New and poorly known species of Bairdoppilata and Paranesidea (Bairdiidae, Ostracoda) from French Frigate Shoals and O’ahu, the Hawaiian Islands

Abstract

Bairdoppilata scaura, n. Sp. and five species of Bairdoppilata and Paranesidea in open nomenclature are described from encrusting communities on French Frigate Shoals and Kane'ohe Bay, Oahu, Hawaiian Islands. Some poorly documented anatomical traits (carapace setae, hingement, antennal claws, genitalia) are examined for their potential taxonomic significance, in order to confirm the coherence of the Genus Bairdoppilata and to explore its diversity.

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Accepted by R. Matzke-Karasz: 9 Nov. 2015; published: 22 Dec. 2015
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New and poorly known species of Bairdoppilata and Paranesidea (Bairdiidae,
Ostracoda) from French Frigate Shoals and O’ahu, the Hawaiian Islands
ROSALIE F. MADDOCKS
Department of Earth and Atmospheric Sciences, Room 312 Science & Research Building 1, University of Houston, Houston, TX
77204–5007. E-mail: rmaddocks@uh.edu
Abstract
Bairdoppilata scaura, n. sp. and five species of Bairdoppilata and Paranesidea in open nomenclature are described from
encrusting communities on French Frigate Shoals and Kane’ohe Bay, Oahu, Hawaiian Islands. Some poorly documented
anatomical traits (carapace setae, hingement, antennal claws, genitalia) are examined for their potential taxonomic signif-
icance, in order to confirm the coherence of the Genus Bairdoppilata and to explore its diversity.
Key words: Ostracoda, Bairdiidae, Bairdoppilata, Paranesidea, Hawaiian Islands, chaetotaxy, zygum
Introduction
The relationships of Pacific Bairdiidae (Ostracoda, Podocopida) are poorly delineated at present, because most
species have been described on the basis of empty carapaces from dried sediment samples, and little is known of
their soft anatomy and life habitats. The taxonomic status of species of Bairdiidae reported from the Hawaiian
Islands was summarized by Maddocks (2013), who also redescribed Neonesidea tenera (Brady, 1886) and
described four new species of Neonesidea from French Frigate Shoals. The following report describes six
additional species of Bairdiidae from the same suite of samples, which are assigned to the genera Bairdoppilata
and Paranesidea.
The samples represent spur and groove environments on the barrier reefs of French Frigate Shoals, as well as
patch reefs and coastal fouling communities of Kâne'ohe Bay, O’ahu. The specimens were collected by hand
scraping from hard surfaces, coralline rubble and macroalgae. The sampled localities are described in Appendix I
and Figure 1. For more maps of the localities and description of collecting methods see Coles et al. (2002a, 2002b),
DeFelice et al. (2002), and Kornicker et al. (2007). The samples are part of the same collection from which
Kornicker et al. (2007) reported the Myodocopina Ostracoda.
French Frigate Shoals is an atoll, consisting of a 32-km long, crescent-shaped reef and lagoon on an almost
submerged seamount in the Hawaii-Midway chain, 900 km northwest of Honolulu and about the same distance
southeast of Midway Island (Fig. 1). It is part of the Hawaiian Islands National Refuge and a National Marine
Sanctuary within the Papahânaumokuâkea Marine National Monument.
Kâne'ohe Bay is a sheltered embayment of about 55 km
2
behind a barrier reef, located on the northeast shore of
the island of O'ahu. This ecosystem has been influenced over the last century by coastal population growth, a
Marine Corps base, shipping and aquaculture, and it now includes a substantial number of invasive, nonindigenous
invertebrate and algal species (Coles et al. (2002a).
Four species of Bairdoppilata and two species of Paranesidea are represented in these collections (Table 1,
Fig. 2). The number of individuals per sample is small, and many are females and juveniles. Unfortunately, all
specimens are partly or entirely decalcified by storage in preservative, and many have secondary crystalline
encrustations and infillings (Figs. 13A, 17K, 20A, 21A). None could be observed dry or by SEM. The
identification of such material is troublesome, but it is unwise to ignore the fragmentary information that they can
provide. It is hoped that entering these incomplete observations into the published record will facilitate more
informed comparisons in the future.

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FIGURE 1. Maps for localities sampled. A, Bathymetry of French Frigate Shoals; numerals indicate stations collected (see
Appendix I). B, Map of O'ahu Island, to show location of Kane'ohe Bay. C, Map of Kane'ohe Bay; numerals indicate stations
collected (see Appendix I). D, French Frigate Shoals and O’ahu are part of the Hawaii-Midway island-seamount chain, which
trends west-northwest across 2,500 kilometers of the North Pacific Ocean.

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TABLE 1. Species distributions of Bairdoppilata and Paranesidea by station on French Frigate Shoals and Kane’ohe
Bay, the Hawaiian Islands, with carapace dimensions. See also Figures 1 and 2 and Appendix I.
Explanation of abbreviations: NAME = name or informal identification; SPECIMEN = Maddocks specimen number;
STATION = collecting locality (see Appendix I for details); LENGTH = carapace length in μm; HEIGHT = carapace
height in μm; M = adult male; F = adult female; J = juvenile (instar).
The genus Bairdoppilata has generally been diagnosed by just two key characters, the supplemental locking
dentition below the hinge and the two terminal claws of the antenna. Because only the first of these features is
visible in the carapace, there was formerly some doubt about the validity of the genus (Morkhoven 1958, 1963). A
broader array of characters is desirable to support a sound genus. Because of homeomorphy, it can be challenging
to discover traits that are reliable indicators of monophyly in bairdiid Ostracoda. Here, several poorly understood
anatomical characters (including the hinge, appendages, genitalia and carapace setae) are reviewed, to evaluate
their consistency and taxonomic potential. A few comparative observations are offered for Paranesidea, as well,
although a systematic treatment of that genus is not possible with the few specimens at hand.
Ecologically, the closest counterparts to these collections are in far-distant settings. Maddocks (1969, 1995)
collected living podocopid Ostracoda by hand from many shallow microhabitats on the subtidal terrace around
Nosy Be, Madagascar, including washings of algae, sea grasses, sponges, corals, coralline rubble and coarse sand
between patch reefs. Bairdiidae were a prominent component (23 species) of those assemblages. Hartmann
sampled encrusting algae from tide pools and other littoral and shallow sublittoral habitats around the coasts of
Africa and Australia (Hartmann 1974, 1978, 1979, 1980, 1981; Hartmann-Schröder & Hartmann 1974). His
collections from algal and coralline substrates of Huahin and Rangiroa in the Society Islands and the Tuamotu
Islands of Polynesia included Neonesidea but no other genera of Bairdiidae, and his descriptions of marine
interstitial faunas of sandy beaches of Hawaii included no Bairdiidae (Hartmann 1984, 1991).
Hartmann (1984, 1988) stated that the most likely mode of long-distance dispersal of shallow benthic

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Ostracoda is by floating seaweeds, a view endorsed by Teeter (1975), Cronin (1988), and Titterton & Whatley
(1988), among others. Algal inhabitants, he said, have lower rates of endemism than sediment-dwellers and are
more suitable for zoogeographic analyses. He stated that species of the “extraordinarily difficult Bairdiidae have
very similar carapace shapes and can be distinguished only by side-by-side comparison of actual specimens, unless
male soft parts are available (1984, pp. 124, 142). For this reason, he considered the reported occurrences of
bairdiids to be of limited usefulness in his analysis of Pacific zoogeography (Hartmann 1988). Whether in
accordance with this prediction or in consequence of the isolated location of the Hawaiian Islands, none of the six
species of Bairdoppilata and Paranesidea described here can be identified with a named species.
Holden (1967) described six species of Bairdia from sediments dredged from drowned terraces of the
Hawaiian Islands, ranging in depth from 476 to 650 m, none of which can be confidently assigned to Bairdoppilata
or Paranesidea. Holden (1976) reported eight Cenozoic species of Bairdoppilata and Paranesidea from two drill
holes on Midway Island. Those generic assignments are poorly supported, and none of the fossil species are
recognized in the material described here.
Depository. All type and identified specimens will be deposited in the Bishop Museum, Honolulu.
Abbreviations. LV = left valve, RV = right valve, MSP = adductor muscle scar pattern, RPC = radial pore
canal, NPC = normal pore canal. Podomeres of limbs are numbered I to VI, proximal to distal.
Methods
The degrees of slope of anterodorsal and posterodorsal margins were measured on external lateral views of the
carapace or LV, with 0
o
being horizontal. If the anterodorsal or posterodorsal corner is indistinct, it was inferred to
be located just outward diagonally from the corresponding end of the hingeline. The maximum range of rotation of
the distal antenna (podomeres IV–VI) was determined from measurements on published drawings of the antenna,
oriented with anterior toward the right. The dorsal margin of Podomere III was set as horizontal (90
o
), and angles
were measured clockwise with respect to vertical (0
o
). The lengths of distal antennal claws were measured on
published drawings, corrected for magnification, and reported as a percentage of carapace length of the same
specimen. The chaetotaxy of thoracic legs was recorded as presence/absence of setae at specified locations in
published drawings.
Taxonomy
Order PODOCOPIDA Müller, 1894
Superfamily BAIRDIOIDEA Sars, 1888
Family BAIRDIIDAE Sars, 1888
Genus Bairdoppilata Coryell, Sample and Jennings, 1935
1935 Bairdoppilata Coryell, Sample and Jennings: 3.
1969 Bairdoppilata (Bairdoppilata) Coryell, Sample and Jennings—Maddocks: 66.
1995 Bairdoppilata Coryell, Sample and Jennings—Maddocks: 215.
History. More than 100 nominal species have been classified in Bairdoppilata (Kempf 1986, 1995, 2004).
Carapaces are recognizable by the accessory (bairdoppilatan) locking dentition, and the living animals have two
scissors-like, terminal antennal claws. Because fossil representatives, including the Miocene type species, have
been identified chiefly by the supplemental dentition, there was some doubt about the value of the genus for several
decades (Morkhoven 1958, 1963). Confusion arose because supplemental dentition is also well expressed in
Glyptobairdia coronata. Shaver (1961, p. 205, fig. 140) recognized Bairdoppilata in the Treatise, but
Glyptobairdia was treated as a synonym of Triebelina. The taxonomic history of living ornate bairdiids has been
reviewed by Bold (1974), Malz & Lord (1988), Maddocks & Wouters (1990), and many others cited therein. Bolz

