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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 146

The Families and Genera of Hyperiidea (Crustacea: Amphipoda)

Thomas E. Bowman and Hans-Eckhard Gruner

SMITHSONIAN INSTITUTION PRESS

City of Washington

1973

ABSTRACT

Bowman, Thomas E., and Hans-Eckhard Gruner. The Families and Genera of Hyperiidea (Crustacea: Amphipoda). Smithsonian Contributions to Zoology, number 146, 64 pages, 82 figures, 1973.—A synopsis of the amphipod suborder Hyperiidea is presented, including diagnoses of its 2 infraorders, 6 superfamilies, 21 families, and 71 genera. Diagnostic keys are given for the families and genera, and illustrations of significant characters are given for each genus. Several changes are made in the classification of the Hyperiidea. Tribes are changed to infraorders and subtribes to superfamilies, with the necessary nomenclatural changes being made. A new superfamily, Lycaeopsoidea, for the family Lycaeopsidae, and a new family, Anapronoidae, for Anapronoe, are proposed. A new genus, Spinoscina, is proposed for Acanthoscina spinosa; and Vibilioides and Parascelus are reduced to synonyms of Vibilia and Thyropus, respectively.

OFFICIAL PUBLICATION DATE is handstampcd in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year. SI PRESS NUMBER 4776. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Bowman, Thomas E.

The families and genera of Hyperiidea.

(Smithsonian contributions to zoology, no. 146)

1. Amphipoda—Classification. I. Gruner, Hans-Eckhard, joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 146.

QL1.S54 no. 146 [QL444.A5] 591′.08s [595′.371] 73–1579

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Contents

Introduction

Morphology

Natural History

Classification

Suborder Hyperiidea H. Milne Edwards, 1830

Infraorder Physosomata Pirlot, 1929

Superfamily Scinoidea, new name

Family Archaeoscinidae Stebbing, 1904

Archaeoscina Stebbing, 1904

Family Mimonectidae Bovallius, 1885

Mimonectes Bovallius, 1885

Pseudomimonectes Vinogradov, 1960

Family Proscinidae Pirlot, 1933

Proscina Stephensen and Pirlot, 1931

Mimoscina Pirlot, 1933

Family Scinidae Stebbing, 1888

Scina Prestandrea, 1833

Acanthoscina Vosseler, 1900

Ctenoscina Wagler, 1926

Spinoscina, new genus

Superfamily Lanceoloidea, new name

Family Microphasmidae Stephensen and Pirlot, 1931

Microphasma Woltereck, 1909

Microphasmoides Vinogradov, 1960

Mimonecteola Woltereck, 1909

Family Chuneolidae Woltereck, 1909

Chuneola Woltereck, 1909

Family Lanceolidae Bovallius, 1887

Lanceola Say, 1818

Megalanceola Pirlot, 1935

Metalanceola Pirlot, 1931

Paralanceola Barnard, 1930

Prolanceola Woltereck, 1907

Scypholanceola Woltereck, 1905

Infraorder Physocephalata, new name

Superfamily Vibilioidea, new name

Family Vibiliidae Dana, 1852

Vibilia H. Milne-Edwards, 1830

Cyllopus Dana, 1852

Family Cystisomatidae Willemoes-Suhm, 1875

Cystisoma Guérin-Méneville, 1842

Family Paraphronimidae Bovallius, 1887

Paraphronima Claus, 1879

Superfamily Phronimoidca, new name

Family Hyperiidae Dana, 1852

Hyperia Latreille, in Desmarest, 1823

Bougisia Laval, 1966

Hyperiella Bovallius, 1887

Hyperietta Bowman, 1973

Hyperioides Chevreux, 1900

Hyperionyx Bowman, 1973

Hyperoche Bovallius, 1887

Iulopis Bovallius, 1887

Lestrigonus Milne Edwards, 1830

Parathemisto Boeck, 1870

Pegohyperia Barnard, 1932

Phronimopsis Claus, 1879

Themistella Bovallius, 1887

Family Dairellidae Bovallius, 1887

Dairella Bovallius, 1887

Family Phronimidae Dana, 1853

Phronima Latreille, 1802

Phronimella Claus, 1871

Family Phrosinidae Dana, 1853

Phrosina Risso, 1822

Anchylomera H. Milne-Edwards, 1830

Primno Guérin-Méneville, 1836

Lycaeopsoidea, new superfamily

Family Lycaeopsidae Chevreux, 1913

Lycaeopsis Claus, 1879

Superfamily Platysceloidea, new name

Family Pronoidae Claus, 1879

Pronoe Guérin-Méneville, 1836

Eupronoe Claus, 1879

Paralycaea Claus, 1879

Parapronoe Claus, 1879

Sympronoe Stebbing, 1888

Anapronoidae, new family

Anapronoe Stephensen, 1925

Family Lycaeidae Claus, 1879

Lycaea Dana, 1852

Brachyscelus Bate, 1861

Pseudolycaea Claus, 1879

Thamneus Bovallius, 1887

Tryphana Boeck, 1870

Family Oxycephalidae Bate, 1861

Oxycephalus H. Milne-Edwards, 1830

Calamorhynchus Streets, 1878

Cranoecephalus Bovallius, 1890

Glossocephalus Bovallius, 1887

Leptocotis Streets, 1877

Rhabdosoma White, 1847

Simorhynchotus Stebbing, 1888

Streetsia Stebbing, 1888

Tullbergella Bovallius, 1887

Family Platyscelidae Bate, 1862

Platyscelus Bate, 1862

Amphithyrus Claus, 1879

Hemityphis Claus, 1879

Paratyphis Claus, 1879

Tetrathyrus Claus, 1879

Family Parascelidae Bovallius, 1887

Thyropus Dana, 1852

Euscelus Claus, 1879

Schizoscelus Claus, 1879

Sources of Illustrations

Literature Cited

Publication in Smithsonian Contributions to Zoology

The Families and Genera of Hyperiidea (Crustacea: Amphipoda)

Thomas E. Bowman and Hans-Eckhard Gruner

__________

Thomas E. Bowman, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Hans-Eckhard Gruner, Zoologisches Museum der Humboldt Universität zu Berlin, Invalidenstrasse 43, 104 Berlin, Deutsche Demokratische Republik.

The Hyperiidea are, therefore, those species among the Amphipoda, in which nature indulges in her widest diversities of development….

—JAMES DWIGHT DANA

Introduction

The hyperiid amphipods are important marine crustacean zooplankters, ranking third in abundance behind the Copepoda and Euphausiacea. Contrary to the statement of Hurley (1969), identification of hyperiids often is a frustrating task, not only because of the unsatisfactory state of our knowledge of such genera as Eupronoe and Lycaea but especially because of the scattered condition of the literature on hyperiid systematics. Much of the essential literature is contained in works of the late 19th century and in reports of the major oceanographic expeditions, which often are not readily available, especially at the smaller and newer marine laboratories. No attempt has been made at a summary of hyperiid systematics since the pioneer monographs of Bovallius (1887a, 1887b, 1887c, 1889, 1890) and Claus (1879a, 1879b), although many important works have since appeared. The need for an up-to-date synopsis has been made more urgent by the increased activity in plankton research in recent years.

A concurrent 2-month visit as consultants at the Indian Ocean Biological Centre, Cochin, India, afforded us the opportunity—free from the usual interruptions at our respective museums—to work closely together in preparing a large part of the present synopsis. Our goal has been to prepare a synopsis that will make it possible for a trained biologist to identify any hyperiid amphipod to genus without having to refer to other publications. We have attempted to write clear and unequivocal diagnoses and we include a section on morphology so that the meanings of terms will be unmistakable. All characters used in our diagnoses are illustrated. We have used published illustrations of other authors freely, sometimes modifying them by eliminating unnecessary or confusing lines or by spreading out overlapping appendages. When adequate illustrations were not available we prepared original drawings.

Sources of the illustration are given in the appendix. The following abbreviations are used in the illustrations:

Our diagnoses are based in part on published accounts, but we have examined numerous specimens in the collections of the Indian Ocean Biological Centre and our own museums in order to verify, correct, or supplement these accounts. Synonymies are omitted, since they will be given fully in a section on Hyperiidea in the Catalogus Crustaceorum, now being prepared. For each family or genus we list a few of the most useful taxonomic works, but we have not attempted to provide an extensive bibliography of the Hyperiidea.

This work is the result of equal effort by each of us, and we should be considered coauthors rather than junior and senior authors.

Morphology

Figure 1 shows a generalized hyperiid with the principal parts labeled.

The body form is quite varied. In some hyperiid amphipods, including the Physosomata and Cystisomatidae, the cuticle is thin and transparent and the muscles are weakly developed. These hyperiids often have rather globular bodies and are weak swimmers. They are mainly bathypelagic. Others, such as Pronoidae (Figures 54–59) and Lycaeidae (Figures 60–64), have rather compact bodies with strong musculature and can swim very rapidly. The plump Platyscelidae and Parascelidae can undergo conglobation (rolling into a ball) by bringing the operculate bases of the left and right pereopods 5 and 6 together ventrally and fitting them against the posteroventral margin of the head (Figure 74). The telson and uropods are tucked under the body and into a unnamed groove on the posterior margin of each pereopod 6 that we propose to call the “telsonic groove.” The function of this conglobation is unknown; possibly it aids these strong swimmers to avoid predators by sinking.

The opposite extreme in body form from the conglobate families is found in the slender elongate Oxycephalidae, culminating in the needle-shaped Rhabdosoma (Figure 70).

The head is typically globular, with most of its surface occupied by the compound eyes. The dorsal midline is free from ommatidia and serves as a line of attachment for a muscle that inserts on the dorsal surface of the midgut. In the Physosomata the eyes are small or absent and situated on the sides of the rather short head. Among the Physocephalata the eyes are small or medium-sized only in Vibilia and Bougisia where they are located on the sides of the cuboidal head. In the Oxycephalidae the anterior part of the head in front of the eyes is drawn out into a short-to-long triangular rostrum.

The pereon comprises 7 segments (pereonites 1–7 = thoracic somites 2–8). Some of the anterior pereonites may be fused in certain genera. The coxae (= coxal plates), or the proximal segments of the thoracic legs, may be fused with the pereonites or separated from them by a suture; they are never enlarged as in many Gammaridea.

The abdomen is formed of the pleon and the urosome. The pleon contains 3 segments or pleonites. The urosome, which is composed of 3 segments or urosomites in most gammarideans is 2-segmented in hyperiids, urosomites 2 and 3 being fused to form the double urosomite. The abdomen ends in a thin triangular piece, the telson, which may be fused with the double urosomite. The telson is never divided or incised at its apex as it is in many Gammaridea.

The 2 pairs of antenna, antenna 1 and antenna 2, may be inserted on either the anterior or the ventral surface of the head. The site of insertion and the structure of antennae 1 and 2 are the major criteria by which the superfamilies of Physocephalata are distinguished (Figure 2). Each antenna consists of a basal part, the peduncle, followed by a flagellum. In the Gammaridea the peduncle is 3-segmented in antenna 1 and 5-segmented in antenna 2, but in the Hyperiidea the number of peduncle segments is often fewer in antenna 1 and always fewer in antenna 2. The second segment of antenna 2, which bears the gland cone on which the antennal gland opens, is fused to the head, hence the peduncle consists of only 3 (or fewer) segments.

The flagellum may be filiform and multisegmented, composed of a few long segments that fold upon each other, or reduced to a single large segment which may be followed by 1 or a few small or rudimentary additional segments. The distal segments may be inserted at the tip of the first segment (terminally) or proximally to the tip (subterminally). A secondary or accessory flagellum, present in many Gammaridea, is never found in Hyperiidea. Both antennae are usually reduced in the female, especially antenna 2, which may be rudimentary or absent. In a few instances antenna 2 is absent in the male.

FIGURE 1.—Diagram of a hyperiid amphipod, based on ♂ Hyperia.

FIGURE 2.—Diagrams of ♂ heads of the 4 superfamilies of Physocephalata: A, Vibilioidea; B, Phronimoidea; C, Lycaeopsoidea: D, Platysceloidea.

The mouthparts of the Hyperiidea (Figure 3) are much reduced compared with those of the Gammaridea. The mandible may or may not have a 3-segmented palp. Frequently the palp is present in the male and absent in the female of the same species. Rarely it is reduced to 1 segment (Pseudomimonectes) or 2 segments (♀ Tryphana). The molar may be rudimentary or absent. Maxilla 1 has an inner lobe in the Physosomata, but not in the Physocephalata, and the palp is 1-segmented. Both maxilla 1 and maxilla 2 may be rudimentary (Lycaeidae) or absent (Oxycephalidae). In the maxillipeds the palp is usually absent, although a 1-segmented rudiment is present in a few instances. The inner lobes are partly or completely separate in most of the Physosomata, but are completely fused into a single median lobe in other hyperiids. In a few species the inner lobe is rudimentary or absent.

FIGURE 3.—Mouthparts of some Hyperiidea: Md, Cyllopus magellanicus; Md’, Chuneola paradoxa (molar and palp absent); Mx.l, Mx.2, Cyllopus magellanicus; Mxp, Proscina magna (inner lobes separate); Mxp′, Parathemisto gaudichaudii (inner lobes fused).

FIGURE 4.—Diagram of an amiphipod pereopod.

FIGURE 5.—Distal segments of simple hyperiid pereopods: A, Lanceola; B, Tryphana; C, Pronoe: D, Paratyphis; E, Thyropus; F, Vibilia; G, Dairella; H, Sympronoe. All drawings are of pereopod 1, except G, which is of pereopod 2.

Each of pereonites 1–7 bears a pair of 7-segmented appendages, the pereopods. The proximal segment, or coxa, forms an immovable flat coxal plate fixed to the lateral surface of the pereonite and sometimes fused to it. The remaining 6 segments forming the functional leg are the basis, ischium, merus, carpus, propus, and dactyl (Figure 4). When a pereopod is not prehensile it is said to be simple (Figure 5). A prehensile pereopod may be either subchelate or chelate. In the Hyperiidea prehension is usually effected by closure of the propus against the expanded carpus, whereas in Gammaridea the dactyl closes against the expanded propus. In a subchelate pereopod (Figure 6) the prehensile surface of the carpus is its widened distal margin or its convex anterior or posterior margin, but in a chelate pereopod the carpus is produced distally into a carpal process against which the propus closes (Figure 7). When the carpal process is small it is difficult to decide whether the pereopod is chelate or subchelate. Here the term “weakly chelate” may be used, but illustrations show the condition far more clearly than words. Rarely, a chela or subchela is formed between the dactyl and propus (Figures 19, 35, 45) and a complex chela is found in Amphithyrus (Figure 7G).

