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SMITHSONIAN INSTITUTION
MUSEUM OF NATURAL HISTORY
a, Uschakoviella echinophora Gurjanova [after Shoemaker, 1964].
b, Stilipes distincta Holmes [after Shoemaker, 1964].
UNITED STATES NATIONAL MUSEUM BULLETIN 271
The Families and Genera of Marine Gammaridean
Amphipoda
J. LAURENS BARNARD
Associate Curator of Crustacea
SMITHSONIAN INSTITUTION PRESS CITY OF WASHINGTON 1969
Publications of the United States National Museum
The scientific publications of the United States National Museum include two series, Proceedings of the United States National Museum and United States National Museum Bulletin.
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This work forms number 271 of the Bulletin series.
FraNK A. TAYLOR Director, United Staies National Museum
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Contents
Introduction .
Acknowledgments P Status of gammaridean ayctementies i Morphological terminology .
Morphology of a gammaridean aapEinod Geographic distribution of marine Gammaridea . The composition of a faunule ig Morphological evolution of the Asnatnipyades Interfamilial gammaridean relationships . Limnetic evolution of the Gammaridea Identification procedures
Diagnosis of the basic marine ganmmaridean (Gene a marine Carmmeridson
Order Amphipoda : Key to the suborders of Avaninporal ; i Nomenclatural changes in gammaridean orenfTas : Key to the families of the suborder Gammaridea . Acanthonotozomatidae Ampeliscidae . Amphilochidae . Ampithoidae . Anamixidae Aoridae .
Argissidae . Astyridae
Atylidae . Bateidae. Beaudettiidae Calliopiidae Cheluridae Colomastigidae . Corophiidae . Cressidae Dexaminidae . Dogielinotidae . Eophliantidae Eusiridae Gammaridae (Goesaing) . Haustoriidae . A. Hyalellidae and Efvalidiae Hyperiopsidae . Isaeidae . Ischyroceridae . Kuriidae
al CONTENTS
Lafystiidae Laphystiopsidae Lepechinellidae . Leucothoidae Liljeborgiidae Lysianassidae Melphidippidae Ochlesidae . Oedicerotidae Pagetinidae Paramphithoidae . Pardaliscidae Phliantidae Geueed) Phoxocephalidae Pleustidae . Podoceridae 4 Prophliantidae evincaye Sebidae . ‘ Siecocepralrdne: Stenothoidae . Stilipedidae Synopiidae (revised)
Superfamily Talitroidea (@auanda Be atelndae. iTalitnidae)
Thaumatelsonidae Vitjazianidae Incertae sedis Supplement Literature cited ae Appendix I. Dissection of an i anpaipodi Appendix II. Amphipod analytical sheet . Appendix III. Glossary of special terms ‘ Index and familial assignment of gammaridean penera :
Page 281 283 286 289 291 294 368 371 373 387 389 397 405 412 421 426 432 435 436 444 451 454 463 473 476 478 481 482 507 512 520 524
The Families and Genera of Marine Gammaridean
Amphipoda
Introduction
Keys and diagnoses, with illustrations, to the families and genera of marine gammaridean Amphipoda are presented here in the form of a handbook. Since 1906, when Stebbing monographed this suborder, the number of families has been increased by 25 percent, the number of genera by 200 percent and the number of species by nearly 200 percent (J. L. Barnard, 1959d). Stebbing’s general arrangement of families and genera remains substantially intact, but the discovery of many additional species has required a considerable emendation of familial and generic limits. The near absence of phyletic monographs and the plethora of faunal compilations widely scattered in the litera- ture have hindered advances in classification of the group. Ready identification of taxa not only by nonspecialists but even by experts has become extremely difficult because of the lack of a modern focal point.
The present work is at best a stopgap. Specialists recognize that much exploration and discovery remain to be accomplished in the Amphipoda (witness the numerous new haustoriid taxa discovered by Bousfield, 1965). No specialist is nearly satisfied with current classifi- cation at familial levels. Many genera are extremely confused, not only for the lack of clear description of their type-species but for the obviously erroneous inclusion of many of their congeners. Few studies on population variability have been published and the criteria to be applied at generic and familial evels have rarely been tested even by dialectical means.
No radical changes in classification are proposed herein, but numerous problems are noted. The purpose of this paper is to assemble the families and genera within the previously arranged general scheme and to provide rapidly usable (and thus simplified) means for their identification. In this way the keys coincide with the published literature and may be of greater help than would be a brief, completely revised system, which might not stand the test of time. Hopefully this paper will encourage recruitment of new taxonomists and relieve some of the burdens of identification now overwhelming the few active
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specialists. Perhaps it will stimulate monographic studies of families and genera. It may be of greatest use to those students who must make generic determinations of their material without adequate library facilities; it should restrict bibliographic needs to small numbers of publications which can be borrowed more easily than can the entire literature. An index to the species of Gammaridea (J. L. Barnard, 1958a) may be of some help. An expanded and more useful index is now in preparation.
Brief sections on distribution, faunules, and evolution are presented in hopes of stimulating further inquiry into these fascinating but poorly studied topics. Our knowledge of anatomy (especially musculature), functional morphology, behaviour, food habits, indeed general ecology is rudimentary.
The identification system proposed herein requires familiarity with a basic gammaridean plan similar to that exemplified by some members of the Gammaridae. These gammarids may be the most primitive grade of living Amphipoda. A knowledge of their structure is vital to an understanding of the unfolding of the complex network of speciali- zations found in other gammaridean families. The lines of evolution from gammarid-like ancestors, so ill understood, are not only manifold but they confuse identification procedures. At times the specialist is confronted with the feeling that most of the ‘missing links” in Amphipoda are still alive. Only a few families are so remote morpho- logically from the Gammaridae that they are instantly recognizable. With a few exceptions the specialization of gammaridean families is a result of the loss of morphological complexity.
The basic gammaridean is illustrated in figures 1 and 2 and described on p. 50. Families other than the Gammaridae are provided with short diagnoses which include only definitive and exclusive differences from the basic gammaridean. Some of these diagnoses are identical among several families. Those taxa are further distinguished by intercomparison of their descriptions and by reference to paragraphs on relationship. Part of each diagnosis is a reference to a related (or superficially similar) family.
Diagnostic characters of the families are illustrated in a group of boxes (figs. 3-53). Instructions for identification procedures using these tools are presented in following pages.
Acknowledgments
This work was compiled during the writer’s tenures at the Allan Hancock Foundation, the Beaudette Foundation of California, and the Smithsonian Institution, to which institutions and their adminis- trative heads my gratitude is expressed. Mrs. Dorothy M. Halmos of
MARINE GAMMARIDEAN AMPHIPODA 3
the Hancock Library and Mr. Jack Marquardt of the Smithsonian Institution have been very helpful in obtaining references. Literature published subsequent to December 1965 has not been incorporated. The figures were redrawn largely from the literature and some of them have been simplified as a means of providing standardization. Four illustrators have assisted with the work over the years, Mr. L. R. Hales, Mrs. D. McLaughlin and, especially, Miss Jacqueline M. Hampton of the Beaudette Foundation; and Miss Naomi D. Mano- witz of the Smithsonian Institution. They were partially supported by grants from the National Science Foundation.
Dr. D. E. Hurley of New Zealand Oceanographic Institute has assisted my thinking with regard to the difficult family Lysianassidae by letting me read his various manuscripts concerning the classifica- tion of that family. Dr. E. L. Bousfield of the National Museum of Canada kindly loaned the paratype of Oldevig’s Haustorioides mun- sternjelmi for this study. Numerous colleagues have helped with friendly advice and encouragement and among those must be men- tioned the foregoing plus Dr. Fenner A. Chace, Jr., Dr. Thomas E. Bowman, and Dr. Olga Hartman. Dr. Richard Cowan, Dr. Donald Squires, Dr. I. E. Wallen, and Dr. Raymond B. Manning have created the ideal environment at the Smithsonian Institution for this work to be brought to its conclusion. Portions of the final draft were com- posed while the writer worked as a Smithsonian fellow to Bishop Museum, Honolulu; and I must thank Dr. Roland W. Force, Director, for providing the facilities and help of that Museum.
This paper is dedicated to Mr. Clarence R. Shoemaker (1876-1958), America’s foremost student of Amphipoda. The frontispiece represents two of the beautiful drawings he did.
Status of Gammaridean Systematics
More than 3300 species in about 670 genera are known currently in the Gammaridea.* The rate of description of new species has increased recently and will presumably exceed the average descrip- tion of 40 per year that occurred between 1906 and 1956. Outside of the littoral north Atlantic, Arctic U.S.S.R., and the northwestern Pacific, knowledge is highly incomplete. The magnificent work of Sars (1895), supported by that of Chevreux and Fage (1925), brought the west European fauna into definition at an early stage. Unfor- tunately, the excellent taxonomic status of north Atlantic amphipods
*About 130 genera and 850 species occur in nonmarine environments and most are not included in this handbook.
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has not been exploited fully in making advances in ecological, physio- logical, and genetical studies, although most of those special studies so far undertaken have been pursued in the northeastern Atlantic.
Intensive work has been accomplished in antarctic seas but the fauna there needs to be placed on a Sarsian basis with full illustra- tions and descriptions. The works of Chevreux (1906c, 1912b) are notable for that region, but numerous sticky problems resulting from other studies have not been clarified there.
The remaining faunas are sketchily known. An imbalanced em- phasis on the deep-sea is occurring in this decade, but this has helped bring into focus many problems in the higher taxa of amphipods. Since World War II research on the marine species has been meager, because more than 90 percent of the printed pages have been pro- duced by fewer than 10 taxonomists (Zoological Record, years 1953-60).
Morphological Terminology
The taxonomic assessment of gammaridean morphology is almost always stated in subjective terms: in words and phrases which, through experience, taxonomists have come to understand and visualize easily, but which are difficult to convey with mensurative | precision. There is no current solution to this difficulty but taxono- — mists may eventually be able to compile an illustrated handbook in which all possible shapes and relative proportions can be figured, named, and coded. Identifications may then be amenable to mechan- ical analysis. Meanwhile one must work with terms such as “strong, large, small, feeble, weak, minute, elongate, shortened” and various adjectives with the modifier ‘sub,’ such as “subquadrate”’ and “Subacute”. I have used these terms everywhere in this handbook, but have attempted to provide illustrations as a demonstration of their extent, for the degree varies in the different families. For in- stance, gnathopods of the genus Apherusa are ‘feeble’ in com- parison to those of its familial congener Calhiopius, but they are scarcely as feeble as the gnathopods of the Acanthonotozomatidae. The term ‘subacute’ apparently has come to mean a shape that has the overall appearance of sharpness but which terminally is softly rounded. The term ‘‘subconical’”’ apparently refers to a 2- dimensional, rather than a geometric cone but the adjective is not necessarily synonymous with “subacute,” for a subconical process may have a subacute apex. The term ‘“quadrate”’ has often been applied to a squared-off shape of an otherwise imperfect rectangle or in simpler fashion as reference to one pair of sides occurring at right angles to each other. ‘SSubquadrate”’ should refer to sides not
MARINE GAMMARIDEAN AMPHIPODA 5
precisely at right angles to each other but also seems to have reference to a quadrate shape with softly rounded corners.
The terms ‘‘vestigial’” and “rudimentary” have their uses, for a mathematical definition in each case might be more involved, con- fusing, and time consuming than is the process of learning these conditions through trial and error.
A decision as to whether a gnathopod is simple or subchelate is occasionally difficult to make and definitions to classify all border- line cases are almost impossible to compose. There are no terms to describe all of the stages between fully subchelate and fully simple and gammaridean students may eventually encounter difficulties arising from dogmatic use of such all-or-none terms. For instance, two subfamilies of Lysianassidae have been described which are partially based on a subchelate or simple gnathopod 1, but there are several borderline cases and several genera in one subfamily which seem to have stronger affinities with genera in the other subfamily than with members of their own group. The contrast between the concepts of cleft telson and entire telson is far too strong to describe all intermediate stages and the two terms tend to condition the human observer to a reliance on the alternatives as highly significant, whereas numerous exceptions to this significance are apparent [e.g., Kusiridae and Calliopiidae].
Names for the gammaridean abdomen and its parts are not stand- ardized. All six segments of the abdomen are frequently called the “pleon’’, but “pleonites’’ 4-6 are often called ‘“urosome” and num- bered as “urosomites’ 1-3. Pleonites 1-3 are rarely called the “‘metasome.”’
Shape and proportion in seemingly infinite array are thus more important to the taxonomist than are qualitative and numerical expressions, but one day we may be able to convert shape and pro- portion to precise formulations.
Morphology of a Gammaridean Amphipod
Amphipods, lke tanaids and isopods, lack a carapace covering the thorax, so that seven definitive thoracic segments (pereonites) are visible (as in other Peracarida the first thoracic segment with its appendage [maxilliped] has become fused to the head). Gammaridean and hyperiidean Amphipoda may be recognized by their possession of three pairs of pleopods (swimmerets) and two or three pairs of uropods on the pleon (abdomen). The consistent presence of at least six pairs of thoracic appendages, five-plus pairs of gills and four pairs of brood lamellae in females are definitive characters of Gammaridea and Hyperiidea.
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The pereon (thorax) almost invariably bears seven pairs of legs. The first two pairs are called gnathopods (or gamopods) and usually are prehensile, having the seventh article (dactyl) folded back on the sixth article (propodus, hand, or palmar article). Rarely in gnathopod 1 is there sexual dimorphism but the male often has greatly enlarged second gnathopods. Gnathopod 2 is believed to be used primarily for erasping the female during copulatory amplexus. The male mounts the dorsal side of the female, projects the gnathopods around her body and hooks them into her fifth coxae. The pair of animals then swims, darts among algae, or rests until the female molts (possibly as much as three days after amplexus), at which time the male emits spermatophores (sacs of spermatozoa) that pass from the ventral side of his seventh pereon segment into the pouch formed by the female brood lamellae. Ecdysis of the female is taking place during this time period and im- mediately thereafter she lays eggs through two genital pores on the sternite of thoracic segment 5. The pores normally are so small and heavily chitinized that the eggs cannot be laid until ecdysis occurs and the pore openings become soft and pliable. How the spermatophores pass from male to female is unknown but they may be assisted by gnathopods or pereopods. The ventral side of pereonite 7 in the male bears two minute penial projections, often spinose. Occasionally they are hidden by small gills attached to the coxae of this segment.
Within a genus of gammaridean amphipods, the taxonomic recogni- tion of species often depends on the shape of male gnathopod 2; hence, it is difficult to identify females specifically because taxonomists have not studied minute differences in females and made them basic to identification. The second gnathopods of juveniles and females often are alike; during maturation the male second gnathopods commence an an increase in size and a morphological differentiation, with changes taking place during each instar, even long after the attainment of sexual maturity. This has resulted occasionally in taxonomic confusion because some of these instars have been described as distinct species. A few species are known to have radically distinct phenotypes espe- cially in the terminal male. The ubiquitous Jassa falcata has dozens of forms, some of these even bridging the concepts of two or more genera.
All thoracic appendages have seven articles (segments), the proximal member of which is the coxa or sideplate. A few gammarideans (e.g., Bateidae) have reduced numbers of articles on various thoracic appendages. Coxae are of greater taxonomic importance in gammar- idean Amphipoda than in other Malacostraca and in many species are structurally more an integral part of the trunk than simply an article of the appendage. They resemble ventral pleuron-like extensions of segments and so contribute to the appearance of lateral compression in the body plan. They are numbered from one to seven, with numbers
MARINE GAMMARIDEAN AMPHIPODA 7
1 and 2 belonging with gnathopods 1-2 and numbers 3-7 with pereopods 1-5.
Gills are thoracic and generally are attached to the medial surfaces of coxae 2-7, occasionally only on 2-6, or ?3-6. In females the medial surfaces of coxae 2—5 (?or 2-6) carry brood lamellae. They are simply buds in young females, but as body growth proceeds they become longer and more heavily setose and are interlocked by their setae to form a cradle enclosing the eggs. As the female increases in age and size, the number of eggs laid after each molt becomes larger. Some amphipods lay such large yolky eggs that a young female can carry only one of them; at maturity she may be able to carry three or four. Not all of the extruded eggs hatch; mortality of about 25-50 percent may occur even before hatching; stunted degenerating eggs often may be seen in the broods (as well as commensals such as spherical copepods and reniform ostracods). Some large Amphipoda lay and carry more than 200 eggs at a time.
Commencing with leg 3, the thoracic appendages are termed pereo- pods (walking legs), so that five pairs of thoracic legs represent pereopods (many specialists commence numbering pereopods with enathopod 1). There is justification in distinguishing gnathopods by name because of their specialization in analogy to the pereopods of other Crustacea (e.g., posterior maxillipeds of decapods). The first two pairs of pereopods are useful in cleaning the gnathopods and other anterior appendages and as a balance when alighting from a swim. The last three pairs appear rather immobile and less adapted for walking than in isopods. Amphipods are poorly balanced for walking, hence their mobility usually depends on swimming. Their body plan, however, permits a motility through dense masses of hydroids and algae. Perhaps this is one of the reasons why amphipods are one of the most abundant macroscopic crustacean groups in algae and other anastomoses.
Some Amphipoda living on the sea-bottom have immensely elon- gated pereopods, which are spread out in the fashion of a spider and prevent the amphipod from sinking into the mud. To lower the center of gravity the body of the amphipod hangs upside down in its cradle of legs.
A few families of Gammaridea have the habit of burrowing into benthic sediments and their pereopods are armed densely with strong spines which aid in burrowing. Elongate flexible setae, as well as stiff spine-like setae may occur on pereopods 3-5 of fossorial Amphipoda. Even though a few other nonfossorial Amphipoda, such as Ampelis- cidae, have these elongate setae, the term “fossorial pereopods’’ is usefully applied to the condition. The glossary contains a precise definition of fossorial pereopods.
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Six or seven families of amphipods construct dwelling tubes. The cylindrical or flattened tubes are spun from strands of material secreted and probably manipulated by pereopods 1-2. The tubes may be limp (Ampelisca) and lay prone on the bottom or they may be stiff and erect and attached to rocks (Photis). Mud is occasionally used to reinforce the walls of the erect tubes.
Glands of domiciliary Amphipoda appear to be concentrated mainly, if not exclusively, in pereopods 1-2. They are found most heavily concentrated in article 2, often in article 4 and occasionally in other articles. Glands usually appear to be composed of densely packed, “volky”’ tissue, often posterior to the main muscles of article 2, often composed of morula-like bodies, and often of different color than the muscles or other tissues, even in specimens preserved in alcohol; they are frequently yellow, ochre, orange, or purple. Apparently a duct carries the secretion to a meatus located subterminally on the dactyl. The meatus is very difficult to see and not definitive of Gammaridea known to be domiciliary because many other Amphipoda seem to have a dactylar meatus but lack conspicuous glands. The presence of these glands may be a useful clue by assisting in the identification of several genera in Isaeidae, Ischyroceridae, Corophiidae, and Am- pithoidae that otherwise resemble nondomiciliary Amphipoda. Com- plete reliance cannot be made on them as familial characters, for all members of domiciliary familes do not necessarily have glandular pereopods 1-2 and very few species have actually been examined for their presence. Some phoxocephalids, haustoriids, and argissids have conspicuous glands in pereopods 3-5.
Most domiciliary amphipods except for the Ampeliscidae have somewhat depressed bodies, shortened pereopods, and have better crawling ability than do nontube builders. They can be observed emerging from their closely crowded tubes, their antennae appar- ently being used digitally in a search for particles of food. Ampeliscid amphipods lie upside down in their tubes and project their strongly setose antennae as filtering organs. Only one benthic gammaridean has been reported as a predator but some pelagic amphipods catch prey.
After hatching, young amphipods are like adults (Gammaridea do not have larvae as do most crustaceans) and are carried about for a few hours or days in the brood pouch. In the early stages molting and growth are rapid; a young amphipod may first molt while still in the brood pouch, within a day or two after hatching. As growth proceeds rates of molting and growth decrease, so that adults may molt every 20-30 days and in some striking instances only every six months or so. Few amphipods have been studied for their molting rates; of those examined it has been determined that sexual ma- turity is reached at about the sixth molt (commencing the seventh
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instar); the animals are fertile yet the secondary sexual characters are rudimentary; male gnathopods may be poorly developed and brood plates are just beginning to develop in the female. Amphipods are known to live through at least 12 instars, the females laying a brood of eggs during the last five or six or in alternative instars; how- ever, terminal adult females (gerontics) may lose their brood plates, apparently fail to lay eggs and develop aberrencies of an andromorphic nature. Occasionally, these stages have been described as distinct species.
Amphipoda have the unfortunate habit of eating their exoskeletons after ecdysis so that it is difficult to trace their molting sequence in the laboratory. One must watch them continuously in order to obtain ecdysial casts. Because the average instar appears to last about 15 days, the average maximum length of life is expected to exceed 6 months but some species in polar regions are estimated to live 5 or 6 years. Often the structure of the next instar may be seen within an appendage. Such replication within the present organ can be confusing to taxonomists and lead to the description of supernumerary parts.
A few species are known to be hermaphroditic, carrying both male and female gonads, and reflecting both sexes in their secondary sexual characteristics.
Paired pleopods on the first three segments of the pleon are biramous, the rami multisegmented and strongly setose. Minute coupling hooks on the medial edges of the peduncles are used to engage the pairs of pleopods for coordinated paddling. Amphipoda usually are good swimmers. Even burrowing amphipods swim well and phoxcephalid males have the habit of leaving their burrows at night and swimming to the seasurface from depths as great as 100 meters. They will swarm around a light suspended in the water. They may be ascending in search of females even though the latter rarely swim to a night-light. Such swimming behavior may be a dispersal mechanism, especially in groups having a low proportion of males. Some pelagic species apparently undergo great vertical migration; several deep-sea amphi- pods caught at night near the surface have been found to have ali- mentary tracts full of benthic sediments.
Variation in pleopods is rarely of sufficient extent to be weed in generic or familial definitions except for Phliantidae, Talitroidea and some Corophiidae. Pleopodal morphology, nevertheless, has been neglected and may afford some help in taxonomic distinctions.
There is justification in restricting the term pleon (=metasome) to the first three abdominal segments bearing pleopods and utilizing the term urosome for the last three abdominal segments bearing uropods. “All” Malacostraca have at least one pair of uropods, that pair of appendages on abdominal segment 6. Malacostracans generally have
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five pairs of pleopods on segments 1—5, but in noncaprellidean amphi- pods the appendages of segments 4 and 5 resemble the terminal uropod and thus are called uropods 1 and 2. Uropods in many mala- costracans are still used for swimming although in different fashion from pleopods, but uropods 1-2 in Amphipoda appear to be used primarily for strengthening the caudal portion of the body to permit jumping or flipping by rapid flexion of the urosome. In many Gam- maridea, the third uropods still bear “‘swimming”’ setae, and may be used for paddling or as rudders. Males especially have natatory third uropods. But the vast majority of Gammaridea probably do not use the third uropods for active swimming and they are often reduced or occasionally absent in sedentary species. Caprellidea have lost all but a vestige of the abdomen and its appendages.
The telson is a flap attached to the sixth pleonite above the anus. It is of primary taxonomic value, depending on whether it is cleft into two lobes, fused into a single flap, elongate, fleshy, or ornate. Possibly, it is a vestige of paired appendages.
The head bears two pairs of antennae. The first three articles of the first pair are known as the peduncle, the remaining smaller articles the flagellum. In many species an accessory flagellum demon- strates the biramous derivation of the appendage; when present, it sprouts from the end of the third peduncular article and may be elongate or reduced to three, two, or one articles. Although appearing to be of minor importance, the condition of the accessory flagellum is crucial to amphipod systematics and is useful especially at familial and generic levels. The second antennae bear five peduncular articles, followed by a single flagellum. The flagella of both antennal pairs may bear, especially in males, sensory appendages, such as aesthetascs and calceoli. Male antennae often are longer than those of females.
Frequently families and genera have been defined as lacking accessory flagella and then have been shown to have some members bearing extremely small 1l-articulate pieces. Allowance should be made by the observer so as to admit to genera and families those species with microscopic remnants of accessory flagella that have heretofore been overlooked.
The mouthparts are composed of the following structures; they are highly variable intergenerically and their morphology is important for classification:
Upper lip: A single lobe or flap anterior to the mouth. In about 10 percent of known species the anterior cephalic surface above the upper lip is produced into a point, keel, or lobe known as the epistome. Its function is unknown. In a few families, especially Lysianassidae, the upper lip has a keel projecting anteriorly and usually separated from the epistomal region by a deep slit or sinus. Occasionally both
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epistome and upper lip are produced together and occasionally they are fully amalgamated.
Lower lip: A bilaterally symmetrical complex forming a partition behind the mouth. It is composed of at least a pair of lateral lobes, having their lateral extremities produced, often acutely and often bearing apicomedially a tiny cusp enclosing the meatus of a salivary duct. About half of the known gammarideans has a pair of medial lobes on the lower lip.
Mandibles: A pair of appendages attached lateral to the mouth; with the upper and lower lips they form a box around the mouth, permitting buccal closure. The mandibles are powerful and difficult to remove because of their large muscles. Mandibles generally have their anterodistal ends (incisors) cut into a series of teeth for biting; just proximal to the distal teeth may be an articulated process, also toothed, the lacinia mobilis (accessory plate) which may occur on only one of the mandibles. A molar with a grinding surface often occurs on the medioventral surface of the mandible. It may be ridged and toothed (triturative), or smooth, or be completely absent, es- pecially in inquilinous amphipods having the mandibles elongated for piercing and sucking. Most Gammaridea have a 3-articulate palp attached to the dorsolateral surface of the mandible, the palp being used to clean the bases of the antennae. Its absence is mod- erately frequent and often of familial importance, but its reduction to two or one articles is uncommon.
First maxillae: These are situated posterior to the lower lip. This pair of appendages is small, each bearing a medial free lobe, an outer lobe with heavy spines, and attached to the outer lobe a palp composed of one or two segments, occasionally reduced in size or absent.
Second mazillae: These are two pairs of lobes behind the first maxillae, each composed of simple medial and lateral plates, occa- sionally reduced to one plate or absent, rarely with the outer lobe attached to the inner by a basal geniculation or extension. Basal articles of maxillae and maxillipeds are present but rarely are of taxonomic importance.
Mazxillipeds: One pair of appendages posterior to the maxillae, each maxilliped is formed of an inner (proximal) lobe, an outer (distal) lobe and a palp of two to four articles, rarely absent in Gammaridea, but always absent in the pelagic and often inquilinous Hyperiidea and in cyamid Caprellidea. The Ochlesidae, lacking such palps, are assigned to the Gammaridea on the basis of their resemblance to Gammaridea in other morphological features and their supposed benthic habits. In peracaridan phylogeny the max- ilipeds were originally the first pair of thoracic legs, but they have
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become incorporated into the cephalic complex; in some talitrids such as Orchestoidea, the lines of fusion of this thoracic segment to the head are still apparent but in most amphipods such external demarcation is obscure.
In comparison to crustacean groups such as the Isopoda, the body plan of the Gammaridea is conservative. Extreme deviations from the laterally compressed body with enlarged coxal plates occur in a few families in which the body has become dorsally depressed (Corophiidae, Cheluridae, Podoceridae). Two or more of the urosomal segments have become fused in the Ampeliscidae, Atylidae, Chelu- ridae, and Kuriidae. Coxae have become deformed or enlarged in some Lysianassidae, Astyridae, and Hyperiopsidae. The body is shortened and puliciform in some Haustoriidae and immensely elobular in some Lysianassidae. In the Podoceridae the first urosomal segment has become elongated and in the EKophliantidae and Colo- mastigidae the body becomes subcylindrical as in tanaids.
Major ornaments, of taxonomic value generally at the specific level, include the frequent occurrence of a rostrum, the differentiation of lateral cephalic lobes, the presence of processes on the peduncles of the antennae (especially the Acanthonotozomatidae and Lepechinel- lidae), and the cuspidation of the pleonal epimera. Dorsal ornamenta- tion is most common among cold-water Gammaridea and occurs in the form of teeth and cusps on the pereonites and pleonites. The first urosomite is often ornamented even in genera without other display.
The absence of eyes is rarely of taxonomic concern other than at the specific level; indeed many sublittoral oculate species are known to have eyeless populations in bathyal depths.
The condition of the head in the Gammaridea is highly variable and thus useful to the taxonomist, but its morphology is often left unde- scribed. The “basic”? gammaridean generally has the head about as long as 1.5 pereonites but it varies in different families and their genera from much shorter than the first pereonite to as long as the first three pereonites combined. The elongate head is especially noticeable in the Ampeliscidae, Phoxocephalidae (including the visor-like rostrum), Synopiidae, and Oedicerotidae. In the latter two families the head is considered ‘‘massive”’ because it is not only elongate but very deep. Other families, such as the Stegocephalidae may have deep heads but they are much shorter than the first three pereonites combined. Recognition of Synopiidae is almost fully dependent on the final confirmation of a ‘‘massive”’ head. Most members of that group have a massive head primarily because pereonites 1-3 are so short that the head appears relatively large by comparison.
The presence and/or condition of the cephalic rostrum is only occasionally conservative at the familial level (e.g., Phoxocephalidae,
MARINE GAMMARIDEAN AMPHIPODA 13
Synoplidae). Its presence is of relatively uniform value at the generic level (example of an exception is Bathymedon) and its shape is often of good specific value.
Gill structures have been used occasionally for specific and generic distinctions but their conditions have been ignored in most Gamma- ridea and require extensive study. Gills are often well known in those genera in which accessory tube-like branchial appendages have been discovered or where the primary gills are extraordinarily plaited or folded. Brood lamellae of the female also have been largely ignored although a great deal of variation occurs in their shape, setosity, and terminal ornamentation of the setae. They and the gills may be of assistance in tracing phylogenetic relationships among families and superfamilies of Gammaridea. Male reproductive appendages gener- ally occur as a small pair of projections on the seventh pereonal sternite; occasionally they are spinose. Other sternal teeth, keels, and flanges appear to be of rare occurrence (e.g., Aoridae, Eophliantidae).
Geographic Distribution of Marine Gammaridea
The distribution of so few species of Gammaridea is well known enough as to offer a precise statement of their geographic distribution, but the distribution of most genera is moderately well outlined be- cause, of course, bits of data afforded by each species are cumulative for genera.
Genera can be sorted out relatively easily by gross geographic zones because faunistic monographers have concentrated primarily within these zones. Only Sars (1895), Chevreux and Fage (1925), and Schellen- berg (1926a) have written faunistic monographs broadly overlapping two zones. Other large scale papers, seemingly monographic, are but obvious collectors’ assemblages. Warm-temperate analyses have suf- fered for the lack of discretion between that zone and either tropics or boreal (=cold-temperate) and I have not taken the time to segre- gate precisely the boundaries between the broad arctic-subarctic regions and cold-temperate in describing the distribution of each species. Arctic Amphipoda of the north polar basin are very poorly known and probably very sparse; most of the northern Siberian, Alaskan, Canadian, Greenlandian, and Norwegian shores are placed in the arctic-subarctic region; Iceland, Kamchatka Peninsula, and Okhotsk Sea are considered as boreal (cold-temperate), and that region extends southward to the Japan Sea, middle California, Cape Hatteras, and the Breton Capes; the warm-temperate includes the ‘southern and Baja Californias, Mediterranean Sea, much of north- west Africa, and the northern Gulf of Mexico; south warm-temperate
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includes the southwestern and southeastern coasts of Australia, South Africa, and Peru to middle Chile; antiboreal includes Tasmania and all of New Zealand, for convenience, and in South America includes all of Schellenberg’s (1931) Magellan and Falkland fauna. South Georgian faunas are thrown into antarctic-subantarctic classification. The tropics of western South America end at approximately 4° S. Warm-temperate of eastern South America is indefinable but of no consequence because of the absence of gammaridean studies along most of that coast except in obvious tropical or cold-temperate regions.
Gammaridean genera have been found to fall relatively easily into these broad classes, the genera either being confined to one class or being of such wide distribution as to be called cosmopolitan. The latter term, however, primarily refers to genera that radiate outwards from tropical regions into boreal regions but not into arctic-antarctic regions, and there has been little point in so splitting the analysis to segregate cosmopolitan genera that extend high into polar regions. Bathyal, abyssal, and hadal faunas are highly discrete, poorly influ- enced by submergent polar faunas, and, thus, are recognizable as distinct from latitudinal considerations; the few deep-sea genera that have been found only in polar regions have been removed from their endemic position in those regions to the deep-sea classifications.
No precise statistical methods have been used in dealing with problem genera, those with distributions partially overlapping two classes, because a bit of subjectivism has been applied in each case and because the principle of ‘‘centralism’”’ has been utilized. Genera are thus considered to be confined primarily to that region in which ‘“‘most”’ of the species occur, to wit: a genus with two boreal and one subarctic species is considered to be boreal but a genus with eight boreal, two warm-temperate and one tropical species is thrown into the cosmopolitan class on the probability that more tropical species remain to be described.
The results of this subjective analysis are presented in table 1; on first sight the data seem to reveal mostly a relationship to study effort, with faunas of low latitudes or southern quartospheres suffering by comparison with well studied boreal-arctic regions. On the other hand the data seem reasonable if one considers that antiboreal regions are few in number, small in size, and low in habitat-diversity com- pared with the extensive boreal regions; except for small oceanic islands the antiboreal region is confined to Tasmania, New Zealand, and two coasts of South America, whereas north boreal regions have four coasts on two continents and a significant disjunct subarctic embayment, the Okhotsk Sea. The rich antarctic shelves are a strong contrast to the polar-arctic impoverishment but perhaps the most striking implication in the data is the low count of tropical endemic
MARINE GAMMARIDEAN AMPHIPODA
15
TaBLE 1.—Geographic classification of gammaridean genera
Class or Zone
Cosmopolitan Antarctic-subantarctic Antiboreal
South warm-temperate Tropical
North warm-temperate Boreal Arctic-subarctic Bipolar !
Biboreal only Biwarm-temperate Bathyal only
Abyssal only
Hadal only
TOTAL FIRST GROUP AMALGAMATION:
Arctic-boreal
Antarctic-antiboreal
North warm-temperate ?
South warm-temperate ?
Cosmopolitan (including bipolar, biboreal, bi-WT) Deep-sea
Tropical
SECOND GROUP AMALGAMATION (genera only):
Antarctic (including bipolar)
Arctic (including bipolar)
North warm-temperate (including cosmopolitan, bi-WT)
South warm-temperate (including cosmopolitan, bi-WT)
Boreal (including cosmopolitan, bipolar, biboreal)
Antiboreal (including cosmopolitan, bipolar, biboreal)
Tropical (including cosmopolitan)
Genera
104
84
183 132
105
Species in those Genera
898 110 75 48 111 61 551 ial 488 64 7
72 4
2723
622 185 68 55 1457 239 111
1 Including arctic-boreal and antarctic-antiboreal. 2? Including biwarm-temperate (bi-WT).
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genera. Even though numerous species of tropical amphipods remain
to be described, my experience in sorting through quantities of
tropical materials suggests that most of these species will be described in known genera. New genera seem to be confined primarily to inquilinous forms probably associated with the vast numbers of
sessile tropical invertebrates. Thus Gammaridea are primarily a
cool-water group in terms of generic diversity; this is reflected in
their strong penetration of the deep-sea. Their body sizes are very strongly associated with thermal conditions in shallow waters, the larger bodies occurring in colder water. This does not apply to deep-
sea benthic Amphipoda, however, for there is strong indication (J. L.
Barnard, 1962d) that body size becomes smaller or remains relatively
static with increase of bottom depth along a latitudinal line.
About 40 percent of the 53 families of Gammaridea can be classified as cosmopolitan in distribution but the other 60 percent are moderately to strongly confined to specific regions or thermal zones (see below). They are primarily cold-water oriented, for only 6 families with 21 genera and 53 species are confined to warm shallow waters of low latitudes. This seems to confirm the orientation of Gammaridea to cool waters.
The geographic distribution of noncosmopolitan gammaridean families, or those almost wholly confined to such classification, are as follows:
ANTARCTIC-ANTIBOREAL: Acanthonotozomatidae, Pagetinidae.
S. WARM-TEMPERATE: Ochlesidae.
TropicaL: Anamixidae, Bateidae, Kuriidae, Leucothoidae.
Boreau: Cressidae, Dogielinotidae, Lafystiidae.
BatuyaL: Astyridae.
ApyssaL: Hyperiopsidae, Lepechinellidae, Vitjazianidae.
CoLD WATER; BIPOLAR SUBMERGENTs: Atylidae, Eusiridae, Haustoriidae, Lysianassidae, Melphidippidae, Oedicerotidae, Pleustidae, Paramphithoidae, Pardaliscidae, Sebidae, Stegocephalidae, Stilipedidae, Synopiidae, Thaumatel- sonidae.
WARM WATER; LOW LATITUDES: Phliantidae, Prophliantidae.
In summary, there is scarcely a coastline where an exploratory taxonomist interested in species diversity cannot make some contribu- tion, as Gurjanova (1962) has so ably shown in her study of boreal- subarctic north Pacific; the taxonomist of western Europe, however, must move into beta-taxonomy and the northwestern Atlantic taxonomist must move in that direction in order to detect the rela- tionships of his fauna to that of western Europe. So also, must the northeast Pacific taxonomist orient himself to the groundwork laid for him in Russian Pacific works. The exploratory taxonomist in- terested in total generic diversity will find the greatest needs for study in all warm-temperate regions, the antiboreal, the tropics, and various islands of low latitudes.
MARINE GAMMARIDEAN AMPHIPODA 17 The Composition of a Faunule
A faunule is the aggregate of species of Gammaridea in one geo- graphic province, region, or habitat. Five megafaunules are well known: cold-temperate of northeast Atlantic, cold-temperate of northwest Pacific (including the arctic-like Okhotsk region); circum- subantarctic encompassing the Norwegian basin and the north coasts of Siberia; the warm temperate of northeast Atlantic including the western Mediterranean and the circum-antarctic-subantarctic faunule. Two faunules secondarily well known are those of the South African warm-temperate and the northeastern Pacific warm-temperate. The faunules of individual islands or small geographic areas also have been more intensively studied than of these larger provinces, examples being Plymouth, England; Naples, Italy; South Georgia Island.
A good model faunule is that of southern California because it is midlatitudinal and relatively well explored by quantitative sampling. The littoral-sublittoral (coastal shelf 0-100 m) faunule comprises 186 reported species of which 59 species live in both littoral and shallow sublittoral depths; thus the littoral (intertidal) faunule totals 138 species, including about 5 species of beachhoppers and the sublittoral (coastal shelf) faunule totals 166 species.
TaBLE 2.—Numober of species and genera of Gammaridea in known faunules
Region Genera Species
Warm-temperate California
intertidal (compiled) 66 138 sublittoral 98 166 total, less common to both ; 112 186 Norway (Sars, 1895), boreal section, mainly sublittoral, 0-80 fms 115 221 0-50 fms 110 203 Plymouth, England (Plymouth Marine Fauna), inter- 63 92 tidal Isle of Man (Bruce et al, 1963) intertidal 40 66 Mediterranean France (Chevreux & Fage, 1925), inter- 41 72 tidal Atlantic France (Chevreux & Fage, 1925), intertidal 69 109 South Georgia Island 88 159 South Georgia Island, intertidal only (Schellenberg, 45 62 1931) Falkland Islands, intertidal (Schellenberg, 1931) 48 63 Magellan continental, intertidal (Schellenberg, 1931) 57 Ut. Indo-Pacific tropics (not including Red Sea with an 97 204 additional 22 species) (J. L. Barnard, 1965)
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TaBLE 3.—Dominant benthic genera of Gammaridea in sublittoral soft-bottom faunule of southern California (+ =yes; 0=no.)
Kind of domination
Genus Number of —@=—<2-—_____— species Specific Frequency Diversity of Individuals Ampelisca ! 12 =P ots Byblis ! 1 0 + Ericthonius 2 1 0 + Eurystheus 2 1 0 + Heterophoxus 2 1 0 + Listriella 4 5 + 0 Metaphoxus 2 2 0 sta Monoculodes 4 0 + Paraphoxus 3 18 ste te Photis 2 5 IP ae (Phoxocephalus)? are 0 + Protomedeia 2 1 0 + Synchelidium * 5 a as Westwoodilla 3 1 0 +
1 Forming limp tubes on or in substrate. 2 Forming stiff or limp tubes on particles or sessile infaunal structures. 3 Burrowers. ‘4 Nestler or semicommensal.
Various other faunules are compiled in table 2. No attempt has been made to modernize certain faunules such as Sars’ Norwegian boreal in order to maintain congruency among the reports; thus each faunule presumably represents a minimal statement on diversity as a result of extensive, but not exhaustive exploration.
Generic domination within a faunule may occur through specific diversity or through high frequencies of individuals in poorly diverse genera. The Californian faunules of tables 2 and 3 thus comprise some dominant genera having numerous species, only a few of which have high frequencies of individuals and other monospecific genera, with extraordinarily abundant individuals per square meter of habitat. Other faunules have been compiled in table 4 on the basis only of intrageneric diversity, as quantitative information is virtually absent except in the California region. These few examples may give the reader an approximation of what to expect in commencing quantita- tive explorations of provinces.
The ecological kinds of Gammaridea within a benthic faunule are to a large extent determined by domiciliary position rather than food-type, as most Gammaridea are presumed to be scavengers or inquilines and their feeding behavior is poorly known. A few true herbivores apparently occur but only one raptorial predator has ever
MARINE GAMMARIDEAN AMPHIPODA 19
TaBLE 4.—Dominant benthic genera of Gammaridea in littoral epifaunal faunule of southern California (+ =yes; 0=n0.)
Kind of domination Number of @=_—-2#2______ Genus species Specific Frequency of Diversity Individuals
*Ampithoe 11 ae at * A oroides i 0 ate Elasmopus 9) aig als *Hricthonius 1 0 + *Eurystheus 4 ap ate Hyale 3 0 oF *Ischyrocerus 2 0 ais *Jassa 2 0 ae Maera 9) ah 0 * Microdeutopus 1 0 +
Parapleustes 4 ote +0 *Photis 6 qr ate
*Forming tubes; others are nestlers.
been described. Of course there are numerous kinds of scavenger feeding. An intertidal faunule of low latitudes in lush algae will be comprised of extremely abundant nestlers such as Hyale, Elasmopus, Maera; protected rock surfaces, interstices and root systems of algae or surf-grass will contain the domicolous amphipods building tubes, such as Ampithoe, Ericthonius, Gammaropsis, Jassa, Ischyrocerus, and Photis. Sessile invertebrates like sponges and tunicates will harbor nestlers and domicilary kinds as well as a few inquilines, such as Leucothoe and Polycheria.
The sublittoral faunule on soft bottoms will be comprised of bur- rowers such as Paraphorus (and other phoxocephalids), haustoriids, oedicerotids; domiciliary kinds building limp tubes on the sediment surface, primarily the Ampeliscidae; various domicilary kinds build- ing tubes on hard particles of the substrate or on projecting tubes of infaunal organisms, thus the Isaeidae (Photidae) ; and a few nestlers or semicommensal organisms like Listriella.
Intertidal zones of high latitudes will have nestlers like Pontogeneia, Paramoera.
The dominant genera of Gammaridea on sublittoral soft bottoms by gross region are as follows (*=genera with low specific diversity but high individual frequency):
CosMopoLitan: Ampelisca, [Gammaropsis], Idunella, Listriella, Lysianassa,
Metaphoxus, Paraphoxus, Podocerus, Synchelidium, Urothoe.
BIPOLAR, COLD-WATER: Aristias, Byblis, Epimeria, HEusirus, Liljeborgia, Mel- phidippa, Orchomene, Podoceropsis, Tryphosella (‘‘Tryphosa, Tmetonyz’’).
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Antarctic: Oediceroides, Pseudorchomene*, Tryphosites*, Uristes.
ANTIBOREAL: Acontiostoma*, Amaryllis, Heterophocus.
Tropica: IJdunella, Platyischnopus.
WARM-TEMPERATE: Listrvella.
ARCTIC-BOREAL: Anonyx, Bathyporeia, Dulichia, flock of haustoriid genera, Lepidepecreum Monoculodes, Oediceros, Onisimus, Paramphithoe, Pontocrates, Pseudalibrotus, Westwoodilla, Unciola.
The dominant genera of Gammaridea in epifaunas of intertidal and shallow sublittoral (0-75 m) zones by gross region (*—genera with low specific diversity but high individual frequency; N—nestler; I=inquiline; D=domicolous tube-dweller).
CosMopoLitaN: Allorchestes N, Ampithoe D, Cerapus D, Colomastiz I, Corophium D, Ericthonius D, Gammaropsis D, Hyale N, Jassa D, Lembos D, Leucothoe I, Maera N, Melita N, Photis D, Podocerus D, Polycheria I, Stenothoe I.
BIPOLAR, COLD-WATER: Aora D, Apherusa N, Halirages N, Liljeborgia N, Paramoera N, Pontogeneia N, Proboloides (Metopoides) N.
ANTARCTIC-ANTIBOREAL: Amaryllis, Andaniotes ?1, Atyloella N, Atylopsis N, Bovallia N*, Djerboa N*, Echiniphimedia, Gnathiphimedia, Haplocheira D, Oradarea, Paradexamine, Pariphimediella, Pontogeniella N, Seba 1, Schraderia N*, Stebbingia N*, Thaumatelson 1.
WARM-TEMPERATE AND TROPICAL: Amphilochus, Anamizis I*, Batea N, Ceradocus N, Cheiriphotis D*, Chevalia D*, Cymadusa D, Elasmopus N, Guitanopsis, Grandidierella D, Leucothoides 1*, Microdeutopus D, Microjassa D, Paragrubia D*, Parelasmopus N, Parhyale N*.
ARCTIC-BOREAL: Amphilochus, Anisogammarus N, Aoroides D*, Atylus N, Calliopius N, Cressa, Gammarus N, Gammarellus N, Gitanopsis, Ischyrocerus D, Leptocheirus D, Metopa, Metopella, Microdeutopus D, Neopleustes N, Para- pleustes N, Pleustes N, Sympleustes N.
The dominant genera of Gammaridea in the pelagic realms, in- cluding some demersal genera, are as follows (*=dominant by frequency of individuals only).
Neritic: Megaluropus, Paraphoxus (males), Synchelidium (pelagic phases), Synopia.
Eprpetacic: Stenopleura*, Synopia.
BaTHY- AND ABYSSOPELAGIC: Andaniexis, Astyra, Cleonardo, Cyclocaris*, Cyphocaris, Huandania*, Euonyz, Eusirella, Eusitrogenes*, Eusirus, Halice, Hirondellea, Hyperiopsis, Ichnopus, Joubinella, Koroga*, Metacyphocaris*, Orchomene, Paracallisoma*, Paralicella, Parandania*, Parargissa, Phippsiella, Rhachotropis, Scopelocheirus.
The dominant genera of bathyal-abyssal benthos are as follows (*=dominant by frequency of individuals only. D—=demersal?). Conservative numbers of species in depths exceeding 200 meters are given following each genus.
BENTHIC: Amathillopsis 6, Ampelisca 25, Anonyx 5 (Pacific boreal), Bathy-
amaryllis 4, Byblis 5, Bonnierella 5, Bathymedon 4, Bruzelia 4, Dulichia 6
(Atlantic boreal), Epimeria 6, Haploops 8, Harpinia 4 (Atlantic boreal), Harpiniopsis 19, ‘“‘Hippomedon’”’ 12, Lepidepecreum 6, Leucothoe 4, Liljeborgia
MARINE GAMMARIDEAN AMPHIPODA 21
6, Melita 5, Metopa 7, Monoculodes 4, Oediceroides 11, Onesimoides 3, Orchomene 14, Proboloides 7, Pseudharpinia 5, Schisturella 5, Stegocephaloides 5, Stenothoe 4, Syrrhoe 5, Syrrhoites 8, Tryphosella 20, Unciola 6 (Atlantic boreal), Uristes 11, Urothoe 5.
DEMERSAL?: Aristias 8, Astyra 5, Cleonardo 8, Eurythenes* 2, Eusirus 9, Halice 8, Lepechinella 13, Pseudotiron 3, Rhachotropis 20, Valettiopsis 3.
Morphological Evolution of the Amphipoda
Amphipoda comprise four unusual suborders: (1) the Gammaridea, primarily benthic, with perhaps 20 percent pelagic and demersal species, but having apparently radiated a half dozen or more times into a gradational suborder, (2) the Hyperiidea, marked by fully pelagic (free or inquilinous) habits, the strange but universal loss of maxillipedal palp, and a spectrum of other minor changes presumably correlated with their habits; perhaps from podocerid gammarideans have evolved (3) the Caprellidea, skeleton shrimps or marine praying mantises, characterized by extremely thin tubular bodies, reduction in abdomen, reduction in two pairs of pereopods, increased cephali- zation and primarily adapted to a sedentary life in epifaunal anas- tomoses; but through secondary body depression within the caprelli- dean scheme (like Temnophlas in the gammaridean scheme) arose the cetacean ectoparasites Cyamidae, essentially comprising a fifth major group of Amphipoda; and finally (4) the Ingolfiellidea, appar- ently undergoing development in association with troglobitic condi- tions but occasionally returning to marine niches yet open to those organisms with vestigial pleopods, often bearing cephalic ‘‘ocular’’ scales and with, perhaps, other special ecological adaptations.
Even though Amphipoda have radiated into nearly 60 families the major diversity can probably be visualized in terms of a score of kinds. These typological centers may be described by the following adjectives: ingolfiellid, cyamid, caprellid, six to eight kinds of hyperiid, gammarid, eusirid, isaeid, lysianassid, colomastigid, eophliantid, phli- antid, talitroid, stenothoid, amphilochid-leucothoid, ampeliscid, and acanthonotozomatid. Some outgrowths of these centers form radical morphs but they fail to qualify as typological centers because (1) their relationships are not discontinuous or (2) they have not radiated strongly. The Cheluridae are an example of a radical morph with presumed relationships to the Isaeidea and low internal diversity; they are not considered as a typological center. The Colomastigidae, though of low diversity, have discontinuous relationships with other Amphipoda and are, therefore, considered as a typological center. The eusirids (not necessarily the type-genus) have clear relationships to another center but have radiated so strongly that they must be considered a typological group. My concept of these centers on cur-
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rent knowledge is weak and open to extensive revision as we come to understand the micromorphology, anatomy, and chemistry of the various Amphipoda.
On numerous occasions parallel adaptation, convergent evolution, and independent evolution of morphofunctional conditions have occurred in Amphipoda. These concern cylindricalization of bodies (Eophliantidae, Podoceridae, Colomastigidae), cylindricalization of heads (Cheluridae and Eophliantidae), dorsoventral flattening of body (Temnophlias, Podocerus and Cyamidae), development of domitubicolous glands (Ampeliscidae and Isaeidea), loss of max- illipedal palps (Hyperiidea, Ochlesidae, Cyamidae, and some Lysi- anassidae), and in a host of minor ways. But, for example, the diversity does have a measure of constriction in that Amphipoda have never evolved as fully as have the Copepoda into numerous parasitic modes or, to our knowledge, have the Amphipoda developed hosts of rapacious or errant predators in the benthic realm. Predators do occur in the nektonic Hyperiidea and Gammaridea but none of the former and few of the latter have returned to a benthic orien- tation. Although few Isopoda have any degree of lateral compression, whereas many Gammaridea do have dorsoventral depression, the Isopoda would seem to be the more highly diversified because they have cylindrical representatives (Astacilla) and fully evolved para- sites (Bopyridae). In contrast, the Amphipoda are far more diverse than certain other orders of Peracarida, such as the Cumacea and the Tanaidacea. Fourteen families of Gammaridea alone are more or less inquilinous.
A microhabitational stress must exist between various Amphipoda and members of the other crustacean orders and phyla, which re- stricts a fuller display of genetic potential than now in existence. The tendency of some Amphipoda to enter the crawling realm of Isopoda, indicates that were Isopoda extinct, Amphipoda could fill many of those niches, even though rudimentarily or imperfectly.
Various members of the Gammaridae have been considered as the most primitive of living amphipods. They display most of the basic gammaridean morphology but the strong development of the lateral shield (Gurjanova, 1962), composed of coxae or pereopodal wings in many gammarids, suggests that they have strongly differentiated from a precursor lacking such a shield. If pereopodal tube-spinning glands represent a secondary development in Amphipoda, then many of the isaeid genera that might be considered as close to a shieldless precursor, have probably undergone a secondary reversion by a reduction of the lateral shield (e.g., Corophiidae). Other groups with reduced lateral shield (Eophliantidae, Podoceridae, Colomastigidae) apparently do not stand close to the primitive amphipod model be-
MARINE GAMMARIDEAN AMPHIPODA 23
cause of vastly modified mouthparts or of so-called pygidization, the solidification of the urosome and its appendages by segmental coales- cence or loss of uropodal rami and peduncles. Of course, other per- acarids without lateral shield may also have tube-spinning glands (tanaidaceans) and the primitive amphipod may have been in the glandular line. There are sufficient intergrades in telsonic morphology between isaeids and gammarids to make this suggestion very attrac- tive. Several isaeids are almost perfect replicas of the basic gammarid- ean except for their fleshy telsonic nobs. One has to balance at least two alternatives, whether the coalesced telson represents a full segment lost in phylogeny or whether the lobed telson of gammarids represents a pair of appendages long lost. This problem is fundamental to other crustacean orders and my impression is that the uncleft, solid telson is the most common, thus suggesting that it is either more primitive or at least more successful functionally. The telson in Amphipoda is not a conservative feature by any means, and our understanding of it as an evolutionary marker will not be clarified until we understand its function. Several cycles of morphological development, whether by advancement or regression are apparent. If the fleshy isaeid telson becomes “‘falsely’’ lobed in advanced species one can almost imagine the small step necessary to convert it into one of the poorly lobed or entire ‘‘subfleshy’’ members of the Gammaridae. Extreme flattening, full clefting, and elongation then occur in other gammarids and various derived families, but coalescence of the lobes and shortening occur again and again. In some haustoriids the telsonic lobes become fully disjunct basally, each lobe appearing as a vestigial appendage. The fully blown pygidization of Gammaridae, Isaeidae, and most gammarideans, in the sense of having the last three pairs of abdominal appendages formed into relatively inflexible, posteriorly directed uropods, obstructs our detection of an ancestor in any other living order of Peracarida, where only the final pair of appendages is formed into uropods. The reduction or loss of uropods, pleopods, and most of the abdomen in Caprellidea is clearly a secondary development. Several good intergrades occur in this procession from Gammaridea to Caprellidea in such taxa as the Podoceridae, Caprogammaridae and Cercops, a caprellidean. Most Caprellidea further have thoracic somite 2 (free segment 1 of other Amphipoda) coalesced with the head, have a reduction in pereopods 1-2, gills and brood plates. If the lateral shield or its functional substitute by means of tubicoly, serve as partial protection for brood and gills in Gammaridea, then Caprellidea, with their complete loss of lateral shield must have some other protection, perhaps reflective of their habitats or behavior. The brood lamellae of caprellids seem to be more strongly cornified than those of gammarideans. The lateral shield has also been suggested
24 U.S. NATIONAL MUSEUM BULLETIN 271
to be a frictional-flotational support for those Amphipoda known to swim in peculiar fashion on their sides. Thus the lateral shield, formed of coxae and articular wings on pereopods, may serve as a kort- nozzle* in reverse by channeling water ahead of the pleopods.
Good swimmers obviously occur in the Hyperiidea, for they are all pelagonts; nevertheless, reduction of the lateral shield is a major trend within that group. Hence, natatorial correlation to the lateral shield is far from universal and the functional morphologist will find many fruits to pick once he directs his attention to this problem. The loss of pleopods in Amphipoda is a mark of the sedentary life of Caprellidea, and the terrestrial habits of a few talitroids. Their reduction in ingolfiellids is unexplained but may have some relation- ship to an interstitial or troglobitic life. A few other inquilinous or sedentary gammarideans have reduced pleopods, but they are other- wise remarkably conservative in most domicolous, fossorial, and inquilinous amphipods, presumably because they are often used for creating water currents.
Function of the last pair of uropods is presumed to be in propulsion and ruddering as in other peracaridans, but uropods 1-2 have seem- ingly little function except as strengtheners for the urosome during explosive flexation that is a part of jumping behavior. Were this the sole function of uropods 1-2 the maintenance of at least a small degree of articular flexibility of the uropodal peduncles and rami would seem incongruous. Progressive coalescence of uropods 1-2 with the ventral margins of their segments would afford a strong ventral pad to be used as a jumping buffer. Perhaps the small degree of flexibility provides a better shock absorber than would a solid uro- some. The increase in ornamentation of uropods on certain fossorial species indicates at least a minimal function in the digging process, but perhaps this ornamentation primarily prevents coarse particles from being lodged in the cracks between uropods and segmental venters. Paddle-like expansion of rami on uropods 1-2 in a few groups (e.¢., Synopiidae) points to a swimming function. A few sketchy observations on swimming suggest that the uropods, despite their restricted flexibility, can be laterally splayed and serve in balance or braking during the end of a gliding motion.
Reduction of uropod 3 and reduction or rigidification of the uro- some occur in various tube dwellers (e.g., Corophiidae), inquilines (some lysianassids, cressids, thaumatelsonids, possibly kurids, pro- phliantids, etc.), tunnel makers (Eophlantidae) and some of those amphipods with semipermanent flexion of the abdomen (Phliantidae). Presumably jumping, swimming, and protection from unwanted par- ticles have been reduced in importance in these species while there
*A tube directing the propeller-wash of ships.
MARINE GAMMARIDEAN AMPHIPODA 25
is substituted an adaptation towards more favorable leverage for burrowing into tissues (plant or animal), maintaining special positions in tubes or channels and in streamlining. Rigidification often occurs without decrease in urosomal or uropodal size. The tube-dwelling Ampeliscidae and the inquilinous Dexaminidae have urosomites 2-3 coalesced while the lignivorous Cheluridae have the urosome greatly increased in size and the uropods greatly enlarged or modified. Jumping ability is very strong in the Cheluridae.
The ecological linkages to urosomal evolution in the Gammaridea are manifold; no single solution to functional adaptation has been necessary. The segmented urosome is a remarkably stable feature of the Amphipoda; that and its three pairs of uropods are a revolu- tionary feature unique to Amphipoda. That it has undergone only one major reversion (Caprellidea), yet has imprinted upon it numerous adaptive features while there is maintained a minimal structural stability, attests to its creation as a major part of the success of amphipods. Like the mandibles, its basic conservativeness has prob- ably been a factor in the successful dispersal of amphipods into many niches. The paradox of the almost complete loss of abdomen in Caprellidea appears to be ameliorated by the recent discovery of the Caprogammaridae (Kudrjaschov and Vassilenko, 1966) in which a podocerid-like metasome and urosome are maintained and by recall of the caprellidean Cercops in which the abdomen, though distinctly vestigial, is clearly macroscopic. The Caprogammaridae seem to demonstrate that the loss of urosome was not the trigger for the development of caprellids; rather, the extreme development of a tubular body with elongate segments, reduced coxae, terminal migration of various legs, reduction in numbers of gills and brood plates, all correlated with numerous behavioral changes were more important. The ultimate radiation of caprellids, of course, may have been assisted by the removal of a relatively useless abdomen and the complete amalgamation of head and pereonite 1.
Development of a dorsal shield in a few Gammaridea has been discussed by Gurjanova (1962). This feature is simply an extreme dorsoventral depression of the body and a splaying of the coxae in the Phliantidae. The abdomen is flexed under the thorax possibly as an additional protection to the ventrum owing to removal of the lateral shield. Many isopods, without flexed abdomen do not appear to require this protection, so there may be other reasons for phliantid flexion. Gurjanova points out possible precursors to phliantids in the calliopiid genera Chosroes and Sancho but those genera must yet be strongly segregated from phliantids because they have fully developed mouthparts and uropods. They probably should be allo- cated to a new family in order to qualify the development of a
26 U.S. NATIONAL MUSEUM BULLETIN 271
dorsal shield as a major, albeit rare, adaptation of Amphipoda. Dorsal shielding is also seen in the calliopiid Amphathopsis and it thereby forms a strong link among other Calliopiidae, the two quixotic genera, and is conducted into the Laphystiopsidae. To some extent body depression also occurs in a few Podoceridae and this seems significant in light of their presumed derivation from tubicolous isaeids having lost the tube-spinning glands. This loss may be correlated with such dorsal shielding because both methods of protection would seem unnecessary together.
Temnophlias, a ‘“‘phliantid,’”’ has also been included by Gurjanova as a member of the progression from Amphithopsis through Sancho into the Phliantidae but another suggestion might be made: that Temnophlias is really a cylindrical organism with secondary pleuroni- zation of the pereonites similar to munnid isopods. It may have affinities with the EKophliantidae, the most strongly developed of the cylindrical gammarideans. Cylindricality and dorsal shielding are difficult to separate as various corophiids, chelurids, and aorids have always been considered to be depressed organisms rather than cylindrioid. Both terms partially apply to these groups. But one may consider that those organisms have substituted domiciliary habits in the form of tube building or burrowing for the true lateral shield and that dorsal depression is a consequence of cylindricaliza- tion that cannot be carried too far without rendering the organism positionally unstable. Some depression is required for the organism to maintain a crawling equilibrium. Gurjanova appears to regard Phliantidae (dorsal shields) and EKophliantidae as cohesive, but eophliantids are strikingly cylindrical. There is now evidence that eophliantids are phycophilous burrowers and this would correspond with the habitats of other families living in diverse kinds of ‘‘tunnels.”’ The spherical heads and cylindrical, rotatable necks of eophliantids are suggestive of limnoriid isopods and are presumably associated with the tunneling habit. They strongly contrast with Phliantidae morphologically but there is one small difficulty in completely segre- gating the two families in that one phliantid is known to be a ligni- vore; one must presume that phliantids are not tunneling lignivores on morphological evidence alone.
The cylindrioid Colomastigidae again may be thought of as tun- nelers or domiciliaries because they probably inhabit tests or tissues of sessile invertebrates. But inquilinous behavior is scarcely confined to cylindrioid amphipods, for anamixids and dexaminids, both with strong lateral shields, are undoubted inquilines because of their mouthparts (Anamixidae) or their known ascidiophilous behavior (Polycheria, Dexaminidae). Piercing and sucking mouthparts are common in many other gammarideans such as Acanthonotozomatidae and various Lysianassidae.
MARINE GAMMARIDEAN AMPHIPODA 2
Stenopody is the presumed primordial condition of the arthropod appendage. How far this logic can be carried into the higher Crustacea as a precedent for any ordinal precursor is wholly philosophical as far as Amphipoda are concerned. Our ‘‘basic gammaridean”’ already has specialized anterior appendages, the gnathopods. A precursor to Amphipoda would already have one pair of maxillipeds. The highest Crustacea, the Decapoda, have those two pairs of gnathopodal homo- logues also attached to the head as maxillipeds. Nonambulatory func- tion of anterior appendages is thus a universally replicated feature of Malacostraca and one might imagine that it was a fundamental cor- ollary of amphipodan development. Amphipoda went one step further than isopods or tanaids in having not just one pair, but two pairs of functional gnathopods. With few presumably secondary exceptions, the second pair became the dominant members. The loss of stenopody signals the development of a nonwalking function as far as the anterior amphipod legs are concerned. In some Amphipoda these changes occur even in legs 3-4 (“‘pereopods 1-2”’).
The loss of stenopody was far from immutably fixed in amphipods. In returning to a slender condition, the marks of a grasping function have been left, with few exceptions. Return to stenopody, hence en- feeblement, of gnathopods is a mixed bag among Amphipoda other- wise recognized as either primitive or advanced. This suggests that gnathopodal evolvement, especially of gnathopod 2, was a primary crystallization of the amphipod plan, but that once upon the scene the original enlargement alone was unnecessary for whatever function the gnathopods were put to. Perhaps enlargement even went through a detrimental cycle of overspecialization.
Reduction in size of gnathopods must have come fairly early in gammaridean evolution for several genera of the Gammaridae have the gnathopodal size reduced. Some of the functional value of size may have been replaced by increased setosity. All but two of the eight families standing near the Gammaridae have the gnathopods enfeebled. Liljeborgiidae retained the enlarged gnathopods, and among other reasons, this fact supports the odd thesis that the inquilinous line of Amphilochidae-Leucothoidae-Stenothoidae has some relationship to the Liljeborgiidae. Those three families, despite the presence of other morphological degradations, have not lost the presence or potentiality of large second gnathopods. The maintenance of enlarged gnathopods in many members of the Eusiridae-Calliopiidae-Pleustidae is also further confirmation of their strong relationships to the basic gam- maridean. The position of the Oedicerotidae, unusually close to the Gammaridae in the scheme of graph 1, reflects their enlarged gnath- opods. But in the 10 families presumed to have evolved out of a eusirid stock, the gnathopods have become enfeebled. Marine tali- troideans maintain the enlarged gnathopod 2 but terrestrial members
285-135 O - 69 - 3
28 U.S. NATIONAL MUSEUM BULLETIN 271
often return to the stenopodous condition even though gnathopod 2 has obviously not returned to a walking function because of its pe- culiar morphology.
The seemingly primitive members of the isaeid stock also maintain the enlarged gnathopod 2 but the advanced members show either an axial reversal, a shift of domination to gnathopod 1, or an enfeeble- ment. Potentiality for enlarged gnathopod 2 is fully maintained in the Ampithoidae and Ischyroceridae and the ultimate peculiarity is reached in the Cheluridae, one species of which has gnathopod 1! ex- panded into a fully prehensile appendage like that of Maera. Although some of the most diverse and, thus, presumably successful shallow- water (and primarily tropical) genera have the fully enlarged and prehensile gnathopod 2, the trend in gammaridean evolution has been a secondary return to stenopody. Retention of the primitively en- larged gnathopod 2 in the Podoceridae and the Caprellidea, even the Cyamidea, is one more mark of their relationship. Those very success- ful tropical gammaridean genera with enlarged gnathopod 2 (in males) occur in several distinct evolutionary lines: Elasmopus, Maera, and Ceradocus in the Gammaridae; Hyale in the Talitroidea; Gammaropsis in the Isaeidae; the axially reversed condition expressed in Lembos of the Aoridae; Podocerus in the Podoceridae; the inquilinous Stenothoe of the Stenothoidae and Leucothoe of the Leucothoidae. These genera clearly have their highest diversity in the tropics and subtropics whether they had their origin there or not. The ecologist’s attention should be drawn to this curious matter.
Nontropical gammarideans with enlarged gnathopod 2 are partic- ularly conspicuous in the demersal eusirids, the liljeborgiids, various stenothoids and the Gammaridae.
Correlation of mandibular functions with morphology are poorly understood. Biting, chewing, grinding, piercing, and rasping functions are obvious, but mandibular variations are far more numerous than just those five categories. Gammaridean amphipods have been thought of primarily as scavengers, feeding on debris and detritus, carrion and dead plant fragments. The basic mandible seems to be adapted to biting off chunks with the incisors and grinding those chunks with the molarial rasp. The lack of emphasis on herbivorous habits of Gam- maridea in the literature is surprising in view of the properly adapted mandibles and the strong infestation of marine plants by amphipods. Macroscopic algae and marine grasses infested with amphipods rarely show gross cropping or evidence of bites having been removed. Stomach contents of a few phycophilous amphipods demonstrate that they probably feed on microscopic epiphytes. Undoubtedly the larger and slower growing algae have evolved mechanisms to limit the success of marine herbivores; poorly adapted macroscopic algae must surely
MARINE GAMMARIDEAN AMPHIPODA 29
become extinct in view of the almost ubiquitous hordes of marine amphipods that are potential macroherbivores. Microscopic epiphytes presumably survive through rapid growth while the total amphipod population may be restricted by the seasonality of epiphytes; probably a balance is thus maintained in ways similar to the diatom-copepod cycle of the pelagic realm. Amphipods may benefit the macrophytes by cleaning their surfaces of infesting epiphytes.
The microherbivorous amphipod with biting-rasping mandible and enlarged male gnathopod 2 may be the basic member of the am- phipodan organization. To suggest that the level bottom scavenging amphipod evolved first and invaded epifloras later would presume that cellulase secretion was not an original part of the amphipod plan. The grossly compressed bodies of the basic amphipod also attest to a preadaptation for nestling and gliding amongst anastomoses. That this ungainly structure later was able to invade a host of other habitats seemingly unsuitable to a flea-like morphology suggests that in the course of their evolution numerous ‘hidden’ adaptations accrued. Some of these may be circumstantial: high prodigality (‘‘success by numerical pressure’’), and low genetic plasticity that maintains a broad adaptability to feeding conditions. A more efficient mechanism than the basic mandible may be imagined for cropping micro-epiphytes without radical changes in the general structure. The maintenance of that basic mandible throughout so many families and genera of am- phipods, which obviously have put it to numerous functions, suggests that one key to success of amphipods is their potential omnivorous habit. A measure of this manifold feeding potential is seen in one’s ability to trap diatom-feeding amphipods by means of a carrion-baited undersea trap.
In the dispersal to level bottoms, amphipodan morphofunction almost invariably changed; the successful Ampeliscidae build tubes; the Phoxocephalidae-Haustoriidae and Oedicerotidae dig burrows and often become much broadened in their bodies; other genera, like Listriella, have obtained special associations with infaunal members of the level bottoms.
Almost all of the greater Isaeidea, the tubicolous amphipods, have maintained the basic mandible, palp included. Even the wood “‘boring”’ Cheluridae are able to rasp wood with the basic mandible. The greater Talitroidea have maintained the rasping mandible although the palp has been lost. Presumably one function of the palp is the cleansing of the anterior cephalic space between the antennae. Amphipods without palp often have few antennal setae projecting into that space to trap particles.
The trend to a distinct change in mandibular morphology is seen in some Gammaridae and even more strongly in some of the families
30 U.S. NATIONAL MUSEUM BULLETIN 271
closely associated with the Gammaridae, such as the Liljeborgiidae, Eusiridae, and Phoxocephalidae-Haustoriidae. Loss of trituration sur- face and reduction in size of molar are universal in the Liljeborgiidae, but the reasons are not yet apparent as the ecology of the group is poorly known; various burrowers in the Phoxocephalidae, Haustori- idae, and Oedicerotidae have smoothed-off molars and several of those genera have enormously enlarged molars covered with a setular velvet. Such molars are also seen in the Synopiidae. They must have some relationship to the fossorial or semifossorial habit of processing mineral grains. But in those large fossorial families the normal mandible is retained by numerous genera.
The strongest changes in mandibular morphology occur in those families presumed to be inquilines. In one way or another these families have adapted to piercing and sucking or possibly to the scraping of slime but even some inquilinous amphipods maintain the biting and grinding functions. The mandible of Polycheria is used chiefly for burrowing into the tests of tunicates for domiciliary purposes rather than for feeding. Indeed, the biting adaptation is rarely lost even in the strongest inquilines; it is maintained in many Acantho- notozomatidae, although others of the family have those incisors developed into stylets. The conformity of the acanthonotozomatid and stegocephalid mouthpart bundle suggests a gross piercing func- tion as if they normally attack some large sessile invertebrate. Pre- sumably some of the ‘‘inquilines,” like those of the greater Steno- thoidea, are grazing predators, biting off coelenterate polyps or consuming sponge and tunicate tissues.
The pardaliscid and stilipedid mandibles are the most paradoxical. They are elytriform like those of some stegocephalids, lack molars but retain palps, yet the mouthpart bundle is rarely coniform and no one has demonstrated an inquilinous behavior. Many of these species are nekters or demersal members of the deep-sea fauna.
Maxillipedal changes mark one of the primary subordinal grades of evolution within the Amphipoda. The loss of palps is a condition of the Hyperiidea. Only two families of Gammaridea and a few genera of two other families have a marked reduction or loss of these palps. Such loss is associated with a nektonic, often inquilinous habit, but numerous pelagic Gammaridea have fully developed palps. Reduction in maxillipedal plates or palps is not perfectly correlated with the inquilinous families or those marked by mandibular changes,but as we should expect, all stages of the perfection of this morphology are ap- parent and the trend is obvious. Plates and palps often evolve in- dependently as if their functions were distinct; in some cases such as Liljeborgiidae and throughout the greater Stenothoidea, the plates become reduced while the palps are maintained or increased in size.
MARINE GAMMARIDEAN AMPHIPODA dl Interfamilial Gammaridean Relationships
Through reeapitulation of the general evolutionary trends within the Gammaridea, one may estimate the pattern of evolution and rela- tive positions of the several families (graph 1). Formulation of this pattern was assisted by organizing data on 3-dimensional stick charts containing plots of characters representing grades of structure, and simple character alternatives; on wheels and triangles so as to demon- strate presence or absence of characters in small familial groups; by determining the fewest logical increments of change necessary to transform one family into another. The final pattern was influenced strongly by ampliative inference in using our knowledge of the ex- treme members of a family plus the implication that some changes require more genetic input than others even though genetic ‘“‘revolu- tions” may occur repeatedly in certain groups and that certain gross characters are very stable.
Only position and change from one condition to another could be incorporated in graph 1; thus each family appears falsely to have equal importance to all others. Estimates of distance might be incorporated as a measure of genetic input; generic and specific diversity of the familial centers could be shown by reducing the graph to an areal projection reflecting the harmony of certain grades of structure (at least in widespread families tested by a wide range of environmental selection) and the low level of diversity in familial ‘‘experiments.”’ The pattern has been organized to fit the space although distance from Gammaridae implies specialization or advancement. At best, the pattern is a straw man.
Graph 1 has been partitioned into more than a dozen blocks of one or more families (or genera). Some of those blocks, like the Colo- mastigidae or Lysianassidae, have uncertain relationships to other Gammaridea. Solid arrows represent estimates of strong and direct relationships; arrows commencing between two families or at long distances from family groups imply relationship only to ancestral pools having grades of structure similar to existing families. Dotted lines imply weak or distant relationships; wavy lines denote families placed close to Gammaridae because of precedential pathways established in adjacent examples. Symbols on the chart are reminders of certain characteristic structures and are cumulative between arrows unless marked otherwise.
The Gammaridae form the base stock and eight other families are enclosed within the block denoting gammarid-like structure, two families, the Phoxocephalidae and Haustoriidae, being so close to Gammaridae as not to warrant extra space on the diagram. They have extreme fossorial adaptations found rudimentarily in several Gammaridae.
32 U.S. NATIONAL MUSEUM BULLETIN 271
The major line of evolution out of the Gammaridae appears to pass through eusirid organisms in the next block below; this trend is first marked by reduction or loss of accessory flagellum and radiates further by various modifications to be discussed in a later paragraph. The point to be made is that the change from gammarids to eusirids seems to be less revolutionary than the changes from gammarids to various families kept within the gammarid block, and each of those probably should be accorded the rank of a block. The mark of their relationship to gammarids is the retention of the accessory flagellum in multiarticulate condition. A presumption that an accessory flagellum could be redeveloped once lost would have to be invoked in order to bring some of those gammarid-like families down into the eusirid block where grades of other structures show some resemblance. I see little objection and a slight precedent to that possibility in the fact that many Gammaridea add articles to the accessory flagellum with body growth; is there a strong difference in metameric potential between 1 and 2 or zero and 1?. If all the gammarid-block families were placed in the eusirid block only two, Liljeborgiidae and Vitjazi- anidae, could be further drawn from eusirids by reestablishing the accessory flagellum. There is no logic in funneling all other gamma- rideans through a eusirid block when many of them can be extended directly from the Gammaridae. Gammaridae are no less diverse than the three major families of the eusirid block. Each of the gammarid- block families shows some tendency in their advanced genera towards reduction in the accessory flagellum, thereby suggesting that the presence of the ramus in those families is primordial.
Melphidippidae have the structure of gammarids in which the coxae have become shortened, and the gnathopods enfeebled lke females of the gammarid genus Cheirocratus, plus an elongation of uropod 3. Cephalic ocular bulges and a strong trend towards reduction of the accessory flagellum are characteristic.
Vitjazianidae have the general appearance and ecology of some pelagic eusirids but their retention of a 3-articulate accessory flagellum and the conjoint base of the primary flagellum indicate a mode of evolution different from that of eusirids and thus the Vitjazianidae are maintained in the gammarid families even though they may be more advanced than eusirids or gammarids. Gnathopod 1 has become simple and the most specialized vitjazianids have extremely reduced coxae. The mouthparts maintain a basic gammarid structure.
Hyperiopsidae form another pelagic theme. The typical genus resembles hyperiid Amphipoda but retains the maxillipedal palps; the second genus, Parargissa, differs so remarkably in its overall appearance that it is debatably an hyperiopsid but mouthparts, antennae, and pereopods seem to confirm the relationships between
285-135 O - 69 (Face p. 32)
ere Oe pereopods glandular Dy,
Isaeidae vy
ettiidae
Aoridae
Ampithoidae
ee U3 a
Tschyroceridae
hotidae
rial Dy
Metoediceros
Kuriidae
| Eophliantidae Ee body cylindrical
Corophiidae
=
Cheluridae Lysianassidae
F egcee
A-1
Phliantidae body depressed
coxae splayed Lidymocheila
285-135 O - 69 (Face p. 32)
BS aexe SES] —— Pardaliscidae C Melphidippidae F eS Sei Cc U-3
v
(Leese a ere 1 a er, fh as ) M BASIC GAMMARIDEAN Bis Oedicerotidae
ees os
aot WS i) pa!
Argissidae Be
a
Beaudettiidae pereopods glandular UN
Ca Isaeidae tf U-3 Talitroidea 6) M
Dogielinotidae
fossorial \
Gammaridae
Haustoriidae — Phoxocephalidae
Synopiidae
Aoridae
Vitjazianidae
Ampithoidae
a US x
= Cal
=
Metoediceros Liljeborgiidae Ischyroceridae =O) Eusiridae -Calliopiidae Pleustidae eS Psevdamphilochus U-3 A
< G-l Paramphithoidae
|
Eophliantidae Ee body
cylindrical
Cheluridae ae eis
ar i Bateidae Va- if YALE Gn. 1 vestigial U C
J Astyridae
Corophiidae
| | Bie a | | | | | |
U-3 Co Sebidae, Pagetinidae
Stenothoidae, Cressidae
Thaumatelsonidae TID. f, Oe Se \
= =
Lepechinellidae Stilipedidae
GZe UV; *
Lysianassidae
oe
Acanthonotozomatidae
VAL
ag
Ampeliscidae
Phliantidae
Colomastigidae ; p . O a body depressed as G-2 body \ =! —> ly dep A-1 cylindrical wv 1% . s Oahisinen: P = sen clara! Didymocheile
=k
Grarx 1.—Pattern of evolution in the Gammaridea (see text for discussion). Symbols: A-1=antenna 1; B=conical mouthpart field; C=coxae 1-3; F=accessory flagellum; G-1 (or Gn. 1), 2=gnathopods 1, 2; H=head; M=mandible, arrows pointing to special features; O=accessory flagellum absent; P=Maxilliped: circles=plates, appendage = palp; p-l, 5=pereopods 1, 5; T==telson, fleshy kind, lateral view; U=urosome; U-2, 3=uropods 2, 3; X=maxilla 1.
(UP Ph
Cy ares eee
MARINE GAMMARIDEAN AMPHIPODA 33
the two genera. Again, some resemblance to the eusirid grade of struc- ture is seen in the elongate enfeebled gnathopods characteristic of some calliopuds, but gnathopod 1 is almost completely simple, the accessory flagellum is 3-articulate and elongate as in some vitjazianids, and, furthermore, the palp on one member of the first maxillae is specially modified, bent and scaly. Article 4 of pereopods 1-2 is enormously elongate, thus giving to the pereopods a strong raptorial function apparently surrendered by the gnathopods.
Argissidae fall as a sidebranch of the gammarid families into the eusirid grade of structure by reduction of the accessory flagellum to 2-articles. Such reduction by itself does not imply any direct rela- tionship to those eusirid families for some marine and nonmarine Gammaridae also have a reduced accessory flagellum; such reduction occurs repeatedly in other completely distinct groups (e.g., isaeids). The peculiar coxal morphology of argissids (fig. 43) is not fully unique, for an analogous condition occurs in a gammarid genus, Megaluropus. Enfeeblement of gnathopods is again a feature of argissids but the quadrilocular eyes of oculate members seem significant. Except for those eyes, the diagnostic characters of argissids, though unique to- gether, are drawn from diverse members of the Gammaridae.
Pardaliscidae are a very difficult puzzle. They are perhaps the most aberrant of the gammarid families, although synopiids rival them. Once the principle has been established that the presence of a well- developed accessory flagellum probably marks a gammarid family, or a noneusiridean branch, the Pardaliscidae are to be kept within the gammarid block. But they are characterized by foliation of the man- dibles and loss of molars, the frequently occurring conjoint condition at the base of the primary flagellum on antenna 1, feeble gnathopods, progressively reduced coxae, and one or more peculiarities of the maxil- lipeds: reduction in overall size of the inner plates, often a reduction in the outer plates, and, occasionally, an elongation of the article carrying the outer plates. The lower lip often has the inner lobes coalesced and forming a convex bridge between the outer lobes. A generalized view of the maxillipeds suggests affinities with the Lil- jeborgiidae in which the maxillipedal palps, like those of the Par- daliscidae, are relatively dominant over the basal plates. The mandibles of Liljeborgiidae show a strong tendency to a complete loss of the molars but liljeborgiid gnathopods are very powerful and the lilje- borgiid with the smallest coxae has larger coxae than any pardaliscid. The lower lip of liljeborgiids could be precursive to that of pardaliscids.
The powerful gnathopods of the Liljeborgiidae are reminiscent of those in the Eusiridae (Rhachotropis) but most lijeborgiids have a strongly developed accessory flagellum and all have a reduced, non- triturative molar. Some eusirids also have this reduced molar. The
34 U.S. NATIONAL MUSEUM BULLETIN 271
EKusiridae seem to be so broadly polyphyletic that there is cause to investigate the interrelationships of liljeborgiids with some of the eusirids. Several grades of structure are congruent between the Liljeborgiidae and certain members of the Pleustidae. The labia of the two groups are very similar and the mandibular molars of pleustids often resemble those of lujeborgiids. Pleustid gnathopods are often enlarged but the family differs from liljeborgiids in their uncleft telson and vestigial accessory flagellum. The outer rami of uropods 1-3 are shortened in Pleustidae and the outer ramus of uropod 3 is uniarticu- late but some liljeborgiids approach these conditions. The pleustid rostrum is a development restricted to a few members only.
Synoptidae are retained in close proximity to the basic gammaridean in view of their elongate accessory flagellum, even though several of their members have the articles reduced to two. Gnathopods are feeble. All but a few synopiids have the head enlarged in relation to the shortened pereonites 1-3. Coxa 3 dominates coxa 4 in most of the genera, and in all but one genus, article 3 of the mandibular palp has become very short, almost vestigial. The mouthparts are otherwise basic except for those genera having the molars extremely enlarged and velvety smooth. This semifossorial condition has also occurred in some haustoriids even closer to the Gammaridae than the Synopiidae. Uropods have undergone the eusirid shortening of the outer rami but uropods 1-2 are far more specialized in the frequent sublamellar con- dition of the inner rami. In all but Synopia, the telson has become elongate. Eyes when present are dorsally coalesced or contiguous.
Oedicerotidae are another group, like the Synopiidae and Parda- liscidae of generally obscure relationships. They have reached the eusirid grade of structure in accessory flagellum, resemble the Syn- opiidae in head and eyes but the telson is a short, thin, ovate or truncato-ovate, uncleft lamina, unlike that of most Synopidae. Uropod 3 has become fully elongate, a tendency seen in several synopiids but the pereopods have become strongly fossorial, the first four pairs by virtue of their long setae and the fifth pair by virtue of its immense articular elongation. On the average, gnathopods are of medium size, thus being more powerful than those of synopiids. Oedicerotidae have a strong resemblance to isaeids especially in pleonal epimera and pigmentation but lack pereopodal glands and have a thin, nonfleshy telson.
Three families, each of them highly distinct from one another, seem to have strong relationships to the Eusiridae-Callioptidae-Pleustidae complex. The least distinct is the Paramphithoidae, a group character- ized by acuminate coxae; several intergrading genera suggest that Paramphithoidae might be incorporated within the broader eusirid amalgam. The Laphystiopsidae carry calliopiid tendencies to a loss of
MARINE GAMMARIDEAN AMPHIPODA 35
molar grinding ridges, miniaturization of coxae, and simplicity of gnathopods to their definitive extremes. Bateidae have the cephalic pleustid aspect but have undergone strong divergence through reduc- tion of gnathopod 1 to a single article or two.
Paramphithoidae have links to more advanced families, either directly or indirectly. Generic intergradations between Paramphi- thoidae and Astyridae are readily apparent (see those families for elaboration). The Lepechinellidae have the acuminate coxae of par- amphithoids, but through coalescence of two urosomal segments resemble a parallel grade, the Atylidae. Their origin is thus plotted in graph 1 as a mixture of characters between Atylidae and Paramphi- thoidae, an unsatisfactory procedure indicating not phyletic affinities but gradational structure.
The body plan of the Stilipedidae conforms remarkably to that of the Astyridae. If the pardaliscid left mandible, adopted by the Stilipedidae, is a character of major evolutionary significance, then one might consider that stilipedids stand close to the Pardaliscidae. But the astyrid mandible is in a condition precursive to the pardaliscid right mandible. It is partially flattened and grossly toothed while retaining a nontriturative molar. It thus resembles the pardaliscid right mandible whereas the stilipedid mandibles resemble the broad- ened untoothed left mandible of pardaliscids. Since pardaliscids combine both kinds in a single individual, it does not stretch one’s imagination to consider that the astyrid mandible evolved into that of stilipedids. That astyrids did not also produce pardaliscids is attested to by the strong accessory flagellum and weak coxae of pardaliscids.
Atylidae have the same general advancements of the eusirids but the fusion of two urosomites marks a line of evolution which some students suggest has led to the inquilinous Dexaminidae lacking mandibular palps; this condition naturally leads to the palpless Prophliantidae in which all urosomites have become coalesced.
A more radical inquilinous specialization in the acanthonotozomatid line seems to have its gravitational balance near the Paramphithoidae. Here a marked change in the mouthpart field from its basic quadrati- form bundle to a conical or triangular field suggests increasing stages towards the functions of piercing and sucking. The individual mouth- parts increasingly become better adapted for those habits in the various acanthonotozomatid genera through incisorial styliformity and reduc- tion of molars; but most acanthonotozomatids live in south polar waters, often reach giant body proportions, and, in many cases, the mandibles have become broad, flattened, and very powerful as if their bearers have evolved toward a state of subpredation, perhaps on giant sessile organisms such as sponges. They might be called predatorial grazers if this habit is confirmed.
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Direct advancement to the Lafystiidae through reduction in maxil- lipedal palps is a consequence of acanthonotozomatid specialization and the ultimate is reached in the Ochlesidae, technically hyperiids, because of the complete loss of palps.
Stegocephalidae are the only other major group of Gammaridea with the strong piercing and sucking mouthpart field of acanthono- tozomatids and even though their coxae are not as acuminate and their mandibular palps and molars have been lost, there is still merit in considering a direct relationship between the two families. Stego- cephalidae have the broad, sublaminar mandibles of several acantho- notozomatids but may have originally evolved as a pelagic group, some members having returned to the benthos in later stages. The breadth of the lateral shield appears to have a relationship to midwater suspension but acumination may still be seen in the anterior coxae. The benthic members may also be predatorial grazers and the only known raptorial predator in the benthic Gammaridea is a member of the Stegocephalidae.
Another line of evolution favoring inquilinous behavior is that commencing with the Amphilochidae and Leucothoidae. Early stu- dents of the Gammaridea noticed similarities between Liljeborgiidae and Leucothoidae in maxillipedal structures; other resemblances such as retention of enlarged gnathopods are so clear that one might say that leucothoids are liljeborgiids in which the accessory flagellum has become vestigial and gnathopod 1 has been transformed into its fully carpochelate condition while the outer plates of the maxillipeds became vestigial. The rudiments of the carpochelate gnathopod may be seen in gnathopod 2 of leucothoids, thus resembling the gnathopods of liljeborgiids. Anamixids carry the inquilinous state to the ultimate by the transformation of mandibles and maxillae into a piercing keel.
Amphilochidae may stand almost completely alone. Their mouth- parts strongly resemble those of liljeborgiids and their gnathopods are usually miniaturized editions, like those of Listriella. But their heads have the appearance of the pleustid-paramphithoid or bateid line. The primary mark of their advancement is the reduction of coxa 1, not. as fully reduced as in the Bateidae. This suggests their direct precursorial relationships to the stenothoids. The peculiar Pseudam- philochus, through its cleft but ovato-acuminate telson, unreduced coxa 1, large rostrum and nonelongate peduncle of uropod 3, stands among the Amphilochidae, Pleustidae, and Liljeborgiidae. Schellen- berg (1931) suggested that it should be assigned to a unique family, the Pseudamphilochidae.
All five families of the stenothoid complex seem to have strong interrelationships by virtue of the mandibular form (see figures) in which at least one mandible has a box-like shape with deeply serrate
MARINE GAMMARIDEAN AMPHIPODA 37
incisor, a lacinia mobilis formed of a thin lamina resembling the incisor and nearly appressed to it, plus a molar bulge clearly moved distal- wards toward the incisor and nearly or partially encroached upon by the spine row. One group of stenothoids has the amphilochid coxa 1, whereas the other has the leucothoid (normal) coxa 1. Sebid enathopods even vaguely resemble gnathopod 1 of leucothoids and numerous other similarities are apparent. Stenothoidae, Thaumatel- sonidae, and Cressidae, those with amphilochid coxa 1, are very closely related among themselves, even though some extremely pygidized species have been used as types of the Thaumatelsonidae and Cressidae. The five stenothoid families are obviously inter- related through the condition of uropod 3. It is uniramous, pre- sumably through loss of the inner ramus, as the remaining ramus is basically biarticulate. Neither the Leucothoidae nor Amphilochidae show a tendency to this condition, except perhaps for the universal genus Pseudamphilochus. If the Amphilochidae were the precursors of the Stenothoidae and the Leucothoidae were precursors to the Sebidae, then the evolution of a common uropod 3 had to occur twice.
