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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3884 (5): 445
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http://dx.doi.org/10.11646/zootaxa.3884.5.5
http://zoobank.org/urn:lsid:zoobank.org:pub:AEF16C1C-5E1D-4A4C-A1A3-096F439C15B5
A review of the Polystira clade—the Neotropic’s largest marine gastropod
radiation (Neogastropoda: Conoidea: Turridae sensu stricto)
JONATHAN A. TODD
1, 3
& TIMOTHY A. RAWLINGS
2
1
Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK. E-mail: j.todd@nhm.ac.uk
2
Department of Biology, Cape Breton University, 1250 Grand Lake Road, Sydney, Nova Scotia B1P 6L2, Canada.
E-mail: timothy_rawlings@cbu.ca
3
Corresponding author
Abstract
The Polystira clade (here comprising Polystira and Pleuroliria) is a poorly known but hyper-diverse clade within the
neogastropod family Turridae (sensu stricto). It has extensively radiated within the tropics and subtropics of the Americas, to
which it is endemic. In this paper we present a synthetic overview of existing information on this radiation together with new
information on estimated species diversity, systematic relationships, a species-level molecular phylogenetic analysis and
preliminary macroecological and diversification analyses, to serve as a platform for further study. We currently estimate that
about 300 species (122 extant) are known from its 36 million year history but this number will undoubtedly increase as we
extend our studies. We discuss the relationships of Polystira to other Neotropical Turridae (s.s.) and examine the taxonomy and
systematics of the geologically oldest described members of the clade. To aid taxonomic description of shells we introduce a
new notation for homologous major spiral cords. Focusing on key publications, we discuss in detail the changing historical
understanding of the taxonomy of the clade and the relationships of its component genus-level taxa: Polystira Woodring, 1928,
Pleuroliria de Gregorio, 1890, Josephina Gardner, 1945 and Oxytropa Glibert, 1955. We designate a neotype for Pleurotoma
(Pleuroliria) supramirifica de Gregorio, 1890, to stabilize our understanding of this, the type species of Pleuroliria.
Application of the name Oxytropa is restricted to the type species. The genus Polystira is conchologically re-described and for
the first time we synthesize available information on the anatomy, feeding and toxinology, reproduction and life history, larval
modes and life habits, and geographic and bathymetric ranges of its species. We give an updated list of the 19 formally
described living species and present the pitfalls of the currently poor species-level taxonomy of Polystira using case examples.
We present a molecular phylogenetic analysis of 22 extant species using three mitochondrial gene fragments (COI, 12S rRNA
and 16S rRNA). This reveals undescribed species and indicates that Recent genetic clades (‘biospecies’) are consistent with
finely divided conchological ‘morphospecies’. Historically, there has been a slow realisation of the high species diversity of the
Polystira clade and we consider that this may be due to inadequate precision of morphological description of shells and a lack
of clear homology statements. We suggest how these both might be improved. Finally, using a data compilation based on
museum specimens we examine species range-size distributions and species abundance distributions for 85 of the 112 extant
western Atlantic species that we have delimited to date. Our results indicate that the majority of species are rare and have short
geographic ranges; only a few are wide-ranging and abundant. This has important implications for surveys of biodiversity.
Key words: Turridae; Polystira; Pleuroliria; systematics; homology; molecular phylogenetics; species hyper-diversity;
radiation; Neotropics; Americas; geographic range; species abundance; diversification dynamics
Introduction to the Polystira radiation
Polystira Woodring, 1928 is an extant genus of marine neogastropod belonging to the family Turridae as it is
currently restricted (Bouchet et al. 2011). The genus is endemic to the Americas where it is largely confined to the
tropics and subtropics today (Powell 1964). Polystira comprises a clade with an extensive history and superb fossil
record throughout the region that extends as far back as the late Eocene (ca 36 Ma) in the southern USA. Over this
long history, Polystira and its close relative Pleuroliria (termed the ‘Polystira clade’ herein) have diversified
extensively. Comprising an estimated 122 discovered living species and >178 known extinct species (see Table 1
for details of current species diversity estimates), Polystira has become the most species-rich marine snail genus in
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the Americas and apparently the most species-rich living genus within the family Turridae (compare with Tucker
2004: table 2). Despite fairly limited morphological variation (=disparity) between taxa within the genus, all
currently known Recent species can be discriminated using shell characters. As such, Polystira may serve as a
model taxon with which to combine palaeontological and neontological approaches to delimiting species and
clades, reconstructing phylogeny, and interpreting evolutionary history. In addition, Polystira offers the
opportunity to examine the factors that may have promoted its enormous species diversification relative to all other
marine snail genera in the region (Todd & Johnson 2013).
Surprisingly little is known of the basic biology of any species of Polystira, even though some species, notably
Polystira albida (Perry) and P. s t a r re tt i Petuch, are among the most common large carnivorous gastropods in
tropical shallow shelf habitats of the western Atlantic. The same is true of P. oxytropis (G. B. Sowerby I) and P.
picta (Reeve) in the eastern Pacific; all these species have ranges spanning thousands of kilometres.
The purpose of the present paper is to place Polystira within its larger systematic context and to provide a
baseline for the more detailed studies underway on this genus by summarizing what is known of its biology,
ecology, distribution, species diversity, fossil history and diversification dynamics.
TABLE 1. Summary of the species diversity counts and estimates that include undescribed species (current in April 2014)
(est.) referred to throughout this paper.
Abbreviations and references
We have given taxonomic authority citations in the references if they deal specifically with Polystira and close
relatives; we exclude papers where new species are introduced if these works are not further discussed. Other
references can be found in Tucker (2004).
Ma = million years ago
My = million years
AMNH: American Museum of Natural History, New York, USA
ANSP: Academy of Natural Sciences of Philadelphia, Philadelphia, USA
FMNH: Field Museum, Chicago, Illinois, USA
FLMNH: (current recommended institutional coden: UF): Florida Museum of Natural History, Gainesville,
Florida, USA
HMNS: Houston Museum of Natural Science, Houston, Texas, USA
LACM: Los Angeles County Museum of Natural History, Los Angeles, California, USA
MHNMC: Museo de Historia Natural Marina de Colombia, Santa Marta, Colombia
Data set Data source N
Total known (extinct + extant) est. (1) + (2) + (3) + (4) below >300
Extant total (WA+ EP) est. (1) (Todd unpubl.) 122
Extant Western Atlantic est. (WA) (Todd & Johnson 2013: p. 3) 112
Extant WA + EP described (including synonyms) this paper 23
Extant WA valid (incl. one preoccupied name) this paper 14
Extant East Pacific (EP) est. (Todd & Johnson 2013: p. 4); this paper >10
Extant EP valid this paper 5
DNA-sequenced WA + EP ‘biospecies’ this paper 22
Extant + extinct SW Caribbean (Costa Rica + Panama) est. (Todd & Johnson 2013: online Table 3) 114
Extant SW Caribbean (Costa Rica + Panama) est. (Todd & Johnson 2013: online Table 3) 26
Extant from Trinidad eastwards to Surinam est. (Todd unpubl.) 15
Extant southern Florida + Florida Keys est. (Todd unpubl.) 10
Extinct Costa Rica + Panama: Miocene–Pleistocene est. (2) (Todd & Johnson 2013: online Table 3) 88
Extinct Dominican Republic: Miocene–Pliocene est. (3) (Todd unpubl.) 60
Extinct Venezuela: Miocene–Pliocene est. (4) (Todd work in progress) >30
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MNHN: Muséum National d’Histoire Naturelle, Paris, France
MZUSP: Museu de Zoologia, Universidade de São Paulo, Brazil
NHM (current institutional coden: NHMUK): Natural History Museum, London, UK
RMNH: Rijksmuseum van Natuurlijke Histoire; collections now held in NCB Naturalis, Leiden, The Netherlands
UMML: University of Miami Marine Laboratory, Rosenstiel School of Marine and Atmospheric Science Miami,
Florida, USA
USNM: National Museum of Natural History; Smithsonian Institution; Washington, DC, USA
The Polystira clade: supraspecific taxonomy and systematics
Polystira and its relationship to other Neotropical Turridae
The genus Polystira Woodring, 1928 has become the name popularly associated with all extant Polystira-like turrid
species in the Neotropics and subtropics. Apart from Polystira, the only other living turrids (as restricted by
Bouchet et al. 2011) inhabiting the same region are species of Gemmula Weinkauff, 1875 and the deep water genus
Cryptogemma Dall, 1918, both of which occur in the eastern Pacific and the western Atlantic.
Regionally, Gemmula is best known through G. hindsiana Berry, 1958—the type species (tropical eastern
Pacific)—and G. periscelida (Dall, 1889) (tropical western Atlantic). Our examination of museum and newly
collected material shows that G. hindisiana consists of multiple species with distinct protoconch and teleoconch
morphologies and that this composite taxon and G. persiscelida do not appear to be closely related to each other
based upon both conchological character states and molecular phylogenetic analysis (see ‘Monophyly of Polystira’
below). In a paper in which two new species of Gemmula were described from the Philippines, Olivera (2004)
described and figured the morphology of a “Caribbean form” of one of his new species, Gemmula sikatunai. He
detailed conchological differences between the Caribbean (Barbados) form and the Philippine type material. We
consider that the specimens from off Barbados very likely belong to an unnamed species. We have also seen a few
more undescribed Gemmula species from the Caribbean.
Here it should be noted that the recently described Gemmula mystica Simone, 2005 living in >500 m water
depth off São Paulo, Brazil, should be excluded from Gemmula. The shell appears to lack a well-developed
principal spiral A otherwise present in the Turridae (see ‘Homology of spiral ornamentation in Turridae’ below),
has an unusually robust rostrum, and the radula has hypodermic marginal teeth that are lacking in the Turridae
(sensu stricto; s.s. hereafter) (Bouchet et al. 2011). This species is of uncertain systematic position but may belong
to the family Borsoniidae.
Regionally, the genus Cryptogemma Dall, 1918 is known from three species in the East Pacific occurring in
water depths of 1,159–2,490 m (McLean 1971) and at least one apparently undescribed species from the Caribbean
off Colombia (see Invemar online database at http://siam.invemar.org.co/siam/). We have not examined specimens
of these species, but at least C. eldorana (Dall, 1908) appears from its teleoconch morphology to belong to the
Turridae and Cryptogemma has been placed in the Turridae based on radular characters by McLean (1971a) and
Bouchet et al. (2011). Cryptogemma polystephanos (Dall, 1908), judging from the figure of the holotype given by
McLean (1971b: fig. 1652), may not belong to the Turridae (s.s.).
Historical considerations: wider relationships of the Polystira clade
Casey (1904: 130) considered the Polystira clade (as Pleuroliria) to be the “American homologue of [his newly
introduced Indo-Pacific genus] Lophiotoma. In contrasting the shells of these genera Casey (1904: 130–131) stated
that Polystira “is composed of much smaller species having a slender form, very characteristic sculpture of two to
three strong spiral carinae, the peripheral bearing the small anal sinus, and a conspicuous system of lines of growth,
bi-oblique toward the peripheral carina and composed of excavated lines, which are less evident in the very early
forms and most conspicuous in the modern species”. Later, Powell (1964: 303, 315) considered Polystira to differ
from Lophiotoma more simply by its “more shallow and broadly V-shaped sinus [see Powell 1942: text fig. E.2, 3],
and protoconch differences” and he thought them to “probably [share] a common ancestry”, so presumably he
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considered them to be sister clades. In contrast, Hickman (1976: 96–97) considered that Polystira and Pleuroliria
were not monophyletic with respect to species membership and implied that together they did not comprise a clade
exclusive of other Turridae. Hickman was pessimistic about the taxonomic value of existing generic names within
the Turridae, and considered that, “the generic names Polystira Woodring, 1929 (sic), Pleuroliria de Gregorio,
1890, and Lophiotoma Casey, 1904, as well as a number of subgeneric names, can be applied with certainty only to
their type species within a large polyphyletic group of fossil and living fusiform turrine species sculptured with
spiral cords and keels and having a siphonal notch located on the periphery.”
Based on a cladistic analysis of foregut characters of a wide range of Turridae (and some other ‘turrids’),
Polystira (P. sp., identified as P. formosissima (E.A. Smith, 1915)) and Cryptogemma (represented by a Japanese
species, C. corneus (Okutani, 1966)) have been suggested to form a sister clade relative to the other Indo-Pacific
taxa studied (Medinskaya 2002). Given the complexity of relationships within ‘Gemmula’ and between it and
other Turridae, however, it remains to be seen if a simple Neotropical/Indo-Pacific biogeographic split within the
extant Turridae will be supported by integrative systematic studies.
Monophyly of Polystira
Recently a preliminary molecular phylogenetic analysis of the Turridae (s.s.) has been undertaken as part of a study
to establish the major clades (families) within the Conoidea (Puillandre et al. 2011a). This study analyzed portions
of three mitochondrial genes (COI, 12S rRNA and 16S rRNA) for 102 genera of conoideans. However, only a
single species exemplar was used for each genus, so for the Turridae (s.s.) just eight species and genera were
analyzed. Maximum likelihood analyses of the concatenated dataset indicated the Turridae to be a well-supported
clade with Polystira being the sister taxon to the rest of the clade as follows: (Ptychosyrinx, (Turridrupa,
(Lophiotoma, Gemmula) (Turris (Xenuroturris, Ioturris)))) (Puillandre et al. 2011a: fig 1). These relationships can
only be considered to be robust for the species sampled; future analyses will require much more complete species
sampling to fully establish intergeneric molecular relationships within the Turridae.
Our molecular phylogenetic analyses of the interspecific relationships of Polystira species uses the same gene
fragments (COI, 12S and 16S) and reveals a strongly supported clade of Polystira to be monophyletic relative to a
turrid (s.s.) outgroup comprising eight species of Gemmula, Ptychosyrinx, Turris, Xenuroturris and
Unedogemmula. We discuss our analysis in ‘Current species diversity’ below.
FIGURE 1. Homology of seven primary (or principal) spiral cords (A–G—notation introduced herein) in four morphologically
disparate species of Polystira. Penultimate and final whorls of (left to right) two undescribed species, P. coltrorum Petuch and
P. tellea (Dall) showing variation in outline shape, strength, spacing and intercalary cord development and, also rostrum length.
Not to scale.
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Principal spiral cord terminology
To aid description throughout this paper we will briefly outline a new terminology (Fig. 1) for labelling
individualized and homologous major (or ‘primary’) spiral cords that will be discussed in more detail elsewhere
(see also ‘Homology of spiral ornamentation in Turridae’, below). In Polystira the following applies:
Spiral A is the most adapical major spiral cord that develops early in ontogeny (herein ‘principal’ spiral cord
A).
B is the principal spiral cord that coincides with the apex of the anal sinus.
Spiral C is the principal cord that develops in the interspace of B and D.
Principal spiral D coincides with the abapical suture and around succeeding whorls attach—at least in early
ontogeny.
Principal spiral E is the first principal cord abapical to D (below the suture) and is covered with inductura
forming (sometimes with F) an angled parietal lip within the aperture.
Principal spiral F lies between spirals E and G.
Principal spiral G delimits the adapical end of the rostrum and coincides with the vertex of the eyestalk sinus.
Oldest known members of the Polystira + Pleuroliria clade
We give a selection of the published discussions on the interrelationships of the named genus-level taxa below
under: “The Polystira clade: Polystira and other names”. The palaeontological record of early Polystira-like
species is quite extensive in the latest Eocene and early Oligocene of the US states along the Gulf of Mexico and
was first described largely by Casey (1904) and later more fully described and figured by Harris (1937). In more
recent years, MacNeil & Dockery (1984) have described some new species and redescribed previously known
species from these strata. Below, we discuss the morphology and systematic relationships of six species-rank taxa
that lay claim to be amongst the earliest members of the Polystira clade. One of us (JAT) has made preliminary
direct observations and we provide notes on these six taxa below. Detailed morphological analyses, including SEM
imaging, have yet to be undertaken so, although we believe our systematic conclusions to be robust, more detailed
morphological observations are necessary for any future taxonomic revision.
1) Pleuroliria crenulosa Casey, 1904: Cook Mountain Formation, Upper Claiborne Group, middle Eocene
(Bartonian) of St. Maurice, Louisiana (Palmer & Brann 1966; Tucker 2004: 263).
Figured syntype, USNM 494343 (Harris 1937: pl. 1, fig. 8a) examined. Principal spiral B is lightly beaded and
unicarinate. Spirals A through G are all present.
