Cave fauna?Galapagos Islands?Santa Cruz Island?
Classification. 4. Crustacea?Galapagos Islands?Santa Cruz Island?Classification. I. Iliffe, Thomas M. II.
Title. III. Series.
QI1.S54 no. 483 591 s [595.3'3] 88-600417
[QL444.08]
Contents
Page
Introduction 1
Collecting Methods 2
Abbreviations 2
Acknowledgments 2
Zoogeography 2
Superorder MYODOCOPA Sars, 1866 2
Order MYODOCOPIDA Sars, 1866 3
Suborder MYODOCOPINA Sars, 1866 3
CYPRIDINIDAE Baird, 1850 3
CYPRIDININAE Baird, 1850 3
CYPRIDININI Baird, 1850 3
Skogsbergia Kornicker, 1974 3
Skogsbergia galapagensis, new species 3
Order HALOCYPRIDA Dana, 1853 24
Suborder HALOCYPRIDINA Dana, 1853 24
Superfamily THAUMATOCYPRIDOIDEA Muller, 1906 24
THAUMATOCYPRIDIDAE Muller, 1906 24
Key to Living Genera of Thaumatocyprididae 24
Danielopolina Kornicker and Sohn, 1976 24
Key to the Species of Danielopolina 25
Danielopolina styx, new species 25
Literature Cited 37

Troglobitic Ostracoda
(Myodocopa: Cypridinidae, Thaumatocyprididae)
from Anchialine Pools on
Santa Cruz Island, Galapagos Islands
Louis S. Kornicker and Thomas M. Iliffe
Introduction
Extensive recent faulting has occurred in the Galapagos
Islands. Among the most notable locations is the south coast
of Santa Cruz Island, where faults may be up to 20 km long
and generally trend in an east-west direction, parallel to the
coast. Some of these faults may have been formed within the
last few thousand years (McBirney and Williams, 1969).
Apparently all the faulting is normal faulting, in which one
crustal block has moved up or down with respect to its neighbor
along a nearly vertical fracture or fault plane (Simkin, 1984).
Such faulting has produced an impressive series of high cliffs
and deep fissures. Some of these fissures nearer to the coast
extend down to anchialine pools, locally referred to as
"grietas." Typically, fissures and grieta pools consist of
opposite-facing sheer rock walls 1-10 m apart and up to 30
m high. Large collapse breakdown blocks, which were
probably generated by the earthquakes forming the fissure,
make up the floor. Occasionally, talus caves occur, both above
and below water, where it is possible to enter totally dark areas
under the breakdown and between fissure walls.
Deep Grieta (Sta 87-005) is located about 250 m east of the
beach at Tortuga Bay and 80 m inland, behind a conspicuous
round lava mound. This large grieta pool runs parallel to the
coast. It is situated between 10 m high cliffs and is about 8 m
wide by 40 m long. Maximum depth in the pool is 12 m. The
rear part of the pool is roofed over with well-wedged collapse
blocks and is thus in darkness. Some root masses extend down
into the water in this section of the pool. Water temperature
measured on 3 May 1987 decreased slightly with depth in the
Louis S. Kornicker, Department of Invertebrate Zoology, National
Museum of Natural History, Smithsonian Institution, Washington,
D.C. 20560. Thomas M. Iliffe, Bermuda Biological Station for
Research, Ferry Reach GE 01, Bermuda.
pool, while salinity increased from 8?/oo to 22 %o at 12 m
(Table 1). Collections had been made previously from this
grieta by Peck and Peck (1986) and Peck and Kulalova-Peck
(1986). In addition to ostracodes, large numbers of the blind
shrimp Typhlatya galapagensis Monod and Cals, 1970, are
present in the pool, especially at the greater depths.
Grieta de Caleta la Torta (Sta 87-018) is located 7 km west
of Puerta Ayora on the south coast of Santa Cruz Island. It is
situated about 100 m inland from a small but well-protected
bay. The grieta is at the western end of a prominent 15 to 20
m high cliff and consists of 3 long pools separated by collapse
barriers. All collections were made in the first (eastern-most)
pool, 3-5 m wide by 100 m long. By passing through a narrow
hole between breakdown blocks at 12 m depth, a large chamber
was reached, which extended to 29 m depths. This chamber
contained clear water in total darkness. Water temperatures in
the pool were nearly constant throughout, while salinity
increased with depth from 9 ?/oo at the surface to 29 ?/oo at 17
m (Table 1). In addition to ostracodes, copepods, amphipods,
several T. galapagensis, and a red hippolytid shrimp (being
studied by Dr. A.J. Bruce, Darwin, Australia) were collected
in the pool.
The grieta north of the trail to Tortuga Bay (Sta 87-02IB)
is probably the same one visited by Peck and Peck (1986:43).
This grieta is located about 100 m past the last house in Puerta
Ayora on the trail to Tortuga Bay. Immediately on the north
side of this trail is a sheer-walled fissure 1 to 2 m wide by 10
m deep. About 30 m along this fissure is a 30 m long pool
divided by breakdown into 2 sections. A small electric pump
is used to take water from the pool, probably for domestic
purposes. In the second part of the pool, a deep crack extends
down underwater past a marked halocline at 9 m before ending
in collapse at 10 m depth. Moderate tidal currents are present
1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 1.?Temperature and salinity depth profiles for grietas on Santa Cruz
Island, Galapagos Islands, from which myodocopid ostracodes were collected.
Locality
Deep Grieta east of Tortuga Bay,
3 May 1987, sta 87-005
Surface
l m
3 m
12 m
Grieta de Caleta la Torta,
30 April 1987, sta 87-018
Surface
l m
3 m
17 m
Grieta north of trail to Tortuga Bay,
18 June 1987.su 87-021B
Surface
3 m
9 m
10 m
Temperature
24.8
24.2
24.0
23.1
22.8
22.8
22.8
22.3
22.7
22.7
227
22.7
Salinity
8
9
10
22
9
9
11
29
1.5
1.5
12.5
13.0
at the surface of the pool. The deepest part of the pool is in
nearly total darkness as are narrow sections of the fissure
extending through the breakdown at either end of the pools.
Water temperatures were uniform, and the salinity changed
sharply at the 9 m deep halocline from 1.5?/oo to 12.5 ?/oo
(Table 1).
COLLECTING METHODS.?Sta 87-005: Ostracodes {Daniel-
opolina styx and Skogsbergia galapagensis) were observed to
be swimming in the water column at 6-12 m depths and were
collected on 3 May 1987 by Thomas M. Iliffe and Jacqueline
Tracey with a suction bottle and a 93 \xm mesh plankton net.
Sta 87-018: Specimens of D. styx were collected with a
plankton net towed by a diver at 17 to 29 m depths; specimens
of S. galapagenis were collected 8-12 June 1987 by Thomas
M. Iliffe with plastic bottle traps (Manning, 1986) baited with
a crushed crab and left overnight at 10-18 m water depth
(most specimens came from traps at 17-18 m depths). Sta
87-021B: Specimens of S. galapagensis were collected 18-19
June 1987 by Thomas M. Iliffe in a plastic bottle trap baited
with a crushed crab and left overnight at 9-10 m water depth.
All specimens have been deposited at the National Museum
of Natural History and have been given USNM numbers.
ABBREVIATIONS.?The following abbreviations are used in
the illustrations: ant., antenna; B.O., Bellonci organ; endop.,
endopodite; exop., exopodite; u.p., unpaired process.
ACKNOWLEDGMENTS.?Collection of specimens in the
Galapagos was supported by National Science Foundation
grant BSR-8215672 to T.M. Iliffe. We thank Jacqueline Tracey
for assistance with collections. Logistical assistance in the
Galapagos was provided by the Charles Darwin Research
Station and the Galapagos National Park Service. We also
thank Thomas E. Bowman, Smithsonian Institution, and Anne
C. Cohen, Los Angeles Museum of Natural History, for
commenting on the manuscript, Jack R. Schroeder, Schroeder
Prints, Inc., Crisfield, Maryland, for inking illustrations of
appendages, Molly Ryan, Smithsonian Institution, for render-
ing the carapace of D. styx, Elizabeth Harrison-Nelson,
Smithsonian Institution, for general technical assistance, and
Don Fisher, Smithsonian Institution Press, for final editing and
preparation of the manuscript for publication. This paper is
contribution No. 1140 of the Bermuda Biological Station for
Research.
Zoogeography
Danielopolina styx is the second example of a troglobitic
species from the Galapagos Islands whose congeners are
otherwise restricted to the Atlantic Ocean. Atlantic species of
Danielopolina are found in caves in the Bahamas, Canary
Islands, Cuba, and Mexico (Yucatan Peninsula) as well as from
a depth of 3459 m in the South Atlantic (Kornicker and Iliffe,
in press a). The other troglobite example from the Galapagos
is Typhlatya galapagensis, a caridean shrimp. The genus
Typhlataya is represented by 8 other species inhabiting caves
and anchialine pools in Ascension Island, Barbuda, Bermuda,
Caicos Islands, Cuba, Dominican Republic, Mexico (Cam-
peche and Yucatan Peninsula), and Puerto Rico (Mona Island).
It seems likely that the ancestors of both Danielopolina and
Typhlatya in the Galapagos reached the Pacific prior to the
closing of the Panama land bridge in the Pliocene. Indeed,
Danielopolina and 6 other troglobitic invertebrate genera
having an amphi-Atlantic distribution may have colonized
caves at an early stage in the formation of the Atlantic, perhaps
during the Mesozoic, and were subsequently dispersed on
plates through sea floor spreading (Iliffe et al., 1984; Hart et
al., 1985; Wagele, 1985; Wilkens et al., 1986). The ancestors
of Danielopolina have been reported from the Permian of Idhra
Island, Greece, and the Lower and Upper Jurassic of Germany
(Kornicker and Sohn, 1976). Thus, Danielopolina is an
apparently ancient form having its origins in the Tethys Sea,
and dispersal of its ancestors was greatly influenced by plate
movement The present distribution of the genus in marine
caves and in the deep sea suggests that the descendants of an
initially shelf form in the Mesozoic later found refuge in caves
and the deep sea; possibly shelf forms could not survive
increased competition from abundant new post-tethyan shelf
taxa. However, other scenarios that cannot be excluded have
been suggested for cave and deep sea invertebrates by others
(Iliffe et al., 1983; Stock, 1986).
Superorder MYODOCOPA Sars, 1866
COMPOSITION.?The Myodocopa comprise 2 orders:
Myodocopida Sars, 1866, and Halocyprida Dana, 1853, both
with representatives in the anchialine caves of the Galapagos
Islands.
NUMBER 483
Order MYODOCOPIDA Sars, 1866
COMPOSITION.?This order contains only the suborder
Myodocopina Sars, 1866.
Suborder MYODOCOPINA Sars, 1866
COMPOSITION.?This suborder comprises 5 families of
which 4, Philomedidae, Sarsiellidae, Ruddermatidae, and
Cylindroleberididae, have been reported from anchialine caves
in Bermuda (Kornicker and Iliffe, in press b). All of the
Bermudan cave myodocopine ostracodes have eyes similar to
the same species, or to closely related species, collected in
shallow waters of the open sea, and it was concluded by
Kornicker and Iliffe (in press b) that none were troglobites.
The troglobitic myodocopine described herein is a member of
the Cypridinidae, a family not collected in Bermudan caves,
nor in open water outside the caves (the latter is possibly
because of insufficient collecting).
CYPRIDINIDAE Baird, 1850
COMPOSITION.?The Cypridinidae comprise 2 subfamilies
of which none has been reported previously from anchialine
caves, but only in the open sea, at shallow to abyssal depths.
Only the Cypridininae is represented in the present collection.
CYPRIDININAE Baird, 1850
COMPOSITION.?The Cypridininae comprise 21 genera in 2
tribes, Cypridinini and Gigantocypridinini, some widespread,
others with limited distribution.
CYPRIDININI Baird, 1850
COMPOSITION.?The Cypridinini comprise 20 genera, of
which only Skogsbergia is represented in the anchialine caves
of the Galapagos Islands.
Skogsbergia Kornicker, 1974
Skogsbergia Poulsen, 1962:162 [nomen nudum].?Komicker, 1974:3 [selected
type species].
TYPE SPECIES.?Skogsbergia minuta Poulsen, 1962, by
Kornicker (1974:3).
COMPOSITION AND DISTRIBUTION.?Ten species are referred
to Skogsbergia, including the new species described herein
(Table 2). Species of the genus are widespread between
latitudes of 60?N and 34?S, and except for 5. galapagensis,
live in the open sea, most at shelf depths.
Skogsbergia galapagensis, new species
FIGURES 1-10
ETYMOLOGY.?From the collecting area.
MATERIAL.?Holotype: Sta 87-018: USNM 193420, 1
adult female in alcohol and on slide.
Allotype: Sta 87-005: USNM 193423, 1 adult male in
alcohol and on slide.
Paratypes: Sta 87-005: USNM 193434, 8 specimens in
alcohol. Sta 87-018: USNM 193422, 4 adult females in
alcohol; USNM 193424, 1 instar I in alcohol and on slide;
USNM 193425, 1 instar II in alcohol and on slide; USNM
193426, 1 instar III in alcohol and on slide; USNM 193427,1
male instar IV in alcohol and on slide; USNM 193428, 1
female instar IV in alcohol; USNM 193429, 1 male instar V
in alcohol and on slide; USNM 193430, 1 female instar V in
alcohol; USNM 193431 1 adult female in alcohol; USNM
193432, 18 specimens in alcohol. Sta 87-021B: USNM
193433, 6 specimens in alcohol.
DISTRIBUTION.?Santa Cruz Island, Galapagos Islands:
Deep Grieta east of Tortuga Bay at 6-12 m water depth; Grieta
de Caleta la Torta at 10-18 m water depth (type locality);
grieta north of trail to Tortuga Bay at 9-10 m water depth.
DESCRIPTION OF ADULT FEMALE (Figures l-4c).?
Carapace oval in lateral view with small incisur at midheight
and narrow but distinct caudal process (Figure la); ventral and
dorsal margins convex; tip of rostrum with minute denticula-
tions (Figure \c). Outer surface of valves smooth.
Infold (Figure Ib-e): Rostral infold with 1 short and 4
long divided hirsute bristles in addition to 2 divided bristles (1
long, 1 short) near inner edge of incisur (Figure \b,c); 1 small
slender divided bristle just ventral to inner end of incisur,
anteroventral infold with 28-34 spinous divided hirsute
brisdes forming row just distal to narrow list (Figure \b)\
ventral infold with 3-6 widely separated short bristles along
narrow list; left valve only with bristle at midwidth of ventral
infold just anterior to caudal process; list broadening anterior
to caudal process and forming inner margin of infold, with 3
or 4 small bristles and numerous pores perpendicular to list
and leading to its posterior edge (Figure \d,e). 7-10 minute
pores forming row along outer edge of infold of caudal process;
2 or 3 minute pores near outer edge of caudal process visible
through infold and leading to slender bristle on outer surface
of valve (Figure \d,e). Right valve with sclerotized bar along
inner edge of posterior infold dorsal to caudal process (Figure
le).