FIGURE 6.—Distal segments of subchelate hyperiid pereopods: A, Phrosina, pereopod 4; B, Anchylomera, pereopod 3; C, Simorynchotus, pereopod 2; D, Eupronoe, pereopod 1; E, Tetrathyrus, pereopod 2; F, Paraphronima, pereopod 1; G, Brachyscelus, pereopod 2.

Pereopods 3 and 4 are usually slender and simple, but in Parathemisto (Figure 43), Phronimopsis (Figure 45), Anchylomera (Figure 6B), and Phrosina (Figure 6A) they are prehensile. Pereopod 5 has a strong subchela in Phronimidae and Anchylomeridae. In pereopods 5 and 6 the basis is often quite broad, especially in the Parascelidae and Platyscelidae, where they form broad opercula covering the other pereopods during conglobation. In these families pereopod 6 is especially enlarged; on its posterior margin is the telsonic groove, and on its lateral surface is usually a pocket-shaped fissure of unknown function (Figure 74). Pereopod 7 is often smaller than the other pereopods, sometimes consisting only of the basipod (Figure 74).

FIGURE 7.—Distal segments of chelate hyperiid pereopods: A, Parapronoe; B, Platyscelus; C, Sympronoe; D, Vibilia; E, Phronimopsis; F, Oxycephalus; G, Amphithyrus; H, Lestrigonus. A–G, pereopod 2; H, pereopod 1.

In the Lanceolidae and Chuneolidae the apex of the propus in some pereopods is expanded into a hood surrounding the base of the dactyl (Figure 8). The dactyl can be flexed posteriad so that it lies within the spoon-shaped cavity within the hood, hence it has been called a “retractile dactyl,” but we propose the term “hooded dactyl,” which more accurately describes the structure.

The gills are hollow sacs arising from the medial surfaces of the coxae. Typically there are 5 pairs, on pereonites 2–6, but they may be reduced to 2, 3, or 4 pairs. The gills may have transverse pleats or folds, which increase the respiratory surface.

Medial to the gills, on mature females, are the oostegites, commonly on pereonites 2–5. Oostegites are thin plates which overlap to form a pouch beneath the pereon in which the eggs and young are brooded. In the Hyperiidea the oostegites usually do not have marginal setae as in the Gammaridea. In Rhabdosoma the oostegites are reduced in size and their function is taken over by the gílls.

FIGURE 8.—Pereopod 6 of Scypholanceola vanhoeffeni, showing hooded dactyl.

Each pleonite bears a pair of biramous swimming appendages, the pleopods. Thus far they have not been shown to be useful in taxonomy, but they have not been closely studied.

The urosome bears 3 pairs of uropods, each consisting of a proximal segment, the protopod, bearing distally an exopod and an endopod (Figure 9).

The exopods and endopods are flattened, moderately to very broad, rarely quite narrow, but never styliform as in some Gammaridea. One or more of the endopods, and less commonly the exopods, may be fused with the protopod. In the Anchylomeridae each uropod is a single leaflike segment, and in the Cystisomatidae and the female Phronimella uropod 2 is entirely absent.

FIGURE 9.—Urosomes: A, Pronoe capito (telson not fused with double urosomite; endopods not fused with protopods); B, Oxycephalus piscator (telson fused with double urosomite; endopods of uropods 2 and 3 fused with protopods).

Natural History

The Hyperiidae are entirely pelagic, mostly oceanic, although a few species are found in coastal waters. The swimming ability, which we have observed in living specimens, varies from feeble movements of the limbs in the weakly muscled Cystisoma to very rapid swimming in Parapronoe, which has strongly developed pleonal muscles. Feeding habits are largely unknown. The appendages lack the rows of plumose setae found in filter-feeding Crustacea. Some species are definitely known to be associated with medusae or salps, e.g., Hyperia (Hollowday, 1947; Pirlot, 1932; Dahl, 1959a, 1959b; Bowman, Meyers, and Hicks, 1963; Laval, 1968a). Dahl (1959a, 1959b) identified nematocysts from the host in the gut of Hyperia galba and concluded that it feeds on the medusa. But the observations of Pirlot (1932) and Bowman, Meyers, and Hicks (1963) suggested that the host tissues are not the only and perhaps not the principal source of nourishment. Possibly the medusa serves as a feeding platform from which the amphipod makes short excursions to collect food particles, including some from food captured by the medusa. Certainly the well-developed grinding surface on the molars of Hyperia mandible is fitted for chewing tougher material than the gelatinous tissues of a medusa.

Observations on living hyperiids have been made in recent years by Laval (1963, 1968a, 1968b, 1972), who raised Lestrigonus schizogeneios and Bougisia ornata with their host, the leptomedusa Phialidium. As long as the medusa was adequately fed (in this case on Artemia) the amphipod shared the food of its host. If the medusa did not capture enough food, the amphipod fed on the host tissues, beginning with the gonads.

The Phronimidae live in transparent “barrels,” open at both ends, made by remodeling pyrosomes or siphonophores. Laval (1968b) describes how a nectophore of the siphonophore Abylopsis is fashioned into a barrel by Phronima curvipes. The Phronima resides in the barrel, capturing prey passing within reach with the 3rd pereopods. It probably feeds mainly on slow-moving and soft-bodied forms. By holding the abdomen outside of one of the barrel entrances and beating the pleopods, the Phronima can propel the barrel through the water.

In the genus Vibilia, ovigerous females attach themselves to salps during the approximately 20 days that the eggs arc being incubated in the marsupium. After the eggs hatch, the mother reaches into her marsupium and grasps a larva with the digitiform dactyls of her 7th pereopods and places it on the surface of a salp. The larva, which cannot swim since the pleopods and uropods have not yet developed, enters the atrium of the salp by way of one of the siphons and feeds on the tissues of its host until it has developed to the stage where it can become planktonic (Laval, 1963).

Members of the deep-water family Lanceolidae are believed to live on deep-water medusae and siphonophores and feed on their tissues. The hooded dactyls of pereopods 5–7 (or of 6 and 7 in Prolanceola) enable them to clasp the host tissue firmly.

Vinogardov (1957) found the guts of Lanceolidae full of a detritus-like mass containing many small bodies that appeared to be nematocysts. In Lanceolidae stomachs, Woltereck (1927) found polychaete worms (Alciopidae), chaetognaths, and even the anterior end of a rather large cephalopod larva. He felt certain that such prey could not have been captured by the slow-moving Lanceolidae and must have been obtained secondarily from the medusa or siphonophore host.

Doubtless many more associations between hyperiids and their hosts, probably mostly gelatinous animals such as coelenterates and thaliaceans, remain to be elucidated. In particular the Platysceloidea, with their reduced mouthparts, must feed on soft tissues. However, some hyperiids appear to be entirely free-living. Parathemisto is predaceous on other plankters (Bigelow, 1926:107; Dunbar, 1946; Bowman, 1960); and unpublished studies of gut contents indicate that Primno is also to a large extent predaceous.

Classification

The classification used here is essentially that of Pirlot (1929) but incorporates the additions of Stephensen and Pirlot (1931). Our primary aim is not to revise the classification of the Hyperiidea, but certain nomenclatural changes have been necessary to bring the classification into harmony with modern taxonomic practice and the International Code of Zoological Nomenclature. Preparation of the diagnoses forced us to examine in detail the characters of all taxa of hyperiids at and above the generic level. As a result we have made the changes given below.

INFRAORDERS AND SUPERFAMILIES.—We consider the names “Hyperiidea Gammaroidea” and “Hyperiidea Genuina” of Woltereck (1909) and “Hyperiidea Physosomata” and “Hyperiidea Eugenuina” of Pirlot (1929) for their tribes (our infraorders; see below) to be singularly inappropriate and meaningless. Moreover, they are double words where single names are required. This legal technicality gives us the opportunity, which we gladly seize to propose the new infraorder name Physocephalata (from “physos,” swollen, plus “cephalon,” head), referring to the large head of most of its members. The name has the advantage of similarity in construction to “Physosomata,” the other infraorder, which is characterized by the swollen pereon.

Tribes have been changed to infraorders and subtribes to superfamilies. These changes were made to conform with the usual rank of the category “tribe,” between family and genus. Since superfamily names are formed by adding a standard ending (usually “oidea”) to the stem of one of the included families, we have replaced the old names of subtribes. Thus, Lanceoliformata and Sciniformata become Lanceoloidea and Scinoidea, using the original stems. We regret that it is necessary to replace Bovallius’ (1890) Recticornia, Filicornia, and Curvicornia, which have been widely used since 1890, but they do not qualify as names of superfamilies. In place of Filicornia we have chosen Phronimoidea rather than Hyperioidea, in order to avoid confusion with Hyperiidea. For Curvicornia we have selected Platysceloidea over other possibilities because Claus (1879b, 1887) included all the families of Curvicornia in his “Platysceliden.” Although the family Lycaeopsidae until now has been included in the Curvicornia, we are convinced that it differs sufficiently from the other families to warrant for it the establishment of a new superfamily, Lycaeopsoidea.

FAMILIES.—Stephensen (1925) assigned Anapronoe to the family Pronoidae without giving his reasons for placing it in that family rather than in the Lycaeidae. The principal criteria by which the two families may be distinguished are found in antenna 2 of the female, mandibular palp, maxillae 1 and 2, and pereopod 7. Of these, only pereopod 7 has been described for Anapronoe, and in this character Anapronoe agrees with the Lycaeidae rather than the Pronoidae. We have been unable to assign Anapronoe to either family, since it shares some characters with each and some with neither. Redefining either family to enable it to encompass Anapronoe would drastically alter that family’s definition and increase the difficulty of distinguishing between the two families; hence, we have erected a new family, Anapronoidae, for Stephensen’s genus.

GENERA.—We have reduced 3 genera to synonyms of older genera because the characters used to separate them seemed insufficient. Vibilioides Chevreux (1905) differs from Vibilia Milne Edwards (1830) only in the reduced number of segments on pereopod 7. Since Hurley (1955) has shown that this character is variable in Cyllopus, the only other genus of Vibiliidae, we do not believe it to have generic value. Vibilioides therefore becomes a synonym of Vibilia, and its type and only species becomes Vibilia alberti (Chevreux).

Similarly, as Stephensen (1925) pointed out, Thyropus Dana, 1852, differs from Parascelus Claus, 1879, only in the longer pereopod 6, a difference of less than generic significance. We are therefore combining the 2 genera under the older name Thyropus. The family name remains Parascelidae, in accordance with Article 40 of the International Code.

Metalycaea was proposed by Stephensen (1925) for a new species, M. globosa, represented by 3 females in the collections of the Thor expedition. We believe that M. globosa is the female of Lycaea serrata Claus, described by Claus (1879b) from the male. Stephensen had 5 males of L. serrata but did not associate them with his M. globosa, perhaps because they did not occur at any of the same stations. Shoemaker’s (1945) 2 males and 1 female of L. serrata were obtained in 3 different net tows, but he recognized that they were conspecific and gave what he thought was the first description of a female L. serrata. His drawing of the whole animal does not resemble closely Stephensen’s habit figures, since the latter are of immature specimens (4 and 5 mm), whereas Shoemaker’s ovigerous female measured 10.5 mm. The appendages drawn by Stephensen were from his 8-mm female and show good agreement with those figured by Shoemaker. Shoemaker’s males agree closely with Claus’ description and illustrations of L. serrata.

We propose 1 new genus, Spinoscina. Acanthoscina spinosa Chevreux, 1914, placed in Ctenoscina by Wagler (1926), has been shown by Vinogradov (1957) to have characters of both Ctenoscina and Acanthoscina. Rather than combine these genera, we have elected to establish the new genus Spinoscina for S. spinosa.

We have omitted from this synopsis the genera Cyllias Bovallius, 1887, and Daira Milne Edwards, 1830 (synonyms: Dairilia Dana, 1953; Eudaira Bovallius, 1889). Cyllias was proposed as a genus of Cyllopodidae (now included in Vibiliidae) by Bovallius (1887b) for Hyperia tricuspidata Streets (1877). Both Dairilia and Eudaira were substituted for Daira in the mistaken belief that the latter name was preoccupied by Daira Leach, a genus of crabs. In fact, no such genus was ever published by Leach, and the crab genus Daira de Haan (1833) is a junior homonym of Daira Milne Edwards. However, the International Commission on Zoological Nomenclature (1957) has placed Daira Milne Edwards on the Official Index, thereby suppressing it, and has placed both Daira de Haan and Dairilia Dana on the Official List, making them the valid names for the crab and amphipod genera respectively. Both Cyllias Bovallius and Daira Milne Edwards were described in vague terms and were not illustrated, and we are unable to assign them even to families.

REVISED CLASSIFICATION.—With the above changes, our classification is as follows:

Order Amphipoda

Suborder Hyperiidea

Infraorder Physosomata

Superfamily Lanceoloidea

Family Lanceolidae

Chuneolidae

Microphasmidae

Superfamily Scinoidea

Family Archacoscinidae

Scinidae

Mimonectidae

Proscinidae

Infraorder Physocephalata

Superfamily Vibilioidea

Family Cystisomatidae

Vibiliidae

Paraphronimidae

Superfamily Phronimoidea

Family Hyperiidae

Dariellidae

Anchylomeridae

Phronimidae

Superfamily Lycaeopsoidea

Family Lycaeopsidae

Superfamily Platysceloidea

Family Pronoidae

Anapronoidae

Lycaeidae

Oxycephalidae

Platyscelidae

Parascelidae

Suborder HYPERIIDEA H. Milne Edwards, 1830

Body form ranging from extremely slender and elongate to nearly globular. Head and pereon generally tumid and round in cross section or somewhat flattened dorsoventrally. Eyes sometimes small, more often enormously developed, frequently covering most of surface of head. Pereonite 1 never fused with head. Coxae small, often fused with pereonites. Pleon usually powerfully developed; urosomites 2 and 3 fused (=double urosomite). Telson entire, never cleft or incised, without setae. Antenna 1 without accessory flagellum. Antennae 1 and 2 often reduced in female. Maxilla 1 with or without inner lobe; palp 1-segmented. Maxilliped with inner lobes usually fused; palp absent. Pereopods 1 and 2 infrequently simple, usually subchelate or chelate or intermediate between these 2 conditions; prehension almost always between carpus and propus rather than between propus and dactyl as in Gammaridea. One or more of pereopods 3–7 also sometimes prehensile. Pereopod 7 often reduced in size and number of segments. Uropods never with 2-segmented rami; without long marginal setae. Gills varying in number from 5 pairs (on pereonites 2–6) to 2 pairs (on pereonites 5 and 6). Gut with 1 pair of digestive caecae and no rectal glands. Heart with 2 (rarely 3) pairs of ostia.

