The Sipuncula: Their Systematics, Biology, and Evolution 9781501723643

Table of contents :
Contents
Preface
Checklist of the Sipunculan Species
Introduction
Glossary
Part I. Systematics
Part II. Sipunculan Biology: A Review
Part III. Zoogeography and Evolution
Appendix 1. Recent Species lnquirenda and lncertae Sedis
Appendix 2. Species lnquirenda and lncertae Sedis as in Stephen and Edmonds, 1972, with Current Status
Bibliography
Taxonomic Index
Subject Index

Citation preview

The Sipuncula

The Sipuncula Their Systematics, Biology, and Evolution

EDWARD

B.

CUTLER

Department of Biology Utica College of Syracuse University

Comstock Publishing Associates a division of Cornell University Press Ithaca and London

Copyright © 1994 by Cornell University All rights reserved. Except for brief quotations in a review, this book, or parts thereof, must not be reproduced in any form without permission in writing from the publisher. For information, address Cornell University Press, Sage House, 512 East State Street, Ithaca, New York 14850. First published 1994 by Cornell University Press. Printed in the United States of America

I 2 3

)

l

Figure 84. Four developmental pathways followed by various Sipuncula. Type 1. Direct development with no pelagic stage. Type 2. Pelagic lecithotrophic trochophore that becomes a vermiform phase. Type 3· The pelagic lecithotrophic trochophore metamorphoses into a lecithotrophic pelagosphera larva that subsequently changes into a vermiform phase. Type 4· The pelagic lecithotrophic trochophore metamorphoses into a planktotrophic pelagosphera larva. After an extended planktonic existence during which it increases in size, this metamorphoses a second time into a vermiform juvenile. (After Rice, 1975c, courtesy of M. E. Rice.)

I. Direct lecithotrophic development with no pelagic stage (D). Found in 3 species from three genera, representing all three families of the order Golfingiiformes. II. One lecithotrophic pelagic stage: trochophore (T). Found in 2 species, each representing one of the two orders in the class Sipunculidea. III. Two pelagic stages: trochophore and lecithotrophic pelagosphera (LP). In 7 species from four genera representing three families in the Golfingiiformes. IV. Two pelagic stages: trochophore and planktotrophic pelagosphera (PP). Found in 10 species; 3 from the class Sipunculidea, representing three genera in two families, one from each order. The remaining 7 examples are from four of the six genera in both orders and families of the class Phascolosomatidea (Table 3).

Reproduction and Regeneration Table 3. Larval developmental types, distributed by taxa Type

II Class Sipunculidea Order Sipunculiformes Family Sipunculidae Sipunculus Siphonosoma Phascolopsis Order Golfmgiiformes Family Golfingiidae Golfingia Nephasoma Thysanocardia Family Phascolionidae Phascolion Family Themistidae Themiste Class Phascolosomatidea Order Phascolosomatiformes Family Phascolosomatidae Phascolosoma Apionsoma Antillesoma Order Aspidosiphoniformes Family Aspidosiphonidae Aspidosiphon

III

IV

xa X

X

XX X

X

X X

X

X

X

XXX

XXX X X XX

Notes: X = one species. No data for five genera. • Somewhat unique pattern.

Species with a trochophore larva commonly spend 2-4 days in this stage (8-10 days in a few species). The lecithotrophic larval stage lasts from 2 days to two weeks, and the planktotrophic stage, when present, lasts on the order of one to three months, occasionally up to six months. The trochophore is fairly typical. It has a broad equatorial prototroch, a ventral metatroch, an apical tuft of sensory cilia, and a complete tripartite mesodermal gut (Figs. 81, 82). The term trochophore can be confusing, and Salvini-Plawen (1973) proposed that it should be restricted to the Annelida and Echiura, a point of view that does not seem to have broad support. The trochophore larva has an ectodermally derived ventral nerve cord, inverted ocelli, and is positively phototaxic (Akesson, 1958, 1961). The details of ciliation and origins of various muscle systems vary from

Sexual Reproduction species to species. The larval cuticle is derived from the egg envelope in most species but is created de novo in others. From a trochophore, some species elongate posterior to the prototroch into a benthic vermiform juvenile. Those that metamorphose into planktonic pelagosphera larvae will later change into the juvenile worm (Figs. 81, 84). The pattern inS. nudus is unique and is considered highly modified. Differences include the fate of the egg envelope (completely cast off during change to pelagosphera rather than becoming the cuticle), and the uniformly ciliated trochophore rather than the more usual narrow prototrochal band (Rice, 1988b). Read Hyman, 1959, for an interesting summary of the research on Sipunculus larvae, which were first named Pelagosphaera and treated as adults of a distinct genus. Much of the early work with these larvae was flawed because it was based on contracted preserved material with withdrawn anterior ends. One of the first to use fresh material was Jagersten ( 1963), who provided excellent drawings of what he characterized as hippopotamus-like heads. Two other Sipunculus species were drawn from living material by Murina (1965). Pelagosphera larvae have well-developed metatrochal cilia and a characteristic head. Besides the obvious behavioral and ecological differences, the internal morphological differences from the adult include the larger number of retractor muscles, eyespots, and body wall muscles. The final metamorphosis from pelagic larva to adult includes organogenesis of several systems, including the introvert and tentacles. The final metamorphosis can be induced in a competent larva by exposing it to sediment previously occupied by adults of the same species. Hall and Scheltema (1966, 1975b) described ten open-ocean planktonic sipunculan larvae. These authors focused on cuticular structures, but they also considered pigmentation and other organ systems such as the body wall musculature. They were unable to relate the larvae to specific adult forms, but among the larvae described were representatives of at least the genera Aspidosiphon, Phascolosoma, and Sipunculus. The presence of multiple retractor muscles in larvae (more pairs than in adults) is intriguing, but the question of whether this difference is due to fusion or to loss during metamorphosis is still unanswered. The larvae described by Hall and Scheltema were kept alive in the laboratory for several months, and a few did undergo metamorphosis. Among the interesting facts gleaned by those authors is that larvae do expend energy to maintain their vertical position in the water column.

308

Reproduction and Regeneration

Larval Dispersal and Settlement Most shallow-water tropical and warm temperate species are widely dispersed by oceanic currents. The teleplanic larval stage lasts two to six months, adequate time to allow transoceanic transport or island colonization (Hall and Scheltema, 1975a; R. Scheltema, 1986a; R. Scheltema and Rice, 1990). During the period 1975-1990, R. Scheltema published seven articles on sipunculan larvae dispersal; the bulk of that work is discussed in Chapter 16 (see Zoogeography). Rice (1986) studied settlement and metamorphosis in a Florida population of Apionsoma misakiana. The settlement-inducing substance (SIS) is species-specific, stable for at least eight days, heat-labile (autoclaving destroys it but freezing has no effect), and has a molecular weight less than 500. Larvae responded to SIS in the seawater by settling, but the response was stronger if sediment was also present, and stronger still if adults were present in the sediment. The possibility that the SIS acts synergistically with bacterial film on the sediment has not been ruled out. Larvae can attach to sediment temporarily using the posterior retractile terminal organ, which has sensory cells and produces an adhesive mucus (Ruppert and Rice, 1983). Ruppert and Rice compared this organ with adhesive organs in other metazoans and concluded that the sipunculan terminal organ evolved independently within the phylum.

Asexual Reproduction Parthenogenesis Parthenogenesis is a "partial" sexual process in that although meiosis occurs and oocytes are formed, no syngamy occurs, and it is thus, by definition, not sexual reproduction. Facultative parthenogenesis, the spontaneous development of unfertilized eggs into normal larvae, appears to be common in Florida Themiste lageniformis, where females outnumber males 24 to I (Pilger, 1987). This mode of reproduction is unknown in other members of the phylum. Budding Unequal transverse fission has been observed in two species: Aspidosiphon elegans (Rice, 1970) and a species belonging to the family Sipunculidae (Rajulu and Krishnan, 1969; Rajulu, 1975).

Asexual Reproduction RM

VNC

ID

lA

Figure 85. The dissected bud and posterior end of Aspidosiphon elegans showing internal organs of parent and offspring. The anterior parts of the offspring will develop from the epidermal invagination. C, collar; E, esophagus; El, epidermal invaginations; lA, ascending intestine; ID, descending intestine; N, nephridium; RM, retractor muscle; S, internal acellular partition across the stricture; SM, spindle muscle; VNC, ventral nerve cord. (After Rice, 1970, courtesy of M. E. Rice,© 1970 by the AAAS.)

In A. e/egans a constriction appears near the end of the trunk, and essential internal organs-including the introvert, retractor muscles, anterior intestine, and nephridia-are replicated in the smaller "daughter" part (Fig. 85). The anterior "parent" only needs to regenerate the posterior body wall as separation occurs. About 15% of the individuals collected in a Caribbean coral community were budding (E. Cutler, pers. observ.). Thus, this seems to be a natural phenomenon and not the result of laboratoryinduced stress. The budding Sipunculidae species was identified as Sipunculus robustus, but my superficial inspection of the specimen indicated that it may

310

Reproduction and Regeneration

well be a Siphonosoma species, most likely S. cumanense. This speculation is supported by personal observations of living members of the latter species collected in Madagascar, which underwent a "pinching off" into subsets when kept for several days in stale seawater. Other than the fact that fission has not been observed inS. cumanense under natural conditions, and seems to occur only in response to environmental stress, it is similar to the sequence in Aspidosiphon. The "daughter" is the smaller posterior part, up to one-third of the trunk. The posterior half may form three to five buds, including lateral ones. The new central nervous system, contractile vessel, set of retractor muscles, and digestive system are produced before separation. The new introvert, tentacles, and anus are formed afterward. Based on thin sections made after three days, the beginning of this process involves the production of an elongate girdlelike "blastema" from coelomocytes that form around the gut. From this blastema the central nervous system develops first, quickly joining with the old ventral nerve cord. The digestive, muscular, and excretory systems develop later, and the final closing of the wound in the parent follows separation. The information on the clusters of lateral buds reported in a few worms is incomplete, and there appear to be minor differences in the sequence of events.

Regeneration Although it is not a mode of reproduction, regeneration is a developmental process. Neuroblasts that function in regeneration have been identified between the two lateral strands of the ventral nerve cord in Golfingia elongata (Akesson, 1961a). Removal of the distal centimeter of introvert from Siphonosoma cumanense was followed one day later by the closure of the cut end by a blastema (Kido and Kishida, 1961). New epithelium grew from the old epithelium as a network over the blastema. Four cell types became evident. Although it proved impossible to follow their subsequent development, two seemed to be coelomic cells. By day 5 a new mouth had formed; muscle layers were partly differentiated by day 6. At this time the epithelium was in good order but no defined cuticle was visible. The nerve cord regeneration string in Phascolion strombus has electrondense granules with diameters of 5000 A (Storch and Moritz, 1970). When the introvert was amputated, the granule-bearing cells migrated anteriorly

Regeneration

311

to form a clublike mass of cells rich in glycogen and lipids. The inclusions were extruded as fibers in the interstitial spaces, and a new cuticle was formed from secretions of these cells. New epithelial cells rich in rough endoplasmic reticulum developed, and amoebocytes produced muscle tissue. This evidence substantiates observations dating back to the late 18oos (BUlow, 1883) on the ability of sipunculans to regenerate lost or removed parts, particularly the introvert, within a few weeks. This regeneration was observed in Phascolion strombus, Nephasoma minutum (Schleip, 1934a, 1934b), Aspidosiphon muelleri, Golfingia vulgaris, Phascolosoma granulatum, and Sipunculus nudus (Wegener, 1938). All but one species replaced the distal end of the introvert. As in so many other areas, the exception isS. nudus, which appears to lack this ability. If a section of the introvert was removed from a worm with a partly retracted introvert, however, the missing segment could be replaced, reconnecting the original head to the body with a new "neck." A repeat of this experiment would be useful since these results are so unlike those reported for other sipunculans. Wegener and Schleip (cited above) observed regenerative tissue associated with the ventral nerve cord. Damaged or cut posterior ends can also be regenerated, although the success rate is higher when the intestine is not involved (Andrews, 189ob; Spengel, 1912; Schleip, 1934a, 1934b).

Part Ill 16

Zoogeography and Evolution

Zoogeography

This chapter gathers together what is known about sipunculan endemism and centers of cladogenesis, including both data from the literature and my own assumptions. Chapter 17 applies this information to the individual genera, and Chapter 18 combines what is known about sipunculan evolution and phylogenetic relationships with a historical overview of the world's oceans in an attempt to weave the threads of our knowledge into the multicolored tapestry of sipunculan evolution through geological time. To paraphraseR. Scheltema (1989), the contemporary spatial distribution of sipunculan species is limited by their ecological history and by past accidents, among other factors. We can explain some of these variables (see Chapter 19), but much is still unknown.* What follows here is descriptive and general. More detailed analyses are planned for the future.

The Quality of the Database The current picture of sipunculan distribution is fuzzy and full of holes, rather like an unfinished Impressionist painting. We have some idea of the overall pattern of sipunculan distribution, but only a few areas have been examined in sufficient detail to engender confidence about the subject. This is the inevitable result of nonuniform, nonrandom sampling by oceanographic expeditions and marine biologists, and it is true of many benthic marine invertebrates, especially the smaller, soft-bodied infaunal taxa. Nor is it only deep-water habitats that have been incompletely sampled. The

* At the time of this writing, a database of specific collection locations of sipunculans that includes latitude, longitude, depth, date, source, and species name is being compiled in the DOS-compatible dBASE III Plus format. Interested readers may request copies from the author.

314

Zoogeography

shallow waters around much of South America, Indonesia, and the Philippines are poorly known as well. When one states where a species lives, what is actually being described is where that species has been collected. If an area has been thoroughly sampled, it is fairly safe to discuss species distributions in that area. To expand from the secure base of a particular bay or transect to describe distributions in the entire world, however, is to leap into partially unknown space. Having stated this, a leap of faith (assuming an ordered world) will now be undertaken.

Species Value In past zoogeographical discussions, equal weight has been given to two rather different kinds of species: (I) taxa regularly collected over decades by more than one biologist and represented by many specimens, and (2) species known from only one or two individuals from a single location and reported by only one biologist. Thus, the available database contains two types of "endemic species." The first type might be called "tested, actual endemics"; that is, they are species whose endemicity has been tested at least once subsequent to the original observation. In addition one might include in this category species that have been reported only once, but by an experienced systematist who understands the biological species concept (i.e., who appreciates that variation is possible within a sipunculan deme), and are based on a significant number of individuals (i.e., more than three or four). The second type, "untested, potential endemics," are species that have been reported only once and are based on fewer than five worms, or species reported by a person lacking experience with sipunculans or one working within a typological species framework. When species in this category are included in zoogeographical analyses, an inflated and false impression of the number of endemic taxa can result. The diminished significance of untested endemics in this analysis and my inclusion of recent taxonomic revisions (resulting in about half the putative species being reduced to junior synonyms) have made the outcome of the present analysis different from earlier ones (Selenka et al., 1883; Herubel, 1903a, 1907; Murina, 197Ic, 1975a; Amor, 1975d; E. Cutler, 1975b).

Endemism and Centers of Origin

315

Endemism and Centers of Origin An endemic taxon is one that lives only in a circumscribed area. The size of the area is arbitrarily defined by the investigator, and marine areas have historically been much larger, with less well defined boundaries, than terrestrial ones (Kay, 1979). Since the days of Darwin and Wallace, biogeographers have used the existence of a high percentage of endemic species in a given area to support the thesis that such an area is a "center of origin." This idea is based on the assumption that land masses have always been where they are today. Previous analyses of sipunculans seem to have implicitly made the same assumption. While the phylum Sipuncula undoubtedly originated in a particular place, it would be a mistake to assume that all sipunculan taxa originated in that same place. It is more useful to think in terms of many centers of origin, including origins of families or genera, not just species. One example of a group with no single center of origin but with rapid dispersal from several different places is the Cenozoic benthic foraminifera of North American waters (Buzas and Culver, 1986). One must also avoid thinking that the world's oceans and land masses in Paleozoic and Mesozoic times, when most of the higher sipunculan taxa came into existence, were like they are today. Three brief examples will serve to illustrate this point (Chapter 18 discusses the issue in greater detail). (1) Although the phylum Sipuncula has existed for 500 million years, the Caribbean has been separated from the Pacific for only the past 3·5 million years (less than I% of the time). (2) The present boundary area between the Pacific and Indian oceans (the Indo-Malayian region) was established less than 20 million years ago (4% of the time the phylum has existed). (3) An Atlantic Ocean extending from Arctic to Antarctic waters did not exist before some 6o million years ago. The work of historical biogeographers thus includes a consideration of time as well as temperature, depth, current direction, and sea level. What is continental shelf today may have been dry land several thousand years ago. What is temperate water today was tropical a few million years ago, and so on. Areas rich in species and including many endemics today may be rather new habitats that are not historically significant as centers of origin or distribution. Alternatively, species-rich areas may represent remnants of suitable habitats that were once much larger, as demonstrated by the post-Miocene

316

Zoogeography

reef corals of Australia and New Zealand (Fleming, 1978). The cor&l, sea grasses, and associated species extended over much larger areas during the Cretaceous when the Tethys Sea existed. The current pattern is the result of extinctions and range shrinkages caused by abiotic and biotic factors (Knox, 1978). The same scenario has been documented for marine crustaceans in the Southern Hemisphere (Newman, 1991). After examining five possible hypotheses, including vicariance events associated with plate tectonics that are relevant for terrestrial and freshwater fauna (e.g., Mayr, 1988, for birds), Newman concluded that the endemic forms are post-Mesozoic relicts of either formerly widespread tropical taxa or remnants left over after the extinction of the northern portions of amphitropical (i.e., occurring on both sides of the equator) barnacle species. Additionally, disjunct distributions may be the outcome of widespread local extinctions resulting from changing conditions such as periodic cooling during Pleistocene glaciations (Fleming, 1978), decline in primary productivity after the Miocene (Vermeij, 1989), or the demise of the Tethys Sea in the Paleogene (Kay, 1979). The likelihood of vicariance events being entirely responsible for bipolar or antitropical disjunct distributions was discounted by Lindberg (I 991 ), who supported multiple mechanisms, including the probability of several biotic interchanges between the hemispheres during times when the tropics were less of a thermal barrier (i.e., during Neogene glaciations). There are other explanations for existing patterns of endemicity as well, especially for marine taxa without a fossil record. Without entering into the debate about the linkage between centers of endemism and centers of origin (see Knox, 1978), I judge the concept to be of such dubious value for sipunculans that I do not use it here. Dispersal, Boundaries, and Biogeographic Units Most sipunculans are capable of dispersing their planktonic larvae over hundreds or thousands of kilometers fairly quickly, and it is thus possible for these infaunal worms to be distributed over large areas. This makes the zoogeographical boundaries of a species's range difficult to define and delimit (Rice, 1981; R. Scheltema, 1975, 1986a, 1986b, 1988; R. Scheltema and Rice, 1990). Most often, ranges appear to be determined by water temperature, but bottom topography and water currents are important fac-

Endemism and Centers of Origin

3I7

tors, especially at bathyal and abyssal depths, where water temperature varies little with latitude. The importance of bottom topography and currents is evident at various points on the Atlantic continental slope but is best documented off Cape Lookout, N.C. (E. Cutler, I968b, I975a; E. Cutler and Doble, I979; E. Cutler and Cutler, I987b). It is difficult to generalize about sipunculans' dispersal ability because some taxa travel much greater distances than others. Although it may be true that the east Pacific barrier (EPB) is the most effective obstruction to the dispersal of contemporary shallow-water tropical fauna, it is not impermeable. Grigg and Hey (I992) found 4% of Pacific reef-building corals and I4% of the molluscs to be amphi-Pacific. It takes free-floating larvae 55-70 days to cross the shortest distance (Christmas Island to the Gahipagos) and up to I55 days to cross elsewhere, and most coral larvae do not live that long. While the EPB is an effective filter barrier (R. Scheltema, I986b), I I shallow-water sipunculan species from seven genera are known to be amphi-Pacific (Sipunculus polymyotus, S. phalloides, S. nudus, Siphonosoma vastum, Phascolosoma (Edmondsius) pectinatum, P. nigrescens, P. perlucens, Antillesoma antillarum, Apionsoma misakianum, Aspidosiphon (Paraspidosiphon) coyi, Lithacrosiphon cristatus). Many more species are present in the western but not the eastern Pacific. Sipunculans do not actively migrate; rather, their larvae are passively transported within the currents that form the highways of the sea. Asymmetrical invasions brought about by passive transport in currents that are largely (but never entirely) unidirectional have been noted in several of the world's oceans; for example, from the Red Sea into the Mediterranean, from the North Pacific to the North Atlantic via the Arctic, from east to west in the North Atlantic, and from west to east in the tropical Pacific (Vermeij, I99Ia, I99Ib). These are generalizations, and there are exceptions. For example the near-shore flora in the Arctic Ocean does not follow the same pattern as the fauna (Dunton I992). More than I20 species of Red Sea marine organisms (plants and animals) have colonized the eastern Mediterranean Sea since the Suez Canal opened in I 869, but migration in the reverse direction has been limited to IO species. This is largely a reflection of the direction and rate of current flow at the time of year when reproduction occurs in many Red Sea species (Agur and Safriel, I98I). I should point out, however, that Por (I975) found no sipunculans or sipunculan larvae in the Suez Canal and

318

Zoogeography

strongly suggested that dispersal of sipunculans along this route was very unlikely-in either direction-because of the extreme euryosmotic habitat and lack of hard substrate for rock-boring taxa. R. Scheltema (1992) set forth several arguments to put to rest the confusion about the role of passive dispersal in determining the present distribution of benthic invertebrates (also see Kay, 1979). For example, the old generalization that most larvae have fixed and short lives was disproved by Scheltema's laboratory and field work. Sipunculan teleplanic larvae retain their competency to metamorphose for long periods (two to six months), and dispersion is not a random process. It occurs along distinct corridors formed by the major current systems. One outcome of this method of dispersal is the widely different degrees of endemism between terrestrial and marine island-dwelling invertebrates (high for the former, low for the latter). Finally, Scheltema challenged the assertion made by vicariance biogeographers that since species distributions are nonrandom, one cannot invoke dispersal processes to explain them (Nelson and Platnick, 1980). This idea is based on the assumption that dispersal is a random process; but clearly it is not. The three-dimensional nature of the oceans adds to the complexity of any analysis. To place a species that has been reported from 88o m off New Zealand (Phascolosoma saprophagicum) on the list of Indo-West Pacific endemics along with a species from intertidal coral off Queensland (Themiste variospinosa) is to ignore significant ecological differences. Adding to this list a species from shelf water in the Red Sea and northwestern Indian Ocean (Sipunculus longipapillosus) means ignoring the gap of thousands of kilometers between them. Such combined lists lose zoogeographical meaning. Meaningful units for analysis must be small, in both vertical and horizontal dimensions-but how small? One cannot neatly separate the shallow from the deep-water species, or warm-water from cold-water taxa. How cold is cold, and how deep is deep, and should one have separate categories for cold-shallow and cold-deep species, etc.? Clearly, marine worms cannot be made to fit into discrete analytical sets as can freshwater or island-dwelling organisms. Subdividing the world's oceans into quadrants or cubes for numerical (i.e., vicariance) analyses is artificial and subjective. Ekman (1967) and Briggs (1974) presented broad pictures of marine zoogeography, but to do so they had to dissect the oceans along arbitrary and ill-defined thermal,

Endemism and Centers of Origin

319

vertical, and horizontal lines. At this writing I find no justification for submitting sipunculan data to this type of segregation. A good reason not to do so is provided by the striking difference between the sipunculans and the other taxa in the Hawaiian Islands. Ekman set these islands apart from the Central Pacific Island subregion based on the large number of endemic species, but there are no endemic sipunculans in this archipelago; most of those present have very wide ranges. The Mediterranean Sea is another example of a restricted region with no endemic sipunculans despite the presence of many endemic invertebrates of other taxa (e.g., the 15 endemic chalimid sponges; see de Weerdt, 1989). Looking at only the shallow-water species in an area, as is so often done, biases the data and could mask valuable evolutionary information. Given all these factors, no formal analysis of endemic species is presented here. Cosmopolitan Species If endemic species are at one end of a zoogeographical continuum, then cosmopolitan species are at the opposite end and are worthy of a brief comment. Taylor (1977), who studied Cambrian trilobites, questioned whether widespread species are widespread because their tolerances are broad (eurytopic) or because suitable habitats are readily available and cover large areas. The eight sipunculan genera with more than six species all contain one to three species that are much more widespread than their congeners. In some genera these species live in habitats typical of the genus: the widely dispersed Nephasoma species all live in cold water; Phascolion strombus is generally confined to cold water; and Siphonosoma cumanense and the three most cosmopolitan Phascolosoma species typically live in warm, shallow habitats. In several cases, however, cosmopolitan species are very eurytopic and live in habitats not typical for the genus as a whole. Widespread species that live in water cooler than most species in their genus prefer include Sipunculus nudus, Phascolosoma stephensoni, Aspidosiphon zinni, and A. muelleri. Alternatively, a few extend into warmer than "normal" habitats; for example, Golfingia margaritacea and Themiste lageniformis. The importance of cosmopolitan species in the evolutionary framework is uncertain. One could argue that they represent ancestral species that gave rise to descendants as they spread around the world, or that their

320

Zoogeography

genes give them a selective advantage. In this phylum it appears that the latter is more often the case in species that live in normal or cooler than normal habitats. For reasons I will present later, however, the pair of species that live in warmer than normal habitats may well represent ancestral taxa. Finally, it is possible that more than one species is present within some very large populations. Cryptic and sibling species exist in other taxa, and there is no reason to assume that there are none in the phylum Sipuncula. Until appropriate objective tests are applied to such taxa, however, the true significance of cosmopolitan species must remain on the list of unsolved mysteries.

17

Generic Analyses: Distribution Summary and Cladogenesis

The analyses presented in this chapter depend on various ad hoc explanations, including dispersal and local extinctions. I have not attempted a vicariance analysis, both for the reasons stated in Chapter 16 and because there is a dearth of species-level cladograms. Given our current limited array of attributes, a severe shortage of synapomorphies exists. Someday this material should be approached again, when more diverse data are available. The analyses in this chapter include untested endemics (UE, as defined in Chapter 16), but these are identified as such. These are of doubtful zoogeographical significance and should be omitted from more quantitative analyses.