Selvage (Figure Ib): Selvage without fringe. Anterodorsal
selvage along rostrum narrow, broadening and becoming
narrowly striate along dorsal and ventral edges of incisur,
dividing at inner end of incisur, narrow along anteroventral and
ventral margins of valve, and absent along outer edge of caudal
process and posterior edge of valve.
Size: USNM 193420, length 2.09 mm, height 1.36 mm;
USNM 193431, length 2.02 mm, height 1.35 mm. USNM
193422,4 specimens: length 2.07 mm, height 1.38 mm; length
2.10 mm, height 1.35 mm; length 2.14 mm, height 1.40 mm;
length 2.04 mm, height 1.41 mm.
First Antenna (Figures \f,g, 3g): 1st joint bare. 2nd joint
spinous. 3rd joint short with medial spines forming rows and
2 short spinous bristles (1 ventral, 1 dorsal). 4th joint with 2
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 1.?Skogsbergia galapagensis, new species, USNM 193420, holotype, adult female: a, complete
specimen from right side, length 2.09 mm, area of central adductor muscle attachments stippled, inner margin
of infold of caudal process as viewed through shell dashed; b, inside view of anterior of left valve; c, detail from
b; d, inside view of caudal process of left valve; e, dorsal end of caudal process of right valve; / , left 1 st antenna,
medial view, only base of sensory bristle of 5th joint shown; g, tip of left 1st antenna showing filaments of
sensory bristle of 5th joint; A, tip of 7th limb; i, lateral eye.
NUMBER 483
FIGURE 2.?Skogsbergia galapagensis, new species, USNM 193420, holotype, adult female: a, distal part of
protopodite and endopodite of right 2nd antenna, medial view; b, joints 2-9 of exopodite of right 2nd antenna,
lateral view, all bristles not shown; c, marginal hairs near midlcngth of bristle of 3rd exopodial joint of right
2nd antenna, ventral edge on top; d, right mandible, medial view; e, base of medial claw and short ventral bristle
on 3rd joint of mandible shown in d;f, left maxilla, lateral view, bristles of endopodial joints not shown, g-j,
right maxilla: g, bristles of 1 st and 2nd endopodial joints, lateral view, only bases of a -bristles shown; h, a-bristles
of 2nd endopodial joint; i, lateral view, neither endites nor all bristles shown; j , medial view of exopodite.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
short spinous bristles (1 ventral, 1 dorsal). Stout sensory bristle
of 5th joint with 9 long filaments followed by 3 short filaments
and 1 smaller filament (long filaments with minute terminal
papillae; short filaments and tip of bristle with minute terminal
spine) (Figure Ig). 6th joint with short medial bristle near
dorsal margin. 7th joint: a-bristle about same length as bristle
of 6th joint, with few marginal spines; b-bristle about 4 times
length of a-bristle, with 3 short filaments followed by 2 longer
filaments (filaments with few marginal spines; bristle and
filaments with terminal spine); c-bristle about lh longer than
sensory bristle of 5th joint, with 8 short filaments followed by
1 longer filament and 1 very small subterminal filament (some
filaments with short marginal spines; filaments and bristle with
terminal spine). 8th joint: d- and e-bristles about same length
as b-bristle, with blunt tips; f-bristle long (tip broken off on
holotype), with 7 marginal filaments, some with marginal
spines); g-bristle long (tip broken off on holotype) with 10
marginal filaments, some with marginal spines.
Second Antenna (Figure 2a-c): Protopodite with short
spinous distal medial bristle. Endopodite 2-jointed: 1st joint
with 3 bare proximal bristles (1 long, 2 short) and 1 spinous
distal bristle (about same length as long proximal bristle); 2nd
joint short, with long terminal filament with blunt tip (Figure
2a). Exopodite (Figure 2b,c): bristle of 2nd joint reaching 9th
joint, ventral margin of bristle with about 6 stout spines (3rd
or 4th spine from proximal end stouter) followed by 2 slender
spines and stout subterminal spine; dorsal margin of bristle
with few slender proximal spines; tip of bristle with narrowly
ringed segment; bristles of joints 3 and 4 with slender proximal
ventral spines and broad spine-like ventral hairs at midlength
followed by slender natatory hairs (Figure 2c); bristles of joints
5-8 with natatory hairs; 9th joint with 4 bristles (3 long with
natatory hairs, 1 small with few short hairs); joints 3-8 with
slender sharply pointed basal spines increasing in length on
distal joints (spine of 8th joint reaching past distal margin of
9th joint) (Figure 2b); 9th joint with small lateral spine near
dorsal margin.
Mandible (Figure 2d,e): Coxale endite spinous with 2 stout
terminal prongs with small process between them; minute
bristle near base of endite. Basale: ventral margin with 3 small
a-bristles (distal unringed), 1 small lateral b-bristle, 2 spinous
c-bristles, 1 long spinous d-bristle, and 1 minute medial bristle
just proximal to c-bristles; dorsal margin with 1 bristle (with
few spines) at midlength and 2 subterminal (bare or with few
spines). Exopodite about 3A length dorsal margin of 1st
endopodial joint, hirsute distally, with 2 subterminal bristles
(proximal bristle spinous, about twice length distal bristle). 1st
endopodial joint with 4 ventral bristles (2 long, 1 short, 1
minute). 2nd endopodial joint: ventral margin with bristles
forming 3 groups comprising 1,1, and 2 short slender bristles
(lateral bristle of distal pair shorter than medial bristle); dorsal
margin and medial surface near dorsal margin with about 14
short bristles (not all shown on illustrated limb), 1 or 2 slightly
longer bristles, and 6 long bristles (some with bases on lateral
side); medial surface with spines forming rows. 3rd endopodial
joint with 3 short claws (medial and lateral claws with stout
proximal spines (Figure 2e); shorter dorsal claw bare) and 4
bristles (ventral bristle short; no bristles swollen near base).
Maxilla (Figure 2/-;): Endites I?III with about 10, 7, and
6 bristles, respectively; 1 short bristle near base of endite III
(on basale?) (Figure 2/). Precoxale and coxale with dorsal
fringe of long hairs; coxale with hirsute dorsal bristle with long
proximal and short distal hairs (Figure 2i). Basale with 3 distal
bristles: 1 short dorsal, 2 ventral (1 medial short, 1 long lateral).
Exopodite with short proximal bristle with base close to 2
terminal bristles (proximal bristle with long proximal hairs;
outer terminal bristle with long hairs followed by shorter hairs)
(Figure 2f). 1st endopodial joint with 2 bare alpha-bristles, 2
beta-bristles (outer pectinate, inner with few hairs), and
smoothly rounded terminal tooth on inner margin (Figure 2g).
2nd endopodial joint with 4 a-bristles with few proximal spines
(Figure 2h), 3 or 4 pectinate b-bristles (presence of 4th b-bristle
on right limb probably aberrancy), 2 pectinate c-bristles, and
3 stout pectinate d-bristles (Figure 2g).
Fifth Limb (Figure 3a-c,f): Sclerotized anterior process
of protopodite elongate, undulate, with smoothly rounded tip
(Figure 3b). Endites I and II each with 5 or 6 bristles; endite
III with 6 or 7 bristles. 1st exopodial joint: anterior side with
1 proximal bristle (with long proximal and short distal spines)
near process of protopodite and 3 distal bristles forming row
(outer bristle stout with long proximal spines and pectinate
distally; inner bristles with long proximal and short distal
spines); main tooth comprising triangular peg followed by 6
pectinate teeth; bristle with long proximal hairs and short distal
spines present proximal to peg of main tooth. 2nd exopodial
joint with 4 stout pectinate unringed a-bristles, 6 pectinate
b-bristles (ringed distally), c-bristle with long proximal and
short distal spines, and long anterior d-bristle with long
proximal and short distal spines. 3rd exopodial joint with 3
bristles on inner lobe and 2 on outer lobe (on left limb of
USNM 193420, both bristles of outer lobe with long proximal
hairs, on right limb only outer bristle with long hairs); fused
4th and 5th joints with 4 or 5 bristles.
Sixth Limb (Figure 3d,e): Endites with total of 16-19
bristles: endite I with 3 or 4 bristles (1 or 2 short proximal, 2
longer terminal); endite II with 3 or 4 bristles (1 or 2 short
medial proximal, 2 terminal); endite III with 4 or 5 bristles (0
or 1 medial proximal, 4 terminal); endite IV with 5-7 bristles
(1 medial proximal, 4-6 terminal). End joint with 13-15
bristles (posterior 2 plumose, others with long proximal hairs
and short distal spines, or only short spines). Stem with 3 or 4
bare epipodial bristles.
Seventh Limb (Figure 1 h): Each limb with 18 or 19 bristles
(12 in terminal group, 6 on each side, and 6 or 7 in proximal
group, 3 or 4 on each side), each with up to 7 bells (most with
4 bells, 2 with only 1 bell). Terminal comb with 11 teeth (3
long teeth in middle, and on each side 4 shorter teeth with
rounded tips and 2 short teeth with square tips). Surface
NUMBER 483
FIGURE 3.?Skogsbergia galapagensis, new species, USNM 193420, holotype, adult female: a, left 5th limb,
posterior view; b, right 5th limb, anterior view, not all bristles shown; c, endites of right 5th limb, medial view
(limb not under cover slip); d, right 6th limb, medial view; e, endiles III and IV of left 6th limb, medial view;
/ , left lamella of furca, anterior process of protopodite of 5th limb, and location of 6th limb; g, anterior of body
showing 1st and 2nd joints of left 1st antenna, Bellonci organ attached to medial eye, and anterior process; h,
posterior of body from left side showing genitalia (stippled) and Y-sclerite (lined).
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
opposite comb with 4 minute teeth (2 on each side; only closer
2 shown on Figure \h).
Furca (Figure 3/): Each lamella with 10 claws; claw 2
broader distally than claw 1 but about same width at base; claw
5 slightly broader than claw 4 but about same length, and
narrower as well as shorter than claw 3; claw 2 with weak basal
suture defined best at anterior and posterior ends; claw 1 with
slender teeth along posterior edge and stouter medial teeth
forming row with stouter teeth at claw midlength; remaining
claws with slender teeth along posterior margin forming lateral
and medial row; right lamella slightly anterior to left and with
few indistinct spines along anterior margin.
Bellonci Organ (Figure 3g): Elongate with tapering tip.
Eyes: Medial eye small, unpigmented (Figure 3g). Lateral
eye indistinct, unpigmented, minute, with globular cells but
no ommatidia (Figure \i).
Upper Lip (Figure Aa,b): Anterior part undivided and with
glandular openings along both anterior and ventral margins;
posterior part paired and with ventral glandular openings; 2
small lobes with few glandular openings present lateral to
posterior end of each paired part; posterior of lip rounded
hirsute.
Genitalia (Figure 3h): Comprising ring on each side of
body.
Anterior of Body (Figures 3g, 4a): Single sclerotized
process with pointed tip present between upper lip and medial
eye.
Posterior of Body (Figure 4c): Bare and without dorsal
process. Posterodorsal margin dorsal to sclerotized internal
girdle slightly scalloped in lateral view with indentations
caused by banded muscles (not segmented).
Y-Sclerite (Figure 3h): Typical for subfamily.
Eggs: No ovigerous females in collection.
Gut Content: 5 specimens (USNM 193420, 193422) with
gut gorged with fine grained unidentified substance appearing
white in reflected light and light amber in transmitted light.
DESCRIPTION OF ADULT MALE (Figure 4d-k).?Carapace
similar to that of adult female but smaller, and rostrum more
rounded (Figure Ad,e).
Infold (Figure 4fg): Bristles of rostrum and caudal process
similar to those of adult female. Anteroventral infold with 24
bristles forming row.
Size: USNM 193423, length 1.76 mm, height 1.10 mm.
First Antenna (Figure 4h,i): Joints 1-6 similar to joints
of adult female except for smaller ventral bristle on 4th joint.
7th joint a-bristle small, similar to that of adult female;
b-bristle about 7 times length of a-bristle, not as long as sensory
bristle of 5th joint, with bulbous base bearing both short
filament (with large transparent sucker at midlength and small
subterminal node) and stem with 2 slender marginal filaments
(proximal with 6, distal with 8 minute suckers), followed by 2
slender marginal filaments (each with about 4 minute marginal
spines), and minute terminal papilla; c-bristle almost twice
length of b-bristle, with bulbous base bearing both short
filament (with large transparent sucker at midlength and small
subterminal node) and stem with 2 slender marginal filaments
(each with 8 minute suckers), followed by 6 long slender
marginal filaments (some with few marginal spines) and short
subterminal filament; tip of stem blunt. 8th joint: d- and
e-bristles slightly shorter than b-bristle, bare with blunt tips;
f-and g-bristles about same length as c-bristle, both with many
long proximal hair-like filaments followed by about 7 stouter
filaments and terminal papilla.
Second Antenna: Protopodite with short spinous distal
medial bristle. Endopodite 2-jointed (similar to that of adult
female): 1st joint with 3 proximal bristles (1 long, 2 short) and
1 spinous distal bristle (slightly shorter than long proximal
bristle); 2nd joint small, with long terminal filament. Exopodite
without medial terminal bristle on 1st joint (also absent on
adult female); bristle of 2nd joint reaching just past 9th joint,
ventral margin with few long proximal hairs followed by 4
stout spines, 2 slender spines, 1 stout subterminal spine, and
terminal papilla; bristles of 3rd and 4th joints with broad flat
spear-like ventral hairs proximal to midlength (similar to those
of same bristles of female); remaining characters similar to
those of adult female.
Mandible: b-bristle of basale absent on left limb of USNM
193423, but present on right limb (possibly, absence of
b-bristle on left limb aberrant); dorsal margin of 2nd
endopodial joint with 21 bristles (14 spinous bristles forming
medial row along edge of joint, 2 longer spinous bristles (about
twice length short bristles) lateral to short bristles, and 6 long
bristles (some with few spines) along edge of joint or on lateral
side near dorsal margin; limb otherwise similar to that of adult
female.
Maxilla: Similar to that of adult female. 2nd endopodial
joint with 4 a-bristles, 3 b-bristles, 2 c-bristles, and 3 d-brisdes.
Fifth Limb: Epipodite with 53 plumose bristles. Endite I
with 6 bristles, endite II with 5 bristles, endite IN with 7
bristles. Exopodite similar to that of adult female.
Sixth Limb: Epipodite with 4 bristles. End joint with 12
bristles. Endite bristles not counted.