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(1969, 1971) reviewed the prevalence of auxiliary dentition in Triassic genera of Bairdioidea. He was the first to
explain its functional significance as a locking mechanism to reduce torque in high-arched carapaces, which
increases the likelihood that this character may be subject to convergence. He emphasized the importance of
considering all morphological features in a taxonomic diagnosis, rather than just one key character: “No single
morphological feature seems alone sufficient to establish higher systematic units” (Bolz 1971, p. 725).
Brady (1880, Pl. 3, figs. 2b, 3a) illustrated the two distal antennal claws and seven furcal setae of B. villosa,
remarking on the prominent barbs of seta 2. Tressler (1949, Figs. 5, 8) illustrated the furca and antenna of B.
cushmani but did not mention the two distal claws. Rome (1960) provided a meticulous description of the soft parts
of G. coronata, which was then classified in Triebelina, but did not evaluate its distinctive features. It was
Kornicker (1961, examining B. carinata, a synonym of B. cushmani) who pointed out distinctive attributes of the
furca and hemipenis that might support a generic diagnosis, although he did not mention the antenna. Kornicker
was also the first to realize the taxonomic significance of the patch pattern of the carapace, which is often preserved
in fossils. Maddocks (1969) proposed an expanded concept for the genus Bairdoppilata, which incorporated these
and other characters of the soft parts and carapace, and which has proved to be sufficiently flexible to
accommodate other new species as they turned up.
Species included. The genus Bairdoppilata in its broadest usage includes at least three ecological groups of
living species. The most diverse cluster consists of relatively small, mostly punctate species in tropical reef and
phytal assemblages. The soft parts have been illustrated (at least in part) for the following shallow-water species of
Bairdoppilata (listed alphabetically by original binomen):
Bairdoppilata (Bairdoppilata) alcyonicola Maddocks, 1969 (Nosy Be, Madagascar)
Bairdoppilata angolensis Hartmann, 1974 (Angola) [The published range (minimum to maximum) for carapace
length is 30 µm, but for height the range is 110 µm. This is probably an error.]
Bairdoppilata balihaiensis Hartmann, 1978 (Northwest Australia)
Bairdoppilata (Bairdoppilata) cratericola Maddocks, 1969 (Nosy Be, Madagascar)
Nesidea cushmani Tressler, 1949 (Florida, Bahamas) [= Bairdoppilata carinata Kornicker, 1961 by Maddocks
1969]
Bairdoppilata cytheraeformis Hartmann, 1974 (Angola)
Bairdoppilata geelongensis Hartmann, 1980 (South Australia)
Bairdoppilata mocamedesensis Hartmann, 1974 (Angola)
Bairdoppilata portsamsonensis Hartmann, 1978 (Northwest Australia) [The identity of this species is in doubt,
because more than one species and genus are represented among the published illustrations. The RV belongs to
a species of Bairdoppilata (Hartmann 1978, Figs. 13, 24), but Fig. 14 shows a LV of Neonesidea with caudal
setae, and Fig. 25 shows a LV of Paranesidea with shield-shaped patch pattern. It is obvious that this RV could
not fit inside either LV. Hartmann compared the hemipenis to that of B. balihaiensis, but his Fig. 31 shows
many differences. The long copulatory tube ending in a tight coil is more appropriate for Neonesidea.]
Bairdia simuvillosa Swain, 1967 (reported from the Gulf of California) [The published illustrations of appendages
are from a living female specimen with scissors-like antennal claws, collected in Scammon Lagoon on the
Pacific side of Baja California. It was illustrated as Fig. 32a–i of Swain (1967) but probably drawn by Kenneth
G. McKenzie. The published illustrations of the valves do not belong to Bairdoppilata but to species of
Neonesidea (Swain, 1967, Figs. 30c, d; Plate I, figs. 2a–f, 8). In a separate paper, McKenzie and Swain (1967,
Pl. 30, fig. 1) reported Bairdia simuvillosa in Scammon Lagoon, providing a photograph and more plausible
carapace dimensions, but they did not mention whether it has bairdoppilatan dentition.]
Bairdoppilata sinusaquilensis Hartmann, 1979 (Southwest Australia, also reported by Hartmann (1980) from
South and Southeast Australia)
Bairdoppilata sp. 2 of Maddocks, 1969 (Northwest Madagascar near Nosy Be)
Bairdoppilata? sp. 2 of Maddocks, 1975 (Ascension Island)
A second species-group of ?Bairdoppilata has been reported from sediment samples in deeper and colder
water. The carapace is large and nearly smooth. The accessory dentition of the hinge is developed in some
populations but inconspicuous or undeveloped in others. Although Maddocks (1969, 1995) suggested that perhaps
these species should eventually be classified in a new genus, it would be difficult to diagnose the genus on the basis

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of present knowledge. The species-level taxonomy is confused. In several cases the anatomical information is
taken from doubtfully identified specimens collected at a great distance from the type locality, published at a time
when taxonomists were inclined to underestimate taxonomic diversity in the deep sea. Brandão (2008) reviewed
the complex history and numerous misidentifications of several of these species in the Southern Ocean and
provided important new anatomical, taxonomic and zoogeographic information. The soft anatomy has been
described, at least in part, for the following nominal species (listed by original binomen):
Bairdia simplex Brady, 1880 (Challenger station 151, off Heard Island, Southern Ocean) [The appendage
descriptions by Maddocks (1969) apply to specimen USNM 121347 from Eltanin station 418, near the
Antarctic Peninsula, and specimen USNM 121348 from Eltanin station 1345, in the Pacific sector of the
Southern Ocean. The identifications of those specimens require verification.]
Nesidea labiata Müller, 1908 (Gauss Station, Southern Ocean)
?Bairdoppilata sp. 1 aff ?B. labiata of Brandão 2008 (Southern Ocean)
?Bairdoppilata sp. 2 aff ?B. labiata of Brandão 2008 (Weddell Sea)
Bairdia hirsuta Brady, 1880 (Challenger station 300, near Juan Fernandez Island in the Southeast Pacific Ocean)
[The appendage descriptions by Maddocks (1969, 1973) apply to specimen USNM 121353 from the Gulf of
Mexico and specimen USNM 139891 from Eltanin station 25, near the Galapagos Islands. The identifications
of those specimens require verification.]
Bairdia villosa Brady, 1880 (Challenger station 149, off Kerguelen Island, Southern Ocean) [The appendage
descriptions by Maddocks (1969) apply to specimen USNM 121344 from Eltanin station 418 near the
Antarctic Peninsula. That identification requires verification.]
?Bairdoppilata sp. 5 of Brandão, 2008 (Knysna Beach, South Africa)
?Bairdoppilata sp. 6 of Brandão, 2008 (Lüderitz Bay, Namibia)
A third cluster is represented by the genus Glyptobairdia, a small group of Neotropical reef-dwelling species with
asymmetrical carapaces, pronounced ridges, deep punctae, bairdoppilatan accessory dentition, and scissors-like
antennal claws. The soft anatomy has been described, at least in part, for the following species (listed by original
binomen):
Bairdia coronata Brady, 1870 (Caribbean, Bermuda, Bahamas, Belize)
Bairdoppilata? sp. 1 of Maddocks, 1975 (Ascension Island; a juvenile)
Carapace size and shape. Carapace lengths of species of Bairdoppilata range from 0.5 mm (B. sp. 2, Ascension
Island) to almost 2 mm (?B. sp. 1 aff. B. labiata, Antarctic Peninsula) (Table 2, Fig. 3). There is a positive
association between carapace length and water depth, which involves an inverse relationship with water
temperature and latitude. Larger species have been collected from bathyal depths and Antarctic waters. Few species
of intermediate size are included in the dataset analyzed here, which is restricted to species whose soft anatomy has
been described. The smallest species are those living in intertidal and shallow-subtidal, algal, sandy and coralline
habitats in the tropical belt (Fig. 4). Unfortunately, for many of these species the dimensions have been reported
only as population ranges (minimum to maximum), which obscures the biological trends.
The carapace height:length proportion ranges from 0.5 to 0.8 (Table 2) and shows no effect of water depth or
latitude. The greatest height is located at 0.47 to 0.51 of length. The carapace thickness:length proportion ranges
from 0.41 to 0.48, and the location of greatest thickness is at 0.46 to 0.5 of length. Males are slightly shorter than
and not as high as females, although the two populations overlap. This is the usual trend in Bairdiidae.
The carapace has conspicuous left-right asymmetry in size and shape, with a larger LV that swells dorsally
above the smaller RV. This is because the functional hinge must remain a straight line, independent of the carapace
inflation and curvature. The higher the dorsal arch of the LV, the greater the difference in height and outline
between the two valves. The LV reaches over the edge of the RV anterodorsally and posterodorsally, as well as
ventromedially (as the bow-shaped process behind the mouth region). The lateral outline of the LV is rounded and
more distinctive taxonomically, while that of the RV is angulate and less diagnostic. In dorsal view the carapace is
moderately compressed, with tapered anterior and posterior ends and gently swollen midsection. The ventral region

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TABLE 2. Carapace size and water depth for 24 records of 21 nominal species of Bairdoppilata. Species are listed
in decreasing order of carapace length. See also Figure 3.
Explanation of abbreviations: NO = identification number in Figure 3, NAME = species name or informal identification,
REF = published reference, LOC = collecting locality; LCAR = carapace length (μm), HCAR = carapace height (μm),
H/L = carapace height:length ratio, DEPTH = water depth (m).
sp1 aff lab = ?Bairdoppilata sp. 1 aff. ?B. labiata, sp2 aff lab = ?B. sp. 2 aff ?B. labiata, sp. 2 = B. sp. 2 (this paper), sp.
3 = B. sp. 3 (this paper) , sp. 4 = B. sp. 4 (this paper).
B2008 = Brandão 2008, H1974 = Hartmann 1974, H1978 = Hartmann 1978, H1979 = Hartmann 1979, H1980 =
Hartmann 1980, M1969 = Maddocks 1969, M1973 = Maddocks 1973, M1975 = Maddocks 1975, R1960 = Rome 1960,
herein = this paper.
ASC = Ascension Island; BAH = Bahama Islands; E25 = Eltanin 25, 04
o
53’N, 80
o
28’W to 04
o
51’N, 80
o
28’W, east of
Galapagos Islands; E418 = Eltanin 418, 62
o
39–40’S, 56
o
8–10’W, Antarctic Peninsula; E1345 = Eltanin 1345, 54
o
50–
51’S, 129
o
46–48’W, Pacific Sector of Southern Ocean; E1418 = Eltanin 1418, 54
o
32’S, 159
o
02’E, Kerguelen Island;
FFS = French Frigate Shoals, the Hawaiian Islands; GAUS = Gauss-Station, Antarctica; GM = Gulf of Mexico; KB =
Kane’ohe Bay, the Hawaiian Islands; NB = Nosy Be, Madagascar; SB = St. Barthelmy Island, Lesser Antilles; SE AUS
= Southeast Australia; W AFR = West Africa; W AUS = West Australia; WED = Weddell Sea, Antarctica.
is not flattened, and the greatest thickness is located a little below mid-height. The exterior surface ranges from
smooth to punctate. The preserved valves of shallow-water species are mostly transparent, except for an oval
opaque patch located centrally over the adductor muscle scar pattern. There may be brown pigmentation either in
two small spots or over much of the lateral surface.
For shallow-water species, the lateral silhouettes of the carapace or LV can be sorted into three inter-
gradational groups, which merely represent combinations of two trends, dorsal inflation and caudal extension:
(1) Oblong, loaf-shaped or scoop-shaped, with low-arched, subtly angulate dorsal margin and smoothly
rounded posterior margin, not caudate: B. scaura n. sp., B. cytheraeformis, B. geelongensis, B. mocamedesensis, B.
sinusaquilensis.
(2) Dorsally arched, semicircular to subtriangular, with continuously rounded outlines, not caudate: B. sp. 2
(herein), B. angolensis, B. balihaiensis.