Family Sipunculidae Sipunculus and Xenosiphon

Of the 13 included species and subspecies, 6 (46%) live in shallow, warm waters of the western Atlantic, Caribbean, or eastern Pacific-the Atlanto-East Pacific (AEP) of Ekman (1967). Four are endemic there: S. marcusi, S. phalloides phalloides, S. polymyotus, and X. branchiatus. Only 2 species are also found outside this area: S. nudus is circumsubtropicalwarm temperate, and S. robustus occurs in the tropical Indian and west Pacific oceans (IWP). Most of the remaining shallow warm-water taxa are quite restricted in distribution. S. longipapillosus is known from the northwestern Indian Ocean and the Red Sea, and S. phalloides inc/usus is from Indonesia and southern Japan. S. (Austrosiphon) indicus is widely distributed in the IWP, X. absconditus is scattered from the Red Sea across to the western Pacific, and S. (A.) mundanus mundanus is limited to the western Pacific. There-

322

Distribution and Cladogenesis

fore, there are seven species or subspecies (54%) in the IWP-five endemic to that region. The two deep cold-water species, anomalies in this family, are S. norvegicus (100-3000 m) and S. lomonossovi (2500-4300 m). Both occur in the North Atlantic and (less commonly) elsewhere, but not in the eastern Pacific, southern Atlantic, or Antarctic. Only the former extends into the Indian Ocean. Center of Cladogenesis. Significant cladogenesis seems to have occurred in two places: the shallow Indo-Malayian region during the early Cenozoic, for a part of the nominate subgenus and S. (Austrosiphon); and, during the later Cenozoic, the American Tethys Sea, which subsequently became the Caribbean-tropical eastern Pacific. This area was important for several members of the nominate subgenus, possibly Xenosiphon, and, in its deeper part, the cold-water Sipunculus species. Siphonosoma

Only one of the 10 shallow-water species is broadly distributed: S. cumanense is almost circumtropical but apparently absent from the eastern sides of the Atlantic and Pacific oceans. Three species have been reported from several localities in the IWP (S. australe, S. rotumanum, and S. vastum), and the last has also been collected on the Pacific coast of Costa Rica. In the western Pacific, S. funafuti and S. boholense have been found at only a few locations. The four remaining species live in very limited areas of temperate shallow water: S. arcassonense (France and Spain), S. ingens (California), S. mourense (central Japan), and S. dayi (Natal, South Africa). The first two species are separated from the other Siphonosoma species by thousands of kilometers, and they are the only two members of the genus whose ranges do not overlap with S. cumanense. They are also the only two sipunculans with the enigmatic fusiform bodies. An analysis of regional endemism in this genus quickly shows the IWP to be favored, with five of the ten species confined to this region. None are restricted to the AEP; only two widely dispersed species live there. Center of Cladogenesis. Siphonosoma clearly underwent most of its radiation in the central IWP. The number of cool-water endemic species (one each in Japan, California, and France) together with the possible descendant Phascolopsis, from the northeastern coast of the United States, is intriguing. A plausible scenario is the following: Each species could be descended from a single widespread polymorphic ancestor which lived in

Family Golfingiidae

323

northern latitudes during the Miocene or Pliocene when the water was considerably warmer. Speciation could have occurred as eurythermal remnants of this hypothetical population were left behind as the main thermophylic Siphonosoma stock was forced southward along with the warm water. As these ancestral demes were probably small, genetic drift may have played an important role. These new allopatric populations would have responded to different selective pressures that were working on different gene pools. Siphonomecus and Phascolopsis

These genera are monotypic, and each has a very restricted range in the western Atlantic. S. multicinctus is known from the southeastern United States (South Carolina to western Florida). P. gouldii lives along the Atlantic coast of Canada and the northeastern United States (rare from Long Island to northern Florida). Center of Cladogenesis. No evident speciation has occurred within these genera. It is safe to assume that these two taxa originated in the western Atlantic where they presently reside, one in cool-temperate and one in subtropical water. It is possible that both taxa evolved from a nowextinct Siphonosoma population after the closing of the Panamanian land barrier in the Pliocene.

Family Golfingiidae Golfingia

Most species in the nominate subgenus inhabit cold subtidal water at depths of 2-6800 m. The unique G. (Spinata) pectinatoides is very different: it lives in tropical coral sands in the IWP. A similar habitat is occupied by G. vulgaris herdmani in shallow Indian Ocean waters. A third exception is the least derived member of its subgenus, G. (G.) elongata, because some populations live in intertidal warm-temperate waters. Of particular interest is the common occurrence of two members of this genus (G. anderssoni and the endemic G. margaritacea ohlini) in the far southern seas at latitudes between 45 and 75° S. Less commonly one also finds G. margaritacea margaritacea and G. muricaudata in the far south. Two endemic species are scattered over the northeastern Atlantic (G. iniqua) and South Africa (G. capensis). Two species based on single records

324

Distribution and Cladogenesis

(untested endemics) come from East Africa (G. mirabilis) and northwestern Pacific (G. birsteini). Eight of the 12 species and subspecies (67%) are found in some part of the Atlantic Ocean. The four non-Atlantic taxa are G. mirabilis (UE), G. birsteini (UE), G. vulgaris herdmani, and G. (S.) pectinatoides (one subspecies, one species from the non-nominate subgenus, and the two untested endemics). The five species in the Indian Ocean seem to be restricted to the eastern, southern, and western borders and are absent from the central and northern parts. The Pacific is home to nine Golfingia taxa (75%). The three that are absent (G. capensis, G. iniqua, and G. mirabilis [UE]) have very restricted ranges in the Atlantic or the western Indian Ocean. As is the case in other genera as well, the deeper-water taxa appear to be more widely dispersed; 6o% of those collected more than once are found in both the Atlantic and Pacific oceans. The fact that Golfingia species live at greater depths in the lower latitudes (known as equatorial submergence) has led some biologists to suggest that some species are bipolar in distribution; however, this may be only an artifact of the difficulty of collecting low-density populations at great depths. For some species, amphitropical would be a more descriptive term than bipolar, since they are more common at middle latitudes than at polar ones. Center ofCladogenesis. It is likely that the ancestral subgenus G. (Spinata) originated in the warm, shallow Panthalassa (precursor of the Pacific, also called the Eo-Pacific) during the early Paleozoic. This taxon produced the nominate subgenus, which appears to have used the far southern seas as a major center of cladogenesis and then probably radiated northward, following the spread of cold water during the late Mesozoic and Tertiary. The absence in the southern and central Pacific Ocean of widespread species such as G. margaritacea and G. muricaudata poses a problem, but this can be explained by local extinctions. Alternatively one would have to assume a northward dispersal route through the Atlantic, across the Arctic (against the prevailing surface currents; perhaps they were benthipelagic larvae?) to the North Pacific. Nephasoma Six of the 27 species and subspecies are known from single reports, including N. abyssorum benhami, N .filiforme (UE), and N. tasmaniense

Family Golfingiidae (UE), plus 3 based on single worms (N. laetmophilum [UE], N. multiaraneusa [UE], and N. vitjazi [UE]). Since most of these are untested endemics, their very limited ranges should not be seriously considered. Despite the fact that 13 "tested" taxa are restricted to one ocean, there does not appear to be any one center of endemism. The richest fauna is in the Atlantic, with 16 taxa (59%). Seven of the 16 live only in northern latitudes, and 6 are endemics: N. bulbosum, N. lilljeborgi, N. minutum, N. multiaraneusa (UE), N. rimicola, and N. wodjanizkii elisae. The seventh North Atlantic species, N. constrictum, has recently been recorded from the southwestern Indian Ocean. None of the other 9 Atlantic taxa is restricted to southern waters, although N. constricticervix has not been collected outside the Atlantic. Five taxa (N. abyssorum abyssorum, N. capilleforme, N. diaphanes corrugatum, and N. eremita) live throughout the Atlantic and in the Pacific, and 3 (N. confusum, N. diaphanes diaphanes and N. pellucidum pellucidum) are found in these two oceans plus the Indian Ocean. Of the 13 taxa living in the Pacific Ocean (48%), 6 are endemic there (22%): N. cutleri, N. laetmophilum (UE), N. novaezealandiae, N. pellucidum subhamatum, N. vitjazi (UE), and N. wodjanizkii wodjanizkii. The remaining 7 species also occur in the Atlantic Ocean. Only 7 species (26%) have been collected in the Indian Ocean, but 4 of these are endemic: N. (Cutlerensis) rutilofuscum, N. filiforme (UE), N. schuttei, and N. tasmaniense (UE). The first is from the African margin (and the non-nominate subgenus), and the other 3 are from the AustraliaIndonesia region (only one tested endemic). Two Nephasoma live in the Antarctic, N. confusum and N. abyssorum benhami (the latter is found nowhere else). The Arctic is home for 3 taxa, but all are common in the North Atlantic and elsewhere as well. It is clear that Nephasoma is the deep-water genus, with 9 species occurring at depths greater than 4000 m and 21 (78%) at depths greater than 1000 m. Of the 6 remaining species, 3 have been collected only once, but 3 (N. minutum, N. rimicola, and N. schuttei) are clearly intertidal and shelf species. A few eurybathyal species fit both catagories: N. (C.) rutilofuscum, 1-1500 m; N. pellucidum, 1-1600 m; N. confusum, 4-4300 m; and N. eremita, 20-2000 m. Center of Cladogenesis. The species richness in the deeper parts of the northern Atlantic and Pacific oceans point to these cold bathyal and abyssal waters as areas of rapid speciation. Despite the probability that most speciation must have occurred after the oceans deepened and cooled in

Distribution and Cladogenesis mid-Tertiary times, the genus probably originated in the Permian or Triassic. Thysanocardia

The three species in this genus appear to have nonoverlapping ranges in cool shelf and upper slope waters. The most restricted is T. procera, which is found only in the northeastern Atlantic. The others are more broadly dispersed: T. catharinae in the rest of the Atlantic Ocean, off East Africa, and off Peru (Murina's 1989 Vietnam record may be a new species); and T. nigra in the northern and western Pacific Ocean. No records of this genus exist from Australian waters. Center of Cladogenesis. This primarily northern genus probably originated in the North Atlantic during the late Tertiary. It underwent limited cladogenesis as it spread, perhaps westward via the Tethys Sea, or possibly via subsurface Arctic currents. The reverse (Pacific origin and migration into the Atlantic via the Arctic) is not unthinkable.

Family Phascolionidae Phascolion

Three species taxa have significant populations in all three of the world's oceans, P. strombus strombus being the most widely distributed and eurytopic. Two deep-water species, P. (Montuga) lutense and P. (M.) pacificum, are close seconds throughout the northern and southern Atlantic and Pacific, but they have been found only in the southern Indian Ocean (i.e., they are absent from low latitudes). An additional three species (P. (lsomya) hedraeum, P. (Lesenka) hupferi, and P. (/.) tuberculosum) occur less broadly in both the Atlantic and Pacific oceans. In addition to the common and widespread occurrence of a few species in the Atlantic Ocean, 6 of the 13 species residing there are endemic along the western side, and these 6 have all been described since 1972: P. (L.) cryptum, P. (/.) gerardi, P. medusae, P. (/.) microspheroidis, P. psammophilus, and P. caupo. The last species also lives in the northeastern Atlantic. Surprisingly, 20 taxa (70%) have been recorded from the IWP, and 12 (43%) are endemic: P. hibridum, P. (/.) lucifugax, P. pharetratum, P. (L.)

Family Phascolionidae

327

rectum, P. strombus cronullae, P. ushakovi (UE), P. (L.) valdiviae summatrense, P. valdiviae valdiviae, P. abnorme, P. (Villiophora) cirratum, P. megaethi, and P. robertsoni; the last 4 live only in the western Indian Ocean and Red Sea. Of the 8 nonendemics, 7 extend into the Atlantic Ocean in deeper water, and I, P. (/.) convestitum, only reaches into the Mediterranean Sea. Aside from the three widespread eurytopic deep-water species noted above, the eastern half of the Pacific Ocean is almost devoid of Phascolion. The single specimen of P. bogorovi (UE) collected from the Peru-Chile Trench is the only one known. Waters near the Indian subcontinent also appear to be unsuitable for Phascolion. This genus was once portrayed as a cold- and deeper-water taxon, but recent data disprove that notion. Almost equal numbers of taxa (14 and 12, respectively) live in shelf waters (1-300 m) and deeper water. Of the 14 shelf taxa, 7 may be intertidal but the remainder have not been collected at depths less than 15m. Six taxa are known from both shelf and continental slope depths (300-3000 m), including P. (/.) hedraeum (7-46oo m) and the eurytopic P. strombus strombus (1-4030 m). Six taxa are known only from slope and deeper waters (300-6900 m), but only P. (M.) lutense and P. (M.) pacificum occur in significant numbers at abyssal depths (>4000 m) as well as on the slope. E. Cutler and Cutler (1985a) described a possible rassenkreis ("race circle") in P. strombus. Certain Japanese populations exhibit two morphs that differ in holdfast shape, hook size, and origin of the ventral retractor muscle. These character states are within the range of variation seen in the diverse North Atlantic population but are at the two extremes of the continuum. Assuming a center of origin in the North Atlantic, it is possible that one population dispersed eastward over the Siberian-Asian Arctic, and a second went westward over the Canadian Arctic, and the two met in the North Pacific, where the two ends of the circle came into contact. It seems clear that gene frequencies shifted as semi-isolated populations spread around the globe, and the Japanese forms may be genetically isolated, although this remains to be tested. An alternative hypothesis is that the North Pacific populations are actually two species that both migrated to the North Atlantic, where they exist as a "superspecies" or a group of cryptic species not yet differentiated. Center of Cladogenesis. If one analyzes this genus by subgenus, the only thing that becomes evident is that the western Pacific Ocean is inhabited by representatives of all subgenera except the most derived-the

Distribution and Cladogenesis monotypic P. (Villiophora). The widespread species are in cold, usually deep, water and are generally absent from lower latitudes. The areas of endemism here are simply too large to be meaningful (e.g., the western Atlantic or IWP). When areas of analysis are defmed more narrowly, scattered clusters of endemism appear along the western side of the Indian, Pacific, and Atlantic oceans. Thus it seems that there are multiple illdefined centers (or fragmentation of a broad Panthalassa population) for this diverse genus, which has probably existed since late Paleozoic times.

Onchnesoma Four of the six species and subspecies live in the Atlantic Ocean. The North Atlantic appears to have the largest populations, although all except 0. squamatum squamatum are also found in southern latitudes, and 0. steenstrupii nuda is known only from one southeastern Atlantic area. The only known extension outside the Atlantic, by the deep-water 0. magnibathum, is a record from the Peru-Chile Trench. The most widespread member of this quartet, 0. steenstrupii steenstrupii, has been collected throughout the Atlantic and in the southwestern Indian and southwestern Pacific oceans. The two taxa not known from the Atlantic (0. intermedium and 0. squamatum oligopapillosum) are represented by very few individuals and are very close to being untested endemics in the northwestern Pacific Ocean. In terms of preferred depths, four species live along the continental slopes (100-2000 m), one is a shelf species (15-250 m), and one is abyssal (3000-5500 m). Center of Cladogenesis. The cold waters of the North Atlantic appear to be the ancestral home of this genus, dating back to the Paleogene. Subsequent speciation probably occurred here also, spreading via the high latitudes into neighboring waters.

Family Themistidae Themiste Four of the six T. (Lagenopsis) species live only in the Australian region, and two of the five T. (Themiste) species are limited to the eastern Pacific. Until recently, T. (Themiste) hennahi was a third eastern Pacific

Family Phascolosomatidae

329

species, but Haldar's (1991) record from Indian waters changed that species's pattern. Unlike the two other uncommon T. (Themiste), which have very limited distributions (T. blanda in Japan and T. alutacea in the westem Atlantic), the two uncommon T. (Lagenopsis) have broad distributions (T. minor minor is from the northwestern and southwestern Pacific and off South Africa, and T. lageniformis is circumtropical-subtropical). Ranges of the two subgenera overlap in three regions: ( 1) Honshu (central Japan), where T. (T.) blanda and T. (T.) pyroides are sympatric with T. (L.) minor; (2) the Caribbean and east coast of Florida, where T. (L.) lageniformis and T. (T.) alutacea coexist; and (3) the Nicobar Islands off India, where T. (L.) lageniformis coexists with T. (T) hennahi. Southern Argentina may be a fourth such place, but the database is too small to ascertain this. Another way to view this genus is as follows: all six of the T. (Lagenopsis) can be found in some part of the IWP (five are endemic), and three of the five T. (Themiste) are found only in the Pacific Ocean (a fourth is in the Indian also). Only two species (one of each subgenus) are found in the Atlantic Ocean; one is restricted to the western Atlantic and the other is nearly circumsubtropical. All Themiste live in intertidal or shallow subtidal water. Center of Cladogenesis. Each of the two subgenera appears to have its own center, one on each side of the Pacific Ocean, which suggests a Pacific origin for the genus. The cool Australian region is the center for T. (Lagenopsis), and the nontropical eastern Pacific coastline has been an active center forT. (Themiste). Since cooler south Australia did not detach from Antarctica until the Eocene, the genus probably postdates that time. Most speciations likely occurred after the Miocene.

Family Phascolosomatidae Phascolosoma

Six of the 19 Phascolosoma species and subspecies appear to have very restricted ranges, 4 in habitats unusual for this genus: P. meteori from the Red and Arabian seas (high salinity, low oxygen tension); P. turnerae from the Gulf of Mexico and off Australia (deep cold water, in wood or near cold-water seeps); P. saprophagicum from New Zealand (deep cold water, from rotting whale skull); and the cold-water P. agassizii kurilense from

330

Distribution and Cladogenesis

the far northwestern Pacific Ocean. In limited but more typical habitats one finds P. maculatum (UE) (Indonesia) and P. glabrum multiannulatum (Tahiti). Eleven of the remaining 13 Phascolosoma taxa, plus the two endemics (P. saprophagicum and P. maculatum [UE]) and P. turnerae, or 74% of all Phascolosoma species, occur in the border region between the Indian and Pacific oceans-the Indo-Malayan subregion. The species richness in this subregion is unparalleled by other sipunculan genera. The five species not found in the Indo-Malayan subregion include three endemics (P. meteori, P. agassizii kurilense, and P. glabrum multiannulatum [UE]) plus P. granulatum, which lives in the colder waters of the northeastern Atlantic Ocean and the Mediterranean Sea, and P. (Fisherana) capitatum, from bathyal waters of the Atlantic Ocean. A second noteworthy feature is that only three species of Phascolosoma live in the Caribbean basin: the restricted P. turnerae and the two circumtropical shallow-water species, P. perlucens and P. nigrescens. The latter two also occur in the central and southern regions of the eastern Atlantic along with P. agassizii agassizii and P. stephensoni. The last species extends through the IWP to Hawaii and also coexists in parts of the northeastern Atlantic with P. granulatum. In the eastern Pacific are found the two shallow-water Caribbean species just mentioned plus the cool-water P. agassizii agassizii and the warm-water P. scolops. These four are also found throughout the IWP. Five species do not extend outside the IWP: P. albolineatum, P. arcuatum, P. glabrum glabrum, P. pacificum, and P. (Fisherana) lobostomum; while P. noduliferum and P. annulatum are restricted to the boundary area between the Indian and western Pacific oceans. Ecologically, Phascolosoma tolerates wide extremes (this in addition to the unique niches of a few species known only from limited populations, noted above), ranging from the warm, very euryhaline mangrove mud habitat of P. arcuatum, through the euryhaline cold intertidal rocks of P. agassizii, to the cold, stenohaline Atlantic where P. granulatum and P. (F.) capitatum live. It is nonetheless true that most species select shallow warm-water habitats. Center ofCladogenesis. The Panthalassa precursor of the Indo-Malayan Archipelago seems to be where this genus originated and where most speciation occurred. The genus is probably of mid-Paleozoic age, but most extant species evolved in Cenozoic times, including the cold-water species.

Family Aspidosiphonidae

331

Antillesoma

The one species in this genus, A. antillarum, is found around the world in tropical and subtropical shallow-water habitats, generally in crevices or burrows in dead coral or other soft rocks. Center of Cladogenesis. Given such an extensive range and only one extant species, it is difficult to be certain, but A. antillarum probably originated in the Mesozoic Panthalassa. Apionsoma

The four species fall into two ecologically different subsets, which are reflected in their distributions. The two deep-water taxa (A. murinae murinae and A. murinae bilobatae) occur in the Atlantic and Pacific oceans at slope to abyssal depths (300-5200 m). The second taxon is also found in the Mediterranean Sea and on both sides of the Indian Ocean at slope depths only (2oo- 1200 m). The three shallow-water species (A. misakianum, A. trichocephalus, and A. (Edmondsius) pectinatum) are also widespread, but in shallow, warm waters. The ftrst is known from the Indian Ocean and both sides of the Pacific, but only the western Atlantic, including the Gulf of Mexico. The second co-occurs in warm-water sandy habitats over most of this range plus the eastern Atlantic Ocean. The third is less common but circumtropical and has been collected on both sides of all three oceans. The broad distribution of Apionsoma in both shallow and deep habitats may be indicative of great age. Center of Cladogenesis. The shallow Paleozoic Panthalassa probably saw the first members of this genus and, according to the hypothesis presented in the next chapter, the first members of the phylum Sipuncula. The deep-water taxa are undoubtedly more recent additions, with origins in the mid-Cenozoic Atlantic Ocean.

Family Aspidosiphonidae Aspidosiphon

Ten of the 19 species (63%) live in the western Atlantic Ocean and Caribbean Sea, bounded by Cape Hatteras on the north and the Amazon delta on the south: A. exiguus, A. gosnoldi, A. (Akrikos) mexicanus, A.

332

Distribution and Cladogenesis

(Paraspidosiphon) parvulus, A. (P.) fischeri, A. (Ak.) a/bus, A. elegans, A. (P.) laevis, A. (P.) steenstrupi, and A. misakiensis. The first four species listed are endemic to the region. The fifth also lives in the eastern Pacific (Panama to the Galapagos). The range of the sixth extends in the other direction, into the eastern Atlantic (Iberia to the Gulf of Guinea). The next three species are circumtropical, and the last is found on both sides of the Atlantic and in the western Pacific Ocean. Two species found in the eastern Atlantic and elsewhere do not live in the western Atlantic: A. (Ak.) venabulum from both sides of Africa, and A. muelleri (see below). A. (Ak.) zinni, the one bathyal-abyssal member of this genus, is also found in the north Atlantic (plus one record from the Mozambique Channel). A. muelleri, the most widespread species, is almost cosmopolitan in temperate to subtropical waters. Two apparent gaps are in the western Atlantic (one record off southern Brazil) and the eastern Pacific (one record off Chile). A. muelleri is also the most eurytopic member of the genus and lives in a wide variety of temperatures and depths. Six species (plus A. muelleri) are widely distributed within the IWP. A. (P.) coyi extends into the eastern Pacific Ocean. Three of the six are also found in the Caribbean (A. elegans, A. (P.) laevis, and A. (P.) steenstrupi). The remaining two do not exist in either Hawaiian waters or the Atlantic Ocean (A. gracilis gracilis and A. (P.) tenuis). Two species (A. [Ak.] thomassini and A. spiralis) are more restricted within the IWP, and A. (P.) planoscutatus (UE) is known only from a single collection (2 specimens) in the Red Sea. Finally, A. gracilis schnehageni is known only from the eastern Pacific Ocean. The roughly equal number of endemic taxa in the AEP (6) and the IWP (4, plus I UE) is noteworthy. Of the I9 taxa, I3 live somewhere in the Atlantic Ocean, I I occupy some part of the IWP, and 6 live in both areas. Although some common widespread members of this genus do bore holes in coral or soft rock (including the entire subgenus A. [Paraspidosiphon] plus one A. [Aspidosiphon]), I I species (58%) do not. These include all the members of the subgenus A. (Akrikos) and all but one A. (Aspidosiphon). The nonboring worms live in empty mollusk shells (8), arenaceous foraminiferan tests (I), or interstitially (2). Most species live between the intertidal zone and the edge of the continental shelf (200m). Center of Cladogenesis. The data suggest two centers of origin and speciation: the IWP and the tropical AEP. The Paleozoic Panthalassa saw the origin of the genus in the form of the ancestral subgenus, A. (Aspi-

Family Aspidosiphonidae

333

dosiphon). Although not obvious, much speciation probably occurred in the Mesozoic Tethys Sea and the Cenozoic IWP. The IWP region was clearly the site of the late Mesozoic origin and Cenozoic speciations in the subgenus A. (Paraspidosiphon). The late Cenozoic warm-water Atlantic was the center for the other derived subgenus, A. (Akrikos). Cloeosiphon and Lithacrosiphon The three species of these two genera are tropical, shallow warm-water, coral-boring worms whose ranges overlap in the western Pacific islands. Cloeosiphon aspergillus is widely distributed in the IWP, from East Africa across the Indian Ocean to northern Australia and from many western Pacific islands west of Hawaii. From this shared space in the western Pacific, the more common Lithacrosiphon (L. cristatus) is found eastward to the eastern Pacific and into the Caribbean. A new subspecies, L. cristatus lakshadweepensis {Halder, 1991), was recorded from the far northwestern comer of the Indian Ocean in the Arabian Sea. The other species (L. maldivensis) fills the gap because its range westward is more continuous across the Indian Ocean into the Red Sea, generally far from continental land masses. Center of Cladogenesis. The mid-Cenozoic Indo-Malayan Archipelago appears to be the place of origin of these taxa.

18

Evolution and Phylogenetic Relationships

Direct Evidence: The Fossil Record Despite recent work on the Ediacarian and Burgess shale faunas (Cloud and Glaessner, 1982; Collins et al., 1983), we have no definitive fossil sipunculan or any fossil that is an acceptable ancestor (Morris, 1985). The best candidate for a sipunculan ancestor among the wormlike Burgess shale fossils may be Ottoia prolifica (Banta and Rice, 1976), but this muddwelling, bilaterally symmetrical worm with a retractable proboscis is not ascribable to any extant phylum. Its posterior anus, posterior ventral hooks, and rows of anterior hooks and spines make it more like the Aschelminthes or Priapulida (Morris, 1989). Another possible ancestor is Hyolitha from the Cambrian of Antarctica and the Ordovician of France. Although this animal has molluscan attributes such as a calcareous cone-shaped exoskeleton and an operculum, the digestive tract with both mouth and anus at the anterior end, the body wall with both circular and longitudinal muscle layers, and a hydrostatic skeleton to evert the "head" of the animal are sipunculan-like. It is possible that this extinct group coexisted with Paleozoic mollusks and sipunculans and that all three taxa shared a common Pre-Cambrian ancestor (Runnegar et al., 1975). One way to address the difficulties of assigning extinct forms such as these to formal taxonomic categories is to create superphyla for metazoan coelomates such as those proposed by Valentine (1973). His system focuses on coelomic architecture, and his group Sipunculata includes unsegmented infaunal burrowers with introverts, which probably fed on detritus. Valentine's five superphyla-Sipunculata, Molluscata, Lophophorata, Deuterostomia, and Metameria-are viewed as ancestral to all modem coelomate phyla. Willmer (1990) supported this approach and favored retaining the Sipuncula as a separate higher taxon because it is monomeric and has no clear links to other protostome taxa.

Direct Evidence

335

The holes made by sipunculans seem to have fossilized much better than the worms themselves did. The ichnogenus Trypanites is absent from Cambrian hard-ground surfaces in Montana, but it is represented by macroborings from the late Cambrian of Labrador and by well-preserved fossil burrows from many Ordovician, Silurian, and Devonian locations. (An ichnogenus is a genus based on traces, such as fossilized burrows, rather than on fossils of the animals themselves. The Greek prefix ichnos means "footprint" or "track.") It is possible that the Cambrian holes were made by a separate group of organisms that became extinct along with their host reef-building organism (archaeocyathid) and that a second group of organisms that produced a very similar burrow appeared in the early Ordovician (Brett et al., 1983). The similarity of fossil borings to those made by modern sipunculans suggests that sipunculans were present by the mid-Paleozoic (Pemberton et al., 1980). Coral assemblages containing coral-boring sipunculans are known from Upper Jurassic, Miocene, Pliocene, and Pleistocene times (Hyman, 1959; Pisera, 1987). The Montana-Wyoming Cambrian sediment contains small, slightly tapered holes that appear to have been made in semilithified micrites (finegrained sediments) prior to their deposition as clasts. It is possible that these holes were produced by "precursors of organisms . . . capable of excavating truly indurated sediment" (Brett et al., 1983:288). The sipunculan origin of these ancient burrows is supported by recent evidence that very similar Quaternary deep-sea burrows (Zoophycos) along the northwest African and Norwegian continental slopes were made by sipunculans belonging to the genus Nephasoma. These lebensspuren correspond to older burrows such as the upper Cretaceous ichnogenus Trichichnus and the Jurassic ichnogenus Ancorichnus from Denmark, both of which may be of sipunculan origin (Wetzel and Werner, 1981; Frey et al., 1984; Romero-Wetzel, 1987). Trichichnus was also reported from the Miocene in Italy by McBride and Picard (1991). The fossil burrows were most common in claystones but also present in sandstone formations. The burrows averaged 0.13 mm in diameter, were up to 160 em long, and were spaced 0.4-50 em apart. McBride and Picard's analysis suggested that the creators of these burrows had a high tolerance to low in situ oxygen levels. There are a few thin deep-water members of the genus Nephasoma that could have constructed burrows with these dimensions. Other possible sipunculan burrows include holes in Miocene deposits at depths of I0003000 m off New Zealand (Hayward, 1976).

