JOURNAL OF CRUSTACEAN BIOLOGY, 22(2): 268–278, 2002
ORIENTATION OF THE MAXILLA OF THE SARSIELLINAE (OSTRACODA)
Louis S. Kornicker
Department of Systematic Biology, Invertebrate Zoology Section, National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560, U.S.A. (e-mail: [email protected])
A B S T R A C T
The maxillae (fourth limbs) of myodocopine Ostracoda are difficult to interpret correctly because
of their complexity, which has led to inaccurate descriptions of the limb in the literature. Maxillae
of species of the Sarsiellinae are described and illustrated herein in their natural orientation on the
body of the animal, and clues are presented to assist in more accurate descriptions of the limb
when compressed under a cover slip. It is shown that the anterior-to-posterior sequence of the three
endites of the protopod in the Sarsiellinae is the reverse of that in the Cypridinidae, Philomedidae,
and Rutidermatidae.
According to Müller (1894: 53) [translation
from German], “The maxilla offers the most
difficulties in anatomical analysis and morphological interpretation.”
The maxilla (also known as fourth limb and
maxillula) (Figs. 1–3) of the Sarsiellinae is
composed of a protopod, endopod, and exopod. The protopod bears three endites termed
herein: endite I (proximal), endite II (middle),
and endite III (distal). The protopod is usually subdivided in morphological descriptions
into a precoxa, coxa, and basis (e.g., Figs.
1–3), but sutures separating them are not always well-defined. Endite I is usually correctly referred to as part of the precoxa, and
endite II is usually correctly referred to as part
of the coxa. Except for Boxshall (1997: 161;
see below), endite III usually also has been
referred to as part of the coxa. The endites
herein are mostly referred to as endites I, II,
and III of the protopod, without attempting to
identify the segments of which they are part.
Most sarsiellids bear a distal coxal bristle on
the anterodorsal margin of the coxa (Fig. 3).
Some sarsiellid species bear a transparent
filmy thumb-like process on the protopod (see
“film” Figs. 1B, 4C, G, 5A). The exopod consists of a small lobe generally bearing two
or three bristles, and the endopod has two
segments, with two bristles on segment 1 and
eight on segment 2 (Figs. 1A, 5A, 6E). The
male, unlike the female, is not a carnivore and
eats little; the maxilla is reduced and not considered further here. The maxillae of males of
the Philomedidae and Rutidermatidae are also
reduced. For spinosity of segments and bristles see description of the sarsiellid maxilla
in Poulsen (1965: 51, 52).
Concerning the maxillula [maxilla],
Boxshall (1997: 161) stated “Four endites
(one precoxal, one coxal and two basal) are
present in copepods, ostracods and mystacocarids.” Boxshall studied the maxilla of the
cypridinid Azygocypridina imperialis, and his
→
Fig. 1. A, Junctichela pax Kornicker et al., 2002, USNM 194595, instar III, sex unknown, lateral view of part of
complete specimen (not under cover slip) with carapace removed showing location of left maxilla relative to other
left appendages; B, Eusarsiella species (Eusarsiella species indeterminate in Kornicker and Iliffe, 2000), USNM
194490, instar IV, lateral view of part of complete specimen (not under cover slip) showing location of left maxilla
relative to other appendages (parts of right mandible, left fifth and sixth limbs) (outer part of left maxilla removed
and shown separately to show location of endites relative to upper lip on main illustration); C, Spinacopia variabilis
Kornicker, 1969, USNM 122550, adult female, medial view of part of complete specimen (not under cover slip and
with carapace removed) showing medial view of maxilla and mandible, and upper lip from left side. Abbreviations
for all figures: bas = basis; bb = basal bristle; cb = coxal bristle; cx = coxa; end = endopod; epip = epipod; ex = exopod; mnd = mandible; mo = mouth; mx = maxilla; precx = precoxa; rt = right; ul = upper lip; I–III = endites I–III.