The superfamily Talitroidea, comprising originally the Talitridae, for nearly a century have been considered as extremely distinct gammarideans, often worthy of even subordinal rank. They are more diverse than they ever have been if one were to add the Dogielinotidae and Kuriidae to their ranks and to suggest that Najna and Metoediceros represent types of new talitroid families. Bulycheva (1957) split the Talitridae into three families, adding the Hyalidae and the Hyalellidae. The three main families are here treated as a superfamily, mainly for convenience in identification. Talitroids have no universally unique characters; several other families have a uniramous uropod 3 and no mandibular palp but among the genera of talitroids are many unusual morphs. Often the cephalic sclerites are clearly marked. Numerous enathopodal, pleopodal, and antennal modifications occur in terrestrial genera; the jumping ability is extreme in those genera. The marine members appear more regularized, some even having vestigial inner rami on uropod 3. Perhaps the structure of fringing setae on the female brood lamellae will prove to be characteristic of the group. Although they may be considered as very distant from the basic gammaridean, their singularity is damaged by discovery of the Beaudettiidae. That monotypic family is composed of a species with clear relationship to Elasmopus, a member of the Gammaridae. Through loss of mandibular palp, reduction of the inner ramus of uropod 3, and telsonic modifica- tion, Beaudettia has come close to the talitroidean grade of structure. Morphological distance of talitroids from gammarids, as a reflection of time or extensive genetic change, is thus lessened by our observation of the revolutionary changes that have occurred in Beaudettia, possibly in relatively recent times.
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The final manifold group in graph 1 is the greater isaeid complex that might be accorded superfamily rank. The primitive isaeid is conceived of as a morphological analogue to the basic member of Gammaridae. Until one examines the fleshy telson and pereopodal glands of primitive isaeids, one is struck by the great similarity of generalized Gammaridae and Isaeidae (—Photidae). Isaeids have become very diverse in many of the same ways as have the gammaridan stock but no highly advanced inquilines have appeared, unless one can link up some of the eophliantids; they instead appear to be lignivores. Even so, the piercing-sucking groups like acanthonotozo- matids, the coelenterate loving groups like the stenothoids and amphilochids, the spongicolous and protochordate inhabiting kinds have not evolved within the isaeid complex, perhaps because they have been primarily adapted to form domiciles of their own out of the pereopodal spinning glands. Identification and relationships are clouded by the frequent loss of those glands in the Podoceridae, some Corophiidae, the Cheluridae, and even the nomenclatorial type, Isaea. A more basic example of the isaeid line is Gammaropsis and its name should ideally be the root of the stock.
The most advanced members of the isaeid group have accumulated two morphological changes, either reversal in gnathopodal domination or development of a partially to fully rigid urosome with loss of uropodal structures. The gnathopodal reversal may be related to a stronger than normal cephalic orientation required of organisms living in tubes open to the anterior end of the animal body. Rigidization of the urosome may assist the organism in maintaining a position within the tube. Some of these tube dwellers have even returned to making burrows (?internally lined) in the substrate.
Perhaps the Podoceridae have come closest to the inquilinous func- tion; their ecology and morphology are poorly known but there has developed the impression that all their members have lost the spinning glands and that many of them are strongly associated with hydroid colonies, as if they were predatorial browsers.
The stability and similarity among the mouthparts of the members of the isaeid complex leads one to the view that podocerids belong with the group. They are frequently mentioned as the root stock of caprel- lids; indeed, since the time (Dec. 1965) that the contents of this paper were formed, a new family, Caprogammaridae (Kudrjaschov and Vassilenko, 1966) has been described with further strong evidence of podocerid-caprellid relationships.
Distinctions between Isaeidae and Aoridae are not clear except that enathopod 1 of aorids is either larger than gnathopod 2 or is that member having sexual dimorphism. The Corophiidae seem to be a polyphyletic group of aorids and isaeids with pygidization. They may
MARINE GAMMARIDEAN AMPHIPODA 39
include some ischyrocerids also. The Ampithoidae are fairly uniform by virtue of their third uropods and the Ischyroceridae, though anal- ogous to ampithoids because of the development of a rudimentarily uncinate condition on the outer rami, are recognizable by the elonga- tion of the peduncle. Ischyroceridae rarely have the reversed gnatho- podal domination. One corophiid, Hricthonius appears to combine features of Corophiidae, Aoridae, and Ischyroceridae, but it is not the only “‘isaeid” genus that gives trouble to the systematist.
The lignivorous Cheluridae have numerous resemblances to the isaeid-corophiid line, but the fully pygidized urosome reveals suture marks indicating that urosomite 3 has become extremely enlarged, a feature unique to this family. There is no better demonstration of the extreme genetic potential of numerous Amphipoda in replicating long lost structures than by considering the enormous, Maera-like enathopod 1 of Chelura insulae Calman.
Families without clear relationships to others are the Lysianassidae, Phliantidae, Colomastigidae, Ampeliscidae, and potential families are represented by Didymocheila, Ceina, and Biancolina.
The Ampeliscidae are a very advanced group, apparently completely divorced from the isaeid complex, which have developed pereopodal glands and spinning tubes of a different form from those of isaeids. Ampeliscid morphofunction is also discrete, as far as we know. Ampeliscids have a mixture of characters represented by argissids and atylids but those two families otherwise bear no relationships. Thus, the line on graph 1 connecting Ampeliscidae to Argissidae and Atylidae represents only a focus on structural grades.
Despite similarities to the Sancho-Chosroes members of the Calliopi- idae, the Phliantidae bear relationships to certain dorsoventrally de- pressed podocerids, like Podocerus and I am inclined to the view that the latter suggestion has great merit and should be examined carefully. Phliantids also have about as much in common with Laphystiopsidae (mostly dorsoventral depression) as they do to Sancho and Chosroes. Apical curls on brood lamellar setae plus many similarities in buccal and urosomal parts suggest phliantid affinities with Talitroidea.
Relationships of the Colomastigidae are difficult to trace. The body form is subcylindrical but the head is not of the spheroid kind found in the Eophliantidae, the rami of uropod 3 are present and elongate, the peduncle is elongate and the mandible apparently lacks a true incisor, that characteristic having been replaced by an enlarged spine row. Affinities with leucothoids are seen in maxillipeds, uropods, and telson.
The Lysianassidae are almost as fully diverse as all of the other Gammaridea put together and comprise about 20 percent of all marine genera and species of the suborder. They are united together by their clearly recognizable gnathopod 2 and furthermore share an almost
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unique antenna 1, with very short and often telescoped articles 2-3. Otherwise they have radiated into species that exhibit all forms of inquilinous specialization such as piercing and sucking mouthparts, coalesced urosomites, reduced uropod 3 and into numerous highly successful pelagonts. The usual nestling and burrowing members also occur in profusion. Some pelagic members are predatorial and some may have developed narcotizing glands. Even wood boring is a pre- sumed habit of some deep-sea lysianassids eating waterlogged coco- nuts. But none has developed the tubicoly of isaeids.
The morphologist cannot as yet recognize any clear links between the Lysianassidae and other gammarideans. There are vague resem- blances to Stegocephalidae, perhaps of convergence and suggesting that Lysianassidae may have developed, like Stegocephalidae, first as a pelagic group that later reinvaded the benthic realm. The obligatorily pelagic genera among the Lysianassidae are very highly specialized but many unspecialized lysianassids of other large genera (e.g., Orchomene) occur prominently in the nekton. Elongation of article 3 on gnathopod 2 is not fully unique to the Lysianassidae, as seen in section B of the written key to families (p. 109). The functional morphology of this lysianassid appendage, with its other unique characters, should be a problem of major concern to gammaridean evolutionists. It is not grossly dimorphic in the two sexes and seems wholly inadequate, like gnathopod 2 in so many other families, of serving as a sexual grasping appendage. Male gnathopod 2 does serve this function in those few Amphipoda that have been observed in amplexus.
Limnetic Evolution of the Gammaridea
That Gammaridae collectively represent the most primitive mem- bers of the Gammaridea and of the living Amphipoda, has some support by knowledge of the occurrence of an enormously diverse faunule in Lake Baikal. Nearly 250 species of Amphipoda live in that Siberian lake (see Bazikalova, 1945; Dybowsky, 1874). One might presume from the extent of radiation that has occurred in the Baikalian gammarids that they entered the lake shortly after its formation in the Triassic. All Gammaridea now living in the lake are classified with the family Gammaridae even though there is justification in re- garding some of the genera, like Hyalellopsis, as members of novel families. Baikalian Gammaridae strikingly reflect many of the morpho- logical conditions now allotted familial importance in marine amphi- pods. Non-Baikalian epigean and troglobitic amphipods also show some of these conditions. The diversity of Amphipoda in Baikal is an enigma in the sense that the human observer has the tendency to allot a great deal of time for the evolution of such a rich display of
MARINE GAMMARIDEAN AMPHIPODA 4]
TABLE 5.—Freshwater or inland saline water Gammaridea .
Family and Locality Genera Species
Gammaridae
Lake Baikal 37 230+
Palearctic—Nearctic 65+ 400+
South Africa 2 11
Australia-New Zealand 5 (+3+) 25+ Hyalellidae
Neotropical 1 28
African 1 iL
Australia-New Zealand 2 5
Miscellaneous 1 4 Bogidiellidae
Tethyan shores 1 9 Hadziidae
E. Tethyan shores 1 1 Caspiellidae
Caspian Sea 1 1 Corophiidae
Tropical—Cold-temperate, mainly river mouths 4 15 Calliopiidae
Tropics—subtropics 2 5 Haustoriidae
Cold-temperate 1 1 Eusiridae
Japan, Australia 2 3+ Metoediceros
Antiboreal 1 1
Totals 127+ 740+
morphs whereas the absence of a solid pre-Miocene fossil record leaves us no confirmation of the group’s antiquity. Those Miocene fossils are amber enclosures of species presumably still alive in European freshwaters. One may become enmeshed in circular thinking that the family Gammaridae must be primitive and the suborder Gammaridea very old because not only is Baikal very old but that Gammaridae came to dominate it and, therefore, Gammaridae were on the scene first as well as very early. There are no current facts to counter that argument but one has to maintain a reserve based on logic. Perhaps only Gammaridae are widely preadapted to freshwater conditions. Only the Hadziidae, Caspiellidae, Bogidiellidae, Hyalellidae, and a few members of other families (table 5) also occupy freshwaters (or inland waters). Gammaridae dominate the freshwaters of the world, except for the Neotropical realm where the distantly related Hya- lellidae replace the Gammaridae. The other minor families mentioned above probably are closely related to the Gammaridae.
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The cold-water orientation of marine Gammaridea and the diversity of brackish water and marine species (the Anisogammarus complex) in the north Pacific Basin, with affinities to Baikalian genera, also sup- port the thesis that Gammaridea may have had a fresh-water origin in the vicinity of Baikal and then invaded the sea. The thesis becomes stronger if one confines the theory only to the living members of the family Gammaridae, for that family is generically impoverished not only in tropical shallows but in all deep-seas of the world. Numerous species of Maera, Elasmopus and Ceradocus are prominent in the tropical sublittoral but generic radiation is comparatively low. Isaeids and hyalids also dominate tropical shallows.
The importance of limnetic environments to the development of Amphipoda is also attested to by the species-flock of Hyalellidae in Lake Titicaca, a lake now of alpine character in the tropics but a lake perhaps of warmer climes in earlier times. It seems more than hap- penstance that most other talitroideans (like the Hyalellidae) are now strongly oriented to the tropics. They include the fully (and only) terrestrial Amphipoda that occur mainly on the islands of the Indo- Pacific region, plus the beachhoppers that also have primarily tropical affinities. Only the aquatic genus Hyalella has successfully invaded the Nearctic realm dominated by Gammaridae (which also dominate Palearctic and occur in African (Ethiopian) realms. Talitroideans are more specialized or more advanced morphologically than members of Gammaridae in uropods, mandibles, and antennae and no one has been able to trace any direct relationships between the two groups, even though one may see in the Beaudettiidae a species presumably derived from Elasmopus (Gammaridae) that has undergone a revolu- tionary change in the direction toward Hyale (Talitroidea).
One may consider that by the time fresh-water Gammaridae had become dispersed to a cordilleran pathway contiguous to South America that the latter continent had become disjunct or that the cir- cumferential Sonoran belt afforded a major barrier. There may be a strong competitional stress between Gammaridae and Hyalel- lidae for the northward migration of one hyalellid into the Nearctic realm does suggest the presence of a migratory pathway now in exist- ence, but only one species has managed to make the crossing.
Dispersal of marine Gammaridae to the Neotropical realm also must have come after the successful invasion by hyalellids, because it would appear that even in tropical waters marine gammarids have been able to invade limnetic environments.
All through the ancient Tethyan Sea marine gammarids invaded underground waters, forming the nipharigid and other troglobitic groups. The South African genus Paramelita may have its origin in marine melitids and have undergone convergence towards the Baika-
MARINE GAMMARIDEAN AMPHIPODA 43
lian gammarus stock, for Schellenberg (1937a,b) has included that genus with the ‘“gammarus” group. If South African ‘‘gammaruses’’ and the poorly known Australian members of the family have indeed been the results of separate fresh-water invasions then the southern hemisphere was completely blocked from an overland dispersal of Kurasian gammarids.
These stocks of ‘‘gammarus” in the southern hemisphere may not be so successful competitively against talitroideans as are Palearctic stocks, for another group of talitroideans, the ‘‘chiltonias’’ have evolved in the African and Australia-New Zealand provinces.
One need not adhere too closely to the thesis that Baikal formed an evolutionary center for the reinvasion of the sea by its products. Baikal and Titicaca may simply be dismissed as habitats open to whatever contiguous marine groups were present. But this also implies, without considering special ecological adaptations of the groups, that if Gammaridae are indeed primitive and represent the survivors of a base stock out of which came talitroideans, that the latter were under- going their evolution on a tropical frontier while the tropicwards dispersal of various marine Gammaridae was stagnating. Talitroids, therefore, penetrated the tropical barrier in the move southward be- fore marine or fresh-water gammarids. The importance of studying ecological stress in the groups is apparent in the limited “‘later’’ success of marine gammarids in partially occupying some of those southern limnetic habitats on the tropical fringe. If there was temporal inde- pendence among all these events in Baikal, South America, South Africa, and Australia, and if marine Gammaridae and Talitroidea were already fully developed before any of these limnetic invasions, then one must entertain a strong and independent ecological success of the Palearctic-Nearctic gammarids, for talitroideans have not been able to populate those realms.
Terrestrial nonaquatic talitroids have never become eminently successful. They occupy no major continent except where they have been locally introduced into gardens. They are confined to Indo- Pacific islands in the tropics, in moist, biotically impoverished environ- ments. Beachhoppers on the strand of large continents have never been able to migrate inland as far as we know.
Only two groups of Amphipoda thus have any clear relationship to historical events that we have as yet been able to discover and on which we may devote a great deal of refinement with promise of results other than speculation. The meager evidence seems to indicate that Gam- maridae had their major radiation in cold northern climes, either marine or fresh water and that they had not reached either realm of the southern hemisphere before those continents were blocked by some kind of environmental barrier, leaving them open to the Talitroidea.
285-135 O - 69 - 4
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The question of interest is whether the Talitroidea existed simul- taneous with these events or whether they presumably came much later, and indeed whether they evolved directly through the gammarid funnel; whether modern Gammaridae truly represent a base stock to all other Gammaridea or whether they are simply a blind-alley sidebranch coincidentally representing today the logical primitive gvammaridean.
That Gammaridae and all other Amphipoda are heavily oriented to cold water seems more than happenstance. We may determine eventually that the tropics have the most numerous species of Am- phipoda of any province but there is little current evidence that we shall also find tropical genera to be the most numerous. Cold waters (but not high polar) undoubtedly have the greater diversity at the higher taxonomic levels. It is occasionally, or at least not always, true that biotic groups have their highest diversity in their centers of origin and I do not wish to imply that diversity and centers of origin are correlated in the Amphipoda. Amphipoda have a bimodal diversity anyway as cold waters obviously occur in two circum- ferential bands. But the facts that the most primitive morphotype has its center of diversity in cold Siberian Seas, that Gammaridae have strongly radiated generically only in a cold Siberian lake and not in isolated tropical lakes, that Nearctic-Palearctic fresh-water Amphipoda clearly are more diverse generically than in tropical fresh waters, that marine gammarid genera are most numerous in cool waters, that a whole suborder of amphipods, the Hyperiidea is confined primarily to cool pelagic waters, that the largest family of marine amphipods, the Lysianassidae is confined largely to cool waters, all suggest that Amphipoda are preadapted to cool waters and have undergone the major “post-amphipodan”’ radiation in such climes.
Identification Procedures
The identification of a gammaridean amphipod even at familial levels so often requires a complete dissection and analysis of all appendages and mouthparts that the procedure is considered to be mandatory. This handbook cannot be utilized successfully by a non- specialist without dissecting appendages (Appendix I) and observing minute characters (checklist of Appendix II). Once the student has gained some experience, however, portions of these procedures may be skipped, for some families and many genera can often be recog- nized without extensive analysis.
The procedures for identification described herein are manifold. They include the memorization of a basic gammaridean plan, illus-
MARINE GAMMARIDEAN AMPHIPODA 45
trated in figures 1 and 2 and diagnosed on p. 50. Almost all other families are defined by combinations of characters expressing either minor specializations or minor simplifications of the basic gammari- dean plan.
THE DIAGRAMMATIC KEY TO FAMILIES.—This key (figs. 1-53) is composed of 54 boxes, 51 of which represent a family or superfamily containing one or more illustrated characters in solid lines that dis- tinguish it from the basic gammaridean (figs. 1-2b). For example, if mouthparts are not illustrated they either resemble those of the basic gammaridean or are so highly variable that they have no diag- nostic differences from the basic gammaridean and are thus not mutually exclusive. Each of these concordant families is noted in the captions of the illustrated key and character differences in broken lines are figured and explained where necessary. The diagrammatic key is, therefore, not an absolute endpoint, for the taxonomist must also check the illustrated boxes of families resembling the first pro- visional identification.
The diagrammatic key is arranged in a way to deal first with those families which have highly characteristic and easily observed morphol- ogy. Thus, Lysianassidae (fig. 3), with their characteristic gnathopod 2 are presented first, followed by the similar Stegocephalidae (fig. 4), which have characteristic coxae, head, mouthparts and mandibles, followed by Ochlesidae (fig. 5) without maxillipedal palp, Lafystiidae (fig. 6) with 2-articulate maxillipedal palp, and Acanthonotozomatidae (fig. 7) with characteristic coxae. Following those groups is a pair of families with cylindrical bodies, succeeded by seven families with uniramous uropod 3. Succeeding that point (Amphilochidae, fig. 19) all families have a biramous uropod 3, rarely with exceptional genera or species. Other characters are abnormal to the basic gammaridean, however, such as the small coxa 1 of Amphilochidae and some Leuco- thoidae and the characteristic gnathopod 1 of Leucothoidae and Anamixidae. There follows a group of families with various combina- tions of coalesced urosomites, and degraded mandibles or acuminate coxae. Commencing with the Oedicerotidae (fig. 27) the identifications become increasingly difficult as the differentiations from the basic gammaridean become less apparent: the Oedicerotidae have elongated pereopod 5 and short, uncleft telson; Paradaliscidae (fig. 28) have characteristic mandibles; Bateidae (fig. 30) have degraded gnathopod 1; Synopiidae (fig. 31) have massive, rostrate heads; the next four families (figs. 32-35) have various combinations of feeble gnathopods, degraded mandibles, special shapes of lower lips and maxillae and various telsonic lengths and clefts; the next three families differ from the basic gammaridean by poorly cleft telsons and degraded accessory flagella; the next group has cleft telesons but degraded accessory
46 U.S. NATIONAL MUSEUM BULLETIN 271
flagella; then a group with normal accessory flagella but with various combinations of degraded molars, characteristic rostra, coxae, and pereopods, until finally the Gammaridae are reached.
After the Gammaridae is the group of isaeid-like families, most of which form tubes, have spinning glands in the pereopods, bear short, uncleft fleshy telsons, posteriorly unexcavate coxa 4, and normal mouthparts but which often have increasingly complex or reduced uropods difficult to separate from the network of families anterior to the Gammaridae. The identifier must keep them in mind. Some genera of the isaeid-like families are very difficult to recognize and all of these family keys and diagnoses should be examined when identifying presumed Corophiidae, Aoridae, Isaeidae, Ischyroceridae, Ampithoidae, and Podoceridae. Perhaps it is wise to become familiar with known variations in characters of these families for it is often difficult to recognize spinning glands in pereopods (and some genera of isaeid-like families lack them). Shallow-water isaeids are usually strongly pigmented in browns and purples but this is not a reliable character, because other families occasionally have strongly pigmented species.
Example 1: Perhaps the observer determines from the completed checklist (Appendix II), that the characters of the box Liljeborgiidae (fig. 41) fit the specimen being identified. All characters match those of the basic gammaridean except for the mandibular molar, which is nontriturative; the accessory flagellum is multiarticulate, the mandible has a 3-articulate palp, the maxillae are normal, maxillipeds have well developed lobes and 4-articulate palps, gnathopod 1 is of normal or enlarged size and is subchelate, gnathopod 2 is enlarged and sub- chelate, all pleonites are free, pereopods are generalized, uropod 3 is large and biramous, and the telson cleft. The caption lists related families or those families with which Liljeborgiidae might be confused and their boxes should be examined for special combinations of char- acters not shared with the specimen at hand. For instance, the Liljeborgiidae caption lists Gammaridae and Eusiridae as congruent families, but the box of Eusiridae (fig. 39) indicates (1) by the absence of a drawing of the mandible that it is either like the basic gam- maridean and therefore has a triturative molar, or is of variable char- acter; and (2) that the accessory flagellum is 0-2 articulate. If the specimen being identified has an accessory flagellum of 3-plus articles then it is not a eusirid; if it has a 2-articulate accessory flagellum and triturative molar then it is not a liljeboriid; if it has a 2-articulate accessory flagellum and a nontriturative molar it may be either a eusirid or liljeborgiid and the reader would turn to the descriptions of those families (p. 213 and p. 291) and compare them with the speci- men. He would read the sections on Relationships to discover that a
MARINE GAMMARIDEAN AMPHIPODA 47
few genera of Eusiridae also have a nontriturative molar and thus resemble Liljeborgiidae; but those genera lack an accessory flagellum entirely; furthermore their gnathopods are unlike those of Liljeborgiidae.
Example 2: Eusiridae (fig. 39) differ from the basic gammaridean only by the reduction of the accessory flagellum to 0-2 articles shown in solid lines and by the elongation of the telson. Also figured in the box is the lower lip in broken lines to show a distinction from the Pleustidae and the Astyridae; uropods 1 and 2 in broken lines show a distinction from the Liljeborgiidae and the illustrated telsons differ from those of the Calliopiidae.
Thus, the vestigial eusirid accessory flagellum is shared with many other families but the special combination of basic gammaridean morphology plus the telson, uropods, and lower lip is distinctive.
Also, an inference is made on the caption that the student refer to the figures of the Vitjazianidae (fig. 34), and the Oedicerotidae (fig. 27) for characters distinguishing those families from the Eusiridae, i.e., the simple first gnathopod and conjoint primary flagella of antenna 1 in the Vitjazianidae and the disproportionately long fifth pereopod in the Oedicerotidae.
Not mentioned are other deviations, such as the occasional reduc- tion of mandibular molars and 1-articulate first maxillary palps. Several other families share these characters and may be partially defined by them, but those families may be distinguished from the Eusiridae by more important characteristics.
Example 3: Calliopiidae (fig. 38) differ from the basic gammaridean only in the reduction of the accessory flagellum to 0-1 article and the coalescence of the telsonic lobes. Also illustrated are the lower lip which differs from that of the Pleustidae, coxa 4 to show a distinction from the Isaeidae (= Photidae) and telsons to show faint distinctions from the Eusiridae. Some calliopiids have isaeid-like coxa 4 but their telsons are not fleshy and their pereopods lack glands.
Example 4: The diagnosis of Amphilochidae (p. 132) includes only: ‘“‘Accessory flagellum absent; coxa 1 very small, partially hidden by following coxae.”’ Next is stated ‘‘See [the family] Pleustidae. .. .”’ The description of that family implies that coxa 1 is ‘‘normal’’ in size and placement. In sequence, the description of the mentioned Steno- thoidae should be examined to see that uropod 3 is uniramous and therefore distinct from uropod 3 of the Amphilochidae.
The Diagrammatic Key to Families is, at best, a method of narrow- ing the search for a familial identification to a few possibilities, each textual diagnosis and description of which must be examined for good- ness of fit. A flexibility has been maintained in the diagrammatic key because of space limitations in presenting a compact visual impression
48 U.S. NATIONAL MUSEUM BULLETIN 271
of a group as well as the possibility that the observer will handle undescribed genera and families. The need to cross-check diagnoses and test keys to genera in several families is a part of the identification process.
The textual diagnoses, descriptions and relationships of the 53 families are presented in alphabetical sequence. The Hyalidae, Hyal- llidae, and Talitridae are combined into the superfamily Talitroidea for ease of identification. Each family has a written key to its genera and each family is accompanied by several sets of figures illustrating: (1) the characters of the generic keys, and (2) the variation in lateral aspects and appendages of the several genera.
Perhaps the key to success in using the diagrammatic keys is the need to memorize the plan of the basic gammaridean and to keep an account of generalized characters, especially the generalization of heads, feeble and strong gnathopods, anterior coxae, and elongation of telsons and pereopods. Thus, Synopiidae are extremely difficult to recognize because the subtle shapes of their heads are primary to their identification. Acanthonotozomatidae, Paramphithoidae, and Stego- cephalidae are recognized mainly by their anterior coxae.
WritTrEN KEys.—If the Diagrammatic Key to Families does not produce results one may utilize the written key to families (pp. 106-116). The written key is not as satisfactory as the Diagrammatic Key be- cause the observer must repeatedly make ‘‘yes or no” decisions on individual characters and no accomodation for undescribed genera and families can be made. Simplicity of the keys requires occasional repetition of taxa in order to account for morphological diversity.
Extraordinarily long keys are occasionally divided into sections in order to improve their usefulness. The establishment of subgroups in this manner is believed to be of assistance in retaining conceptual images while the observer is proceeding through the maze of an extremely complicated key.
DiaGnosrs.—Familial diagnoses are compatible with the Diagram- matic Key to Families and are not mutually exclusive, for the diagnoses only differentiate the families from the basic gammaridean. Inter- related families are listed as a part of the diagnoses and are discussed under ‘‘Relationships.’”’ These discussions are not necessarily repeated in all possible places and the reader may have to turn from the Synopiidae to the Liljeborgiidae, for example, in order to find a dis- cussion of the interrelationships between the two families.
Genera are diagnosed in as brief and consistent form as possible, in direct relationship to the keys. Where two genera are mutually dis- tinguished by a character of no taxonomic importance to other genera of a family, the alternatives of the character are italicized and the related genus is stated. Such character alternatives are omitted from
MARINE GAMMARIDEAN AMPHIPODA 49
the diagnoses of other genera. Character alternatives that are known as “combining characters,’’ because of their usage in keys, are occasion- ally listed in diagnoses and italicized; they reflect the necessity to state a general character that is diagnostic only because it occurs in unique combination with all other characters stated for the genus under question; the character is not necessarily restricted to the taxon in question.
Each diagnosis has as its framework the conditions known for the type-species or type-genus. In some families with few genera, the diagnoses are confined to the keys. No doubt, the range of variation permitted in the diagnoses is too narrow; time has not been allotted to analyze the literature of each species in each genus in order to widen the generic limitations because the literature is occasionally so defec- tive that one would waste time to ponder, analyze, and argue possible truths or to speculate on omissions. Many described species are un- doubtedly misclassified.
Brief synonymies of recently established families are given but other- wise such information may be found in Stebbing (1906) or J. L. Barnard (1958a).
Type-species of each genus and their modes of selection are listed below the generic synonymies. A modern reference, if available, is given for the type-species. The number of species is included, along with generalized distributional notes on the genus. ‘‘Littoral’’ includes sublittoral, to 300 m; ‘‘bathyal” includes depths from 300 to 2000 m (here) ; ‘‘abyssal’”’ exceeds 2000 m, and “‘hadal’’ 6000 m. Some species in depths exceeding 300 m are pelagic but have not been so designated because of uncertain data on collecting methods. Reference to terms such as ‘arctic, antarctic, boreal,’’ is very imprecise because no exact definitions are followed. If a genus is centered generally in the antarctic as well as the subantarctic the terminology is simplified to “antarctic.” The “arctic” includes the Norwegian Basin and its fringes as well as the polar basin. ‘“‘Biboreal’’ denotes occurrence in northern and south- ern hemispheres; ‘‘amphiboreal”’ denotes occurrence in both oceans of the northern hemisphere.
Occasionally depths in meters are given where they have some inter- est or precision. Distributional information in parentheses indicates rarity in those situations.
50 U.S. NATIONAL MUSEUM BULLETIN 271
Diagnosis of the Basic Marine Gammaridean (Many Marine Gammaridae)
Figures 1, 2
Accessory flagellum well developed, with four or more articles. Primary flagellum of antenna 1 not basally conjoint.
Mouthpart field quadratiform from lateral view. Each mandible with 3-articulate palp, article 3 longer than article 1; molar present, with grinding surface composed of ridges and teeth (=triturative). Lower lip with principal lobes undivided (unnotched) and not widely separate. Each maxilla 1 bearing inner lobe, outer spinose lobe, and strong unflexed palp of two articles. Each maxilla 2 with two well- developed, setose lobes. Each side of maxillipeds with large inner (proximal) lobe and outer (distal) lobe, and 4-articulate palp, article 4 claw-shaped (unguiform).
Gnathopods well developed and subchelate (‘‘powerful”), non- lysianassid. Gnathopod 2 larger than gnathopod 1 in male, gnathopod 1 never larger than 2 in female, article 3 of gnathopod 2 short. Article 4 of pereopods 1-2 not extensively elongate. Pereopods 3-5 of con- gruent structure and successively slightly longer.
Three pairs of large and subequal uropods present, all biramous, rami subequal in length, lanceolate. Peduncle of uropod 3 not elongate. Telson deeply cleft, of medium length.
Coxae forming elongate, rectangular plates with quadrate or rounded distal edges, coxae 1—4 of uniform shape or slightly increasing in size consecutively, coxa 4 excavate posteriorly.
All body segments free. Metasome only as long as last five pereonites combined. Head subcuboidal, not ‘‘massive,’’ rostrum small.
MARINE GAMMARIDEAN AMPHIPODA 51
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Ficure 1.—BASIC GAMMARIDEAN, part I, lateral aspect. Note the following char- acteristics of the basic gammaridean: elongate antennal peduncles; head of medium size, rostrum small or obsolete; eyes when present paired, lateral, composed of ommatidia and lacking cuticular lenses; accessory flagellum multiarticulate; coxae 14 subequal to each other in size, subquadrate in outline, coxa 4 posteriorly excavate; gnathopod 2 equal to or larger than gnathopod 1; no articles of pereopods 1-2 differentially elongated; pereopods 3-5 subequal to each other in length; all body segments free; uropods all biramous, no uropod elongated; telson free. See diagnosis.
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Figure 2A.—BASIC GAMMARIDEAN. part II: a, head, lateral aspect [mouthparts ar- ranged in quadrate bundle]; b, urosome, left lateral aspect, with uropods 1-3 and telson [all segments free, none extremely elongate, no uropods extremely disproportionate, all uropods biramous, telson freely articulate and not thickened dorsoventrally]; c, uropod 3 [well developed but of medium length, rami subequal to each other, no parts greatly elongate]; d, telson [of medium length, cleft about three fourths of its length]; ¢, gnathopod 2 [subchelate, equal to or larger than gnathopod 1].
Symbols: 1, 2, 3, etc.=articles of appendage; c=coxa; |=lacinia mobilis; i=incisor;
ip=inner plate; ir—inner ramus; op outer plate; or=-outer ramus; p=palp.
MARINE GAMMARIDEAN AMPHIPODA 53
Ficure 2B.—BASIC GAMMARIDEAN. part II: f, mandible [with 3-articulate palp, molar present and strongly triturative]; g, lower lip [lateral lobes close together, not perfectly oval, inner lobes if distinct, not amalgamated]; h, maxilla 1 [well developed, spinose, palp 2-articulate]; 7, maxilla 2 [well developed]; j, coxae 1-4 [all present, generally sub- equal in size or consecutively slightly larger, quadrate in outline, coxa 4 excavate pos- teriorly]; k, accessory flagellum [multiarticulate]; J, maxilliped [inner and outer plates large, palp 4-articulate]. See fig. 2A for symbols.
54 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 3.—Lysiannassidae: a, antenna 1, note very short peduncle [see Sebidae]; b, gnatho- pod 2, and an extreme variation of articles 6-7, as offset; note elongation of article 3 [highly characteristic overall shape, but see Sebidae, Stegocephalidae].
Stegocephalidae always lack molars and palps on mandibles, have some foliaceous portions
on maxillae, always have a 1—2~articulate accessory flagellum and a characteristic shape of
coxae 14. The combination of those characters never occurs in Lysianassidae.
MARINE GAMMARIDEAN AMPHIPODA 55
Ficure 4.—Stegocephalidae: a, head, left lateral aspect showing subconical grouping of mouthparts, shortening of head, enlargement of epistomal region (epi); 6, mandible, note absence of palp and molar and weakly developed styliform shape [see Acantho- notozomatidae];.c, accessory flagellum [see Acanthonotozomatidae]; d,e, variation in telson; f, coxae 1-4, left to right, note slight acumination of anterior coxae [overall shape
not occurring in Lysianassidae].
56 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 5.—Ochlesidae: a, head, left lateral aspect showing conical grouping of mouth- parts [stippled portions are body and coxae]; b, accessory flagellum absent; c, maxillipeds, note absence of palps [see Acanthonotozomatidae, Lafystiidae]; d, coxae 1-4, left to right, note subacumination; e, mandible, note styliform shape; f, telson.
MARINE GAMMARIDEAN AMPHIPODA 57
Figure 6.—Lafystiidae: a. head and conical mouthpart bundle [stippled area is body]; b, accessory flagellum absent; c, mandible, molar absent, note styliform shape; d, max- illiped, note only two palp articles [differing from nearly all other Gammaridea]; ¢, coxae 1-4, left to right [see Acanthonotozomatidae]; f, telson.
Gnathopod 1 is simple, gnathopod 2 is weakly subchelate. Similar to Acanthonotozomatidae
except for reduced maxillipedal palp, lack of pereonal and metasomal sculpture, and shapes
of coxae 1-4.
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Ficure 7.—Acanthonotozomatidae: a, head, note mouthparts arranged into conical bundle [see Paramphithoidae, Stilipedidae]; b, accessory flagellum absent or vestigial; c, man- dible, note styliform shape [see Stegocephalidae]; d,e, variation in coxae 1-4, left to right, note acumination of one or more coxae ventrally [see Pardaliscidae, Lafystiidae, Stili- pedidae]; f,g, variation in telson.
Ochlesidae have no maxillipedal palps whereas Acanthonotozomatidae have 3—4-articulate
palps. Lafystiidae have 2-articulate maxillipedal palps.
MARINE GAMMARIDEAN AMPHIPODA 59
Ficure 8.—Eophliantidae: body cylindrical, coxae short: a,b, mandibles, note absence of molars and palps [see Talitroidea (except Najna), Kuriidae, Corophiidae, Podoceridae]; c, accessory flagellum absent; d, maxilliped [see Colomastigidae]; ¢,f,g, variations of uro- pod 3 [see Prophliantidae]; A, telsons, cleft weakly or into two pieces [see Colomastigi- dae]; 1, body, cylindrical, coxae very short [see Phliantidae, Talitroidea].
285-135 O - 69-5
60
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Figure 9.—Colomastigidae: a, mandible, note absence of palp, molar (stippled) and deeply serrate cutting edge [see Eophliantidae, Corophiidae, Podoceridae]; b, accessory flagellum absent; c, uropod 3 [see Kuriidae, Eophliantidae]; d, maxilliped, note coalesced inner lobes [see Eophliantidae]; ¢, body, subcylindrical, coxae short; f, telson [see Eophliantidae. Prophliantidae].
MARINE GAMMARIDEAN AMPHIPODA 61
Ficure 10.—Phliantidae: a, mandible, note absence of palp and molar, projection to left is ?tendon [see Podoceridae, Talitroidea, Kuriidae, Dexaminidae, Pagetinidae]; }, ac- cessory flagellum absent; c, maxilla 1, note degradation of palp [see Dexaminidae]; d, gnathopod 1 (and 2), note feeble condition (except Ceinina) [see Prophliantidae]; note elongated article 3; e¢, body, depressed, rugose, coxae splayed [see Eophliantidae, Colomas- tigidae]; f, uropod 3 of Phlias, probably erroneous; g,h, variation in uropod 3 [see Acanthonotozomatidae, Prophliantidae]; 7,7, variation in telson, condition in figure 7 rarely occurring.
Kurtidae have poorly developed outer plates of the maxillipeds.
62 U.S. NATIONAL MUSEUM BULLETIN 271
Figure 11.—Dogielinotidae: a, mandible, note absence of palp [see Haustoriidae]; b, ac- cessory flagellum absent [see Haustoriidae]; c, antenna 1, note spinosity [see Talitroidea]; d, maxilla 1, note degradation of palp; e, epistome and upper lip, left lateral aspect, note pendant epistome [see Talitoridea]; f, any of pereopods 3-5, note heavy spinosity [see Talitroidea]; g, uropod 3, uniramous [see Haustoriidae]; h, telson.
MARINE GAMMARIDEAN AMPHIPODA 63
Ficure 12.—Talitroidea: a, mandible, note absence of palp; b, accessory flagellum absent; ¢,d,e, variations of uropod 3, essentially uniramous, rarely with scale-like inner ramus; f,g, extremes of telson.
Dexaminidae, Prophliantidae, and Kuriidae have coalescence of two or three urosomal
segments. Dogielinotidae have aspects of extreme fossorial adaptation such as spinose an-
tennae and pereopods, and a nasiform epistomal projection. Eophliantidae have cylindrical bodies, short coxae and degraded mandibular molars. Phliantidae have depressed, rugose
bodies and splayed coxae. Note the convergence of Beaudettiidae, figure 42.
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Ficure 13.—Kurtidae: a, mandible, note absence of palp and presence of molar [see Pro- phliantidae]; b, accessory flagellum absent; c, uropod 3, uniramous [see Prophliantidae, Dexaminidae]; d, urosome, left lateral diagrammatic, all segments coalesced [see Tali- troidea]; ¢, telson.
Eophliantidae have cylindrical bodies. Phliantidae have depressed, rugose bodies and splayed coxae.
MARINE GAMMARIDEAN AMPHIPODA 65
Ficure 14.—Pagetinidae: a, mandible, note palp [see Eophliantidae. Phliantidae]; b, ac- cessory flagellum absent [see Sebidae]; c, maxilliped, note absence of outer plate [see Sebidae]; d, coxae 1-3, left to right [see Stenothoidae, Cressidae, Thaumatelsonidae]; eé, uropod 3, uniramous, ramus 1—2-articulate [see Pardaliscidae]; f, telson short [see Steno- thoidae, Cressidae, Thaumatelsonidae]; g, side of urosome, segments 2-3 apparently coalesced.
Sebidae have chelate gnathopods.
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Ficure 15.—Sebidae: a, mandible, note absence of molar; b, accessory flagellum; c, uro- pod 3, uniramous and elongate [see Lysianassidae, Leucothoidae, Anamixidae, Amphilo-
chidae]; d, maxilliped, [see Pagetinidae]; ¢, gnathopod 1, note chelateness [see Leucothoi- dae]; f, gnathopod 2, note chelateness and elongation of article 3 [see Pagetinidae]; g, telson; h, antenna 1, note elongation of peduncle [see Lysianassidae].
Stenothoidae, Thaumatelsonidae, Cressidae have a reduced coxa 1.
MARINE GAMMARIDEAN AMPHIPODA 67
Ficure 16.—Cressidae: a, mandible, note obsolescent molar; b, accessory flagellum absent; c, coxae 1-4, left to right, note coxa 4 often not completely shield-like [see Thaumatel- sonidae, Stenothoidae, Pagetinidae]; d, maxilliped, note reduced outer plate; ¢, urosome, left, note telson (T) fused with segment 3 [see Stenothoidae, Thaumatelsonidae]; f, uropod 3, uniramous [see Amphilochidae]; g, telson.
Article 2 of pereopod 3 is always expanded in Cressidae, unlike Stenothoidae.
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Figure 17.—Thaumatelsonidae: a, mandible, note small, nontriturative molar; b, accessory flagellum absent or vestigial; c, coxae 1-4, left to right [see Pagetinidae]; d, urosome, left side (U=urosome, T=telson), note fusion of segments and thickened telson [see Stenothoidae, Cressidae]; e¢, maxilliped, note smallness of outer plate; f, uropod 3, unira- mous [see Amphilochidae]; g, telson.
MARINE GAMMARIDEAN AMPHIPODA 69
Ficure 18.—Stenothoidae: a, mandibles, note variations in palp, absence of triturative molar; b, accessory flagellum vestigial or absent; c, maxilliped, note obsolescence of outer plate [see Amphilochidae]; d, uropod 3, uniramous [see Amphilochidae]; ¢, coxae 1-4, left to right [see Leucothoidae, Anamixidae, Pagetinidae, Phliantidae]; f, telson.
Thaumatelsonidae often have some of the urosomal segments coalesced and the telson
strongly thickened. Cressidae have the telson fused with pleonite 6, and always have an
expanded article 2 of pereopod 3, whereas Stenothoidae always have a slender article 2.
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Ficure 19.—Amphilochidae: a, coxae 1-4, left to right [see Leucothoidae, Pleustidae, Calliopiidae, Stegocephalidae]; b, accessory flagellum absent [see Stegocephalidae]; c, uropod 3, note biramous condition [Stenothoidae, Cressidae, Thaumatelsonidae]; d, telson, entire [with one exception].
Stegocephalidae have mandibles always without molars and palps, whereas Amphilochidae
have mandibles with either one or both occurring together. Anamixidae have a ventral
cephalic keel replacing mandibles and maxillae.
MARINE GAMMARIDEAN AMPHIPODA 71
Ficure 20.—Anamixidae: a, oblique ventral view of head, showing absence of maxillae and mandibles, presence of ventral keel (cross-lined), maxillipeds removed to show maxil- lary protuberances [see Leucothoidae, Amphilochidae]; b, accessory flagellum absent; c, gnathopod 1, note chelate condition [and absence of article 7 in this case] [see Amphilo- chidae], occasionally this appendage is absent; d, maxillipeds, note amalgamated inner lobes and absence of outer lobes; e¢, coxae 1-4, left to right [see most Leucothoidae]; f, telson.
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Ficure 21.—Leucothoidae: a, mandible, note absence of molar; b, extremes of accessory flagellum, always very small; c, maxilliped, note obsolescence of outer plate; d, gnathopod 1, note complex subchelation [see Cressidae, Thaumatelsonidae, and Stenothoidae, all of which also have uniramous uropod 3]; e¢, telson, entire.
Sebidae have a uniramous uropod 3 and chelate gnathopod 2. Anamixidae, resembling
Leucothoidae strongly, lack maxillae and recognizable mandibles, and have a strongly
reduced coxa 1.
MARINE GAMMARIDEAN- AMPHIPODA 73
Ficure 22.—Prophliantidae: a, mandible, note absence of palp and recognizable molar [see Atylidae]; b, accessory flagellum absent; c, uropod 3 [see Kuriidae, Talitroidea, Phliantidae]; d, urosome, left lateral aspect, note amalgamation of all segments and telson [see Tali- troidea, Dexaminidae]; ¢, telson.
Eophliantidae have cylindrical bodies and reduced coxae.
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Ficure 23.—Dexaminidae: a, mandible, palp absent [molar evanescent in Anatylinae]; 3, accessory flagellum absent; c, urosome, left lateral aspect, note amalgamation of uro- somites 2-3 [see Prophliantidae, Talitroidea]; d, telson.
Phliantidae have the palp of maxilla 1 degraded, and depressed rugose bodies with splayed
coxae.
MARINE GAMMARIDEAN. AMPHIPODA 75
Figure 24.—Lepechinellidae: a, accessory flagellum [see Pleustidae, Ampeliscidae]; ), coxae 1-4, left to right, note acumination [see Atylidae, Ampeliscidae]; c, urosomites 1-3, left to right, note amalgamation of segments 2-3, presumably occurring in all lepechinel- lids [see Pleustidae, Paramphithoidae]; d, telson.
Presumably all lepechinellids have coalesced inner lobes on the lower lip and thus differ
from Paramphithoidae. Kuriidae, Dexaminidae, and Prophliantidae lack mandibular palps,
285-135 O - 69 - 6
76 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 25.—Atylidae: a, head [see Ampeliscidae]; b, extremes of accessory flagellum; c, urosome, left lateral view, showing amalgamation of urosomites 2-3; d, telson. Lepechinellidae have acuminate coxae 1-4. Shallow-water Ampeliscidae have cuticular
cephalic lenses. Dexaminidae, Prophliantidae, and Kuriidae lack a mandibular palp.
MARINE GAMMARIDEAN AMPHIPODA aoe
Figure 26.—Ampeliscidae: pereopods glandular: a, head, left lateral aspect, showing cuticular lenses and dotted lines indicating internal pigment (lenses unique in Gam- maridea but absent in deep-sea ampeliscids) [see Atylidae]; b, accessory flagellum absent; c, pereopod 4 [often longer than pereopod 5 and of different shape]; d, pereopod 5; ¢,f, variations in telson; g, urosome, left lateral aspect, note amalgamation of urosomites 2-3.
Atylidae have ommatidial eyes and lack the special combination of pereopods 4 and 5
shown here.
78 U.S. NATIONAL MUSEUM BULLETIN 271
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Ficure 27.—Oedicerotidae: a,b,c, pereopods 3, 4, 5, note elongation of pereopod 5 [see Calliopiidae, Pleustidae, Synopiidae, Isaeidae and related families, Laphystiopsidae]; d, accessory flagellum absent [see Synopiidae, Pardaliscidae]; ¢, telson; f, uropod 3, note elongated peduncle [see Calliopiidae, Pleustidae, Laphystiopsidae].
Isaeid-like families have a thick, fleshy telson. Oculate oedicerotids mostly have dorsally contiguous or coalesced eyes.
MARINE GAMMARIDEAN AMPHIPODA
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Ficure 28.—Pardaliscidae: a, mandibles, left with palp not illustrated, note absence of molars and foliate cutting edges [see Laphystiopsidae]; b, accessory flagella, usually multiarticulate, absence extremely rare [see Stilipedidae]; c, coxae 1-7, very short [see Stilipedidae, Acanthonotozomatidae]; d, maxilliped, note obsolescence of inner plate [see Acanthonotozomatidae, Astyridae, Laphystiopsidae]; ¢,f, extremes of telson.
79
80 U.S. NATIONAL MUSEUM BULLETIN 271
TT; / Hl
Kn
SS
SS
Sa
RS
SS ISS ISS ~N
YU,
YU
i (
Ficure 29.—Stilipedidae: a, mandible [see Synopiidae, Hyperiopsidae, Astyridae, Laphy- stiopsidae]; b, extremes of accessory flagellum, vestigial or absent [see Pardaliscidae which usually have multiarticulate condition]; c,d, maxillae 1, 2, foliaceous [see Par- daliscidae, never having foliaceous maxilla 2, Laphystiopsidae]; ¢, gnathopod 1, simple (gnathopod 2 also); f, telson.
Acanthonotozomatidae have their mouthparts grouped as a conical bundle and only one
acanthonotozomatid genus has foliaceous maxillae.
MARINE GAMMARIDEAN AMPHIPODA 81
Ficure 30.—Bateidae: a, head (H) and pereonites 1-3 with coxae 2-4 [coxa | vestigial or absentl: b, accessory flagellum absent; c, gnathopod 1, composed of coxa and article 2, other articles absent [exclusively characteristic]; d, telson.
82 U.S. NATIONAL MUSEUM BULLETIN 271
Figure 31.—Synopiidae: a,b,c, variations of head, head very large, usually galeate, with large downturned rostrum, except a few genera with head like fig. c; head like fig. ¢ also occurring in Astyridae [see Vitjazianidae, Eusiridae]; d, accessory flagella, absence prob- ably erroneous, usually more than 3-articulate [see Eusiridae]; ¢, gnathopods feeble; f, mandible, note short article 3 of palp (except in Bruzeliopsis) [see Astyridae, Par- daliscidae, Eusiridae]; g,h, telsons [see Astyridae, Oedicerotidae].
Astyridae have a uniarticulate accessory flagellum, and widely spaced outer lobes of the
lower lip. Argissidae have a unique combination of shapes of coxae 1-4. Pardaliscidae have
very short coxae. Oedicerotidae have elongated fifth pereopods. Hyperiopsidae have bent maxillary palps and elongated fourth articles of pereopods 1-2. Occasionally Astyridae and
Pardaliscidae have heads like figure b. Oculate synopiids have dorsally contiguous or co-
alesced eyes, occasionally with anteroventral accessory ommatidia.
MARINE GAMMARIDEAN AMPHIPODA
\ A A
~ ~.
~ Se SSS =
SESSA
\
ts Yi Vl “fl Wiyt Mitt
Ficure 32.—Astyridae: a, mandible, note setose and conical but nontriturative molar [see Synopiidae, Pardaliscidae, most Eusiridae, Vitjazianidae]; b, accessory flagellum [see Synopiidae, Oedicerotidae, Vitjazianidae, Hyperiopsidae]; c, lower lip, note widely separated outer lobes [highly characteristic] [see Paramphithoidae]; d, gnathopod 1
(2 is similar) [see Liljeborgiidae, most Oedicerotidae, Vitjazianidae]; e,f, extremes of telson.
Eusiridae generally have an elongate telson, and often enlarged gnathopods. Hyperiopsidae have bent maxillary palps, and elongated fourth articles of pereopods 1-2. Haustoriidae generally have either a multiarticulate accessory flagellum, or a large or triturative molar. Astyrid cephalons may resemble those of Synopiidae (see fig. 31). Two paramphithoid genera, with nontriturative molars and weakly acuminate coxae, resemble Astyridae closely.
83
84 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 33.—Hyperiopsidae: a,b, maxilla 1, note enlarged condition of palps, one of them bent; c, accessory flagellum [see’ Ampeliscidae and Stilipedidae]; d,e, gnathopods 1, 2; f, pereopod 1 (and 2 similar), note elongation of article 4 and either slender or inflated; g,h, extremes of telson.
Combination of elongate article 4 of pereopods 1-2 and bent maxillary palps differentiates
Hyperiopsidae from Astyridae, Synopiidae, Pardaliscidae, Vitjazianidae, Calliopiidae, and
Eusiridae.
MARINE GAMMARIDEAN AMPHIPODA 85
Ficure 34.—Vitjazianidae: a, peduncle (stippled) of antenna 1, very short, and base of flagellum conjoint (a long article 1 of antenna 1 flagellum ) [see Eusiridae, Gammaridae, Melphidippidae]; b, accessory flagellum [see Astyridae, Stilipedidae]; c, gnathopod 1 [see Liljeborgiidae]; d, telson.
Maxilla 1 normal [see Hyperiopsidae, Stilipedidae]. Article 4 of pereopods 1-2 of normal
length [see Hyperiopsidae]. Uropod 3 of normal length [see Melphidippidae]. Mandibular
molar large and triturative [see Astyridae, Stilipedidae]. Lower lip normal [see Astyridae].
Head small and poorly rostrate [see Synopiidael].
86
U.S. NATIONAL MUSEUM BULLETIN 271
VLA
/
z MLM SL ML
Se: Nie: S S yo!
Ficure 35.—Laphystiopsidae: a, mandible, note simple molar (general shape differing from that of Stilipedidae) [see Isaeidae and related families]; b, accessory flagellum [see Astyridae]; c, coxa 4, left aspect, of only type-genus; d,e, gnathopods 1, 2 (simple) [see Oedicerotidae, Isaeidae and related families]; f, telson.
One genus perhaps erroneously included here, see text. Probably a compound and artificial
family. Lower lip resembling Pleustidae but type-genus Laphystiopsis distinguished by
having reduced coxae and simple gnathopods. Prolaphystius has elongated urosomite 1.
Family scarcely distinct from Calliopiidae but Laphystiopsis bearing large spatulate rostrum. See lower lip of Astyridae. Oedicerotidae usually have subchelate gnathopods, large coxae,
and elongated peduncles of uropod 3.
MARINE GAMMARIDEAN AMPHIPODA 87
Ficure 36.—Paramphithoidae: (incl. Amathillopsidae): a, coxae 1-4, left to right, note acumination [see Pleustidae, Eusiridae, Calliopiidae]; b, extremes of accessory flagellum; c, gnathopod 2, note feeble condition; d,e, extremes of telson.
Acanthonotozomatidae lack a distinct mandibular molar and have their mouthparts ar-
ranged in a conical bundle. Lepechinellidae have pleonites 5-6 amalgamated, extremely
slender second articles of pereopods 3-5 and coalesced inner lobes of the lower lip. Haustori- idae have multiarticulate accessory flagella and strongly spinose pereopods.
Astyridae have nontriturative mandibular molars and widely separate outer lobes of the
lower lip. Two genera of Paramphithoidae have nontriturative molars and may be examined
in the key to the genera of Astyridae.
88 U.S. NATIONAL MUSEUM BULLETIN 271
Figure 37.—Pleustidae: a, lower lip [see Calliopiidae]; b, accessory flagellum, absent or uniarticulate [see Liljeborgiidae]; c, coxa 4, left aspect, posteriorly excavate [see Isaeidae and related families, Laphystiopsidae]; d, telson [see Liljeborgiidae, Eusiridae, Lepechinel- lidae].
Paramphithoidae and Lepechinellidae have one or more of coxae 1-4 cuspidate or pointed
| Mesopleustes appears to be assignable to either Paramphithoidae or Pleustidae]. Oediceroti-
dae have a disproportionately elongated pereopod 5 and short telson. Amphilochidae have
a very small coxa 1. Laphystiopsidae may be synonymous with Pleustidae.
MARINE GAMMARIDEAN AMPHIPODA
SL att gf yp yy,
“
you w S
Ki
‘e ba EON + Y 4 sending,
Ficure 38.—Callioptidae: a, lower lip, with or without inner lobes [see shape of lower lip of Pleustidae]; b, accessory flagellum either absent or uniarticulate; c, coxa 4, left aspect, posterior edge excavate [see Isaeidae and related families]; d,e,f, variations in telson [see Eusiridae, Gammaridae].
Oedicerotidae have disproportionately long fifth pereopods (but see Paracalliope). Laphys-
tiopsidae have a poorly developed mandibular molar. Paramphithoidae have one or more
of coxae 1-4 cuspidate or pointed. Amphilochidae have reduced first coxae. Some calliopiids
have nonexcavate coxa 4 but lack pereopodal glands of isaeid-like families ana do not have fleshy telsons.
89
90 U.S. NATIONAL MUSEUM BULLETIN 271
hi H
i
1
'
t
'
{ A \ 7) ' hi 1 4 i A
x.
‘ >-—-8-_-—>
a a te
B-s>-
Ficure 39.—Eusiridae: a, lower lip, with or without inner lobes [see shape of lower lip of Pleustidae, Astyridae]; b, accessory flagellum, absent, uni- or biarticulate [see Gammaridae which have a few species with 2-articulate accessory flagella]; c,d, uropods 1, 2, note shortened outer rami [most Gammaridae do not]; ¢,f, extremes of telson [see Calliopiidae, Pleustidael].
Astyridae have nontriturative mandibular molars but all Eusiridae except Eusirella have
strongly triturative molars. Vitjazianidae have simple first gnathopods and basally con-
joint primary flagella of antenna 1. Synopiidae have massive heads with deflexed rostra, usually have multiarticulate accessory flagella and feeble gnathopods whereas most Eusiridae have powerful gnathopods. Oedicerotidae have pereopod 5 disproportionately elongate in comparison with pereopods 3 and 4. Paramphithoidae always have the unique combina- tion of feeble gnathopods and one or more acuminate coxae 1-4. Liljeborgiidae have the unique combination of powerful gnathopods and nontriturative mandibular molar. Gam- maridae have pleonites 1-3 together not exceeding length of 5 pereonites together, whereas
Eusiridae have these pleonites (metasome) exceeding six pereonites in length; all Eusiridae
except one genus have elongate telsons, whereas Gammaridae have short telsons not or
scarcely exceeding the length of pleonite 6.
MARINE GAMMARIDEAN AMPHIPODA 91
Ai
i at wit
Ficure 40.—Melphidippidae: a, extremes of accessory flagellum; b, coxae 1-7, left to right, note shortness; c, gnathopod 1, feeble (gnathopod 2 also feeble); d,e,f, uropods 1, 2, 3, [note extreme elongation of uropod 3, see Gammaridae]; g, telson.
Melphidippidae differ from Vitjazianidae by the normally elongated peduncle of antenna 1.
See couplet 7, Section A of written key to families (p. 107) for definition of elongate uropod 3.
285-135 O - 69 - 8
92 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 41.—Liljeborgiidae: a, mandible (molar weak and nontriturative) [see Gammaridae, Haustoriidae and most genera of Eusiridae]; b, extremes of accessory flagellum [see Pleustidae]; c, gnathopods 1-2 (powerful) [see Haustoriidae, Astyridae, Vitjazianidae]; d, telson [see Pleustidae].
Article 1 of mandibular palp in Liljeborgiidae often elongate.
MARINE GAMMARIDEAN AMPHIPODA 93
Figure 42.—Beaudettiidae: a, mandible (palp absent) [see Gammaridae]; b, accessory flagellum (2-articulate); c, coxa 4, left aspect (not posteriorly excavate); d, uropod 3 (inner ramus vestigial); ¢, telson, very short, entire [see Gammaridae].
94 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 43.—Argissidae: a, accessory flagellum [see Ampeliscidae]; b, coxae 1-4, left to right [see Gammaridae, especially Megaluropus; Haustoriidae]; c, gnathopod 1 [2 is similar], feeble; d, telson.
Shallow-water Ampeliscidae have external cephalic cuticular lenses and amalgamated
urosomites 2-3. Shallow-water Argissidae have four internal, bigeminous, lenticular, ocular
bodies.
MARINE GAMMARIDEAN AMPHIPODA 95
Ficure 44.—Phoxocephalidae: a,b, lateral and dorsal views of head, note wide or narrow (stippled) rostrum in fig. b; c, accessory flagellum; d, pereopod 4 (elongated) [see Gam- maridae]; ¢, pereopod 5; f, telson.
Combination of head with large rostrum and configurations of pereopods 4 and 5 is char-
acteristic [see Haustoriidae].
96 U.S. NATIONAL MUSEUM BULLETIN 271
gnathopods feeble
Figure 45.—Haustortidae: a,b, head, dorsal views showing small and large rostra (occa- sionally genera occur with no rostra; those with large rostra (fig. b) have pereopod 5 like fig. e¢, thus not shorter than pereopod 4; genera with large rostra and pereopod 5 like fig. f are assigned to Phoxocephalidae); c, variational extremes in accessory flagellum [see Ampeliscidae, Astyridae, Paramphithoidae]; d, pereopod 4, note spinosity and occasional absence of article 7; ¢,f, extremes of pereopod 5, note spinosity [see Paramphithoidae]; g,h, telsons; 1, gnathopods highly variable but feeble [see most Gammaridae, Liljeborgiidae].
Ampeliscidae have amalgamated urosomites 2—3. Dogielinotidae lack mandibular palps
and have a uniramous uropod 3. Astyridae have a characteristic lower lip. Argissidae have
a special configuration of coxae 1-4.
MARINE GAMMARIDEAN AMPHIPODA 97
Figure 46.—Gammaridae, marine: a,b, two kinds of coxa 4, left aspect; c,d, two extremes of accessory flagellum, 2- or multiarticulate, rarely 1-articulate; ¢,f,g, three kinds of telson (arrows showing possible combinations with kinds of coxae, thus poorly excavate coxa 4 never combined with uncleft telson, except in Falklandella); h, uropod 3 (common kind); 1, uropod 3 (rare kind). Figures a,d,e and h represent a combination similar to the basic
gammaridean.
98 U.S. NATIONAL MUSEUM BULLETIN 271
' ' 1 ' 1 ( t ‘ ' \ '
SS
Sy Sea oe <9 ~---------
Ficure 47.—Isaeidae (incl. Photidae), pereopods usually glandular: a, coxa 4, note lack of posterior excavation (coxa 4 may be long or short) [see some Gammaridae, Pleustidae, Calliopiidae and related families]; b, extremes of accessory flagellum; c,d,e, variations in uropod 3; f, telson, thick, short, and fleshy [see Calliopiidae, Pleustidae, Oedicerotidae].
Five isaeid genera have uniramous uropod 3 and might be confused with numerous other
gammaridean families having uncleft telsons. But Cressidae, Stenothoidae, and Thauma-
telsonidae have reduced first coxae; Eophliantidae have cylindrical bodies; Phliantidae,
Talitroidea, and Dogielinotidae lack mandibular palps. Sebidae have subequally large,
chelate gnathopods. Pagetinidae have evanescent mandibular molars and poorly developed
outer plates of maxillipeds. See Aoridae, Ischyroceridae, Ampithoidae, Corophiidae, Podo- ceridae, and Cheluridae to follow for diagnostic differences.
MARINE GAMMARIDEAN AMPHIPODA 99
Ficure 48.—Aoridae, pereopods glandular, like Isaeidae but gnathopod 1 larger than gnathopod 2 and bearing male sexual modifications: a, coxa 4, note absence of posterior excavation; b, extremes of accessory flagellum; c,d, gnathopods 1, 2, note 1 larger than 2 [see Isaeidae]; ¢,f,g,h, variation in uropod 3 [see Ischyroceridae, Corophiidae, Cheluridae]; i, telson, short, thick, fleshy.
See Isaeidae for characters differentiating genera with uniramous uropod 3 from families
with uniramous uropod 3. Podoceridae have elongate urosomite 1. Cheluridae have coalesced
urosomites 1-3. Corophiidae have depressed urosomes. Ischyroceridae and Ampithoidae have elongate peduncles of uropod 3. Ampithoidae have characteristic lower lips.
100 U.S. NATIONAL MUSEUM BULLETIN 271
Ay NK : We.
Figure 49.—Ischyroceridae, pereopods glandular, like Isaeidae and Aoridae but uropod 3 with elongate peduncle and outer ramus with minute distal ornaments: a, coxa 4, note lack of posterior excavation; b, extremes of accessory flagellum; c,d, uropod 3 and enlarged rami, note elongate peduncle and minute nonarticulate ornaments on outer ramus [see Isaeidae, Aoridae, Corophiidae, Ampithoidae]; ¢, telson, short, thick, fleshy.
Ampithoidae have characteristic lower lips. Podoceridae have elongate urosomite 1.
MARINE GAMMARIDEAN AMPHIPODA 101
Figure 50.—Ampithoidae, pereopods glandular, like Isaeidae and Aoridae but peduncle of uropod 3 elongated and lower lips with characteristic shape, usually with notched outer lobes: a, coxa 4, note lack of posterior excavation; b, extremes of accessory flagellum; c,d, lower lips, outer lobes usually notched or excavate medially [see Ischyroceridae, Isaeidae, Aoridae, Corophiidae]; ¢, uropod 3, note elongate peduncle, short rami, presence of one or two large distal spines on outer ramus [see Ischyroceridae, Isaeidae, Aoridae, Corophiidae, Cheluridae]; f, telson.
Podoceridae have elongate urosomite 1. See text for Jassa (p. 279), the ischyrocerid often
confused with ampithoids.
102 U.S. NATIONAL MUSEUM BULLETIN 271 |
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b
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a Ue ite: G4ggg 438s!
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Figure 51.—Corophiidae, pereopods usually glandular; like Isaeidae, Aoridae, Ischyroceri- dae, but urosome depressed; uropod 3 almost always uniramous: a, extremes in accessory flagellum; 5, urosome, left lateral aspect to show depression and occasional fusion of segments; c, coxa 4, note lack of posterior excavation [shape highly variable]; d,e,f,g,h, variations in uropod 3, usually uniramous [see Isaeidae for distinctions from distantly related families]; 7,7, extremes in telson.
Genera with uropod 3 like figs. e and h are probably assignable to Ischyroceridae. Genera
with enlarged gnathopod 1 are probably assignable to Aoridae, and others, except Corophium,
are assignable to Isaeidae. Podoceridae have extremely elongated urosomite 1. Colo- mastigidae and Eophliantidae have degraded mandibles.
MARINE GAMMARIDEAN AMPHIPODA 103
Ficure 52.—Podoceridae, pereopods probably never glandular: a, urosome, left lateral aspect, showing elongated urosomite 1, typically depressed condition and frequent amalgamation of urosomites 2 and 3 (=2) [see Isaeidae, Ischyroceridae, Aoridae, Coro- phiidae, Cheluridae, Ampithoidae]; b, extremes of accessory flagellum; c, uropod 1; d,e,f, variation in uropod 2; g, coxa 4, note unexcavate posterior edge; h,7, extreme variations of uropod 3; 7, telson.
Colomastigidae and Eophliantidae have degraded mandibles.
104 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 53.—Cheluridae: a, urosome, left lateral aspect showing amalgamation of all segments and depressed condition, uropods 1-3 (U-1, 2, 3) and telson (T) [see Podoceridae, Corophiidae]; b, accessory flagellum; c, coxa 4, note absence of posterior excavation; d,e,f, uropods 1, 2, 3 [see Corophiidae, Podoceridae]; g, telson.
MARINE GAMMARIDEAN AMPHIPODA 105
Order Amphipoda
Draenosis.—Peracarid Malacostraca lacking carapace, thus having all but one or two of the thoracic segments freely visible; one thoracic segment carrying maxillipeds fused to head, occasionally next thoracic segment carrying gnathopods also fused to head (Caprellidea), followed by seven (occasionally six) visibly articulated thoracic seg- ments, each bearing paired appendages, followed by six abdominal segments or their macroscopic remnants (except Caprellidea), first three (pleon) usually bearing paired biramous pleopods, remaining three (urosome) bearing paired biramous uropods; telson freely articu- late in primitive and majority of members; head with two pairs of antennae, first occasionally biramous; maxillipeds lacking exopodites; heart mainly thoracic; respiration thoracic with gills attached to coxae (or their remnants) of segments 2-7 (variable); eyes sessile or rarely borne on unstalked cephalic scale; eggs carried in female brood pouch on ventral thorax formed of 2 to 4 (or 5) pairs of lamellae attached to coxae 2-6.
There are no radically degenerate and fully endoparasitic Gam- maridae, although a number are inquilines, ectoparasites, and com- mensals which have sucking mouthparts and prehensile mechanisms on their appendages.
Gammaridea, especially the Gammaridae, lie closest to the logical, primitive stem of the Amphipoda and almost all evolutionary lines from Gammaridae-like ancestors are based on simplification of primi- tive parts. The Caprellidea have reduced but a vestige of the abdomen (except Cercops and Caprogammarus) and the Hyperiidea have lost the maxillipedal palps. Reduction of coxae is common in both of those suborders. Ingolfiellidea have lost all pleopods but some have devel- oped a movable scale on which the eyes are borne when present.
One family of Gammaridea, the Ochlesidae, have lost the maxilli- pedal palps but their resemblance to Gammaridea in coxae and body shape and their presumed benthic habits have caused their assignment to the Gammaridea.
Key to the Suborders of Amphipoda
1. Abdomen normally vestigial, usually lacking large pleopods or uropods, except for microscopic vestiges not exceeding two pairs, gills two or 3 pairs and brood lamellae two pairs each, head and segment bearing gnathopod 1 usually immovably coalesced.
CAPRELLIDEA (skeletonshrimps, whalelice, 250 species)
Abdomen well developed, with six segments or their gross vestiges apparent, bearing some gross appendages, gills and brood lamellae exceeding three pairs.each, headiand thoraxdrees e-news ete co lye copes 2
106 U.S. NATIONAL MUSEUM BULLETIN 271
2.
3.
Maxillipeds absent . . . . . . . HYPERIIDEA (pelagic, ?500 species) Maxillipeds bearing palps (except one rare family) . ae 3 Movable compound dactyl of gnathopods 1-2 formed of antieles 6-7 tayuelnen: head occasionally with articulate ocular scale; pleopods when present, all reduced in size, leaf-like. INGOLFIELLIDEA (caves and marine, 15+ species) Dactyl of gnathopods 1—2 formed only of article 7; head never bearing ocular scale; pleopods biramous, elongate, rarely reduced except in terrestrial species.
GAMMARIDEA (marine and limnetic scuds; beachhoppers; 3300 species)
Nomenclatural Changes in Gammaridean Families
Amathillopsidae are combined with the Paramphithoidae. Anatylidae are combined with the Dexaminidae. Hyalellidae are included with the superfamily Talitroidea. Hyalidae are included with the superfamily Talitroidea. Photidae are combined with the Isaeidae.
Pontogeneiidae are combined with the Eusiridae. Talitridae are included with the superfamily Talitroidea. Tironidae are combined with the Synopiidae.
Key to the Families of the Suborder Gammaridea '!
(“‘exceptional”’ refers to rare occurrence of familial members in this
category) SECTION A Article 3 of gnathopod 2 elongate (fig. 3). ........ . . Section B Article 3 of gnathopod 2 not elongate ... . ER CERE NC ENIEAN G" 30/24
Urosomites 1-2 coalesced, urosomite 3 free (fig. 105). some species of PHLIANTIDAE; Kamaka (Corophiidae) ; and Chevalia (Isaeidae) Urosomites 2-8 or 1-8 coalesced. . . .-.....4. .. . . Section © Urosomites separate .... A Vale Coxa 1 small or absent, mostly Ridden ray a Ralllowmne coxa , one of following coxae longer than wide) (occasionally gnathopod 1 absent or vestigial). Section D Coxa 1 small or large but usually subequal to coxa 2 and never hidden by following coxae (occasionally coxa 1 partially hidden but all following
coxae wider than long), (gnathopod 1 always fully developed) . . . . 4 Uropod 3 uniramous, or lacking rami, or absent. . . .. . . . Section E Uropod 3 biramous, inner ramus often reduced .... . oy STEM eo: Mandibular palp absent (use alternate if Section F Tea HaACtORy, because
of uncertain loss of palp) ... . cA gest Cpa SeCtONmn Mandibular palp present (with very rare , exesoviion cesent) 2. of Sa eciee ete inan
Mandibular molar absent or if present not triturative, lacking numerous ridges and teeth (occasionally with articulate spines), occasionally large or immense and dominating mandible (fig. 166d) . . . . . . Section G
1 Gurjanova (1962) also has a modern key to the families.
10.
11.
12.
MARINH GAMMARIDEAN AMPHIPODA 107
Mandibular molar well developed, triturative, bearing ridges and teeth, never extremely large or dominating mandible... . . SUMATRAN Lrg, Peduncle of uropod 3 elongate, as long as or longer than rami of uropods 1-2 (fig. 40) and more than twice as long as telson and peduncle of uropod 2, uropod 3 also greatly exceeding apices of either uropods 1 or 2 and rami elongate, subequal and nearly as long as rami of uropods 1-2 (uropod 3 is usually missing on individuals of this category but melphidippids are also recognized by the presence of dorsal teeth and serrations on the pleon, in combination with short, subequal and evenly quadrate anterior coxae, plus strong hemispherical lateral ocular bulges on head). ...... 82 If peduncle of uropod 3 elongate as above then uropod 3 not strongly exceeding apices of uropods 1—2; if peduncle of uropod 3 twice as long as telson then uropod 3 not exceeding apices of uropods 1-2 or peduncle not as long as rami of uropods 1-2; if uropod 3 greatly exceeding apices of uropods 1—2 then inner ramus short and scale-like or peduncle not elongate and not more than 1.2 times as long as peduncle of uropod 2, or rami much shorter than peduncle; thus uropod 3 not like that of first part of this couplet... . 2 9 Coxae 2-3 longer ben. pron Dleon Hoteally aroun mercoued I very conspicuously longer than 3 or 4, head without lateral ocular bulges (head often strongly rostrate) . . (exceptional) OEDICEROTIDAE (p. 373) Coxae 2-3 very short and broader than long, pleon dorsally toothed and serrate, pereopods 3-5 very long but subequal in length to each other, head with strong hemispherical ocular bulges on sides. MELPHIDIPPIDAE (p. 368) Coxae 2-3 about as long as broad, pleonite 4 with one dorsal tooth, pereopods 3-5 long but subequal in length, head without lateral ocular bulges. Casco (Gammaridae) (p. 238) Coxae 1-3 successively and very distinctly shortened (fig. 43), (and see mimic Megaluropus in Gammaridae and Casco with anteriorly acuminate coxa
lin Gammaridae) ... . . . . ARGISSIDAE (p. 157) Coxae 2 or 3 always as long as or Neceer hen c Coxaelen ss Ory Renee ss tl Telson entire, or emarginate, very short, coxa 4 not excavate posteriorly
(pereopods often glandular) . .. . Hae aaa SCION: le
Telson entire or cleft, short or long, coxa 4 ignellie excavate posteriorly or acuminate (coxa 4 always excavate posteriorly or acuminate when telson entire, but see Parapherusa in Section F and in Gammaridae), (pereopods not glandular) ...... Bi atone red
Head massive (see dismaliion § in rlgssay he or eh Eronels downturned rostrum, or “‘shark-nose’’ rostral projection ie 3l1c), [gnathopods weak GSI aciico anneal: PARRY HDD
Head not massive or rose cf estan) on massive rend Hot downturned (Hyperiopsidae have massive head, no rostrum; Pleustidae and some Phoxocephalidae have downturned rostrum on small head)... . 13
Pereopod 5 conspicuously longer than pereopods 3-4, at least 1.5 times as long as pereopod 4, accessory flagellum absent, or l-articulate and short if present, peduncle of uropod 3 always elongate and telson always half or less as long as that peduncle and usually entire or rarely emarginate.
OEDICEROTIDAE (p. 373)
Pereopod 5 subequal to pereopod 4 in length, though both often elongate, accessory flagellum always present and usually multiarticulate; when 1- articulate, accessory flagellum elongate; peduncle of uropod 3 rarely
2 See also the calliopiid Metaleptamphopus.
285-135 O - 69 - 7
108 U.S. NATIONAL MUSEUM BULLETIN 271
elongate but telson always (except Synopia with large accessory flagellum) longer than and as twice as long as peduncle of uropod 38, cleft or
entine se ariens . . . SYNOPIIDAE (p. 454)
13. Pereopods 3-5 atronele spinose or 2 aeiOse ond rahi Gea setae (‘‘fossorial,”’ see glossary) (figs. 97-99) . aa it Ea at Mat elie gunn ab Pereopods 3-5 poorly unos ¢ or SOU or noe Toscana ee epnet aia meter: 1)
14. Pereopod 5 conspicuously elongate, at least 1.5 times longer than pereopods SAGs Nsw te + » . OEDICEROTIDAE (p. 373) Pereopod 5 not pongoreonsley Smeets, subequal to, shorter than or slightly longer than pereopod4.... . RES PR eer! (a Lis)
15. Telson elongate, nearly twice as long as epednmcle Of hinonod 3 or urosomite Sey asus eed . . . (exceptional) sYNoPIIDAE (p. 454) Telson not Soni, Rearcel: eased length of peduncle on uropod 3 or urosomite3..... BM eriti ed 0)
16. Base of primary de eee a tena 1 camnictint, ane article 1 of flagellum longer than peduncle (fig. 34a) . (exceptional) VITJAZIANIDAE (p. 476) Base of primary flagellum of antenna 1 multiarticulate, article 1 of flagellum less than half as long as peduncle... . . echapaeeh aeaenalledt 17. Rostrum usually evanescent, pereopod 5 ennai Solan ig Herconnd 4 in structure; when pereopod 5 shorter than and of different structure than pereopod 4 then rostrum absent (fig. 45). (See Sancho in Calliopiidae) 18 Head with distinct, and depressed or downturned, rostrum, pereopod 5 always shorter and of different structure than pereopod 4 (fig. 44). PHOXOCEPHALIDAE (p. 412) 18. Check the individual through the family key of Haustoriidae. Most haus- toriids will be detected in the first 15 couplets. If one of the following characters does not apply to the species, proceed to Gammaridae: (1) pereopod 5 much shorter than 4 and article 2 lamellar (Pontoporeia and Urothoides) ; (2) mandible immense, triturating surface large and smooth, distal, palp relatively small and thin, and nearly asetose (Carangolia); (3) article 2 of maxilla 1 palp shorter than article 1 (Phozocephalopsis); (4) mandibular incisor unproduced and teeth absent or obsolescent and article 6 of pereopods 1-2 with stout spines (Urothoe and Urothoides). HAUSTORIIDAE (p. 248) and GAMMARIDAE (p. 231) 19. Some anterior coxae acuminate midventrally (fig. 36), and accessory flagel- lum less than 3-articulate. PARAMPHITHOIDAE (p. 389) and ACANTHONOTOZOMATIDAR (p. 117) 3 Anterior coxae not acuminate midventrally, or if acuminate then accessory flagellum more than 2-articulate . ... . Biveea ings X0) 20. Article 4 of pereopods 1-2 extremely elongate Polntive fo ened articles, palp of one member of first maxillae distinctly geniculate and scaled. (fig. 33) HYPERIOPSIDAE (p. 261) Article 4 of pereopods 1-2 not ee palp of maxilla 1 neither strongly
geniculate nor scaled . ... . Pepe pret s(n val: 21. Telson entire (often with minute notch or Spoilers auindiantion) - ican 4 Melsonnelefty seek yi os in Aouce: bay hele die gh Men hey (elie cate Ian ocean ek Seem, OF
3 See also Halirages stebbingi Schellenberg (1931) (=H. huzleyanus Stebbing, 1888, not Bate) in family Calliopiidae. Note that almost all acanthonotozomatids have a nontriturative mandibular molar or no molar whereas almost all Paramphithoidae have triturative molars. Acanthonotozomatidae usually have conically grouped mouth parts from lateral view whereas most Paramphithoidae have mouth parts grouped in a quadrate bundle. See text for further discussion.
22.
23.
24.
25.
26.
27.
28.
29.
30.
ol.
32.
33.
34.
MARINE GAMMARIDEAN AMPHIPODA 109
Atecessory flagellum 2-articulate’. 29.059. 5 2 63 8 28 Accessory flagellum l-articulate or absent ... . ROU cra
Rami of uropod 3 cylindroconical, much shorter anon Sones peduncle. Bathyphotis (Ischyroceridae) (p. 275)
Rami of uropod 3 lanceolate or flabellate, much longer than peduncle.
(exceptional) GAMMARIDAE (p. 231) Lower lip with unpointed, tilted, oval lobes astride partially coalesced inner lobes (fig. 37) .... . . . PLEUSTIDAE (p. 421) Lower lip with tilted or mntlted putes lobes! but with distinct mandibular extensions, inner lobes when present not coalesced ae 38), (examine also
key to Eusiridae if telsonic condition dubious) .. . Sumeineumetnin ssa7 4) Telson longer than wide ... . ee CALLTOPITDAE = (DLO) Telson as wide as or wider than lone . . . (exceptional) GAMMARIDAE
Article 1 of primary flagellum on antenna | as long as peduncle. VITJAZIANIDAE (p. 476) and (exceptional) male SYNOPIIDAE (p. 454) Article 1 of primary flagellum on antenna 1 not longer than half of pe-
clummcley rane oe sao cee ie Parl Telson elongate (fig. 39), vice as s long : as peduncle at Groped Sue ee Telson rarely longer than peduncle of uropod 3, never twice as long as peduncle of uropod3 ..... een ee se COU,
Urosomite 3 twice as long as masonic 2, jhendl oa lenterodorcal margin extended as blunt plow (fig. 66d) . Pseudotiron (Synopiidae) (p. 461) Urosomite 3 less than 1.5 times as long as urosomite 2, head of normal
dimensions as in basic gammaridean . . . sy Seelass caste NAS) Accessory flagellum 3-+--articulate . (xccutionel) SYNOPIIDAE (p. 454) Accessory flagellum O-2-articulate Gane also key to Calliopiidae if telsonic cleft dubious) ...... . . . BUSIRIDAE (p. 213)
Inner ramus of uropod 3 short and puale: ie (Ge. 96c). (exceptional) GAMMARIDAE (p. 231)
Innerramustof unopod’3 elongate 4. vs 2)". fa 5 2 el. 3 Jol Rami of uropod 3 foliaceous ... . coos GAMMARIDAE (p. 231) Rami of uropod 3 lanceolate . .. . Loatiis SA Sh ele ees tl ue dea
Mandibular palp article 2 shorter than onttialts ie (exceptional) GAMMARIDAE (p. 231)
Mandibular palp article 2 longer than article 1 . .... . dd
Gnathopod 1 simple ..... . . (exceptional) GAMMARIDAE ©. 231)
Gnathopod 1 subchelate . .. . EAA SUS As MVC aie SD Rc oe Ye
Accessory flagellum 2+ - aceite ee i oO GAM MAR ED Amys 2 oi)
Accessory flagellum 1-articulate or absent . . . . . BUSIRIDAE (p. 213) SECTION B
Head and body strongly depressed, rugose, coxae splayed (fig. 10).
PHLIANTIDAE (p. 405)
Head subglobular, body cylindrical (fig. 82), coxae short, not splayed. (exceptional) EOPHLIANTIDAE (p. 209) Head compressed or subglobular, ae compressed laterally, rarely rugose, coxae never splayed . ... . Reese Article 2 of antenna 1 shorter han or sapell to article iL nronod 3 usually biramous but when uniramous then article 2 of antenna 1 not elongate or gnathopod 1 not chelate .. . 3 Article 2 of antenna 1 much longer ahem, or Secniell fa onfalle 1 (Ges 15), nropods always uniramous, gnathopods strongly chelate . . . SEBIDAE (p. 435)
110 U.S. NATIONAL MUSEUM BULLETIN 271
10.
Lis
12.
13.
. Article 4 of pereopods 1—2 enormously elongate relative to other articles and
often inflated (fig. 338f) ..... Ree Aaa Article 4 of pereopods 1—2 of normal lenedh wala to Saher aatiialles - eee)
. Palp of maxilla 1 claviform, slightly geniculate (figs. 33a, b.).
HYPERIOPSIDAE (p. 261)
Palp of maxilla 1 not geniculate .. . . . proceed to couplet 5 . Gnathopod 2 with form of typical lestamaseicl saittiom (fig-13) ee Gnathopod 2 not typical of lysianassids .. . . SAS AEE ea
. Accessory flagellum absent and pleonites 1-3 wails han dowel processes,
articles 5 and 6 of gnathopod 2 lacking scales or minute coarse setules. (exceptional) ACANTHONOTOZOMATIDAE (p. 117)
If accessory flagellum absent then pleonites 1—3 dorsally smooth; accessory flagellum otherwise present, articles 5 and 6 of gnathopod 2 bearing scales
or minute coarse setules . .... . . . (fig. 3) LYSIANASSIDAE (p. 294)
. Urosomiteszally coalescedg ne 2) iic50 ieee obs eer oe es ea a ee Wrosomites Separate i056 Gs sepia tec (ee aeo ule pels weary ikea te Won ea
. Uropod 3 biramous. ...... . . . (fig. 22) PROPHLIANTIDAE (p. 432) Uropod 3 uniramous .. . See oo 6 (aalegs 113) aqenenaanyeus, (>, 2EO))
. Uropod 3 formed only of Simmel, Agelle- like peduncle.
Didymocheila (incertae sedis) (p. 478)
Uropod 3 bearing one or two rami (inner often reduced) . . avai uo Uropod 3 with one ramus . . . .. . . Microprotopus Gsacidse) (p. 274) Uropod 3 with tworami .. . dace seem hl
All mouthparts except emilline dt repliced Be sentra (buecell reel and bulbs. (fig. 20) ANAMIXIDAE (p. 145)
Mandibles, lower lip and maxillae present. . .. . shaven bt ede Accessory flagellum 1-2 articulate or absent, moaminaaNts om lateral view usually grouped in conical bundle, pereopods not strongly spinose. . 13
Accessory flagellum always 1-+-articulate, mouthparts from lateral view usually. quadrately grouped, pereopods strongly spinose and setose (‘‘fos- sonial’y)m (filet 4 5) se . . (exceptional) HAUSTORIIDAE (p. 248)
Mandibular palp present, Picon snoaesstieenous (fig. 7).
(exceptional) ACANTHONOTOZOMATIDAE (p. 117)
Mandibular palp absent, pleon never processiferous (fig. 4).
STEGOCEPHALIDAE (p. 436)
SECTION C Body plan cylindrical or depressed (tanaid-like, or cuit or extremely broad and flat)... . Pe aN a: 2 Body plan compressed laterally. @ocmal cemmmenidlennn lia) 6 Mandible lacking palp and molar degraded .... . Bee mene Gul (Re Mandible with palp of one to three articles, molar ftomattire 4
Body cylindrical, tanaid-like, smooth (fig. 8), coxae very small. COLOMASTIGIDAE and EOPHLIANTIDAE (pp. 182 and 209) Body strongly depressed, very broad and rugose (fig. 10), coxae large and splayed or body, if not rugose, with pereonites laterally discontiguous. PHLIANTIDABR (p. 405) Urosomites 1-3 coalesced . . .. . Meal sea 25) Urosomites 2—3 coalesced and urosoraies Ls very leloneate (Ge 52). PODOCERIDAE (p. 426) Uropods 1-2 of very diverse structure (fig. 53) . . . CHELURIDAE (p. 180) Uropods 1-2 similar to each other and of normal gammaridean structure. (exceptional) COROPHIIDAE (p. 184)
10.
Wik
12.
13.
14.
15.
16.
MARINE GAMMARIDEAN AMPHIPODA 111
WropodkescuniramOuUsice sta vst keeenel MEA CtoN. ay ths Dns OTN Seale 07 Uropod 3 biramous. . . . aa Cerne Ulu aeraganaay SETAC) Coxa 1 visible, not covered By folloeane | CORE) iii. 5, BUR Ce i ee ee tS Coxa 1 small and hidden by large following coxae .. . Bing a ah Sons Mandible lacking palp, bearing molar, inner plates of mamillipeds obsolescent,
telson cleft. .... ... . (fig. 13) KURIIDAE (p. 280) Mandible bearing palp, lacking! molar inner plates of maxillipeds fully
developed, telson entire. . . ... . . (fig. 14) PAGETINIDAE (p. 387)
Telson and pleonite 6 coalesced even though telson recognizable and un- thickened, article 2 of pereopod 3 expanded. (fig. 16) CRESSIDAE (p. 198) Telson thickened dorsoventrally and distinct from urosome, but urosomites
partially coalesced, article 2 of pereopod 3 linear. (fig. 17) THAUMATELSONIDAE (p. 473)
Mandibularspalpyabsentba each) eae oe ene 28 UL Mandibular palp present. . . Se asa, WEEE Se Only urosomites 2-3 coalesced (fig. 23) . . . . . DEXAMINIDAE (p. 200) Urosomites 1-3 coalesced (fig. 22)... . . . PROPHLIANTIDAE (p. 432) Pereopod 5 of different structure and almost always shorter than pereopod 4
(igs i226) kis ae . . . . AMPELISCIDAE (p. 128) Pereopod 5 similar nl pabedual oF or elonger than pereopod 4 (or these
pereopods' often brokenjorsmissing) ioe Pe Se ae es 1S Telson entire orslightly emarginate ................ 14 Telson cleft or deeply emarginate ........ mee Was LG Urosomite 1 elongate (fig.52) .... . . . . PODOCERIDAE (p. 426) Urosomite 1 not elongate, or all eroeorntce ponlesecd Sr SERA ERY aay os oe RRR) EE
Uropods 1 and 2 of extremely diverse structure (fig. 53). CHELURIDAE (p. 180) Uropods 1 and 2 similar to each other and of normal gammaridean structure (nike 399) Dae uae ne . . . COROPHIIDAE (p. 184)
Coxae 1-4 rounded, fuadente or Eplunt Penirallen
(fig. 25) ATYLIDAE (p. 161) One or more of coxae 1-4 acuminate ventrally and often bifid (fig. 24). LEPECHINELLIDAE (p. 286)
1 See Paracalliope in supplement.
SECTION D Gnathopod 1 reduced to one or two articles or absent (figs. 20,30) . . . 2 Gnathopod 1 present, comprising six or seven articles... . 3 Mouthparts all present and normal . . . . . (fig. 30) BATEIDAE “3 (Ge 163)
Mandibles and maxillae absent or coalesced into sharp keel (fig. 20) ANAMIXIDAE (p.145) Pereopods 3-5 fossorial, article 4.0f antenna 2 strongly expanded, antennae with plumose setae longer than any antennal article. (exceptional) HAUSTORIIDAE (p. 248) Pereopods 3-5 usually not fossorial but if fossorial then article 4 of antenna 2 not expanded, setae of antennae not elongate plumes. ...... 4 Gnathopod 1 strongly carpochelate, carpus larger than propodus (or ap- pearing as propodochelate because of loss of dactyl) (fig. 20)... .. 5 Gnathopod 1 subchelate or simple, or carpus smaller than propodus (oc- casionally carpus with long lobe but not distinctly chelate) . . ... 6 Mouthparts except maxilliped replaced by a ventral keel and bulbs. (fig. 20) ANAMIXIDAE (p. 145)
112
10.
11.
12.
U.S. NATIONAL MUSEUM BULLETIN 271
Mandibles, lower lip and maxillae present. (exceptional) LEUCOTHOIDAE (p. 289)
Uropod 3 biramous:: 12)) ihe eanreeh Gras tly Si sea ese Ucar enact aerial NU armG| Uropod 3 uniramous . . aU as Sree ree ae eyes ih (0) Coxa 4 not excavate aire. poreapods ie 2 eflesmdtcllon GET LOR Ose Coxa 4 excavate posteriorly, pereopods 1-2 not glandular ...... 9
Rami of uropod 3 shorter than peduncle, outer ramus with distal hook or distolateral denticles, lacking setae. (exceptional) ISCHYROCERIDAE (p. 275) Rami of uropod 3 longer than peduncle, outer ramus setose or spinose but lacking special hooks or denticles. (exceptional) ISAEIDAE (= Photidae) (p. 264) Article 4 of pereopods 1-2 exceptionally elongate (fig. 33f), base of primary flagellum on antenna 1 conjoint and forming an article longer than pe- duncle, palp of maxilla 1 geniculate and scaled (figs. 102a, 6). Parargissa (Hyperiopsidae) (p. 264) Article 4 of pereopods 1—2 not elongate, base of primary flagellum on antenna 1 not conjoint, palp of maxilla 1 not geniculate or scaled. AMPHILOCHIDAE (p. 132) Gnathopods simple and slender, with dense setae as long as fifth and sixth articles, all coxae at least as broad as long. (exceptional) ISAEIDAE (=Photidae) (p. 264) Gnathopods usually subchelate and one pair usually stout, if not then their setae very short and sparse, some coxae very much longer than broad . 11 Article 2 of pereopod 3 slender and linear (like fig. 162 v) . . ..... 12 Article 2 of pereopod 3 expanded like pereopods 4—5 (like fig. 162s). CRESSIDAE (p. 198) Urosomal segments coalesced fully or partially, telson thickened dorso- ventrally (fig. 17), or gnathopod 2 carpochelate. (exceptional) THAUMATELSONIDAE (p. 473) Urosomal segments separate, telson depressed (fig. 18), gnathopod 2 never carpochelate ............. . . . STENOTHOIDAE (p. 444)
SECTION E
Uropod 3 essentially uniramous, lacking rami or occasionally absent; when present inner ramus scalelike and outer ramus cylindrical; Cheluridae partially included herein although outer ramus flat; observer urged to verify that one ramus of uropod 3 not accidentally broken off.