Evolution and Phylogeny Many fossils of the Devonian tabulate coral Pleurodictyum contain overgrown gastropod shells, most of which were occupied by a secondary resident, possibly a sipunculan like the modem Aspidosiphon (Brett and Cottrell, 1982). Solitary corals with sipunculan symbionts are known from the upper Cretaceous (Wadeopsammia from Texas and Tennessee) and the Miocene (Symbiongia from Florida). The sipunculan is clearly an Aspidosiphon, probably A. muelleri (a taxon that now includes A. corallicola and A. jukesii). The modem hosts of this worm, the corals Heterocyathus and Heteropsammia, are known from the Miocene in France and the Neogene in the western Pacific (Gill and Coates, 1977). Although direct evidence is lacking, the above data support the following points: (I) a common sipunculan-molluscan ancestor existed in Ediacarian or earliest Paleozoic times; (2) sipunculans were living in softbottom burrows at least by the mid-Paleozoic (Devonian) and probably earlier (Cambrian); and (3) some sipunculans have lived in association with corals since mid-Paleozoic times and throughout the Mesozoic and Cenozoic. The Sipuncula thus seems to be an ancient taxon with an unknown history of divergence and retrenchment (escalation and extinction). It is a group that underwent early but conservative cladogenesis, and its members occupied diverse niches early in its history (hard/soft, shallow /deep, warm/cold) and persist in these niches at the present time.

Indirect Evidence The phylum Sipuncula is usually considered most closely related to the annelids and mollusks, but there is no clear consensus as to its true sister group. First, a point about the clustering of phyla into yet higher taxa. The historically accepted constructs Protostomia and Deuterostomia have been broadly criticized in recent decades (e.g., Siewing, 1976). Siewing dismissed these two subkingdoms, as well as the concept of acoels and pseudocoels, and instead proposed the Archicoelomata as the ancestral group that gave rise to three modem groups: Spiralia, Chordata, and Pogonophora. The Spiralia includes the Sipuncula and most of the groups formerly placed in the Protostomia. Although other biologists also support a distinct status for the Pogonophora, no consensus has yet been reached on that issue (E. Cutler, 1975c; Ivanov, 1983, 1988).

Indirect Evidence

337

Nevertheless, some biologists continue to use Deuterostomia and Protostomia (e.g., Lake, 1990). In the following section I use the older termsthose used by the authors whose work is being discussed-when it is appropriate to do so. This is not meant to diminish the value of the Spiralia construct. The descriptive taxon Spiralia is used by many biologists, including Willmer (I990), even though he believes that sipunculans evolved from a hypothetical Protocoelomate group and considers them monomeric coelomates. The assertion that sipunculans are segmented (Ruppert and Carle, I983) seems have its root in Siewing's (I976) idea that the tentacular coelom, which extends into the contractile vessel, is derived from the mesocoel. According to this unusual interpretation, sipunculans would be oligomerous. Siewing's system represents evolution within the Spiralia as follows. An ancestral Spiralian underwent cladogenesis to give rise to the early Sipuncula and its sister group, an ancestral Deutomere. The Deutomere gave rise to the Mollusca and an ancestral Articulata. The latter entity was the precursor of the Annelida and Arthropoda. We will return to this below. A paper presented at the I970 Sipuncula symposium held in Kotor, Yugoslavia, proposed four eumetazoan phyla: Ameria, Polymeria, Oligomeria, and Chordonia (Hadzi, I975). The most advanced class within the Oligomeria was the Sipunculidea, which had evolved from the annelids via the echiurans. This hypothesis has not received support from other biologists. Comparative Immunology The sipunculan immune system is thought to be intermediate between the most primitive systems and the most advanced (see Chapter 10). Ionescu-Varo and Tufescu (I 982) used an iterative analysis (assuming no homoplasy) to survey 12 immunological characters of 9 invertebrate phyla and 1 2 vertebrate taxa. They postulated that the I 2 characters arose sequentially as follows: Recognition of self Rejection of xenograft 3· Specialized leukocytes 4. Rejection of allograft I.

2.

Evolution and Phylogeny 5· 6. 7. 8. 9· IO. I I.

I2.

Immunological memory Type T lymphocytes Circulating antibodies Organs (e.g., thymus and spleen) Plasmocytes Type B lymphocytes Lymph nodes Bursa fabricii, or Peyer's plates

Sipunculans exhibit the first 7 characters. From this primary matrix the authors generated a secondary matrix using percentage similitude and differentiation, then used these data to plot a dendrogram of immune evolution along polar coordinates, giving seven evolutionary levels or stages. The analysis placed the sipunculans in stage 3, together with the Annelida and two non-Spiralia taxa, the Tunicata and Echinodermata. The sipunculans are the only protostomes with circulating antibodies, and no other invertebrate is known to have a more complex immune system. According to Ionescu-Varo and Tufescu's analysis, the mollusks and arthropods have regressed from stage 3 to a lower stage, closer to the coelenterates. This analysis is very interesting, but different interpretations of the data are possible. The arrangement of characters may be suspect because the reasoning used to create it was somewhat circular. The dendrogram suggests that the sipunculans share a common ancestor with annelids, echinoderms, and tunicates. If the arthropods and mollusks evolved from this ancestral group, as Ionescu-Varo and Tufescu believe, some regressive selective pressures must be postulated to have led to the loss of a useful defense mechanism. It is just as reasonable to propose that the arthropods and mollusks split off from a common stock before the sipunculan-annelid line evolved the more complex immune system, and that the deuterostome immune systems evolved independently but in a parallel manner. Siewing's (I976) phylogeny requires either three separate origins for the same defense mechanism or one very early origin and at least two subsequent losses. Neither of these possibilities is very parsimonious, and it is usually best to seek the simplest explanation. The suggestion that sipunculans are more advanced than other invertebrates, while perhaps true with regard to immune systems, may not apply more broadly. If they separated from the other Spiralia as early as Siewing suggested, however, the sipunculans have had sufficient time to evolve many unique attributes.

Indirect Evidence

339

Comparative Biochemistry and Physiology A number of biochemists and physiologists have looked at sipunculan systems (see Part 2), but the database for comparative work is not extensive. Clark's review of the systematics and phylogeny of sipunculans, echiurans, and annelids, in Chemical Zoology (1969), focuses on their developmental biology and supports separate phylum status for each of the three taxa, perhaps within the superphylum Trochozoa. Florkin reviewed the existing biochemical evidence for the phylogeny of the Sipuncula in 1970 (published as Florkin, 1975). Based largely on hemerythrin biochemistry, but also considering nitrogen metabolism and the lack of chitin in the group, Florkin concluded that the sipunculans are a distinct collateral evolutionary line of a preannelid stock. In her excellent review of the role played by physiology and biochemistry in the understanding of phylogeny, Mangum (1990) illustrated how simple generalizations become less credible as knowledge accumulates. Situations that were formerly clear dichotomies become less clear polychotomies. The more we learn, the less certain we are about absolute truths. The example Mangum used was the assertion, considered to be true well into the 196os, that all invertebrates use arginine phosphate in ATP synthesis and all vertebrates use creatine phosphate. By 1970 exceptions had begun to accumulate, and this idea is no longer credible. New information about proteins, amino acid sequences, DNA hybridization, etc., has expanded our understanding of the evolution of molecules, but Mangum properly cautioned readers about equating the evolution of molecules with the evolution of taxa. In fact, many biochemical studies are applicable only to lower-level groupings of organisms such as demes, populations, or species. Mangum suggested that 16-18 S RNA studies may prove helpful at higher levels, but more time is needed to evaluate these methods. With these caveats as background, then, I will proceed. The fact that the level of carbonic anhydrase activity in the red blood cells is fairly high in sipunculans and annelids but not in mollusks led Henry (1987) to propose that mollusks are evolutionarily the more primitive group. A comparison of phospholipids from 59 species of invertebrates from seven phyla showed similarities between sipunculans and other marine worms, including annelids and echiurans (Kostetskii, 1984). The study did not include mollusks, however, and thus does not shed any light on the

340

Evolution and Phylogeny

sipunculan-mollusk relationship. Having similar phospholipids is not necessarily an indication of common ancestry. The types and directions of chemical changes (e.g., the replacement of polar groups, the types of bonds, and the relative amounts of different lipids) are known to respond to environmental factors such as temperature (Kostetskii and Shchipunov, I983). Therefore, a similar phospholipid composition found in two taxa may reflect a similar habitat rather than similar phylogenetic histories. At least two sipunculan species have both actin and myosin regulation of muscle contraction, and thus are like many arthropods, annelids, and nematodes (Lehman and Szent-Gyorgyi, I975). The mollusks, brachiopods, echinoderms, nemertines, and echiurans have lost the actin and have only myosin control, a more derived condition. Single control by actin, the system present in vertebrate striated muscles, occurs in fast muscles of decapods, in mysids, and in one sipunculan, Themiste pyroides (see Chapter 9). The interesting implication of these data, in an evolutionary context, is that sipunculans are similar to annelids but not mollusksand not their presumed closest relatives, the echiurans. Evolution seems to be proceeding in two different directions, but both are away from dual- and toward single-control systems. The selective advantages of single-control over dual-control systems are unclear. The chromatin of eukaryotes is composed of repeating sequences known as nucleosomes. A DNA molecule can be cleaved into its nucleosomes, whose lengths can then be measured (see Chapter 1 1). The nucleosomal DNA repeat length (number of base pairs) in Sipunculus nudus red blood cells is I 77, versus 2 I 2 for chickens, 200 for frogs, and 203 for trout (Wilheim and Wilheim, I 978). Other vertebrate tissues have values in the range of I95-2IO. On the basis of this information, Wilheim and Wilheim asserted that sipunculans are primitive eukaryotes, and that "the small repeat of S. nudus could be correlated to the fact that this marine invertebrate forms an isolated ancestral phylum." A broader sipunculan database (i.e., more than one species) plus data on annelids and mollusks might make this information more useful for determining phylogenetic relationships. Aerobic respiration involves a variety of pyruvate oxidoreductases. Sipunculans-most invertebrates, in fact-use lactate dehydrogenase, among others. They also have alanopine and strombine dehydrogenase, as do annelids and mollusks but not arthropods or echinoderms (Livingstone et al., I983). The one difference Livingstone et al. noted between sipunculans and annelids was the absence of octopine dehydrogenase from the latter, as well as from arthropods and echinoderms. This distribution of

Indirect Evidence

34I

enzymes suggests that sipunculans are more closely related to mollusks than to annelids. Two types of biochemical information, both derived from single sipunculan species and described in Chapter I I, are of no use within this context. The work on amino acid sequences is not useful due to the lack of comparative data, and the electrophoresis of gene loci revealed too much polymorphism. Lake (I990) applied rate invariant analysis of I8 S ribosomal RNA sequences as a means to understand phylogenetic relationships of I I metazoan phyla and classes from the Cnidaria to the Chordata. His analysis was based on data generated by others and included only one sipunculan species, the phylogenetically enigmatic Phascolopsis gouldii, whose distribution is restricted to the northeastern coast of the United States (where it is endemic and not of great age), and whose developmental path and karyotype are of a derived type. Whether it is wise to extrapolate from one such species to the entire phylum is questionable, but Lake's conclusions were that sipunculans are the sister group of mollusks and that the annelids and the sipunculan-molluscan group share a common ancestor. In summary, biochemical and physiological information accumulated since I970 indicates the following phylogenetic relationships: I. Carbonic anhydrase: Mollusks are more primitive than sipunculans, and annelids are similar to sipunculans. 2. Phospholipids: Sipunculans are like other marine worms, and similarities in phospholipid concentrations may reflect ecological, not phylogenetic, similarities. 3· Actin and myosin control: Sipunculans are like annelids and unlike echiurans and mollusks. 4· Nucleosomal DNA: Sipunculans form an isolated ancestral phylum. s. Pyruvate oxidoreductases: The respiratory enzymes of sipunculans are like those of most other invertebrates, but they have three enzymes lacking in arthropods and echinoderms and one enzyme that annelids lack. They show no differences from the mollusks. 6. Amino acid sequence and protein electrochemistry: No value. 7· Ribosomal RNA Sequence: Annelids diverged from a molluscansipunculan ancestor and the latter two are sister groups.

These data generally point to a close relationship among sipunculans, annelids, and mollusks (i.e., all three probably evolved from a common ancestor) and suggest that mollusks may be less derived than sipunculans

342

Evolution and Phylogeny

and annelids. Alternatively, the differences that mollusks evince in items 1 and 3 above could mean that mollusks are more derived. On the other hand, if rRNA sequencing is all that its proponents claim, then item 7 is all one needs to consider. In this regard, a larger database would generate much more confidence in these conclusions. Comparative Fine Structure The electron microscope has revealed details of cell structure that are useful for determining the degree of relatedness among taxa (Barnes, 1985). The evidence that concerns sipunculans is of three types, discussed below. Cells that line lumens or exterior surfaces have a belt around their apical circumference known as an intercellular junction. The precise nature of this junction varies (13 types have been identified), but in sipunculans and annelids it is a pleated septate junction of the "lower invertebrate" variety. Mollusks and arthropods have a "protostome" septate junction (C. Green and Berquist, 1982). The same authors interpreted this to mean that sipunculans are in the deuterostome lineage. Actually, the data could support the idea that sipunculans and annelids are closely related and more primitive than mollusks. It is also possible that the mollusks added this character after separating from the sipunculan-molluscan group. Nielsen's (1987) analysis of the feeding and swimming cilia in 15 phyla of invertebrates showed that the nature and position of the accessory centriole, which is perpendicular to the basal body and on the downstream side, links sipunculans to annelids and mollusks. The nature of the striated sperm anchoring fiber apparatus was mentioned in Chapter 15. In the present context it is worth noting Klepel's (1987) assertion that the sipunculan arrangement is like that of the original protostomial type. The anatomical data point to the conclusion that sipunculans are a primitive group related to both annelids and mollusks and probably less derived than either group (and therefore consistent with Siewing, 1976). Comparative Embryology The reproductive biology of sipunculans is discussed in Chapter 15, but there are several points worth reviewing here. While an understanding of developmental pathways can provide a good context in which to analyze

Indirect Evidence

343

relationships among taxa, linking ontogeny and phylogeny too rigidly can lead to false conclusions. One type of evidence that is universally considered to indicate monophyly is the manner in which the egg undergoes cleavage. Sipunculan eggs follow the spiral route, thus are placed in the Spiralia along with the annelids and mollusks (Siewing, 1976). Even though it has been suggested that the trochophore is probably a derived larval form that could have evolved independently more than once (Ivanova-Kazas, 1985), animals with this larval stage are still assumed to be related (Rice, 1985a). Strathmann (1978) expressed a similar concern with regard to cilia used as feeding structures in larvae and asserted that larval morphology should not be used to suggest a close relationship between sipunculans and annelids or mollusks. Rice (1985a) pointed to the following similarities as evidence of sipunculans' closer relationship to annelids: the prototroch and metatroch ciliary bands, the development of the larval cuticle from the egg envelope, and the development of the nervous system (see Chapter 16, Sense Organs, for details about the latter). One interesting similarity between sipunculan and molluscan embryology is the radial position of the cross cells in the apical plate. A major difference from the annelids is sipunculans' lack of metamerism at any stage in their ontogeny. Based on these points, Rice proposed that sipunculans are a primitive phylum that arose from an annelid-mollusk stem. Two other considerations of developmental attributes reached a different conclusion. Freeman and Lundelius (1992) proposed a close relationship between sipunculans and Mollusca (class Aplacophora) based on the mode of D quadrant specification (both taxa have unequal cleavage). Their argument that induction is the primitive mode of D quadrant specification rests on a series of assumptions, as follows: equal cleavage can be equated with induction; the more derived cytoplasmic localization is linked to unequal cleavage (as in sipunculans); the database is sufficiently complete (they included three sipunculans); the phylogenetic relationships of the groups they discussed were accurate (these were not complete). Freeman and Lundelius were unable to relate sipunculans and aplacophorans to other metazoan taxa in more than a tenuous manner, and they went on to say that these two groups are the only ones that do not fit their developmentevolutionary scenario. In other words, both taxa have unequal cleavage, which would translate into a derived mode, but Freeman and Lundelius resisted that conclusion.

Evolution and Phylogeny

344

In an article devoted largely to the proposition that these same wormlike aplacophorans comprise a primitive taxon within the Mollusca, A. Scheltema (1993) postulated that Sipuncula and Mollusca are sister groups. This argument is based on similarities in early development (the molluscan cross) and a few transitory features of pelageosphera larvae (lip gland and buccal organs). Scheltema also postulated that, like mollusks, the sipunculans must have appeared early in the evolution of the metazoans. The fact that sipunculans and some mollusks, which are known to have originated in the Cambrian, all use hemerythrin as an oxygen transport molecule supports an early origin for sipunculans.

Conclusions Although the evidence is not totally congruent, there is consensus that there was an ancestral form common to the sipunculans, annelids, and mollusks in existence by the earliest Paleozoic. From this point there are three possibilities, each supported by some part of the data: ( 1) the annelids separated from an ancestor that later gave rise to the sipunculans and mollusks; (2) the molluscan stock diverged first, followed by the sipunculan-annelid separation; or (3) the sipunculans diverged from a stock that subsequently became the common ancestor to the mollusks and annelids. Table 4. Possible phylogenies for Annelida, Mollusca, and Sipuncula

Modell ASM

Model2 MSA

Model3 SAM

~

~

~

+

+ + +

?

Paleontology Immunology Biochemistry Fine Structure Embryology

X

+

X

+

X

?

X

+ + +

Notes: A = Annelida, M = Mollusca, S = Sipuncula; X most strongly, + = permits (no contradiction).

?

X

+

= supports

Conclusions

345

The third model is consistent with Siewing, 1976. The third and second models are each supported by one, and permitted by the remaining four types of evidence just discussed. The frrst model is supported by the paleontological, biochemical, and embryological data and is permitted by the immunological and fmestructure data (see Table 4). Based on my evaluation of the evidence, model 1 aopears to be the most probable, although I acknowledge the limited nature of some parts of the database. I must also restate that this is really linking the most primitive of the molluscan taxa (according to A. Scheltema, 1993) with the least derived of the sipunculans as defined in Chapter 19.

19

Within-Phylum Relationships

Phylogenetic relationships among the taxa within the phylum Sipuncula are considered in E. Cutler, 1980, and E. Cutler and Gibbs, 1985. A formal presentation of the resulting classification, with a few corrected spellings, followed a short time later (Gibbs and Cutler, 1987). Readers interested in the philosophy and details of the numerical methods used to determine relationships should see the 1985 work. An abbreviated discussion of the characters used in the analysis is presented below, together with some new information and a reevaluation of some assumptions about character state polarities (Table 5). The suprageneric taxa as used earlier remain unchanged, but the proposed historical relationships among taxa are radically different in a few of the aspects described below. When the plesiomorphic/apomorphic (i.e., ancestral/derived) character states of the 12 morphological characters used in earlier analyses were described, the polarities were rooted in a hypothetical ancestral sipunculan (HAS). That model is redefined here. The nature of sipunculans imposes severe limitations on character analysis. Their elastic, soft bodies have almost nothing meaningful to measure or count, there is no fossil record, there is no good out-group to help root characters when attempting to polarize, and the number of useful characters is modest. All the available morphological information on sipunculans has been recoded and used as input for the PAUP phylogenetic analysis program. In general, the end products (cladograms) do not differ from already published configurations (e.g., E. Cutler and Gibbs, 1985:fig. 1). My computer analyses used five different data sets: family-level data, from the six families, followed by runs for each of the four orders at the subgeneric level. The later runs used more restricted and appropriate data (see Tables 6-10). The paucity of characters available for analysis resulted in dendrograms

Morphological Data

347

Table 5. Attributes used in cladistic analyses 1.

2. 3. 4. 5. 6. 7. 8.

Tentacles: 0, nuchal only; 1, nuchal and peripheral; 2, peripheral only; 3, dendritic peripherals Nephridia: 0, pair, bilobed; 1, pair, unilobed; 2, single Coelomic extensions: 0, none; 1, pouches; 2, canals Introvert-trunk junction: 0, straight; 1, angle Postesophagealloop: 0, absent; 1, present Anus location: 0, anterior of trunk; 1, on introvert Anal shield: 0, none; 1, simple (Aspidosiphon); 2, massive (Lithacrosiphon); 3*, Cloeosiphon Spindle muscle: 0, attached posteriorly to body wall; 1, ends within gut posteriorly; 2, absent

Homoplastic characters, for within-family analyses Introvert hooks: 0, in rings, with basal spinelets; 1, in rings, no spinelets; 2, in rings of very young, absent in adults; 3, none in adult or young; 4, none in rings, replaced with scattered hooks 10. Body wall muscle layers: 0, both continuous; 1, longitudinal layer divided into bundles, some anastomosing; 2, both layers with anastomosing bundles; 3, both layers divided into distinct bands 11. Contractile vessel villi: 0, absent; 1*, many short digitiform units; 2 *, few long stringy tubular units 12. Introvert retractor muscles:a 0, two equal pairs; 1, ventral pair only; 2, fused ventral pair only; 3, all there but fusion of dorsal to dorsal and ventral to ventral; 4, fusion and reduction of ventrals; 5, incomplete fusion of all four; 6, complete fusion of all four. 13. Nephridiopores relative to anus (in Sipunculidae ): 0, anterior; 1, posterior 9.

Note: Character state polarity coding; asterisk indicates unordered components (o is always the ancestral, or plesiomorphic, state). apossible sequences: 0-1-2, 0-3-4, 0-3-5-6.

Table 6. Character states of sipunculan families

Attribute

Sipunculidae Golfingiidae Themistidae Phascolionidae Phascolosomatidae Aspidosiphonidae

2 1, 2 3 2 0 0

2

3

4

5

6

7

8

1 1 1 2 0, 1 1

1, 2 0 0 0 0 0

0 0 0 0 0 0, 1

0, 1 0 0 0 0 0

0 0 0 0, 1 0 0

0 0 0 0 0 1, 2, 3

0, 1 1 1 2 0 0

Note: Attribute numbers are numbers 1-8 in Table 5. Character states are those used in Table 5.

Table 7. Character states of sipunculiformes genera and subgenera Attribute

3 2 2 3 1 1

Sipunculus S. (Austrosiphon) Xenosiphon Siphonosoma Siphonomecus Phascolopsis

8

5

1

0 0 0 0

0

0 0

9

10

12

13

3 3 3

3 3

0

3

1 2

2 2 1

0 0 0 0

1 1

0

0 0 0

1

Note: Attribute numbers are those listed in Table 5.

Table 8. Character states of Golfingiiformes genera and subgenera Attribute

Golfingia G. (Spinata) Nephasoma N. (Cutlerensis) Thysanocardia Themiste T. (Lagenopsis) Phascolion P. (Jsomya) P. (Montuga) P. (Villiophora) P. (Lesenka) Onchnesoma

2 2 2

2

6

8

9

11

12

1

0 0 0 0 0 0 0 0 0 0

1 1 1 2 1 1 1 2 2 2 2 2 2

4* 0 4* 3 3 4* 4* 4 4 4 3 3* 3

0 0 0 0

0 0

1 2 1

1 1 1

0 0 0

3

0

4

1

3 3

1 1

2 2 2 2 2 2

2 2 2 2 2

2

0* 1

2

4

3

5

0 0

5 6

Note: Attribute numbers are those listed in Table 5. * = Polymorphic, but most species exhibit indicated state. States 1, 2, or 3 are present in one to several species for character 9.

Table 9. Character states of Phascolosomatiformes genera and subgenera Attribute

Phascolosoma P. (Fisherana) Antillesoma Apionsoma A. (Edmondsius)

2

8

9

10

11

0 0 0 0

1

I 1

1 2

0

0 0

0

0 0 1 0 0

0 0

1

Note: Attribute numbers are those listed in Table 5.

1

Morphological Data

349

Table 10. Character states of Aspidosiphoniformes genera and subgenera Attribute

4 Aspidosiphon A. (Paraspidosiphon) A. (Akrikos) Lithacrosiphon Cloeosiphon

1 1 0

7

9

10

1 2

1 4 1 1

0 1 0 1 0

3

Note: Attribute numbers are those listed in Table 5.

that are not worth presenting here. The many polychotomies (i.e., unresolved branch points) and the artificial and arbitrary nature of the methodology, as pointed out by E. Cutler and Gibbs (I985), are other reasons for not presenting dendrograms. As more information becomes available the conclusions presented below should be tested and, if necessary, modified. I encourage others to use the data in Tables 6- I o in appropriate analyses.

Morphological Data Changes in applicability and character state polarity from that presented in Cutler and Gibbs, I985, are as follows. Tentacles: Nuchal tentacles now considered ancestral to peripherals. Spindle muscle: Posterior attachment (complete) is now ancestral; unattached and absent are derived states. 3· Introvert hooks: (A) Complex hooks in rings are ancestral; scattered or absent hooks are derived and homoplastic (i.e., they evolved more than once in different lineages; this may result in either parallel or convergent evolution, but most importantly such characters are not homologous and thus do not indicate a shared common ancestor). (B) Hooks with basal spinelets are ancestral. 4. Longitudinal muscle bands: Presence is homoplastic above family level. 5· Contractile vessel villi: Presence is homoplastic above family level. 6. Introvert retractor muscles: Loss or fusion is homoplastic above family level. I.

2.

350

Within-Phylum Relationships

Broadly Useful Characters The following eight morphological attributes can be used to analyze relationships among all of the families and genera. It is assumed that they originated only once and therefore are not homoplastic. 1. Tentacle arrangement. The oral disks, with their tentacular crowns, are diverse, but when the misleading descriptions are corrected, two general types are recognizable: type P, in the class Phascolosomatidea, and type S, in the class Sipunculidea. Type P tentacles are simple and small, arranged in a dorsal arc around the nuchal organ, and number 10-30 in most species. TypeS tentacles are arranged peripherally on the oral disk encircling the mouth. They are especially well developed in Thysanocardia, reduced in other genera, and significantly modified in Themiste. The dorsal nuchal organ in some typeS arrays may be encircled by an arc of small nuchal tentacles. The variations on these themes are described in the sections on morphological characters throughout Part I. It had been proposed that type S is ancestral to type P (E. Cutler and Gibbs, 1985). Alternatively, the peripheral tentacles may represent a later adaptation for feeding, and the reverse polarity is proposed here. The cephalic collar below the oral disk in the Phascolosomatidea is now considered to be the precursor of the peripheral tentacles, not the remnant. The evolutionary sequence now proposed is from an ancestor with only nuchal tentacles to a form with both peripheral (feeding) and nuchal (chemoreception) tentacles, to forms with only peripheral tentacles; that is, a gradual reduction of one set and elaboration of the other. The external feeding apparatus is subject to direct selection pressures (predation), and its efficacy directly affects the success of the genotype. The wide variety of types between and within genera suggests a faster rate of change in tentacles than in the general body plan. This opens the way for convergent or parallel trends as well as reductions in complexity or reemergence of previously suppressed complex phenotypes. 2. Nephridia number. Most sipunculans are bilaterally symmetrical and have two nephridia. The apomorphic loss of one nephridium has occurred in Phascolion and Onchnesoma. 3· Coelomic extensions. In most Sipunculidae, the coelom connects to epidermal canals or sacs via small pores through the muscle layers. The nature of the synapomorphy (i.e., shared derived character) varies among the four genera. For more details see Chapter 2, in this volume, and E. Cutler, 1986).