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(3) Caudate to sinuate, with smoothly arched to sinuous dorsal margin: B. sp. 2 (herein), B. sp. 3 (herein), B. sp.
4 (herein), B. alcyonicola, B. cratericola, B. cushmani.
These shape tendencies help to distinguish species of Bairdoppilata from some other genera of Bairdiidae:
Neonesidea (teardrop-shaped, with more symmetrical LV and RV), Aponesidea (flatiron-shaped, with flat venter),
Havanardia (angulate, flat venter with ventrolateral keel), Mydionobairdia (box-shaped, tuberculate); Triebe lina
(rhomboidal box-shaped, punctate, ridged). Confusion arises because similar carapace outlines are found in some
species of Paranesidea, which can be distinguished from Bairdoppilata only by close attention to details of
hingement, patch pattern, surface ornament, appendages and genitalia.
Hinge. The functional hinge is a straight median bar where the thickened edge of the RV fits into a groove
beneath a ledge in the LV. The anterior and posterior ends of the bar are expanded ventrally and may be slightly
elevated. The corresponding terminal sockets of the LV are shelf-like, without a ventral confining ridge.
In calcified valves of some species, the dorsal surfaces of both the bar and the groove may be finely to coarsely
striate. This serrate texture was first illustrated by Morkhoven (1958, Pl. 46, figs. 4–6) for G. co ro n at a, where it is
dramatically expressed. In that thick-walled species the “exceedingly minute striations under favorable lighting”
(Stephenson 1946, p. 346) are vertical, regular, and sharply incised. The band of striations thins at both ends to
wrap dorsally over the LV sockets, where it is reflected as complementary indentations across the terminal hinge
teeth of the RV (Morkhoven 1958, Pl. 46, figs. 5, 6). A somewhat similar effect occurs in Macrocyprididae (Triebel
1960, Pl. 14, figs. 4–10; Maddocks 1990, Pl. 60, figs. 1–9, Pl. 61, figs. 1–9). Crenulate hinge texture in Bairdiidae
was illustrated for Neonesidea schulzi (Hartmann, 1964), Neonesidea michaelseni Hartmann, 1984, and
Bairdoppilata mocamedesensis by Hartmann (1964, Pl. 5, figs. 20, 21; 1974, Pl. 20, figs. 150a, b; 1984, Pl. I, fig.
9). Titterton & Whatley illustrated striate hinges for four more species of Neonesidea and Bairdoppilata (1988, Pl.
1, figs. 8, 15; Pl. 2, figs. 6, 16).
This striate zone is a microstructure within the carapace wall, rather than an articulation surface. It marks the
uncalcified connective zone along the midline where only the chitin ligament connects the valves (Maddocks 1990,
1995). The term ligament was re-established for Ostracoda by Kornicker (1969). Its distinct ultrastructure as an
independent element of the carapace was demonstrated by Yamada (2007b), who sectioned the hinges of
Neonesidea oligodentata (Kajiyama) and Triebelina sp. On this basis, he classified the hinge of Bairdioidea as
“exterior type,” because the overlap structure (edge of LV) develops dorsal to the ligament.
In the decalcified carapaces studied here, the contact zone between the valves displays as two narrow cords or
ribbons of chitin (Fig. 9G–J, 17P, 21K). This connective band is strong. The dissected specimens ripped through
the fabric of the adjacent valve wall rather than separating along the midline (Figs. 12E–F; 16A; 20A, L). In some
Bairdiidae these cords appear to be straight (Paranesidea sp. 2, Fig. 21K–L; unpublished images of Neonesidea
tenera). In B. scaura, B. sp. 2 and B. sp. 4 each ribbon is sinusoidally rippled from anterior to posterior (Figs. 9I–J,
12A, 17P). It is likely that this rippled band of chitin confers the striate texture to the hinge in a calcified specimen.
Hartmann published an unusual SEM view of the hinge region in a well calcified valve of B. sinusaquilensis (1979,
Pl. I, figs. 16–18). It shows a row of tiny globular projections within the calcified fabric of the hinge zone, which
may be the lateral meander-edges of these zig-zag ribbons.
At the anterior and posterior ends of the hinge zone in dorsal view in B. scaura, this chitinous band expands
across the midline into the LV and becomes more coarsely scalloped (Figs. 9G, I–J). These scallops are interpreted
as tiny, crenulate teeth, just beyond and dorsal to the terminal shelf-sockets of the LV hinge. A similar, crenulate,
dorsal end-tooth is visible in B. sp. 2 (Fig. 12A, at the far right of the image, beyond the anterior socket-shelf). In B.
sp. 4 these terminal teeth are evident but more subtle (Fig. 17O). A five-part LV hinge with unusually deep,
loculate terminal sockets and tiny, crenulate terminal teeth, as well as a serrate median element, was illustrated for
Neonesidea michaelseni by Hartmann (1984, Pl. 1, figs. 6–11). The differentiation of terminal hinge teeth and
sockets is a common device in Ostracoda to minimize valve offset, and one may speculate that Bairdiidae living in
high-energy environments would benefit thereby.
Accessory bairdoppilatan dentition. In calcified specimens, accessory bairdoppilatan dentition is clearly
seen in both valves. On the anterodorsal and posterodorsal marginal infold (duplicature) of the LV, beneath the
dorsal overhang, the surface swells into a small crescentic platform, in which are four to six depressions. Small
teeth project from the anterodorsal and posterodorsal edges of the RV and fit into these depressions when the
valves are closed. The posterior platform of the LV is located at (beneath) the slight concavity in the posterodorsal

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silhouette, where the posterodorsal margin meets the caudal process. The anterior platform is located at (beneath)
the corresponding point in the anterodorsal silhouette, although most species of Bairdoppilata have a continuous
anterodorsal slope rather than a concavity. In the decalcified specimens studied here, the chitinous fabric retains
sharp outlines of the teeth and sockets (Figs. 8E–F, 9H, 12E–F, L, 16L, 17A).
Normal pore canals. Smith & Kamiya (2002, Fig. 5) documented the ontogenetic increase of normal pore
canals for the LV of Neonesidea oligodentata, from 10 pores in the A-7 instar to 1485 in the A-1 instar and 2145
pores in the adult. Over the central regions of the valve, newly added pores are sprinkled between the pores
inherited from the previous instar, but along the free margin pores are crowded together densely, nearly touching in
the adult.
Simple normal pores of many sizes are abundant in Bairdoppilata. The larger pores, which probably arose
earlier in ontogeny, are fewer in number and have distinct muri (9C, F, 17P). Pores of smaller diameter have
narrower muri or none (8D, 16H). Simple (rimless), tunnel-shaped (with narrow rims) and funnel-shaped (with
broad sloping rims) NPC were reported in Bairdiidae by Puri & Dickau (1969, types A’ and A), Puri (1974), and
Keyser (1980).
Radial pore canals. The radial pore canals of Bairdoppilata are exceptionally numerous, remarkably straight,
and very closely spaced (Figs. 8G–I, 9E–F, 16K). Each canal originates in the vestibule (lined with epidermis) and
ends in a pore with an associated seta. Some lead to marginal pores and setae, but many are so-called false radial
pore canals, leading to pores and setae clustered near but not quite at the valve edge (Figs. 8H–I). There are no
differences in thickness of the canals, and the distinction between true and false radial pore canals is probably not
significant. Perfectly straight RPC are also evident in Paranesidea sp. 1 (Fig. 20C, L–M, S).
Exterior carapace setae. Numerous simple setae (sensilla) of many sizes have been observed in these species
of Bairdoppilata (Figs. 8A, D, 12H; 13B, F, 16A–B; D–E, H), but no branching (polyfurcate) setae, no barbed or
thorny setae, and no anchor setae (Maddocks 2013). All have limited flexibility and taper to a sharp point. Most are
oriented more or less posteriorly. The color is light to medium brown in these specimens. Because the setae are less
conspicuous than in some species of Neonesidea, although their numbers and density are comparable, shallow-
water species of Bairdoppilata are less likely to have been described as "hirsute."
Marginal carapace setae. Three kinds of marginal setae (sensilla) have been observed in the species of
Bairdoppilata studied here: simple setae, eyelash setae and plumose setae. No caudal setae have been seen. A fifth
kind (pappose setae) has not been seen in Bairdoppilata but is present in a species of Paranesidea. These terms, of
which two were defined by Broodbakker & Danielopol (1982, in context of appendage chaetotaxy) and three were
proposed by Maddocks (2013), are used here in the descriptive sense, with no implications regarding homology.
Because of the large number and crowded spacing of pores and setae in the marginal zone, individuals are difficult
to recognize, and it is not known whether an individual seta may change its morphologic expression during
ontogeny.
Simple setae are setae on the carapace exterior (so-called outer lamella), some of which happen to be located
near the valve edge (Figs. 8H–I, 9C–F, 16D–E, K). They include some of the ontogenetically oldest (hence longest,
thickest and darkest) carapace setae. They originate from false radial pore canals at pores with larger diameters and
distinct, circular muri, near the edge of the valve on the exterior surface. They are stiff, smooth, nearly straight and
taper to a point. Their diameter is greater than eyelash setae or plumose setae, and they are longer and darker in
color.
Eyelash setae are short, thin, straight or weakly curved setae, which are aligned at regular intervals in a single
row and parallel, like pickets of a fence (Figs. 9C–D, F; 16D–E). They are light-colored and taper to a point. They
originate from pores at the edge of the calcified zone, along the base of the selvage-ridge, but inside the chitinous
flange, if one is present (Figs. 9C–D, 12E). They are about half as long (or less) as any nearby plumose setae. Their
form is the same as simple setae, with a basal constriction and ring-like collar. Although many pores are crowded
close together in this marginal zone, the pores belonging to eyelash setae may be recognized by their location
closest to the edge and their regular spacing. They are thought to open on the interior surface, although that fact has
little significance for homology, because the marginal infold is a continuous fabric (Harding 1964, Kornicker 1969,
Yamada 2007a). An early illustration showing relationships of simple setae, eyelash setae and radial pore canals is
that of Müller (1894, Pl. 15, fig. 9, for a species of Neonesidea). Hartmann (1974, Pl. 17, fig. 127) illustrated these
two sizes of marginal setae for B. angolensis. For additional illustrations see Maddocks (2013, Figs. 8G–H, 18H,
24G–H, 25F, 29C, 30C–E).

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Plumose setae are short, thin setae, edged by two rows of fine, regularly spaced, parallel barbs or vanes that
give them a plume-like or feathery appearance (Figs. 9F, 12F, K, 16K, 17L–M). They are light-colored and
somewhat flexible. Where present, they are spaced at regular intervals and have consistent lengths (twice as long as
nearby eyelash setae, or longer). They open from regularly spaced pores near the edge of the calcified zone on the
inner surface, along the outward-facing edge of the selvage-ridge. The barbs are fine, barely detectable
microscopically, nearly straight, uniformly parallel, and oriented at about 45
o
to the shaft. Some suspected plumose
setae are thought to have barbs that are too small to be seen; these plumose setae are discernable by their beaded
aspect, gentle curvature, relative length and tendency to attract detritus (Figs. 12E, 17A).
The term plumose seta was defined by Broodbakker & Danielopol (1982, p. 106, Fig. 2C) as a division of
plumed seta in which “the setules are hairy and have a flexible appearance.” Their analysis of homologies was
documented by SEM photos of appendage setae. Published drawings of plumose setae as marginal carapace setae
of Bairdiidae include Müller (1894, Pl. 15, figs. 8, 10) and Maddocks (2013, Figs. 5E–F).
Plumose setae were documented for two species in the Neonesidea pateriformis species-group by Maddocks
(2013), N. tenera Brady (1886) and N. plumulosa Maddocks (2013). In that publication they were labeled as
“caudal setae” (perhaps a less-developed form), but it now seems appropriate to distinguish them by a more precise
term to avoid speculation concerning homology. In those two species, both simple setae and plumose setae are
located in close proximity at the caudal region, on the exterior and interior surfaces, respectively, but there are no
caudal setae present (Maddocks 1995, Figs. 5A–D; 8D–E; 19A–B, D).
In the four species of Bairdoppilata studied here, about eight plumose setae are visible at the top of the
posteroventral free margin, in the caudal region of both RV and LV. In B. sp. 2, additional plumose setae are
suspected to occur on the anterior margin (Fig. 12E). Where eyelash setae and plumose setae occur together along
the same margin, the plumose setae are thicker, longer, arise a little farther from the margin, and probably
originated earlier in ontogeny.
Caudal setae are large, fan-shaped carapace setae, which open from wide but rimless normal pores on the
dorsal exterior surface of the caudal process (Maddocks 2013). The adjacent valve surface may be slightly swollen
to form what is called a “humped caudal process.” They have unusually thick basal shafts and broad, flat, terminal
fans with fringed edges. They are well developed in the Neonesidea schulzi species-group, just dorsal to the
posterior spine. Caudal setae were documented for one Hawaiian species (Neonesidea holdeni Maddocks 2013,
Figs. 25F–I, 26H).
Caudal setae have not been observed in Bairdoppilata. In the species studied here, the simple setae on the
exterior dorsal surface of the caudal process have no visible barbs. The normal pores in this location are not
enlarged, and the surrounding surface is not swollen. The LV with caudal setae illustrated for B. portsamsonensis
by Hartmann (1978, Pl. 1, fig. 14) belongs to a species of Neonesidea.
Pappose setae were defined by Broodbakker & Danielopol (1982, p. 106) for plumed setae in which the
setules arise on all sides of the shaft. On carapaces of Bairdiidae, they were first illustrated by Kornicker (1961,
Text-figs. 10A, D) for Paranesidea gigacantha. They are not restricted to the caudal region but occur around the
free margin. In Paranesidea sp. 2, the suspected pappose setae are thin, short (about 30 µm), originate at pores on
the outer face of the selvage-ridge, and reach across the chitinous flange but not beyond it (Figs. 21M–Q). They are
not visible from the exterior. The shaft is thicker at the base and tapers. The barbs or setules appear to arise all
around the shaft, rather than in just two lateral rows, giving it a tree-like or root-like rather than feather-like
appearance. None have been observed in Bairdoppilata.
In Fig. 21M for Paranesidea sp. 2, about 9 pappose setae emerge from regularly spaced, arched, tunnel-like
pores at the crest of the selvage-ridge, along the anterior margin. Several more pappose setae (out of focus) emerge
at lower elevations on the selvage-ridge. The six curved setae at the base of this image are eyelash setae, which
emerge at the base of the selvage-ridge and extend beyond the chitinous flange. A separate nerve canal (RPC) leads
to each seta. In total, this image includes at least 15 pappose setae plus 6 eyelash setae. Figs. 21N–Q are
enlargements of this same region, at different levels of focus, showing details of these moss-like pappose setae.
Antennule. The antennules of Bairdoppilata are constructed according to the normal pattern for the family
(Figs. 6A–B). As usual, the three distal podomeres are severely reduced and carry a tassel of about 13 very long,
flexible, tactile setae. So far as known, they do not provide information useful for discriminating species and
genera.