2) Pleuroliria crenulosa crescens Harris, 1937: Middle or Upper Claiborne Group, middle Eocene of Sabine
River, Sabine Parish near Columbus, Louisiana (Palmer & Brann 1966). Given subspecies rank by Palmer & Brann
(1966) (see Tucker 2004: 263).
Figured holotype, PRI 2361 (Harris 1937: pl. 1, fig. 11) examined. This is a worn shell with a broken
protoconch and rostrum. The protoconch comprises more than 3 whorls and is smooth until its last ca ¾ whorl
which, though worn, is seen to be axially ribbed (ca 8 axial ribs seen per half-whorl). There is a clear protoconch/
teleoconch junction followed by principal spirals A, B and C, with C rising from its position at the basal suture
almost immediately; the suture presumably then following spiral D. Spiral B is vaguely gemmulate for ca 1¾
whorls then becomes smooth. Spirals A, B, C and D are +/- equidistant throughout growth. In the last 1¼ whorls
weak intercalary spirals become clearly visible between the primary spirals. In the last whorl weak, raised axial
ornament/growth stops begin which continue until the broken aperture.
3) Pleuroliria simplex Casey, 1904: Cook Mountain Formation, Upper Claiborne Group, middle Eocene
(Bartonian) of St. Maurice, Louisiana (Palmer & Brann 1966; Tucker 2004: 908)
Figured holotype, USNM 494342 (Harris 1937: pl. 1, fig. 6) examined. Early whorls have relatively strong
principal spirals A, B and D. Spiral B is rounded and for its first ca 2–2½ whorls is slightly irregularly noded where
crossed by irregularly raised collabral growth lines. Axial ornament is very fine and irregular; occasionally as
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slightly stronger growth pauses that generate vague nodes where they cross spiral B. After about 2½ teleoconch
whorls an intercalary spiral appears in the spiral B-C interspace and rapidly becomes equally as strong as B and C,
and these three spirals become equally spaced. The final whorl has spirals A, B, B-C intercalary spiral, D, a weaker
E and F, stronger E-F intercalary spiral, and a strong G.
As first noted by Harris (1937), taxa 1–3 all have a ‘crenulate’ carina (= spiral B herein) which immediately
separates them from all other members of the Polystira clade, with the sole exception of Pleuroliria jacksonella
Casey (see below). MacNeil & Dockery (1984: 176) noted that Casey (1904: 131) keyed out taxa 1 and 3 from his
other species [of what Casey and MacNeil & Dockery considered Pleuroliria] by their having “lines of growth less
pronounced, uneven and never deeply incised”. Casey also noted them to be of small size and geologically older
(Casey 1904: 131). This led MacNeil & Dockery to consider that “if these species are actually referable to
Pleuroliria they should probably be set aside as a distinct subgenus”. This should not be confused with Casey’s
own informal division of Pleuroliria into two groups of species based on possession of multispiral versus
paucispiral protoconchs, a division he considered to be “almost subgeneric in value” (Casey 1904: 131). MacNeil
& Dockery’s suggested new subgeneric-rank taxon falls within Casey’s group characterized by its multispiral
protoconch. We have no hesitation in placing MacNeil & Dockery’s putative supraspecific taxon within the
Turridae (s.s.) based on the presence in Pleuroliria crenulosa and its unnamed ‘varieties’ (Harris 1937: pl. 7, figs
8–10)—and probably P. crenulosa crescens too, of; 1) seven strong principal spirals seemingly homologous to A
through G of Polystira and the Turridae 2) the vertex of the anal sinus lying on spiral B, and 3) a multispiral
protoconch, the first (four?) whorls of which are smooth and the last with axial ribbing. This putative supraspecific
taxon shares a number of characters with Polystira and Pleuroliria, including the suture tracking spiral D and the
anal sinus centred on a unicarinate, wide-based, spiral B. However it differs in lacking conspicuous, regular and
well-defined raised axial ornament on the flanks of the whorls, though very fine axial ornament may be present on
the base and rostrum. It is possibly a sister taxon to Polystira +Pleuroliria.
4) Pleuroliria jacksonella Casey, 1904; Moodys Branch Formation, Lower Jackson Group, late Eocene
(Priabonian) of Montgomery Landing, Red River, Louisiana (Palmer & Brann 1966; Tucker, 2004: 510).
Holotype, USNM 494344 (Harris 1937: pl. 1, fig. 7; Harris & Palmer 1947: pl. 57, fig. 1) examined. This
juvenile shell has principal spirals A through G preserved. Spiral B is unicarinate and has rather low, poorly
preserved, ‘beads’ for ca 1 whorl. Axial ornament is very thin but well pronounced in the spiral A–B and B–C
interspaces. Axial ornament is seen to extend over the whole rostrum on the last whorl. Outer lip bows out slightly
over spiral F. Aperture shows three strong axial lirae. MacNeil & Dockery (1984: 176) thought that this species
was likely to be “the earliest true Pleuroliria from America”, and we concur that it appears to be the earliest and
most basal member of the Polystira + Pleuroliria clade.
5) Pleuroliria tenuis MacNeil in MacNeil & Dockery, 1984: 178–179; pl. 8, fig. 3. Red Bluff Formation, early
Oligocene (Rupelian) of Mississippi. Non-type specimens examined.
This species has very fine axial ornament, even-strength smooth and simple principal spirals A–G, a shallow V-
shaped anal sinus and a short rostrum. A number of other extinct US species (described and undescribed) share
these character state combinations. Together they may comprise a supraspecific clade of Turridae (s.s.) distinct
from Polystira + Pleuroliria.
6) Pleuroliria subsimilis Casey, 1904; Red Bluff Formation, Early Oligocene (Rupelian) of Red Bluff, Mississippi
(Tucker 2004: 960). Examined: figured holotype (acc. Harris 1937: pl. 1, fig. 4, USNM 494345) and non-type
specimens.
Pleuroliria subsimilis possesses a combination of teleoconch character states that include: 1) fine, regular and
distinctly raised collabral ornament; 2) principal spirals A and B that become morphologically more complex and
that develop subsidiary spirals upon them through their ontogeny; and; 3) a spiral B that is smooth-topped, has a
wide base, and which lacks axial crenulations or gemmulations. These states are characteristic of the clade that
includes all Recent Polystira species (and the type species of Pleuroliria; see below).
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The Polystira clade: Polystira and other names
Despite common use of the generic name Polystira, by ourselves included, a number of other genus-group names
are nomenclaturally available for taxa within what we here call the “Polystira clade”. Indeed, one of these names,
Pleuroliria de Gregorio, 1890, has nomenclatural priority over Polystira Woodring, 1928, were these names to be
considered synonymous. The Polystira clade is large enough in terms of numbers of species and conchologically
diverse (=disparate) enough for future systematic subdivision to be appropriate. We are working on providing
detailed hypotheses of the phylogenetic relationships within the Polystira clade based on an integrative taxonomy
combining shell morphological, molecular phylogenetic and other available comparative systematic data
(Rawlings et al. 2003; Todd & Rawlings 2003). Only after the type species of fossil genus-level taxa have been
accommodated within a phylogenetic framework will we be able to confidently and accurately assign the available
generic/subgeneric names to individual subclades within the Polystira clade.
We use the term Polystira clade because this is the most widely used genus-rank name for Recent species and
because our own work has started with establishing the inter-relationships of Recent species. Currently, we are still
extending our studies backwards in geological time to encompass species more usually assigned to Pleuroliria and
to establish the relationship of the Polystira clade to other turrids, both living and extinct.
Available genus-level taxa comprising the Polystira clade
Below we discuss at some length the names available at the genus-level applicable to the Polystira clade. This
focuses on the following areas: 1) providing original diagnoses and descriptions; 2) clearing-up any ambiguities
that exist with regard to their type species and their systematic content; 3) a selected discussion of published
commentary on the appropriate hierarchical rank these names should be given; as well as; 4) pertinent published
comments on the perceived relationships between these groups.
Pleurotoma (Pleuroliria) de Gregorio, 1890: 38, pl. 2, figs 46–48. Type species by subsequent designation
(Cossmann 1893: 43): Pleurotoma (Pleuroliria) supramirifica de Gregorio, 1890 = Pleurotoma cochlearis Conrad,
1848a (1848b: 115, pl. 11, fig. 23). Byram Formation, Late Oligocene, Vicksburg, Mississippi, USA (MacNeil &
Dockery 1984: 76); see discussion below for identity of the type species and its type locality.
Original description: A combined description of the new subgenus and species was provided in Latin; de
Gregorio also placed another new (and difficult to interpret) species, P. ( P.) ti z i s , in Pleurotoma (Pleuroliria).
Casey (1904: 132) synonymized P. tizis under P. cochlearis, others have regarded it as a distinct species (Harris
1937: 30; Palmer & Brann 1960): we consider it best regarded as a nomen dubium.
Powell (1966: 51) treated Pleuroliria at generic rank and briefly diagnosed it as, “Smaller than Polystira, with
a less prominent peripheral keel, and a protoconch of about four whorls, first very small, last 2½ with numerous
axial riblets.”
Problem of the type species: Uncertainty about the identity of Pleurotoma supramirifica has arisen because the
type material, at first housed in de Gregorio’s private collection, then housed at the Università Palermo, was
subsequently lost (Palmer & Brann 1965: 12–13). We follow Casey (1904: 128, 132), Gardner (1937: 288),
Woodring (1970: 363) and MacNeil & Dockery (1984: 176–177) in considering Pleurotoma supramirifica to be
synonymous with P. cochlearis Conrad, rather than a variety of Conrad’s species as supposed by de Gregorio and
upheld by Dall (1918: 330) and Harris (1937: 30). Pleuroliria cochlearis (Conrad) is known from a variety of
specimens and has been fully redescribed and figured, with a lectotype chosen (ANSP 13420) by MacNeil &
Dockery (1984: 176–177; pl. 35, figs 8, 9; pl. 59, fig. 12). Despite this attention, we give further clarification of the
identity of supramirifica in order to definitively close the case, because MacNeil & Dockery (p. 176) left open
some doubt in stating, “…if P. supramirifica should prove to be the Vicksburg P. cochlearis Conrad.”
The close similarity of de Gregorio’s illustration with specimens of P. cochlearis (e.g. MacNeil & Dockery
1984: pl. 35, fig. 8) is evident in terms of size, proportions, details of spiral and axial ornamentation, shape of the
anal sinus, short rostrum and presence of apertural lirae. This synonymization implies that the supposed Claiborne,
Alabama, type locality is mistaken and the specimen illustrated by de Gregorio was obtained from the Byram
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Formation, Oligocene of Mississippi. Strong support for this supposition was first given by Gardner (1945: 245)
who stated, “There is no example of de Gregorio’s type [i.e., P. supramirifica] in our extensive collections [at the
U.S. Geological Survey/USNM] from the Claiborne group. The suspicion must arise that he had before him an
example of Pleurotoma cochlearis from the Byram marl of Vicksburg, Mississippi, but there is no proof.” Even
firmer support for this interpretation came from Woodring (1970: 363) who briefly commented that, “there is no
reasonable doubt that it is the Oligocene species Pleurotoma cochlearis. The known Eocene species are much
smaller”. Indeed, the lectotype of P. cochlearis has been determined from its preservation and attached matrix to
have been obtained from the Byram Formation of Vicksburg, Mississsippi (MacNeil & Dockery 1984: 177).
Finally, MacNeil (in MacNeil & Dockery 1984: 9) ascertained that de Gregorio likely obtained his material from a
commercial supplier of scientific items and that it contained a number of undoubted Vicksburg species.
FIGURE 2. Pleuroliria cochlearis (Conrad, 1848); lectotype, ANSP 13420, Byram Formation, Late Oligocene, Vicksburg,
Mississippi, USA. Herein designated the neotype of Pleurotoma (Pleuroliria) supramirifica de Gregorio, 1890, which is the
type species of Pleurotoma (Pleuroliria) de Gregorio, 1890.
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MacNeil (in MacNeil & Dockery 1984) undertook lectotypification of Pleurotoma cochlearis to stabilize this
name, and we offer a few comments on his actions. Two figures of Pleurotoma cochlearis Conrad are shown in the
original plate of Conrad (reprinted as appendix 1 in Dockery 1982: 232–233). The larger specimen (on the left hand
side) has a similar form to the rostrum and the same proportions as ANSP 13420 (l. = 42.3 mm, w. = 10.8 mm), the
lectotype selected by MacNeil (pp. 9, 177 of MacNeil & Dockery 1984). The smaller specimen on the right hand
side of the plate is presumably one of the three smaller original specimens of Conrad (ANSP 13421, see Moore
1962), only one of which was designated as a paratype (sic, actually a paralectotype) by MacNeil (p. 177). We have
re-examined these paralectotype specimens; one (l. = 21.4 mm, w. = 6.6 mm) is a specimen of P. cochlearis but the
other two belong to another undescribed species belonging to the Polystira clade. In order to unambiguously
document P. cochlearis w e p r o v i d e a n i m a g e o f t h e l e c t o t y p e h e r e ( F i g . 2 ) .
Neotype designation for Pleurotoma (Pleuroliria) supramirifica
To clarify the identity of this species and prevent the possibility of any further taxonomic instability, we select
the lectotype of Pleurotoma cochlearis Conrad (ANSP 13420) to serve as the neotype of Pleurotoma (Pleuroliria)
supramirifica de Gregorio. This action effectively stabilizes the usage of Pleuroliria which otherwise might be
based on differing interpretations of a species described from a single shell from an uncertain locality that is
interpretable only from de Gregorio’s inadequate illustration. The qualifying conditions for selection of a neotype
(ICZN 1999: Article 75.3) are met in our discussion of “problem of the type species” above, taken together with the
full description of P. cochlearis provided by MacNeil & Dockery (1984: 176–177).
Rank and relationships: Woodring (1928: 145) did no more than imply that Pleuroliria might be an ancestor of
Polystira, but Powell (1966: 51) went further in stating that it was “very similar to the Miocene-Recent Polystira,
and evidently the forerunner of it” and “this genus and the derived Miocene-Recent Caribbean-Panamic Polystira
have their parallel in the Indo-Pacific Lophiotoma”. By 1970, Woodring felt confident in stating (p. 363), “Though
Polystira was named at the generic level, it is proposed to treat it as a subgenus of Pleuroliria. It is interpreted to be
the direct descendent of Pleuroliria and the two groups have similar basic features.” and, interestingly, “too much
reliance was based on [the protoconch features of] type species when Polystira was named—a procedure that has
been justly criticized (MacNeil 1960: 100)”.
Polystira Woodring, 1928: 144–145; Type species by original designation: Pleurotoma albida Perry, 1811.
“Recent, West Indies and Florida”.
Original diagnosis: “Shell relatively large, fusoid. Nucleus stout, cylindrical, consisting of almost two whorls,
the last quarter whorl bearing a few axial riblets. Aperture narrow. Anterior canal long, narrow, unemarginate.
Siphonal fasciole slightly or rather strongly inflated. Between it and the inner lip lies a narrow umbilical groove or
a relatively wide umbilical opening. Anal sinus moderately deep, shaped like a V with a rounded apex, which lies
on the peripheral keel. Interior of outer lip bearing far within aperture fine ridges or fluting. Sculpture consisting of
spiral keels and threads, the peripheral keel strongest, and of strong growth threads.”
Problem of the type species: Some modern authors have doubted the identity of Perry’s “Pleurotoma albida”
and have followed Bartsch (1934: 8) in trusting Perry’s statement (1811: pl. 32, fig. 4: explanation) that it is “from
the South Seas, being frequently found at New Zealand and Lord Howe’s Island” (e.g., Glibert 1960; MacNeil &
Dockery 1984: 176). If this is correct then it is remarkable that no one has been able to offer a plausible identity for
the shell in Perry’s figure amongst Indo-Pacific turrids, especially given our rapidly growing knowledge of the
fauna. Having examined Perry’s illustration and his description we have no doubt that his species corresponds
exactly with Polystira albida as illustrated in many well-known and easily accessible sources of regional (e.g.,
Morris 1973; Abbott 1974; Pointier & Lamy 1998; Diaz Merlano & Hegedus 1994) and worldwide scope (e.g.,
Eisenberg 1981; Abbott & Dance 1982). Woodring (1928, 1970) was quite correct in considering Perry’s locality to
be erroneous and he amended it to the known tropical American range of this species.