Morphological Data

351

4. Introvert-trunk junction. The anterior-posterior axis ofthe introvert is a continuation of the main trunk axis in most genera, and the ancestral condition. The anal shield present in Aspidosiphon and Lithacrosiphon forces the introvert ventrally to an angle ranging from 40 to 90°. 5· Postesophagealloop. The esophagus leads directly into the double helix of the gut coil in most genera. A derived condition is seen in the genus Sipunculus, which has a separate and distinct anterior loop between the straight esophagus and the double-coiled gut. 6. Anus location. The anus is located mid-dorsally very near the anterior end of the trunk in most genera. In Onchnesoma and four species of Phascolion the anus is located on the introvert, at least 20% of the distance toward its tip, an apomorphic condition. 7. Anal shield. A horny or calcareous shieldlike structure occurs at the anterior end of the trunk in three coral-inhabiting genera. The form of this shield varies considerably, and homoplasy is likely. Aspidosiphon represents one independent line, with Lithacrosiphon being a modification of this apomorphic state. The anal shield in Cloeosiphon is very different from the other two and is assumed to have evolved independently. 8. Posterior attachment of spindle muscle. The threadlike spindle muscle extends through the gut coil to the posterior end of the trunk in some genera (ancestral), but in others it terminates within the coil (derived). This reduction or loss continues to a second derived state in Phascolion and Onchnesoma, in which its total loss has left only fixing muscles to anchor the gut. Limited-Use and New Characters The following characters have been determined to be misleading if used to determine relationships above the family level. One previously unused character is presented here as well. 9· Introvert hooks. Various kinds of hooks and spinelike structures grow on the distal half of the introvert. The phascolosomatid type of hook, which exhibits an internal complexity and is arrayed in distinct rings, was considered apomorphic (E. Cutler and Gibbs, 1985), but now there is good reason (as tentatively proposed in E. Cutler and Cutler, 1988) to consider the reverse more likely; that is, complex hooks in rings are plesiomorphic. The primary reason for this reversal is the discovery of such hooks in very young members of species previously thought to be hookless, including one Themiste, the single Antillesoma species, and Phascolosoma meteori. Phascolopsis gouldii juveniles have hooks, but the arrangement is not

352

Within-Phylum Relationships

clearly in rings. In some polymorphic genera (e.g., Siphonosoma or Apionsoma), species without hooks have rings of small papillae where hooks are found in congeners. Furthermore, the presence of hooks in an ordered array appeared early (in the evolutionary and paleontological sense) in related taxa such as the enigmatic Cambrian Ottoia and several groups of extant worms such as some acanthocephalans, kinorhynchs, and priapulids. The loss of regular rings of hooks in adults has probably occurred several times (homoplasy) and via different genetic mechanisms, because in modern sipunculans the loss occurs at different times during the ontogeny and results in diverse end products. The loss is often, but not always, followed ontogenetically by replacement with a scattered array of some other type of hook. In some species, though, the animal remains hookless for the remainder of its life. While the absence of hooks in rings may be apomorphic, this character should not be used as an indication of common ancestry (synapomorphy) above the genus level. In an analysis based on phenetic (rather than cladistic) methods, the synplesiomorphy (i.e., shared ancestral character state) of hooks in rings might be of value. Another major change that concerns hooks is the inclusion of hooks with basal spinelets as plesiomorphic (Fig. 54). This position is counter to earlier assertions that these hooks and the bilobed nephridia and very long introverts found in the Apionsoma species, as well as the monotypic subgenus Golfingia (Spinata), are unique derived character states (E. Cutler, 1979; E. Cutler and Cutler, 1987a; N. Cutler and Cutler, 1990). Rather than considering these traits to be recently evolved, specialized traits that arose independently and convergently in two different generaand therefore omitting them from phylogenetic analyses-it is now proposed that these traits are ancestral and have been retained in a few "living fossils." The presence of complex ornamented or pectinate hooks, teeth, and spines in other living worms, such as some polychaetes, priapulids, and kinorhynchs, and in some Burgess shale fossils (e.g., the proboscis spinules of Ottoia), supports this character polarity. Figure 86 suggests how an Ottoia-type structure might have undergone a folding along the midline to become an Apionsoma type of hook. The presumed evolutionary sequence is thus from ringed hooks with basal spinelets, to ringed hooks without spinelets, to the loss of hooks in adults, followed by loss in both juveniles and adults, which then either stay hookless or develop new scattered hooks. 10. Body wall muscle layers. The two layers of musculature in the body

Morphological Data

353

., ~ . .

~

.....

I

•:

..

::

~~-:

. .

A

B

c

Figure 86. Possible sequence in the early evolution of sipunculan hooks. A. Proboscis spinules of the Cambrian Ottoia (after Banta and Rice, 1976). B. Hypothetical intermediate stage with the lateral edges folding together. C. Apionsoma hook with basal spinelets (see also Fig. 54B).

wall are continuous layers in the ancestral state. In some genera, however, the longitudinal muscle layer of the body wall has split into more or less distinct bands. Although this attribute is considered apomorphic and is used as an indication of common ancestry, it has undoubtedly occurred more than once (homoplasy). The circular layer may further divide into partially separated bundles, an even more derived state. Within the Sipunculidae both layers form distinct, separate muscle bands as the most derived condition. I I. Contractile vessel villi. The contractile vessel is spacious and has digitiform villous outpouchings in a number of species. It seems likely that this is an apomorphic but homoplastic condition that has appeared independently, along with complex and voluminous tentacular crowns, in five of the six families. The simple contractile vessel without villi is plesiomorphic. In Themiste, two types of villi evolved. One subgenus, T. (Themiste ), has few long, thin, threadlike extensions, and E. Cutler and Cutler (I988) questioned the presumption of homology. T. (Lagenopsis) has the same type of contractile vessel villi as those found in the other genera that possess them. I2. Introvert retractor muscles. The extended introvert is retracted by muscles whose origins are on the trunk wall and insertions are behind the cerebral ganglia. The plesiomorphic state is two equal-sized pairs-a ventral and a dorsal. A number of genera have only one pair. This reduction probably occurred at least once in each class, possibly four times altogether. Muscle fusion also occurs, commonly in Phascolion, and involves fu-

354

Within-Phylum Relationships

sion of dorsal to dorsal or ventral to ventral. In a few speciesOnchnesoma, for example-so much fusion (and reduction?) has occurred that only a single muscle is apparent. I3. Nephridiopores-anus relationship. For most taxa this relationship is not of phylogenetic value. Within the family Sipunculidae, however, the nephridia open just anterior to the anus in all but three species, where this relationship is reversed in the derived state. Characters Not Used in Numerical Analyses Epidermal Glands. One rather general observation not used in previous discussions but which supports these phylogenetic conclusions was made by Akesson (I958). In the context of a detailed commentary on sipunculan epidermal organs, he identified three groups: (I) the Golfingia group, with two types of cells and secretory products; (2) the Phascolosoma group, with only one type of cell and product; and (3) the Sipunculus group, with separate sensory and secretory cells and glands of two types like group I (bi- and multicellular). A plausible and parsimonious evolutionary sequence could begin with the simplest (second) type (Phascolosoma) as the ancestral form, which then led to the apomorphic type I (Golfingia), which in turn could have given rise to the most derived type, the third (Sipunculus).

Karyological Data "Evolution is essentially a cytogenetic process," and ignoring this fact "makes for a weak and incomplete analysis." With these words of M. White (I973:759) setting forth the consensus viewpoint, the little that is known about sipunculan genetics is presented below. This is fertile ground for future work. The chromosomal morphology of I4 species of sipunculans as determined by J. Silverstein (Ig86, and pers. comm., I99I) is summarized in Table I 1. The diploid number is 20 for all five members of the class Phascolosomatidea included in the table, and for six of the nine sipunculideans. Most species show a gradual transition from small to large chromosomes; a few exhibit a bimodal pattern. The four species in the order Phascolosomatiformes show a strong tendency toward asymmetrical arm length; that is, 70-100% of the chro-

7

4

6 1 15

7

6

8 9 10 10

-

Metacentric

1

-

3 3 3

2

-

1

3

-

Submetacentric

2 1

1 2

4

few

+

6

7

Subtelocentric

Chromosomal morphology

9

most

4

+

Telocentric

Source: Data provided by J. Silverstein. Note: Collection locations were as follows: I. Sesoko, Okinawa, Japan (beach near marine lab); 2. Oki Island, on Japan Sea; 3. Ft. Pierce, Fla. (near Harbor Branch Lab); 4. same as 3 and Curac;:ao; 5. same as 1; 6. Shimoda, Japan (near marine lab); 7. Ushimado, Okayama, Japan; 8. Santa Barbara, Calif. (near Pt. Conception); 9. Hollister Ranch near Gaviota, Calif.; 10. Carmel Pt., Monteray Bay, Calif.; 11. Woods Hole, Mass.; 12. Suva, Fiji; 13. Ushimado, Okayama, Japan; 14. same as I; 15. same as 1.

Sipunculidea Golfingiiformes (2N = 20) 6. Golfingia margaritacea 7. Thysanocardia nigra 8. Themiste hennahi 9. Themiste dyscritta 10. Themiste pyroides Sipunculiformes (2N = 18-34) 11. Phascolopsis gouldii 12. Siphonosama australe 13. Siphonosoma cumanense, Okayama 14. Siphonosoma cumanense, Okinawa 15. Sipunculus nudus

Phascolosomatidea (2N = 20) Phascolosomatiformes I. Phascolosoma pacificum 2. Phascolosoma scolops 3. Phascolosoma perlucens 4. Antillesoma antillarum Aspidosiphoniformes 5. Aspidosiphon steenstrupii

Table 11. Karyotypes of sipunculans

Within-Phylum Relationships mosomes are telocentric or subtelocentric, and none are metacentric. The single Aspidosiphoniformes species analyzed has so% telocentric or subtelocentric and so% metacentric or submetacentric chromosomes. In contrast, 80-100% of the chromosomes in the nine species from six genera in the class Sipunculidea are metacentric or submetacentric; that is, they exhibit much greater symmetry of arm length. An apparent anomaly exists in one population of the widespread Siphonosoma cumanense. The Okinawa subpopulation of this species (2N = 24) appears to have mostly telocentric chromosomes, while the Okayama subpopulation (2N = 18) has none. The latter group is like all three Themiste species, which also have no telocentric or subtelocentric chromosomes. The five Golfingiiformes species are much more stable (all with 2N = 20) than their four Sipunculiformes counterparts, of whom only Phascolopsis gouldii has 10 pairs of chromosomes. In addition to S. cumanense mentioned above, S. australe has 22, and Sipunculus nudus appears to have 34 miniaturized chromosomes. It would be interesting to know the karyotypes of two highly derived members of this order, the hermaphroditic Nephasoma minutum and the parthenogenetic Themiste lageniformis. Given that telocentric chromosomes have only one arm and metacentric chromosomes have two, a comment about arm number and recombination is in order. The frequency of chiasmata formation and crossing over during Prophase I of meiosis is different in the two configurations. The probability of genetic recombination increases with the number of arms. A pair of telocentric chromosomes, with only two arms per homologous pair, is less likely to experience crossing over than is a similar pair of metacentric chromosomes with four arms (M. White, 1973). Genetic recombination rarely produces new species. More commonly it provides morphological or physiological variation (polymorphism) within a species. Extending this generalization to the sipunculan data may explain the great physiological polymorphism in the eurytopic sand- and muddwelling Sipunculidea. Although the worms whose karyotypes are known are all intertidal species, the rock-boring Phascolosomatidea taxa all live in thermally stable habitats and are exposed to insignificant fluctuations in salinity. One could speculate that the Sipunculidea is a more rapidly evolving group, able to respond to changing conditions, and therefore more common in geologically more recent (cold, deep) habitats. Most members of the Phascolosomatidea, on the other hand, are slower evolving and largely restricted to geologically older (warm, shallow) habitats.

Embryological Data

357

Finally, a correlation between chromosomal symmetry and asymmetry and apomorphic versus plesiomorphic character states is fairly well established for plants and some insects (M. White, 1973). It seems clear that ancestral taxa have high proportions of telocentric chromosomes (8 of 9 in grasshoppers), and more derived taxa have high numbers of metacentric chromosomes (20 of 23 in ladybird beetles). Assuming this to be true for sipunculans as well, and incorporating the information presented above, I propose the following evolutionary hypothesis for this phylum, one that is consistent with other information: The ancestral population had little or no chromosomal symmetry (i.e., they were phascolosomatids). Some of these telocentric units experienced a redistribution of material in a more symmetrical fashion. These mutated chromosomes produced new taxa within the Phascolosomatidea (e.g., the aspidosiphonids). Larger differences in chromosomal morphology (more chromosomes with equal arm length) led to larger differences in adult worm morphology and even a new class, the Sipunculidea, with mostly metacentric chromosomes.

Embryological Data Rice's (1985a) model for the evolution of sipunculan larval forms begins with a presipunculan ancestor having an egg with a simple envelope and low yolk content that developed into a planktotrophic trochophore larva. From this stock evolved an ancestral primitive sipunculan with a moderately yolky egg, a thick egg envelope, a nonfeeding trochophore with a persistent egg envelope (retarding planktotrophy), and a planktotrophic pelageosphera stage. The increased yolk and thickness of the egg envelope were the two major evolutionary trends leading to the original pelagosphera. From this hypothetical starting point Rice envisioned a bifurcating evolutionary street to the extant forms, one branch leading toward an increase in yolk and a decrease in the length of the pelagic stage, the other leading to a decrease in yolk and an increase in the length of planktotrophic pelagic life. I suggest that there is a less complex, more parsimonious model (Table 12, Fig. 87).It requires a single nonbranching path, with no reversals, from a single presipunculan ancestor, in the direction of gradual increases in

Within-Phylum Relationships Table 12. Summary of sipunculan developmental pathways

Type• Number of species Vermiform juvenile Pelagosphera larva Planktotrophic Lecithotrophic Trochophore Planktonic life Egg Size Yolk content Envelope Cladogenic events b

PP (IV)

LP (III)

T (II)

D (I)

lO X

7 X

2 X

3 X

X X Weeks X Medium Medium Medium

X Days X Medium Medium Medium

None X Large High Thick

X X Months X Small Low Thin A

B

c

Note: Presented in an evolutionary context presuming a loss or simplification at each step. • PP = planktotrophic pelagosphera; LP = lecithotrophic pelagosphera; T = trochophore; D = direct. The number in parentheses is Rice's (1985a) type number. b A = yolk content of egg increases, egg envelope thickens, egg size increases, larva unable to feed on plankton, shorter time in plantkon; B = yolk content increases, egg envelope thickens, egg size increases, loss of pelagosphera, shorter time in plankton; C = yolk content of egg increases, egg envelope thickens, egg size increases, direct development, no larval life.

yolk content, thickening of the egg envelope, and decrease in time of planktonic existence. Larval lifestyles went from long-lived planktotrophic, to shorter lecithotrophic pelagosphera, to trochophore only; and in a few special cases on to direct development (Rice's types IV, III, II, and 1). This model requires the extension of the larval life of the presipunculan through the addition of the novel pelagosphera-as does Rice's modelbut without the immediate increase in yolk and thickening of the egg envelope that Rice's model postulates (1985a). The available data, reinterpreted this way and combined with other, nonembryological, research, lend support to the new model. Four of the six genera in the less-derived class, Phascolosomatidea, are known to have the least derived ontogeny (Rice's type IV), including the species Phascolosoma agassizii, P. perlucens, P. nigrescens, Antillesoma antillarum, Apionsoma misakianum, Aspidosiphon parvulus, and A.fischeri. No members of this class are known to have any type of larvae other than Rice's type IV (long-lived planktotrophic). Representatives of the class Sipunculidea exhibit all four developmental types. Two members of the family Sipunculidae (Sipunculus nudus and Siphonosoma cumanense) have type IV development, as does the

Embryological Data

+

359

I

t

·I~

.

.

t

.

®

t

~

Figure 87. Proposed evolutionary sequence of sipunculan developmental patterns, from left to right, incorporating text discussion and Table I I. Each drawing in this figure is represented by an X in Table I I.

golfingiid Nephasoma pellucida. Three other golfingiids (G. vulgaris, G. elongata, and Thysanocardia nigra) and two themistids (T. alutacea, T. lageniformis) exhibit type III development (no planktotrophic stage). Phascolion strombus has type II development (trochophore only), as does the enigmatic Phascolopsis gouldii, whose familial affinity is ambiguous but is currently considered to be Sipunculidae (formerly Golfingiidae). The most derived path, type I (direct development), occurs in Themiste

Within-Phylum Relationships pyroides, Phascolion cryptus, and the hermaphroditic Nephasoma minuta; each is in a different family within the order Golfingiiformes. It seems very probable that the class Sipunculidea began with type IV development, and that types Ill, II, and I each evolved more than once, within different genera, during subsequent cladogenic events (homoplasy). Therefore, developmental pathways can be used as a guide to the evolution of sipunculan higher taxa, but they must be used with caution and preferably in conjunction with other kinds of information.

Zoogeographical Data: Paleo-Oceanographic Analysis As I asserted above, some early sipunculans probably existed in Paleozoic times, more than 500 million years ago (Ma). (In pre- 1980s literature, Ma was abbreviated as MYBP, million years before present, and a few recent works use Ma BP). The genesis of clades was not instantaneous, however, and the present distribution patterns around certain geologically important regions are informative. Theories about the size, shape, position, and movements of the land masses on this planet have been produced under the rubric of plate tectonics or continental drift, mostly since 1960. Although most of this work has focused on land masses, it is possible to infer information about the surrounding oceans, and since 1970 a few authors have concentrated their studies on the ocean basins. Authors' opinions and conclusions vary, and the literature is not consistent with regard to dates and shapes. The precision lessens as one goes back in time. Table 13 presents the names and dates of the geological time units along with major tectonic events (also see Fig. 88). The following overview is based partly on A. Smith et al., 1981, and Weijermars, 1989, which incorporate and modify data published only a decade earlier in works such as Fleming, 1978, and Grant-Mackie, 1978. The literature on the probable times and rates of extinction and speciation in different parts of the oceans is far too complex to be adequately considered in this small space. Part of the confusion centers on definitions of terms such as old and young and whether one is dealing with plants or animals; plankton, nekton, or benthos; infauna or epifauna. An Eocene event is old if one is talking about the deep Atlantic Ocean but very young in terms of life in the Pacific. A simplistic summary is this: taxa found in high-stress, unstable habitats

Zoogeographical Data Table 13. Geological time in the Phanerozoic eon and zoogeographically significant paleo-oceanographic events Paleozoic periods Ediacarian Cambrian Ordovician Silurian Devonian Carboniferous Permian Mesozoic periods Triassic Jurassic Cretaceous

675 570 505 438 408 360 286

248 213 140

Cenozoic periods and epochs Tertiary (Paleogene) 65 Paleocene Eocene

55

Oligocene

38

(Neogene) Miocene

25

Pliocene

5

Quaternary Pleistocene Holocene

2 0.01

Pangea forms.

Pangea splits and Tethys Sea forms; Madagascar splits from Africa. Early: Atlantic begins to form; Tethys at maximum size; Madagascar arrives at present location; India splits from Africa. Late: N and S Atlantic join; Bering Strait closed by land bridge.

New Zealand breaks away from Australia-Antarctica; North Atlantic opens to Arctic; deep sea warming. Australia splits from Antarctica; deep water connections between N and S Atlantic form; India arrives at present location; present biogeographic provinces begin to form along with polar ice; land north of Australia fragmenting and adjacent seas forming. Atlantic reaches present depth; Australia arrives, and there is deep water between it and Antarctica; Antarctic sea ice forms. Early: Drake Passage opens; Africa meets Eurasia, closing eastern Mediterranean; Antarctic ice cap grows and deep Pacific cools. Late: broad IWP shelf; upwelling off SW Africa; Iberian portal closes. Panama isthmus closes; Bering Strait reopens; glaciation with permanent ice at both poles. Biogeographic provinces well formed; periodic glaciation. Continued temperature and sea level fluctuations.

Note: Numbers are million years since the start of the period or epoch.

(temperate, intertidal) are younger than those found in more stable areas (deep sea, tropical). Differing viewpoints do exist, of course. Some authors suggested that the tropics are no older or more stable than the Arctic. Both areas underwent significant Miocene thermal changes, and Valentine

80W

40W 40E

80E

120E

160E

160W

Figure 88. Map of present world, with historically important regions labeled.

0

120W

80W

Zoogeographical Data (I984:649) argued that "if there are any shallow-sea regions likely to harbor particularly large numbers of old species, they might be the temperate and subtropical zones." Many references to the antiquity of marine taxa apply only within Cenozoic, or even Neogene, times. A safe guideline is to assume this to be what the author meant unless it is specifically stated otherwise. Paleozoic (570-248 Ma) During the Paleozoic period the land was formed into three separate continents: Gondwanaland (the southern land masses), Laurasia (North America and Europe west of the Urals), and eastern Eurasia. These masses migrated over the surface of the earth to the Southern Hemisphere, where eventually they coalesced into a single supercontinent, Pangaea, sometime late in the Paleozoic (Boucot and Gray, I983). Mesozoic (248-65 Ma) Pangaea persisted for at least IOO million years (28o-I8o Ma), during the Triassic and part of the Jurassic. A single continent meant a single surrounding ocean, the Eo-Pacific, or Panthalassa, which was much larger than today's Pacific. The Pacific Ocean has been shrinking since its formation, partly because it is surrounded by subduction zones (where one tectonic plate slides down and under another) that consume ocean floor 24 cm/yr faster than it is being produced along the spreading zones. The change in dimensions has occurred asymmetrically because the subduction along the eastern margin is faster than on the western end, and the principal spreading zone that was the Mid-Pacific Ridge of 65 Ma is now the East Pacific Rise. Viewed on a geological time scale, the ocean floor has not been a static habitat for benthic invertebrates. There is no place where the floor is older than I 8o million years (Jurassic), and 6o% of it is less than 65 million years old (end of Cretaceous). About I 8o Ma Pangaea split into two parts: Laurasia (northern) and Gondwanaland (southern). The two land masses were separated by a shallow, warm ocean called the Tethys Sea. This sea reached its maximum size during the Cretaceous, about I40-I35 Ma. Its growth ceased when Gondwanaland broke apart, beginning the creation of the Atlantic Ocean. During this period the Tethys linked the Gulf of Mexico to the northern Pacific

Within-Phylum Relationships Ocean. It was not until the end of the Mesozoic that the Bering land bridge separated the Arctic regions from subtropical waters (Dunton, I992). While there is some disagreement over the matter, depending on which data are more heavily weighted, the Tethys probably had a net eastward flow (Barron and Peterson, I989; P. Smith and Westermann, I990; Follmi et al., I99I). The Tethys shrank but persisted until the Miocene, about I8 Ma (earlier authors placed this at 36 Main the Oligocene), when Africa collided with Eurasia along a complex subduction line in the Mediterranean region. Fragments of the uplifted (obducted) Mesozoic Tethys Sea floor (I 8o85 Ma) appear as ophiolites along the Alpine chain between the western Mediterranean and the Indian Ocean. Much of the widespread Cretaceous Tethys marine fauna disappeared before the Miocene, but many Pacific islands functioned as refugia for some of these Mesozoic taxa (Kay, I 979). Elsewhere, in the late Jurassic (I65 Ma) Madagascar split off from East Africa in the region of Kenya-Somalia and moved southeast, arriving at its present position 125 Ma, in the early Cretaceous (Rabinowitz et al., I983). India began a longer move at about this time. It broke away from southeastern Africa in the Cretaceous (I45-I20 Ma), moved northward, and collided with Asia during the Eocene (50-40 Ma). However, its long isolation did not result in significant development of endemic marine species (Briggs, 1989). Cenozoic (65 Ma-Present) Much more information is available about the Cenozoic period. Rather than examining the entire planet, I focus below on subsets of the world ocean system and track events in those regions independently-keeping in mind their interdependence. First, however, I present an overview. On a geological time scale, the present array of temperatures and levels of the world's oceans are neither of long duration nor likely to remain as they are indefinitely. In other words, global warming and cooling are not new phenomena, as even a cursory inspection of the more recent literature on global dynamics shows. The overall pattern of temperature changes has been known for decadt;s (see Ekman, I967; Briggs, I974). For example, the Mesozoic polar seas were temperate (I 6- I 7°C), and the early Tertiary European Atlantic Ocean shelf fauna was clearly tropical, as evidenced by fossil remains of

Zoogeographical Data the reef-building corals, echinoderms, mollusks, etc. During the Miocene and Pliocene, however, there was a dramatic shift in the fauna to more temperate forms. This coincided with the growth of the polar ice caps, which had begun in the Oligocene, and the partitioning of the Tethys Sea, which resulted in the separation of the Indian and Atlantic oceans by the Mediterranean. The present thermally defined biogeographical provinces began to form around 50-40 Ma along with the polar ice but did not become well established with a polar fauna and cold, deep water until about 2 Ma (Berggren and Hollister, 1974; Benson et al., 1984). Cenozoic Temperature. An abrupt but short-lived deep-sea warming occurred 57 Ma at the end ofthe Paleocene (Kennett and Stott, 1991). During the Paleocene and Eocene (65-38 Ma) the surface water of the Antarctic was 13-14°C and the bottom water was about I0°C, significantly warmer than today's near-zero water (sea water freezes at -2°C}. The decoupling of the benthic and planktonic ecosystems is indicated by the extinction of 72% of the larger benthic foraminiferans and the lack of impact on their shallow-water counterparts. A similar warming and mass extinction of benthic forams occurred at the same time in the far northern Atlantic as the seaway opened between Norway and Greenland. There is evidence to suggest global warming, or a greenhouse effect, resulting from the buildup of C02 as a side effect of volcanic activity associated with plate tectonics. The rapid warming, which took less than 3000 years, resulted from salty, dense Tethys Sea water replacing the colder polar water present in the deep sea. The resulting drop in oxygen and rise in salinity and temperature formed a combination lethal for many taxa. The change in ocean circulation and the increasing instability of the water column's abiotic attributes might have resulted from an early greenhouse warming event. If benthic forams were so dramatically affected, it is hard to imagine that sipunculan populations survived unscathed, despite Vermeij's suggestion that warming generally causes fewer extinctions than comparable cooling (Vermeij, 1987). These events of 57 Ma were the reverse of the extinction events seen a relatively short time earlier, at the beginning of the Tertiary (K-T boundary, 65 Ma), when the shallow fauna was greatly affected and the deep fauna experienced very little change. Zinsmeister and Feldmann's (1984) historical analysis of five classes of marine invertebrates approaches the Tertiary ocean system from a different