Seventh Limb: Left limb of USNM 193423 with 23 bristles
FIGURE 4.?Skogsbergia galapagensis, new species, USNM 193420, holotype,
adult female: a, anterior of body from right side showing anterior process at
top and upper lip at bottom; b, upper and lower lips from left side and
esophagus (dashed); c, posterior of body showing scalloped posterodorsal
margin dorsal to stippled girdle. USNM 193423, allotype, adult male: d,
complete specimen from left side, length 1.76 mm, central adductor muscle
attachments indicated by dashed oval, inner margin of infold of caudal process
as viewed through shell dashed; e, outside view of anterior of right valve; /,g,
inside view of caudal process of left and right valves, respectivly, not all bristles
of list shown; h, left 1st antenna, medial view, all bristles of 7th and 8th joints
not shown; i, proximal suckers of b- and c-bristles and proximal part of g-bristle
of left 1st antenna, medial view; j , posterior of body showing claws 1-3 of
right lamella of furca, right copulatory structures, and internal sclerites; k,
anterior of body from left side showing left lateral eye, medial eye and Bellonci
organ, anterior process at midheight, and upper lip (note 4 lateral glandular
processes).
NUMBER 483
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
(15 in terminal group, 7 or 8 on each side, and 8 in proximal
group, 3 on comb side, 5 on opposite side); bells on bristles
similar to those of adult female. Comb and surface opposite
comb similar to adult female.
Furca (Figure 4/): Similar to that of adult female except
claws appearing slightly more slender.
Bellonci Organ (Figure 4k), Posterior of Body, Y-
Sclerite: Similar to those of adult female.
Eyes: Medial eye small unpigmented (Figure 4k). Lateral
eye indistinct, transparent, with small amber-colored cells
(Figure 4k).
Upper Lip (Figure 4k): Four small glandular openings
forming row proximal to small posterior lateral lobe; lip
otherwise similar to that of adult female.
Genitalia (Figure 4/): Copulatory organ well developed,
lobate.
Anterior of Body (Figure 4k): Single process with 2 small
triangular processes on tip.
Gut Content: Gut with unrecognizable minute light
amber-colored particles.
DESCRIPTION OF INSTAR I (Sex unknown) (Figures 5-6c).?
Carapace similar to that of adult female except being more
elongate and with relatively broader caudal process (Figure
5a). Tip of rostrum digitate as on adult female. Narrow curved
line (ridge?) extending from upper edge of inner end of incisur
and intersecting anteroventral valve edge as on adult female.
Infold: Infold of rostrum similar to that of adult female,
with 4 long bristles, 1 short bristle near inner end of incisur,
and paired bristles near valve edge at inner end of incisur. Inner
edge of infold and list well defined, similar to those of adult
female. Anteroventral infold with 1 bristle near inner end of
incisur and 11 bristles forming row just distal to list. Broad
list of caudal process similar to that of adult female. Ventral
infold with 2 bristles (1 near middle, 1 near caudal process).
Right valve with bar dorsal to list of caudal process similar to
that of adult female.
Selvage: With lamellar prolongation similar to that of adult
female.
Size: USNM 193424, length 0.78 mm, height 0.53 mm.
First Antenna (Figure 5b,c): 1st joint bare. 2nd joint with
distal medial and lateral spines. 3rd joint with 2 short bristles
(1 ventral, 1 dorsal). 4th joint bare. Sensory bristle of 5th joint
without marginal filaments and with minute terminal papilla.
6th joint with short medial bristle with few marginal spines.
7th joint: a-bristle about same length as bristle of 6th joint,
with few marginal spines; b-bristle bare, about 4 times length
of a-bristle; c-bristle long, bare. 8th joint d- and e-bristles
longer than b-bristle, bare with blunt tips; f-bristle long, bare,
bent dorsally; g-bristle long bare.
Second Antenna (Figure 5c,d): Protopodite with well-
developed distal medial bristle. Endopodite 2-jointed (Figure
5d): 1st joint bare; 2nd joint about same length as 1st joint,
with long terminal filament. Exopodite: joints 3-8 with minute
spines forming row along distal margins; bristle of 2nd joint
reaching midlength of 8th joint, with few ventral spines; 9th
joint with 2 bristles, limb otherwise similar to that of adult
female.
Mandible: Coxale endite spinous similar to that of adult,
with small bristle near base. Ventral margin of basale with 2
a-bristles (proximal about half length distal), no b-bristle, 2
c-bristles, and no d-bristle; dorsal margin with 1 midbristle and
2 subterminal bristles. Exopodite almost reaching distal margin
of 1st endopodial joint, with 2 bristles (proximal about 4 times
length of distal). 1st endopodial joint with 2 long ventral
bristles. Ventral margin of 2nd endopodial joint with distal
spines and only 1 bristle (distance of base of bristle from distal
margin of joint equal to width of joint); dorsal margin with 5
bristles (3 long, 2 short). End joint with 3 claws (lateral and
medial stout and with proximal spines, dorsal more slender and
bare, all about same length) and 1 ventral bristle about same
length as claws.
Maxilla (Figure 5e,f): Endites I?III each with 5 spinous
bristles (Figure 5e). Coxale with long slender dorsal bristle
(relatively longer than coxale bristle of adult). Basale with long
terminal ventral bristle. 1st endopodial joint with 1 slender
alpha-bristle and 1 pectinate beta-bristle. 2nd endopodial joint
with 2 a-bristles (with few proximal spines), 1 pectinate
b-bristle, and 2 pectinate claw-like d-bristles; cutting tooth
with 2 prongs (Figure 5/). Exopodite similar to that of adult.
Fifth Limb (Figure 5g,h): Sclerotized anterior process of
protopodite very small or absent. Endites I?III with 2, 3, and
4 bristles, respectively; posterior bristle of each endite minute,
bare, others spinous. 1st exopodial joint: anterior side with
stout proximal bristle with long proximal spines and pectinate
distally; main tooth with triangular peg followed by 1 complex
pectinate tooth; spinous posterior bristle proximal to teeth. 2nd
exopodial joint with 1 stout pectinate a-bristle (ringed distally),
1 long b-, or d-bristle (with long proximal spines and pectinate
distally; distal teeth stronger than on d-bristle of later instars)
on proximal outer edge of joint, and c-bristle with long
proximal spines. 3rd exopodial joint: inner lobe not well
defined, without bristles, outer lobe hirsute, with 2 bristles
with long proximal hairs; fused 4th and 5th exopodial joints
hirsute, with 2 terminal bristles (posterior with short marginal
spines, anterior with long proximal and short distal spines).
Epipodite with about 30 plumose bristles.
Sixth Limb (Figure 5i): Flap-like, hirsute, without bristles.
Seventh Limb: Absent.
Furca (Figure 6a-c): Each lamella with 5 claws; claws 2-5
without basal suture, but narrow ledge along inner edge of
lamellar medial to claws 3-5 appears to be basal suture when
viewed in transmitted light; claws 3 and 4 small, straight, with
broad spinous base; claw 5 small blunt with terminal spines
(Figure 6c); claws 1 and 2 with spines forming medial and
lateral rows along posterior margins (distal medial spines of
claw 1 larger); minute spines forming row along lamella
posterior to claw 5; right lamella slightly anterior to left.
Bellonci Organ: Elongate, similar to that of adult female
NUMBER 483 11
FIGURE 5.?Skogsbergia galapagensis, new species, USNM 193424, paratype,
instar I (sex unknown): a, complete specimen from right side, length 0.78
mm, central adductor muscle attachments indicated by dashed oval, inner
margin of infold of caudal process as viewed through shell dashed; b, tip of
right 1st antenna, lateral view, only proximal part of f-, c-, and g-bristles
shown; c, anterior part of body from left side showing location of lateral eye,
1st antenna, and protopodite of 2nd antenna; J. distal part of prolopodite and
endopodite of left 2nd antenna, medial view; e, endites of right maxilla, medial
view;/, left maxilla, lateral view, endites not shown; g, right 5th limb, anterior
view, not all bristles shown; h, left 5th limb, posterior view; i, right 6th limb,
anterior to right; /, anterior of body from right side showing anterior process
at upper right and part of upper lip at bottom.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 6.?Skogsbergia galapagensis, new species, USNM 193424, paratype, instar I (sex unknown): a,
posterior of body from left side showing hirsute dorsal process, left lamella of furca, and fused Y-sclerite and
girdle; b, right lamella of furca; c, detail from b. USNM 193425, paratype, instar II (sex unknown): d, complete
specimen from right side, length 0.92 mm, central adductor muscle attachments indicated by dashed circle, inner
margin of infold of caudal process as viewed through shell dashed, e, anterior of body from left side showing
location of lateral eye, 1 st antenna, and anterior process; /, distal part of protopodite and endopodite of right 2nd
antenna, medial view; g, tip of left maxilla, medial view, a-bristles not shown; h, left 5th limb, posterior view,
endues I and II not shown; i, right 6th limb, anterior to right; j , right 7th limb; k, posterior of body from right
side showing posterodorsal hairs, right lamella of furca, and internal sclerites; /, right lateral eye.
NUMBER 483 13
but filled with cells or globules.
Eyes: Medial eye small unpigmented. Lateral eye elongate,
indistinct, filled with cells or globules (Figure 5c).
Upper Lip: Similar to that of adult female.
Anterior of Body (Figure 5/): Anterior process between
medial eye and upper lip with 3 small processes.
Posterior of Body (Figure 6a): Posterodorsal corner with
small process with few long hairs.
Y-Sclerite (Figure 6a): Posterior end fused to other
sclerites; anterior end without ventral branch.
Gut Content: USNM 193424 with mostly unidentifiable
particles but also with few fragments bearing claws suggesting
arthropod origin.
DESCRIPTION OF INSTAR II (Sex unknown) (Figure 6d-l).?
Carapace similar to that of adult female except caudal process
relatively broader (Figure 6d). Tip of rostrum with minute
single process (not digitate as on adult and instar I).
Infold: Anteroventral infold with 13 bristles. Bristles of
infold of rostrum, ventral margin, and caudal process similar
in number to those of adult female.
Selvage: Lamellar prolongation similar to that of adult
female.
Size: USNM 193425, length 0.92 mm, height 0.58 mm.
First Antenna (Figure 6e): 1st joint bare. 2nd joint with
distal medial spines. 3rd joint with medial spines and 2 bristles
(1 ventral, 1 dorsal). 4th joint with 1 dorsal bristle; sensory
bristle of 5th joint with 3 long stout proximal filiments
followed by 1 short slender filament and 1 smaller subterminal
filament. 6th joint with short medial bristle. 7th joint a-bristle
slightly shorter than bristle of 6th joint; b-bristle about 3 times
length of a-bristle, with 1 proximal filament; c-bristle long,
with 4 long filaments followed by 1 short subterminal filament.
8th joint: d- and e-bristles about 1/3 longer than b-bristle, bare
with blunt tips; f-bristle long, with 3 long filaments followed
by 1 short subterminal bristle; g-bristle long, with 4 long
filaments followed by 1 short subterminal filament.
Second Antenna (Figure 6/): Protopodite with medial
distal bristle with short marginal spines. Endopodite 2-jointed:
1st joint with fairly long distal bristle; short 2nd joint with
long terminal filament. Exopodite: bristle of 2nd joint reaching
8th joint, with about 7 ventral spines (proximal 3 stouter than
others; subterminal spine long slender); 9th joint with 3 bristles
(1 long with natatory hairs, 1 short and 1 minute, both with
short marginal hairs); bristles of other joints and basal spines
similar to those of adult female.
Mandible: Coxale endite similar to that of adult Basale
with 3 a-bristles, 1 b-bristle, 2 c-bristles, 1 minute medial
bristle just proximal to c-bristles, and no d-bristle; dorsal
margin of basale with 3 bristles (1 near midlength, 2
subterminal). 1st endopodial joint with 3 ventral bristles. 2nd
endopodial joint ventral margin with spines, and bristles
forming 2 groups (1 in proximal group, 2 in distal group);
dorsal margin with 7 bristles (3 long, 4 medium or short). End
joint with 3 claws (2 stouter claws with ventral spines) and 3
bristles.
Maxilla (Figure 6g): Coxale with long slender dorsal
bristle. Endite I with 7 bristles, endites II and III each with
about 6 bristles; 1 short bristle on basale some distance from
proximal end of endite III (an equivalent bristle on adult female
is close to endite III). Basale with additional long ventral bristle
near ventral margin and no dorsal bristle (proximal ventral
corner obscured on both limbs of USNM 193424, and long
ventral bristle could be on endite III). 1st endopodial joint with
I slender alpha-bristle and 1 pectinate beta-bristle; cutting tooth
with 2 prongs. 2nd endopodial joint with 3 a-bristles (not
shown), 2 b-bristles, 1 c-bristle, and 3 d-bristles; exopodite
similar to that of adult female.
Fifth Limb (Figure 6/i): Sclerotized anterior process of
protopodite elongate with smoothly rounded tip. Endites I and
II each with 5 bristles; endite III with 6 bristles. 1st exopodial
joint: anterior side with 2 distal bristles forming row (outer
bristle stouter, with long proximal hairs and pectinate distally;
inner bristle with long proximal and short distal hairs); main
tooth of 1st exopodial joint with 1 triangular peg and 2
pectinate teeth; spinous bristle proximal to peg. 2nd exopodial
joint with 3 pectinate a-bristles (proximal ringed distally), 4
pectinate b-bristles (ringed distally), c-bristle with long
proximal and short distal spines, and anterior d-bristle with
long proximal and short distal spines. Inner and outer lobes of
3rd exopodial joint each with 2 bristles; fused 4th and 5th
joints with 2 bristles; outer lobe of 3rd joint and fused 4th and
5th joints hirsute.
Sixth Limb (Figure 60: Single endite with 1 long bristle.
Seventh Limb (Figure 6j): Short bare thumb-like.
Furca (Figure 6k): With 5 claws; claw 2 without basal
suture; claw 3 narrower and longer than claw 4.
Bellonci Organ: Similar to that of adult female.
Eyes: Medial eye small unpigmented. Lateral eye small
with few light amber-colored cells, no ommatidia (Figure 6e,l).
Upper Lip: Similar to that of adult female.
Anterior of Body (Figure 6c): With anterior process similar
to that of adult female.
Posterior of Body (Figure 6k): With small dorsal process
with few long hairs similar to that on instar I.
Y-Sclerite (Figure 6k): Posterior end not fused to other
sclerites as on instar I; anterior end without ventral branch.
Gut Content: Unidentifiable minute particles amber in
transmitted light
DESCRIPTION OF INSTAR III (Sex unknown) (Figure la-g).?
Carapace similar to that of adult female (Figure 7a). Tip of
rostrum minutely digitate.
Infold: Anteroventral infold with 15 bristles just distal to
list; infold otherwise similar to that of adult female.
Selvage: Lamellar prolongation similar to that of adult
female.
Size: USNM 193426, length 1.19 mm, height 0.70 mm.
First Antenna (Figure lb,g): Joints 1-3 similar to those
of instar II. 4th joint with 2 bristles (1 ventral, 1 dorsal).