1.
Mandibular molar bearing grinding oe (triturative) or if not triturative
then molar cupshaped. ... . CN rea ete Mandibular molar smooth or pearing a fou erticulate snes, or molar
absent) or not:cUpshbaped va voy ae ae) Leen eee eee) Mandibular palp' presenti oo hey 2 elena eae ee tes ae a Mandibular palp absent. ... . aetna opie 20 (0) Urosomite 1 elongate, longer than nelccomnlis 3 and more nan twice as
long as urosomite 2 (fig. 52)... . . . . . . PODOCERIDAE (p. 426) Urosomite I notelongate as’aboves..- <1. 6 ee ee ee All urosomites‘coalesced) Neco eee astve usin oe een ee rar Urosomites separate . . HHMI aN ene mien 1 8S
Uropods 1-2 of strongly diverse perucitine Ge 53) . CHELURIDAE (p. 180) Uropods 1-2 normally styliform, although occasionally with rami or ped- uncles reducedin size . ...... . . (fig. 51) coROPHIIDAE (p. 184)
MARINE GAMMARIDEAN AMPHIPODA 113
6. All urosomites coalesced (fig. 13)... ... =... . KURIIDAE (p. 280) Urosomites separate .... : SA bee mE
7. Antennae and pereopods strongly splines ¢ or ecioee spines anal setae long, epistome nasiform (fig. 11) .... . . . .DOGIELINOTIDAE! (p. 207)
Antennae and pereopods poorly spinose or setose (spines if numerous very small), epistome rounded or truncate in front. (fig. 12) TALITROIDEA (p. 463) 8. Lower lip lacking inner lobes, outer lobes obsolescent (fig. 173g), gnathopods small, chelate, slender, equal, sixth article elongate (figs. 173d, e). Didy mocheila (incertae sedis) (p. 478) Lower lip with inner lobes and strong mandibular se ee not
taking special form of figs. 173d,e ....... Be Prams cy NO
9. Urosome depressed... .. . . . COROPHIIDAE (p. 184) Urosome not depressed . . (Cxeeptional) AORIDAE (p. 147) and ISAEIDAE
(p. 264)
LOM Viandibularipalp present’ soc. ol Lees ele SOR hea ee es) LD Mandibular palp absent ... . Bi ca ceo ead iy
11. Gnathopods subchelate, outer plates oF aeciiped very painall: inner obsolete.
(fig. 14) PAGETINIDAE 2 (p. 387)
Gnathopods chelate (fig. 15e) inner and outer plates of maxilliped well formed.
(exceptional) SEBIDAE (p. 435)
12. Body cylindrical (fig. 87a), coxae small . . (fig. 8) EOPHLIANTIDAE (p. 209) Body compressed (normal), coxae large, not splayed.
Najna (Talitroidea) (p. 470)
Body depressed, rugose, coxae splayed laterally, or if body not rugose pereon-
ites laterally discontiguous . . . . . . . (fig. 10) PHLIANTIDAE (p. 405)
1 Metoediceros Schellenberg (1931) resembles this family but its epistome is unknown and its telson is nearly a perfect, uncleft circle, whereas Dogielinotidae have a rectangular, cleft telson. Metoediceros is pres- ently classified with the Oedicerotidae but it probably should form the type of a new family and is therefore removed to ‘‘incertae sedis.”’
2 Urosomites 2-3 are presumed to be coalesced.
SECTION F le bodvyaplan cylindrical (fig. Sia). ie teenies Mae eee Tees aie oe 2, Body plan compressed or depressed . . . . . 5)
2. Palp of maxilla 1 larger than outer plate, inner etnies of Peeained! dermaded: (fig. 9) COLOMASTIGIDAE (p. 182) Palp of maxilla 1 oa inner plates of maxilliped well developed, Separate .... . . . . (fig. 8) EOPHLIANTIDAE (p. 209) 3. Mandibular molar of ted sizes trituratives(ige2f yet ts Bowes A
Mandibular molar enormous, smooth (fig. 166d). Jeddo (Synopiidae) (p. 461) Mandibular molar evanescent or absent, lacking grinding ridges ... . 6
4. Uropod 3 with subequal rami (fig. 50).
Sunamphitoe (Ampithoidae) (p. 145)
Uropod 3 with inner ramus reduced or absent . . 5 5. Accessory flagellum absent. ... . . (fig. 12) TALITROIDEA (os 463) Accessory flagellum present .... . . (fig. 42) BEAUDETTIIDAE (p. 166)
6. Mouthparts from lateral view conically grouped below head, uropod 3 bi- ramous and well developed, coxae 1—4 compressed, elongate, forming lateral shield, body compressed, smooth . . (fig. 4) STEGOCEPHALIDAE (p. 436)
114 U.S. NATIONAL MUSEUM BULLETIN 271
Mouthparts from lateral view not conically grouped below head, uropod 3 degraded, coxae 1-4 not Pas lateral shield, splayed laterally, body depressed, rugose . . . .. . . (fig. 10) PHLIANTIDAE (p. 405)
Mouthparts from lateral view nor conically grouped below head, uropod 3 biramous and well developed, coxae 1-4 compressed, very short, not forming lateral shield, body compressed, smooth.
Parahalice (Pardaliscidae) (p. 403)
SECTION G 1. Maxillipedal palp with fewer than four articles ........... 2 Miaxaillipedal\palpy with; foursarticles| iio nes ar ae ae eS
2. Maxillipedal palp 3-articulate. (fig. 7) (exceptional) ACANTHONOTOZOMATIDAE (p. 117) Maxillipedal palp 2-articulate (fig. 6)... . . . . LAFYSTIIDAE (p. 281) Maxilliped lacking palp (fig. 5) .... . . . OCHLESIDAE (p. 371) 3. Mouthparts from lateral view conically prouned belowsheadi see Mouthparts from lateral view not conically grouped below head . . . . 5 4. Anterior coxae long, some acuminate ventrally (fig. 7). ACANTHONOTOZOMATIDAE (p. 117) Coxae short, not acuminate ventrally (fig. 28). Halicella (Pardaliscidae) (p. 402)
5. Gnathopod 1 carpochelate (fig. 21) . . . . . . . LEUCOTHOIDAE (p. 289) Gnathopod 1 propodochelate, subchelate, or simple ......... 6 6. Mandibular molar present (though occasionally very minute) ..... 7 Mandibular molar absent . . . PA ere S22
7. Pereopod 5 shorter and of diferent sinew inert merconad 4, head with strongly depressed large rostrum.
(fig. 44) (exceptional) PHOXOCEPHALIDAE (p. 412)
Pereopod 5 of structure similar to pereopod 4, rarely shorter than 4, rostrum
if present not depressed (not dorsoventrally flattened) ....... 8 8. Telson elongate, more than twice as long as urosomite 3 ....... 9 Telson short, subequal to or shorter than urosomite 3 (seen laterally) . . 10
9. Accessory flagellum l-articulate; antennae with large calceoli; gnathopods large and strongly subchelate (fig. 92h).
Eusirella (p. 225) and Eusiropsis (Eusiridae) (p. 225)
Accessory flagellum multiarticulate; antennae lacking calceoli; gnathopods
feeble (fig. 167) .... . . . SYNOPIIDAE (p. 454)
10. Pereopods 3-5 fossorial Gone fees of tinoxe pecconods at least half as long
as longest article of any pereopod, fig. 45)... . . é oa e ele:
Pereopods 3-5 not fossorial (setae and spines very dhar, seancelly exceeding
article 3 of those pereopods) ... . Aire isk: Goel
11. Accessory flagellum l-articulate and oxtroinaly eral) or “alpsen: pereopod
5 very elongate, at least 1.5 times as long as pereopods 3-4 (uropod 3
always elongate, rami very slender and lanceolate, subequal to peduncle,
but uropod 3 often broken, telson always short, linguiform and entire or
emarginate). ........ . . (exceptional) OEDICEROTIDAE (p. 373)
Accessory flagellum rarely vestigial (if so, uropod 3 very short, flabellate,
rami highly unequal), pereopod 5 shorter than or usually scarcely longer
than pereopod 4 (uropod 3 usually with short peduncle, unequal rami
one of which longer than peduncle, if not then accessory flagellum multi- articulate, telson cleft even minutely, usually elongate).
(exceptional) HAUSTORIIDAE (p. 248)
12.
13.
14.
15.
16.
I2fe
18.
19.
20.
21.
22.
MARINE GAMMARIDEAN AMPHIPODA 115
MelSONRCLELG Ama ean ie Magica? seen eer S NENA rie Cet h aan Hea deneeeh Mayr ath be 1S Telson entire .. . Ses ic Tae BY Gnathopods porreneeilly piticnelete (fig. Ae pat like pe 609) MEP eR LA: Gnathopods feeble (fig. 32d)... . . 15 Accessory flagellum 2-+--articulate, Tes ina pbsolescent 4s cleft 75 percent of itslength ....... (fig. 41) LILJEBORGIIDAE (p. 291) Accessory flagellum absent, rostrum neslly as long as article 1 of antenna 1, telson cleft 40 percent of its length. Pseudamphilochus (Amphilochidae) (p. 139) Accessory flagellum 0-1 articulate, rostrum obsolescent, telson cleft 25 percent of itslength .. . . . . Austropleustes (Pleustidae) (p. 422) Mandibular molar a conical or trapezoidal, setose lamina ..... . 16 Mandibular molar a large setulose tuberosity. Synopia (Synopiidae) (p. 462) Accessory flagellum present ...... . . (fig. 32) asTyRIDAE (p. 157) Accessory flagellum absent.
Eclysis and Epimeriella (Paramphithoidae and Astyridae) (pp. 161, 389) Coxa 3 with about 3 times surface area of coxa 4, maxillipeds foliaceous, article 3 of mandibular palp vestigial, subequal to article 1 (fig. 165e).
Synopia (Synopiidae) (p. 462) Coxa 4 usually slightly larger than, subequal to, or scarcely smaller than coxa 3, maxillipeds not foliaceous, article 3 of mandibular palp much
longer than articlel .... Srey te a Coxae 1-4 wider Garcnomnosterion) ian Tene (Gormaleyentrsl) (fig. TIVST GAs <a ie Bh Alan heNan. Pee te AO Snead phere yO) Coxae 1-4 longer diner mide Ge 37) aac ale RRs: CUR Te Shine, SEES 20
Coxae contiguous or slightly overlapping. (exceptional) CALLIOPIIDAE (p. 167) Coxae not touching serially (fig. 111a). Laphystiopsis (p. 285) and Prolaphystiopsis ena (p. 285)
Mandibular molar bulbous, ovate or cylindrical. . . pees 2 Mandibular molar asetose conicallamella . . (fig. 32), ASTYRIDAE (p. 157) Article 4 of pereopods 1-2 enormously elongate (fig. 33f), accessory flagellum
long and atleast 3-articulate . .. . . . . .HYPERIOPSIDAE (p. 261)
Article 4 of pereopods 1-2 of normal aimensions! accessory flagellum short or absent and less than 3-articulate.
PLEUSTIDAE (p. 421) and CALLIOPIIDAE (p. 167)
Accessory flagellum usually (probably always) present and 3-+-articulate,
maxillae not foliaceous (occasionally palp of maxilla 1 slightly foliaceous).
(fig. 28) PARDALISCIDAE (p. 397)
Accessory flagellum absent, vestigial or 2-articulate, extremely small, max-
illae foliaceous. ....... =... . (fig. 29) stmrpEDIDAE (p. 451)
SECTION H
Pereopod 5 exceeding 1.5 times length of pereopods 3-4 ! (fig. 27). OEDICEROTIDAE (p. 373) Pereopod 5 scarcely longer than or shorter than pereopod4...... 2 Outer lobes of lower lip notched apically or medially excavate, outer ramus of uropod 3 bearing one or two giant, hooked, articulate spines, rami flattened. (fig. 50) AMPITHOIDAE (p. 141)
1 Consult Aoridae if telson fleshy and pereopods 1-2 glandular.
116 U.S. NATIONAL MUSEUM BULLETIN 271
oo
10.
11.
12.
Outer lobes of lower lip not notched nor medially excavate, outer ramus of uropod 3 not bearing giant hooked spines (except Jassa), occasionally bearing small distal hook, or bent and serrate spine, rami lanceolate or styliform. ... aD aap. 3
. Peduncle of uropod 3 aien snail as Mlonere as or rlonee ‘hora podiums of uropod
2, rami of uropod 3 much shorter than peduncle, outer ramus of uropod
3 distally hooked (hook a giant spine in Jassa), or with minute distolateral denticles, pectinae or serrations (and see Parapherusa in Gammaridae).
(fig. 49) ISCHYROCERIDAE (p. 275)
Peduncle of uropod 3 usually much shorter than peduncle of uropod 2, or
rami of uropod 3 subequal to or longer than peduncle, or outer ramus of
uropod 3 not distally hooked and lacking distolateral ornamentation . 4
- Urosome depressed) 3 jeoi. she) SOON Oe Ce) Share ea earat
Urosome not depressed . . 7
. All urosomites coalesced, nropede ie 3 OP fehl divers Aine (Gig 53).
CHELURIDAE (p. 180) Urosomites rarely all coalesced, uropods 1-2 (but rarely 3) similar to each other and of normal gammaridean form. ............. 6
. Urosomite 1 elongate (fig. 52) . . .... =... . PODOCERIDAE (p. 426)
Urosomite 1 not elongate (fig. 51) .... . . . COROPHIIDAE (p. 184)
. Gnathopods 1-2 feeble, subequal to each cer! simple or nearly so, article 3
of mandibular palp vestigial (fig. 165e), article 1 of primary flagellum of antenna 1 elongate, plates of maxilliped foliaceous.
Synopia (Synopiidae) (p. 462)
Gnathopods 1-2 strong, usually dimorphic and subchelate, article 3 of man-
dibular palp strong, article 1 of primary flagellum of antenna 1 not elongate,
plates of: maxilliped not: foliaceous:isc 43 n) ee eS
. Gnathopod 1 larger than 2 (fig. 48) ...... SMA esi Gee oO)
Gnathopod 2 larger than 1 or gnathopods equalinsize. ....... 10
. Telson thick and fleshy, perepods glandular. . . . . . AORIDAE (p. 147)
Telson thin dorsoventrally, pereopods not glandular. Falklandella 2? (Gammaridae) (p. 241) Telson thick and fleshy, pereopods usually glandular. (fig. 47) ISAEIDAE (p. 264) Telson thin dorsoventrally, pereopods not glandular ......... Ii Accessory flagellum exceeding three articles. (exceptional) GAMMARIDAE (p. 231) Accessory flagellum 2-articulate or less, often absent ......... 12 Coxae 1-4 strongly or weakly acuminate midventrally or coxae midventrally excavate to form weak or strong anterior and posterior acuminations. (exceptional) PARAMPHITHOIDAE (p. 389) Coxae 1—4 rounded or quadrate ventrally. (exceptional) CALLIOPIIDAE (p. 167)
2 Schellenberg (1931) writes that coxa 4 is distinctly excavate posteriorly but no figure is given and such excavation must be deep and distinct to avoid placing a genus in section H.
MARINE GAMMARIDEAN AMPHIPODA 117
Acanthonotozomatidae
Ficures 54-56
Dracnosis.—Mouthparts grouped into cone-shaped bundle pro- jecting from ventral surface of head; accessory flagellum absent or vestigial. See Paramphithoidae, Ochlesidae, Pardaliscidae, La- fystidae.
Description.—Accessory flagellum absent, 1- or 2-articulate when present, very small and short; body usually with dorsal processes; rostrum well-developed; mouthparts rather linear in shape (to fit conical bundle and probably adapted for piercing and sucking); mandible especially linear, usually lacking molar or having only small molar protuberance, although triturative molar present in some genera; palp of mandible always present; lower lip variable, with or without inner lobes, apices of outer lobes occasionally incised; maxilliped with well-developed plates but palp with three or four articles and long or short; coxae 1-4 usually acuminate or subacuminate; gnathopod 1 and usually gnathopod 2 very feeble, often minutely chelate, article 3 of gnathopod 2 occasionally elongate; rami of uropod 3 longer than peduncle, rami flattened-lanceolate; telson entire or slightly cleft; article 2 of pereopods 3-5 usually with posterior teeth.
RELATIONSHIP.—The mandibular molar of Astyridae is larger than but not morphologically distinct from the protuberances occurring in some acanthonotozomatids; the accessory flagellum of astyrids is 1-articulate but long, and the mouthparts are not arranged in a conical bundle.
The Lafystidae have a 2-articulate maxillipedal palp, but otherwise the family is so similar to the Acanthonotozomatidae that the two families might be joined together.
The Stilipedidae resemble Acanthonotozomatidae but apparently the mouthparts are not grouped in a conical bundle; the maxillae are strongly foliaceous but this is true also of Mazilliphimedia.
With one exception the mouthparts of Pardaliscidae are not grouped in a conical bundle; usually pardaliscids have a multiarticulate accessory flagellum and the inner plates of the maxillipeds are obsoles- cent. One of the mandibles is especially flattened and heavily toothed.
The Stegocephalidae lack a mandibular palp.
Mouthparts of the Paramphithoidae are not arranged in a conical bundle but this distinction is confused by the acanthonotozomatid Bathypanoploea ausiralis (Chilton, 1912) (see Schellenberg, 1931). The mouthparts of B. australis not only are grouped in a quadrate bundle but the molar is absent and the gnathopods are styliform as in most acanthonotozomatids. The type genera of the two families differ in the following ways:
118 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 54.—Acanthonotozomatidae: a, Echiniphimedia hodgsont (Walker), original draw- ing; b, Acanthonotozoma cristatum (Ross) (Sars, 1895, pl. 131).
MARINE GAMMARIDEAN AMPHIPODA 119
Paramphithoe Acanthonotozoma Mouthparts bluntly bundled conically bundled Mandible stout substyliform Mandibular molar large, ridged obsolescent, conical Maxillae stout substyliform Gnathopods small, subchelate styliform
Intergradations of these conditions occur in the genera of both
families. If the sea yields more genera similar to Bathypanoploea, the fusion of the Paramphithoidae and Acanthonotozomatidae may be required.
Acanthonotozomella oates: K. H. Barnard (1930) is transferred to
Acanthonotozomoides.
10.
12.
13.
Key to the Genera of Acanthonotozomatidae
Ralproremaxillayl uniarticulate (fig! 56d)) 2. 2 Palp on maxilla biarticulates(fig.;56c)) - S24 Ses ee Gnathopod 2 slender and chelate (fig. 560). . . ......2.2.~. 3 Gnathopod 2 stout and subchelate (fig. 56p) ... . is fa Odius Palp article 2 of maxilliped strongly produced medially along article 3
(e562). Sas . . . . Pariphimedia Palp article 2 of raeeilleed not mroduced (fig. 56h) ess Barichinhime dis Palp of maxilla 1 very short, not reaching end of outer plate (fig. 56d) . . 5 Palp of maxilla 1 reaching end of outer plate (fig. 56c) ........7 Gnathopods 1 and 2 simple (figs. 56k,n). . . . . . . . Panoploeopsis
Gnathopod 1 minutely chelate (figs. 561, m), prachened 2 not simple. . 6 Palp article 2 of maxilliped produced along inside of third (fig. 562), lower
lip usually incised (fig. 56a) . . . . . . . Panoploea Palp artile 2 of maxilliped not produecd plone: neide of third (fig. 56g), lower lip not incised (fig. 566) . . . . . . . Anchiphimedia Mandible tapering to a smooth, pramenecnicall ecoopeshaned apex (fig. 55g). 8 Mandible not as above (e.g., fig. 55k) . 5 9 Upper lip rounded, not incised (fig. 55c); owes ie fet saciaedl Ge, 560). Labriphimedia
Upper lip quadrangular, incised (fig. 55d); lower lip incised (fig. 56a). Maoriphimedia Gnathopod simple: (figs 56/4)! sete ay oh ene ciacee e See not. enema ml \() Gnathopod 1 chelate (fig. 561)... ....... carte eens LS) Telson entire, rounded (fig. 56g) .. . se “Acanthonotozomoides Telson distally notched or emarginate (figs. 56r YS) Meena iinet ka |
Gnathopod 1 feeble, very slender, gnathopod 2 stouter ‘fhom i
Acanthonotozoma Gnathopods 1 and 2 similar to each other, well developed. . . . .. 12 Palp of maxilliped shorter than outer plate; mandible pointed (fig. 55l). Bathypanoploea Palp of maxilliped longer than outer plate; mandible distally broad (fig. 55k). Acanthonotozomella
Cutting edge of mandible drawn out in a long, needle-like apex (fig. 557). Parapanoploea
120 U.S. NATIONAL MUSEUM BULLETIN 271
Mandible not drawn out in a long, needle-like apex . . . . 14
14. Palp of maxilla 1 enormously expanded into an ovate leant Tne lip asymmetrically incised (fig. 55e), lower lip not incised (fig. 56b).
Maxilliphimedia
These characters not combined . . . BMH oo 55
15. Palp article 2 of maxilliped distinctly anedtnced siarays medial adlge af article
3 (fig. 562), outer lobes of lower lip ae and strongly notched (fig.
56a) Pe twe te CLG
Palp article 2 8 menlliped andlithaclliy or ner anedingedl allo medial edge
of article 3 (figs. 56g,h), outer lobes of lower lip unnotched or very weakly
mnotcheds. 23 va) ae sone tay IL 16. Pereonite 1 as long as est ihtce sans combined Ge 55m), mandibular incisor apparently absent... . . ... . . Cypsiphimedia Pereonite 1 not longer than each of next ines sesTnemti (fig. 55a), mandibular incisor strong . . . . . . Iphimedia! 17. Upper lip deeply merce iGo “BEO), Past wor articles of maxillipedal palp extraordinarily broadened (fig. 56f) ... . . . . Pseudiphimediella Upper lip entire or emarginate, first two articles of maxillipedal palp not broadened (fig. 56g) . . . . : eutinatat teres sit Ra ea NLS, 18. Some or all coxae with salnamaehiell teeth RU aaa ‘Echiniphimedia Coxae lacking submarginal teeth .. . Ree si Whe)
19. Primary cutting edge of mandible elndlentete Ges. ih, ®. Pariphimediella Primary cutting edge of mandible 2- or less dentate (fig. 551) . . . .. 20 20. Mandible short, stout, Pee cutting edge very broad, smooth, molar conical (fig. 557). ... . . . . . . Gnathiphimedia Mandible slender, long, primary arin ede tapering, subacute, molar if present, not conical (fig. 55/) ........... .. Iphimediella
1See also Panoploea spinosa Thomson (Hurley, 1954c) which keys here because of its scarcely shortened palp of maxilla 1.
Genera of Acanthonotozomatidae
Acanthonotozoma Boeck
Acanthonotus Ross, 1835 (homonym, Pisces). Acanthonotozoma Boeck, 1876 (new name for Acanthonotus) —Stebbing, 1906.
Type-species: Acanthonotus cristatus Ross, 1835 (original designa- tion). See Sars, 1895.
Upper lip incised; mandible with narrow, rounded apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching or exceed- ing end of outer plate; maxillipedal palp exceeding outer plate, 4-articulate, article 2 not produced; gnathopods simple, gnathopod 1 very slender, feeble, gnathopod 2 stouter than 1; telson apically incised. Species: 4, arctic-boreal, littoral (rarely to 700 m).
MARINE GAMMARIDEAN AMPHIPODA 121
Figure 55.—Acanthonotozomatidae: a, Odius carinatus (Bate) (Sars, 1895, pl. 133). Accessory flagellum: b, Iphimedia haurakiensis Hurley (1954c). Upper lip: c, Labriphi- media vespuccii K. H. Barnard (1932); d, Acanthonotozoma serratum (Fabricius) (Sars, 1895, pl. 131); e, Odius. Mandible: f, Acanthonotozoma; g, Labriphimedia, apex; h, Pariphimedia integricauda Chevreux (1906c); i, Gnathiphimedia mandibularis K. H. Barnard (1930); 7, Parapanoploea oxygnathia Nicholls (1938); k, Maoriphimedia hinemoa Hurley (1954c); 7, Odius. Front of body: m, Cypsiphimedia gibba K. H. Barnard (1955).
122 U.S. NATIONAL MUSEUM BULLETIN 271
Acanthonotozomella Schellenberg Acanthonotozomella Schellenberg, 1926a.
Type-species: A. alata Schellenberg, 1926a (monotypy).
Upper lip ?incised [‘‘eingekerbt’’]; mandible with narrow, rounded apex, minutely serrate; lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 4-articulate, article 2 not produced; gnathopods simple, alike; telson apically incised. Species: 1, antarctic, littoral.
Acanthonotozomoides Schellenberg Acanthonotozomoides Schellenberg, 1931.
Type-species: A. sublitoralis Schellenberg, 1931 (monotypy).
Upper lip incised; mandible with narrow, slightly toothed apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 4-articulate, article 2 slightly produced medially; gnathopods simple, alike; telson apically rounded, entire. Species: 2, antiboreal, littoral.
Anchiphimedia K. H. Barnard Anchiphimedia K. H. Barnard, 1930.
Type-species: A. dorsalis K. H. Barnard, 1930, 1932 (monotypy).
Upper lip incised; mandible with narrow, acute apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, very short; maxillipedal palp exceeding outer plate, 3-articulate, article 2 not produced; genathopod 1 minutely chelate, gnathopod 2 not simple; telson slightly incised. Species: 1, antarctic, bathyal (to 550 m).
Bathypanoploea Schellenberg
Epimeriopsis K. H. Barnard, 1931 (void ab znitzo).
Iphimediopsis Schellenberg, 1931 (homonym, not Della Valle, 1893).
Bathypanoploea Schellenberg, 1939 (footnote p. 137; new name for Iphimediopsis Schellenberg) .
Pseudiphimediopsis Ruffo, 1949.
Type-species: Acanthonotozoma australis Chilton, 1912 (monotypy). K. H. Barnard (1932, p. 182) noted that H’pimeriopsis is void ab initio because of its basis on a misidentified specimen of the designated type-species, A. australis Chilton.
Upper lip incised; mandible with broad, toothed apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp shorter than outer plate, 4-articulate, article 2 not produced; gnathopods simple, alike; telson emarginate. Species: 1, subantarctic, abyssal.
MARINE GAMMARIDEAN AMPHIPODA 123
Cypsiphimedia K. H. Barnard
Cypstphimedia K. H. Barnard, 1955.
Type-species: C. gibba K. H. Barnard, 1955 (original designation).
Upper lip incised; mandibular incisor feeble or ?absent; lobes of lower lip notched; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 3-articulate, article 2 medially produced along article 3; gnathopod 1 chelate, gnathopod 2 subchelate; pereonite 1 as long as next three segments combined; telson slightly emarginate. Species: 1, South Africa, littoral.
Ficure 56.—Acanthonotozomatidae: Lower lip: a, Iphimedia obesa Rathke (Sars, 1895, pl. 132); b, Acanthonotozoma serratum (Fabricius) (Sars, 1895, pl. 131). Maxilla 1: c, Acanthonotozoma; d, Odius carinatus (Bate) (Sars, 1895, pl. 133). Maxilla 2: e, Acantho- notozoma. Maxilliped: f, Pseudiphimediella nodosa (Dana) (Schellenberg, 1931); g, Labriphimedia vespuccur K. H. Barnard (1932); h, Acanthonotozoma; 1, Iphimedia; j, Pariphimedia integricauda Chevreux (1906c).° Gnathopod 1: k, Acanthonotozoma; 1, Iphimedia; m, Odius. Gnathopod 2: n, Acanthonotozoma; 0, Iphimedia; p, Odius. Tel- son: gq, Paritphimedia; r, Iphimedia; s, Acanthonotozoma; t, Odius. Uropod 3: u, Acanthonotozoma.
285-135 © = 69 - 9
124 U.S. NATIONAL MUSEUM BULLETIN 271
Echiniphimedia K. H. Barnard Echiniphimedia K. H. Barnard, 19380.
Type-species: Iphimedia hodgsont Walker (see 1907) (present selection).
Upper lip entire or slightly emarginate; mandible with narrow, rounded apex; lobes of lower lip not incised or minutely so; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 4-articulate, article 4 extremely minute, article 2 scarcely or not produced; gnathopods 1 and 2 chelate; telson with shallow, broad emargination; some or all coxae with submarginal teeth. Species: 3, antarctic, bathyal to littoral (to 824 m).
Gnathiphimedia K. H. Barnard Gnathiphimedia K. H. Barnard, 1930.
Type-species: G. mandibularis K. H. Barnard, 1930 (present selection).
Upper lip entire; mandible short, broad, apex smooth, molar conical; lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 3-articulate, article 2 indistinctly or not produced; gnathopod 1 chelate; telson ?emarginate. Species: 3, antarctic, bathyal (littoral) (to 824 m).
Iphimedia Rathke
Iphimedia Rathke, 1843.—Stebbing, 1906. Microcheles Kréyer, 1846.
Type-species: J. obesa Rathke, 1843 (monotypy). See Sars, 1895.
Upper lip slightly emarginate or truncate; mandible with medium- broad, slightly toothed apex; lobes of lower lip notched; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 3-articulate, article 2 produced along article 3; gnathopods both chelate or gnathopod 2 subchelate; telson incised or emarginate. Species: 8, tropical Pacific to antarctic, littoral (one bathyal).
Iphimediella Chevreux Iphimediella Chevreux, 1911e, 1912a, 1912b.
Type-species: J. marguerite. Chevreux 1912a (designated by Chevreux) ; 1912b.
Upper lip entire or weakly incised; mandible medium-broad, apex scarcely toothed; lobes of lower lip not incised; palp of maxilla 1
MARINE GAMMARIDEAN AMPHIPODA 125
biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 3-articulate, article 2 scarcely or not produced; gnathopods both chelate; telson cleft one third. Species: 4, antarctic, bathyal (110-457 m).
Labriphimedia K. H. Barnard
Labriphimedia K. H. Barnard, 1931.
Type-species: JL. vespuccw K. H. Barnard, 1931 (original designa- tion).
Upper lip entire; mandible with smooth, scoop-shaped, tapering apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, composed of three articles tipped with a very minute fourth, article 2 not produced; gnathopods both ?chelate (type-species not described) ; telson notched. Species: 2, antarctic, littoral.
Moaoriphimedia Hurley Maortphimedia Hurley, 1954c.
Type-species: M. hinemoa Hurley, 1954c (original designation).
Upper lip slightly incised; mandibular apex broad, untoothed, scoop-shaped; lobes of lower lip slightly incised apically; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 3-articulate, article 2 not produced; gnathopods both chelate; telson apically notched. Species: 1, New Zealand, littoral.
Maxilliphimedia K. H. Barnard
Mazilliphimedia K. H. Barnard, 1930.
Type-species: Iphimedia longipes Walker (see 1907) (monotypy).
Upper lip broad, incised; mandible with broad, toothed apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, article 2 enormously expanded; maxillipedal palp exceeding outer plate, 3-articulate, article 2 not produced; gnathopod 1 chelate, (gnathopod 2 not described) ; telson deeply notched. Species: 1, antarctic, bathyal, (183-379 m).
Odius Liljeborg
Otus Bate, 1862 (homonym, Lepidoptera). Odius Liljeborg, 1865.—Stebbing, 1906.
Type-species: Otus carinatus Bate, 1862 (monotypy). Upper lip narrow, incised; mandible with narrow, dentate apex; lobes of lower lip not incised; palp of maxilla 1 uniarticulate, tiny,
126 U.S. NATIONAL MUSEUM BULLETIN 271
not reaching end of outer plate; maxillipedal palp exceeding outer plate, 4-articulate, article 2 not produced; gnathopod 1 slender, chelate, gnathopod 2 stout, subchelate; telson notched. Species: 2, arctic-boreal, N. Atlantic, littoral.
Panoploea Thomson
Panoploea Thomson, 1880.—Stebbing, 1906. Iphimediopsis Della Valle, 1893.
Type-species: P. spinosa Thomson, 1880 (present selection) ; Chevreux and Fage (1925) erroneously cite as type-species, Iphimedia eblanae Bate (1857d), a species not included originally in the genus by Thomson.
Upper lip narrow, ?incised or not; mandible with very narrow, scarcely dentate apex; lobes of lower lip incised or not; palp of maxilla 1 biarticulate, not reaching end of outer plate; maxillipedal palp exceeding outer plate, 3-articulate, article 2 produced medially; gnathopod 1 chelate, gnathopod 2 slightly chelate or subchelate; telson broadly or minutely incised or cleft. Species: 8, amphiboreal, littoral (one in bathyal antarctic to 550 m).
Panoploeopsis Kunkel Panoploeopsis Kunkel, 1910.
Type-species: P. porta Kunkel, 1910 (monotypy).
Upper lip narrow, possibly not incised; mandible with narrow, dentate apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, not reaching end of outer plate; maxillipedal palp barely exceeding outer plate, 3-articulate, article 2 slightly produced medially; gnatho- pods simple; telson bilobate. Species: 1, Bermuda, littoral.
Paranchiphimedia Ruffo Paranchiphimedia Ruffo, 1949.
Type-species: P. monodi Ruffo, 1949 (original designation).
Upper lip deeply emarginate; mandible subpyramidal, incisor un- toothed; [lobes of lower lip unknown]; palp of maxilla 1 uniarticulate, short; maxillipedal palp exceeding outer plate, 3-articulate, article 2 not produced; gnathopods chelate; telson emarginate. Species: 1. antarctic, ?littoral.
MARINE GAMMARIDEAN AMPHIPODA 127
Parapanoploea Nicholls
Parapanoploea Nicholls, 1938.
Type-species: P. orygnathia Nicholls, 1938 (original designation).
Upper lip broad, faintly emarginate; mandible with needle-like apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 4-articulate, article 2 slightly produced medially; gnathopods chelate; telson notched. Species: 1, antarctic (220 m).
Pariphimedia Chevreux
Pariphimedia Chevreux, 1906a.
Type-species: P. integricauda Chevreux, 1906a (original designa- tion).
Upper lip broad, faintly emarginate; mandible with narrow, dentate apex; lobes of lower lip not incised; palp of maxilla 1 uniarticulate, not reaching end of outer plate; maxillipedal palp exceeding outer plate, 3-articulate or with minute article 4, article 2 strongly produced medially; gnathopods chelate; telson entire. Species: 2, antarctic, littoral.
Pariphimediella Schellenberg
Pariphimediella Schellenberg, 1931.
Type-species: Iphimedia serrata Schellenberg, 1926a (original designation).
Upper lip broad, faintly emarginate; mandible with narrow, toothed apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 3-articulate, article 2 faintly produced; gnathopods chelate; telson broadly emarginate or cleft. Species: 5, antarctic, littoral to bathyal.
Pseudiphimediella Schellenberg Pseudiphimediella Schellenberg, 1931.
Type-species: Amphitoé nodosa Dana, 1853 (original designation).
Upper lip broad, incised; mandible with broad, toothed apex; lobes of lower lip not incised; palp of maxilla 1 biarticulate, reaching end of outer plate; maxillipedal palp exceeding outer plate, 3-articulate, article 2 not produced, articles 1-2 broadened; gnathopod 1 chelate, enathopod 2 slightly chelate; telson entire or faintly emarginate. Species: 1, subantarctic, littoral.
128 U.S. NATIONAL MUSEUM BULLETIN 271
Ampeliscidae
Ficures 57, 58
Diaenosis.—Accessory flagellum absent; pereopod 5 shorter than and of different structure from 4; article 4 of pereopods 1—2 elongate; head elongate; eyes when present bearing maximum of two pairs of anterolateral cuticular lenses; urosomites 2—3 coalesced; pereopods 1-2 glandular. See Atylidae, Haustoriidae, Argissidae.
Description.—Accessory flagellum absent; head elongate, tall, compressed; body lacking dorsal processes, except on pleonite 4 and occasionally low carinae on pleonites 1-3; antennae and their pe- duncles elongate, often with long setae, male primary flagellum conjoint basally ; anterior coxae long, coxa 1 often broader than 2, 2 occasionally tapering; mouthparts basic but mandibular palp article 3 of many species much shorter than article 2 and mandibular lobes of lower lip obsolescent; gnathopods feeble, subchelate or nearly simple; article 4 of pereopods 1-2 elongate; pereopod 5 always shorter and of different morphology from pereopod 4; pereopods with a few elongate setae, especially on article 2 of pereopod 5 and article 4 of pereopods 1-2, pereopods 3-4 often with strong submarginal spines on articles 4-5; rami of uropod 3 lanceolate, usually elongate, occasionally foliaceous or shortened; telson either elongate or very short, usually cleft, cleft often short.
RELATIONSHIP.—The shiny cuticular lenses of oculate ampeliscids are unique to this family, except for their rare occurrence in a few lysianassids. Oculate ampeliscids also have bright red, brown, or black pigmentary masses in the head towards which, perhaps, the cuticular lenses direct light.
Pereopods of this family are definitively fossorial but presumably they are used not in digging but in clinging to or rearranging the insides of their tubes spun by pereopods 1-2 or in creating water currents and feeding.
The Phoxocephalidae always bear a rostrum and a multiarticulate accessory flagellum. The Haustoriidae bear a multiarticulate accessory flagellum. Both of those families form burrows in sediments whereas Ampeliscidae are provided with glands in the pereopods for the construction of tubes lying on the substrate surface.
The Atylidae have ommatidial eyes and pereopod 5 is usually longer than pereopod 4 and of similar structure; the pereopods are not glandular.
MARINE GAMMARIDEAN AMPHIPODA
129
IN &
Sa >
N \
SNS
Figure 57.—Ampeliscidae: a, Ampelisca tenuicornis Liljeborg (Sars, 1895, pl. 58); b, Byblisoides arcillis J. L. Barnard (1961).
130 U.S. NATIONAL MUSEUM BULLETIN 271
Key to the Genera of Ampeliscidae
1. Pereopod 5: anterior edge of posterior lobe on article 2 lacking setae near its
junction with article 3 (fig. 581)... . . ERP henge 102) Pereopod 5: anterior edge of posterior lobe on amills 2 penne setae up to
its junction with article 3 (figs. 58m,n,o) . . 4
2. Flagella of antennae poorly developed, with 2 46 4 amiclles Ge. 57b) [eniene- ventral corner of head produced (fig. 57b)] . .. . =... . Byblisoides Flagella of antennae long, usually with more than 10 articles [anteroventral cornerorheadirarelyjproduced| aes) sites e alee eS
3. Anteroventral corner of head produced. ... . Joon ee eee Oo dos Anteroventral corner of head sloping, Anorodecdt | .... . . Ampelisca
4. Posterior lobe of article 2 on pereopod 5 strongly expanded distally, posterior edge oblique (fig. 58m) ...... Ale By Dlis Posterior lobe of article 2 on pereopod 5 noe etoamdledl dis! ally, posterior edge vertical (igs. 58730) 2 2 iss a ee ee enloons
Genera of Ampeliscidae
Ampelisca Kr¢yer
Ampelisca Krgyer, 1842.—Stebbing, 1906. Pseudophthalmus Stimpson, 1853. Araneops Costa, 1853c, 1857.
Tetromatus Bate, 1856, 1857a, 1857c.
Type-species: A. eschrichtia Krgyer, 1842 (monotypy). See Sars, 1895.
Pereopod 5 with posterior lobe of article 2 greatly expanded distally, posterior edge oblique, anterior edge of posterior lobe lacking setae near its junction with article 3; palp article 3 of mandible variable in length; antenna 2 with more than five flagellar articles; anteroventral corner of head unproduced. Species: 94, primarily littoral cosmo- politan, about 20 bathyal and one abyssal species.
Byblis Boeck Byblis Boeck, 1871.—Stebbing, 1906.
Type-species: Ampelisia (lapsus) gaamardu Krgyer, 1846 (mono- typy). See Sars, 1895.
Pereopod 5 with posterior lobe of article 2 greatly expanded distally, posterior edge oblique, anterior edge of posterior lobe bearing setae near its junction with article 3; palp article 3 of mandible shorter than article 2; antenna 2 with more than five flagellar articles; antero- ventral corner of head unproduced. Species: 21, cosmopolitan, littoral to abyssal.
MARINE GAMMARIDEAN AMPHIPODA 131
Soa
Ficure 58.—Ampeliscidae: Upper lip: a, Ampelisca typica (Bate) (Sars, 1895, pl. 57): Mandible: b, Ampelisca; c, Byblis gaimardi (Kr@yer) (Sars, 1895, pl. 64). Lower lip: d, Ampelisca; e, Byblis. Maxillae 1-2: f,g, Ampelisca. Maxilliped: h, Ampelisca. Gnathopods 1-2, Pereopods 1,5: 1,7,k,/, Ampelisca. Pereopod 5: m, Byblis; n, Haploops setosa Boeck (Sars, 1895, pl. 68); 0, Haploops tubicola Liljeborg (Sars, 1895, pl. 67). Uro= pod 3: p, Ampelisca. Telson: g, Ampelisca; r, Byblis.
132 U.S. NATIONAL MUSEUM BULLETIN 271
Byblisoides K. H. Barnard
Byblisoides K. H. Barnard, 1931, 1932.
Type-species: B. juaticorns K. H. Barnard, 1931 (original designation).
Pereopod 5 with posterior lobe of article 2 greatly expanded distally, posterior edge oblique, anterior edge of posterior lobe lacking setae near its junction with article 3; palp article 3 of mandible shorter than article 2; antenna 2 with less than five flagellar articles; anteroventral corner of head produced. Species: 4, tropics to antarctic, bathyal.
Haploops Liljeborg Haploops Liljeborg, 1856.—Stebbing, 1906.
Type-species: H. tubicola Liljeborg, 1856 (original designation). See Sars, 1895.
Pereopod 5 with posterior lobe of article 2 not expanded distally, usually narrow, posterior edge vertical, anterior edge of posterior lobe bearing setae near its junction with article 3; palp article 3 of mandible as long as article 2; antenna 2 with more than five flagellar articles; anteroventral corner of head unproduced. Species: 13, cosmopolitan, cold-water, littoral to abyssal, primarily bathyal.
Triodos K. H. Barnard Triodos K. H. Barnard, 1916.
Type-species: 7. insignis K. H. Barnard, 1916 (monotypy).
Pereopod 5 with posterior lobe of article 2 greatly expanded distally, posterior edge oblique, anterior edge of posterior lobe lacking setae near its junction with article 3; palp article 3 of mandible as long as article 2; antenna 2 with more than five flagellar articles; anteroventral corner of head produced. Species: 1, S. Africa, littoral.
Amphilochidae
Fiaures 59, 60
Dracnosis.—Accessory flagellum absent; coxa 1 very small, par- tially hidden by following coxae. See Pleustidae, Calliopiidae, Leuco- thoidae, Anamixidae, Stenothoidae, Thaumatelsonidae, Cressidae, Stegocephalidae.
Description.—Accessory flagellum absent; rostrum conspicuous; coxa 1 very small in all but one genus and partially hidden by fol-
134 U.S. NATIONAL MUSEUM BULLETIN 271
lowing coxae, coxa 2 occasionally as small as coxa 1 but often as large as coxa 3 and not hidden; coxae 3-4 enlarged, either overlapping or, when immensely enlarged, with contiguous margins abutting; upper lip usually deeply incised, rarely slightly excavate; mandible usually with 3-articulate palp, rarely absent, molar well developed and trit- urative or formed of a smooth protuberance or evanescent; lower lips of two kinds (see figures); maxilla 1 with 1- or 2-articulate palp; maxilla 2 rarely reduced in size, in one genus formed of a single plate; maxillipeds normal; gnathopods of medium size or small, subchelate or nearly simple, often incompletely carpochelate; uropod 2 shortened ; uropod 3 biramous, peduncle elongate (except Pseudamphilochus) ; telson entire in all but one genus, often elongate and triangular, also short and linguiform.
RELATIONSHIP.—The Pleustidae and Calliopiidae have large first coxae, not hidden by following coxae. Leucothoidae have completely carpochelate first gnathopods because article 6 is narrow.
The Stegocephalidae always lack a mandibular molar and palp; no amphilochid lacks both at the same time; the mouthparts of stego- cephalids project in a conical bundle, an accessory flagellum is present and the first coxa is never hidden by the following coxae.
The Stenothoidae, Thaumatelsonidae, and Cressidae have uni- ramous third uropods.
The Anamixidae have a completely carpochelate first gnathopod or the appendage is absent, and a ventral cephalic keel replaces the mandibles and maxillae.
Pseudamphilochus Schellenberg is an aberrant amphilochid in its normal coxa 1 and cleft telson. It tends to fall into the Eusiridae in the various keys but a special place has been made for it also in Gammaridea Family Key, Section G. It also appears to be a pleustid with cleft telson even though its lower lip is not fully typical of pleustids. It can also be confused with liljeborgiids even though the accessory flagellum is absent in Pseudamphilochus.
Incomplete carpochelation of gnathopods is ignored in the keys and diagnoses to follow and the terms “simple” and ‘‘subchelate”’ refer only to the condition of the propodus (article 6).
Neocyproidea peninsulae Hurley (1955) is removed to Peltopes.
Key to the Genera of Amphilochidae
1. Coxae 3-4 not immensely broadened, with contiguous margins overlapping, not concealing coxa 2 (fig. 59a) . . . (Amphilochinae, new subfamily) 2 Coxae 3-4 immensely broadened, contiguous margins abutting, concealing the vestigial first two coxae (fig. 59b) . (Cyproideinae, new subfamily) 10
2. Mandibular molar large, with ridged and toothed triturating surface (fig. COG re ABER EE TN et CAN Ca area tea ab En GTS CO LE EARS ee a 3
10.
12.
13.
14.
15.
16.
MARINE GAMMARIDEAN AMPHIPODA 135
Mandibular molar small, or absent, unarmed or bearing one to three spines
ieGOd) ae is |: BOA Be teny te Se" Ab. SRD terete at 0 Palp of maxilla 1 ith ‘ares Pericles (fig. 60;) . ...... . . Gitanopsis Palp of maxilla 1 with one article (fig. 60k) ...... ee ee A: Outer plate of maxilliped strongly excavate medially, prticle 1 of palp much
longer than other palp articles (fig. 60p) ... . . . . Gitanogeiton Outer plate of maxilliped straight or slightly oun medially, article 1 of
palpssubequal-to article!2 (figiGQo) 9) Vrs 259° IR) Garg foc s Rob Gnathopod 2 large, subchelate (fig. 60v) . ... . .. . Amphilochopsis Gnathopod 2 small, nearly simple (fig. 60w) . . . . 3 . . Gitana Maxilla 2 composed of only one elongate plate (fig. 60m) . : Agauhiloehelia Maxilla 2 composed of two plates (fig. 60/) .... ape N, Maxilla 2 degraded, plates tiny, subequal in width Ge 60n) . Ry eS Maxilla 2 normal, inner plate much broader than outer (fig. 60/1) . . . . 9 Posterolateral angles of pleonite 6 not produced. . . . . Amphilochoides Posterolateral angles of pleonite 6 produced, reaching apex of telson. Cyclotelson Melsonventire) (fig. G0z) . -oegek 2 »sgease «>. ev.) Amphilochus Telson.cleft (fig; 60y).... =»... ... .. . . Pseudamphilochus Article 2 of pereopods 4-5 flac. Blender Shires: SA cep cai srige al Article 2 of pereopod 5 and usually pereopod 4 sanded sa Rue Drentas Ss Palm of gnathopod 2 transverse, urosomite 3 vaulting over telson, telson
Smallest ser hae ... . . Cyproidea Palm of gnathopod 2 oblique TOSS 3 not ~sanlGing over telson, telson
NWS eR Ay ee: i de une ee ay IP aracyproiged' Urosomite 1 wealeacled, toner or hast Soh NALCO OBEN ERENT a arrue Lem pa aeea NA ka) Urosomite 1 dorsally keeled, elongate . . . eee eee Gnathopod 2 simple; uropod 2 shortened, not senrahtaye end of uropod 3
(igse 590,0)) i) 2) fa ... . . Stegoplax Gnathopod 2 subchelate, ogi ene alan mroparl 2 reaching end of
uropod 3 (figs. 60bb,cc) . . . Rie lege: Se ls SPeltocoxa Article 2 of pereopod 3 slender, tere Bee Serna Par ca FO ae anes ae oe MON al 10) Article 2 of pereopod 3 expanded .... . Hie dee tar mae Sal 6)
Palp of maxilla 1 biarticulate, outer plate of card lived nendatlag to end of palp article 2, article 7 of gnathopod 1 apparently fitting oblique palm.
Hoplopleon
Palp of maxilla 1 uniarticulate, outer plate of maxilliped reaching to end of
palp article 1, article 7 of gnathopod 1 greatly overlapping short, trans-
verse palm ... . ee ew ww ee es .). . . Hoplopheonoides Mandibular palp cen PR aete Ceotan Sonat ae tae See ger obese Pe LLODeS Mandibular palp absent .............. ... Neocyproidea
Genera of Amphilochidae
Subfamily Amphilochinae, new subfamily
Type-genus: Amphilochus Bate. Coxae 2-4 not immensely broadened, with contiguous margins
overlapping.
i36). U.S. NATIONAL MUSEUM BULLETIN 271
Amphilochella Schellenberg Amphilochella Schellenberg, 1926a.
Type-species: <A. simplicarpus Schellenberg, 1926a (monotypy).
Mandibular molar absent; palp of maxilla 1 biarticulate; maxilla 2 composed of one elongate plate; outer plates of maxillipeds not excavate; palp article 1 shorter than article 2; gnathopod 2 small, slender, scarcely subchelate; telson apparently entire. Species: 1, antarctic, littoral.
Amphilochoides Sars Amphilochoides Sars, 1895.—Stebbing, 1906.
Type-species: Amphilochus odontonyx Boeck, 1871 (original desig- nation). This species is figured by Sars erroneously under his Amphilo- choides pusillus which becomes a synonym of A. odontonyx; Sars’ description and figures of A. odontonyx are proposed by him on p. 690 to have the name A. boeckiit which Stebbing (1906) finds is a junior synonym of Probolium serratipes Norman (1869a).
Mandibular molar obsolete; palp of maxilla 1 biarticulate; lower lip atypical for family, with tilted oval outer lobes astride distinct inner lobes; maxilla 2 degraded but with two distinct subequal lobes; outer plates of maxillipeds not excavate, palp article 1 subequal to article 2; gnathopod 2 large, subchelate; telson entire. Species: 4, N.E. Atlantic, littoral.
Amphilochopsis Stephensen Amphilochopsis Stephensen, 1925a.
Type-species: A. hamatus Stephensen, 1925a (monotypy).
Mandibular molar large, triturative; palp of maxilla 1 uniarticulate; maxilla 2 normal, outer plate narrower than inner; outer plates of maxillipeds not excavate, palp article 1 subequal to article 2; gnathopod 2 large, subchelate; telson entire. Species: 1, subarctic, bathyal to abyssal (825-2702 m).
Amphilochus Bate
Amphilochus Bate, 1862.—Stebbing, 1906. Callimerus Stebbing, 1876. Type-species: A.manudens Bate, 1862 (monotypy). See Sars, 1895. Mandibular molar small, not triturative; palp of maxilla 1 biarticu- late; maxilla 2 normal, outer plate narrower than inner; outer plates of maxillipeds not excavate, palp article 1 subequal to article 2; enathopod 2 large, subchelate; telson entire. Species: 11, cosmo- politan, littoral (one species to 913 m).
MARINE GAMMARIDEAN AMPHIPODA ey
Ficure 60.—Amphilochidae: Upper lip: a, Amphilochus manudens Bate (Sars, 1895, pl. 74); b, Pseudamphilochus shoemakeri Schellenberg (1931). Mandible: c, Gitanopsis bispinosa (Boeck) (Sars, 1895, pl. 76); d, Amphilochus; e, Neocyproidea otakensis Hurley (1955); f, Pseudamphilochus. Lower lip: g, Gitanopsis; h, Amphilochoides odontonyx (Boeck) (Sars, 1895, pl. 76); 1, Amphilochus. Maxilla 1:7, Amphilochus; k, Gitana sarsi Boeck (Sars, 1895, pl. 78). Maxilla 2: 1, Amphilochus; m, Amphilochella simplicarpus Schellenberg (1926a); », Amphilochoides. Maxilliped: 0, Amphilochus; p, Gitanogeiton sarsi Stebbing (1910). Gnathopod 1: g, Amphilochus; r, Gitana; 5, Hoplopheonotdes obesa Shoemaker (1956a); t, Peltopes productus K. H. Barnard (1930). Gnathopod 2: u, Gitana; v, Amphilochus; w, Hoplophenonoides. Telson: x, Amphilochus; y, Pseudam- philochus; x, Cyclotelson purpureum Potts (1915). Uropod 3: aa, Amphilochus. Pleon and Telson, lateral: bb, Hoplopheonoides; cc, Peltocoxa marioni Catta (Chevreux and Fage, 1925).
138 U.S. NATIONAL MUSEUM BULLETIN 271
Cyclotelson Potts Cyclotelson Potts, 1915.
Type-species: C. purpureum Potts, 1915 (monotypy).
Mandibular molar absent; palp of maxilla 1 biarticulate; maxilla 2 degraded but with two distinct subequal lobes; outer plates of maxilli- peds not excavate, palp article 1 subequal to article 2; gnathopod 2 small, subchelate; telson entire; lateral angles of urosomite 3 strongly produced (combining character). Species: 1, Gulf of Mexico, littoral.
Gitana Boeck Gitana Boeck, 1871.—Stebbing, 1906.
Type-species: G. sarst Boeck, 1871 (designated by Sars, 1895, p. 229, ‘‘Remarks’’).
Mandibular molar large, triturative; palp of maxilla 1 uniarticulate; maxilla 2 normal, outer plate narrower than inner; outer plates ‘of maxillipeds slightly excavate, palp article 1 subequal to article 2; gnathopod 2 small, nearly simple; telson entire. Species: 4, boreal, littoral to bathyal (475 m).
Gitanogeiton Stebbing Gitanogeiton Stebbing, 1910.
Type-species: G. sarsi Stebbing, 1910 (monotypy).
Mandibular molar large, triturative; palp of maxilla 1 uniarticulate; maxilla 2 normal, outer plate narrower than inner; outer plates of maxillipeds strongly excavate, palp article 1 much longer than other articles; gnathopod 2 small, subchelate [telson damaged]. Species: 1, off Manning R., Australia, littoral.
Gitanopsis Sars Gitanopsis Sars, 1895.—Stebbing, 1906.
Type-species: Amphilochus bispinosus Boeck, 1871 (original desig- nation).
Mandibular molar large, triturative; palp of maxilla 1 biarticulate; maxilla 2 normal, outer plate narrower than inner; outer plates of maxilliped rarely excavate, palp article 1 subequal to article 2; enathopod 2 small, subchelate; telson entire. Species: 12, cosmo- politan, littoral (2 species in bathyal to 1096 m).
MARINE GAMMARIDEAN AMPHIPODA 139
Pseudamphilochus Schellenberg Pseudamphilochus Schellenberg, 1931.
Type-species: P. shoemakert Schellenberg, 1931 (monotypy).
Mandibular molar obsolete; palp of maxilla 1 biarticulate; maxilla 2 normal, outer plate narrower than inner; outer plates of maxilliped not excavate, palp article 1 subequal to article 2; coxa 1 not reduced; enathopod 2 small, subchelate; telson deeply cleft. Species: 1, S. Georgia Is., littoral.
Subfamily Cyproideinae, new subfamily
Type-genus: Cyproidea Haswell. Coxae 3-4 immensely broadened, with contiguous margins abutting, hiding coxae 1 and 2.
Cyproidea Haswell
Cyproidea Haswell, 1880a.—Stebbing, 1906. Cyproidia [sic] Haswell, 1880c. Gallea Walker, 1904.
Type-species: Cyproidia [sic] ornata Haswell, 1880c (present selection). See Walker, 1904 (as Gallea tecticauda).
Article 2 of pereopods 3-5 linear, slender; gnathopod 2 subchelate, palm transverse; uropod 2 long; urosomite 1 not elongate, not keeled; urosomite 3 vaulting over telson; telson normal; mandibular molar absent or present, palp 3-articulate. Species: 2, tropics to antiboreal, EK. Hemisphere, littoral.
Hoplopheonoides Shoemaker Hoplopheonoides Shoemaker, 1956a.
Type-species: H. obesa Shoemaker, 1956a (monotypy).
Article 2 only of pereopod 5 expanded, of pereopods 3-4 slender and linear (Hoplopleon); gnathopod 2 subchelate, tending to be chelate; uropod 2 long; urosomite 1 elongate, dorsally keeled; urosomite 3 not vaulting over telson; telson normal; mandibular molar well developed, triturative, palp absent; palp of mazilla 1 uniarticulate (Hoplopleon). Species: 1, Florida, littoral. Hoplopleon similis Schellenberg is inter- mediate between this and the next genus.
285-135 O - 69 - 10
140 U.S. NATIONAL MUSEUM BULLETIN 271
f Hoplopleon K. H. Barnard Hoplopleon K. H. Barnard, 1932.
Type-species: Peltocoxa australis K. H. Barnard, 1916 (original designation).
Article 2 of pereopods 4-5 expanded, of pereopod 3 slender (Hoplo- pheonoides); gnathopod 2 subchelate, palm oblique or transverse; [uropod 2 unknown]; urosomite 1 elongate, keeled; urosomite 3 not vaulting over telson; telson normal; mandibular molar triturative, palp absent; palp of maxilla 1 biarticulate (Hoplopheonoides). Species: 3, antarctic to S. W. Africa, littoral.
Neocyproidea Hurley Neocyproidea Hurley, 1955.
Type-species: Cyproidea otakensis Chilton, 1900 (original designation).
Article 2 of pereopods 3-5 somewhat expanded; gnathopod 2 nearly simple; uropod 2 long; urosomite 1 elongate, dorsally keeled, slightly vaulting to base of telson or less; urosomite 3 not vaulting over telson; telson normal; mandibular molar strong, palp absent. Species: 2, New Zealand, littoral.
Paracyproidea Stebbing Paracyproidea Stebbing, 1899d, 1906.
Type-species: Cyproidia [sic] lineata Haswell, 1880c (original designation).
Article 2 of pereopods 3-5 slender, linear; gnathopod 2 subchelate, palm oblique; uropod 2 long; urosomite 1 not elongate, not keeled; urosomite 3 not vaulting over telson; telson huge; mandibular molar triturative, palp 3-articulate. Species: 1, E. Australia, littoral.
Peltocoxa Catta Peltocoza Catta, 1875.—Stebbing, 1906.
Type-species: P. marioni Catta, 1875 (original designation). See Chevreux and Fage, 1925.
Article 2 of pereopods 4-5 expanded, of pereopod 3 slender and linear; gnathopod 2 subchelate, palm transverse; uropod 2 long; urosomite 1 short, unkeeled; urosomite 3 not vaulting over telson; telson huge; mandibular molar triturative, palp 3-articulate. Species: 2, E. Atlantic, littoral.
MARINE GAMMARIDEAN AMPHIPODA 141
Peltopes K. H. Barnard Peltopes K. H. Barnard, 1930.
Type-species: P. productus K. H. Barnard, 1930 (monotypy).
Article 2 of pereopods 3-5 expanded; gnathopod 2 simple; uropod 2 long; urosomite 1 elongate, dorsally keeled, process vaulting over following segments; urosomite 3 not vaulting over telson; telson normal; mandibular molar [unknown in type] strong, palp 3-articulate. Species: 2, New Zealand, littoral.
Stegoplax Sars Stegoplax Sars, 1882.—Stebbing, 1906.
Type-species: SS. longirostris Sars, 1882 (original designation). See Sars, 1895.
Article 2 of pereopods 3-5 expanded; gnathopod 2 simple; uropod 2 shortened; urosomite 1 short, unkeeled; urosomite 3 slightly vaulting over telson; telson normal; mandibular molar triturative, palp 3- articulate. Species: 1, N. Atlantic, bathyal.
Ampithoidae
FIGURE 61
Draenosis.—Accessory flagellum present or absent; telson entire, short, fleshy ; lower lip with anterior lobes notched or medially excavate (figs. 61e,f); coxa 4 not excavate posteriorly; pereopods glandular; uropod 3 with very small, short rami, shorter than peduncle, rami quadrate, blunt, outer armed with one or two hooks. See Isaeidae, Ischyroceridae, Oedicerotidae, Calliopiidae, Pleustidae.
Description.—Accessory flagellum varying from absent to long and multiarticulate; body smooth; rostrum absent; coxae medium in size, quadrate or rounded, coxa 4 not excavate posteriorly; mouthparts basic except for one genus lacking mandibular palp and another with reduced molar; lower lip with lateral (outer) lobes projecting, notched (bilobed) or medially excavate; gnathopods usually powerful, sub- chelate, in one genus feeble and chelate, gnathopod 1 usually smaller than 2, occasionally larger than 2; uropod 3 with very short quadrate rami, shorter than peduncle, outer ramus armed with one or two hooks; telson short, entire, nearly circular, square or triangular, fleshy.
ReELATIONSHIP.—The notched outer lobes of the lower lip distinguish this family from its relatives, the Isaeidae, Aoridae, Ischyroceridae, and Corophiidae. The very short quadrate rami with a hook or hooks on the outer are distinctive (but see Jassa in Ischyroceridae).
142 U.S. NATIONAL MUSEUM BULLETIN 271
The Ischyroceridae also have short rami but the peduncle is more elongate, the rami are superficially lanceolate and poorly setose, and the outer is armed with denticles, not large hooks (except Jassa).
The genus Amphitholina Ruffo was transferred to Eophliantidae by Gurjanova (1938) ; it bridges these two families in its intersimilar third
Ficure 61.—Ampithoidae: a, Ampithoe rubricata (Montagu) (Sars, 1895, pl. 206). Upper lip: b, Ampithoe. Mandible: c, Ampithoe; d, Sunamphitoe pelagica (Milne Edwards) (Sars, 1895, pl. 208, as S. conformata). Lower lip: e, Ampithoe; f, Pleonexes gammaroides (Bate) (Sars, 1895, pl. 207). Maxillae 1-2, Maxilliped: g,h,i, Ampithoe. Gnathopod 1: 3, Ampithoe; k, Macropisthopus stebbingi K. H. Barnard (1916). Gnathopod 2: J, Ampithoe. Pereopod 5: m, Ampithoe; n, Pleonexes. Uropod 3: 0, Ampithoe. ‘Telson: p, Ampithoe; q, Pleonexes.
MARINE GAMMARIDEAN AMPHIPODA 143
uropods, but the degeneration of mouthparts and coxae and the sub- cylindrical body mark it as an eophliantid.
The shape of uropod 3 distinguishes the Ampithoidae from the Calliopiidae, Pleustidae, and Oedicerotidae.
Nomenclatural Changes in Ampithoidae
Amphitholina Ruffo (1953) is removed to Kophliantidae. Amphithoides patrizi. Maccagno (1936) is removed to Cymadusa.
Key to the Genera of Ampithoidae
1. Mandible lacking palp (fig. 61d) .......... .. . . Sunamphitoe Mandibleypearimp=palp (fig. Gillie): 2k on SA aoe, Se a ARN ZepAntenna,l lacking accessory flagellum: 64 os si. hee. ). Bole e. 8 Antenna | bearing accessory flagellum. . . pla har ene eens arse eee 0) 3. Gnathopod 1 larger and stouter than apetingned oy. . . . . . Exampithoe Gnathopod 1 smaller and more slender than gnathopod2 ....... 4 4. Gnathopods feeble, chelate (fig. 61k). . . ... =. +... . .Macropisthopus Gnathopods large, subchelate (figs. 617,1). . . . 5
5. Article 6 of pereopods 3-5 widened apically, siripemalieastlle, Gis. 61n), ‘iin bearing large striated spines, telson with apical pair of reverted, elongate, cornified processes (fig. PUD) Aiea etre . .. . . Pleonexes
Article 6 of pereopods 3-5 widened or mee Pidened rarely subprehensile (fig. 61m), apical cornified processes of telson if present, obsolescent (fig.
Gil) te Panis Si nane me ages Jom eon AIM DIthOe
6. Outer ramus of ameyoudl 3 Beane one Thaliels ..... . . . Amphithoides Outer ramus of uropod 3 bearing two hooks (fig. 6lo) ......... 7
7. Gnathopod 1 larger than gnathopod 2 ......... .. . Paragrubia Gnathopod 1 smaller than gnathopod. ........... . Cymadusa
Genera of Ampithoidae
Amphithoides Kossmann Amphithoides Kossmann, 1880.—Stebbing, 1906.
Type-species: A. longicornis Kossmann, 1880 (monotypy).
Antenna 1 with accessory flagellum; mandible with palp; gnatho- pods large, subchelate, gnathopod 2 equal to or larger than 1; article 6 of pereopods 3-5 not apically widened; outer ramus of uropod 3 with one hook. Species: 1, Red Sea, littoral.
Ampithoe Leach
Ampithoe Leach, 1814a.—Stebbing, 1906.
Type-species: Cancer (Gammarus) rubricatus Montagu, 1808 (monotypy). See Sars, 1895.
144 U.S. NATIONAL MUSEUM BULLETIN 271
Antenna 1 lacking accessory flagellum; mandible with palp; gnatho- pods large, subchelate, gnathopod 2 equal to or larger than 1; article 6 of pereopods 3-5 scarcely widened apically, rarely prehensile (Pleo- nexes); outer ramus of uropod 3 with two hooks; apical cornified processes of telson, wf present, obsolescent (Pleonexes). Species: 35, cosmopolitan, littoral.
Cymadusa Savigny
Cymadusa Savigny, 1816. Grubia Czerniavski, 1868. Acanthogrubia Stout, 1912.
Type-species: OC. filosa Savigny, 1816 (monotypy). See Chevreux and Fage, 1925 (as Grubia hirsuta), Shoemaker, 1935 (as Grubia filosa).
Antenna 1 with 1- or 2-articulate accessory flagellum; mandible with palp; gnathopods large, subchelate, gnathopod 2 equal to or larger than 1; article 6 of pereopods 3-5 not apically widened; outer ramus of uropod 3 with two hooks. Species: ca. 10, generally tropical- amphiboreal, littoral.
Exampithoe K. H. Barnard Exampithoe K. H. Barnard, 1925.
Type-species: HH. natalensis K. H. Barnard, 1925 (monotypy).
Antenna 1 lacking accessory flagellum; mandible with palp, molar reduced; gnathopods large, subchelate, gnathopod 1 stouter than 2; article 6 of pereopods 3-5 apically widened; outer ramus of uropod 3 with two hooks. Species: 1, 8. Africa, littoral.
Macropisthopus K. H. Barnard Macropisthopus K. H. Barnard, 1916.
Type-species: MM. stebbingi K. H. Barnard, 1916 (monotypy).
Antenna 1 lacking accessory flagellum; mandible with palp; gnatho- pods feeble, chelate; article 6 of pereopods 3-5 scarcely widened distally; outer ramus of uropod 3 with two hooks. Species: 1, S. Africa, littoral.
Paragrubia Chevreux Paragrubia Chevreux, 1901a.
Type-species: P. vorax Chevreux, 1901a (monotypy).
Antenna 1 with accessory flagellum; mandible with palp; gnathopods large, subchelate, gnathopod 1 larger than 2; article 6 of pereopods 3-5 not apically widened; outer ramus of uropod 3 with two hooks. Species: 1, tropical Pacific, littoral.
MARINE GAMMARIDEAN AMPHIPODA 145
Pleonexes Bate
Anisopus Templeton, 1836 (homonym, Diptera). Pleonexes Bate, 1857a.
Type-species: P. gammaroides Bate, 1857a (monotypy). See Sars, 1895.
Antenna 1 lacking accessory flagellum; mandible with palp; enathopods large, subchelate, gnathoped 2 equal to or larger than 1; article 6 of pereopods 3-5 apically widened and subprehensile (Ampithoe) ; outer ramus of uropod 3 with two hooks; telson with apical pair of reverted, elongate, cornified . processes (Ampithoe). Species: 3, N.E. Atlantic, New Zealand, littoral.
Sunamphitoe Bate Sunamphitoe Bate, 1857a.
Type-species: Amphithoe pelagica Milne Edwards, 1830 (desig- nated by Chevreux and Fage, 1925). See Sars, 1895 (as S. conformata).
Antenna 1 lacking accessory flagellum; mandible lacking palp; enathopods large, subchelate, gnathopod 2 equal to or larger than 1; article 6 of pereopods 3-5 not apically widened; outer ramus of uropod 3 with two hooks. Species: 2, N.K. Atlantic, epipelagic.
Anamixidae
FIGURE 62
Diaenosis.—Mandible, lower lip, and maxillae absent; buccal regions with ventrally projecting keel; outer plates of maxillipeds absent, inner plates small, fused together; coxa 1, when present, hidden by the shield-like coxae 2—4; telson entire. See Leucothoidae, Amphilochidae, Cressidae, Sebidae, Thaumatelsonidae.
Dzscrietion.—Body smooth, lacking dorsal processes; mouth- parts aberrant as in diagnosis; accessory flagellum absent; gnathopod 1, when present, small and complexly chelate, having a chelate process on article 5, articles 6 and 7 (when present) together forming dactyl; gnathopod 2 immense, article 5 forming a false chela but articles 6 and 7 large, and article 7 alone forming the true dactyl; uropod 3 with styliform rami as long as the elongate peduncle; telson entire.
RELATIONSHIP.—The Leucothoidae bear mandibles and maxillae.
Key to the Genera of Anamixidae
Pee EGmathopodlepresemti cies 8 fr eels ved ok eA Ce a ese inca a ein hae Anamixis Cnathopodwltalosemt yeu nee ila mien eo ee ae aia ele iy aun Paranamixis
146 U.S. NATIONAL MUSEUM BULLETIN 271 Genera of Anamixidae
Anamixis Stebbing Anamixis Stebbing, 1897; 1906.
Type-species: A. hanseni Stebbing, 1897 (monotypy). Species: 4, Indo-Pacific tropics and subtropics, littoral (sponges).
Paranamivis Schellenberg
Paranamixis Schellenberg, 1938.
Type-species: P. bocki Schellenberg, 1938 (monotypy). Species: 1, Gilbert Islands, littoral.
Figure 62.—Anamixidae: a, Anamixis hanseni Stebbing (1897). Anamixis linsleyi J. L. Barnard (1955b): 6, oblique ventral view of head showing keel presumably formed of mandibles and maxillae: c, maxilliped; d, e, gnathopods 1, 2; f, uropod 3; g, telson.
MARINE GAMMARIDEAN AMPHIPODA 147
Aoridae
FIGURES 638, 64
Draenosis.—Accessory flagellum variable, multiarticulate, or ab- sent; telson entire, short, fleshy; coxa 4 not excavate posteriorly; pereopods glandular; wropod 3 rarely projecting beyond uropods 1 and 2, at least one of rami as long as or longer than peduncle; gnathopod 1 larger than gnathopod 2, in rare cases only as large as gnathopod 2 but with male secondary sexual modifications. See Isaeidae, Coro- phiidae, Ischyroceridae.
Description.—Accessory flagellum varying from absent to long and multiarticulate; body smooth; rostrum vestigial; coxae rounded or quadrate below, varying from long to short, fourth not excavate posteriorly; mouthparts basic; gnathopods powerful, subchelate or complexly subchelate, occasionally nearly simple, but then often extremely setose, first always larger or more complex than second in both sexes and bearing most of male secondary sexual modifications; pereopod 5 usually conspicuously elongate; uropod 3 short, rami usually as long as or longer than peduncle, occasionally inner ramus reduced in size or both rami reduced; telson short, nearly circular or square, entire, occasionally falsely (?secondarily) cleft.
RELATIONSHIP.—Like the Isaeidae but having an enlarged first gnathopod; otherwise Aoridae are related to other families in the same way as are the Isaeidae. Keys of both families and the Coro- phiidae should be checked when examining any genera with abnormally enlarged or complexly subchelate gnathopods. One may understand the difficulties of segregating Aoridae and Corophiidae when examining the small differences between Grandidierella (Corophiidae) and Neomicrodeutopus (Aoridae).
Nomenclatural Changes in Aoridae
The female of Dryopoides was first assigned to the Aoridae by Stebbing (1888). Its first gnathopod is larger than the second. In 1910, Stebbing discovered the male, with gnathopod 2 larger than 1, but the species was transferred to the Corophiidae. Because the genus may yet prove to be an aorid, the female is retained in the key to Aoridae as if it were a male.
Paradryope and Dryopoides might be placed in the family Ischyro- ceridae, especially Paradryope because of the short rami of its uropod 3 and the slightly elongate peduncle. The uropodal peduncle of Dryopoides is not greatly elongate. Ischyroceridae would thus receive genera with gnathopod 1 larger than 2, of which Bonnierella is an
U.S. NATIONAL MUSEUM BULLETIN 271
148
AQ S NS
S35 S S535 SScasen
See. oF a > => = a
J A “le {/' H
Aora:
Figure 63.—Aoridae: a, Aora typica Krgyer (Sars, 1895, pl. 193, as 4. gracilis). b, upper lip; c, mandible; d, lower lip; e, f, maxillae 1-2; g, maxilliped. Gnathopod 1:
h, female Aora; 1, male Aora; 7, Xenochetra fasciata Haswell (Pirlot, 1938); k, Lembos
websteri Bate (Sars, 1895, pl. 194); 2, Rudilemboides stenopropodus J. L. Barnard (1959a);
m, Lembopsis spinicarpus Pearse (1912). Uropod 3: n, Aora; 0, Dryopoides westwood
Stebbing (1888); p, Paraoroides unistilus Stebbing (1910); 9, Acuminodeutopus heteruropus Telson: r, Aora.
J. L. Barnard (1959a).
MARINE GAMMARIDEAN AMPHIPODA 149
intermediate genus. This procedure would also justify combining the remaining Aoridae with the Isaeidae (Photidae) and might eventually result in the Ischyroceridae also being joined to the Isaeidae, perhaps as a subfamily.
10.
Key to Genera of Aoridae
[MALES AND MALE-LIKE FEMALES]
Article 4 of gnathopod 1 produced into a long tooth (merochelate) (fig. 632)! 2 Article 4 of gnathopod 1 not produced into a one tooth Oe onay with
ashort tooth). ... : Die CeO Accessory flagellum long, compeced of dine 6 or more amides Gee AOra Accessory flagellum absent . . . j . .. . . Aoroides Article 5 of gnathopod 1 with one or more icone distal or subposterior teeth
(carpochelate) (fig. 64j) .... Niece saga Article 5 of gnathopod 1 lacking a enone dicen Gemn (Ge. 631) Geen tata eel lg Uropod 3 uniramous (fig. 63p). .... =. . =... . Neomicrodeutopus Uropodrs biramousy. (figs; GSq)) se Ne i ya eg BO) Gnathopod I of both sexes alike. ............. . Hansenella Gnathopod 1 differing in each sex . . . » 6
Gnathopod 2 heavily setose on full anterior adee of percle 5 Ge 649) . 7 Gnathopod 2 sparsely setose on anterior edge of article 5 or setae compacted into a distal bundle only . . a eas 10 Article 6 of gnathopod 1 as long ond: as Tprondl ¢ as pide 5 Ge. 63m). Lembopsis Article 6 of gnathopod 1 shorter and/or narrower than article 5 (fig. 631) . 8 Article 5 of gnathopod 2 longest (fig. 64g), female gnathopod 1 simple (like fi G4f) os oo: ees Neue peules Article 6 of paathopod 2 lowes female eaathopod 1 subchelate (fig. 63h) . Head elongate, antenna 2 attached halfway back on ventral cephalic margin, gnathopod 2 of male as stout as 1, of female weakly setose anteriorly. Amphideutopus Head not elongate, antenna 2 attached to anteroventral corner of head in front of eye, gnathopod 2 of male much more slender than 1, of female
densely setose anteriorly . .. . Be ROR Goremanas Inner ramus of uropod 3 less than half as iene as OHS: ramus (fig. 63q). Acuminodeutopus
Rami of uropod 3 subequal ne 63n)....... . .. . Microdeutopus Uropod 3 lacking rami .. . Cea On mince en ee ne ee OM CInOtALSUS Uropod 3 with at least oneramus .. . BIER etary NEGA LY Rami of uropod 3 minute, less than half as fone as erredinclc (ae 630) ..13 2 Rami of uropod 3 not minute, as long as or longer than peduncle .. 14 Pleonite 6 evanescent dorsally, artiels 3 of antenna 1 shorter than article 1, accessory flagellum vestigial . . . ... . . .. .. . Dryopoides, female
1 Note that juvenile males of Aora and Aoroides have a very short tooth and thus may key to Lembos but gnathopod 1 of Lembos is always strongly subchelate, whereas gnathopod 1 of Aora and Aoroides is essentially simple.
2 These genera may belong with the Ischyroceridae.
150 U.S. NATIONAL MUSEUM BULLETIN 271
Pleonite 6 visible dorsally, article 3 of antenna 1 ened than article 1, acces-
sory flagellum well developed. . . Berea Eavadtvone
14. Gnathopod 1 with article 6 equal to or 2 EEE 6 in Jength and breadth than article SiGich O37) eae eee ba tasers ALG Gnathopod 1 with article 6 shorten amcor narrower ‘hem price 5 (fig. GSE) yk a Sone PDs seni mame etn ites OS ILS)
15. Uropod 3 uniramous, _ aoeeREny feeeliacn albsents .. . . . . Paraoroides Uropod 3 biramous, accessory flagellum present. . . . . Wheto eer)
16. Article 3 of antenna | subequal to article 1, gnathopods 1-2 eouel: in size and morphology, subchelate ... . ... . . Aorcho
Article 3 of antenna 1 shorter than ariel i aamoped 1 larger than gnatho-
pod 2 or gnathopod 2 extremely setose, occasionally gnathopod 2 simple. 17
iver Gnathepod 2:subchelate swan Aeon ay eee | ieee ee eNOS
Gnathopod 2 simple (fig. 1061) ..... . been tees) Itc)
18. Coxae 1 and 2 short, wider than long, pabequall in size, _gulsomadbnite coxa 2 not concealing coxa 1, article 5 of gnathopod 2 very bulbous (fig. 64d).
: Xenocheira
Coxa | smaller than and partially to fully hidden by coxa 2, latter longer than wide, article 5 of gnathopod 2 not bulbous. ... . . . Leptocheirus
19. Gnathopods fully subchelate . .. . ... .. . . Lemboides Gnathopods scarcely subchelate (figs. 631, 64/) . . . . . Rudilemboides
Genera of Aoridae
Acuminodeutopus J. L. Barnard Acuminodeutopus J. L. Barnard, 1959a.
Type-species: A. heteruropus J. L. Barnard, 1959a (original designation).
Article 3 of antenna 1 shorter than article 1, accessory flagellum present; male gnathopod 1 with large distal tooth on article 5, article 6 shorter and narrower than article 5, simple, female gnathopod 1 sub- chelate; male gnathopod 2 nearly simple, poorly setose, article 5 longest; outer ramus of uropod 3 longer than peduncle, inner ramus less than half as long as outer. Species: 1, California, littoral.
Amphideutopus J. L. Barnard Amphideutopus J. L. Barnard, 1959a.
Type-species: A. oculatus J. L. Barnard, 1959a (original designation).
[Antenna 1 unknown]; male gnathopod 1 with distal tooth on article 5, article 6 shorter and narrower than 5, simple, female gnathopod 1 subchelate; male gnathopod 2 with moderately setose anterior edge on article 5, article 6 slightly the longest, subchelate; male gnathopod 2 as stout as gnathopod 1; head elongate, antenna 2 attached halfway back on ventral cephalic margin; female gnathopod 2 weakly setose anteriorly (Coremapus) ; rami of uropod 3 equal, longer than peduncle. Species: 1, California, littoral.
MARINE GAMMARIDEAN AMPHIPODA IVS
Ficure 64.—Aoridae: a, Microdeutopus gryllotalpa Costa (Sars, 1895, pl. 192); b, Aorcho delgadus J. L. Barnard (1961). Gnathopod 2: c, Aora typica Krgyer female (Sars, 1895, pl. 193, as 4. gracilis); d, Xenocheira fasciata Haswell (Pirlot, 1938); e, dora male; f, Rudilemboides stenopropodus J. L. Barnard (1959a); g, Neomegamphopus roosevelt: Shoe- maker (1942); h, Lembopsis spinicarpus Pearse (1912); 1, Lembos websteri Bate (Sars, 1895, pl. 194). Gnathopod 1: 7, Microdeutopus.
152 U.S. NATIONAL MUSEUM BULLETIN 271
Aora Kr¢éyer
Aora Krgyer, 1845.—Stebbing, 1906. Lalaria Nicolet, 1849. Lonchomerus Bate, 1857a. Type-species: A. typica Krgyer, 1845 (monotypy). See Sars, 1895. Article 3 of antenna 1 shorter than article 1, accessory flagellum present; male gnathopod 1 complexly subchelate, with long tooth on article 4, articles 5-6 lacking teeth, article 6 slightly shorter and some- times narrower than article 5, simple, female gnathopod 1 scarely subchelate; male gnathopod 2 subchelate, not heavily setose on anterior edge of article 5, articles 5 and 6 subequal; rami of uropod 3 subequal, longer than peduncle. Species: 2, Atlantic, amphiboreal, littoral, bathyal.
Aorcho J. L. Barnard Aorcho J. L. Barnard, 1961.
Type-species: A. delgadus J. L. Barnard, 1961 (original designa- tion).
Article 3 of antenna 1 as long as article 1, accessory flagellum present; male gnathopod 1 lacking teeth, scarcely larger than and similar to gnathopod 2, [female unknown]; gnathopod 2 subchelate, poorly setose, articles 5-6 subequal; rami of uropod 3 subequal, longer than peduncle. Species: 1, Tasman Sea (610 m).
Aoroides Walker Aoroides Walker, 1898.—Stebbing, 1906.
Type-species: A. columbiae Walker, 1898 (monotypy). See J. L. Barnard, 1954.
Article 3 of antenna 1 shorter than article 1, accessory flagellum absent; male gnathopod 1 complexly subchelate, with long tooth on article 4, articles 5-6 lacking teeth, article 6 shorter and narrower than 5, simple, female gnathopod 1 scarcely subchelate; male gnatho- pod 2 not heavily setose, articles 5-6 subequal, subchelate; rami of uropod 3 subequal, as long as peduncle. Species: 2, Pacific boreal, littoral.
Coremapus Norman Coremapus Norman, 1905.
Type-species: Lembos versiculatus Bate, 1857a (original designa- tion). See Chevreux and Fage, 1925.
Article 3 of antenna 1 shorter than article 1, accessory flagellum present; male gnathopod 1 with distal tooth on article 5, article 6
MARINE GAMMARIDEAN AMPHIPODA 153
shorter and narrower than 5, simple, female gnathopod 1 subchelate; male and female gnathopod 2 with heavily setose anterior edge of article 5, article 6 longest, subchelate; male gnathopod 2 much more slender than gnathopod 1; head not elongate, antenna 2 attached at antero- ventral cephalic corner in front of eye (Amphideutopus); rami of uropod 3 equal, longer than peduncle. Species: 1, N. Atlantic boreal, littoral. Dryopoides Stebbing
Dryopoides Stebbing, 1888; 1906.
Type-species: D. westwoodi Stebbing, 1888 (monotypy). See Stebbing, 1910.
Article 3 of antenna 1 shorter than 1, accessory flagellum minute; male gnathopod 1 subchelate, articles 4-6 lacking teeth; female enathopod 1 subchelate; male gnathopod 2 subchelate, not heavily setose, article 6 longest; rami of uropod 3 minute, equal, much shorter than peduncle; pleonal segment 6 evanescent dorsally (Paradryope). Species: 1, Australia, littoral. Also assigned to Corophiidae.
Hansenella Chevreux Hansenella Chevreux, 1909.
Type-species: H. longicornis Chevreux, 1909 (original designation).
[Male unknown, but probably like female]; article 3 of antenna 1 shorter than article 1, accessory flagellum present; gnathopod 1 like that of male aorid, with distal tooth on article 5, article 6 shorter and narrower than 5, poorly subchelate; gnathopod 2 subchelate, article 6 longest, article 5 poorly setose anteriorly; rami of uropod 3 equal, slightly longer than peduncle. Possibly synonymous with Microdeutopus and representing aberrant female. Species: 1, N. Atlantic (1,360 m).
Lemboides Stebbing
Lemboides Stebbing, 1895; 1906.
Type-species: JL. afer Stebbing, 1895 (original designation).
Article 3 of antenna 1 shorter than article 1, accessory flagellum present; male gnathopod 1 subchelate, articles 4-5 lacking teeth, 6 occasionally with chela, article 5 longer and broader than 6, female gnathopod 1 subchelate, male gnathopod 2 subchelate, article 5 longest, poorly setose; rami of uropod 3 equal, longer than peduncle. Species: 4, antiboreal, littoral.
Lembopsis Pearse Lembopsis Pearse, 1912.
Type-species: L. spinicarpus Pearse, 1912 (original designation). Article 3 of antenna 1 shorter than article 1, accessory flagellum
154 U.S. NATIONAL MUSEUM BULLETIN 271
present; male gnathopod 1 with long posterior tooth on article 5, article 6 longer and as broad as 5, subchelate; male gnathopod 2 heavily setose on anterior edge of article 5, articles 5-6 subequal; rami of uropod 3 equal, longer than peduncle. Species: 1, Gulf of Mexico, littoral.
Lembos Bate
Lembos Bate, 1857a.—Stebbing, 1906. Autonoe Bruzelius, 1859. Bemlos Shoemaker, 1925.
Type-species: JL. webstert Bate, 1857a (designated by Chevreux and Fage, 1925). See Sars, 1895.
Article 3 of antenna 1 shorter than article 1, accessory flagellum present; male gnathopod 1 subchelate, with articles 4—5 lacking teeth, article 6 not narrower but usually longer than 5, female gnathopod 1 subchelate; male gnathopod 2 subchelate, with article 5 longest, poorly setose on anterior edge; rami of uropod 3 equal, longer than peduncle. Species: 31, cosmopolitan, primarily tropical, littoral (2 bathyal, 1 abyssal).
Leptocheirus Zaddach
Leptocheirus Zaddach, 1844.—Stebbing, 1906. Ptilocheirus Stimpson, 1853. Boeckia Malm, 1871.
Type-species: JL. pilosus Zaddach, 1844 (original designation). See Sexton, 1911; Chevreux and Fage, 1925.
Article 3 of antenna 1 shorter than article 1, accessory flagellum 3+-articulate; male gnathopod 1 subchelate, articles 4-5 lacking teeth; female gnathopod 1 subchelate and extremely setose; gnathopod 2 simple in both sexes and extremely setose on articles 2 and 5 but article 5 not especially bulbous; rami of uropod 3 equal or unequal, one or both rami longer than peduncle; coza 1 often very small and occasionally large but always slightly smaller than coxa 2 and often tri- angular, hidden fully or partially by enlarged coxa 2, latter longer than wide (Xenocheira). Species: 10, boreal N. Atlantic, littoral.
Microdeutopus Costa
Microdeutopus Costa, 1853c.—Stebbing, 1906. Stimpsonia Bate and Westwood, 1863 (homonym, Nemertea). Stimpsonella Della Valle, 1893.
Type-species: MM. gryllotalpa Costa, 1853c (monotypy). See Sars, 1895.
Article 3 of antenna 1 shorter than article 1, accessory flagellum present; male gnathopod 1 with long distal tooth on article 5, article
MARINE GAMMARIDEAN AMPHIPODA 155
6 shorter and narrower than article 5, simple, female gnathopod 1 subchelate; male gnathopod 2 subchelate, articles 5-6 variable in length, article 5 not heavily setose anteriorly; rami of uropod 3 sub- equal, as long as or longer than peduncle. Species: 10, cosmopolitan, littoral.
Neomegamphopus Shoemaker Neomegamphopus Shoemaker, 1942.
Type-species: WN. rooseveltt Shoemaker, 1942 (original designation).
Article 3 of antenna 1 as long as article 1, accessory flagellum small; male gnathopod 1 with long distal tooth on article 5, article 6 simple, as long but narrower than article 5, female gnathopod 1 simple; male enathopod 2 with article 5 longer than 6 and heavily setose on anterior edge; rami of uropod 3 subequal, slightly longer than peduncle. Species: 1, E. Pacific tropical, littoral.
Neomicrodeutopus Schellenberg Neomicrodeutopus Schellenberg, 1925b.
Type-species: N. cabindae Schellenberg, 1925b (monotypy).
Article 3 of antenna 1 shorter than article 1, accessory flagellum present; male gnathopod 1 with long distal tooth on article 5, article 6 shorter and narrower than article 5, simple, [female unknown]; male enathopod 2 subchelate, stout, article 5 longer than 6, article 5 poorly setose anteriorly; uropod 3 with a single ramus twice as long as pe- duncle, latter with a broad distomedial lobe. Species: 2, W. Africa, littoral. Differs from Grandidierella in Corophiidae by normally short article 1 of mandibular palp.
Paradryope Stebbing Paradryope Stebbing, 1888; 1906.
Type-species: P. orguion Stebbing, 1888 (monotypy).
Article 3 of antenna 1 longer than article 1, accessory flagellum present; male gnathopod 1 subchelate, articles 4—5 lacking teeth, article 6 longer and broader than 5; male gnathopod 2 subchelate, not heavily setose, articles 5-6 equal; rami of uropod 3 minute, equal, much shorter than peduncle. Species: 1, N. Pacific, abyssal (4,200 m).
Paraoroides Stebbing Paraoroides Stebbing, 1910.