Within-Phylum Relationships perspective. The authors noted an early warming trend in the southern mid-latitudes and suggested that the shallow Antarctic waters functioned as a Paleocene "holding tank" for ancestral taxa. During cooler times in the Oligocene and Miocene (38-5 Ma), these taxa migrated northward toward the equator, diversifying as they went. This latitudinal and taxonomic expansion continued more broadly into the Pliocene and Pleistocene, and some previously shallow taxa adapted to deeper habitats. The deeper Atlantic Ocean was clearly warmer than it is today until the early Oligocene (35 Ma), when Antarctic sea ice formed (Hammond, 1976). The growth of the Antarctic ice cap caused the cooling of the deep tropical western Pacific and a distinct change in benthic foraminiferan communities there (Woodruff et al., 1981). The Arctic surface water cooled to below soc about 12 Ma (Dunton, 1992). Cenozoic cooling was significant in the lower latitudes, especially in shallow waters, which dropped from around 25°C to around 15°C between 55 and 35 Ma (Valentine, 1984). Tropical waters remained cool until about 20 Ma, when they began warming back to the present temperature of 28300C. One important conclusion to be drawn from this information is that today's tropical regions are not geologically old; like high-latitude climates, they are mostly of Neogene age. Later in the Miocene (IO Ma), along the coast of Namibia (southwestern Africa), an upwelling of cold, nutrient-rich water provided a biologically productive environment. Microfossil populations indicate that this productivity dropped sharply at the end of the Miocene (Siesser, 1980), suggesting that most deep-sea species are of geologically recent age. Briggs (1974) supported this idea on the basis of the assumption that earlier, during the Mesozoic, the deep sea was largely anaerobic and therefore not a suitable habitat for most metazoans. The inflow of warm Tethian water from the Mediterranean into the deep Atlantic slowed and was finally cut off about 6 Ma during the Pliocene when the Iberian portal formed. The resultant cooling of the Atlantic in the east was complemented from the west when the Panamanian connection between the tropical Atlantic and Pacific closed (about 3·5 Ma). The subsequent Pleistocene glaciations between 3·5 and 1.8 Ma and the permanent sea ice at both poles accelerated the cooling process. As a result of this cooling, the deeper ostracods became more cosmopolitan in their distribution as they became separated from the more restricted shallow-water species. A similar historical sequence is likely for sipuncu-

Zoogeographical Data lans-that is, deeper-water species became widespread. Also during this time (in the Pliocene, 5-2 Ma) 50-75% of the North Atlantic bivalve species became extinct (Jablonski and Bottjer, 1991). Periodic glaciations continued during the Pleistocene (2-o Ma), separated by periods of warmth similar to present conditions. These repeated fluctuations resulted in local extinctions and disjunct distributions (Fleming, 1978). The sea ice and periodic glaciation in the Arctic between 2 and 0.7 Ma were major cladogenic forces (Dunton, 1992). During the late Cenozoic, oscillations between glacial and interglacial thermal and water circulation patterns occurred in the eastern Atlantic with a periodicity of 30,000 to 40,000 years (Jansen et al., 1986). A view of temperature and circulation patterns in the Atlantic (tropical and Caribbean), determined by examining fossil foraminiferans, indicates three incursions of cool water over the last 135,000 years (Prell et al., 1976). The impact of such climatic catastrophes on sipunculans may well have been dampened by the fact that they are low-energy infauna. Many species feed largely on decomposing organic material rather than depending directly on living plants, and they are generally small-bodied. Thus, they are less likely to be affected by short-term fluctuations and possibly less prone to extinction (see Vermeij, 1987). Cenozoic Sea Level. Ninety-six sea level changes occurred during the 6oo million years of the Phanerozoic eon, and these can be grouped into three levels of magnitude and frequency (Vail et al., 1978). The sea was at its highest level in the late Cretaceous (about 350m higher than present), creating extensive epicontinental seas. Before and after this the level was much lower-about 150-250 m below present levels in the early Jurassic, the mid-Oligocene, and the late Miocene. These fluctuations are attributed to geotectonic and glacial events. As a result of Quaternary glaciations, the Arctic Ocean experienced periodic changes of sea level every I0,000-20,000 years. The level dropped about 85 m each time, exposing large areas of continental shelf and resulting in local extinctions of shelf fauna (Dunton, 1992). Using fossils of Pleistocene ostracods and pollen from along the east coast of the United States as indicators, Cronin et al. (1981) identified at least five warm intervals over the past 500,000 years when the sea level was 6-7 m higher than present. The glaciated Canadian and New England Atlantic coast continued to undergo change during the past 15,000 years, and today's shallow subtidal

Within-Phylum Relationships configuration was not reached until about 3000 years ago (Bousefield and Thomas, 1975). During the previous 12,000 years the sea level was much lower and the temperature was cooler. Cenozoic Subregional Events The Atlantic Ocean. The Atlantic first appeared during the Cretaceous and achieved significant size about 100 Ma when the North and South Atlantic oceans merged at shallow depths. During the early Cenozoic, the Atlantic continued to spread outward from the Mid-Atlantic Ridge, further separating South America from Africa, and North America from Europe (Boucot and Gray, 1983). That spreading continues today. In the far north, between Norway and Greenland, the Atlantic became connected with the Arctic Ocean about 57 Ma. Deep-water connections between the North and South Atlantic did not form until about 50 Ma (Hammond, 1976). Mediterranean Sea and Northeastern Atlantic. The modem Mediterranean Sea is a remnant of the Tethys Sea. It began to form when the eastern end of the Tethys closed, about 18 Ma. The western end of the Mediterranean was at least partially closed by the formation of the Iberian portal about 6 Ma, near the end of the Miocene. This effectively cut off access to and from the eastern deep-water Atlantic Ocean (Keigwin, 1982; Benson et al., 1984). The Mediterranean may have been totally closed off between 6 and 5 Ma, when much evaporation occurred (the Messinian salinity crisis). The basin refilled when the Gibraltar gate opened at the end of this period (Hammond, 1976; de Weerdt, 1989). Shallow-water marine taxa that lived in the Tethys during Paleogene times could still persist today in both the eastern Atlantic-Mediterranean and the western Indian oceans (see Table 14). Although some taxa may have migrated around the tip of South Africa, it is safe to assume that most Indo-Atlantic species are older than 18 million years. Conversely, species now found on only one side of this 18-million-year-old land barrier probably evolved after the barrier formed; that is, are younger taxa. This last view is based on the assumption that there have been no abiotir or biotic changes since that time that were lethal to past generations of the taxa under study. Such local extinctions are known to have occurred in other marine taxa, and these restricted sipunculan populations may actually represent relicts of once broadly distributed, older species (see Valentine, 1984).

Zoogeographical Data South Atlantic. At the beginning of the Miocene (25 Ma), the Drake Passage opened between Antarctica and South America, allowing an eastward-flowing circumpolar current, an event that had global significance. The current gained in intensity until the Pleistocene (Fleming, I 978; Vermeij, 1991a). The fossil record of mollusks and echinoderms illustrates how, during the late Cenozoic, as temperatures cooled, some taxa with planktonic larvae successfully migrated from the eastern Atlantic and Mediterranean, across the Indian Ocean, and into the western Pacific. The same record shows how other taxa took advantage of this late Cenozoic cooling to migrate from western Europe or western North America southward past Africa or South America, eastward via the circumpolar current, and then northward past Australia to Japan (Fleming, 1978). Western Atlantic and Eastern Pacific. The connection between the westem Atlantic and eastern Pacific oceans closed during the Pliocene (about 3·5 Ma) when Central America uplifted and formed the Panamanian land barrier. Applying the same logic as was used above, taxa now on both sides of Central America may be presumed to have been in existence for more than 3·5 million years (see Table 14). The significance of taxa restricted to one side of this barrier is less certain, however, since the present habitats are different. On the Pacific side, for example, there is coastal upwelling of cold water, a paucity of coral reefs and sedimentary rock, and a scarcity of macroalgae. Nevertheless, in the absence of a fossil record, these patterns may be useful for dating cladogenic events. Thus, taxa on only one side of the Panamanian isthmus may be among the youngest in the phylum. One exception that may prove the rule was reported by Laguna (1987). Electrophoretic studies of two closely related trans-Panamic, endemic barnacle species show a greater genetic distance between the two than expected. The molecular clock suggests speciation in the upper Miocene, well before the formation of the land barrier. Laguna offered no mechanism by which this sympatric speciation might have occurred. After studying invertebrate taxa such as crabs and echinoderms, which have a fossil record that can be dated, Ekman (1967) asserted that there are more amphi-American genera than species, and that amphi-American species are demonstrably older than species found on only one side of the Panamanian barrier. If it is safe to extrapolate from benthic foraminiferans to sipunculans, then shelf-dwelling species are younger than deeper taxa along the east

Within-Phylum Relationships

370

Table 14. Sipunculans living in historically significant areas from shallow (1-100 m) or upper slope (100-1000 m) depths

1.

Eastern Atlantic and western Indian Oceans Antillesoma antillarum, Apionsoma murinae bilobatae,• Apionsoma trichocephalus, Apionsoma (Ed.) pectinatum, Aspidosiphon muelleri,b Aspidosiphon (Pa.) laevis, Aspidosiphon (Pa.) steenstrupii, Golfingia vulgaris,b Phascolosoma nigrescens, Phascolosoma perlucens, Phascolosoma stephensoni,b Phascolosoma (Fi.) capitatum,• Phascolion (Is.) convestitum,b Sipunculus nudus,b Sipunculus norvegicus•

A.

Eastern Atlantic and Mediterranean but not NW Indian Ocean Apionsoma murinae murinae,• Phascolosoma (Fi.) capitatum,• Aspidosiphon (Ak.) a/bus, Aspidosiphon (Ak.) venabulum, Aspidosiphon (Ak.) zinni,• Golfingia elongata, Onchnesoma steenstrupii,a.b Onchnesoma squamatum,•·b Phascolosoma granulatum, Phascolion (Is.) tuberculosum,•·b Phascolion (Le.) hupferi, Siphonosoma arcassonense,c Thysanocardia procerac

B.

NW Indian Ocean and/or Red Sea, but not Mediterranean or eastern Atlantic Apionsoma misakianum, Aspidosiphon coyi, Aspidosiphon elegans, Aspidosiphon gracilis, Aspidosiphon (Pa.) planoscutatus,c Cloeosiphon aspergillus, Lithacrosiphon maldivensis, Nephasoma rutilofuscum,c Phascolosoma albolineatum, Phascolosoma meteori,c Phascolosoma pacificum, Phascolosoma scolops, Phascolosoma (Fi.) lobostomum, Phascolion abnorme, Phascolion robertsoni, Phascolion (Le.) valdiviae sumatrense,• Phascolion (Vi.) cirratum,c Sipunculus longipapillosus, Sipunculus robustus, Sipunculus (Au.) indicus, Siphonosoma australe, Siphonosoma cumanense, Themiste (La.) lageniformis

2.

Both sides of Central America (all are also amphi-Pacific) Antillesoma antillarum,d Apionsoma misakianum, Apionsoma trichocephalus,d Apionsoma (Ed.) pectinatum,d Aspidosiphon (Pa.) fischeri, Lithacrosiphon cristatus, Phascolosoma nigrescens,d Phascolosoma perlucens,d Sipunculus nudus,d Sipunculus phalloides, Sipunculus polymyotus, Xenosiphon branchiatus

A.

Atlantic side of Central America, but not the Pacific Phascolosoma (Fi.) capitatum,• Aspidosiphon exiguus,c Aspidosiphon gosnoldi,c Aspidosiphon elegans, Aspidosiphon misakiensis, Aspidosiphon (Ak.) mexicanus, Aspidosiphon (Ak.) a/bus, Aspidosiphon (Ak.) zinni,• Aspidosiphon (Pa.) parvulus,c Golfingia elongata, Nephasoma pellucidum, Phascolion caupo,c Phascolion medusae,c Phascolion (Is.) microspheroidis,• Phascolion (Le.) cryptum,c Sipunculus norvegicus,• Sipunculus robustus, Siphonomecus multicinctus,c Siphonosoma cumanense, Themiste alutacea,c Themiste (La.) lageniformis, Thysanocardia catharinae•

B.

Pacific side of Central America, but not the Atlantic Aspidosiphon gracilis schnehageni,c Siphonosoma vastum • Primarily a slope species. bIn the Mediterranean also. cEndernic. d Also in group 1, above.

coast of the United States. Furthermore, northern North American species are likely to be younger than southern ones (Buzas and Culver, 1984). The idea that extinction and cladogenesis of higher taxa occur more rapidly in stressful habitats or where long-term stasis is disturbed is not

Zoogeographical Data

371

new (see Littler et al., 1985; Ross and Allman, 1991). The same principle may support the proposition that temperate, shallow-water species within eurytopic sipunculan genera are younger than those that live in stable habitats such as the tropics or the deep sea. This does not contradict the fact that stable areas have greater taxonomic diversity (see Sanders, 1968; Sanders and Hessler, 1969). The debate about the antiquity of taxa in the deep sea continues, but Vermeij (1987) is among those making a strong case for the deep sea serving as a haven for ancestral or relict species that were driven out of shallower habitats by biotic and/or abiotic forces. He pointed to the high number of adaptively anachronistic species that are defensively inferior and went on to assert that these deep-sea stocks do not serve as sources of genetic material for the reinvasion of shallower habitats. The infaunal habitat itself is a kind of haven, and deep-sea infaunal sipunculans thus are especially well protected. Endolithic animals (e.g., Lithacrosiphon or Cloeosiphon) are also well protected and are presumed to have evolved from infaunal ancestors. Indo-Malayan Region and the Pacific Ocean. An area of particular interest to marine biogeographers is the current Indo-West Pacific (IWP), especially the Indo-Malayan Archipelago, which is asserted to be the center of origin and dispersal for many taxa. If one traces this region back through geological time, one sees significant changes from the early Mesozoic, when it was the open western part of the Panthalassa. Australia to Southeast Asia. Of particular interest is the movement of Australia, which was part of the Antarctic land mass until the end of the Paleocene (55 Ma). New Zealand had broken off at least 5 million years before Australia began its 20-million-year journey northward through temperate seas. The separation from Antarctica became complete enough for deep currents to run south of Australia about 30 Ma. The Tasman and Coral Sea basins were formed by this time, and additional fragments of the Australian land mass began to splinter off. During the Eocene and Oligocene (45-29 Ma) the New Hebrides, Norfolk, and South Fiji basins formed; the North Fiji basin formed since the late Miocene, less than 10 Ma. Much ofthis activity involved Sumatra and Java as well as the Malay Peninsula (Grant-Mackie, 1978). The string of islands from Australia-Papuasia to Southeast Asia developed in conjunction with a southern movement of Southeast Asia (Mayr, 1988). It was not until the upper Miocene, however, less than 10 Ma, that the present broad, shallow shelf and archipelago configuration was in

372

Within-Phylum Relationships

place to form the incomplete and permeable boundary between the Pacific and Indian oceans (Newman, 1991). Thus, the Indian Ocean, as a region distinct from the western Pacific, is geologically recent. The formation of the Andaman Sea and the opening of the Sunda Strait (between Java and Sumatra) did not occur until 2 Ma. Therefore, any cladogenic events in this tropical archipelago cannot be much older than a few million years. An interesting observation about the exaggerated importance of the Indo-Malayan region as a center of origin of marine taxa was made by Ekman (1967:chap. 4), whose examination of the fossil record suggested that before the end of the Cretaceous there were no significant differences in diversity between the IWP and the Atlanto-East Pacific (AEP). The present-day difference is the result of significant climatic cooling experienced by the AEP during the Miocene, which led to local extinctions and emigration of many taxa. The IWP, which experienced no such trauma, preserved its earlier diversity and added to it in later times. This statement assumes a barrier between the western and eastern Pacific regions. Eastern Pacific. The Eastern Pacific Barrier (EPB) is acknowledged to be a very effective filter but not an impassible barrier. As an example, only 4% of the Hawaiian mollusks, and 16% of the Hawaiian coral fauna, reaches the west coast of the Americas (Vermeij, 1991a). The question of the EPB 's antiquity was addressed by Grigg and Hey (1992). Their analysis of Mesozoic and Cenozoic fossil corals showed that no barrier existed throughout the Cretaceous, and dispersal was from east to west. This dispersal was aided by stepping-stones-central Pacific islands that subsequently drowned (guyots). Hamilton's (1956) work on tropical corals provided the first demonstration of this Cretaceous phenomenon, but the complete explanation had to await an understanding of plate tectonics. The present island groups are not good stepping-stones, given their placement relative to the main current systems. Thus, the EPB did not exist before Cenozoic times, and isolation permitting allopatric speciation was less likely then, despite the wider ocean basin. North Pacific, Arctic, and Far North Atlantic. In the mid-Pliocene (43·5 Ma), when the Panamanian isthmus formed a barrier in the tropics, the Bering land bridge between Asia and North America was breaking up. The bridge, a barrier between the Pacific and Arctic oceans, had existed since 65 Ma. When it ceased to exist, a new migration route was opened. Most species seem to have migrated from the northern Pacific to the

Conclusions and Assumptions

373

Atlantic Ocean (Venneij, I99Ia, I99Ib; Dunton, I992). The Arctic Ocean fauna located between the two major oceans is young and of mixed origins. Most of the nearshore fauna (but not the flora) has Pacific ancestry. This youth (less than 3·5 million years) is attributed to repeated Pleistocene glaciations that were lethal to inhabitants (Dunton, I992).

Conclusions and Assumptions Based on the paleo-oceanographic information and the zoogeographical data summarized earlier and in Table 14, at least four of the five Phascolosomatidae genus groups are more than I 8 million years old, and probably much older (Phascolosoma [Fisherana] being a possible exception; i.e., younger). In the Aspidosiphonidae, only two of the three Aspidosiphon subgenera have pre-Miocene origins, but considerable cladogenesis has occurred since then. The subgenus A. (Akrikos) is less than 18 million and possibly less than 3 million years old. Lithacrosiphon evolved in the interval between I8 and 3 Ma, and Cloeosiphon is at least that young, probably first appearing less than 3 Ma (Fig. 89). Within the class Sipunculidea, it appears that Sipunculus is more than 18 million years old. Active cladogenesis occurred between I 8 and 3 Ma, including the genesis of the closely related Xenosiphon. The proposed early appearance (400 Ma) of Siphonosoma is not supported by these data unless one invokes local extinctions in the eastern Atlantic and eastern Pacific, where this genus is absent. The remaining genera of the order Sipunculifonnes-the monotypic Phascolopsis and Siphonomecus-are the youngest, probably less than 3 million years old. The Golfingiifonnes genera Golfingia and Phascolion are much older, from the Paleozoic, but speciation was probably common during the Neogene. Thysanocardia, Themiste, and many shallow-water Nephasoma appear younger, on the order of 3-I5 million years old. Most of the Nephasoma and Onchnesoma species are found in cold, deep water. It appears likely that while Nephasoma originated in the late Paleozoic, significant speciation in that genus and the first appearance of Onchnesoma probably occurred IS-3 Main the Neogene. In summary, I propose that the phylum Sipuncula had its origins in the earliest Paleozoic, and that by the late Paleozoic representatives of five of the six extant families existed, living in all of the then-available oceanic habitats. By the mid-Mesozoic, eight of the modem genera existed, and

374

Within-Phylum Relationships

this situation persisted until mid-Cenozoic times. By the end of the Miocene, all except three genera were present, and these appeared during the Pliocene (Fig. 89). Without a fossil record we cannot know whether the single species in a monotypic genus represents a remnant of a once polytypic genus or is the only one that ever existed. A genus with a single extant species may well have contained 10 species before the end of the Cretaceous. While the data cannot be used to propose extinction events or rates, there is no reason to believe that sipunculans are immune to the environmental changes that have had negative impacts on other benthic marine invertebrates. The fact that they are infaunal or endolithic animals may provide some protection, but one can only assume that what we see today is a very incomplete picture of the diversity present in the phylum throughout time.

20

Evolutionary Hypothesis

In this chapter items currently considered to have the opposite polarity of that presented in Cutler and Gibbs, 1985, are marked with an asterisk, and new items are marked with a double asterisk. What follows is a synthesis of all the material presented in this book.

Ancestor The revised hypothetical ancestral sipunculan (RHAS) had a body wall with a continuous longitudinal muscle layer and no epidermal coelomic extensions. The anterior end of the trunk bore the anus, was without a horny shield, and tapered into the introvert along the same axis. The introvert carried *regular rings of sculptured proteinaceous hooks with **basal spinelets, and the tentacular crown consisted of a *crescent of small nuchal tentacles plus a circumoral collar (cuticular fold) that was the precursor of a set of peripheral tentacles. Internally, the contractile vessel was small and did not have villi. Also present were two pairs of unfused, equal-sized introvert retractor muscles,

two nephridia, *possibly bilobed, and a complete spindle muscle *attached to the posterior end of the trunk. **The epidermal organs consisted of only one cell type and one secretory product. This Cambrian population lived in shallow, warm seas and **had 10 pairs of mostly telocentric chromosomes (2N = 20). The RHAS produced an egg with very little yolk and a thin egg envelope that developed into a trochophore. **This larva grew into a rather long-lived planktotrophic pelageosphera stage (type IV) and eventually settled to become a juvenile worm. Process A plausible and parsimonious evolutionary scenario beginning with the RHAS is shown as a dendrogram in Figure 89. That figure stops at the

Evolutionary Hypothesis Genus Extant species Holocene Pleistocene Quaternary Pliocene Miocene Oligocene Eocene Paleocene Tertiary CENOZOIC Cretaceous Jurassic Triassic MESOZOIC Permian Carboniferous Devonian Silurian Ordovician Cambrian Ediacarian PALEOZOIC

Li As Cl

An Ph Ap

2 19 1

I 16

Xe Si Sm Sh

Ps

Go Ne Ty

Th

Pn On

2 10 I

1

12 23 3

10

23 4

6

10

?

-(l)

(o)

(F)

(I)

-

(N)

(P)

(o)

(L)

-(H)

(M)f--1--(K)------' -(G)-

(B) (A)------1

RHAS

Abbreviations: Li = Lithacrosiphon, As = Aspidosiphon, Cl = Cloeosiphon, An = Antillesoma, Ph= Phascolosoma, Ap = Apionsoma, Xe = Xenosiphon, Si = Sipunculus, Sm = Siphonomecus, Sh = Siphonosoma, Ps = Phascolopsis, Go = Gol.fingia, Ne = Nephasoma, Ty = Thysanocardia, Th = Themiste, Pn = Phascolion, On= Onchnesoma.

Figure 89. Plausible historical representation of cladogenic events leading to extant sipunculan genera. The events occurred at nodes labeled with letters and are described in text.

generic level, but the text takes the process one step further, to subgenera. Although the scenario below is written as if it were fact, it is, at present, a working hypothesis. Early in the Paleozoic (node A on Fig. 89) a major cladogenic event resulted in the production of peripheral tentacles around the mouth and a reduction of nuchal tentacles. Concurrently the posterior spindle muscle shortened, terminating within the gut coil, and the epidermal glands became more complex with two types of cells and secretions. Significant alterations of the genetic material involving a replacement of telocentric chromosomes with chromosomes of more equal arm length (metacentric) also occurred. These changes led to a golfingiid ancestor of the class Sipunculidea.

Evolutionary Hypothesis

377

The group that retained the ancestral traits was the ancestor of the class Phascolosomidea (node A). Within this class, a major change associated with the occupation of new niches (empty mollusk shells, soft rock, and coral) was the development of a hardened, operculum-like shield at the anterior end of the trunk. At the same time this Ordovician stock experienced the loss of the dorsal retractor muscles and the loss of the basal spinelets on the hooks. Underlying these changes was the conversion of several telocentric to metacentric chromosomes. This gave rise to the ancestor of the order Aspidosiphoniformes, family Aspidosiphonidae (node B). The anterior trunk papillae, which produce the shield matrix, increased their activity, but not evenly, so that eventually the introvert axis shifted ventrally. The shield eventually came to consist of separate hardened, noncalcareous units (Aspidosiphon). Much later, a Neogene population lost the hooks in rings, giving rise to the subgenus A. (Akrikos), some of whose members remained hookless while others produced scattered hooks. Within the main stock of A. (Aspidosiphon), another line developed into the subgenus A. (Paraspidosiphon) when the longitudinal muscle layer split into more or less separate bundles during the Mesozoic. From a mid-Cenozoic member of this last subgenus a type of shield evolved in which the secreted material produced a single solid calcareous mass (Lithacrosiphon, node C). A Pliocene branch of the family lacking longitudinal muscle bands (LMBs) developed a very different anal shield made up of thick, separate, diamond-shaped, calcareous units dispersed in an ordered manner that allowed the introvert to remain on the same axis as the trunk (Cloeosiphon, node D). The main branch in the class Phascolosomidea retained more of the ancestral attributes and led to the order Phascolosomatiformes, family Phascolosomatidae (node B). The stock that changed least from the RHAS became the present-day genus Apionsoma. The development of muscle banding in some part of this stock led to the monotypic subgenus A. (Edmondsius). A split occurred when part of the early Paleozoic Apionsoma stock lost the basal spinelets on the hooks and the secondary nephridial lobes, thus leading to Phascolosoma (node E). An early dichotomy occurred within this genus when the nominate subgenus developed LMBs, leaving the small subgenus P. (Fisherana) with the plesiomorphic trait. The monotypic genus Antillesoma evolved during the Mesozoic from Phascolosoma by losing the adult hooks and gaining the linked attributes of contractile vessel villi and a larger array of tentacles (node F). Only modest changes in the developmental sequence occurred within

Evolutionary Hypothesis this class; most retained the type IV mode. Some species might have produced eggs with more yolk, had a shorter pelagosphera life, or both. The other main stock at node A was the ancestor to the class Sipunculidea. This Paleozoic stock split when one branch leading to the order Sipunculiformes (node G) experienced the partial division of the body wall muscles into anastomosing bands and developed epidermal organs that had not only two types of secretory cells and products but, eventually, separate sensory and secretory cells. Coelomic extensions into the epidermis began to develop, and a wide variety of chromosomal configurations appeared within the family (e.g., 2N ranged from 18 to 34). One part of the Mesozoic Sipunculidae stock (node H) reexpressed (or redeveloped) the posterior attachment of the spindle muscle, leading to Siphonosoma, which retains all four retractor muscles. Two monotypic genera developed from Neogene Siphonosoma populations. Siphonomecus resulted from the loss of the dorsal retractors. At about the same time (node I) Phascolopsis arose from another Siphonosoma ancestor whose circular muscle layer was still an undivided sheet with no coelomic extensions. In this stock the adult hooks were lost, leaving hooks in ill-defined rings only in juvenile worms, and larval stages were limited to a trochophore only. This cladogenesis probably occurred while the marine habitat was fluctuating during the late Cenozoic glaciations. The cooling forced the stenothermal warm-water Siphonosoma ancestor to retreat into warmer water, leaving behind this relict. This situation resembles that of the corals reported in Jablonski and Bottjer, I 991. It must be noted, however, that as good a case can be made for Phascolopsis having a golfingiid ancestor (see below). The Mesozoic clade that diverged at node H lost all hooks but developed distinct, separate longitudinal and circular muscle bands and more extensive epidermal coelomic canals. This line gave rise to Sipunculus, which developed the postesophagealloop. From the nominate stock developed the small subgenus S. (Austrosiphon), in which the nephridia shifted posterior to the anus and the anterior attachment of the spindle muscle shifted from the body wall to the rectum. This latter Neogene stock also gave rise to Xenosiphon by losing the postesophageal loop and changing the nature of the coelomic extensions (node J). Returning to node G, the Paleozoic Sipunculidea stock that did not develop LMBs was ancestral to the order Golfingiiformes. Within this group the chromosomal number remained more constant (2N = 20), but a greater variety of developmental options with shorter larval lives appeared at different times in various lineages (types III, II, and 1).