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 7.?Skogsbergia galapagensis, new species, USNM 193426, paratype, instar III (sex unknown): a,
complete specimen from left side, length 1.19 mm, central adductor muscle attachments indicated by dashed
oval, inner margin of infold of caudal process as viewed through shell dashed; b, anterior of body from left side
showing location of 1 st antenna, protopodite of 2nd antenna, and lateral eye; c, endopodite of left 2nd antenna,
lateral view; d, end joint of endopodite of right maxilla, a-bristles not shown, medial view; e, left 7th limb,
anterior to left; / , posterior of body from left side showing hirsute posterodorsal process, left lamella of furca,
and internal sclentes (Y-sclerite, girdle); g, anterior of body from right side showing 1st joint of 1st antenna,
Bellonci organ, and anterior process. USNM 193427, paratype, male instar IV: h, complete specimen from right
side, length 1.30 mm, central adductor muscle attachments indicated by dashed oval, inner margin of infold of
caudal process as viewed through shell dashed; i, endopodite of right 2nd antenna, medial view; j , anterior of
body from left side showing joints 1 and 2 of left 1st antenna, protopodite of 2nd antenna, and lateral eye; k,
part of left maxilla showing dorsal bristles of coxale and basale; /, posterior of body showing right lamella of
furca, right copulatory organ, and internal sclentes; in, anterior of body from right side showing joints 1 and 2
of right 1 st antenna, Bellonci organ, lateral eye, and upper lip.
NUMBER 483 15
Sensory bristle of 5th joint with 4 long stout filaments followed
by 2 short slender filaments and smaller subterminal filament.
6th joint with short medial bristle. 7th joint: a-bristle spinous,
about same length as bristle of 6th joint; b-bristle 4 or 5 times
length of a-bristle, with 2 short marginal filaments (1 proximal,
1 near midlength); c-bristle long, with 4 marginal filaments
followed by 1 long filament and 1 short subterminal filament.
8th joint d- and e-bristles bare with blunt tips; f-bristle long,
tip broken off on specimen examined, with 5 filaments on
remaining part, some with marginal spines; g-bristle long, with
5 marginal filaments followed by 1 longer filament and short
subterminal filament.
Second Antenna (Figure lb,c): Protopodite with distal
medial bristle. Endopodite 2-jointed (Figure 7c): 1st joint with
2 bristles; 2nd joint with long terminal filament. Exopodite:
bristle of 2nd joint reaching 7th joint; ventral margin of bristle
of 2nd joint with few slender marginal spines followed by 4
stout spines followed by 4 or 5 slender spines (distal spine
longer); 9th joint with 3 bristles (1 long with natatory hairs, 1
about 1/2 length of long bristle, also with natatory hairs, 1 short
bare, or with few short hairs); other bristles and joints similar
to those of adult female.
Mandible: Coxale endite, basale, exopodite, similar to
those of adult female. 1st endopodial joint with 3 bristles (2
long, 1 short). 2nd endopodial joint ventral margin spinous,
with bristles forming 2 groups (1 in proximal group, 2 in distal
group); dorsal margin with 7 bristles (4 long, 1 medium, 2
short). End joint with 3 claws and 3 bristles.
Maxilla: Endite I with 9 bristles; endite II with 6 bristles,
endite III with 4 terminal bristles; 1 short bristle near base of
endite III. Coxale with fringe of dorsal hairs; dorsal bristle not
observed. Basale with 1 long bristle near ventral margin and 1
short dorsal bristle. Exopodite similar to that of adult female.
1st endopodial joint with 2 slender alpha-bristles and 2
beta-bristles (inner short, outer long pectinate), and poorly
defined cutting tooth. 2nd endopodial joint with 3 slender
a-bristles, 2 pectinate b-bristles (proximal ringed), 2 pectinate
c-bristles, and 3 pectinate d-bristles (Figure Id).
Fifth Limb: Sclerotized anterior process of protopodite
elongate with smoothly rounded tip. Endite I with 6 bristles;
endite II with 5 bristles; endite III with 7 bristles. 1st exopodial
joint: anterior side with 1 proximal bristle near process of
protopodite and 3 distal bristles forming row (outer bristle stout
with long proximal spines and pectinate distally); main tooth
comprising triangular peg and 3 pectinate teeth; bristle present
proximal to peg. 2nd exopodial joint with 3 stout pectinate
unringed a-bristles, 6 pectinate b-bristles (ringed distally),
c-bristle with long proximal and short distal spines, and
anterior d-bristle (with long proximal and short distal spines)
proximal to 3rd joint. 3rd exopodial joint with 3 bristles on
inner lobe and 2 on outer lobe. Fused 4th and 5th joints with
3 bristles.
Sixth Limb: Endite I with 1 long spinous bristle; endite II
with 4 spinous bristles; endites III and IV each with 3 spinous
bristles. End joint spinous, with 2 anterior bristles with long
proximal and short distal spines, and 2 stouter posterior
plumose bristles; 1 epipodial bristle present.
Seventh Limb (Figure le): Elongate, bare.
Furca (Figure If): Each lamella with 7 claws; claw 2 with
weak basal suture; claw 5 stouter and longer than claw 4.
Bellonci Organ (Figure Ig): Similar to that of adult female.
Eyes: Medial eye small, unpigmented. Lateral eye small
with several minute amber-colored cells (Figure 7b).
Upper Lip, Y-Sclerite (Figure If): Similar to those of adult
female.
Anterior of Body (Figure Ig): Anterior process similar to
that of adult female.
Posterior of Body (Figure If): With dorsal process bearing
long hairs.
Gut Content: Gut gorged with unidentifiable particles
amber-colored in transmitted light.
DESCRIPTION OF MALE INSTAR IV (Figure lh-m).?
Carapace similar in shape to that of adult female (Figure Ih).
Infold: Bristles of rostrum and caudal process similar to
those of adult female. 1 small bristle just ventral to inner end
of incisur. Anteroventral infold with 18-21 bristles forming
row.
Size: USNM 193427, length 1.30 mm, height 0.81 mm.
First Antenna (Figure lj,m): Joints 1-4 similar to those of
instar III. Sensory bristle of 5th joint with 5 long stout filaments
followed by 3 short slender filaments and smaller subterminal
filament Sixth joint with short medial bristle. 7th joint:
a-bristle about same length as bristle of 6th joint; b-bristle
about 5 times length of a-bristle, with 3 marginal filaments;
c-bristle long, with 6 short filaments followed by 1 long
filament and short subterminal filament 8th joint d- and
e-bristles bare with blunt tips; f-and g-bristles long, with about
5 short filaments followed by 1 long filament and 1 short
subterminal filament.
Second Antenna (Figure li,j): Protopodite with distal
medial bristle. Endopodite 2-jointed: 1st joint with 3 bristles;
2nd joint with long terminal filament. Exopodite: bristle of 2nd
joint reaching 8th joint, with 8 ventral spines (proximal 4 stout,
distal slender spine longer than other slender spines); 9th joint
with 3 bristles (middle bristle about twice length of dorsal
bristle; ventral bristle almost twice length middle bristle, all
bristles with natatory hairs); remaining bristles and joints
similar to those of adult female.
Mandible: Coxale, endite, basale, and exopodite similar
to those of adult female. 1 st endopodial joint with 3 or 4 bristles
(2 long, 1 or 2 small). 2nd endopodial joint: ventral margin
with bristles forming 3 groups comprising 1, 1, and 2 short
slender bristles; dorsal margin with 9 bristles (4 long, 2
medium, 3 short). End joint with 3 claws and 4 bristles (ventral
bristle minute).
Maxilla (Figure Ik): Coxale and precoxale with dorsal
fringe; coxale with very long slender dorsal bristle. Endite I
with 10 bristles, endites II and III each with 6 bristles; 1 short
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
bristle near base of endite III (on basale?). Basale with 3 distal
bristles (1 short dorsal, 1 long ventral, 1 short medial near
ventral margin). Exopodite similar to that of adult female. 1st
endopodial joint with 2 slender alpha-bristles and 2 beta-
bristles (outer longer and pectinate), and evenly rounded
cutting tooth. 2nd endopodial joint with 4 bare a-bristles, 3
pectinate b-bristles, 2 pectinate c-bristles, and 3 pectinate
d-bristles.
Fifth Limb: Sclerotized anterior process of protopodite
similar to that of adult female. Endites I and II each with 5
brisdes; endite III with 7 bristles. 1st exopodial joint: anterior
side with 1 proximal bristle near process of protopodite and 3
distal bristles forming row (outer bristle stout and with long
proximal and short distal hairs); main tooth with smooth
triangular peg and 4 pectinate teeth; spinous bristle present
proximal to peg. 2nd exopodial joint with 3 pectinate a-bristles,
6 pectinate b-bristles, 1 posterior c-bristle, and 1 anterior
d-bristle (proximal to 3rd exopodial joint). 3rd exopodial joint
with 3 bristles on inner lobe and 2 on hirsute outer lobe. Fused
4th and 5th joints hirsute, with 4 bristles.
Sixth Limb: Endite I with 2 bristles; endite II with 3
bristles; endite IN with 4 bristles; endite IV with 6 bristles. End
joint with 5 anterior bristles followed by space and 3 or 4
bristles along edge of posterior half of joint (1 or 2 posterior
bristles plumose, 2 anterior bristles short and with short
marginal spines, others with long proximal and short distal
spines). 2 epipodial bristles present
Seventh Limb: Bristles strongly tapered. Each limb with 5
or 6 bristles (4 in terminal group, 2 on each side, and 1 or 2
in proximal group, 1 on comb side, 0 or 1 on process side);
each bristle with 1 bell. Terminal comb with 7 teeth (3 long
teeth in middle, and 2 short teeth on each side). Surface
opposite comb with small triangular process.
Furca (Figure It): Each lamella with 9 claws; claw 2 with
weak basal suture; claw S broader than claw 4.
Bellonci Organ (Figure 7m): Similar to that of adult
female.
Eyes: Medial eye small unpigmented. Lateral eye small
with 3-5 cells (Figure 7/,m).
Upper Lip (Figure 7m), Y-Sclerite (Figure 70: Similar to
adult female.
Genitalia (Figure 70: Well-defined copulatory organ on
each side of body anterior to furca.
Anterior of Body (Figure 7m): With anterior process
similar to that of adult female.
Posterior of Body: Small dorsal process, but without hairs
that were present in younger instars.
Gut Content: Gut gorged with unidentifiable paniculate
matter amber-colored in transmitted light.
DESCRIPTION OF FEMALE INSTAR IV (Figure Sa,b).?
Carapace similar in shape to that of male instar IV (Figure 8a).
Infold: Bristles similar to those of adult female; anteroven-
tral infold with 18 bristles forming row.
Selvage: Similar to that of adult female.
Central Adductor Muscle Attachments: Comprising both
elongate and oval individual scars (not all muscles shown in
Figure Sb.)
Size: USNM 193428, length 1.39 mm, height 0.88 mm.
First Antenna (Figure Sb): Joints 1-4, and 6 similar to
those of male instar IV. Sensory bristle of 5th joint with 5 long
stout filaments followed by 3 short slender filaments and
smaller subterminal filament. 7th joint: a-bristle about same
length as bristle of 6th joint; b-bristle with 3 marginal
filaments; c-bristle long, with 5 short filaments followed by 1
long filament and short subterminal filament. 8th joint: d- and
e-bristles bare with blunt tips; f-bristle long, with about 5
marginal filaments and shorter subterminal filament; g-bristle
long, with about 8 marginal filaments and short subterminal
filament.
Second Antenna (Figure Sb), Mandible: Similar to those
of male instar IV.
Maxilla, Fifth and Sixth Limbs: Not examined in detail
but seemingly similar to those of instar IV male.
Seventh Limb: Proximal group with 2 bristles, 1 on each
side; terminal group with 4 bristles, 2 on each side. Terminal
comb with 2 short teeth on each side of 3 longer teeth; side
opposite comb with small triangular process.
Furca: Each lamella with 8 claws; claw 2 with weak basal
suture; claw 5 broader than claw 4 but about same length.
Bellonci Organ, Upper Lip, V-Sclerite: Similar to those
of adult female.
Eyes: Lateral eye small, unpigmented, with few minute
amber-colored cells (Figure Sb). Medial eye small, unpigmen-
ted.
Genitalia: None observed.
Anterior of Body: Anterior process similar to that of adult
female.
Posterior of Body: Posterodorsal corner without tuft of
hairs.
Gut Content: Gut gorged with unidentifiable fine-grained
particles, amber-colored in transmitted light.
DESCRIPTION OF INSTAR V MALE (Figure Sd-g).?Carapace
similar in shape to that of adult female (Figure Sd).
Infold: Bristles of rostrum and caudal process similar to
those of adult female. Anteroventral infold with 23-27 bristles.
Selvage: Similar to that of adult female.
Size: USNM 193429, length 1.83 mm, height 1.16 mm.
First Antenna (Figure 8/): Joints 1-4 and 6 similar to
those of adult female. Sensory bristle of 5th joint with 7 long
stout filaments followed by 3 short slender filaments and 1
smaller subterminal filament. 7th joint: a-bristle about same
length as bristle of 6th joint; b-bristle short, with 3 marginal
filaments; c-bristle long, with 5 short slender filaments
followed by 1 long slender filament and 1 short subterminal
filament. 8th joint: d- and e-bristles bare with blunt tips;
f-bristle long, with 6 marginal filaments followed by 1 shorter
subterminal filament; g-bristle long, with about 6 short
NUMBER 483 17
FIGURE 8.?Skogsbergia galapagensis, new species, USNM 193428, paratype, female instar IV: a, complete
specimen from left side, length 1.39 mm, central adductor muscle attachments indicated by dashed oval, inner
margin of infold of caudal process as viewed through shell dashed; b, anterior of body from left side showing
joints 1 and 2 of left 1st antenna, protopodite of 2nd antenna, lateral eye, and some central adductor muscle
attachments. USNM 193430, paratype, female instar V: c, complete specimen from right side, length 1.75 mm,
central adductor muscle attachments indicated by dashed oval, inner margin of infold of caudal process as viewed
through shell dashed. USNM 193429, paratype, male instar V: d, complete specimen from right side, length
1.83 mm, central adductor muscle attachments indicated by dashed oval, inner margin of infold of caudal process
as viewed through shell dashed; e, posterior of body showing right lamella of furca;/, anterior of body showing
1st joint of left 1st antenna and lateral eye; g, right copulatory organ.
filaments followed by I shorter subterminal filament.
Second Antenna: Same number of bristles as that of adult
female.
Mandible: Coxale, basale, 1st endopodial joints similar to
those of adult female. 2nd endopodial joint: bristles of ventral
margin similar to those of adult female; dorsal margin with
10 bristles (5 long, 2 medium, 3 short). End joint similar to
that of adult female.