Type-species: P. unastilus Stebbing, 1910 (monotypy). Article 3 of antenna 1 shorter than article 1, accessory flagellum
285-135 O-69-11
156 U.S. NATIONAL MUSEUM BULLETIN 271
minute, vestigial; male gnathopod 1 subchelate, articles 4—5 lacking teeth, articles 5-6 subequal, female gnathopod 1 scarcely subchelate; male gnathopod 2 subchelate, poorly setose, article 5 slightly longer than 6; uropod 3 uniramous, ramus equal to peduncle. Species: 1, S.E. Australia, littoral.
Rudilemboides J. L. Barnard Rudilemboides J. L. Barnard, 1959a.
Type-species: . stenopropodus J. L. Barnard, 1959a (original designation).
Article 3 of antenna 1 shorter than article 1, accessory flagellum present; male gnathopod 1 simple, lacking teeth, article 5 much broader and longer than article 6, female gnathopod 1 subchelate; male gnathopod 2 nearly simple, poorly setose, article 5 longest; rami of uropod 3 equal, longer than peduncle. Species: 1, California, littoral.
Uncinotarsus L’Hardy and Truchot Uncinotarsus L’ Hardy and Truchot, 1964.
Type-species: U. pellucidus L’ Hardy and Truchot, 1964 (original designation).
Article 3 of antenna 1 shorter than article 1, accessory flagellum 2-articulate; male gnathopod 1 subchelate, articles 4—5 lacking teeth, article 6 broader and longer than article 5, female gnathopod 1 sub- chelate; male gnathopod 2 subchelate, not heavily setose, article 6 longer than 5; uropod 2 with one ramus; rami of uropod 3 absent. Species: 1, Atlantic France, littoral. Differs from Concholestes in Corophiidae by slender antenna 2 being shorter than antenna 1, by enathopod 1 being larger than 2, by the short article 3 of antenna 1, acuminate coxae 5-7, slender pereopods 1—2, and elongate pereopods o-9.
Xenocheira Haswell Xenocheira Haswell, 1880b.—Stebbing, 1906.
Type-species: X. fasciata Haswell, 1880b (monotypy). See Pirlot, 1938.
Article 3 of antenna 1 shorter than article 1, accessory flagellum present; male gnathopod 1 subchelate, articles 4—5 lacking teeth; female gnathopod 1 subchelate; male gnathopod 2 simple, female minutely subchelate, article 6 longest, article 5 strongly setose anteriorly and very bulbous; gnathopod 2 extremely setose; rami of uropod 3 equal, longer than peduncle; corae 1-2 subequal i size, short, wider than long, coxa 1 not concealed by coxa 2 (Lep tocheirus). Species: 2, tropical Pacific, littoral.
MARINE GAMMARIDEAN AMPHIPODA 157 Argissidae FIGURE 65
Draenosis.—Accessory flagellum 2-articulate; coxae 1 and 4 long, coxae 1-3 successwely smaller, coxa 4 larger than coxa 1; gnathopods feeble, simple or poorly subchelate. Monogeneric. See Gammaridae, Haustoriidae, Synopiidae, Ampeliscidae, Vitjazianidae.
Description.—Accessory flagellum 2-articulate; when present, primary flagellum basally conjoint in male; dorsal teeth only on urosome, rostrum obsolescent; coxa 1 large, coxae 2 and 3 successively smaller, coxa 4 larger than 1; mouthparts basic, upper lip slightly incised; mandibular palp with article 3 longer than article 2; gnatho- pods feeble, simple, except one species having gnathopod 1 poorly subchelate; uropod 3 biramous; telson of medium length, cleft.
RELATIONSHIP.—Argissids are distinguished by the consecutive reduction in size of coxae 1-3. They closely resemble Gammaridae, especially the genus Megaluropus in which coxae 1-4 also are irregular but in which coxa 2 is larger than 1 and coxa 3 smaller than 2.
Despite their definitive similarity to Gammaridae, argissids bear external resemblance to Haustoriidae and were at one time included in that family. They call attention to the close relationship of haus- toriids to gammarids. Again the unusual coxae distinguish them from haustoriids. The peculiar eyes of Argissa hamatipes, four bigeminous lenticular bodies imbedded at the periphery of a common pigmentary mass, seem unique to the family but the other species of Argissa, A. stebbingi, lacks eyes, so that they are not diagnostic.
Sars (1895) has noted the similarity of the genus Argissa to the Ampeliscidae (at that time Argissa was placed in the Haustoriidae). Resemblance occurs in the antennae, pereopods 1-2 and 5, the uropods, and the telson. Argissa nevertheless has some primitive characters which place it between the Ampeliscidae and the Gammaridae: the accessory flagellum, the intermediately modified pereopods 3 and 4, and a weakly 2-articulate outer ramus of uropod 3. The eyes of Argissa may represent a stage in the development of the ampeliscid corneal lens. Of course, Argissa cannot stand on the direct line between Ampeliscidae and Gammaridae because of its specializations in coxae but it may well be a close relative of the organisms that occurred directly on the evolutionary pathway. The most important specializa- tions of the Ampeliscidae have been the development of tube-spinning glands in pereopods 1-2, a function apparently not developed in Argissa, and the amalgamation of the last two pleonal segments.
Article 2 of pereopods 3-5 in Argissa does have conspicuous glands.
Vitjazianids have many characters in common with argissids but Argissa may be recognized by the special configuration of coxae.
158 U.S. NATIONAL MUSEUM BULLETIN 271
Nomenclatural Changes in the Argissidae
Parargissa Chevreux (1908c; Pirlot, 1934) is removed to the Hyperiopsidae.
Phylluropus K. H. Barnard (1932) is a synonym of Megaluropus (Gammaridae).
Phylluropus capensis K. H. Barnard (1932) is a synonym of Megalu- ropus agilis (fide K. H. Barnard, 1940).
\
| Si HN
\\
/ AK LN
N
Ficure 65.—Argissidae: Argissa hamatipes (Norman) (Sars, 1895, pl. 48, as 4. typica): a, upper lip; b, lateral; c, mandible; ¢, lower lip; f,g, maxillae 1, 2; h, maxilliped; 7, uropod 3; 7,k, gnathopods 1, 2; /, uropod 3. Argissa stebbingi Bonnier (1896): d, mandible; m, gnathopod 1.
MARINE GAMMARIDEAN AMPHIPODA 159 Genera of Argissidae
Argissa Boeck
Argissa Boeck, 1871.—Stebbing, 1906. Chimaeropsis Meinert, 1890 (homonym, Pisces).
Type-species: Syrrhoe hamatipes Norman, 1869a (monotypy and subsequent synonymy). See Sars, 1895.
Species: 2 (possibly synonymous), N. Hemisphere and possibly cosmopolitan, eurybathic (30-1919 m).
Astyridae
FIGURE 66
Diaenosis.—Accessory flagellum 1- or 2-articulate, slender (or absent in dubious genera); mandibular molar nontriturative; lower lip with outer lobes very widely separated; gnathopods feeble. Mono- generic. See Eusiridae, Calliopiidae, Pleustidae, Liljeborgiidae, Haustoriidae, Synopiidae, Vitjazianidae, Hyperiopsidae, Paramphi- thoidae, Stilipedidae.
Description.-—Accessory flagellum a single, medium-sized article or 2-articulate; body smooth; rostrum inconspicuous, head usually normal in appearance but occasionally massive and with “‘shark nose” appearance (as in some Synopiidae [=Tironidae] and some Par- daliscidae) ; coxae of medium size or long, rounded or quadrate below but 2 and 4 occasionally subacuminate; mandibular molar a conical, setose, nontriturative lamina; outer lobes of lower lip very widely separated, with or without small erect inner lobes; remainder of mouthparts basic, occasionally maxillae subfoliaceous; gnathopods feeble, barely subchelate; rami of uropod 3 very elongate; telson short, entire or cleft.
ReEwatTionsuHip.— Usually, the Synopiidae strongly differ from the Astyridae only by their multiarticulate accessory flagella (except the genus Jeddo). The combination of very broadly separated lobes of the lower lip, the fact that coxa 4 is usually larger than coxa 3 and the peculiar, sharp, conical, nontriturative molar serve to distinguish Astyridae.
The short coxae and the absence of a mandibular molar distinguish the Pardaliscidae.
Oedicerotidae have strongly setose pereopods, lack an accessory flagellum, have relatively short rami and therefore a relatively long peduncle of uropod 3 and a non-astyrid lower lip.
Most Eusiridae have either a ridged mandibular molar, do not bear
160 U.S. NATIONAL MUSEUM BULLETIN 271
the astyrid lower lip or have an elongate telson. Most Calliopiidae have triturative molars and the nontriturative molars of Pleustidae are bulbous, not conical.
In terms of accessory flagellum, mandibular molar, maxillae, maxillipeds, gnathopods, uropods, and telson the Astyridae resemble Paramphithoidae closely, especially to the genera Helysis and Epi- meriella, which have the same mandibular molar and gnathopods, maxillae and maxillipeds. Astyridae differ from the Paramphithoidae
spp howe x ON
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Ficure 66.—Astyridae: a, Astyra abyssi Boeck (Sars, 1895, pl. 73); 6, upper lip; c, mandible; e, lower lip; g,h, maxillae 1, 2; 7, maxilliped; j,k, gnathopods 1, 2; m, telson; 0, uropod 3; d, head of Astyra gardinert (Walker) (Pirlot, 1934, as Parastyra longidactyla); f, lower lip of A. bogorovi Birstein and Vinogradov (1955); /, telson of A. zenkevitchi Birstein and Vinogradov (1955); n, uropod 3 of A. gardinert.
MARINE GAMMARIDEAN AMPHIPODA 161
only by the peculiar lower lip. That this character is sufficient to remove Astyridae from Paramphithoidae is doubtful, especially because of the subacuminate condition of coxae 2 and 4 in astyrids and the distally expanded and rounded-quadrate coxa 1 of Eelysis.
The Laphystiopsidae lack an accessory flagellum, have normal lower lips, and entire telsons.
The Haustoriidae do not have the characteristic astyrid lower lip and have fossorial (spiny and setose) pereopods 3-5.
Astyroides Birstein and Vinogradova is synonymized with Alez- andrella in the Stilipedidae. Stilipedids differ from Astyra in the absence of a mandibular molar, the broadening of the mandibular body, the greater foliaceousness of the maxillae, but the two families are strongly related by the condition of coxae 1-2, of which coxa 1 is broader than 2 and 2 tends to taper distally. Stedipes stands between Astyra and Alexandrella in the condition of its upper lip.
Key to the Genera of Astyridae
(PE CoxalesubaGUMINALG inh coisas uel elena ase) SOR SESS [Epimeriella] ! Coxaylsrounded-subquadrate distally 2) ie 2 ann ee eee Ag 2, Pee Ne cessorysiagellumepnresemt ..9.. 25) 3 Ge eae ee ies cl pel Astyra NCcCeSsonyehacellummalbsenmtim.) coin nie pies cea ore iyi Me les [Eclysis] !
1See these genera in the Paramphithoidae (pp. 394, 395).
Genera of Astyridae
Astyra Boeck
Astyra Boeck, 1871.—Stebbing, 1906. Chagosia Walker, 1909. Parastyra Pirlot, 1934. Type-species: A. abyss1 Boeck, 1871 (monotypy). See Sars, 1895. Species: 5, cosmopolitan, bathypelagic (100-2000 m).
Atylidae
FIGURES 67, 68
Driacnosis.—Accessory flagellum vestigial or absent; urosomites 2-3 coalesced. Monogeneric. See Ampeliscidae, Dexaminidae, Pro- phliantidae, Lepechinellidae.
Description.—Accessory flagellum vestigial, 1-articulate or absent; body with dorsal carinae or teeth; rostrum prominent; urosomites
162 U.S. NATIONAL MUSEUM BULLETIN 271
2 and 3 coalesced; mouthparts basic, lower lip lacking distinct inner lobes; gnathopods small, subchelate; uropod 3 biramous; telson of medium length, cleft.
RELATIONSHIP.—The Ampeliscidae have cuticular visual lenses and pereopod 5 is shorter and of different structure than pereopod 4. Other nonisaeid families with coalesced urosomites 2-3 are the Dexa-
Ficure 67.—Atylidae: a, Atylus falcatus Metzger (Sars, 1895, pl. 164); b, Atylus carinatus, (Fabricius) (Sars, 1895, pl. 166).
MARINE GAMMARIDEAN AMPHIPODA 163
minidae, Lepechinellidae, and members of the Eophliantidae, Pro- phliantidae, and Kuridae.
The Dexaminidae lack a mandibular palp.
The Lepechinellidae have acuminate or very short coxae and a spine-like accessory flagellum.
The Eophliantidae, Prophliantidae, and Kuriidae all lack mandib- ular palps and have other mouthparts departing from the basic configuration.
Atylus aberrantis J. L. Barnard (1962d) is removed to
Lepechinella. Genera of Atylidae
Atylus Leach
Atylus Leach, 1815.—Stebbing, 1906.—J. L. Barnard, 1956. Nototropis Costa, 1853c, 1857.
Epidesura Boeck, 1861.
Paratylus Sars, 1895.
Type-species: Gammarus carinatus J. C. Fabricius, 1793 (mono- typy). See Sars, 1895. Species: 22, cosmopolitan, littoral (2 boreal bathyal).
Ficure 68.—Atylidae: Atylus carinatus (Fabricius) (Sars, 1895, pl. 166); a, upper lip; d lower lip; c,d, maxillae 1, 2; e, gnathopod 1; f, maxilliped; g, mandible; h, telson; 7, uropod 3; 7, gnathopod 2. Atylus swammerdami (Milne Edwards) (Sars, 1895, pl. 163): k, mandible.
164 U.S. NATIONAL MUSEUM BULLETIN 271
Bateidae
FIGURE 69
DraGnosis.—Coxa 1 vestigial or absent, remainder of gnathopod 1 degraded to a single article; accessory flagellum absent. See Gam- maridae, Kusiridae.
Description.—Accessory flagellum absent; rostrum conspicuous; back often with carinae; mouthparts basic, lower lip with or without inner lobes; gnathopod 1 and its coxa degraded; gnathopod 2 feeble, subchelate; others of first 3 coxae rounded ventrally; uropod 3 with elongate, lanceolate rami; telson of medium length, longer than peduncle of uropod 3, cleft.
RELATIONSHIP.—This is the only family with a degraded first gnathopod except the genus Paranamizis in the Anamixidae but that family is far removed from the Bateidae. Except for gnathopod 1, bateids resemble the Eusiridae.
Bateidae have a grade of structure similar to the Amphilochidae and the two families casually resemble one another in overall appearance.
Key to the Genera of Bateidae
1. Back with carinae, palp article 1 of maxilla 1 elongate . . . Carinobatea Back smooth, palp article 1 of maxilla 1 of normal dimensions. . . Batea
Genera of Bateidae
Batea Muller Batea Miller, 1865.—Stebbing, 1906.
Type-species: B. catharinensis Miiller, 1865 (monotypy). See Shoemaker, 1926. Species: 4, Caribbean, tropical eastern Pacific, littoral.
Carinobatea Shoemaker Carinobatea Shoemaker, 1926.
Type-species: C. cuspidata Shoemaker, 1926 (original designation). Species: 2, Caribbean, littoral.
MARINE GAMMARIDEAN AMPHIPODA 165
Ficure 69.—Bateidae (all figures after Shoemaker, 1926): a, Carinobatea cuspidata Shoe- maker. Head: b, Batea catharinensis Muller. Mandible: c, Batea catharinensis. Lower lip: d, Carinobatea; e, Batea. Mouthparts, Batea: f, maxilla 1; g, maxilla 2; f, maxilliped. Gnathopod 1: 1, Batea. Telson: 7, Batea transversa; k, Batea rectangulata; 1, Batea catharinensis. Uropod 3: m, Batea catharinensis. Gnathopod 2: n, Batea catharinensis.
166 U.S. NATIONAL MUSEUM BULLETIN 271
Beaudettiidae
Figure 70
Draenosis.—Accessory flagellum 2-articulate; mandible lacking palp; inner ramus of uropod 3 reduced; coxa 4 not excavate posteri- orly; telson very short, entire (but emarginate). Monogeneric. See Gammaridae.
Description.—Body like that of Gammaridae, dorsally unorna- mented; rostrum obsolete; antennal peduncles elongate, accessory flagellum 2-articulate; mandible with molar but lacking palp; mouth- parts otherwise basic; gnathopod 2 larger than 1; coxa 4 unexcavate posteriorly; uropod 3 short, inner ramus half as long and one third as broad as outer ramus; telson very short, entire (but emarginate).
RELATIONSHIP.—This monotypic family apparently is derived from the Gammaridae. It is like the genus Elasmopus but has lost the man- dibular palp, has a reduced inner ramus of uropod 3, and a short, emarginate telson.
Ficure 70.—Beaudettiidae: Beaudettia palmeri J. L. Barnard (1965): a, lateral aspect; b, mandible; c, lower lip; d,e, maxillae 1, 2; f, maxilliped; g, telson; A, upper lip; 7, uropod 3.
MARINE GAMMARIDEAN AMPHIPODA 167 Genera of Beaudettiidae
Beaudettia J. L. Barnard Beaudettia J. L. Barnard, 1965.
Type-species: B. palmera J. L. Barnard, 1965 (original designation). Species: 1, Micronesia, littoral.
Calliopiidae Figures 71-73, 146 7-k
Diacnosis.—Accessory flagellum absent or composed of a tiny article or scale; telson entire, apically emarginate or apically notched, never deeply split. See Eusiridae, Pleustidae, Isaeidae, Laphystiopsi- dae, Oedicerotidae, Amphilochidae, Ampithoidae, Paramphithoidae.
Description.—Accessory flagellum vestigial or absent; rostrum usually inconspicuous; upper lip usually poorly incised or not; mouthparts basic, but mandibular molar occasionally nontriturative; lower lip with or without inner lobes, never taking the form of Pleu- stidae (see p. 421); coxae of medium length or short; occasionally coxa 4 not posteriorly excavate; gnathopods powerful or feeble, usually subchelate, occasionally simple; uropod 3 with elongate lanceolate rami, outer occasionally shorter than inner; telson of medium length, entire, acuminate, emarginate, or notched.
RELATIONSHIP.—EKusiridae are close to this family but have a distinctly cleft telson; when only notched apically the telson is elongate in eusirids. Interfamilial gradation is seen by comparing telsons of Chosroes, Atylopsis, and Halirages in Calliopiidae and Pontogeneoides in EKusiridae.
That the Callopiidae are simply Eusiridae-Pontogenelidae with fused telsonic lobes is well demonstrated in the fact that some callio- pid genera are paired with some eusirid genera. Djerboa (Eusiridae) and Metaleptamphopus (Calliopiidae) share pectinate dactyls of pere- pods; Sancho (Calliopiidae) has eusirid gnathopods; Harpinioides (Calliopiidae) and Harpinioidella (Eusiridae) have similar gnathopods; Stenopleura (Calliopiidae) and Metewsiroides (Eusiridae) are very similar in all characters except telsons.
Calliopiids no longer differ from other families by their incon- spicuous rostra because several have been discovered lately that have a longer rostrum than do some eusirids and pleustids.
Pleustidae differ from Calliopiidae only by the special configuration of their lower lips. The genera Mesopleustes (Pleustidae) and Harpini- oides (Calliopiidae) have lower lips which are very difficult to classify
168
U.S. NATIONAL MUSEUM BULLETIN 271
SEQ 2 ee:
co Sas SS
iS ry H A ry i\ Kt a aN \\ A nt K Nt \\ \\ Ny i\ i \ i
Ficure 71.—Calliopiidae: a, Calliopius laeviusculus (Kréyer) (Sars, 1895, pl. 158); b, Cleippides quadricuspis Heller (Sars, 1885). Upper lip: ¢, Calliopius. Lower lip: d, Calliopius; e, Halirages fulvocinctus (M. Sars) (Sars, 1895, pl. 154). Mandible and palp: f, Calliopius; g, Leptamphopus sarsi Vanhoffen (Sars, 1895, pl. 162, as L. longimanus); h, Oradarea walkeri Shoemaker (Walker, 1903).
Maxilla 1: 7, Calliopius; j, Laothoes meinerti Boeck (Sars, 1895, pl. 160).
Maxilla 2: k, Calliopius.
MARINE GAMMARIDEAN AMPHIPODA 169
Ficure 72.—Calliopiidae: a, Apherusa tridentata (Bruzelius) (Sars, 1895, pl. 156). Pectinate dactyl of pereopod: b, Metaleptamphopus pectinatus Chevreux (1912b). Maxilliped: c, Calliopius laeviusculus (Krgyer) (Sars, 1895, pl. 158). Telson: d, Calliopius; e, Har- pinioides drepanocheir Stebbing (1888); f, Stenopleura atlantica Stebbing (1888); g, Atylopsis emarginata Stebbing (1888). Gnathopod 1: h, Leptamphopus sarsi VanhOffen (Sars, 1895, pl. 162, as L. longimanus); 1, Regalia fascicularis K. H. Barnard (1930); 7, Sancho platynotus Stebbing (1897). Gnathopod 2: k, Leptamphopus; l, Laothoes meinerti Boeck (Sars, 1895, pl. 160); m, Haliragoides inermis Sars (1895, pl. 153); n, Calliopius; 0, Harpinioides. Uropod 3: p, Calliopius.
170 U.S. NATIONAL MUSEUM BULLETIN 271
because they are almost perfect intergrades between the lower lips of pleustids and calliopiids.
Calliopiids (and pleustids) may be confused with isaeids, ischyroce- rids, and ampithoids unless one notes the absence of pereopodal glands in Calliopiidae. The isaeid-like families have a fleshy telson which is usually much shorter than broad, and generally have unexcavate posterior margins on the fourth coxae.
The Laphystiopsidae, very closely related to the Calliopiidae, differ primarily by the poorly developed mandibular molar. One laphystiopsid genus, Prolaphystius, however is included in the key to the Calliopiidae because of its uncertain position in the Laphystiop- sidae. It has a moderately well developed molar.
The Vitjazianidae are distinguished by their distinctly simple first gnathopods. Pleustidae have their characteristic lower lips.
The Paramphithoidae are characterized by cuspidate or pointed anterior coxae. Occasionally these acuminations are extremely weak and coxae 1-4 are scarcely excavate ventrally.
One calliopiid, Halirages regis (Stebbing, 1914) (=H. stebbingi Schellenberg, 1931; =H. huzleyanus Stebbing, not Bate, 1862) has subacuminate anterior coxae and thus keys to Paramphithoe, type- genus of Paramphithoidae. Halirages regis has strongly developed enathopods with ovate sixth articles and thus differs from most of its congeners and Paramphithoe. Superficially, this species forms the perfect intergrade between Calliopiidae and Paramphithoidae and distinctions between the two families are completely confounded; further study of the problem is warranted. Some species of Cleippides also have subacuminate coxae.
Some genera of the Gammaridae, such as Falklandella, might be placed in the Calliopiidae, except that Falklandella has short rami of uropod 3, not characteristic of Calliopiidae. Gammarellus and Weyprechtia with entire telson, and Parapherusa are not assignable to Calliopiidae because of their multiarticulate accessory flagella.
The Oedicerotidae have disproportionately long fifth pereopods and densely setose coxae and pereopods.
Most synopiids have a multiarticulate accessory flagellum but a few have an accessory flagellum similar to that of eusirids. The massive head of synopiids is characteristic, especially because of its strongly deflexed rostrum. Synopiid gnathopods are feeble, whereas those of eusirids are usually powerful.
Gnathopods within a genus are highly variable as to proportions of articles 5 and 6; departures from a ‘‘normal theme” are seen in Oradarea and Leptamphopus, which have exceptionally elongate sixth articles of gnathopod 2. Cleippides has a relatively elongate second gnathopod. Harpinioides has peculiar, bent sixth articles.
MARINE GAMMARIDEAN AMPHIPODA 171
Even the type-genus, Calliopius, has enlarged gnathopods, not typical of the remaining members of the family. Extremes of variation are shown in a group of gnathopodal figures presented herein.
Apherusa, Bouvierella, Callioprella, Halirages, and Haliragoides are often very difficult to separate. Their type-species are easily dis- tinguished by conditions of lower lip, anteroventral cephalic corners, coxa 1 and accessory flagella, but those characters are highly variable among other species of the five genera.
Gurjanova (1962) has pointed out the unusual character of Sancho and Chosroes. They have broadly vaulted, extremely depressed bodies and heads and splayed coxae like those of Phliantidae but they also have subfossorial pereopods and gnathopod 1 like that of fossorial families Phoxocephalidae, Haustoriidae, and Oedicerotidae.
a
Figure 73.—Calliopiidae: Gnathopod 2: a, Sancho platynotus Stebbing (1897); b, Clarencia chelata K. H. Barnard (1932); c, Stenopleura atlantica Stebbing (1888); d, Cleippides bicuspis Stephensen (Gurjanova, 1951); ¢, Atylopsis dentatus Stebbing (1888).
The mouthparts of both genera have a distinct look of the Oedi- cerotidae. Neither genus has the fully developed pereopod 5 nor elongate peduncle of uropod 3 found in the Oedicerotidae and Chosroes has an elongate telson. Gnathopod 2 of Sancho distantly resembles that of the phoxocephalid Joubinella (and various eusirids) ; and since Phoxocephalidae and Haustoriidae seem to have strong affinities one might consider Sancho to be an aberrant member of the Haustoriidae, were it not for the uncharacteristic short peduncle of antenna 1, the large male gnathopod 2, and the shortened outer rami of the uropods. The conditions of the telson and pereopod 5 exclude the genera from the Phoxocephalidae. But the short outer rami of the uropods of Sancho and Chosroes like those of the Calliopiidae and Eusiridae are not characteristic of the fossorial families. Probably a new family
285-135 O - 69 - 12
172 U.S. NATIONAL MUSEUM BULLETIN 271
should be created for these two genera which combine in a remarkable way the characteristics of phliantids, oedicerotids, and eusirids. This problem is also discussed under the Phliantidae (p. 405).
Remarks on the key to genera: Differences among genera are minor, for example: couplets 5, 6, 9, 16. Stebbing’s (1906) key to the genera utilized such quantitative features as vaulting of the dorsum of the body, the dentition of the segments, and ornaments of the telson, of which the present key resorts only to the latter. Nevertheless, Halirages and Bouwerella are not clearly separated by qualitative characters and the validity of the genera Calliopiella, Atylopsis, or Amphithopsis is not convincing, the latter being related closely to Halirages.
Clarencia is not firmly assignable to the Calliopiidae because the uropods and telson were missing on the unique type-specimen.
- Metaleptamphopus appears to be a primitive melphidippid in its elongate uropod 3 and especially resembles Melphisana in the uncleft telson, obsolete accessory flagellum, slightly enlarged coxae and well-developed rostrum.
Almost all of the species, except types, need refiguring of mouth- parts and accessory flagella.
Nomenclatural Changes in Calliopiidae
Atylopsis latupalpus Walker and Scott (1903) should be removed to a eusirid genus. It appears to resemble some species of Pontogeneia.
Pseudopontogeneia Oldevig (1959) is removed to the Eusiridae.
Leptamphopus litoralis Gurjanova (1938) is removed to Calliopiella.
Key to the Genera of Calliopiidae
(Paracalliope and Paraleptamphopus occupy freshwater and are not included in the key.)
1.) -Gnathopod: 2’ chelate (figs 73b) ais Vea) Pe Se es Clarencia Gnathopod 2 subchelate or simple ... . 2 ie Caley,
2. Articles 5-6 of gnathopod 2 together amen cl Solenente Sender rectan- gular, sixth article, 75%, as long as article 2) (fig. W2k) 4) 2) Seu Hither article 5 or 6 of gnathopod 2 elongate and slender, article 6 never more than 60% aslongasarticle2 ... . ras el sie 5 747 Neither articles 5 or 6 elongate and Slondles, net foceanes ecently elongate, usually slightly inflated oroval ..... SR ae ieee)
3. Article 3 of mandibular palp shortened, Afout about ae nid as long as article 2 (fig. 71h), accessory flagellum present, lower lip with inner lobes Oradarea !
Article 3 of mandibular palp subequal to or about four fifths as long as article
2, relatively slender, faleate or pointed terminally (fig. 71g), accessory flagellum absent, lowerliplackinginnerlobes ........... 4
! See footnote 1 on p. 173.
15.
16.
Wie
18.
MARINE GAMMARIDEAN AMPHIPODA 173
Outer ramus of uropod 3 half or less as long asinnerramus . Leptamphopus ! Outer ramus of uropod 3 three fourths or more as long as inner ramus. Bouvierella !
Mowerlip withanner lobes! (igs (id) Want ee he: Fe eae AG Lower lip lacking inner lobes (fig. 7le). . . . Soden lA Article 5 of gnathopods with distinct posterior lobe Ge SRY Ae et Senders Article 5 of gnathopods without distinct posterior lobe (fig. 727) . . . 10 Article 6 of gnathopods slender, bent distally (fig. 720), essentially simple, mandibular molar nontriturative .. . . . . . . Harpinioides Article 6 of gnathopods stout, not bent, spandtiballen molar triturative. . 8 Body dorsally processiferous, pereopod 5 greatly elongate, article 5 as long as articles 2, 3,4, and 6 together ... . . . . Stenopleuroides Body dorsally smooth or without sharp arTeRSeS, pereopod 5 slightly longer than 4, article 5 not disproportionately elongate. . .. . a or Mees) Antenna 1 longer than or equal to antenna 2, telson srovteallls notched or pointed (fig. 72f).. . . . . . . . Stenopleura Antenna 2 slightly longer ‘Pham 1, telson poically somndad (fig. 72d). Calliopius Gnathopods bearing grossly distinct palms (fig. 72m) . . Seep eanedeala| Gnathopods not bearing distinct palms, essentially simple Ge C20) tee els, Articulate accessory flagellum present. ... . . . . . Atylopsis 3 Accessory flagellum absent, occasionally senneasatatl ibys OROCESSiieys Ah eae U2 Coxa 4 slightly longer than coxa 1 (fig. 71a) . . Halirages and Apherusa ? Coxa 4 not longer than coxa 1 (fig. 71b) . . . ... .. . Haliragoides Article 5 of gnathopods much longer than 6 (fig. 73d) . . . . Cleippides Articles 5 and 6 of gnathopods subequal in length (fig. 727). . . Regalia Palp of maxilla 1 reduced, reaching only halfway along outer plate (fig. 71)). Laothoes Palp of maxilla 1 of normal length, Nea to or ae apex of outer platen Giese) 3) cages Ai hah em oa Be tay: a Ee) Dactyls of pereopods mechinate Ge 72b) . Ae aah Metaleptamphonus Dactyls of pereopods weakly serrate or smooth but not pectinate . . . 16
Gnathopod 2 of the EHusirus kind, a large suboval sixth article attached to a slender fifth article (fig. 73a), (body and head extremely flattened).
Sancho
Gnathopod 2 not of the Husirus kind .... . ys ine eer ate Wh Accessory flagellum as long as article 1 of ane “leeallenn (body mod- erately depressed) .... ....... . . Amphithopsis Accessory flagellum scale-like or “sibson pa Has ni Sean aR ecw rene Article 2 of pereopods 1 and 2 smooth anteriorly, body comprcesed lat- erallyaeeie eee BA eS) Article 2 of mer opods 1 and 2 sft paterion mroceet! body premely flattened. Chosroes
1 See also Atylopsis megalops Nicholls (1938); and a few species of Apherusa, Halirages, and Laothoes. Except for A. megalops, none of these species has article 6 of the gnathopods more than 60% as long as article 2. Lao- thoes is characterized by the condition of maxilla 1 palp.
2 Pontogeneoides (Eusiridae) resembles Atylopsis but has an especially elongate telson; it also keys to Stenopleura but has powerfully subchelate gnathopods like Eusiroides, large coxae and elongate telson.
3A few species of Rhachotropis (Eusiridae) may be keyed to this point because of obsolescent telsonic clefts but they may be distinguished by their extremely elongate telsons being more than twice as long as the peduncle of uropod 3.
174 U.S. NATIONAL MUSEUM BULLETIN 271
19. Accessory flagellum absent or represented by a process on article 3 of
antennal ... ENE RER: sn AAD) Accessory flagellum fence of a | eeaneall artmoulleis seals aaiigs Calliopiella
20. Pereopods 1-5 minutely subcheliform ...... .... . Bouvierella Pereopods: 1-5) simple teas votive Nine! psa Gee die ie Waa AONE DED RANE Renae ates So aL Mie en
21. Male antennae with caleeoli ........ 2.2... +... . Halirages Male antennae without calceoli ............ . . Apherusa
22. aeNccessony flagellum presenti. dee erenae tne eal ate Aone Sh fn ieee enna) Accessory flagellum absent. . . . ATER ae
23. Article 5 of gnathopods elongate, aaron faneen and diene thicker than emo G64) bc . . . (exceptional) Cleippides Articles 5 and 6 hoeethen slamainie andl of similar thickness. . . Oradarea
24. Lower lip withsmallinnerlobes . . somespecies of Apherusa and Halirages Lower lip lacking inner lobes. . . . : Div. As eee 74)
25. Outer ramus of uropod 3 half as fone as inner ramus. . iiveprariphonie Outer ramus of uropod 3 three fourths or more aslong asinnerramus . . 26
26. Palp of maxilla 1 reduced, not reaching end of outer plate . . Laothoes Palp of maxilla 1 normal, reaching or exceeding outer plate .... . 27
27. Gnathopod 2 perfectly simple, inner plate of maxilla 1 without setae, palp of maxilliped not as long as outer plate and its parent article combined. Prolaphystius (Laphystiopsidae)
Gnathopod 2 subchelate, inner plate of maxilla 1 setose, palp of maxilliped
nearly twice as long as article and its lobe of outer plate... . . . 28 284e<Body= dorsally smooth: 2.) (005%. 30. 2 we Cae aoe a Bouvierella Body dorsally processiferous . ... . . . . some species of Halirages
Genera of Calliopiidae
Amphithopsis Boeck Amphithopsis Boeck, 1861.—Stebbing, 1906.
Type-species: A. longicaudata Boeck, 1861 (selected by Boeck, 1876). See Sars, 1895.
Antennae equal in length; accessory flagellum as long as article 1 of primary flagellum; lower lip without inner lobes; gnathopodal palms distinct, article 5 slightly longer than article 6, articles together not elongate, article 5 of gnathopod 1 with slight posterior lobe; telson rounded apically, outer ramus of uropod 3 less than half as long as inner, body moderately depressed. Species: 1, subarctic Atlantic (40-1,505 m).
Apherusa Walker
Gossea Bate and Westwood, 1863 (homonym, Coelenterata).
Apherusa Walker, 1891b.—Stebbing, 1906.
not Phaedra Bate, 1858b (possibly type-species a composite of two specimens, head and body).
Type-species: Amphithoe jurinei Milne Edwards, 1830 (monotypy and subsequent synonomy). See Sars, 1895.
MARINE GAMMARIDEAN AMPHIPODA 175
Antenna 1 shorter than antenna 2; accessory flagellum absent; lower lip with slight inner lobes; gnathopodal palms distinct, article 5 slightly longer than article 6 and occasionally elongate, articles 5-6 together not elongate, article 5 of gnathopod 1 lacking posterior lobe; telson pointed, toothed, emarginate or rounded; coza 4 slightly longer than coxa 1 (Haliragordes) ; calceolr absent on male antennae (Halirages). Species: 16, biboreal, littoral.
Atylopsis Stebbing Atylopsis Stebbing, 1888; 1906.
Type-species: A. emarginatus Stebbing, 1888 (present selection).
Antenna 1 shorter than antenna 2 in type but otherwise variable in length; accessory flagellum shorter than article 1 of primary flagellum ; lower lip with inner lobes; gnathopodal palms distinct, articles 5 and 6 together not elongate, article 5 on gnathopod 2 shorter than 6, article 5 of gnathopod 1 as long as 6 or shorter than 6, lacking posterior lobe; enathopod 2 occasionally slightly elongate; telson with deep notch, broad or narrow. Species: 3, antiboreal, subantarctic, littoral (one species to 567 m).
Bouvierella Chevreux Bouvierella Chevreux, 1900.
Type-species: Paramphithoe carcinophilus Chevreux, 1889 (mono- typy).
Antenna 1 slightly longer than antenna 2; accessory flagellum absent; lower lip without inner lobes; gnathopodal palms distinct, article 5 of gnathopod 1 slightly shorter than article 6, with weak posterior lobe, articles 5 and 6 together on gnathopods slightly elongate; telson emarginate; pereopods 1-5 minutely subcheliform (Apherusa, Calliopiella, and Halirages). Very close to Leptamphopus but gnathopod 2 not fully elongate and outer ramus of uropod 3 nearly as long as inner. Species: 1, Atlantic bathyal (620-1,386 m).
Calliopiella Schellenberg Calliopiella Schellenberg, 1925b.
Type-species: Calliopiella michaelsent Schellenberg, 1925b (mono- typy).
Antenna 1| slightly shorter than antenna 2; accessory flagellum scale- like (type) or absent; lower lip without inner lobes; gnathopodal palms distinct, article 5 shorter than article 6, together not elongate, article 5 with slight posterior lobe; telson rounded. Species: 3, W. Africa, N. Pacific temperate, littoral.
176 U.S. NATIONAL MUSEUM BULLETIN 271
Calliopius Liljeborg
Calliope Bate, 1857a (homonym, Aves). Calliopius Liljeborg, 1865 (new name).—Stebbing, 1906.
Type-species: Amphithoe laeviuscula Krgyer, 1838 (monotypy and subsequent synonymy). See Sars, 1895.
Antennae subequal; accessory flagellum absent or article 3 of antenna 1 with a process; lower lip with inner lobes; gnathopods stout, palms distinct, article 5 shorter than article 6, together not elongate, article 5 with distinct posterior lobe; telson rounded. Species: 2, boreal, littoral.
Chosroes Stebbing
Chosroes Stebbing, 1888; 1906.
Type-species: C. incisus Stebbing, 1888 (monotypy).
Body and head extraordinarily depressed; antenna 1 slightly shorter than antenna 2; accessory flagellum absent; lower lip lacking inner lobes; gnathopodal palms distinct, articles 5 and 6 subequal, together not elongate, article 5 lacking posterior lobe; telson notched or cleft; article 2 of pereopods 1 and 2 with large anterior process. Species: 2, antarctic, littoral (1 to 342 m).
Clarencia K. H. Barnard
Clarencia K. H. Barnard, 1931; 1932.
Type-species: C.chelata K. H. Barnard, 1931 (original designation) ; 1932.
Antenna 1 very short, stout, with few articles; accessory flagellum absent; lower lip with tiny fused inner lobes; gnathopod 1 with distinct palm, articles 5 and 6 subequal; gnathopod 2 large, chelate, article 5 tiny and masked by articles 4 and 6, article 6 very large, bearing a thumb; pleonite 4 elongate; [telson unknown]. Species: 1, antarctic (342 m).
Cleippides Boeck Cleippides Boeck, 1871.—Stebbing, 1906.
Type-species: Acanthonotus tricuspis Krgyer, 1846 (monotypy). See Bate, 1862; Gurjanova, 1951.
Antenna 1 much longer than antenna 2; accessory flagellum about half as Jong as article 1 of primary flagellum; inner lobes of lower lip obsolescent; gnathopodal palms indistinct, article 5 much longer than 6, together slightly elongate, article 5 with weak or no posterior lobe; telson rounded. Close to Leptamphopus but inner lobes of lower lip slightly developed and accessory flagellum present. Species: 3, subarctic, littoral to bathyal (1,960 m).
MARINE GAMMARIDEAN AMPHIPODA eee
Halirages Boeck Halirages Boeck, 1871.—Stebbing, 1906.
Type-species: Amphithoe fulvocuncta M. Sars, 1858 (selected by Boeck, 1876). See Sars, 1895.
Antennae subequal or antenna 1 short; accessory flagellum absent or article 3 of antenna 1 with process; lower lip with small or large inner lobes; gnathopodal palms distinct, articles 5 and 6 subequal, together not elongate or article 5 elongate, article 5 lacking distinct posterior lobe; telson emarginate, toothed or pointed; telson and peduncle of uropod 3 elongate, body usually dorsally processiferous (Bounmerella); calceol present on male antennae (Apherusa). Species: 10, bipolar, littoral to abyssal (2,500 m). See note on H. regis (Stebbing, 1914) in “Relationship” of Calliopiidae.
Haliragoides Sars Haliragoides Sars, 1895.—Stebbing, 1906.
Type-species: Halirages vnermis Sars, 1882 (monotypy). See Sars, 1895.
Antenna 1 shorter than antenna 2; accessory flagellum absent; lower lip with inner lobes; gnathopodal palms distinct, article 5 longer than article 6, together not elongate, article 5 lacking posterior lobe; telson pointed or toothed; cora 4 not longer than cora 1 (Apherusa). Species: 3, bipolar, littoral to abyssal (2,450 m).
Harpinioides Stebbing Harpinioides Stebbing, 1888; 1906.
Type-species: H. drepanocheir Stebbing, 1888 (monotypy).
Antenna 1 longer than antenna 2; accessory flagellum more than half as long as article 1 of primary flagellum; mandibular molar tumid, ovate, not triturative; lower lip with slight inner lobes; sizth articles of gnathopods slender, simple, apically bent, tapering, article 7 half as long as article 6, article 5 much shorter than article 6, with distinct posterior lobe; telson notched. This genus also assigned to Pleustidae. Species: 1, Indonesia, Kerguelen (108-496 m).
Laothoes Boeck
Laothoes Boeck, 1871.—Stebbing, 1906.
Thoelaos Della Valle, 1893. Type-species: L. meinerti Boeck, 1871 (monotypy). See Sars 1895. Antenna 1 shorter than antenna 2; accessory flagellum absent; lower
lip lacking inner lobes; gnathopodal palms distinct, articles 5 and 6
equal, together not elongate or slightly elongate, article 5 lacking
178 U.S. NATIONAL MUSEUM BULLETIN 271
posterior lobe; telson slightly emarginate; palp of maxilla 1 reduced, reaching halfway along outer plate. Species: 4, subarctic, littoral to bathyal (1,500 m).
Leptamphopus Sars
Leptamphopus Sars, 1895.—Stebbing, 1906.
Type-species: JL. sars1 Vanhéffen, 1897 (monotypy; L. sarsi is a new name for Sars’ misidentification of Amphithopsis longimana Boeck). For stability, this arrangement should be preserved.
Antenna 1 slightly longer than antenna 2; accessory flagellum absent; lower lip without inner lobes; gnathopodal palms distinct, articles 5 and 6 together elongate; telson notched; article 3 of mandibular palp sub- equal to article 2 or very slightly shorter, relatwely slender, falcate or pointed (Oradarea); outer ramus of uropod 3 half as long as inner (Bouvierella). Species: 2, ?amphiboreal, bipolar, littoral to bathyal (1,505 m).
Metaleptamphopus Chevreux
Metaleptamphopus Chevreux 1911c; 1912a; 1912b.
Type-species: M. pectinatus Chevreux, 1912a (designated by Chevreux) ; 1912b.
Antenna 1 longer than antenna 2; accessory flagellum absent (type) or uniarticulate; lower lip without inner lobes; gnathopodal palms scarcely distinct, articles 5 and 6 equal, together not elongate; article 5 lacking posterior lobe; telson rounded; dactyls of pereopods strongly pectinate. Species: 2, littoral subantarctic to S. Atlantic bathyal (GRY ian).
Oradarea Walker Oradarea Walker, 1903.—Shoemaker, 1930.
Type-species.—O. walkert Shoemaker, 1930 (monotypy; O. walkeri is new name for Walker’s 0. longimana which is a homonym of Boeck’s Amplhithopsis longimana later transferred to Oradarea).
Antenna 2 probably longer than antenna 1; accessory flagellum cylindrical but shorter than article 1 of primary flagellum; lower lip with inner lobes; gnathopodal palms distinct, articles 5 and 6 together elongate; telson truncate, rounded but with minute notch or apically pyramidal; article 3 of mandibular palp short, stout, two thirds as long as article 2 (Leptamphopus). Species: 9, primarily antarctic (6), one bathyal Indonesia, one boreal, littoral to bathyal (15165 mm);
Regalia K. H. Barnard
Regalia K. H. Barnard, 1930. Type-species: fF. fascicularis K. H. Barnard, 1930 (monotypy).
MARINE GAMMARIDEAN AMPHIPODA 179
[Antennae unknown; accessory flagellum unknown]; lower lip with slight inner lobes; gnathopodal palms indistinct, articles 5 and 6 equal, together not elongate, article 5 lacking posterior lobe; telson short, broad, slightly emarginate. Species: 2, one pelagic at New Zealand and one at 304 m in Indonesia.
Sancho Stebbing Sancho Stebbing, 1897; 1906.
Type-species: WS. platynotus Stebbing, 1897 (monotypy).
Body and head extremely depressed; antennae subequal, peduncle of antenna 1 very short; accessory flagellum as long as article 1 of primary flagellum; lower lip without inner lobes; gnathopods not elongate, male gnathopod 1 with transverse palm, article 5 longer than article 6, gnathopod 2 much larger, Eusirus-like; female gnathopod 2 like male gnathopod 1; pereopods spinose as in Haustorvidae; telson tapering but apically truncate. Species: 1, E. Australia, littoral.
Stenopleura Stebbing Stenopleura Stebbing, 1888; 1906.
Type-species: S. atlantica Stebbing, 1888 (monotypy). See Stephensen, 1915; Birstein and Vinogradov, 1958; J. L. Barnard, 1962d.
Antenna 1 slightly longer than or equal to antenna 2; accessory flagellum absent; lower lip with inner lobes; gnathopods moderately stout, palms indistinct, article 5 much shorter than 6 and with distinct posterior lobe, articles 5 and 6 together not elongate, article 7 as long as article 6, latter not bent (Harpinioides); telson apically toothed. Species: 1, probably cosmopolitan epipelagic in tropics. Compare diagnosis of Stenopleuroides.
-
Stenopleuroides Birstein and Vinogradov Stenopleuroides Birstein and Vinogradov, 1964.
Type-species: S. macrops Birstein and Vinogradov, 1964 (mono- typy).
Antenna 1 much longer than antenna 2; accessory flagellum absent; [lower lip unknown]; gnathopods moderately stout, palms indistinct, article 5 much shorter than 6 and with distinct posterior lobe, articles 5-6 together not elongate, article 7 nearly as long as article 6, latter not bent (Harpinioides); telson entire, wnornamented; body dorsally cuspidate (pereonite 7 and pleonites 1-4); pereopod 5 very elongate, article 5 as long as articles 2, 3, 4, and 6 together (Stenopleura). Species: 1, Indian Ocean, probably epipelagic.
180 U.S. NATIONAL MUSEUM BULLETIN 271
Cheluridae
FIGURE 74
DraGcnosis.—Body depressed; segments of urosome immovably fused together, marked ventrally by sutures, segment 3 very large; uropods 1, 2, and 8 radically dissimilar to each other in structure and size; telson foliate, entire; accessory flagellum present; flagellum of antenna 2 clavate, composed largely of one article in the adult. See Corophiidae.
Description.—Accessory flagellum 1l-articulate, stout; flagellum of antenna 2 largely composed of one clavate article; head either bulbous and lacking supra-antennal line, or quadrate and bearing such line; mouthparts basic; coxae short, not overlapping; gnathopod 1 either feeble and slightly chelate, or large and subchelate; gnathopod 2 feeble and slightly chelate; uropods dissimilar, uropod 1 composed of normal peduncle and two short rami; uropod 2 composed of foli- aceous peduncle, with or without rami; uropod 3 composed of short peduncle, a greatly enlarged outer ramus and a scale-like or no inner ramus; telson entire.
ReLatronsuip.— Unlike the Cheluridae, uropods 1 and 2 of the Corophiidae are similar to each other, the second antennal flagellum is distinctly multiarticulate, the third urosomal segment, although occasionally fused to the others, is never greatly enlarged, and the third uropods are never immensely enlarged as in the Cheluridae.
Key to the Genera of Cheluridae
1. Uropod 3 bearing a scale-like inner ramus (fig. 741) ....... Chelura Wropod)3 Jacking an immeriramUs:) 9 is fons eee rte ee ene 2
2. Uropod 2 with two rami (fig. 74k), gnathopod 1 large, subchelate (fig. 747). Tropichelura
Uropod 2 lacking rami (fig. 740), gnathopod 1 small, slightly chelate (fig. 74h). Nippochelura
Genera of Cheluridae
Chelura Philippi Chelura Philippi, 1839—Stebbing, 1906.
Type-species: (. terebrans Philippi, 1839 (monotypy). See Sars, 1895.
Supra-antennal line absent except in defining the lateral ocular lobes; gnathopods small; uropod 2 with two rami; uropod 3 with small inner ramus. Species: 1, cireumamphiboreal, littoral, bores wood.
MARINE GAMMARIDEAN AMPHIPODA 181
Nippochelura J. L. Barnard Nippochelura J. L. Barnard, 1959c.
Type-species: Chelura brevicauda Shiino, 1948 (original designa- tion), 1957.
Supra-antennal line absent, anterodorsal part of head supporting a large process; gnathopods small; uropod 2 lacking rami; uropod 3 lacking inner ramus. Species: 1, Japan, littoral, bores wood.
Ficure 74.—Cheluridae: Chelura terebrans Philippi (Sars, 1895, pl. 225): a, lateral; b, upper lip; c, mandible; d, lower lip; e,f, maxillae 2, 1; g, maxilliped; h,i, gnathopods 1, 2; 7,k,1, uropods 1, 2, 3; m, telson. Tropichelura insulae (Calman) (J. L. Barnard, 1959c): n, male gnathopod 1. Nippochelura brevicauda (Shiino) (J. L. Barnard, 1959c): 0, uropod 2.
182 U.S. NATIONAL MUSEUM BULLETIN 271
Tropichelura J. L. Barnard Tropichelura J. L. Barnard, 1959c.
Type-species: Chelura insulae Calman, 1910 (original designation).
Supra-antennal line present; gnathopod 1 large; uropod 2 with two rami; uropod 3 lacking inner ramus. Species: 1, cireumtropical, littoral, bores wood.
Colomastigidae
FIGguRE 75
Draenosis.—Accessory flagellum absent; body subcylindrical, urosomites 2 and 3 coalesced; coxae very short but overlapping; mandibular palp absent, primary incisor cut into a series of long spine-like teeth; molar problematical, large but nontriturative in some species, absent or unknown in others; inner plates of maxillipeds very small, fully or partially coalesced, unarmed or acuminate; uropod 8 biramous; telson entire. Monogeneric. See Eophliantidae, Phliantidae, Prophliantidae.
Description.—Accessory flagellum absent; body subcylindrical; coxae short but overlapping; epistome projecting acutely in front; upper lip bilobed; mandible lacking palp, its body cut into several long spine-teeth, molar large and unridged, small or ‘‘absent’’; lower lip poorly known; maxillae stunted, maxilla 1 with 1-articulate palp larger than outer plate; lobes of maxilla 2 partially coalesced; inner plates of maxilliped very small, fused, or split but forming a sub- conical piece, outer lobes of medium size but appearing large because the articles they are attached to are elongate; gnathopod 1 simple, usually armed apically with spines or setae forming a brush, some males lacking these setae; gnathopod 2 in male enlarged, subchelate, in female small and subchelate; uropod 3 biramous, occasionally outer ramus considerably reduced in size; telson short to medium in length, entire, often apically ornamented.
ReLatTionsuip.—The Eophliantidae also have cylindrical bodies but differ from colomastigids by the well-developed inner lobes of the maxillipeds and the cleft telson; the mandible is not cut into a series of long projecting teeth and the palp of maxilla 1 is degraded.
Phliantidae and Prophliantidae have compressed, or depressed, not cylindrical bodies. The latter have cleft telsons. Probably all phliantids have a reduced inner ramus of uropod 3 and prophliantids have all urosomal segments coalesced.
MARINE GAMMARIDEAN AMPHIPODA 183
Possibly the primary cutting edge of the colomastigid mandible is missing and the digitations represent enlarged and basally fused spines of the normal gammaridean spine-row.
Corophiidae and Podoceridae may resemble Colomastigidae be- cause of depressed and subcylindrical bodies but the former two families have mandibular palps and normal maxillipeds.
Figure 75.—Colomastigidae: a, Colomastix pusilla Grube (J. L. Barnard, 1955a). Telson: b, C. pusilla; c, Colomastix subcastellata Hurley (1954d). Mandible: d, C. pusilla. Maxilla 1: e, C. subcastellata; f, C. pusilla. Maxilla 2: g, C. subcastellata; h, C. pusilla. Maxillipeds: i, C. subcastellata; 7, C. pusilla. Uropod 3: k, C. pusilla; l, C. magnirama Hurley (1954d). Gnathopod 1, C. pusilla: m, female; n, male. Gnathopod 2, C. pusilla:
o, male; p, female.
184 U.S. NATIONAL MUSEUM BULLETIN 271
Genera of Colomastigidae
Colomastix Grube
Colomastix Grube, 1861; 1864a.—Stebbing, 1906. Cratippus Bate, 1862. Exunguia Norman, 1869b. Type-species: Colomastiz pusilla Grube, 1861 (monotypy). See J. L. Barnard, 1955a. Species: 9, cosmopolitan, littoral.
Corophiidae
FigurEs 76-80
Diacnosis.—Accessory flagellum variable, often absent; urosome visibly depressed; wrosomal segment 2 subequal to segment 1 in length; telson entire; pereopods often glandular. See Isaeidae, Aoridae, Podoceridae, Ischyroceridae, Cheluridae.
DescriptTion.—Accessory flagellum multiarticulate, 1-articulate or absent; body often subcylindrical, generally depressed, especially urosome, some or all of urosomites often coalesced; body rarely carinate and only on urosome; coxae long, of medium length, or short, often not touching each other, occasionally acuminate; mouthparts basic, except mandibular palp occasionally reduced to one or two articles and upper lip occasionally bilobed; gnathopods variable, either pair the larger, often complexly subchelate; uropods 1 and 2 usually normal, rarely with inner rami reduced, rarely with inner ramus on uropod 2 absent; uropod 3 variable: (1) bearing two rami, the inner reduced in size; (2) bearing one ramus, either longer than peduncle or very short; or (3) rami absent; telson entire, fleshy, circular or symmetrically trapezoidal, or very broad and short, occasionally armed with hooks; pereopod 5 occasionally very elongate and bearing long setae like fossorial families.
ReELATIONSHIP.—The Corophiidae are a mixture of isaeid, aorid, and ischyrocerid kinds having the urosome especially depressed and usually losing the inner ramus of the third uropods, although this condition is found also in some isaeids. It is very difficult to distinguish these three families, as exemplified by the genus Aorcho, originally assigned to Corophiidae but now provisionally assigned to Aoridae (see discussion under the Isaeidae). Because of its tall urosome, Unciolella is another genus of doubtful assignment to the Corophiidae. Neomicrodeutopus (Aoridae) and Grandidierella (Corophiidae) are very closely related.
MARINE GAMMARIDEAN AMPHIPODA 185
Ficure 76.—Corophiidae: a, Cerapus crassicornis (Bate) (Sars, 1895, pl. 217); b, Corophium affine Bruzelius (Sars, 1895, pl. 221); c, Gaviota podophthalma J. L. Barnard (1958b); d, Kamaka kuthae Derjavin (Gurjanova, 1951).
186 U.S. NATIONAL MUSEUM BULLETIN 271
Because of its peculiar mandibular palp and immensely thickened male antenna 2, Corophium, the type-genus, is not typical of most other genera assigned to the family. Dr. E. L. Bousfield (in litt.) has pointed out to me the probability that Hricthonius, with its elongate peduncle of uropod 3, belongs with the Ischyroceridae.
The Podoceridae differ from the Corophiidae by their elongate urosomal segment 1.
Corophiidae may resemble Eophliantidae and Colomastigidae be- cause of their depressed, subcylindrical bodies and short, serially gaping coxae but the latter two families have degraded mandibles.
Nomenclatural Changes in Corophiidae
Aorcho J. L.. Barnard (1961) is removed to the Aoridae.
Bogenfelsia J. LL. Barnard (1962d) is removed to the Ischyroceridae.
Dryopoides Stebbing (1888) was in Aoridae in Stebbing (1906) and transferred to Corophiidae by Stebbing (1910). Female keys out to the Aoridae.
Runanga J. L. Barnard (1961) removed from Podoceridae.
Key to the Genera of Corophiidae
(All species having gnathopod 1 larger than 2 should also be keyed out in the Aoridae.)
1. Uropod 3 lacking rami (fig. 80n) . . ....... .. . . Concholestes Uropod:3) bearingal—2 rami (figs 80b—2)) 3 a. 5 a 2 ee
2. Mandibular palp 3-articulate (fig. 789). . . . . Lig 2a a2 ee ne Mandibular palp less than 3-articulate (figs. 78e, f). Peer del xh Pall
3. Inner ramus of uropod 8 distinct (fig. 80h)... ... REN Stine Inner ramus of uropod 3 absent or indistinct (uropod 3 Stee one ramus only) (higstiSObS gu renieiie SON plead voi ahh Wate. ineepeee a aaa
4. Coxae not touching erally, Ges, 76b 7a) OR aC) Coxae touching serially (figs. 76c,d) .... . SY Sahay take 1
5. Gnathopod 1 scarcely larger than and similar to pnethonod 2, waalice smooth, inner ramus of uropod 3 distinct but much shorter than outer ramus (fig. 80h), third pleonal epimeron smoothly quadrate ... .. . Camacho
Gnathopod 1 much larger than 2, palm sculptured, inner ramus of uropod 3 usually represented by a large peduncular process (fig. 80z), rarely distinct or marked off, when distinct scarcely shorter than outer ramus, third
pleonal epimeron bearing posteroventral tooth . . ... . . . Unciola 6. Inner ramus of uropod 3 much shorter than outer ramus, gnathopod 2 mero- chelate... . . . Paracorophium
Rami of uropod 3 sapmally lone, poathonedl 2 not meroehelare: male of Dryopoides
10.
Jhaik,
12.
13.
14.
15.
16.
IT
18.
I).
20.
21.
MARINE GAMMARIDEAN AMPHIPODA 187
Wropodyawithioneramusx(i gS 0) vee ter eee Se ee ne a8 Uropod 2 with tworami (fig. 80h) ...... 5 1@ Pleopod 3 with one ramus, [article 3 of antenna 1 loner chan antici 1]. Runanga Pleopod 3 with two rami, [article 3 of antenna 1 rarely longer than article Laan ceo 9 Male peetnonad 2, mate paethonod) 1, pampleniy, Baenelere Ge 791), head with thorn-like rostrum andeyes. . . . . . Cerapus Male gnathopod 1, not gnathopod 2, complexly Sapenelits head with ob- solescent rostrum and no eyes . . . .. . . . Chevreuxius Inner rami of uropods 1—2 less than half as Tener as outer rami (fig. 807). Pseudericthonius Inner rami of uropods 1—2 usually subequaltoouterrami ....... Ii PMILenmagelackine accessory tagellum™ 9. 9, foe eee ees MD Antenna 1 bearing accessory flagellum .. . is 1) Male gnathopod 2 complexly subchelate Gaol) i 79k). Ericenonine Male gnathopod 2 not complexly subchelate ... . ; ficial o gaelic} Ficonitest4—5srcoalescedeise othe Cota oye o.oo a. a 7s Kamaka Pleonites 4-5 separate... . Btacor lah Ue eect mantat shenmiienl Ul Male gnathopod 2 chelate Gugpododnens) - . . .. . . . Cerapopsis Male gnathopod 2 subchelate (fig. 793) . .......... .. Gaviota PNCEESSOLYAlagellUmMeMNUltiantiCulate pala ie eee eee ha os ae iG PNCCeSSOlveHacellum: dearticulater, ws enue smi eae een ations 1G Coxae angular .. . 2 SURED ROA E PR SAR eR COLE ORME ANG: Coxae with softly semadled Cornersi aye Ck. mapasgedles,
Article 6 of gnathopod 2 an elongate nagiemele. Prarvenclae or palm trans- verse, antennae usually much shorter than body and flagellum of antenna 2 scarcely longer than peduncular article 5, peduncle of uropod 3 often expanded medially very eran antenna 2 generally considered stout, often strongly so... . . . Unciola
Article 6 of gnathopod 2 ouate or ene ordeal eepanded palm oblique, antennae usually as long as or longer than body, flagellum of antenna 2 nearly as long as all articles of peduncle, peduncle of uropod 3 rarely expanded medially and usually poorly so, antenna 2 very slender.
Neohela
Inner plate of maxilla 1 with one seta, ramus of uropod 3 about 1.5 times aslongas peduncle. ... . . .. . . Unciolella Inner plate of maxilla 1 with about 10 seine lesadiays medial edge, ramus of uropod 3 about twice as long as peduncle. . . ... . . . Parunciola Male gnathopod 1 complexly subchelate. . . .. . . . . Grandidierella Male gnathopod 1 simply subchelate .. . eee 20
Male antenna 2: articles 3-5 stout (fig. 76b), loneen Paani uropod 3 with asymmetrically prolonged peduncle (fig. 802), or uropod 2 with vestigial inner ramus . . . . . Unciola
Male antenna 2: articles 3-5 Blender Ges. 76a, a, 77a), monedl 2 with sym- metrical peduncle (fig. 80f), rami of uropod 2 subequal.
Unciolella (juvenile)
Mandibular palp l-articulate (fig. 78e). . .... . . . . Siphonoecetes
Mandibular palp 2-articulate (fig. 78f) . .... .. .. . . Corophium
285-135 O - 69 - 13
188 U.S. NATIONAL MUSEUM BULLETIN 271 Genera of Corophiidae
Camacho Stebbing
Camacho Stebbing 1888; 1906.
Type-species: C. bathyplous Stebbing, 1888 (monotypy). See J. L. Barnard, 1961.
Accessory flagellum multiarticulate; article 3 of antenna 1 much shorter than article 1, flagellum very long; antenna 2 much shorter than antenna 1, slender, flagellum shorter than article 5 of peduncle; mandibular palp 3-articulate; coxae short, not touching serially; enathopods normally subchelate; uropod 2 biramous; uropod 3 with outer ramus longer than peduncle, inner ramus one third as long as outer. Species: 1, antiboreal, eurybathic, littoral to abyssal.
Cerapopsis Della Valle Cerapopsis Della Valle, 1893.—Stebbing, 1906.
Type-species: C. longipes Della Valle, 1893 (monotypy).
Accessory flagellum absent; article 3 of antenna 1 equal to article 1; flagellum slightly longer than peduncle; antenna 2 slender, slightly longer than antenna 1, flagellum slightly longer than peduncle; mandibular palp 3-articulate; coxae large, touching serially; gnatho- pods normally subchelate in female, gnathopod 2 chelate in male, enlarged, with digitate thumb; uropod 2 biramous; uropod 3 with its single ramus as long as peduncle; head with long lateral, ocular lobes. Species: 1, Mediterranean, littoral.
Cerapus Say
Cerapus Say, 1817.—Stebbing, 1906. Cerapodina Milne Edwards, 1840.
Type-species: C. tubularis Say, 1817 (monotypy). See J. L. Barnard, 1962a.
Accessory flagellum absent; article 3 of antenna 1 shorter than or 120% as long as article 1, flagellum as long as article 3; antenna 2 slender, equal to antenna 1, flagellum as long as article 5; mandibular palp 3-articulate; coxae short, not touching serially; gnathopod 1 normally subchelate, gnathopod 2 in female simple even though article 6 slightly inflated, in male very large, essentially carpochelate, with teeth on article 5, article 6 simple and more slender than 5, dactyl very large and folding back to middle of article 5; uropod 2 with one ramus; uropod 3 with one vestigial, hooked ramus; head with thorn-like rostrum. Species: 5, (possibly less), cosmopolitan, littoral (one
bathyal).
MARINE GAMMARIDEAN AMPHIPODA 189
Figure 77.—Corophiidae: a, Unciolella lunata Chevreux (191la); 6, Paracorophium excavatum (Thomson, 1884).
190 U.S. NATIONAL MUSEUM BULLETIN 271
Chevreuxius Bonnier Chevreuxius Bonnier, 1896.—Stebbing, 1906.
Type-species: C. grandimanus Bonnier, 1896 (monotypy).
Accessory flagellum 2-articulate; article 3 of antenna 1 slightly shorter than article 1, flagellum slightly longer than article 3; antenna 2 slender, equal to antenna 1, flagellum shorter than article 5; mandib- ular palp 3-articulate; coxae short, not touching serially; male gnathopod 1 much larger than gnathopod 2, carpochelate, article 6 stout and subchelate but narrower than article 5; gnathopod 2 sub- chelate; uropod 2 with one ramus; uropod 3 with its single ramus as long as peduncle. Species: 1, N. Atlant-c, bathyal (950 m).
Concholestes Giles Concholestes Giles, 1888.—Stebbing, 1906.
Type-species: C. dental Giles, 1888 (monotypy).
Accessory flagellum absent; article 3 of antenna 1 subequal to article 1, flagellum equal to article 3; antenna 2 of medium stoutness, longer than antenna 1, flagellum much shorter than article 5; [mandible not described]; coxae not touching serially; gnathopod 2 normally sub- chelate, gnathopod 1 nearly simple; uropod 2 with ?one ramus; uropod 3 lacking rami. Species: 1, Bay of Bengal, littoral, in shell of Dentalium. See Uncinotarsus in Aoridae.
Corophium Latreille
Corophium Latreille, 1806.—Stebbing, 1906. Audouinia Costa, 1851b (nom. nud.).
Type-species: Oniscus volutator Pallas, 1776 (monotypy and sub- sequent synonymy). See Sars, 1895 (as C. grossipes).
Accessory flagellum absent; article 3 of antenna 1 shorter than article 1, flagellum longer than article 3; antenna 2 equal to or longer than antenna 1, stout, especially in male flagellum much shorter than article 5, male article 4 usually with a distal tooth; mandibular palp 2-articulate; coxae short, not touching serially; gnathopods small, gnathopod 1 subchelate, gnathopod 2 simple, extremely setose, articles 4 and 5 elongate and anteroposteriorly apposed; uropod 2 biramous; uropod 3 flattened, its single ramus equal to peduncle; urosomites coalesced or free. Species: 47, cosmopolitan, open-sea littoral, brackish, fresh-water and Caspian Sea.
Dryopoides Stebbing Dryopoides Stebbing, 1888.—Stebbing, 1906.
Type-species: D. westwood: Stebbing, 1888 (monotypy). See Stebbing, 1910.
MARINE GAMMARIDEAN AMPHIPODA 191
u f
=
eX AR ?
e/a mr
Figure 78.—Corophiidae: a, Bogenfelsia incisa J. L. Barnard (1962d); b, Camacho bathy- plous Stebbing (J. L. Barnard, 1961). Upper lip: c, Neohela monstrosa (Boeck) (Sars, 1895, pl. 224); d, Corophiwm volutator (Pallas) (Sars, 1895, pl. 219 as C. grossipes). Man- dible: e, Siphonoecetes colletti Boeck (Sars, 1895, pl. 218); f, Corophium; g, Ericthonius brasiliensis (Dana) (Sars, 1895, pl. 215, as E. abditus). Mouthparts, Corophium: h, lower lip, 2,7, maxillae 1, 2, &, maxilliped. Maxilliped: /, Ericthonius.
192 U.S. NATIONAL MUSEUM BULLETIN 271
Accessory flagellum absent; article 3 of antenna 1 much shorter than article 1, flagellum longer than peduncle; antenna 2 slender, slightly shorter than antenna 1, flagellum shorter than article 5; mandibular palp 3-articulate; coxae of medium size, touching serially; enathopods normally subchelate, gnathopod 2 larger than 1 in male, smaller than 1 in female; uropod 2 biramous; uropod 3 biramous, rami shorter than peduncle. Female assigned to Aoridae. Species: 1, E. Australia, littoral.
Ericthonius Milne Edwards
Ericthonius Milne Edwards, 1830.—Stebbing, 1906. Pyctilus Dana, 1852b.
Type-species: HH. difformis Milne Edwards, 1830 (selected by Stebbing, 1888, p. 142!). See Sars, 1895.
Accessory flagellum absent; article 3 of antenna 1 longer than article 1, flagellum subequal to peduncle; antenna 2 slender, equal to antenna 1, flagellum as long as peduncle; mandibular palp 3-articulate ; coxae short, scarcely touching serially; male gnathopod 2 larger than 1, carpochelate, article 6 simple, female gnathopods normally sub- chelate; uropod 2 biramous; uropod 3 with its single, medium length, uncinate ramus shorter than peduncle. Species: 8, mainly N. Hemisphere and tropics (one species), littoral (one extends to 1,134 m).
Gaviota J. L. Barnard Gaviota J. L. Barnard, 1958b.
Type-species: G. podophthalma J. L. Barnard, 1958b (original designation).
Accessory flagellum absent; article 3 of antenna 1 equal to article 1, flagellum longer than article 3; antenna 2 slender, equal to antenna 1, flagellum longer than article 5; mandibular palp 3-articulate; coxae long, touching serially; gnathopods normally subchelate; uropod 2 biramous; uropod 3 with its single ramus shorter than peduncle; head with vmmense, pedunculate lateral ocular lobes. Species: 1, California, littoral.
Grandidierella Coutiére Grandidierella Coutiére, 1904.
Type-species: G. mahafalensis Coutiére, 1904 (monotypy). See Ruffo, 1958.
Accessory flagellum 1l-articulate, usually elongate; article 3 of antenna | shorter than article 1, flagellum nearly as long as peduncle; antenna 2 slender (rarely stout in male), nearly equaling antenna 1, flagellum often shorter than article 5; mandibular palp 3-articulate;
MARINE GAMMARIDEAN AMPHIPODA 193
——=—
— a eS= Ss
Ficure 79.—Corophiidae: a, Neohela monstrosa (Boeck) (Sars, 1895, pl. 224). Gnathopod 1: b, Ericthonius brasiliensis (Dana) (Sars, 1895, pl. 215, as FE. abditus); c, Unciola leucopis (Krgyer) (Sars, 1895, pl. 222); d, Siphonoecetes colletti Boeck (Sars, 1895, pl. 218); e, Corophium volutator (Pallas) (Sars, 1895, pl. 219, as C. grossipes); f, male Cerapopsis longipes Della Valle (1893). Gnathopod 2: g, Cerapopsis; h, Corophium; 1, Unciola; 7, Siphonoecetes; k, Ericthonius; 1, Cerapus crassicornis (Bate) (Sars, 1895, pl. 217).
194 U.S. NATIONAL MUSEUM BULLETIN 271
coxae short, scarcely touching serially; male gnathopod 1 large, carpochelate, article 5 bearing a tooth (or two), article 6 simple, enathopod 2 smaller, normally subchelate or slightly chelate, and articles 4-5 sometimes like Corophium; uropod 2 biramous; uropod 3 with its single ramus longer than peduncle. Species: 15, circum- tropical and warm-temperate, littoral, especially brackish water. Differs from Neomicrodeutopus in Aoridae by elongate article 1 of mandibular palp, that article as long as article 2.
Kamaka Derjavin Kamaka Derjavin, 1923.—Gurjanova, 1951.
Type-species: K. kuthae Derjavin, 1923 (monotypy).
Accessory flagellum absent; antenna 1 short, article 3 slightly shorter than article 1, flagellum equal to peduncle; antenna 2 much longer than antenna 1, slender, flagellum equal to or shorter than article 5, often about 2-articulate; mandibular palp 3-articulate; coxae of medium size, touching serially ; gnathopods normally subchelate and male gnathopod 2 with large palmar tooth; uropod 2 biramous; uropod 3 with its single ramus slightly shorter than (or subequal to) peduncle; pleonites 4-5 coalesced. Species: 3, boreal N. W. Pacific, littoral.
Neohela Smith
Hela Boeck, 1861 (homonym, Decapoda). Neohela Smith, 1881 (new name).—Stebbing, 1906. Helella Sars, 1882.
Type-species: Hela monstrosa Boeck, 1861 (monotypy). See Sars, 1895.
Accessory flagellum multiarticulate; article 3 of antenna 1 longer than article 1, flagellum longer than peduncle; antenna 2 slender, slightly longer than 1, flagellum longer than peduncle; mandibular palp 3-articulate; coxae short, some sharp, not touching serially; enathopods normally subchelate, gnathopod 1 slightly larger than enathopod 2; uropod 2 biramous; uropod 3 with its single ramus equal to or twice as long as peduncle; telson slightly enlarged and apparently coalesced with urosomite 3; inner plate of maxilla 1 unth about five setae mostly terminal (Parunciola). Species: 3, subarctic-boreal, littoral to abyssal (2,288 m).
Paracorophium Stebbing
Paracorophium Stebbing, 1899b, c; 1906.
Type-species: Corophium excavatum Thomson, 1884 (monotypy). See Hurley, 1954b.
MARINE GAMMARIDEAN AMPHIPODA 195
Figure 80.—Corophiidae: a, Runanga coxalis J. L. Barnard (1961). Uropod 2: b, Cerapus crassicornis (Bate) (Sars, 1895, pl. 217). Uropod 3: c, Cerapus; d, Ericthonius brasili- ensis (Dana) (Sars, 1895, pl. 215, as E. abditus); e, Corophium crassicorne Bruzelius (Sars, 1895, pl. 220); f, Siphonoecetes colletti (Boeck) (Sars, 1895, pl. 218); g, Neohela monstrosa (Boeck) (Sars, 1895, pl. 224); h, Camacho bathyplous Stebbing (J. L. Barnard, 1961); 2, Unciola leucopis (Krdyer) (Sars, 1895, pl. 222). Urosome, dorsal: 7, Pseudericthonius gausst Schellenberg (1926a); k, Corophium volutator (Pallas) (Sars, 1895, pl. 219, as C. grossipes); 1, Cerapus; m, Corophium crassicorne; n, Concholestes dentalii Giles (1888). Telson: 0, Cerapus; p, Ericthonius difformis Milne Edwards (Sars, 1895, pl. 216); gq, Neohela; r, Unciola; s, Corophium.
196 U.S. NATIONAL MUSEUM BULLETIN 271
Accessory flagellum absent; article 3 of antenna 1 shorter than article 1, flagellum longer than peduncle; antenna 2 stout, shorter than antenna 1, flagellum slightly longer than article 5; mandibular palp 3-articulate; coxae of medium size, touching serially; gnathopod 1 normally subchelate; gnathopod 2 subchelate or chelate, article 4 greatly prolonged as in Corophium but diverging from article 5 and strongly setose; uropod 2 biramous; uropod 3 with outer ramus slightly shorter than peduncle, inner ramus half as long as outer. Species: 2, New Zealand, fresh-water.
Parunciola Chevreux
Parunciola Chevreux, 1911a.
Type-species: P. seuratt Chevreux, 191la (monotypy).
Accessory flagellum multiarticulate; article 3 of antenna 1 much longer than article 1 [flagellum broken]; antenna 2 slender, equaling antenna 1 [flagellum broken]; mandibular palp 3-articulate; coxae small, with softly rounded corners, but touching serially; gnathopods normally subchelate; uropod 2 biramous; uropod 3 with its single ramus twice as long as peduncle; inner plate of maxilla 1 with about 10 medial setae (Neohela, Unciolella). Species: 1, Algeria, littoral.
Pseudericthonius Schellenberg Pseudericthonius Schellenberg, 1926a.
Type-species: P. gaussi Schellenberg, 1926a (monotypy).
Antenna | not described but apparently as in Hricthonius; antenna 2 slender; mandibular palp 3-articulate; coxae short, not touching; enathopods normally subchelate; uwropods 1 and 2 with inner rami much shorter than outer; uropod 3 with its single elongate uncinate ramus shorter than peduncle. Species: 1, antarctic, littoral (to 342 m).
Runanga J. L. Barnard
Runanga J. L. Barnard, 1961.
Type-species: R&. coxalis J. L. Barnard, 1961 (original designation).
Accessory flagellum scale-like; article 3 of antenna 1 much longer than article 1, flagellum longer than article 3; antenna 2 slender, equal to antenna 1, flagellum longer than article 5; mandibular palp 3- articulate; coxae short, not touching serially; gnathopods normally subchelate; uropod 2 with one ramus; uropod 3 with two vestigial rami, outer ramus hooked; pleopod 3 with one ramus. Species: 1, Tasman Sea, bathyal (610 m).
MARINE GAMMARIDEAN AMPHIPODA 197
Siphonoecetes Kréyer Stphonoecetes Kréyer, 1845.—Stebbing, 1906.
Type-species: S. typicus Krgyer, 1845 (monotypy). See Stephensen, 1944a.
Accessory flagellum absent; article 3 of antenna 1 subequal to article 1, flagellum subequal to or longer than article 3; antenna 2 slender or slightly thickened, longer than antenna 1, flagellum shorter than article 5; mandibular palp 1l-articulate; coxae short, not touching serially; gnathopods weakly or normally subchelate; uropod 2 with 1-2 rami; uropod 3 with medially expanded pedunele, its single ramus shorter than peduncle. Species: 9, arctic-boreal to ?antisubtropics, littoral to bathyal (565 m).
Unciola Say
Unciola Say, 1818.—Stebbing, 1906. Glauconeme Krgyer, 1845 (homonym, Bryozoa). Dryope and Driope Bate, 1862 (Dryope=homonym, Diptera).
Type-species: U. irrorata Say, 1818 (monotypy). See Shoemaker, 1945b.
Accessory flagellum elongate, multiarticulate or 1-articulate; article 3 of antenna 1 equal to article 1, flagellum subequal to peduncle; antenna 2 of medium to strong stoutness, slightly shorter than antenna 1, flagellum subequal to article 5; mandibular palp 3-articulate; coxae short, not touching serially; gnathopod 1 larger than gnathopod 2, both normally subchelate or gnathopod 2 simple or parachelate; uropod 2 biramous; uropod 3 often with medially expanded pedunele, its single ramus slightly shorter than peduncle. Species: 15, arctic- boreal, littoral to abyssal.
Unciolella Chevreux Unciolella Chevreux, 1911la.
Type-species: U. lunata Chevreux, 191la (monotypy).
Accessory flagellum 1-articulate, elongate or multiarticulate; article 3 of antenna 1 equal to or shorter than article 1, flagellum equal to peduncle; antenna 2 slender, shorter than antenna 1, flagellum shorter than article 5; mandibular palp 3-articulate; coxae short, touching or not touching serially, with softly rounded corners; gnathopods nor- mally subchelate, gnathopod 1 shghtly the larger; uropod 2 biramous; uropod 3 with its single ramus about 1.5 times longer than peduncle; inner plate of maxilla 1 with one terminal seta (Neohela, Parunciola). Species: 2, Mediterranean and S. Atlantic, littoral (to 392 m).
198 U.S. NATIONAL MUSEUM BULLETIN 271
Cressidae Ficures 81, 82
Diacnosis.—Accessory flagellum absent; mandibular molar evanes- cent; coxa 1 very small, always partially covered by following coxae; coxa 4 large, either excavate posteriorly or shield-like and not excavate; uropod 3 uniramous; outer lobes of maxilliped vestigial; telson entire and fused with pleonite 6, latter often coalesced with pleonite 5; article 2 of pereopod 3 expanded. See Stenothoidae, Amphilochidae, Thaumatelsonidae, Pagetinidae, Leucothoidae, Anamixidae, Phliantidae.
if g y
Ficure 81.—Cressidae: Cressa dubia (Bate) (Sars, 1895, pls. 98-99): a, lateral; b, upper lip; c, lower lip; d, telson; ¢,f, gnathopods 1, 2; g, maxilliped; h,7, maxillae 1, 2; 7, mandible.
MARINE GAMMARIDEAN AMPHIPODA 199
DescripTion.—Rostrum inconspicuous; accessory flagellum absent; body smooth or with slight dorsa] teeth; coxa 1 small, hidden by follow- ing coxae; coxa 4 large, either shield-like or excavate posteriorly; upper lip incised; mandible with weak, sparsely spinose molar, not triturative, palp long, 3-articulate; lower lip with coalesced inner lobes, outer lobes with blunt extremities; palp of maxilla 1 uniartic- ulate; maxilla 2 small, lobes blunt; maxillipeds slender, outer lobes vestigial; gnathopods small, first simple, second subchelate; uropod 3 uniramous, the single ramus 2-articulate; telson of medium length, entire, fused with pleonite 6; article 2 of pereopod 3 expanded.
a
SS ase ae 4
LEE
\
Figure 82.—Cressidae: a, Cressina monocuspis Stephensen (1931); b, uropod 3 of Cressa minuta Boeck (Sars, 1895, pl. 99); c, pleonite 3 and urosome, left to right, of Cressina, showing absence of urosomite 3.
RELATIONSHIP.—The Stenothoidae have all segments of the urosome and the telson distinct, coxa 4 is always shield-like, not excavate posteriorly (like Cressina, however) and article 2 of pereopod 3 is always linear.
All Thaumatelsonidae have coxa 4 shield-like, not excavate pos- teriorly. Although the segments of the urosome may be coalesced the telson is not clearly fused with them and is usually thickened dorso- ventrally; article 2 of pereopod 3 is always linear.
The Phliantidae have depressed, rugose bodies, and relatively equal coxae.
200 U.S. NATIONAL MUSEUM BULLETIN 271
Key to the Genera of Cressidae
1. Coxa 4 rectangular, excavate posteriorly (fig. 8la) ........ Cressa Coxa 4 shield-like, not excavate posteriorly (fig. 82a). ..... Cressina
Genera of Cressidae
Cressa Boeck
Danaia Bate, 1857a (homonym, Anthozoa). Cressa Boeck, 1871.—Stebbing, 1906.
Type-species: Danaia dubia Bate, 1857a (selected by Boeck, 1876, as Cressa schiodte. Boeck, 1871). See Sars, 1895.
Species: 6, arctic-boreal, N. Atlantic, littoral to bathyal.
Cressina Stephensen Cressina Stephensen, 1931.
Type-species: C. monocuspis Stephensen, 1931 (monotypy). Species: 1, subarctic, bathyal (682—1,096 m).
Dexaminidae
[including Anatylidae Bulycheva, 1955] Figures 83-85
Draenosis.—Accessory flagellum absent; urosomites 2 and 3 coalesced; mandible lacking palp, molar well developed or vestigial; uropod 3 biramous; telson deeply cleft; body laterally compressed; gnathopods feeble but subchelate. See Atylidae, Prophliantidae, Phlian- tidae. See Sphaerophthalmus in ‘‘Incertae Sedis.”’
Description.—Accessory flagellum absent; body laterally com- pressed and often dorsally carinate or toothed; urosomites 2 and 3 coalesced; mandible lacking palp, molar well developed, triturative (Dexamininae) or evanescent (Anatylinae) ; lower lip with or without mandibular processes, inner lobes well developed or vestigial, occa- sionally fused; palp of maxilla 1 uni- or biarticulate; maxillipeds with 3- or 4-articulate palp, inner plates occasionally vestigial; gnathopods feeble, subchelate; pereopods occasionally cheliform; uropod 3 biramous; telson long or short, deeply cleft.
Rexationsuie.—The Talitroidea have fully separated urosomal segments.
Phliantidae and Prophliantidae have the mandibular molar evanes- cent. The Phliantidae, furthermore, have the palp of maxilla 1 de- graded or absent. The Prophliantidae have all urosomal segments coalesced.
MARINE GAMMARIDEAN AMPHIPODA 201
Ficure 83.-—Dexaminidae: a, Polycheria antarctica (Stebbing, 1888); b, Dexamonica redun - cans J. L. Barnard (1958c).
202 U.S. NATIONAL MUSEUM BULLETIN 271
Probably Sphaerophthalmus Spandl is assignable to the Dexamin-
idae. See its diagnosis under ‘“‘Incertae Sedis.”’
co
(oy)
“I
Key to Subfamilies of Dexaminidae Mandibular molar large, triturative (fig.847) ... . . . . Dexamininae Mandibular molar evanescent (fig. 85e) ...... . . . .Anatylinae
Key to the Genera of Dexaminidae
(expanded after Sheard, 1938)
Palp of maxilliped with three articles (fig. 84d). ........... 2.2 Palp of maxilliped with four articles (fig. 84e) ............ .4 Inner lobes of lower lip well developed (fig. 84h) . . . . . . Dexaminella Inner lobes of lower lip vestigial (fig. 847) . . . . BY ke
Pereopods 1—5: article 4 shorter than articles 5-6 combined Ge 85n). Dexamine
Pereopods 1-5: article 4 longer than articles 5-6 combined (fig. 850). Tritaeta Pereopods 1-5 cheliform (fig. 85p) ........... .... .Polycheria Pereopods 1-5 simple. . . . Sane ACME INEM CCE Gy AG ison) 6 Palp of maxilla 1 uniarticulate Ge. 84m) PURER. VR OI UU ara Palp of maxilla 1 biarticulate (fig. 847) .... Manes itiay nats) Inner plates of maxillipeds vestigial, short and bud-like Gee, 84c) . Pee n ats 276 Inner plates of maxillipeds of medium size (fig. 84d) ......... .8 Inner lobes and mandibular lobes of lower lip distinct; palp article 4 of maxilliped claw-shaped (fig. 84e) ... . . . . . .Dexaminoides Inner lobes of lower lip coalesced, remelirale lobes obsolescent (fig. 84f); palp article 4 of maxilliped stout, blunt (fig. 84c). . . . . Dexamonica Mandibular processes of lower lip absent (fig. 84g). . . . . Syndexamine Mandibular processes of lower lip present (fig. 847). . . . . Paradexamine
Palp article 2 of maxilla 1 large (fig. 841); inner plates of maxillipeds well developed (fig. 857); coxa 5 small, coxa 4 not posteriorly excavate.
Anatylus
Palp article 2 of maxilla 1 small (fig. 84k) or large; inner plates of maxillipeds
vestigial, short and bud-like (fig. 84c); coxa 5 larger than anterior coxae,
coxa) 4) posteriorly excavate 7Ms 099525 6 a0 4s ees Gennes
Genera of Dexaminidae
Anatylus Bulycheva
Anatylus Bulycheva, 1955.
Type-species: A. pavlovskii Bulycheva, 1955 (original designation). Mandibular molar evanescent; lower lip bearing inner lobes, man-
dibular processes distinct, palp of maxilla 1 biarticulate, artrcle 2 long (Guernea) ; maxillipedal palp 4-articulate, inner plates well developed; coxa 4 not excavate posteriorly. Species: 1, boreal N. W. Pacific, littoral.
285-135 O - 69 - 14
MARINE GAMMARIDEAN AMPHIPODA 203
Dexamine Leach
Dexamine Leach, 1814b.—Stebbing, 1906. Amphithonotus Costa, 1851b.
Type-species: Cancer spinosus Montagu, 1813 (monotypy). See Sars, 1895.
Lower lip with poorly developed inner lobes, mandibular processes distinct; palp of maxilla 1 uniarticulate; maxillipedal palp 3-articulate, inner plates well developed; article 4 of pereopods shorter than articles 5-6 combined (Tritaeta). Species: 5, boreal N. E. Atlantic, Mediter- ranean, Ceylon, littoral.
Dexaminella Schellenberg Dexaminella Schellenberg, 1928b.
Type-species: D. aegyptiaca Schellenberg, 1928b (monotypy).
Lower lip with well-developed inner lobes, mandibular processes distinct; palp of maxilla 1 uniarticulate; maxillipedal palp 3-articulate, inner plates absent. Species: 1, Red Sea, littoral.
Dexaminoides Spandl Dexaminoides Spandl, 1923.
Type-species: D. orientalis Spandl, 1923 (monotypy). See J. L. Barnard, 1965.
Lower lip with well-developed inner lobes, mandibular processes distinct; palp of maxilla 1 uniarticulate; maxillipedal palp 4-articulate, article 4 claw-like (Dexamonica), inner plates vestigial, bud-like. Species: 1, Indo-Pacific, tropical, littoral.
Dexamonica J. L. Barnard Dexamonica J. L. Barnard, 1958c.
Type-species: D. reduncans J. L. Barnard, 1958c (original designation).
Lower lip with distinct, fused inner lobes, mandibular processes obsolescent; palp of maxilla 1 uniarticulate; maxillipedal palp 4- articulate, article 4 short, not claw-like (Dexaminoides), inner plates vestigial, bud-like. Species: 1, California, littoral.
Guernea Chevreux
Helleria Norman, 1868 (homonym, Isopoda). Guernea Chevreux, 1887 (new name).—Stebbing, 1906. Prinassus Hansen, 1887.
Type-species: Helleria coalita Norman, 1868 (monotypy). See Chevreux and Fage, 1925.
204
U.S. NATIONAL MUSEUM BULLETIN 271
B
{\
LOM) Zi al
TT. AR EN Th 7U Le rk Da \ HY “nN =
iN IN 4 \ N
\\\ Bi NN Xs 1 \ a) \ i \ a ey \\ V Th \ \)
a=
p Figure 84.—Dexaminidae: a, Dexamine spinosa (Montagu)(Sars, 1895, pl. 167). lip: b, Dexamine.
Upper Maxillipeds: c, Guernea coalita (Norman) (Chevreux and Fage, 1925); d, Dexamine; e, Polycheria antarctica (Stebbing, 1888). Lower lip: f, Dexamonica redun- cans J. L. Barnard (1958c); g, Syndexamine carinata Chilton (1914); 4, Dexaminella aegyptica Schellenberg (1928); 7, Dexamine. Mandible: 7, Dexamine. Maxilla 1: k, Guernea; 1, Polycheria; m, Dexamine. Maxilla 2: n, Dexamine. Gnathopods 1, 2: 0, 9, Dexamine.
MARINE GAMMARIDEAN AMPHIPODA 205
t
Figure 85.—Dexaminidae: Anatylus pavlovskii Bulycheva (1955), parts as follows: a, body without appendages; b,c, antennae 1, 2; d, upper lip; ¢, mandible; f, lower lip; g,h, maxillae 1, 2; 7, maxilliped; 7, uropod 3; k, telson; /,m, gnathopods 1, 2. Pereopod 4: n, Dexamine spinosa (Montagu) (Sars, 1895, pl. 167); 0, Tritaeta gibbosa (Bate) (Sars, 1895, pl. 168). Pereopod 3: p, Polycheria antarctica (Stebbing, 1888). Uropod 3: gq, Dexamine. Telson: 7, Guernea coalita (Norman) (Chevreux and Fage, 1925); 5, Dexamine.
Mandible: #, Guernea.