Arabic numerals 1 and 2 indicate individual joints of each limb; Arabic numbers 5 and 6 indicate 5th and 6th limbs,
respectively; arrows on illustrations indicate anterior of body; illustrations may not show the rings on all bristles that
possess rings; individual lower-case letters a, c and symbols α, β on illustrations are those routinely used in species
descriptions to identify certain bristles on the second endopodial segment having the same location on all sarsiellid
species; sclerites stippled; muscles striated.
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KORNICKER: ORIENTATION OF MAXILLA OF SARSIELLINAE
269
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JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 22, NO. 2, 2002
Fig. 2. A, Horizontal cross section of the maxilla of a species belonging in the Sarsiellinae. The limb is interpreted
here to be a dorsal view of a left limb, but could also be interpreted to be a ventral view of a right limb. The location of the cross section (×-sect) on the limbs is indicated in B, C, D, and E. Simplified views of maxilla: B, medial
view; C, lateral view; D, anterior view; E, posterior view.
interpretation of the distribution and number
of endites is clearly plausible for that species
(see Boxshall, 1997: Figs. 13.1, 13.2b). Examples of cypridinids that do not have endite III as part of the basis are present in some
species of the genus Cypridinodes (see
Poulsen, 1962: Figs. 129a, 134a).
In the Sarsiellinae, whether endite III is
part of the coxa or basis is not obvious. An
example of a species that appears to have endite III as part of the coxa is Parasarsiella
globulus (Brady and Norman, 1896) (see
Poulsen, 1965: Fig. 21a); whereas, an example of a species appearing to have endite
III as part of the basis is Sarsiella crispata
(Scott, 1905) (see Poulsen, 1965: Fig. 18h).
No attempt is made herein to relate endite III
to either the coxa or basis. Unless a small
bristle on the basis (adjacent to the exopod)
(see e.g., Fig. 4A–C, F, G) is interpreted to
be an endite, which is not so interpreted
herein, the maxilla of the Sarsiellinae bears
a total of only three endites on the protopod.
In the natural position of the maxillae in
place on the body of the Cypridinidae (females and males), the Philomedidae (females), and the Rutidermatidae (females),
when viewed either laterally or medially, the
anterior edge of endite I is posterior to the anterior edge of endite II, and the anterior edge
of endite II is posterior to the anterior edge
of endite III (for Cypridinidae see Poulsen,
1962: fig. 6; Kornicker, 1997: fig. 1). The
relative positions of the anterior edges of the
KORNICKER: ORIENTATION OF MAXILLA OF SARSIELLINAE
271
Fig. 3. Maxilla of species of Sarsiellinae (all limbs viewed while not compressed under a cover slip): Spinacopia
variabilis Kornicker, 1969, USNM 122550, adult female, left limb: A, medial view; B, lateral view; C, anterior
view; D, posterior view; E, ventral view; F, Eusarsiella species A (Eusarsiella species indeterminate in Kornicker
and Iliffe, 2000 (drawn while attached to body)), USNM 194490, instar IV, right limb, lateral view. Figure 3A–D
matches the diagrammatic illustrations shown in Fig. 2B–E, respectively. Not all bristles shown in illustrations.
endites of the maxillae in place on the body
of the Sarsiellinae (females), when viewed either laterally or medially, are the reverse of
those three families; i.e., the anterior edge of
endite I is anterior to the anterior edge of endite II, and the anterior edge of endite II is
anterior to the anterior edge of endite III
(Figs. 1B, C, 2B, C, 3A, B). The difference
in the relative locations of the endites of the
Sarsiellinae and other families suggests that
members of the former handle food somewhat
differently. Females of the Sarsiellinae and
Rutidermatidae are predators, whereas the
Philomedidae are detritus feeders and the
Cypridinidae have multiple feeding methods
(Kornicker, 1975: 38, table 16).