Evolutionary Hypothesis

379

The late Paleozoic Golfingiiformes stock split when one group lost the nuchal tentacles and one nephridium, the spindle muscle underwent vast reduction or complete loss, and the retractor muscles experienced significant fusion, leading to the family Phascolionidae (node K). A series of changes occurred within Phascolion that eventually led to five subgenera. Most of the changes involved the retractor muscles and probably are of Cenozoic age. The least derived extant taxon is P. (lsomya), which exhibits very little fusion between the equal-sized dorsal and ventral muscles. The nominate subgenus resulted from a significant reduction in the diameter of the fused ventral retractor to only half to one-fourth that of the dorsal. A significant amount of fusion of the dorsal and ventral retractors into an almost solid column led to the subgenus P. (Montuga). These three subgenera and part of P. (Lesenka) exhibit apomorphic scattered hooks. The remainder lack hooks altogether. The complete fusion of the retractors into a single muscle column produced the subgenus P. (Lesenka). From a part of this subgenus that shifted its anus out on the introvert, the monotypic subgenus P. (Villiophora) developed by adding contractile vessel villi. From some Miocene Phascolion stock, possibly a hookless P. (Lesenka), the genus Onchnesoma appeared (node L) when the dorsal retractors were lost, the ventral pair fused for almost their entire length, and the anus shifted out toward the distal end of the very long introvert. At node K, the Golfingiiformes stock that retained the spindle muscle and both nephridia gave rise to the family Golfingiidae. The main stock led to the modem genus Golfingia and its two subgenera: the monotypic G. (Spinata), which retains the plesiomorphic bilobed nephridia and hooks in rings with basal spinelets; and the nominate (but derived) subgenus, which has unilobed nephridia and some species with no hooks, some with scattered hooks, and a few with hooks in rings. This golfingiid branch divided during the late Paleozoic (node M) when a clade lost the dorsal retractor muscles, producing the ancestor to the diverse genus Nephasoma, which has many external morphological parallels to Golfingia, including the same amount of hook polymorphism. As I noted above, it is possible that Phascolopsis had a Neogene Golfingia with deciduous hooks as an ancestor; the only change required (at node N) is for the longitudinal musculature to partially divide into anastomosing bundles. From the Nephasoma stock two more groups arose. The monogeneric family Themistidae originated in the Neogene when unique stemlike extensions carrying dendritically branched peripheral tentacles appeared (node 0). The genus underwent rapid cladogenesis and divided into sub-

Evolutionary Hypothesis genera having two different types of contractile vessel villi: T. (Themiste), with a few long, threadlike tubules, unique in this phylum; and T. (Lagenopsis), with the more common numerous digitiform villi. The possibility that the genus is not monophyletic (i.e., that the subgenera might have had separate origins) is not out of the question, especially given the largely disjunct distributions. Finally (node P), Thysanocardia arose from a Neogene Nephasoma stock after acquiring contractile vessel villi. This clade developed an extensive tentacular crown with an elaborate array of peripheral tentacles in addition to well-developed nuchal tentacles. This chapter contains some speculation; nevertheless, it is based on a broad synthesis of the existing knowledge interpreted by a mind that has had 30 years of experience with thousands of these animals, living and dead. Most of the patches in this patchwork quilt are real. However, there may be other ways to arrange the patches to create different end results. This compendium of information and ideas is still incomplete, and there is the need for biologists to give more attention to this small, one might say peanut-sized, group of worms. Especially helpful would be the application of newer biochemical and genetic approaches by students of evolutionary biology. Relatively few phyla exist that are small enough to be treated in their entirety as a natural group. Much phylogenetic work is necessarily restricted to one family or order; not so here-this phylum is of manageable size. I hope that the clues presented here will encourage others to unravel the remaining evolutionary mysteries.

Appendix I Recent Species lnquirenda and lncertae Sedis

The following is a list of recent species inquirendo (A) and incertae sedis (B) determined since Stephen and Edmonds, 1972. The list is alphabetized by species name and includes only the original description, the ftrst use of subsequent combinations, and the publication where the current status was ftrst proposed. (A) Phascolosoma anguineum Sluiter, 1902:36. Golfingia anguinea.Stephen and Edmonds, 1972:85.-E. Cutler and Cutler, 1987a:756. (A) Phascolosomum approximatum Roule, 1898b:385. Golfingia (Golfingiella) approximata Stephen and Edmonds, 1972:II9.-E. Cutler et al., 1983:670. (B) Sipunculus bonhourei Herubel, 1904a:479· Siphonosoma bonhourei Stephen and Edmonds, 1972:64.-E. Cutler and Cutler, 1982:755. (B) Phascolion botulus Selenka, 1885:18. Phascolion botulum Stephen and Edmonds, 1972:173.-E. Cutler and Cutler, 1985a:838. (B) Diesingia Chamissoi de Quatrefages, I865b:6o6.-Saiz, 1984a:41. (B) Phascolosoma chuni W. Fischer, 1916:15. Golfingia chuni.-Stephen and Edmonds, 1972:136. Nephasoma chuni N. Cutler and Cutler, 1986:567. (B) Physcosoma corallicola ten Broeke, 1925:90. Phascolosoma corallicolum.-Stephen and Edmonds, 1972:298-299.-N. Cutler and Cutler, 1990:701. (A) Phascolosoma coriaceum Keferstein, 1865b:432-433. Golfingia (Thysanocardia) coriaceum Stephen and Edmonds, 1972:122. ?Themiste coriacea Gibbs et al., 1983:301-302. Herein, p. 141. (B) Diesingia cupulifera de Quatrefages, 1865b:6o7.-Saiz, 1984a:41. (A) Aspidosiphon cylindricus Horst, 1899:195-198.-E. Cutler and Cutler, 1989:837.

Appendix 1 (B) Phascolosoma delagei Herubel, 1903a:10o. Golfingia delagei.-Stephen and Edmonds, 1972:139-140. Nephasoma delagei N. Cutler and Cutler, 1986:567. (B) Physcosoma demanni Sluiter, 1891:121. Phascolosoma (Satonus) demanni Stephen and Edmonds, 1972:283.-E. Cutler and Cutler, 1983: 184. (A) Phascolosoma depressum Sluiter, 1902:39-40. Golfingia depressa.Murina, 1964a:227. Nephasoma depressum N. Cutler and Cutler, 1986:567. (B) Phymosoma falcidentatus Sluiter, 1881a:150. Physcosoma falcidentatus Sluiter, 1902:13. Phascolosoma (Satonus) falcidentatum Stephen and Edmonds, 1972:284.-E. Cutler and Cutler, 1983:185. (A) Phascolosoma fimbriatum Sluiter, 1902:34-35. Golfingia fimbriata. -Stephen and Edmonds, 1972:143. Nephasomafimbriatum N. Cutler and Cutler, 1986:567. (A) Phascolion ikedai Sato, 1930:20-23.-E. Cutler and Cutler, 1985a: 838. (A) Phascolosoma immunitum Sluiter, 1902:40. Golfingia (Siphonoides) immunita.-Murina, 1967c: 1334. Golfingia (Golfingiella) immunita.Cutler and Murina, 1977:180. Golfingia (Apionsoma) immunita.Cutler et al., 1983:670. Apionsoma immunitum Herein, p. 190. (A) Phascolosoma innoxium Sluiter, 1912:13. Golfingia (Golfingiella) innoxia.-Stephen and Edmonds, 1972:II9.-E. Cutler et al., 1983:671. (B) Sipunculus joubini Herubel, 1905b:51-54· Siphonosoma joubini Stephen and Edmonds, 1972:66.-E. Cutler and Cutler, 1982:757. (B) Phascolosoma lagense W. Fischer, 1895:13-14· Golfingia lagensis.Stephen and Edmonds, 1972:93.-E. Cutler and Cutler, 1987a:756. (A) Phascolosoma macer Sluiter, 1891:I14-II5. Golfingia macra.Stephen and Edmonds, 1972:149.-E. Cutler and Murina, 1977:183. Aspidosiphon macer.-N. Cutler and Cutler, 1986:568; E. Cutler and Cutler, 1989:838. (B) Phascolion manceps Selenka et al., 1883:44-45.-E. Cutler and Cutler 1985a:838. (B) Physcosoma mauritaniense Herubel, 1924:IIO. Phascolosoma (Satonus) mauritaniense Stephen and Edmonds, 1972:286.-E. Cutler and Cutler, 1983:186. (A) Phascolion moskalevi Murina, 1964b:255-256.-E. Cutler and Cutler 1985a:839.

Appendix 1 (A) Golfingia (Thysanocardia) neimaniae Murina, 1976:62-63. ?Themiste neimaniae Gibbs et al., 1983:302. Herein, p. 141. (B) Phymosoma nigritorquatum Sluiter, 188Ia:151-152. Physcosoma nigritorquatum.-Sluiter, 1902:13. Phascolosoma nigritorquatum.Stephen and Edmonds, 1972:286.-N. Cutler and Cutler, 1990:701. (A) Phascolosoma papilliferum Keferstein, 1865b:433. Fisherana papillifera.-Stephen and Edmonds, 1972:332. Golfingia (Apionsoma) papillifera.-E. Cutler, 1979:174-176.-Herein, p. 193. (A) Phascolion parvus Sluiter 1902:30-31. Phascolion parvum Stephen and Edmonds 1972:185.-E. Cutler and Cutler, 1985a:839. (A) Sipunculus pellucidus Sluiter, 1902:9-10. Siphonosoma pellucidum Stephen and Edmonds, 1972:69.-E. Cutler and Cutler 1982:758. (B) Dendrostoma pinnifolium Keferstein, 1865b:429. Themiste pinnifolia.-Stephen and Edmonds, 1972:209.-Gibbs and Cutler, 1987:53. (B) Phascolosoma quadratum Ikeda, 1905:170-171. Golfingia (Siphonides) quadrata Murina, 1967b:1335.-E. Cutler et al. 1983:673. (A) Phascolosoma reconditum Sluiter, 1900:II-12. Golfingia recondita. -Stephen and Edmonds, 1972:105. Golfingia (Apionsoma) recondita. -Cutler, 1979:372.-Herein, p. 193. (B) Phascolosoma reticulatum Herubel, 1925a:262. Golfingia reticulata. -Stephen and Edmonds, 1972:105.-E. Cutler and Cutler 1987a:756. (B) Phascolosoma rueppellii Griibe, 1868b:643. Physcosoma ruppellii Shipley, 1902:135. Phascolosoma (Rueppellisoma) rueppellii Stephen and Edmonds, 1972:275.-E. Cutler and Cutler, 1983:181. (B) Phascolosoma rugosum var. mauritaniense Herubel, 1925a:262. Golfingia (Golfingia) rugosa mauritaniensis Stephen and Edmonds, 1972:107.-E. Cutler and Cutler, 1987a:752. (A) Phascolion sandvichi Murina, 1974b:283-284.-E. Cutler and Cutler 1985a:839. (B) Onchnesoma Sarsii Koren and Danielssen, 1877:143-144. Phascolosoma Sarsii Theel, 1905:83. Golfingia sarsi Gibbs 1982:121. (B) Phascolosoma scutiger Roule, 1906:81-86. Golfingia scutiger.Murina, 1975c:I085-I08g.-E. Cutler and Cutler, 1987a:756. (A) Physcosoma sewelli Stephen, 1941b:405-407. Phascolosoma sewelli. -Stephen and Edmonds, 1972:276. ?Nephasoma sewelli E. Cutler and Cutler, 1983:181-182. (B) Dendrostoma spinifera Sluiter, 1902:41. Themiste spinifera.-Stephen and Edmonds, 1972:212.-E. Cutler and Cutler, 1988:741.

Appendix 1 (B) Phascolosoma vitreum Roule, 1898b:386. Golfingia vitrea.-Stephen and Edmonds, 1972:158-159. Nephasoma vitreum N. Cutler and Cutler, 1986:568. (A) Sipunculus zenkevitchi Murina, I969b:1733-1734·-E. Cutler and Cutler 1985b:240.

Appendix 2 Species lnquirenda and lncertae Sedis as in Stephen and Edmonds, 1972, with Current Status

Names are as presented in Stephen and Edmonds, I972:339-340, not always as in the original description. Sipunculus clavatus de Blainville, I 827-same. Sipunculus corallicolus Pourtales, I 85 I -same. Sipunculus echinorhynchus Delle Chiaje, 1823-same. Sipunculus gigas de Quatrefages, 1865b-Sipunculus nudus. Sipunculus glans de Quatrefages, I865b-Antillesoma antillarum. Sipunculus javensis de Quatrefages, 1865b-Phascolosoma noduliferum and P. pacificum (part in each). Sipunculus macrorhynchus de Blainville, I827-same. Sipunculus microrhynchus de Blainville, 1827-same. Sipunculus rapa de Quatrefages, I865b-Themiste hennahi. Sipunculus rubens Costa, 186o-same. Sipunculus rufojimbriatus Blanchard, I849-same. Sipunculus saccatus Linnaeus, 1767-same. Sipunculus vermiculus de Quatrefages, 1865b-Phascolosoma perlucens. Sipunculus violaceus de Quatrefages, I 865b--Siphonosoma vastum. Phascolosoma ambiguum (Brandt, 1835)-same. Phascolosoma carneum Leuckart and Rtippell, 1828-P. scolops. Phascolosoma cochlearium (Valenciennes, 1854)-Aspidosiphon muelleri. Phascolosoma constellatum de Quatrefages, I865b-same. Phascolosoma exasperatum Simpson, 1865-same. Phascolosoma fasciolatum (Brandt, I835)-same. Phascolosoma guttatum (Quatrefages, 1865b)-P. scolops. Phascolosoma johnstoni (Forbes, 1841)-same. Phascolosoma leachii (de Blainville, 1827)-same.

Appendix

2

Phascolosoma longicolle Leuckart and Riippell, I 828-Goljingia vulgaris. Phascolosoma loricatum (de Quatrefages, 1865b)-same. Phascolosoma nordfolcense (Brandt, I 835)-same. Phascolosoma orbiniense de Quatrefages, 1865b-Themiste alutacea. Phascolosoma placostegi Baird, 1868-nomen dubium. Phascolosoma plicatum (de Quatrefages, 1865b)-P. nigrescens. Phascolosoma pourtalesi (Pourtalt~s. 1851)-same. Phascolosoma pygmaeum (Quatrefages, I865b)-same. Phascolosoma semicinctum Stimpson, I855-same. Themiste ramosa (de Quatrefages, I865b)-T. hennahi. Themiste lutulenta (Hutton, 1879)-same. Aspidosiphon coyi de Quatrefages, I865b-now a valid senior synonym including A. truncatus. Aspidosiphon eremitus Diesing, 1859-A. muelleri. Aspidosiphon laevis de Quatrefages, 1865b-now a valid senior synonym for A. cuimingii, A. klunzingeri, and others. Aspidosiphon rhyssapsis Diesing, I 859-same.

Bibliography

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Taxonomic Index

Currently valid genus and species group names are in roman type. Names not currently considered valid senior synonyms, or archaic spellings of current names, are in italics. Subgenera are in parentheses, and names of suprageneric taxa are in capital letters. Descriptions of currently valid taxa begin on the pages listed; when more than one page number is given, the description begins on the page number in boldface type. abnorme, Phascolion, I I4, 126 abnorrnel -is, Phascolosoma/Goljingia, 198 absconditus, Xenosiphon, 43 abyssorum: Nephasoma, Phascolosorna, Golfingia, 78, 83, 89, 96 abyssorurn: Physcosorna, Phascolosorna, Apionsoma, I68 adelaidensis, Golfingia rnargaritacea, 7I adenticulaturn: Physcosoma scolops, Phascolosoma scolops, I84 adriatica, Golfingia, 76 (Aedematosornurn), Sipunculus, I88 aeneus, Sipunculus, 50 aequabilis, Sipunculus, 36 africanurn, Phascolion, I30 agassizii: Phascolosoma, Phymosoma, Physcosoma, I63, 173 (Akrikos), Aspidosiphon, 20I, 205, 211 a/berti, Phascolion, I 30 albidurn, Phascolosoma, 7I albolineatum: Phascolosoma, Phyrnosoma, Physcosoma, I63, 174, I84 albus, Aspidosiphon, 211, 2I5 alticonus, Lithacrosiphon, 229 alutacea/-urn, Themiste/Dendrostornum, 145, 148, I49. I5I amorniensisl -e, Sipunculus/ Siphonosoma, 53 arnbiguurn, Phascolosorna, 385 arnbonense: Phascolosoma, Physcosorna, 176 arnbonensis, Aspidosiphon, 227 arnbonensis: Aspidosiphon steenstrupii, Paraspidosiphon steenstrupii, 227 anceps, Phascolosoma, 93, 97 andamanensis, Phascolosorna, I74 anderssoni: Golfmgia, Phascolosorna, 63, 64, 65,68

angasii, Sipunculus, 39 anguineuml-a, Phosco/osomo/Golfingia, 38I angulatus, Aspidosiphon, 223 annulatum, Phascolosoma, 175 anomalus, Phascolion, 13I antarctica/ -urn, Goljingia margaritaceal Phascolosoma margaritacea, 72 antarcticum, Phascolosorna, 72 antillarum: Antillesoma, Phyrnosoma, Physcosoma, Phascolosoma, I 57. I 59. 186 Antillesoma, 186; antillarum, 157, I59, 186 (Antillesoma), Phascolosoma, I59. I86 Apionsoma, I57, I59, I67, 189, 194; abyssorum, I68; capitata, I68; irnrnuniturn, 190, 382; misakianum/-a, I95: murinae bilobatae, I 96; murinae murinae, I 95; papilliferurn, 193, 383; pectinatum, I57. 197; reconditurn, I 93; trichocephalus, I 96 (Apionsorna), Golfingia, I89 appendiculata/ -urn, Golfingia/ Phascolosoma, 74 approxirnaturnl -a, Phascolosoma/Gol.fingia, 381 arcassonense/-is, Siphonosoma/Sipuncu/us, 45.48,49 arcuatum/-us, Phascolosoma/Sipunculus, 163, 167, 176 armaturn, Aspidosiphon, 2 I 8 artificiosus/ -urn, Phascolion, I 30 aspergillus/-urn: Cloeosiphon, Loxosiphon/Echinosiphon, 232 asperurn: Phascolosoma, Phyrnosoma, 182 Aspidosiphon, 200; albus, 2II, 215; arnbonensis, 227; angulatus, 223; armatum, 218; brasiliensis, 223; brocki, 214; carolinus, 2I4; clavatus, 218; corallicola, 2I8; coyi, 221, 386; cristatus, 229; curningii, 222; cy-

Taxonomic Index

440 Aspidosiphon (cont.): lindricus, 38I; elegans, 205, 214, 226; e/egans elegans, 214; eiegans yapensel-is, 214; eremita/-us, 2I8, 386; exhaustum/-us, 2I8; exhaustus mirus, 2I9; exiguus, 2I5; exilis, 2 I4; exostomuml -us, 226; fischeri, 222; fischeri cubanus, 222; formosanus, 227; fuscus, 225; gerouidi, 2I7; gigas, 223; gosnoldi, 2I5; gracilis gracilis, 2I6; gracilis schnehageni, 2I6; grandis, 223; grandis obliquoscutatus, 223; hartmeyeri, 2I I, 2I7; havelockensis, 227; heteropsammiarum, 218; hispitrofus, 2I9; homomyarium/-us, 2I4; imbellis, 2I8; inquilinus, 2I8; insularis, I82; jukesii, 206, 2I8; kiunzingeri, 223; kovaleskii, 2I9; laevis/-e, 205, 22I, :z:z:z, 386; levis, 227; longirhyncus, 2 I :z; macer, 382; major, 223; makoensis, 225; mexicanus, 203, 205, 212; michelini, 2I8; mirabilis, 218; misakiensis, 217, 222; mokyevskii, I88; muelleri, 205, 218, 220, 23 I; ochrus, 226; pachydermatus, 223; parvulus, 2I5, 224; pygmaeus, 218; planoscutatus, 225; quatrefagesi, 223; ravus, 2I4; rhyssapsis, 386; rutilofuscus, 87; semperi, 225; speciosus, 223; speculator, 217, 225; spinalis, 214, 2I5; spinososcutatus, 224; spinosus, 214; spiralis, 220; steenstrupii, 22S, 227; steenstrupii fasciatus, 225; tenuis, 226; thomassini, 205, 212; tortus, 2I9; trinidensis, 226; truncatus, 22 I; uniscutatus, 229; venabulum/-us, 2I2; zinni, 203, 213 ASPIDOSIPHONIDAE, I99 ASPIDOSIPHONIFORMES, I99 aspidosiphonoides, Phascolosoma, 87 asser: Phymosoma, Physcosoma, Phascoiosoma, I88 astuta: Phascolosoma vulgare, Golfingia vulgaris, 75 australe, Siphonosoma, 45, so australe/-is, Phascolosoma!Sipunculus, 50 australel -is, Golfingia eremitalPhascolosoma eremita, 94, 99 (Austrosiphon): Sipunculus, Xenosiphon, 3I, 40

balanophorus, Sipunculus, 37 barentsii, Stephanostoma, 7I beklemischevi, Phascolion, I I7 benhami: Nephasoma abyssorum, Phoscolosoma, Golfingia, 90 bernnhardus, Sipunculus, I 30 billitonense/ -is, Siphonosomal Sipunculus, 52 bilobatae: Apionsoma murinae, Golfingia murinae, 196

birsteini, Golfingia, 62, 64, 65, 69 blanda/-um, Themiste/Dendrostomum, 149, I5I bogorovi, Phascolion, I I4, 127 boholense/-is, Siphonosoma/Sipunculus, 45,

so

bonhourei: Sipunculus, Siphonosoma, 38I borealel-is, Phascolosoma!Sipunculus, 94 botulusl-um, Phascolion, 381 branchiatus, Xenosiphon, 30, 35, 4I, 44 brasiliensis: Aspidosiphon, Paraspidosiphon, 223 brocki, Aspidosiphon, 214 brotzkajae, Phascolion, I 3 I bulbosum, Nephasoma, 81, 90 bulbosuml-a, Phascolosoma!Golfingia, 90 caementarius, Sipuncuius, I30 californica, Golfingia eremita, 94 californiensis, Golfingia margaritacea, 7I cantabriensis, Golfingia, 72 canum, Phascolion, I24 capensis/-e, Goifingia/Phascolosoma, 69 capilleforme/-is, Nephasoma/Golfingia, 69, 90. 97 capitata: Fisherana, Golfingia, Apionsoma, I68 capitatum: Phascolosoma, Physcosoma, I68 capitatus, Sipunculus, 130 capsiforme, Phascolosoma, 71 caribaeum, Xenosiphon, 44 carneum, Phascolosoma, I84, 385 carolinense, Siphonosoma, 52 carolinum, Cloeosiphon, 232 carolinus, Aspidosiphon, 2I4 catharinae: Thysanocardia, Phascolosoma, Golfingia, 103, 105, 106 caupo, Phascolion, I I4, I 22, 127 Centrosiphon, 6I; herdmani, 76 Chamissoi, Diesingia, 38 I charcoti: Phascolosoma, Golfingia, 70 chuni: Phascolosoma, Golfingia, Nephasoma, 38I cinctal-um, Golfingia/Phascolosoma, 93 cinereal -um, Golfingia/ Phascolosoma, 99 cirratum/-us, Phascolion, II4, IJ2 clavatus/ -um, Aspidosiphon/Pseudaspidosiphon, 2I8 clavatus, Sipunculus, 385 claviger, Sipunculus, 52 Cloeosiphon, 230; aspergillus/-um, 232; carolinum, 232; japonicum, 232; javanicum, 232; mol/is, 232 cluthensis, Phascolosoma, 70 cochlearium, Phascolosoma, 385 cochlearius, Sipunculus, 2I8 collare, Phascolion, I I4, 121

Taxonomic Index commune, Phascolosoma, 75 communis, Sipunculus, 75 concharum, Sipunculus, IJO confusum, Nephasoma, 78, 91 confusuml-a, Phascolosoma!Golfingia, 91 constellatum, Phascolosoma, 385 constricticervix, Nephasoma, 78, 8r, 84, 91, 93, IOI constrictum, Nephasoma, 83, 84, gr, 92 constrictuml-a, Phascolosoma!Golfingia, 92 (Contraporus), Sipunculus, 40 convestitum/-us, Phascolion, II4, 117 corallicola, Aspidosiphon, 218 corallicolal -urn, Physcosomal Phascolosoma, 381 corallicolus, Sipunculus, 385 coriacea, ?Themiste, 38 I coriaceuml -a, Phascolosoma/Golfingia, 99, 154. 381 corrugatum, Nephasoma diaphanes, 94 coyi, Aspidosiphon, 221, 386 crassum, Siphonosoma, 55 cristatus, Lithacrosiphon/Aspidosiphon, 229 cronullae, Phascolion strombus, I 3 I cryptum/-us, Phascolion, I 14, I 15, 122 cubanus, Aspidosiphon fischeri, 222 cumanense: Siphonosoma, Phoscolosoma, Sipunculus, 45, 48, 51, 54 cumingii, Aspidosiphon, 222 cupulifera, Diesingia, 38 I (Cutlerensis), Nephasoma, 78, 87 cutleri: Nephasoma, Golfingia, 69, 78, 92 cylindrata/ -urn, Golfingia/ Phascolosoma, 70 cylindricus, Aspidosiphon, 38 r cymodoceae: Themiste, Dendrostomum, 143, 152 (Dasmasiphon), Siphonosoma, 44 dayi, Siphonosoma, 45, 50, 53 deani: Phymosoma, Phascolosoma, 176 deformis, Sipunculus, 51 dehamata/-um, Themiste/Dendrostomum, 143. 153 delagei: Phascolosoma, Golfingia, Nephasoma, 70, 382 delphinus, Sipunculus, 37 demanni: Physcosoma, Phascolosoma, 382 Dendrostoma!-um, 140; alutaceum, 148; blandum, 151; cymodoceae, I52; dehamatum, 153; dyscritum, 149; ellipticum, 153; fisheri, 153; fuscum, 154; hexadactylum, 151; huttoni, rs6; lissum, 150; minor, 154; mytheca, rso; perimeces, rso; peruvianum, I 50; petraeum, I 5 I; petrico/urn, 148; pinnifolia, 383: pyroides, rsr; ramosum, 150; robertsoni, 154; rosaceum, 148; schmitti, 150; signifer, 153; spinifer,

441 383; stephensoni, 69, I54: tropicum, I54: zosterico/um, 150 dentalii, Sipunculus, I 30 dentalico/a/ -urn, Phascolion, II 8 dentigerum: Phascolosoma, Physcosoma, 182 depressa, Golfingia, 382 depressum: Phascolosoma, Nephasoma, 382 derjugini: Phascolosoma, Golfingia, 70 diaphanes: Nephasoma, Phascolosoma, Golfingia, 84, 89, 93, 97, roo diaphanes: Phymosoma, Phasco/osoma, r8o Diesingia, 381; Chamissoi, 381; cupulifera, 38I digitatum, Phasco/osoma, 94 diptychus, Sipunculus titubans, 37 discrepans, Sipunculus, 40 dissors, Phascolosoma, 193 dogieli, Phascolion, r 38 dubium, Phascolosoma, 75 dunwichi, Phascolosoma, I84 duplicigranulatum: Phascolosoma, Phymosoma, Physcosoma, r8o (Dushana), Golfingia, 6r dyscrita/-um, Themiste/Dendrostomum, r 49 echinorhynchus, Sipunculus, 385 Echinosiphon aspergillum, 232 (Edmondsius): Apionsoma, Phascolosoma, 159, 167, Igo, 197 edule, Siphonosoma, 51 edulis: Sipunculus, Lumbricus, 51 e/achea, Golfingia, 94 elegans: Aspidosiphon, Sternapsis, Sipunculus, Loxosiphon, Phascolosoma, 205, 214, 226 elisae: Nephasoma wodjanizkii, Golfingia, Nephasoma, 78, 102 elliptical -urn, Themiste/ Dendrostomum, I 53 elongata/-um, Goifingia/Phascolosoma, 6r, 62, 64, 69. 100 eniwetoki, Siphonosoma, 54 eremita: Nephasoma, Sipunculus, Phascolosoma, Golfingia, 78, go, 94, g8 eremital-us, Aspidosiphon, 218, 386 esculenta/ -urn, Physcosomal Phascolosoma,

