Table Of ContentTHE UNIVERSITY OF KANSAS
PALEONTOLOGICAL CONTRIBUTIONS
MONOGRAPH 2
Living and Fossil Macrocyprididae
(Ostracoda)
Rosalie F. Maddocks
The University of Kansas Paleontological Institute
The University of Kansas, Lawrence, Kansas
February 27, 1990
ISSN 0278-9744
Library of Congress Cataloging-in-Publication Data
Maddocics, Rosalie F.
Living and fossil Macrocyprididae (Ostracoda) / Rosalie F.
Maddocks.
p. cm.— (The University of Kansas paleontological contribu-
tions. Monograph, ISSN 0278-9744; 2)
Includes bibliographical references.
1. Macrocypridida—Classification. 2. Macrocyprididae, Fossil.
I. Title. II. Series.
QL444.086M33 1990� 595.3'3—dc20� 89-78041
CONTENTS
PAGE
Aas-rRAcr �1
INTRODUCTION �1
Taxonomic principles and objectives �2
Procedures and conventions �4
History of project �6
Acknowledgments �7
ANATOMY OF MACROCYPRIDIDAE �8
Carapace �8
Appendages� 13
Body and genitalia �18
INTERPRETIVE MORPHOLOGY � 19
DEVELOPMENTAL BIOLOGY � 24
BIOGEOGRAPHY � 26
EVOLUTION � 38
SYSTEMATIC DESCRIPTIONS � 41
Family Macrocyprididae Müler, 1912� 41
Genus Macrocypris Brady, 1868 � 42
Genus Macromckenziea n. gen. � 48
Genus MacroMxis n. gen. � 57
Genus Macrocypria Sars, 1923 � 80
Genus Macrocypnssa Triebel, 1960 � 84
Genus Macrosansa n. gen. � 87
Genus Macroscapha n. gen. � 95
Genus Macrocyprina Triebel, 1960 � 108
REFERENCES � 131
APPENDIX I: Some Other Species Referrable to Macrocyprididae � 143
APPENDIX II: Some Species Excluded from Macrocyprididae � 145
APPENDIX III: List of Collections Studied � 149
APPENDIX IV: List of Numbered Specimens � 161
EXPLANATION OF FIGURES � 179
EXPLANATION OF PIATES � 273
INDEX� 403
ILLUSTRATIONS
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TEXT-FIGURES PAGE
1. Postulated homologies of macrocypridid, pontocypridid, cypridid, bythocy-
pridid, and bairdid adductor muscle scars � 13
2. Locations of BLM collecting stations on the middle Atlantic outer continental
shelf � 26
3. Locations of colecting stations in the northwestern Gulf of Mexico� 28
4. Collecting stations and geographic distributions of species in the western
Atlantic, Caribbean, Gulf of Mexico, and central eastern Pacific oceans� 29
5. Collecting stations and geographic distributions of species in the southeastern
Atlantic Ocean� 30
6. Collecting stations in the southwestern Atlantic and southeastern Pacific
oceans � 31
7. Geographic distributions of species in the southwestern Atlantic and south-
eastern Pacific oceans� 32
8. Collecting stations and geographic distributions of species in Antarctic waters 33
9. Bathymetric and latitudinal ranges of Macroscapha turbida � 33
10. Collecting stations and geographic distributions of species in the Mozam-
bique Channel and western Indian Ocean� 33
11. Colecting stations around Nosy Be, northwestern Madagascar � 34
12. Colecting stations in the Strait of Magelan and nearby region � 35
13. Collecting stations and geographic distributions of species around Australia
and New Zealand � 35
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FIGURES 1-80 following page 189
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PLATES 1-114 following page 285
TABLES
1. Numbers of living adult males and females in each species and genus of
Macrocyprididae � 21
2. Total number of specimens of three species of Macrocyprididae at BLM
stations on the middle Atlantic continental shelf for al seasons� 27
3. Total number of specimens of three species of Macrocyprididae at BLM
stations Al through G7 on the middle Atlantic continental shelf for each of
the eight collecting seasons � 27
4. Total number of specimens of three species of Macrocyprididae at BLM
stations Hl through L6 on the middle Atlantic continental shelf for each of
the four colecting seasons � 28
5. Total number of living specimens of 10 species at stations in the southeastern
Atlantic Ocean � 30
6. Numbers of species of Macrocyprididae found in 479 living and dead (but not
fossil) samples � 37
7. Bathymetric distribution of 207 samples yielding live Macrocyprididae for this
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study 37
vi
Now, clearly if we are going to get sense into
the classification of
ostracods we must use both the characters of the carapace and
the characters of the soft parts, and the
mystery lying behind
classification must always be which characters are
evolving and
which are remaining static in the phylogeny;
we need to know
which are the conservative characters, the
ones which mark
common ancestry, and which characters are evolving quickly
and therefore adapting themselves to environment
and likely to
produce convergences. Now as far as I can make
out, there is
not one character in the ostracods
which does not sometimes
behave in one way and sometimes in the
other. You cannot lay
down general rules of what is important in classification and
what is not.