The angles at which various parts of the
maxilla join each other have resulted in many
mistakes in descriptions of maxillae in the literature, especially if the descriptions are
based on a maxilla compressed under a cover
slip, because the relative positions of parts
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may differ as a result of distortions caused by
the flattening process. The present note attempts to show how to recognize the orientation of various parts of the maxilla of members of the Sarsiellinae, and is a sequel to a
previous similar note concerning the maxilla
of the Cypridinidae (Kornicker, 1997: 654).
Figure 2 is a mechanical-type drawing of
a maxilla viewed from four directions while
attached to the body of a sarsiellid. The cross
section shown in Fig. 2A is designated herein
to be of a left limb viewed dorsally, but could
be interpreted to be a right limb viewed ventrally. The medial, lateral, anterior, and posterior views, respectively, are shown in Fig.
2B–E. The endopod and endites are slightly
oblique to the long axis of the body, but it is
expedient to refer to their outer sides as lateral and inner sides as medial, and to the anteromedial and posterolateral edges as anterior and posterior edges, respectively. Note
that in Fig. 2B, C, which represent the medial and lateral views of a left maxilla, respectively, the anterior edge of the endopod
is separated from the endites by the width of
the endopod, whereas, in Fig. 2D, E, which
represent the anterior and posterior views of
the limb, respectively, the anterior edge of the
endopod is adjacent to the endites. The reverse is true for the exopod, which is adjacent to the endites in medial and lateral views
(Fig. 2B, C), and is separated from the endites by the width of the endopod in anterior
and posterior views (Fig. 2D, E).
Fig. 3A–E illustrates the left maxilla of
Spinacopia variabilis Kornicker, 1969, either
in its natural position on the body, or removed
from the body but shown in the natural orientation. Fig. 3A–D matches the diagrammatic illustrations shown in Fig. 2B–E, respectively. Figure 3A is a medial view, Fig.
3B a lateral view, Fig. 3C an anterior view,
and Fig. 3D a posterior view. Figure 3E is a
ventral view of the maxilla (a view not shown
in Fig. 2); Fig. 3F is a lateral view of the right
maxilla of Eusarsiella species (instar IV) (Eusarsiella species indeterminate in Kornicker
and Iliffe, 2000: 45) drawn while attached to
the body. Not all bristles are shown in illustrations of Fig. 3.
Identification of Limb.—The locations of certain morphological characters on maxillae
segments are useful in orienting the segments
(Figs. 1, 3).
Endopod: For origin of letters used to identify endopodial bristles see Poulsen (1965:
52). Segment 1: The distal margin bears 1
long anterior α-bristle, and 1 long posterior
β-bristle. Segment 2: Two short a-bristles are
always lateral (the bristles, especially the bristle farthest from midwidth are located in the
posterior half of the segment) (Fig. 3B); a
single small c-bristle is always medial (Figs.
3A, 6F).
Because of the greater width of the endopod,
when compressed under a cover slip it usually
is oriented with its medial or lateral side uppermost. The location of the a- and c-bristles
of the second endopodial segment permits
identification of the limb as either a left or
right limb, and whether the medial or lateral
side is uppermost.
Exopod: The exopod is generally a small
lobe located posterior and lateral to the base
of the endopod (Fig. 3B, D, F).
Protopod: Coxa: The distal dorsal margin
generally bears a short anterior bristle (Fig.
3). Precoxa: In some orientations of the limb,
endite I is clearly seen as part of the first protopodial segment (precoxa) (e.g., Fig. 3C, D).
Basis: The lateral side of the basis generally
bears a small distal bristle just posterior to the
exopod (Fig. 3B, D, F).
Endites: Endite II lies between endites I
and III (Fig. 3D). Endite I: As mentioned
above, in some views endite I is clearly part
of the first protopodial segment (Fig. 3). Endite III is adjacent to the exopod; it generally is fairly broad, bears 4–6 bristles, with
one or two being clawlike; the bristles are
located along the ventral margin and have two
or three anterior bristles separated by a space
from two or three posterior bristles (Fig. 4).