206 U.S. NATIONAL MUSEUM BULLETIN 271
Lower lip with well-developed inner lobes, mandibular processes absent; palp of maxilla 1 biarticulate, artecle 2 short (Anatylus) except in G. petalocera Ruffo; maxillipedal palp 4-articulate, inner plates vestigial, bud-like; coza 5 larger than anterior coxae (Anatylus). Species: 3, boreal-tropical in N. and E. hemispheres, littoral.
Paradexamine Stebbing Paradexamine Stebbing, 1899d; 1906.
Type-species: Dexamine pacifica Thomson, 1879 (original designa- tion). See Stephensen, 1927.
Lower lip with well-developed inner lobes, mandibular processes distinct; palp of maxilla 1 uniarticulate; maxillipedal palp 4-articulate, inner plates of moderate size. Species: 9, antarctic-antiboreal, littoral.
Polycheria Haswell Polycheria Haswell, 1880c.—Stebbing, 1906.
Type-species: Dexamine antarctica Stebbing, 1875 (present selec- tion, denoted by synonymy of Chilton, 1912). See Stebbing, 1888 (as Tritaeta antarctica and T. kerguelent).
Lower lip with well-developed inner lobes, mandibular processes absent; palp of maxilla 1 ?biarticulate (uniarticulate=auct. other than Stebbing); maxillipedal palp 4-articulate, inner plates well developed; pereopods cheliform. Species: 4, Antarctic to N. Pacific, littoral.
Syndexamine Chilton
Syndexamine Chilton, 1914.
Type-species: SS. carinata Chilton, 1914 (monotypy).
Lower lip with small but distinct inner lobes, mandibular processes absent; palp of maxilla 1 uniarticulate; maxillipedal palp 4-articulate, inner plates small. Species: 1, New Zealand, littoral.
Tritaeta Boeck
Lampra Boeck, 1871 (homonym, Lepidoptera). Tritaeta Boeck, 1876 (new name).—Stebbing, 1906.
Type-species: Atylus gibbosus Bate and Westwood, 1863 (mono- typy). See Sars, 1895.
Lower lip with tiny imner lobes, mandibular processes distinct; palp of maxilla 1 uniarticulate; maxillipedal palp 3-articulate, imner plates well developed; article 4 of pereopod 5 longer than articles 5-6 combined (Dexamine). Species: 2, EK. Atlantic, littoral.
MARINE GAMMARIDEAN AMPHIPODA 207
Dogielinotidae
FIGURE 86
Diacnosis.—Accessory flagellum absent; epistome with a pendant lobe; mandible lacking palp; palp of maxilla 1 vestigial; uropod 3 uniramous or lacking a ramus; telson broad, cleft shortly; antennae and pereopods densely spinose and setose (fossorial). See Talitroidea, Haustoriidae, Kurtidae, Phliantidae.
Description.—Accessory flagellum absent; body not carinate or toothed; rostrum small; coxae of medium length; mandible lacking palp, molar large, triturative; palp of maxilla 1 vestigial, 1-articulate; enathopods of medium size, subchelate; uropod 3 uniramous or ramus absent, ramus small, bearing apical setae; telson short, broad, shortly cleft; antennae and pereopods densely spinose as in phoxocephalids and haustoriids.
RELATIONSHIP.—This remarkable family combines characters of Talitroidea and Phoxocephalidae-Haustoriidae. Its two species have these characters of Talitroidea: no accessory flagellum; uniramous [or aramous] third uropod; the telson of a Hyale; no mandibular palp; and a vestigial first maxillary palp. Dogielinotids have these characters of the Phoxocephalidae-Haustoriidae: shape of head and rostrum; short, very spinose antennae; similar gnathopods and pereopods; long peduncular setae of the first and second uropods; and the strong tooth at the posteroventral corner of the third pleonal epimeron.
Metoediceros Schellenberg (1931), now assigned erroneously to Oedicerotidae probably should form the type of a new family with resemblance to Dogielinotidae (see Incertae Sedis). Metoediceros differs from dogielinotids in its circular, uncleft telson. Its epistome is unknown.
Dogielinotids are distinguished from Talitroidea by their phoxo- cephalid characters and vice-versa, and from the Haustoriidae by the uniramous uropod 3 and the absence of a mandibular palp.
The Kurtidae have coalesced urosomal segments.
The Prophliantidae and Phliantidae have evanescent mandibular molars.
Key to the Genera of Dogielinotidae
i ee Oropodisiwithtonewramusys ca5 45 yas estes ee ae see Dogielinotus Wropodis lacking, rami un mus eyy ices ate Pewter bea Neen § Haustorioides
208 U.S. NATIONAL MUSEUM BULLETIN 271 Genera of Dogielinotidae
Dogielinotus Gurjanova
Dogielinotus Gurjanova, 1953.
Type-species: Allorchestes moskvitina Dershavin, 1930 (original designation). Species: 1, boreal Pacific, littoral.
Ficure 86.—Dogielinotidae: Dogielinotus sp., California, original figures: a, lateral aspect; b, mandible; c, maxilla 1; d, maxilliped; ¢, telson; f, uropod 3; g, lower lip; h, anterior view of epistome pendant in front of upper lip; 7, maxilla 2.
MARINE GAMMARIDEAN AMPHIPODA 209
Haustorioides Oldevig Haustorioides Oldevig, 1958.
Type-species: H. munsterhjelmi Oldevig, 1958 (monotypy). Species: 1, boreal Pacific, littoral.
Eophliantidae
(Sheard, 1936a, elevated by Gurjanova, 1958] FicursEs 87, 88
Diacnosis.—Accessory flagellum absent; body cylindrical; all coxae short, generally not touching; mandible lacking palp, molar vestigial or absent; palp of maxilla 1 degraded or absent; gnathopods usually feeble; uropod 3 becoming degenerate; telson short, weakly to fully cleft. See Prophliantidae, Phliantidae, Talitroidea, Kuriidae, Colomastigidae.
Description.—Body vermiform, head globular, lacking rostrum, urosome tending to be reduced and its segments coalesced, uropod 3 becoming degenerate, with two, one or no rami; accessory flagellum absent; mandible lacking palp and molar obsolescent; palp of maxilla 1 small, 1-articulate or absent; remaining mouthparts basic; gnatho- pods feeble, simple or weakly subchelate or parachelate, article 3 of enathopods occasionally elongate; telson short, weakly to fully cleft; pleopods biramous or uniramous, requiring further study in the several genera.
Uropod 3 and its relationship to the urosome need careful study; it is necessary to distinguish clearly pleonite 6, the peduncle of uropod 3, and its ramus. Sometimes pleonite 6 is fused to 5 but a peduncle of uropod 3 may be confused with the segment if one mistakes a ramus for a peduncle.
ReLationsuip.—The Phliantidae differ from the Eophliantidae by their depressed or compressed bodies with long coxae.
The Talitroidea differ by their strongly triturative mandibular molars (except Najna) and compressed bodies.
The Colomastigidae have cylindrical bodies but the inner plates of the maxillipeds are very small, the telson is cleft, and the mandib- ular cutting edge is divided into large teeth.
Eophliantidae may resemble Corophiidae and Podoceridae because of their depressed, subcylindrical bodies but the latter two families have mandibular palps and molars.
210 U.S. NATIONAL MUSEUM BULLETIN 271
f g
Figure 87.—Eophliantidae: a, Wandelia crassipes Chevreux (1906c); b, Ceinina japonica Stephensen (Gurjanova, 1951). Upper lip: c, Wandelia. Head: d, Bircenna fulva Chilton (Nicholls, 1939) (arrow marks ventral flange). Mandible: e, Amphitholina cuniculus (Stebbing) (Chevreux and Fage, 1925); f, Wandelia; g, Wandelia japonensis (Nicholls, 1939). Lower lip: h, Bircenna ignea Nicholls (1939).
MARINE GAMMARIDEAN AMPHIPODA P11
Key to the Genera of Kophliantidae
1. Uropod 3 biramous (figs. 88l,0,q) 2 Uropod 3 not biramous (figs. 88m,n,p) . 4
2. Maxilla 1 lacking palp (fig. 88a) . Apes Senne Oiy eT REN ¢ 5 3 Maxilla 1 with small palp (fig. 88c) .. . Se frophiianticn
3. Uropod 3 with short, broad rami less than half as longs as peduncle (fig. 880), gnathopod 2 powerful (fig. 887). . . . . . Amphitholina Uropod 3 with slender rami as long as s pedunele (iw 88), gnathopod 2 feeble HeAS Sa) iiss ta) é . . . Biancolina
4. Pleonites 5-6 small emt Shatin. tas alae Nar LR a ania as ee OLN RN eM Pleonites 5-6 coalesced. . . . . . BV US HMO gd
5. Antennae half as long as head, avila, 1 lackine alo: mena lacking mn@lene 55) 6) 6 . . . . (theoretical position) Ceinina Antennae as tones as heel rts les 1 with minute palp, mandible bearing small molar (one member at least). . . . . . . =... . . Eophliantis*
6. Pleopods uniramous (fig. 88k) ........... . .. Cylindryllioides Pleopods biramous. . . aaica hey See MT t canaene east Mes SRSA
7. Pereonite 1 with a ential fiance Ge. 87d). ieee eee ee ea TRE ene Rereonitemilackineayventraliflamgel cis ais) ye) aS ee 8
8. Coxae tiny, serially noncontiguous, flagellum of antenna 2 biarticulate, mandibular palp absent... . .... .+. Ceinina Coxae small, serially contiguous, Abyscllhonn of POA 2 about 6-articulate, mandible with vestigal palp. . ........... =... Wandelia
1 Sheard (1936a) and Nicholls (1939) have different concepts of uropod 3 on Eophliantis.
Genera of Eophliantidae
Amphitholina Ruffo
Amphitholina Ruffo, 1953. Type-species: Amphithoe cuniculus Stebbing, 1874 (original des- ignation). See Chevreux and Fage, 1925 (as Biancolina cuniculus). Maxilla 1 lacking palp; pleonites 5-6 apparently separate; uropod 3 with two short broad rami, half as long as peduncle, gnathopod 2 powerful (Biancolina). Species: 1, Mediterranean, littoral.
Biancolina Della Valle Biancolina Della Valle, 1893.—Stebbing, 1906 (in part).
Type-species: B. algicola Della Valle, 1893 (monotypy).
Maxilla 1 lacking palp; pleonites 5-6 separate; uropod 3 with slender rami as long as peduncle, gnathopod 2 feeble (Amphitholina). Species: 2, Mediterranean, Australia, littoral.
212 U.S. NATIONAL MUSEUM BULLETIN 271
Bircenna Chilton Bircenna Chilton, 1884.—Stebbing, 1906.
Type-species: B. fulvuus Chilton, 1884 (monotypy). See Sheard, 1936a; Nicholls, 1939.
Maxilla 1 lacking palp; pleonites 5-6 coalesced; uropod 3 with a single vestigial ramus or none; pereonite 1 with a ventral flange. Species: 4, 5S. Australia and New Zealand, littoral.
r
a
Figure 88.—Eophliantidae: Maxilla 1: a, Wandelia crassipes Chevreux (1906c); b, Am- phitholina cuniculus (Stebbing) (Chevreux and Fage, 1925); c, Eophliantis tindale: Nicholls (1939). Maxilliped: d, Wandelia. Gnathopod 1: e, Wandelia; f, Amphitholina; g, Biancolina algicola Della Valle (1893). Gnathogod 2: h, Wandelia; 1, Amphitholina; j, Biancolina algicola. Pleopod: k, Cylindryllioides mawsont Nicholls (1938). Uropod 3: 1, Biancolina australis Nicholls (1939); m, Eophliantis; n, Bircenna nichollst Sheard (Nicholls, 1939). Telson, uropod 3 and dorsum of urosome: 0, Amphitholina; p, Wan- delia; q, Biancolina algicola. Urosome, lateral: r, Eophiiantts.
MARINE GAMMARIDEAN AMPHIPODA 213
Ceinina Stephensen Ceinina Stephensen, 1933.
Type-species: C. japonica Stephensen, 1933 (monotypy). See Gurjanova, 1951; Nicholls, 1939 (as Wandelia gaponensis).
Maxilla 1 lacking palp; pleonites apparently coalesced; uropod 3 with one ramus; antennae half as long as head, mandible lacking molar (Hophliantis) ; coxae tiny, serially noncontiguous (Wandelia). Species: 1, Japan, littoral.
Cylindryllioides Nicholls
Cylindryllioides Nicholls, 1938.
Type-species: C. mawsoni Nicholls, 1938 (original designation). Maxilla 1 lacking palp; pleonites 5-6 coalesced; uropod 3 lacking rami; pleopods uniramous (combining character). Species: 1, antarctic, littoral. Eophliantis Sheard
Eophliantis Sheard, 1936a.
Type-species: . tindalei Sheard, 1936a (original designation).
Maxilla 1 with small palp; pleonites 5-6 separate; uropod 3 with two blunt rami (Sheard), no rami (Nicholls, 1939) ; antennae as long as head, mandible bearing small molar (Ceinina). Species: 1, Tasmania, littoral.
Wandelia Chevreux Wandelia Chevreux, 1906a; 1906b.
Type-species: W. crassipes Chevreux, 1906a (original designation) ; 1906b.
Maxilla 1 lacking palp; pleonites 5-6 coalesced; uropod 3 lacking rami; coxae small, serially contiguous (Ceinina). Species: ?2, antarctic to N. Pacific, littoral.
Eusiridae including Pontogeneiidae]
Fiaures 89-93
Diacnosts.—Accessory flagellum either a single styliform article or two short articles, a scale or absent; telson cleft. See Calliopiudae, Pleustidae, Liljeborgiidae, Gammaridae, Vitjazianidae, Astyridae, Lepechinellidae, Paramphithoidae.
Descriptron.—Accessory flagellum absent, or composed of a scale or one long article or two short articles; rostrum present or obsolete; upper lip not incised (bilobed in one genus); mandibles bearing 3-
214 U.S. NATIONAL MUSEUM BULLETIN 271
—
SESE TASS BEISDERIBPAIIES N
a oe
Ficure 89.—Eusiridae: a, Rhachotropis aculeata (Lepechin) (Sars, 1895, pl. 149); b, Eustrus cuspidatus Krdéyer (Sars, 1895, pl. 146). Kinds of accessory flagella: c, Liouvillea oculata Chevreux (1912b); d, Rhachotropis; e, fused process, Prostebbingia gracilis (Chevreux, 1912b); f, Pontogeneoides abyssi Nicholls (1938); g, no accessory flagellum on Pontogeneia inermis (Krdyer) (Sars, 1895, pl. 159); h, Djerboa furcipes Chevreux (1906c); 2, Eusirus propinquus Sars (1895, pl. 147). 7, Calceolus.
MARINE GAMMARIDEAN AMPHIPODA 215
articulate palp, molar triturative (except in a few genera); lower lip with or without inner lobes; other mouthparts basic, except one genus with shortened palp of maxilla 1; gnathopods either powerful and subchelate, occasionally small, or feeble, occasionally nearly simple (Harpinioidella); coxae of medium length or short; uropod 3 with elongate, lanceolate rami; telson often elongate, much longer than peduncle of uropod 3, or increasingly shortened, cleft deeply or slightly cleft only apically, occasionally appearing to be emarginate; in Eusirogenes telson shorter than peduncle of uropod 3.
Revationsuip.—The Astyridae differ from the Eusiridae by the nontriturative mandibular molar and the characteristic shape of the lower lip. Some astyrids have a massive head and all have a very short telson.
The Vitjazianidae are distinguished by their distinctly simple first enathopods and the elongate article 1 of the flagellum on antenna 1 which equals the peduncle in length. Pleustidae have their character- istic lower lips.
Since Calliopiidae appear to be Eusiridae-Pontogeneiidae with fused telsonic lobes and some Eusiridae have nearly entire telsons, it must be pointed out that Sancho in the Calliopidae, with entire telson and with 1-articulate accessory flagellum as long as article 1 of the primary flagellum has gnathopod 2 similar to the “‘Kusirus” gnathopods and might be placed more logically in Eusiridae than in Calliopiidae. See the latter family for further discussion.
Most synopiids have a multiarticulate accessory flagellum but a few have an accessory flagellum similar to that of eusirids. The massive, galeate head of most synopiids is characteristic, especially because of its strongly deflexed rostrum. Synopiid gnathopods are feeble whereas those of eusirids are usually powerful.
Rhachotropis aculeata bears resemblance to Paramphithoidae in its slightly acute first four coxae but differs by its powerful gnathopods. Gnathopods of Paramphithoidae are feeble and one or more of the first four coxae are acuminate.
All Liljeborgiidae have a nontriturative mandibluar molar, but eusirid resemblance to this family is close because some liljeborgiids have a 2-articulate accessory flagellum and a few eusirid genera have a poorly triturative mandibular molar.
Most Gammaridae differ from Eusiridae by their multiarticulate accessory flagella but a number of genera have 1-2-articulate accessory flagella (see couplets 6-17 in Gammaridae key). Eusiridae differ from those genera of Gammaridae by their elongate telsons and the slight elongation of the metasome. Nearly half of those gammarid genera have short inner rami on uropod 3, and most of the remainder have either foliaceous rami on uropod 3, and elongate article 2 of the
216 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 90.—Eusiridae: a, Eurymera monticulosa Pfeffer (Chevreux, 1906c). Upper lips: b, Eusirus propinquus Sars (1895, pl. 147); c, Pontogeneia inermis (Krgyer) (Sars, 1895, pl. 159). Mandible: d, Eusirus; e, Pontogeneia; f, Liouvillea oculata Chevreux (1912b). Lower lip: g, Pontogeneia; h, Eusirus; 1, Bovallia gigantea Pfeffer (Chevreux, 1906c); 7, Pontogeneoides abyssi Nicholls (1938); k, Eurymera; 1, Meteusiroides keyensis Pirlot (1934).
MARINE GAMMARIDEAN AMPHIPODA Dilvy,
mandibular palp or a simple gnathopod 1. Only Elasmopus, Liagocera- docus, and Anelasmopus do not fit these statements.
Though originally assigned to Amphilochidae, Pseudamphilochus Schellenberg falls to the Eusiridae, especially through the illustrated family key. That genus has the head of an amphilochid or pleustid, enathopods of an amphilochid, a lower lip faintly resembling that of Pleustidae, but coxa 1 is not reduced as it is in Amphilochidae and the cleft telson is unlike either pleustids or amphilochids. Pseudamphilo- chus has been given a special place in Gammaridea Family Key G (p. 114) and is mentioned also under the Amphilochidae.
Lepechinellids have acuminate coxae and apparently always have pleonites 5-6 coalesced.
Key to the Genera of Eusiridae !
1. Article 6 of gnathopods attached to the strongly produced or extended apex of article 5 (‘‘Eusirus gnathopods’’), article 6 as broad as long (figs.
COP TY Ost) Vi ance aber ae TSA Me eae, Article 6 of Pahonods with nerraal deehment. Bericle 6 longer than broad (figs. 92a,c,d,i,m) .. . 5
2. Articles 4 and 6 of gnathopods saucehiars e. 92h), ariniennaes sla fcalecol (fig. 897) .. . . . Eusiropsis eis 4 and 6 aie pontnepadst not Vouching| pCparated ie a lobe of article
5 (figs. 92b,7), antennae lacking calceoli. . . 3
3. Nettie 6 of gnathopod 1 larger than article 6°of pannaped 2, alto amielle 3 of mandible shorter or scarcely shorter than article 2 (fig. 90f) . Eusirogenes Article 6 of gnathopod 1 equal to or smaller than article 6 of gnathopod 2, palp article 3 of mandible equal to or longer than article 2 (fig.90d). . 4
4. Posterior border of article 5 of gnathopods forming a very narrow projecting lobe, much narrower than length of tapered distal end of article 5 (fig.
Q2b) Ree eB ee SUSInUS Posterior border of price 5 of Pee honoda foamed of a » NONE shallow setose lobe, its length greater than the tapered distal end of article 5 (fig. 92e).
Pareusirogenes 5. Palp of maxilla 1 short, article 1 longer than 2 (fig. 91f). . . . . Eusirella Palp of maxilla 1 long, article 2 longer than 1 (fig. 91g)... . 6 6. Articles 4, 5, 6 of pereopods 3-5 each 1.5 times as long as omits 2 (if lees broken proceed to couplet 9) (fig. 89a)... . dl Articles 4, 5, 6 of pereopods 3-5 each as long as aheie poco preteen Gt SOO) ee ee Ea SSA NMRA ENE US IanC Rr Si li OU ae OR an SU se nee) 7. Gnathopods not pubenerire RO en ie aN pence mh aga ey ALCL GO Gnathopods subchelate .. . sea CES
8. Body carinate (often weakly), coxa 4 Dodie’ Aarawnendl Ge 89a). Rhachotropis Body not carinate, coxa 1 not producedforward. .... . . . Cleonardo 9. Epistome with large anterior process (seen from lateral view) . . . . . 102
1 Pseudopontogencia Oldevig (1959) and Pontogenevides Nicholls (1938) are not sufficiently described for inclusion in the following key.
2 See also small process of Afetewsiroides and broadly flabelliform (dorsal-ventral) but shallow process of Chosroes and footnote 3 to couplet 31.
218
13.
14.
15.
16.
Mas
28.
U.S. NATIONAL MUSEUM BULLETIN 271
Hpistome lacking largelanterion processie. vei) co bike eu ened Postantennal cephalic angle acutely produced. . ... .. . .. Atyloella Postantennal cephalic angle presumably rounded. . . . . . Liouvillea Both articles 5 and 6 of gnathopod 2 long and sublinear (fig. 92k) Se Miele Either article 5 or 6 of gnathopod 2notlongandsublinear. ...... 14 Convex side of dactyli on pereopods 3-5 with accessory spine or spines (fie OSC) Maw ig : Reese Nasa... Ue} Dactyli of soreenodls 3 cl facie Dontenion EOSIN sine . . Schraderia Pereopods 1 and 2 with dactylar spine, article 2 of first antennal peduncle shorter than article]... .. . . . . . Djerboa Pereopods 1 and 2 lacking dectlen g spine, _ amelie 2 of first antennal peduncle longer than article]... . . . . Bathyschraderia Gnathopods not subchelate (ike Pleoeneneoniies tt in Calhionide) (figs. 92d,p). 15
Gnathopods distinctly subchelate. . . . «ee hG Probably each of articles 4-6 of syarsonronls au 5 nor fonees neva article 2. Harpinioidella
Each of articles 4-6 of pereopods 3-5 longer than article 2. . . Harcledo Accessory flagellum distinctly articulate, usually cylindrical, often as long as article 1 of primary flagellum (figs. 89h,z) . . . . BER al) esi dy ALT Accessory flagellum usually absent, when present foomed on a fused scale or process, not articulate (figs. 89e,f,g) .... . ; . 26
Inner plate of maxilla 1 with more than four ne. seine penal lining medial edge and apex (figs. 91d,e). Paramoera (and Atylopsis in Calliopiidae)
Inner plate of maxilla 1 with four or fewer terminal setae (figs. 91c,g). . 18 Article 6 of gnathopod 2 twice as long as article5. ......2.2.2. «19 Article 6 of gnathopod 2 subequal to article 5inlength ....... 24 Article 6 of pereopods 3-5 subequal to or shorter than article 2... . 20 Article 6 of pereopods 3-5 longer than article2........... =. 23 Accessory flagellum as broad as long (fig. 89f) .......2.2.=.. 2i1 Accessory flagellum long, rectangular (fig. 89h) ..... . 22 Melsonedeephy, cleft) (igs Ol) eerie ne dooce (nie wee nnn ees air Necederioenn
Telson apically notched (fig. 91m). Paramoera (and Atylopsis in Calliopiidae)
Upper lip bilobed .. . wi nachauny visti an crip a A Unocal NeMtOR Ag Ses ee OTE CO: Upper lip rounded outa y eA ie he Hoe usss ae) UeStroides Body ecarinate, accessory flagellum 2 darticulate . . . . . . Rhachotropis Body smooth, accessory flagellum l-articulate . . ... . . . Cleonardo Body. canimate Wan le Woah clas voles) Soleo seul eC LEONG ODSIS Body smooth. . . . BINA Han eee cane Meee Gy ys) PAD) Article 6 of Paainonods narrower 3 dhvam fifth . ..... . . Pseudomoera Article 6 of gnathopods as broad as fifth . . . . . Accedomoera (in part) Anteroventral cephalic angle with large tooth, coxa 1 produced forward to anterior edge of eye (fig. 93b) ... . . . . . . Rozinante Anteroventral cephalic angle with small ‘root or none, coxa 1 not pro- ducediats once WU es ane oiten POU Inner plate of eile 1 it tour or fear avast sone Mieaiicrb eo" cs Ze: Inner plate of maxilla 1 with more than four setae and some setae lining medial edge (fig. 9le) . . . BENET GE OY sig EAE oR
Articles 4, 5, and 6 of nrorasoods 3- 5 cera 15 ines as long as article 2 Harcledo
Articles 4, 5, and 6 of pereopods 3-5 not longer than article2. .. . . 29
MARINE GAMMARIDEAN AMPHIPODA P19
29. Gnathopods simple (like Harpinioides in Calliopiidae) (figs. 92d,p).
Harpinioidella
Gnathopods subchelate .. . so BW
30. Outer lobes of lower lip broadly conamtcd: inner Nlobes farce (Ge 901), telson extending three fourths along rami of uropod 3... . . . Meteusiroides
Outer lobes of lower lip with small medial gap, inner lobes obsolescent or absent (fig. 90g), telson extending one third or less along rami of uropod
31. Inner lobes of lower lip small but distinct, inner plate of maxilla 2 with poiae lining medial edge, upper lip rounded or truncate below, gnathopod astavalliliman ie a sie ... . . Pontogeneia 3
Inner lobes of lower iD Apeent inner yale on jedi & with one medioter- minal seta, upper lip with small incision on ventral margin, gnathopod 2
very large (like Rhachotropis) Ane . . . . . Dautzenbergia
32. Dactyls of pereopods 3 and 4 as long as erncle 6, pereopods very setose and oedicerotid-like (fig. 93g) ... . .. . . . Zaramilla Dactyls of pereopods 3 and 4 very shorts nereopode anon setose, slender . 33
SORE loweripnwivasimmer lObesiey kl ke he Rn eA ela nie Wowerliplackingimner lobes) 394 4%) = 2 =. 2) ee 3 enone
34. Coxa 4 not excavate posteriorly. ............ prostebbineia Coxamtvexcavate POSteLlOnlyamea mae iis (chee is til, alien BT vy re MGURE oD
Se bodyaumibbonate (ie 90G)ea.h feswer es eee oc on. ae. Burnymera Body not umbonate. . . . Pera Seles 0)
36. Inner lobes of lower lip very ene inner alate a soneallle 1 oath setae mostly eT MMII Aall ess ey eens 5 .. . . . Pontogeneia
Inner lobes of lower Ep tere: inner plate of oceaill 1 with setae lining full medialedge .... ... . . Pontogeneiella
37. Coxa 4 deeply ae agosto, upper 2 iti rounded below, telson cleft halfway, maxillipedal palp artiele 4 shorter than article3 . . . . Bovallia
Coxa 4 scarcely excavate posteriorly, upper lip slightly incised, telson cleft one fifth, maxillipedal palp article 4 longer than article 3. (theoretical possibility) Dautzenbergia
3 Also see Chosroes decoratus K. H. Barnard (1932), which is related to the type, C. incisus (Calliopiidae), but which has a cleft telson and so inadvertently keys to Pontogeneia.
Genera of Eusiridae
Nore: ‘“‘eustrip” gnathopods have article 6 as broad as long and attached to the produced apex of article 5.
Accedomoera J. L. Barnard Accedomoera J. L. Barnard, 1964c.
Type-species: Pontogeneia tricuspidata Gurjanova, 1938 (original designation).
Accessory flagellum articulate, short; lower lip with small inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; enathopods not eusirid, subchelate, article 5 not lobate, shorter than 6, 6 as broad as 5 (Pseudomoera); pereopods 3-5 with each of articles 4-6 not distinctly longer than article 2. Species: 3, subantarctic, boreal, N.W. Pacific, littoral.
285-135 O - 69 - 15
NATIONAL MUSEUM BULLETIN 271
220 U.S.
a (| LY a =, ea *y R 7h, F f f 4 i \ fill 7 of Ww # qf Ae 1i\ D MEY Z\\’ fh ‘ _—, —A
{ ff : (AU = Cal) Re
» i oA : \ \ : »
= SSI L/ ome ee
BZ
Y (\ \\ ih i
k m
Maxilla 2: h, Eusirus; 1,
Ficure 91.—Eusiridae: a, Pontogeneia inermis (Krdyer) (Sars, 1895, pl. 159); b, Cleonardo appendiculatus (Sars, 1885). Maxilla 1: c, Eusirus propinquus Sars (Sars, 1895, pl. 147); d, Eurymera monticulosa Pfeffer (Chevreux, 1906c); ¢, Pontogeneia; f, Eusirella elegans Telson: 1, Pontogeneia; m, Ponto-
Chevreux (1908b); g, Pseudomoera gabrieli (Sayce, 1901).
Maxillipeds: 7, Pontogeneia; k, Eusirus. geneoides abyssi Nicholls (1938); n, Eusirus minutus Sars (1895, pl. 147); 0, Eusirus
Pontogeneia. propinquus; p, Rozinante fragilis Goés (Gurjanova, 1951); g, Liouvillea oculata Chevreux
(1912b).
MARINE GAMMARIDEAN AMPHIPODA 221
n
Ficure 92.—Eusiridae: Gnathopod 1: a, Bovallia gigantea Pfeffer (Chevreux 1906c); b, Eusirus propinquus Sars (1895, pl. 147); c, Djerboa furcipes Chevreux (1906c); d, Harcledo plumipes (Birstein and Vinogradov, 1955); e, Pareusirogenes carinatus Birstein and Vinogradov (1955); f, Rhachotropis aculeata (Lepechin) (Sars, 1895, pl. 149); g, Pontogeneia inermis (Kroyer) (Sars, 1895, pl. 159); h, Eusiropsis riisei Stebbing (1897). Gnathopod 2: 1, Bovallia; j, Eusirus; k, Djerboa; 1, Rozinante fragilis (Goés) (Gurjanova, 1951); m, Rhachotropis; n, Pontogeneoides abyssi Nicholls (1938); 0, Pontogeneia; p, Harcledo.
222 U.S. NATIONAL MUSEUM BULLETIN 271
Atyloella Schellenberg Atyloella Schellenberg, 1929a.
Type-species: Atylopsis magellanicus Stebbing, 1888 (monotypy).
Accessory flagellum articulate, short; lower lip with inner lobes; inner plate of maxilla 1 with four or five terminal and medial setae; gnathopods not eusirid, subchelate, article 5 not lobate, shorter than 6; pereopods 3-5 with each of articles 4-6 not longer than article 2; postantennal cephalic angle acutely produced (Laouvillea) ; epistome unth anterior process.* Species: 3, antarctic, littoral (to 441 m).
*See footnote with Meteusiroides (p. 227).
Bathyschraderia Dahl Bathyschraderia Dahl, 1959.
Type-species: B. magnifica Dahl, 1959 (monotypy).
Accessory flagellum very short but articulate; lower lip lacking inner lobes; inner plate of maxilla 1 densely furnished with terminal and medial setae; gnathopods not eusirid, scarcely subchelate, both articles 5 and 6 of both gnathopods elongate; [pereopods 3-5 not described for lengths of articles]; pereopods 3-5 with a row of spines on convex side of dactyli, not present on pereopods 1-2 (combining character and Djerboa). Species: 1, Kermadec Trench, hadal.
Bovallia Pfeffer Bovallia Pfeffer, 1888.—Stebbing, 1906.
Type-species: B. gigantea Pfeffer, 1888 (monotypy). See Chev- reux, 1906b.
Accessory flagellum absent; lower lip lacking inner lobes; inner plate of maxilla 1 with numerous terminal setae and setae partially — lining medial edge; gnathopods not eusirid, subchelate, article 5 strongly lobate, shorter than article 6; pereopods 3-5 with articles 4-6 each not longer than article 2; coxa 4 deeply excavate posteriorly, upper lip rounded ventrally; telson cleft halfway; mazillipedal palp article 4 shorter than article 3 (Dautzenbergia). Species: 1, antarctic, littoral.
Cleonardo Stebbing Cleonardo Stebbing, 1888; 1906.
Type-species: C. longipes Stebbing, 1888 (monotypy).
Accessory flagellum articulate, short; lower lip with inner lobes; inner plate of maxilla 1 with less than four terminal setae; gnathopods not eusirid, scarcely subchelate, article 5 strongly lobate, shorter than
MARINE GAMMARIDEAN AMPHIPODA Dae
article 6; pereopods 3-5 with articles 4-6 each much longer than article 2; body dorsally smooth (Rhachotropis). Species: 7, N. Hemi- sphere largely, bathyal-abyssal, pelagic or ?demersal.
Cleonardopsis K. H. Barnard
Cleonardopsis K. H. Barnard, 1916. Amathillopleustes Pirlot, 1934.
Type-species: C. carinata K. H. Barnard, 1916 (monotypy).
Accessory flagellum articulate and long; lower lip with broad semifused inner lobes; maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, subchelate, article 5 lobate, as long as article 6; pereopods 3-5 with articles 4-6 each not longer than article 2; body carinate (Pseudomoera). Possibly belongs in the Pleustidae because of the lower lip. Species: 1, probably cosmopolitan, bathypelagic.
Dautzenbergia Chevreux Dautzenbergia Chevreux, 1900.
Type-species: Amphithopsis grandimana _ Chevreux, 1887 (monotypy).
Accessory flagellum absent; lower lip lacking inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, subchelate, article 5 of gnathopod 2 with slender pos- terior lobe, article 6 in male very large, about 5 times as long as article 5; pereopods 3-5 with articles 4-6 each not longer than article 2; coxa 4 scarcely excavate posteriorly, upper lip slightly incised, telson cleft one fifth, maxillipedal palp article 4 longer than article 3 (Bovallia). Species: 3, subarctic, N. Atlantic to ?Indian Ocean, bathyal to abyssal.
Djerboa Chevreux Djerboa Chevreux, 1906b.
Type-species: D. furcipes Chevreux, 1906b (monotypy).
Accessory flagellum articulate, long; lower lip lacking inner lobes; inner plate of maxilla 1 densely lined with setae on medial edge; genathopods not eusirid, subchelate, article 5 not lobate, on gnathopod 2 especially, both articles 5 and 6 greatly elongated and sublinear; pereopods 3-5 with articles 4-6 each not longer than article 2; pereopods 1-5 with accessory spine on conver side of dactyli (combining character and Bathyschraderia). Species: 1, antarctic, littoral.
Eurymera Pfeffer Eurymera Pfeffer, 1888.—Stebbing, 1906.
Type-species: E. monticulosa Pfeffer, 1888 (monotypy). See Chevreux, 1906b.
DIA U.S. NATIONAL MUSEUM BULLETIN 271
Accessory flagellum absent; lower lip with inner lobes; inner plate of maxilla 1 densely lined with setae on medial edge; gnathopods not eusirid, subchelate, article 4 not lobate, slightly shorter than 6; pereopods 3-5 with articles 4-6 each not longer than article 2; body umbonate. Species: 1, antarctic, littoral.
Ficure 93.—Eusiridae: a, Djerboa furcipes Chevreux (1906c); b, Rozinante fragilis Goés (Shoemaker, 1930). Uropod 3: c, Eusirus propinquus Sars (1895, pl. 147); d, Eusirus
minutus Sars (1895, pl. 147). End of pereopod 3: e, Djerboa. Right coxa 4: f, Prosteb- bingia gracilis (Chevreux, 1906c). Pereopod 5: g, Zaramulla kergueleni Stebbing (1888).
MARINE GAMMARIDEAN AMPHIPODA 225
Eusirella Chevreux
Eusirella Chevreux, 1908b.
Type-species: LE. elegans Chevreux, 1908b (original designation).
Accessory flagellum absent; mandibular molar not triturative; [lower lip unknown]; palp of mazilla 1 short, article 1 much longer than article 2; gnathopods not eusirid, scarcely subchelate but article 6 not fully linear, article 5 not or scarcely lobate, shorter than article 6; pereopods 1 and 2 with article 6 bearing long plumose posterior setae; pereopods 3-5 with articles 4-6 each longer than article 2. Species: 4, cosmopolitan, bathypelagic.
Eusirogenes Stebbing Eustrogenes Stebbing, 1904.
Type-species: £. dolichocarpus Stebbing, 1904 (monotypy).
Accessory flagellum articulate, very long; lower lip with inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods eusirid, subchelate, articles 4-6 each not longer than article 2; palp article 3 of mandible shorter than article 2 (Eusirus, Pareusirogenes); antennae lacking calceoli (Eusiropsis); article 6 of gnathopod 1 larger than that of gnathopod 2. Species: 5, cosmopolitan (N. Hemisphere), bathypelagic.
Eusiroides Stebbing Eusiroides Stebbing, 1888; 1906.
Type-species: Atylus monoculoides Haswell, 1880c (indicated by Della Valle, 1893, through erroneous synonymy and thus by synonymy of Stebbing, 1906, but firmly selected by Chevreux and Fage, 1925).
Accessory flagellum articulate, of medium size; lower lip with small inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, subchelate, article 5 lobate, shorter than article 6, palms armed with very stout spines; pereopods 3-5 with articles 4-6 each not longer than article 2. Species: ca. 6, ?cosmo- politan, littoral to ?bathyal.
Eusiropsis Stebbing Eustropsis Stebbing, 1897; 1906.
Type-species: LE. riisei Stebbing, 1897 (monotypy).
Accessory flagellum articulate, minute; mandibular molar not trit- urative; lower lip with inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods eusirid, articles 4-6 touching; pereopods 3-5 with articles 4-6 each much longer than article 2; antennae with calceoli (Eusirus, Eusirogenes, Pareusirogenes). Species: 1,S. Hemisphere (?cosmopolitan), bathypelagic.
226 U.S. NATIONAL MUSEUM BULLETIN 271
Eusirus Kréyer Eusirus Kr¢yer, 1845.—Stebbing, 1906.
Type-species: EE. cuspidatus Kroyer, 1845 (monotypy). See Sars, 1895.
Accessory flagellum articulate, of medium size; lower lip with inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods eusirid, articles 4-6 not touching; pereopods 3-5 with articles 4-6 each not longer than article 2; antennal calceoh absent (Eusiropsis); posterior border of gnathopod article 5 forming a projecting lobe much narrower than length of tapered distal end of article 5 (Pareusirogenes). Species: 17, bipolar and tropical submergent, primarily bathyal to abyssal.
Harcledo J. L. Barnard Harcledo J. L. Barnard, 1964c.
Type-species: Meteusiroides plumipes Birstein and Vinogradov, 1955 (original designation). Birstein and Vinogradov (1964) synony- mized M. plumipes with M. curvidactyla (Pirlot, 1934).
Accessory flagellum presumably absent; lower lip unknown, but presumably with inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, not distinctly sub- chelate, article 6 tapering and with article 7 folded back on its setose posterior edge, article 5 slightly lobate and shorter than 6; pereopods 3-5 with articles 4-6 each much longer than article 2. Species: 1, N. W. Pacific, ?3500 m (has eyes), possibly epipelagic.
Harpinioidella Schellenberg Harpinioidella Schellenberg, 1926a.
Type-species: H. fissicauda Schellenberg, 1926a (monotypy).
[Accessory flagellum unknown]; lower lip with inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, simple, article 6 long, bent distally, article 5 much shorter than 6, apparently lobate; [pereopods 3-5 unknown]. Species: 1, antarctic to S. Atlantic, bathyal to ?abyssal (4,898 m), possibly bathypelagic.
Liouvillea Chevreux Liouvillea Chevreux, 1911c; 1912a; 1912b.
Type-species: L. oculata Chevreux, 1912a (designated by Chevreux) ; 1912b.
Accessory flagellum articulate, short; lower lip with inner lobes; inner plate of maxilla 1 densely lined with setae on medial edge;
MARINE GAMMARIDEAN AMPHIPODA 237
gnathopods not eusirid, subchelate, article 5 nearly as long as 6, not lobate; pereopods 3-5 with articles 4-6 each not longer than article 2: postantennal cephalic angle presumably rounded (Atyloella); rostrum reaching end of article 1 of antenna 1 (Paramoera). Species: 1, antarctic, littoral.
Meteusiroides Pirlot
Meteusiroides Pirlot, 1934.
Type-species: MM. keyensis Pirlot, 1934 (original designation).
Accessory flagellum absent; lower lip with large inner lobes, outer lobes very widely spread (Pontogeneia)*; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, subchelate, article 5 lobate, shorter than article 6; pereopods 3-5 with articles 4-6 each not longer than article 2. Species: 2, Indonesia, N. Atlantic, bathyal to abyssal.
*Apparently inner lobes shown by Pirlot (1934), are horizontal basal processes unnaturally flattened and outer lobes therefore excessively spread; small inner lobes present as in Pontogeneia; information from antarctic individual of Mete- usiroides (possible new species in colls. of El Tanin expeds.). Meteusiroides has telson extending 3/4 along rami of uropod 3, whereas telson of Pontcgeneza extends only 1/3 along rami of uropod 3. Also Meteusiriodes has small adze-shaped process on epistome and might be confused with Atyloella.
Paramoera Miers
Paramoera Miers, 1875.—Stebbing, 1906 (in part, not Atylozdes Stebbing, 1888).— Schellenberg, 1929a.—K. H. Barnard, 1932.
Stebbingza Pfeffer, 1888.
Aucklandia Walker, 1908.
Type-species: Amphithoe fissicauda Dana, 1852a (monotypy and subsequent synonymy). See Schellenberg, 1931; Atyloides australis and A. assimilis Stebbing, 1888.
Accessory flagellum articulate, of medium length; lower lip with inner lobes indistinct or absent; inner plate of maxilla 1 densely setose along medial edge; gnathopods not eusirid, subchelate, article 5 not lobate, shorter than article 6; pereopods 3-5 with articles 4-6 each not longer than article 2; rostrum inconspicuous (Liouvillea). Species: 28, bipolar, coldwater, littoral.
Pareusirogenes Birstein and Vinogradov Pareusirogenes Birstein and Vinogradov, 1955. ~Type-species: P. carinatus Birstein and Vinogradov, 1955 (original designation).
Accessory flagellum absent; lower lip with inner lobes; inner plate of maxilla 1 with four or fewer terminal setae, hairy along medial edge;
228 U.S. NATIONAL MUSEUM BULLETIN 271
enathopods eusirid, articles 4-6 not touching; pereopods 3-5 with articles 4-6 each much longer than article 2; antennal calceoli absent (Eusiropsis); posterior border of gnathopodal article 5 formed of a broad, setose lobe, its width greater than tapered distal end of article 5 (Eusirus). Species: 1, N.W. Pacific, abyssopelagic.
Pontogeneia Boeck Pontogeneia Boeck, 1871.—Stebbing, 1906.
Type-species: Amphithoe inermis Krgyer, 1838 (monotypy). See Sars, 1895.
Accessory flagellum absent; lower lp with small inner lobes; inner plate of maxilla 1 with four or more terminal or subterminal setae; gnathopods not eusirid, article 5 not lobate, sightly longer than article 6; pereopods 3-5 with articles 4-6 each not longer than article 2, article 4 slender (Zaramilla); outer lobes of lower lip not broadly separated (Meteusiroides). Species: 25, cosmopolitan, littoral.
Pontogeneiella Schellenberg Pontogeneiella Schellenberg, 1929a.
Type-species: Atyloides brevicornis Chevreux, 1906a, 1906b (pres- ent selection).
Accessory flagellum absent; lower lip with inner lobes; inner plate of maxilla 1 densely lined with setae on medial edge; gnathopods not eusirid, subchelate, article 5 not lobate, nearly as long as article 6; pereopods 3-5 with articles 4-6 each not longer than article 2, article 4 one third as wide as article 2 (Zaramilla); antennae subequal, coxa 4 strongly excavate posteriorly (Prostebbingia). Species: 2, antarctic, littoral.
Pontogeneoides Nicholls Pontogeneoides Nicholls, 1938.
Type-species: P. abyssi Nicholls, 1938 (original designation).
Accessory flagellum articulate, short; lower lip with small inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, subchelate, article 5 lobate on gnathopod 2, shorter than article 6, pereopods unknown; telson minutely notched. Species: 2, antarctic, bathyal (480—-1,590 m).
Prostebbingia Schellenberg
Prostebbingia Schellenberg, 1926a.
Type-species: Stebbingia gracilis Chevreux, 1912a, 1912b (present selection).
MARINE GAMMARIDEAN AMPHIPODA 929
Accessory flagellum absent; lower lip with inner lobes; inner plate of maxilla 1 densely lined with setae on medial edge; gnathopods not eusirid, subchelate, article 5 not lobate, slightly shorter than article 6; pereopods 3-5 with articles 4-6 each not longer than article 2; antenna 1 longer than antenna 2, coxa 4 scarcely excavate posteriorly (combining character). Species: 2, antarctic, littoral (to 400 m).
Pseudomoera Schellenberg
Pseudomoera Schellenberg, 1929a.
Type-species: Atyloides gabrieli Sayce, 1901 (monotypy).
Accessory flagellum articulate, of medium length; lower lip with inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, subchelate, article 5 lobate, equal to article 6 in length, 5 slightly broader than 6 (Accedomoera); pereopods 3-5 with articles 4-6 each not longer than article 2; body smooth (Cleo- nardopsis). Species: 1, Australia, freshwater.
Pseudopontogeneia Oldevig Pseudopontogeneia Oldevig, 1959.
Type-species: P. intermedia Oldevig, 1959 (monotypy).
[Accessory flagellum and lower lip not described]; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods [not distinctly described]; article 6 longer than 5; pereopods 3-5 with articles 4-6 each not longer than article 2. Species: 1, Bering Island, littoral.
Rhachotropis Smith
Tritropis Boeck, 1871 (homonym, Reptilia). Rhachotropis Smith, 1883 (new name).—Stebbing, 1906.—Shoemaker, 1930. Gracilipes Holmes, 1908.
Type-species: Oniscus aculeatus Lepechin, 1780 (monotypy). See Sars, 1895.
Accessory flagellum 2-articulate, short; lower lip with small inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, subchelate, article 5 lobate, shorter than article 6; pereopods 3-5 with articles 4-6 each longer than article 2; body carinate or toothed (Cleonardo). Species: 30, bipolar, tropical submergents, primarily bathyal-abyssal.
Ronco J. L. Barnard Ronco J. L. Barnard, 1965. Type-species: R. sosa J. L. Barnard, 1965 (original designation).
230 U.S. NATIONAL MUSEUM BULLETIN 271
Accessory flagellum articulate, of medium length; upper lip bilobate; lower lip with inner lobes; inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, subchelate, article 5 lobate, shorter than article 6; pereopods 3-5 with articles 4-6 each not longer than article 2. Species: 1, Micronesia, littoral.
Rozinante Stebbing Rozinante Stebbing, 1894; 1906.
Type-species: Paramphithoé fragilis Goés, 1866 (original desig- nation). See Shoemaker, 1930.
Accessory flagellum absent; lower lip with inner lobes obsolete (examined by me); inner plate of maxilla 1 with four or fewer terminal setae; gnathopods not eusirid, subchelate, article 5 not lobate, equal to or slightly longer than article 6; pereopods 3-5 with articles 5-6 (not article 4) each longer than article 2 (examined by me); antero- ventral cephalic corner with large tooth projecting forward to end of small rostrum, coxa 1 produced forward to anterior edge of eye. Species: 1, arctic-boreal, N. Atlantic, littoral (to 372 m).
Schraderia Pfeffer
Schraderia Pfeffer, 1888.—Stebbing, 1906.—K. H. Barnard, 1932. Atyloides Stebbing, 1888.
Type-species: SS. gracilis Pfeffer, 1888 (monotypy).
Accessory flagellum articulate, of medium length; lower lip lacking inner lobes; inner plate of maxilla 1 densely lined with setae along medial edge; gnathopods not eusirid, subchelate, article 5 not lobate, equal to article 6 in length, in gnathopod 2 especially articles 5-6 greatly elongate and sublinear; pereopods 3-5 with articles 4-6 each not longer than article 2. Species: 2, antarctic, littoral.
Zaramilla Stebbing Zaramilla Stebbing, 1888; 1906.
Type-species: Z. kergueleni Stebbing, 1888 (monotypy).
Accessory flagellum absent; lower lip with scarcely discrete inner lobes; inner plate of maxilla 1 densely lined with setae on medial edge; enathopods not eusirid, subchelate, article 5 not lobate, slightly longer than article 6; pereopods 3-5 with articles 4—6 each not longer than article 2, article 4 broadly expanded (especially Bovallia and Pontogeneia), article 4 two thirds as wide as article 2 (Pontogenevella) ; antennae subequal in length, coxa 4 strongly excavate posteriorly (Pros- tebbingia) ; dactyli of pereopods 3-4 as long as article 6, pereopods re- sembling those of oedicerotids (combining character). Species: 1, Kerguelen, littoral. Probably referrable to Haustoriidae and included in that family key (p. 252); absence of accessory flagellum in contrast to Haustoriidae.
MARINE GAMMARIDEAN AMPHIPODA 231
Marine Gammaridae Figures 94-96
Dracnosis.—Like the basic gammaridean, from which all other diagnoses in this paper are used to distinguish families, but accessory flagellum occasionally reduced to one long article, maxillipedal palp occasionally 3-articulate, gnathopod 1 rarely larger than 2, telson occasionally uncleft, inner ramus of uropod 3 reduced, coxae 1-4 rarely of nonuniform shape, coxa 4 occasionally not posteriorly excavate and pereopod 5 rarely elongate.
Description.—The Gammaridae are variable and many genera form links to other families. Accessory flagellum always present but varying from one long article to more than 20 short articles; peduncle of antenna 1 elongate; rostrum present or absent; upper lip not incised; mandible always bearing strong triturative molar and 3-articulate palp; lower lips variable, never ampithoid or pleustid in structure; maxillae fully developed, not strikingly foliaceous; palp of first maxilla 2-articulate; maxillipeds with well-developed plates, palp 3- or 4- articulate; gnathopods usually powerful and subchelate, occasionally slender and simple, gnathopod 1 rarely larger than 2; coxae of medium length, occasionally shortened; coxa 4 occasionally not excavate posteriorly; uropod 3 highly variable but rami never shorter than peduncle (except Parapherusa, Falklandella, and Gammarella) and usually flattened, not cylindrical, generally lanceolate, foliaceous or subquadrate, inner ramus occasionally very short; telson not elongate (except Parapherusa), usually deeply cleft but occasionally broader than long, uncleft or emarginate.
Re.ationsHips.—Gammaridae are difficult to define precisely be- cause so many genera have exceptional morphology. Four of the genera have uncleft telsons and thus resemble Calliopiidae but all have well-developed accessory flagella. They may also resemble some of the isaeid-like families but their telsons are not fleshy (not thickened dorsoventrally). The most difficult member of this quartet is Para- pherusa Stebbing which confounds all possible simplifications made in the keys and diagnoses of families and genera. The unique species of this genus has a third uropod resemblnig that of ischyrocerids or ampithoids (fig. 96d), unexcavate coxa 4, and its telson is entire but elongate, uropod 1 has a ventrodistal peduncular tooth like that of various ischyrocerids and isaeids. The absence of pereopodal glands apparently demonstrates its correct assignment to the Gammaridae, rather than to the glandular families. The other marine gammarid genera with uncleft telson have deep posterior excavations on coxa 4 and thus cannot be confused with isaeid-like genera. One ischyrocerid, Bathyphotis, however, has a strongly excavate coxa 4 and has a special
250, U.S. NATIONAL MUSEUM BULLETIN 271
Poy] |/K » K j YH —A/ i) YZ H|= ff
Ficure 94.—Gammaridae: a, Maera lovent (Bruzelius) (Sars, 1895, pl. 182); b, Megaluropus tlongimerus Schellenberg (J. L. Barnard, 1962b). Accessory flagella: c, Marinogam- marus marinus (Leach) (Sars, 1895, pl. 175); d, Elasmopus rapax Costa (Sars, 1895, pl. 183). Coxa 4: e, Maera othonis (Milne Edwards) (Sars, 1895, pl. 182); f, Gammarellus
homari (Fabricius) (Sars, 1895, pl. 172).
MARINE GAMMARIDEAN AMPHIPODA 233
place in Gammaridea Family Key A (p. 109). Its uropod 3 is even more typically that of an ischyrocerid than is uropod 3 of Parapherusa. Another 13 genera of Gammaridae (in couplets 6-18) have vestigial accessory flagella and cleft telsons and thus resemble Eusiridae. All Eusiridae except Eusirogenes Stebbing have elongate telsons and that genus is easily recognizable as a eusirid in its gnathopodal morphology. Eusiridae also have slightly enlarged metasomal segments, the meta- some having the length of 6 pereonal segments, whereas gammarids have a metasome the length of only 5 pereonal segments. This is a
Ficure 95.—Gammaridae: a, Melita dentata (Krdyer) (Sars, 1895, pl. 181). Mouthparts of Elasmopus rapax Costa (Sars, 1895, pl. 183): b, upper lip; c, lower lip; ¢, mandible; f,g, maxillae 1, 2; h, maxilliped. Lower lip: d, Gammarellus homari (Fabricius) (Sars, 1895, pl. 172). Gnathopod 2: 1, Elasmopus; j, Cheirocratus sundevalli (Rathke) (Sars, 1895, pl. 184). Gnathopod 1: k, Elasmopus; 1, Cheirocratus.
234 U.S. NATIONAL MUSEUM BULLETIN 271
difficult character analysis to practice however, and one must use simply a process of elimination by checking generic keys and diagnoses in both families for genera which are not obviously gammarid or eusirid. Five of the genera in couplets 6-18 have reduced inner rami of uropod 3 (melita-like uropod 3) and thus are easily eliminated from consideration as eusirids, for in the latter family, the inner ramus of uropod 3 is always as long as or longer than the outer.
Key to the Genera of Marine Gammaridae
Only those genera of Gammaridae with marine representatives are included in this key (except Falklandella). Some generic pairs, such as Maeropsis-Paraceradocus and Ceradocus-Ceradocoides have not been separated by firm distinctions. Other genera are cited twice in the key because of their intergrading or variable structures.
One must pursue both pathways in couplet 6 if the species at hand has three accessory flagellar articles. Since other characters are as variable as accessory flagella and more often poorly known, couplet 6 has been found to be more satisfactory than conceivable substitutes until some of the poorly described genera are clarified.
1.) Belson entire (fig. 96g) but often emarginate. ©7250 2 2 2°. 2.025 292 Telson cleft (fig. 96h)... . . BRENDAN NEMS" (6)
2. Urosomal segments with puonlles of Glove eines paoesommies with deep serrations on posterodorsal margins . . . . .. . . Mesogammarus Urosomal segments lacking dorsal bundles of spines, metasome lacking serrations but often with carinae. .. . Le) oh Sale ea
3. Rami of uropod 3 at least 1.5 times as long as 2 pede - wos eho ee Rami of uropod 3 equal to or shorter than peduncle ......... 5
4. Body dorsally carinate (even weakly), article 3 of mandibular palp shorter than article 2, lower lip without inner lobes . . . . . . Gammarellus !
Body dorsally smooth, article 3 of mandibular palp longer than article 2, lower lip with inner lobes. . .. . . . . . . Weyprechtia
5. Accessory flagellum 1-articulate, telson Beendes near long, emarginate, lower lip without inner lobes, coxa 4 posteriorly excavate. . . Falklandella
Accessory flagellum 8-articulate or more, telson longer than broad, not emarginate, lower e with strong inner lobes, coxa 4 not excavate pos-
teniorlye yuan a). . . . . . Parapherusa
6. Accessory flagellum a or D artrenlate! “eoeionallly Bi EGE En gy Accessory flagellum 4- or more articulate, occasionally 3. ..... . 19
7. Inner ramus of uropod 8 scale-like and much smaller than outer (fig. 96f). 8 Inner ramus of uropod 3 nearly as large as outer . . . Mea ca ts LP
8. Apex of uropod 3 not exceeding extent of uropods 1-2 (fig. 96a) . Melitoides Apex of uropod 3 greatly exceeding extent of uropods 1-2. ...... 9
9. One or more of coxae 1-4 very a longer (almost GO) than coxae Mp oviiae Cony renee Aca taal: KAS gine cui VEC Ital All coxae selocamell 3 in Tort very Short Ae i A Gay gL Ea ected a ame LO)
1 Gammaracanthus has a telson broader than long but so deeply emarginate as to be considered cleft.
10.
be
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
MARINE GAMMARIDEAN AMPHIPODA 235
Maxilla 2, medial edge of inner plate lined with setae (fig. 95g), article 2 of outer ramus of uropod 3 subequal to article 1 in length (fig. 96f) . Eriopisa Maxilla 2, medial edge of inner plate unarmed, article 2 of outer ramus of
uropod 3 much shorter than article 1 or absent. ........ . Il Outer ramus of uropod 3 biarticulate. ....... .... . Eriopisella Outer ramus of uropod 3 uniarticulate. .. . . . . . Netamelita Rami of uropod 3 foliaceous, oval, apically Poured (Gig. 96e), either coxa 2
or 3 much shorter thancoxal..... . . . . . Megaluropus Rami of uropod 3 lanceolate, or apices Aanoete es 96b,d), not foliaceous,
coxae 2-3 aslongascoxal.. . RS ie eat aN Ne ee NAIR ee U) Mandibular palp article 2 shorter ‘them penele ioe . . . . Parelasmopus Mandibular palp article 2 equal to orlongerthanarticlel....... 14 Cuathopodwlysubchelatemuh maieweePikts wheel hier el Rey sees 2 LO Gnathopod 1 simple (fig. 957). . . . DEV us wie Wie: ENE iL aie Inner plate of maxilla 1 poorly seen (2- 5 ‘gaind) . . . . Elasmopus Inner plate of maxilla 1 densely setose (11 setae) ... Bee OU LG Uropod 3 strongly exceeding uropod 1, outer ramus Dearticulate:
Liagoceradocus Uropod 3 not exceeding uropod 1, outer ramus l-articulate. Anelasmopus ” Female gnathopod 2 simple (fig. 951). . . ... =.=. +... . Cheirocratus Hemaleynathopod 2; sulbehelate ims 5) san yi Bem ele We ee ue GA sla Ween We) Antennal nearly as long as;antennaiy2, 2 5... . .). 4. Gheirgecatella Antenna 1 reaching only to end of peduncle of antenna2... . . Casco All pereonites carinate. ... . . . . . . Gammaracanthus Pereonites not carinate, or pecanionally aumealboe 6-7 are carinate . . . 20 Urosome with two or three segments panape bundles or rows of articulate
Spines). . ; ea urea varie Urosome saansitonellhy fli seniierad syne nec on fll {pene and not ar-
ranged in bundles, spines usually absent . ... . spe Wausi er Geis aoe
Some segments anterior to pleonite 4 bearing dorsal prcalerte spines. “echinogammarus” members of Anisogammarus No segments anterior to pleonite 4 bearing dorsal articulate spines (oc-
casionally bearing teeth or serrations) .. . is er aR Cea le at 274 Metasome with posterodorsal teeth and/or someone, La MALL oc adlves De Metasome lacking posterodorsal teeth and serrations . . bate 5 ills AD) Telsonic apices quadrate, lobes closely appréssed and peoerted by a slit.
Maerella Telsonic apices subacute, lobes gaping. . . MOV io niet iO
Metasomites with a single dorsoposterior feothe ‘“Marine Carinate Gammarus”
Metasomites multiserrate dorsoposteriorly . . .. . . . Mesogammarus Gnathopod 1 slightly larger than 2, both gnathopodal palms bearing two or
more chisel-shaped spines in rows. . . . . . . . Anisogammarus Gnathopod 1 smaller than or oe to 2, spalizes bearing one or no chisel-
shaped spines. ... . . . . . Gammarus Inner ramus of uropod 3 sone, very “ sirocill Ge 96f) IM aed aa Set, [nner ramus.of ureped 3 nearly,as long as outer. . 6... ...,.,. 28 Uropod 3 strongly exceeding apex of uropod1 ....... ... Melita Uropod 3 not exceeding apex of uropod1........ .. .. Melitoides Rami of uropod 3 foliaceous, oval, apices rounded (fig. 96e). . . . . . 29 Rami of uropod 3 lanceolate, square, or apices truncate or pointed . . . 30
2 Not clearly described. See diagnoses. Hornellia Walker resembles Anelasmopus but the inner plate of maxilla 1 has not been described.
285-135 O - 69 - 16
236 U.S. NATIONAL MUSEUM BULLETIN 271
29. Kyes four in tae inner plate of maxilla 1 with medial edge fully lined
Walthesetaer nies cl: .. . . . . Quadrivisio
Eyes two in number; inner wallet oi snes, 1 Ruin only terminal setae. Paraceradocus 30. Maxilla 2, medial edge of inner plate not lined with setae ...... 383i Maxilla 2, medial edge of inner plate lined with setae ........ 388 31. Palp of maxilliped with three articles ......... =... Maerella Palp of maxilliped with four articles. . . . LH as ED MEET, RED eR Ny 32. Mandibular palp article 3 stout, falcate (fig. 95e) . .. . . . Elasmopus Mandibular palp article 3 slender, not faleate. . . . . . . . . Maera
33. Palm and posterior margin of article 6 on gnathopod 2 nee distinct from one another (fig. 951), article 7 of gnathopod 2 nearly as long as article 6,
rami of uropod 3 equal to peduncle in length. . . . . . Gammarella Mheseicharacters not: combined aia cee ey yer eee kee Oe A See
34. Lower lip lacking inner lobes... . BS chil oa) Lower lip bearing inner lobes (presumptive in @oradeeories and Maera- cunha). . Aiba eho)
35. Mandibular alo cntiiale 3 naed on meal eee! Saw of Peeailined about 75% as long as article 3, claw-shaped, coxa 4 not excavate posteriorly.
Ceradocopsis Mandibular palp article 3 setose medially, dactyl of maxilliped 25% as long as article 3, setose, coxa 4 excavate posteriorly ... . Bathyceradocus 36. Maxilla 1, medial edge of inner lobe not lined with setae. . ..... 37 Maxilla 1, medial edge of inner lobe lined with setae. . ...... . 38 37. Outer ramus of uropod 3 bearing minute article2 ... . . Maeracunha Outer ramus of uropod 3 lacking article2....... . .. . Maeropsis 38:3) | Uropod 3) with’ 2-articulate outer ramus si) an eee Uropod 3 with l-articulate outer ramus .. . OAS aan A)
39. Mandibular palp article 3 subequal to article 1, haaing only two setae. Maeracunha
Mandibular palp article 3 slightly longer than article 1, multisetose.
Metaceradocoides 40. Male gnathopod 2 very small and slender. . ... . . . Metaceradocus Male gnathopod 2 enlarged (fig. 95a). . . . Sian ASL 41. Mandibular palp article 3 as long as article 2, elietiily digdeshoned, article 1 simple. . . . . . Elasmopoides
Mandibular niin pcticle 3 les chen 7 5% andl penerally half or less as long as article 2, article 1 with distomedial process or spines. Ceradocus and Ceradocoides 4
3 Not clearly described.
(Genera of Marine Gammaridae
The term ‘‘NORMAL GNATHOPODS” indicates that both pairs of gnathopods are subchelate and gnathopod 2 is larger than 1 in the male.
Anelasmopus Oliveira Anelasmopus Oliveira, 1953.
Type-species: A. kraui Oliveira, 1953 (monotypy).
MARINE GAMMARIDEAN AMPHIPODA 237
Accessory flagellum 2-articulate; lower lip [unclear], apparently with inner lobes; inner plate of maxilla 1 densely setose (11 setae), inner plate of maxilla 2 setose partially on medial edge; gnathopods normal; uropod 3 short, not exceeding uropod 1, rami short, rec- tangular, inner slightly shorter than outer, outer 1-articulate; telson deeply cleft; body lacking dorsal carinae, teeth or spines. Species: 1, Brazil, littoral.
Anisogammarus Dershavin
Antsogammarus Dershavin, 1927.—Gurjanova, 1951. Eogammarus Birstein, 1933 [a valid subgenus].
as
h
Figure 96.—Gammaridae: a, Gammaracanthus loricatus (Sabine) (Sars, 1895, pl. 174, as G. relictus). Uropod 3: b, Cheirocratus sundevalli (Rathke) (Sars, 1895, pl. 184); ¢, Melita dentata (Krgyer) (Sars, 1895, pl. 181); d, Elasmopus rapax Costa (Sars, 1895, pl. 183); e, Gammaracanthus; f, Eriopisa elongata (Bruzelius) (Sars, 1895, pl. 181). Tel- son: g, Gammarellus homari (Fabricius) (Sars, 1895, pl. 172); h, Elasmopus; 1, Gammaracanthus.
238 U.S. NATIONAL MUSEUM BULLETIN 271
Type-species: Gammarus pugettensis Dana, 1853 (monotypy and subsequent synonymy). See J. L. Barnard, 1954.
Accessory flagellum more than 3-articulate; lower lip with small or obsolete inner lobes; inner plates of maxillae both densely setose medially; gnathopod 1 slightly larger than 2, palms bearing many chisel-shaped spines; uropod 3 exceeding uropod 1, outer ramus long, 2-articulate (? rarely 1-articulate), inner ramus long or very short; telson deeply cleft; urosome and occasionally metasome armed with dorsal bundles of spines, except one species [type of Anzsogammarus] having a tooth on urosomite 2 instead of spines. Species: 18, boreal Pacific, littoral and freshwater. Includes former marine members of Echinogammarus. Possibly this genus divisible into 4 subgenera.
Bathyceradocus Pirlot
Bathyceradocus Pirlot, 1934
Type-species: B. stephenseni Pirlot, 1934 (original designation).
Accessory flagellum multiarticulate; lower lip lacking inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods small, subchelate, slender; uropod 3 exceeding uropod 1, rami of me- dium length, subequal, broadly lanceolate, outer 1-articulate; telson deeply cleft; metasome and urosome toothed and carinate, urosome with scattered dorsal spines; palp article 4 of maxilliped short, blunt, apically setose (Metaceradocoides). Species: 1, Indo-Pacific, bathyal- abyssal (1,165-4,930 m).
Casco Shoemaker
Casco Shoemaker, 1930.
Type-species: Cheirocratus bigelowi Blake, 1929 (original desig- nation).
Accessory flagellum 2-articulate; lower lip with inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopod 1 essentially simple, but dactyl large and folded back on article 6, gnathopod 2 slightly larger than 1 and slightly subchelate; uropod 3 exceeding uropod 1, rami subequal, long, lanceolate, outer 1-articulate; telson deeply cleft; urosome with a tooth and scattered spines; antenna 1 reaching only to end of peduncle of antenna 2 (Cheirocratella); antero- ventral corner of head strongly produced. Species: 1, boreal W. Atlantic, littoral.
Ceradocoides Nicholls
Ceradocoides Nicholls, 1938. Type-species: ©. chiltoni Nicholls, 1938 (original designation).
MARINE GAMMARIDEAN AMPHIPODA 239
Accessory flagellum more than 4-articulate; [lower lip unknown]; inner plates of maxillae 1-2 densely setose medially; gnathopods apparently normal; uropod 3 not exceeding uropod 1, rami equal, lanceolate [outer ramus articles not described]; telson cleft shortly; hody lacking dorsal teeth or spines (distinctions from Ceradocus by Nicholls, 1938, unclear). Species: 1, antarctic (420 m).
Ceradocopsis Schellenberg Ceradocopsis Schellenberg, 1926a.
Type-species: C. kergueleni Schellenberg, 1926a (monotypy).
Accessory flagellum more than 4-articulate; lower lip lacking inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods normal; uropod 3 not exceeding uropod 1, rami equal, short, outer 2-articulate; telson deeply cleft; body lacking dorsal teeth or spines. Species: 1, Kerguelen Island, littoral.
Ceradocus Costa Ceradocus Costa, 1853c, 1857.—Stebbing, 1906.
Type-species: C. orchestiipes Costa, 1853c (monotypy). See Chevreux and Fage, 1925.
Accessory flagellum 3- or more articulate; lower lip with inner lobes; inner plate of maxilla 1 densely setose medially, of maxilla 2 moderately to strongly setose medially; gnathopods normal; uropod 3 exceeding uropod 1, rami equal, lanceolate, broad or slender, outer l-articulate; telson deeply cleft; palp article 3 of mandible half as long as article 2 (Metaceradocus, Elasmopoides), article 1 of palp with medial process or spines or cusp. Species: 18, cosmopolitan, littoral (and 3 bathyal).
Cheirocratella Stephensen Chetrocratella Stephensen, 1940.
Type-species: C. thori Stephensen, 1940 (original designation).
Accessory flagellum 2-articulate; [lower lip unknown]; inner plates of maxillae 1-2 apparently densely setose medially; gnathopod 1 simple; gnathopod 2 larger and subchelate (only female is known); [uropod 3 unknown]; telson deeply cleft; metasomal and urosomal segments dorsally toothed, urosome with scattered spines; antennae subequal in length (Casco). Species: 1, Iceland, littoral.
Cheirocratus Norman Chetrocratus Norman, 1867a.—Stebbing, 1906.
Type-species: Gammarus assimilis Liljeborg, 1851 (monotypy and subsequent synonymy). See Sars, 1895.
240 U.S. NATIONAL MUSEUM BULLETIN 271
Accessory flagellum 2-, occasionally 3-articulate; lower lip with inner lobes; inner plates of maxillae 1-2 densely setose medially; female gnathopods simple, slender, male gnathopod 1 simple, male gnathopod 2 powerfully subchelate; uropod 3 exceeding uropod 1, rami equal, lanceolate, outer 1l-articulate; telson deeply cleft; uro- some with dorsal teeth and setae. Species: 4, boreal N.E. Atlantic, littoral.
Elasmopoides Stebbing
Elasmopoides Stebbing, 1908b.
Type-species: E. chevreuxi Stebbing, 1908b (monotypy).
Accessory flagellum exceeding 20 articles; lower lip with inner lobes; inner plate of maxilla 1 densely setose medially, of maxilla 2 moder- ately to strongly setose medially; gnathopods normal; uropod 3 not exceeding uropod 1, rami equal, broad, outer 1-articulate; telson deeply cleft, lobes separated basally; palp article 3 of mandible as long as article 2 and slightly sickle-shaped; article 1 simple (Ceradocus). Species: 1, S. Africa, sublittoral.
Elasmopus Costa
Elasmopus Costa, 18538c, 1857.—Stebbing, 1906. Neogammaropsis Stout, 1913.
Type-species: EH. rapax Costa, 1853c (monotypy). See Sars; 1895; Chevreux and Fage, 1925; J. L. Barnard, 1962b.
Accessory flagellum 3- or more, occasionally 2-articulate; lower lip with inner lobes; inner plates of maxillae 1-2 with only terminal setae; gnathopods normal; uropod 3 variable in length, rami equal, rectangular, outer l1-articulate; telson deeply cleft; urosome occasionally with dorsal teeth; palp article 3 of mandible falcate, stout (Maera). Species: 35, tropico-warm-temperate, cosmopolitan, littoral (one bathyal record).
Eriopisa Stebbing
Eriopis Bruzelius, 1859 (homonym, Coleoptera). Eriopisa Stebbing, 1890 (new name); 1906.
Type-species: riopis elongata Bruzelius, 1859 (monotypy). See Sars, 1895.
Accessory flagellum 2-articulate; lower lip with slight inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods normal; uropod 8 greatly exceeding uropod 1, immensely elongate, outer ramus with two very long, subequal articles, or article 2 oc- casionally short, inner ramus very short, scale-like; telson deeply cleft; body lacking dorsal teeth or spines. Species: 6, cosmopolitan, littoral to bathyal (752 m).
MARINE GAMMARIDEAN AMPHIPODA 241
Eriopisella Chevreux
Eriopisella Chevreux, 1920.
Type-species: EH. pusilla Chevreux, 1920 (monotypy).
Accessory flagellum 2- articulate or less; lower lip with scarcely defined inner lobes fused with outer lobes; inner plates of maxillae 1-2 setose only terminally; gnathopods small, subchelate, equal in size; uropod 3 greatly exceeding uropod 1, immensely elongate, outer ramus 2-articulate, article 2 much shorter than article 1, inner ramus very short, scale-like; telson deeply cleft; body lacking dorsal teeth or spines. Species: 3, France, Seychelles, India, littoral.
Falklandella Schellenberg
Falklandella Schellenberg, 1930, 1931.
Type-species: FF. obtusa Schellenberg, 1930, 1931 (monotypy).
Accessory flagellum 1-articulate; lower lip lacking inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopod 1 larger than gnathopod 2, subchelate; uropod 3 short, not exceeding uropod 1, rami semirectangular, outer l-articulate, inner half as long as outer; telson short, emarginate; body lacking dorsal teeth or spines. Species: 2, Falkland Islands, freshwater close to sea.
Gammaracanthus Bate
Gammaracanthus Bate, 1862.—Stebbing, 1906.
Type-species: Gammarus loricatus Sabine, 1821 (original designa- tion!, Bate, 1862, p. 203). See Gurjanova, 1951.
Accessory flagellum 3- or more articulate; lower lip with inner lobes obsolescent; inner plate of maxilla 1 setose only terminally, of maxilla 2 densely setose medially; gnathopods large, nearly equal in size, nearly eusirid in structure (fig. 96a); uropod 3 slightly exceeding uropod 1, rami subequal, foliaceous or subfoliaceous; outer 1-artic- ulate; telson very short, cleft, lobes gaping; body carinate through- out. Species: 1, arctic and Caspian Sea, littoral.
Gammarella Bate, new synonymy
Pherusa Leach, 1814a.—Stebbing, 1906 (homonym, Polychaeta). Gammarella Bate, 1857a. Pherusana J. L. Barnard, 1964c (new name for Pherusa).
Type-species: Pherusa fucicola Leach, 1814a (monotypy and subsequent synonymy). See Walker, 1891la, b; Chevreux and Fage, 1925.
Accessory flagellum 4- or more articulate; lower lip with inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods
242 U.S. NATIONAL MUSEUM BULLETIN 271
normal; uropod 3 short, not exceeding uropod 1, rami not longer than peduncle, outer l-articulate, inner slightly shortened; telson cleft; urosome dorsally carinate; palm and posterior margin of article 6 on gnathopod 2 not separate, article 7 nearly as long as article 6 (com- bination of characters). Species: 1, subtropical E. Atlantic and Mediterranean, littoral.
Gammarellus Herbst
Gammarellus Herbst, 1793.—Stebbing, 1906.
Amathia Rathke, 1837 (homonvm, Hydrozoa).
Grayia Bate, 1862.—Bate and Westwood, 1863.
Amathilla Bate and Westwood, 1863 (new name for Amathza).
Type-species: <Astacus homari J. C. Fabricius, 1779 (type indicated by Stebbing, 1899a, but firmly selected by Chevreux and Fage, 1925). See Sars, 1895.
Accessory flagellum more than 3-articulate; lower lip lacking inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods normal; uropod 3 short, not exceeding uropod 1, rami lanceolate, outer l-articulate; telson entire or emarginate; body dorsally carinate. Species: 2, arctic and high boreal, littoral.
Gammarus Fabricius
Gammarus Fabricius, 1775.—Stebbing, 1906.
Marinogammarus Schellenberg, 1937a—Sexton and Spooner, 1940 [subgenus, often considered a full genus].
Pectenogammarus Reid, 1940 [valid subgenus].
Type-species: Cancer locusta Linné, 1758 (selected by Boeck, 1876). See Sars, 1895.
Accessory flagellum 3- or more articulate; lower lip with inner lobes obsolescent or absent; inner plates of maxillae 1-2 densely setose medially; gnathopods normal or gnathopod 1 slightly larger than 2; uropod 3 exceeding uropod 1, outer ramus 2-articulate, inner variable, long or short; telson deeply cleft; no dorsal teeth or carinae, urosome with dorsal spine-bundles. Marine and brackish species: 13-15, arctic (boreal in Atlantic), littoral. See ‘“Marine Carinate Gammarus” (p. 244).
Hornellia Walker
Hornellia Walker, 1904.
Type-species: H. incerta Walker, 1904 (monotypy).
Accessory flagellum 2-articulate; inner lobes of lower lip obsoles- cent; [inner plate of maxilla 1 undescribed], inner plate of maxilla 2 densely setose on medial edge; gnathopods normal; uropod 3 not exceeding uropod 1, rami equal, broadly lanceolate, outer 1-articulate;
MARINE GAMMARIDEAN AMPHIPODA 243
telson long and deeply cleft; metasome and urosome dorsally serrate. Species: 1, Ceylon, littoral.
Liagoceradocus J. L. Barnard Liagoceradocus J. L. Barnard, 1965.
Type-species: L. pusillus J. L. Barnard, 1965 (original designation)
Accessory flagellum 2-articulate; lower lip lacking inner lobes; inner plates of maxillae 1-2 densely setose medially, gnathopods feeble, subchelate, uropod 3 exceeding uropod 1, rami long, equal, broadly lanceolate, outer 2-articulate; telson deeply cleft; body lack- ing dorsal teeth or spines. Species: 1, Micronesia, littoral.
Maera Leach
Maera Leach, 1814a.—Stebbing, 1906.—Schellenberg, 1938. Mulleria Leach, 1814a (nomen nudum).
Leptothoe Stimpson, 1853.
Linguimaera Pirlot, 1936.
Type-species: Cancer grossimanus Montagu, 1808 (monotypy). See Chevreux and Fage, 1925.
Accessory flagellum 3- or more articulate; lower lip with inner lobes; inner plates of maxillae setose only terminally; gnathopods normal; uropod 3 long or short, rami equal, lanceolate or slightly rectangular, outer 1-articulate; telson deeply cleft; urosome occa- sionally toothed; mandibular palp article 3 slender, not falcate (Elas- mopus). Species: 35, cosmopolitan, primarily tropical, littoral to bathyal (1,238 m).
Maeracunha Stephensen Maeracunha Stephensen, 1949.
Type-species: MM. tristanensis Stephensen, 1949 (monotypy).
Accessory flagellum more than 4-articulate; lower lip with small inner lobes; inner plate of maxilla 1 with seven partially medial setae, inner lobe of maxilla 2 densely setose medially; gnathopods normal; uropod 3 short, rami subequal, sublanceolate, outer 2-articulate; telson deeply cleft; metasome and urosome untoothed; mandibular palp article 3 subequal to article 1, bearing only two setae (combining character). Species: 1, Tristan da Cunha, littoral.
Maerella Chevreux
Maerella Chevreux, 191 1a.
Type-species: Gammarus tenuimanus Bate, 1862 (monotypy). See Chevreux and Fage, 1925.
244 U.S. NATIONAL MUSEUM BULLETIN 271
Accessory flagellum 4- or more articulate; lower lip with inner lobes; inner plates of maxillae setose only terminally; gnathopods normal; uropod 3 exceeding uropod 1, rami equal, outer 1-articulate; telson cleft shortly; metasome and urosome dorsally toothed, urosome with scattered spines; mazillipedal palp with only three articles. Species: 1, temperate N. E. Atlantic, littoral.
Maeropsis Chevreux
Maeropsis Chevreux 1919; 1927.
Type-species: MM. perrieri Chevreux, 1919, 1927 (monotypy).
Accessory flagellum more than 4-articulate; lower lip with inner lobes; inner plate of maxilla 1 setose only terminally, of maxilla 2 densely setose medially; gnathopods normal; uropod 3 not exceeding uropod 1, rami equal, rectangular, outer l-articulate; telson deeply cleft; body lacking dorsal teeth or spines. Species: 1, N. Atlantic, bathyal (698 m).
**Marine Carinate Gammarus”’
Species: Gammarus mucronatus Say, 1818.
Accessory flagellum 3- or more articulate; lower lip with inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods normal, subequal in size; uropod 3 exceeding uropod 1, rami long, outer 2-articulate, inner slightly shorter than outer; telson deeply cleft; each metasomite with one dorsal tooth; urosomites with dorsal bundles of spines. Species: 1, Atlantic and Gulf United States, littoral lagoonal. Formerly included with Baikalian Carinogammarus but having affinities with freshwater Rivulogammarus. See Meso- gammarus.
Megaluropus Hoek
Megalonoura Herdman, 1889 (nomen nudum). Megaluropus Hoek, 1889.—Stebbing, 1906.—K. H. Barnard, 1940. Phylluropus K. H. Barnard, 1932.
Type-species: M. agilis Hoek, 1889 (monotypy). See Chevreux and Fage, 1925.
Accessory flagellum 2-articulate; lower lip with inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods feeble, poorly subchelate; uropod 3 exceeding uropod 1 or not, rami equal, foliaceous, outer 1-articulate; telson deeply cleft; metasome and urosome often dorsally serrate. Species: 3, bi-boreal, littoral (pri- marily neritic).
MARINE GAMMARIDEAN AMPHIPODA 9A5
Melita Leach, new synonymy
Melita Leach, 1814a.—Stebbing, 1906.—Shoemaker, 1941b.
Boscia Leach, 1814a (nomen nudum).
Megamoera Bate, 1862 (type by present selection, Gammarus dentatus Krgyer, 1842).
Ps Stout, 1913.
Dulichiella Stout, 1918.
Type-species: Cancer palmatus Montagu, 1804 (monotypy). See Sars, 1895.
Accessory flagellum 2- or more articulate; lower lip with slight inner lobes; inner plate of maxilla 1 with largely terminal setae, of maxilla 2 densely setose medially; gnathopods normal; uropod 3 greatly exceeding uropod 1, outer ramus very long, 1-articulate, inner short and scale-like; telson deeply cleft; metasome often toothed, urosome often toothed and bearing scattered dorsal spines. Species: 45, cosmopolitan, littoral to abyssal.
Melitoides Gurjanova
Melitoides Gurjanova, 1934.
Type-species: MM. makarovt Gurjanova, 1934 (monotypy). See Gurjanova, 1951.
Accessory flagellum 4-articulate; lower lip with inner lobes; inner plates of maxillae densely setose medially; gnathopods normal; uropod 3 not exceeding uropod 1, outer ramus long, 1-articulate, inner ramus short, scale-like; telson deeply cleft; urosome dorsally toothed. Species: 1, subarctic, littoral.
Mesogammarus Tzvetkova
Mesogammarus Tzvetkova, 1965.
Type-species: MM. melitoides Tzvetkova, 1965 (original desig- nation).
Accessory flagellum more than 3-articulate; lower lip with scarce indication of inner lobes; inner plates of maxillae 1-2 lined medially with setae; gnathopod 1 slightly larger than 2; uropod 3 apparently exceeding uropod 1, rami long, lanceolate, subequal in length, outer ramus with short article 2; telson short, broad, cleft halfway, lobes gaping [possibly interpreted as deeply emarginate]; pleonites 1-3 with posterodorsal margins cut into several medium-sized teeth; urosome armed dorsally with rows of spines. Species: 1, Japan Sea, near Vladivostok, littoral.
Metaceradocoides Birstein and Vinogradova
Metaceradocoides Birstein and Vinogradova, 1960.
246 U.S. NATIONAL MUSEUM BULLETIN 271
Type-species: MM. vitjazi Birstein and Vinogradova, 1960 (original designation).
Accessory flagellum greatly more than 4-articulate; lower lip with inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopod 2 longer and more slender than gnathopod 1, both sub- chelate; uropod 3 not exceeding uropod 1, rami equal, lanceolate, outer 2-articulate; body lacking dorsal teeth or spines; palp article 4 of maxilliped long, claw-like (Bathyceradocus). Species: 1, N.W. Pacific, hadal (7,210 m).
Metaceradocus Chevreux
Metaceradocus Chevreux, 1925.
Type-species: M. perdentatus Chevreux, 1925 (monotypy).
Accessory flagellum 3- or more articulate; lower lip with inner lobes; inner plate of maxilla 1 mostly setose terminally, of maxilla 2 densely setose medially; gnathopods feeble, slender, subchelate; uropod 3 exceeding uropod 1, rami long, equal, lanceolate, outer l-articulate; telson deeply cleft; pleonites dorsally serrate and with scattered spines; mandibular palp articles 2 and 3 subequal (Ceradocus). Species: 2, E. Atlantic, E. Pacific, littoral.
Netamelita J. L. Barnard
Netamelita J. L. Barnard, 1962b.
Type-species: N. cortada J. L. Barnard, 1962b (original desig- nation).
Accessory flagellum 1-articulate; lower lip with inner lobes; inner plates of maxillae 1-2 setose only terminally; gnathopods small, subchelate, equal in size; uropod 3 exceeding uropod 1, outer ramus long, 1-articulate, inner short, scale-like; telson deeply cleft; body lacking dorsal teeth or spines. Like Eriopisella but outer ramus of uropod 3 uniarticulate. Species: 1, California, littoral.
Paraceradocus Stebbing
Paraceradocus Stebbing, 1899a; 1906.
Type-species: Megamoera miersii Pfeffer, 1888 (original designa- tion). See K. H. Barnard, 1932.
Accessory flagellum 4- or more articulate; lower lip with imner lobes; inner plate of maxilla 1 only terminally setose, of maxilla 2 densely setose medially; gnathopods normal; uropod 3 exceeding uropod 1, rami foliaceous, equal, long, outer 1-articulate; telson deeply cleft; some urosomal segments dorsally toothed. Note: P. micramphopus Stebbing, 1910, is probably referrable to Metaceradocus. Species: 2, subantarctic, Australia, littoral (to 310 m).
MARINE GAMMARIDEAN AMPHIPODA 247
Parapherusa Stebbing
Harmomia [lapsus for Harmonia] Haswell, 1880c (Harmonia =homonym,
Coleoptera).
Chloris Haswell, 1880a (homonym, Aves). Parapherusa Stebbing, 1906 (new name for Harmomia).
Type-species: Harmonia crassipes Haswell, 1880c (monotypy). See Stephensen, 1949.
Accessory flagellum 4- or more articulate; lower lip with large inner lobes; inner plates of maxillae 1-2 densely setose medially; gnathopods normal; uropod 3 short, not exceeding uropod 1, rami equal, shorter than peduncle, outer 1-articulate and bearing enlarged serrate spine; telson simple, elongate; body lacking dorsal teeth or spines. Species: 1, antiboreal, Australia-New Zealand, littoral.
Parelasmopus Stebbing
Parelasmopus Stebbing, 1888; 1906.
Type-species: Gammarus suluensis Dana, 1852a (monotypy). See Chevreux, 1901b (as P. setiger).
Accessory flagellum 2- or 3- articulate; lower lip with inner lobes; inner plate of maxilla 1 setose only terminally, of maxilla 2 generally terminally setose; gnathopods normal; uropod 3 not exceeding uropod 1, rami short, broad, lanceolate, outer l-articulate; telson deeply cleft; metasome and urosome dorsally carinate, no spines; mandibular palp article 2 shorter than article 1. Species: 3, tropical Pacific, littoral (to 522 m).
Quadrivisio Stebbing
Quadrivisio Stebbing, 1907. Pseudoceradocus Shoemaker, 1933b. Type-species: Q. bengalensis Stebbing, 1907 (monotypy). Accessory flagellum greatly more than 4-articulate; lower lip with inner lobes; inner plates of maxillae 1-2 densely setose medially ; enathopods normal; uropod 3 exceeding uropod 1, rami equal, foli- aceous, outer l-articulate; telson deeply cleft; body lacking dorsal teeth or spines; eyes four in number. Species: 2, circumtropical, littoral.
Weyprechtia Stuxberg
Weyprechtia Stuxberg, 1880.—Stebbing, 1906.
Type-species: Amathilla heuglini Buchholz, 1874 (monotypy and subsequent synonymy). See Gurjanova, 1951.
Accessory flagellum 4- or more articulate; inner lobes of lower lip obsolescent (in drawings they appear distinct); inner plates of maxillae
248 U.S. NATIONAL MUSEUM BULLETIN 271
1-2 densely setose medially; gnathopods slender, feeble, poorly sub- chelate; uropod 3 not exceeding uropod 1, rami equal, lanceolate, outer l-articulate; telson entire; body lacking dorsal teeth or spines. Species: 2, arctic, littoral.
Haustoriidae
Figures 97-100
Dracnosis.—Antenna 1 usually with accessory flagellum; rostrum usually absent but when present then pereopod 5 longer than 4; otherwise pereopod 5 shorter and of different structure than pereopod 4; pereo- pods strongly spinose and setose (fossorial); gnathopods feeble. See Gammaridae, Phoxocephalidae, Liljeborgiidae, Ampeliscidae, Param- phithoidae, Astyridae, Argissidae.
Description.—Accessory flagellum absent (rare) or 1- to multi- articulate; peduncles of antennae elongate or not; antenna 2 peduncle occasionally spinose, often with elongate plumose setae; rostrum usually inconspicuous, occasionally present; upper lip not strongly incised; mandible always with 3-articulate palp and molar rarely absent but often nontriturative; lower lip with inner lobes; maxillae basic; maxilliped with well-developed plates, palp 3-4 articulate, occa- sionally of peculiar shape; gnathopods feeble, varying from minutely chelate, through simple to lacking article 7; coxae of medium length or long, highly variable, occasionally acuminate; uropod 3 variable but one or both rami longer than or equal to peduncle, lanceolate, rectangular, or subcylindrical, rarely flabellate; telson short or medium in length, deeply cleft or subentire.
RELATIONSHIP.—The family is so neatly linked to the Gammaridae by the genera Pontoporeia and Bathyporeia that on gross morpho- logical grounds there seems little reason to maintain it; nevertheless taxonomists seem to have been able to recognize its members (ex- cepting Hurinia Tucolesco). Of particular significance are the feeble enathopods of haustoriids and the strong development of spines and setae for these digging animals. Certainly the fossorial habit dis- tinguishes them from most Gammaridae.
Liljeboriids have a weak mandibular molar, poorly spinose append- ages and very powerful gnathopods.
Priscillina and Haustorius resemble the Paramphithoidae through their subacute first four coxae but the excessive spination and seta- tion of appendages and multiarticulate accessory flagella are distinc- tive of Haustoriidae.
The Phoxocephalidae are closely related and merge into Haus- torlidae through genera such as Platyischnopus and Zobracho. Most
MARINE GAMMARIDEAN AMPHIPODA QAO
haustortids lack a distinct rostrum. All phoxocephalids have pereopod 5 shorter than 4 and of different structure, and a large visor-like rostrum. Some haustoriids have a small rostrum and pereopod 5 shorter than 4 and of different structure. With some genera, one must flip a coin to choose the proper family.
g
Ficure 97.—Haustoriidae: a, Priscillina armata (Boeck) (Sars, 1895, pl. 42); b, Urothoe elegans Bate (Sars, 1895, pl. 47, as U. norvegica). Upper lip: c, Haustorius arenarius (Slabber) (Sars, 1895, pl. 46). Mandible: d, Urothoe; e, Haustorius; f, Carangolia mandt- bularis J. L. Barnard (1962d). Lower lip: g, Haustorius; h, Pontoporeia femorata Krgyer (Sars, 1895, pl. 41).