Peter C. Sylvester-Bradley, 1969, p. 245
LIVING AND FOSSIL MACROCYPRIDIDAE (OSTRACODA)
Rosalie F. Maddocks
Department of Geosciences, University of Houston, Houston, Texas 77204-5503
ABSTRACT
This comprehensive taxonomic revision of the family Macrocyprididae is based on
the analysis of 138 species, of which 123 are living and 15 are fossil. Thirty-six previously
named species are redescribed from type and additional specimens, 54 new species are
named, and 48 additional species are described in open nomenclature. Appendage and
genital characters are described for 73 of these species. A review of more than 100
additional species that have at one time or another been classified in Macrocyprididae
suggests that at least 23 are likely to belong there, bringing known diversity to at least 161
species. Seven species names (Macrocyprina decora, Mn. hieroglyphica, Mn. maculata,
Macrocypris cala bra, M. compressa, M. elongata, M. trigona) should be abandoned as namina
dubia.
Eight genera are recognized, of which four are existing or elevated from subgeneric
rank (Macrocypris, Macrocy pria, Macrocyprissa, Macrocyprina) and four are new (Macropyxis,
Macramckenziea, Macrosarisa, Macroscapha). Paramacrocyprisi s shown to be a synonym of
Macrocyprissa. Pseudamacrocypris is referred to the Pon tocyprididae as a close relative of
Propontocypris. Novocypris is a marine cypridid related to Paracypris. Premacrocyprisa nd the
Acratiinae, so far as known at present, lack the diagnostic features of Macrocyprididae as
here revised.
The Macrocyprididae are morphologically homogeneous and conservative with com-
pellingly cypridacean rather than bairdiacean affinities. No Paleozoic or Triassic and
only two Jurassic species can be identified, but Cretaceous and Cenozoic species abound.
There are many more species with correspondingly narrower geographic ranges than
previously recognized. There may be several hundred living species of Macrocyprididae,
which are most abundant on coral reefs, near the edges of continental shelves, in the
oxygen-minimum zone, in bathyal and abyssal depths, and beneath cold or upwelling
surface currents, wherever high biological productivity produces food-rich bottom sedi-
ments. Adult and juvenile anatomy, development of late instars, sexual dimorphism,
size-depth clines, and geographic and bathymetric distributions are analyzed.
the entire family, and several of the constituent species
INTRODUCTION
were reported worldwide. Unlike the burgeoning com-
The family Macrocyprididae is the smallest and most plex of Paracypridinae, the Macrocyprididae have not
homogeneous of the three families that make up the su- adapted to low or fluctuating salinities, and they are poorly
perfamily Cypridacea. Although globally ubiquitous in represented in most coastal habitats. There is also no
nearly all euhaline environments, the family is rarely rep- evidence for any commensal or ectoparasi tic associations
resented at any one location by more than one or two with larger invertebrates, such as those displayed by some
species, and those species are never predominant and of the Pon tocyprididae. Instead, they are solitary, sluggish,
rarely common constituents of the local podocopid ostra- sediment-ingesting bottom-crawlers. They tend to be rela-
code fauna. The external morphological features are re- tively large and include the largest known marine
markably constant, so much so that until 1960 a single podocopid ostracodes.