Infraorder PHYSOSOMATA Pirlot, 1929

Head small, usually shorter than pereonite 1. Eyes small, composed of few facets, often completely absent. Mandible without molar. Maxilla 1 with inner lobe. Maxilliped almost always with inner lobes separate. Pereopods 1 and 2 simple; rarely, pereopod 1 chelate.

References: Stephensen and Pirlot, 1931:501; Vinogradov, 1957:186.

The infaorder is divided into 2 superfamilies. Since the characters used to separate the superfamilies are sometimes difficult to see, we give a key to the seven families of the infraorder.

Key to the Families of Physosomata

Superfamily SCINOIDEA, new name

Synonym: Sciniformata Stephensen and Pirlot, 1931.

Male always with body of normal form, not inflated. Mandible with well-developed lacinia mobilis. Maxilla 2 with only a few distal spines on inner and outer lobes. Maxilliped with inner lobes either completely separate or completely fused. Pereopods 5–7 very rarely with hooded dactyls. Gills on pereonite 2, when present, as large as those of following pereonites; gills rarely absent on pereonite 2 or 3. Oostegites on pereonites 2–5, usually with marginal setae.

Four families (Archaeoscinidae, Mimonectidae, Proscinidae, Scinidae).

I. Family ARCHAEOSCINIDAE Stebbing, 1904

Body form very different in male and female. Male slender, similar to Scina; total length close to 2 mm. Female with pereonites 1–5 inflated into a spherical form; head immersed in pereonite 1; pereonites 6 and 7 and pleon narrow; total length up to 6 mm. Cuticle transparent. Coxae distinct from pereonites. Telson very small, triangular, not fused with double urosomite. Eyes very small or absent. Antenna 1 with large, densely setose 1st flagellar segment; distal segments short, slender. Antenna 2 about half as long as antenna 1, 5- or 6-segmented. Mandible with long, 3-segmented palp; incisor very narrow; lacinia mobilis well developed. Maxilliped with large outer lobes; inner lobes small, fused proximally, separate distally. All pereopods simple, without chela or subchela; dactyl not hooded; carpus of pereopods 1 and 2 not broadened distally. Uropods long and very narrow; all rami free. Gills on pereonites 3–6.

One genus.

1. Archaeoscina Stebbing, 1904

FIGURE 10

Type-species: Archaeoscina bonnieri Stebbing, 1904.

With the characters of the family.

References: Bovallius, 1889:70; Woltereck, 1906: 190; Stephensen and Pirlot, 1931:534; Vinogradov, 1956:200.

II. Family MIMONECTIDAE Bovallius, 1885

Body form different in male and female. Male slender; head as high as pereonite 1. Female with pereonites 1–6 strongly inflated; pereonite 7 and pleon slender; head much smaller than pereonite 1. Total length 6–24 mm. Eyes very small and composed of few facets or completely lacking. Coxae distinct from pereonites. Telson small, triangular, not fused with double urosomite. Antenna 1 much longer than head, inserted in middle of anterior margin of head; with short peduncle, large 1st flagellar segment and a few short distal flagellar segments. Antenna 2 in male composed of few segments, not more than one-third as long as antenna 1; in female rudimentary, with inflated proximal segment and 1 or 2 stubby distal segments. Mandible without molar or palp, rarely with rudimentary 1-segmented palp; lacinia mobilis well developed. Maxilliped with large rounded outer lobes (sometimes with 1-segmented rudimentary palp); inner lobes small, separate. Pereopods all simple, without chela or subchela; dactyls not hooded. Uropods long, narrow; exopods and endopods free. Gills on pereonites 2–6.

FIGURE 10.—Archaeoscina.

Two genera.

Key to the Genera of Mimonectidae

2. Mimonectes Bovallius, 1885

FIGURE 11

Type-species: Mimonectes loveni Bovallius, 1885 (by present designation).

Mandible without palp. Pereopods 6 and 7, propus slender; dactyl at most weakly curved.

References: Stephensen and Pirlot, 1931:503; Vinogradov, 1957:205.

3. Pseudomimonectes Vinogradov, 1960

FIGURE 12

Type-species: Pseudomimonectes robustus Vinogradov, 1960a. (The genus is based on a single, sexually immature specimen.)

FIGURE 11.—Mimonectes.

FIGURE 12.—Pseudomimonectes.

Mandible with rudimentary 1-segmented palp. Pereopods 6 and 7 with broad propus and strong, claw-shaped dactyl forming a prehensile organ.

Reference: Vinogradov, 1960a:219.

III. Family PROSCINIDAE Pirlot, 1933

Body in both male and female normal, not at all or only slightly inflated; total length 4.5–18 mm. Coxae separate from pereonites. Telson small, not fused with double urosomite. Eyes absent. Antenna 1 much longer than head, inserted on dorsal part of anterior surface of head, with large 1st flagellar segment and a few short distal segments. Antenna 2 in male slender, about as long as antenna 1; in female short or rudimentary. Mandible without palp or molar; incisor serrate. Maxilliped with large outer lobes; inner lobes separate. Pereopods 1 and 2 simple; carpus not broadened. Pereopods 5–7 sometimes with hooded dactyls. Uropods long and slender, with free exopods and endopods. Gills on pereonites 2–6.

Two genera.

Reference: Pirlot, 1939:24.

Key to the Genera of Proscinidae

4. Proscina Stephensen and Pirlot, 1931

FIGURE 13

Type-species: Parascina stephenseni Pirlot, 1929. Both sexes with normal body, not inflated. Antenna 1 with 1st flagellar segment long and conical. Antenna 2 of female reduced to an unsegmented knob. Lacinia mobilis of left mandible as broad as incisor. Pereopods 5–7 without hooded dactyls.

References: Pirlot, 1939:25; Vinogradov, 1956:202; 1957:208; 1960a:222; 1964:127.

5. Mimoscina Pirlot, 1933

FIGURE 14

Type-species: Mimoscina gracilipes Pirlot, 1933.

Body of female weakly inflated. Antenna 1 with 1st flagellar segment very long, with nearly parallel margins. Antenna 2 of female composed of 4 free segments. Lacinia mobilis of left mandible not as broad as incisor. Pereopods 5–7 with sharply curved, hooded dactyls.

References: Pirlot, 1933:2; 1939:29; Vinogradov, 1960a: 222; 1962:13.

IV. Family SCINIDAE Stebbing, 1888

Body slender, usually flattened dorsoventrally; total length including antennae and uropods, 5–30 mm. Head small, wider than long. Eyes small, sometimes absent. Pereon always broader than pleon, both sometimes with middorsal spines. Pereonites 1 and 2 occasionally fused. Coxae distinct. Telson quite small. Antennae 1 as long as pereon or as pereon and pleon combined, rod-shaped, inserted on dorsal anterior margin of head, 2- or 3-segmented, with very long 2nd segment, 3rd segment minute or absent. Antenna 2 inserted on ventral anterior margin of head; in male long, slender, and folded beneath body, or sometimes rudimentary; in female always rudimentary. Mandible without palp or molar. Maxilliped with outer lobes well developed, rarely reduced; inner lobes completely fused, sometimes absent. Pereopods slender, without spoon-shaped propus and hooded dactyls; pereopod 1 simple, rarely chelate; pereopod 2 always simple. Pereopod 5 the longest; basipod of pereopod 5 often broadened and armed with marginal teeth. All uropods with endopods fused with protopods; exopods short, spiniform. Gills on pereonites 2–6, 3–6, or 4–6. Oostegites with marginal setae.

FIGURE 13.—Proscina.

FIGURE 14.—Mimoscina.

References: Wagler, 1926:319; 1927:85; Vinogradov, 1960a: 224.

Four genera.

Key to the Genera of Scinidae

6. Scina Prestandrea, 1833

FIGURE 15

Type-species: Scina ensicorne Prestandrea, 1833.

Body segments without middorsal spines. Pereonites 1 and 2 separate. Antenna 2 of male well developed, folded beneath body. Maxilliped with inner lobe present; outer lobes well developed. Pereopod 1 simple. Pereopod 5 without fusion of segments. Gills on pereonites 2–6 or 3–6.

References: Wagler, 1926:324; 1927:85; Hurley, 1956:4; Vinogradov, 1960a:224.

FIGURE 15.—Scina.

7. Acanthoscina Vosseler, 1900

FIGURE 16

Type-species: Scina acanthodes Stebbing, 1895.

Body segments with middorsal spines, 6 on pereon, 3 on pleon. Pereonites 1 and 2 fused. Second segment of male antenna 1 expanded at base. Antenna 2 of male and female represented by a pair of long sharp spines. Maxilliped without inner lobe; outer lobes reduced to two small oval plates. Pereopod 1 simple. Pereopod 5 with carpus, merus, and propus fused. Gills on pereonites 2–6.

Reference: Wagler, 1926:424.

8. Ctenoscina Wagler, 1926

FIGURE 17

Type-species: Ctenocina tenuis Wagler, 1926 (by present designation).

Body segments with middorsal spines, 5 or 7 on pereon, 3 or 4 on pleon. Pereonites 1 and 2 separate. Antenna 2 of male and female represented by a pair of long sharp spines, except in the male C. macrocarpa, which has antenna 2 composed of 5 short segments. Maxilliped without inner lobe; outer lobes well developed. Pereopod 1 chelate. Pereopod 5 without fusion of segments. Gills on pereonites 4–6.

FIGURE 16.—Acanthoscina.

FIGURE 17.—Ctenoscina.

References: Wagler, 1926:430; Vinogradov, 1964: 140.

9. Spinoscina, new genus

FIGURE 18

Type-species: Acanthoscina spinosa Chevreux, 1914.

Body segments with middorsal spines, 6 on pereon, 4 on pleon. Pereonites 1 and 2 fused. Antenna 2 of female (male unknown) represented by a pair of long sharp spines. Maxilliped with inner lobe present; outer lobes well developed. Pereopod 1 chelate. Pereopod 5 without fusion of segments. Gills on pereonites 3–6.

Reference: Vinogradov, 1957:219.

Superfamily LANCEOLOIDEA, new name

Synonym: Lanceoliformata Stephensen and Pirlot, 1931.

Body in both sexes more or less inflated. Mandible usually with weakly developed lacinia mobilis. Maxilla 2 with numerous, usually very large spines on both lobes. Maxilliped with inner lobes fused proximally, separate distally. Pereopods 6 and 7 with spoon-shaped propus and hooded dactyl, or some of posterior pereopods subchelate; rarely all pereopods simple. Gills on pereonites 2–6 or 4–6, those of pereonite 2 very small. Oostegites on pereonites 2–5, usually without marginal setae.

FIGURE 18.—Spinoscina.

Three families (Microphasmidae, Chuneolidae, Lanceolidae).

V. Family MICROPHASMIDAE Stephensen and Pirlot, 1931

Pereonites 2–4 strongly inflated; pereonites 5–7 and pleon slender; total length 3–25 mm. Cuticle transparent. Coxae separate from pereonites. Telson small, triangular, not fused with double urosomite. Head about as long as pereonite 1, with massive mouthpart region. Eye very small or absent. Antenna 1 flagellum with large 1st segment and a few very small distal segments, the latter sometimes missing. Antenna 2 5- or 6-segmented, shorter or slightly longer than antenna 1. Mandible with long, 3-segmented palp; incisor broad, lacinia mobilis very narrow. Maxilliped with large outer lobes; inner lobes fused proximally, separate distally. Pereopods 1 and 2, carpus broadened distally; propus conical. Pereopods 3–5 and 7 often prehensile. Pereopods never with hooded dactyls. Uropods narrow, with free exopods and endopods.

Three genera.

Key to the Genera of Microphasmidae

10. Microphasma Woltereck, 1909

FIGURE 19

Type-species: Microphasma agassizi Woltereck, 1909.

Eye very small. Antenna 1 short, inserted on and extending beyond anterior margin of head. Antenna 2 much shorter than antenna 1; proximal segment much inflated; gland cone very large. All pereopods with carpus distinctly shorter than propus. Pereopods 3–5 subchelate; propus broad, distal part of closing margin with row of strong spines; dactyl curved, on pereopods 3 and 4 almost half as long as propus, on pereopod 5 somewhat shorter.

References: Woltereck, 1909:153; Stephensen and Pirlot, 1931:539; Shoemaker, 1945:218.

FIGURE 19.—Microphasma.

11. Microphasmoides Vinogradov, 1960

FIGURE 20

Type-species: Microphasmoides vitjazi Vinogradov, 1960a.

Eye absent. Antenna 1 small, inserted on lateral surface of head and not reaching anterior margin of head. Antenna 2 shorter than antenna 1; proximal segment much inflated; gland cone very large. All pereopods with carpus distinctly shorter than propus. Pereopods 3 and 4 with long narrow propus. Pereopod 5 subchelate; propus with subparallel margins; distal margin with row of strong spines; dactyl short, hook-shaped. Pereopods 6 and 7 with propus narrowing distally.

Reference: Vinogradov, 1960a:213.

FIGURE 20.—Microphasmoides.

12. Mimonecteola Woltereck, 1909

FIGURE 21

Type-species: Mimonecteola diomedeae Woltereck, 1909.

Eye lacking or very small. Antennae 1 and 2 very long, inserted on and extending far beyond anterior surface of head. Antenna 2 somewhat longer than antenna 1; proximal segment not inflated; gland cone small. All pereopods with carpus as long as or nearly as long as propus. All pereopods usually simple; pereopods 5 and 7 rarely subchelate, but propus only slightly broadened.

References: Woltereck, 1909:153; Shoemaker, 1945:224; Vinogradov, 1964:118.

FIGURE 21.—Mimonecteola.

VI. Family CHUNEOLIDAE Woltereck, 1909

Body weakly arched, somewhat flattened dorsoventrally, slightly broader than high; total length 5–41 mm. Coxae not fused with pereonites. Telson short, triangular, not fused with double urosomite. Head with blunt to triangular rostrum, somewhat longer than pereonite 1. Eyes very small or absent. Antenna 1 somewhat shorter than head, inserted in lateral depression of head beneath rostrum; peduncle 3-segmented; flagellum with very large proximal segment and sometimes a few minute distal segments. Antenna 2 shorter than antenna 1, with large inflated proximal segment (containing opening of antennal gland) and minute distal segment. Mandible without palp or molar; lacinia mobilis broad. Maxilliped with large, rounded outer lobes; inner lobes small and completely separate. All pereopods with hooded dactyls; pereopods 3–7 with spoon-shaped propus. Uropods with free exopods and endopods. Gills on pereonites 2–6. Oostegites on pereonites 2–5.

One genus.

13. Chuneola Woltereck, 1909

FIGURE 22

Type-species: Chuneola paradoxa Woltereck, 1909.