250 U.S. NATIONAL MUSEUM BULLETIN 271
re p
HN P09
Ficure 98.—Haustoriidae: a, Pontoporeia femorata Krdyer (Sars, 1895, pl. 41); b, Zobracho canguro J. L. Barnard (1961). Maxilla 1: c, Haustorius arenarius (Slabber) (Sars, 1895, pl. 46). Maxilla 2: d, Pontoporeia; e, Haustorius. Maxilliped: f, Haustorius; g, Prisctl- lina armata (Boeck) (Sars, 1895, pl. 42). Urosome: h, Urohaustorius halei Sheard (1936b).
MARINE GAMMARIDEAN AMPHIPODA 251
All Ampeliscidae have pereopod 5 shorter than 4 and of different structure, similar to all Phoxocephalidae and some Haustoriidae. Unlike the burrowing Phoxocephalidae and Haustoriidae, the Am- peliscidae have glands in pereopods 1-2 that secrete a substance used for the formation of tubes. Ampeliscidae never have multiple ommatidia in the eyes as in Phoxocephalidae and Haustoriidae but form cuticular Jenses. They have elongate, laterally compressed heads. Unlike phoxocephalids and haustoriids, Ampeliscidae lack any trace of an accessory flagellum. The accessory flagellum of Carangolia is vestigial but the animal is otherwise clearly an haustoriid.
Oedicerotidae, with fossorial pereopods and always with vestigial or absent accessory flagella resemble a few haustoriids, but Oedicero- tidae always have pereopod 5 very elongate, uropod 3 with elongate peduncle, elongate subequal rami and linguiform, entire (rarely emareginate) telson.
A major division of the key to genera concerns the absence of pereopodal dactyls. The ends of those legs often are highly spinose and if one of the ‘‘spines” is not elongate nor obviously distinct from the others then dactyls are considered to be absent.
Nomenclatural Changes for Haustoriidae
See Bousfield (1965) for specific nomenclatural changes.
Haustoriopsis Schellenberg (1938) is removed to the Prophliantidae.
Urothoe simplignathia J. L. Barnard (1962d) is removed to Par- daliscella, Pardaliscidae.
Note the comments in the Calliopiidae on the genus Sancho. Haustoriids are perhaps the most interesting group of amphipods
and so diversified that by necessity many genera have been illus-
trated herein.
Key to the Genera of Haustoriidae
ieeeAlleotepereopods, lo lacking dactyls (ig. 990) 2 2 ae i) eee) ee Some of pereopods 1-5 bearing dactyls (fig. 98a) ! . ........ aa)
2. Pereopod 2 smaller than and morphologically distinct from pereopod 1 (its orientation apparently reversed)... .. . . . . .EHohaustorius Pereopods 1-2 subequal in size and morphologically intersimilar. . . . 3
See Vera pedal pal) with four anticless 49 =. eee) 0s ne celica uel ran Maxillipedalspalp: wathathreevarticles 3s Sens G. ce se se 5
1 See explanation in text (p. 251).
285-135 O- 69-17
252
17.
18.
9:
20.
U.S. NATIONAL MUSEUM BULLETIN 271
Head with large rostrum (fig. 986), telson longer than broad and deeply
cleft (fig. 99f) . . . . . . . Zobracho 2 Head lacking rostrum, ialaon BE andes bem lone amd seam cleft (fig. 99q). Carangolia
Inner ramus of uropod2absent. ........ ... . .. Neohaustorius Inner ramus of uropod 2 present. ... . AME aN pul (0) Maxillipedal palp article 3 weakly penientate! or clavate, anion plate of maxilla 2 scarcely enlarged (fig. 98d) .... . 7 Maxillipedal palp article 3 strongly geniculate (Gee 98f), autier mate of maxilla 2 greatly enlarged (fig. 98e) ... . 8 Rami of uropod 1 expanded distally, sostanadansall ‘bowls: of dleoatke 3 forming a lobe overhanging urosome. .. . . . . Pseudohaustorius Rami of uropod 1 not expanded distally, posterodorsal border of pleonite 3 not forming lobe overhanging urosome. ... .. . Protohaustorius Pleonal epimeron 3 with large posteroventral tooth . . . Acanthohaustorius
Pleonal epimeron 3 rounded posteroventrally. Haustorius and Parahaustorius
Antenna 1 geniculate between articles 1 and 2 (fig. 99c) ...... . 10 Antenna 1 not geniculate, or occasionally between articles 2 and3. . 11 Gnathopod 2 lacking dactyl (fig. 1002)... ... .. . .. Bathyporeia Gnathopod 2 bearing dactyl. .... . . . . . Amphiporeia Coxa 1 vestigial, less than one fourth as lone. AS /COKA Gi en ernie Coxa’ l not) vestigial’ at least half asilong as coxa: 3)... eee ie Coxa 2 vestigial, less than one fourth as long as coxa 3. . Urohaustorius Coxa 2 not vestigial, almost as large as coxa 3... . .. =. +. . Cardenio Coxai 2 obtusely, pointed below, (fies Oia). | 5 nas oe ee Coxa: 2 rounded or, quadrate below, (igs 100g) 593-7.) 9 | eo Telson/split nearly its fulllength) 97> 5 3. 2 ee Zobracho Telson split: one third its length: 3°00. <2 =. i.e. aye Briscillina Gnathopods chelate (fig. 100d) ............ . Platyischnopus Gnathopods subchelate or simple . ... . Pee ses ae La) Inner ramus of uropod 3 scale-like, half as fonee as een ramus (fig. 100m). Carangolia Inner ramus of uropod 3 subequal to outer. . . . Rime hi Lah MIU Mandibular palp large, borne on basal process (like ie. 1509), article 3 clavate, molar of medium size, ridged ... . . .. . Phoxocephalopsis Mandibular palp small, directly attached to mandibular body, article 3 linearsmolar large or, medium; smooth) 0) 2) a a eS Outersramus of uropodia umiarticulatem 0) een oe 6) Outer ramus of uropod 3 biarticulate . .. . Ae O) Accessory flagellum present, usually 2- cnouliaite: aanaerarl 5 shorter and of different structure than pereopod 4... . . . . . . . Pontoporeia
Accessory flagellum absent; pereopod 5 similar to Dereon 4, Zaramilla (Eusiridae)
Article 4 of pereopod 4 not expanded, article 2 of pereopod 5 not greatly
produced downward .. . . . . Urothoe Article 4 of pereopod 4 coronmledl Aericle 2 of neieapod 5 prem produced downward) suey Aieiek slot ica a, Wenner Mian wee ne gO ROUMOLLES
2 Zobracho J. L. Barnard (1961). Note that the basal article of the maxillipedal palp was not inked on the original drawing; the palp has four articles.
MARINE GAMMARIDEAN AMPHIPODA 253
Ayo
Rw [K H
A JU oe € i g h
Ficure 99.—Haustoriidae: a, Carangolia puliciformis J. L. Barnard (1961); b, Haustorius arenarius (Slabber) (Sars, 1895, pl. 46). Antenna 1:c, Bathyporeia guilliamsoniana (Bate) (Sars, 1895, pl. 43, as B. norvegica). Telson: d, Bathyporeia: e, Priscillina armata (Boeck) (Sars, 1895, pl. 42); f, Pontoporeia femorata Krgyer (Sars, 1895, pl. 41); g, Carangolia; h, Haustorius.
254 U.S. NATIONAL MUSEUM BULLETIN 271
Genera of Haustoriidae
Acanthohaustorius Bousfield
Acanthohaustorius Bousfield, 1965.
Type-species: A. millsi Bousfield, 1965 (original designation).
Antenna 1 not geniculate; mandibular palp not on basal process, molar ridged; outer plate of maxilla 2 greatly enlarged; maxillipedal palp 3-articulate, article 3 geniculate; coxae 1-2 of normal size, coxae 1-3 obtusely pointed below; gnathopod 1 simple, article 5 longer than 6, gnathopod 2 minutely chelate; pereopods lacking dactyls; uropod 3 slightly elongate, rami subequal; telson short, cleft nearly to base; pleonal epimeron 3 with spine-like posteroventral process (Parahaus- torius, Haustorius); lower lip lacking mandibular lobes. Species: 4, N.W. Atlantic, littoral.
Amphiporeia Shoemaker
Amphiporeia Shoemaker, 1929.
Type-species: A. lawrenciana Shoemaker, 1929 (original desig- nation).
Antenna 1 geniculate between articles 1 and 2; mandibular palp attached to basal process, molar ridged; outer plates of maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxae 1-2 of normal size, rounded-quadrate below; gnathopods subchelate, article 5 of gnath- opod 1 slightly longer than article 6; gnathopod 2 bearing article 7 (Bathyporeia); pereopods dactylate; inner ramus of uropod 3 half as long as outer; telson fully cleft. Species: 2, N.W. Atlantic, littoral.
Bathyporeia Lindstrom
Bathyporeia Lindstrom, 1855.—Stebbing, 1906. Thersites Bate, 1857a, 1857b (homonym, Mollusca).
Type-species: B. pilosa Lindstrom, 1855 (monotypy). See Sars, 1895.
Antenna 1 geniculate between articles 1 and 2; mandibular palp attached to basal process, molar ridged; outer plate of maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxae 1-2 of normal size, rounded-quadrate below; gnathopods simple, gnathopod 2 lacking article 7 (Amphiporeia), article 5 of gnathopod 1 longer than article 6; at least pereopods 1-2 dactylate; inner ramus of uropod 3 short, scale-like; telson fully cleft. Species: 11, boreal N. Atlantic, littoral.
MARINE GAMMARIDEAN AMPHIPODA 955
Ficure 100.—Haustoriidae: a, Platyischnopus herdmani Walker (Pillai, 1957). Gnathopod 1: b, Bathyporeia guilliamsoniana (Bate) (Sars, 1895, pl. 43, as B. norvegica); c, Pontoporeia femorata Krgyer (Sars, 1895, pl. 41); d, Platyischnopus mirabilis Stebbing (1888); ¢, Priscillina armata (Boeck) (Sars, 1895, pl. 42); f, Urothoe; g, Haustorius arenarius (Slabber) (Sars, 1895, pl. 46); h, Carangolia puliciformis J. L. Barnard (1961). Gnathopod 2: i, Bathyporeia; 7, Haustorius. Uropod 3: k, Carangolia; 1, Haustorius; m, Priscillina; n, Pontoporeia; 0, Urothoe.
256 U.S. NATIONAL MUSEUM BULLETIN 271
Carangolia J. L. Barnard Carangolia J. L. Barnard, 1961.
Type-species: C. mandibularis J. L. Barnard, 1961 (original designation).
Antenna 1 not geniculate; mandibular palp not on process, molar large, smooth, terminal in position; outer plate of maxilla 2 not en- larged, maxillipedal palp 4-articulate; gnathopods nearly simple, article 7 immensely elongate, articles 5 and 6 of gnathopod 1 subequal in length; pereopods dactylate (except ?pereopod 1); uropod 3 greatly reduced in size, rami very short, subcircular, inner shorter than outer; telson short, broad, minutely cleft. Species: 2, antiboreal, bathyal (610-1,861 m).
Cardenio Stebbing
Cardenio Stebbing, 1888; 1906.
Type-species: (C. paurodactylus Stebbing, 1888 (monotypy).
Antenna 1 not geniculate; mandibular palp not on basal process, molar ridged or toothed; outer plates of maxilla 2 not enlarged; maxillipedal palp 3-articulate; coxa 1 very small, coxae rounded- quadrate below; gnathopods simple, article 5 of gnathopod 1 longer than article 6; pereopods 3-5 dactylate, pereopods 1—2 apparently not dactylate; uropod 3 short, rami equal in length; telson long, deeply cleft; lower lip apparently lacking mandibular processes. Species: 1, subantarctic, littoral.
Eohaustorius J. L. Barnard Eohaustorius J. L. Barnard, 1957b.
Type-species: Haustorius washingtonianus Thorsteinson, 1941 (original designation).
Antenna 1 not geniculate; mandibular palp not on basal process, molar smooth; outer plates of maxilla 2 not enlarged; maxillipedal palp 3-articulate, article 3 not geniculate; coxae 1 and 2 much smaller than and partially hidden by coxae 3-4; gnathopod 1 simple, article 5 longer than article 6, gnathopod 2 minutely chelate; pereopod 2 similar to pereopod 3 in orientation, and smaller than pereopod 1; uropod 3 short, rami equal in length; telsonic lobes completely detached basally; outer lobes of lower lip lacking mandibular processes. Species: 4, boreal Pacific, littoral.
Haustorius Muller
Haustorius Muller, 1775.—Stebbing, 1906. ?Lepidactylis Say, 1818 (see Bousfield, 1965). Pterygocerus Latreille, 1825.
MARINE GAMMARIDEAN AMPHIPODA 257
Bellia Bate, 1851 (homonym, Decapoda). Sulcator Bate, 1854.
Type-species: Oniscus arenarius Slabber, 1769 (monotypy). See Sars, 1895.
Antenna 1 not geniculate; mandibular palp not on basal process, molar ridged; outer plate of maxilla 2 greatly enlarged; maxillipedal palp 3-articulate, article 3 geniculate; coxae 1—2 of normal size, coxae 1-3 obtusely pointed below; gnathopod 1 simple, article 5 longer than article 6, gnathopod 2 minutely chelate; pereopods lacking dactyls; uropod 3 short, rami equal in length; telson short, partially cleft; pereopods 4-5 urth articles 4-5 three fourths as broad as article 2 of pereopod 5 (Zobracho and Priscillina); pleonal epimeron 3 rounded posteroventrally (Acanthohaustorius); outer lobes of lower lip lacking mandibular processes. Species: 2, N. Atlantic, littoral.
Neohaustorius Bousfield
Neohaustorius Bousfield, 1965.
Type-species: N. schmitzi Bousfield, 1965 (original designation) .
Antenna 1 not geniculate; mandibular palp not on basal process, molar triturative; outer plate of maxilla 2 greatly enlarged; maxil- lipedal palp 3-articulate, article 3 geniculate; coxae 1-2 of normal size, to some extent coxae 1-2 obtusely pointed below; gnathopod 1 simple, article 5 longer than 6, gnathopod 2 minutely chelate; pereo- pods lacking dactyls; uropod 3 short, rami subequal; telson short, uncleft; inner ramus of uropod 2 absent; lower lip lacking mandibular lobes. Species: 2, N.W. Atlantic, littoral.
Parahaustorius Bousfield
Parahaustorius Bousfield, 1965.
Type-species: P. longimerus Bousfield, 1965 (original designation).
Antenna 1 not geniculate; mandibular palp not on basal process, molar triturative; outer plate of maxilla 2 greatly enlarged; maxil- lipedal palp 3-articulate, article 3 geniculate; coxae 1-2 of normal size, coxae 1-2 obtusely pointed below; gnathopod 1 simple, article 5 longer than 6, gnathopod 2 minutely chelate; pereopods lacking dactyls; uropod 3 slightly elongate, rami subequal; telson short, cleft about halfway; pleonal epimeron 3 rounded posteroventrally (Acanthohaustorius); lower lip lacking mandibular lobes. Species: 3, N.W. Atlantic, littoral.
Phoxocephalopsis Schellenberg
Phoxocephalopsis Schellenberg, 1931. Haustoriella K. H. Barnard, 1931.
258 U.S. NATIONAL MUSEUM BULLETIN 271
Type-species: P. zvmmeri Schellenberg, 1931 (monotypy).
Antenna 1 not geniculate; mandibular palp on basal process, large, article 3 clavate (Urothoe, Urothoides), molar ridged; outer plates of maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxa 1 nearly obtusely pointed below, others rounded-quadrate; gnathopods sub- chelate or nearly simple, article 5 of gnathopod 1 longer than article 6; pereopods dactylate; uropod 3 short, rami subfoliaceous, subequal in length; telson short, deeply cleft. Species: 2, subantarctic-anti- boreal, littoral.
Platyischnopus Stebbing
Platyischnopus Stebbing, 1888; 1906.
Type-species: P. mirabilis Stebbing, 1888 (monotypy).
Antenna '1 not geniculate; mandibular palp not on basal process, molar “ridged,’”’ smooth or absent; outer plates of maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxae of medium length, rounded-quadrate below; gnathopods chelate, article 5 of gnathopod 1 longer or shorter than article 6; pereopods dactylate; uropod 3 (outer ramus only) immensely elongate, inner ramus short, scale-like; telson partially cleft or emarginate; head with “‘shark-nose’’ rostrum. Species: 6, warm N. temperate through tropics to antiboreal, littoral.
Pontoporeia Kr¢yer
Pontoporeia Kr¢yer, 1842.—Stebbing, 1906.
Type-species: P. femorata Kroyer, 1842 (monotypy). See Sars, 1895.
Antenna 1 not geniculate; mandibular palp not on basal process, molar ridged; outer plate of maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxae 1-2 of normal size, rounded-quadrate below; gnathopod 1 subchelate, articles 5-6 subequal in length, gnathopod 2 slightly chelate; at least pereopods 1, 2, 5 dactylate; rami of uropod 3 equal in length; telson cleft halfway or more. Species: 5; 1 Caspian, 4 subarctic-boreal and glacial relicts, littoral.
Priscillina Stebbing
Priscilla Boeck, 1871 (homonym, Coleoptera). Priscillina Stebbing, 1888 (new name); 1906.
Type-species: Pontoporeia armata Boeck, 1861 (monotypy). See Sars, 1895.
Antenna 1 not geniculate; mandibular palp not on basal process, molar ridged; outer plate of maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxae 1-2 of normal size, coxae 1-4 obtusely pointed below; gnathopods subchelate, articles 5-6 of gnathopod 1 subequal
MARINE GAMMARIDEAN AMPHIPODA 259
in length; some pereopods dactylate; uropod 3 short, inner ramus scale-like; telson cleft one third of its length; articles 4-5 of pereopods 4-6 one third as broad as article 2 of pereopod 5 (Haustorvus). Species: 1, arctic-subarctic, littoral.
Protohaustorius Bousfield
Protohaustorius Bousfield, 1965.
Type-species: P. deichmannae Bousfield, 1965 (original designa- tion).
Antenna 1 semigeniculate; mandibular palp not on basal process, molar triturative; outer plate of maxilla 2 not enlarged; maxillipedal palp 3-articulate, article 3 not geniculate; coxae 1-2 of normal size, not obtusely pointed below; gnathopod 1 simple, article 5 longer than 6, gnathopod 2 minutely chelate; pereopods lacking dactyls; uropod 3 short, rami subequal; telson short, slightly emarginate; rami of uro- pod 1 not distally expanded; posterodorsal border of pleonite 3 not form- ing lobe overhanging urosome (Pseudohaustorius); lower lip. lacking mandibular lobes. Species: 2, N.W. Atlantic, littoral.
Pseudohaustorius Bousfield Pseudohaustorius Bousfield, 1965.
Type-species: P. caroliniensis Bousfield, 1965 (original designa- tion).
Antenna 1 not geniculate; mandibular palp not on basal process, molar ridged; outer plate of maxilla 2 not enlarged; maxillipedal palp 3-articulate, article 3 clavate; coxae 1-2 of normal size, 1-3 subobtusely pointed below; gnathopod 1 simple, article 5 longer than 6, gnathopod 2 minutely chelate; pereopods lacking dactyls; uropod 3 short, rami subequal; telson short, slightly emarginate; rami of uropod 1 distally expanded; posterodorsal border of pleonite 3 forming lobe overhanging urosome (Protohaustorius); lower lvp lack- ing mandibular lobes. Species: 3, N.W. Atlantic and Gulf of Mexico, littoral.
Urohaustorius Sheard Urohaustorius Sheard, 1936b.
Type-species: U. halet Sheard, 1936b (original designation).
Antenna 1 not geniculate; mandibular palp apparently not on basal process, molar large and projecting but apparently lacking ridges or teeth; outer plates of maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxae 1 and 2 vestigial; gnathopod 1 simple, article 5 longer than article 6, gnathopod 2 minutely chelate; pereopods dactylate but some dactyls resembling spines; rami of uropod 3 sub- equal in length; telson short, entire. Species: 2, S. Australia, littoral.
260 U.S. NATIONAL MUSEUM BULLETIN 271
Urothoe Dana
Urothoe Dana, 1852b.—Stebbing, 1906. Egidia Costa, 1853c, 1857.
Type-species: Urothoe irrostratus Dana, 1853 (indicated by elimi- nation and ipso-facto monotypy of Boeck, 1876, but firmly selected by Stebbing, 1891). Type-species remains dubious and genus herein based provisionally on U. elegans Bate, 1857a (Sars, 1895, as U. norvegica Boeck, 1861).
Antenna 1 not geniculate; mandibular palp not on basal process, small, article 3 linear (Phoxocephalopsis), molar large, poorly ridged; outer plates of maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxae 1-2 of normal size, rounded-quadrate below; gnathopods sub- chelate or simple, article 5 of gnathopod 1 longer than article 6; pereopods dactylate; rami of uropod 3 equal in length; telson fully cleft; article 4 of pereopod 4 not strongly expanded, article 2 of pereopod 5 not greatly produced downward (Urothoides). Species: 22, cosmo- politan, littoral to abyssal.
Urothoides Stebbing
Urothoides Stebbing, 1891; 1906.
Type-species: Urothoe lachneessa Stebbing, 1888 (monotypy).
Antenna 1 not geniculate; mandibular palp not on basal process, small, article 3 linear (Phoxocephalopsis), molar large, poorly ridged; outer plates of maxilla 2 not enlarged; maxillipedal palp 4-articulate; coxae 1-2 of normal size, rounded-quadrate below; gnathopods subchelate, article 5 of gnathopod 1 longer than article 6; pereopods dactylate; uropod 3 of medium length, inner ramus half as long as outer; telson deeply cleft; article 4 of pereopod 4 expanded, article 2 of pereopod 5 greatly produced downward (Urothoe). See other remarks by K. H. Barnard, 1932. Species: 1+ ?1, subantarctic, bathyal (236-720 m).
Zobracho J. L. Barnard
Zobracho J. L. Barnard, 1961.
Type-species: Z. canguro J. U. Barnard, 1961 (original designation).
Antenna 1 not geniculate; mandibular palp not on basal process, molar small and slightly toothed; outer plates of maxilla 2 not en- larged; maxillipedal palp 4-articulate; coxa 1 strongly reduced in size, coxae 1—2 obtusely pointed below; gnathopods subchelate, article 5 of gnathopod 1 longer than article 6; some pereopods presumably dactylate but dactyls resembling spines; uropod 3 short, inner ramus half as long as outer; telson long, nearly fully cleft; articles 4-5 of pereopods 4-5 one third as broad as article 2 on pereopod 5 (Haustorius) ; head with large rostrum. Species: 1, Australia, bathyal (875 m).
MARINE GAMMARIDEAN AMPHIPODA 261
Hyalellidae and Hyalidae
[see Superfamily Talitroidea]
Hyperiopsidae Ficures 101, 102
Diacnosis.—Accessory flagellum well developed; article 4 of pereopods 1-2 greatly elongate; palp of first maxilla claviform, slightly geniculate; gnathopods feeble, simple. See Vitjazianidae, Astyridae, Stilipedidae, Synopiidae.
Description.—Accessory flagellum well developed, its articles very elongate; body variable, smooth or carinate, head globular (resembling hyperiids), subglobular or nearly normal, but rarely bearing distinct lateral lobes; coxae variable, first four small and uniform or coxa 4 enlarged and covering anterior coxae; upper lip with minute incision; mandibular body very short, poorly toothed, molar very small, weakly triturative or not, palp immense; lower lip without inner lobes, outer lobes often alate; palp of maxilla 1 biartic- ulate, article 2 slightly or greatly bent medially (at least on one side) ; maxilla 2 and maxilliped normal; gnathopods feeble or nearly simple; uropod 3 biramous; telson cleft or entire, with apical ornamentation.
It is difficult, at first glance, to justify a familial association of the two hyperiopsid genera. Hyperiopsis resembles an hyperiid amphipod. It has a globular head, smooth body and short uniform coxae. Parar- gissa has the normal appearance of a gammaridean and although the head is subglobular it is rather small and may have lateral lobes; its fourth coxa is enlarged and covers the smaller coxae 1-3. Nevertheless, Hyperiopsis and Parargissa share some features that are unusual in the Gammaridea: (1) the peculiar first maxillary palps of which the second articles are claviform and slightly bent on at least one of the maxillae; (2) the very short body of the mandible with a nearly untoothed cutting edge and a very small, poorly developed molar; (3) the great elongation of article 4 on pereopods 1 and 2, a condition rarely occurring in other families (cf. Ampeliscidae); (4) the elonga- tion of article 1 of the first antennal flagellum (not uncommon in some other families, viz. Lysianassidae, Synopiidae).
RELATIONSHIP.—The combination of the elongate article 4 of pere- opods 1 and 2 with the bent first maxillary palp separates this family from the following similar families: Astyridae, Synopiidae, Pardalis- cidae, and Vitjazianidae.
The Ampeliscidae resemble hyperiopsids because of the elongate fourth articles of pereopods 1-2, but ampeliscids lack accessory flagella and have well-developed mandibular molars.
U.S. NATIONAL MUSEUM BULLETIN 271
h 1
Figure 101.—Hyperiopsidae: a, Hyperiopsis voringt Sars (1885); b, Parargissa galatheae J. L. Barnard (1961). Accessory flagellum: c, Hyperiopsis laticarpa Birstein and Vino- gradov (1955). Lower lip: d, Parargissa arquata (Birstein and Vinogradov, 1955); e, Hyperiopsis. Mandible: f, Hyperiopsis gibbosa Pirlot (1934); g, Parargissa arquata.
Telson: h, Parargissa arquata; 1, Hyperiopsis gibbosa.
MARINE GAMMARIDEAN AMPHIPODA 263 Alezandrella, in the Stilipedidae, has a bent first maxillary palp
but hyperiopsids have well-developed accessory flagella and mandi- bular molars.
Key to the Genera of Hyperiopsidae
1. Coxae 1-4 small, similar in size and shape (fig. 101a) . . . . MHyperiopsis Coxa 4 greatly enlarged, concealing the small first three coxae (fig. 1016) Parargissa
Figure 102.—Hyperiopsidae: Maxilla 1: a,b, Parargissa arquata (Birstein and Vinogradov, 1955); c, Hyperiopsis gibbosa Pirlot (1934); d, other palp of Hyperiopsis. Maxilla 2: e, Hyperiopsis. Pereopods 1 and 2: f, Hyperiopsis laticarpa Birstein and Vinogradov (1955); g, Parargissa arquata. Maxilliped: h, Hypertopsis.
Genera of Hyperiop sidae
Hyperiopsis Sars Hyperiopsis Sars, 1885.—Stebbing, 1906.
Type-species: H. voringvi Sars, 1885 (monotypy). See Stephensen ; 1934. Species: 7, cosmopelagic, abyssal and bathyal.
264. U.S. NATIONAL MUSEUM BULLETIN 271
Parargissa Chevreux
Parargissa Chevreux, 1908¢e.—Birstein and Vinogradov, 1958. Protohyperiopsis Birstein and Vinogradov, 1955.
Type-species: P. nasuta Chevreux, 1908c (original designation). Species: 5, cosmopelagic, abyssal.
Isaeidae
[including Photidae] Ficures 103-106
Diacnosis.—Accessory flagellum variable in length or absent; telson entire, short, fleshy; coxa 5 not excavate posteriorly; pereopods 1-2 olandular; wropod 3 rarely projecting beyond uropods 1 and 2, rami (at least one) as long as or longer than peduncle, occasionally uropod 3 uniramous. See Aoridae, Corophiidae, Ischyroceridae, Calliopidae, Laphystiopsidae, Oedicerotidae, Pleustidae.
Description.—Accessory flagellum varying from absent to long and multiarticulate; body smooth, rostrum absent; coxae rounded or quadrate ventrally, long or short, fourth not excavate posteriorly; mouthparts basic; gnathopods usually powerfully subchelate or extremely setose; uropod 3 short, rami as long as or longer than pe- duncle, not uncinate, occasionally inner ramus reduced or absent; telson entire, short, fleshy, nearly circular or square, occasionally falsely cleft.
ReELATIONSHIP.—The Isaeidae do not warrant distinction from the Photidae on the basis of prehensile or partially subchelate pereopods and so they must be joined, Isaeidae (Isaeinae Dana, 1853) taking precedence. This move is unfortunate because of the long use of the name Photidae for a diversity of genera and the relatively infrequent use of the name Isaeidae.
The Laphystiopsidae have a nontriturative mandibular molar and are otherwise like calliopiids and pleustids.
See Calliopiidae for calliopiid-pleustid differences.
The Gammaridae usually have a cleft telson; when entire, the telson is not circular or subcircular and coxa 4 is strongly excavate posteriorly ; some Gammaridae have a small, square, unexcavate coxa 4 but in those species the telson is cleft; Gammaridae lack glandular pereopods, do not build tubes, usually have uropod 3 projecting beyond uropods 1 and 2 and the rami are enlarged, either by dorsoventral depression, elongation, or foliaceousness.
The Aoridae have gnathopod 1 larger than gnathopod 2. Linkage is shown to that family by Aorcho, in the Aoridae, having gnathopods 1
MARINE GAMMARIDEAN AMPHIPODA 265
y SK
aE ey \\ \ > Of fe IT
S\ Lo) c) ye rN SI w Ke,
Ml Y s ps L> S ‘Ss aS
Ficure 103.—Isaeidae: a, Gammaropsis (=Eurystheus) maculata (Johnson) (Sars, 1895, pl. 198, as G. erythrophthalma); 6, Photis reinhardi Krg¢yer (Sars, 1895, pl. 202); c, Megamphopus cornutus Norman (Sars, 1895, pl. 200).
J
Ficure 104.—Isaeidae and Aoridae: a, Isaea montagui Milne Edwards (Chevreux and Fage, 1925); b, Leptocheirus pilosus Zaddach (Sars, 1895, pl. 197) [=Aoridae]. Pereopod 1: c, Ampelisciphotis tridens Pirlot (1938). Telson: d, Microprotopus maculatus Norman (Sars, 1895, pl. 201); ¢, Photis reinhardi Krdyer (Sars, 1895, pl. 202). Uropod 3: f, Micro- protopus; g, Leptocheirus; h, Photis;1, Megamphopus cornutus Norman (Sars, 1895, pl. 200); J, Cheiriphotis megacheles (Giles) (J. L. Barnard, 1962a).
MARINE GAMMARIDEAN AMPHIPODA 267
and 2 equal in size. One must also examine Neomegamphopus, now assigned to Aoridae, as an intermediary between Isaeidae and Aoridae. All Isaeidae with male gnathopod 1 having secondary sexual modi- fications have been removed to the Aoridae.
The Ischyroceridae have the rami of uropod 3 shorter than the elongate peduncle and usually have one or more distinct hooks or an uncinate condition on the outer ramus.
The Ampithoidae have an even more stunted third uropod than do isaeids, the outer ramus of which usually bears hooks; the ampithoid lower lip has notched or medially excavate outer lobes.
The Corophiidae have a depressed urosome although this is not clearly distinct in some intergrading genera; all but a few corophiids have aberrant third uropods lacking one ramus, although a few Isaeidae also demonstrate this; some corophiids and isaeids are not clearly distinct and the taxonomist must use both family keys for doubtful genera.
Camacho and Parunciola in the Corophiidae and Aorcho, now assigned to the Aoridae, show the difficulty in clearly distinguishing among isaeids, aorids, and corophiids. These genera, with their moderately or fully depressed urosomes, equal-sized gnathopods, biramous third uropod [or uniramous third uropod with long ramus], and circular or quadrate telsons, lacking hooks, could be assigned to any of the three families.
The loss of one ramus of uropod 3 in five isaeid genera confounds technical distinction of Isaeidae from several other, distantly re- lated families having uncleft telsons. Cressidae, Stenothoidae, and Thaumatelsonidae are distinguished from isaeids by their small first coxae. Eophliantidae have cylindrical bodies and Phliantidae, Talitroidea, and Dogielinotidae lack mandibular palps. Sebidae have chelate gnathopods and obsolescent mandibular molars and Pagetinidae have evanescent mandibular molars and poorly developed outer plates of the maxillipeds.
Nomenclatural Changes in Isaeidae
Amphideutopus J. L. Barnard (1959a) is removed to the Aoridae.
Audulla Chevreux (1901a) is assigned in the Isaeidae as subgenus to Gammaropsis.
Bathyphotis Stephensen (1944b) is removed to the Ischyroceridae.
Bonnierella Chevreux (1900) is removed to the Ischyroceridae but is also assigned in the Isaeidae to Megamphopus.
Eurystheus dentatus Holmes (1908) is removed to Protomedeia (Cheirimedeia).
Leptocheirus Zaddach (1844) is removed to the Aoridae.
285-135 O - 69 - 18
268 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 105.—Isaeidae: a, Chetriphotis megacheles (Giles) (J. L. Barnard, 1962a); b, Chevalia aviculae Walker (J. L. Barnard, 1962a). Head: c, Ampelisciphotis tridens Pirlot (1938). Accessory flagella: d, Gammaropsis (= Eurystheus) maculata (Johnston) (Sars, 1895, pl. 199, as G. melanops); e, Microprotopus maculatus Norman (Sars, 1895, pl. 201).
MARINE GAMMARIDEAN AMPHIPODA 269
Neomegamphopus Shoemaker (1942) is removed to the Aoridae. Pseudeurystheus Schellenberg (1931) is assigned in the Isaeidae as
10.
11.
18.
subgenus to Gammaropsis.
Key to the Genera of Isaeidae
Mropod auuminramousndienMO4p) ie eo ea en Me. 2 Uropod 3 biramous (figs. 104g-j). . . . EAS Ua ONG Lateral cephalic lobes pedunculate, article 6 oi Beopode 1-2 elongate (igss 1O5e 104) ine . . . . . Ampelisciphotis Lateral cephalic lobes and Artiele 6 of percopods 1-2 notelongate. ... 3 Gnathopod 1 simple (fig. 106h). . ....... . . . . Kuphocheira Cnathopod lisubbchelateq sks. see eu oa Ne cle 3) aa eae Matbennamtewithvaccessony Hagellumy. 402). were ae eee eu yiueery =) iO Antenna 1 lacking accessory flagellum... . . . . . . Microphotis Coxae 1-4 similar to each other in size and shape (fe. 103a) . Microprotopus Coxae 1—4 of varying shapes and sizes (fig. 105a) . . . . . Cheiriphotis Uropod 3 with one ey shortened ramus (less than 60% as long as outer) (fig 04h) 2 xo: Sa ME RS 1 Sc SU CO a Uropod 3 with subequal rami Ge 104i) SES CNG sigh areal Sam Oa Taam gaa sk Sees AL) Gnathopods 1-2 simple. . . . . . .. . . . Haplocheira Gnathopod 1 subchelate, pneehoped 2 ‘simple . . Leptocheirus (Aoridae) Gnathopods 1-2 subchelate. . .. . Sah Ace ta eat: Antenna 1 lacking accessory dopelum or bearing a otra. anticls or scale. Photis
Antenna 1 with multiarticulate accessory flagellum. ......... 9 Uropod 3 scale-like, peduncle plate-like (fig. 1047). . . . . . Cheiriphotis Uropod 8 cylindrical (fig. 104g) . . . Protomedeia (subgenus Cheirimedeia) Gnathopod 1 complexly subchelate (fig. 64a), gnathopod 2 with well- developedpalm. .. . BN tener eae tenn os ee (Aoridae) These characters not combined. Aslan eval BAe tae aie aa eens eal FL Article 3 of antenna 1 as long as article 1 or lone: Cent spi vatantec! ioe reheat cue Leet Article 3iof antenna 1) shorter than article) 2.0.0.4... 3 .-,-,2h6 Accessory flagellum of antenna 1 absent... . . . . . Podoceropsis Accessory flagellum of antennal present. ........ ay eta eenet alte Accessory flagellum composed of ascale ........ - Kermystheus Accessory flagellum composed of one or more long articles. . ..... 14
Accessory flagellum composed of one or two articles. Megamphopus (Segamphopus and Audulla), Bonnierella !
Accessory flagellum composed of three or more articles . . .... . .15 Pereopods 1-5 slightly subchelate (fig. 104a)... . Noa ea SAR Pereopods 1-5simple .. . . . Gammaropsis (eubeeaue Pseudeurystheus) Accessory flagellum absent. . ... . . . . . Dryopoides? and Goesia Accessory flagellum present. . .. . ' aaa uch Li Gnathopod 2 subchelate, coxae 1 and ‘fils aaa of Donraalle balanced
dimensions. . . AU semi lye gal lke) Gnathopod 2 simple Ge 1061), « coxa aL very ral aad cov percd by enlarged
COXAw2 a iene HAY aie a cota at . . . Leptocheirus (Aoridae) Pleonites 4 and 5 apace OH Sek va al eae Protomedeia Pleonites 4 and 5 coalesced (fig. 1058) . Beet ss ae Te ecm a oe ee CHOVEIIB
1 See this genus in the Ischyroceridac. 2 See male, Stebbing (1910) and Aoridae (p. 153).
270 U.S. NATIONAL MUSEUM BULLETIN 271
n ay
m
Ficure 106.—Isaeidae and Aoridae: Mouthparts, Photis reinhardi Kréyer (Sars, 1895, pl. 202): a, upper lip; b, mandible; c, lower lip; d,e, maxillae 1, 2: f, maxilliped. Max- illiped: g, Isaea montagui Milne Edwards (Chevreux and Fage, 1925). Gnathopod 1: h, Kuphocheira setimanus K.. H. Barnard (1932); 7, Photis tenuicornis (Sars, 1895, pl. 203); j, Leptocheirus pilosus Zaddach (Sars, 1895, pl. 197) [=Aoridae]; k, Megamphopus cornutus Norman (Sars, 1895, pl. 200). Gnathopod 2: /, Leptocheirus; m, Megamphopus; n,
Photts tenuicornis, female; 0, Kuphocheira; p, Photis tenuicornis, male.
MARINE GAMMARIDEAN AMPHIPODA o71 Genera of Isaeidae
Ampelisciphotis Pirlot Ampelisciphotis Pirlot, 1938.
Type-species: A. tridens Pirlot, 1938 (original designation).
Lateral cephalic lobes and article 6 of pereopods 1—2 elongate (com- bining character); accessory flagellum absent; article 3 of antenna 1 equal to or slightly longer than article 1; gnathopods subchelate; uropod 3 uniramous, ramus shorter than peduncle. Species: 1, Indonesia, littoral.
Cheiriphotis Walker
Cheiriphotis Walker, 1904.
Type-species: Melita megacheles Giles, 1885 (monotypy). See J. L. Barnard, 1962a.
Accessory flagellum 3- or more articulate; article 3 of antenna 1 shorter than article 1; gnathopods subchelate; wropod 3 scale-like, peduncle plate-like (Cheirimedeia), outer ramus equal to or shorter than peduncle, inner ramus half as long as outer or absent; cozrae 1-4 of varying sizes and shapes (Microprotopus). Species: 2, tropical Indo-Pacific, littoral.
Chevalia Walker
Chevalia Walker, 1904. Neophotis Stout, 1913.
Type-species: C. aviculae Walker, 1904 (monotypy). See J. L. Barnard, 1962a.
Accessory flagellum 2-articulate; article 3 of antenna 1 slightly shorter than article 1; gnathopods subchelate, gnathopod 1 poorly; uropod 3 short, biramous, rami subequal to each other, equal to pe- duncle in length; pleonites 4 and 5 coalesced. Species: 1, tropical Indo-Pacific, littoral.
Gammaropsis Liljeborg, new synonymy
(Gammaropsis) Liljeborg, 1855 [subgenus].
Eurystheus Bate, 1857a.—Stebbing, 1906.
Paranaenia Chilton, 1884.
Maeroides Walker, 1898.
Audulla Chevreux, 1901a [a valid subgenus]. Fimbriella Stout, 1913.
Pseudeurystheus Schellenberg, 1931 [a valid subgenus].
Type-species: Gammarus maculatus Johnston, 1827 (selected by Boeck, 1876, with subsequent synonymy). See Sars, 1895.
Accessory flagellum 3- or more articulate; article 3 of antenna 1 equal to or longer than article 1; gnathopods subchelate; uropod 3
DD U.S. NATIONAL MUSEUM BULLETIN 271
biramous, rami equal to each other, variable in length, generally equal to or longer than peduncle. Species: 53, cosmopolitan, littoral
to bathyal. Subgenus Audulla Chevreux (190la): Like Gammaropsis but flagellum of antenna 2 stout. Species: 1, tropical Indian Ocean,
littoral.
Subgenus Pseudeurystheus Schellenberg (1931): Like Gammaropsis but article 5 of male gnathopod 2 at least 1.6 times as long as article 6. Species: 1, subantarctic, littoral to 310 m.
Goesia Boeck
Goesia Boeck, 1871.—Stebbing, 1906.
Type-species: Autonoe depressa Goés, 1866 (monotypy). See Stephensen, 1942, 1944a.
Accessory flagellum absent; article 3 of antenna 1 shorter than ar- ticle 1; gnathopods subchelate, gnathopod 2 poorly; uropod 3 short, biramous, rami equal to each other, subequal to peduncle. Species: 1, arctic-subarctic, littoral.
Haplocheira Haswell
Haplocheira Haswell, 1880b.—Stebbing, 1906.
Type-species: Gammarus barbimanus Thomson, 1879 (monotypy and subsequent synonymy). See Stebbing, 1888.
Accessory flagellum 2- or more articulate; article 3 of antenna 1 shorter than article 1; gnathopods simple; uropod 3 biramous, outer ramus subequal to peduncle, inner ramus very short. Species: 3, subantarctic-antiboreal, littoral to bathyal (457 m).
Isaea Milne Edwards
Isaea Milne Edwards, 1830.—Stebbing, 1906.
Type-species: J. montagui Milne Edwards, 1830 (monotypy). See Chevreux and Fage, 1925.
Accessory flagellum 3- or more articulate; article 3 of antenna 1 equal to article 1 in length; gnathopods subchelate; uropod 3 biramous, rami equal to each other, slightly longer than peduncle; pereopods 1-6 slightly prehensile or subchelate, or article 6 widened distally. Species: 4, N. Atlantic, littoral, some species inhabit decapods.
Kermystheus J. L. Barnard
Kermystheus J. L. Barnard, 1962a.
Type-species: Podoceropsis kermadeci Stebbing, 1888 (original designation).
MARINE GAMMARIDEAN AMPHIPODA DHS)
Accessory flagellum a short, 1-articulate scale; article 3 of antenna 1 equal to or longer than article 1; gnathopods subchelate, gnathopod 1 occasionally poorly so; uropod 3 apparently biramous, rami sub- equal to each other and to peduncle. Species: 2, Kerguelen Island and California, littoral to bathyal (1,152 m).
Kuphocheira K. H. Barnard Kuphocheira K. H. Barnard, 1931; 19382.
Type-species: KK. setimanus K. H. Barnard, 1931 (original des- ignation); 1932.
Accessory flagellum absent; [article 3 of antenna 1 undescribed]; enathopods simple, dactyl of gnathopod 2 reduced in size and setose; uropod 3 uniramous, ramus shorter than peduncle. Species: 1, 5. Orkney Islands, littoral.
Megamphopus Norman
Megamphopus Norman, 1869a.—Stebbing, 1906.
(Segamphopus) J. L. Barnard, 1962a [subgenus].
[Bonnierella Chevreux, 1900], as possible synonym, but also assigned to Ischyro- ceridae.
Type-species: MM. cornutws Norman, 1869a (original designation). See Sars, 1895.
Accessory flagellum 1- or 2-articulate, article 1 elongate; article 3 of antenna 1 equal to or longer than article 1; gnathopods subchelate, occasionally poorly; uropod 3 biramous, rami subequal to each other, longer than peduncle. Species: 15, biboreal, littoral to bathyal (385 m).
Subgenus Bonnierella Chevreux (1900): Like Megamphopus but article 2 of pereopods 3-5 with parallel edges. This subgenus also is assigned as a genus to the Ischyroceridae. Species: 5, bathyal to abyssal.
Subgenus Segamphopus J. L. Barnard (1962a): Like Megamphopus but article 5 of gnathopod 2 at least 1.6 times as long as article 6. Species: 1, S. Georgia Island, littoral.
Microphotis Ruffo Microphotis Ruffo, 1952.
Type-species: MM. blachei Ruffo, 1952 (monotypy).
Accessory flagellum absent; article 3 of antenna 1 equal to article 1; gnathopods subchelate; uropod 3 uniramous, ramus shorter than peduncle. Species: 1, S. China Sea, littoral.
274. U.S. NATIONAL MUSEUM BULLETIN 271
Microprotopus Norman
?Dercothoe Dana, 1852b. Microprotopus Norman, 1867b.—Stebbing, 1906. Orthopalame Hoek, 1879.
Type-species: MM. maculatus Norman, 1867b (monotypy). See Sars, 1895.
Accessory flagellum 2-articulate, article 1 elongate; article 3 of antenna 1 shorter than article 1; gnathopods subchelate; uropod 3 uniramous, ramus slightly longer than peduncle. Species: 2, boreal to subtropical E. Atlantic and Mediterranean, littoral.
Photis Kr¢éyer
Photis Kroyer, 1842.—Stebbing, 1906. Eiscladus Bate and Westwood, 1863. Also Heiscladus, lapsus by Norman, 1869a.
Type-species: P. reinhardi Kroyer, 1842 (monotypy). See Sars, 1895.
Accessory flagellum absent or formed of a vestigial article or scale; article 3 of antenna 1 equal to or longer than article 1; gnathopods subchelate; uropod 3 biramous, outer ramus equal to or shorter than peduncle, inner ramus much shorter than outer ramus. Species: 41, cosmopolitan, littoral (rare to abyssal).
Podoceropsis Boeck
Podoceropsis Boeck, 1861.—Stebbing, 1906. Naenia Bate, 1862 (homonym, Lepidoptera). Xenoclea Boeck, 1871. Type-species: P. sophia Boeck, 1861 (monotypy). See Sars, 1895. Accessory flagellum absent; article 3 of antenna 1 equal to or longer than article 1; gnathopods subchelate; uropod 3 biramous, rami equal to each other, equal to peduncle in length. Species: 6, Atlantic, littoral to abyssal.
Protomedeia Kr¢yer ©
Protomedeia Krgyer, 1842.—Stebbing, 1906. Cheirimedeia J. L. Barnard, 1962a [valid subgenus]. Type-species: P. fasciata Kroyer, 1842 (monotypy). See Sars, 1895. Accessory flagellum 3- or more articulate; article 3 of antenna 1 shorter than article 1; gnathopods subchelate; uropod 3 biramous, rami subequal to each other, subequal to peduncle. Species: 14, subarctic, boreal Pacific (one Atlantic), littoral (?to bathyal). Subgenus Cheirimedeia J. L. Barnard (1962a): Like Protomedeia but inner ramus of uropod 3 half as long as outer ramus. Species: 2, boreal Pacific.
MARINE GAMMARIDEAN AMPHIPODA 275
Ischyroceridae
FicurEs 107, 108
Diacnosis.—Accessory flagellum variable; telson short, entire, fleshy; coxa 4 not excavate posteriorly except in one genus; pereopods glandular; uropod 3 rarely projecting beyond uropods 1 and 2, peduncle elongate, rami styliform, shorter than peduncle, outer ramus usually uncinate, either with hooked tip, or hooked spine at tip, or denticles. See Isaeidae, Ampithoidae, Calliopiidae, Pleustidae, Corophiidae.
Description.—Accessory flagellum varying from uniarticulate to multiarticulate; body smooth; rostrum absent; coxae rounded or quadrate below, varying from long to short, fourth not excavate posteriorly (with exception in Bathyphotis); mouthparts basic, possibly epistome always acutely produced anteriorly; gnathopods usually powerfully subchelate; structure of uropod 3 as in diagnosis but merging with conditions typical of Isaeidae by tendencies to elongation of rami and lack of uncination; telson entire, subcircular or nearly square, thick.
RELATIONSHIP.—The Isaeidae (= Photidae) differ from the Ischy- roceridae by the lack of uncination on the outer ramus of uropod 3, the longer rami and shortened peduncles, but some species now assigned to Isaeidae bear third uropods merging with the Ischyro- ceridae.
Calliopiids and pleustids have a nonfleshy telson and a short peduncle of uropod 3.
Ericthonius in the Corophiidae has an ischyrocerid uropod 3 without an inner ramus. Dryopoides and Paradryope in the Aoridae have third uropods which technically are ischyrocerid. Parapherusa in the Gammaridae lacks pereopodal glands but has uropod 3 similar to that of ischyrocerids.
Key to the Genera of Ischyroceridae
1. Coxae minute, not touching. .. . .. . . . Bogenfelsia
Coxae small or large, overlapping or banal foocite AS esis a Em Ie oa MESA hte Gnathopod 1 distinctly larger than 2
Dryopoides (Aoridae) and Paradryope (Aoridae)
Gnathopod 1 smaller than 2 or gnathopods subequal to each other . . . 3
3. Second articles of pereopods 3-5 narrowly linear, palp article 4 of maxilliped
claw-like, longer than article3.... . . . . . . Bonnierella
Second articles of pereopods 3-5 suboval or br wudly Peoria ular: palp article 4
of maxilliped shorter than article 3, blunt,! or subconical,! (setose) (fig.
OMS) Mee ANE Cc RE RAR EMRE RES Mt MORON RN ERNIE cor MeN) jhe aN ENA A aa
i)
1 Not including terminal spine.
276 U.S. NATIONAL MUSEUM BULLETIN 271
4. Article 5 of gnathopod 1 much longer than article6.......... 5 Article 5 of gnathopod 1 shorter than or subequal to article6. ..... 6
5. Spines on outer plate of first maxilla nine or more, article 4 of pereopod 3 not wider than on pereopods 4-5 .... . . . . . . Pseudischyrocerus Spines on outer plate of maxilla 1 reduced i four, article 4 of pereopod 3 twice as wide as on pereopods 4-5 . ... . . . . . . Bathyphotis
6. Accessory flagellum a small scale, articulate or Hot (Gg 107c). . Parajassa Accessory flagellum elongate (figs. 107d,e) . . . . se) ie Rie Sua,
7. Coxa 5 subequal to 6 in length, coxa 1 half as long as coxa 2. . Microjassa Coxa 5 much longer than 6, coxa 1 three fourths as long as coxa 2. . . . 8
8. Pereopods subchelate (fig. 108d) ............ .. .. .Isaeopsis Pereopods simple (fig. 107b) . ... . Reiner 8)
9. Outer ramus of uropod 3 with basally ranma, hooked femiaal spine, distolateral margin of ramus with one to three large, reverted, sharp scales or several denticles (fig. 108k); male gnathopod 2 usually with large tooth on proximal portion of palm and with tooth on distal portion of palm.
Jassa ?
Outer ramus of uropod 3 with or without basally immersed terminal spine, never hooked, distolateral margin of ramus with 0-7+ perpendicular, blunt, usually extremely small denticles (fig. 1087); male gnathopod 2 lacking proximal palmer tooth but often with distal palmar process. Ischyrocerus ”
2 These genera extremely difficult to distinguish except in adult males.
Genera of Ischyroceridae
Bathyphotis Stephensen Bathyphotis Stephensen, 1944b.
Type-species: JB. tridentata Stephensen, 1944b (monotypy).
Accessory flagellum multiarticulate, elongate; article 5 of gnathopod 1 longer than article 6; coxa 1 more than three fourths as long as coxa 2, coxa 4 subequal to coxa 6 in length, coxa 4 excavate posteriorly; article 4 of pereopod 8 tuice as wide as on pereopods 4-5; outer plate of maxilla 1 with four spines (Pseudischyrocerus). Species: 1, subarctic Atlantic, bathyal (1,096 m).
Bogenfelsia J. L. Barnard Bogenfelsia J. L. Barnard, 1962d.
Type-species: 8B. incisa J. L. Barnard, 1962d (original designation).
[Accessory flagellum unknown]; article 5 of gnathopod 1 longer than article 6; coxae minute, not touching, equal in length. Species: 1, antiboreal Atlantic, abyssal (4,893 m).
Bonnierella Chevreux Bonnierella Chevreux, 1900.
Type-species: Podoceropsis abyssi Chevreux, 1887 (monotypy).
MARINE GAMMARIDEAN AMPHIPODA ORT
g Di \ OE
Ficure 107.—Ischyroceridae: a, Microjassa macrocoxa Shoemaker (1942); b, Ischyrocerus anguipes Kréyer (Sars, 1895, pl. 209). Accessory flagella: c, Parajassa pelagica (Leach) (Sars, 1895, pl. 214, as Janassa capillata); d, Jassa falcata (Montagu) (Sars, 1895, pl. 212); e, Ischyrocerus megacheir (Boeck) (Sars, 1895, pl. 211). Mouthparts, Ischyrocerus : f, upper lip; g, mandible; h, lower lip; 7,7, maxillae 1, 2; &, maxilliped.
278 U.S. NATIONAL MUSEUM BULLETIN 271
Accessory flagellum 2-articulate, elongate; article 6 of gnathopod 1 longer than or equal to article 5; coxa 1 as long as coxa 2, coxa 5 longer than coxa 6; article 2 of pereopods 3-5 narrow, linear, palp article 4 of marilliped claw-like, longer than article 3 (combining char- acter). Species: 5, Atlantic, E. Pacific, bathyal to abyssal.
Isaeopsis K. H. Barnard Tsaeopsis K. H. Barnard, 1916.
Type-species: J. tenaz K. H. Barnard, 1916 (monotypy).
Accessory flagellum 1-articulate; article 6 of gnathopod 1 longer than article 5; coxa 1 “‘small,”’ coxa 5 longer than coxa 6; pereopods pre- hensile. Species: 1, S. Africa, littoral.
Ficure 108.—Ischyroceridae: Gnathopod 1: a, Ischyrocerus anguipes Krgyer (Sars, 1895, pl. 209); b, Jassa falcata (Montagu) (Sars, 1895, pl. 212); c, Pseudischyrocerus denticauda Schellenberg (1931). Pereopod 1: d, Isaeopsis tenax K.. H. Barnard (1916). Gnathopod 2: ¢, Jassa, note sharp distal palmar tooth and poor development of proximal tooth as an exception to couplet 9 of key; f, Ischyrocerus. Pereopods 3, 4, 5: g,h,1, Microjassa macro- coxa Shoemaker (1942). Uropod 3:7, Ischyrocerus; k, Jassa. Telson: 1, [schyrocerus.
MARINE GAMMARIDEAN AMPHIPODA 279
Ischyrocerus Kr¢yer Ischyrocerus Kr¢yer, 1838.—Stebbing, 1906.
Type-species: J. anguipes Krgyer, 1838 (monotypy). See Sars, 1895.
Accessory flagellum 2-articulate, elongate; article 6 of gnathopod 1 longer than article 5; coxa 1 more than three fourths as long as coxa 2, coxa 5 longer than coxa 6; article 6 of male gnathopod 2 lacking large posteroproximal tooth; outer ramus of uropod 3 with or without basally immersed, unhooked terminal spine, distolateral margin usually with perpendicular, blunt, mostly very minute denticles (Jassa). Species: 32, possibly cosmopolitan but primarily boreal, littoral to bathyal (2,090 m).
Jassa Leach
Jassa Leach, 1814a.—Stebbing, 1906. Lusyta Nardo, 1847.
Cratophium Dana, 1852b.
Macleayia Haswell, 1880a.
Wyvillea Haswell, 1880c.
Bruzeliella Norman, 1905.
Hemiyassa Walker, 1907.
Type-species: Cancer falcatus Montagu, 1808 (selected by Chevreux and Fage, 1925; indicated by subsequent montoypy through elimination by Stebbing, 1899a, and thereafter synonomy of species). See Sars, 1895 (as Podocerus falcatus).
Accessory flagellum 2-articulate, elongate; article 6 of gnathopod 1 longer than article 5; coxa 1 more than three fourths as long as coxa 2, coxa 5 longer than coxa 6; article 6 of male gnathopod 2 usually bearing a large posteroproximal tooth; outer ramus of uropod 3 with distal, basally immersed, hooked spine, distolateral margin with 1-3 large, sharp, reverted scales (Ischyrocerus). Species: 7, cosmopolitan, littoral.
Microjassa Stebbing
Microjassa Stebbing, 1899b; 1906.
Type-species: Podocerus cumbrensis Stebbing and _ Robertson, 1891 (monotypy). See Chevreux and Fage, 1925.
Accessory flagellum 1-articulate, very small; article 6 of gnathopod 1 longer than article 5; coxa 1 less than half as long as coxa 2; coxa 5 not longer than coxa 6. Species: 2, E. Atlantic, E. Pacific, littoral.
Parajassa Stebbing
Janassa Boeck, 1871 (homonym, Pisces). Parajassa Stebbing 1899a; 1906 (new name for Janassa).
280 U.S. NATIONAL MUSEUM BULLETIN 271
Type-species: Jassa pelagica Leach, 1814a (monotypy of Boeck and subsequent synonymy). See Sars, 1895 (As Janassa capillata). Accessory flagellum a small tubercle; article 6 of gnathopod 1 longer than article 5; coxae 1 and 2 equal in length, coxa 5 longer than coxa 6. Species: 4, Atlantic, E. Pacific, amphiboreal, littoral.
Pseudischyrocerus Schellenberg Pseudischyrocerus Schellenberg, 1931.
Type-species: P. denticauda Schellenberg, 1931 (present selection).
Accessory flagellum multiarticulate, elongate; article 5 of gnathopod 1 longer than article 6; coxae 1 and 2 equal in length; coxa 5 probably longer than coxa 6; article 4 of pereopod 3 wider than on pereopods 4-5; outer plate of maxilla 1 with nine spines (Bathyphotis). Species: 1, antiboreal, littoral.
Kuriidae [J. L. Barnard, 1964c]
FicureE 109
Driaenosis.—Accessory flagellum absent; body laterally compressed ; urosomites all coalesced, mandible lacking palp, molar well developed, triturative; gnathopods feeble, subchelate; uropod 3 uniramous; telson deeply cleft. Monogeneric. See Phliantidae, Talitroidea, Dogielinotidae, EKophlhiantidae, Prophliantidae, Pagetinidae.
DescripTion.—Body laterally compressed, all urosomal segments coalesced; mandible lacking palp, molar well developed; [lower lip and maxillae unknown]; maxilliped with outer lobes strongly reduced ; egnathopods feeble, slender, subchelate; uropod 3 uniramous; telson short, deeply cleft.
RELATIONSHIP.—The Talitroidea have discrete urosomal segments and well-developed outer plates on the maxillipeds.
The molar of the mandible is evanescent in the Prophliantidae and Pagetinidae.
Eophhantidae have cylindrical bodies.
The Phliantidae have evanescent mandibular molars and well- developed outer plates of the maxillipeds. Probably the Kuriidae should eventually be assigned as a subfamily to Phliantidae to demon- strate their close relationship. The Kuriidae show a link between Phliantidae and Talitroidea.
The Dexaminidae have only the last two urosomal segments co- alesced and possess biramous third uropods but the Kuriidae other- wise demonstrate a link between the Dexaminidae and the Talitroidea.
MARINE GAMMARIDEAN AMPHIPODA 281 Genera of Kuriidae
Kuria Walker and Scott
Kuria Walker and Scott, 1903.
Type-species: K. longimana Walker and Scott, 1903 (monotypy). Species: 1, Abd-el-Kuri, littoral.
ye
Ficure 109.—Kuriidae: Kurita longimanus Walker and Scott (1903): a, lateral aspect, note overlap of pereopods 3-5 probably erroneous; b, mandible; c, maxilliped; d, uropod 3; e, telson; f,g, gnathopods 1, 2.
Lafystidae FicurE 110
Diacnosis.—Accessory flagellum absent; mouthparts arranged into a conical bundle; mandibular molar absent; palp of maxilla 1 reduced to a single tiny article; maxilliped with 2-articulate palp; coxa 4 acu-
282 U.S. NATIONAL MUSEUM BULLETIN 271
minate, cozae 1-3 quadrate; gnathopod 1 simple; telson oval, short, entire. Monogeneric. See Laphystiopsidae, Acanthonotozomatidae.
Description.—Accessory flagellum absent; body broad, not carin- ate, urosomite 1 elongate; rostrum large, mouthparts arranged into a conical bundle; mandible lacking molar and spine row; lower lip lack- ing inner lobes; palp of maxilla 1 reduced to a single article; palp of maxilliped reduced to two small articles; coxae 1-3 quadrate, coxa 4 acuminate; gnathopods feeble, 1 simple, 2 weakly subchelate; uro- pod 3 biramous; telson small, oval, entire.
ReELATIONSHIP.—This family is so closely related to Acanthonoto- zomatidae that they might be joined together as subfamilies.
The Laphystiopsidae bear a 4-articulate maxillipedal palp, a man-
Ficure 110.—Lafystiidae: Lafystius sturionis Krgyer (Sars, 1895, pl. 134): a, lateral; b, dorsal view of head; c, upper lip; d, mandible; ¢, lower lip; f,g, maxillae 1, 2; 4,maxilliped; 1,7, gnathopods 1, 2; k, uropod 3; J, telson.
MARINE GAMMARIDEAN AMPHIPODA 283
dibular molar, a large palp on the first maxilla and an incised, broad- ened upper lip.
Superficially, the Lafystiidae resemble several other families through the Laphystiopsidae [which should be consulted]. As discussed therein the Laphystiopsidae are still considered distinct from Lafystiidae, necessitating the removal of Prolaphystiopsis to the Laphystiopsidae.
Genera of Laphystiidae
Lafystius Kr¢gyer
Lafystius Krgyer, 1842.—Stebbing, 1906. Darwinia Bate, 1857a. ?Dermophilus Beneden and Bessels, 1870. ?Ichthyomyzocus Hesse, 1878. Type-species: JL. sturionis Krgyer, 1842 (monotypy). See Sars,
1895. Species: 1, boreal N.E. Atlantic, littoral, occurring on fishes.
Laphystiopsidae
FicurE 111
DiaGcnosis.—Accessory flagellum absent; mandibular molar non- triturative; gnathopods feeble, simple; telson entire. See Isaeidae, Stilipedidae, Oedicerotidae, Calliopiidae, Pleustidae, Lafystiidae.
Description.—Accessory flagellum absent; body broad, carinate only on a few pleonites; urosomite 1 elongate or not; rostrum large and spatulate or absent; upper lip incised; mandible bearing molar process, simple or slightly triturative; lower lip with or without inner lobes; palp of maxilla 1 large, 1- or 2-articulate; maxilliped with small 4-articulate palp; coxae variable, either very short and rectan- cular and serially disjunct or of medium length, quadrate or slightly acuminate and overlapping; gnathopods feeble, simple; outer rami of uropods 1-2 slightly shortened; uropod 3 biramous, peduncle short; telson short or of medium length, oval, entire.
Reationsurp.—The Laphystiopsidae are simply Calliopiidae- and Pleustidae-like organisms with feeble, simple gnathopods, feeble, poorly setose maxillipedal palps and poorly developed mandibular molars. Even the lower lip of the type-genus is similar to that of Pleustidae, although Prolaphystius lacks inner lobes. Even Cleippides and Regalia in the Calliopiidae have simple gnathopods.
The Lafystiidae faintly resemble Laphystiopsidae but the latter have lost the mandibular molar and spine row, two articles of the
285-135 O - 69 - 19
284 U.S. NATIONAL MUSEUM BULLETIN 271 maxillipedal palp, and have the first maxillary palp reduced in size. The unique species of lafystiid, however, has retained the subchelate condition of its second gnathopod, although it is feeble.
The uniformity of the Laphystiopsidae is complicated by the loss of the rostrum in Prolaphystvus and that genus warrants further in- vestigation on its familial assignments. Prolaphystius has an elongate urosomite 1 like Podoceridae but otherwise bears no resemblance to that family. It has been included in the key to Calliopiidae where it
seems to have strong affinities. The Astyridae bear a small accessory flagellum and broadly sepa-
rated outer lobes on the lower lip. The Oedicerotidae warrant close inspection of their slightly sub-
chelate gnathopods as a feature distinguishing them from laphysti-
= =
> —— —— ee a oo ry
Cs Laphystiopsis planifrons Sars (1895, pl. 135): a, lateral;
Figure 111.—Laphystiopsidae: b, upper lip; c, mandible; d, lower lip; e,f, maxillae 1, 2; g, maxilliped; h,7, gnathopods 1, 2; Prolaphystius tsopodops K. H. Barnard (1930): /, head, right side,
j, uropod 3; k, telson. stippled; m, maxilla 1; n, mandible; 0, telson.
MARINE GAMMARIDEAN AMPHIPODA 285
opsids. Oedicerotids have an elongate peduncle of uropod 3, equal rami of uropods 1-2, unnotched upper lips, strongly setose appendages and disproportionately long fifth pereopods.
Stilipedidae have a broad, pardaliscid mandibular incisor, no molar and foliaceous maxillae.
Isaeids and their relatives have strongly triturative mandibular molars and usually have subchelate and/or large gnathopods.
Key to the Genera of Laphystiopsidae
(Ae AGlarcelrostrumaspresent, (liga vWblla) ior fa bo: fee meters) a Poa ol Waele eas 2 Rostrumyalosemte chica IWl)) newer eens ici ae nts lees Prolaphystius Zealpiotimaxilla libiarticulate: ies Tila) yo one. Prolaphystiopsis Halprommaxitia leumanticulate: (ig: ile). 29. 8 2 Laphystiopsis
Genera of Laphystiopsidae
Laphystiopsis Sars Laphystiopsis Sars, 1895.—Stebbing, 1906.
Type-species: L. planifrons Sars, 1895 (monotypy).
Rostrum well developed; mandibular molar conical, unridged; palp of maxilla 1 uniarticulate; telson short, oval; pleonite 3 not forming a projecting shelf; coxae not touching serially. Species: 3, boreal to China Sea, littoral to bathyal (900 m).
Prolaphystiopsis Schellenberg Prolaphystiopsis Schellenberg, 1931.
Type-species: P. platyceras Schellenberg, 1931 (monotypy).
Rostrum well developed; mandibular molar truncate, poorly ridged; palp of maxilla 1 biarticulate; telson short, oval; pleonite 3 not forming a projecting shelf; coxae not touching serially. Species: 1, subantarctic, littoral.
Prolaphystius K. H. Barnard Prolaphystius K. H. Barnard, 1930.
Type-species: P. isopodops K. H. Barnard, 1930 (monotypy).
Rostrum absent; mandibular molar truncate, poorly ridged; palp of maxilla 1 biarticulate; telson long, linguiform; pleonite 3 forming a shelf-like projection; coxae of medium size, touching serially. Species: 1, antarctic, bathyal (406 m).
286 U.S. NATIONAL MUSEUM BULLETIN 271
Lepechinellidae
Ficure 112
DraGnosis.—One or more and usually all of coxae 1-4 acutely pointed below or anteriorly, occasionally bifid; accessory flagellum composed of one spine-like article; pleonites 5-6 ?coalesced. See Paramphithoidae, Atylidae, Eusiridae, Pleustidae.
Description.—Body processiferous; accessory flagellum 1-articu- late, spine-like; rostrum conspicuous; upper lip probably always incised slightly; molar of mandible triturative, palp 3-articulate, slender, short or long; lower lip apparently with coalesced inner lobes, remaining mouthparts basic; gnathopods feeble, slender, subchelate; coxae of medium length or shortened, usually acuminate, often bifid; rami of uropod 3 greatly elongate, very slender, 3-6 times as long as peduncle; telson short, cleft, lobes diverging.
ReLationsuip.—Originally this family was a part of the Param- phithoidae, because the type-genus Lepechinella was placed there by Stebbing (1908a); the junior synonym Dorbanella Chevreux (1914) was placed in the Tironidae but Schellenberg (1925a) removed that to a family Dorbanellidae, later to take the name of the senior synonym of the type-genus. Lepechinellidae resemble Paramphithoidae but presumably all species have pleonites 5-6 coalesced. The inner lobes of the lower lip of lepechinellids apparently are always fused together in contrast to the Paramphithoidae.
The coxae of Atylidae are elongate and not acuminate distally and their gnathopods are shorter and stouter than those of Lepechinellidae. Coxae 2-4 of Paralepechinella are scarcely, if at all acuminate.
No pleustid has coalesced urosomal segments nor a_ spine-like accessory flagellum. That family has characteristic lower lips.
Dexaminidae, Kuriidae, Prophliantidae all lack mandibular palps.
Key to the Genera of Lepechinellidae
1. Mandibular palp article 3 much shorter than article 2 (fig. 112c). Lepechinella Mandibular palp article 3 immense, much longer than article 2 (fig. 112h). Paralepechinella
Genera of Lepechinellidae
Lepechinella Stebbing
Lepechinella Stebbing, 1908a. Dorbanella Chevreux, 1914.
Type-species: L. chrysotheras Stebbing, 1908a (monotypy). Species: 13, cosmopolitan, bathyal and abyssal.
MARINE GAMMARIDEAN AMPHIPODA 287
Ficure 112.—Lepechinellidae: a, Lepechinella bierii J. L. Barnard (1957c). Accessory flagellum: b, L. bierii. Mouthparts, Lepechinella chrysotheras Stebbing (1908a): c, man- dible; d, lower lip; ¢,f, maxillae 1, 2; g, maxilliped. Mandible: h, Paralepechinella longt- palpa Pirlot (1933a). Gnathopods 1-2: 7,7, L. bierit. Uropod 3: k, L. bierit. Telson: 1, L. chrysotheras.
288 U.S. NATIONAL MUSEUM BULLETIN 271
Paralepechinella Pirlot Paralepechinella Pirlot, 1933a.
Type-species: P. longipalpa Pirlot, 1933a (original designation). Species: 1, Indonesia, bathyal (1,301 m).
Ficure 113.—Leucothoidae: a, Leucothoe spinicarpa (Abildgaard) (Sars, 1895, pl. 100); b, Leucothoella bannwarthi Schellenberg (1928); c, Leucothoides pottst Shoemaker (1933a). Upper lip: d, Leucothoe. Mandible: e, Leucothoe; f, Leucothoides.
MARINE GAMMARIDEAN AMPHIPODA 289
Leucothoidae
Figures 113, 114
Diaenosis.—Accessory flagellum vestigial, 1- or 2-articulate, very small; mandible lacking molar; outer plates of maxilliped very small, probably never larger than inner plates; telson entire; gnathopod 1 carpochelate. See Anamixidae, Sebidae, Amphilochidae, Cressidae, Thaumatelsonidae.
DescripTion.—Body lacking dorsal processes; accessory flagellum vestigial; epistome probably always produced in front; upper lip probably always asymmetrically incised; mandibles lacking molar, palp 1- or 3-articulate; maxillae basic; maxilliped with very small outer plates, partially fused inner plates and long slender palp; gnathopod 1 carpochelate, immovable finger long and narrow, gnatho- pod 2 larger than 1, subchelate, article 5 elongate and guarding article 6 posteriorly; coxae variable; uropod 2 shortened; uropod 3 with styliform rami, peduncle elongate, rami not longer than peduncle; telson entire, not longer than peduncle of uropod 3.
ReuatTionsHip.—The Anamixidae lack maxillae and recognizable mandibles.
The Cressidae, Thaumatelsonidae, and Stenothoidae have unira- mous third uropods and subchelate first gnathopods.
The Sebidae have uniramous third uropods and chelate second enathopods.
Key to the Genera of Leucothoidae
1. Coxa 2 covering most of coxa 1 (fig. 113c), mandibular palp 1-articulate Gils, WISI) ie os . . . Leucothoides
Coxa 1 not concealed, Sibu male Be icuiate ae NBT AOD Masse ties Ais 074
2. Coxa 2 very short, broader than long, acute anteriorly (fig. 1130). Leucothoella
Coxa 2 at least as long as broad, rounded below and anteriorly (fig. ll3a). .3
3. Outer plate of maxilliped reaching less than halfway along palp article 1 Gig, WMA SG a . . . . . Leucothoe
Outer plate of Tealliped P aehine almnoet 46 ond of palp article 1. Paraleucothoe
Genera of Leucothoidae
Leucothoe Leach
Leucothoe Leach, 1814a.—Stebbing, 1906. Cuvieria Leach, 1814a (nomen nudum). Lycesta Savigny, 1816. Type-species: Gammarus spinicarpus Abildgaard, 1789 (monotypy and subsequent synonymy). See Sars, 1895.
290 U.S. NATIONAL MUSEUM BULLETIN 271
Coxa 2 at least as long as broad, rounded ventrally and anteriorly, coxa 1 not concealed; mandibular palp 3-articulate; outer plate of maxilliped reaching less than halfway along palp article 1. Species: 25, cosmopolitan, littoral to abyssal.
Ficure 114.—Leucothoidae: Leucothoe spinicarpa (Abildgaard) (Sars, 1895, pl. 100): a, lower lip; b,c, maxillae 1, 2; d, maxilliped; ¢, uropod 3; f, telson; g,h, gnathopods 1, 2. Leucothoides pottst Shoemaker (1933a): 7, telson; 7, gnathopod 2.
Leucothoella Schellenberg Leucothoella Schellenberg, 1928b.
Type-species: ZL. bannwarthi Schellenberg, 1928b (monotypy)-
Coxa 2 very short, broader than long, acute anteriorly, coxa 1 not concealed; mandibular palp 3-articulate; outer plate of maxilliped reaching less than halfway along palp article 1. Species: 1, tropics, Red Sea to Fiji, littoral.
Leucothoides Shoemaker Leucothoides Shoemaker, 1933a.
Type-species: L. pottsi Shoemaker, 1933a (monotypy).
Coxa 2 at least as long as broad, rounded ventrally and anteriorly, coxa 2 covering most of coxa 1; mandibular palp l-articulate; outer plate of maxilliped reaching less than halfway along palp article 1. Species: 2, tropical Pacific, Caribbean, littoral.
MARINE GAMMARIDEAN AMPHIPODA 9291
Paraleucothoe Stebbing
Paraleucothoe Stebbing, 1899d.—Stebbing, 1906.
Type-species: Leucothoe novaehollandiae Haswell, 1880c (original designation).
Coxa 2 at least as long as broad, rounded ventrally and anteriorly, coxa 1 not concealed; mandibular palp 3-articulate; outer plate of maxilliped reaching almost to end of palp article 1. Species: 1, Australia, littoral.
Liljeborgiidae Fiacures 115, 116
Draanosis.—Molar of mandible poorly developed, not triturative, accessory flagellum 2- or more articulate. See Gammaridae, Eusiridae, Pleustidae, Haustoriidae, Astyridae, Vitjazianidae.
Descriprion.—Accessory flagellum always present, 2- or more articulate; rostrum inconspicuous; coxae long or of medium length; upper lip possibly slightly emarginate, not conspicuously so; mandibles always bearing 3-articulate palp with article 1 often elongate and as long as article 3 (except Jdunella forming a link to Gammaridae); molar of mandible obsolescent, nontriturative; remaining mouthparts basic; gnathopods powerfully subchelate; uropod 3 with flattened, lanceolate rami; telson deeply cleft, not greatly elongate, apices of lobes bifid.
RELATIONSHIP.—In the Gammaridae only the genus Parelasmopus has an elongate mandibular palp article 1 but its mandibular molar has a well-developed grinding surface, unlike that of Liljeborgiidae. Apart from the mandible the Liljeborgiidae are like the Gammaridae.
The Astyridae differ from the Liljeborgiidae by the overall appear- ance and the presence of inner lobes or a broad medial space on the lower lip (see figures 32 and 41). Astyrids have feeble gnathopods, whereas those of liljeborgiids are powerfully developed. The rami of uropod 3 of Astyridae are much more elongate than those of Lilje- borgiidae.
Some genera of Pleustidae have poorly developed mandibular molars as in Liljeborgiidae and the lower lips are similar. Pleustids, however, have vestigial or no accessory flagella and usually have uncleft telsons, but one species of Austropleustes confounds the defini- tion. See Gammaridean Family Key G for specific distinctions.
The Eusiridae always have the accessory flagellum 2-articulate or less and this conflicts only with the genus Listriella in the Liljeborgiidae. But in contrast to the Eusiridae, Listriella usually has a 2-articulate
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U.S. NATIONAL MUSEUM BULLETIN 271
outer ramus of uropod 3 with an elongated article 1 of the mandibular palp. Lusirella and Eusiropsis are the only eusirids having an obsoles- cent mandibular molar and they are clearly eusirids for their possession of calceoli. Eusirids have the outer ramus of uropods 1 and 2 shortened; in liljeborgiids this shortening is slightly evident only on uropod 2.
One genus of Amphilochidae, Pseudamphilochus, bears resemblance to Liljeborgiidae for its unreduced coxa 1, cleft telson, weak mandi- bular molar, and moderately powerful gnathopods. Pseudamphilochus
presumably lacks an accessory flagellum, has a large rostrum, and nonbifid apices of the telson.
oe an as ee a
—eN <5 ol BE Se
Ficure 115.—Liljeborgiidae: a, Liljeborgia brevicornis (Bruzelius) (Sars, 1895, pl. 187, as L. pallida); b, Idunella aequicornis (Sars, 1895, pl. 190).
MARINE GAMMARIDEAN AMPHIPODA 293
Key to the Genera of Liljeborgiidae
1. Gnathopod 1 larger than gnathopod 2 (fig. 115b) . . . . .. . +. =. Idunella
Gnathopod 1 smaller than or equal to gnathopod 2 (fig. 115a) . ES AD,
2. Article 5 of gnathopods 1-2 strongly produced, slender and elongate (fig. 1169) ;
outer ramus of uropod 3 always simple (fig. 1167) .... . . Liljeborgia
Article 5 of gnathopods 1-2 weakly produced, thick and blunt (fig. 116)); outer ramus of uropod 3 usually composed of two articles (fig. 116k)
Listriella
1
Ficure 116.—Liljeborgiidae: Liljeborgia brevicornis (Bruzelius) (Sars, 1895, pl. 187, as L. pallida): a, upper lip; b, mandible; c, lower lip; d,e, maxillae 1, 2; f, maxilliped; g, gnathopod 2; h, telson; 1, uropod 3. Gnathopod 2: 7, Listriella goleta J. L. Barnard (1959b). Uropod 3: k, Listriella diffusa J. L. Barnard (1959b); l, Listriella ertopisa J. L. Barnard (1959b).
Genera of Liljeborgiidae
Idunella Sars
Idunella Sars, 1895.—Stebbing, 1906.—J. L. Barnard, 1959b. Sextonia Chevreux, 1920.
Type-species: Liljeboria aequicornis Sars, 1876 (monotypy). See Sars, 1895.
Species: 4, subarctic, warm-temperate of N. Atlantic, tropical Indian Ocean, Japan, littoral to bathyal (763 m).
294 U.S. NATIONAL MUSEUM BULLETIN 271
Liljeborgia Bate
Iduna Boeck, 1861 (homonym, Aves). Liljeborgia Bate, 1862.—Stebbing, 1906.—K. H. Barnard, 1932. Microplax Liljeborg, 1865 (homonym, Hemiptera). Lilljeborgiella Schellenberg, 1931. Type-species: Gammarus pallidus Bate, 1857a (monotypy of Bate, 1862). See Bate and Westwood, 1863. Species: 28, bipolar and amphiboreal, littoral to abyssal.
Listriella J. L. Barnard
Listriella J. L. Barnard, 1959b.
Type-species: JL. goleta J. L. Barnard, 1959b (original designation). Species: 8, tropical and warm-temperate Atlantic and E. Pacific, littoral, often inhabiting tubes of maldanid polychaetes.
Lysianassidae Figures 117-132
Diacnosis.—Accessory flagellum usually present and multiarticu- late; article 3 of gnathopod 2 elongate (fig. 3a), gnathopod 2 other- wise of the typical lysianassid kind and with a fuzz of scales or setules on articles 5 and 6 (fig. 129); peduncle of antenna 1 short and stout. See Sebidae, Ochlesidae, Stegocephalidae, Didymocheila (incertae sedis).
Description.—Accessory flagellum usually present and more than 3-articulate but occasionally vestigial or few in segments; peduncle of antenna 1 short and stout, articles 2 and 3 usually much shorter than 1 and occasionally scarcely distinguishable because of tele- scoping; pereon generally lacking processes, often urosomal segment 1 processiferous; body compact, chitin usually very smooth, porcel- lanous; mouthparts enormously variable, either chewing or piercing, often with reduction in mandibular molars, variable in position of mandibular palps; upper lip and epistome invaluable for identification, either or both often produced in variable configurations; maxilla 1 generally with 2-articulate palp but occasionally reduced or absent; maxilla 2 occasionally poorly developed; maxilliped occasionally with reduced plates and palps; gnathopods usually quite small, 2 mitten- shaped terminally, with article 3 elongate, article 5 generally with a “pineapple cushion”’, i.e., posterior margin bulbous and with micro- scopic texture of a pineapple, structure composed of scales or coarse setules, article 6 generally covered with coarse setules or tasseled setules; pereopods generally stout, occasionally prehensile; coxae usually long, first one or two or three occasionally reduced in size or
MARINE GAMMARIDEAN AMPHIPODA 295
hidden by following coxae; rami of uropod 3 lanceolate, outer often 2-articulate, inner occasionally reduced in size, rarely both absent; telson cleft, emarginate or entire, long or short; rarely absent.
Gnathopod 2 is an unmistakable mark of this family. Nevertheless, Sebidae have a chelate gnathopod 2 with elongate article 3 but they are distinguished from Lysianassidae by the elongate peduncle of antenna | and the elongate and uniramous uropod 3. Some talitroideans have a mitten-shaped gnathopod 2 but article 3 is not elongate. Some Stegocephalidae also have long third articles but all of them have a characteristic shape of coxae 1-4, the first acuminate, the second and third narrower than 1, the fourth extremely broad, all four coxae fitting together tightly, forming a ventral semicircle and not over- lapping. Stegocephalid mouthparts are highly characteristic although a few Lysianassidae have mouthparts resembling them. Stegocephalid mandibles usually have some foliaceous parts, rarely occurring in Lysianassidae. Stegocephalid mouthparts are always formed into a conical bundle, rarely occurring in Lysianassidae. Gnathopod 2 is always simple, whereas few Lysianassidae have a completely non- subchelate gnathopod 2.
Nomenclatural Changes in Lysianassidae
Alicella scotiae Chilton (1912) is removed to Paralicella.
Aruga Holmes (1908) is synonymized with Lysianassa.
Arugella Pirlot (1936) is synonymized with Lysianassa.
Lakota adversicola K. H. Barnard (1925) is removed to Schisturella.
Lakota rotundatus K. H. Barnard (1925) is removed to Schisturella.
Lysianassa anomala Nicholls (1938) is removed provisionally to Parambasia.
Lysianopsis Holmes (1903) is synonymized with Lysianassa.
[Pronannonyz] Schellenberg (1953) is also synonymized with Lysian- assa.
Shoemakerella Pirlot (1936) is synonymized with Lysianassa.
Socarnopsis obesa Chevreux (1927) is removed to Socarnes in view of Hurley’s (1963) classification, because uropod 3 has the outer ramus 2-articulate.
Tryphosites cozalis J. Li. Barnard (1962d) is transferred to Tryphosella Bonnier, and is reflected in Key I, triplicut 7 (p. 304).
Key to the Genera of Lysianassidae
A basic key partitions the genera into several groups further developed in keys A to Q. This method brings together genera or portions of genera in different places in order to signal their complicated, net-like
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U.S. NATIONAL MUSEUM BULLETIN 271
relationships and obviates the use of a long key with more than 100 couplets. Key N is provided as a separate entity for a special group of
genera. BASIC LYSIANASSID KEY
ie aelsonvabsents (hess W22 ail 24 a) Ne eet sine cnet re ee enh Once eae av
Telson present . . RT AN Hee ses Ae uA SEM ALE tr ert
2. Mandible without distineth incisor iGo 126c) ..... . . .Kerguelenia
Mandible with distinct incisor (figs. 126a,b,d) .... . ii) CaaS
3. From lateral view, mouthparts group strongly and comically ancien below
head, more or less styliform (fig. 123e) ... . Pan aeersenune aan 2" 6}
Mouthparts group usually not strongly or eonielhy projects below head,
parts not styliform [if so, uropod 3 lacking rami ae SVAN iis eects ncn Ae
4. Gnathopod 1 chelate (figs. 129a,e,h,0). . . . : 4 Visuaoaae KCN
Gnathopod 1 subchelate or simple, rarely appearing areen ele Ben ice aD
5. Mandibles: middle of incisor sharply dentate (fig. 1260) ... . .Key D
Mandibles: middle of incisor not dentate .............. 6
Oo Maxilla tilackane: palpy (ies 127b))i ie AGIAN e ms bond (ONY, IS
Maxilla 1 bearing 1- or 2-articulate palp (fig. 126a) . Sioa 1 2 gol Meee TE
7. Both coxae 1 and 2 (and occasionally 3) very small or distinctly smaller
10.
Ii.
12.
13.
than following coxae, largely hidden by following coxae (fig. 122c). Key F Neither coxae 2 nor 3 distinctly reduced, both visible (fig. 122d) (occasionally coxa 1 reduced alone) . . . . . . . Choose either couplet 8 or 21! Telson entire or emarginate, no syniteall slit (iigss 128pp)) ae ee yaGr Telson cleft enough to provide slit or facing edges at angle of less than 45 degrees (figs. 128n,0,t) . . . SPUN GAA On ae aE ear Article 2 of pereopod 3 deeply anGlantaniadl (Ge 1310) ... . Glycerina Article 2 of pereopod 3 not deeply indentured . . use either couplet 10 or 11 [Branchiae with large plaits (fig. 127s) (use Key H for rapid identification of select group of genera only, otherwise continue to couplet 11) . . Key H] [Branchiae lacking plaits (if in doubt use this couplet). ...... 414] Article 7 of gnathopod 1 minute, shrouded among a large and long tuft of setae or cirri attached to either or both propodus and dactyl (figs. 129n, ES O70) a ne ene peer os od Sy IEG
Article 7 of paaoped 1 leary eaiple! not ehroudcd:
use either couplets 12 or 19 Upper lip produced forward lobately and usually conspicuously in front of epistome (epistome occasionally also projecting forward alongside dorsal
edge of labral lobe) (figs. 125a-d) ... .. . Mena anints ) IK LY Upper lip not conspicuously produced forward lobe, or epistome and upper lip coalesced even if produced. . . pseu lay Epistome produced forward acutely and seoematalhy ioral appee lip (fig. 1250). 14
Epistome, if produced, not acute or conical or epistome and upper lip co- alescedieven though produced sac) ied iach)
1 Couplet 8 leads to characters of ease in identification whereas couplet 21 leads to a possible natural group- ing by use of mandibular structures.
2 Key I generally contains taxa with produced upper lip (labrum) even if coalesced with epistome as long as labral portion recognizable by its position; couplet 13 generally contains taxa with epistome and upper lip together lobate and separated by a notch even though some of these taxa occur in key I for comparative purposes.
14.
15.
16.
17.
18.
19.
20.
ile
22.
23.
24.
MARINE GAMMARIDEAN AMPHIPODA 297
Mandibular palp attached level with molar or at its proximal corner. . 15 Mandibular palp attached strongly proximal to molar.
(exceptional) Orchomene Coxa 1 tapering slightly and partially covered by coxa 2, inner ramus of
WEOPOG) eZ uasimiple rns) ae me . .. . Tryphosella Coxa 1 fully expanded distally and: freely, “eels, t inner ramus of uropod 2 deeply constricted... . . Tryphosites Telson cleft less than one fonean ai Ais fener Gre. 128n, OGM) sche ) WEN] Telson cleft one third or more of its length (figs. 128s,t,v) . . . ... 17
Coxa 2 concealing anteroventral corner of coxa 1; or otherwise shielding coxa 1; or coxa 1 reduced in size or subtriangular (figs. 119b, 122d, 1236).
Key L Coxae 1 and 2 both large and visible and not triangular (figs. 117a, 118a@) . 18 Mandible lacking palp...,.......... . . .Stephensenia Mandible bearing palp. . . . Gai wee NE
(from couplet 11.) Coxa 1 dlicitiy or Perrone pedieca in size or tapering distally (figs. 119b, 122d, 1236), often partially covered by coxa 2. Key O Coxa 1 of normal size, anterior and posterior edges parallel or coxa 1 distally
expanded... . nS CERT SLI Baa MeL Meili ir carne CoN P Rng Snare ene AD Mandible lacking snl CATENIN epUNE OS Retccict Saas ame Sara _ Stephensenia Mandible bearing palp. . . . Patan Stardaan ee (eye) SOV ANNE (from couplet 7.) Mandible neta walls . 2... . . . . Stephensenia Mandible bearing palp .... aber ee Article 7 of gnathopod 1 minute, simondles among eciae or cirri attached to
AGCICIES a/MOL iO elec: Sete Oe oe erage . Key K Article 7 small or large, Higiislle, neues, Sona ey OA, oe Ry 8)
Pereopod 1 prehensile and resembling a large patnonod) Endevoura and Ensayara Pereopod 1 simple ..... GaGa ur Pao Mandibular palp attached diecinedly -onodianell 40 mola or ait molar afer palp distinctly proximal in position. . . . pea le eae Mandibular palp attached level with or distal io inoler, or if molar absent palp attached distinctly toward distal end of mandible. . . . . . Key Q
KEY A
These are highly aberrant genera, not certainly placed in the Lysianas- sidae. Shoemaker (1945a) believed Thoriella to be in the family Talitridae but the second gnathopod is distinctly that of a lysianassid. Chevreuriella and Danaella bear resemblance to cyphocarids (key F) by reason of their coxae.
Il,
to
1.
Maxillipedal palp absent (fig. 1282); pereonites 3-6 hugely inflated, forming a
slobular body, m4 (heal Zila) aan 5 ene . . . . . Danaella
Maxillipedal palp present even if degraded: Wercomires 3-6 not larger than
other body segments (fig. 1176). . . . . EO Oca aa as ye,
Urosome composed of two segments (fig. 1240) . SPO een oE Uoheweuxiells
Urosome composed of three segments (fig. 122a). . . ... . . . Thoriella KpyY BA
i)
HKyes and gnathopod 1 together powerfully developed (fig. 123a) . Eyes and gnathopod 1 not together powerfully developed .