Comparisons and relationships: Woodring (1928: 145) stated that, “The genus is the American tropical
representative of the Indo-Pacific Turris (“Bolten”) Roeding…, which also has the interior of the body whorl
fluted. The sinus of Turris is deep and narrow and lies behind the peripheral keel on a flat band”. He considered
that, “Polystira apparently does not extend further back than lower Miocene time. A similar genus, Pleuroliria de
Gregorio…which is much smaller and has a less prominent peripheral keel, is found in the Eocene and Oligocene
deposits of southeastern United States. No specimens of P. supramirifica are available, but according to de
Gregorio’s figures, Pleurotoma cochlearis Conrad, a species from the upper Oligocene Byram marl, is very similar
to it. This species has a nucleus different from that of Polystira, consisting of about four whorls, the first one of
which is very small and the last two and a half of which are sculptured with many axial riblets.”
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By 1970, Woodring (p. 363) was more circumspect concerning the putative differences between these taxa:
“Polystira reaches a larger size than Pleuroliria s.s. and has a less rapidly enlarging protoconch. The protoconch of
Polystira generally has fewer whorls and fewer whorls sculptured with axial riblets. The number of protoconch
whorls and of sculptured protoconch whorls, however, overlap in the two groups and not all of the species of
Polystira are large. Some species of Polystira, including the type species, have a blunt protoconch, but that of
others is acute…Both Pleuroliria s.s and Polystira embrace American species. The age range of Polystira is early
Miocene to the present time and the living species occur in western Atlantic and eastern Pacific waters. They are
characterized by strong, coarse, exaggerated growth threads [sic], at least on early post-protoconch whorls, if not
also on later whorls. This may seem to be a minor feature but it can be traced back through the fossil species of
Polystira to the Oligocene species that doubtless is the type species of Pleuroliria [i.e., P. cochlearis].”
Subdivision of Polystira: According to Olsson (1964: 89), “In the Recent West Indian fauna and in many
Miocene areas, two distinct groups of Polystira can be recognized. First, the typical or carinate species belonging
to the P. albida-barretti stock with strongly emphasized spiral sculpture, keel-like on the periphery; and secondly
the non-carinate species such as P. tellea-haitensis with rounded whorls, no peripheral keel and composite crowded
spirals”. Woodring (1970: 364) pointed out that, “If…so much emphasis is to be placed on the anal sinus, the
western Atlantic species described by Dall as Pleurotoma albida var. tellea (Dall, 1889: 73) is not a typical species
of Polystira. The sinus of mature shells of that species (but not of immature shells) is decidedly asymmetric and
shallow” and that its mature sculpture was “unusual” too.
Pleuroliria (Josephina) Gardner, 1945: 246. Type species by original designation: Pleuroliria tenagos Gardner,
1937: 288, pl. 38, figs 25, 26. Shoal River Formation, Middle Miocene, Florida, USA.
Original diagnosis: “The section is proposed to include those Pleuroliria of medium or relatively large
dimensions with protoconchs of four or more whorls and conchs with spiral cords so prominent that they contour
the whorls”.
This was earlier expanded upon by Woodring (1928: 145) who stated, “A branch of this early Tertiary genus
[i.e., Pleuroliria] is represented by a species from the Middle Miocene Shoal River formation of Florida, described
in manuscript by Gardner [i.e., tenagos]. It has a nucleus resembling that of [P.] cochlearis, though the number of
whorls is reduced to a little less than four, but the shell is larger and the peripheral keel is stronger, so that except
for the nucleus it resembles the still larger species of Polystira”.
Rank and relationships: Woodring (1928: 145) considered that, “If the arrangement here proposed is worth
anything, this species should be placed in a subgenus under Pleuroliria. This same subgenus is represented by
living species in the Panamic and Mazatlanic regions (“Pleurotoma” picta Reeve and “Pleurotoma” albicarinata
Sowerby), but the number of nuclear whorls is again reduced to three or a little less than three. Both these species
are smaller than the West Indian species and they are comparable in size to the Shoal River fossils. So far as can be
discovered all the living West Indian species fall in Polystira, though “Pleurotoma albida var.” tellea Dall has
relatively weak keels.”
Powell (1966: 52) followed Woodring in stressing the value of protoconch morphology and treated Josephina
as a subgenus of Pleuroliria. However he noted that it “occupies an apparent transitional position between the
Eocene-Oligocene Pleuroliria, with its multispiral protoconch and relatively weak spiral keels, and the Miocene-
Recent Polystira, which has a paucispiral protoconch and prominent spiral keels. However [P.] tenagos, by its
protoconch is a Pleuroliria and there seems to be little justification for recognizing an intermediate taxon that relies
for its distinction, only upon stronger spiral sculpture, approaching that of Polystira.”
Later Woodring (1970: 364) re-evaluated the characters used for taxonomic assignation used to discriminate
these taxa. By implicitly considering teleoconch ornament to be more important than protoconch morphology, he
considered Josephina to fall under Pleuroliria (Polystira), noting, “One of the species of Polystira that has an acute
protoconch—Pleuroliria tenagos …was designated as the type of Josephina (Gardner 1945: 246), proposed as a
section of Pleuroliria. That proposal followed my suggestion (Woodring 1928: 145), which overemphasized the
distinction between blunt and acute protoconchs.”
Turris (Oxytropa) Glibert, 1955: 6. Type species by original designation: Pleurotoma oxytropis G. B. Sowerby I,
1834: 135. Recent, tropical East Pacific.
“Plésiogénotype fossile: Turris (Oxytropa) konincki Nyst, sp. 1843” Glibert (1955: 6) (N.B. originally spelt
koninckii). Early-late Oligocene and “Lattorfian”; Belgium, Germany, Netherlands (Glibert 1960).
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Original description (translated from French): “Protoconch of the Gemmula type formed from two smooth
whorls and one or two carinate whorls ornamented with thin axial ribs, prominent, almost straight, widely spaced.
Spire slender, with a conical outline. Whorls carinate towards the anterior, slightly concave behind the carina, and
more or less flat in front of the carina. At least the first three post-embryonic whorls are entirely smooth, the
following whorls are generally ornamented with uniform, closely-spaced spiral cords. Several quite strongly
projecting spiral cords on the base of the last whorl, at least in the juvenile. Sinus with a very deep V-shape, with a
rounded apex, positioned on the projecting carina. Carina smooth and convex, unadorned. Lacking axial
ornamentation after the protoconch. Interior of outer lip smooth. Canal long and slender, well defined at its base,
ornamented by the ridges of twisted spirals which progressively become attenuated from the posterior to the
anterior”.
Original (differential) diagnosis (translated from French): “Separated from Turri s s.s. by its sinus positioned on
the carina; from Lophiotoma by its; shallower, non-rectangular sinus, by the sinus rib (carina) being simply
rounded, by its costate protoconch and by the absence of interior liration within the labrum; from Polystira by the
weakness of its spiral ornamentation, by its smooth labrum, by its strongly excavated base; from Gemmula by the
absence of nodules on the sinus rib (carina); from α-Gemmula by its narrower and much deeper sinus, by the sinus
rib (carina) being narrowly rounded and by the absence of post-embryonic axial ornamentation”.
Later Glibert (1957: 77) noted that (translated from French), “The closest species to our fossil [i.e., Turris
(Oxytropa) koninckii (Nyst, 1843)] is T. oxytropis (Sowerby), type species of the subgenus Oxytropa Glibert, 1955
(p. 5), which can only be distinguished by its slightly wider apical angle (35°), its more prominent carina which is
narrower in the adult, its finer and more compact spiral ornamentation, its base ornamented with larger and much
wider spaced spirals, its shallower whorls, and less voluminous protoconch”.
How should Oxytropa be interpreted? Given the clarity of Glibert’s statement (1957) about what he perceived
as the close relationship between P. oxytropis and P. koninckii we think that his selection of P. oxytropis as type
species was not injudicious but was intended to anchor the name Oxytropa using a living type species—a
commendable aim. Oxytropa (whether as a genus or subgenus) has not received wide usage subsequent to Glibert’s
papers (1955, 1957 and 1960). After 1960 those taxonomists treating Polystira species have either been unaware of
the name Oxytropa or have considered it unnecessary. We are unaware of other authors using this name in
association with Polystira oxytropis or any other members of the Polystira clade (see Tucker 2004: 720). Hickman
(1976: 97) noted that Glibert’s usage of Oxytropa for “European Oligocene species with a pronounced constriction
of the anterior canal and bicarinate whorl profile” was inconsistent with his designation of Pleurotoma oxytropis as
type species which Hickman correctly noted, “fits more readily into Polystira”. We agree with her view that there
is no close relationship between P. oxytropis and P. koninckii or other European fossil species. In brief, in contrast
to Polystira, Pleurotoma koninckii lacks a well-developed principal spiral A throughout its ontogeny, and spiral B
lies at or just above the basal suture (see Gründel 1989: pl. 2; text-figs 6, 7; Moths 2000: pl. 8, fig. 7). It should be
noted that Glibert’s selection of a “plésiogénotype” or a “species related to the genotype (belonging to the same
genus, subgenus and section) [= type species] but occupying a different geologic formation or zoologic province”
(Frizzell 1933: 662) has no official nomenclatural standing (ICZN 1999: Article 72) and does not affect how one
properly interprets the taxonomic name. We are aware of only comparatively few usages of the name Oxytropa by
European authors for the fossil species (and a few others) placed there by Glibert (e.g., Amitrov 1971; Báldi 1963,
1973; Gründel 1989, 1997; Moths 2000). Amitrov (1973) later changed his opinion and placed the non-gemmulate
koninckii within his own very wide concept of Gemmula! Gründel (1989: 119) mistakenly thought the type species
of Oxytropa to be Turris (O.) pseudovolgeri Glibert, 1955 and assigned this species (as a possible synonym of P.
koninckii) and P. koninckii to Oxytropa. Finally, Hickman (1976: 96) saw a similarity between her Pleuroliria
oregonensis (probably a Kuroshioturris, see above) and Oxytropa pseudovolgeri (Glibert), although she
perceptively suggested (p. 98) that “the geographic and temporal discontinuities involved raise the possibility of
parallel development from separate ancestral turrine stocks”.
Powell (1966: 52) considered Oxytropa to be a subjective synonym of Polystira (sensu lato) and recently this
opinion has been followed in a summary classification of the Turridae (Bouchet et al. 2011: 297). Unfortunately
synonymization has led to Oligocene European species that correspond with Glibert’s (1955) intended
circumscription (e.g., Oxytropa koninckii (Nyst, 1845) and O. pseudovolgeri (Glibert, 1955)), but which we
consider to lie outside the Polystira clade, subsequently having been assigned erroneously either to Polystira (e.g.,
Janssen 1978; Müller 1983) or Pleuroliria (Jannsen 1979; listed by Schnetler & Beyer 1987, 1990). The result is
that the European species that have been assigned to Oxytropa lack an available generic-level name. We do not
think that restricting the published concept of Oxytropa to the living type species from the East Pacific will
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markedly affect the nomenclatural stability of the European species as there has been little agreement among
authors on their correct systematic position (e.g. for P. koninckii see Tucker 2004: 530).
The systematics of P. koninckii and related species cannot be dealt with here; we consider this to be one of a
number of groups of fossil and Recent Turridae (s.s.) that require detailed revision. As their study progresses,
erection of further genus-level taxa will be necessitated.
Traditional diagnoses and circumscriptions
In examining original and subsequent diagnoses of the supraspecific taxa comprising the Polystira clade, including
the selected examples above, one is struck by a number of related features that continually recur;
Firstly, the breadth of conchological variation within museum collections that are known to have been
available to these and other systematists had been overlooked entirely. This resulted in very few, and usually only
the largest-shelled, species having been recognized by these workers. This is brought home by Olsson’s recognition
within Polystira (s.s.) of just two groups of species, both comprising large-bodied representatives, and Woodring’s
observation about the “unusual” morphology of Polystira tellea (Dall)—in fact a number of what we now
recognize as (putative) species-level taxa, fossil and living, share similar character states to those he mentioned
(see Todd & Johnson 2013: fig. 1). It was Hickman (1976: 96) who first noted that the available supraspecific
taxonomy (Pleuroliria versus Polystira) seemed inadequate to account for the morphological disparity within the
clade and that, “many species, however, do not fit either branch of the above dichotomy.” However, the apparent
paucity of morphological variation (and the resultant very few species) recognized by authors throughout the
twentieth century remained the current situation until we began studying Polystira in depth in 2000.
Secondly, following on from the above, taxa appear to be based solely upon the type species rather than, in the
absence of any phylogenetic analysis, the entirety of species taxa included within them.
Thirdly, there is a much greater focus on the supposedly correct application of names than there is on the
detailed description of (shell) morphology: the range of characters and character states displayed by these taxa
themselves. By contrast, in a modern phylogenetic context, it is only through detailed description and analysis of
character state distributions that permits the groups (putative clades) to be identified and subsequently named.
Naming is secondary (in both senses) to establishing the reality of clades, a natural classification. This contrast is
precisely the difference between Linnaean classification and systematics, for example as characterized by Ebach &
Williams (2010).
Fourthly, much is made of apparent ancestor-descendent relationships between supraspecific taxa—though no
detailed morphological analysis of the supposed phylogenetic relationships is offered. Apparently, the consensus is
that protoconchs decrease in number of whorls (and numbers of costate whorls) and spiral sculpture (particularly
the carina) increases in strength from Pleuroliria (Eocene-Oligocene) to Josephina (Middle Miocene to Recent—if
recognized) to Polystira (Late (?) Miocene to Recent) in a presumed lineal sequence. This is a good example of the
traditional palaeontological perspective whereby evolutionary patterns are reconstructed by identifying supposed
lines of ancestry (ancestor-descendent sequences) through a literal reading of the rock record, an approach that the
cladistic revolution of the 1970s to 1980s has largely discredited (e.g., Forey 2004). Our reading across a range of
recently published taxonomic papers shows this approach still remains common within the molluscan
palaeontological community.
Homology, character recognition and description: a way ahead
“It is apparent that no satisfactory classification can be derived for this group of turrids [i.e., Polystira, Pleuroliria,
Lophiotoma and others] using shell characters alone, and it is highly possible that anatomical or immunological
studies of living species in the group will sort out forms that would not have been placed together on the basis of
shell characters.” (Hickman 1976: 96).
Considering the continuing flux in generic-level taxonomy of the Turridae (s.s.) based primarily on molecular
sequence analyses (Heralde et al. 2007, 2010; Olivera et al. 2008), the above statement may seem as true now as it
did to Hickman when it was written some thirty-eight years ago. However, it is our contention that the proximate
cause of non-recognition of species diversity and difficulties in delimiting supraspecific taxa is likely not a taxon-
specific problem, that is, limited to the Turridae. Rather it lies in two related and more general problems within the
fields of molluscan taxonomy and systematics.
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The first is the sometimes rudimentary level of morphological description of gastropod shells and other
molluscan taxa with accretionary shells. Hickman hypothesizes that “shell characters can be much more useful in
turrid [here, sensu lato] taxonomy if they are more carefully chosen and their states more rigorously defined”
(Hickman 1976: 22). With the perceived exception of the turrids (s.s.) that she studied, this indeed proved to be the
case in her own work on extinct conoideans. Despite this, to date very little work has been undertaken in this area.
In the present case some conchological descriptions of Polystira species are so generalized that they do not
adequately discriminate species, especially given what we now believe is the ‘real’ species diversity. For example,
it is instructive to contrast the level of conchological detail and precision contained within the descriptions of new
Polystira species provided by Petuch (1987, 1988, 2001, 2002) with those descriptions of fossil and Recent
Turridae (s.s.) presented by e.g., Gardner (1937: Polystira, Pleuroliria, Hemipleurotoma [=“Gemmula”]), MacNeil
& Dockery (1984: Pleuroliria), Tracey (1996: Gemmula), Sysoev (2002) and Kantor et al. (2008: Ioturris) amongst
many others.
No matter how detailed a taxonomic description might be in delimiting the features of a taxon and permitting
its accurate identification, its usefulness for systematics is determined by how well it delimits and describes
homologous parts that can be identified across a wider range of taxa. The lack of clear homology statements
contained in molluscan shell descriptions (Merle & Houart 2004: 162, 167; Papadopoulos et al. 2004: 224; Merle
2005) is a second and much more general problem that deeply afflicts molluscan systematics, though its
importance remains greatly underappreciated. It is in this area—more precisely, the identification of detailed shell
homologues—that there is great opportunity to improve our understanding within the Conoidea. Our current lack
of knowledge can be contrasted with the recent advances made in other gastropod taxa, most notably the Muricidae
(reviewed in Merle 2001, 2005; Merle & Houart 2003, 2004)—though methodological challenges remain. Here,
we can only briefly mention work that is currently underway in identifying shell homologues across the Turridae
(s.s.) and notions of what we will here term “deep shell homology” across the Conoidea.