With the characters of the family.

References: Pirlot, 1930:3; Vinogradov, 1956:196; 1957:201; 1960b:251.

FIGURE 22.—Chuneola.

VII. Family LANCEOLIDAE Bovallius, 1887

Body greatly inflated in larva, only slightly so in adult, total length 6–70 mm. Eyes small, rudimentary, or absent. Coxae separate from pereonites. Antenna 1 with 1st flagellar segment large, robust, nearly straight or slightly curved; distal segments small, few in number. Antenna 2 usually longer and more slender than antenna 1. Mandible with 3-segmented palp, without molar. Pereopod 1 with carpus usually more or less expanded distally. Pereopods 6 and 7 and usually pereopod 5 with spoon-shaped propus and hooded dactyl.

Key to the Genera of Lanceolidae

Six genera.

14. Lanceola Say, 1818

FIGURE 23

Type-species: Lanceola pelagica Say, 1818.

Head usually with well-developed rostrum; interantennal lobe rounded, rudimentary to well developed. Eyes small, round to oval. Mandible with 3rd segment of palp shorter than 2nd. Maxilla 1 with well-developed palp and broad inner lobe. Pereopod 1 with carpus expanded distally. Pereopods 5–7 with spoon-shaped propus and hooded dactyl.

References: Bovallius, 1887c:28; Stebbing, 1888: 1301; Stephensen, 1918:8; Woltereck, 1927:60; Barnard, 1932:254; Shoemaker, 1945:206; Vinogradov, 1957:189; 1960a:200; 1926:6; 1961:109.

FIGURE 23.—Lanceola.

15. Megalanceola Pirlot, 1935

FIGURE 24

Type-species: Lanceola stephenseni Chevreux, 1920.

Large amphipods, up to 70 mm in length. Head short; rostrum very small; interantennal lobe rather narrow, with concave lateral surface, its distal margin flaring laterad to form a rim. Eyes oval, without pigment. Mandible with lacinia mobilis nearly as broad as incisor. Maxilla 1 with narrow inner lobe. Pereopod 1 with carpus only slightly expanded distally. Pereopods 5–7 with spoon-shaped propus and hooded dactyl.

References: Pirlot, 1935:passim; 1939:8; Shoemaker, 1945:212; Vinogradov, 1964:114.

FIGURE 24.—Megalanceola.

16. Metalanceola Pirlot, 1931

FIGURE 25

Type-species: Metalanceola chevreuxi Pirlot, 1931.

Head with inconspicuous rostrum. Eyes absent. Antenna 1 with well-developed distal segments. Antenna 2 short, only slightly longer than peduncle of antenna 1. Mandible with 3rd segment of palp slightly longer and not much narrower than 2nd segment. Maxilla 1 with palp shorter than outer lobe. Maxilliped with very small inner lobes. Pereopod 1 with carpus not expanded. Pereopods 3 and 4 with strong, recurved dactyls. Pereopods 5–7 with spoon-shaped propus and hooded dactyls. Gills on pereonites 4–6.

FIGURE 25.—Metalanceola.

References: Pirlot, 1931:passim; 1939:12; Vinogradov, 1960a:210.

17. Paralanceola Barnard, 1930

FIGURE 26

Type-species: Paralanceola anomala Barnard, 1930.

Head with inconspicuous rostrum. Eyes minute, composed of only 4 facets. Antenna 1 very short and stout. Antenna 2 with last segment of peduncle elongate and slightly enlarged distally; flagellum very long, filiform, without visible sutures. Mandibular palp with long 3rd segment. Pereopod 1 with carpus not expanded; propus produced into lobe above insertion of dactyl so that dactyl is inserted subterminally. Pereopods 5–7 with spoon-shaped propus and hooded dactyl.

FIGURE 26.—Paralanceola.

Reference: Barnard, 1930:398.

18. Prolanceola Woltereck, 1907

FIGURE 27

Type-species: Prolanceola vibiliformis Woltereck, 1907.

Body not tumid. Head only slightly shorter than pereonite 1, with short rostrum. A row of 4 eyespots extending dorsally from usual eye on side of head. Antenna 1 with 1st flagellar segment tapering and becoming very slender distally. Antenna 2 much longer than antenna 1; flagellum 6-segmented, longer than peduncle. Mandible with rather narrow incisor and well-developed lacinia mobilis; palp with long 3rd segment. Pereopod 1 subchelate, with triangular carpus; propus produced distally into lobes above and below insertion of dactyl. Pereopods 6 and 7, but not pereopod 5, with hooded dactyls and spoon-shaped propus.

References: Woltereck, 1909:157; Vinogradov, 1957:198.

19. Scypholanceola Woltereck, 1905

FIGURE 28

Type-species: Scypholanceola vanhoeffeni Woltereck, 1909.

Large amphipods, body length up to about 60 mm. Head with well-developed rostrum; lateral surface with 2 concavities separated by ridge running obliquely posteroventrad from interantennal lobe. Eyes without refracting elements or pigment, in form of band with ends lying in concavities and connected across ridge. Coxae 2–5 sharply pointed anteriorly. Mandible with 3rd segment of palp shorter than 2nd. Maxilla 1 with well-developed palp and broad inner lobe. Pereopod 1 with carpus expanded distally. Pereopods 5–7 with spoon-shaped propus and hooded dactyl.

FIGURE 27.—Prolanceola.

FIGURE 28.—Scypholanceola.

References: Woltereck, 1909:161; 1927:65; Chevreux, 1920:7; Shoemaker, 1945:215; Vinogradov, 1956:197; 1960a:207.

Infraorder PHYSOCEPHALATA, new name

Head large, longer than pereonite 1. Eyes large (small to moderately large in Vibilia), occupying most of head surface (excluding rostrum in Oxycephalidae). Maxilla 1 without inner lobe. Maxilliped with inner lobes completely fused. Pereopods 1 and 2 usually chelate or subchelate, less commonly simple.

Four superfamilies.

Key to the Superfamilies of Physocephalata

Superfamily VIBILIOIDEA, new name

Antenna 1 inserted on anterior surface of head; flagellum with large straight 1st segment and one or a few small or rudimentary additional segments inserted terminally. Antenna 2 inserted on anterior or ventral surface of head, short, composed of a few segments or rudimentary. Pereopod 5 never with large subchela.

Three families (Vibiliidae, Cystisomatidae, Paraphronimidae).

Key to the Families of Vibilioidea

VIII. Family VIBILIIDAE Dana, 1852

Body compact, moderately slender; total length from about 5 to 20 mm. Head small to moderately large, subquadrangular or rounded. Eyes small to moderate-sized and separated, or large and occupying most of head surface. Pereonites all separate. Coxae separate from pereonites. Telson short, triangular or semicircular. Antenna 1 with short 3-segmented peduncle; 1st segment of flagellum large, straight, spatuliform or conical; remaining segments rudimentary. Antenna 2 filiform, subequal to antenna 1. Mandible with palp and molar process. Maxilliped with short rounded inner lobe; outer lobes rounded at apices, with straight medial margins and rounded lateral margins. Pereopod 1 simple; pereopod 2 chelate. Pereopods 5 and 6 the longest; pereopod 7 short, segments sometimes reduced in number, distal segment always digitiform. Uropods with free exopods and endopods. Gills on pereonites 2–6. Oostegites on pereonites 2–5.

Two genera.

Key to the Genera of Vibiliidae

20. Vibilia H. Milne-Edwards, 1830

FIGURE 29

Type species: Vibilia peronii H. Milne-Edwards, 1830.

Head rather small, subquadrangular. Eyes small to moderately large but never occupying most of head surface. Antenna 1 with broad, laterally flattened flagellum. Antenna 2 inserted on anterior surface of head. Pereopod 7 usually with complete number of segments, rarely with only 3 segments distal to basipod. Protopod of uropod 3 shorter than that of uropod 1.

References: Behning, 1913, 1925; Stephensen, 1918:33; Chevreux and Fage, 1925:382; Pirlot, 1929:91; Laval, 1963.

FIGURE 29.—Vibilia.

21. Cyllopus Dana, 1852

FIGURE 30

Type-species: Cyllopus magellanicus Dana, 1852. Head large, almost globular. Eyes large, occupying most of head surface. Antenna 1 with slender conical flagellum. Antenna 2 inserted on ventral surface of head. Pereopod 7 with basipod longer than remaining segments combined, the latter comprising 5 or 3 segments. Protopod of uropod 3 longer or shorter than that of uropod 1.

References: Barnard, 1930:405; Hurley, 1955:129.

FIGURE 30.—Cyllopus.

IX. Family CYSTISOMATIDAE Willemoes-Suhm, 1875

Body large, cuticle thin, almost completely transparent; musculature very weak. Total length up to 120 mm. Head very large, rounded dorsally; ventral surface flat or slightly concave, with dentate border. Eyes comprising 2 oval areas on dorsal surface of head. Short middorsal spines often present on pereonites and pleonites. Pereonites 1 and 2 or even 1–4 fused. Coxae fused with pereonites. Telson very small. Antenna 1 with 1 or 2 poorly defined peduncle segments; flagellum composed of a slender lanceolate proximal segment and a minute distal segment. Antenna 2 represented by a pair of small spines on ventral surface of head. Mandible with molar; palp absent. Maxilla 2 without inner lobe. Maxilliped with long inner lobe, widening distally; outer lobes pointed at apices. Pereopod segments with serrate margins. Pereopods 1 and 2 distinctly chelate. Pereopod 5 the longest. Pereopod 7 sometimes prehensile, with dactyl closing against distal end of propus. Uropod 2 absent. Endopod of uropods 1 and 3 fused with protopod. Gills on pereonites 2–6 or 4–6. Oostegites on pereonites 2–5, those on pereonites 4 and 5 rudimentary.

One genus.

22. Cystisoma Guérin-Méneville, 1842

FIGURES 31, 32

Type-species: Cystisoma neptunus Guérin-Méneville, 1842.

With the characters of the family.

References: Stephensen, 1918:56; Barnard, 1932:268; Woltereck, 1903:passim; 1904:passim; Willemoes-Suhm, 1874: passim.

X. Family PARAPHRONIMIDAE Bovallius, 1887

Body rather slender, transparent; total length 6–15 mm. Head large, almost cubical. Eyes large, occupying most of head surface. Pereonites all separate. Coxae fused with pereonites. Telson very short. Antenna 1 shorter than head; peduncle 3-segmented; first segment of flagellum enlarged, the others rudimentary. Antenna 2 inserted on ventral margin of head, rudimentary in female, slightly longer than antenna 1 in male, composed of a few segments. Mandible without palp and without molar. Maxilliped with inner and outer lobes fused into a single broad plate. Pereopods 1 and 2 much shorter than following pereopods. Pereopod 1 weakly subchelate, merus produced into short spoon-shaped lobe beneath carpus. Pereopod 2 with propus produced into a thin hollowed process on either side of dactyl. Pereopods 3–6 slender, subequal; pereopod 7 shorter. Uropods with free exopods and endopods. Gills on pereonites 3–6. Oostegites on pereonites 2–5.

FIGURE 31.—Cystisoma: diagram of head, ventral.

FIGURE 32.—Cystisoma.

One genus.

23. Paraphronima Claus, 1879

FIGURE 33

Type-species: Paraphronima gracilis Claus, 1879a.

With the characters of the family.

References: Claus, 1879a:64; Bovallius, 1889:22; Stebbing, 1888:1335; Vosseler, 1901:94.

FIGURE 33.—Paraphronima.

Superfamily PHRONIMOIDEA, new name

Antenna 1 inserted on anterior surface of head; flagellum of male with long straight 1st segment and shorter, terminally inserted distal segments; flagellum of female 1-segmented. Antenna 2 inserted on anterior surface of head; flagellum of male long, multisegmented; flagellum of female short, composed of a few segments, sometimes rudimentary or absent. Pereopod 5 sometimes with large subchela.

Four families (Hyperiidae, Dairellidae, Phronimidae, Phrosinidae).

Key to the Families of Phronimoidea

XI. Family HYPERIIDAE Dana, 1852

Body compact, form rather variable, most often subglobular in female, more slender in male; total length, 2–30 mm. Head usually large, globular. Eyes usually large and occupying most of head surface. Pereonites all separate, or up to 5 anterior pereonites fused. Coxae separate or fused with pereonites. Telson small to moderate-sized, rounded or triangular. Antenna 1 with long filiform flagellum in male, with short 1-segmented flagellum in female. Antenna 2 similar to antenna 1, but often more reduced in female. Mandible with molar; palp present in both sexes or absent in female. Maxilliped quite variable; outer lobes usually well developed, sometimes fused medially; inner lobe well developed, reduced, or even absent. Pereopod 1 simple, subchelate, or chelate. Pereopod 2 chelate. Pereopods 3–7 varying in relative lengths, but pereopod 7 not reduced. Uropods with free exopods and endopods. Gills on pereonites 2–6. Oostegites on pereonites 2–5.

Thirteen genera.

Key to the Genera of Hyperiidae

24. Hyperia Latreille, in Desmarest, 1823

FIGURE 34

Type-species: Cancer medusarum O. F. Müller, 1776.

Large species, 10–30 mm in length. Head deeper than long. Pereon more or less dilated in female; all pereonites free. Coxae not fused with pereonites. Antenna 1 with 3-segmented peduncle. Mandible with serrate incisor; palp present in both sexes. Maxilla 1 with outer lobe having 5 apical spines; palp with robust spine at mediodistal corner. Maxilliped with inner lobe well developed, usually with 2 terminal spines. Pereopod 1 subchelate or barely chelate; carpal process only slightly developed. Pereopod 2 chelate; carpal process spoon-shaped, bearing spines along edge of spoon. Pereopods 1 and 2, carpus, and propus with many spines. Pereopods 5–7 shorter than pereopods 3 and 4. Uropod 3 of male with broad endopod.

References: Bovallius, 1889:129; Sars, 1890:6; Bowman, 1972:5.

FIGURE 34.—Hyperia.

25. Bougisia Laval, 1966

FIGURE 35

Type-species: Bougisia ornata Laval, 1966.

Head with well-developed rostrum and acute lateral lobes. Eyes small, lateral. Pereonites 1 and 2 fused. Coxal plates separate from pereonites. Antenna 2 of female composed of a single segment. Mandible with palp in both sexes. Maxilla 1 with rather broad palp. Maxilliped with very short inner lobe. Pereopod 1 subchelate; pereopod 2 chelate, with short spoon-shaped carpal process. Pereopods 3–7 subequal; dactyl of pereopod 5 prehensile, closing against expanded distal margin of propus.

Reference: Laval, 1966.

FIGURE 35.—Bougisia.