In their natural position on the body the endites are medial to the endopod, and generally are in the same relative location when the
limb is flattened under a cover slip, but the
flattening process may distort the limb so that
the endites are not medial to the endopod.
Analysis.—The orientation of parts of 10 sarsiellid maxillae that had been compressed under a cover slip are analyzed below to demonstrate method (Figs. 5, 6). Not all bristles are
shown in illustrations of Figs. 5, 6.
Eusarsiella absens (Kornicker, 1981),
USNM 158116, adult female, right limb (Fig.
5D); Eusarsiella costata (Kornicker, 1958),
USNM 158334, adult female, right limb (Fig.
KORNICKER: ORIENTATION OF MAXILLA OF SARSIELLINAE
273
Fig. 4. Endite III of maxilla of species of Sarsiellinae: A, Eusarsiella dispar Kornicker, 1986, USNM 158033,
adult female, left limb, lateral view; B, Eusarsiella maurae (Kornicker, 1977), USNM 156739, adult or A-1 female,
left limb, lateral view; C, Eurypylus hapax Kornicker and Iliffe, 2000, USNM 194494, instar IV male, right limb,
lateral view; D, Anscotiella vertex Kornicker, 1991, USNM 158317, adult female, left limb, medial view; E, Ancohenia robusta (Brady, 1890), USNM 193623, instar III female, left limb, lateral view; F, Spinacopia menziesi Kornicker, 1969, USNM 122086, adult female, right limb, lateral view; G, Eusarsiella styx Kornicker and Iliffe, 1989,
USNM 193367, adult female, left limb, lateral view. (Figure 2 in L. S. Kornicker, 2001, Exopod and protopodial endite III of the maxillae of Sarsiellinae (Crustacea: Ostracoda: Myodocopa). Proceedings of the Biological Society of
Washington 114. Used with permission.)
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Fig. 5. Maxilla of species of Sarsiellinae compressed under cover slip: A, Metasarsiella vibex Kornicker, 1991,
USNM 158310, adult female, right limb; B, Chelicopia arostrata Kornicker, 1958, USNM 158335, adult female, left
limb; C, Chelicopia arostrata Kornicker, 1958, USNM 158335, adult female, right limb; D, Eusarsiella absens (Kornicker, 1981), USNM 158116, adult female, right limb.
KORNICKER: ORIENTATION OF MAXILLA OF SARSIELLINAE
275
Fig. 6. Maxilla of species of Sarsiellinae compressed under cover slip: A, Spinacopia rex Kornicker, 1995, USNM
193868, adult female, right limb; B, Eusarsiella bex Kornicker, 1994, USNM 193866, adult female, right limb; C,
Eusarsiella gettlesoni Kornicker, 1986, USNM 156826, adult female, right limb; D, Eusarsiella costata (Kornicker,
1958), USNM 158334, adult female, right limb; E, Eusarsiella bex Kornicker, 1994, USNM 193866, adult female,
left limb; F, Ancohenia robusta (Brady, 1890), USNM 193623, Instar III female, right limb.
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6D): The a-bristles of the second endopodial
segment of both illustrated limbs are uppermost, indicating a lateral view of the endopod. The same bristles are posterior to midwidth of the segment, indicating that the
illustrations are a lateral view of the endopod of a right limb with the anterior towards
the right. The location of the anterodorsal
bristle of the coxa indicates a lateral view of
that segment. The small bristle on the basis
adjacent to the lateral exopod indicates a lateral view of the basis. The orientation of the
illustrations are comparable to left limbs illustrated in Figs. 2C and 3B, F, which are
not flattened under a cover slip.