genus was considered sufficient to comprehend almost Like other podocopid Ostracoda, populations of
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2 The University of Kansas Paleontological Contributions—Monograph 2
macrocypridid species show considerable individual vari- family rank, coequal with the Pontocyprididae and the
ation, analysis of which must precede species identification. Cyprididae (Alm, 1915; Sars, 1922-1928; Maddocics, 1969a;
The right and left valves are distinctly asymmetrical, and Danielopol, 1976). Indeed, their conservatism of mor-
the shape of the larger right valve is more distinctive and phology and habitat provides a refreshing contrast to the
representative of the species than is that of the smaller left unruly Cyprididae, which have swarmed over nearshore
valve, most characters of which can be predicted from the marine, brackish-water, freshwater, subterranean, and
right valve. Both sexes are present, and male and female damp subaerial habitats. The extraordinary breadth of
carapaces may show either size or shape dimorphism or adaptive radiations and frequency of convergence within
both. Juveniles develop through a series of eight instars the Cyprididae have generated disproportional inflationary
before reaching the adult size and shape (Müller, 1894), tendencies in the classification of Cypridacea, such as the
up to six of which have been recognized in the material proposed segregation of marine representatives into a
studied here. Late instars have a fairly broad calcified separate "Family Paracyprididae" (see McKenzie, 1979,
inner lamella, broader than that found in comparable 1982 for rationale) or the proposed isolation of supralit-
developmental stages of other podocopid ostracodes toral invaders into a "Family Terrestricyprididae" (Schor-
though not as wide as in the adult. This makes it easy to n ikov, 1980). The Pon tocyprididae also show prolific eco-
mistake isolated juveniles for adults of the same or a differ- logical diversification, encompassing phytal, soft-bottom
ent species and has contributed to past misidentifications. crawling and burrowing, ectoparasitic or symbiotic, inter-
Species of Macrocyprididae also show variability in size stitial, anchialine, xylophile, and many other specializa-
and shape between populations from different depths and tions; they occupy the full range of marine habitats from
geographic locations, which has further contributed to the strandline to the deep sea. By contrast, the Macrocy-
difficulties in identification. This unreliability of species prididae seem monotonous, even humdrum, in their ar-
recognition has hindered use of the family in paleoeco- chaic morphology and lack of ecomorphotypic innova-
logic and biostratigraphic interpretation. tion. Nevertheless, from the perspective of one who works
This is a family of respectable antiquity. There are with the Macrocyprididae the tripartite division of the
probable records in the Jurassic, and by the Cretaceous it Cypridacea is plain.