Homology of spiral ornamentation in Turridae
One of us (JAT) has directly examined a wide range of specimens of species belonging to the following living and
extinct generic-level taxa that we consider to belong to the Turridae (s.s.): Annulaturris Powell, 1966; Cinguliturris
Powell, 1964; Coronia de Gregorio, 1890; Coroniopsis MacNeil in MacNeil & Dockery, 1984; Gemmula
Weinkauff, 1875; Lophiotoma Casey, 1904; Pinguigemmula MacNeil,1960; Pleuroliria de Gregorio, 1890;
Polystira Woodring, 1928; Ptychosyrinx Thiele, 1925; Sinistrella Meyer, 1887; Trypanotoma Cossmann, 1893;
Turridrupa Hedley, 1922; Turris Batsch, 1789; Unedogemmula MacNeil, 1961 [including Lophioturris Powell,
1964]; Veruturris Powell, 1944; and Xenuroturris Iredale, 1929 [including Ioturris Medinskaya & Sysoev, 2001] as
well as other extinct taxa that are currently included in the same family (Powell 1964, 1966). A few genera remain
to be examined. Observations indicate that the six or seven major spiral cords that were previously delimited for
Polystira (see ‘Principal spiral cord terminology’) may be identified as a basic ground plan in the family, lettered
alphabetically from top (adapically) to bottom (abapically), A through G, (see Fig. 2) though some taxa may not
possess them all. Use of these identifying letters does not necessarily imply homology of these spiral cords with
those in taxa where a similar notation system is used, for example the Turritellidae (Marwick 1957) and Neptunea
(Buccinidae) (Nelson 1978). In Turridae, spiral cords are identified on the basis of their position relative to: a) the
suture; b) each other; c) biological landmarks; d) relative strength, and e) ontogenetic appearance. One of us (JAT)
will give a much fuller explanation of the criteria used to identify these homologues elsewhere. Use of such a
system will enable shell description and variation to be made more precise. Critically, through identification of
shell homologues, we can identify conchological ground plans that will allow us to develop a much more rigorous
taxonomy for the family. In turn this will allow us to examine patterns of shell evolution and their relationship to
molecular, morphological and toxinological evolution throughout the family and maybe wider within the
Conoidea.
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FIGURE 3. Morphology of a typical, a) multispiral protoconch (see text) and, b) paucispiral protoconch (see text); c, d)
protoconch-teleoconch boundaries of a) and b) respectively, showing contrasting depth of the sinusigera, the
protoconch–teleoconch boundary. Not to scale.
Biology, ecology, and diversity of Polystira species
Conchology
Polystira and Pleuroliria have very similar shell morphologies, but the following is a preliminary description of
Polystira pending more detailed study of the species comprising Pleuroliria.
Turrid (sensu Bouchet et al. 2011) gastropod; adult shell medium-sized (11 mm) to very large (120 mm),
showing determinate growth. Protoconch paucispiral (globose) or multispiral (bullet-shaped) of 1.6 to 3.3 whorls
(Van Osselaer 1999: “Practical method”), macroscopically smooth initially, with as few as three (paucispiral) to a
whorl or more (multispiral) of flexed backwards, arcuate, axial ribs; protoconch sharply terminating with a shallow
symmetrical to deep asymmetrical sinusigeral notch (Fig. 3), the lower limb being longer than the upper as seen on
the whorl side. Teleoconch fusiform, moderately thick-shelled, with a straight to slightly curved, short to long,
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abapically tapering rostrum and siphonal canal, aperture comprising 40–58% of shell height. Overall whorl profile
usually slightly to moderately convex, sometimes almost flat-sided; usually dominated by moderate to strong spiral
ornament. Principal (or 'primary') spiral ornament (see above) of three major spiral cords (A, B, D)—principal
spiral C usually ontogenetically delayed and weaker—more or less equally spaced on whorl side; spiral B at or
slightly above whorl mid-height and usually forming periphery. Spirals A (usually) and B and D (always) present
immediately at start of teleoconch and producing strongly tricarinate earliest teleoconch whorls, then maintaining
or diminishing in relative strength to adulthood. Suture tracks around principal spiral D, which defines junction of
whorl flank with whorl base. Sutures adpressed. Base of whorl poorly to moderately well delimited, with two
additional principal spiral cords, E and F; spiral G delimits the beginning of a poorly to moderately-well delimited
rostrum uniformly ornamented with very fine to medium spiral cords, often alternating in strength. Spiral B usually
sharp, sometimes becoming flat-topped but never bicarinate; crest coincident with apex of anal sinus or slightly
below. Growth line chord orthocline over entire whorl and between upper and lower suture in spire whorls; growth
line prosocline in upper part of whorl, swinging abaperturally (backwards) to make a shallow to moderately deep
V-shaped anal sinus with vertex coinciding with crest of spiral B, then swinging adaperturally (forwards) slightly to
moderately to spiral D, E or F from where it then remains orthocline or then forms a smaller sinus centred on spiral
G (Fig. 4). Other orders of spiral may be intercalated throughout growth. Frequently spiral A, and less often spiral
B, becomes complex with subsidiary spirals on its flanks. Axial ornament comprises numerous faint to prominent,
even-strength commarginal threads or riblets, most prominent on whorl side and base, not crossing summits of
major spirals; frequently diminishing in relative strength through ontogeny. Adult apertural modifications
(Papadopoulos et al. 2004) usually present and may include: suture rising relative to spiral D, narrowing or
widening anal sinus, prominent, closely spaced and thickened growth pauses, and an internally thickened outer lip.
Inner face of outer lip may have spiral lirations developed at adulthood or earlier in ontogeny or may remain
smooth. The operculum is leaf-shaped and with a terminal nucleus; its attachment area covers nearly the whole
surface, surrounded by a narrow raised border (Dall 1889: 73; Powell 1966: 52).
FIGURE 4. Outer lip and anal sinus morphology in adult shells of six extant Polystira species. Note variability in depth and
shape of anal sinus, and of the shapes of the upper (posterior) and lower (anterior) limbs of the outer lip. Not to scale.
Anatomy
The gross external morphology of Polystira has yet to be described, although images of living animals can be
found on a few websites. Published anatomical work is scattered and focused on aspects of the alimentary system.
Published studies of the entire gut of any species are also lacking though an unpublished Master’s degree thesis
(Leviten 1970) provides a detailed anatomical and histological study of the gut of P. albida (Perry, 1811) (possibly
here including another closely related and undescribed species too; Todd unpublished data). Some of this
information has been abstracted and referred to in wider systematic studies of conoidean foregut anatomy (Taylor
et al. 1993). García-López et al. (2007) describe the histology of the digestive tract (buccal cavity, oesophagus,
stomach and intestine) of Polystira albida and note the specializations in foregut histology compared to other
prosobranchs. As part of a larger study on the comparative anatomy of the foregut of Turridae, Medinskaya (2002)
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describes the foregut of an undescribed species of Polystira from eastern Brazil, identified under the name
Polystira formosissima, and compares it with other turrids. Powell (1964: pl. 247) illustrates a pair of marginal
radula teeth from a specimen identified as P. albida. One of these teeth was apparently re-figured by Powell in
1966 (text-fig. C, fig. 47), a work in which he also figures a marginal tooth identified as being from P. albicarinata
(G. B. Sowerby II, 1870) (text-fig. C, fig. 46) and two teeth from P. pi c t a (Reeve, 1843) (text-fig. C, fig. 48).
Confusingly, McLean in Keen (1971b: fig. 1648) re-figured the marginal tooth of albicarinata as given by Powell
(1966) and associated it with an image of the holotype of P. artia (Berry, 1957), a distinct species then thought to be
synonymous. A photograph of the radula of a specimen identified as P. oxytropis (G. B. Sowerby I, 1834) is given
by McLean (1971a). For proper documentation of radular morphology, we clearly require SEM images for all
species; this will help prevent future misidentifications.
Feeding and toxinology
The sole study of prey preference is by Leviten (1970) who notes that the guts of many specimens of Polystira
“albida” (see above) that he studied contained partially digested remains and setae from unidentified polychaetes.
Though P. oxytropis has been kept in an aquarium over an extended period no observations were made of
individuals feeding (F. Rodriguez pers. comm.) and analyses of stomach contents have yet to be made.
Polystira albida (Perry) was one of the first two species of Turridae (s.s.) from which venom polypeptides
were purified and characterized (Lopez-Vega et al. 2004), using specimens collected from the Bay of Campeche,
on the Gulf coast of Mexico, and whose species identity we can confirm. Unlike conotoxins derived from Conidae
(s.s.), these polypeptides contained a large number of methionine residues—leading to them being dubbed
‘turritoxins’—and indicating the presence of a yet wider range of biochemical strategies in the Conoidea. Extracts
from P. albida were shown by Rojas et al. (2008) to contain toxins with anticholinergic and unusual
antihistaminergic properties that had not been described previously from other conoideans. More recently, venom
of P. albida has yielded a conopeptide, pal9a, which is the first P-conotoxin-like turritoxin characterized from a
member of the family Turridae in the western Atlantic (Aguilar et al. 2009). This toxin shows low sequence
similarity to framework IX-toxins from other Turridae (s.s.), including three species of Lophiotoma, and four
species of Gemmula, Terebridae (Hastula hectica), and various species of Conidae from the Indo-Pacific and the
western Atlantic.
To evaluate the role of trophic specialization in a “turrid” radiation such as Polystira we need both studies from
species within different clades across the taxonomic breadth of the radiation and more detailed studies of selected
species and clades. Future research on Polystira feeding preferences, specialization and fine-scale toxinological
evolution, as initiated by Heralde et al. (2010) for Indo-Pacific “Gemmula”, might usefully target those species
groups whose individuals are known to be both abundant and show high levels of species sympatry.
Reproduction and larval modes
Overall life history parameters remain unknown for any Polystira species. The typical presence of adult
modifications, such as a constricted anal sinus, flared outer lip, or a rising suture on the last whorl, and increasingly
crowded growth pauses (Papadopoulos et al. 2004) indicates that Polystira has determinate growth and presumably
stops or almost stops shell extension upon reaching maturity or some time afterwards when it diverts resources into
reproduction (Stearns 1992). We do not know how long individuals may live, but specimens identified as Polystira
oxytropis obtained as adults or near adults have survived for three years in aquaria before problems in maintaining
water quality apparently led to their death (Felix Rodriguez pers. comm. 2011). Veliger larvae and egg capsules of
P. albida (misidentified as P. b a rre t t i ) have been described and figured by Penchaszadeh (1982). The oval egg
capsules are up to 10 mm long, with a flat base and convex upper side with a central slit-like escape aperture
covered by a thin membrane. In an aquarium they were laid directly onto a sandy substrate and attached via their
mucilaginous base and thin rim to sand grains. Capsules contained a mean number of 73 eggs (mean diameter 438
µm) which develop into free-swimming veliger larvae. We have seen morphologically similar but smaller egg
cases attached to the shells of living Polystira shells, including ones laid onto the shells of P. oxytropis that had
been kept separated from other snail species in an aquarium (F. Rodriguez pers. comm.). This style of egg capsule
appears to be widespread in the ‘turrid’ conoids (see Gustafson et al. 1991: 48–49; table 3).
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Our SEM and light microscopy examination of the shell apices of 55 living morphologically delimited species
from the western Atlantic with preserved larval shells shows that there is remarkably little variability in larval shell
morphology among them. Two visually discrete morphologies predominate, and exceptions are rare. Using a
binocular microscope, graticule and Van Osselaer’s (1999) recommended “practical method” (“method C”) of
counting whorls, these protoconch types can be characterized as follows:
Multispiral (Fig. 3a): protoconch I and II together comprising 2.3 to 3.3 larval whorls; protoconch I small and
acutely pointed, with a diameter of 250–400 µm. The protoconch has a bullet shaped apex and more or less evenly
increases in diameter throughout its length. It is smooth with a macrosculpture of at least one whorl of arcuate axial ribs
(= “brephic axials” of turrid literature) and has a deep sinusigeral notch at its junction with the teleoconch (Fig. 3c).
FIGURE 5. Diameter of protoconch I plotted against total number of protoconch whorls for 55 living morphospecies of
Western Atlantic Polystira showing paucispiral (green circles, + indicates multiple species with same measurement) and
multispiral clusters (red triangles, x indicates multiple species with same measurement).
Paucispiral (Fig. 3b): protoconch I and II together comprising 1.6 to 2.4 larval whorls; protoconch I large and
globose, with a diameter of 300–530 µm. The first whorl of the protoconch (pc I) is large, hemispherical to globular
and the protoconch usually does not increase in diameter after this. There is usually no more than one-half whorl of
typically three to six arcuate axial riblets, and the sinusigeral notch is typically shallow and arcuate (Fig. 3d).
Larval whorl number was plotted against protoconch I diameter for the same 55 living morphospecies. Two
clusters were produced, corresponding exactly to the visually separated groups of multispiral and paucispiral
morphologies (Fig. 5). A multispiral protoconch almost certainly reflects a planktotrophic veliger larva and a
paucispiral protoconch indicates a non-planktotrophic larva, but live examples of the latter have yet to be studied.
Extinct species assigned to Pleuroliria, such as P. cochlearis (Conrad) (see MacNeil & Dockery 1984: pl. 59,
fig. 12) and P. subsimilis (Casey, 1904) (Todd pers. obs.) may have multispiral protoconchs of up to 4½ (4¾ using
Van Osselaer’s “method C”) whorls of a conical shape, with a tiny protoconch I and then a sequence of weakly
inflated whorls with the periphery at their base, of which at least two whorls have arcuate axial ornament. In overall
shape these appear to resemble protoconchs of European Eocene “Gemmula” species (e.g., Tracey 1996: pl. 6, figs
59–67 [as Gemmula]) more closely than those of extant Polystira, but detailed studies have yet to be undertaken.
PoDWA database
We have made a Polystira Data: Western Atlantic occurrence database (PoDWA v.1) that currently (Dec. 2012)
comprises 1559 species/occurrence records of well localized specimens identifiable to species level and deposited
(or planned to be deposited) in public museums. We have directly examined and re-identified all the specimens that
we have entered; other metadata have been obtained from specimen labels, museum databases and our own
collection records (for specimens housed in NHM, London). This data compilation is referred to in the sections
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below dealing with: life habits, geographic range, bathymetric ranges and substrates, frequent sympatry, and under
macroecological and diversification patterns. Data can be provided upon request by JAT.
Life habits
Despite species of Polystira frequently being locally abundant throughout the wide geographic range of the genus
today in both the western Atlantic (e.g., Penchaszadeh 1982; Buitrago et al. 2006; Torruco et al. 2007; pers. obs.) and
eastern Pacific (Olsson 1971: 42–43; pers. obs.), published observations on the ecology and habits of any species of
Polystira appear to be almost non-existent. Some observations have been made of individuals of P. oxytropis from the
Gulf of Panama kept alive in a continuous circulation seawater aquarium at Naos Marine Lab, Smithsonian Tropical
Research Institute, Panama (F. Rodriguez pers. comm.). Specimens were seen to be more active and crawling on the
sediment surface by night; in daylight hours they remained buried in the muddy sediment from which they were
dredged. The presence of live-collected specimens in the PodWa database does allow some limited generalizations
about depth and substrate preferences (see “Bathymetric ranges and substrates” below).
Geographic range
Polystira today has a dominantly tropical and subtropical distribution in the Americas ranging from off North
Carolina, USA, in the North Atlantic, throughout the Gulf of Mexico and Caribbean to Uruguay in the South Atlantic
and in the East Pacific from the northern Sea of Cortez (Gulf of California), Baja California (Mexico) southwards to
Ecuador. During warmer climatic intervals in the Neogene Polystira sensu lato (including Pleuroliria) has had an
even wider latitudinal range, notably during the Middle Miocene Climate Optimum, a global warming event at about
15 Ma. This clade occurs in Argentina at 40°S in the Gran Bajo del Gualicho Formation of latest Early Miocene/
earliest Middle Miocene age (Reichler 2010), and at 39°N in the Kirkwood Formation of late Early Miocene age in
New Jersey, USA (Pilsbry & Harbison 1933; Ward 1998). Eastern Pacific fossil occurrences are known as far north as
35°N, the Middle Miocene Upper Olcese Sand of California (Addicott 1970).