I?6 evisceratum: Phascolosoma, Physcosoma, r8o exasperatum, Phascolosoma, 385 exhaustuml-us, Aspidosiphon, 219 exiguus, Aspidosiphon, 2 I 5 exilis, Aspidosiphon, 2 I4 eximioclathratus, Sipunculus, 37 exostomum!-us, Aspidosiphonl Paraspidosiphon, 226 extortum: Phascolosoma, Physcosoma, r8o

442 falcidentatum, Phascolosoma, 382 falcidentatus: Phymosoma, Physcosoma, 382 farcimen, Lesinia, 2 I 8 fasciatum, Phascolosoma, 178 fasciatus: Aspidosiphon steenstrupii, Paraspidosiphon steenstrupii, 225 fasciolatum, Phascolosoma, 385 filifonne, Nephasoma, 84, 95 filiformel-is, Phascolosoma!Golfingia, 95 fimbriatal-um, Golfingia!Nephasoma, 382 fimbriatum, Phascolosma, 382 finmarchica: Phascolosoma, Golfingia margaritacea, 7 I fischeri, Aspidosiphon, 222 Fisherana, 159, 167; capitata, r68; lobostoma, r6g; papillifera, 193, 383; wasini, 169 (Fisherana), Phascolosoma, Golfingia, 159, 167 fisheri: Dendrostomum, Themiste, 153 flagriferum, Nephasoma, 6o, 68, 81, 83, 84, 90, 95 flagriferum!-a, Phascolosoma/Golfingia, 95 flavus, Sipunculus, 178 forbesi, Phascolosoma, 70 formosanus: Aspidosiphon, Paraspidosiphon, 227 formosense: Phascolosoma, Physcosoma, I73 formosum, Siphonosoma, 52 fulgens, Phascolosoma, ?I funafuti: Siphonosoma, Sipunculus, 45, 48, 51, 53 funafutiense: Physcosoma, Phasco/osoma, 177 fuscal-um, Themiste! Dendrostomum, I 54 fuscum, Phascolosoma, 72, r88 fuscus: Aspidosiphon, Paraspidosiphon, 225 galapagensis, Sipunculus, 38 gaudens: Physcosoma, Phascolosoma, I88 genuensis, Sipunculus, 178 georgianum, Phascolosoma, 72 gerardi, Phascolion, I 17 gerouldi, Aspidosiphon, 2I7 gigas: Aspidosiphon, Paraspidosiphon, 223 gigas, Sipunculus, 37, 385 glabrum: Phascolosoma, Physcosoma, I63, I64, 177 glaciale: Onchnesoma, Phascolosoma, 96 glacialis, Golfingia, 90, 96 glans: Sipunculus, Phascolosoma, r88, 385 glauca: Golfingia, Themiste, 154 glaucum: Phascolosoma, Physcosoma, 154, I73 glossipapillosa!-um,

Taxonomic Index Golfingia! Phascolosoma, 72 Golfingia, 61, 67, 68, 159; abnormis, Ig8; abyssorum, 89; adriatica, 76; anderssoni, 63, 64, 65, 68; anguinea, 38I; appendiculata, 74; approximata, 38I; benhami, 90; birsteini, 62, 64, 65, 69; bulbosa, 90; cantabriensis, 72; capensis, 69; capilleformis, 90; capitata, I68; catharinae, 103; charcoti, 70; chuni, 381; cincta, 93; cinerea, 99; confusua, 91; confusa zarenkovi, 91; constricta, 92; constricticervix, 9I; coriaceal-um, 99, 140, 154, 381; cutleri, 92; cylindrata, 70; delagei, 382; depressa, 382; derjugini, 70; diaphanes, 93; elachea, 94; elisae, 102; elongata, 61, 62, 64, 69, roo; eremita, 94; eremita australe, 94, 99; eremita californica, 94; eremita scabra, 94; filiformis, 95; fimbriata, 382; flagrifera, 95; glacialis, go, g6; glauca, 154; glossipapillosa, 72; hespera, 195, 196; hozawai, ros; hudsoniana, 74: hyugensis, ros; immunita, 190, 382; improvisa, 93. 97; incomposita, 89; iniqua, 64, 70; innoxia, 382; kolensis, 76; /aetmaphila, 96; lagensis, 382; lanchesteri, 154; lilljeborgii, 96; liochros, n; lobostoma, r6g; longirostris, 195; macginitiei, ros; mackintoshii, 76; macra, 382; margaritacea, 69, 71, 74; margaritacea adelaidensis, 71; margaritacea antarctica, 72; margaritacea californiensis, 7 I; margaritacea finmarchica, 71; margaritacea hanseni, 71; margaritacea ikedai, 72; margaritacea meridiana/is, 72; margaritacea ohlini, 73; margaritacea sibirica, 71; margaritacea trybomi, 72; martensi, 103; mawsoni, 73; mexicana, 212; minuta, 88, 89, 93, 97, roo; mirabilis, 62, 65, 74; misakiana, 195; makyevskii, r88; mucida, 95; multiaraneusa, 98; muricaudata, 62, 64, 65, 74; murinae, 195; murinae bilobatae, 196; murinae unilobatae, 195; mutabilis, 7I; neimaniae, 141, 382; nicolasi, ror; nigra, IOS; nordenskjoldi, 72; nota, 72; novaezealandiae, 98; okinoseana, 72; onagawa, 105; owstoni, 76; papillifera, I93. 383; pavlenkoi, 105; pectinatoides, 62, 64, 67; pellucida, 98; procera, ro6; profunda, 72; pudica, 73: pugettensis, ros; pusil/a, I97; pyriformis, 154; quadrata, 383; recondita, I93, 383; reticulata, 383; rimicola, roo; rugosa, 70; rugosa mauritaniensis, 383; rutilofusca, 87; sanderi, 76; sarsii, 383; savalovi, I68; schuettei, roo; scutiger, 383; sectile, 93; semperi, 103; sewelli, 383; signa, 72; sluiteri, 99; solitaria, 76;

Taxonomic Index soya, 72; tasrnaniensis, wo; tenuissirna, I95: trichocephala, I96; verrillii, 99; vitjazi, IOI; vitrea, 384; vulgaris, 62, 65, 74, 7s; vulgaris astuta, 75; vulgaris herdmani, 62, 76; vulgaris rnultipapillosa, 75; vulgaris rnurinae, 73; vulgaris queenslandensis, 77; vulgaris selenkae, 75; vulgaris tropica, 75; vulgaris vesiculosus, 75; wodjanizkii, I o I; zenibakensis, I 05 GOLFINGIAFORMES, 6o (Golfingiella), Golfingia, 6I GOLFINGllDAE, 6o GOLFINGIIFORMES, 6o golikovi, Phascolosorna, I74 gosnoldi, Aspidosiphon, 2 I 5 gouldii: Phascolopsis, Sipunculus, Phascolosorna, Golfingia, 57 gracilis/ -e, Aspidosiphon/ Pseudaspidosiphon, 2I6 grandis: Aspidosiphon, Paraspidosiphon, 223 granulatum: Phascolosoma, Physcosorna, I77 gravieri, Sipunculus, 39, 43 grayi, Phascolosorna, I8I gurjanovae, Lithacrosiphon, 229 guttatus/ -urn, Sipunculus/ Phascolosorna, I84, 385 harnulaturn, Phascolosorna, I30 hanseni: Phascolosorna rnargaritacea, Go/fingia rnargaritacea, 7I hartrneyeri, Aspidosiphon, 2rr, 2I7 harveyi: Syrinx, Phascolosorna, 75 hataii, Siphonosorna, 52 havelockensis, Aspidosiphon, 227 hawaiense, Siphonosorna, 54 hebes: Physcosorna, Phascolosorna, 52 hedraeum, Phascolion, I I4, 118 hennahi, Themiste, I49, 150 herdmani: Centrosiphon, Golfingia vulgaris, 62,76 heronis, Phascolosorna, I85 herouardi, Physcosorna, I78 hespera: Phascolosorna, Go/fingia, I95, I96 (Hesperosiphon), Siphonosorna, 44 heterocyathi, Sipunculus, 2I8 heteropapillosurn, Phascolion, I 27 heteropsarnrniarurn, Aspidosiphon, 2I8 hexadactylal -urn, Therniste/ Dendrostornurn, I5I hibridum/-us, Phascolion, rr4, 128 hirondellei, Phascolion, I20 hispitrofus, Aspidosiphon, 2I9 Hornalosorna laeve, 7I hornornyariurnl-us, Aspidosiphon, 2I4 horsti: Physcosorna, Phascolosorna, I8o hozawai: Phascolosorna, Golfingia, I05

443 hudsoniana/ -urn, Go/fingial Phascolosorna, 74 hupferi, Phascolion, I I4, 122 huttoni: Themiste minor, Phascolosorna, Dendrostornurn, I54, 155 hyugensis: Phascolosorna, Go/fingia, I05 ikedai, Golfingia rnargaritacea, 72 ikedai, Phascolion, 382 irnbellis, Aspidosiphon, 2I8 irnrnodestus/ -urn, Sipunculusl Phascolosorna, I88 irnrnuniturnl-a, Phascolosorna/Go/fingia, I90, 382 irnprovisual -urn, Golfingial Phascolosorna, 93.97 inclusus, Sipunculus phalloides, 38 incornposita, Golfingia, 89 incornpositurn: Phascolosorna, Nephasorna, 89 indicus, Lithacrosiphon, 229 indicus, Phascolion, I22 indicus, Sipunculus, 30, 40 indicus, Xenosiphon (Xenopsis), 40 infrons, Sipunculus, 36 ingens: Siphonosoma, Siphonornecus, 45, 48, 53. 54 iniqua/-urn, Golfingia/Phascolosorna, 64, 70 innoxiurn/-a, Phascolosorna!Go/fingia, 382 inquilinus, Aspidosiphon, 2I8 insularis: Aspidosiphon, Paraspidosiphon, I82 interrnedial -urn, Phascolionl Phascolosorna, I30 intermedium, Onchnesoma, I34. 136 (lsomya), Phascolion, I I6 japonicurn, Cloeosiphon, 232 japonicurn: Phascolosorna, Phyrnosorna, Physcosorna, I73 javanicurn, Cloeosiphon, 232 javenense/ -is, Phascolosorna/ Sipunculus, I8I, I82, 385 jejfreysii, Phascolosorna, I78 johnstoni, Paraspidosiphon, 223 johnstoni, Phascolosorna, 385 joubini: Sipunculus, Siphonosorna, 382 jukesii, Aspidosiphon, 206, 2 I 8 kapalurn, Phascolosorna, I85 klunzingeri, Aspidosiphon, 223 kolensel-is, Phascolosorna/Golfingia, 76 koreae, Siphonosorna, 52 kovaleskii, Aspidosiphon, 2I9 kukenthali, Lithacrosiphon, 229 kurchatovi, Phascolion, rr8

444 kurilense: Phascolosoma agassizii, Physcosoma, I74

lacteum: Phasco/osoma, Phymosoma, Physcosoma, I 8o laetmophilum/-a, Nephasoma/Goifingia, 78, 96 laeve, Homalosoma, 7I laeve, Phascolosoma, I78, I85 laevis/-e, Aspidosiphon, 205, 22I, 222, 386 lageniformis, Themiste, I43, I44, 153 (Lagenopsis), Themiste, I42, I43. I44, 152 /agensel-is, Phascolosoma/Goifingia, 382 lakshadweepensis, Lithacrosiphon cristatus, 230 lanchesteri, Goifingia, I54 lanzarotae, Physcosoma, I78, I85 leachii, Phascolosoma, 385 (Lesenka), Phascolion, I 2 I Lesinia farcimen, 2 I 8 levis: Aspidosiphon, Paraspidosiphon, 227 levis, Sipuncu/us, I78 lilljeborgi: Nephasoma, Phascolosoma, Goifingia, 83, 89, 93, 96 liochros, Goifingia, 77 lissal-um, Themiste/Dendrostomum, ISO Lithacrosiphon, 227; alticonus, 229; cristatus, 229; cristatus lakshadweepensis, 230; gurjanovae, 229; indicus, 229; kukenthali, 229; maldivensis, 230; odhneri, 229; poritidis, 229; uniscutatus, 229 lobostoma: Phascolosoma, Fisherana, Goifingia, I69 lobostomum, Phascolosoma, I 69 lomonossovi, Sipunculus, 33 longico/le, Phascolosoma, 75, 386 longipapillosus, Sipunculus, 25, 27, 30, 3I, 35. 43.44 /ongirhyncus, Aspidosiphon, 2I2 longirostris, Goifingia, I95 Iordi, Phymosoma, I73 /oricatum, Phascolosoma, 386 loveni, Phascolosoma, 178 Loxosiphon aspergillus, 232; elegans, 2I4 lucifugax, Phascolion, I I4, 119 Lumbricus edulis, 5I; phalloides, 38 lurco: Phascolosoma, Phymasoma, Physcosoma, 176 lutense, Phascolion, I09, I 14, I I5, 124 luteum, Phascolosoma, 75 lutulenta, Themiste, 386 lytkenii, Tylosoma, I20 macer: Phasco/osoma, Aspidosiphon, 382 macginitiei, Goifingia, 105 mackintoshii, Goifingia, 76

Taxonomic Index macra, Golfingia, 382 macrorhynchus, Sipunculus, 385 maculatum: Phascolosoma, Phymosoma, Physcosoma, I78 magnibathum/-a, Onchnesoma, I34, 137 major, Aspidosiphon, 223 makoensis: Aspidosiphon, Paraspidosiphon, 225 malaccensis, Phascolosoma arcuatum, I76 ma/accensis: Phymasoma lurco, Physcosoma lurco, 176 maldivensis, Lithacrosiphon, 230 manceps, Phascolion, 382 maoricus, Sipunculus, 41 marchadi, Siphonosoma, 52 marcusi, Sipunculus, 30, 35 margaritacea: Golfingia, Phascolosoma, 69, 71 margaritaceum: Golfingia, Phascolosoma, 7I margaritaceus, Sipuncu/us, 7I marinki, Nephasoma, 95, 96 martensi: Phascolosoma, Goifingia, I03 mauritaniensel -is, Phascolosoma rugosum/Goifingia rugosa, 383 mauritaniense: Physcosoma, Phascolosoma, 382 mawsoni: Phascolosoma, Goifingia, 73 mediterraneum, Phascolion, I I7 medusae, Phascolion, 114, 115, 128 megaethi, Phascolion, I I4, 128 meridionalis: Phascolosoma margaritacea, Goifingia margaritacea, 72 meteori: Phascolosoma, Phymosoma, Physcosoma, r6I, 179 mexicanus/-a, Aspidosiphon/Goifingia, 203, 205, 212 michelini, Aspidosiphon, 2 I 8 microdentigerum: Phascolosoma, Physcosoma, I82 microdontoton: Phascolosoma, Phymosoma, I74. I77 microrhynchus, Sipunculus, 385 microspheroidis/-e/-es, Phascolion, 114, 119 minor: Themiste, Dendrostomum, I4I, 154 minuta, Goifingia, 97 minutum: Nephasoma, Phascolosoma, Petalosoma, 88, 89, 93, 97, IOO minutum: Physcosoma, Phascolosoma, I8o mirabilis, Aspidosiphon, 2 I 8 mirabilis, Golfingia, 62, 65, 74 mirus, Aspidosiphon exhaustus, 2I9 misakiana: Apionsoma, Goifingia, 195 misakianum: Apionsoma, Phascolosoma, I95 misakiensis, Aspidosiphon, 214, 222 (Mitosiphon), Goifingia, I89, I90, I94 mogadrense, Phascolion, I30

Taxonomic Index mokyevskii: Golfingia, ?Aspidosiphon, 188 mollis, Cloeosiphon, 232 (Montuga), Phascolion, 124

moskalevi, Phascolion, 382 mossambiciense: Phymosoma scolops, Physcosoma scolops, Phascolosoma scolops, 175. 183 mourense, Siphonosoma, 45, 54

mucidal -urn, Golfingial Phascolosoma, 95

muelleri, Aspidosiphon, 205, 218, 220, 231

multiannulata, Phascolosoma, 174, 177

multiaonulatum, Phascolosoma glabrum, 177 multiaraneusa, Nephasoma, 78, 98 multicinctus, Siphonomecus, 56

multipapillosalum, Golfingia vulgaris/Phascolosoma vulgare, 75 multisulcatus, Sipunculus, 38 multitorquatus, Sipunculus, 178 mundanus: Sipunculus, Xenosiphon, 30, 40 muricaudata/-um, Golfingia/Phascolosoma, 62, 64, 65, 74 murinae: Apionsoma, Golfingia, 195 murinoe, Golfingia vulgaris, 73 murrayi, Phascolion, 123 mutabile/-is, Phascolosoma/Golfingia, 71 mytheca, Dendrostomum, 150 nahaense: Phymosoma, Phascolosoma, 184 natans, Sipunculus, 39 neimaniae: Golfingia, ?Themiste, 141, 383

Nephasoma, 77, 88; abyssorum, 78, 83, 89, 96; abyssorum benhami, 90; bulbosum, 81, 90; capillefonne, 69, 90, 97; chuni, 381; confusum, 78, 91; constricticeiVix, 78, 81, 84, 91, 93, 101; constrictum, 83, 84, 91, 92; cutleri, 69, 78, 92; delagei, 382; depressum, 382; diaphanes, 84, 89, 93, 97, 100; diaphanes corrugatum, 94; eremita, 78, 90, 94, 98; filifonne, 84, 95; fimbriatum, 382; flagriferum, 6o, 68, 81, 83, 84, 90, 95; incompositum, 89; laetmophilum, 78, 96; lilljeborgi, 83, 89, 93, !)6; marinki, 95, 96; minutum, 88, 89, 93, 97, 100; multiaraneusa, 78, 98; novaezealandiae, 78, 84, 98; pellucidum, 92, 98, 99, Ioo; pellucidum subhamatum, 99; rimicola, 78, 100; rutilofuscum, 78, 84, 87; schuettei, 78, 94, 100; sewelli, 383; tasmaniense, 84, 100, 10 I; vitjazi, 78, 83, 84, 101; vitreum, 384; wodjanizkii, 83, 84, 101; wodjanizkii elisae, 78, 102 nicolasi, Golfingia, 101 nigra, Thysanocardia, 103, 105 nigral-um, Golfingia/Phascolosoma, 105 nigrescens: Phascolosoma, Phymasoma, Physcosoma, 161, 163, 179

445 nigriceps, Phascolosoma, 188 nigritorquatum: Phymosoma, Physcosoma, Phascolosoma, 383 nitidus, Sipunculus, 36 noduliferum/-us, Phascolosoma/Sipunculus, 164. 181

nodulosus, Sipunculus, 181 nordenskjoldi: Golfingia, Phascolosoma, 72 nordfolcense, Phascolosoma, 386 noiVegicus, Sipunculus, 30, 35, 36

notal -o, Golfingia/ Phascolosoma, 72 novaepommeraniae: Sipunculus, Siphonosoma, 52

novaezealandiae, Nephasoma, 78, 84, 98 nudum, Onchnesoma steenstrupii, 134, 139 nudus, Sipunculus, 30, 31, 35. 36, 39 nudus, Syrinx, 37 nudus, Xenosiphon branchiatus, 43

obliquoscutatus, Aspidosiphon grandis, 223 obscurum/-us, Phascolosoma/Sipunculus, 70, 75

ochrus, Aspidosiphon, 226 odhneri, Lithacrosiphon, 229 oerstedii, Phascolosoma, 71

ohlini: Golfingia margaritacea, Phascolosoma, 73

okinoseana/ -urn, Golfingia/ Phascolosoma, 72

oligopapillosum, Onchnesoma squamatum, 134. 138 onagawa: Phascolosoma, Golfingia, 105 Onchnesoma, 133; glaciale, 96; intennedium, 134, 136; magnibathum/-a, 134, 137; sarsii, 383; squamatum, 134, 136, 137; squamatum oligopapillosum, 134, 138; steenstrupii, 133, 134, 138; steenstrupii nudum, 134. 139

onomichianum: Phymosoma, Physcosoma, Phascolosoma, 188 opacal-um, Siphonosoma cumanense, 51 opacus, Sipunculus cumanense, 51 orbiniense, Phascolosoma, 148, 386 orbiniensis: Sipunculus, Themiste, 148, 386 owstoni: Golfingia, Phascolosoma, 76 oxyurum, Phascolosoma, 70

pachydermatus, Aspidosiphon, 223 pacificum, Phascolion, 114, 125 pacificum: Phascolosoma, Phymosoma, Physcosoma, 161, 164, 177, 181 pallidum, Phascolion, 120 papillifera: Fisherana, Golfingia, 193, 383 papilliferum: Apionsoma, Phascolosoma, 193. 383

papillosum, Sipunculus, 75, 178

Taxonomic Index Paraspidosiphon, 20I, 22I; ambonensis, 227; angulatus, 223; brasiliensis, 223; cumingii, 222; exostomus, 226; jischeri, 222; jischeri cubanus, 222; formosanus, 227; gigas, 223; grandis, 223; insularis, I 82; johnstoni, 223; klunzingeri, 223; levis, 227; makoensis, 225; pachydermatus, 223; pygmaeus, 2I8; schnehageni, 2I6; semperi, 225; speciosus, 223; speculator, 2I7, 225; spinososcutatus, 224; steenstrupii, 225; steenstrupii fasciatus, 225; tenuis, 226; trinidensis, 226; truncatus, 22 I (Paraspidosiphon), Aspidosiphon, 200, 20S, 221

parvulus, Aspidosiphon, 2 I 5, 224 parvum, Siphonosoma, 55 parvus/ -urn, Phascolion, 3S3 pavlenkoi: Phascolosoma, Goljingia, 105 pectinatoides, Golfingia, 62, 64, 67 pectinatum: Apionsoma, Phymosoma, Physcosoma, Phascolosoma, I57, 197 pellucidal -us, Goljingia/ Sipunculus, 9S pellucidum: Nephasoma, Phascolosoma, 92, 98, 99, IOO pellucidus/ -urn, Sipunculus/ Siphonosoma, 3S3 pelma: Phymosoma, Physcosoma, ISS pelmum, Phascolosoma, ISS perimeces: Themiste, Dendrostomum, ISO perlucens, Phascolosoma, I63, I64, I65, 182 peruvianum, Dendrostomum, ISO pescadolense, Siphonosoma, so Petalosoma minutum, 97 petraeum, Dendrostomum, ISI petricola/ -urn, Themistel Dendrostomum, I4S phalloides: Sipunculus, Lumbricus, 3S pharetratum, Phascolion, I !4, 129 (Phascolana), Golfingia, IS9, I94 Phascolion, IOS, I24, I2S; abnonne, 114, 126; africanum, I 30; alberti, I 30; anamalus, I3I; artificiosus/-um, I30; beklemischevi, I I7; bogorovi, I I4, 127; botulusl-um, 3SI; brotzkajae, I3I; canum, I 24; caupo, II4, I 22, 127; cirratum/ -us, I I4, 132; collare, 114, 121; convestiturn/-us, II4, 117; cryptum/-us, II4, liS, 122; dentalicolal-um, uS; dogieli, I3S; gerardi, 117; hedraeum, II4, 118; heteropapillosum, I 27; hibridum/-us, II4, 128; hirondellei, 120; hupferi, II4, 122; ikedai, 3S2; indicus, I22; intermedia, I30; kurchatovi, I IS; lucifugax, I I 4, 119; lutense, I09, I I4, liS, 124; manceps, 3S2; mediterraneum, I q; medusae, II4, 115, 128; megaethi, I I4, 128; microspheroidis/e/-es, II4, 119; mogadrense, I30; mo-

skalevi, 3S2; murrayi, I23; pacificum, II4, us; pallidum, I20; parvuml-us, 3S3; pharetratum, I !4, 129; psammophilus, I29; rectum/-us, I !4, 123; robertsoni, 114, I I5, 12S, 130; sandvichi, 3S3; spetsbergense, I30; squamatum, I37; strombus/-i, 109, 112, 114, 115, 11S, I22, I2S, I27, 130; strombus cronullae, I3I; temporariae, I20; tortum, I3I; tridens, I22; tuberculosurn, I09, 114, 115, I 17, u8, II9, 120, I 32; tubiculum, I30; ushakovi, I !4, 132; valdiviae, I I4, 123; valdiviae sumatrense, I I4, 123 PHASCOLIONIDAE, I07 (Phascoloides), Goljingia, 77 Phascolopsis, 57; gouldii, 57 Phascolosoma, IS9. I69; abnorme, I9S; abyssorum, S9; abyssorum punctatum, 70; agassizii, I63, 173; agassizii kurilense, I74; albidum, 7I; albolineatum, I63, 174, IS4; ambiguum, 3S5; ambonense, q6; anceps, 93, 97; andamanensis, I74; andersoni, 6S; anguineum, 38I; annulatum, I75; antillarum, 1S6; appendiculatum, 74; approximatum, 3SI; arcuatum, I63, I67, 176; arcuatum malaccensis, 176; asperum, 1S2; aspidosiphonoides, S7; asser, ISS; australe, so; benhami, 90; boreale, 94; bulbosum, 90; capense, 69; capitatum, I6S; capsiforme, 71; carneum, 1S4, 3S5; catharinae, 103; charcoti, 70; chuni, 3S1; cinctum, 93; cinereum, 99; cluthensis, 70; cochlearium, 3S5; commune, 75; confusum, 91; constel/atum, 3S5; constrictum, 92; cora/licolum, 3SI; coriaceum, I40, 3S1; cumanense, SI; cylindratum, 70; deani, I76; delagei, 70; demanni, 382; dentigerum, I 82; depressum, 382; derjugini, 70; diaphanes, I8o; digitatum, 94; dissors, 193; dubium, 75; dunwichi, 1S4; duplicigranulatum, 1So; elegans, 214; elongatum, 69; eremita, 94; esculental-um, I76; evisceratum, 180; exasperatum, 385; extortum, 180; falcidentatum, 382; fasciatum, 178; fasciolatum, 385; filiforme, 95; fimbriatum, 382; flagriferum, 95; forbesii, 70; formasense, I73; fulgens, 7I; funafutiense, 177; juscum, 72, 188; gaudens, I88; georgianum, 72; glabrum, 163, I64, I77; glabrum multiannulatum, 177; glaciale, 96; glans, I88; glaucum, 154, I73; glossipapillosum, 72; golikovi, 174; gouldii, 57; granulatum, 177; grayi, I8I; guttatum, I84, 3S5; hamulatum, 130; harveyii, 75; hebes, 52; heronis, 1S5; hespera, I95, I96; horsti, ISo; hozawai,