was already very diverse. The family has remained stub-
bornly conservative, with relatively few morphological
TAXONOMIC PRINCIPLES AND OBJECTIVES
innovations. The adductor muscle-scar pattern, for ex-
ample, remains curiously devoid of taxonomic significance Zoological classification attempts to find order in the
below the family level. Though complex, with more con- diversity of animals and to express their inferred evolu-
stituent scars than any other cypridacean taxon, this scar is tionary relationships (Mayr, 1969). Although the func-
invariant in its general features, while displaying vigorous tional unit in nature and in taxonomy is the species, which
but apparently meaningless variability in its minor details. is essentially a genetic and ecologic phenomenon, for
It offers neither the distinctive rearrangements of a few ostracode species we know little of the genorne except its
discrete scars seen in the Cyprididae and Pon tocyprididae morphologic expression. By their functions, skeletons are
nor the repetitive fusion or fission of individual scars within both conservative and subject to homcomorphy, so that
a constant spatial deployment that characterizes Cyther- evolutionary hypotheses based on only skeletal characters
acea and Bairdiacea. are likely to oversimplify the true history. In Ostracoda,
It is this deceptively primitive scar pattern that has fortunately, the skeleton is part of the external integu-
caused some workers to link the Macrocyprididae with the ment of the body and therefore likely to have more nu-
Bairdiacea rather than the Cypridacea. There is no other merous biological functions and more taxonomically use-
anatomical evidence for such a classification, of course, in ful characters than a mere external secretion or test. Even
the appendages, genitalia, or carapace characters. None- so, there are all too many podocopid ostracodes with
theless, some macrocypridid scar configurations invoke smooth, elongate carapaces that are almost featureless
homologies with basic pontocypridid, cypridid, and bairdiid externally. For these cypridiform taxa, extra emphasis
scar patterns. If real, these homologies would sustain a must be given to the few available distinguishing features,
unitary, five-clade concept of the living Podocopida (Mad- chiefly valve overlap, hingement, muscle scars, pore ca-
docks, 1976, 1982), as contrasted to the two- or three-clade nals, and marginal structures. In the case of the Macrocy-
model espoused by McKenzie et al. (1983) and Grande' prididae these internal details are nearly invariant as well,
(1969, 1978). In other words, it. is the Macrocyprididae while the ineffable shape differences are extremely difficult
through which it is easiest to trace connections of Cypri- to define, communicate, and evaluate. It is also not easy to
dacea with the other four living branches of the recognize male, female, and juvenile carapaces, all of which
Podocopida. must be included in a natural species concept. As a result,
Within the Cypridacea the Macrocyprididae stand well most species have not been consistently recognizable, and
apart from the other two main branches and deserve full the diversity of Macrocyprididae has been underestimated.
�
Maddocks—Living and Fossil Macrocyprididae
3
For optimal taxonomy, both carapace and soft-part the classic problems of ostracode
taxonomy. Develop-
characters should be used, and it cannot be denied that mental stages and sexual dimorphs have
not always been
the species and genera of Macrocyprididae are most easily clearly distinguished within local
populations of a species,
recognized by using a combination of both. A primary while depth and other components of geographic variabil-
strategy of this revision has been to incorporate as much ity among populations of a species
have never been ana-
information as possible from the external soft anatomy, lyzed. Of the four genera and subgenera
(Macrocypris,
mostly appendages and genitalia, to supplement that from Macrocypria, Macrocyprissa, Macrocyprina) established before
the carapace. In this way it has been possible to distin- this study, one had only a single fossil species
(Mc. cy/in-
guish males and females, adults and juveniles, and prob- dracea), and another contained only a
single living species
able species with much less uncertainty than if carapace (Ma. Hirst) (see p. 5 for abbreviations). For the third and
traits alone were used. With this larger pool of characters, fourth, only two species had been fully described for each
it then becomes possible to calibrate, as it were, those (M. minna, Mx. sapeloensis, Mn. propinqua, Mn. succinea),
carapace traits that show high correlations with the soft- although partial or inaccurate soft-part information had
part features and to use them as reliable diagnostic charac- been published for a few other species. The relatively
ters for fossils. small number of species, as was then supposed, their con-
Experience with Bairdiidae and Pontocyprididae has servative carapace characters, and their very broad re-
suggested that trends or relationships marked by obvious ported geographic ranges made it seem feasible to at-
carapace features are always corroborated by soft-part tempt a comprehensive, global review of this family.