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Antenna. The antennae (second cephalic limbs) of Bairdiidae are the anterior walking legs, assisted by the
three thoracic legs. The sharply pointed, curved distal claws provide a secure toe-hold on steep surfaces, like the
crampons of a climber on a glacier. In females of Neonesidea, the tip of the main claw is obliquely beveled with a
sharp point, and this feature is repeated in the distal claws of the thoracic legs. This beveled configuration
resembles the ice tool (technical axe) used by mountaineers to climb a vertical wall of ice. In Aponesidea and
Havanardia the claw tip is less abruptly bent. In Triebelin a the tips of both the antennal claw and the thoracic legs
are barbed. In Paranesidea and Mydionobairdia the pointed tip merely follows the natural curvature of the claw,
like a scimitar. In Bairdoppilata and Glyptobairdia the distal claws are relatively broad and bladelike, curving
smoothly to rounded or pointed tips.
The antenna of Bairdoppilata and Glyptobairdia has two terminal claws of the same or nearly the same size,
resembling a pair of scissors (Fig. 6C–D, 10G, 13G, 14H–I, 16I–J, 17G). In these shallow-water species, the
anterior accessory claw has the same curvature and almost the same length and thickness as the main distal claw. In
three species of ?Bairdoppilata described by Brandão (2008) from the Southern Ocean (?B. labiata, ?B. sp. 1 aff.
?B. labiata, and ?B. sp. 2 aff. ?B. labiata), the anterior accessory claw is noticeably shorter and thinner than the
main distal claw. By contrast, in Paranesidea and other genera (Neonesidea, Aponesidea, Triebelina, Havanardia
and Mydionobairdia) the homologue of this anterior claw is a thin accessory seta (Fig. 18A–B; 20J, Q–R; 21E–F).
This seta is flexible and sexually dimorphic, longer in females than in males. It is occasionally damaged or
overlooked in published drawings.
The accessory claw does not assume this form until the final molt, being represented only by a flame-shaped
anlage in the A-1 (pre-adult) instar (Figs. 6L, 11K, 13I–J; see also Maddocks 1973, p. 63, Fig. 6B). In spite of its
late emergence in ontogeny, at the same time as many sexually dimorphic traits, it has the same configuration in
adult males and females. None of the distal antennal claws are dimorphic in Bairdoppilata, although they are in
some other bairdiid genera.
Although the accessory claw of Bairdoppilata is usually described as enlarged, relative to its development in
other living genera of Bairdiidae, this might be a primitive or relict condition rather than derived. Bairdoppilata is
well represented in Cretaceous and younger strata by fossil species (identified by accessory locking dentition), and
possible relatives are known in the Triassic.
The bairdiid antenna has 6 podomeres, of which I and II belong to the protopodite and III–VI belong to the
endopodite. Podomere I is often omitted or unclear in illustrations, because of damage during dissection, or
because its curvature makes it unstable in lateral profile. Between podomeres I and II, a stove-pipe joint allows
podomere II to rotate up and in, until the dorsal (outer) edge becomes parallel to the inclined surface of the
forehead, near the concave upper portion of the S-shaped strut of the head capsule, a little below the base of the
antennule (Fig. 13C). A second stove-pipe joint between podomeres II and III provides additional lift and
accommodation around the forehead. The exopodite scale, carrying one long and two tiny setae, is tightly attached
to the outer proximal edge of podomere III and remains with it if the protopodite is torn away.
At the “elbow joint” between podomeres III and IV, the rigid distal part of the limb can rotate through
approximately 74
o
. It can be raised as high as 126
o
, and it rotates down past vertical to an orientation of 200
o
. The
limit to which the limb can be raised is set by the anterodorsal marginal overlap of the LV (Fig. 16F). At maximum
extension in preserved animals, the distal claws, all of podomere VI and about half of podomere V may be exposed
outside of the carapace. When the antenna withdraws into the domicilium, the distal claws are held just below the
upper lip, parallel to and just within the anteroventral margin of the closed carapace (Fig. 13C). The rather straight
course of the anteroventral margin in the RV may indicate the orientation of the antennal claws within.
Podomeres IV and V are laterally compressed and aligned in a stiff arc that resists distortion. There is no
flexibility at the joint between them, which is set at an oblique angle to the trend of the limb and braced by a claw-
like ventrodistal seta. A strong tendon within podomere V confers additional rigidity. The joint between podomeres
V and VI has very little movement, because the robust ventrodistal claw of podomere V fits against a basal notch in
podomere VI and acts as a stop. The amalgamation of the posterodistal fused claw into podomere VI provides a
brace to limit movement of the terminal claws. In this way, the entire distal limb (podomeres IV, V, VI and terminal
claws) behaves as a rigid grappling unit, which is operated in lever fashion by the muscles in podomere III
(somewhat like a backhoe).
Podomere IV is always shorter than podomere V, but the proportional relationship between them is
inconsistent, ranging from 0.58 in G. coronata to nearly equal (0.97) in B. hirsuta. This is unexpected, because

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these two podomeres are immovably sutured and function as a unit. This relationship should be investigated in
other genera of Bairdiidae.
The antenna is the most frequently illustrated limb and is thought to present diagnostic details for the
identification of species. Yet, it is surprisingly difficult to extract consistent chaetotaxial information from
published drawings because of missing information and probable errors. For a few species, only the most distal
podomeres and claws have been figured. For six species, the appendage drawings were published without
magnification scales, and their dimensions cannot be compared (B. angolensis, B. balihaiensis, B. cytheraeformis,
B. geelongensis, B. mocamedesensis, B. sinusaquilensis). For some species, the illustrated legs belong to a different
species than the illustrated valves (B. portsamsonensis, B. simuvillosa). Verbal assessments of shape (long, thin)
have limited usefulness, because they lack a context for comparison.
Within these limitations, the antennal chaetotaxy (presence/absence of setae) of Bairdoppilata appears to
follow the standard pattern for the family: podomere I: 2 long posterodistal setae; podomere II: 1 long anterodistal
and 1 shorter posterodistal setae; podomere III: 3 medium posteromedial aesthetascs and 2 posterodistal setae (both
long, or one long and one medium); podomere IV: 1 or 2 posterodistal setae (1 long claw, or 1 long claw and 1 short
seta); podomere V: 2 or 3 anterior filaments and 1 short posterodistal claw; podomere VI: 1 long anterodistal
(accessory) claw, 1 long terminal (main) claw, 1 short terminal aesthetasc, and 1 short posterodistal (fused) claw.
Beyond presence/absence, the relative proportions of these structures may carry taxonomic information.
Lengths and widths of antennal podomeres, setae and claws were measured and compiled for the illustrated species
of Bairdoppilata. Because of numerous gaps (missing information), the full data set (36 illustrations, 22 species, 21
characters) proved unsuitable for systematic analysis. A subset of these data is presented as Table 3. Associative
plots (such as Fig. 5) suggest that the following dimensions tend to vary together:
(1) Length of accessory claw compared to length of main claw. In the shallow-water species, they are very
nearly the same length (always within 10 percent, usually within measurement error).
(2) Length of distal claw of podomere V compared to length of fused claw of podomere VI, and (3) Length of
distal claw of podomere IV compared to length of podomere V, and (4) Length of distal claw of podomere V
compared to length of podomere VI. Each of these claws acts as a brace to prevent excessive movement of the
following podomere. The consistent size relationships support this functional hypothesis.
(5) All widths of distal podomeres and widths of claws.
These results suggest that drawings showing only the distal claws (like Figs. 6D and 14I) do not provide as
much independent taxonomic information as is commonly supposed. More proximal setae and podomeres might be
more distinctive, but they are also more frequently damaged or omitted from illustrations.
TABLE 3. Lengths of distal antennal claws, measured on 25 illustrations of 12 species of Bairdoppilata,
converted to percentage of carapace length. Species are listed in decreasing order of carapace length. See also
Figure 5.
Explanation of abbreviations: NO = number, NAME = species name or informal identification, SPEC =
catalog number of specimen, SEX = sex of illustrated specimen, M = male, F = female, REF = published
reference, FIG = figure number of measured illustration, LOC = locality, LCAR = carapace length (µm), HCAR =
carapace height (µm), H:L = Height:Length Ratio of carapace, DEPTH = water depth in meters, MAIN = length
of main (terminal) claw, ACC = length of anterior accessory claw, FUS = length of fused claw of podomere 6, P5C
= length of posterodistal claw of podomere 5.
sp1 aff lab = ?Bairdoppilata sp. 1 aff. ?B. labiata; sp2 aff lab = ?B. sp. 2 aff ?B. labiata; sp. 2 = B. sp. 2 (this
paper), sp. 3 = B. sp. 3 (this paper).
B2008 = Brandão 2008, M1969 = Maddocks 1969, M1973 = Maddocks 1973, M1975 = Maddocks 1975,
R1960 = Rome 1960, Herein = this paper.
E25 = Eltanin 25, 04
o
53’N, 80
o
28’W to 04
o
51’N, 80
o
28’W, east of Galapagos Islands; E418 = Eltanin 418,
62
o
39–40’S, 56
o
8–10’W, Antarctic Peninsula; E1345 = Eltanin 1345, 54
o
50–51’S, 129
o
46–48’W, Pacific Sector of
Southern Ocean; E1418 = Eltanin 1418, 54
o
32’S, 159
o
02’E, Kerguelen Island; FFS = French Frigate Shoals, the
Hawaiian Islands; FL = Florida Keys; GAUS = Gauss-Station, Antarctica; GM = Gulf of Mexico; NB = Nosy Be,
Madagascar; SB = St. Barthelmy Island, Lesser Antilles; BAH = Bahama Islands, ASC = Ascension Island; WED
= Weddell Sea, Antarctica.