Rare extralimital records, such as Pleuroliria oregonensis Hickman, 1976 from the Late Eocene Keasey
Formation of Oregon, USA (NW Pacific at 45°N) have been wrongly assigned to this clade; this species probably
belongs to Kuroshioturris Shuto, 1961. Indeed, Hickman (1976: 98) noted how similar her new species was to
Shuto’s (1961) figures of Polystira kurodae (sic) (Makiyama). This Japanese Early Pliocene species originally
described as Turris kurodai Makiyama, 1927, was transferred to Polystira by Shuto (1961: 77). Again we consider
this species to probably belong to Kuroshioturris; Shuto (1961: 78) already suggested a probable close relationship
to this taxon. Donn Tippett (unpublished data) came to the same conclusion about the affinities of these two species
many years ago. As a final example, Glibert (1960: 4) assigned Pleurotoma septemlirata Harris, 1897 from the
Middle Miocene of Victoria, Australia to Pleuroliria (Polystira). We have examined topotypic specimens in the
NHM (London) collections and can refute this placement; a strongly bicarinate spiral B may suggest that this
species is more closely related to Lophiotoma, rather than Turris where Powell (1964, 1966) among others (see
Tucker 2004: 898) has placed it.
Bathymetric ranges and substrates
Species of Polystira are almost always found on soft bottoms (PoDWA data, pers. obs.) and the genus has a wide
bathymetric range, living in as little as 1 m of water in oolitic sand associated with Thalassia sea grass beds in the
Bahamas (P. bayeri Petuch, 2001) and occasionally intertidally on muddy sand and sand in the Gulf of Panama and
the Pacific coast of Colombia (P. “oxytropis”). The deepest well-documented record of living animals in our
database is 476–658 m (260–360 fathoms) for P. macra Bartsch from off Puerto Rico (Bartsch 1934: 11), but it is
possible that living Polystira elsewhere occur as deep as 1030 m, though records deeper than 450 m are rare. Some
common, widespread, well sampled species such as P. albida have particularly wide bathymetric ranges. This
eurytopic species has been collected alive in water depths between 9 m and 192 m on a range of soft bottoms from
calcareous sand and mud, coarse shelly sand to fine sand, muddy sand and occasionally fine mud (PoDWA data), as
well as rubble and other sediments associated with drowned palaeo-reefs (Buitrago et al. 2006).
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Wherever sampling of the Recent fauna has been sufficient such that depth transects can be reconstructed (e.g.,
Florida Keys), Polystira species show pronounced bathymetric zonation; preliminary examination suggests that
deeper water species appear to replace shallower taxa at around 100 m depth. Interestingly, a similar pattern, yet to
be studied in detail, is present in the Mio-Pliocene sedimentary basins of the south-western Caribbean, including
the Caribbean coast of Panama and Costa Rica. For example, superb geological exposures arrayed across the Bocas
del Toro Basin, Panama display partly contemporaneous 3–4 Ma Pliocene sediments (Cayo Agua, Shark Hole
Point and Escudo de Veraguas formations) deposited across the shelf in palaeodepths of ca 50–175 m (Coates
1999; Collins 1999). Their diverse and bathymetrically segregated Polystira faunas point to the antiquity of this
ecological pattern.
Frequent sympatry
Polystira species frequently show unusually high levels of sympatry, and thus high alpha diversity. In areas of the
western Atlantic where Polystira is abundant today, including the Straits of Florida, off Isla Margarita and the Paria
Peninsula (Venezuela) and off Santa Marta (Colombia) a single dredge haul may contain as many as five
morphospecies (PoDWA database). In the Neogene of the Caribbean, the most finely sampled geological horizons
(or beds) sampled by the Panama Paleontology Project (Coates & Collins 1999) often contain similar numbers of
species. As a result many museum lots, both fossil and Recent, obtained from a single site contain a mixture of
species, many obviously differing in maximum size, sculpture and protoconch morphologies, and coloration
patterns.
Morphological discrimination of species
Shell morphology in the genus Polystira appears to be highly conserved and at first glance shells from across the
clade seem to possess limited morphological variability (Pls 1, 2). However, shells are moderately morphologically
complex, which allows Polystira clades and their component morphospecies to be recognizable on the basis of
PLATE 1. Type and representative specimens of smaller living Polystira species
1a, b; Polystira artia (Berry, 1957); paratype, USNM 612212 (smaller specimen); adult; off Angel de la Guarda Island, Baja
California, Gulf of California, Mexico, 67 fathoms (123 m).
2a, b; P. bayeri Petuch, 2001; non-type, UF 164636 (one of 12 specimens); adult; near Cat Cay (type locality), Bimini Islands,
Bahamas, 1–6 fathoms (2–11 m).
3a, b; P. florencae Bartsch, 1934; holotype, USNM 429760; adult; off north coast of Puerto Rico, between 18°30´20"N;
66°22´05"W and 18°30'30"N; 66°23'05"W, 33–40 fathoms (60–73 m).
4; P. formosissima (E. A. Smith, 1915); syntype, NHMUK 1915.4.18.309; broken aperture and apex, off Rio de Janeiro, Brazil,
22°56'S; 41°34'W, 40 fathoms (73 m).
5; P. g r un e ri (Philippi, 1848), lectotype of P. phillipsi Usticke, 1969, AMNH 195454; adult; Reuters Bay, Water Island, St.
Thomas, U.S. Virgin Islands.
6a, b; P. lindae Petuch, 1987; holotype, USNM 859895; adult; off Punto Fijo, Paraguana Peninsula, Gulf of Venezuela,
Venezuela, 12°N; 70°W, 35 m.
7a, b; P. macra Bartsch, 1934; holotype, USNM 430395; adult; off northwest coast of Puerto Rico (north of St. John, U.S.
Virgin Islands), between 18°40'30"N; 64°50'W and 18°45'40"N; 64°48'W, 190–300 fathoms (348–549 m).
8; P. starretti Petuch, 2002; non-type, small and narrow adult; UMML 30.7450; east of Florida Keys, USA, 25°33'N; 80°04'W,
96 m.
9a, b; P. “oxytropis” (G. B. Sowerby I, 1834); syntype of P. albicarinata (G. B. Sowerby II, 1870) NHMUK 1874.12.11.295
(larger specimen); adult; Manzanilla, W. Mexico (Pacific Ocean).
10a, b; P. parthenia (Berry, 1957); paratype, USNM 612211; adult with marked apertural modifications; off Islas Tortugas,
Gulf of Nicoya, Costa Rica (Pacific Ocean), 10 fathoms (18 m).
11a, b; P. sunderlandi Petuch, 1987; holotype, USNM 859896; adult; 50 km south of Apalachicola, Florida, USA (Gulf of
Mexico), 29N; 85°W, 150 m.
12a, b; P. vibex (Dall, 1889); syntype, USNM 87385 (largest of three specimens), adult; off Havana, Cuba, 28°N; 82.5°W,
80–127 fathoms (146–232 m).
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suites of conchological characters including size, proportions, and details of ornamentation. When shells are
studied in detail, particularly in the light of sculptural homology (Fig. 2), then a wealth of characters and states is
seen to be potentially taxonomically informative. Identification of homologous parts of the shell facilitates more
precise taxonomic description. As a result comparison among individuals and taxa can be rooted in homology and
relationship rather than overall phenetic resemblance. Taxonomically informative character complexes have been
previously listed by Todd & Johnson (2013) and are given again here; 1) protoconch morphology, sculpture and
size (Fig. 3), 2) teleoconch size, 3) spiral angle and shape, 4) shape and proportions of spire whorls and shoulder
morphology, 5) shape of base of whorl and basal spiral ornamentation patterns, 6) length, shape and ornamentation
of rostrum, 7) strength, spacing and micromorphology of axial ornamentation and its ontogenetic development, 8)
size, strength, profile, spacing, complexity and micromorphology of primary and subsidiary spiral ornamentation
and their ontogenetic development (Fig. 2), 9) shape of outer lip and anal sinus (Fig. 4) and its changes through
ontogeny, 10) growth patterns and presence and nature of adult modifications, 11) presence and nature of sexual
dimorphism, and 12) shell colour patterns.
Establishing current species diversity
Taxonomic revisions of Polystira clades and species have yet to be published and previous recognition of Polystira
species diversity has been limited. For example, Abbott (1974) listed a total of just nine species for its entire Pacific
and Atlantic range and Rosenberg’s web-based systematic inventory of Recent western Atlantic molluscs (2009
release) recognizes thirteen species. The Census of Marine Life listed a total of eight species distributed within the
Caribbean (Diaz & Miloslavich in Miloslavich et al. 2010). Most recently, the WoRMS data compilation
(Appletans et al. 2012) lists 15 described Recent species occurring in the western Atlantic (of which we recognize
14 as distinct) and five in the tropical eastern Pacific, all of which we recognize.
Our application of sequence-based approaches for delimitation of Polystira species (e.g., Pons et al. 2006;
Puillandre et al. 2011b) has been limited by the availability of suitable fresh or ethanol-preserved tissue for
analysis. While there are large collections of Polystira in major museums in the USA and Europe, as well as in a
number of smaller collections, the majority of these consist of dry shell material alone; of those wet collections,
most contain specimens that were first preserved in formalin, with typically only more recently collected
specimens fixed in ethanol. Despite these constraints, we have attempted to assess the correspondence between
genetically distinct lineages of Polystira and our newly assessed morphospecies. First we sorted ethanol-preserved
specimens in collections of western Atlantic (our primary focus) and eastern Pacific Polystira into morphospecies
delimited with reference to the character complexes listed above. We then sampled DNA from selected specimens
PLATE 2. Type and representative specimens of larger living Polystira species.
1a, b; Polystira antillarum (Crosse, 1865 non d’Orbigny, 1848); non-type, MNHN; adult; off Port Louis, Guadeloupe, 130 m.
2a, b; P. jelskii (Crosse, 1865); holotype of P. hilli Petuch, 1988, USNM 859949; small adult; St James, Barbados, 175–225 m.
3a, b; P. starretti Petuch, 2002; non-type, HMNS 10143; large, broad adult; SSE of Key West, Florida Keys, USA, 114 fathoms
(209 m).
4a, b; P. coltrorum Petuch, 1993; non-type, MZUSP 32.708 (one of seven); medium-sized adult, off Alcobaça, Espirito Santo
Province, Brazil.
5a, b; P. p i ct a (Reeve, 1843); syntype (largest of three), NHMUK MOEA 20120043; adult with repaired outer lip; “Bay of
Panama”, Panama, (Pacific Ocean), 13–20 fathoms (24–37 m).
6; P. tellea (Dall, 1889); non-type, RMNH 81123; subadult with short rostrum; off Guyana, 07°44'N; 57°03'W, 120–200m.
7a, b; P. tellea (Dall, 1889); syntype, USNM 93912; fairly narrow, very large adult with very long rostrum and prominent
growth pauses; between the delta of the Mississippi and Cedar Keys, Florida, Gulf of Mexico, USA; 28.6000°N;
85.5833°W, 111 fathoms (203 m).
8; P. nobilis (Hinds, 1843); non-type, NHMUK MOEA 20120042, juvenile; off Islas de las Perlas, Gulf of Panama, Panama,
08°26'24"N; 079°09'14"W, 66–68 m.
9a, b; P. nobilis (Hinds, 1843); non-type, LACM 152628; very large and broad adult with thickened outer lip and prominent
growth pauses; Puertecitos, Baja California, Gulf of California, Mexico.
10a, b; P. albida (Perry, 1811); non-type, UMML 30.10633; medium-sized adult with thin outer lip; off Martinique, 14°54'N;
61°04'W, 47m.
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comprising all of these morphospecies. These molecular samples were also supplemented with freshly collected
material obtained from our own field work in the Florida Keys and Pacific coast of Panama (see Puillandre et al.
2011a, for details) and relevant turrid (s.s.) outgroups from the Neotropics and Indo-Pacific. Again, we first sorted
the specimens into morphospecies before undertaking DNA sampling. This dataset included 88 ingroup specimens,
including what we identified as representatives of 10 described species of Polystira (P. albida, P. coltrorum, P.
nobilis, P. tellea, P. picta, P. vibex, P. parthenia, P. oxytropis, P. starretti and P. artia). Our taxonomic reassessment
of specimens in this collection based on our fine scale examination of shell morphology and comparison with type
material enabled us to delimit a total of at least 22 distinct morphospecies, including 16 from the western Atlantic
(11 undescribed) and 6 from the eastern Pacific (one undescribed), increasing the putative diversity of this sample
set two-fold.
To explore genetic divergence and phylogenetic relationships among these morphospecies, we amplified and
sequenced fragments of three fast evolving mitochondrial genes: 1) cytochrome oxidase subunit I (COI): aligned
fragment length 658bp; 2) 12S ribosomal RNA (rrnS) (and neighbouring tRNA valine): aligned fragment length
690bp; and 3)16S ribosomal RNA (rrnL): aligned fragment length 528bp, using protocols outlined in Puillandre et
al. (2011a). A Bayesian analysis was undertaken on the combined 3-gene dataset, with independent (“unlinked”)
models of molecular evolution applied to each gene region. The tree was rooted to the outgroup taxa, comprising
five genera and eight species of Turridae (s.s.). The resulting 50% consensus phylogram is shown in Fig. 6, with
branch support illustrated for major clades where posterior probabilities were ≥ 90%. While a detailed exploration
of the phylogenetic relationships of living Polystira will be provided elsewhere, our molecular phylogeny, in
general, showed a poor degree of correspondence between molecular clades and traditional species identifications
as given on museum labels (Rawlings & Todd, unpubl. data). As an example, specimens labelled as P. tellea—a
large and generally “well known” species of Polystira - were sprinkled within well supported clades of P. R-COL-
10, P. R-VEN-4, and P. starretti, in addition to the P. tellea clade. Likewise, our molecular analyses provided strong
evidence of seven deeply divergent clades comprising either a single or multiple undescribed morphospecies (10 in
total); 1) P. R-GUY-9 + P. R-GAN-3 + P. R-GOH-4 (three morphospecies); 2) P. R-GAN-2; 3), P. R-GAN-10 (=P.
R-MIS-1), 4) P. R-PAN-10, 5) P. R-VEN-4, 6) P. R-COL-10, and 7) P. R-GUY-1 (= P. R-VEN-10) + P. R-GAN-26
(two morphospecies) within our sample. In contrast, congruence was much stronger between genetically defined
lineages and our morphospecies assessments. Of the 22 morphospecies assessed prior to our molecular analyses, 17
(77%) of these corresponded to lineages separated by large genetic distances (and typically high support indices) in
our Bayesian phylogeny; these lineages were also identified using the Automatic Barcode Gap Discovery method
of Puillandre et al. (2011b) using pairwise genetic distances of COI. Morphospecies may also comprise
geographically and genetically distinct lineages with smaller levels of divergence (e.g., P. R-GUY-3, P. aff.
parthenia, and the three morphospecies comprising the P. R-GUY-9 + P. R-GAN-3 + P. R-GOH-4 clade)
suggesting that these too should be considered distinct species. Continuing study on the eastern Pacific species is
likely to show that what we have labelled as P. oxytropis in Fig. 6 represents a clade comprising distinct
morphospecies (discussed below). One morphospecies (P. R-GAN-26) was not supported by genetic differentiation
based on the mtDNA gene regions sampled and warrants further study. Specimen and DNA sequence details for the
88 ingroup and eight outgroup individuals depicted in the phylogram (Fig. 6) are given in the Appendix.
FIGURE 6. Molecular-based phylogenetic relationships among alcohol-preserved Polystira sampled from museum collections
(ANSP, FMNH, LACM, MHNMC, MZUSP, NHMUK, RMNH, UF, UMML and USNM). The phylogram illustrated represents
the 50% majority-rule consensus tree from a Bayesian analysis of a combined gene dataset (COI, rrnS-trnV, and rrnL gene
fragments), based on a sampling of 6002 trees (4,000,000 generations, sample frequency = 100, burnin = 0.25, heat = 0.2).
Support values are provided for clades where posterior probabilities are ≥90%. Terminal labels for the 22 ingroup species
recognized correspond to the state (in the US) or country adjacent to where each specimen was collected (see below for
abbreviations). Species names indicate the correspondence of shells of analyzed specimens to currently described
(conchological) species; in many cases these taxon assignments differ from those under which they are catalogued. Codes in
square brackets after P. sp., e.g. R-PAN-10, are taxon labels (sensu Schindel & Miller 2010) here used to indicate undescribed
Atlantic species only. Geographic abbreviations are as follows: US states: FL: Florida; LOUIS: Louisiana; TX: Texas;
Countries: BAH: Bahamas; BRAZ: Brazil; COL-C: Colombia-Caribbean coast; COL-P: Colombia-Pacific coast; CRICA-P:
Costa Rica-Pacific; DOM-R: Dominican Republic; ECUAD: Ecuador; GUY: Guyana; MART: Martinique; MEX-C: Mexico-
Caribbean; MEX-P: Mexico-Pacific; NIC-C: Nicaragua-Caribbean; PAN-C: Panama-Caribbean; PAN-P: Panama-Pacific;
SUR: Suriname; TRIN: Trinidad; TOB: Tobago; VENEZ: Venezuela. See Appendix for specimen details.