26. Hyperiella Bovallius, 1887

FIGURE 36

Type-species: Hyperiella antarctica Bovallius, 1887b.

Body small to moderate-sized; pereon rather plump; pereonites all separate. Antenna 1 and antenna 2 of female 4-segmented. Mandible with palp in both sexes; incisor with serrate margin. Maxilla 1 with inner lobe having 5 terminal spines. Maxilla 2 with 1 terminal spine on inner lobe, 2 or 3 on outer lobe. Maxilliped with inner lobe well developed, with 2 terminal spines; outer lobes separate. Pereopod 1 subchelate; pereopod 2 chelate; posterior margin of propus serrate in pereopods 1 and 2. Pereopods 3 and 4 with posterior margins of carpus and propus serrulate; posterior margins of carpus bearing a few slender spines. Pereopod 5 (or pereopods 5 and 6) much longer than pereopods 3 and 4; pereopod 7 (or pereopods 6 and 7) subequal to pereopods 3 and 4. Dactyls of pereopods 3–5 very long. Uropod 3 elongate, except in H. macronyx.

References: Bovallius, 1889:242; Stebbing, 1888: 1403; Walker, 1907:7; Bowman, 1973:26.

FIGURE 36.—Hyperiella.

27. Hyperietta Bowman, 1973

FIGURE 37

Type-species: Hyperia luzoni Stebbing, 1888.

Small species, with body rather compressed laterally. Head rather short anteroposteriorly; eyes occupying most of its surface. Pereonites 1 and 2 fused in both sexes. Coxae fused with pereonites. Telson inserted distinctly anterior to insertion of protopod of uropod 3. Female telson at least half as long as protopod of uropod 3. Epistome small, inconspicuous. Antenna 1 of female 2-segmented. Antenna 2 of female 1-segmented, rudimentary; gland cones rather inconspicuous, in anterior view converging medially. Mandible with smooth incisor and dentate lacinia; molar narrow; with palp in male, without in female. Maxilla 1 with outer lobe having 3 terminal spines. Maxilliped with outer lobes fused medially; inner lobe rudimentary. Pereopod 1 simple, weakly subchelate or barely chelate. Pereopod 2 chelate, with spoon-shaped carpal process bearing marginal spines. Pereopods 3 and 4 of female with 1 conspicuous spine on posterior margin of merus, 2 spines on posterior margin of carpus; pereopods 3 and 4 of male with shorter spines in same positions. Distal margin of propus of pereopods 6 and 7 and sometimes of pereopod 5 produced into spinose lobe medial to base of dactyl; dactyl unarmed. Uropods slender, margins of exopods and endopods smooth or with extremely fine serrations.

FIGURE 37.—Hyperietta.

References: Stebbing, 1888:1382; Vosseler, 1901: 64; Stephensen, 1924:83; Yang, 1960:33; Bowman, 1973:55.

28. Hyperioides Chevreux, 1900

FIGURE 38

Type-species: Hyperioides longipes Chevreux, 1900.

Small species with body rather compressed laterally. Head globular, produced anteriorly above insertion of antenna 1; eyes occupying most of its surface or limited entirely to dorsal surface. Pereonites 1 and 2 fused in both sexes. Coxae fused with pereonites. Posterior elevation of pleonite 1 of male pronounced. Telson rather short. Antenna 1 of female 2- or 3-segmented. Antenna 2 of female 1-segmented, moderately long, with characteristic sinuous shape in lateral view; gland cone distally rounded. Mandible with dentate incisor; with palp in male, without in female. Maxilla 1 with outer lobe having 3 large terminal spines and a smaller subterminal spine. Maxilla 2 with outer lobe having 1 terminal and 1 subterminal spine; inner lobe with 1 terminal spine. Maxilliped having inner lobe well developed, with 2 terminal spines. Pereopods 1 and 2 chelate, with spoon-shaped carpal process bearing marginal spines. Pereopods 5 and 6 distinctly longer than pereopods 3 and 4 and pereopod 7; dactyls of pereopods 5–7 long, with comb of fine setules on proximal 3rd of anterior margin. Uropods, exopods notched on lateral margins.

FIGURE 38.—Hyperioides.

References: Chevreux, 1900:143; Stebbing, 1888:1379; Vosseler, 1901:60; Bowman, 1973:30.

29. Hyperionyx Bowman, 1973

FIGURE 39

Type-species: Hyperia macrodactyla Stephensen, 1924.

Small species. Head globular, eyes occupying most of its surface. Pereonites 1–3 fused in both sexes. Coxae fused with pereonites. Telson very short. Antenna 1 of female 2-segmented. Antenna 2 of female 1-segmented, slender, moderately long; gland cone small, triangular. Mandible reduced [?]; with palp in male, without palp in female. Maxilla 1 with outer lobe having 1 terminal spine; palp with very few marginal setae. Maxilla 2 with few setae; outer lobe with 2 terminal spines; inner lobe very short. Maxilliped with very few setae; inner lobe slender; outer lobes obovate. Pereopod 1 subchelate. Pereopod 2 chelate, with spinose, spoon-shaped carpal process. Pereopods 3–7 with strong, curved, unarmed dactyls. Pereopods 3 and 4 and pereopod 6 subequal; pereopod 5 much shorter; pereopod 7 somewhat longer.

References: Stephensen, 1924:90; Yang, 1960:35; Bowman, 1973:71.

FIGURE 39.—Hyperionyx.

30. Hyperoche Bovallius, 1887

FIGURE 40

Type-species: Metoecus medusarum Krøyer, 1838.

Pereonites all separate. Coxae separate from pereonites. Antenna 1 with 3-segmented peduncle. Mandible with palp in both sexes; molar laminate, triangular, nearly smooth. Maxilla 1 with broad palp. Maxilliped with inner lobe well developed; outer lobes separate. Pereopods 1 and 2 distinctly chelate; carpal process laterally compressed, knife-shaped. Pereopods 3–7 subequal; pereopods 3 and 4 with posterior margin of carpus produced into thin ridge which may end distally in a tooth.

FIGURE 40.—Hyperoche.

References: Bovallius, 1889:83; Senna, 1906:154; Steuer, 1911:674.

31. Iulopis Bovallius, 1887

FIGURE 41

Type-species: Iulopis loveni Bovallius, 1887b (by present designation).

Body and pereopods covered with fine setae, giving the species a furry appearance. Pereonites all free. Pereonites 2–7 raised into transverse rounded folds. Coxae not fused with pereonites. Antenna 2 much reduced in female. Mandible with palp in male, without palp in female. Maxilliped with obsolete inner lobe and sparsely spinose outer lobes. Pereopod 1 subchelate; pereopod 2 chelate; carpal process narrowly hollowed in both. Pereopods 3–7 subequal; pereopods 3 and 4 may be slightly shorter or slightly longer than pereopods 5–7.

References: Bovallius, 1889:116; Spandl, 1927: 159; Shoemaker, 1945:238.

FIGURE 41.—Iulopis.

32. Lestrigonus Milne Edwards, 1830

FIGURE 42

Type-species: Lestrigonus fabrei Milne Edwards, 1830.

Small species with rather plump pereon. Head globular; eyes occupying most of its surface. Some of anterior pereonites fused, first 2 to first 5 in female, first 2 to first 4 in male, always more in female. Coxae fused with pereonites. Telson of moderate size. Antenna 1 of female 2–segmented. Antenna 2 of female 1-segmented, usually very small; gland cone conspicuous, with rounded or pointed apex. Epistome prominent, strongly convex anteriorly. Mandible with dentate incisor, with palp in male, without palp in female. Maxilla 1 with outer lobe having 3 large terminal spines and usually a smaller subterminal spine. Maxilliped with outer lobes separate, tapering distally; inner lobe well developed. Pereopod 1 subchelate or barely chelate. Pereopod 2 distinctly chelate; carpal process spoon-shaped with spines along margin of spoon. Pereopods 5–7 usually longer than pereopods 3 and 4; pereopods 5 and 7 subequal, slightly shorter than pereopod 6.

References: Vosseler, 1901:66; Yang, 1960:15; Laval, 1968a:passim; Bowman, 1973:33.

FIGURE 42.—Lestrigonus.

33. Parathemisto Boeck, 1870

FIGURE 43

Type-species: Parathemisto abyssorum Boeck, 1870.

Body rather compressed laterally. Pereonites all free. Coxae not fused with pereonites. Antenna 1 with 3-segmented peduncle. Mandible with palp in both sexes. Pereopod 1 simple. Pereopod 2 chelate, with spoon-shaped carpal process. Pereopods 3 and 4 with dilated carpus, forming prehensile organ with propus. Pereopods 5–7 longer than pereopods 3 and 4.

References: Stephensen, 1924:94; Barnard, 1930: 419; Hurley, 1955:151; Bowman, 1960:passim; Kane, 1966:passim.

FIGURE 43.—Parathemisto.

34. Pegohyperia Barnard, 1932

FIGURE 44

Type-species: Pegohyperia princeps Barnard, 1932.

Body rather elongate. Head short; eyes without facets. Pereonites all free, with rounded transverse folds. Coxae not fused with pereonites; coxae 3–5 much deeper than others. Telson nearly as long as protopod of uropod 3. Antenna 1 of female with large laminate flagellum. Mandible with palp in both sexes; molar laminate, triangular. Pereopods 1 and 2 distinctly chelate; carpal process laterally compressed, knife-shaped. Pereopods 5 and 6 somewhat longer than pereopods 3, 4, and 7. Protopods of uropods broad.

References: Barnard, 1932:277; Hurley, 1960:112.

FIGURE 44.—Pegohyperia.

35. Phronimopsis Claus, 1879

FIGURE 45

Type-species: Phronimopsis spinifera Claus, 1879a.

Body slender in male; pereon somewhat inflated in female. Head higher than long. Pereonites 1 and 2 fused. Coxae fused with pereonites. Telson very short. Antenna 1 of female with 1-segmented peduncle produced into long spine beneath flagellum. Mandible with palp in male, without palp in female. Pereopod 1 simple; dactyl covered with short setae. Pereopod 2 chelate; dactyl closing against anteroventral process of strongly swollen propus. Pereopods 3–7 prehensile; dactyl closing against expanded distal margin of propus. Pereopods 5–7 subequal, longer than pereopods 3 and 4. Uropods slender.

FIGURE 45.—Phronimopsis.

References: Claus, 1879a:63; Bovallius, 1889:318; Vester, 1900:passim; Vosseler, 1901:51.

36. Themistella Bovallius, 1887

FIGURE 46

Type-species: Themistella steenstrupi Bovallius, 1887b.

Small species with rather broad pereon. Head rather broad; eyes occupying most of its surface. Pereonites 1–5 fused in both sexes. Coxae fused with pereonites. Telson very short. Antenna 1 of female 2-segmented, rather long. Antenna 2 of female 1-segmented, rudimentary; gland cone small. Mandible with serrate incisor; with palp in male, without palp in female. Maxilliped, outer lobes narrow, tapering distally; inner lobe almost completely absent, apparently represented by slight unarmed bulge on basal segment. Pereopods with rather broad segments. Pereopods 1 and 2 chelate; carpal process gauge-shaped, with marginal spines. Pereopod 5 much longer than pereopods 6 and 7. Dactyls of pereopods 6 and 7 with flexure slightly distal to midlength.

References: Bovallius, 1889:312; Bowman, 1973: 63.

FIGURE 46.—Themistella.

XII. Family DAIRELLIDAE Bovallius, 1887

Body moderately slender, but pereon of female somewhat expanded, transparent; total length 6–10 mm. Head large, globular in female, somewhat smaller in male. Eyes large, occupying most of head surface. Pereonites 1 and 2 fused dorsally. Coxae fused with pereonites. Telson rounded, very short. Antenna 1 with short 3-segmented peduncle; flagellum of male long, filiform, multisegmented; flagellum of female composed of a single cylindrical segment. Antenna 2 of male long, multisegmented; absent in female. Mandible without palp, with molar. Maxilliped with inner and outer lobes fused into a single subquadrate plate. Pereopods 1 and 2 slender, simple, only slightly shorter than pereopods 3–7, which are subequal. Uropods with free exopods and endopods. Gills on pereonites 2–6. Oostegites on pereonites 2–5.

One genus.

37. Dairella Bovallius, 1887

FIGURE 47

Type-species: Paraphronima californica Bovallius, 1885.

With the characters of the family.

The mouthparts of Dairella have been described by Stebbing (1888) and Bovallius (1889). The small size makes dissection of these mouthparts difficult, and it is not surprising that both authors misinterpreted maxilla 2 and maxilliped. The maxilliped is a single subquadrate plate with a slightly concave distal margin bearing a few spines at the distolateral corners. Stebbing considered this plate to be the base only of the maxilliped. The pyriform maxilla 2 lies directly in front of and close to the maxilliped. Both Bovallius and Stebbing believed it to be the outer lobe of the maxilliped, but careful dissection showed it to be completely separate from the latter throughout its entire length. The reason for Stebbing’s failure to find maxilla 2 is now obvious, but the nature of the structure that Bovallius described as maxilla 2 is not clear.

References: Stebbing, 1888:1342; Bovallius, 1889:332.

FIGURE 47.—Dairella.

XIII. Family PHRONIMIDAE Dana, 1853

Body slender, elongate; total length 5–40 mm. Head subconical, widest dorsally, narrowing and prolonged ventrally. Eyes composed of a larger dorsal part and a smaller bulging ventrolateral part. Pereonites all separate or pereonites 1 and 2 fused. Coxae fused with pereonites. Telson very small. Antenna 1 in male long, multisegmented, first segment of flagellum distinctly enlarged; antenna 1 of female very short, 2-segmented. Antenna 2 of male long, multisegmented, sometimes absent; antenna 2 of female reduced to a tubercle. Mandible without palp in both sexes. Maxilliped with slender outer lobes and short inner lobe. Pereopods 1 and 2 simple, subchelate, or chelate depending on development of carpal process; propus produced into thin triangular process on either side of dactyl. Pereopods 3, 4, 6, and 7 simple, slender; pereopod 7 not reduced. Pereopod 5 ending in subchelate claw. Uropods slender, with free exopods and endopods. Gills on pereonites 4–6. Oostegites on pereonites 2–5.

Two genera.

Key to the Genera of Phronimidae

38. Phronima Latreilie, 1802

FIGURE 48

Type-species: Cancer sedentarius Forskål, 1775. Body moderately slender. Pereonites all free. Pereopod 5 with carpus markedly widened distally; anterodistal angle forming strong pointed process; anterior margin smooth. Uropod 2 present in both sexes; endopod sometimes reduced but never missing.

FIGURE 48.—Phronima.

References: Claus, 1879a:passim; Vosseler, 1901:1; Stephensen, 1924:112; Shih and Dunbar, 1963: passim; Shih, 1969:passim.