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Mandibles. So far as known, the mandibles of Bairdoppilata and Paranesidea are consistent with the familial
pattern (Figs. 6E–G, 14B, 15B, 18D), although this limb has not been illustrated for many species. The terminal
claw of the palp is distally fringed or pectinate, which offers a potential site for taxonomic or diet-related variation
(Fig. 6F, 14B, 21B). The palp is thought to move food into the mouth. The basal teeth are inserted into the atrium
from the sides and chop or partition the food into bites.
Maxillae. The palp of the maxilla (maxillule) and the three masticatory processes (endites) are held nearly
horizontal along the boat-shaped ventral surface of the lower lip (Figs. 13C, F) and may act as a brush to sweep
food toward the atrium. The outer integument may have a prickly texture with rows of papillae or tiny setules (Figs.
10H–I, 11A). As far as known, the chaetotaxial pattern described below for Bairdoppilata is the same as in
Paranesidea and other genera of Bairdiidae. There is inconsistency in published illustrations, especially in smaller
species, either because the number of setae is reduced or because of difficulties of observation. Details of this
configuration might have taxonomic significance, but there is insufficient documentation at present.
The slender, unjointed palp carries a cluster of three setae at the anteromedial indentation, a posteromedial
seta, a terminal seta, and a stout, fringed or pectinate terminal claw (6K, 10I, 15A, 18E, 20H, 21B). The first
masticatory process carries a posteromedial seta and 6 or more terminal setae, of which one is a bristle (6J, 10H,
15A, 20G, 21C). The second masticatory process has a posteroproximal seta, a fringed or pectinate terminal claw,
which may be spatulate or spoon-shaped, another claw, a bristle, and 3 or more other setae (6J, 10H, 15A, 20F,
21D). The third masticatory process has a posteroproximal seta, two fringed or pectinate terminal claws, which
may be spatulate or spoon-shaped, another claw, and up to three additional setae (6J, 10H, 15A, 20E, 21D).
The vibratory plate has been illustrated for only two species of Bairdoppilata, and two more are added herein.
B. alcyonicola has 25 plumose setae (Maddocks 1969, Fig. 38A). Brady (1880, Pl. 5, fig. 2d) drew 22 setae for B.
villosa. B. scaura n. sp. has at least 20 setae (Fig. 6H, damaged). B. sp. 3 has 23 setae (Fig. 14G). In Neonesidea
this structure has either 25 or 26 setae (Maddocks 2013). Smith et al. (2005) suggested a range of 24 to 26 for
genera of Bairdiidae.
The ventral accessory plate of the maxilla in Bairdoppilata carries 6 stiff reflexed setae with wedge-shaped
terminations, as usual for this family (Fig. 6I).
Thoracic legs. The vibratory plate of the first thoracic leg in Bairdoppilata is thought to have 13 plumose
seta, as in Neonesidea and other genera of Bairdiidae (Fig. 7A, Table 4). For four species, only 12 plumose setae
have been reported, however, as also is the case in Paranesidea sp. 1 (Fig. 19D).
The four reflexed setae are segregated anteriorly, well separated from the plumose setae. Hartmann (1978)
reported only 3 reflexed setae in B. balihaiensis. As far as known, they are similar in males and females, not
sexually dimorphic as in Neonesidea. In Bairdoppilata the first reflexed seta may be a little shorter than the others
(Figs. 7A–B). In Paranesidea sp. 1 the first seta is actually longer than the others (Fig. 19D). They are smooth,
somewhat stiff and end in oblique, wedge-shaped, flat tips, as described for Neonesidea (Maddocks 2013).
On the second and third thoracic legs, the corresponding, vestigial epipodite consists of two thin, unequal
setae, fused at their base.
TABLE 4. Chaetotaxy of thoracic legs of 14 species of Bairdoppilata, based on 16 published illustrations.
The numeral indicates the number of setae at each location. Italic font and gray tone indicate no information or
suspected errors. It appears that this chaetotaxy follows a consistent pattern, and most deviations represent errors
or missing data.
Explanation of abbreviations: ALCY = B. alcyonicola, BALI = B. balihaiensis, CRAT = B. cratericola,
HIR1 = B. hirsuta, USNM 121353, HIR2 = B. hirsuta, USNM 139891; MOCA = B. mocamedesensis; SIMP = B.
simplex; VILL = B. villosa; SP2A = B. sp. 2, Ascension; SP2L = ?B. sp. 2 aff. B. labiata; CORM = G. coronata;
CORR = G. c o r o na t a ; SCAU = B. scaura; SP 2 = B. sp. 2; SP 3 = B. sp. 3; SP 4 = B. sp. 4.
T5, T6, T7 = first, second or third thoracic leg; I–V = podomeres; anteroproximal (etc.) = location of seta;
vib = vibratory plate; plumose = number of plumose setae; reflexed = number of reflexed setae; shorter = whether
the first reflexed seta is shorter than others.
B2008 = Brandão 2008, H1974 = Hartmann 1974, H1978 = Hartmann 1978, M1969 = Maddocks 1969,
M1973 = Maddocks 1973, M1975 = Maddocks 1975, R1960 = Rome 1960, here = this paper.

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The chaetotaxy of the thoracic legs for Bairdoppilata follows the standard pattern for the family with no sexual
dimorphism (Table 4; Figs. 7A, C–D, 14C–E). Claw lengths are relatively shorter and width:length proportions of
podomeres are less elongate in the shallow-water species of Bairdoppilata than in some other genera. Because of
inconsistencies in published illustrations (missing setae and discrepancies in relative lengths of setae), these limbs
do not contribute much to identification of species and genera at the present time. Paranesidea sp. 1 follows the
same chaetotaxial pattern, although both podomeres and setae appear to have more elongate proportions (Figs.
19A–D).
The brush-shaped appendage of the male is asymmetrical in Bairdoppilata, as usual in the family (Fig. 7E).
Zygum. The two furcal rods, the single post-abdominal bristle, and the paired external genitalia (hemipenes or
genital lobes) of Bairdiidae articulate to a median plate or frame, the zygum, which is carried on the midline at the
posterior end of the body. This central frame is lozenge- or shield-shaped and heavily muscularized (Figs. 10A–B,
11B). It has been illustrated for males many times (McKenzie & Swain 1967, Text-Fig. 3h; Maddocks 1969, Fig.
6B, 7D, 9H, 10B, 11A, 12F, 17B, 21E, 24E, 25C, 28F, 30G, 37C, 39H, 41C; Danielopol 1972, Fig. 3; Maddocks
1975, Fig. 3C; Maddocks & Iliffe 1986, Fig. 7G, 9A, 11A; Maddocks 2013, Figs. 10A–B, 14E, 19G, 31E, 36M),
but its presence in females has rarely been mentioned (Maddocks 1969, Figs. 31G; 1975, Fig. 7A). The skeletal
frame of the zygum consists of two narrow, flat, arcuate, lateral bars and a short posterior cross-bar. In some
specimens the lateral bars may be less well defined along the anterior edges of the plate, or they may be replaced by
converging chitin fibers (Fig. 10B, 11B). In the specimens examined, there is no skeletal connection betweem the
zygum and the lateral framework of chitinous struts, from which the thoracic legs are suspended.
In males the zygum is posterodorsal, and the hemipenes are suspended from it on either side (Fig. 8C, 10A–B).
This U-shaped apparatus is separate from the posteroventral region of the body and hangs in saddlebag fashion,
with the distal ends of the hemipenes pointing posteriorly (Fig. 9A–B, 10A; see also Maddocks 2013, Figs. 10A–B,
35C, for Neonesidea). The anterior connection to the body is a wrinkled tube (Maddocks 2013, fig. 20B, for
Neonesidea). In use the connecting tube inflates, the whole apparatus extrudes posteriorly (but not the
posteroventral part of the body), and the hemipenes unfold through 180
o
.
In females the zygum is oriented approximately vertically, behind the swollen rump containing the internal
female organs (Figs. 12G, 13A–B, 16F, 17A, E). The genital lobes arise close to the end of the zygum, but no direct
skeletal articulation has been observed (Figs. 15D, 17P; see also Maddocks 2013, Figs. 9F–G for Neonesidea).
Less musculature is apparent, because in females the zygum is not extrudable.
In Cytheroidea, this rectangular frame between the hemipenes was first illustrated for Hemicytheridae by
Skogsberg (1928, Pl. 6, fig. 1). The term zygum was designated by Hart & Hart (1969, 1974). In Entocytheridae
three additional skeletal structures are located anterior to the zygum: a pair of small bars (sterinx), a single long bar
(tropis) and a pair of lateral bars (pastinum) (Hart & Hart 1969, Fig. 1; 1974, Fig. 10). Each pastinum connects to
the lateral framework of chitinous rods, from which the thoracic legs are suspended. Schulz (1976, Figs. 71, 78)
illustrated a connecting rod in this position for Sclerochilus sp. and Paradoxostoma sp. (Bythocytheridae,
Paradoxostomatidae). Tsukagoshi & Parker (2000, Fig. 10A) illustrated a U-shaped zygum for Callistocythere
(Leptocytheridae). The sterinx, tropis and pastinum have not been seen in the Bairdiidae studied.
Furca. The familiar term “furca” is used here without reference to any hypothesis about its segmental origin.
Meisch (2007) showed that the “furcal rami” in Ostracoda may have been transformed from vibratory plates on the
uropods (appendages of the posterior-most segment), whereas true furcae develop from the telson. He stated further
(p. 197), “The transformation of a pair of vibratory plates into a pair of variously shaped structures adapted for
locomotion and/or feeding demonstrates the powerful evolutionary potentiality of the epipodites of ostracodes.” In
the case of Bairdioidea, however, the furcae play no role in either feeding or locomotion. In resting position (in
preserved specimens), the furcae wrap horizontally around the genitalia (hemipenes or genital lobes) (Figs. 9A–B,
17A, E). This suggests that the setae of the furcae (especially seta 2) may have a cleaning function. The
disproportionate length of seta 2 (more than twice the length of any other seta, though not long enough to extend
outside of the domicilium) further implies a special function.
The setal armature of the furca is distinctive for several genera within Bairdiidae, though not sexually
dimorphic. Bairdoppilata has seven setae, all fairly long (Fig. 7G, 10B, 11B, 15D, 17F). Neonesidea, Aponesidea,
and Havanardia have seven setae, of which setae 6 and 7 (most proximal) are somewhat shorter than the others.
Paranesidea and Triebe lina have six setae, of which the two most proximal setae are shorter than others (18C).
Mydionobairdia apparently has only five setae.

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FIGURE 2. Carapace dimensions for the species of Bairdoppilata and Paranesidea described here. The assemblages of B.
scaura, B. sp. 2 and B. sp. 3 each include two late instars as well as adults; the others include only adult specimens. See also
Table 1.
FIGURE 3. Plot of carapace length and height vs. log water depth, for 21 species of Bairdoppilata. These data include
adults only, of species for which the soft parts have been described. See also Table 2.

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FIGURE 4. Carapace dimensions for 15 shallow-water species of Bairdoppilata. These data include adults only, of species
for which the soft parts have been described. See also Table 2.
FIGURE 5. Lengths of the four distal claws of the antenna, converted to percentages of carapace length, plotted for 25
illustrations of 12 nominal species of Bairdoppilata. The data include only those species for which the antenna was illustrated
at a known scale and dimensions were provided for the carapace. The species are listed from left to right in decreasing order of
carapace length. See Table 3 for species names and dimensions.

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FIGURE 6. Bairdoppilata scaura, n. sp., A–K, holotype male specimen 3860M; L, instar A–1 3959J. A, antennule with long
setae; B, podomeres of antennule; C, antenna; D, distal claws of antenna; E, mandible; F, pectinate claw of mandible palp; G,
teeth of mandible; H, vibratory plate of maxilla; I, first reflexed seta of maxilla, to show chisel-like termination; J, masticatory
processes of maxilla; K, palp of maxilla; L, distal claws of juvenile A-1 antenna, showing anlage of accessory claw. Scale bar =
100 µm.

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FIGURE 7. Bairdoppilata scaura, n. sp., holotype male specimen 3860M. A, first thoracic leg; B, reflexed setae of vibratory
plate of first thoracic leg, to show chisel-like terminations; C, second thoracic leg; D, third thoracic leg; E, brush-shaped organ;
F, plate of masticatory organ; G, furca; H, hemipenis. Scale bar = 100 µm.