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To date, we have delimited 122 living morphospecies of Polystira using shell morphology (Dec. 2012), though
our work has so far largely focused on the western Atlantic species. The strong correspondence between
genetically distinct lineages and our morphospecies assessments, as demonstrated through the limited subset of
Polystira analyzed above, provides supportive evidence that fine scale shell differences are taxonomically
meaningful within this genus. Likewise, these results also support our hypothesis that numerous undescribed
species of Polystira remain in museum collections, either classified erroneously based on superficial similarities to
other named taxa, hidden away in mixed lots of specimens, or simply catalogued as “Polystira sp.”.
DNA sequence data coupled with conchological studies suggests that there may be considerable diversity
currently concealed under the ‘umbrella species’ P. oxytropis. Two of its component species have been described
by Berry (1957) as Pleuroliria artia and Pl. parthenia (figured in Keen 1958: figs 909 and 912, respectively; also
Hertz 1984) but these were later synonymised by McLean (in Keen 1971) and have generally been disregarded.
Both shell morphology (Pl. 1, figs 1, 9, 10) and sequence data (Fig. 6) confirm that these three named species are
specifically distinct from each other and the phylogram indicates that they lie in deeply divergent clades.
Why has species diversity in Polystira taken so long to be recognized?
On one hand it may seem remarkable that the huge morphological diversity of Polystira species held in major
museums has until recently gone completely unrecognized. However Polystira is far from unique in this regard.
Failure to recognize the magnitude of taxonomic diversity of fossil and Recent species represented in molluscan
research collections, including those of major natural history museums, is both very widespread and extremely
deep-rooted. Here we should explain that we are talking about species recognition—whether using formal
nomenclature or ‘taxon labels’ (Schindel & Miller 2010; Todd & Johnson 2013; see Fig. 6 herein)—not species
description; the latter continues at an increasing rate (Bouchet 2006). Understandably, lack of recognition has been
worst in the case of morphologically tightly structured species comprising highly species-rich clades such as
Polystira or the freshwater cerithioid Lavigeria (Michel et al. 2004). The perceived difficulties taxonomists may
have in recognizing or delimiting species or higher taxa using conchological features contrasts with the increasing
power of molecular techniques in rapidly uncovering those very taxa. Over the past 20 years, molecular sequence
data have revealed that much species diversity across the living world, including marine metazoans, is structured in
morphologically similar cryptic, semi-cryptic or sibling species (e.g., general reviews by Knowlton 1993, 2000; for
gastropods: Allmon & Smith 2011). And these are mostly narrowly distinguishable using traditional morphological
characters. Over a huge diversity of organisms across the globe, the generality of this pattern is becoming
increasingly obvious (Bickford et al. 2007) and it appears that the majority of taxonomists and systematists accept
the species-status of so-called cryptic species. Rapid delimitation of supposed cryptic taxa generally has not been
followed by their subsequent formal description (for a notable exception in conoideans see Puillandre et al. 2010),
a scenario that has contributed to an increasing decoupling of morphological systematics and molecular
phylogenetics. Perhaps as a consequence, for species discovery, morphology is increasingly seen as superseded if
not untrustworthy.
Taxonomic list of extant species
We have made detailed studies of holotype, syntype and paratype specimens for 18 of the 23 described living
species (19 accepted herein–below) excepting; 1) P. albida Perry (type lost—Petit 2003; images of syntypes of
synonymous P. v irg o Lamarck, 1816 examined); 2) P. antillarum (Crosse) and 3) P. jelskii (Crosse) (types lost); 4)
P. s t a r re t t i Petuch (type series unavailable for study when FLMNH collection visited); and 5) P. coltrorum Petuch
(holotype and paratype deposited at Museu Oceanográfico Professor Eliézer de Carvalho Rios, Rio Grande, Brazil:
unexamined). For all of these, we have examined topotypes or other material collected geographically close to the
type locality. This has allowed us to reassess the taxonomic diversity of the valid named living species, a minority
of the living species diversity. The most current detailed list of the western Atlantic species is that of Rosenberg
(2009) which provides a very useful bibliography and summaries of bathymetric and distributional ranges. Full
details of our synonymies will be presented with future taxonomic revisions; they differ from Rosenberg’s and thus
his summarized occurrence data will require revision.
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Western Atlantic species: 14 named and accepted species
P. albida (Perry, 1811) [= Pleurotoma virgo Lamarck, 1816; = P. barretti (Guppy, 1866) sensu Petuch 1981,
Penchaszadeh 1982 and others; unpublished data]
P. antillarum (Crosse, 1865 non d’Orbigny, 1848); [date for d’Orbigny’s authorship provided by G. Rosenberg,
pers. comm.]
P. bayeri Petuch, 2001
P. coltrorum Petuch, 1993
P. florencae Bartsch, 1934
P. formosissima (E. A. Smith, 1915)
P. gruneri (Philippi, 1848) [= P. phillipsi Usticke, 1969, see Boyko & Cordeiro 2001: 103; considered
synonymous herein based on unpublished data]
P. jelskii (Crosse, 1865) [= P. h i ll i Petuch, 1988; considered synonymous herein based on unpublished data]
P. lindae Petuch, 1987
P. macra Bartsch, 1934
P. starretti Petuch, 2002
P. sunderlandi Petuch, 1987
P. tellea (Dall, 1889)
P. v ib e x (Dall, 1889)
To date, an additional approximately 98 living species so far undescribed have been recognized by us based on
shell features, and molecular data when available (11 present in Fig. 6); work on their description and systematic
relationships is underway.
Eastern Pacific species: 5 named and accepted species
P. artia (Berry, 1957) [species status confirmed herein]
P. nobilis (Hinds, 1843)
P. oxytropis (G. B. Sowerby I, 1834) [this ‘umbrella taxon’ comprises multiple species; currently includes P.
albicarinata (G. B. Sowerby II, 1870)]
P. parthenia (Berry, 1957) [species status confirmed herein]
P. p ic t a (Reeve, 1843)
We have studied the eastern Pacific species less intensively but we have seen at least five more undescribed
species (one is present in Fig. 6). Examination of putative type material of the described species suggests that some
taxonomic revision may be required.
Pitfalls of poor taxonomy
Historically, within the Polystira clade, species groupings and inferred relationships have been based on overall
shell similarities, focusing on; 1) number and sculpture of the protoconch whorls; 2) relative strength of the major
spiral cord or carina (spiral B), and whether or not it forms the periphery; 3) presence of axial sculpture; 4) depth of
the sinus; and 5) length of the rostrum (siphonal canal); for example, see treatments in Olsson (1964) and Woodring
(1970). Excepting Olsson’s 1964 faunal monograph, almost all works have treated just one or a few species, and
therefore no reliable overview of diversity has been available. Consequently, when describing new species,
typically workers have made relatively few comparisons with other taxa; where comparisons have been made these
have often been between morphologically quite different and rather distantly related species.
In addition, it has not helped that some species descriptions of common taxa were not accompanied by
adequate illustrations when first introduced. The best example of this relates to the taxa now known as P. tellea
(Dall) and P. v ib e x (Dall), the first of which was only very briefly described (Dall 1889: 73–74) and neither of
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FIGURE 7. Polystira vibex (Dall, 1889); syntype, USNM 87385 (largest of three specimens); off Havana, Cuba, 28°N;
82.5°W, 80–127 fathoms (146–232 m), showing characteristic stripes of brown periostracum present within and between
principal spiral cords.
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which were illustrated. Images of correctly identified specimens of Polystira tellea are widely available (e.g.,
Morris 1973: pl. 68, fig. 13; Abbott 1974; Abbott & Dance 1982: 239; Kaicher 1984: card 3893). Despite this,
some specimens illustrated under this name, even very recently, are incorrectly identified (e.g., Pointier & Lamy
1998: 160; Tunnell et al. 2010: 251; Daccarett & Bossio 2011: fig. 889). A far worse state of affairs exists for P.
vibex: we are aware of just a single correctly identified image of this species ever having been published in its 122
year history (Williams 2005: left hand specimen only, a syntype mistakenly identified as the holotype; see below).
Unfortunately this image is of poor quality.
Careless taxonomic treatments and the lack of an accessible data source for type illustrations means that there
is little appreciation of the morphological and species diversity present within Polystira. Consequently, popular and
semi-popular treatments, such as shell identification manuals and guide books (which is all most workers have had
to go on), have frequently misidentified even the commoner species. Many of the species identifications offered
over the Web by shell dealers of their imaged specimens are similarly erroneous. In turn, this situation has led to
misidentified material entering museum collections from both amateur and professional sources. Increasingly, the
metadata associated with these specimens is being web-served as aggregated data showing supposed geographical
(Global Biodiversity Information Facility (GBIF): www.gbif.org/) and geological distributions (Paleobiology
Database: paleodb.org/). Todd & Johnson (2013: 896) give an example of the misinformation that may result from
uncritical data harvesting.
A history of misidentification: P. v i b ex and P. starretti
As an example of such misidentification, the specimens figured as Polystira vibex (Dall) by Morris in his
‘Peterson’s Field Guide’ (1973: pl. 68, fig. 8) and by Abbott (1974: fig. 2940) in his very widely used compendium
‘American Seashells 2
nd
Edition’ are very probably specimens of a species not described until 2002, P. s t a rre t t i
Petuch—as discussed further below. The supposed syntype of P. vi b ex , USNM 87386 figured by Kaicher (1984:
card 3950), is misidentified and is a juvenile specimen of P. te l l e a (Dall). In fact, Dall’s syntype lots of P. v i be x
have registration numbers USNM 87835 and 87837. Kaicher’s error has gone unnoticed (see Rosenberg & Petit
2003), so we provide images of a syntype (USNM 87835) here (Fig. 7). As a result of these errors, the numerous
museum specimens labelled as P. vi b e x are almost all misidentified, and the true P. v ib ex is a much less frequently
collected species. Specimens bearing this name in other publications (e.g., Takeda & Okutani 1983 as “vipex” (sic);
Daccarett & Bossio 2011: fig. 890), on many web resources, and shell dealers’ sites belong instead to a variety of
very different Polystira species.
In a paper describing an apparently novel gastropod fauna living in 300–400 m water depth off the Bimini
Islands, Bahamas, Petuch (2002) described five, dead, worn and broken shells as representing a new species of
Polystira, P. starretti, which he considered to be a “Bahamian deep water endemic” (Petuch 2002: appendix 1).
Petuch likely did not examine major museum collections in the USA to establish whether or not this species had
been previously collected and whether or not it was truly endemic to the Bimini Shelf. This is unfortunate because,
by the year of its description, 2002, what we interpret to be P. s t ar r et t i was represented by over 1,680 specimens,
hundreds of which are undamaged whole specimens, including live-collected material, in numerous US museum
collections including the AMNH, ANSP, HMNS, UF, UMML and USNM, as well as NHM (London) and NCB
Naturalis (Leiden), albeit under a range of misidentified species names (PoDWA database). Indeed, not only does
this species appear to be the most abundant Polystira species occurring in shelf waters off Florida, Florida Keys
(Todd & Rawlings, pers. obs.) and the Bahamas (live animals recorded from 37–229 m; dead shells to 457 m), but
overall it is the most numerous species of western Atlantic Polystira in major museum collections. We provide
figures of whole, unworn and live-collected specimens (Pl. 1, fig. 8; Pl. 2, fig. 3a, b).
We have provided two plates of specimens, mostly from type series, of each of the 19 described living species
listed above (Plates 1, 2). We hope this will allow at least some Recent species to be accurately identified and allow
rectification of the more obvious misidentifications. It must be stressed, however, that species-level conchological
differences in this genus are frequently subtle and are likely to be overlooked prior to the taxonomic re-descriptions
of species and the clades they comprise. We urge users not to shoehorn their specimens into the small number of
named taxa we have illustrated. One of us (JAT) is pleased to provide identifications of material submitted to him
for examination.
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Macroecological and diversification patterns
Species’ range size distributions
Geographic ranges of individual living species vary from those so far found at only one site, which might be
interpreted as either reflecting poor sampling or point endemism, through to Polystira starretti and P. tellea with
perhaps the widest distributions. Polystira lindae Petuch is an example of a species that we believe to be known
from just two specimens from a single site off the Paraguana Peninsula, Venezuela (Petuch 1987), though very
similar, but probably distinct, species occur elsewhere in the Caribbean. By contrast, the well documented species
P. albida today ranges from eastern peninsular Florida, USA, southwards to the Florida Keys, around the Gulf of
Mexico (López-Vega et al. 2004; Torruco et al. 2007), and southwards around the western Caribbean to Panama
(PoDWA data), Colombia (Diaz Merlano & Hegedus 1994), and the Venezuelan coast (Macsotay & Villarroel
2001). In the east, it extends from Cuba, Jamaica, Hispaniola (PoDWA data), and then southeastwards through the
Antilles to Guadeloupe (Pointier & Lamy 1998), Martinique to Tobago (PoDWA data). However in compiling data
for this the best known species, care must be taken, as some museum records from Colombia and Venezuela and all
those from further east along the northern coast of South America—from Guyana, Suriname, and northern
Brazil—belong to closely related but undescribed species (see Massemin et al. 2009: 212–213). Specimens of P.
albida from Colombian and Venezuelan waters have been erroneously identified as the extinct species P. b a r re tt i
(Guppy) by Petuch (1981: fig. 6a) and this identification has been followed by some subsequent authors (e.g.,
Penchaszadeh 1982; Rosenberg 2009) and on shell dealers’ websites. Diaz Merlano & Hegedus (1994) first noted
that this was a misidentification and we can confirm this.
FIGURE 8. Linear geographic range (longest distance between sampled localities) plotted against rank for 85 living Western
Atlantic morphospecies of Polystira. This hollow curve shows that very few species have large geographic ranges. In contrast,
50 are currently known from just a single site (“spot” occurrence).
We examined geographic ranges of 85 (of a total of 112) western Atlantic morphologically delimited species of
Polystira using PodWA data. We have measured geographic ranges as maximal linear extent measured as a
straight-line distance between the two most distant recorded localities (see Gaston 1994: table 2.1) and have
considered single spot records, for example a single dredging site, to have an arbitrary ‘range’ of 5 km. Our results
(Fig. 8) show that two species have almost identical large ranges of >5000 km: Polystira starretti (5,204 km) and P.
tellea (5,113 km). A further 18, including P. albida, can be considered to be widely distributed (1000–4280 km).
However, a clear majority, 50 of the 85 species (59%), are currently represented in our database by spot
occurrences (<5km), essentially a single site in each case. When plotted with untransformed axes, a ‘hollow curve’
(i.e., right-skewed curve) is produced with few species with large ranges and a long tail of species with very short
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ranges. At large spatial scales like ours, a hollow curve of species ranges is very commonly found among
ecological or taxonomic groupings (Gaston 1994), and has even been described as “a persistent characteristic of
life on Earth” (Gaston 2003: 79); few marine examples of this pattern have been demonstrated so far, however
(Gaston 2003: 79). In our case, we assume that this represents a real pattern in species ranges, though sampling
density is very uneven across the region. Further sampling is likely to uncover yet more species with small ranges
while also increasing the ranges of those that are under-recorded at present. Hence, the overall pattern across the
genus will probably remain similar.
FIGURE 9. Species abundance (log scale) plotted against rank for 85 living Western Atlantic morphospecies of Polystira.
Only five species are represented by more than 100 specimens in the major museum collections examined (see text). The most
numerous species is P. starretti Petuch (1685 specimens), the next two most numerous are P. albida (Perry) (770) and an
undescribed species, Polystira R-GUY-1 (743 specimens). In contrast, 28 are known from just a single specimen (=singleton).
Species-abundance distributions
We analysed PoDWA data to examine variation in species-abundance patterns in major holdings of this genus in
museum collections. The species represented by the largest number of specimens is P. s ta r re t t i with 1685
specimens, the next two most numerous are P. albida (770 specimens) and Polystira R-GUY-1 (see Appendix)
(743 specimens). Only two more species are represented by more than 100 specimens. In contrast, 28 species
(33%) are known from singletons (single specimens) and 29 are known from between two and eleven specimens
(Fig. 9). Plotting of these data reveals a hollow curve, with P. s t ar r et t i being represented by more than twice the
number of specimens than the second most abundant species, P. albida, and a long tail of apparently rare species.