(es)
298 U.S. NATIONAL MUSEUM BULLETIN 271
2. Coxa 1 half as long as coxa 2 (fig. 123a), palp of maxilla 1 very short (fig.
LTE) wie ane ee : Trischizostoma Coxa 1 nearly as Ns & as coxa 2, jello Bf saocaill 1 noneie . . . Normanion 3. Uropod 3biramous . . . SAY UAE CS pha ce VEC Seats as ale eR Uropod 3 not biramous, rami aia aalsie PUTA ee mime Oa Ud See Ny 4. Maxilla 1: palp well developed, 2-articulate ......2.2.2.2.2.2.. 5 Maxilla 1: palp vestigial (fig. 127g)... ....... . . Acidostoma 5. Maxilliped: article 4 of palp well developed. .......2.2.2.2.2. 6 Maxilliped: article 4 of palp vestigial (fig. 128d) ........2.2.2.7 6. Telson deeply cleft, gnathopod 1 stout. ....... Shackletonia Telson entire, gnathopod 1 very slender, flagellate. . . . . Azotostoma 7. Gnathopod 1 simple. .. . be betes tea Ee ie EP HOxOSstona Gnathopod 1 strongly pubenelates POE OE SIN edo as Des NOR MaAnion 8: "Gnathopod df ‘simples. ee an he Oe Ron pa Gnathopod -1 ‘chelate’; 4.050% 50.0. . bo ee ee Se Deriugiana o> Dactyl of gnathopod'2) present) (yy, 20 es eee ieee) Dactyl of gnathopod 2 absent... . . . . Ocosingo
10. Coxae 1-2 strongly reduced in size and nivel iidden Ph large coxa 3. Lepidepecreella Coxae 1-2 large and visible. . . . . . Acontiostoma and Stomacontion
1 See also Parawaldeckia kidderi (Smith) (p. 357).
KEY C
The first gnathopods of Valettia, Normanion, Cheirimedon and occa- sionally of Orchomene and Hirondellea and females of Onesimoides are weakly chelate. These genera are also included in other keys. Gnatho- pod 1 of Microlysias is vaguely chelate but the genus is not included here.
1.” Uropod! dijlackingrami 2.3 us, 618 aa oe ee ee enjugiana Uropod 3 withrami .. . 2
2. Mandible: cutting edge ohenly touted (though sue minutely) (ig P26 Oi es ae Pre Rea SAE 4g : 3 Mandible: einen pds net toothed! Giz 126h) . HPs eye ee eS
3. Pereopod 3: article 2 deeply indentured (fig. 131b) . . . . . . Podoprion Pereopod, ajarticle 2 not indentured: 4 vis) 6) ve ee
4. Inner ramus of uropod 8 vestigial, telson entire . . . . . . Onesimoides Rami of uropod 3 subequal to each other, telson cleft. . . . . . Valettia
5. | Maxillipedall palp)3-articulate (fig. 1289). a) ts = Maxillipedalpalp 4-articulate 22.) s gmk tk
6. Melsoniclefts ox mencan Whatua ay atte ronnie hy runs Dien ene OC ODEONICES Telson entire .. . LiSeh si gS Aiea eal Sn RCIA NSt AR dele a atc aN ae le
7. Maxilla 1 lacking Palpe slated al de Bec gun cle Mt | et hee ene ee eae aC Lite lle Maxilla 1 bearing palp .... : ae 8
8. Pereopods 3-5: article 2 strongly andienined pal p of Peeciined nen eeaadiing end of outer plate. .... . . . . . Podoprionella Pereopods 3-5: article 2 not Sadana: wal of maxilliped shorter than outer plate (fig. 128g) .... . . . Normanion
9. Coxa 1 reduced in size or tapering distally andl santelie hidden by coxa 2. 10 Coxa 1 of normal size and subquadrate shape, and not hidden by coxa 2. 14
MARINE GAMMARIDEAN AMPHIPODA 299
10. Inner plate of maxilla 1 with large sickle-shaped setae (fig. 127d).
Hirondellea
Inner plate of maxilla 1 with normal setae... . sii ep beh eae a
11. Mandibular molar obsolescent or absent, gnathopod 1 eeronely: chelate or lleWeuey wae ens ctl elie bY Mandibular polar Ww ell developed: rieed or asnelloxs, Eatinonod 1 weakly chelate (fig. 129a) . . . ; Aenea li
12. Gnathopod 1 very slender, anal aries (fig. 1290), anions si aionoaly. Euonyx
Gnathopod 1 powerful, article 6 very broad, chela formed by deeply excavate palm (fig. 129e), epistome unproduced . . . . Opisa
Gnathopod 1 powerful, article 6 very broad and disiellly wexonniling palm protuberant, not excavate (fig. 1291), epistome and labrum each with
shallow rounded lobe ... . . . . Cheirimedon latimanus Sars, 1895
13. Epistome with anterior bulb separated from protrusion of upper lip by long,
flat margin... ... . . Aristiopsis Epistome rounded anal papresced t0 Proton oF Mane lip (exceptional).
Schisturella
14. Mandibular molar well developed, cuboid or ovate, setulose or ridged . . 15
Mandibular molar conical and unridged, obsolete or absent. . . .. . 19
SMe SOmsaclehts neu eeumalsunesen lewis: taal = aanae sole is .... . Gainella
Telson entire... . . . . some species of Orchomene
16. Gnathopod 2 tapering tor a point (but Rate chelate) (fig. 1312). (alternate position) Gainella
Gnathopod 2 distally blunt. . . OS RSS ERE oh er Ta ocd LG 17. Maxillipedal palp article 4 cestisial Ge. 128¢) 1 aoe ae ne Pachynus Maxillipedal palp article 4 normal, claw-shaped. . .. . ee ON car ae LSS 18. Outer plate of maxilliped large, exceeding end of palp article 2 Pg een ase gs Nadas)
Outer plate of maxilliped small, not or scarcely exceeding palp article 1 . 20 19. Eyes very large (losing pigment in alcohol), accessory eae 4+ -articulate,
telson longer than broad. ... . Jon Moroga
Eyes absent, accessory flagellum 1- maaeiee, telson as broad as long. Figorella 20. Spines on outer plate of maxilla 1 reduced to two or three fused members, outer ramus of uropod 3 uniarticulate ... . . . . . . Sophrosyne Spines on outer plate of maxilla 1 of normal humber (9) and articulate, Guberramus of uropodss biarticulate!. 0... 62) 4.00.0. 4. eiyska
KEY D
Most lysianassids have the primary cutting edge of the mandible bearing at least one notch subtended by a tooth but these notches are always lateral and never in the middle of the cutting edge; the following species have teeth along the middle margin of the primary mandibular plate.
Valettia is also included in Key B, because of its slightly chelate first gnathopod. Podoprion and Onesimoides in Key C and Aristiopsis in Keys C and N also have a toothed incisor. Clepidecrella may have an extremely minute incisorial dentition. Nicholls’ (1938) figure of Adeliella has a slight indication of incisorial teeth.
285-135 O - 69 - 20
300 U.S. NATIONAL MUSEUM BULLETIN 271
1) ¢Gnathopod Aysimple a evs OM nnn Chin MG has aes ate Alice lal Gnathopod 1 subchelate. . . . . ia SLM Daa) ni aan Ni Ato 2. Telson entire, inner ramus of anopod 3 sjestioval ... . . . Onesimoides Telson cleft, rami of uropod 3 subequal to each other. ........ 3
3. Coxa 2 perfectly rectangular and elongate, partially covering coxa 1, latter shortened, subtriangular; inner plates of maxillae 1-2 fully lined with setae medially. .... .. . . . Valettiopsis
Coxa 2 short, broad, cupreumded soareely sonesaline any portion of coxa 1, latter not shorter than coxa 2 and of similar shape; inner plates of maxillae 1-2 with mostly terminal setae, medial edges scarcely or not setose.
Valettia
KEY E Maxilla 1 lacks a palp, but the mouthparts are arranged in a quadrate bundle, gnathopod 1 is not chelate, and the mandibular cutting edge
is not toothed. See Key A, Acidostoma in Key B and Prachynella in Key C.
1. Gnathopod 1 very large, subchelate (fig. 130a), ‘‘telson entire’ . Pachychelium
Guathopod 4d smallisimpley {telson clebti@. 4 92) se ee
2. Antenna 1: article 2aslongasarticle1] ...... . . . Bathyamaryllis
Antenna I article: 2;abouthalfaswongyas Wei) ea en eee
3. Anteroventral corner of coxa 4 acutely produced ....... . . Vijaya
Anteroventral corner of coxa 4 rounded .......... .. Amaryliis KEY F
These are the cyphocarids, characterized by the distinctly reduced first and second coxae (and occasionally the third), often hidden behind coxa 3 (fig. 122c) but not always, depending on the anterior prolongation of the third coxae (or fourth). Coxae 3-4 of Metacyclocaris do not fit this key because of their small size but the genus is included for other comparisons. See note of Key A.
1. Urosomites 2 and 3 coalesced (fig. 122c). . . . . . . .Crybelocyphocaris Wrosomites.2 andis separatel, m0 suis hy way ene ne ce nuit le dae eam: 2.5 Mandible lacking: paler tai ce wives sie Ree Miele eeu enn aS. Mandible bearing palp ... . at SES eae
3. Pereopods 4—5 prehensile, article 6 of Pneihopod 1 enone then article 5. Mesocyclocaris Pereopods 4-5 not prehensile, articles 5 and 6 of gnathopod lsubequal . . 4 4. Pelson entired sn, 0 ie to es Sl am ee 2 ee = Cry belocephalus Telson cleft... . . . . . . . Metacyphocaris 5. Uropod 3: inner ramus speomeley neetneadl or aneent (fig 32h) 2) eee Uropod 3: rami subequal to each other, well developed. . ..... =. 8 6. Pereopods simple (fig. 117a). .......... . . . Lepidepecreella Pereopods prehensile a LSEG) ORRIN Oe SS ot 8a ae (eel SOnnClehtineiie Manin (chu iain ninn Winer aaah ea Me SoCyDROeanls Telson entire... . . . . Cebocaris 8. Coxa 2 much larger pan 1 eval covering Hf bout coxae 90 4 small, not covering anterior coxae. . . . . . . . Metacyclocaris
Some of coxae 1-2 or 1- 3 porered ener a 3 or i coxa 4 large) 5) oul
MARINE GAMMARIDEAN AMPHIPODA 301
9. Article 2 of pereopod 3 deeply indentured (fig. 1316) or with very long, simple posterodistal tooth; coxae 1-3 all small and covered by coxa 4.
Cyphocaris
Article 2 of pereopod 3 not deeply indentured, only coxae 1—2 reduced and COMELEMUDYACOKAC ES LOGLA Mie We aia eu) styrcliby sr hes il eea nls Meant AI ae ganna KO) NOsaSomespercopods premensile yi ism seep sega) ccna etnies fol) Baracehocnnie Pereopods simple ... . foc aaaunrasis ella!
11. Mandible lacking distinct nals or aria wonicclarminets amd unridged anallar. Cyclocaris
Mandible bearing triturative molar (fig. 126g) . . . . . . . Procyphocaris
KEY G
The genus ‘“‘Onisimus” auct. (=Boeckosimus, new genus, herein) has always been poorly defined and confusing (see Stebbing, 1906, key p. 9, couplet 14). Some of its species overlap with ‘““Pseudalibrotus”’ auct. (=Onisimus as senior synonym) to which I believe they should be transferred. The genus also appears in Keys J and M.
“Onisimus’”’ auct. comprises a series ranging from entire telson (as in some species of ‘‘Pseudalibrotus” auct.) to slightly split (as in Key J) to half split (in Paronesimus and Paratryphosites (= Hippomedon) in Key M). The proportions of articles 5 and 6 of gnathopod 1 vary slightly, enough to cross couplets in Key M.
1. Pereopod 1 formed into a subchelate grasping organ similar to a gnathopod
(ieanlG0D) sate olen sonia nse) el ey 2 Mndevoura andy Ensayara Pereopod 1simple ... . Sve eee
2. Upper lip formed into a lanes ibeel: iftke lobe wrens stronely projecting in front of epistome (fig. 125c) . ... . LNG Neg evan
Upper lip usually small, obscure, andlor scaroaly mroicctine in front of epistome (figs. 125g,n,0), or epistome and upper lip coalesced, upper lip
if lobate equalled in extent by lobe of epistome. . . . pte gaiecne Ml
3. Uropod 3: inner ramus less than half as ie as outer, invele 5 of male antenna 2 dilated (fig. 124j)}. .............. Parawaldeckia Uropod 3: inner ramus subequal to or more than half as long as outer, [article 5 of male antenna 2 not dilated, article 4 rarely dilated] . . . . 4
4. Gnathopod 1 with broadly transverse palm (fig. 129a) .. . . . Koroga Gnathopod 1 minutely but distinctly subchelate (fig. 129b) . ..... 5 Gnathopod ysimple (ig 129m) ta ne ee ey cree HeArticle 4-ofantenmar2 tumiducnen set ae ee iysianella Article 4 of antenna 2 slender. ..... =.=. +... .. . Paralysianopsis
6. Article 4 of male antenna 2 dilated. ......... .. . Lysianella Article 4 of male antenna 2 not dilated... . . ERM in phn Noes an ooeed ana
7. Uropod 3: peduncle expanded distally into a narrow, olavelike projecting siiveliige (fiicasl'322))) ines ee BN Nera AUR ATE I Uropod 3: peduncle simple, sabeyiindriedl Ge 1329) . . . . Lysianassa !
8. Maxilla 1 with two kinds of spines on inner plate. .......... 9 Maxilla 1 with one kind ofspineoninnerplate. .........-. .- 10
9. Inner ramus of uropod 2 constricted (fig. 117a). .. . . . [Arugella‘] Inner ramus of uropod 2 unconstricted. . . .. . . . [Pronannonyx']
10. Mazxilla 2: inner plate nearly twice as broad as outer . . [Shoemakerella ']
Maxilla 2: inner plate equal to outer in width . [Lysianopsis '] and [Aruga ']
1 Tam amalgamating all of these genera in Lysianassa, but Pronannonyz is questionable and also retained as a genus in alphabetical order.
13.
14.
15.
16.
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U.S. NATIONAL MUSEUM BULLETIN 271
Uropod 3: inner ramus vestigial orabsent. ... . Bios tania. euiey iy Uropod 3: inner ramus more than half as long as outers TFAMNUS eee eet Uropod 3: inner ramus vestigial, article 1 of antenna 1 flagellum 5 times as long as next article. .... . . . . . Onesimoides 2 Uropod 3: inner ramus absent or nprented Bye a spine, article 1 of antenna 1 flagellum not longer than next article. . . . . . . . . Paronesimoides Uropod 3: inner ramus vestigial, article 1 of antenna | flagellum not longer than wnextiarticle:: ia) ha ee see Saar nee Clepidectellags Gnathopod 1 simple. .. . oy eo et ae Gnathopod 1 distinctly, fyoueh Biten Prince pubehelate SS OLS Ry Dia Hy 1.7 Mandibular palp attached level with molar (fig. 126m), coxa 4 scarcely excavate posteriorly (fig. 117a) . . . . . Paralibrotus and Menigrates Mandibular palp attached well proximal of molar (fig. 126h), coxa 4 excavate posteriorly (fig. 119b) .... ia Se eiecu mM Rea inet tetany Boi ry 3 DLED) Palp article 4 of maxilliped tiny Gre, 128k) BG Sd tee oN TTT ONYX Palp article 4 of maxilliped well developed (fig. 128c). . . ...... 16 Epistome and upper tear together strongly produced and separated by a Siti ean ke ‘ . . . Aruga (in part) [—Lysianassa]
(and ORs [Pronannonyx] and see Adeliella) Epistome and upper lip together not strongly produced, and apparently
comlesced yaar Spelotel ah oak Lailvics ast Mules eauley eat ARAM ASI Maxillipedal palp 3- smowilaie ete ere imma eis 3 a. LS Maxillipedal palp 4-articulate. . . . SESE Oe 2 ot Nas eae Ne Maxilliped: inner plates obsolescent (Gis, 128)) . oa. oe Rerrierella Maxilliped: inner plates as long as palp (fig. 128g). . . . . . Normanion Gnathopod 1: palm excavate (mouthparts unknown) . . . Pseudambasia Gnathopod 1: palm not excavate .. . PUREE Ni io, oe) AU) Mandibular palp attached level with mole Ge. 126m) . Foyle) ag vee eaareammead| Mandibular palp attached proximal to molar (fig. 126h) or molar absent. 23 Maxilla 2 strongly setose (fig. 127p), coxa 1 visible . ..... 2... 22 Maxilla 2 poorly setose (fig. 127k), coxa 1 nearly concealed by coxa 2. Adeliella Mandibular molar small, conical, Cia: setulose, gnathopod 1 article 6 enlarge dime cae : I ee os oroga
Mandibular molar of eesti size, etimatioe’ gnalionod 1 not enlarged. “Pseudalibrotus’”’ (—Onzsimus) and ‘‘Onisimus”’ [—Boeckosimus (in part) | Rami of uropod 3 reduced in size, inner ramus about half as long as peduncle
Gare NAVE) gg . . Nannonyx Rami of uropod 3 as iene as or longer shea me danele (Ge 1324) . ese ue: Article 6 of gnathopod 1 very stout, enlarged (fig. 1291) ....... 25
Article 6 of gnathopod 1 small, rectangular (fig. 129d). some species of Orchomene and [Allogaussia] Mandibular molar of medium size, triturative, article 4 of maxillipedal palp
Shortaeeueiaas .. . . . Pseudokoroga Mandibular molar sell conical etnonetadl, eeralece article 4 of maxilli- pedal palpi loma:s ee Sy ieee ik Vik edn pets aiRu lem neSl aes a nen O ee
2 See also Adeliella. 3 See also Parawaldeckia kidderi (Smith) (p. 357), distinguished from Clepidecrella by extreme proximal location of mandibular palp.
MARINE GAMMARIDEAN AMPHIPODA 303 KEY H
These genera have gills bearing swellings or plaits. Since so many lysianassids have not been investigated for this characteristic and because gills of the following genera are not always clearly plaited the genera have been added to subsequent keys. Hence key H simply represents a quick way for the taxonomist to determine familiar genera with obviously plaited gills. If this key is unsatisfactory one must proceed to couplet 11 of the Basic Key. Many species of Hip- pomedon have indistinctly plaited gills. Gills of Socarnella are unknown.
1. Coxa 1 reduced and hidden by coxa 2 (fig. 123b) . . . . . . Schisturella Coxa 1 of normal dimensions and not hidden by coxa 2 . : Bilas aia) 7 2. Gnathopod 1 subchelate 3 Gnathopod 1 simple 7 3. Mandibular palp attached rexel! to mole Gin, 1267) . 4 Mandibular palp attached level with molar (fig. 126e) : 5)
4. Article 5 of male antenna 2 extremely tumid (like article 4 of fie 124h). Microlysias Article 5 of male antenna 2 slender or not relatively stouter than article 4. Orchomene 5. Mandibular molar imperfectly conical or attenuate, setulose, not ridged Giger 267) Ge .. . Anonyx Mandibular molar cylindrical or ncaboical: mdeed @e 126g) . Beene 6 6. Dactyl of gnathopod 1 very small, similar to dactyl of gnathopod 2, Alp article 4of maxilliped stout, short... . . . Pseudoanonyx Dactyl of gnathopod 1 and palp article 4 of meciliped long and claw-like. Hippomedon
7. Dactyl of gnathopod 1 setose or spinose on posterior edge (fig. 18072), upper lip poorly delineated (fig. 125g), mandibular palp attached level with molar. Ichnopus
Dactyl of gnathopod 1 not setose or spinose on posterior edge, upper lip strongly delineated (figs. 125a,c), mandibular ae attached distinctly
proximal) tojmolar =.) a eee site
8. Outer ramus of uropod 3 Drareiculates or eicompoccd of one syareicle and large terminal spine .. . SUES a er Ua emai aoe Ue SOCALNES Outer ramus of uropod 3 amieeticulates AEM Es ca Sera ie eee)
9. Telson emarginate; mandible long and narrow, malp aricley2 Thoin 5 times as long as article 1 and more than 2 times as long as article 3. . . Socarnella
Telson cleft more than halfway; mandible short and stubby, palp article 2 only 3 times as long as article 1 and about 1.5 times as long as article 3. Socarnopsis
KEY I
Genera of Key H are included here owing to possible confusion in Key G concerning gills. Koroga is repeated in case its telson is con- sidered as slightly cleft.
1. Coxa 2 concealing anteroventral corner and most of coxa 1 (fig. 1196), coxa 1
slightly or strongly reduced in size or not quadrate. ........ 2
304
10.
11.
12.
13.
14.
U.S. NATIONAL MUSEUM BULLETIN 271
Coxa 2 not concealing anteroventral corner of coxa 1 (fig. 117a), coxa 1 normal and quadrate or distally expanded . 8 Coxa 1 half as long as its article 2 (fig. 130k) . . Late, By aac ne eee Coxa 1 as long as or longer than its article 2 (fig. 180g). ....... 6 Gnathopod 1 subchelate (often ee OT MR haa 0 4 Gnathopod 1 simple 5 Lobes of maxilla 2 equally Broad or inner Tope: narrower “hema amie, “qacmadtion lar molar short, blunt, or weakly laminate (fig. 126e), inner plate of maxilla 1 with two setae... . . . . .Metambasia and Schisturella Inner lobe of maxilla 2 soul Tprond a than outer, mandibular molar conico- laminate (fig. 126f), inner plate of maxilla 1 densely setose (4 or more
Setaey te rs ... . . Aristias
Lobes of maxilla 2 not or eerecly. eesostin inner sae Orowdes than outer. Metambasia
Lobes of maxilla 2 gaping (fig. 1277), inner much broader than outer.
Aristias Maxilliped: outer plate lacking spines . ........ . . . Ambasiopsis Maxilliped: outer plate spinose .. . Peary les titty enc Telson deeply cleft (three fourths or aRGr), outer lobe of smassle 1 poorly spinose (fig. 127c), inner ramus of uropod 2 simple. . . . . Neoambasia Telson deeply cleft, outer lobe of maxilla 1 normally spinose, inner ramus of uropod 2simple.... . . . . . . Tryphosella Telson cleft one third, outer lobe Gr smosalla 1 nonnalls spinose, inner ramus of uropod 2 constricted .... . . . .Schisturella (—Pseudonesimus) Guuthopod Teimple) Ue) kh GE) aia a alt Gnathopod 1 subchelate .. . Be ice iran vin aR amaelsn iG: AL IL Pereopod 5 much longer than neetonod 4 GC A ae Gia ae Te hnopus Pereopod 5 not longer than pereopod4......... Boner gh tO) Inner ramus ofjuropod!2inciseds =) iis ae a I Soenrnideet
Inner ramus of uropod 2 simple. Waldeckia, Socarnes, and Socarnella (see Key P, couplet 20) Head lacking distinct lateral lobes, anterolateral cephelic margins straight
(fig. 121b); coxa 5 as long as coxa 4... .. . .. .. .Lepidepecreoides Head bearing lateral lobes, anterolateral ae margins bulbous or convex; coxa 5 shorter than coxa 4 .. 2 AGUA CR Nt ee Article 4 of antenna 2 very tumid Go 124k) . Lise a Ae iLysianella Article 4 of antenna 2 of normal dimensions. . . . Ste Aes sete) Les}
Mandibular molar large but not strictly Cabordain sien subconical or attenuated or basally bulbous and covered densely with setules (fig. 1267)
14
Mandibular molar large or of medium size, cuboidal, ridged, setules if rarely present very sparse (fig. 1269)... . Bae is auccee ILS) Article 6 of gnathopod 1 enlarged (fig. 1291), felcon apical DGtened Korsga Article 6 of gnathopod 1 not enlarged, telson deeply cleft. . . . . . Anonyx Uropod 2 with inner ramus strongly incised, epistome with extremely large, acute anterior process... . . .. . . . Tryphosites
Uropod 2 with simple inner ramus, eaions produced! “Tmetonyx”’ nobilis (Stimpson) (see Shoemaker, 1930) and Hippomedon Uropod 2 with inner ramus strongly incised, epistome and upper lip coalesced (not separated by notch from lateral view), apparent upper lip portion strongly produced and acute. ‘“Tryphosa’”’ carinata Schellenberg, 1926a
1 And see “Tryphosa”’ carinata Schellenberg, 1926a (couplet 15) and Orchomene (p. 353).
MARINE GAMMARIDEAN AMPHIPODA 305
KEY J
Both Koroga and Orchomene (=?Allogaussia) are repeated here, in the event their quantitatively expressed characters were overlooked in previous keys.
iy 1 @oxail nearly concealed by coxa 2-°%)0-0-0.0.5. 23.0... 2 4. Adeliella Coxa fully) visible.) 5 <7. Be ne Cer ee RR nO
2. Mandibular palp attached level with molar (fg. tea). Ns ae Wea Resale ra SUR Mandibular palp attached proximal to molar (fig. 1267). . .......5
3. Gnathopod 1 subchelate, inner ramus of uropod 3 as long as article 1 of outer. 4
Gnathopod 1 simple, imner ramus of uropod 3 vestigial. . . . Clepidecrella
4. Mandibular molar triturative........ “Onisimus”’ [= Boeckosimus] Mandibular molar simple... . .... . . .Menigrates
5. Gnathopod 1: article 6 not wider fae article 2) SRN Y: See O
Gnathopod 1: article 6 distally twice as wide as article 3 (fig. 1291). Koroga and Pseudokoroga (see Key G for distinction) 6. Gnathopod 1 subchelate ...... . . . . Orchomene [=? Allogaussia] Gnathopod 1 simple. ........ .. .. .. .Socarnella and Menigrates
KEY K
Gnathopod 1 is slender, the dactyl hidden in a tuft of setae or cirri (fig. 130m). Ichnopus bears resemblance to this group as a genus with a posterior or distal brush of setae on the dactyl. Pseudoanonyz is only marginally assignable to this group.
1. Gnathopod 1 stout and minutely and transversely subchelate or chelate, palp
article 4 of maxilliped stout and subclavate. . . . . . . Pseudoanonyx Gnathopod 1 slender and simple, or palm very oblique, palp article 4 of maxilliped slender and claw-like. . . . 2
2. Mandible lacking molar, article 6 of pereouods) 12 2 slightly longer har articles 4 and 5 combined (fig. 131c) and slightly prehensile.
Scopelocheiropsis
Mandible bearing molar, article 6 of pereopods 1-2 shorter than or equal to articles 4 and 5 combined (fig. 117a). . ... : A oho
3. Lower lip with major lobes a excavate aeinlie forming a medial gap dig: 125s) 5. . . . Bathycallisoma Lower lip with ede adles of major Nokes Elosely appressed ( ve WAR A) os e:
4. Gnathopod 2 minutely chelate (fig. 131f). ........ Bice He ameepeta) Gnathopodli2isubchelatenCipadoily)ire st ei ahkie nr Weert (i700
5. Epistome flat in front (fig. 1250) .. . Neon ei each et hee AT OUT Epistome grossly lobate in front (fig. 12510) .... . . . Scopelocheirus
6. Inner plate of maxilla 1 setose only terminally and subterminally, dactyl of gnathopod 1 not distinct from cirri (fig. 129m) . . . . Paracallisomopsis
Inner plate of maxilla 1 setose medially, dactyl of gnathopod 1 distinct from
cirri (fig. 130m) (see other characters under Paracallisomopsis diagnosis) . 7
7. Article 2 of gnathopod 1 linear, not glandular (fig. 130m), coxa 5 much larger than and partially covering coxa6.. . . . . Paracallisoma Article 2 of gnathopod 1 swollen, glandular fe 1299), coxa 5 not much larger than and not covering coxa6.... . ..... . . . Eucallisoma
306
tw
10.
14.
U.S. NATIONAL MUSEUM BULLETIN 271
KEY L Pereopods 1—5 with long dactyl folded back on an inflated article 6, thus prehensile (fig. 130d) .... . .. . . . Metacyclocaris Pereopods 1—5 normal, dactyls lnc anal path ticles unexpanded. . . 2
Maxilla 2: lobes gaping and/or inner much broader than outer (fig. 1277); mandible with molar, combined with nearly simple gnathopod 1 . Aristias Maxilla 2: lobes appressed and of similar shape, other characters not in combination. . . [MENTE ERM AGAS GEE RO. UE ya ne en am eS Mandible lacking moles (pat hopodl 1 Emplel SNE 4 Mandible bearing molar, gnathopod 1 subchelate Gun in iNfciambascom 5) Article 2 of maxillipedal palp as broad as or broader than outer plate. Ambasiella Article 2 of maxillipedal palp less than half as broad as outer plate. Ambasia Inner plate of maxilla 1 with one or two immense, falcate setae (fig. 127d). Hirondellea !
Inner plate of maxilla 1 with small, slender setae. . ......... 6 Maxillayisinner plates with tivetoramore Sevaciis sich) cat icicles enn) Maxilla 1: inner plate with three or fewer setae. . . . AR Ae seh tS) Mandibular molar immense (fig. 126d), inner plate of aT 1 setose only
terminally, article 3 of gnathopod 1 not elongate .. . . . Eurythenes
Mandibular molar slender but strongly projecting, inner plate of maxilla 1 setose medially and terminally, article 3 of gnathopod 1 elongate [see 2
species of Cheirimedon] . . . . Se IE ene tal ee aralice lla
Inner ramus of uropod 2 Coneenictca AGRA URM ENS SER cE Mom tM mast ti ph i a <i v8)
Inner ramus of uropod 2 simple. . .. . | Vase Rates: Weal Gnathopod 1 slightly chelate, epistomal margin daneoyenteallly elongate.
Aristiopsis
Gnathopod 1 subchelate or simple (Metambasia), epistomal margin dorso-
ventrally compressed even though lobate (normal) ........ 10
Mandibular molar poorly developed, weakly triturative or fuzzy, palp attached proximal to molar, gnathopod 1 subchelate.
“Orchomenella’”’ groenlandica (Hansen)
Mandibular molar strongly triturative, palp attached level with molar,
gnathopod 1 subchelate . .. . .. . . . Schisturella Mandibular molar weakly tritur ative or 4 fiom bat very large, palp attached
level with molar, gnathopod 1 simple. . ..... . . . Metambasia Outer ramus of uropod 3 uniarticulate . . .. . . . . . . Tryphosoides Outer ramus of uropod 3 biarticulate ........ Sok een Lez, Mandibular molar lacking setules =) 2.5 24 5 2. hepidenecreunace Mandibular molar bearing setules. . . . Berea ists alot AL)
Gnathopod 1 with article 6 enlarged and dieealle qantas, nearly twice as broad as article 2, palm protuberant, long, and slightly chelate. Cheirimedon latimanus Sars, 1895 Gnathopod 1 with article 6 unexpanded, palm not chelate . .... . 14 Head very small,? epistome and upper oe weakly developed and incon- spicuous. . . .. . . . Uristes [—Uristoides] Head of normal size, , aotone andl | upper lip strongly developed, epistome generally shightly lobateici) 2h) os ie fee rei Ona ra Se eee to)
1 See note, Key N, couplet 23 (p. 310). 2 Occasionally head ventrally extended and covering base of antenna 2 as in ‘‘ Uristes’”’ lepidus J. L. Barnard (1964a).
15.
10.
Il,
12.
13.
MARINE GAMMARIDEAN AMPHIPODA 307
Article 7 of gnathopod 1 with accessory tooth. some species of Tryphosella [—‘‘Tmetonyzx’’ auct. ]
Article 7 of gnathopod 1 simple . . . . Tryphosella [—‘‘7ryphosa”’ auct.| KEY M Palp of mandible attached proximal to molar (fig. 1267) . .. . A
Palp of mandible attached level with molar or molar absent (figs. 126j,k) . Article 1 of antenna 1 nee produced distally and dorsally La
Gigs 4d) es irks 2 . . . . . Lepidepecreum Article 1 of antenna 1 yearly or nee menaced discal, not dorsally crested
(tags WOO A So aides San ORMOND MMU ae LUC rss IOUS DE Nadel ts hanciteareD Gnathopod 1 simple andl oun sai ed ass detec Suse Meee Gnathopod 1 subchelate (or very large aed coominely sfimoralls) Seale Vaan eee Outer ramus of uropod 3 uniarticulate ...... =... . .Socarnopsis Ourerramus,omunropodsoppiarpiculave sii ly nceyle Westie nel euiu ian lesa. O Inner ramus of uropod 2 constricted. ....... .. . . .Socarnoides Inner ramus of uropod 2 simple. . . . .6
Telson cleft one third of its length, Soma Doris not onomninend Ge 1254), articles 2-3 of antenna 1 peduncle half as long as article 1 of primary flagellum .... . . . . .Menigrates
Telson cleft faleeete or tore mreoucen rpms anouainent (fig. 1250), articles 2-3 of antenna 1 as long as article 1 of primary flagellum.
Waldeckia and Socarnes (see Key P, couplet 21)
Article 3 of gnathopod 1 elongate and slender as in typical lysianassid gnathopod 2, articles 5-6 also elongate and slender (fig. 1297).
Pseudorchomene Articles 3,5,6 of gnathopod 1 stout, not relatively elongate (fig. 129d) . .8 Article 4 of antenna 2 very tumid, article 5 slender . . . . . . Lysianella
Article 5 of [male] antenna 2 very tumid, stouter and longer than article 4. Microlysias ! Neither of articles 4-5 of antenna 2 expanded . . i Articles 5 and 6 of gnathopod 1 not immensely cnlemaed Ge. 1294) (also check Lepidepecreum). Orchomene [=Orchomenella] and juveniles of Pseudokoroga
Articles 5 and 6 of gnathopod 1 immensely enlarged (fig. 1291) . . . .10 Maxillae slender (fig. 1271), maxillipedal palp article 4 short, mandibular molar triturative ... . . . . . Pseudokoroga Maxillae stout (fig. 1270), ranselltraadlel sopalfay ontniells 4 elongate, mandibular molar nontriturative . .. . ee A ECOROZa Mandibular molar laminate, ponical nee euiotcely Grince simple, lacking ridges, often weakly setulose, or molar absent . ... . sete GULP
Mandibular molar cuboidal or columnar, surface usually beseine conspicuous ridges and spines, or distinct punctations, or densely setulose if not fully cuboidal and slightly attenuate (figs. 126e,9,7,m) . . . arcuate ser 0)
Gnathopod 1: article 6 very large and expanded, more ‘her 3 times as long as article 5 (fig. 1291). . .. . . . . . .Koroga and Cheirimedon
Gnathopod 1: article 6 small, never more than 1.5 times as long as article 5. 13
Inner lobe of maxilla 1 bearing five or moresetae.......... 14
Inner lobe of maxilla 1 bearing two or fewer setae. ......... 15
1 Possibly article 5 of Microlysias has been misidentified.
308 U.S. NATIONAL MUSEUM BULLETIN 271
14.) Gnathopodsisimple) 22s Meeat itey el) on) Mice eae eee Ce lia Gnathopods subchelatee ane ey one ee aralicella
152 sGnathopod Lsimipleyy ia eau aac: Saeeayeia bel ant ean ipa Guile eres Meatee ammaa Gnathopod 1 subchelate. ... . SBE etsy aia. lve 11 7/
16. Rami of uropod 3 subequal to each onhen in Tenet» . . . . . Ichnopus ? Inner ramus of uropod 8 vestigial, scale-like. ... . . . . Clepidecrella
17. Head globular, maxilla 1 with sickle-shaped setae on inner lobe (fig. 127d). Hirondellea
Head other than globular, maxilla 1 with normal setae on inner lobe. . 18
18. Maxillipedal palp article 4 reduced in size (fig. 128a). . . . Centromedon Maxillipedal palp article 4 of normal size. . . . Set) tera ee peeteas LG)
19. Coxa 1 normally quadrate, untapering, ganhelion leasing Tater lobes or strongly truncate anteriorly (fig. 121b), coxa 5 as long as coxa 4.
Lepidepecreoides
Coxa 1 normally quadrate, untapering, cephalon with lateral lobes, coxa 5 much shorter than coxa4 . . . . . proceed to 24
Coxa 1 tapering distally, slightly “-qnosioncdl ‘aaellon with lateral lobes, coxa 5 much shorter than coxa 4 . . Lepidepecreopsis and Tryphosella
20. Coxa 1 very slightly triangular, tapering distally (figs. 129c,k) .... 21 Coxa 1 quadrate, or rounded, expanded or unexpanded distally (figs. 129
CH TOS aU Cie Ne nie MATAR DI NMR cad Ye 3 Say
21. Outer ramus of meopad 3 aantieulate SAR UGA NsERE AN eh aaa iunyphocuides Outer ramusofuropod:s biarticulates ~ 1. 4.40070) fai) oO) een ee
22. Head small, epistome and upper lip inconspicuous. Uristes ? [= Uristoides] ?some species of Cheirimedon Head large, epistome and upper lip prominent from lateral view .. . 23 23. Dactyl of gnathopod 1 with inner accessory tooth. some species of ‘‘Tmetonyx” auct. [= 7 ryphosella] Dactyl of gnathopod 1 lacking accessory tooth. Tryphosella 3 [=‘‘Tryphosa’’ auct.] 24. Telson not longer than broad, always cleft less than halfway. Rifcus and Boeckosimus [=‘“‘Onisimus’’ auct.]
Telson longer than broad (or if short, then cleft halfway or more). . . 25 25. Gna thopod jh Ssimiple nity ay spin) ease mnie Neola UUs OM Rr he DO Gnathopod 1 subchelate .... : Baer eA AAS)
26. Gnathopod 1 perfectly simple, srecallinaedtal dace samntied. - Menigratopsis Gnathopod 1 very slightly subchelate, palm small, maxillipedal dactyl
clongate ea wn. Bou Minuet eny ese BONA MES BPA TF
27. Uropod 2 with simple sh innerramus .. . ie iparoncsitnns (in part) Uropod 2 with incised inner ramus . i onmonyace cicadoides Stebbing, 1888
28. Mandibular molar densely covered with setules . .. . BS ei ia KG) Mandibular molar ridged or weakly punctate, setules aqanernllly absent or inconspicuous . . . pes ler oO)
29. Coxa 1 expanded dligiclt. npenion dated of rametinonadl 1 rigireilby lacking accessory tooth, upper lip slightly or strongly protruding in front of epistome, inner ramus of uropod 2 often incised, palp article 4 of maxilliped UIT GUE OLIN pe ie sane eee) Rul ave CU ey ee UIA Cin su aN Meta Fis eee ANT O TTLVAXG
2 Menigrates angustipes Gurjanova (1962) may key to this position inadvertently but note diagnosis of Ichnopus does not otherwise fit Menizrates. 3 See also ‘“‘Orchomene’’ takoradia J. L. Barnard (1961); ‘‘Uristes’”’ velia J. L. Barnard (1961).
30.
Bl,
32.
33.
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MARINE GAMMARIDEAN AMPHIPODA 309
Coxa 1 unexpanded distally, unguiform dactyl of gnathopod 1 bearing inner accessory tooth, epistome slightly protruding in front of upper lip, inner ramus of uropod 2 simple, palp article 4 of maxilliped unguiform.
type-species of Tmetonyx
Coxa 1 slightly expanded distally, dactyl of gnathopod 1 with accessory tooth but reduced to size of dactyl on gnathopod 2, prebuccal parts apparently like Anonyz of triplicut 29, inner ramus of uropod 2 simple,
palp article 4 of maxilliped tumid, subclavate. . . . . . Pseudoanonyx Dactyl of gnathopod 1 very small and similar in size and morphology to dactyl of gnathopod2 ... . . . . . Pseudoanonyx Dactyl of gnathopod | long, alanis, thus aan Seng santana leas Wea inaeae Article 5 of gnathopod 1 longer than article6 .. . a hea Article 6 of gnathopod 1 longer than or subequal to mols 5 SG ks GO Head without distinct lateral lobes, anterolateral margin convexly truncate, coxa Daslongascoxa4..... . . . . . Lepidepecreoides
Head with small but distinct lateral lobes, .¢ coxa 5 shorter than coxa 4. Hippomedon [= Paratryphosites, ? Paracentromedon, ? Elimedon| Outer ramus of uropod 3 uniarticulate ...... .. . . Tryphosoides Outer ramus of uropod 3 biarticulate . . . . te aie aa OA Articles 5 and 6 of gnathopod 1 subequal in fenet “atiels 6 not saoaacled!, palm obsolescent ... . . . .[Paracentromedon] and Paronesimus Article 6 of gnathopod 1 large, distally expanding, more than 3 times as long as article 5 (fig. 1291), palm strong ... . . . . .Cheirimedon (type)
KEY N
All lysianassid genera with coxa 1 distinctly reduced in size and/- or often largely hidden by coxa 2 are assembled here for ease of com- parison. Those species of Key F, in which both coxae 1 and 2 are small and hidden by following coxae, are not included. See also notes on
Stomacontion. 1. Pereopods 1-5 with long dactyls folded back on inflated article 6, thus pre- hensile (fig. 130d)... . . . . . . . Metacyclocaris Pereopods 1-5 normal, dactyls shen’ ail sein articles unexpanded. . . 2
ou
Upper lip strongly lobate and projecting in front of epistome (figs. 125c,d) . 3
_ Upper lip scarcely or not projecting in front of epistome, latter often large
and dominating upper lip (figs. 125h,7,k) or epistome and upper lip not distinct from one another . US Rae 8)
Coxa 1 half as long as its second paticle Ge. 130) . SOs IG ety Boren Coxa 1 as long as or longer than its second article (fig. 1309) . Bi Ree Nn Ie) Gnathopod 1 subchelate even ee weakly... . . . . Schisturella CGnathopodi la simples. (9 4) .0). . . . . . Metambasia Outer plates of Se lacking spines, “messi 1 with seven spines on
GQUIEE OND 5 6 4 es .. . . . Ambasiopsis Outer plates of maillyped spinose, masa 1 nora with 9+ spines on outer
Ko) oveypre i enAe Oe .. . . . Schisturella Outer plates of anenclliaed spinose, peceilln 1 ant spines on outer lobe
REGUCeGtOptiVe; 4.2. ..... . . . Neoambasia Mandibular incisor walthicleretiarta (fig. 1269). A rieh islet La ceed ea Le NGO END TE Mandibular incisor not dentate... . 8
Mandible lacking molar, gnathopod 1 cnetnte: seule 2 of Dercopad 3 deeply imdentunedi (fies SND) ae Wo re ol oes o Podoprion
310
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U.S. NATIONAL MUSEUM BULLETIN 271
Mandible with triturative molar, Chia 1 subchelate, article 2 of pereo-
pod 3 not deeply indentured. ... . ... . . Valettiopsis Lobes of maxilla 2 broadly separated, inner lobe aoineine medially and of
different shape or much broader than outer lobe (fig. 1277) . . .... 9 Lobes of maxilla 2 appressed and similar in shape (fig. 12717)... .. . 10 Telson cleft... . . . Ambasiella and Aristias (see Key O, couplet 2) Telson entire... . . . Perrierella Eyes and panenopods 1 sarventtallly developed Ge. 123) . 2 eit GD Eyes and gnathopod 1 not together powerfully developed. . .... . 13
Gnathopod 1 with complete torsion distal from article 4 (fig. 130h), dactyl and palm thus posterior, mandibular molar needle-like (fig. 126b),
telson searcely longer than broad (fig. 128w) . . . . . Trischizostoma Gnathopod 1 untwisted, normal, mandibular incisor broad, flat, telson elongate, muchlongerthan broad .. . Berita Wer 74 Gnathopod 1 chelate, palm thus deeply oats (Ge. 1296) . . . Opisa
Gnathopod 1 cheliform only, palm convex and protuberant (fig. 1291). some species of Cheirimedon
Palp article 4 of maxilliped vestigial (figs. 128d,e,7) ......... 14 Palp article 4 of maxilliped well developed, claw-like. ........ 17 Lobes of maxilla 2 strongly setose, telson deeply cleft... ..... 15 Lobes of maxilla 2 with one and four setae, respectively (fig. 127k), telson
scarcely.) cleft ei adi cult Gre Ri MAAC UV aa 9) REA ep aerate sme Mem@Ar elke lta) Mt pal ley TL waaeloy VOM Gh ee are oh aka ey oo be eT A eG Maxilla 1 lacking palp ... . Prot! feoutinite to couplet 21
Epistome and labrum aoelenead, iabral aonvion not projecting (epistome often projecting), mandible without molar . Ambasia and Ambasiella Epistome and labrum separated by notch, labral portion slightly projecting, mandible with molar .. . . . . Centromedon Mandibular molar distinctly agounedl al riiecal (Ge, 1269) ii se ees Mandibular molar untoothed and unridged (fig. 1267), or ridges forming marginal girdle only (fig. 126d), often fuzzy, or molar absent ... . 19
Inner ramus of uropod 2 incised, outer ramus of uropod 38 biarticulate. Aristiopsis and Schisturella !
Inner ramus of uropod 2 simple, outer ramus of uropod 3 uniarticulate.
Tryphosoides Gnathopod 1 chelate (chela and gnathopod slender) . Podoprion and Euonyx Gnathopodt subchelaterorsimyple sit Wai ye ins e o) cele nua an nee, () Gnathopod 1 simple. ... . Mapes Nh bee fie 24dl Gnathopod 1 subchelate (often ofeawoteely <0) or r chelate . Rees ret ey hey) 1) Anteroventral corner of coxa 4 acutely produced ....... . Vijaya Anteroventral corner of coxa 4rounded or quadrate. . ....... =. 22 Article 2 of antenna 1 half as long as article 1 ... . . . . Amaryllis Article 2 of antenna 1 fully as long as article 1 . . . . . Bathyamaryllis
Inner plate of maxilla 1 with one or two immense falcate setae (fig. 127d). (cf. Cheirimedon pectinipalma K. H. Barnard, 1925) Hirondellea Inner plate of maxilla 1 with normal setae 24 Inner plate of maxilla 1 with five or more setae Ganlts) Inner plate of maxilla 1 with three or fewer sctae .
ww we “JI or
| And see Key L, couplet 10, for ‘“Orchomenella’’ groenlandica (Hansen).
25.
26.
27.
28.
29.
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MARINE GAMMARIDEAN AMPHIPODA 311
Setae of inner plate on maxilla 1 mostly terminal, mandibular molar ex- tremely broad even though laminate and subconical, with girdle of ridges or setules (pleonite 3 with dorsal notch), mandible when flattened, with deep S-shaped insinuation between palp and incisor (fig. 126d).
Eurythenes
Setae of inner plate on maxilla 1 medial only or medial and terminal, man- dibular molar ‘‘absent”’ or long, narrow, conical and lacking distinct girdle (apparently pleonite 3 dorsally smooth), mandible when flattened
with shallow coneavity between palp and incisor. . . . piteg Nau a7A0) Coxa 1 longer than broad, ovatorectangular, as long as its prorele 2, lobes of maxilla 2 equally broad. ... . . . . . . Paralicella
Coxa 1 as broad as long and auipaniadiante, andl oni wo thirds as long as its article 2, inner lobe of maxilla 2 much broader than outer (lobes often
ORNS OM OVER) Serer) Gaara Pape Cee hens Ure not PATISUAS Mandibular molar ineaine aetulese ... . . . . . . . Lepidepecreopsis Mandibular molar strongly setulose . .. . St Nv Geel! mae a Ao} Head small, epistome and upper lip small antl inconspicuous . . . Uristes Head of normal size, epistome and upper lip a and conspicuous, epistome
usually slightly iopane ir eelaa ESE NEI ROD VNB EA Gah a Ce ih Pam a ae)
Article 7 of gnathopod 1 with accessory “oot some species of ‘“‘Tmetonyx” auct. [= 7ryphosella]
Article 7 of gnathopod 1 simple . . . . Tryphosella [=‘“7ryphosa” auct.] KEY O
Pereopods 1-5 prehensile (fig. 130d)... .... .. . . Metacyclocaris
Pereopods 1—5 not prehensile. . . . . Py eRe aise
Lobes of maxilla 2 gaping, and/or inner ane accel proaden than outer (fig. 1277), mandible with molar and gnathopod 1 nearly simple. Aristias Lobes of maxilla 2 appressed and of intersimilar shape, characters not in
SimMilarmcomtoimatiomexts joke eis teAuiery arma Pernciie EDS Poa hRiheeetns. ee Gnathopod 1 simple .. . gripes, suk A TR Se kd Aree ema sca To ee Gnathopod 1 subchelate, even einonen WwW DT BANG Oe se a ee Ser Mandible lacking molar (or if rudiment present, palp aiiecinad Sosa to
TOMOMENE)) gs 6 oe i eV het tier on SAeU NR Tae ae Fad Toe (Wy SA let coca ds bese tad) Mandible bearing olan ue ed ened a)
Article 2 of maxillipedal palp ec) ‘ahem ell? as locale as outer “ile. Centromedon and Ambasia Article 2 of maxillipedal palp much broader than outer plate . Ambasiella Outer plates of maxilliped with spines. . . ... . . . . Metambasia! Outer plates of maxilliped lacking spines. . . .. . . . . Ambasiopsis Inner plate of maxilla 1 with one or two immense faleate setae (fig. 127d). Hirondellea ?
Inner plate of maxilla 1 with small, slender setae. ..........8 Inner ramus of uropod 2 dorsally notched or constricted. . ...... 9 IMnMereramUusyOL UTOpOG..2) simple wy Meade tee sey mend igeumeets ta) LoL Gnathopod 1 subchelate. .. . A UIMOLU URC SIE ES Mee APR gh iti ti Seda WC] Gnathopod 1 weakly chelate Ge. 1292). Po en 2 BAe VAPIstiopsis
Coxa 1 tryphosid in outline (fig. 129k). “Orchomenella’’ groenlandica (Hansen)
1 See Schisturella diagnosis (p. 361). 2 See note, Key N, couplet 23.
312
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U.S. NATIONAL MUSEUM BULLETIN 271
Coxa 1 hemioval or subtriangular (fig. 129c). Schisturella [= Pseudonesimus]
Outer lobe of maxilla 1 degenerate and Sane distally, bearing only
five or six spines. . . BEE ean) mien cam angel Outer lobe of maxilla 1 aun) distill bearing seven ormore spines. . .13 Apices of maxilla 2 lobes densely setose, with nine or more setae each. Neoambasia Apices of maxilla 2 lobes sparsely setose, inner with one seta, outer with about four setae (fig. 127k)... . . . . . . . Adeliella Medial edge of inner lobe of maxilla 1 fully aeiase, enfticlle 3 of poenoned 1 elongate, as long as article 6..... . . . . . Paralicella Only distal end of inner lobe of maxilla 1 aeines article 3 of gnathopod 1 not elongate... . Pua Sees
Mandibular molar an immense, [mend panier seumlese, girdled lemma (fig. 126d), article 1 of antenna 2 enlarged and circular in adults (fig. 1247), inner plate of maxilla 1 with five or more setae. . . . . . Eurythenes
Mandibular molar of medium or small size, cuboid, subspheroid, or conically laminate, article 1 of antenna 2 not subcircular or eae inner plate of
maxilla 1 with three or fewer setae. . . Heat es UNS alo Article 4 of maxillipedal palp vestigial (fig. 1284) . . . . .Centromedon Article 4 of maxillipedal palp not vestigial. . .. . Reet a ICG)
Mandibular molar very small, sublaminate, conical or ‘hava, article 6 of gnathopod 1 enlarged (fig. 1291). some species now assigned to Cheirimedon Mandibular molar of normal size and cuboidal dimensions, article 6 of gnatho- pod small es Atte artes rear le Gh Outer ramus of uropod 3 cinfantenlate, Tedibulen molar nideed) Tryphosoides Outer ramus of ee 3 biarticulate, mandibular molar unridged but setulose;. 9.23 Sia Sabin ttc lke Epistome from tera view oxiondlins anh fareher hen nel projecting convexly in front of upper lip (fig. 1251). Tryphosella (‘‘Tryphosa’’? auct. and some species of “‘7'’metonyx’’ auct.) Epistome small, short, unprojecting, occasionally dominated by upper lip (figs. DEO) Ne J NSE reek) Upper lip and epistome sane Mreoncpicnee Ges. l2soa)) some species of Uristes
Upper lip dominating epistome (fig. 125n) (head ?large, coxa 1 an elongate
triangle in type-species, variable in others). . . . . . . Ambasiopsis KEY P
Gnathopod! I subchelateseven minutely.) 915.2240 ee Gnathopod 1 simple. . . . Reimann ws cous ld Hither article 4 or 5 of smieama 2 Fer nnd (Ge 124k). Moe Sis 4) ch i@) Neither article: 4 or 5) of, antenna: 2 tumids jos ese shee Broad epistomal lobe dominating labrum. . . .. . . . . . Microlysias Labrum strongly projecting in front of epistome ... . . . . Lysianella Epistome and upper lip amalgamated even though their parts produced,
parts not separated by notch (fig. 125k). . .. . Bee even 0) Epistome and upper lip separated by deep notch (figs. 125¢ if) Bina parity Ce
Maxillipedal palp 3-articulate, peduncle of uropod 3 elongate, nearly as long as pleonite 4 and peduncles of uropods 1-2 and longer than rami of uro- DOGS ess Sr Ee SERS Tet as sen Ue re un ove ea NOT STRATE TY
10.
11.
12.
13.
14.
15.
16.
21.
22.
MARINE GAMMARIDEAN AMPHIPODA 313
Maxillipedal palp 4-articulate, peduncle of uropod 3 short, subequal to pleonite 6 and much shorter than peduncles of uropods 1-2. ... . 6 Epistomal portion of prebuccal complex slightly produced, or flush, inner ramus of uropod 2 simple, inner ramus of uropod 3 half or less as long as outerramus... . ... . . Nannonyx Labral portion of erepuccall someon neately) mrodced! inner ramus of uropod 2 constricted, inner ramus of uropod 3 exceeding three fourths
length of outer ramus. .. . . . . . . Paralysianopsis Inner ramus of uropod 2 gemsanciedl Ge 131d) Ly CN ol eee oe Mike em Mir eet Aor ) Ininersramus of Uropodi2 simples sgn fea eye ete es ean Coxa 1 normally quadrate . . . . . . Pseudokoroga
Coxa 1 tapering distally and Partially elton ie coxa 2. “Orchomenella”’ groenlandica (Hansen) Article 1 of antenna 1 dorsodistally crested or toothed (fig. 124d), body often crested dorsally. . .. . . . . . . Lepidepecreum Article 1 of antenna 1 simple, body spoon or wiesios only on pleonite 4. 10 Article 3 of gnathopod 1 normally short and stout, articles 5 and 6 normally
stout (ig. 129d)... . . . . Orchomene Article 3 of gnathopod 1 dloneste andl Bender like osnls 3 of prerhonod Ds
articles 5-6 elongate and slender (fig. 1297) . . . . . Pseudorchomene Telson entire or emarginate ... . dg Rami etak RS US es Ae a aK A Pst NL Telson cleft one third or more .. . cihike Bias) Sr ae ne ge LD) Inner ramus of uropod 3 half or less ironed af Outer Ms Parawaldeckia Inner ramus of uropod 3 three fourths or more length of outer . . . . 13
Upper lip prominently lobate, occasionally epistome also lobate (fig. 125c). Lysianassa, ?Socarnella, [Pronannonyx]
Upper lip not prominently lobate ... . Slee Santee We Telson distinctly longer than wide, inner ramus ai moped 2 constricted.! Parambasia Telson not longer than wide, inner ramus of uropod 2 simple.
Menigratopsis Article 2 of pereopod 3 deeply indentured (fig. 131b), article 3 of gnathopod 1 elongate and slender like article 3 of gnathopod 2... . . Glycerina Article 2 of pereopod 3 with small teeth or none, article 3 of gnathopod 1 not elongate .. . Aad aces eat ALG) Coxa 1 tapering aictally or cnonrened andl ear ial frdden by coxa 2 (Aig eI Obie eae aes Ane ba cleat a bh Coxa 1 large, subquadrate or pearcely tapering, riety Sr eiples icon Van cl Ka) Palp article 2 of maxilliped half or less as broad as outer plate . . Ambasia
Palp article 2 of maxilliped as broad as or broader than outer plate. Ambasiella Inner ramus of uropod 2 constricted... ...... . . . Socarnoides Innerjranis ofirepod 2rsimple i. fv je) ol ie) Cena tyae a (LO Outer ramus of urepod Ss) umiarticulate i.) Gena sa ee 20 Outer ramus of uropod 3 biarticulate. . . . . iimedclstneee ly Pall Palp article 2 of mandible about 3 times as lone as eral 1, article 3 two thirds length of article 2... .°. . . . . . Socarnopsis Palp article 2 of mandible about 5 Aime as fone as article 1, article 3 about halfas lone as article 2,204) 0 ks. wale ect ye.) Socarnella Gillsistrongly:plaited) (figs 2s). 5 ot ye ee Socaknes Gills minutely ridged or smooth . 22
Telson about as long as wide, cleft less (han hear pills suave
Menigrates Telson about twice as long as wide, cleft more than halfway, some gills with basalzaccessoryslobes 8 24) Win ay ye Waldeckia
! Assumed in type-species.
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U.S. NATIONAL MUSEUM BULLETIN 271
KEY Q Mandibular molar distinctly triturative and usually columnar, cylindrical or cuboidal, bearing ridges or cusps for grinding (figs. 126d,g,m) . . . . 2 Mandibular molar not triturative or absent, often setulose or bearing a few articulate spines (figs. 126e,h,7,j) . . . Bete Nance a 17 Coxa 1 noticeably reduced and partially nvidldlen onl coxa o or coxa | tapering Or wrlenoepene 55 4G spe! SLA aie cane eam Coxa 1 large, visible, nacional ¢ or alistetlse spared. Si ei lela a Mandibular molar with triturative aac molar immense and not columnar (ficemlt2 OG) eee . . . . . Eurythenes
Mandibular molar nor anol aoitnnmen or 2 adie raped and medium in size. 4 Telson emarginate or cleft one sixth or less, maxilla 2 with setae reduced to
One, to fours pernloben (igen 27/1) i weenie .. .. . Adeliella Telson cleft one third or more, maxilla 2 si ‘iipibes strongly setose CATR AED CSN cS ae eeuG Ue MAUNA Og 3 | &
Inner ramus of uropod 2 simple. Tryphosella, Tryphosoides, and ‘‘Pseudotryphosa’”’ (?= Uristes)
Inner/ramus) of uropod!2) constrictediy 4) ey sed eee ee Epistomal region not dorsoventrally elongate. . . . . . . . Schisturella Epistomal region dorsoventrally elongate . ..... . .. . . Aristiopsis Gnathopod di simple wh esa ea eS Gnathopod 1 subchelate. . . . Be dessa Hessen el 1h G) Telson as broad as long, oremnate BAI a Vonicinmele Pecudaieroeial Telson longer than broad, deeply cleft . .... . . . . Menigratopsis
Telson as broad as long, cleft halfway. Rifcus and Paronesimus (sec diagnoses) Inner ramus of uropod 3 vestigial or absent ane 132k), always less than
half length of outerramus... . Rete Manta NA i oT) Inner ramus of uropod 3 subequal to outer 7 LATO See ibis eee ihie dk! Inner ramus of uropod 3 present, article 1 of both flagella of antenna 1
elongate and that of accessory flagellum expanded . . . . Onesimoides Inner ramus of uropod 3 absent, article 1 of both flagella of antenna 1 of
normal size .. . . 2 a. ss a ss . . . Paronesimoides Accessory flagellum 2- Perle: eRe eM ta erie gg!» LLP Accessory flagellum 3+--articulate . . . Maer eee oh lee Spines on outer plate of maxilla 1 Poomeed 6 eae or six. . . . .Rifcus Outer plate of maxilla 1 with nine or more spines. . . a oe nae ie) Hand of gnathopod 1 enlarged and expanding distally . Ponenimedan Gane Hand of gnathopod 1 of normal unexpanded dimensions. . . Tryphosoides
Telsonic apices with seven or more spines each (fig. 128s). Hippomedon [= Paratryphosites] Telsonic apices with three or fewer spines or telsonentire. ..... . .15 Pelson cleft jhaliway ermorel oo pani ais. th See erie dae Telson cleft one third or less. Boeckosimus and Onisimus [= ‘“‘Onisimus’’ and ‘‘Pseudalibrotus’’} Inner ramus of uropod 2 simple, epistome unproduced and slightly dominated by labrum (from lateral view) (fig. 125n). Hippomedon ! and [Paracentromedon] ! Inner ramus of uropod 2 deeply constricted, epistome acutely produced ig 12Gb) i as ina ae a co Ge Oe an a ie Ree Gu nM OSITes
! And see Cheirimedon femoratus (Pfeffer) and C. fougneri Walker distinguished from these genera by the transverse palm of gnathopod 1 (sce pp. 334-335).
17.
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MARINE GAMMARIDEAN AMPHIPODA 315
Gnathopod 1 simple?. . . By Pt od eae ey cua LTH Ga crt ont Ame ne) Gnathopod 1 distaois pubchelate ra eas Brea RUC Coxa 1 reduced or distinctly eerie from nase aul Sian strongly hidden lOs7 COkG Boo 65 co Sa tes eee! Re) Coxa 1 large, TaviemeEe: aml not fiddent fener if resent occurring in onlysdistalithird of ‘coxays 2 2a Pia oe hf ud ae seisunae) vase 26 Rereopods) prehensile) sive aie sy eis se eh ee victacelacaris Pereopods simple. . . Ha) We pbuivees see «20 Maxillipedal palp with iheeel catiotes telecon eaaie) . .. . . . Perrierella Maxillipedal palp with four articles, telson cleft. . ......... .2!21 Maxillipedal palp article 4 vestigial (fig. 128a). .......... .22 Mazxillipedal palp article 4 well developed and claw-like. . . .... . .238 Mandibular molar present, upper lip and epistome coalesced, epistomal region strongly produced ..... ... . .Centromedon Mandibular molar absent, upper lip and soitome separated by notch, upper lip slightly protruding .. . . . . .Ambasia Inner plate of maxilla 2 much broader Aten oniar plate aaa usually gaping mediad. .... ... . . .Aristias Plates of maxilla 2 seemed in Raden nal not pironely, CApINGeees eet. 1. 24: Upper lip and epistome not differentially produced, small and inconspicuous. Uristes Upper lip projecting as lobe in front of epistome (from lateral view) . . 25 Outer plates of maxilliped lacking spines, gnathopod 1 weakly subchelate, coxa 1 nearly as long as article 2 of gnathopod 1... . . Ambasiopsis Outer plates of maxilliped spinose, gnathopod 1 perfectly simple, coxa 1 about half as long as article 2 of gnathopod1 .... . . Metambasia Inner plate of maxilla 1 fully lined medially with eight or more setae. Alicella Inner plate of maxilla 1 with three or fewer terminal setae . . . . . . 27 Inner ramus of uropod 3 vestigial, less than one third length of outer. Clepidecrella Inner ramus of uropod 3 subequal to outer. . ......2.2.2.2.~. 28 shelsonicleftxone| third or more! 292 >. S3ee si. Bt. aie. ee eee 29 Telson entire or emarginate . . ahh. coer 0) Gnathopod 1 perfectly simple, pieader “oiibgiviitemn, dacel strong and bearing posterior setal brush near base (fig. 1307) . . . . . . Ichnopus Gnathopod 1 perfectly simple, slender, dactyl strong and lacking posterior setal brush. .... . . . . . Menigratopsis
Gnathopod 1 weakly pubele lates ee raediuen sienna, dactyl strong but lacking posterior setal brush although bearing small accessory tooth.
Paronesimus Labral region of prebuccal complex with strongly acute anterior lobe [telson longer than broad]. ...... . . . . . Paralysianopsis
Labral region of prebuccal complex aieh convened rounded keel or unpro- duced [telson as broad as long in Paralibrotus only].
Menigrates (in part), Paralibrotus
Coxa 1 reduced or distally tapering from base and often strongly hidden by
COXA) 275 a ee SAS OES, Oe Coxa 1 large, receaanlee inal aah hidden by coxa 2, panes if Berea occurring only in distal third of coxa... . NSRORILGE., Na mim nada cman ASD Inner plate of maxilla 1 with three or more Botae UNS ee SB RU SeR en OO
Inner plate of maxilla 1 with two or fewer setae (or inner plate absent). 35
2 Some latitude is expressed in this division as certain genera are found in both pathways.
285-135 O - 69 - 21
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48.
U.S. NATIONAL MUSEUM BULLETIN 271
Maxillipedal palp with three articles (fig. 1287) .... . . . Perrierella Maxillipedal palp with four articles . . . ; URE: Plates of maxilla 2 subequally broad and nae capstan, mtiiclle 1 of antenna 2
very enlarged and subspherical (adults), telson elongate . . Eurythenes
Inner plate of maxilla 2 much broader than outer, inner plate often strongly gaping mediad (fig. 1277), article 1 of antenna 2 normally small, telson
not elongate... . WR bo) a APIStaSs Palp article 4 of amped ood rests Ge 128k) . . . . . Centromedon 2 Palp article 4 of maxilliped well developed and claw-like. ..... . 36
Inner plate of maxilla 1 with at least one large falcate seta (fig. 127d). Hirondellea 4
Inner plate of maxilla 1 with normal setae ... . MeEESe DERE ay Cai aI Setae of maxilla 2 reduced to one-two and four ee inner ianid outer lobes
CED eyes seis ceils eae eo deliell a! Setae of maxilla 2 numerous 4 per tobe): BU ae is PVRS LER NEC tee ool Pea teS Outer plate of maxilla 1 with spines reduced to five or fomee . Neoambasia Outer plate of maxilla 1 with seven or more spines... . Sots Oe esa)
Gnathopod 1 enlarged, article 5 very short and with small, narrow posterior lobe, article 6 expanding distally, very broad, palm transverse or nearly
chelate (fig. 1291) . . . .. . . . . Cheirimedon latimanus Sars, 1895 Gnathopod 1 of normal size, article 5 rarely lobate, article 6 not expanding distally even though palm rarely transverse. . . . ic iente 40 Labrum with distinct lobe projecting in front of epierome (irom lateral view). Ambasiopsis Labrum not dominating epistome . . . Be a ole vege tice a “Head small’ (fig. 1236), epistome ond mpsa el gine, inconspicuous, neither dominant . . . “ana e) SanUEIStes “Head large’’ (fig. 122d), Prebacenl complex pone pienGne: epistome usually dominating upper lip ..... .. . . . Tryphosella Inner plate of maxilla 1 with eight or more aeuae le medial edge. Paralicella Inner plate of maxilla 1 with three or fewer setae... . foe: Auentee tee Coxa dS as\longias4 (fe 21D) ) >. = eee Tepidepeceenidas Coxa 5 much shorter than4 .... . BBY es SOG LAM RS ek Telson entire or cleft less than one sein its lenenh PR MMR i ekey dk oo) Telson cleft one eighth or moreitslength. ....... Seer Senet aC Outer ramus of uropod 3 uniarticulate ......... Ubeeudarabacis Outer ramus of uropod 3 biarticulate . . . . Sei eG
Labrum rounded anteriorly and scarcely produced nerond iepietoue’ inner ramus of uropod 2 simple, article 6 of gnathopod 1 coe and stout, palm transverse (fig. 129a) . . . . ... . . Koroga
Labral area of prebuccal complex nanialy ‘prodiiead| tt in front of epistomal area, inner ramus of uropod 2 constricted, article 6 of gnathopod 1 slender,
palm obsolescent, oblique. . . . . . . Paralysianopsis Palp article 4 of maxilliped very stout ind supeluate. dactyl of gnathopod 1 as short and stout as dactyl of gnathopod 2. . . . . . Pseudoanonyx Maxillipedal palp article 4 typically unguiform (fig. 128c). .... . 48 Epistome with acute anterior process dominating upper lip. . Tryphosites Hpistomedlationroundedvanteriorly 6 45 ane ee
3 Compare Adeliella. 4 See note, Key N, couplet 23.
MARINE) GAMMARIDEAN AMPHIPODA 317
49. Gills plaited (deeply folded), labral lobe slightly dominating epistome. Anonyx Gills simple (at most striate), epistome equivalent to or dominating labral IWoY YS): sisi ced aa te anes ISIC eam MR ANS Sous sg ated Aer Tmetonyx (type only) Gills simple (at most striate), labral lobe slightly dominating epistome. Paronesimus
Genera of Lysianassidae
The following characters are not mentioned in the diagnoses unless they have classificatory value; if not mentioned they conform to the typical condition as stated in sequel: gnathopod 2 either minutely chelate or subchelate; mazilla 1 with 2-articulate palp, inner lobe poorly setose; maailliped with well-developed lobes and 4-articulate palp, the fourth article large and claw-like; coxae 1 or 2 large, visible; uropod 8 of medium length, outer ramus 2-articulate, inner slightly shorter than outer, rami lanceolate; accessory flagellum multiarticulate; inner ramus of uropod 2 unconstricted or unnotched; mouthparts not formed into a cone-shaped bundle; pereopod 3, article 2 not deeply indentured.
Acidostoma Liljeborg
Acidostoma Liljeborg, 1865.—Stebbing, 1906.
Type-species: Anonyx obesus Bate, 1862 (monotypy). See Sars, 1895.
Mouthparts forming a ventral conical bundle, styliform; antenna 1 very stout in male; upper lip and epistome continuous; mandible usually lacking molar, small when present, palp attached quite proximally; palp of maxilla 1 vestigial, 1-articulate, outer plate with hook-like spines; palp of maxilliped scarcely exceeding outer plate, article 4 vestigial; gnathopod 1 simple, article 6 slightly longer than 5; gnathopod 2 lacking article 7; uropod 3 short, outer ramus 1-articulate; telson short, cleft or emarginate. Species: 7, boreal, littoral to bathyal (1,200 m).
Acontiostoma Stebbing
Acontiostoma Stebbing, 1888; 1906.
Type-species: A. marionis Stebbing, 1888 (original designation).
Mouthparts forming a ventral conical bundle, styliform; mandible lacking molar, palp attached quite proximally; palp of maxilla 1 uniarticulate; maxillipedal palp scarcely exceeding outer plate, article 4 very small; gnathopod 1 simple, article 6 slightly longer than 5; coxa 1 projecting over side of head; uropod 3 lacking rami; telson short, entire or emarginate. See Stomacontion for remarks. Species: 1, subantarctic, littoral.
318 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 117.—Lysianassidae: a, Onisimus (=Pseudalibrotus) litoralis (Kréyer) (Sars, 1895, pl. 35); b, Stomacontion pepinii (Stebbing, 1888), note coxa 1 overlapping head, and eye dimly showing through.
MARINE GAMMARIDEAN AMPHIPODA 319
all Zipp [fh yf
NN
Figure 118.—Lysianassidae: a, Lepidepecreum umbo (Goés) (Sars, 1895, pl. 39); b, Kergue- lenia borealis Sars (1895, pl. 40).
320 U.S. NATIONAL MUSEUM BULLETIN 271
Adeliella Nicholls
Adeliella Nicholls, 1938.
Type-species: A. laticornis Nicholls, 1938 (original designation).
Upper lip and epistome not differentially produced; mandible with “distinct molar”, palp apparently attached level with molar; maxilli- pedal palp article 4 rather small; maxillae poorly spinose or setose; gnathopod 1 subchelate, short, articles 5 and 6 equal in length; telson short, emarginate or notched apically. Species: 1, antarctic, bathyal (420 m).
Alicella Chevreux
Alicella Chevreux, 1899.—Stebbing, 1906.
Type-species: A. gigantea Chevreux, 1899 (original designation). See Chevreux, 1935.
Upper lip and epistome not differentially produced; mandibular cutting edge with small teeth, molar conicolaminate, setulose, palp attached level with molar; inner plate of maxilla 1 densely setose on medial edge; maxillipedal palp article 4 very small; gnathopod 1 feeble, simple, articles 5 and 6 equal in length; article 6 of gnathopod 2 as long as article 5, both very elongate and linear; rami of uropod 3 long, equal, lanceolate, apparently outer ramus 1l-articulate; telson of medium length, deeply cleft. Species: 1, N. Atlantic, abyssal.
[Allogaussia Schellenberg] Allogaussia Schellenberg, 1926a.—K. H. Barnard, 1932.
Type-species: A. paradoxa Schellenberg, 1926a (selected by Stasek, 1958).
Type-species distinct from type-species of Orchomene by entire telson, produced posterior lobe of coxa 5, asymmetrical posterior lobe of pereopod 3. Species that have been assigned on basis of telson alone: 6 (also included in Orchomene).
Amaryllis Haswell Amaryllis Haswell, 1880b.—Stebbing, 1906.
Type-species: A. macrophthalmus Haswell, 1880b (selected by Pirlot, 1933b; indicated by Stebbing, 1888, p. 702!, through monotypy by synonymy).
Epistome apparently projecting in front of upper lip; mandibular molar feeble, setulose, palp attached near proximal end of molar; maxilla 1 lacking palp; gnathopod 1 simple, article 6 longer than 5; coxa 1 partially hidden, following coxae increasingly larger; inner ramus of uropod 2 constricted; telson of medium length, half or
MARINE GAMMARIDEAN AMPHIPODA 321
deeply cleft; article 2 of antenna 1 half as long as article 1 (Bathya- maryllis) ; anteroventral corner of coxa 4 rounded (Vyaya). Species: 2, antiboreal, littoral.
Ambasia Boeck
Ambasia Boeck, 1871.—Stebbing, 1906.
Type-species: Gammarus atlanticus Milne Edwards, 1830 (mono- typy and subsequent synonymy). See Sars, 1895.
Epistome and upper lip coalesced, but epistomal part greatly and sharply produced in front; mandibular molar absent, palp attached to middle of mandible; marillipedal palp article 2 half or less as broad as outer plate (Ambasiella), article 4 very small; gnathopod 1 simple, article 5 slightly longer than 6; coxa 1 slightly shortened and partially hidden by coxa 2; inner ramus of uropod 3 half as long as outer; telson of medium length, deeply cleft. Species: 1, subarctic N. Atlantic, littoral and bathyal (to 1,400 m).
Ambasiella Schellenberg
Ambasiella Schellenberg, 1935.
Type-species: Ambasia murmanica Briiggen, 1905 (monotypy). See Gurjanova, 1951.
Epistome and upper lip not differentially produced; mandibular molar absent, palp attached proximally; mazillipedal palp article 2 much broader than outer plate (Ambasia, Hirondellea), article 4 vestig- ial; gnathopod 1 simple, article 5 longer than 6; coxa 1 slightly shortened, coxa 2 concealing most of it; telson of medium length, deeply cleft. Species: 1, arctic, littoral.
Ambasiopsis K. H. Barnard
Ambasiopsis K. H. Barnard, 1931; 1932.
Type-species: A. georgiensis K. H. Barnard, 1931 (original designation) ; 1932.
Upper lip lobately produced in front of epistome; mandibular molar large, truncate, not ridged, setulose, palp attached level with molar; outer plate of maxilliped lacking spines (Neoambasia) ; gnathopod 1 poorly subchelate; coxa 1 two thirds as long as coxa 2 and partially hidden by it, but as long as article 2 of gnathopod 1; telson deeply cleft. Species: 2, subantarctic, littoral (to 342 m).
U.S. NATIONAL MUSEUM BULLETIN 271
vik ; A/ i ‘
a
SEF V7 RY) ae ti
i Ce
fa” hye Mee VS
=
Ficure 119.—Lysianassidae: a, Eucallisoma glandulosa J. L. Barnard (1961); b, Ambasia
atlantica (Milne Edwards) (Sars, 1895, pl. 17, as 4. danielssent); c, Pachychelium davidis Stephensen (1925a).
MARINE GAMMARIDEAN AMPHIPODA 323
VAN Uh Tp x , YY) Ns) 5} f a
4 [ ) [> Soe lim (| cl WS | Ol Lyf ft ff y, Seated «| al Figure 120.—Lysianassidae: a, Acid b (Bate) (Sars, 1895, pl. 114); b, Tryphos-
esum Barnard, 1962d).
324 U.S. NATIONAL MUSEUM BULLETIN 271
Anonyx Kr¢yer
Anonyz Kr¢gyer, 1838.—Stebbing, 1906.—Gurjanova, 1962.—Hurley, 1963. Chironesimus Sars, 1895. Lakota Holmes, 1908.
Type-species: Cancer nugax Phipps, 1774 (elimination and syn- onymy to result in monotypy by Boeck, 1871, but firmly selected by Boeck, 1876, with subsequent synonymy). See Sars, 1895.
Upper lip projecting slightly or strongly in front of epistome; mandibular molar large, or of medium size, subconical, unridged (Boeckosimus), setulose, palp attached level with molar; gnathopod 1 subchelate, articles 5 and 6 variable in length; inner ramus of uropod 2 constricted or not; telson of medium length, deeply cleft; gills plaited except in Lakota; eyes present; pereopods 1-2 unrth distal locking spines on article 6 at base of dactyls; article 4 of mazillipedal palp slender and claw-like (Pseudoanonyz). Species: 30+, arctic- boreal, littoral to abyssal.
Aristias Boeck Aristias Boeck, 1871.—Stebbing, 1906.
Type-species: Anonyx tumidus Krgyer 1846 (monotypy). See Sars, 1895.
Upper lip either projecting or not projecting beyond epistome, occasionally lobate, angular or poorly produced; mandibular molar a long unridged conical lamina, palp attached level with molar; inner lobe of maxilla 1 strongly setose (5+ setae) and spinose; lobes of maxilla 2 often gaping, inner much broader than outer; gnathopod 1 nearly simple or with minute palm, article 7 overlapping palm, articles 5 and 6 equal, article 3 not elongate (Paralicella); coxa 1 short, half as long as its article 2, partially hidden by coxa 2; telson of medium length or short, deeply cleft. Differs from Eurythenes by: shorter telson, broader inner plate of maxilla 2, narrow mandibular molar, longer article 2 of pereopod 8. Species: 16, primarily bipolar, amphiboreal (one tropical), littoral to bathyal.
Aristiopsis J. L. Barnard Aristiopsis J. L. Barnard, 1961.
Type-species: A. tumidus J. L. Barnard, 1961 (original desig- nation).
Both epistome and upper lip with short anterior humps separated by a long, flat, unnotched margin; mandible with well-developed, ridged molar, palp attached over posteroproximal corner of molar;
MARINE GAMMARIDEAN AMPHIPODA 325
gnathopod 1 stout, slightly chelate; coxa 1 short, half as long as its article 2, partially concealed by coxa 2; uropod 2 with inner (not outer = error of J. L. Barnard, 1961) ramus slightly constricted; telson short, cleft one third its length. Species: 1, Pacific, bathyal- abyssal.
Ficure 121.—Lysianassidae: a, Danaella mimonectes Stephensen (1925b); b, Lepidepecreoides xenopus K. H. Barnard (1932).
326 U.S. NATIONAL MUSEUM BULLETIN 271
VES POOR i Ȥ Cee oe
Ficure 122.—Lysianassidae: a, Thoriella islandica Stephensen (1915), offset showing dorsal urosome; b, Onesimoides chelatus Pirlot (J. L. Barnard, 1961); c, Crybelocyphocarts tattersalli Shoemaker (1945a); d, Tryphosella sarsi Bonnier (Sars, 1895, pl. 27, as Tryphosa nana).
MARINE GAMMARIDEAN AMPHIPODA 327
Ficure 123.—Lysianassidae: a, Trischizostoma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi); b, Uristes umbonatus (Sars, 1895, pl. 29); c, Hippomedon denticulatus (Bate) (Sars, 1895, pl. 20); d, Centromedon pumilus (Liljeborg) (Sars, 1895, pl. 34); e, Acidostoma obesum (Bate) (Sars, 1895, pl. 14).
328 U.S. NATIONAL MUSHUM BULLETIN 271
Aroui Chevreux Arouti Chevreux, 191 1a.
Type-species: A. setosus Chevreux, 191la (monotypy).
Upper lip and epistome not differentially produced, epistome flat in front (Scopelocheirus); mandibular molar small, cylindrical, ridged, palp attached slightly proximal to molar; inner plate of maxilla 1 extremely setose; lobes of maxilla 2 subcircular, outer shorter than inner; article 6 of gnathopod 1 terminated by short shroud of setae; article 7 vestigial; gnathopod 2 minutely chelate (Paracallisoma); telson of medium length, deeply cleft. Species: 1, Algeria, littoral.
Azotostoma J. L. Barnard Azotostoma J. L. Barnard, 1965.
Type-species: <A. fusta J. L. Barnard, 1965 (original designation).
Mouthparts forming a ventral conical bundle, styliform; mandible lacking molar, palp attached very proximally; palp of maxilla 1 large, exceeding outer plate, bent, 2-articulate; inner plate of maxilla 2 expanded proximally, distal tapered portion with three stout spines; inner plates of maxilliped styliform, outer very large and flagellate, palp 4-articulate, scarcely exceeding outer plate, article 2 produced distally; gnathopod 1 simple, flagelliform, article 6 much longer than 5, article 7 elongate but not as long as article 6; uropod 3 biramous, rami equal, 1-articulate; telson short, entire. Species: 1, Micronesia, littoral.
Bathyamaryllis Pirlot
Bathyamaryllis Pirlot, 1933a.
Type-species: JB. perezii Pirlot, 1933a (original designation).
Epistome projecting slightly in front of upper lip; mandible with long, subconical, setulose molar, palp attached level with molar; palp of maxilla 1 absent; gnathopod 1 simple, article 6 longer than 5; coxa 1 small, partially hidden, subsequent coxae increasing in size; uropod 3 with l-articulate outer ramus; telson of medium length, deeply cleft; article 2 of antenna 1 equal to article 1 (Amaryllis). Species: 4, probably cosmopolitan, bathyal to abyssal.
Bathycallisoma Dahl
Bathycallisoma Dahl, 1959. Synonym of Scopelocheirus fide Birstein and Vinogradov, 1964. See also Paracallisoma.
Type-species: B. pacifica Dahl, 1959 (monotypy). Upper lip and epistome not differentially produced; mandibular
MARINE) GAMMARIDEAN AMPHIPODA 329
molar small, conical, unridged, palp attached level with molar but both rather proximal; lower lip with major lobes excavate medially, forming medial gap (combining character); inner plate of maxilla 1 densely setose; gnathopod 1 simple, article 7 small, article 6 tipped with short setal shroud; gnathopod 2 subchelate but not distinctly chelate (Scopelocheirus); telson of medium length, deeply cleft. Species: 1, central Pacific, hadal.
Ficure 124.—Lysianassidae: a, Chevreuxiella metopoides Stephensen (1915), offset showing dorsal urosome (3, 4, 5-6; U=uropod). Antenna 1: b, Bathyamaryllis perezit Pirlot (1933a); c, Lysianassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); d, Lepidepecreum longicorne (Bate and Westwood) (Sars, 1895, pl. 38, as L. carinatum); e, Ambasia atlantica (Milne Edwards) (Sars, 1895, pl. 17, as 4. danielsseni); f, Socarnotdes kergueleni Stebbing (1888); g, Menigrates obtusifrons (Boeck) (Sars, 1895, pl. 38); h, Normanion sarsi Stebbing (Sars, 1895, pl. 13, as N. quadrimana). Antenna 2: 1, Eurythenes gryllus (Lichtenstein) (Sars, 1895, pl. 30); 7, Parawaldeckia kidderi (Smith) (Tattersall, 1922); k, Lystanella petalocera Sars (1895, pl. 18).
330 U.S. NATIONAL MUSEUM BULLETIN 271
Boeckosimus, new genus
Type-species: Anonyx edwardsi Krgyer, 1846 (present selection). This genus embraces those species formerly included in the genus Onisimus Boeck, 1871, since the time of Sars (1895) and Stebbing (1906). The genus Onisimus must include those species having gone in the genus Pseudalibrotus Della Valle, 1893, since the time of Steb- bing (1906). Boeck (1876) designated Anonyzx litoralis Krgyer, 1845, as type of Onisimus Boeck, 1871, and Pseudalibrotus Della Valle is a junior synonym of Onisimus because of monotypy, A. litorahs also being the type-species of Pseudalibrotus. Sars (1895) designated
Uy
Ficure 125.—Lysianassidae: Upper lip-epistome complex, lateral view, left sides: a, Socarnoides kerguelent Stebbing (1888); b, Tryphosites longipes (Bate and Westwood) (Sars, 1895, pl. 28); c, Lystanassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); d, Menigrates obtusifrons (Boeck) (Sars, 1895, pl. 38); e, Eurythenes gryllus (Lichtenstein) (Sars, 1895, pl. 30); f, Orchomene batei Sars (1895, pl. 22); g, Ichnopus spinicornis Boeck (Sars, 1895, pl. 15); 4, Ambasia atlantica (Milne Edwards) (Sars, 1895, pl. 17, as 4. dani- elsseni); 1, Lepidepecreum longicorne (Bate and Westwood) (Sars, 1895, pl. 38, as L. carinatum); 7, Orchomene amblyops Sars (1895, pl. 25); k, Nannonyx goest (Boeck) (Sars, 1895, pl. 24); 1, Orchomene minuta (Kr¢yer) (Sars, 1895, pl. 24); m, Scopelochetrus crenatus Bate (Sars, 1895, pl. 19); », Hippomedon denticulatus (Bate) (Sars, 1895, pl. 20); 0, Orcho- mene obtusa (Sars, 1895, pl. 26). Lower lips: p, Hippomedon; q, Nannonyx; r, Normanton; 5, Eurythenes; t, Ichnopus; u, Opisa eschrichti (Krgyer) (Sars, 1895, pl. 14); v, Trischizo- stoma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi). Upper lip, anterior view: w, Acidostoma obesum (Bate) (Sars, 1895, pl. 14); x, Trischizostoma.
v
MARINE GAMMARIDEAN AMPHIPODA 331
A. edwardsti as type of Onisimus, erroneously, and I follow his prece- dent in using A. edwardsii as type of Boeckosimus. Gurjanova (1951) listed Anonyx plautus Kroyer, 1845, as type of Onisemus but this is also erroneous. The genus Alibrotus Milne Edwards, 1840, may be a senior synonym either of Boeckosimus or Onisimus but its status has remained unclear. Stebbing (1906) lists it as a dubious genus. Its type-species, Lysianassa chauseica Milne Edwards, 1830 (mono- typy), probably cannot be a member of either Onisimus (= Pseudali- brotus) or Boeckosimus (=‘‘Onisimus”’ auct.) because no species of either genus has been recorded as far south as Isles Chauseay in the English Channel.
Upper lip and epistome not prominent; mandibular molar ridged (Anonyzx), palp attached level with molar; gnathopod 1 subchelate or poorly subchelate, articles 5 and 6 subequal or article 6 longer than 5; inner ramus of uropod 2 constricted or not; telson short (Hippome- don), variable, entire or cleft one third; outer plate of maxilla 1 with nine or more spines; palp of mazilliped elongate and slender, outer plate of maxilliped reaching to end of palp article 2 (Rifcus). Species: 16, arctic-boreal, mostly N. Atlantic, littoral to abyssal.
Note: This genus (‘Onisimus” auct.) and Onisimus (= Pseudali- brotus auct.) are very close. Sars (1895) distinguished them on the more powerful gnathopod 1 and uropod 3 and longer antennae of “Pseudalibrotus”’ and Gurjanova (1962) distinguished them on the absence of bent spines on the apices of the maxillipedal outer plates in ‘“Pseudalibrotus.’’ Most species of ‘“Onisimus’’ have slightly cleft telsons and various species of ‘‘Pseudalibrotus’”’ have the epistomal portion of the prebuccal complex dominating the labral portion. All of these seem to be very weak generic characters. If the genera were synonymized the unfortunate nomenclatural problems would be solved.
Cebocaris J. L. Barnard Cebocaris J. LL. Barnard, 1964a.
Type-species: OC. grutesca J. L. Barnard, 1964a (original designa- tion).
Epistome large, broad, convex, but not projecting beyond upper lip from lateral view; head deformed; mandibular molar absent; gnathopod 1 simple, articles 5 and 6 subequal; article 5 of gnathopod 2 only slightly longer than article 6, appendage simple; coxae 1 and 2 small, but coxa 3 only slightly enlarged and covering coxa 2 only partially; article 2 of pereopod 3 not indentured; pereopods 1-5 strongly prehensile; telson short, entire. Species: 1, Caribbean, abyssopelagic.
285-135 O - 69 - 22
Ficure 126.—Lysianassidae: Mandible: a, Centromedon pumilus (Liljeborg) (Sars, 1895, pl. 34); b, Trischizostoma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi); c, Kergu- elenia borealis Sars (1895, pl. 40); d, Eurythenes gryllus (Lichtenstein) (Sars, 1895, pl. 30); e, Tryphosella sarst Bonnier (Sars, 1895, pl. 27, as Tryphosa nana); f, Aristias neglectus Hansen (Sars, 1895, pl. 17, as A. audouinianus); g, Hippomedon denticulatus (Bate) (Sars, 1895, pl. 20); h, Lystanassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); 1, Orchomene batei Sars (1895, pl. 22); 7, Anonyx nugax (Phipps) (Sars, 1895, pl. 31); &, Acidostoma obesum (Bate) (Sars, 1895, pl. 14); 7, Menigrates obtusifrons (Boeck) (Sars, 1895, pl. 38); m, Boeckosimus edwardsi (Krgyer) (Sars, 1895, pl. 36); n, Normanion sarst Stebbing (Sars, 1895, pl. 13, as MN. quadrimana); 0, Alicella gigantea Chevreux (1899); p, Ichnopus spinicornis Boeck (Sars, 1895, pl. 15); g, Valettiopsis multidentata J.L. Barnard (1961), distal portion to show teeth of incisor.
MARINE GAMMARIDEAN AMPHIPODA 333
Centromedon Sars
Centromedon Sars, 1895.—Stebbing, 1906.
Type-species: <Anonyx pumilus Liljeborg, 1865 (monotypy).
Upper lip and epistome small, but separated by a notch, upper lip slightly projecting; mandibular molar conicolaminate, unridged, palp attached level with molar; maxillipedal palp article 4 very small; enathopod 1 poorly subchelate, articles 5 and 6 equal; coxa 1 slightly tapering distally; telson of medium length, deeply cleft; head small, lateral lobes acute (Hirondellea), mouthparts hidden by coxae. Species: 1, subarctic and boreal N. Atlantic, littoral.
Cheirimedon Stebbing Cheirimedon Stebbing, 1888; 1906.
Type-species: C. crenatipalmatus Stebbing, 1888 (monotypy).