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Seta 2 (counting from the distal end) is always much longer than the others, with a basal shaft that is as long as
the other setae, followed by a long distal bristle. The proximal part of the bristle (near the shaft) may be sharply
bent and perhaps flattened. The five or six most proximal barbs of the bristle may be noticeably larger than the
others and might have taxonomic value (Figs. 11J, 15D, 17B, H–I). The finer distal spinules are nearly invisible in
small species and often omitted from drawings.
Hemipenis. The male copulatory apparatus of Bairdiidae, where known, is thought to be distinctive for each
species. General resemblances at the generic level are suspected but not yet reliably delineated. This complex
appendage was described by Müller (1894, for Neonesidea) and Rome (1960, for G. c or o na ta ), and Danielopol
interpreted homologies of the parts (1972, for Neonesidea).
Each hemipenis is composed of three articulated parts, usually termed the basal, medial and terminal
segments. In fact, the basal segment consists of two overlapping branches, joined posteriorly, which swing apart or
can easily be torn apart as a V (Fig. 10B). The dorsal branch of the basal segment is longer, oblong, and attached
to the posterolateral edge of the zygum. The ventral branch of the basal segment is shorter, more or less
quadrangular, and strengthened by a diagonal ridge, which runs from the posterior end of the dorsal branch to the
dorsal edge of the median capsule (Fig. 10E–F; 11B–D, F–G). Both branches house bundles of muscles. The
medial segment (also known as the median capsule) displays several configurations in Bairdoppilata, ranging
from subcircular and complex to crescentic and streamlined. It is filled with two groups of outward-fanning
muscles (Figs. 10C–F, 11C–H). In the shallow-water species the distal end carries various conical, lamellar and
hook-shaped prominences, of which the most ventral one may flare as a hood-like shield over the others in some
species. The terminal segment is relatively small and lamelliform, ending in a ridged projection above an incised
groove, which receives the copulatory tube (Figs. 10C–F, 11C–H).
The sleeve of the copulatory tube in shallow-water species of Bairdoppilata arises fairly close to the posterior
end of the median capsule and arches gracefully through 180
o
to 360
o
(Figs. 10C–F, 11C–H). It is held in place by
distal latches but not constrained within a channel, as it is in Neonesidea.
Some of the deep-water species of ?Bairdoppilata have a more linear, more conspicuously hinged hemipenis
with a crescentic medial segment, few or no distal prominences, and a shorter, nearly straight copulatory tube
(Brandão 2008, Figs. 8M–Q, 9A, G).
Genital lobe. The external form of each genital lobe is a pendant ovoid (Fig. 12H–I, 13H, 17C–D, F). A small
copulatory opening leads through a nearly straight canal into the seminal receptacle. A larger orifice opens out of
the oviducts. A spirally coiled canal connects the seminal receptacle with the latter orifice, and a set of muscles
pumps sperm through the spiral canal for fertilization of the eggs (Müller 1894). Müller (1894) considered the
number of coils and thickness of the spiral canal to be useful for distinguishing some species.
The genital lobes have not been illustrated for many species of Bairdoppilata. Their appearance changes with
viewing aspect and focus, and their taxonomic potential remains undetermined. In the species examined here, they
have subquadrate outlines with a flattened venter (Fig. 12G–I, 13H, 17C–D, F, O). The internal tubes make
relatively few coils and occupy only a small part of the volume.
In Paranesidea sp. 1 and sp. 2 the genital lobes are more elongate, twisted in lateral outline and taper ventrally
(Figs. 20N–P, 21H–I). The coiled tubes are thicker, more complexly coiled, occupy more of the internal volume,
and their internal surfaces are striated internally (Figs. 20P, 21H–J). These internal, radially arranged, longitudinal
striations might be sculptural lines or contractile fibers. This feature has not been observed in Neonesidea or
Bairdoppilata.
Masticatory organ. All living Bairdiidae have a unique masticatory organ (chewing structure) at the top of the
esophagus (Figs. 7F, 11I, 12J, 15C, 16C, 17J). It is often ignored but deserves more attention, and Maddocks
(2013) suggested that it might have taxonomic significance. This structure is illustrated here for the four species of
Bairdoppilata, as it was for the five Hawaiian species of Neonesidea (Maddocks 2013). Its function and taxonomic
potential will be evaluated in a separate paper.
Median eye. The median eye of Bairdiidae has been described chiefly for species of Neonesidea (Müller 1894,
Tanaka 2005, Maddocks 2013). Müller stated that it is poorly developed in other species-groups. The four species
of Bairdoppilata investigated here all have medium-sized to large, red-brown eyes, somewhat more conspicuous
than those of Neonesidea. The median eye is situated below the carapace on the dorsal midline of the body, at the
anterior edge of the isthmus, just behind the base of the antennules and in front of the midgut (Figs. 10A, 12L,
13C–E). In lateral view, if the anterior food ball is large, the eye may be obscured. In B. scaura the eye is especially

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large and dark-colored, with long, cone-shaped lateral cups (Figs. 10A, J). In the species of Paranesidea examined
here the eye is somewhat smaller and lighter in color than in Bairdoppilata (Figs. 20D, K).
In the shallow-water species of Bairdoppilata reviewed here, there is no eye-spot or ocular lens embedded in
the carapace, as described for the Paleocene-Eocene genus Oculobairdoppilata by Itterbeeck et al. (2007). They
reviewed reports of fossil bairdiids with eye-spots and noted that such species occur lower in the photic zone,
below an estimated depth of 50–175 m. None have been reported living.
Systematic descriptions
Bairdoppilata scaura n. sp.
(Figs. 6–11)
Etymology. Latin scaurus, with swollen ankles, club-footed; referring to the short antennal claws.
Material. Five specimens, including two males and three juveniles.
Types. Holotype male specimen 3360M, paratype male specimen 3864M, juvenile instar A-1 specimen 3959J,
and two undissected juveniles.
Type locality. Station FFS-TC-06 on French Frigate Shoals.
Dimensions. Holotype male specimen 3860M, entire carapace measured before dissection: Length 897 µm,
height 500 µm. See also Table 1 and Figures 2 and 4.
Diagnostic description. Carapace smooth; oblong-subovate in lateral outline, with subangulate dorsal margin,
greatest height located at rounded anterodorsal corner, at 0.49 of length; rounded posterodorsal corner located at
0.84 of length; anterodorsal brim located at 0.02 of length and 0.61 of height; posterodorsal brim located at 0.98 of
length and 0.39 of height; highest point of ventral indentation located at 0.41 of length and 0.08 of height. Slope
(measured from horizontal) of anterodorsal margin 26
o
, of mid-dorsal margin 21
o
, of posterodorsal margin 55
o
, of
dorso-caudal margin 55
o
, of hinge 15
o
. No caudal process.
Antennal claws very short, about 0.38 of length of podomere 5; accessory claw slightly longer than main claw
(1.10) and very nearly as wide (0.88); pointed tips narrowly rounded; fused claw smooth, blunt; ventrodistal claw
of podomere IV stout, with sharply beveled tip.
Hemipenis heavily sclerotized, complex. Basal segment broad, subrectangular, with flaring, rounded
posteroventral corner and diagonal chitinous ridge. Medial segment subcircular; seminal vesicle and base of
copulatory tube located posterodorsally, at 0.81 of length. Terminal segment short, located at dorsal inner edge;
lamellar, with short lobate extension above incised groove. Distal protuberances include spatulate lamella and
irregular triangle connected by low ridge, thin spine, and thick hook-shaped claw. Sclerotized sleeve of copulatory
tube short, stout, tapered, looped across posterior end, held in place by curved grooves on two prominences; distal
part short, thin, flexible; apparently trifurcate but perhaps snarled.
Comparisons. In size, B. scaura is closest to two Australian species, B. geelongensis and B. sinusaquilensis,
which also share the scoop-shaped carapace outline (shape group 1). The (decalcified) carapace of B. scaura does
not display the finely punctate texture of those two species. The antennal claws of B. scaura resemble those of B.
sinusaquilensis, whereas those of B. geelongensis are more pointed. Other points of similarity to B. sinusaquilensis
include the three-part hinge with dentate terminal elements and serrate median element, and the enlarged proximal
barbs of seta 2 of the furca.
The hemipenis of B. scaura resembles that illustrated for B. sinusaquilensis by Hartmann (1980). Similarities
include the rounded, expanded posteroventral corner of the basal segment; the short, posteriorly displaced, broadly
elevated copulatory tube; and the general configuration of posterodistal appendages of the medial segment. All of
these posterodistal prominences are shorter and more delicate in B. scaura, however. The hemipenis of B. scaura
lacks the flared, hood-like posteroventral shield of B. alcyonicola, B. angolensis, B. cratericola, and B.
mocamedesensis. The heavily sclerotized posteroventral corner of the medial segment is not prolonged into a
lobate or angulate prominence, as it is in B. cushmani and B. cytheraeformis.
Remarks. B. sinusaquilensis was described by Hartmann (1979, 1980) from Eagle Bay at Cape Naturalist,
southwest Australia and from three localities on the warm-temperate coast of south and southeast Australia. Direct
dispersal between these localities and the Hawaiian Islands is unlikely.

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FIGURE 8. Bairdoppilata scaura, n. sp., holotype male specimen 3860M. A, RV exterior; B, LV exterior; C, left exterior of
whole animal before dissection, with soft body and partly detached coil of vas deferens, focused on hinge and supplemental
dentition; D, LV exterior with smooth surface, MSP with adhering muscle fragments, NPC, and setae of many sizes; E–F, RV
exterior, anterodorsal and posterodorsal margins with supplemental dentition; G, RV exterior, posteroventral region with
vestibule, selvage, flange, RPC, simple and eyelash setae; H–I, LV and RV exteriors, anterior margins, with vestibule, selvage,
flange, true and false RPC, simple and eyelash setae. Scale bar = 50 µm.

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FIGURE 9. Bairdoppilata scaura, n. sp., all male specimen 3864M. A–B, right and left exteriors of whole animal before
dissection, with soft body, food balls and genitalia; C, anterior margin of RV exterior with LV interior below, with eyelash and
simple setae, selvage and flange; D, LV interior, anteroventral margin (located just below fig. C); E, LV interior, RPC at ventral
indentation (bow-shaped process) just behind mouth; F, LV interior, posteroventral region (rotated 40
o
counterclockwise),
showing RPC, simple, eyelash and plumose setae; G, dorsal region of decalcified carapace with striate and scalloped hinge,
viewed from interior; H, posterior extension of fig. G, to show supplemental dentition on posterodorsal edge of RV; I–J,
enlarged views of fig. G to show striate hinge, with sinusoidal ribbons and scalloped terminal extensions. Scale bar = 50 µm.

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FIGURE 10. Bairdoppilata scaura, n. sp., all holotype male specimen 3860M. A, dorsal region of soft body, left side, with
median eye, endoskeleton of head capsule, adductor muscles, and hemipenes; B, dorsal view of both hemipenes, zygum, both
furcae, and post-abdominal bristle; C–D, exterior of left hemipenis, focused higher and lower; E–F, exterior of right hemipenis,
focused higher and lower; G, distal claws of antenna with aesthetasc; H, masticatory processes of maxilla, with rough texture; I,
palp of maxilla; J, median eye with conical eye-cups and middle connecting body; K, spiral coil of vas deferens. Scale bar = 50
µm.

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FIGURE 11. Bairdoppilata scaura, n. sp. A–J, male specimen 3864M; K, juvenile specimen 3959J. A, palp and masticatory
processes of maxilla, showing rough texture of cuticle; B, dorsal view of both hemipenes, zygum, furcae and post-abdominal
bristle; C–E, exterior of posterior region of right hemipenis, focused from high to low, to show distal appendages and trifurcate
tip of copulatory rod; F–H, exterior of posterior region of left hemipenis, focused from high to low, to show distal appendages
and trifurcate tip of copulatory rod; I, masticatory organ; J, middle region of seta 2 of furca, with a few tiny barbs; J, distal
claws of juvenile antenna, with anlage of accessory claw. Scale bar =50 µm.