Range size and abundance clearly are not independent variables, and Polystira starretti and P. albida
exemplify a well-known and pervasive pattern that species with wide ranges tend to be abundant (e.g., Gaston
1994, 2003); at the other extreme, our large number of singletons can only have spot occurrences. We chose not to
analyse our data further due to great inequality of sampling effort over the region. Nevertheless, there are clearly
great differences in relative abundance between species and this can be seen within sites when comparing
sympatric species as well as between sites. A similar hollow curve in abundance is seen in the vast deep-water
turrid (sensu lato = Conoidea minus Conidae and Terebridae) fauna of New Caledonia. Of the enormous fauna of
1409 species made during quantitative collecting surveys by Philippe Bouchet and team, 586 (41%) are known
from singletons (Bouchet et al. 2009: fig. 2), and 483 (82%) of these from an empty shell. Our figure for singletons
(33%) is roughly comparable to this and suggests that the overall pattern of rarity we have recorded is likely to be
real. We think that the pattern may become stronger still with more rigorous and denser sampling. It seems likely
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that this hollow curve of abundance exists over a wide range of scales of the taxonomic hierarchy in tropical marine
molluscs and over a wide range of bathymetries (Bouchet et al. 2002, 2009).
Why rarity matters
The realisation that most living species of Polystira are probably rare or point endemics has wide-ranging
implications. It means that estimation of species diversity in hyper-diverse taxa, such as Polystira, is absolutely
dependent upon sampling intensity. One way to obtain an accurate estimate of total living diversity from limited
sampling might be to measure diversity within a number of well sampled regions and then to estimate spatial
turnover within and between them. At present perhaps only two regions in the western Atlantic—1) southern
Florida and the Florida Keys (10 morphospecies) and; 2) the South American coastline eastwards from Trinidad to
Suriname (15 morphospecies)—can be regarded as well sampled. These regions share only two species in
common, P. starretti and P. tellea, the two species with the largest ranges in the tropical western Atlantic (PoDWA
database). For Polystira, much of the tropical western Atlantic must be regarded as still rather poorly sampled;
indeed, our data show that previously unknown species are inevitably found whenever a previously poorly
collected region becomes better sampled. Accurate estimates of the species diversity of Polystira living in the
western Atlantic will thus require more and better controlled sampling.
Most rare species within the western Atlantic will likely continue to remain biologically poorly known.
Species limits—the boundary between intraspecific and interspecific variation—will be difficult to assess for rare
and widely geographically separated taxa (see Lim et al. 2012), because of problems determining where these
boundaries lie. Consequently, our accounting for the total living species diversity of this relatively shallow-
dwelling gastropod genus will remain rather imprecise for the foreseeable future.
The completeness of our knowledge of species diversity for the living Polystira, therefore, may not be so
different from that obtainable for fossil faunas, where common wide-ranging species will obviously be more likely
preserved, found and collected at outcrop, whereas short-ranged species will have a much poorer chance of
recovery. Over the past two decades extensive new collections of mollusc faunas have been made by the Panama
Palaeontology Project (Collins & Coates 1999) from Miocene to Pleistocene age sediments in basins bordering the
south-western Caribbean. As a result, our sampling intensity of the Pliocene-aged rocks, in particular, is a few
orders of magnitude greater than our comparable sampling of the Recent fauna of this part of the Caribbean
(Jackson et al. 1999; Johnson et al. 2007) and consequently our knowledge of the Pliocene Polystira fauna is that
much more complete.
The inter-dependence of the geographic range-size and abundance of a species introduces a potential source of
bias into macroevolutionary studies. Marine gastropod species with restricted geographic ranges tend to show low
abundances (Russell & Lindberg 1988)—a pervasive macroecological pattern (Gaston 2003)—and are thus
inherently less likely to be represented and discovered in the fossil record. Among benthic gastropods there is
expected to be correlation of geographic range-size and abundance with larval ecology, in which alternate modes of
larval development (planktotrophic versus non-planktotrophic) convey very different dispersal potentials (e.g.,
Jablonski 1986; Krug 2011). Consideration of larval mode is clearly central to better understanding and
interpreting the fossil record of gastropod species radiations.
Species durations, evolution and extinction rates
Among living Polystira species, P. albida has the longest duration. It first occurs in the Cayo Agua Formation of
Atlantic Panama in strata dated to the Early Pliocene, 3.5–5.0 (midpoint estimate 4.25) Ma. This extended duration
is very much an exception among extant species of Polystira, however (Todd & Johnson 2013).
The high diversity and rapid chronological turnover of Polystira species is illustrated by the fauna of a single
geological formation in the Southern Limon Basin of Caribbean Costa Rica. So far we have delimited 16 species of
Polystira from the shallow lagoonal, inter-reef and fore-reef mud facies (Coates 1999a, b: sections 34–38; McNeill
et al. 2000; Todd & Collins 2005) of the Moin Formation of Early Pleistocene age (exposures mostly dated
between 2.1–1.5 Ma: McNeill et al. 2000). Of these, only two, Polystira albida and P. te ll e a , have so far been
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found living in the Caribbean today, yielding an apparent extinction of 88% of the local standing diversity over the
past ca 2 My within this small part of the south-western Caribbean. This pattern strongly contrasts with 71% of the
prosobranch gastropod fauna (total fauna = 306 species) of the Moin Formation reported to still be extant
(Robinson 1993). Much of this extinction of Polystira was focused during the past 1.6 My, a poorly sampled
interval due to limited exposure of shelf strata of this age (Todd & Johnson 2013). In other macrofauna too,
extinction rates peaked a million years or more after isthmian seaway closure (ca 3 Ma) during a time of profound
regional oceanographic change (O’Dea et al. 2007). Over the past 2 My Polystira shows a comparable level of
extinction to that recorded in pectinid bivalves (95%) over a longer period (4–0 Ma) in the same region (Smith &
Jackson 2009). The Pectinidae extinction mostly occurred between 4–3 Ma, with extinction of only 10 of the
standing diversity of 26 species (38%) in the last 2 Myr (Smith & Jackson 2009: fig. 4A). Discounting the
possibility that this extraordinarily high level of turnover in Polystira is due not to regional extinction but to faunal
migrations within the Caribbean—for which we have only limited evidence (Robinson 1993)—the corollary is that
if the south-western region is unexceptional then the large majority of the ca 112 living Polystira species known
from the Caribbean are very likely to be less than 1.6 million years old (Todd & Johnson 2013).
Recently, we have estimated speciation rates from a combined dataset of Recent and fossil Polystira
occurrences in the south-western Caribbean (Todd & Johnson 2013). Our taxic analyses reveal that Polystira shows
faster origination rates (0.585–0.935 My
–1
) than those so far determined for Conus—previously celebrated as the
most rapidly diversifying marine gastropod (Kohn 1990; Stanley 2008; Puillandre et al. 2014). Further studies are
required to facilitate comparison between these taxa and more importantly to determine whether either is truly
exceptional with respect to diversification dynamics within the conoidean radiation as a whole. It is clear that
further studies integrating analyses of living and extinct species are necessary before we can accurately outline the
major patterns of conoidean diversity change through space and time and begin to understand the interaction of
intrinsic and extrinsic driving forces in generating hyperdiversity (Todd & Johnson 2013).
Acknowledgements
This research was supported by NERC Standard Research Grant GR3/13110 (Co-PIs: JAT and Richard H Thomas),
Systematics Association Research Grant (22/03/2000) (P-I: JAT) and the Natural History Museum, London.
Dredging cruises in the Gulf of Panama (2000) and the Lower Florida Keys (2001) were on R/V Urraca and R/V
Bellows respectively and we thank the captain and crew of each and the respective expedition leaders Harilaos
Lessios (STRI, Panama) and Timothy Collins (FIU, Miami). Our involvement in the latter cruise was supported by
a grant from the Florida Institution of Oceanography. Many thanks to all those who have helped us to obtain fresh
or alcohol-preserved specimens of Polystira and other turrids including: Nestor Ardila, Rüdiger Bieler, Paul
Callomon, Henry Chaney, Rachel Collin, Tim Collins, Emilio Garcia, Ivan & Charlotte Goodbody, Jeroen Goud,
Paul Greenhall, Lindsey Groves, Jerry Harasewych, Edgar Heimer de la Cotera, Kirstie Kaiser, Dominique Lamy,
James McLean, Paula Mikkelsen, Brian Morton, Baldomero Olivera, Colin Redfern, Gary Rosenberg (also for
bibliographic information), Luiz Simone, John Slapcinsky, John Taylor, Ole Tendal, Fred Thompson, Nancy Voss,
and others. We would like to thank curators and collection managers in the following institutions for kindly giving
us access to their collections: AMNH, ANSP, FMNH, HMNS, LACM, MHNMC (INVEMAR), MNHN, MZUSP,
NHMB, NMNH, NHMUK, RMNH, FLMNH (UF), UMML & ZOOLOGISK MUSEUM (Copenhagen). Karl
Bates (then NHM, London) helped provide SEM images, measurements of protoconchs and estimates of ranges.
Zoë Hughes (NHM) made some images of type specimens and Ken Johnson (NHM) helped support this work.
Felix Rodriguez (STRI, Panama) helped us collect specimens in Panama, loaned collections from STRI, Panama
City and provided us with unpublished information on living specimens. Finally we would like to acknowledge the
late Donn Tippett for having very kindly provided us with his extensive notes on the taxonomy of Polystira
species. Warren Allmon, an anonymous reviewer, and editor Thomas Duda are thanked for their helpful comments
that have much improved this paper.
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APPENDIX: Polystira specimens and associated Turridae (s.s.) outgroup taxa used in the molecular phylogeny.
Museum registration numbers for voucher specimens are provided as well as locality information (including field
number, collector and date, where available) and GenBank accession numbers for the three gene regions analyzed. All
sequences have been newly submitted apart from one marked *. Specimens are listed in the order presented in the
phylogram (Fig. 6).
GenBank Accession #s
Species Registration # Country Locality, Collector & Year rrnS rrnL cox1
Polystira albida (Perry, 1811)
NHMUK MOEA
20140676
Panam a-
Caribbean
San Blas Islands, Panama (SB02-
03); 44m; 9° 31.49’N, 78°
57.49’W; R. Collin; 2002
KM218461 KM218558 KM218654
NHMUK MOEA
20140678
USA-FL S of Marquesas Rock, Florida
Keys (FK-613); 82.5m; 24°
23.441’N, 82° 14.081’W; R.
Bieler, P. M. Mikkelsen, T. A.
Rawlings; 2002
KM218462 KM218559 KM218655
NHMUK MOEA
20110066
USA-FL W of Looe Key Reef, S. of Bahia
Honda Key (FK-539); 30-34m;
24° 34.24’N, 81° 16.64’W; R.
Bieler, P. M. Mikkelsen, T. A.
Rawlings; 2001
KM218463 JF276963* KM218656
USNM 751627 USA-LO South of Marsh Island, Louisiana,
(R/V Oregon-Sta. 3801); 49-51m;
28° 27’N, 92° 51’W; 1962
KM218464 KM218560 KM218657
USNM 663946 Mexico-
Caribbean
Campeche Bay, Mexico, (R/V
Oregon, Sta. 723); 46m; 20°
48.2’N; 91° 45.7’W; 1952
KM218465 KM218561 KM218658
USNM 751657 Colombia -
Caribbean
Gulf of Uraba, Colombia, (R/V
Oregon II, Sta. 11234); 51m; 8°
49’N; 76° 53’W; 1970
KM218466 KM218562 KM218659
FMNH 297362 USA-FL Ramrod Key, SW corner of Looe
Key National Marine Sanctuary,
Florida Keys (R-21, Sta. FK-
336); 43.4m; 24.5264°N
81.4322°W; R. Bieler, P.M.
Mikkelsen; 2000
KM218467 KM218563 KM218660
USNM 751641 USA-TX Off Galveston, Texas, (R/V
Oregon-Sta. 3831); 51m; 28°
22’N; 94° 02’W; 1962
KM218468 KM218564 KM218661
NHMUK MOEA
20140675
Panam a-
Caribbean
San Blas Islands, Panama (SB02-
31); 32m; 9° 34.26’N, 78°
43.9’W; R. Collin; 2002
KM218469 KM218565 KM218662
NHMUK MOEA
20140677
Panam a-
Caribbean
San Blas Islands, Panama (SB02-
11); 45m; 9° 30.95’N, 78°
57.7’W; R. Collin; 2002
KM218470 KM218566 KM218663
UMML 30.6467 Venezuela NE of San Juan de las Goldonas,
Peninsula de Paria, Venezuela;
(P-710); 47m; 10° 47’N; 62°
55’W; 1968
KM218471 KM218567 KM218664
UMML 30.11479 Dominican
Republic
Bahia de Neiba, S coast of
Dominican Republic; (P-1294);
48m; 18° 15’N; 70° 58’W; 1970
KM218472 KM218568 KM218665
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GenBank Accession #s
Species Registration # Country Locality, Collector & Year rrnS rrnL cox1
UMML 30.11479 Dominican
Republic
Bahia de Neiba, S coast of
Dominican Republic; (P-1294);
48m; 18° 15’N; 70° 58’W; 1970
KM218473 KM218569 KM218666
UMML 30.10633 Martinique NE of Basse Pointe, Martinique;
(P-913); 47m; 14° 54’N; 61°
04’W; 1969
KM218474 KM218570 KM218667
Polystira coltrorum Petuch, 1993
MZUSP 116827 Brazil Off Guarapari, Espirito Santo
State, Brazil; 30-35m; local
fisherm en; 2000
KM218475 KM218571 KM218668
Polystira R-GUY-3
RMNH 81125 Surinam Off Surinam (Luymes OCPS-II,
Sta. M88); 46m; 06° 42.8’N; 53°
58.8’W; 1969
KM218476 KM218572 KM218669
RMNH 81104 Surinam Off Surinam (Luymes OCPS-II,
Sta. M73); 55m; 06° 55’N; 53°
54.5’W; 1969
KM218477 KM218573 KM218670
Polystira nobilis (Hinds, 1843)
NHMUK MOEA
20140684
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-34); 66—68m;
08° 26.24’N; 79° 9.14’W; J. A.
Todd; 2000
KM218478 KM218574 KM218671
NHMUK MOEA
20140683
Panama-Pacific Gulf of Panama (GP97-16, CTPA
00454, sample 1400); 94m; 08°
14.9’N; 79° 15.2’W; H.
Fortunato; 1997
KM218479 KM218575 KM218672
NHMUK MOEA
20140682
Panama-Pacific Islas de las Perlas , Gulf of
Panama (JTD-00-34); 66—68m;
08° 26.24’N; 79° 9.14’W; J. A.
Todd; 2000
KM218480 KM218576 KM218673
Polystira R-GUY-9
RMNH 81114 Surinam Off Surinam (Luymes OCPS-II
Expedition, 1969, Sta. J112);
88.5m; 06° 52.8’N; 54° 41’W;
1969
KM218481 KM218577 KM218674
RMNH 81062 Surinam Off Surinam, (Sta. M97); 130m;
07° 18.5’N, 53° 49’W; 1969
KM218482 KM218578 KM218675
UMML 30.7327 Trinidad &
Tob ago
Off E coast of Tobago (R163; P-
842); 70m; 11 11’N, 60° 31’W;
1969
KM218483 KM218579 KM218676
Polystira R-GAN-3
UMML 30.6276 Bahamas Off NW tip of Little Bahama
Bank (R109; G-394); 223m; 27°
22’N, 79° 11’W; 1964
KM218484 KM218580 KM218677
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Appendix continued
GenBank Accession #s
Species Registration # Country Locality, Collector & Year rrnS rrnL cox1
UMML 30.7805 Bahamas Cay Sal Bank, Bahamas (G-985);
217m; 24° 06’N, 80° 12’W; 1968
KM218485 KM218581 KM218678
Polystira R-GOH-4
UMML 30.6112 Honduras Bahia de Omoa, Honduras (R162;
P-168); 113m; 15° 46’N, 88°
10’W; 1967
KM218486 KM218582 KM218679
UMML 30.6124 Honduras Off N coast of Honduras (R146,
P-628); 47m; 15° 57’N, 86°
15’W; 1967
KM218487 KM218583 KM218680
Polystira R-GAN-2
UMML 30.6275 Bahamas Off NW tip of Little Bahama
Bank, Bahamas, (G-393); 169m;
27° 22’N, 79° 11’W; 1964
KM218488 KM218584 KM218681
Polystira tellea (Dall, 1889)
NHMUK MOEA
20140700
USA-FL S of Marquesas Rock, Florida
Keys (FK-610); 106-110m; 24°
23.413’N, 82° 12.720’W; R.