Maxilla: Coxale with dorsal bristle. Endite I with about
10 bristles, endite II with about 7 bristles, endite III with 6
bristles; 1 short bristle near base of endite III. Basale with 3
distal bristles: 1 short dorsal, 2 ventral (1 medial short, 1 lateral,
long). Bristles of exopodite and 1st endopodial joint similar
to those of adult female; cutting tooth of 1st endopodial joint
evenly rounded; 2nd exopodial joint with 4 a-bristles, 3
b-bristles, 2 c-bristles, and 3 d-bristles.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 2.?Comparison of carapace, furca, and lateral eyes of species of Skogsbergia (nd = no data).
Species
S. costal Kornicker, 1974
S. curvata Poulsen, 1962
S. galapagensis, new species
S. hesperida (Miiller, 1906)
5. lerneri (Kornicker, 1958)
5? megalops (Sars, 1872)
S. menezi Kornicker, 1970
S. minuta Poulsen, 1962
S? tarsi (Muller, 1912)
S. squamosa (Muller, 1894)
Carapace length (mm)
female
3.14-3.9
1.70-1.80
2.04-2.14
2.00-2.12
1.40-2.11
3
1.61
0.93-1.20
2.62
3.3
male
2.7
1.50-1.70
1.76
nd
1.50-1.76
nd
1.49
1.13
nd
2.6
Caudal
process
narrow
narrow
narrow
long
narrow
narrow
narrow
narrow
pointed
narrow
Furcal claws
total
10-11
8
10
9-10
7-9
11
8
8
9-10
7
fused*
2
2
2t
0
2
nd
2
2
0?
2,3
Lateral
eyes
large
large
vestigia]
large
large
large
large
large
large
large
?The number of the claw or claws fused to lamella.
tCkw with weak suture at base.
Note: Date for S. hesperida from Muller, 1906, and Poulsen, 1962; S. lerneri from Cohen, 1983, and Kornicker,
1984; S? sarsi from Sars, 1888, and Muller, 1912; S. squamosa from Muller, 1894, and Kornicker, 1974;
remaining species from original descriptions.
Fifth Limb: Sclerotized anterior process of protopodite
similar to that of adult female. Endite I with 5 bristles; endite
II with 6 bristles; endite III with 7 bristles. 1st exopodial joint:
anterior side with 4 anterior bristles similar to those of adult
female; main tooth comprising triangular peg followed by 5
pectinate teeth; bristle with long proximal and short distal
spines present proximal to peg of main tooth. 2nd exopodial
joint with 4 stout pectinate a-bristles, 5 pectinate b-bristles,
c-bristle with long proximal and short distal spines, and
anterior d-bristle with long proximal and short distal spines.
3rd exopodial joint with 3 bristles on inner lobe and 1-2 on
outer lobe; fused 4th and 5th joints hirsute, with 5 bristles.
Sixth Limb: Endites with total of 18-20 bristles: endite I
with 3-4 bristles (2 short proximal, 1 or 2 long terminal);
endite II with 4 bristles (2 short, medial, proximal, 1 long and
1 short, terminal); endite III with S bristles (1 medial, proximal,
4 terminal); endite IV with 6 or 7 bristles (1 short, proximal,
5 or 6 terminal). End joint with 10 bristles (posterior 2 plumose,
others with long proximal hairs and short distal spines, or only
short spines. Stem with 2 or 3 epipodial bristles.
Seventh Limb: Limb with 18 tapered bristles (terminal
group with 7 on comb side and S on opposite side; proximal
group with 3 on comb side, 4 on opposite side); 4 or 5 proximal
bristles with only 1 bell, others with 3; several terminal bristles
with 1 bell, others with 3 or 4 bells. Surface opposite comb
not examined in detail, but of similar type to that of adult
female.
Furca (Figure 8e): Each limb with 9 claws; claw 2 with
weak basal suture; claw 5 stouter than claw 4 but about same
length.
Bellonci Organ, Upper Lip, Anterior and Posterior of Body,
Y-Sclerite: Similar to those of adult female.
Eyes: Lateral eye (Figure 8/) and medial eye similar to
that of adult female.
Genitalia (Figure 8g): Small copulatory organ on each
side (not identified with certainty).
Gut Content: Gut gorged with unidentifiable particles
amber-colored in transmitted light.
DESCRIPTION OF FEMALE INSTAR V (Figure 8c).?Carapace
similar in shape to that of adult female (Figure 8c).
Size: USNM 193430, length 1.75 mm, height 1.18 mm.
Seventh Limb: Bristles tapered, many with only 1 bell.
Furca: Each lamella with 9 claws; claw 2 of left lamella
with weak basal suture, of right lamella with strong suture
(aberration?). Claw 5 stouter than claw 4 but about same length.
Gut Content: Gut gorged with unidentifiable particles
amber-colored in transmitted light.
COMPARISONS.?Skogsbergia galapagensis differs from
previously described species referred to Skogsbergia in lacking
well-developed lateral eyes. Some important morphological
characters of the species of Skogsbergia are compared in Table
2. Two species that according to Poulsen (1962:162) probably
should be referred to Skogsbergia are included in the table
with a question, because they are insufficiently known.
REMARKS CONCERNING SEVENTH LIMB.?The 7th limbs of
the preserved specimens vary in length and degree of curl.
Some adult females and juveniles have fairly short 7th limbs
having either curled (Figure 9c,d) or fairly straight tips. At least
1 adult female and 1 juvenile were observed to have 7th limbs
about 1.5 times the length of the short type, and their tips were
uncurled (Figure 9b). The single adult male in the collection
has a 7th limb about 1.6 times the length of the limb of adult
females having short 7th limbs, and the tip of the male limb
is straight (Figure 9a). The different types of 7th limbs are
shown in Figure 9 (limbs drawn at same magnification; only
proximal bristles indicated on limbs). The coils are on the
posterior side of the limb (clockwise when viewed from left
side and counterclockwise when viewed from right side).
NUMBER OF GROWTH STAGES.?Skogsbergia galapagensis
has 5 juvenile instars, the same number reported for S. lerneri
NUMBER 483 19
FIGURE 9.?Relative total length and degree of curling of 7th limbs of
Skogsbergia galapagensis, new species: a, long limb of adult male, USNM
193423 (allotype); b. long limb of adult female, USNM 193431 (paratype); c4,
short curled limbs of adult females, USNM 193432 (paratype) and 193420
(holotype), respectively.
by Cohen (1983:250); all growth stages of S. galapagensis are
present in the collection. Five juvenile instars have been
reported for 3 additional species in 3 other genera of
Cypridinini: Doloria pectinata Skogsberg, 1920, by Kornicker
(1969, table 1), Macrocypridina castanea Skogsberg, 1920,
by Poulsen (1962:126), and Vargula hilgendorfii (Miiller,
1908), by Hiruta (1980:145). No members of the Cypridinini
have been reported with other than 5 juvenile growth stages.
This suggests that the number of juvenile instars is constant
in the Cypridinini, and that the troglobitic habit of S.
galapagensis has not affected the number of growth stages.
ONTOGENY AND SEXUAL DIMORPHISM.?Except for the
caudal process being slightly broader, the shape of the juvenile
carapace is similar to that of the adult The adult carapace bears
an unusually low number of bristles on the rostral infold, only
7, and that number is present on all instars including the 1st.
The number of bristles on the anteroventral infold increases
from 11 on instar I to 24-34 on the adult (Table 9). The shape
of the adult male carapace is similar to that of the adult female
but is smaller (Table 10).
The order of appearance of appendages is similar to that
reported for many myodocopids (Table 3): the anterior 5 limbs
are already well developed in the 1st instar; in instar I the 6th
limb appears as an anlage without bristles and the 7th limb is
TABLE 3.?Order of appearance of appendages of Skogsbergia galapagensis
(x = present; 0 = absent; anlage = appendage present but without bristles).
Growth
stage
I
II
III
rv
V
Adult
1st and 2nd antennae, mandible, maxilla,
5th limb, furca, Bellonci organ
X
X
X
X
X
X
6th limb
anlage
X
X
X
X
X
7th limb
0
anlage
anlage
X
X
X
absent; the latter appears as a small bare bud in instar II and
is longer but remains bare in instar III; bristles appear on the
6th limb in instar II and on the 7th limb in instar IV. The
copulatory limb of the male was observed to appear in instar
IV, but the female genitalia was observed only in the adult.
First Antenna (Table 4): All instars have no bristles on
joints 1 and 2, 2 bristles (1 ventral, 1 dorsal) on joint 3, 1
sensory bristle on joint 5,1 bristle (medial) on joint 6,3 bristles
on joint 7, and 4 bristles on joint 8. The 4th joint bears no
bristles in instar 1,1 dorsal bristle in instar II, and 2 bristles (1
ventral, 1 dorsal) in later instars. The sensory bristle of the 5th
joint is without sensory filaments in instar I, and the number
of filaments gradually increase to 13 in the adult (Table 5).
The b- and c-bristles of the 7th joint, and the f- and g-bristles
of the 8th joint are without filaments on instar I, and have
filaments in later instars. The adult male has suckers on the
b-and c-bristles of the 7th joint, and abundant hair-like
proximal filaments on the f- and g-bristles of the 8th joint
Second Antenna (Table 5): In all instars the protopodite
bears a single medial bristle, and exopodial joints 2-8 each
bear a single bristle, of which those on joints 3-8 are natatory.
The endopodite is 2-jointed on all instars, but the 2nd joint is
longer on younger instars, and bears a terminal filament in all
instars; the 1st joint is without bristles in instar I, and the
number of bristles increases gradually to 4 in instar V. The
number of bristles on the 9th exopodial joint is 2 in instar I,
and the number increases gradually to 4 in instar V. The
appendage is similar in males and females.
Mandible (Table 6): The coxale endite is similar and the
dorsal margin of the basale bears 3 bristles in all instars. The
ventral margin of the basale bears 2 a-bristles in instar I and 3
in later instars, and 2 c-bristles in all instars; the lateral b-bristle
and a small medial bristle proximal to the c-bristles is absent
in instar I but present in other instars; the d-bristle appears first
in instar III. The 1st endopodial joint bears 2 ventral bristles
in instar I, and the number increases gradually to 4 in instar
IV or V. The ventral margin of the 2nd endopodial joint bears
1 bristle in instar I, and this increases to 3 in instars II and III,
and to 4 in later instars. The dorsal margin has 5 bristles in
instar I, and this number increases gradually to 10 in instar V,
then jumps to about 22 in the adult The end joint bears 3 claws
in all instars but the number of bristles increases from 1 in
instar I to 4 in instar IV. The appendage is similar in males and
females.
Maxilla (Table 7): Endite I bears 5 bristles in instar I, and
the number increases gradually to 10 in instar IV; endite II
bears 5 bristles in instar I, 6 in instars II-IV, and 7 in later
instars; instar III bears 5 bristles in instar I and 6 in later instars.
The exopodites are similar in all instars. The basale bears 1
bristle in instar I, and the number increases gradually to 4 in
instar IV. The 1st endopodial joint bears 1 alpha-bristle and 1
beta-bristle in instar I, and increases to 2 in instar III and later
instars; the cutting tooth bears 2 prongs in instars I and II and
is evenly rounded in instars III or IV and later instars. The 2nd
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 4.?Number of bristles on joints of 1st antenna of Skogsbergia galapagensis.
Growth
stage
I
II
III
rv
V
Adult
Female
Male
1st
joint
0
0
0
0
0
0
0
2nd
joint
0
0
0
0
0
0
0
3rd joint
ventral
1
1
1
1
1
1
1
dorsal
1
1
1
1
1
1
1
4th joint
ventral
0
0
1
1
1
1
1
dorsal
0
1
1
1
1
1
1
5th
joint
1*
1
1
1
1
1
1
6th
joint
1
1
1
1
1
1
1
7th
joint
3*
3
3
3
3
3
3t
8th
joint
4*
4
4
4
4
4
4*
*Bristles without marginal filaments.
tb- and c-bristles with suckers.
$f- and g-bristles with abundant long hair-like proximal filaments.
TABLE 5.?Distribution of filaments on the sensory bristle of the 5th joint of
the 1st antenna, and number of bristles on the endopodite and 9th joint of the
exopodite of the 2nd antenna of Skogsbergia galapagensis.
Growth
i
i i
in
IV female
IVmale
Vmale
Adult
female
male
Filaments of sensory
bristle of 1 st antenna
long proximal
0
3
4
5
5
7
9
9
short distal
0
2
3
4
4
4
4
4
2nd antenna
Endopodite
1st joint
0
1
2
3
3
4
4
4
Exopodite
9th joint
2
3
3
3
3
4
4
4
TABLE 6.?Distribution of bristles and claws on joints of mandible of Skogsbergia
galapagensis.
Growth
stage
I
II
III
IVmale
Vmale
Adult
female
male
a
2
3
3
3
3
3
3
Basale (ventral)*
b
0
1
1
1
1
1
0-1
c
2
2
2
2
2
2
2
d
0
0
1
1
1
1
1
1st joint
2
3
3
3-4
4
4
4
Endopodite
2nd joint
ventral
1
3
3
4
4
4
4
dorsal
5
7
7
9
10
22
22
3rd joint
4
6
6
7
7
7
7
?Letters a-d represent a-, b-, c-, d-bristles.
NUMBER 483 21
TABLE 7.?Distribution of bristles on maxilla of Skogsbergia galapagensis.
Growth
stage
I
II
III
IVmale
Vmale
Adult
female
male
I
5
7
9
10
10
10
10
Endites
II
5
6
6
6
7
7
7
III
5
6
6
6
6
6
6
Coxale
1
1
0
1
1
1
1
Basale
1
2*
3*
4*
4*
4*
4*
Exopodite
3
3
3
3
3
3
3
1st joint
alpha
1
1
2
2
2
2
2
beta
1
1
2
2
2
2
2
Endopodite
a
2
3
3
4
4
4
4
2nd joint
b
1
2
2
3
3
3
3
c
0
1
2
2
2
2
2
d
2
3
3
3
3
3
3
?Includes 1 short bristle close to base of endite III.
TABLE 8.?Distribution of bristles on the 5th limb of Skogsbergia galapagensis (ant. = anterior).
Growth
stage
I
II
III
IVmale
Vmale
Adult
female
male
I
2
5
6
5
5
6
6
Endites
II
3
5
5
5
6
6
5
III
4
6
7
7
7
7
7
ant.
1
2
4
4
4
4
4
1st
main tooth
peg + 1 tooth
peg + 2 teeth
peg + 3 teeth
peg + 4 teeth
peg + 5 teeth
peg + 6 teeth
peg + 6 teeth
a
1
3
3
3
4
4
4
Exopodial joints
2nd*
b
0
4
6
6
5
6
6
c
1
1
1
1
1
1
1
d
1
1
1
1
1
1
1
inner
0
2
3
3
3
3
3
3rdt
outer
2
2
2
2
1-2
2
2
4th+5th
2
2
3
4
5
4-5
4
?Letters a-d represent a-, b-, c-, and d-bristles.
tBristles on inner and outer lobes.