Taxonomic Index I05; hudsonianum, 74: huttoni, 155: hyugensis, I05; immodestum, rSS; immunitum, 190, 3S2; improvisum, 97; incompositum, S9; iniquum, 70; innoxium, 3S2; intermedium, 130; japonicum, 173; javenense, rSr, rS2; jeffreysii, 17S; johnstoni, 3S5; kapalum, rS5; kolense, 76; lacteum, rSo; laeve, 17S, rS5; lagense, 3S2; /eachii, 3S5; lilljeborgii, 96; lobostomum, 169; longicolle, 75, 3S6; loricatum, 3S6; loveni, I7S; lurco, 176; luteum, 75; macer, 3S2; maculatum, 17S; margaritacea, 71; margaritacea antarcticum, 72; margaritacea hanseni, 71; margaritacea meridiana/is, 72; margaritacea trybomi, 72; martensi, 103; mauritaniense, 3S2; mawsoni, 73; meteori, r6r, 179; microdentigerum, rS2; microdontoton, 177; minutum, 93, 97, r8o; misakianum, 195; mucidum, 95; multiannulatal-um, 174, 177; muricaudatum, 74; mutabile, 71; nahaense, 1S4; nigrescens, 161, 163, 179; nigriceps, rSS; nigritorquatum, 3S3; nigrum, 105; noduliferum, 164, 181; nordenskjoldi, 72; nordfolcense, 3S6; noto, 72; novae-zealandiae, 9S; obscurum, 70, 75; oerstedii, 71; ohlini, 73; okinoseanum, 72; onagawa, 105; onomichianum, rSS; orbiniense, 14S, 3S6; owstoni, 76; oxyurum, 70; pacificum, r6r, 164, 177, 181; papilliferum, 193, 3S3; papillosum, 193, 3S3; pavlenkoi, I05; pectinatum, 197; pellucidum, 9S; pelmum, r8S; perlucens, 163, 164, 165, 182; placostegi, 3S6; planispinosum, rSo; plicatum, 179, 3S6; pourtalesi, 3S6; procerum, 103, ro6; profundum, 72; psaron, 184; pudicum, 73; punctatissimum, 75; puntarenae, 179; pusillum, 197; pygmaeum, 3S6; pyriformis, 154; quadratum, 383; radiata, 218; rapa, 150; reconditum, 193, 383; reticulatum, 3S3; rhizophora, 176; riisei, 99; riukiuensis, 1S4; rottnesti, 184; rueppellii, 3S3; rugosa mauritaniense, 3S3; rugosum, 70; sabellariae, 93, 97; sanderi, 76; saprophagicum, r6r, 163, 167, 183; sarsii, 383; schmidti, r8S; schiittei, roo; scolops, 163, 174, 183; scolops adenticulatum, 184; scolops mossambiciense, r 83; scutiger, 383; semicinctum, 3S6; semirugosum, 52; semperi, 103; sewelli, 383; signum, 72; simile, r88; sluiteri, 99; socium, 72, 184; solitarium, 76; soyo, 72; spengeli, I So; spinicauda, rS5; spinosum, 182; spongicolum, 1S4; squamatum, 137; stephensoni, 163, 165, 178, 184, 185; subhamatum, 99;

447 tasmaniense, 175; tenuicinctum, 70; teres, 70; thomense, rS2; truncatum, 221; tubicula, 130; tumerae, 163, 185; validum, 76; varians, r8o; vermiculusl-um, rS2; verrillii, 99; violaceum, 55: vitreum, 3S4; vulgare, 75; vulgare astuta, 75; vulgare multipapillosum, 75; vulgare selenkae, 75; vulgare tropicum, 75; wasini, 169; we/doni, rS8; yezoense, 173; zenibakense, 105 PHASCOLOSOMAFORMES, 156 PHASCOLOSOMATIDAE, 156 PHASCOLOSOMATIDEA, 156 PHASCOLOSOMATIFORMES, 156 PHASCOLOSOMIDA, 156 Phascolosomum, 159 Phymosoma!-um, 159; agassizii, 173; albolineatum, 174; antillarum, I 86; asperum, IS2; asser, r8S; deani, 176; dentigerum, r82; diaphanes, 1So; duplicigranulatum, 180; falcidentatus, 3S2; japonicum, 173; lacteum, I So; Iordi, 173; lurco, 176; lurco malaccensis, 176; maculatum, 178; meteori, 179; microdontoton, 174, 177; nahaense, 1S4; nigrescens, I79; nigritorquatum, 383; onomichianum, r8S; pacificum, 1Sr; pectinatum, 197; pelma, rS8; psaron, 184; sco/ops, 183; scolops mossambiciense, 183; spengeli, r8o; varians, rSo Physcosomal-um, 159; abyssorum, r6S; agassizii, 173; albolineatum, 174; ambonense, 176; antillarum, r86; asser, rSS; capitatum, r68; corallicola, 381; demanni, 3S2; duplicigranulatum, r8o; esculenta, I76; evisceratum, r So; extortum, I So; falcidentatus, 382; formosense, 173; funafutiense, 177; gaudens, rSS; glabrum, 177; glaucum, 173; granulatum, 177; hebes, 52; herouardi, I7S; horsti, r8o; japonicum, 173; kurilense, 174; lacteum, rSo; lanzarotae, 17S, 185; lurco, 176; lurco malaccensis, 176; maculatum, 17S; mauritaniense, 3S2; meteori, 179; microdentigerum, rS2; minutum, rSo; nigrescens, 179; nigritorquatum, 383; onomichianum, rSS; pacificum, 181; pectinatum, 197; pelmum, rS8; psaron, 1S4; rueppellii, 383; scolops, 183; scolops adenticulatum, 1S4; scolops mossambiciense, 175, 183; scolops tasmaniense, 175; simi/is, rS8; socium, rS4; spengeli, rSo; spongicola, 184; stephensoni, 185; thomense, 182; varians, rSo; we/doni, r8S; yezoense, 173 pinnifolia!-um, Dendrostomum!Themiste, 3S3 placostegi, Phascolosoma, 386 planispinosum, Phascolosoma, rSo planoscutatus, Aspidosiphon, 225

448 plicatusl -urn, Sipunculus/ Phascolosoma, 179, 386 polymyotus, Sipunculus, 29, 30, 3 I, 39 poritidis, Lithacrosiphon, 229 porrectus, Sipunculus, 40 pourtalesi, Phascolosoma, 386 priapuloides, Sipunculus, 36 procera: Thysanocardia, Phascolosoma, Golfingia, I 03, 1o6 procerum, Phascolosoma, 106 profundal-um, Golfingia/ Phascolosoma, 72 psammophilus, Phascolion, 129 psaron: Phymosoma, Physcosoma, Phascolosoma, 184 Pseudaspidosiphon, 200; gracile, 216; clavatum, 218 pudical-um, Golfingial Phascolosoma, 73 pugettensis, Golfingia, 105 punctatissimuml -us, Phascolosoma/Sipunculus, 75 punctatum, Phascolosoma abyssorum, 70 puntarenae: Phascolosoma, Sipunculus, 179 pusillum/-a, Phascolosoma/Golfingia, 197 pygmaeum, Phascolosoma, 386 pygmaeus: Aspidosiphon, Paraspidosiphon, 218

pyriformis: Phascolosoma, Golfingia, Themiste, 154 pyroides: Themiste, Dendrostomum, 145, 149, rso, 151, 155 quadratuml-a, Phascolosoma/Golfingia, 383 quatrefagesi, Aspidosiphon, 223 queenslandensis, Golfingia vulgaris, 77 radiata, Phascolosoma, 218 ramosal-um, Themiste/Dendrostomum, 150, 386 rapa: Sipunculus, Phascolosoma, 150, 385 ravus, Aspidosiphon, 214 recondita, Golfingia, 193, 383 reconditum, Apionsoma, Phascolosoma, 193, 383 rectum/-us, Phascolion, I14, 123 reticulatuml -a, Phascolosoma!Golfingia, 383 rhizophora, Phascolosoma, 176 rhyssapsis, Aspidosiphon, 386 rickettsi, Siphonides, 198 riisei, Phascolosoma, 99 rimicola: Nephasoma, Golfingia, 78, 100 riukiuensis, Phascolosoma, 184 robertsoni: Dendrostomum, Themiste, 154 robertsoni, Phascolion, II4, II5, 128, 130 robustus, Sipunculus, 30, 39 rosaceal-um, Themiste!Dendrostomum, 148

Taxonomic Index rotumanum/-us, Siphonosoma/Sipunculus, 45.54 rottnesti, Phascolosoma, 184 rubens, Sipunculus, 385 rueppellii: Phascolosoma, Physcosoma, 383 (Rueppellisoma), Phascolosoma, 159 rufofimbriatus, Sipunculus, 385 rugosa/ -urn, Golfingial Phascolosoma, 70 rutilofuscum, Nephasoma, 78, 84, 87 rutilofuscus I-a, Aspidosiphonl Golfingia, 87

sabellariae, Phascolosoma, 93, 97 saccatus, Sipunculus, 385 sanderi: Phascolosoma, Golfingia, 76 sandvichi, Phascolion, 383 saprophagicum, Phascolosoma, 161, 163, 167, 183 sarsii: Onchnesoma, Phascolosoma, Golfingia, 383 (Satonus), Phascolosoma, 159, 197 savalovi, Golfingia, 168 scabra: Phascolosoma eremita, Golfingia eremita, 94 schmidti, Phascolosoma, 188 schmitti: Themiste, Dendrostomum, 150 schnehageni: Aspidosiphon gracilis, Paraspidosiphon, 216 schuettei: Nephasoma, Golfingia, 78, 94, 100 schuttei, Phascolosoma, roo scolops: Phascolosoma, Phymosoma, Physcosoma, 163, 174, 183 scutatus, Sipunculus, 2 I 8 scutiger: Phascolosoma, Golfingia, 383 sec tile, Golfingia, 93 selenkae: Phascolosoma vulgare, Golfingia vulgaris, 75 semicinctum, Phascolosoma, 386 semirugosum: Phascolosoma, Siphonosoma cumanense, 51, 52 semperi: Aspidosiphon, Paraspidosiphon, 225 semperi: Phascolosoma, Golfingia, 103 sewelli: Phascolosoma, Golfingia, Nephasoma, 383 sibirica: Phascolosoma margaritacea, Golfingia margaritacea, 7 r signal-urn, Golfingial Phascolosoma, 72 signifer, Dendrostomum, 153 simi/isl-e, Physcosoma/Phascolosoma, 188 Siphonides rickettsi, 198 (Siphonoides), Golfingia, 6r Siphonomecus, 55; ingens, 53; multicinctus, 56 Siphonosoma, 44; amamiense, 53; arcassonense, 45, 48, 49; australe, 45, so; australe takatsukii, so; billitonense, 52; bo-

Taxonomic Index holense, 45, 48, so; bonhourei, 8r; carolinense, 52; crassum, 55: cumanense, 45, 48, 51, 54; cumanense opacal-um, sr; cumanense semirugosum, 52; cumanense vitreum, 51; cumanense yapense, 51; dayi, 45. so, 53; edule, sr; eniwetoki, 54;/ormosum, 52; funafuti, 45, 48, 51, 53; hataii, 52; hawaiense, 54; ingens, 45, 48, 53, 54: joubini, 382; koreae, 52; marchadi, 52; mourense, 45, 54; novaepommeraniae, 52; parvum, 55; pellucidum, 383; pescadolense, so; rotumanum, 45. 54; vastum, 45, 48, 55 SIPUNCUUDA, 24 SIPUNCULIDAE, 24 SIPUNCULIDEA, 24 SIPUNCULIFORMES, 24 Sipunculus, 28; aeneus, so; aequabilis, 36; amamiense, 53; angasii, 39; arcassonensis, 49; arcuatuu, 176; australis, so; balanophorus, 37; bernnhardis, 130; billitonensis, 52; boholensis, so; bonhourei, 381; borealis, 94; caementarius, 130; capitatus, 130; clavatus, 385; claviger, 52; communis, 75; cochlearius, 218; concharum, 130; corallicolus, 385; cumanense, sr; cumanensis opacus, 51; deformis, 51; delphinus, 37; denta/ii, 130; discrepans, 40; echinorhynchus, 385; edulis, 51; e/egans, 214; eremita, 94; eximioclathratus, 37; flavus, 178; funafuti, 53; galapagensis, 38; genuensis, 178; gigas, 37; glans, r88, 385; gouldii, 57; gravieri, 39, 43; guttatus, I 84; heterocyathi, 2 I 8; immodestus, 188; indicus, 30, 40; infrons, 36; javenensis, r8r, r82, 385; joubini, 382; levis, 178; lomonossovi, 33; longipapillosus, 25, 27, 30, 31, 35, 43, 44; macrorhynchus, 385; maoricus, 41; marcusi, 30, 35, 37; margaritaceus, 71; microrhynchus, 385; multisulcatus, 38; multitorquatus, 178; mundanus, 30, 40; natans, 39; nitidus, 36; noduliferus, r8r; nodulosus, r 8 r; norvegicus, 30, 35, 36; novaepommeraniae, 52; nudus, 30, 31, 35, 36, 39; nudus tesselatus, 37; obscurus, 70, 75; orbiniensis, 148; papillosum, 75, 178; pellucidus, 98, 383; phalloides, 38; phailoides inc!usus, 38; plicatus, I 79, 386; polymyotus, 29, 30, 31, 39; porrectus, 40; priapuloides, 36; punctatissimus, 75; puntarenae, 179; rapa, 150, 385; robustus, 30, 39; rotumanus, 54; rubens, 385; rufofimbriatus, 385; saccatus, 385; scutatus, 218; spinicauda, 185; strombus, 130; ti-

449 grinus, 37; titubans, 37; titubans diptychus, 37; tuberculatus, I 80; vastus, 55; vermiculus, I82, 385; verrucosus, 178; violaceus, 55, 385; vulgaris, 75; zenkevitchi, 384 sluiteri: Phascolosoma, Golfingia, 99 socium: Physcosoma, Phascolosoma, 72, 184 solitarial -urn, Golfingia/ Phascolosoma, 76 soya/ -o, Golfingia! Phascolosoma, 72 speciosus: Aspidosiphon, Paraspidosiphon, 223 speculator: Aspidosiphon, Paraspidosiphon, 217, 225 spengeli: Phascolosoma, Phymosoma, Physcosoma, r8o spetsbergense, Phascolion, 130 spinalis, Aspidosiphon, 2 I 4, 2 I 5 (Spinata), Golfingia, 65, 67 spinicaudal -urn, Sipunculus/ Phascolosoma, 185 spinifera: Dendrostoma, Themiste, 383 spinososcutatus: Aspidosiphon, Paraspidosiphon, 224 spinosum, Phascolosoma, I82 spinosus, Aspidosiphon, 214 spiralis, Aspidosiphon, 220 spongicola/ -urn, Physcosomal Phascolosoma, 184 squamatum: Onchnesoma, Phascolosoma, Phascolion, 134, 136, 137 steenstrupii, Aspidosiphon, 225, 227 steenstrupii, Onchnesoma, 133, 134, 138 Stephanostoma barentsii, 71 stephensoni, Dendrostoma, 69, 154 stephensoni: Phascolosoma, Physcosoma, 163, r6s. 178, r84, 185 (Stephensonum), Themiste, 61, 140 Sternaspis elegans, 214 strombus, Sipunculus, 130 strombus/-i, Phascolion, 109, II2, II4, II5, II8, 122, 125, 127, 130 subhamatal-um, Golfingia/Phascolosoma, 99 subhamatum, Nephasoma pellucidum, 99 sumatrense, Phascolion valdiviae, r 14, 123 Syrinx, 28; harveyii, 75; nudus, 37; tesselatus, 37 takatsukii, Siphonosoma australe, 50 tasmaniense/-is, Nephasoma/Golfingia, 84, 100, 101 tasmaniense: Phascolosoma, Physcosoma scolops, 175 temporariae, Phascolion, 120 tenuicinctum, Phascolosoma, 70 tenuis: Aspidosiphon, Paraspidosiphon, 226

450 tenuissima, Golfingia, 195 teres, Phascolosoma, 70 tesselatus, Syrinx, 37 Themiste, 140, 148; alutacea, 145, 148, 149, rsr; blanda, 149. rsr; cymodoceae, I43. 152; dehamata, I43. 153; dyscrita, 149; elliptica, I 53; fisheri, I 53; fusca, I 54: glauca, I54; hennahi, I49, 150; hexadactyla, ISI; lageniformis, I43. I44, 153; Iissa, I 50; /utu/enta, 386; minor, I4I, 154; minor huttoni, I 54. 155; neimaniae, I4I, 383; orbiniensis, I48; perimeces, I 50; petrico/a, I48; pinnifolium, 383; pyriformis, 154; pyroides, I45, I49, 150, 151, ISS; ramosa, I so. 386; robertsoni, 154; rosacea, 148; schmitti, rso; spinifer, 383; stephensoni, 68, 154; tropica, I54; variospinosa, I 55; zosterico/a, I 50 THEMISTIDAE, I40 thomassini, Aspidosiphon, 205, 212 thomense: Phasco/osoma, Physcosoma, I82 Thysanocardia, I 02; catharinae, 103, I 05, ro6; nigra, 103, 105; procera, 103, 1o6 tigrinus, Sipunculus, I78 titubans, Sipunculus, 37 tortum, Phascolion, I3I tortus, Aspidosiphon, 2I8 trichocephalus/-a, Apionsoma/Golftngia, 196 tridens, Phascolion, I22 trinidensis: Aspidosiphon, Paraspidosiphon, 226 tropica, Golftngia vulgaris, 75 tropical -urn, Themiste/ Dendrostomum, I 54 tropicum, Phascolosoma vulgare, 75 truncatum, Phascolosoma, 22I truncatus, Aspidosiphon/Paraspidosiphon, 22! trybomi: Phascolosoma, Golfingia, 72 tuherculatus, Sipunculus, 181 tuberculosum, Phascolion, 109, I I4, I IS, 117, II8, II9, 120, I32 tubiculal -um, Phascolosomal Phascolion, r 30 tumerae, Phascolosoma, I63, 185 Tylosoma lytkenii, I 20 unilobatae, Golfingia murinae, 195 uniscutatus: Aspidosiphon, Lithacrosiphon, 229 ushakovi, Phascolion, I 14, 132

Taxonomic Index valdiviae, Phascolion, I 14, 123 validum, Phascolosoma, 76 varians: Phasco/osoma, Phymosoma, Physcosoma, r8o variospinosa, Themiste, I 55 vastum/-us, Siphonosoma/Sipunculus, 45, 48, 55 venabulum/-us, Aspidosiphon, 2I2 vermiculus: Sipunculus, Phascolosoma, I82, 385 verrillii: Phascolosoma, Golfingia, 99 verrucosus, Sipunculus, 178 vesiculosus, Golfingia vulgare, 75 (Villiophora), Phascolion, I 32 violaceum, Phascolosoma, 55 violaceus, Sipunculus, 55, 385 vitjazi: Nephasoma, Golfingia, 78, 83, 84, 101 vitrea, Golfingia, 384 vitreum: Phascolosoma, Nephasoma, 384 vitreum, Siphonosoma cumanense, 51 vulgare, Phascolosoma, 75 vulgaris: Golfingia, Sipunculus, 62, 65, 74, 75

wasini, Phascolosoma, 169 we/doni: Physcosoma, Phascolosoma, r88 wodjanizkii: Nephasoma, Golfingia, 83, 84, 101 (Xenopsis), Xenosiphon, 40 Xenosiphon, 4I; absconditus, 43; branchiatus, 30, 35, 41, 44; branchiatus nudus, 43; caribaeum, 44; indicus, 40; mundonum, 40 yapensel-is, Aspidosiphon elegans, 214 yapense, Siphonosoma cumanense, 51 yezoense: Phasco/osoma, Physcosoma, 173 zarenkovi, Golftngia confusa, 9I zenibakense/ -is, Phascolosomal Golftngia, 105 zenkevitchi, Sipunculus, 384 zinni, Aspidosiphon, 203, :z13 zostericolal -um, Themistel Dendrostomum, ISO

Subject Index

actin control, 254-255 amoebocytes, 256, 268-269 amphi-Arnerican taxa, 369 amphitropical taxa, 324 anaerobic metabolism, 272-273 ancestral sipunculan (RHAS), revised hypothetical, 375 Ancorichnus, 335 Annelida, 253, 286, 303, 306, 336-344 antibacterial activity, 269 antibody production, 268-270 antiquity of taxa, 363, 371-374 Aphrodite, 242 Aplacophora, 343-344 Archicoelomata, 336 Arenicola, 255 arginine kinase, 275 asexual reproduction, 308-310 barriers, zoogeographical, 317, 323, 368369, 372 behavior, 239-241, 246 Bering land bridge, 364, 372 bilirubin, 258 bioerosion of reefs (coral boring), 237-238 biomedical uses of urn cells, 267-268 boundaries, zoogeographical, 316, 330, 372 Brachiopoda, 257, 258, 260, 261, 263, 340 breeding cycles, 300-301 budding, 308-309 burrows, burrowing, 237, 240-241 calcium carbonate, 249. 251 calcium ions, 254-255, 280 Cambrian, 319, 334-336, 344. 352, 375 carbonic anhydrase, 339 catch muscle, 252, 254-255 Caulalotilus, 244 Cenozoic, 322, 330, 331, 333, 336, 374, 377, 378, 379; sea level, 367-368; sea temperatures, 365-367; subregional oceanic events: Australia to Southeast Asia, 37 I; Eastern Pacific Barrier, 372; Mediterra-

nean Sea and northeastern Atlantic, 368; North Pacific, Arctic, and Far North Atlantic, 372-373; Soutlr Atlantic, 369; western Atlantic and eastern Pacific, 369371 center of origin, 315-316, 371-372 cerebral organ, 289 character state polarities, 346-354 chemoreception, 289 chromatin, 273-274, 340 chromosomal number/morphology, 354-357 cilia, 251 cladogenesis/speciation centers and events: Aspidosiphoniformes, 332; Golfingiiformes, 324-329; Phascolosomatiformes, 330-331; Sipunculiformes, 322-323 cleavage, 301 coelomic cells, 256-257 Collostoma, 247 commensals, 246-247; mollusks, 246; polychaetes, 247; small metazoans, 246 corals, 238, 245, 331, 332, 335-336, 372, 378 cosmopolitan species, 319-320 crabs, 244, 246 Cretaceous, 316, 363, 364, 367, 368, 372; fossils, 335, 336 Crustacea, 242, 289, 291, 316 currents/ocean circulation, 365, 367, 369, 371-372 cuticle, 249-251 cytochromes, 264-265 cytotoxic activity, 270 Daphnia, 298 dermis, 251 Devonian, 335. 336 digestive system: anatomy, 282-283; physiology, 283-285 disjunct distributions, 316, 367, 388 dispersal, 3 I 6-3 I 9, 372 Drupa, 244

Subject Index

452 Echinodermata, 338, 340, 365, 369 Echiura, 306, 337, 339-340 encapsulation, 268-269 endemic taxa, 3I4-3I6, 3I8-3I9; Phascolosomatidea, 330, 332; Sipunculidea, 32I-329 Entoprocta, 246 environmental deterioration/pollution indicators, 239 Eocene, 36o, 364-365, 37I erythrocytes, 256-258, 264-265 Eudendrium, 246 excretory system: anatomy, 276-279; physiology, 279-28I extinctions, 36o, 365, 370; local, 3 I6, 32 I, 324, 367-368, 372-373 f~ng, 239, 24I-243 food for predators, 243-244 Foraminifera, 3I5, 332, 365, 366, 367, 369 fossil record, 334-336 Fronsella, 246 fusiform bodies, 293-296

gametes, 298-299 gas exchange, 27I-272 gastrulation, 303 genetic variability (enzymes), 274 glands, epidermal, 25 I -252 glycogen storage, 264 Gondwanaland, 363 granulocytes, 256-257, 268 gravity reception, 290 guanidine compounds, 275 habitat, 236-239 hemerythrins: function, 261 -264; oxidation levels, 264; structure, 258-26I hermaphroditism, 298 Heterocyathus, Heteropsammia, 238, 245 histones, 273-274 hooks, 249, 252 Hydrozoa (hydroids), 246 Hyolitha, 334 immune system, 268-270 mactivation, 263-269 mtegument/epidermis, 249-252 intercellular junction, 342 intestine, 282-283 irrigation of shelter, 239, 245

karyotypes, 354-357 Keferstein bodies, 292-293

larval development, 304-307 Laurasia, 363 leucocytes, 256-257, 270 Lingula, 263 Loxosomella, 246 Lumbricus, 255 Menestho, 247 Mesozoic, 3I5, 363-364, 371, 372; cladogenic events, 324, 33I, 333, 373. 377, 378 Miocene: cladogenic events, 323, 329, 374, 379; fossils, 335-336; oceanic conditions, 3I6, 36I, 364-372 Mollusca, 246, 288, 296, 303, 317, 334, 336-345. 365, 369, 372 Montacuta, 246-247 mucus-stimulating substance, 267-268 muscle physiology and biochemistry, 254255, 340 muscle systems, 252; body wall, 252; mtestinal fasteners, 253-254; introvert retractors, 253; protractors, 253 Mustellus, 244 mutualism, 244-245 NADH diaphorase, 264 Nematoda, 248, 340 Neogene, 366, 373, 377, 378-380 nephridia, 276-279 nerve transmission, 288 nervous system, structure of, 286-288 neurosecretion, 291-292 niches, 236, 330, 336 nicotinic type receptor, 254 Nipponmysella, 246 nuchal organ, 289 numerical (phylogenetic) analysis, 346-349 Oligocene, 364-367, 371 Oligochaeta, 255, 270 Ordovician, 334-335, 377 osmoregulation, 279-281 Ostracoda, 366-367 Ottoia prolifica, 334, 352 Paleocene, 365, 366, 37I Paleogene, 316, 328, 368 Paleozoic: cladogenic events, 324, 328, 330332, 373, 376-379; fossils, 334-336; oceanic conditions, 36o, 363 Panamanian area, 323, 366, 369, 372 Pangaea, 363 Panthalassa (Eo-Pacific), 324, 328, 330-332, 363, 371 paramyosin, 254-255

Subject Index parasitism, 243, 247-248 parthenogenesis, 298, 308 passive transport of larvae, 3I7-3I8 pelagosphera larva, 305-307 Perigonimus, 246 Permian, 326 phagocytosis, 256, 267-269 pheromone, 296 phospholipids, 339-340 phosphorus metabolites, 264 photoreception, 290 phototaxis, 239-240 plate tectonics, 300-364, 368-373 Pleistocene conditions, 3I6, 335, 366-367, 369, 373 Pleurodictyum, 336 Pliocene: cladogenic events, 323, 335, 374, 377; oceanic conditions, 365-367, 369, 372 Pogonophora, 25I, 336 Polychaeta, 237-238, 242-243, 244, 247, 25I, 255. 278 prey items, 243-244 Priapulida, 257, 258, 334. 352 Protoctista, 247, 269 Protostomia, 299, 336-338, 342 pyruvate catabolism, 340 Quaternary, 335, 367 quick muscle, 255 recognition of self/nonself, 268-270 regeneration, 3I0-3II respiration, 27 I -272 ribosomal RNA, 339, 34I

453 sea level, 367-368 sense organs, 288-290 settlement of larvae, 308 sexual reproduction, 308-31 o Silurian, 335 species concepts, 3 I 4 species-rich areas, 3 I 5 sperm anchoring fiber, 299, 342 Spheciospongia, 238 Spiralia, 336-338, 343 sterility, 248, 299 superoxide dismutase, 264 superphyla, 334, 339 Symbiongia, 336 symbiotic relationships, 244-248 temperature changes, 364-367 Tertiary, 324, 326, 364, 365 tested endemics, 3I4 Tethys Sea, 3I6, 322, 333, 363-365, 368 Triassic, 326, 363 Trichichnus, 237, 335 Tritice/la, 246 trochophore, 300-307 trophic dynamics, 24I-244 Trypanites, 335 untested endemics, 3I4, 321, 324, 325, 328 urn cell complex (UCC), 265-268; biomedical applications, 267-268 vicariance, 3I6, 3I8, 32I Wadeopsammia, 336