Bieler, P. Mikkelsen, & T. A.
Rawlings; 2002
KM218489 KM218585 KM218682
NHMUK MOEA
20140699
USA-FL SW of Sombrero Key Light,
Florida Keys (JTD-01-16); 129-
156m; 24 32.99’N, 81 08.33’W;
J. A. Todd & T. A. Rawlings;
2001
KM218490 KM218586 KM218683
MHNMC MAC-
6291
Colombia-
Caribbean
Off Punta Gallinas, Bahia Honda
(E10, Sta. G31); 310-314m; 12°
34’18”N, 71° 50’0”W; N. Ardila;
1998
KM218491 KM218587 KM218684
RMNH 81110 Surinam Off Surinam (Luymes OCPS-II
Expedition 1969, Sta. K102);
81m; 7° 11.3’N, 54° 23’W; 1969
KM218492 KM218588 KM218685
RMNH 81123 Guyana Off Guyana (Luymes Guyana
Shelf Expedition, Sta. 15049);
120-200m; 07° 44’N, 57° 03’W;
1970
KM218493 KM218589 KM218686
MHNMC MAC-
6422
Colombia-
Caribbean
Bahia Portete, Colombia (E15,
Sta. G32); 304-310m; 12° 24’
1.8’’N; 72° 15’ 0.6”W; N. Ardila;
1998
KM218494 KM218590 KM218687
MHNMC MAC-
6536
Colombia-
Caribbean
N of Dibulla, La Guajira
Province, Colombia (E23, Stn
P31); 298-300m; 11° 29’25.8”N,
73° 22’49.8”W; N. Ardila; 1998
KM218495 KM218591 KM218688
UF 28756 USA-FL SW of Boca Grande Key, Straits
of Florida, Florida; 102m; 24°
23.76’N, 82° 02.12’W; K.
Auffenberg; 1980
KM218496 KM218592 KM218689
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GenBank Accession #s
Species Registration # Country Locality, Collector & Year rrnS rrnL cox1
USNM 751746 Surinam Off Paramaribo, Surinam (R/V
Oregon II, Sta. 10622); 201m; 7°
27’N, 54° 30’W; 1969
KM218497 KM218593 KM218690
UMML 30.11477 Trinidad &
Tob ago
Off N coast of Trinidad, (P- 848);
146m; 11° 23’N, 61° 26’W; 1969
KM218498 KM218594 KM218691
NHMUK MOEA
20140698
USA-FL S of Sombrero Key Light, Florida
Keys (JTD-01-11); 112-118m; 24
34.16’N, 81 05.91’W; J. A. Todd
& T. A. Rawlings; 2001
KM218499 KM218595 KM218692
UMML 30.11484 Surinam Off Surinam (O-4182); 73m; 7°
02’N, 54° 00’W; 1963
KM218500 KM218596 KM218693
Polystira R-GAN-10 [=R-MIS-1]
UMML 30.7805 Bahamas Cay Sal Bank, Bahamas (G-985);
217m; 24° 06’N, 80° 12’W; 1968
KM218501 KM218597 KM218694
UMML
30.11476
Nicaragua E of Cayos Miskitos (P-1354);
228m; 14° 21’N, 81° 55’W; 1971
KM218502 KM218598 KM218695
Polystira picta (Reeve, 1843)
ANSP A6646 Mexico-Pacific San Carlos, Guaymas, Sonora,
(ANSP—A6646); 35m; 27°
59’N, 110° 54’W; L. & F.
Poorman; 1975
KM218503 KM218599 KM218696
ANSP A9742D Ecuador 1 mile W of jetty at Esmeraldas;
(T357, Sta, 25); 6-12m; 01° 01’N,
79° 43’W; P. Skoglund & V. O.
Maes; 1981
KM218504 KM218600 KM218697
ANSP 358094 Mexico-Pacific 1 mile E of Punta Doble, Sonora;
(ANSP A10200, T357); 23-33m;
27° 55’N, 111° 05’W; V. O. Maes
et al.; 1982
KM218505 KM218601 KM218698
NHMUK MOEA
20140694
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-41); 24-25m,
08° 31.93’N, 79° 5.60’W; J. A.
Todd; 2000
KM218506 KM218602 KM218699
ANSP A9755 Costa Rica-
Pacific
W side of Bahia Culebra, off
Nacascola (T35713); 23m; 10°
37’ 15”N, 85° 41’ 20”W; P.
Skoglund; 1982
KM218507 KM218603 KM218700
ANSP 358094 Mexico-Pacific 1 mile E of Punta Doble, Sonora;
(ANSP A10200, T357); 23-33m;
27° 55’N, 111° 05’W; V. O. Maes
et al.; 1982
KM218508 KM218604 KM218701
NHMUK MOEA
20140693
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-46); 24-25m;
08° 31.37’N, 79° 5.79’W; J. A.
Todd; 2000
KM218509 KM218605 KM218702
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GenBank Accession #s
Species Registration # Country Locality, Collector & Year rrnS rrnL cox1
Polystira R-PAN-10
NHMUK MOEA
20140681
Panam a-
Caribbean
San Blas Islands, Panama (no
other locality details available);
R. Collin; 2002
KM218510 KM218606 KM218703
Polystira R-VEN-4
UMML 30.6707 Venezuela N of Puerto Píritu, Anzoátegui
State, Venezuela (P-727); 64m;
10° 20’N, 65° 02’W; 1968
KM218511 KM218607 KM218704
UMML 30.6498 Venezuela W of Los Testigos Archipelago,
Venezuela (P-714); 59m; 11°
29’N, 63° 24’W; 1968
KM218512 KM218608 KM218705
Polystira R-COL-10
MHNMC MOL-
1921
Colombia -
Caribbean
NW of Islas de San Bernardo,
Gulf of Morosquillo (E-73); 268-
280m; 09° 57’ 40.8”N; 76° 7’
57”W; N. Ardila; 1999
KM218513 KM218609 KM218706
Polystira vibex (Dall, 1889)
UMML 30.7453 Bahamas N of North Bimini, Bahamas (G-
637); 183m; 26° 05’N, 79° 13’W;
1965
KM218514 KM218610 KM218707
Polystira aff. parthenia (Berry, 1957)
ANSP 358119 Mexico-Pacific 2 miles E of Punta Doble, Sonora;
20-30m, 27° 55’N, 111° 04’W;
V.O. Maes et al.; 1982
KM218515 KM218611 KM218708
Polystira parthenia (Berry, 1957)
LACM 1984-53.11 Panama-Pacific Bahia Montijo, SW end of Isla
Gobernadora, Veraguas Province;
intertidal; 7° 33.30’N; 81° 13.70’
W; T. Bratcher; 1984
KM218516 KM218612 KM218709
LACM-1934-
127.30
Costa Rica-
Pacific
Puerto Culebra, Guanacaste
Province (R/V Velero III); 18m;
10° 37 30’N, 85° 40.0’W; 1934
KM218517 KM218613 KM218710
Polystira oxytropis (G.B. Sowerby I, 1834)
NHMUK MOEA
20140688
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-35); 72-76m;
08° 25.81’N, 79° 11.03’W; J. A.
Todd; 2000
KM218518 KM218614 KM218711
NHMUK MOEA
20140689
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-35); 72-76m;
08° 25.81’N, 79° 11.03’W; J. A.
Todd; 2000
KM218519 KM218615 KM218712
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GenBank Accession #s
Species Registration # Country Locality, Collector & Year rrnS rrnL cox1
NHMUK MOEA
20140685
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-31); 15-17m;
08° 19 15’N, 79° 06.88’W; J. A.
Todd; 2000
KM218520 KM218616 KM218713
NHMUK MOEA
20140686
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-46); 24-25m;
08° 31.37’N, 79° 05.79’W; J. A.
Todd; 2000
KM218521 KM218617 KM218714
NHMUK MOEA
20140691
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-34); 66-68m;
08° 26.24’N,79° 09.14’W; J. A.
Todd; 2000
KM218522 KM218618 KM218715
NHMUK MOEA
20140690
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-34); 66-68m;
08° 26.24’N,79° 09.14’W; J. A.
Todd; 2000
KM218523 KM218619 KM218716
NHMUK MOEA
20140692
Panama-Pacific Islas de las Perlas, Gulf of
Panama, (JTD-00-34); 66-68m;
08° 26.24’N,79° 09.14’W; J. A.
Todd; 2000
KM218524 KM218620 KM218717
NHMUK MOEA
20140687
Panama-Pacific Gulf of Chiriqui (GC 89-1, CTPA
00410, sample #1564); H.
Fortunato; 1989.
KM218525 KM218621 KM218718
ANSP 358093 Mexico-Pacific 1 mile E of Punta Doble, Sonora;
23-33m, 27° 55’N, 111° 05’W;
V.O. Maes et al.; 1982
KM218526 KM218622 KM218719
ANSP 358119 Mexico-Pacific 2 miles E of Punta Doble, Sonora;
20-30m, 27° 55’N, 111° 04’W;
V.O. Maes et al.; 1982
KM218527 KM218623 KM218720
ANSP
A9741F
Ecuador 3 miles SW of Esmeraldas (Sta.
28); no depth information given;
0° 99’N, 79° 44’W; P. Skoglund
& V.O. Maes; 1981
KM218528 KM218624 KM218721
Polystira starretti Petuch, 2002
RMNH 81120 Surinam Off Surinam (Luymes, Guyana
Shelf Expedition, 1970, Sta.
15035); 200m; 7° 23’N, 53°
54’W; 1970
KM218529 KM218625 KM218722
MHNMC MAC-
6536
Colombia-
Caribbean
N of Dibulla, La Guajira Province
(E23, P31); 298-300m; 11°
29’25.8”N, 73° 22’49.8”W; N.
Ardila; 1998
KM218530 KM218626 KM218723
RMNH 81114 Surinam Off Surinam (Luymes OCPS-II
Expedition, 1969, Sta. J112);
88.5m; 06° 52.8’N; 54° 41’W;
1969
KM218531 KM218627 KM218724
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GenBank Accession #s
Species Registration # Country Locality, Collector & Year rrnS rrnL cox1
MHNMC MAC-
6291
Colombia-
Caribbean
NW of Punta Gallinas, Bahia
Honda (E10, Stn G31); 310-
314m; 12° 34’18”N, 71° 50’0”W;
N. Ardila; 1998
KM218532 KM218628 KM218725
NHMUK MOEA
20140697
USA-FL SE of Sombrero Key Light,
Florida Keys (JTD-01-04); 111-
124m; 24 34,29’N, 81 05.19’W;
J. A. Todd & T. A. Rawlings;
2001
KM218533 KM218629 KM218726
RMNH 81123 Guyana Off Guyana (Luymes Guyana
Shelf Expedition, Sta. 15049);
120-200m; 07° 44’N, 57° 03’W;
1970
KM218534 KM218630 KM218727
NHMUK MOEA
20140696
USA-FL SW of Sombrero Key Light,
Florida Keys (JTD-01-18); 123-
147m; 24° 32.96’N, 81° 09.08’W;
J. A. Todd & T. A. Rawlings;
2001
KM218535 KM218631 KM218728
RMNH 81115 Surinam Off Surinam (Luymes OCPS-II
Expedition, 1969, Sta. I116);
60m; 07° 00’N; 54° 54’W; 1969
KM218536 KM218632 KM218729
NHMUK MOEA
20140695
USA-FL SE of Sombrero Key Light,
Florida Keys (JTD-01-08); 140-
146m; 24° 34.010’N, 81°
03.090’W; J. A. Todd & T. A.
Rawlings; 2001
KM218537 KM218633 KM218730
RMNH 81104 Surinam Off Surinam (Luymes OCPS-II
Expedition, 1969, Sta. M73);
55m; 06° 55’N; 53° 54.5’W; 1969
KM218538 KM218634 KM218731
UMML 30.11477 Trinidad &
Tob ago
Off N coast of Trinidad(P-848);
146m, 11° 23’N, 61° 26’W; 1969
KM218539 KM218635 KM218732
Polystira artia (Berry, 1957)
LACM 1950-28-5 Mexico-Pacific Baja California Sur, Mexico; (R/
V Velero IV); 66-71m; 26°
52.68’N; 114° 2.30’W; 1950
KM218540 KM218636 KM218733
NHMUK MOEA
20140679
Panama-Pacific Gulf of Panama (GP 97-7; CTPA
-00434, Sample 1378); 88m; 07°
44.9’N; 78° 35.2’W; H.
Fortunato; 1997
KM218541 KM218637 KM218734
NHMUK MOEA
20140680
Panama-Pacific Gulf of Panama (GP 97-7; CTPA
-00434, Sample 1378); 88m; 07°
44.9’N; 78° 35.2’W; H.
Fortunato; 1997
KM218542 KM218638 KM218735
ANSP
A6642
Mexico-Pacific S.W. of Point Doble, Guaymas,
Sonora, Mexico; 80-90m; 27°
59’N, 110° 54’W; 1950
KM218543 KM218639 KM218736
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GenBank Accession #s
Species Registration # Country Locality, Collector & Year rrnS rrnL cox1
Polystira R-GUY-1 [=R-VEN-10]
UMML 30.6467 Venezuela NW of San Juan de las Goldonas,
Peninsula de Paria, Venezuela ( P-
710); 47m; 10° 47’N; 62° 55’W;
1968
KM218544 KM218640 KM218737
RMNH 81115 Surinam Off Surinam (Luymes OCPS-II
Expedition, 1969, Sta. I116);
55m; 07° 00’N; 54° 54’W; 1969
KM218545 KM218641 KM218738
RMNH 81104 Surinam Off Surinam (Luymes OCPS-II
Expedition, 1969, Sta. A68);
55m; 06° 55’N; 53° 54.5’W; 1969
KM218546 KM218642 KM218739
RMNH 81125 Surinam Off Surinam (Luymes OCPS-II
Expedition, 1969, Sta. M88);
46m; 06° 42.8’N; 53° 58.8’W;
1969
KM218547 KM218643 KM218740
RMNH 81100 Surinam Off Surinam (Luymes OCPS-II
Expedition, 1969, Sta. A68);
37.5m; 06° 32’N; 55° 16’W; 1969
KM218548 KM218644 KM218741
Polystira R-GAN-26
UMML 30.11479 Dom inican
Republic
Bahia de Neiba, S coast of
Dominican Republic (P-1294);
48m; 18° 15’N; 70° 58’W; 1970
KM218549 KM218645 KM218742
Outgroups
Gemmula hindsiana Berry, 1958
NHMUK MOEA
20140702
Panama-Pacific Islas de las Perlas, Gulf of
Panama (JTD-00-35); 72-76m;
08° 25.81’N, 79° 11.03’W; J. A.
Todd; 2000
KM218550 KM218646 KM218743
Gemmula periscelida (Dall, 1889)
UF 290962 USA Off the lower Florida Keys,
Florida; 494-530m; 24 14.959’N,
82 48.104’W; E. Rothrock; 2001
KM218551 KM218647 KM218744
Ptychosyrinx ?carynae (Haas, 1949)
USNM 832922 USA Off Virginia, North Atlantic
Ocean; 3188m; 38 00 12’N. 070
29 44’W; R. Carney; 1978
KM218552 KM218648 KM218745
Gemmula diomedea Powell, 1964
NHMUK MOEA
20140701
Philippines Aligway, Philippines; B. Olivera;
2001
KM218553 KM218649 KM218746
Turris babylonia (Linnaeus, 1758)
NHMUK MOEA
20140704
Philippines Olango Island, Cebu, Philippines;
B. Olivera; 2001
KM218554 KM218650 KM218747
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GenBank Accession #s
Species Registration # Country Locality, Collector & Year rrnS rrnL cox1
Xenuroturris albina (Lamarck, 1822)
NHMUK MOEA
20140703
Philippines Olango Island, Cebu, Philippines;
B. Olivera; 2001
KM218555 KM218651 KM218748
Xenuroturris legitima Iredale, 1929
NHMUK MOEA
20140705
Philippines Olango Island, Cebu, Philippines;
B. Olivera; 2001
KM218556 KM218652 KM218749
Unedogemmula leucotropis (Adams & Reeve, 1850)
NHMUK MOEA
20140706
China: Hong
Kong
Off Hong Kong, South China Sea
(Sta. 75); B. Morton; 2001
KM218557 KM218653 KM218750