TABLE 9.?Distribution of bristles on 6th and 7th limbs of Skogsbergia
galapagensis, and number of bristles forming row along anteroventral infold
of carapace (nd = no data).
Growth
stage
I
II
III
IVmale
Vmale
Adult
female
male
Endites
0
1
11
15
18-20
16-19
nd
6th limb
End joint
0
0
4
8-9
10
13-15
12
Epipodite
0
0
1
2
2-3
3-4
4
7th limb
absent
0
0
5-6*
18*
18-19
24
Carapace
11
13
15
18-21
23-27
28-34
"Bristles tapering distally.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
cndopodial joint bears 5 bristles in instar I, and the number
increases gradually to 12 in instar V. The appendage is similar
on males and females.
Fifth Limb (Table 8): Endites I-111 bear a total of 9 bristles
in instar I, 16 bristles in instar II, and 17 to 19 in later instars.
The number of anterior bristles on the 1st exopodial joint
increases from 1 in instar I to 4 in instar III and later instars;
the main tooth bears a proximal smooth triangular peg in all
instars, 1 pectinate tooth in instar I, then the number of
pectinate teeth increases by 1 tooth in each later instar (adult
with 6 pectinate teeth); a spinous bristle is present on all instars
proximal to the triangular peg. The 2nd exopodial joint bears
3 bristles in instar I, and the number increases gradually to 12
in the adult. The outer lobe of the 3rd exopodial joint bears 2
bristles in all instars, whereas the inner lobe bears no bristles
in instar I, 2 in instar II and 3 in later instars. The fused 4th
and 5th exopodial joints bear 2 bristles in instars I and II, 3 in
instar III, and 4 or 5 in later instars. The appendage is similar
in males and females.
Sixth Limb (Table 9): The limb in instar I is flap-like with
marginal hairs but no bristles. In instar 11 it bears a single bristle
on an endite. The number of endite bristles increases to 11 in
instar III and 18-20 in instar V. The number of bristles on the
end joint increases from 4 in instar III to 12-15 in the adult.
The number of epipodial bristles increases from 1 in instar III
to 3 or 4 in the adult. The appendage is similar in males and
females.
Seventh Limb (Table 9): The limb is absent in instar I,
minute in instar II, elongate but bare in instar III, elongate with
tapering bristles in instars IV (5 or 6 bristles) and V (18
bristles). The adult bears 18-24 cylindrical bristles. The distal
part of the limb is straight in the adult male, a small number
of adult females, and a small number of instars IV and V, and
is curled as well as being shorter in others. The adult male may
have more bristles than the adult female (24 compared to 18
or 19, but more specimens would have to be examined to rule
out variability).
Furca (Table 10): Claw 1 has a distinct basal sutures in
all instars. Claw 2 is either without a basal suture (instar I) or
has a weakly developed suture (instar II to adult). Except in
instar I remaining claws have a distinct basal suture; in instar
I claws 3-5 are without a basal suture. The number of claws
increases from 5 in instar I to 10 in the adult (instars I and II
have 5 claws, instar III 7 claws, instar IV 8 or 9 claws, instar
V 9 claws, and the adult 10 claws). The furca is similar in
males and females.
Bellonci Organ: Similar in all instars and in males and
females.
Eyes: The medial eye is similar in all instars and in males
and females. The lateral eye is vestigial in all instars, but is
relatively slightly larger in instar I.
Upper Lip: Similar in all instars and in the adult male and
female, except 4 posterior, lateral, glandular processes
observed on adult male, but not on others.
Genitalia: A reduced male copulatory organ is present in
instars IV and V, but female genitalia were observed only in
the adult
Anterior of Body: The anterior process has a pointed tip
in juvenile instars and in the adult female but the tip bears 2
triangular processes in the male; however, as only 1 male was
examined, the difference could be due to intraspecific
variability.
Posterior of Body: The posterodorsal corner bears a tuft
of long hairs in instars I?III, absent in later instars.
Y-Sclerite: The posterior end of the sclerite is fused to
other sclerites in instar I but is separated from them in later
instars. The distal ventral branch is absent in instars I and II
and present in later instars.
Gut Content: The gut of the only 1st instar examined
(xl500 magnification) contained small fragments bearing
claws suggesting arthropod origin; no recognizable fragments
were observed in later instars, but as these were examined at
lower magnification (x300) it is possible that minute fragments
were not resolved.
COMPARATIVE ONTOGENY.?A reason for the detailed study
of the ontogeny of S. galapagensis herein is to compare the
ontogeny of a troglobitic species with that of an open water
species of Skogsbergia. Skogsbergia lerneri (Kornicker, 1958)
was recently redescribed by Kornicker (1984:14), and its
ontogeny studied by Cohen (1983). It is a species that has been
collected in the Gulf of Mexico and Caribbean Sea (Kornicker,
1984:5), and has been shown by Cohen (1983:242) to be a
scavenger of dead animals. Unlike S. galapagensis, which
have vestigial lateral eyes, all instars of 5. lerneri have
well-developed lateral eyes. Both species have 5 juvenile and
1 adult instars.
Carapace: The carapace of S. galapagensis is larger than
that of 5. lerneri but the growth curves of both species are
parallel (Figure 10). The rostral infold of the adult S. lerneri
bears 32-42 bristles compared to only 7 for S. galapagensis.
The anteroventral infold of S. lerneri bears 15-20 bristles
inside the list compared to none on S. galapagensis. For both
species, the infolds of the caudal process are similar and the
carapaces of adult males are smaller than those of adult
females.
First Antenna: The distribution of bristles during ontogeny
is similar in both species with 1 exception: the 4th joint of the
adult male S. galapagensis bears 2 bristles (1 ventral, 1 dorsal)
whereas none were reported on S. lerneri by Cohen (1983, table
1). The distribution of filaments on adult females are, in
general, similar for both species. The adult male of S. lerneri
has 1 or 2 small suckers on each distal filament of the b- and
c-bristles (Cohen, in litL, 1988) compared to 6-8 for S.
galapagensis.
Second Antenna: The addition of bristles on the 9th
exopodial joint is similar for both species up to the adult stage,
where 5. galapagensis has 4 bristles and S. lerneri 5. The
addition of endopodial bristles is similar for both species.
NUMBER 483 23
1.5
1.4
1.3
1.2
I.I
i
E
1
 0.9 -
0 8
0.7
0.6
0.5
0.4
0.3
S. galapagensis
Sex Unknown x
Females ?
Males o
S. galapagensis y
adult
/
adult Zs **/
0 . 6 0 .7 0 . 8 0 .9 1.0 I.I 1.2 1.3 1.4 1.5 1.6 1.7 1.8 1.9 2 . 0 2 .1 2 . 2 2 . 3 2 . 4
Length (mm)
FIGURE 10.?Length-height curve of ontogenetic series of carapaces of Skogsbergia galapagensis, new species,
compared to that of 5. lerneri (Kornicker, 1958). Latter curve from Cohen (1983, fig. 3).
TABLE 10.?Carapace lengths and distribution of claws on each lamella of furca of Skogsbergia galapagensis
(X = claw with basal suture; A = claw with basal suture and stouter than previous claw; x = claw with either
weak basal suture or no basal suture).
Growth
stage
I
II
III
IV
IV
V
V
Adult
Adult
Sex
unknown
unknown
unknown
female
male
female
male
female
male
Carapace
length (mm)
0.78
0.92
1.19
1.39
1.30
1.75
1.83
2.02-2.14
1.76
1
X
X
X
X
X
X
X
X
X
2
X
X
X
X
X
X
X
X
X
3
X
X
X
X
X
X
X
X
X
4
X
A
X
X
X
X
X
X
X
Distribution of claws*
5
X
X
A
A
A
A
A
A
A
6
X
X
X
X
X
X
X
7
X
X
X
X
X
X
X
8
X
X
X
X
X
X
9
X
X
X
X
X
10
X
X
*Each number represents the position of a claw counting from distal end of lamella.
Mandible: The development of bristles of the coxale,
basale, exopodite, ventral margin of the 1st endopodial joint,
and the number of claws and bristles on the 2nd endopodial
joint are similar for both species. The 1st endopodial joint of
instar III of S. galapagensis has 3 bristles compared to 4 for S.
lerneri (Cohen, in litt., 1988). The number of dorsal bristles
on the 2nd endopodial joint of the adult, is 22 on S.
galapagensis and about 17 on S. lerneri.
Maxilla: The increase in numbers of alpha- and beta-
bristles on the 1st endopodial joints during ontogeny is similar
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
for both species. The number of bristles on the 2nd endopodial
joint of instar II 5. lerneri is 9 not 6 as was listed in Cohen
(1983, table 1) (Cohen, in litt., 1988).
Fifth Limb: Only small differences occur between the 2
species in the number of bristles on the fused 4th and 5th
exopodial joints. Additional differences may occur in the
number of bristles on the 1st and 2nd exopodial joints, but the
data presented herein and that by Cohen (1983, tables 1, 2) are
not directly comparable.
Sixth Limb: The increase in the number of bristles in
instars I and II are similar for both species, and differs only
slightly in later instars.
Seventh Limb: Bristles appear first in instar IV for both
species, and the rate of increase of bristles in later instars differs
only slightly.
Furca: For both species each lamella of the furca in instar
I has 5 claws, with only claw 1 having a basal suture, and in
instar II each lamella bears 5 claws with all except claw 2
having a distinct basal suture. In later instars S. galapagensis
bears 1 or 2 more claws than 5. lerneri (the adult S.
galapagensis has 10 claws, whereas S. lerneri has only 8).
Lateral Eye: The lateral eye of S. galapagensis is vestigial
on all instars, but it is relatively larger in instar I (compared
to size of protopodite of 2nd antenna (Figure 5c)). The lateral
eye of the embryo as well as juveniles and adults of 5. lerneri
are well developed (Cohen, in litt., 1988).
Upper Lip: The upper lip was fully developed in instar I
for both species.
Posterior of Body: For both species a tuft of hairs is
present on the posterodorsal margin of instars I?III, and absent
in later instars.
CONCLUSION.?Except for the vestigial lateral eye on the
troglobite S. galapagenis, its morphology and ontogeny are
quite similar to that of the open water species S. lerneri.
Order HALOCYPRIDA Dana, 1853
COMPOSITION.?The Halocyprida comprise the suborders
Halocypridina Dana, 1853, and Cladocopina Sars, 1866. Only
the former is present in the anchialine caves of the Galapagos
Islands.
Suborder HALOCYPRIDINA Dana, 1853
COMPOSITION.?The Halocypridina comprise the super-
families Thaumatocypridoidea Miiller, 1906, and Halocypri-
doidea Dana, 1853 (Kornicker and Sohn, 1976, fig. 2). Only
the former is represented in the collection from the Galapagos
Islands.
Superfamily THAUMATOCYPRIDOIDEA Miiller, 1906
COMPOSITION.?The superfamily comprises the family
Thaumatocyprididae Muller, 1906.
THAUMATOCYPRIDIDAE Muller, 1906
COMPOSITION.?This long-ranging family (Permian to
Holocene) comprises 5 genera: 2 known only as fossils
(Thaumatomma Kornicker and Sohn, 1976, from the Permian
of Idhra Island, Greece, and Pokornyopsis Kozur, 1974, from
the Upper and Lower Jurassic of southwestern and western
Germany), and 3 known only from the Holocene (Thaumato-
cypris Muller, 1906, from off Sumatra (1100 m) and Indonesia
(2000 m), Thaumatoconcha Kornicker and Sohn, 1976, widely
distributed in the Atlantic, Pacific, and Indian Oceans at depths
ranging from 150 to 4758 m, and Danielopolina Kornicker and
Sohn, 1976, from marine caves (Cuba, Canary Islands,
Bahamas, Yucatan peninsula, Mexico, Galapagos Islands) and
also from the deep sea (South Atlantic, 3459 m) (Kornicker
and Iliffe, in press a)).
2.
Key to Living Genera of Thaumatocyprididae
Each valve of carapace with long spine-like process (about xl% length of carapace)
at anteroventral margin Thaumatocypris
Each valve of carapace with only short process (less than l/4 length of carapace) at
anteroventral margin 2
Carapace surface smooth or punctate Thaumatoconcha
Carapace surface reticulate or spinous Danielopolina
Danielopolina Kornicker and Sohn, 1976
TYPE SPECIES.?Danielopolina carolynae Kornicker and
Sohn, 1976.
COMPOSITION AND DISTRIBUTION.?The genus comprises 6
species: D. carolynae Kornicker and Sohn, 1976, from the
South Atlantic near the equator at a depth of 3459 m, D.
orghidani (Danielopol, 1972), from a saline grotto in Cuba, D.
wilkensi Hartmann, 1985, from a marine lava-tunnel in the
Canary Islands, D. bahamensis Kornicker and Iliffe, in press
a, from a marine cave in Eleuthera, Bahamas, D. mexicana
Kornicker and Iliffe, in press a, from a marine cave on the
Yucatan Peninsula, Mexico, and D. styx, new species, from a
marine pool in the Galapagos Islands.
NUMBER 483 25
Key to the Species of Danielopolina
1. Carapace with surface spines D. mexicana
Carapace with surface reticulations 2
2. Carapace with walls of reticulations formed of minute papillae 3
Carapace with walls of reticulations formed by continuous or discontinuous ridges
5
3. Carapace longer than 1.5 mm D. carolynae
Carapace shorter than 1.0 mm 4
4. Each valve with single posterodorsal process; each lamella of adult furca with 3
short nonarticulated claws D. orghidani
Each valve without posterodorsal process; each lamella of adult furca with more
than 3 short nonarticulated claws D. styx, new species
5. Adult furca with 3 short nonarticulated claws on each lamella . . . D. bahamensis
Adult furca with 6 short nonarticulated claws on each lamella D. wilkensi
Danielopolina styx, new species
FIGURES 11-17
ETYMOLOGY.?From the Latin and Greek Styx (river in the
nether world).
MATERIAL.?Holotype: Sta 87-005: USNM 193439, 1
instar IV, appendages on 1 slide, carapace in alcohol.
Paratypes: Sta 87-005: USNM 193421, 1 instar III,
appendages on 1 slide, carapace in alcohol. Sta 87-018: USNM
193436A,B, 2 instar III, undissected, in alcohol; USNM
193437, 1 instar II, appendages on slide, carapace in alcohol;
USNM 193438, 1 instar I, appendages and 1 furcal lamella on
slide, carapace and 1 furcal lamella in alcohol.
DISTRIBUTION.?Santa Cruz Island, Galapagos Islands:
Deep Grieta east of Tortuga Bay, at 6-12 m water depth (type
locality); Grieta de Caleta la Torta at 17-29 m water depth.