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FIGURE 12. Bairdoppilata sp. 2, all female specimen 3969F. A, interior of LV, striate hinge; B, right exterior of whole animal
before dissection; C, LV exterior with brown pigmentation; D, MSP and NPC in central region of fig. C; E, LV exterior,
anterior margin; F, LV exterior, posterior margin, focused low on adhering shreds of RV hinge and setae; G, left posterior region
of soft body, showing ovaries, zygum, genital lobes and furcae; H–I, genital lobe, after and before dissection; J, masticatory
organ; K, right exterior of whole animal, to show plumose setae in posterior caudal region; L, right exterior, anterodorsal region
of fig. B, showing food ball, median eye and accessory dentition. Scale bar = 50 µm.

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FIGURE 13. Bairdoppilata sp. 2. A–E, female specimen 3966F; F, juvenile specimen 3970J; G–H, female specimen 3972F; I,
juvenile specimen 3965J (molting); J, juvenile specimen 3964J. A, right exterior of whole animal; B–C, right exterior, of whole
animal, posterior region and anterior region of whole animal; D, anterodorsal region of fig. C, enlarged to show median eye and
isthmus; E, median eye, focused on right eye-cup, enlarged from fig. D; F, right exterior of whole animal; G, distal claws of
adult antenna; H, right genital lobe before dissection; I, distal claws of juvenile antenna, molting, with adult claws being
withdrawn from interior; J, distal claws of juvenile antenna, with anlage of accessory claw. Scale bar = 50 µm.
Bairdoppilata sp. 2
(Figs. 12–13)
Material. Twelve specimens, including six females and six juveniles representing two instars.
Dimensions. Adult female specimen 3966F, length 869 µm, height 547 µm. See also Table 1 and Figures 2 and
4.
Descriptive comments. Carapace semicircular in lateral outline, nearly symmetrical (anterior to posterior),
with continuously curved, high-arched dorsal margin, lacking angles or sinuosities; greatest height 0.64 of length,
located at 0.48 of length; anterodorsal corner not distinguishable, located approximately at 0.30 of length and 0.93
of height; posterodorsal corner not distinguishable, located at approximately 0.67 of length and 0.93 of height;
anterodorsal brim located at 0.54 of height; posterodorsal brim located at 0.45 of height; greatest height of ventral

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indentation located at 0.41 of length and 0.09 of height. Slope (measured from horizontal) of mid-dorsal margin 0
o
,
of anterodorsal margin 39
o
, of posterodorsal margin 45
o
, of dorso-caudal margin 44
o
, of hinge 3
o
. Very weakly
caudate, with barely distinguishable posterodorsal change in slope; length of caudal process 0.06 of length. Surface
nearly smooth, with fine microgranular texture, no visible punctae; brown pigmentation over much of central
lateral surface. Striate hinge, well developed plumose setae.
Comparisons. B. sp. 2 is approximately the same size as B. alcyonicola, B cratericola, and B. geelongensis
and is at the lower end of the published range for B. angolensis (Fig. 4). The high-arched, non-caudate to very
weakly caudate lateral outline is shared with B. angolensis and B. balihaiensis (group 2), but both of those species
have more distinctly peaked mid-dorsal angles and somewhat subtriangular outlines. All members of group 3 (B.
alcyonicola, B. cratericola, B. cushmani) have a more sinuous dorsal margin with more prominent caudal process.
The surface of B. sp. 2 appears to be smooth, lacking the distinctly punctate texture of these and most species of
Bairdoppilata.
Bairdoppilata sp. 3
(Figs. 14–16, 17A–F)
Material. Five specimens, including two females and three juveniles representing two instars.
Dimensions. Adult female specimen 3963F, carapace measured before dissection: length 995 µm, height 631
µm. See also Table 1 and Figures 2 and 4.
Descriptive comments. Carapace rounded-subtriangular in lateral outline, with very high-arched dorsal
margin and nearly straight, steeply sloping anterodorsal and posterodorsal margins, lacking angles; greatest height
0.69 of length, located at 0.54 of length; anterodorsal corner not distinguishable, located approximately at 0.37 of
length and 0.92 of height; posterodorsal corner not distinguishable, located approximately at 0.57 of length and
0.89 of height; anterodorsal brim located at 0.04 of length and 0.59 of height; posterodorsal brim located located at
0.91 of length and 0.59 of height; greatest height of ventral indentation located at 0.42 of length and 0.08 of height.
Slope (measured from horizontal) of mid-dorsal margin 3
o
, of anterodorsal margin 37
o
, of posterodorsal margin 47
o
,
of dorso-caudal margin 34
o
, of hinge 3
o
. Caudate, with weak concavity separating dorsal-caudal margin from
posterodorsal margin; length of caudal process 0.06 of length. Surface smooth, except for faint granular texture.
Numerous simple setae, none very long. Seta 2 of furca with 1 or 2 unusually large proximal barbs (each side),
followed by about 8 much smaller, regularly arranged barbs of decreasing size.
Comparisons. B. sp. 3 is about the same size as B. balihaiensis and B. cushmani (Fig. 4). The sinuous, high-
arched lateral outline with distinct caudal process is shared with B. alcyonicola, B. cratericola and B. cushmani
(group 3), but the (decalcified) surface is smooth, lacking the punctate texture of these and most species. The
almost subtriangular LV outline resembles that of B. angolensis but has a more distinct caudal process.
Bairdoppilata sp. 4
(Figs. 17G–P)
Material. One adult female specimen 3963F.
Dimensions. Adult female 3963F, carapace measured before dissection: Length 1084 µm, height 732 µm. See
also Table 1 and Figs. 2 and 4.
Descriptive comments. Carapace rounded-subtriangular in lateral outline, with very high-arched dorsal
margin and nearly straight, steeply sloping anterodorsal margins, lacking angles; greatest height 0.67 of length,
located at 0.64 of length; anterodorsal corner not distinguishable, located approximately at 0.33 of length and 0.89
of height; posterodorsal corner not distinguishable, located approximately at 0.67 of length and 0.84 of height;
anterodorsal brim located at 0.69 of height; posterodorsal brim located at 0.43 of height; greatest height of ventral
indentation located at 0.38 of length and 0.7 of height. Slope (measured from horizontal) of mid-dorsal margin 1
o
,
of anterodorsal margin 38
o
, of posterodorsal margin 44
o
, of dorso-caudal margin 35
o
, of hinge 1
o
. Caudate, with
weak concavity separating dorsal-caudal margin from posterodorsal margin; length of caudal process 0.06 of
length. Plumose setae with barely visible barbs. Hinge striate, ends only slightly expanded and scalloped. Surface
apparently smooth, with distinct reticular pattern of micropapillae. Seta 2 of furca with a proximal cluster of about
8 lateral barbs (on each side), followed by much tinier barbs or nearly smooth distally.

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FIGURE 14. Bairdoppilata sp. 3, female specimen 3923F. A, first thoracic leg, some setae of vibratory plate are obscured; B,
fringed claw of mandible palp; C, basal podomere (I) of second thoracic leg, left side; E, basal podomeres (I, II) of third
thoracic leg, right side; E, terminal podomeres (II–V) of third thoracic leg; F, reflexed setae of maxilla; G, vibratory plate of
maxilla; H, terminal podomeres (IV–VI) of antenna; I, distal claws of antenna with aesthetasc. Scale bar = 100 µm.
Comparisons. B. sp. 4 is significantly larger than most shallow-water species (Fig. 4). The broadly arched
dorsal margin and gently extended caudal process place B. sp. 4 in shape group 3, together with B. alcyonicola, B.
cratericola and B. cushmani. From all of these it differs by the smooth surface with micropapillate micro-
ornament, which may reflect cell arrangement in the underlying epidermis.

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FIGURE 15. Bairdoppilata sp. 3, female specimen 3923F. A, palp and masticatory processes of maxilla; B, teeth of mandible;
C, masticatory organ; D, zygum, genital lobe, furca and post-abdominal bristle. Scale bar = 100 µm.
Genus Paranesidea
1969 Paranesidea Maddocks: 39.
1995 Paranesidea Maddocks—Maddocks: 206
Species included. The genus Paranesidea should be restricted to a fairly short list of distinctive species inhabiting
tropical coralline and phytal environments. The following list includes only named species whose soft parts have
been described, at least in part, listed alphabetically by original binomen:
Paranesidea algicola Maddocks, 1969 (Nosy Be, Madagascar)
Bairdia arostrata Kornicker, 1961 (Bahama Islands, Florida Keys)
Paranesidea fracticorallicola Maddocks, 1969 (Nosy Be, Madagascar)
Bairdia gigacantha Kornicker, 1961 (Bahama Islands)
Paranesidea harpago Kornicker, 1961 (Bahama Islands)
Paranesidea onslowensis Hartmann, 1978 (Northwest Australia)
Paranesidea parva Hartmann, 1978 (Northwest Australia)
Paranesidea posidonicola Hartmann, 1979 (Southwest Australia)
Paranesidea spongicola Maddocks, 1969 (Nosy Be, Madagascar)
Paranesidea sterreri Maddocks in Maddocks & Iliffe, 1986 (Bermuda)
Comparative comments. Because the two available species of Paranesidea are represented by only three,
poorly preserved, female specimens (Table 1), they contribute little to our existing understanding of the genus.
Observations and comparisons, where possible, have been inserted above within the discussion of Bairdoppilata.

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FIGURE 16. Bairdoppilata sp. 3, all female specimen 3923F. A, RV exterior with attached dorsal fragment of LV; B, LV
exterior, dorsal margin is torn; C, masticatory organ; D–E, LV exterior, anterior and posterior regions; F, right exterior of whole
animal before dissection; G, left side of mouth region, with upper lip, atrium and lower lip; H, LV exterior with smooth surface,
MSP with adductor muscles, NPC and carapace setae (detail of fig. B); I–J, distal claws of both antennae; K, right exterior of
whole animal, posterior end, showing RPC, simple and plumose setae; L, right exterior of whole animal, posterodorsal region
to show accessory dentition, higher focus than fig. K. Scale bar = 50 µm.

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FIGURE 17. A–F, Bairdoppilata sp. 3, female specimen 3923F; G–P, Bairdoppilata sp. 4, female specimen 3963F. A, right
exterior of whole animal before dissection, posterior region; B, enlarged proximal barbs of seta 2 of furca; C–D, genital lobes;
E, left side of posterior body with zygum, furca and genital lobes; F, dorsal view of genital lobes, zygum, furcae, and post-
abdominal bristle; G, distal claws of antenna; H–I, tapering proximal barbs of seta 2 of furca; J, masticatory organ; K, right
exterior of whole animal, filled with secondary crystals; L, valve fragment with setae and micropapillae; M–N, left exterior of
whole animal, caudal region, showing plumose setae at high and low focus; O, left genital lobe and edge-bar of zygum; P,
dorsal view of hinge region of decalcified carapace with sinusoidally folded chitin ribbons. Scale bar = 50 µm.

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FIGURE 18. Paranesidea sp. 1, all female specimen 3968F. A, antenna; B, distal claws of antenna with delicate aesthetasc, the
fused seta is only faintly serrate, as usual for females; C, furca, seta 2 is incomplete; D, mandible; E. palp of maxilla with
fringed claw. Scale bar = 100 µm.
Paranesidea sp. 1
(Figs. 18–20)
Material. Four adult females.
Dimensions. Adult female 3968F, carapace measured before dissection: Length 984 µm, height 660 µm. See
also Table 1 and Figures 2 and 4.
Descriptive Comments. Carapace smooth, high-arched; subtriangular-subcircular in lateral outline, with
continuously curving dorsal margin, lacking anterodorsal and posterodorsal corners; greatest height located at 0.47
of length; anterodorsal brim located at 0.04 of length and 0.61 of height; posterodorsal brim located at 0.97 of
length and 0.39 of height; highest point of ventral indentation located at 0.38 of length and 0.05 of height. Slope
(measured from horizontal) of anterodorsal margin 37
o
, of posterodorsal margin 52
o
, of dorso-caudal margin 49
o
, of
hinge 11
o
. Caudal extension short, broad, nearly triangular, with indistinct posterodorsal break in slope. No patch
pattern is visible in the decalcified carapace, but pigment spots are evident. The fused antennal claw is distinctly
pectinate.
Comparisons. P. sp. 1 lacks the conspicuously punctate surface texture and caudate outlines of the several
species of Paranesidea described from Nosy Be. The carapace dimensions of P. sp. 1 would be at the uppermost
edge of the size range for females of P. spongicola and too large for most of the others.