Eusarsiella bex Kornicker, 1994, USNM
193866, adult female, left limb (Fig. 6E): The
c-bristle of the second endopodial segment
is uppermost, indicating a medial view of the
endopod. The posterior location of the a-bristles of the second endopodial segment indicates that the anterior of the endopod is towards the right, which identifies the limb as
a left limb. The anterior bristle of the coxa
conforms with the anterior of the endopod,
indicating a medial view of the coxa. The exopod is posterior to the endopod, indicating
a medial view of the exopod. The posterior
location of the small bristle of the basis,
which is adjacent to the exopod, indicates a
medial view of the basis. The endites are
more difficult to identify. The endite to the
left in the illustration has bristles typical of
endite III (see Fig. 4), and therefore is identified as that endite; the adjacent endite is then
endite II, and the right endite is endite I,
which is anterior to the other two endites. The
endites are interpreted to be medial views.
The orientation of the limb is comparable to
the left limbs illustrated in Figs. 2B and 3A,
which are not flattened by a cover slip.
Eusarsiella bex Kornicker, 1994, USNM
193866, adult female, right limb (Fig. 6B):
The c-bristle of the second endopodial segment is uppermost, identifying the segment
as being a medial view; the a-bristles of the
same segment are posterior to midwidth,
identifying the left side of the segment as being anterior. The location of the a- and c-bristles identify the limb as a right limb. The
overlapping stout claws of the second endopodial segment indicate that the endopod
is mounted in an oblique orientation with the
posterior edge of the segment somewhat medial to the anterior edge. The posterior loca-
tion of the single stout bristle of the second
endopodial joint identifies it as the β-bristle;
the α-bristle is missing from the specimen.
Endite I is recognized by the well-defined suture separating it from endites II and III. Endite III is uppermost on the slide relative to
the endopod, indicating a medial view of the
endites. The exopod is not identified with certainty; it is not lobate and bears only one bristle, but it is in the expected location for an exopod. The usual anterodorsal coxa bristle was
not observed. The limb is interpreted to be a
more-or-less posterior view of a right limb.
The orientation of the limb is comparable to
the left maxillae shown in Figs. 2E and 3D,
which are not flattened under a cover slip.
Spinacopia rex Kornicker, 1995, USNM
193868, adult female, right limb (Fig. 6A):
The c-bristle of the second endopodial segment is medial, and the a-bristles are lateral
and towards the posterior of the segment.
Therefore, the illustration is a medial view
of an endopod of a right limb with the anterior towards the left. The anterior location of
the anterodorsal bristle of the coxa indicates
a medial view of the coxa. Endite III is identifiable by the typical arrangement of bristles.
It is beneath endites I and II, indicating a medial view of the endites. Endite I is anterior
to endites II and III. The exopod is visible underneath the limb (on the lateral side of the
limb), and the medial side of the exopod is
visible. The medial side of the small basal
bristle, which is on the lateral side of the
limb, is visible just posterior to the exopod.
The orientation of the limb is comparable to
the left limbs illustrated in Figs. 2B and 3A,
which are not flattened under a cover slip.
Ancohenia robusta (Brady, 1890), USNM
193623, Instar III female, right limb (Fig.
6F): The location of the c-bristle indicates a
medial view of the endopod. The posterior location of the c-bristle indicates the anterior of
the endopod is towards the left. These characters identify a right limb with the anterior
of the endopod towards the right. The anterior bristle of the coxa indicates a medial view
of the coxa, which on this illustration is hidden underneath the precoxa. Endite III is recognized by its typical bristles. Endites I and
II (bristles not shown) are anterior to endite
III, indicating that the endites have their medial surfaces uppermost. The exopod is not
visible on this illustration. The orientation of
the limb is comparable to the left limbs il-
KORNICKER: ORIENTATION OF MAXILLA OF SARSIELLINAE
lustrated in Figs. 2B and 3A, which are not
flattened under a cover slip.
Chelicopia arostrata Kornicker, 1958,
USNM 158335, adult female, right limb (Fig.