39. Phronimella Claus, 1871

FIGURE 49

Type-species: Phronima elongata Claus, 1862.

Body and pereopods extremely slender. Pereonites 1 and 2 fused. Pereopod 5 with carpus elongate, only slightly widened distally, without clearly defined anterodistal angle; anterior margin dentate. Uropod 2 absent in female, rudimentary in male, sometimes with small exopod.

References: Same as for Phronima.

FIGURE 49.—Phronimella.

XIV. Family PHROSINIDAE Dana, 1853

Body rather compact; total length 4–30 mm. Head large, globular; eyes occupying most of head surface. Pereonites all separate or pereonites 1 and 2 fused. Coxae separate from pereonites. Telson well developed, oval or triangular. Antenna 1 with long filiform flagellum in male; with 1-segmented flagellum in female, sometimes reduced. Antenna 2 similar to antenna 1 in male, rudimentary in female. Mandible with palp in male, without palp in female. Maxilliped with slender outer lobes and well-developed inner lobe.

Pereopods 1 and 2 simple; one or more of following pereopods prehensile, subchelate; pereopod 5 always the longest, with large subchela; pereopod 7 reduced in length or number of segments. Uropods composed each of a single leaflike segment only. Gills on pereonites 2–6. Oostegites on pereonites 2–5.

Three genera.

Key to the Genera of Phrosinidae

40. Phrosina Risso, 1822

FIGURE 50

Type-species: Phrosina semilunata Risso, 1822.

Head produced into 2 rostral points. Pereonites 1 and 2 fused. Telson round-triangular, nearly as wide as urosome. Antenna 1 of female very short. Antenna 2 absent in female. Pereopods 3 and 4 subchelate, with sharp carpal process; distal margin of carpus dentate, with larger teeth in pereopod 4. Pereopods 5 and 6 subchelate; carpus relatively short and broad, with dentate distal margin; propus much longer than carpus. Pereopod 7 composed of broad basis and sometimes a rudimentary second segment. Uropods rounded, with smooth margins. Gills without folds.

References: Stebbing, 1888:1424; Vosseler, 1901:89.

FIGURE 50.—Phrosina.

41. Anchylomera H. Milne-Edwards, 1830

FIGURE 51

Type-species: Anchylomera blossevillii H. Milne-Edwards, 1830.

Head globular, without rostrum. Pereonites 1 and 2 fused. Telson round-triangular, nearly as wide as urosome. Antenna 1 of female very short. Antenna 2 of female reduced to a tubercle. Pereopods 3 and 4 subchelate, with sharp carpal process; distal margin of carpus smooth. Pereopod 5 with broad basis, having greatest width proximally; carpus with broad distal margin with rounded teeth. Pereopod 6 prehensile, with swollen and rounded carpus. Pereopod 7 with all segments present; dactyl digitiform. Uropods rounded, with smooth margins. Gills with folds.

References: Bovallius, 1889:408; Vosseler, 1901:88.

FIGURE 51.—Anchylomera.

42. Primno Guérin-Méneville, 1836

FIGURE 52

Type-species: Primno macropa Guérin-Méneville, 1836.

Head produced into single, very short rostrum. Pereonites 1 and 2 separate. Telson much narrower than urosome. Antenna 1 of female longer than head. Antenna 2 of female reduced to a tubercle. Pereopods 3, 4, and 6 simple, with some teeth on margin of carpus. Pereopod 5 prehensile, entire anterior margin of carpus dentate; propus shorter than carpus. Pereopod 7 with all segments present, dactyl digitiform. Uropods pointed; uropods 2 and 3 with teeth on outer margins. Gills without folds.

References: Stebbing, 1888:1440; Bovallius, 1889:397; Vosseler, 1901:87.

FIGURE 52.—Primno.

LYCAEOPSOIDEA, new superfamily

Antenna 1 inserted on anterior surface of head; flagellum of male with large triangular 1st segment and a few small distal segments inserted subterminally; flagellum of female short, composed of a few segments. Antenna 2 absent in female; in male inserted on ventral surface of head, very short, curved, composed of a few segments. Pereopod 5 never with large subchela. Pereopod 7 reduced in size, but all segments present.

One family (Lycaeopsidae).

XV. Family LYCAEOPSIDAE Chevreux, 1913

Body slender, somewhat compressed laterally; total length 3–6 mm. Head globular, higher than long; eyes large, occupying most of head surface. Coxae not fused with pereonites. Telson usually rather narrow, not fused with double urosomite. Mandible with palp in male, without palp in female. Maxilliped with outer lobes pointed, only slightly longer than inner lobe.

Pereopods 1 and 2 simple. Pereopod 5 long and very slender in male, normal in female and young male. Pereopod 6 long, with broad segments, especially in male. Uropods 1–3 having exopods and endopods not fused with protopods. Uropod 3 with endopod modified in male. Gills on pereonites 5 and 6, without folds. Oostegites on pereonites 2–5.

One genus.

43. Lycaeopsis Claus, 1879

FIGURE 53

Type-species: Lycaeopsis themistoides Claus, 1879b.

With the characters of the family.

References: Pirlot, 1930:27; 1939;42; Spandl, 1927:213.

Superfamily PLATYSCELOIDEA, new name

Antenna 1 inserted on ventral surface of head; flagellum of male with large curved 1st segment and a few small distal segments inserted terminally, or more commonly subterminally, on 1st segment; flagellum of female usually curved, composed of a few segments. Antenna 2 inserted on ventral surface of head; in male composed of 5 usually long segments (3 peduncular, 2 flagellar) that fold on one another in zigzag fashion; in female short, composed of a few segments, or absent. Pereopod 5 never with large subchela. Pereopod 7 reduced in size and sometimes in number of segments.

Six families (Pronoidae, Anapronoidae, Lycaeidae, Oxycephalidae, Platyscelidae, Parascelidae).

References: Claus, 1887; Spandl, 1927.

FIGURE 53.—Lycaeopsis.

Key to the Families of Platysceloidea

XVI. Family PRONOIDAE Claus, 1879

Body compact, more or less laterally compressed; total length 3–16 mm. Head globular; eyes large, occupying most of head surface. Coxae not fused with pereonites. Telson triangular, separate from or very rarely fused with double urosomite. Antenna 2 of female short, with few segments. Mandible with palp in both sexes. Maxillae 1 and 2 well developed. Pereopod 1 simple, sometimes subchelate. Pereopod 2 simple or chelate. Pereopod 5 longer than pereopod 6; basis somewhat broadened. Pereopod 6 having basis much broader than that of pereopod 5, but not operculate; distal segments inserted subterminally. Pereopod 7 with well-developed basis followed by 1 or 2 rudimentary segments; all segments may be present in juveniles. Uropods usually with free rami; endopods of uropods 2 and 3 rarely fused with protopods. Gills on pereonites 2–6, with or without folds. Oostegites on pereonites 2–5.

Five genera.

References: Claus, 1887:47–55; Spandl, 1927:216–227.

Key to the Genera of Pronoidae

44. Pronoe Guérin-Méneville, 1836

FIGURE 54

Type-species: Pronoe capito Guérin-Méneville, 1836.

Body rather slender; double urosomite about as wide as long. Telson very short, not fused with double urosomite. Antenna 2 of male short, folded only once or twice. Pereopod 1 simple; basis oval, almost as broad as long. Pereopod 2 simple. Pereopod 5 distinctly longer than pereopod 6. Pereopod 7 composed of basis and very small second segment. Uropod 1 with exopod and endopod pointed. Uropod 2 with exopod pointed, endopod rounded. Uropod 3 much longer than telson. Gills without folds.

References: Claus, 1887:48; Stebbing, 1888:1507.

FIGURE 54.—Pronoe.

45. Eupronoe Claus, 1879

FIGURE 55

Type-species: Eupronoe maculata Claus, 1879b.

Body rather stout; anterior pereonites short, appearing as though compressed together longitudinally; double urosomite about as wide as long. Telson moderately long, not fused with double urosomite. Antenna 2 of male long, with very short terminal segment. Pereopod 1 simple or subchelate; basis twisted in male. Pereopod 2 chelate; carpal process pointed or rounded. Pereopod 5 distinctly longer than pereopod 6. Pereopod 7 with pear-shaped basis and small elliptical second segment. Uropod 1 with exopod and endopod pointed. Uropods 2 and 3 with exopod and endopods rounded, very delicate; uropod 3 much longer than telson. Gills with folds.

References: Claus, 1887:50; Spandl, 1927:222; Hurley, 1955:175.

FIGURE 55.—Eupronoe.

46. Paralycaea Claus, 1879

FIGURE 56

Type-species: Paralycaea gracilis Claus, 1879b. Body rather stout; double urosomite somewhat broader than long; telson long, fused with or separate from double urosomite. Antenna 2 of male with long terminal segment. Pereopods 1 and 2 simple. Pereopod 5 distinctly longer than pereopod 6. Pereopod 7 with narrow basis; distal segments sometimes separate, usually fused into short recurved segment. Uropod 2 having endopod fused with or separate from protopod. Uropod 3 having endopod fused with protopod. Uropods 1 and 2 having exopods and endopods pointed. Uropod 3 slightly longer than telson; exopod pointed; endopod obtusely rounded. Gills without folds.

References: Claus, 1887:63; Pirlot, 1930:30.

FIGURE 56.—Paralycaea.

47. Parapronoe Claus, 1879

FIGURE 57

Type-species: Parapronoe crustulum Claus, 1879b.

Body rather slender; double urosomite distinctly longer than wide; telson long, not fused with double urosomite. Antenna 2 of male moderately long, with very short terminal segment. Pereopod 1 simple. Pereopod 2 chelate; carpal process pointed, about as long as propus. Pereopod 5 much longer than pereopod 6; basis of pereopod 6 with concave distal margin. Pereopod 7 with broad basis and very small additional segments. Uropods with pointed endopods and exopods; uropod 3 only slightly longer than telson. Gills with folds.

References: Claus, 1887:53; Stebbing, 1888:1521; Barnard, 1930:427; Pirlot, 1939:52.

FIGURE 57.—Parapronoe.

48. Sympronoe Stebbing, 1888

FIGURE 58

Type-species: Parapronoe parva Claus, 1879b.

Body rather slender; double urosomite about twice as long as wide; telson very short, not fused with double urosomite. Antenna 2 of male long, with very short terminal segment. Pereopod 1 simple. Pereopod 2 chelate; carpal process rounded, about half as long as propus. Pereopod 5 much longer than pereopod 6; basis of pereopod 6 with rather narrow distal margin produced into broadly rounded anterior lobe and narrow, pointed posterior process. Pereopod 7 with pear-shaped basis and 2 very small additional segments. Uropods 1 and 2 wtih pointed endopods and exopods; uropod 3 much longer than telson, endopod rounded, exopod bluntly pointed. Gills with folds.

References: Stebbing, 1888:1533; Spandl, 1927:225.

FIGURE 58.—Sympronoe.

XVII. ANAPRONOIDAE, new family

Body rather plump, not laterally compressed. Head globular; eyes large, occupying most of head surface. Double urosomite shorter than wide. Telson triangular, not fused with double urosomite. Antenna 2 of female short, with few segments. Mandible without palp in both sexes. Maxillae 1 and 2 moderately well developed. Pereopod 1 subchelate; pereopod 2 chelate. Pereopod 5 longer than pereopod 6; basis of both broad, but not operculate. Pereopod 6 with distal segments inserted on medial surface of basis proximal to its distal end, their combined length subequal to that of basis. Pereopod 7 with all segments present.

One genus.

49. Anapronoe Stephensen, 1925

FIGURE 59

Type-species: Anapronoe reinhardti Stephensen, 1925.

With the characters of the family.

References: Stephensen, 1925:163; Spandl, 1927:217.

FIGURE 59.—Anapronoe.

XVIII. Family LYCAEIDAE Claus, 1879

Body rather compact, laterally compressed or somewhat flattened dorsoventrally; total length 3–22 mm. Head globular or somewhat flattened dorsoventrally; eyes large, occupying most of head surface. Coxae not fused with pereonites. Telson triangular, separate from or fused with double urosomite. Antenna 2 rudimentary or absent in female. Mandible with palp in male, without palp in female (except 2-segmented palp in female Tryphana). Maxillae 1 and 2 reduced but present. Pereopods 1 and 2 simple or subchelate. Pereopod 5 slightly or distinctly longer than pereopod 6; basis of both somewhat broadened but not operculate. Pereopod 6 with distal segments inserted on margin of basis or on medial surface proximal to margin. Pereopod 7 with all segments present. Uropod 3 having endopod sometimes fused with protopod. Gills on pereonites 2–6, with folds. Oostegites on pereonites 2–5.

Five genera.

References: Claus, 1887:55–65; Chevreux and Fage, 1925:426.

Key to the Genera of Lycaeidae

50. Lycaea Dana, 1852

FIGURE 60

Type-species: Lycaea ochracea Dana, 1853.

Body rather plump, especially in female. Telson fused with double urosomite. Antenna 2 of male with very short terminal segment. Pereopods 1 and 2 subchelate, margins of carpus and propus smooth or finely serrulate; basis of pereopod 1 not markedly broadened. Pereopod 5 distinctly longer than pereopod 6. Uropod 1 with protopod much longer than endopod. Uropod 3 slightly longer than telson; endopod not fused with protopod.

References: Claus, 1887:61; Barnard, 1930:428; Shoemaker, 1945:243.

FIGURE 60.—Lycaea.

51. Brachyscelus Bate, 1861

FIGURE 61

Type-species: Brachyscelus crusculum Bate, 1861.

Body compact, moderately slender. Telson not fused with double urosomite. Antenna 2 of male with short terminal segment. Pereopods 1 and 2 subchelate, margins of carpus and propus dentate; basis of pereopod 1 not markedly broadened. Pereopod 5 slightly longer than pereopod 6. Uropod 1 with protopod and endopod subequal in length. Uropod 3 slightly longer than telson; endopod not fused with protopod.

References: Claus, 1887:56; Stephensen, 1925:172.

FIGURE 61.—Brachyscelus.

52. Pseudolycaea Claus, 1879

FIGURE 62

Type-species: Pseudolycaea pachypoda Claus, 1879b.

Body moderately plump. Telson fused with double urosomite. Antenna 2 of male with very short terminal segment. Pereopods 1 and 2 simple; basis of pereopod 1 not markedly broadened. Pereopods 5 and 6 subequal in length. Uropod 1 with protopod much longer than endopod. Uropod 3 slightly longer than telson; endopod fused with protopod.

References: Claus, 1887:64; Chevreux and Fage, 1925:430; Spandl, 1927:215.