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FIGURE 19. Paranesidea sp. 1, female specimen 3968F. A, third thoracic leg; B, second thoracic leg; C, first thoracic leg; D,
vibratory plate of first thoracic leg; E, left side of mouth region, with forehead, upper lip, lower lip and atrium; F, genital lobe.
Scale bar = 100 µm.
P. sp. 1 cannot be identified with any of the species of “Bairdia” reported by Holden (1967) from Hawaii or
with any of the four species of Paranesidea reported by Holden (1976) from the Lower Miocene of Midway Island.
Its anteroventral margin slopes more steeply than that of P. sp. 1 of Holden (1976, Pl. 9, fig. 6), and the pigment
spots are smaller and round, rather than dorsoventrally elongated. Its upright, nearly triangular lateral outline
resembles that illustrated for P. sp. 2 of Holden (1976, Pl. 9, fig. 5), but it is substantially larger, with less produced
anterodorsal and posterodorsal angles. It is larger than P. onslowensis and smaller than P. poseidonicola.

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FIGURE 20. Paranesidea sp. 1. A–K, female specimen 3968F; L–S, female specimen 3973F. A, RV exterior, with attached
dorsal fragment of LV; B, LV exterior, dorsal margin is torn; C, LV exterior, posterior margin; D, right exterior of entire animal
before dissection, filled with secondary concretions; E–H, masticatory processes and fringed claw of palp of maxilla; I, genital
lobe; J, distal claws of antenna; K, right exterior, anterodorsal region of fig. D, with median eye, isthmus and antennule; L–M,
RV and LV exteriors, fragmentary; N–P, genital lobes before and after dissection; Q, serrate fused claw of antenna; R, distal
claws of antenna; S, RV exterior, posterior region (detail of fig. L). Scale bar = 50 µm.

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FIGURE 21. Paranesidea sp. 2; all female specimen 3961F. A, left exterior of whole animal before dissection, filled with
secondary concretions; B–D, fringed claws of palp and masticatory processes of maxilla; E–F, distal claws of antenna; G,
pectinate distal claw of mandible palp; H–I, genital lobes; J, detail of Fig. I showing longitudinal striations within tubes; K–L,
hinge region of carapace, showing straight connecting ribbons and micropapillate surface texture; M, anterior margin with
RPC, selvage, flange, pappose and eyelash setae; N–Q, details of Fig. M at high and low focus, to show pappose setae; R,
fragment of decalcified carapace, showing NPC and reticular pattern of micropapillae; S, detail of left exterior before
dissection, with setae and micropapillae. Scale bar =50 µm.

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Paranesidea sp. 2
(Fig. 21)
Material. One adult female.
Dimensions. Adult female 3961F, carapace fragmentary and dimensions approximate: Length 916 µm, height
630 µm. See also Table 1 and Figures 2 and 4.
Descriptive comments. Carapace apparently smooth; rounded-subhexagonal (bairdian) in lateral outline, with
narrowly arched dorsal margin, indistinct anterodorsal and posterodorsal corners; greatest height located at 0.47 of
length; anterodorsal brim located at 0.04 of length and 0.60 of height; posterodorsal brim located at 0.98 of length
and 0.43 of height; highest point of ventral indentation located at 0.46 of length and 0.07 of height. Slope
(measured from horizontal) of anterodorsal margin 39
o
, of posterodorsal margin 51
o
, of dorso-caudal margin 17
o
, of
hinge 4
o
. Distinct caudal process. No patch pattern or pigmentation is visible in the decalcified carapace. The
assignment to Paranesidea is based on the moderately pectinate character of the fused antennal claw, as usual for
females of this genus. The apparently smooth surface with a reticulate pattern of tiny micropapillae (which may
reflect cells in the underlying epidermis) is noteworthy.
Comparisons. The carapace dimensions of P. sp. 2 would fall in the upper part of the size range for P.
spongicola and close to that of P. onslowensis. It is much smaller than P. poseidonicola and it lacks the expanded
caudal flange and marginal denticles of that species. The upright, distinctly caudate shape cannot be equated to any
of the species of “Bairdia” or Paranesidea described by Holden (1967, 1976) from Hawaii or Lower Miocene of
Midway.
Acknowledgments
I am grateful to Dr. Louis S. Kornicker, long-time Curator and Scientist, Department of Invertebrate Zoology,
Smithsonian Institution, for suggesting this study; and to Dr. S.L. Coles, Bishop Museum, Honolulu, for
permission to study these collections. The suggestions of two reviewers, Dr. Dan L. Danielopol and Dr. Karel
Wouters, are greatly appreciated.
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APPENDIX I. Descriptions of sampling stations and specimens of species collected.
FRENCH FRIGATE SHOALS
French Frigate Shoals stations, Northwestern Hawaiian Islands Rapid Assessment Monitoring Program (NOW-RAMP), Sep
2000, specimens collected by R. DeFelice ("FFS-R") and D. Minton ("FFS-TC"). TC = Townsend Cromwell.
FFS-TC-06
(23
o
52.5 N, 166
o
17.68 W), 11 Sep 2000
N forereef, 1.5 km N of Tern Island, depth 5–12 m
Spur and groove habitat with moderate current and surge. Top of spurs at 5 m depth were scoured carbonate pavement with
encrusting Porites. Some areas of dead coral matrix on spur top and slopes. Most common macro invertebrate was the boring
urchin Echinostrephus aciculatus. Grooves at 10–12 m with carbonate sediments and rubble deposits shifting with surge.
3 Bairdoppilata scaura n. sp.
FFS-TC-7
(23
o
52.0 N, 166
o
18.28 W), 11 Sep 2000
N forereef, 2 km W of Tern Island, depth 5–10 m
Spur and groove habitat with very little current or surge. Spurs about 5 m high and 5 m wide with encrusting Porites lobata
dominating upper exposed surfaces. Undercut spur base showed evidence of occasional scour from intermittent surge. Groove
floors at 10 m depth were about 5 m wide. Groove channels with abundant carbonate sediments and coral rubble. Unlike the
previous station, Echinometra were relatively rare.
1 Bairdoppilata scaura n. sp.
FFS-TC-13
(23
o
43.957 N, 166
o
10.375 W), 14 Sep 2000
Central lagoon patch reef, 0.5 km W of Gin Islands, depth 3–10 m
Isolated patch reef with base at about 10 m depth surrounded by flats of shifting coarse sediment with some rubble deposits.
Patch reef heavily eroded with abundant turf algae and blue-greens. Top of reef with encrusting Porites compressa.
4 Bairdoppilata sp. 3
FFS-TC-16
(23
o
38.265 N, 166
o
10.768 W), 15 Sep 2000
S perimeter forereef, 1 km S of Disappearing Island, depth 12–18 m
Sloping forereef terrace with moderate relief of mounds, short ledges, shallow grooves and holes with some sand and rubble
deposits. Dominant corals were Porites lobata and Pocillopora spp.
1 Bairdoppilata scaura n. sp.
KÂNE’OHE BAY, O’AHU
Station 1, North Channel
(21
o
30’22.1”N, 157
o
50’57.1”W), 17 Nov 1999
North Channel, depth 1–6 m.

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Zootaxa 4059 (2) © 2015 Magnolia Press
Wave-turbulent area along south side of main channel near Buoy Marker 5, on reef area with abundant live coral extending
from 4 to 6 m depth next to dredged channel. Outside of live coral area bottom is consolidated limestone with coarse sand and
moderate Halimeda algae cover.
1 Paranesidea sp. 2
Station 5, Waiahole Reef
(21
o
29’14.6, 157
o
49’55.8), 17 Nov 1999
Waiahole Reef, near Channel Marker 4, depth 7.5 m.
Very abundant (75–90% coverage) of Porites compressa, Montipora capitata, and Sarcothelia edmondsoni and moderate cover
of Dichtyosphaeria cavernosa in 1.5 m depth. Other corals present to moderately abundant were Pocillopora damicornis,
Pocillopora meandrina, Porites lobata and Montipora patula. High Porites compressa coverage extends down to ca. 7 m.
1 Bairdoppilata sp. 4
Station 12, He’eia Kea Pier
19 Jan 2000
(21
o
26’37.8”N, 157
o
48’40.3”W), 19 Jan 2000
He’eia Kea Pier, depth 0–7 m.
Northwest side of main He’eia pier and on a shallow reef flat separated from dock by a narrow channel. Pier pilings only
sparsely covered with fouling organisms, mostly filamentous algae. Channel has abundant algae on bottom, and reef flat is
covered with coarse sand and heavy growths of algae, especially introduced Gracilaria salicornia, Hypnea musciformis,
Kappaphycus spp. and Acanthophora spicifera.
4 Bairdoppilata sp. 2, 2 Paranesidea sp. 1.
Station 15, Coconut Island Reef
(21
o
26’38.5”N, 157
o
47’47.2”W), 8 Nov 1999
Coconut Island Reef, depth 6 m.
West of the dolphin pens of Hawai’I Institute of Marine Biology (HIMB). This area is directly leeward of where nonindigenous
algae Kappaphycus spp. were introduced in 1974-76 and escaped from the pens when they broke up in a storm. Consequently,
this is the area most heavily impacted by these algae, which have formed an essential monoculture covering large areas of the
reef flat, edge and slope with dense growths with algal thalli many cm thick. The introduced alga Acanthophora spicifera is
also very abundant along with Gracilaria salicornia, Dictyosphaeria cavernosa and Dictyosphaeria versluysii. The habitat
space is so completely monopolized that few benthic invertebrates were noted.
1 Bairdoppilata sp. 2, 1 Paranesidea sp. 5
Station 18, Makani Kai Marina
(21
o
25’06.0”N, 157
o
47’23”W), 8 Nov 1999
Makani Kai Marina, depth 0–3 m.
Floating docks and concrete wall in a harbor area nearly isolated from the bay by a rock jetty and much affected by fresh water
runoff. Introduced oysters Crassostrea sp., fanworm Sabellastarte spectabili and bryozoan Amathia distans are abundant along
the concrete wall along with numerous other fouling invertebrates. Few macro-organisms were seen on the floating docks. Only
a single coral species Montipora capitata occurred at 2 m depth, well below freshwater lens.
1 Bairdoppilata sp. 2
Station 21, Kâne’ohe Yacht Club patch reef
(21
o
25’09.5”N, 157
o
46’22.6”W), 5 Nov 1999
Kâne’ohe Yacht Club patch reef, depth 1–5 m.
Located between entrance channels to the Yacht Club, the reef extends from 0.5 m down to 10 m depth. Reef top is mixed
Montipora capitata and Porites compressa corals in coarse sand with abundant macroalgae Gracilaria salicornia,
Dictyosphaeria cavernosa and Kappaphycus spp. Introduced invertebrates, e.g. the polychaete Sabellastarte spectabilis, the
ascidian Phallusia nigra and the hydroid Pennaria disticha were abundant, along with the synaptid holothurian Opheodesoma
spectabilis.
1 Bairdoppilata sp. 2
Station 22, Marine Corps Base Fuel Dock
(21
o
26’15.3”N, 157
o
45’55.1”W), 15 Jan 2000
Marine Corps Base Fuel Dock, depth 0.5–8.0 m.
Concrete and iron pilings of the main docking area for the Kâne’ohe Marine Base on the southwest side of Môkapu Peninsula
and east of the base runways. Surfaces are densely fouled with heavy coverage of sponges and other invertebrates to the bottom
in 9 m depth. Sponges are sediment coated, and bottom is silty fine sediment with numerous burrow openings.
5 Bairdoppilata sp. 2