5C): The location of the a-bristles indicates
that the endopod is towards the left, and the
location of the c-bristle indicates a medial
view. Therefore, the maxilla is a right limb
with the anterior towards the left. The anterior location of the anterodorsal coxa bristle
indicates a medial view of the coxa. The short
suture separating endites II and III identifies
the endites; endite I is clearly part of the precoxa. Endite I is anterior to endites II and
III, which indicates a medial view of the endites. The dashed exopodial lobe (three bristles not shown) is visible beneath the limb.
The orientation of the limb is comparable to
the left limbs shown in Figs. 2B and 3A.
Chelicopia arostrata Kornicker, 1958,
USNM 158335, adult female, left limb (Fig.
5B): The location of the a-bristles of the second endopodial segment indicates a lateral
view of the endopod with the anterior towards
the left, which identifies the limb as a left
limb. The anterior location of the anterodorsal bristle of the coxa indicates a lateral view
of the coxa. The exopod is posterior to the endopod 1 indicating a lateral view of the exopod. The short suture separating the two endites towards the left of the illustration, and
the fact that the third endite towards the right
is clearly part of the precoxa, identifies the
right endite as endite I, and the left endites
as endites II and III, with endite II between
endites I and III. The exopod is uppermost
relative to endite III. The posterior location
of endite I relative to the other two endites
is interpreted to indicate that the endites have
been folded out from their normal location
medial to the endopod, and that their medial
side is uppermost. There is no direct counterpart to the orientation of this limb on Figs.
2 or 3. The location of the endopod and exopod are similar to those segments in Fig. 2C,
and the location of those two segments as
well as the coxa are similar to those segments
in Fig. 3B. The location of the endites is closest to that shown for the endites in Figs. 2D
and 3C, which show anterior views of the endites.
Eusarsiella gettlesoni Kornicker, 1986,
USNM 156826, adult female, right limb (Fig.
6C): The location of the c-bristle of the second endopodial segment indicates a medial
277
view of the limb. The posterior location of the
a-bristles on the segment indicates that the anterior of the endopod is towards the left in the
illustration, which identifies the limb as a
right limb. Endite I is identified by its obviously being part of the precoxa. In their natural position endites are medial to the endopod and endite I is anterior to endites II and
III, contrary to the relative location of the endites on the illustrated limb. Therefore, the illustrated limb is interpreted as having the endites folded back to the right, indicating a
more-or-less anterior view of them. There is
no comparable view of this limb on Figs. 2
or 3. The locations of the illustrated endopod and coxa are similar to those in Figs. 2B
and 3A. The location of the illustrated endites
is similar to those in Figs. 2D and 3C.
Metasarsiella vibex Kornicker, 1991,
USNM 158310, adult female, right limb (Fig.
5A): The location of the a-bristles indicates
a lateral view of the second endopodial segment. The posterior location of the bristles
indicates that the anterior of the endopod is
towards the right, which identifies the illustration as a right limb. The anterior location
of the anterodorsal bristle of the coxa indicates a lateral view of that segment. The uppermost location of the exopod relative to the
endites indicates a lateral view of the exopod.
Endite III is identified by its typical number
and location of attached bristles. The anterior location of endite I relative to endites II
and III, and the location of the endites beneath the endopod suggest that the endites are
in their natural location, and therefore their
lateral sides are uppermost. The orientation
of this limb is comparable to the left limb illustrated in Figs. 2C and 3B.
ACKNOWLEDGEMENTS
I thank Molly K. Ryan (Smithsonian Institution) for
drawing Fig. 2, using the computer program Adobe Illustrator, and Jack Schroeder for inking my camera lucida drawings of appendages. I also thank Elizabeth Harrison-Nelson (Smithsonian Institution) for assisting in
preparing the paper for publication.
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RECEIVED: 3 February 2001
ACCEPTED: 4 September 2001.