characters, when the latter become available, and that Therefore, this taxonomic revision of the family
Macro-
trends determined from soft parts are always con fi rmed by cyprididae was undertaken with the following
objectives:
carapace traits, albeit often subtle ones that had been (1) to describe and analyze the ontogenetic development
overlooked previously or that are ambiguous when taken of soft parts and carapace, especially to find criteria
for
alone. The conservative carapace of Macrocyprididae reliable distinction of adults from late instars, and also to
makes the family ideal for further testing of this thesis. evaluate the likelihood of postmaturational molting;
(2)
Indeed, any macrocypridid species diagnosed by soft-part to describe and recognize sexual dimorphism, if
present,
characters also turns out to be recognizable from the in carapace and soft parts; (3) to examine the variability of
carapace features, and vice versa. This familiar but often carapace and soft parts within local populations in order
queried premise has been recon fi rmed for the 73 macro- to evaluate the relative taxonomic usefulness of carapace
cypridid species represented by soft parts in this study, and soft-part characters; (4) to examine the
between-
making it possible then to describe many additional spe- population variability within species, especially to analyze
cies from empty and fossil carapaces with only slightly less the possible influence of depth on size
and shape; (5)
confidence. from these insights, to develop regionally applicable spe-
Genera are clusters of species thought to share com- cies concepts that permit
accurate recognition of biological
mon ancestry, but most ostracode genera have been pro- species; (6) to redescribe in full, from type specimens
and
posed initially with too few species. Even though most additional material, the carapace and soft parts of the
names have proved to be useful later on, as newly discov- previously named species of living Mac
rocyprid idae; (7) to
ered species leafed out this branch of the evolutionary describe in full the carapace and soft parts
of a sufficient
tree, rarely have the original taxonomic diagnoses proved number of new living species to provide a
satisfactory
satisfactory. Undue emphasis on one or a few key taxo- estimate of taxonomic diversity and trends in
this family;
nomic characters has hindered phylogenetic analysis. (8) to establish sound generic concepts, both by revising
Indeed, only after most of the member-species are known previously named genera and by establishing new genera,
can the shared-derived attributes of a taxon be listed; and as necessary and appropriate to express the taxonomic
because each lineage evolves independently of others, the and probable evolutionary affinities of these species; (9)
more numerous the species the fewer attributes are likely to apply numerical taxonomic methods to the resulting
to be shared by all. Again, the twigs-on-a-branch analogy compilation of character traits for a large number of spe-
reminds us that a genus should be defined, not by the cies; (10) to determine which carapace characters are
boundaries of the morphological space those species oc- suitable for accurate, consistent identification and generic
cupy but by their common origin. To attempt to specify, a classification of fossil species; (11) to test the applicability
priori, which anatomical structures should have taxonomic of these findings by redescribing and reclassifying a
few
value at any particular hierarchical level (Delleckker, 1979) previously named fossil species; (12) to compile ecological
is futile. Most ostracode genera and higher taxa prove to information for living species, for possible future applica-
be polythetic, with rampant mosaic evolution, as Sylvester- tion in paleoecologic interpretations; (13) to ascertain the
Bradley (1969) , Dan iclopol (1976), Maddocks (1976), geographic distributions of living species and their pos-
McKenzie (1982), and others have warned. sible historical implications; (14) to review the stratigraphie
The family Macrocyprididae presents in a microcosm occurrences of fossil species, with possible biostratigraphic
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4 The University of Kansas Paleontological Contributions—Monograph 2
and evolutionary implications; and (15) to evaluate pres- tain more than three points of contact of a specimen with
ent evidence relevant to the origin and evolution of the stage, producing charging and poor resolution. Elec-
Macrocyprididae, and especially to resolve, finally, the tronic problems caused distortion of the left edge of many
question of whether they have cypridacean or bairdiacean images, distorting lateral outlines slightly, and therefore
affinities. most of the SEM photos taken are not published here. As
for most smooth ostracodes, SEM micrographs proved to
be less satisfactory than transmitted-light photographs.