FIGURE 62.—Pseudolycaea.

53. Thamneus Bovallius, 1887

FIGURE 63

Type-species: Thamneus rostratus Bovallius, 1887b.

Body moderately plump, flattened dorsoventrally. Head somewhat flattened, with short, broad, upturned rostrum in male. Telson not fused with double urosomite. Antenna 2 of male much shorter than head, 4-segmented. Pereopods 1–6 covered with numerous setae on distal segments. Pereopods 1 and 2 subchelate, margins of carpus and propus dentate; basis of pereopod 1 not markedly broadened. Pereopods 5 and 6 subequal in length. Uropod 1 with protopod and endopod subequal in length. Uropod 3 distinctly longer than telson; endopod not fused with protopod.

Reference: Stebbing, 1888:1558.

FIGURE 63.—Thamneus.

54. Tryphana Boeck, 1870

FIGURE 64

Type-species: Tryphana malmi Boeck, 1870.

Body robust. Telson not fused with double urosomite. Antenna 2 of male with long terminal segment. Mandible of female with 2-segmented palp. Pereopods 1 and 2 simple; basis of pereopod 1 oval, nearly as broad as long; pereopod 2 with slender distal segments, propus and dactyl covered with short setae. Pereopods 3–6 stout, with prominent glands in carpus, distally prehensile with dactyl closing against produced distal margin of propus. Pereopod 5 distinctly longer than pereopod 6. Uropod 1 with protopod and endopod subequal in length. Uropod 3 much longer than telson; endopod not fused with protopod.

Reference: Sars, 1890:16.

FIGURE 64.—Tryphana.

XIX. Family OXYCEPHALIDAE Bate, 1861

Body form variable, sometimes compact, usually elongate, sometimes extremely long and slender; total length 3–120 mm. Head sometimes globular, usually produced anteriorly into long rostrum, sometimes with distinct neck; eyes large, occupying most of head surface except rostrum and neck. Coxae usually separate from pereonites, sometimes fused. Telson fused with or separate from double urosomite. Antenna 2 absent in female. Mandible with palp in male, without palp in female. Maxillae 1 and 2 absent. Pereopod 1 rarely simple, usually subchelate or chelate. Pereopod 2 subchelate or chelate. Pereopod 5 subequal to or distinctly longer than pereopod 6; basis of both rarely slender, usually broadened but not operculate. Pereopod 6 with distal segments inserted terminally or rarely subterminally on basis, their combined length greater than that of basis. Pereopod 7 usually with all segments present; distal segments sometimes reduced in number or absent. Uropods 2 and 3 having endopods sometimes fused with protopods. Gills on pereonites 2–6, sometimes rudimentary or absent on one or more of pereonites 2–4. Oostegites on pereonites 2–5, sometimes on pereonites 3–5, sometimes reduced in size.

Nine genera.

References: Bovallius, 1890; Fage, 1960; Pillai, 1966a.

Key to the Genera of Oxycephalidae

55. Oxycephalus H. Milne-Edwards, 1830

FIGURE 65

Type-species: Oxycephalus piscatoris H. Milne-Edwards, 1830.

Body elongate. Head somewhat shorter than pereon, produced into long, pointed rostrum. Coxae fused with pereonites. Double urosomite longer than wide, sometimes twice as long, fused with telson. Pereopods 1 and 2 chelate. Pereopod 7 with all segments present. Uropods 2 and 3 having endopods fused with protopods. Uropod 3 slightly shorter, equal to, or slightly longer than telson.

FIGURE 65.—Oxycephalus.

56. Calamorhynchus Streets, 1878

FIGURE 66

Type-species: Calamorhynchus pellucidus Streets, 1878.

Body elongate. Head longer than pereon, produced into long, pointed rostrum expanded into broad lateral flanges. Coxae fused with pereonites. Double urosomite more than 3 times as long as wide, fused with telson. Pereopods 1 and 2 chelate. Pereopod 7 with all segments present. Uropod 3 shorter than telson; endopod fused with protopod.

FIGURE 66.—Calamorhynchus.

57. Cranocephalus Bovallius, 1890

FIGURE 67

Type-species: Cranocephalus goesi Bovallius, 1890.

Body rather compact, only moderately elongate. Head somewhat shorter than pereon, produced into pointed rostrum. Coxae fused with pereonites but demarcated by indentations. Pereonite 5 produced into backward-pointed processes on either side. Double urosomite somewhat longer than wide, fused with telson. Pereopods 1 and 2 subchelate; pereopod 2 sometimes almost chelate. Pereopods 5–7 having broad basis, with row of 3 or 4 pitlike glands. Pereopod 7 with all segments present. Uropod 3 subequal to or shorter than telson.

FIGURE 67.—Cranocephalus.

58. Glossocephalus Bovallius, 1887

FIGURE 68

Type-species: Glossocephalus milne-edwardsi Bovallius, 1887b.

Body very slender, elongate. Head much shorter than pereon, produced into short rounded rostrum. Coxae separate on pereonites 1–6, fused with pereonite 7. Double urosomite about as long as wide, fused with telson. Pereopods 1 and 2 chelate. Pereopods 3 and 4 very slender. Pereopods 5 and 6 paddle-like, with broad segments; dactyl very small. Pereopod 7 with all segments present. Uropod 3 somewhat longer than telson. Gills on pereonite 2 absent in male, rudimentary in female; on pereonite 3 rudimentary in both sexes. Oostegites on pereonites 3–5.

FIGURE 68.—Glossocephalus.

59. Leptocotis Streets, 1877

FIGURE 69

Type-species: Leptocotis spinifera Streets, 1877.

Body elongate. Head longer than pereon, produced into long, pointed rostrum. Coxae not fused with pereonites. Double urosomite more than 3 times as long as wide, fused with telson. Pereopods 1 and 2 chelate. Pereopod 7 with all segments present. Uropod 1 having endopod very small. Uropod 2 having endopod fused with protopod in female, separate in male. Uropod 3 shorter than telson; endopod fused with protopod; exopod very small.

FIGURE 69.—Leptocotis.

60. Rhabdosoma White, 1847

FIGURE 70

Type-species: Oxycephalus armatus H. Milne-Edwards, 1840.

Body extremely slender and elongate. Head much longer than pereon, with long neck and very long, needle-shaped rostrum. Coxae fused with pereonites. Double urosomite more than 3 times as long as wide, not fused with telson. Pereopods 1 and 2 very short, chelate. Pereopods 3–6 very slender. Pereopod 7 very small, usually consisting only of basis, sometimes with one or more additional rudimentary segments. Uropods very slender; uropods 2 and 3 having endopod fused with protopod, exopod very short. Gills on pereonites 2–6 or 5 and 6 in female, on pereonites 4–6 or 5 and 6 in male. Oostegites reduced in size, on pereonites 2–6.

FIGURE 70.—Rhabdosoma.

61. Simorhynchotus Stebbing, 1888

FIGURE 71

Type-species: Simorhynchus antennarius Claus, 1871.

Body compact. Head much shorter than pereon, globular, with short, rounded rostrum in male. Coxae separate. Double urosomite slightly shorter than wide, fused with telson. Pereopod 1 simple. Pereopod 2 subchelate. Pereopod 7 with all segments present. Uropods 2 and 3 having endopod fused with protopod. Uropod 3 slightly longer than telson.

FIGURE 71.—Simorhynchotus.

62. Streetsia Stebbing, 1888

FIGURE 72

Type-species: Streetsia challengeri Stebbing, 1888.

Body elongate. Head variable in length, from somewhat shorter to distinctly longer than pereon, produced into pointed rostrum. Coxae not fused with pereonites. Double urosomite from slightly longer to 2.5 times as long as wide, fused with telson. Pereopod 1 subchelate; pereopod 2 subchelate or chelate. Pereopod 7 with all segments present. Uropod 3 shorter than telson.

FIGURE 72.—Streetsia.

63. Tullbergella Bovallius, 1887

FIGURE 73

Type-species: Tullbergella cuspidata Bovallius, 1887b.

Body rather compact, only moderately elongate. Head much shorter than pereon, produced into short, pointed rostrum. Coxae not fused with pereonites. Pleonite 3 produced into long, backward-pointing spines on either side. Double urosomite slightly shorter than wide, fused with telson. Pereopods 1 and 2 subchelate. Pereopod 6 with distal posterior corner of basis produced into triangular process. Pereopod 7 composed of basis and 3 small distal segments. Uropod 3 as long as telson; endopod fused with protopod.

FIGURE 73.—Tullbergella.

XX. Family PLATYSCELIDAE Bate, 1862

FIGURE 74

Body wide, stout, conglobate, usually somewhat depressed dorsoventrally. Pleon distinctly narrower than pereon and capable of folding under pereon. Total length 2–24 mm. Head short, wider than pereon, ventral margin (viewed from front) produced into median round-triangular process. Eyes large, occupying most of head surface. Coxae not fused with pereonites. Telson broadly triangular, fused with double urosomite. Antenna 2 of female short, of few segments. Mouthparts in the form of a broad, short cylinder. Mandible short, with broad incisor; with palp in male, wtihout palp in female. Maxillae 1 and 2 with large broad lobes.

Pereopods 1 and 2 simple, subchelate, or chelate. Pereopods 5 and 6 with bases transformed into broad opercula, always longer in pereopod 6; distal segments inserted subterminally in basis of pereopod 5, much farther proximally in pereopod 6; posterior margin of basis of pereopod 6 with longitudinal groove (telsonic groove) into which telson and uropods fit during conglobation; outer surface of basis of pereopod 6 often with fissure. Pereopod 7 with narrow, curved basis and sometimes one or more rudimentary additional segments. Uropods 2 and 3 having endopods sometimes fused with protopods. Gills on pereonites 2–6. Oostegites on pereonites 2–5.

Five genera.

FIGURE 74.—Platyscelidae.

Key to the Genera of Platyscelidae

64. Platyscelus Bate, 1862

FIGURE 75

Type-species: Typhis ovoides Risso, 1816.

Antenna 2 of male having 2 distal segments much shorter than half the length of preceding segment (third segment from end). Pereopods 1 and 2 chelate, with pointed carpal process. Pereopod 6 having basis with nearly straight fissure.

References: Claus, 1887:31; Stebbing, 1888:1480; Spandl, 1927:240; Shoemaker, 1945:255.

FIGURE 75.—Platyscelus.

65. Amphithyrus Claus, 1879

FIGURE 76

Type-species: Amphithyrus bispinosus Claus, 1879b (by present designation).

Cuticle with polygonal pattern of surface sculpturing. Antenna 2 of male with two distal segments subequal in length to preceding segment. Pereopods 1 and 2 complexly chelate; carpus with pointed process forming chela; propus with concave distal margin forming subchela. Pereopod 6 having basis relatively short and wide, with semicircular fissure.

References: Claus, 1887:41; Pirlot, 1930:43; Spandl, 1927:246.

66. Hemityphis Claus, 1879

FIGURE 77

Type-species: Hemityphis tenuimanus Claus, 1879b.

Antenna 2 of male having two distal segments more than half the length of preceding segment. Pereopods 1 and 2 chelate, with pointed carpal process. Pereopod 6 having basis with short, curved fissure.

References: Claus, 1887:38; Stebbing, 1888:1471; Spandl, 1927:233; Pirlot, 1930:37; Shoemaker, 1945:259.

67. Paratyphis Claus, 1879

FIGURE 78

Type-species: Paratyphis maculatus Claus, 1879b.

Antenna 2 of male having 2 distal segments slightly less than half or somewhat more than half the length of preceding segment. Pereopod 1 simple. Pereopod 2 weakly chelate, with short carpal process. Pereopod 6 having basis with nearly straight fissure.

FIGURE 76.—Amphithyrus.

FIGURE 77.—Hemityphis.

FIGURE 78.—Paratyphis.

References: Claus, 1887:39; Stebbing, 1888:1476; Spandl, 1927:243; Pirlot, 1930:39.

68. Tetrathyrus Claus, 1879

FIGURE 79

Type-species: Tetrathyrus forcipatus Claus, 1879b.

Antenna 2 of male with two distal segments more than half the length of preceding segment. Pereopods 1 and 2 subchelate; dactyl closing against concave distal margin of propus. Pereopod 6 having basis with long ventral margin, without fissure.

References: Claus, 1887:40; Stebbing, 1888:1480; Spandl, 1927:240; Barnard, 1930:439.

FIGURE 79.—Tetrathyrus.

XXI. Family PARASCELIDAE Bovallius, 1887

Body wide, stout, conglobate, usually somewhat depressed dorsoventrally. Pleon distinctly narrower than pereon and capable of folding under pereon. Total length 2.5–7 mm. Head short, wider than pereon, ventral margin (viewed from front) broadly rounded. Eyes large, occupying most of head surface. Coxae not fused with pereonites. Telson broadly triangular, fused with double urosomite. Antenna 2 of female short, of few segments. Mouthparts pointed, in the form of a cone. Mandible elongate, with narrow incisor; with palp in male, without palp in female. Pereopods 1 and 2 simple or chelate. Pereopods 5 and 6 with basis transformed into broad operculum, always longer in pereopod 6; distal segments inserted subterminally on basis of pereopod 5, much farther proximally in pereopod 6; posterior margin of pereopod 6 with longitudinal groove (telsonic groove) into which telson and uropods fit during conglobation; outer surface of pereopod 6 with or without fissure. Pereopod 7 with all segments present. Uropods 2 and 3 having endopods sometimes fused with protopods; exopods much shorter than endopods. Gills on pereonites 2–6. Oostegites on pereonites 2–5.

Three genera.

References: Claus, 1887:42; Stebbing, 1888:1491; Spandl, 1927:250.

Key to the Genera of Parascelidae

69. Thyropus Dana, 1852

FIGURE 80

Synonym: Parascelus Claus, 1879.

Type-species: Thyropus diaphanus Dana, 1852.

Pereopods 1 and 2 simple. Pereopod 6 having basis very long, narrowed in distal half, with or without short fissure. Uropods 1–3 having exopods and endopods not fused with protopod.

References: Claus, 1887:45; Stebbing, 1888:1492; Stephensen, 1925:208; Spandl, 1927:258; Shoemaker, 1945:260.

70. Euscelus Claus, 1879

FIGURE 81

Type-species: Euscelus robustus Claus, 1879b.

Pereopods 1 and 2 chelate; carpal process narrow, curved, with truncate and dentate distal margin. Pereopod 6 having basis without fissure. Uropods 1–3 with exopods oval, much shorter than endopods; exopods and endopods not fused with protopods.

References: Claus, 1887:43; Spandl, 1927:251.

FIGURE 80.—Thyropus.

FIGURE 81.—Euscelus.