MATURITY OF SPECIMENS.?Previously described species
of Thaumatocyprididae have on each lamella of the furca of
the adult 2 articulated claws on the anterior margin followed
by 3-6 nonarticulated ventral claws. The furcae of the 4 stages
of D. styx in the present collection have only 1 articulated
anterior claw. Therefore, the furca is a different type than
previously described. Because of this, the number of nonarticu-
lated claws cannot be used with certainty to estimate the age
of the specimens. However, because the number of nonarticu-
lated ventral claws increases by 1 in each of the 4 stages, it is
reasonable to believe that the stages are in sequence. In 2
TABLE 11.?Order of appearance of appendages of Danielopolina styx
(x = present; 0 = absent).
Growth
stage
I
II
III
rv
1 st and 2nd antennae, mandible,
maxilla, 5th limb, furca
X
X
X
X
6th limb
0
0
X
X
7th limb
0
0
0
X
previously described species of Thaumatocyprididae, D.
bahamensis and Thaumatoconcha radiata, the 6th limb
(bearing bristles) appeared first in instar III, and for the latter
species the 7th limb appeared first in instar IV. Instar IV is not
known for D. bahamensis, but since the 7th limb is absent in
instar III and present in instar V, it may also be present in instar
IV (Kornicker and Iliffe, in press a, table 2). By using the first
appearance of the 6th and 7th limbs on the 4 stages of D. styx,
the stages are interpreted to be instars I to IV (Table 11).
The total number of growth stages in the ontogeny of D.
styx is unknown. Within the Thaumatocyprididae, species with
5 and 7 growth stages have been described (Kornicker and
Iliffe, in press a). If D. styx is assumed to have either 5 or 7
growth stages, its instar IV would be either the A-1 or A-3
stage, but it is not known which. The ventral edge of each
lamella of the furca of instars I?III, but not instar IV, bears a
small triangular process that could be the site of a claw present
on the following stage. Its absence on instar IV may indicate
that instar IV is an adult, but because no genitalia were
observed that conclusion cannot be made at this time; however,
the possibility cannot be eliminated, because female genitalia
are small and could be overlooked.
DESCRIPTION OF INSTAR IV (Figures 11, 12).?Carapace
similar in shape, ornamentation, selvage, and central adductor
muscle attachments to those of instar III described herein.
Size: USNM 193439, length with anterior process 0.68
mm, length without anterior process 0.59 mm, height 0.51 mm.
First Antenna (Figure 1 \a,b): Limb with 8 joints. 1st joint
with 1 bare dorsal bristle and 1 longer, bare, backward pointing,
lateral bristle near ventral margin. 2nd joint with 1 bare dorsal
bristle and distal medial spines. 3rd joint defined from 4th by
slight indentation in ventral margin but without medial or
lateral sutures, and without separation of sclerotized frame at
ventral and dorsal margins, with spines forming 2 rows on
ventral margin. 4th joint with none or 1 short ventral bristle;
ventral edge of 3rd joint longer than ventral edge of 4th joint,
but dorsal margins of 3rd and 4th joints without discernible
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE II.?Danielopolina styx, new species, USNM 193439, holotype, instar IV, length 0.68 mm: a, left 1st
antenna, lateral view; b, joints 2-5 of right 1 st antenna, medial view; c, left 2nd antenna, medial view, only
base of endopodite shown; d, endopodite of left 2nd antenna, medial view; e, coxale endite of left mandible,
medial v iew;/ coxale endite of right mandible, lateral view; g, basalc and endopodite of left mandible, lateral
view; h, left maxilla, medial view, bristles of endites not shown; i, right maxilla, lateral view, bristles of endiies
not shown; j , an endite of maxilla; k, left 7th limb; /, right lamella of furca and unpaired process (u.p.).
NUMBER 483 27
boundary separating them. 5th joint about same length as
ventral margin of 4th joint, with 3 ventral bristles (2 long, 1
minute, medial) (Figure 1 Ib). 6th joint bare, slightly shorter
than 5th joint. 7th joint slightly longer than 6th, with 2 long
ventral b-, and c-bristles, and 1 short, dorsal a-bristle with
small marginal spines. 8th joint about half length of 7th, with
3 bristles (2 long ventral e-, and f-bristles, 1 shorter dorsal
d-bristle).
Second Antenna: Protopodite with long spines at pos-
teroventral corner (Figure lie). Endopodite (Figure lid): 1st
joint elongate with 1 distal bristle; 2nd joint slightly shorter
than 1st, with 1 short, dorsal, lateral bristle and 4 terminal
ventral bristles (3 long, 1 short); 3rd joint short, with 4 terminal
bristles (2 short, 2 minute). Exopodite with 9 joints (Figure
lie): 1st joint divided by medial suture into long proximal and
short distal parts; joints 2-8 each with long bristle with distal
natatory hairs; 9th joint small, with 2 bristles (1 long, 1
medium), both with distal natatory hairs; some long bristles
with few widely separated marginal spines.
Mandible: Coxale endite similar to that of instar 111 (Figure
llej). Basale (Figure llg): tooth of endite with 5 triangular
cusps with small marginal teeth; posterior edge of endite with
2 distal tubular bristles; anterior margin with 1 distal bristle;
lateral side of endite with 3 long bristles near posterior margin,
1 long bristle near anterior margin, and 1 short distal knife-like
bristle; medial side with 1 proximal bristle near dorsal margin
(only proximal part shown in illustrated limb) and proximal
and distal hairs. Endopodite (Figure llg): 1st joint spinous,
with 1 dorsal bristle at midlength; 2nd joint spinous, with 2
distal dorsal bristles, and 3 or 4 distal ventral bristles; 3rd joint
spinous, with 6 bristles.
Maxilla (Figure Uh-f): Coxale with stout plumose dorsal
bristle. Basale with slender medial tubeformed bristle. Endopo-
dite: 1st joint with proximal ventral bristle with long spines, 3
dorsal bristles, and 3 distal bristles on or near ventral margin;
end joint with 1 stout, unringed, nonarticulated, terminal claw
and 5 articulated ringed bristles. Endites obscure but each with
about 6 or 7 bristles, some tubular.
Fifth Limb (Figure 12a-c): Epipodite with bristles form-
ing 3 groups, each with 4 or 5 hirsute bristles (Figure 12c).
Protopodite and endopodite with 16 or 17 bristles (Figure 12a);
distal endopodial joint with additional short tooth-like medial
bristle. Exopodite 3-jointed: 1st joint divided into proximal
and distal parts by weak medial suture; proximal part with 3
ventral bristles at joint midlength, 1 medial bristle at joint
midwidth, and 1 long terminal dorsal bristle; distal part with
2 terminal ventral bristles and 1 medial bristle at joint
midwidth; 2nd joint with 1 ventral bristle at joint midlength,
and 1 medial bristle at joint midwidth; end joint with 3 bristles
(midbristle 50-59 percent and smallest bristle 35-42 percent
length of longest bristle) (Figure lib).
Sixth Limb (Figure I2d,e): Epipodite with 14 bristles
forming 3 groups (proximal dorsal group and distal ventral
group, each with 5 bristles, middle group with 4 bristles).
Protopodite with 3 distal, spinous, ventral bristles. Basale with
2 spinous ventral bristles. Small endopodite with 2 long
spinous bristles. Exopodite: 1st and 2nd joints fused (bare on
right limb (Figure I2d)), left limb with 2 distal bristles (1
ventral, 1 at midwidth (Figure 12e)); end joint with 2 bristles
(smaller bristle 71-72 percent of longer bristle).
Seventh Limb (Figure 1 Ik): Elongate with 2 long spinous
terminal bristles.
Furca (Figure 11/): Each lamella with 1 long anterior
articulated claw and 4 short ventral nonarticulated claws; all
claws with minute, anterior and posterior spines; each lamella
with medial and lateral spines (not shown in illustration;
similar to those on furca of instar III); stout unpaired process
(with minute marginal spines) on posterior of body just
proximal to lamellae.
Bellonci Organ: Absent.
Lip: Upper lip hirsute, in lateral view projecting posteri-
orly (Figure I2f-h); posterior edge with 2 tubular bristles and
2 stout spines (Figure 12/i). Esophagous narrow then broadens
at anterior gut (Figure 12/). Lower lip comprising lateral flap
on each side of mouth (Figure 12/).
Gut Content: Unrecognized granular particles.
DESCRIPTION OF INSTAR I (Figure 13).?Carapace similar
in shape (illustrated carapace somewhat distorted (Figure 13a)),
ornamentation, and central adductor muscle attachment scar
to that of instar III.
Size: USNM 193438 (shell distorted), length with anterior
process 0.29 mm, length without anterior process 0.27 mm,
height 0.27 mm.
First Antenna (Figure 13b): Limb with 8 joints similar in
proportions to those of instar II. 1st joint bare. 2nd joint with
distal medial spines. 3rd and 4th joints similar to those of
instars II and III. 5th joint with small ventral bristle not
reaching distal end of 7th joint. 6th joint bare. 7th joint with 2
bristles (1 long ventral, 1 short dorsal); 8th joint with 2 long
bristles.
Second Antenna (Figure 13c): Protopodite bare. Endopo-
dite 3-jointed but with 2nd and 3rd joints fused: 1st joint
elongate, bare; fused 2nd and 3rd joints with 1 filamentous
short unringed ventral bristle and 4 ringed bristles (2 long, 2
short). Exopodite similar to that of instar III, 8th joint with 2
bristles.
Mandible: Coxale endite similar to that of instar 111 (Figure
13d). Basale (Figure 13e): 4 terminal teeth and 2 posterior
tubular bristles similar to those of instar III; anterior margin
with 1 long distal bristle; lateral side with 1 long bristle distal
to midlength; indistinct short proximal medial bristle near
dorsal margin. Endopodite (Figure 13e): 1st joint bare; 2nd
joint with 1 dorsal bristle; 3rd joint with 4 bristles.
Maxilla (Figure 13/,g): Endite I with about 6 bristles;
endite II with 1 proximal and 4 terminal bristles; endite III
with 1 proximal and 3 terminal bristles; at least 1 tubular bristle
on each endite. Coxale with spinous dorsal bristle. Basale with
1 or 2 bristles near ventral margin (1 medial, 1 lateral).
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 12.?Danielopolina styx, new species, USNM 193439, holotype, instar IV, length 0.68 mm: a, left 5th
limb, lateral view, epipodite not shown; b, tip of right 5th limb, lateral view; c, epipodite of 5th limb, dorsal
lobe at top, not all bristles shown on ventral lobe; d, left 6th limb, lateral view; e, tip of right 6th limb, lateral
view; / , posterior view of esophagus, mouth, and upper lip, compressed under cover slip; g, posterior view of
upper lip, not under cover slip; h, detail of edge of upper lip from/; i, lower lip, anterior to left.
Endopodite: 1st joint with no or 1 proximal ventral bristle and
2 distal bristles (1 ventral, 1 dorsal); 2nd joint with 1 stout
nonarticulated claw-like bristle, and 2 or 3 slender bristles.
(Sutures between joints indistinct so that exact location of
some bristles uncertain.)
Fifth Limb (Figure I3h,i)' Epipodite with bristles forming
3 groups (not all bristles shown on Figure 130- Protopodite
and endopodite with 7 or 8 bristles including short tooth-like
bristle on distal endopodial joint Exopodite 3-jointed but with
1st and 2nd joints fused: 1st joint with 1 long terminal dorsal
bristle, and 2 bristles (1 ventral, 1 medial near ventral margin);
2nd joint bare; 3rd joint small with 1 terminal bristle.
Sixth and Seventh Limbs: Absent.
Furca (Figure 13/): Each lamella with 1 long anterior
articulated claw followed by 1 short nonarticulated claw on
anteroventral corner of lamella; a small triangular process at
midlength of ventral margin of each lamella; lateral and medial
surfaces of each lamella with spines forming rows; stout
unpaired process on posterior of body just proximal to
lamellae.
NUMBER 483 29
proximal
set J distal, sets
triangular '
tooth /
u.p.
FIGURE 13.?Danielopolina styx, new species, USNM 193438, paratype, instar I: a, complete specimen from
right side (shell distorted), length including anterior process 0.29 mm, central adductor muscle attachments
indicated by dashed oval; b, right 1st antenna, lateral view; c, endopodite of right 2nd antenna, medial view; J,
coxale endite of right mandible, lateral view; e, basale and endopodite of right mandible, lateral view; f,g,
maxillae; h, left 5th limb, medial view; i, right 5th limb, lateral view, not all epipodial bristles shown: j , left
lamella of furca and stout unpaired process (u.p.).
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Bellonci Organ: Absent.
DESCRIPTION OF INSTAR II (Figure 14).?Carapace similar
in shape, ornamentation and central adductor muscle attach-
ment scar to that of instar III (Figure I4a,b).
Size: USNM 193437: left valve, length with anterior
process 0.50 mm, length without anterior process 0.46 mm,
height 0.40 mm; right valve, length with anterior process 0.48
mm, length without anterior process 0.46 mm, height 0.40 mm.
First Antenna (Figure 14c,d): Limb with 8 joints. 1st joint
with 1 bare dorsal bristle. 2nd joint with distal medial spines
and distal low bulbous process (with small spines) extending,
past ventral margin of joint (Figure 14d). 3rd joint defined from
4th by slight step in ventral margin but without medial or lateral
sutures and without separation of sclerotized frame at ventral
and dorsal margins (slight sinuate curvature in dorsal margin
probably indicates location of division between 3rd and 4th
joints), with spines forming 2 rows on ventral margin, and
spines forming distal row on dorsal margin. 4th joint bare.
Ventral edge of 3rd joint longer than ventral edge of 4th joint
but reverse on dorsal margin. 5th joint smaller than 4th, with
short ventral bristle with minute terminal papilla. 6th joint
smaller than 5th and without bristles. 7th joint longer than 6th,
with 2 bristles (1 long ventral with few widely separated
minute spines, 1 short dorsal with small marginal spines). 8th
joint smaller than 7th, with 3 bristles (1 long lateral, 1 medium
dorsal medial, 1 short ventral medial, all with widely separated
minute marginal spines and terminal papilla).
Second Antenna: Protopodite bare. Endopodite 3-jointed
but with 2nd and 3rd joints fused (Figure 14e): 1st joint
elongate with 1 short distal bristle; fused 2nd and 3rd joints
with 5 bristles (3 long, 1 short, 1 minute). Exopodite similar
to that of instar III.
Mandible: Coxale endite folded on illustrated limb (Figure
14/), but similar to that of instar III. Basale similar to that of
instar III except for absence of short distal lateral bristle, and
for 1 of paired anterior bristles being short (Figure 14g).
Endopodite with 1 dorsal bristle on 1st joint, 2 dorsal bristles
on 2nd joint, and 4 bristles on end joint (Figure \4h).
Maxilla (Figure I4i,j): Similar to that of instar III but with
fewer terminal bristles. Endites not examined in detail.
Fifth Limb (Figure 14*): Epipodite present but exact
number of bristles not determined because bristles obscured.