PROCEDURES AND CONVENTIONS
The drawings were done with a mirror mounted on a
Dissections.—Most of the specimens collected living had Leitz research petrographic microscope with a high-inten-
been preserved in alcohol and were not stained. First the sity xenon lamp, which projects the image onto the draw-
assemblage was transferred to a drop of glycerin, and all ing surface. In a darkened room the desired structures
specimens were counted, measured, and the sex deter- can be accurately traced by pencil and then inked. All
mined if possible. One or more representative specimens proportions and structures shown in the drawings are
of each sex, growth stage, and geographic population traced from actual specimens. However, all legs are shown
were then selected for dissection. With fine needles, the from the left side to facilitate comparisons, and for com-
appendages and external genitalia were teased from the plex structures or crumpled specimens some structures
body, in the same drop of glycerin that became the mount- may have been moved slightly, separated, or straightened
ing medium for the dissection-mount slide. The valves out in the drawing to clarify their relationships. Because
were washed with water, dried, measured again separately, all drawings were done after mounting, some structures
and stored in paper micropaleontological slides. may be slightly displaced or distorted from life position by
Decalcified or very fragmented carapaces were mounted the overlying cover glass. Nevertheless, the resulting
on the dissection slide. Years later, when much of the configurations are standard among such mounted mate-
glycerin had evaporated from these unsealed slides, it was rial and readily comparable with each other. Broken or
necessary to remove the cover slip and add additional missing structures are represented by dotted or dashed
glycerin or in some cases to transfer all body parts to a new lines. Some minute structures that could not be drawn
slide mount. The slides were then sealed with Murrayite. accurately at the scale of the drawings, such as the fine
Photographs and drawings.—The carapace and soft parts secondary hairs or barbs that characterize certain setae,
of most dissected specimens were photographed with a have been omitted. To assure standard orientations and
Leitz Orthomat automatic camera microscope, to provide comparability of shapes, nearly all drawings of valves are
a permanent record. Kodak Panatomic-X film was devel- external views, with internal features shown as seen from
oped in Kodak Microdol-X or D-19, the latter to increase the exterior in transmitted light. All drawings were done
contrast for soft parts. Only a small number of those many at a constant magnification, so that the range of sizes on
thousands of photographs could be included in this re- the figures presents the true size ranges of these species.
port. Photographs were taken in plane and cross-polar- Contrary to usual taxonomic procedures, in this mono-
ized transmitted light, reflected light, a combination of graph the drawings and photographs are grouped by struc-
these, or Zernike phase contrast illumination. For wet ture rather than by species. The morphologic conserva-
valves, the transparency and depth of field improve with tism of the Macrocyprididae is such that in isolation many
immersion in glycerin. Dry valves may be wetted with of the species look pretty much alike. Only a close com-
water or ethanol and immersed again in glycerin, but it is parison shows subtle differences of shape or details of soft
then more difficult to prevent air bubbles from being parts. To facilitate such comparisons and to emphasize
trapped in the vestibules, especially in specimens of Macro- the differences, or in some cases the lack of significant
/xis with capacious air-filled radial pore canals. Such air differences, the illustrations of the same structures of closely
bubbles can be seen in the plates, as well as surfaces that related species are grouped together on the same figures
resisted wetting, dirt, and cracks or other flaws, especially and plates, approximately in taxonomic sequence. As
in the older type specimens. Adhering epidermis and much as possible, legs are illustrated in standard, lifelike
uncalcified inner lamella also are visible in many photo- orientations and as viewed from the left side. For valves,
graphs of live specimens. These unretouched photographs right valves are always on the left-hand pages and left
were printed at standard sizes but varying magnifications valves of the same species in corresponding positions on
to facilitate comparison of shapes among species and geo- the facing right-hand pages. The photographs of valves
graphic populations. Within each species, however, the and genitalia include a selection of specimens from differ-
magnifications of the published figures are constant or ent geographic localities, to illustrate geographic variabil-
ity in size and shape as well as sexual dimorphism, and to
nearly so.
SEM micrographs were taken with a Cambridge ma- portray a population concept for the species.
chine at 5 kV and 00 . The relatively large size of deep-sea Specimen catalog and repositories.—A total of 6733 speci-
macrocypridids and the greatly protruding hinge elements mens was examined. Specimens borrowed from museums
and ventral bow-shaped processes made it difficult to oh- or individuals have been returned to that museum or