71. Schizoscelus Claus, 1879

FIGURE 82

Type-species: Schizoscelus ornatus Claus, 1879b.

Pereopod 1 simple. Pereopod 2 chelate, with pointed carpal process. Pereopod 6 having basis subrectangular, with long, slightly curved fissure. Uropods 2 and 3 having endopods fused with protopods.

References: Claus, 1887:43; Stebbing, 1888:1503; Spandl, 1927:225.

FIGURE 82.—Schizoscelus.

Sources of Illustrations

Figure 1: Original.

Figure 2: Original.

Figure 3: Md, original. Md′, Pirlot, 1930, fig. 3. Mx.1, Mx.2, original. Mxp, Stephensen and Pirlot, 1931, fig. 17. Mxp′, original.

Figure 4: Original.

Figure 5: Original.

Figure 6: Original.

Figure 7: Original.

Figure 8: Shoemaker, 1945, fig. 22J.

Figure 9: A, original. B, Fage, 1960, fig. 8.

Figure 10: ♂, Woltereck, 1909, pl. 5, fig. 13; ♀, fig. 12. A.1, A.2, Md, P.1, P.2, Vinogradov, 1956, fig. 2. Us, Stephensen and Pirlot, 1931, fig. 13.

Figure 11: ♂, Woltereck, 1909, pl. 3, fig. 8. ♀, Stephensen and Pirlot, 1931, fig. 7; Md, Mxp, fig. 8; P.1, P.2, P.7, Us, fig. 9.

Figure 12: Whole animal, Vinogradov, 1960a, fig. 12; Md. P.7, fig. 13.

Figure 13: A.1, Vinogradov, 1956, fig. 4. A.2, Md. P.1, P.2.Stephensen and Pirlot, 1931, fig. 17; Us. fig. 18.

Figure 14: ♀, Pirlot, 1933, fig. 1. A.1, A.2, Vinogradov, 1960a, fig. 14. Md, Mxp, Pirlot, 1933, fig. 2. P.1, P.2, P.7, Vinogradov, 1960a, fig. 14. Us, Pirlot, 1933, fig. 3.

Figure 15: ♂, Chevreux and Fage, 1925, fig. 387. A.2, P.1, P.5, Scina borealis, off California, original. Md, Mxp, Scina curvidaotyla, off Washington, original. Us, Scina tullbergi, off California, original.

Figure 16: ♀, Vosseler, 1901, pl. 10, fig. 1. A.1, Wagler, 1926, fig. 51b; Mxp, fig. 52d; P.1, P.5, fig. 52a.

Figure 17: ♀, Wagler, 1926, fig. 53; H (ventral), fig. 54; Mxp, fig. 56d; P.1, P.5, fig. 56a. P.5 (dactyl, enlarged), original.

Figure 18: ♀, Mxp, Vinogradov, 1957, fig. 18. P.1, P.5, Chevreux, 1914, fig. 4.

Figure 19: Whole animal, Woltereck, 1909, pl. 4, fig. 11. A.1, A.2, P.1, P.3, P.5, P.6, Us, Stephensen and Pirlot, 1931, fig. 16.

Figure 20: Whole animal, Vinogradov, 1960a, fig. 9; A.1, A.2, Md, Mxp, Us, fig. 10; P.1, P.3, P.5, P.6, fig. 11.

Figure 21: Whole animal, Woltereck, 1909, fig. 10. Head, P.1, P.2, Shoemaker, 1945, fig. 29; P.5, P.7 (left), fig. 30. P.5, P.7 (right), Vinogradov, 1964, fig. 10.

Figure 22: ♀, Pirlot, 1930, fig. 1; A.1, A.2, Md, Mxp, fig. 3; P.1, P.2, P.3, P.7, fig. 4; Us, fig. 5. H (dorsal), Vinogradov, 1960b, fig. 2.

Figure 23: ♂, Stephensen, 1918, fig. 4. H, Md, Mxp, Shoemaker, 1945, fig. 15; Mx.1, fig. 20; P.1, P.2, fig. 19. Md incisor and lacinia, Lanceola sayana, off New Jersey, original.

Figure 24: ♀, original. A.1, A.2, Pirlot, 1935, fig. 3; Md, Mx.1, fig. 1. P.1. P.2, Shoemaker, 1945, fig. 21.

Figure 25: ♂, Pirlot, 1931, fig. 1; Md, Mxp, P.3, fig. 2. A.1, A.2, Mx.1, P.1, P.2, Vinogradov, 1960a, fig. 6.

Figure 26: Barnard, 1930, fig. 52.

Figure 27: Whole animal, Woltereck, 1909, fig. 14. A.1, A.2, Md, Mx.1, vinogradov, 1957, fig. 7; P.1, P.2, P.5, fig. 8.

Figure 28: ♂, Woltereck, 1909, fig. 21. H, original. Md, Mx.1, Mxp, P.1, P.2, Shoemaker, 1945, fig. 22.

Figure 29: ♀, Vosseler, 1901, pl. 11, fig. 6. Md, Mxp, Us, Vibilia viatrix, off California, original. P.7, V. cultripes, off California, original.

Figure 30: ♀, Hurley, 1955, fig. 23. P.7, Us, Cyllopus magellanicus, off New Zealand, original.

Figure 31: Original.

Figure 32: ♀, Spandl, 1927, fig. 8. Md, Mxp, P.1, P.2, original. Us, Stephensen, 1918, fig. 21.

Figure 33: ♂, Vosseler, 1901, pl. 8, fig. 22. Md. Mxp, P.1, P.2, Us, Paraphronima gracilis, off California, original.

Figure 34: ♀, A.1, A.2, Bowman, 1973, fig. 18a–c; Md, Us, fig. 2h,o; Mxp, fig. 15e,f; P.1, P.2, fig. 4a,b.

Figure 35: Laval, 1966.

Figure 36: ♀, Bowman, 1973, fig. 22a; Mxp, fig. 20i; P.1, P.2, fig. 23a,b; Us, fig. 21a.

Figure 37: ♀, Vosseler, 1901 pl. 5, fig. 16. A.1, A.2, Mxp, P.3, Bowman, 1973, fig. 46d,e,i,j,m; Md, P.1, P.2, fig. 43d,i,j; P.6, fig. 41o; Us, fig. 41q.

Figure 38: ♀, Vosseler, 1901, pl. 7, fig. 6. Md, original. Mxp, Bowman, 1973, fig. 24j and original. P.1, P.2, Bowman, 1973, fig. 24k,l. Us, original.

Figure 39: ♀, P.1, P.2, P.5, Yang, 1960, fig. 9. Mx.1, Mxp, P.5 dactyl, Bowman, 1973, fig. 52b,d–f. Us, Stephensen, 1924, fig. 35.

Figure 40: ♀, Md, Hyperoche martinizii, Gulf of Guinea, original. Mx.1, Mxp, H. medusarum, Antarctic, original. P.1. P.2, P.3, H medusarum, off California, original.

Figure 41: ♀, Bovallius, 1889, pl. 8, fig. 1. Mxp, P.1, P.2, Iulopis mirabilis, off California, original.

Figure 42. ♀, P.1, P.2, Bowman, 1973, fig. 32a,o,p; A.2, fig. 37j; Md, Mxp, fig. 36c,f. A.1, original.

Figure 43: ♀, ♂, Chevreux and Fage, 1925, figs. 407, 408. Md, Bowman, 1960, fig. 1; P.1, P.4, P.5, fig. 16; P.2, fig. 2.

Figure 44: ♀, original. A.1, P.1, P.2, Barnard, 1932, fig. 162b,d,e; Md, Mxp, fig. 163a,f; Us, fig. 164c.

Figure 45: ♀, Chevreux and Fage, 1925, fig. 406. ♂, Us, Bovallius, 1889, pl. 14. Md, Mxp, P.1, P.2, P.6, original.

Figure 46: Bowman, 1973, fig. 51a,e,f,g,j,k,p,r,s.

Figure 47: ♂J, H ♀, Bovallius, 1889, pl. 15. Md, Mx.1, Mx.2, Mxp, P.1, P.2, Us, original.

Figure 48: ♀, ♂, Vosseler, 1901, pl. 2. Md, Mxp, Us, Phronima sedentaria, original.

Figure 49: ♀, P.5, Shih and Dunbar, 1963, fig. 9. Us ♂, Us ♀, original.

Figure 50: ♀, Bovallius, 1889, pl. 18. fig. 3; P.3, fig. 6; P.4, fig. 7; P.5, fig. 8; P.6, fig. 9; P.7, fig. 10 (distal segment added); Us, fig. 11.

Figure 51: ♀, Stebbing, 1888, pl. 177. Md, Mxp. P.1, P.2, P.3, P.5, P.6, P.7, Us, original.

Figure 52: Original.

Figure 53: ♂, Stebbing, 1888, pl. 181. A.1, P.1, P.2, P.5, P.6, P.7, Us ♀, Pirlot, 1930, fig. 9; Us ♂, fig. 9 (but Up.3 added from Stebbing, 1888, pl. 180).

Figure 54: ♂, A.2, Stebbing, 1888, pl. 186. P.1, P.2, P.7, Us, original.

Figure 55: ♂, Claus, 1887, pl. 13, fig. 7. A.2, P.1 ♂, P.1 ♀, P.2, P.7, original. Us, Stebbing. 1888, pl. 187.

Figure 56: ♀, P.1, P.2, P.7, Us, original. A.2♂, Claus, 1887, pl. 20, fig. 2.

Figure 57: ♀, Stebbing, 1888, pl. 191. A.2 ♂, P.1, P.2, P.6, P.7, Us, original.

Figure 58: ♂, Stebbing, 1888, pl. 192. A.2, P.1, P.2, P.6, P.7, Us, original.

Figure 59: ♀, P.1, P.2, P.6, P.7, Anapronoe sp., off California, original. A.1♀, A.2♀, A.2♂, Md, Mx.1, Us, A. reinhardti, Gulf of Guinea, original.

Figure 60: ♀, Us, Shoemaker, 1945, fig. 38; P.1, P.2, fig. 39. A.2 ♂, P.1 ♂, original.

Figure 61: ♂, Stebbing. 1888, pl. 195. A.2, P.1, P.2, Us, original.

Figure 62: ♂, P.1, P.2, Chevreux and Fage, 1925, fig. 420. P.7, Us, Stephensen, 1925, fig. 64. A.2 ♂, original.

Figure 63: ♂, ♂ H, dorsal, A.2 ♂ original. P.1, P.6, Pillai, 1966b, fig. 16. Us, Stebbing, 1888, pl. 198.

Figure 64: ♂, A.2 ♂, Stebbing, 1888, pl. 194. P.1, P.2, P.5, Us, original.

Figure 65: ♀, Fage, 1960, fig. 5a. P.1, P.2. P.7, Us, original.

Figure 66: ♀, Pillai, 1966a, pl. 1; P.1, P.2, P.7, fig. 8. Us, Fage, 1960, fig. 19.

Figure 67: ♀, Shoemaker, 1945, fig. 44A. US ♂, Us ♀. Fage, 1960, fig. 51. P.1, P.2, original.

Figure 68: ♀, P.1, P.2, Up.3, Us, Shoemaker, 1945, fig. 45.

Figure 69: ♂, Pillai, 1966a, pl. 1; P.7, fig. 7. Up.2 ♀, Fage, 1960, fig. 22; Us ♂, fig. 21. P.1, P.2, original.

Figure 70: ♀, Bovallius, 1890, pl. 7, fig. 20. P.1, P.2, Pillai, 1966a, fig. 15; P.7, fig. 16. Us, Fage, 1960, fig. 63.

Figure 71: ♂, Pillai, 1966a, pl. 1. P.1, P.2, P.7, Us, original.

Figure 72: ♀, Pillai, 1966a, pl. 1; Us, fig. 12. P.1, P.2, P.7, original.

Figure 73: ♀, Pillai, 1966a, pl. 1; P.1, P.2, P.6, P.7, fig. 6. Us, original.

Figure 74: Platyscelus ovoides, original.

Figure 75: A.2 ♂, Claus, 1887, pl. 2, fig. 6. P.1, P.2, P.6, Spandl, 1927, fig. 45.

Figure 76: Amphithyrus bispinosus. Head and pereon, dorsal, original. A.2, P.1, P.2, P.6, Claus, 1887, pl. 6.

Figure 77: A.2 ♂, Claus, 1887, pl. 4. P.1, P.2, P.6, Spandl, 1927, fig. 47.

Figure 78: A.2 ♂, Claus, 1887, pl. 5; P.1, P.2, P.6, pl. 7.

Figure 79: A.2 ♂, Claus, 1887, pl. 5. P.1, P.2, P.6, Spandl, 1927, fig. 48.

Figure 80: ♂, A.2 ♂, P.7, Claus, 1887, pl. 8. P.1, P.2, P.6, Us, Spandl, 1927, fig. 53. Mouth cone (labrum, Md, Mxp), original.

Figure 81: ♂, Spandl, 1927, fig. 51.

Figure 82. Spandl, 1927, fig. 52.

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In synonymy, use the short form (taxon, author, date:page) with a full reference at the end of the paper under “Literature Cited.” Begin each taxon at the left margin with subsequent lines indented about three spaces. Within an entry, use a period-dash (.—) to separate each reference. Enclose with square brackets any annotation in, or at the end of, the entry. For references within the text, use the author-date system: “(Jones, 1910)” and “Jones (1910).” If the reference is expanded, abbreviate the data: “Jones (1910:122, pl. 20: fig. 1).”

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they should not contain rules. Formal tables must be submitted as pages separate from the text, and each table, no matter how large, should be pasted up as a single sheet of copy.

Use the metric system instead of, or in addition to, the English system.

Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed throughout the printed text. They will be termed Figures and should be numbered consecutively; however, if a group of figures is treated as a single figure, the components should be indicated by lowercase italic letters on the illustration, in the legend, and in text references: “Figure 9b.” If illustrations (usually tone photographs) are printed separately from the text as full pages on a different stock of paper, they will be termed Plates, and individual components should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the numbering system of text illustrations with that of plate illustrations. Submit all legends on pages separate from the text and not attached to the artwork. An instruction booklet for the preparation of illustrations is available from the Press on request.

In the bibliography (usually called “Literature Cited”), spell out book, journal, and article titles, using initial caps with all words except minor terms such as “and, of, the.” For capitalization of titles in foreign languages, follow the national practice of each language. Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume, number, and page citations: “10(2):5-9.” Spell out such words as “figures,” “plates,” “pages.”

For free copies of his own paper, a Smithsonian author should indicate his requirements on “Form 36” (submitted to the Press with the manuscript). A non-Smithsonian author will receive 50 free copies; order forms for quantities above this amount with instructions for payment will be supplied when page proof is forwarded.