Protopodite obscured on specimen examined. Distal endopo-
dial joint with short tooth-like bristle, 4 ventral bristles (1 stout
pectinate, 3 slender). Exopodite 3-jointed: 1st joint with 1 very
long terminal dorsal bristle, 1 ventral bristle, and 1 medial
bristle at joint midwidth; 2nd joint elongate, without bristles;
end joint small with 2 or 3 bristles (1 long, 1 or 2 short, all
with marginal spines).
Sixth Limb: Not observed.
Seventh Limb: Absent.
Furca (Figure 14/): Each lamella with 1 long anterior
articulated claw followed by 2 short nonarticulated ventral
claws and small triangular process; all claws with anterior and
posterior spines; lateral and medial surfaces of lamella with
spines forming rows; stout unpaired process on posterior of
body just proximal to lamellae.
Bellonci Organ: Absent.
Lip: Similar to that of instar III.
Gut Content: Unrecognized amber-colored particles.
DESCRIPTION OF INSTAR III (Figures 15-17).?Carapace
subround in lateral view with straight dorsal margin in vicinity
of hinge and also straight margin between anterior and
anteroventral processes (Figures 15, 16a,b); ventral and
posterior margins as well as anterior margin dorsal to anterior
process evenly rounded; valves broadest at about midlength
and midheight, in vicinity of central adductor muscle
attachments. Short anterior and anteroventral processes with
bases just lateral to valve edge; each process bearing fragile
spine-bearing frill that easily breaks off at slight touch with
dissecting needle leaving smaller firm triangular protuberance.
Ornamentation (Figures 15, \6a,b): Surface finely reticu-
late with reticulation walls formed of minute pustules, most
with blunt tips, but few with pointed tips; on anterior part of
shell pustules form rows (about 8) paralleling straight edge of
shell between anterior and anteroventral processes. (Pustules
break off easily, and on most specimens in collection pustules
missing on parts of shell, especially near center.)
Adductor Muscle Attachment Scar (Figures 15, 16a): Scar
central in location; subround with greatest diameter trending
towards posterodorsal margin of shell; scar consisting of 7 or
8 wedge-shaped scars more or less radially arranged.
Crescent-shaped mandibular scar located anteroventral to
adductor muscle scar (Figure 16a).
Size: USNM 193421, length with anterior process 0.57
mm, length without anterior process 0.52 mm, height 0.41
mm; USNM 193436A, length with anterior process 0.59 mm,
length without anterior process, 0.55 mm, height 0.46 mm;
USNM 193436B, length with anterior process 0.56 mm, length
without anterior process 0.53 mm, height 0.44 mm.
First Antenna (Figure 16c): Limb with 8 joints. 1st joint
with 1 bare dorsal bristle. 2nd joint with 1 bare dorsal bristle
and distal medial spines. 3rd joint defined from 4th by slight
step in ventral margin but without medial or lateral sutures,
and without separation of sclerotized frame at ventral or dorsal
margins (slight sinuate curvature in dorsal margin probably
indicates location of division between 3rd and 4th joints), with
spines forming 1 or 2 ventral rows, and spines forming distal
row near dorsal margin. 4th joint bare; ventral edge of 3rd joint
longer than dorsal edge of ventral 4th joint, but relationship
reversed on dorsal margin. 5th joint smaller than 4th, with 1
long ventral bristle with few minute widely separated spines.
6th joint slightly smaller than 5th and without bristles. 7th joint
about same length as 6th, with 2 bristles (1 long ventral and
with few small widely separated marginal spines, 1 short dorsal
with small marginal spines), 8th joint smaller than 7th, with 2
long ventral bristles and 1 shorter, dorsal bristle.
Second Antenna (Figure 16a): Protopodite with long
NUMBER 483 31
FIGURE 14.?Danielopoima styx, new species, USNM 193437, paratype, instar II: a, complete specimen from
right side, length including anterior process 0.50 mm; b, oblique view of right valve from inside, note list along
ventral and posterior infold; c, right 1 st antenna, lateral view; d, parts of joints 2 and 3 of left 1st antenna showing
spines, medial view; e, endopodite of right 2nd antenna, lateral view; /, coxale endite of left mandible (endite
folded); g, basale and 1st endopodial joint of left mandible, medial view, dorsal bristle of 1st endopodial joint
not shown; h, endopodite of right mandible, lateral view; i, left maxilla, medial view, endite bristles not shown;
/, right maxilla, lateral view, endite bristles not shown; k, left 5th limb, medial view; /, right lamella of furca and
unpaired process (u.p.).
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 15.?Danielopolina styx, new species, USNM 193421, paratype, ins tar
III, complete specimen from right side, length including anterior process,
0.57 mm.
spines at posteroventral corner (Figure \6d). Endopodite
3-jointed, but with 2nd and 3rd joints fused: 1st joint elongate,
with 1 bristle; 2nd joint about 3/o length of 1st, with 2 terminal
bristles (1 lateral, short, filamentous, 1 medial, long, ringed,
bare); 3rd joint short, with 4 bristles (2 long, 1 short, 1 minute).
Exopodite with 8 joints: 1st joint undivided; bristles of joints
2-7 long, ringed, with few widely separated minute marginal
spines, and distal natatory hairs; 8th joint with 2 bristles (1
long, 1 medium), both with few minute widely separated
marginal spines.
Mandible (Figure \6e-g): Coxale endite with proximal
and distal sets of teeth separated by space (Figure 16e);
proximal set comprising 4 broad cusps plus triangular tooth
close to distal set of teeth; surface between cusps and medial
and lateral surfaces just proximal to cusps with slender spines;
1 spinous bristle with base just distal to triangular tooth; distal
set of ?jeeth consisisting of 2 flat teeth, each having 4 or 5
pointed cusps; cusps of proximal flat tooth pectinate; distal
flat tooth with small proximal medial tooth on medial cusp.
Basale (Figure 160: tooth of endite with 5 triangular cusps
with small marginal teeth; posterior edge of endite with 2
tubular bristles (1 proximal, 1 distal, each with terminal
papilla); anterior margin of endite with 2 ringed bristles (1
with medial base, 1 with lateral base); lateral side of endite
with 2 slender proximal ringed bristles and 1 shorter distal
ringed bristle; medial side with 1 proximal ringed bristle near
dorsal margin. Endopodite 3-jointed with 1st and 2nd joints
about same length and 3rd joint shorter (Figure 16/,g): 1st joint
with 1 dorsal bristle at midlength; 2nd joint spinous, with 2
dorsal bristles (dorsal margin of joint narrower distal to
bristles); 3rd joint spinous, with 4 bristles.
Maxilla (Figure \la-c): Endite I with 6 bristles; endite II
with 7 bristles; endite III with about 4 bristles; 1 or 2 tubular
bristles on each endite. Coxale with long stout plumose dorsal
bristle. Basale with slender medial tubular bristle. Endopodite:
1st joint with 2 distal bristles on or near dorsal margin and
distal bristle at joint midwidth or near ventral margin. End joint
with 1 anterior, stout, linear, sclerotized nonarticulated claw,
and 4 articulated slender bristles.
Fifth Limb (Figure Yld.e): Epipodite with bristles forming
3 groups (Figure \ld). Protopodite and endopodite with about
11 bristles; distal endopodial joint with additional short
tooth-like bristle. Exopodite 3 jointed: 1st joint with 1 very
long terminal dorsal bristle, 1 ventral bristle, and 1 medial
bristle at joint midwidth; 2nd joint elongate with 1 bristle on
ventral margin; end joint small with 3 bristles (middle bristle
45 percent and smallest bristle 32 percent length of longest
bristle).
Sixth Limb (Figure llf.g): Epipodite with bristles forming
3 groups (Figure 17/). Protopodite obscured on limb examined.
Basale with 1 ventral bristle (this bristle could be on
protopodite). Small endopodite with 2 long bristles. Exopodite:
1st and 2nd joints fused, bare; end joint short, with 2 weakly
developed bristles.
Seventh Limb: Not observed.
Furca (Figure \7h): Each lamella with 1 long anterior
articulated claw followed by 3 short nonarticulated ventral
claws; claws with minute spines along anterior and posterior
edges; 1 or both lamellae with small triangular process
posterior to posterior ventral claw; anterior margin of lamella
with few short spines; lateral surface of lamella with minute
spines forming rows, more and longer spines on medial surface
(some medial spines extending past ventral edge of lamella);
stout unpaired process (with minute marginal spines) on
posterior of body just proximal to lamellae.
Bellonci Organ: None observed.
Lip: Upper lip with distal spines and a short tubular bristle
on posterior edge on each side of midwidth (Figure \6h).
Esophagus narrow; broad anterior part of gut separated from
smaller and narrower posterior part by slight restriction; anus
small (Figure 160-
Gut Content (Figure 16/): Unrecognized brown particles.
COMPARISONS.?The carapace of D. styx resembles those
of D. orghidani and D. carolynae in having reticulations with
walls formed by papillae. It differs from both species in not
having a small posterodorsal process on each valve. The
carapace of D. styx is much smaller than that of D. carolynae.
Danielopolina styx bears more than the 3 short nonarticulated
claws present on each lamella of the furca of D. orghidani.
Danielopolina styx differs from previously described species
of the Thaumatocyprididae in having only 1 instead of 2
articulated anterior claws on each lamella of the furca.
ONTOGENY.?As in Thaumatoconcha radiata and D.
NUMBER 483 33
FIGURE 16.?Danielopolina styx, new species, USNM 193421, paratype, insiar III: a, complete specimen from
left side, length including anterior process 0.57 nun; b, oblique view of left valve from inside, note central
adductor muscle bundle; c, left 1 st antenna, lateral view; d, endopodite and distal part of protopodite of right
2nd antenna, medial view; e, coxale endite of left mandible, medial view; /, basale and 1 st endopodial joint of
left mandible, medial view, dorsal bristle of 1st endopodial joint not shown; g, endopodite of left mandible,
medial view; h, upper lip compressed under cover slip; i, gut viewed from right side, ventral side to left.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
7
FIGURE 17.?Dan'ulopcAina styx, new species, USNM 193421, paratype, instar III: a,b, maxillae, bristles of
endites not shown; c, endites of maxilla shown in b; d, epipodite of 5th limb, dorsal lobe at top, not all bristles
shown on ventral lobe; e, right 5th limb, medial view, epipodite not shown;/, epipodite of 6th limb, dorsal lobe
at top, not all bristles shown on ventral lobe; g, 6th limb, epipodite not shown; h, furcal lamellae and unpaired
process (up).
NUMBER 483 35
bahamensis the 1st and 2nd antennae, mandible, maxilla, 5th
limb, and furca are already present in the 1st instar (Table 11);
the 6th and 7th limbs are not present; the Bellonci organ is
absent on all instars. The 6th limb appears, but with just a few
bristles, in the 3rd instar, and the 7th limb in the 4th instar.
The carapace of the single instar I in the collection is distorted,
but all instars appear similar except for size.
First Antenna: The number of bristles on the 1st antenna
increases from 5 on instar I to 12 or 13 on instar IV (Table
12). The ventral bristle of the 5th joint of does not reach past
the 8th joint on instars I and II but is extremely long on instar
III.
Second Antenna: The protopodite of the 2nd antenna is
bare on instars I and II but bears long spines on the posterior
ventral corner on instars III and IV. The exopodite has 8 joints
on instars I to III and 9 joints on instar IV. The exopodial
bristles bear distal natatory hairs on all instars. The 1st
endopodial joint is bare on the 1st instar but bears 1 bristle on
later instars (Table 13). The 2nd and 3rd endopodial joints are
fused on all instars, but can be differentiated by the location
of bristles on instars III and IV. The total number of bristles
on the 2nd and 3rd joints increases from 5 on instar I to 9 on
instar IV (Table 13). All 4 terminal bristles of the 3rd
endopodial joint of instar IV are short, whereas 2 of 4 terminal
bristles of instar III are very long, possibly the difference is
sexual, but sex of the instars was not determined.
Mandible: The terminal teeth of the coxale and basale are
similar on all 4 instars. The number of lateral bristles of the
basale as well as the number of bristles of 1st and 2nd
endopodial joints are higher on later instars, but the number
of bristles of the 3rd endopodial joint remains at 4 (Table 13).
Fifth Limb: The number of exopodial bristles increases
from 4 on instar I to 13 on instar IV (Table 14).
Sixth Limb: The 6th limb appears first on instar III, and
bears rather short and weakly developed bristles, but the
number of bristles is only slightly less than on the limb of
instar IV (Table 14).
Seventh Limb: The limb appears first on instar IV and
bears 2 terminal bristles, the same number observed on adults
of other species of the genus.
Furca: All 4 instars have only 1 articulated claw on the
anterior margin of each lamella. The number of nonarticulated
claws increases from 1 on instar I to 4 on instar IV (Table 15).
The distribution of furcal claws on other Thaumatocyprididae
are presented for comparison in Table 15.
TABLE 12.?Number of bristles on joints of 1st antenna of Danielopolina styx.
I
II
III
IV
1st
joint
lateral
0
0
0
1
dorsal
0
1
1
1
2nd
joint
dorsal
0
0
1
1
3rd
joint
0
0
0
0
4th
joint
ventral
0
0
0
0-1
5th
joint
ventral
1
1
1
3
6th
joint
0
0
0
0
7th
joint
ventral
1
1
1
1-2
dorsal
1
1
1
1
8th
joint
2
3
3
3
TABLE 13.?Number of bristles on 2nd antenna and mandible of
Danielopolina styx.
Growth
stage
I
II
III
rv
Second antenna
Endopodite
1st
joint
0
1
1
1
2nd + 3rd
joints
5
5
6
9
Basale
lateral
bristles
1
2
3
5
Mandible
1st
joint
0
1
1
1
Endopodite
2nd
joint
1
2
2
5-6
3rd
joint
4
4
4
4
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 14.?Number of bristles on exopodites of 5th and 6th limbs of Danielopolina styx.
Growth
stage
I
II
III
rv
1st
joint
3
3
3
8
Fifth limb
2nd
joint
0
0
1
2
3rd
joint
1
2-3
3
3
1st
joint
3
4
Sixth limb
lst-2nd
joints (fused)
limb absent
limb absent
0
0-2
3rd
joint
2
2
TABLE 15.?Number of claws on each lamella of furca of Danielopolina styx,
D. bahamensis, and Thaumatoconcha radiata (long articulated anterior
claws:short nonarticulated ventral claws; nd = no data; ns = species without
this stage).
Growth
stage
I
II
III
IV
V
VI
Adult
D.styx
1:1
1:2
1:3
1:4
nd
nd
nd
D. bahamensis*
2*:0
2:l(anlage)
2:1
nd
ns
ns
2:3
T. radiata^
2*:0
2:1
2:2
2:3
2:4
2:5
2:6
?Data from Kornicker and Iliffe (in press a).
+Data from Kornicker and Sohn (1976, table 7).
^Second long anterior claw not articulated.
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Islands. Mitteilungen aus dem Hamburgischen Zoologischen Mu-
seum und I nst'Uut, 83:223-230, figures 1-3.
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