Upper lip and epistome not differentially produced; mandibular molar very small, sharp or blunt, ridged (type) or poorly ridged and sublaminate, palp attached level with molar; gnathopod 1 subchelate, article 6 broadly expanded (type) or poorly expanded, occasionally slightly chelate in appearance; coxae normal (type) or coxa 2 con- cealing part of slightly shortened (and occasionally strongly narrowed) coxa 1; telson of medium length, deeply cleft (type) or cleft as little as one third. Some species resembling Anonyz, Boeckosimus (=“Onisimus” auct.), Schisturella (= Pseudonesimus), and Tryphosella (inel. “‘7ryphosa”’ auct.). Species: 5, amphiboreal, littoral to bathyal.
The type-species, Cheirimedon crenatipalmatus Stebbing (1888) is characterized by: lateral outline of mouthpart group quadrate; epistome and upper lip not differentially produced; head normal; mandibular palp attached level with molar, molar process columnar, triturative; other mouthparts normal; coxa 1 quadrate, not narrowed distally or covered by coxa 2; gnathopod 1 subchelate, with article 6 longer than 5 and slightly expanded distally; telson cleft more than three fourths of its length; uropod 3 normal, outer ramus 2-articulate.
Cheirimedon crenatipalmatus thus differs from Anonyz in (1) the nonsetulose mandibular molar which in Anonyx is bulging or sub- laminate but lacking ridges and teeth and (2) in the expanded article 6 of gnathopod 1. It differs from Tryphosella (=“Tryphosa’”’ auct.) in the nontapering coxa 1, and the distally expanding article 6 of enathopod 1. The type-species is joined by C. femoratus (Pfeffer) (=C. dentumanus Chevreux, 1906b) which is distinguished generically only by the telson being cleft halfway and by the poor expansion of gnath- opod 1.
The other species that have been assigned to this genus depart radically from the above brief diagnosis. Cheirimedon latimanus
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Sars, 1895, has the expanded article 6 of gnathopod 1 but coxa 1 tapers strongly to a subacute point and is partially covered by coxa 2; the mandibular molar is like that of some species of Anonyzx in its sublaminar condition lacking teeth and ridges. The telson is deeply cleft. Except for gnathopod 1 this species could be assigned to Uristes, although Uristes has a mandibular molar more like that of Cheirimedon crenatipalmatus. Cheirimedon latimanus also bears comparison to Centromedon pumilus on technical grounds, and is very close to “Tryphosa” auct. except for the first gnathopodal expansion.
Figure 127.—Lysianassidae: Maxilla 1: a, Lysianassa heterodonta (Pirlot, 1936); », Bathyamaryllis perezii Pirlot (1933a); c, Trischizostoma nicaeense (Costa) (Sars, 1895, pl. 12, as T. rascht) d, Hirondellea trioculata Chevreux (1900); e, Scopelochetrus crenatus Bate (Sars, 1895, pl. 19); f, Lystanassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); g, Acidostoma obesum (Bate) (Sars, 1895, pl. 14); h, Aristias neglectus (Bate) (Sars, 1895, pl. 17); 1, Orchomene batei Sars (1895, pl. 22). Maxilla 2: 7, Perrierella; k, Adeliella laticornis Nicholls (1938); 1, Orchomene; m, Trischizostoma; n, Acidostoma; 0, Scopelocheirus; p, Lysianassa plumosa. Coxa1: q, Anonyx nugax (Phipps) (Sars, 1895, pl. 31). Coxa 4: r, Paralibrotus setosus Stephensen (1923). Gills: 5, Ichnopus spinicornis Boeck (Sars, 1895, pl. 15). Pleonites 3-6 (5-6 fused), lateral: ¢, Crybelocyphocaris tattersalli Shoemaker (1945a).
MARINE GAMMARIDEAN AMPHIPODA 335
Cheirimedon fougnerr Walker (1903) differs from the type-species of Cheirimedon by the short cleft of the telson and thus resembles some species of “‘Onisimus’’ auct. and Paronesimus. Cheirimedon pectini- palma K. H. Barnard (1925) resembles C. latimanus in coxa 1 and mandible but has a poorly cleft telson, and is distinguished from Uristes primarily by the telson. Because these species assigned to Cheirimedon confound its crisp definition and bridge so many other genera it would seem prudent to isolate at least C. latimanus in a genus of its own and assign C. pectinipalma to it provisionally. The latter also has affinities with Hirondellea except in its head. The three species then remaining in Cheirimedon, C. crenatipalmatus, C. fougneri, and C. femoratus should be examined from time to time in light of a viewpoint that they simply represent minor departures in the evolutionary scheme of certain genera such as Anonyz (C. crenatipalmatus and C. femoratus) and ‘‘Onisimus’”’ (C. fougnerr) while those species of the new genus, with C. latimanus as type, should be considered as possible subgeneric members of Uristes and ‘‘Tryphosa.’’ A parallel may be seen in Pseudokoroga rima J. L. Barnard which is simply an Orchomene with enlarged first gnathopod. Perhaps many of the lysianassids with enlarged first gnathopods are the rubble left behind in the differentia- tion of genera, now so remote from their ancestors, that their origins are unrecognizable.
Cheirimedon is mentioned often in the foregoing keys to Lysianas- sidae because the several species confound the couplets at vital points.
Chevreuxiella Stephensen
Chevreuxiella Stephensen, 1915.
Type-species: C. metopoides Stephensen, 1915 (monotypy).
Accessory flagellum absent; upper lip and epistome not differen- tially produced but both large and helmet-shaped; mandible lacking molar and palp; maxilliped of diverging interpretation: possibly composed of small inner and outer plates and an immense opercular palp composed of one article supporting a second minute proximal article; possibly the opercular palp article is the outer plate and the second article is the palp; gnathopod 1 simple, short, article 5 longer than 6, article 7 with apical spine; gnathopod 2 simple; coxae 1 and 2 much smaller than coxae 3 and 4, coxa 2 partially hidden by 3 and coxa 1 partially hidden by 2; coxa 4 large, not posteriorly ex- cavate; uropods 1 and 2 flattened, inner ramus less than half of outer; telson and uropod 3 absent; urosome composed of two segments. Species: 2, Atlantic, antarctic, bathy-abyssopelagic.
Frcure 128.—Lysianassidae: Maxilliped: a, Centromedon pumilus (Liljeborg) (Sars, 1895, pl. 34); b, Kerguelenia borealis Sars (1895, pl. 40); c, Lystanassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); d, Acidostoma obesum (Bate) (Sars, 1895, pl. 14); e, Pachynus chelatum Bulycheva (Gurjanova, 1962); f, Lystanassa heterodonta (Pirlot, 1936); g, Normanion sarsi Stebbing (Sars, 1895, pl. 13, as N. quadrimana); h, Trischizos- toma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi); 1, Ambasia atlantica (Milne Ed- wards) (Sars, 1895, pl. 17, as 4. danielsseni); 7, Perrierella audouiniana (Bate) (Chevreux and Fage, 1925); k, Nannonyx goest (Boeck) (Sars, 1895, pl. 24), /, Danaella mimonectes Stephensen (1925b), palp absent. Telson: m, Acidostoma; n, Onisimus edwardsi (Kr¢yer) (Sars, 1895, pl. 36); 0, Orchomene batei Sars (1895, pl. 22); p, Lysianassa; q, Ichnopus spinicornis Boeck (Sars, 1895, pl. 15); 7, Opisa eschrichti (Krdyer) (Sars, 1895, pl. 14); 5, Hippomedon abysst (Goés) (Gurjanova, 1962); t, Eurythenes gryllus (Lichtenstein) (Sars, 1895, pl. 30); u, Hippomedon denticulatus (Bate) (Sars, 1895, pl. 20); 2, Boeckosimus normani (Sars, 1895, pl. 36); w, Trischizostoma.
Ficure 129.—Lysianassidae: Gnathopod 1: a, Koroga megalops Holmes (Stephensen, 1923); b, Paracentromedon crenulatus (Chevreux) (Chevreux and Fage, 1925); c, Schisturella (=Pseudonesimus) abyssi (Chevreux, 1926); d, Orchomene bate: Sars (1895, pl. 22); e, Opisa eschrichti (Krdyer) (Sars, 1895, pl. 14); f, Nannonyx goesi (Boeck) (Sars, 1895, pl. 24); g, Eucallisoma glandulosa J. L. Barnard (1961); h, Gainella chelata Chevreux (1912b); 7, Pseudorchomene coatst (Chilton, 1912); 7, Tryphosella sarsi Bonnier (Sars, 1895, pl. 27, as Tryphosa nana); k, Uristes umbonatus (Sars, 1895, pl. 29); 7, Chetrimedon latimanus Sars (1895, pl. 13); m, Menigrates obtusifrons (Boeck) (Sars, 1895, pl. 38); », Paracal- lisomopsis beljaevi Gurjanova (1962); 0, Euonyx chelatus Norman (Sars, 1895, pl. 40).
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Clepidecrella J. L. Barnard Clepidecrella J. L. Barnard, 1962d.
Type-species: C. cabinda J. L. Barnard, 1962d (original designa- tion).
Head small, partially hidden by coxa 1; article 1 of antenna 1 carinate; upper lip and epistome not differentially produced; mandib- ular molar vestigial, palp very long, attached level with molar; outer plates of maxilliped rather small, palp very long; gnathopod 1 simple, articles 5 and 6 equal; uropod 3 very short, inner ramus short, scale-like; telson short, apically emarginate. Species: 1, S. Atlantic, abyssal.
Crybelocephalus Tattersall Crybelocephalus Tattersall, 1906.
Type-species: C. megalurus Tattersall, 1906 (original designa- tion). See Shoemaker, 1945a.
Accessory flagellum absent; head somewhat deformed; mandible lacking both molar and palp; gnathopod 1 simple, articles 5 and 6 subequal; gnathopod 2 simple; coxae 1 and 2 small and fully hidden by coxa 3; pereopods 1 and 2 subprehensile; telson large, entire. Spe- cies: 3, cosmopolitan, bathy-abyssopelagic.
Crybelocyphocaris Shoemaker Crybelocyphocaris Shoemaker, 1945a.
Type-species: C. tattersalli Shoemaker, 1945a (original designa- tion).
Accessory flagellum absent; head deformed; mandible lacking molar, palp attached rather proximally; gnathopod 1 simple, articles 5 and 6 equal; gnathopod 2 simple; coxae 1 and 2 small and partially hidden by coxa 3; pereopods 1 and 2 prehensile, pereopod 3 subprehensile; uropod 3 small, outer ramus equal to peduncle, 2-articulate, inner ramus one third as long as outer; urosomites 2 and 3 coalesced. Spe- cies: 1, Atlantic, bathypelagic.
Cyclocaris Stebbing
Cyclocaris Stebbing, 1888; 1906.
Type-species: C. tahitensis Stebbing, 1888 (monotypy).
Apparently upper lip lobately produced in front of epistome; head somewhat deformed and globular; mandible flattened, possibly with elongate, lanceolate, setulose molar or none, palp attached medially
MARINE GAMMARIDEAN AMPHIPODA 339
or level with molar; gnathopod 1 slender, simple; coxae 1 and 2 small and partially hidden by coxa 3; telson long, deeply cleft. Species: 2, arctic to Pacific tropics, epi- to ?abyssopelagic.
Cyphocaris Boeck
Cyphocaris Boeck, 1871.—Stebbing, 1906.
Type-species: C. anonyx Boeck, 1871 (monotypy). See Stebbing, 1888 (as C. micronyz); Schellenberg, 1926b, 1927.
Epistome and upper lip not differentially produced; head deformed; mandibular molar of medium size, ridged, palp attached level with molar; gnathopod 1 simple, articles 5 and 6 equal; gnathopod 2 nearly simple, article 5 only slightly longer than article 6; coxae 1, 2, and 3 small, partially hidden by large coxa 4; article 2 of pereopod 3 deeply indentured or not, or with long posterior cusp; pereopods 1-4 decreasingly prehensile or not prehensile; telson long, deeply cleft. Species: 7, cosmopolitan, bathy-abyssopelagic.
Danaella Stephensen Danaella Stephensen, 1925b.
Type-species: D. mimonectes Stephensen, 1925b (monotypy, origi- nal designation in title).
Body globular, with pereonites 3-6 especially enlarged; accessory flagellum absent; epistome and upper lip not prominent; mandible simple, lacking molar and palp; maxilliped composed of large oper- cular outer plates covering smaller curved inner lobes; gnathopod 1 simple, dactyl very short; gnathopod 2 simple; coxae 1 and 2 small and hidden by following coxae; inner rami of uropods 1 and 2 short and spiniform; uropod 3 a vestigial microscopic bud; telson probably absent; urosome possibly consisting of two segments with a vestigial third. Species: 1, Atlantic, bathypelagic.
Derjugiana Gurjanova Derjugiana Gurjanova, 1962.
Type-species: JD. insolita Gurjanova, 1962 (monotypy).
Mouthpart group conically produced, mouthparts styliform; man- dible lacking molar, palp attached proximally; maxillipedal palp not exceeding outer plate, article 4 short and not claw-shaped; gnathopod 1 chelate, article 6 longer than 5; inner ramus of uropod 2 slightly constricted, uropod 3 lacking rami; telson short, entire. Species: 1, Okhotsk Sea, littoral.
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Ficure 130.—Lysianassidae: Gnathopod 1: a, Pachychelium davidis Stephensen (1925a); c, Valettiopsis dentatus Holmes (1908); ¢, Lysianassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); f, Valettia coheres Stebbing (1888); g, Neoambasia tumicornis (Nicholls, 1938) ; h, Trischizostoma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi); 1, Ichnopus spinicornts Boeck (Sars, 1895, pl. 15); 7, Kerguelenia borealis Sars (1895, pl. 40); k, Aristias neglectus Hansen (Sars, 1895, pl. 17, as 4. audouinianus); 1, Sophrosyne robertsoni Stebbing and Robertson (1891); m, Scopelocheirus crenatus Bate (Sars, 1895, pl. 19). Pereopod 1: b, Endeavoura mirabilis Chilton (1921); d, Paracyphocaris praedator Chevreux (Shoemaker, 1945a).
MARINE GAMMARIDEAN AMPHIPODA 341
Ficure 131.—Lysianassidae: Pereopod 3: a, Paracyphocaris praedator Chevreux (Shoe- maker, 1945a); b, Glycerina tenuicornis (Haswell) (Pirlot, 1936). Pereopod 2: c, Scopelo- chetropsis abyssalis Schellenberg (1926a). Uropod 2: d, Tryphosites longipes (Bate and Westwood) (Sars, 1895, pl. 29); e, Orchomene groenlandica (Hansen) (Sars, 1895, pl. 26). Gnathopod 2: f, Scopelocheirus crenatus Bate (Sars, 1895, pl. 19); g, Hirondellea brevicau- data Chevreux (1910); 4, Lysianassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); 1, Gainella chelata Chevreux (1912b); 7, Tryphosites; k, Trischizostoma nicaeense (Costa) (Sars, 1895, pl. 12, as T. raschi); 1, Lepidepecreum longicorne (Bate and Westwood) (Sars, 1895, pl. 38, as L. carinatum); m, Anonyx debruynii (Hoek) (Sars, 1895, pl. 37); n, Hippo- medon denticulatus (Bate) (Sars, 1895, pl. 20). Urosome: 0, Danaella mimonectes Stephen- sen (1925b), showing long uropod 1, shorter uropod 2 and arrow pointing to vestigial uropod 3 tubercles.
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Endevoura Chilton Endevoura Chilton, 1921.
Type-species: EH. mirabilis Chilton, 1921 (original designation).
Antennal flagella very short, 2- to 5-articulate; [upper lip and epistome not described]; mandibular molar well developed, ridged, palp attached proximal to molar; maxillipedal palp article 4 bulbous; enathopod 1 simple, dactyl very short; pereopod 1 prehensile, subche- late, large; telson entire, short. Species: 1, Bass Strait, littoral.
Ensayara J. L. Barnard Ensayara J. L. Barnard, 1964d.
Type-species: EE. ramonella J. L. Barnard, 1964d (original desig- nation).
Antennal flagella very short, about 5-articulate; upper lip and epistome inconspicuous, latter projecting shghtly; mandibular molar smooth, palp attached about level with molar; maxillipedal palp article 4 claviform; gnathopod 1 simple, dactyl short; pereopod 1 prehensile, subchelate, large; telson entire, short. Species: 1, Baja California, littoral.
Eucallisoma J. L. Barnard Eucallisoma J. L. Barnard, 1961.
Type-species: Ff. glandulosa J. L. Barnard, 1961 (original desig- nation).
Head very small but not deformed; antenna 1 immense, stout, conical, article 1 of flagellum elongate, flagellum bearing only three articles, accessory flagellum appressed to body of antenna (Paracal- lisomopsts) ; epistome and upper lip each convex in front; mandibular molar small, conical, unridged, palp attached level with molar; enathopod 1 simple, not elongate, article 7 vestigial, shrouded by short cirri, article 2 immensely swollen; telson of medium length, deeply cleft. Species: 1, tropical Atlantic, abyssal.
Euonyx Norman
Euonyx Norman, 1867b.—Stebbing, 1906. Leptochela Boeck, 1876 (homonym, Decapoda).
Type-species: EE. chelatus Norman, 1867b (monotypy). See Sars, 1895.
Epistome acutely produced in front of upper lip; mandibular molar obsolete or absent, palp attached distally; gnathopod 1 slender,
MARINE GAMMARIDEAN AMPHIPODA 343
chelate, thumb narrow; article 7 of gnathopod 2 rather large and curved; coxa 1 short, nearly hidden by coxa 2; telson of medium length, deeply cleft. Species: 7, cosmopolitan, epi- to bathypelagic.
Eurythenes Smith
Eurytenes Liljeborg, 1865 (homonym, Hymenoptera). Eurythenes Smith, 1884.—Stebbing, 1906.
Euryporeia Sars, 1895.
Katius Chevreux, 1905c.
Type-species: Gammarus gryllus Lichtenstein, 1822 (monotypy). See Sars, 1895.
Article 1 of antenna 2 tumid in adults; epistome broadly lobate in front of upper lip; mandibular molar immense, extremely broad, laminate, apically ridged, palp attached level with molar; gnathopod 1 subchelate, article 7 overlapping palm, article 6 longer than 5, article 3 not elongate (Paralicella); coxa 1 shortened, partially con- cealed by coxa 2; telson long, deeply cleft. See Arzstias. Species: 2, cosmopolitan, bathyal to abyssal.
Figorella J. L. Barnard Figorella J. L. Barnard, 1962d.
Type-species: F. tanidea J. L. Barnard, 1962d (original desig- nation).
Accessory flagellum 1-articulate; [upper lip and epistome not described]; mandibular molar absent, palp attached rather distally; palp of maxilla 1 uniarticulate; gnathopod 1 slightly chelate, article 6 somewhat enlarged, much longer than article 5; uropod 3 very short, inner ramus shortened; telson short, apparently entire. Species: 1, S. Atlantic, abyssal.
Gainella Chevreux Gainella Chevreux, 1911c; 1912a; 1912b.
Type-species: G. chelata Chevreux, 1912a, 1912b (designated by Chevreux, 1912a).
Epistome and upper lip not differentially produced; mandibular molar small but ridged, palp attached proximal to molar; inner plates of maxilliped styliform, palp very long, slender, article 4 densely setulose apically; gnathopod 1 chelate, thumb narrow and short, article 6 much longer than 5; gnathopod 2 styliform but mi- nutely chelate; uropod 3 short, inner ramus a short scale, one third as long as outer ramus; telson short, deeply cleft. Species: 1, antarc- tic, littoral (297 m).
344. U.S. NATIONAL MUSEUM BULLETIN 271
i ee A\N
Ficure 132.—Lysianassidae: a, Eurythenes gryllus (Lichtenstein) (Sars, 1895, pl. 30); b Metacyphocaris helgae Tattersall (1906). Uropod 3: c, Danaella mimonectes Stephensen (1925b); d, Stomacontion pepinii (Stebbing, 1888); ¢, Aristias tumidus (Kr@yer) (Sars, 1895, pl. 18); f, Hippomedon denticulatus (Bate) (Sars, 1895, pl. 20); g, Lysianassa plumosa Boeck (Sars, 1895, pl. 16, as L. costae); h, Lepidepecreella ctenophora Schellenberg (1926a); 1, Lysianassa alba (Holmes) (Shoemaker, 1933b); 7, Acontiostoma marionis Stebbing (1888); k, Nannonyx goesi (Boeck) (Sars, 1895, pl. 24).
MARINE GAMMARIDEAN AMPHIPODA 345
Glycerina Haswell
Glycera Haswell, 1880b (homonym, Polychaeta). Glycerina Haswell, 1882.—Stebbing, 1906.
Type-species: Glycera tenuicornis Haswell, 1880b (monotypy). See Pirlot, 1936.
Upper lip acutely produced in front of epistome; mandibular molar prominent, setulose, palp attached proximal to molar; gnathopod 1 simple, slender, article 5 slightly longer than 6; article 2 of pereopod 3 deeply indentured ; inner ramus of uropod 2 constricted; telson deeply cleft. Species: 1, Coral Sea to E. Australia, littoral.
Hippomedon Boeck
Hippomedon Boeck, 1871.—Stebbing, 1906.—Gurjanova, 1962. Platamon Stebbing, 1886.
Paratryphosites Stebbing, 1899d.
?Paracentromedon Chevreux and Fage, 1925.
?Hlimedon J. L. Barnard, 1962d.
Type-species: Anonyx holboll1 Krgyer, 1846 (selected by Boeck, 1876; note, the A. holbolli of Boeck, 1871 and 1876=Hippomedon denticulatus [Bate, 1857a] but the type-species should stand as A. holbolla Krgyer). See Sars, 1895.
Head small, largely hidden by cora 1 (‘‘Tryphosa’’ auct.), with distinct lateral lobes (Lepidepecreoides); upper lip either distinctly or not lobate in front of epistome but both projections small, incon- spicuous; mandibular molar large, ridged, palp attached level with molar; gnathopod 1 subchelate or nearly simple, article 5 longer than or equal to 6; telson short or medium, half or deeply cleft. See also Paronesimus. Species: 41, cosmopolitan, cold water, littoral to abyssal.
Hirondellea Chevreux
Mirondellea Chevreux, 1889.—Stebbing, 1906.—K. H. Barnard, 1930. Tetronychia Stephensen, 1923.
Type-species: Hirondellea trioculata Chevreux, 1889 (original designation). See Chevreux, 1900.
When visible, eyes triple; head globular or with large lateral lobes rounded, palp article 4 of mazilliped claw-shaped (Centromedon) ; mandibular molar conical, setulose, palp attached at proximal end of molar; inner plate of mazilla 1 with at least one enlarged sickle-seta or seta with strong attenuation or constriction, palp article 2 often notched medially; mazillipedal palp article 2 half or less as broad as outer plate (Ambasiella); gnathopod 1 subchelate or slightly chelate, articles 5 and 6 equal; coxa 1 rather small, often partially hidden by
346 U.S. NATIONAL MUSEUM BULLETIN 271
coxa 2; inner ramus of uropod 2 constricted or not; telson of medium length or short, deeply or shortly cleft. Species: 6, cosmopolitan, primarily bathyal to abyssal, pelagic.
Ichnopus Costa
Ichnopus Costa, 1853c; 1857.—Stebbing, 1906.
Type-species: J. taurus Costa, 1853c (monotypy). See Chevreux and Fage, 1925.
Upper lip slightly produced in front of epistome; mandibular molar laminate, scarcely ridged, palp attached level with molar; gnathopod 1 simple, article 7 with a posterior or distal bundle of setae; telson of medium length, cleft; gills plaited on both sides, pereopod 5 much longer than pereopod 4 (Socarnes). Species: 5, probably cosmopolitan, littoral to abyssal, pelagic.
Kerguelenia Stebbing Kerguelenia Stebbing, 1888; 1906.
Type-species: KK. compacta Stebbing, 1888 (monotypy).
Upper lip and epistome small, inconspicuous; mandible lacking distinct cutting edge and molar, palp attached distally; maxillae and maxilliped poorly developed, stunted; gnathopod 1 simple, styliform; uropod 3 tiny, rami usually shorter than peduncle, inner often vestigial or absent, outer 1- or 2-articulate; telson short, entire. Species: 6, bipolar, amphiboreal, littoral to bathyal.
Koroga Holmes Koroga Holmes, 1908.
Type-species: K. megalops Holmes, 1908 (original designation). See Gurjanova, 1962.
Upper lip projecting lobately in front of epistome; mandibular molar short, conical or laminate, setulose, palp attached slightly proximal to molar; mazillae stout, mavzillipedal palp article 4 long (Pseudokoroga); gnathopod 1 subchelate, palm broad, transverse, article 6 slightly broadened, much longer than article 5; telson apically notched. Species: 1, cosmopolitan, epipelagic.
Kyska Shoemaker Kyska Shoemaker, 1964.
Type-species: K. dalli Shoemaker, 1964 (original designation). Upper lip not projecting in front of epistome; mandibular molar of
MARINE GAMMARIDEAN AMPHIPODA 347
medium size, conical, setulose, lacking ridges, palp attached distal to molar; gnathopod 1 chelate, article 6 and chela slender, article 6 much longer than 5; inner ramus of uropod 2 unconstricted; telson of medium length, deeply cleft; gills plaited on both sides. Like Anonyz but enathopod 1 chelate. Species: 1, Alaska, littoral.
Lepidepecreella Schellenberg
Lepidepecreella Schellenberg, 1926a. Paracyclocaris K. H. Barnard, 1930. Type-species: JL. ctenophora Schellenberg, 1926a (monotypy). Epistome forming immense anterior keel on head, projecting in front of upper lip but together coalesced; mouthpart field thus giving conical appearance, mandible slender, molar bulge present, palp attached proximal to molar; gnathopod 1 simple, styliform, article 7 apically (and marginally) setose; coxae 1 and 2 small, partially hidden by coxa 3; outer ramus of uropod 3 shorter than peduncle, 1- or 2-articulate, inner ramus one third (rarely one half) as long as outer or absent; telson short, entire. Species: 5, bipolar, littoral to bathyal.
Lepidepecreoides K. H. Barnard
Lepidepecreoides K. H. Barnard, 1931; 1932.
Type-species: L. xenopus K. H. Barnard, 1931 (original desig- nation); 19382.
Head deeply concavotruncate from lateral view (Hippomedon); upper lip lobately projecting in front of epistome; mandibular molar small, palp attached level with molar; gnathopod 1 subchelate, article 5 longer than 6; article 7 of gnathopod 2 rather large, palm slightly chelate; coxa 5 as deep as coxa 4 (Hippomedon); telson of medium length, deeply cleft. Species: 1, subantarctic, littoral.
Lepidepecreopsis Stephensen Lepidepecreopsis Stephensen, 1925a.
Type-species: L. biloba Stephensen, 1925a (monotypy). See Tmetonyz Stebbing and Tryphosella Bonnier.
Epistome broadly rounded and dominating upper lip, latter slightly produced, both separated by deep incision; mandibular molar sub- columnar, not triturative (Tryphosella), palp attached level with molar or at proximal corner; coxa 1 slightly tapering distally and partially hidden by coxa 2; gnathopod 1 subchelate, articles 5 and 6 subequal in length or article 5 shorter than 6; telson of medium length, deeply cleft. Species: 1, subarctic Atlantic, abyssal.
285-135 O - 69 - 23
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Lepidepecreum Bate and Westwood
Lepidepecreum Bate and Westwood, 1868.—Stebbing, 1906.
Type-species: Anonyx longicornis Bate and Westwood, 1863 (monotypy and subsequent synonymy). See Sars, 1895.
Article 1 of antenna 1 usually carinate (combining characteristic) but occasionally not; accessory flagellum absent (type) or present; epistome broadly produced in front of upper lip; body often dorsally carinate or toothed; mandibular molar a setulose, quadrate lamina, palp attached proximal to molar; gnathopod 1 subchelate, articles 5 and 6 subequal; telson long, deeply cleft. Some species approach Orchomene complex. Species: 19, bipolar, amphiboreal, littoral to bathyal.
Lysianassa Milne Edwards, provisional synonymy
Lysianassa Milne Edwards, 1830.—Stebbing, 1906.
Lystanassina Costa, 1867.
Lysianax Stebbing, 1888.
Lysianopsis Holmes, 1903; 1905. Type-species: L. alba Holmes, 1903 (mono- typy). See Shoemaker, 1933b.
Aruga Holmes, 1908. Type-species: A. oculata Holmes, 1908 (monotypy).
Arugella Pirlot, 1936. Type-species: A. heterodonta Pirlot, 1936 (original designa- tion).
Shoemakerella Pirlot, 1936. Type-species: Lysianassa nasuta Dana, 1853 (original designation). Hurley (1963) notes that Pirlot’s diagnosis of Shoe- makerella fits Lysianax cubensis Stebbing, 1897, and not -L. nasuta. However, the latter should remain as type-species because it was so designated.
?Pronannonyx Schellenberg, 1953. Type-species: P. minimus Schellenberg, 1953 (monotypy). [See also in alphabetical order.]
Type-species: JL. costae Milne Edwards, 1830 (denoted by elimina- tion of Milne Edwards, 1840, and selected by Boeck, 1876). Species obscure. Genus based here on L. plumosa Boeck (L. costae of Sars, 1895, pl. 16, fig. 1).
Upper lip strongly lobate in front of epistome or epistome pro- duced forward along with upper lip; mandibular molar large, coni- colaminate, unridged or nearly obsolete, palp attached proximal to molar; gnathopod 1 simple, articles 5 and 6 subequal; inner ramus of uropod 2 unconstricted (L. plumosa), or constricted (L. longicornas) ; outer ramus of uropod 3 uniarticulate; telson entire, emarginate, or minutely notched. Species, including synonymous genera: 24, cosmopolitan, littoral (rarely in bathyal). Species with aberrant maxilla 2: L. hypocrita Ruffo.
MARINE GAMMARIDEAN AMPHIPODA 349
Lysianella Sars Lysianella Sars, 1882.—Stebbing, 1906.
Type-species: JL. petalocera Sars, 1882 (original designation). See Sars, 1895.
Article 4 of antenna 2 tumid, article 5 slender; upper lip lobately produced in front of epistome; mandibular molar vestigial, palp at- tached proximal to molar; gnathopod 1 subchelate, often weakly, articles 5 and 6 equal; inner ramus of uropod 2 constricted or not; telson of medium length, entire or cleft 25 percent. Species: 3, boreal and subtropical Atlantic and Mediterranean, littoral to abyssal.
Menigrates Boeck Menigrates Boeck, 1871.—Stebbing, 1906.
Type-species: Anonyz obtusifrons Boeck, 1861 (monotypy). See Sars, 1895.
Upper lip and epistome not differentially produced, inconspicuous; mandibular molar obsolescent, an undifferentiated lamina, palp at- tached slightly proximal to molar; gnathopod 1 simple, article 6 longer than 5; telson short, cleft one third (type) or a quarter or less; peduncular articles 2 and 3 of antenna 1 short, half as long as article 1 of flagellum (Waldeckia and Socarnes). See Paralibrotus. Species: 3, subarctic, littoral.
Menigratopsis Dahl Menigratopsis Dahl, 1945.
Type-species: MM. svenillsoni Dahl, 1945 (monotypy).
Epistome and upper lip indistinct from each other; mandibular molar well developed, but possibly not triturative, palp attached level with molar; gnathopod 1 simple, article 5 slightly longer than article 6; telson of medium length, deeply cleft. Species: 1, the Sound between Sweden and Denmark, littoral.
Mesocyclocaris Birstein and Vinogradov Mesocyclocaris Birstein and Vinogradov, 1964.
Type-species: MM. gracilis Birstein and Vinogradov, 1964 (mono- typy).
Accessory flagellum absent; mandible lacking molar and palp; gnathopod 1 simple, article 5 longer than 6; coxae 1 and 2 much smaller than coxae 3 and 4 and probably partially hidden by coxa 3; pereopods 1-5 prehensile; uropod 3 elongate, rami interequal, outer 1-articulate; [telson unclear]. Species: 1, Indian Ocean, bathypelagic.
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Mesocyphocaris Birstein and Vinogradov Mesocyphocaris Rirstein and Vinogradov, 1960.
Type-species: M/. longicaudatus Birstein and Vinogradov, 1960 (original designation).
Accessory flagellum 1-articulate; head somewhat deformed; man- dible lacking molar, palp attached medially on mandibular body; enathopod 1 simple, articles 5 and 6 subequal; coxae 1 and 2 much smaller than coxae 3 and 4 and probably partially hidden by coxa 3; pereopods 1-4 prehensile; uropod 3 elongate, outer ramus 2-articulate, inner very short and scale-like; telson short, cleft one third its length. Species: 1, Pacific, abyssopelagic.
Metacyclocaris Birstein and Vinogradov Metacyclocaris Birstein and Vinogradov, 1955.
Type-species: MM. polycheles Birstein and Vinogradov, 1955 (orig- inal designation).
Mandible lacking molar, palp attached rather distally; gnathopod 1 simple, article 5 longer than 6, gnathopod 2 slender, nearly simple; coxa 2 much larger than coxa 1 and covering it, coxae 3 and 4 not enlarged as in other cyphocarid genera; pereopods all prehensile; telson long, deeply cleft. Species: 1, N.W. Pacific, abyssopelagic
Metacyphocaris Tattersall Metacyphocaris Tattersall, 1906.
Type-species: MM. helgae Tattersall, 1906 (original designation).
Head deformed; accessory flagellum 1-articulate; mandible with very small, smooth molar, palp absent; gnathopod 1 simple, articles 5 and 6 equal; gnathopod 2 simple; coxae 1 and 2 small, partially hidden by coxa 3; pereopods 1-3 prehensile; outer ramus of uropod 3 elongate, 2-articulate, inner ramus very short, scale-like; telson of medium length, cleft one fourth. Species: 1, cosmopolitan, bathypelagic.
Metambasia Stephensen Metambasia Stephensen, 1923.
Type-species: MM. faeroensis Stephensen, 1923 (monotypy).
Article 3 of male antenna 2 tumid; upper lip projecting slightly in front of epistome; mandibular molar a long conical lamina, palp attached level with molar; gnathopod 1 simple, article 5 longer than 6; coxa 1 short, half as long as its second article, partially hidden by coxa 2; inner ramus of uropod 2 constricted; telson cleft halfway or more. See Schisturella. Species: 1, N. Atlantic, bathyal.
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Microlysias Stebbing Microlysias Stebbing, 1918.
Type-species: MM. xenokeras Stebbing, 1918 (monotypy). See K. H. Barnard, 1937.
Article 4 of male antenna 2 very tumid; epistome slightly protruding in front of upper lip; mandibular molar feeble, palp attached quite proximally; maxillipedal palp 3-articulate, possibly with a minute fourth article; gnathopod 1 subchelate, article 6 slightly longer than 5; telson of medium length, deeply cleft; gills plaited. Species: 2, S. Africa, S. Arabian coast, littoral.
Nannonyx Sars Nannonyz Sars, 1895.
Type-species: Orchomene goesit Boeck, 1871. (monotypy). See Sars, 1895.
Mouthparts substyliform; epistome and upper lip coalesced, epistomal portion projecting slightly in front of upper lip; mandibular molar vestigial, palp attached proximally; maxillipedal palp scarcely exceeding outer plate, article 4 very small (except N. spinimanus Walker); gnathopod 1 appearing simple but with microscopic palm, articles 5 and 6 short and stout; rami of uropod 3 equal to or shorter than peduncle; telson entire or slightly emarginate. Species: 4, boreal-subtropical N.E. Atlantic and Mediterranean, Kerguelen Island, littoral.
Neoambasia Dahl Neoambasia Dahl, 1959.
Type-species: Ambasiopsis tumicornis Nicholls, 1938 (monetypy).
Upper lip projecting slightly in front of epistome; mandibular molar apparently well developed, setose, palp attached level with molar; outer plate of maxilla 1 poorly spinose (Pseudambasia) ; outer plate of maxilliped spinose (Ambasiopsis); gnathopod 1 subchelate, article 6 slightly longer than 5; coxa 1 not greatly shortened, but almost concealed by coxa 2, about as long as article 2 of gnathopod 1; rami of uropod 3 not longer than peduncle; telson short, deeply cleft. Species: 1, antarctic, bathyal.
Normanion Bonnier
Normania Boeck, 1871 (homonym, Ostracoda). Normanion Bonnier, 1893 (new name).—Stebbing, 1906.
Type-species: Opis quadrimana Bate and Westwood, 1868 (mono- typy). See Sars, 1895. Even though Boeck’s identification of O.
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quadrimana was in error, the type should remain that species for purposes of stability. Boeck’s material was named Normanion sarsi by Stebbing (1906).
Mouthparts forming pseudoconical bundle or not; epistome appla- nated, not distinct from upper lip; mandibular molar conical or nearly obsolete, not ridged (?except N. abyssi Chevreux), palp attached proximal to molar; outer plates of maxilliped almost or ereatly exceeding palp, palp essentially 3-articulate, scarcely or slightly exceeding inner plates, article 4 obsolete; gnathopod 1 mod- erately powerful, subchelate, palm transverse; peduncle of uropod 3 elongate; telson short, quadrate, entire. Species: 3, N.E. Atlantic, Mediterranean, littoral to abyssal (some fish commensalism).
Ocosingo J. L. Barnard Ocosingo J. L. Barnard, 1964b.
Type-species: 0. borlus J. L. Barnard, 1964b (original designation).
Mouthparts substyliform, forming ventral subconical bundle; man- dible bearing vestigial molar ridge, palp attached quite proximally; palp of maxilla 1 uniarticulate; maxillipedal palp scarcely exceeding outer plate, 3-articulate, ultimate article claviform; gnathopod 1 simple, article 6 longer than 5; coxa 1 projecting over side of head; uropod 3 lacking rami; telson small, entire. Species: 1, California, littoral.
Onesimoides Stebbing Onesimoides Stebbing, 1888; 1906.
Type-species: O. carvnatus Stebbing, 1888 (monotypy).
Upper lip and epistome not differentially produced, mandibular molar ridged, palp attached level with molar; gnathopod 1 subchelate or slightly chelate, article 5 much longer than 6, gnathopod 1 rather enlarged in males; outer ramus of uropod 3 biarticulate, inner ramus half or less as long as outer; telson entire; article 1 of first antennal flagellum 5 times as long as neat article (Paronesimoides). Species: 3, Indo-Pacific tropics, bathyal to hadal.
Onisimus Boeck, new synonymy
Onisimus Boeck, 1871. Pseudalibrotus Della Valle, 1893.—Stebbing, 1906.
Type-species: Anonyzx litoralis Krgyer, 1845 (selected by Boeck, 1876). Stebbing, Sars, and Della Valle apparently overlooked the fact that Boeck (1876) selected Anonyz litoralis as type of Onisumus. Della Valle erected Pseudalibrotus and cited only A. litoralis, thus making it type of the genus by monotypy. All species of Pseudali-
MARINE GAMMARIDEAN AMPHIPODA 353
brotus, having gone under that name for over 60 years, must now be transferred to Onisumus and the 15 or more species of Onisimus, having gone under that name for nearly 100 years, must now be transferred to a new appellation, which I designate as Boeckosimus in honor of Axel Boeck, one of the first students of Amphipoda to understand the importance of type designations.
The genus Alibrotus Milne Edwards (1840) with its monotype, A. chausercus Milne Edwards (1840) may be a senior synonym of either Onisimus (auct. =Pseudalibrotus) or Boeckosimus, new name, but the problem has never been clarified and may be insoluble. The problem should be reviewed at least once more by a student of arctic- boreal faunas and perhaps submitted to ICZN in order to cancel the genus Alibrotus.
Epistome broadly rounded and projecting slightly in front of upper lip; mandibular molar ridged, palp attached level with molar; gnath- opod 1 subchelate, articles 5 and 6 equal; inner ramus of uropod 2 constricted or not; telson short, entire. See Boeckosimus. Species: 7, arctic littoral, Caspian Sea, glacial relicts.
Opisa Boeck
Opis Krgyer, 1842 (homonym, Mollusca). Opisa Boeck,1876.—Stebbing, 1906.
Type-species: Opis eschrichtia Kroyer, 1842 (monotypy and sub- sequent synonymy; Boeck (1876) cited O. typica Krgyer as type- species, probably as a technical error for he recognized it was a junior synonym of O. eschrichtir).
Upper lip rounded, slightly projecting in front of epistome; mandib- ular molar obsolete, palp attached proximally; gnathopod 1 powerful, chelate, chela broad, palm excavate to form thumb; coxa 1 slightly shortened and partially hidden by coxa 2; telson long, deeply cleft. Species: 2, boreal, littoral (to 432 m).
Orchomene Boeck, new synonymy
Orchomene Boeck, 1871.—Stebbing, 1906.
Tryphosa Boeck, 1871.—Stebbing, 1906.
Orchomenella Sars, 1895.*—Stebbing, 1906.
Orchomenopsis Sars, 1895.—Stebbing, 1906.
?Allogaussia Schellenberg, 1926a. [See also in alphabetical order.]
Type-species: Anonyx serratus Boeck, 1861 (selected by Boeck, 1876). See Sars, 1895* (as Orchomenella ciliata). Regrettably Sars and Stebbing overlooked the fact that Boeck (1876) had selected the type- species of Tryphosa as Anonyx nanus Krgyer (1846) a species un-
*This portion of Sars’ monograph was published in 1891.
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doubtedly congeneric with the type-species of Orchomenella, Anonyx minutus Kroyer, 1846 (original designation). Sars had incorrectly identified Anonyz nanus and his material later was named Tryphosella sarsi by Bonnier (1893); indeed Sars had in hand Anonyz nanus but he erected for the material the name Orchomenella ciliata, later shown by Stebbing (1906) to be a synonym of A. nanus. Unfortunately Stebbing did not therefore synonymize Orchomenella with ‘‘Tryphosa,” since the two type-species were congeneric. Thus, Orchomenella must fall to Tryphosa and all species of Tryphosa (auct.), so well known since the time of Sars (1895) and Stebbing (1906) must be given a different generic name, which is 7ryphosella Bonnier.
Epistome generally broadly lobate in front, occasionally acute, or flat and unproduced, lobe of upper lip generally prominent but rarely projecting far in front of epistome; mandibular small, poorly ridged or weakly setulose, palp attached proximal to molar; gnathopod 1 subchelate, article 6 longer than 5; telson variable, entire (Adllogaussia) or minutely and deeply cleft. See Lepidepecreum for intergrading species. Species (including Allogaussia): 61-+-, cosmopolitan, cold- water, littoral to abyssal. ‘“Orchomenella’”’ groenlandica (Hansen) is treated in Keys L, N, O, and P as a distinct generic taxon.
Pachychelium Stephensen Pachychelium Stephensen, 1925a.
Type-species: P. davidis Stephensen, 1925a (original designation).
Antenna 1 very stout, body slender; mandibular molar absent, palp attached rather distally; maxilla 1 lacking palp; maxilla 2 with vestigial inner plate; maxilliped lacking inner plate or inner plate very small; enathopod 1 enlarged, subchelate, article 6 large, ovate; telson short, entire. Species: 3, bipolar, littoral to bathyal.
Pachynus Bulycheva Pachynus Bulycheva, 1955.
Type-species: P. chelatum Bulycheva, 1955 (original designation).
Upper lip and epistome apparently not differentially produced; mandibular molar absent, palp attached rather distally; maxillipedal palp 3-articulate, or with a small fourth article, article 3 elongate or not; gnathopod 1 chelate; telson short, entire. Species: 2, boreal Pacific, littoral to bathyal.
Paracallisoma Chevreux Paracallisoma Chevreux, 1903.
Type-species: P. alberti Chevreux, 1903 (original designation).
MARINB GAMMARIDEAN AMPHIPODA 355
[Upper lip and epistome not studied]; mandibular molar small, simple, palp attached level with molar, both molar and palp attached proximally; inner plate of maxilla 1 densely setose; gnathopod 1 simple, with apical setal shroud, article 7 vestigial; gnathopod 2 subchelate (Aroui, Scopelocheirus); telson long, deeply cleft. Species: 2, Atlantic-Pacific, bathypelagic.
Paracallisomopsis Gurjanova Paracallisomopsis Gurjanova, 1962.
Type-species: P. beljaevi Gurjanova, 1962 (monotypy).
Head lacking lateral lobes; upper lip and epistome not differenti- ally produced; mandibular molar conical, unridged, palp attached level with molar; gnathopod 1 simple, article 7 vestigial, shrouded with cirri; telson of medium length, deeply cleft.
Differs from Eucallisoma J. L. Barnard (1961) by: the larger head, smaller first antenna with its less elongate flagellar base, the acces- sory flagellum conspicuous (in Hucallisoma it is appressed to inner face of the immense antenna and is brush-like); the less styliform first gnathopod, the cirri more dense, dactyl not distinct from cirri; more spines on the outer plate of the maxilliped; the shorter second article on the outer ramus of uropod 3. Species: 1, Bering Sea, ?epipelagic.
[Paracentromedon Chevreux and Fage]
Paracentromedon Chevreux and Fage, 1925. Presumed synonym of Hippomedon. Elimedon J. L. Barnard, 1962d.
Type-species: Centromedon crenulatum Chevreux, 1900 (original designation).
Head small, partially hidden by coxa 1 (Tryphosella); upper lip and epistome not differentially produced or inconspicuous; mandibular molar ridged, palp attached level with molar; gnathopod 1 nearly simple, articles 5 and 6 equal, article 7 greatly overlapping minute palm; telson of medium length, deeply cleft. See Paronesimus. Spe- cies: 3, Atlantic, bathyal-abyssal.
Paracyphocaris Chevreux Paracyphocaris Chevreux, 1905a.
Type-species: P. praedator Chevreux, 1905a (original designation).
Head deformed; mandible lacking molar, palp attached medially; enathopod 1 simple or very minutely subchelate, article 5 slightly longer than 6; gnathopod 2 nearly simple; coxae 1 and 2 small, par- tially hidden by coxa 3; pereopods 1-4 (or 5) prehensile; telson long, deeply cleft. Species: 2, Atlantic-Pacific, bathy-abyssopelagic.
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Paralibrotus Stephensen Paralibrotus Stephensen, 1923.
Type-species: P. setosus Stephensen, 1923 (monotypy).
Epistome and upper lip not differentially produced; mandibular molar small but distinct, possibly triturative, palp attached over proximal end of molar; gnathopod 1 simple, rather stout, article 6 slightly longer than 5; article 2 of outer ramus on uropod 3 minute; telson short, entire. Merges with some species of Menigrates. Species: 1, subarctic, littoral.
Paralicella Chevreux Paralicella Chevreux, 1908a.
Type-species: P. tenuipes Chevreux, 1908a (original designation).
Upper lip and epistome not described; mandible lacking molar in type-species or bearing conical setulose lamina in other species; palp attached rather distally; inner plate of mavilla 1 densely setose, with 10 or more setae (combining character); gnathopod 1 subchelate, article 6 slightly longer than 5, article 3 elongate, nearly as long as article 6 (Aristias, Eurythenes); telson of medium length, deeply cleft. Species: 4, Atlantic-Pacific, bathyal-abyssal.
Paralysianopsis Schellenberg
Paralysianopsis Schellenberg, 1931. Austronisimus K. H. Barnard, 1931.
Type-species: P. odhneri Schellenberg, 1931 (monotypy).
Upper lip produced acutely in front of epistome, but almost fully coalesced with epistome; mandibular molar obsolescent, palp attached slightly proximal to molar; gnathopod 1 nearly simple, with minute oblique palm; inner ramus of uropod 2 constricted; telson entire, with minutely notched apex. Species: 1, subantarctic, littoral.
Parambasia Walker and Scott Parambasia Walker and Scott, 1903.
Type-species: P. forbesi Walker and Scott, 1903 (monotypy).
Diagnosis based partially on P. rossi Stephensen (1927) assuming it to be congeneric. [Epistome and upper lip poorly known], probably fused together; mandibular molar small, setulose, palp attached proximal to molar; gnathopod 1 simple, article 5 slightly longer than article 6; inner ramus of uropod 2 constricted (possibly not on type) ; outer ramus of uropod 3 uniarticulate; telson entire. Species: 2, Arabian Sea, Auckland Islands, littoral. Possibly congeneric: Lyst- anassa anomala Nicholls, 1938, subantarctic.
MARINE GAMMARIDEAN AMPHIPODA 357
Parawaldeckia Stebbing
Parawaldeckia Stebbing, 1910.
Type-species: Nannonyz thomson Stebbing, 1906 (monotypy).
Article 5 of male antenna 2 slightly dilated and longer than article 4; [prebuccal area and mandible unknown but presumably similar to Waldeckia]; gnathopod 1 simple, articles 5 and 6 equal; uropod 3 short, outer ramus not longer than peduncle, peduncle sublamelliform, inner ramus scarcely half as long as outer; telson entire or slightly emarginate. Species: 2,* antiboreal, littoral.
*Parawaldeckia kiddert (Smith).—Generic diagnosis: to some extent mouth- parts bundled conically, upper lip and epistome confluent, slightly convex ante- riorly, lacking sinus; mandibular molar conical, unridged, distally setose, palp attached very proximally; gnathopod 1 simple; uropod 3 with vestigial inner ramus; telson short, entire. Diagnosis based on Monod (1926) and cotypes examined in U.S. National Museum; general appearance of cotypes, without dissection, similar to Monod’s figures and not to concepts of Chilton (1909, 1921), Stephensen (1927), and Tattersall (1922) in the following characters: general appearance including coxae and pleon, (antenna 2 slightly shorter than shown by Monod), prebuccal region, telson, and uropod 3.
Paronesimoides Pirlot
Paronesimoides Pirlot, 19338a.
Type-species: P. lignivorus Pirlot, 1933a (original designation).
Apparently upper lip projecting and dominating epistome; man- dibular molar not strongly produced, setulose, poorly ridged, palp attached at proximal end of molar; gnathopod 1 subchelate, palm transverse, article 6 slightly enlarged and much larger than article 5; uropod 3 very small, outer ramus 2-articulate, equal to peduncle, inner ramus reduced to a minute spiniform process; telson short, entire; articles 1 and 2 of flagellum of antenna 1 equal in length (Onesimoides). Species: 1, Indonesia, abyssal.
Paronesimus Stebbing
Paronesimus Stebbing, 1894; 1906.
Type-species: P. barentsi Stebbing, 1894 (monotypy). See Gurjanova, 1962.
[Upper lip and epistome not described, presumably like Onisimus]; mandibular molar small, weakly ridged, palp attached level with molar; gnathopod 1 with very small palm, appearing simple, articles 5 and 6 subequal; telson of medium length, cleft about halfway. Very close to Hippomedon. See Paracentromedon. Species: 2, arctic, littoral.
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Perrierella Chevreux and Bouvier
Perrierella Chevreux and Bouvier, 1892. Pararistias Robertson, 1892.
_ Type-species: Lysianassa audowimana Bate, 1857a (original des- ignation and subsequent synonymy). See Sars, 1895.
Upper lip and epistome not differentially produced; mand bular molar a vestigial hump, palp attached level with molar; lobes of maxilla 2 slightly gaping, inner broader than outer; maxillipedal palp not exceeding outer plate, 3-articulate or with vestigial fourth article, inner plates vestigial; gnathopod 1 nearly simple, with poorly defined palm; coxa 1 very small and largely hidden by coxa 2; telson of me- dium length, entire, apically truncate. Species: 1, boreal and warm- temperate E. Atlantic and Mediterranean, littoral.
Phoxostoma K. H. Barnard Phozostoma K. H. Barnard, 1925.
Type-species: P. algoense K. H. Barnard, 1925 (monotypy).
Mouthparts forming conical bundle; upper lip and epistome con- tinuous; mandible slender, molar obsolete, palp attached quite prox- imally; maxillipedal palp slightly exceeding outer plate, article 4 very small; gnathopod 1 simple, article 6 longer than 5; telson short, deeply insinuate. Species: 1, 8. Africa, littoral.
Podoprion Chevreux Podoprion Chevreux, 1891.—Stebbing, 1906.
Type-species: P. bolwari Chevreux, 1891 (monotypy). See Chevreux and Fage, 1925.
Epistome and upper lip not differentially produced; mandible lack- ing molar, palp attached rather distally, ?cutting edge dentate; gnath- opod 1 chelate, thumb narrow; article 2 of pereopod 3 deeply indentured; coxa 1 shorter than coxa 2 and partially hidden by it; telson of medium length, deeply cleft. See Huwonyx. Species: 1, warm-temperate E. Atlantic, littoral.
Podoprionella Sars
Podoprionella Sars, 1895.—Stebbing, 1906.
Type-species: P. norvegica Sars, 1895 (monotypy).
Epistome and upper lip not differentially produced; mandible lack- ing molar, palp attached rather proximally; maxillipedal palp shorter than outer plate, 3-articulate; gnathopod 1 chelate; article 2 of pere-
MARINE GAMMARIDEAN AMPHIPODA 359
opods 38-5 deeply indentured; outer ramus of uropod 3 uniarticulate; telson entire. Species: 1, boreal N. E. Atlantic, littoral.
Podoprionides Walker Podoprionides Walker, 1906a.
Type-species: P. incerta Walker, 1906a (monotypy). See Walker, 1907.
Epistome and upper lip not differentially produced; mandible lack- ing molar, palp attached rather proximally; maxillipedal palp not ex- ceeding outer plate, 3-articulate; gnathopod 1 chelate; article 2 of pereopods 3-5 deeply indentured; telson of medium length, cleft more than halfway. Species: 1, antarctic, littoral.
Prachynella J. L. Barnard Prachynella J. L. Barnard, 1964b.
Type-species: P. lodo J. L. Barnard, 1964b (original designation).
Upper lip and epistome small, not differentially produced; man- dibular molar absent, palp attached about in middle of mandibular body; maxilla 1 lacking palp; maxillipedal palp with three articles, article 3 long and claw-like; gnathopod 1 chelate; telson short, entire. Species: 1, California, littoral.
Procyphocaris J. L. Barnard Procyphocaris J. L. Barnard, 1961.
Type-species: P. primata J. L. Barnard, 1961 (original designa- tion).
Head not deformed; epistome and upper lip not differentially produced; mandibular molar ridged, palp attached level with molar; enathopod 1 scarcely subchelate, palm very oblique; coxae 1 and 2 small, largely hidden by coxa 3; telson long, deeply cleft. Species: 1, Australia, bathypelagic.
[Pronannonyx Schellenberg] Pronannonyx Schellenberg, 1953. Presumed synonym of Lysianassa.
Type-species: P. minimus Schellenberg, 1953 (monotypy).
Upper lip lobately produced forward (possibly including the epi- stome); mandibular molar vestigial, palp attached quite proximally; maxilla 1 unth two kinds of spines on the outer plate (combining charac- ter); gnathopod 1 simple, article 6 longer than 5; uropod 3 with l-articulate outer ramus equal to peduncle, peduncle with plate-like expansion; telson short, entire; inner ramus of uropod 2- simple. Species: 1, S. W. Africa, littoral.
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Pseudambasia Stephensen Pseudambasia Stephensen, 1927.
Type-species: P. bipartita Stephensen, 1927 (monotypy).
Article 2 of antenna 1 elongate, two thirds as long as article 1; epistome obtusely produced in front of upper lip; [mandible and maxilla 1 undescribed]; gnathopod 1 subchelate, article 6 slightly longer than 5, palm excavate (combining character); outer ramus of uropod 3 uniarticulate, rami equal to each other and peduncle; telson of medium length, entire; urosomal segments apparently coalesced. Species: 1, Auckland Islands, littoral.
Pseudoanonyx Kudrjaschov
Pseudoanonyx Kudrjaschov, 1965.
Type-species: P. caecus Kudrjaschov, 1965 (original designation):
Apparently upper lip projecting slightly in front of epistome [from analogy to Anonyx]; mandibular molar large, unridged (Onisimus), setulose, palp apparently attached level with molar; gnathopod 1 subchelate, articles 5 and 6 stout, subequal in length, dactyl very small and like dactyl of gnathopod 2; inner ramus of uropod 2 simple; telson of medium length, deeply cleft; gills plaited; eyes absent; pereopods 1-2 without distal locking spines on article 6 at base of dactyl; article 4 of mazillipedal palp very short, tumid (Anonyz). Species: 1, Okhotsk Sea, littoral.
Pseudokoroga Schellenberg
Pseudokoroga Schellenberg, 1931.
Type-species: P. barnardi Schellenberg, 1931 (monotypy).
Upper lip and epistome produced forward together; mandibular molar small, ridged, palp attached proximal to molar; mazilae slender, article 4 of mazillipedal palp short (Koroga); gnathopod 1 subchelate, article 6 enlarged, much longer than article 5; inner ramus of uropod 2 constricted; telson of medium length, entire. Species: 2, subantarctic, Baja California, littoral.
Pseudorchomene Schellenberg
Pseudorchomene Schellenberg, 1926a.
Type-species: Orchomenopsis coats Chilton, 1912 (monotypy).
Epistome broadly lobate in front of upper lip; mandibular molar small, poorly ridged, palp attached proximal to molar; gnathopod 1 poorly subchelate, articles 5 and 6 equal, article 3 as long as 5 or 6,
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all these articles slender and elongate (combining character); telson deeply cleft. Species: 1, antarctic, littoral.
Rifcus Kudrjaschoy Rifcus Kudrjaschov, 1965.
Type-species: FR. auspicatus Kudrjaschov, 1965 (original designa- tion).
Upper lip and epistome apparently not prominent; accessory flagel- lum 2-articulate; mandibular molar ridged (Anonyz), palp attached level with molar; gnathopod 1 weakly subchelate, articles 5 and 6 subequal in length; inner ramus of uropod 2 not constricted; telson short (Hippomedon), cleft less than halfway; outer plate of maxilla 1 with only five spines (Tryphosoides); palp of mazilliped very stout and short, outer plate of mazilliped reaching nearly to end of palp article 3 (Boeckosumus). Species: 1, Okhotsk Sea, littoral.
Schisturella Norman, new synonymy
Schisturella Norman, 1900a. Pseudonesimus Chevreux, 1926.
Type-species: Zryphosa pulchra Hansen, 1887 (monotypy). See Shoemaker, 1930.
Upper lip lobately produced in front of epistome or not (Pseudo- nesimus) but both separated by a notch; mandibular molar well developed or small, blunt, ridged, palp attached level with or at proximal end of molar; gnathopod 1 weakly or strongly subchelate, article 5 longer than or equal to 6; coxa 1 slightly or strongly shortened, often half as long as its article 2, largely hidden by coxa 2; inner ramus of uropod 2 constricted; telson of medium length, cleft one third to three quarters of its length; (article 3 of antenna 2 slightly tumid). See Metambasia. Species: 3, Atlantic-Pacific, coldwater, littoral to hadal.
Scopelocheiropsis Schellenberg Scopelocheiropsis Schellenberg, 1926a.
Type-species: S. abyssalis Schellenberg, 1926a (monotypy).
Epistome lobately produced slightly in front of upper lip; mandible lacking molar, palp attached rather distally; inner plate of maxilla 1 densely setose; maxillipedal palp article 4 vestigial; gnathopod 1 linear, simple, articles 5 and 6 equal, dactyl vestigial and shrouded with a short tuft of setae; pereopods 1-2 subprehensile, article 6 longer than articles 4-5 combined (combining character); telson deeply cleft. Species: 1, tropical Atlantic, abyssal (?pelagic).
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Scopelocheirus Bate
Callisoma Costa, 185la, b (homonym, Coleoptera) [type species: Callisoma hopei Costa, 185la, b, selected by Boeck, 1876].
Scopelocheirus Bate, 1857a.—Stebbing, 1906.—Birstein and Vinogradov, 1964.
Bathycallisoma Dahl, 1959 [see also this genus in alphabetical order].
Type-species: SS. crenatus Bate, 1857a(monotypy). SeeSars, 1895.
Epistome slightly and lobately produced in front of upper lip (Arouz); mandibular molar conical, simple, palp attached level with molar; inner plate of maxilla 1 densely setose on inner edge; gnathopod 1 simple, linear, article 7 shrouded in setae or cirri, article 6 longer than 5; gnathopod 2 minutely chelate (Paracallisoma, Bathycallisoma) ; telson medium to long, deeply cleft. Species: 3 or 4, probably cosmopolitan, bathy-abyssopelagic.
Shackletonia K. H. Barnard Shackletonia K. H. Barnard, 1931; 1932.
Type-species: S. robusta K. H. Barnard, 1931 (original designa- tion) ; 1932.
Mouthparts presumed to be produced in a ventral conical bundle; epistome and upper lip not differentially produced, apparently coalesced; mandibular molar slightly conical, palp attached slightly proximal to molar; outer plate of maxilla 1 with hook-like spines; gnath- opod 1 simple, stout; rami of uropod 3 ovate, outer with a minute, spine-like article 2; telson of medium length, deeply cleft. Species: 1, antarctic, deep littoral.
Socarnella Walker Socarnella Walker, 1904.
Type-species: S. bonniera Walker, 1904 (monotypy).
[Upper lip and epistome not described]; mandibular molar absent, palp attached proximally; gnathopod 1 simple, article 6 longer than 5; outer ramus of uropod 3 uaiarticulate; telson short, emarginate or slightly cleft; mandibular palp article 2 about 5 tumes as long as article 1 (Socarnopsis). Related to Lysianassa [=Lysianopsis, Arugella, etc.] Species: 1, Ceylon, littoral.
Socarnes Boeck
Socarnes Boeck, 1871.—Stebbing, 1906 (in part, not Ephippiphora White, see Waldeckia herein).
Type-species: Lysianassa vahlii Krdyer, 1838 (monotypy). See Sars, 1895.
MARINE GAMMARIDEAN AMPHIPODA 363
Diagnosis based only on type-species. Upper lip lobately produced in front of epistome or both prominently produced; mandibular molar conicolaminate, unridged, palp attached proximal to molar; gnathopod 1 simple, articles 5 and 6 equal; outer ramus of uropod 3 biarticulate (Socarnopsis) ; telson of medium length, cleft slightly more than half- way; gills plaited on one side only (Ichnopus and Socarnopsis), [or two sides in S. obesa (Chevreux)]; pereopod 5 not longer than pereopod 4 (Ichnopus). See Waldeckia. Species: 5 or 6, amphiboreal, subtropical, littoral to bathyal.
Socarnoides Stebbing
Socarnoides Stebbing, 1888; 1906.—Schellenberg, 1931. Acidostomella Schellenberg, 1926b.
Type-species: S. kerguelent Stebbing, 1888 (monotypy).
Epistome forming broad, flabellate lobe in front of somewhat lobate upper lip (type) or upper lip forming lobe exceeding unproduced epistome; mandibular molar low, moderately long, apparently ridged (type) or setulose, palp attached proximal to molar; maxillipedal palp scarcely exceeding outer plate (type only) ; gnathopod 1 simple, articles 5 and 6 subequal; inner ramus of uropod 3 constricted; telson of medium length, cleft one third (type) or deeply. Species: 4 or 5, possibly cosmopolitan (except polar), littoral.
Socarnopsis Chevreux Socarnopsis Chevreux, 191la.
Type-species: 8S. crenulata Chevreux, 1911a (monotypy).
Upper lip and epistome together produced forward lobately; man- dibular molar blunt, ridged, palp attached slightly proximal to molar; enathopod 1 simple; outer ramus of uropod 3 uniarticulate; telson of medium length, deeply cleft; gills plaited on both sides (Socarnes) ; mandibular palp article 2 about 3 times as long as article 1 (Socarnella). Species: 1, eastern Atlantic, littoral.
Sophrosyne Stebbing
Sophrosyne Stebbing, 1888; 1906. Paropisa Stebbing, 1899d.
Type-species: S. murrayi Stebbing, 1888 (monotypy).
Upper lip and epistome not differentially produced; mandibular molar absent, palp attached rather distally; outer plate of maxilla 1 poorly spinose; gnathopod 1 slightly chelate, article 6 longer than 5; telson short, deeply cleft. Species: 3, boreal-antiboreal, littoral to bathyal, coldwater.
285-135 O - 69 - 24
364 U.S. NATIONAL MUSEUM BULLETIN 271
Stephensenia Schellenberg Stephensenia Schellenberg, 1928a.
Type-species: S. haematopus Schellenberg, 1928a (monotypy).
Upper lip and epistome not differentially produced; mandibular molar large, blunt, palp absent; gnathopod 1 nearly simple, article 5 longer than 6; outer ramus of uropod 3 uniarticulate; telson of medium length, deeply cleft. Species: 1, antiboreal, littoral.
Stomacontion Stebbing Stomacontion Stebbing, 1899d; 1906.
Type-species: <Acontiostoma pepini Stebbing, 1888 (original designation).
Mouthparts forming conical bundle, styliform; mandibular molar absent, palp attached very proximally; palp of maxilla 1 biarticulate but very small; maxillipedal palp article 4 vestigial; gnathopod 1 simple, article 6 longer than 5; coxa 1 projecting over side of the head; uropod 3 uniramous, ramus vestigial; telson short, apically notched. [Based only on type-species. |
Note: Species assigned to this genus intergrade with Acontiostoma and two of the species have coxa 1 ventrally subconical. Stomacontion capense K. H. Barnard (1916) lacks any first maxillary palp and S. insigne K. H. Barnard (1932) has the first maxillary palp 1-articulate but combined with a vestigial fourth maxillipedal palp article, and these characters obfuscate the generic limits of Acontiostoma and Stomacontion. Both species, however, have a subconical first coxa, perhaps in itself a useful generic or subgeneric character. Species: 5, subantarctic, antiboreal, littoral.
Thoriella Stephensen Thoriella Stephensen, 1915.
Type-species: 7. islandica Stephensen, 1915 (original designation).
Accessory flagellum absent; flagellum of antenna 2 with especially inflated articles; apparently epistome broadly rounded and produced in front of upper lip; mandibular molar obsolete, palp absent; maxil- liped opercular, inner plates apparently large and subrectangular, outer plates narrower and triangular, palp apparently composed of single large opercular article with deep lateral incision [open to dif- ferent interpretation]; gnathopod 1 simple, short; gnathopod 2 simple; coxae small, short, not touching serially, coxa 4 not excavate pos- teriorly ; uropod 3 lacking rami; telson absent. Species: 1, N. Atlantic, Indian, abyssopelagic.
MARINE GAMMARIDEAN AMPHIPODA 365
Tmetonyx Stebbing
Hoplonyzx Sars, 1895 (homonym, Coleoptera). Tmetonyx Stebbing, 1906 (new name).
Type-species: Oniscus cicada O. Fabricius, 1780 (original desig- nation). See Sars, 1895.
This genus was synonymized with Tryphosa by J. L. Barnard, 1962d, but Tryphosa must be synonymized with Orchomenella, of which Orchomene is a senior synonym, because Boeck designated as type-species of 7ryphosa a taxon later proved to be congeneric with the designated type-species of Orchomenella. The Tryphosa concept is now given the next available name, 7ryphosella Bonnier, but 7'met- onyx is reinstated on the condition of coxa 1 to include only its type-species.
Epistome broadly rounded, upper lip broadly rounded, both pro- jecting together slightly (Anonyzx), but separated by distinct notch; mandibular molar subcolumnar, slightly attenuated, apparently mostly setulose and poorly or not ridged, palp attached level with molar; cora 1 perfectly quadrate, neither expanded nor tapered distally (Anonyz, Tryphosella); gnathopod 1 subchelate, articles 5 and 6 subequal in length; telson of medium length, deeply cleft; head of normal size (Uristes); dactyl of gnathopod 1 with wnner tooth (Anonyz). Species: 1 or 2, N.K. Atlantic Ocean, littoral to abyssal.
Trischizostoma Boeck
Guerinia Costa, 1853b, 1857 (homonym, Diptera). Trischizostoma Boeck, 1861.—Stebbing, 1906. Guerina Della Valle, 1893 (new name for Guerinia).
Type-species: Guerinia nicaeensis Costa, 1853b (monotypy and subsequent synonymy). See Sars, 1895.
Mouthparts forming a ventral conical bundle, styliform; epistome and upper lip not differentially produced; mandibular molar absent, palp attached about medially; lobes of lower lip sublanceolate, lack- ing mandibular processes; palp of maxilla 1 very short; gnathopod 1 very powerful, articles 6 and 7 generally inverted in adult so that article 7 closes on article 6 from below; coxa 1 short, dominated by coxa 2; telson short, entire or cleft to middle; branchiae apparently plaited; eyes enormous. Species: 6, cosmopolitan, bathy-abyssopelagic.
Tryphosella Bonnier (=‘“‘Tryphosa’’ auct.)
Tryphosella Bonnier, 1893 [not Stebbing, 1906]. Tryphosa.—Sars, 1895.*—Stebbing, 1906 (not Boeck, 1871).
*Sars’ book is cited herein under the date 1895 but it was printed in parts from 1891 to 1895 and the section on Tryphosa was printed in 1891, two years before Bonnier’s work.
366 U.S. NATIONAL MUSEUM BULLETIN 271
Type-species: Tryphosella sarsi Bonnier, 1893 (present selection), (=Tryphosa nana of Sars, 1895, not of Boeck, 1861). Bonnier in- tended the name Tryphosella to replace Sars’ usage of the name Tryphosa for a group of species distinct from Orchomenella Sars (=Orchomene Boeck). Sars did not realize that those species he called Orchomenella were congeneric with Jryphosa. Stebbing (1906) also failed to recognize Bonnier’s perception and restricted Tryphosella to a single species, Tryphosa barbatipes Stebbing, 1888, which later has been considered to be congeneric with Uristes Dana. Thus J. L. Barnard’s (1962d) move of Tryphosella to Uristes was illegal because the type-species of Tryphosella had never been selected. Legaliza- tion of Bonnier’s intent by selection of the type-species that he obviously would choose through his comments and his listing of it in primary position now permits reinstatement of a “tryphosa”’ group to be called a “‘tryphosella”’ group of species. They are close to Tmetonyx Stebbing, 1906, but have coxa 1 distinctly tapering distally. Lepidepecreopsis Stephensen may be asynonym of Tryphosella as Gurjanova (1951) removed the former to Tryphosa. See discussion with T'metonyz.
Epistome broadly rounded and dominating upper lip, both sepa- rated by a notch; mandibular molar subcolumnar or cuboid, weakly setulose and poorly ridged if at all (Lepidepecreopsis), palp attached level with molar; cora 1 slightly shortened, distinctly tapering (Anonyx, Tmetonyx) and partially hidden by coxa 2; gnathopod 1 subchelate, articles 5 and 6 subequal in length; telson of medium length, deeply cleft; head of normal size (Uristes) ; (dactyl of gnathopod 1 with or without inner tooth). Species: ca. 63, cosmopolitan cold- water, littoral to hadal, in low latitudes primarily bathyal-abyssal. Composition: all species formerly known under ‘‘Tryphosa”’ and ‘““Tmetonyz’”’ since 1895 except their type-species.
Tryphosites Sars Tryphosites Sars, 1895.—Stebbing, 1906.
Type-species: Anonyzx longipes Bate and Westwood, 1863 (orig- inal designation).
Epistome acutely produced in front of upper lip, upper lip also slightly lobate; mandibular molar ridged, palp attached level with molar; gnathopod 1 subchelate, article 5 longer than 6; inner ramus of uropod 2 constricted; telson of medium length, deeply cleft. Species: 5, bipolar, coldwater submergent, littoral to abyssal.
MARINE GAMMARIDEAN AMPHIPODA 367
Tryphosoides Schellenberg
Tryphosoides Schellenberg, 1931.
Type-species: T. falcata Schellenberg, 1931 (monotypy).
Epistome not produced (apparently not as in “7ryphosa’’) ; mandib- ular molar ridged, palp attached level with molar; gnathopod 1 subchelate, article 6 longer than 5; coxa 1 scarcely narrowed, slightly beveled anteroventrally; outer ramus of uropod 3 uniarticulate (‘‘Try- phosa’”’); accessory flagellum vestigial, 2-articulate; telson of medium length, deeply cleft. Species: 1, antiboreal S. America, littoral. Possibly synonymous with Uristes. See Rifcus.
Uristes Dana
Uristes Dana, 1849.—Stebbing, 1906. ?Pseudotryphosa Sars, 1895. Uristoides Schellenberg, 1931.
Type-species: U. gigas Dana, 1852a; 1853 (monotypy of Dana, 1852a). See Stebbing, 1888 (as Tryphosa antennipotens).
Head small, mouthparts largely covered by corae 1-2 (Tmetonyx and Tryphosella); epistome and upper lip generally small and incon- spicuous, often slightly produced differentially (Tmetonyzx and Try- phosella); mandibular molar setulose, palp attached level with molar; enathopod 1 subchelate or nearly simple, article 6 longer than 5 (Tmetonyx usually); coza 1 narrowing slightly distally (Hippomedon) ; telson of medium length, deeply cleft. Species: 23, cosmopolitan, coldwater submergent, littoral to abyssal.
Valettia Stebbing
Valettia Stebbing, 1888; 1906.
Type-species: V. coheres Stebbing, 1888 (monotypy).
Apparently epistome and upper lip not prominent; mandibular molar ridged, primary cutting edge of mandible toothed, palp attached level with molar; setae of inner plate of mazilla 1 mostly terminal (Valettiopsis); outer plates of maxilliped acutely produced apically; cora 2 short, broad, subrounded, scarcely concealing coxa 1, latter not Shorter than coxa 2 and of similar shape (Valettiopsis); gnathopod 1 weakly chelate, palm transverse; gnathopod 2 rather stout, article 5 shorter than 6; inner ramus of uropod 2 slightly constricted; telson short, deeply cleft. Species: 1, antarctic, abyssal.
368 U.S. NATIONAL MUSEUM BULLETIN 271
Valettiopsis Holmes Valettiopsis Holmes, 1908.
Type-species: V. dentatus Holmes, 1908 (original designation).
Apparently upper lip and epistome not prominent; mandibular molar ridged, primary cutting edge of mandible toothed, palp attached level with molar; inner edge of inner plate of maxilla 1 densely setose; coxa 2 rectangular and elongate, almost covering coxa 1 completely, latter short, subtriangular (Valettia); gnathopod 1 subchelate; telson of medium length, deeply cleft. Species: 3, Atlantic-Pacific, bathyal-abyssal.
Vijaya Walker Vijaya Walker, 1904.—Gurjanova, 1962.
Type-species: V. tenuipes Walker, 1904 (monotypy).
[Epistome and upper lip unknown]; mandibular molar presumably feeble, palp attached ?level with molar; maxilla 1 presumably lacking palp; gnathopod 1 simple, article 6 longer than 5, article 3 slightly elongate; coxa 1 presumably partly hidden, following coxae increas- ingly larger; inner ramus of uropod 2 possibly constricted; telson of medium length, cleft one third; article 2 of antenna 1 half as long as article 1 (Bathyamaryllis); anteroventral corner of coxa 4 acutely pro- duced (Amaryllis). Species: 1, Ceylon, littoral.
Waldeckia Chevreux
Ephippiphora White, 1847b (homonym, Lepidoptera). Charcotia Chevreux, 1905b (homonym, Mollusca). Waldeckia Chevreux, 1906b (new name).
Type-species: Charcotia obesa Chevreux, 1905b (monotypy).
Epistome broadly rounded and slightly produced in front of upper lip, both together prominent; mandibular molar subconical, smooth, palp attached proximal to molar; gnathopod 1 simple, article 6 longer than 5; telson long, deeply cleft; gills wnplaited but with accessory lobes (Menigrates). Species: 2, antiboreal, antarctic, littoral. See Socarnes.
Melphidippidae FI@uRE 133
Draenosts.—Accessory flagellum vestigial or multiarticulate; pe- duncle of uropod 3 greatly elongate, as long as or longer than rami of uropods 1-2, and more than twice as long as telson and peduncle of
MARINH GAMMARIDEAN AMPHIPODA 369
uropod 2, uropod 8 also greatly exceeding apices of uropods 1-2, rami of uropod 3 elongate, subequal and nearly as long as rami of uropods 1-2; gnathopods feeble; body posterodorsally carinate and toothed. See Gammaridae, Oedicerotidae.
Description.—Accessory flagellum vestigial or multiarticulate; body slender, posterodorsally carinate and toothed; rostrum poorly developed; head with lateral ocular bulges; coxae short and scarcely touching serially; mouthparts basic; gnathopods feeble, intermediate between subchelate and simple; pereopods greatly elongate; uropod 3 immensely elongate as in diagnosis above and Gammaridea Key A, couplet 7 (p. 107); telson of medium length, deeply cleft or emarginate.
ReLationsHip.—Uropod 3 usually falls off of preserved individuals of this family as it often does on individuals of Oedicerotidae and Megaluropus (Gammaridae). Some oedicerotids have uropod 3 elon- gate as in melphidippids but they lack posterodorsal pleonal teeth. Melphidippids may also be recognized by the lateral ocular bulges of the cephalon and the very short coxae. Gammaridae with elongate uropod 3 have one or more of the following exceptions to the melphi- dippid pattern: inner ramus of uropod 8 shortened and scale-like; uropod 3 not exceeding apices of uropods 1-2; peduncle of uropod 3 shorter than rami of uropods 1-2 and not twice as long as telson or urosomite 3; rami of uropod 3 much shorter than peduncle. Melphi- dippids lack sexual dimorphism in the gnathopods common to most Gammaridae.
The Vitjazianidae have a short peduncle of antenna 1.
My interpretation of Walker’s description of Hornellia is that it lacks the melphidippid condition of uropod 3, for he mentions the medium length of the uropods and that the inner ramus of uropod 3 is twice as long as the peduncle; if the peduncle were of the normal melphidippid length the total uropod would be two-thirds the length of the body, a condition so remarkable that Walker would have mentioned it. Indeed, Hornellia Walker was removed to the Gam- maridae by Pillai (1957).
The calliopiid Metaleptamphopus may belong in the Melphidippidae.
Key to the Genera of Melphidippidae
1. Accessory flagellum long, 2- or more articulate (fig. 133b) . . Melphidippa Accessory flagellum very short, l-articulate (fig. 138c)......... 2 Zomeehelson. sclekt: i(figs USI): annual Meath aloo a oh be Ge Melphidippella Melson’ entire! ‘emarginate (ig 33) ek oe a cere Melphisana
370 U.S. NATIONAL MUSEUM BULLETIN 271
“tly Pod Iit ZC ip oa)
“il
Ficure 133.—Melphidippidae: a, Melphidippa macrura Sars (1895, pl. 170). Accessory flagella: b, Melphidippa goesi Stebbing (Sars, 1895, pl. 169, as M. spinosa); c, Melphidip- pella macra (Norman) (Sars, 1895, pl. 171). Mouthparts, M. goest: d, upper lip; e, mandible; f, lower lip; g,h, maxillae 1, 2; 7, maxilliped. M. goesi: j,k, gnathopods 1, 2; 1, uropod 3; m, telson. Telson: n, Melphisana bola J. L. Barnard (1962b).
MARINE GAMMARIDEAN AMPHIPODA Si Genera of Melphidippidae
Melphidippa Boeck Melphidippa Boeck, 1871.—Stebbing, 1906.
Type-species: Melphidippa goest Stebbing, 1899a (=new name for homonym, Gammarus spinosus Goés, 1866) (type indicated by Boeck, 1876, considered as type by Sars, 1895, but selected by Gurja- nova, 1951). See Sars, 1895.
Accessory flagellum long, 2- or more articulate; mandibular palp article 3 twice as long as article 1; maxillipedal palp article 4 claw- shaped; gnathopod 2 with linear fifth and sixth articles; telson cleft. Species: 7, bipolar, littoral to bathyal (750 m).
Melphidippella Sars
Melphidippella Sars, 1895.—Stebbing, 1906.
Type-species: Atylus macer Norman, 1869a (original designation).
Accessory flagellum short, 1-articulate; mandibular palp article 3 equal to article 1; maxillipedal palp article 4 claw-shaped; gnathopod 2 with linear fifth and sixth articles; telson cleft. Species: 2, boreal N. E. Atlantic, Japan, littoral.
Melphisana J. L. Barnard
Melphisana J. L. Barnard, 1962b.
Type-species: MM. bola J. L. Barnard, 1962b (original designation).
Accessory flagellum short, 1l-articulate; mandibular palp article 3 equal to article 1; maxillipedal palp article 4 short, stout, not claw- shaped; gnathopod 2 with articles 5 and 6 intermediate between linear and oval; [uropod 3 unknown]; telson entire, emarginate. Species: 2, California, Japan, littoral.
Ochlesidae
FIGURE 134
Driacenosis.—Accessory flagellum absent; palp of marilliped absent; mouthpart group projecting subconically; mandible lacking molar, coxae 1-4 subacuminate; telson entire. Monogeneric. See Lysi- anassidae, Acanthonotozomatidae.
Description.—Body massive, processiferous; accessory flagellum absent; flagella of antennae reduced to a few articles; rostrum large; coxae 1-4 subacuminate; mouthparts projecting in a subconical
3/2 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 134.—Ochlesidae: a, Ochlesis levetzowi Schellenberg (1953). Antennae 1, 2: 5, O. innocens Stebbing (1910). Lower lip: c, O. lenticulosus K. H. Barnard (1940). 0. in- nocens (after Pirlot, 1936): d, mandible; ¢, maxilla 1; f, maxilla 2; g, maxilliped; h,2, gnathopods 1, 2. Uropod 3: 7, O. innocens Stebbing (1910). Telson: k, O. innocens (after Pirlot, 1936); J, O. lenticulosus.
MARINE GAMMARIDEAN AMPHIPODA 373
bundle, elongate; upper lip elongate; mandible lacking molar, palp long, 3-articulate, article 1 elongate; lower lip lacking inner lobes, outer lobes apically acuminate; maxilla 1 with minute, vestigial palp or none; maxilla 2 elongate; maxillipeds lacking palps, inner plates acuminate, outer lobes operculiform; gnathopods simple or mero- and carpochelate; uropod 3 biramous; telson of medium length, entire.
ReELATIONSHIP.—Of the suborder Gammaridea, only the Ochlesidae and the genus Danaella (see also Thoriella and Chevreuziella) in the Lysianassidae lack a maxillipedal palp. This lack is characteristic of all members of the suborder Hyperiidea but Ochlesidae appear in other respects to be so closely related to gammarideans that they are retained therein. The large coxae are especially characteristic of gammarideans but the general body shape resembles that of the Acanthonotozomatidae and Stilipedidae.
Genera of Ochlesidae
Ochlesis Stebbing
Ochlesis Stebbing, 1910.
Type-species: 0. innocens Stebbing, 1910 (monotypy). Species: 3, S. Africa, Indonesia, Australia, littoral.
Oedicerotidae
FIGURES 135-138
Diacnosis.—Accessory flagellum absent or vestigial; telson entire or emarginate, very short; eyes when present usually coalesced dorsally; head often massive and galeate (see glossary); anoculate members assigned to the family on basis of aspects similar to shallow water genera having coalesced eyes; pereopod 5 immensely longer than pereopod 4; rami of uropod 3 scarcely longer than the elongate peduncle. See Calliopiidae, Pleustidae, Isaeidae, Ampithoidae, Syn- oplidae, Laphystiopsidae.
Description.—Accessory flagellum absent; head often galeate and massive, conspicuous rostrum usually present but occasionally completely absent; mouthparts basic except for mandibular molar which varies from strongly triturative to nontriturative and nearly obsolescent; one genus lacking mandibular palp; gnathopods powerful to weak, highly variable; coxae of medium size; rami of uropod 3 narrowly lanceolate, scarcely longer than the elongate peduncle;
374 U.S. NATIONAL MUSEUM BULLETIN 271
telson entire or emarginate; pereopods strongly setose (fossorial), pereopod 5 immensely longer than pereopod 4.
ReELATIONSHIP.—In gross morphology, this family is scarcely dis- tinct from the Calliopiidae and Pleustidae in which the telson is
ay ry. ae
//
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a lh 7 samred
foaointhic Ni x SW) | 2. a ae ee ‘ eee >
Ficure 135.—QOedicerotidae: a, Oediceros saginatus Kroyer (Sars, 1895, pl. 102); b, Synchel- idium halpocheles (Grube) (Sars, 1895, pl. 112); c, Monoculodes carinatus (Bate) (Sars,
1895, pl. 105).
MARINE GAMMARIDEAN AMPHIPODA 375
also entire and the accessory flagellum vestigial or absent. The rel- atively longer peduncle of uropod 3 in oedicerotids is a reliable character as is the aspect of those oedicerotids having massive heads and projecting rostra; often they have pereopods adapted for bur- rowing by enlarged and spinose articles and pereopod 5 is much longer than pereopod 4. Since the eyes are lost in many species the taxono- mist requires some experience in differentiating these families.
Ficure 136.—Oedicerotidae: a, Oediceroides apicalis K. H. Barnard (J. L. Barnard, 1961); b, Aceroides latipes Sars (1895, pl. 120). Telson: c, Paroediceroides sinuata Schellenberg (1931); d, Monoculodes borealis Boeck (Sars, 1895, pl. 106); ¢, Monoculodes tesselatus Schneider (Sars, 1895, pl. 106); f, Oediceros saginatus Kr@yer (Sars, 1895, pl. 102). Uro- pod 3: g, Oediceros. Coxa 4: h, Perioculodes longimanus (Bate and Westwood) (Sars, 1895, pl. 110).
376 U.S. NATIONAL MUSEUM BULLETIN 271
The Synopiidae and Pardaliscidae usually have a well-developed accessory flagellum in contrast to its absence in oedicerotids.
The Isaeidae (=Photidae) differ from Oedicerotidae in their glandular pereopods, fleshy telson, and tube-dwelling habits but some Oedicerotidae, lacking rostra, are morphologically similar to photids and differ from them by disparate shortening of pereopods 3-4 in relation to pereopod 5.
Oedicerotids need extensive generic revision. Ocular shapes are useful for those genera having oculate type-species but deep-sea species lacking eyes often are difficult to classify. If eyes were to be ignored various genera might be combined, such as Oediceropsis and Paroediceroides (but note that the type-species of Oediceropsis has a swollen article 1 of antenna 1); Paroediceroides might be included with Oediceroides except for its acutely produced coxa 4 (a feature which is present or absent in species of other genera); Paraperioculodes might be congeneric with Oediceroides and Paroediceroides as coxa 4 is not specifically described, or with Oediceropsis as antenna 2 article 1 is not described; Oediceroides brevirostris may belong with Paraperi- oculodes (fide Ruffo, 1949); Oediceropsis proxima may belong with Paroediceroides. Numerous other examples could be cited.
Metoediceros Schellenberg (1931) lacks a mandibular palp, uropod 3 has a simple peduncle without rami, appears to have lateral, unco- alesced eyes and probably should form the type of a new family with affinities near the Dogielinotidae.
Key to the Genera of Oedicerotidae '
This key attempts to minimize the characteristics of eyes but several genera cannot remain separate without some reference to eyes, as seen in the later portion of the key.
1 (Gnathopod:2 chelate (igs l38k) ho) a. ee ee ee Gnathopod 2 subchelate or simple . . . 3
2. Mandibular molar triturative (fig. 137k), Panthened 2 ef lone tebe on article 5 guarding article 6 ..... .. . . . Pontocrates Mandibular molar degraded, not triturative, ehen beanies articulate spine(s), article 5 of gnathopod 2 not lobate (fig. 1371) . . . . . . Synchelidium
3. Primary cutting edge of mandible short, poorly projecting, untoothed Gigs lS 7Tk)iea eae 4
Primary cutting edge of sapritialle paonely oneieaans, footed Ge 1371) . 8 4. Gnathopod 2 with posterior lobe of article 5 partially or fully guarding
article 6 (figs. 1387,7,l). . . . 6 Gnathopod 2 with posterior lobe af article) 5 jornectiins at saeth sipettag ner guarding article 6 (figs) 1S8dih mm) vey na eae
1 Oedicerina Stephensen (1931) is not included; many of its features are not described because the type is a broken specimen.
11.
12.
13.
14.
15.
16.
17.
18.
WG
20.
21.
22.
23.
24.
25.
26.
MARINE GAMMARIDEAN AMPHIPODA 377
Gnathopod 1 palm oblique . . . . . . . Westwoodilla and Bathymedon Gnathopod 1 palm transverse .. . ...... . Carolobatea Mandibular molar unridged, with a fen nO. .... . . . Perioculopsis Mandibular molar large, ridged. . . 7 “Mandibular palp article 2 strongly sree” (Ge. 137h), arial 2 of prrcane: 1 as long as or longer than article]. . . . .. . . Arrhis “Mandibular palp article 2 straight’ (fig. 1371) (type species), article 2 of antenna 1 shorter than article] .... . . . Aceroides Uropod 2 reaching only to end of peduncle at Tropa 3. ER en St O Uropod 2 reaching halfway or more ane rami of uropod 3 (or rami of uropod 3 absent) ... . RANG ata MON YS oly re uiereae else MN ANZ, Gnathopods 1-2: Metiele sie ronely lobate: Loupe ar ciitibnnn Glee canmiecug. IE Gnathopods 1-2: article 5 not lobate (figs. ee BN ee Pe eas wy Seen UO) Gnathopod 2 simple (fig. 1387). . ..... Uae ina Nes ibaihwnoreiapis Gnathopod 2 subchelate ... . . . . . . Parhalimedon Eyes contiguous, appearing as one Body, BE rcapods 1 and 2 with article 7. Halicreion Eyes not contiguous, pereopods 1 and 2 lacking article 7 . . . Exoediceros
Mandible lacking palp, uropod 3 lacking rami. Metoediceros (Incertae Sedis)
Mandible bearing palp, uropod 3 bearing rami. . . . Bie tie eat) Mandibular molar lacking triturating surface, often ah preiculate spine(s)
(Giles Wa) Son ip BS eA aeons cdnaeanig Leg Mandibular molar hearts teeth ema dee: (Ge. 137k) . aS Nay ap iase's Coe agiae ho) Wowe»r lip: imner lobes coalesced (fig. 137t). .0 4. su.) Sa TS lower lips innerclobes separate; (fig-13/w)> ci. ee i a 16 Melsonventire (fie 136f). 50202 2 8. ee ws) Perioculodes Telson emarginate (fig. 186a) . ............ . . Perioculopsis Gnathopods—2 structurally salike: ys (uaa, ie gees ee a Ng, Gnathopods 1-2 structurally dissimilar. . . ... . . . . Paroediceros Gnathopods lacking posterior lobe on article5 . .. . . . Exodiceropsis Gnathopods bearing posterior lobe on article 5 . . Bb oicties ei OME Weo)
Gnathopods 1-2: lobe of article 5 long, fully guarding article! 6. Arrhinopsis Gnathopods 1-2: lobe of article 5 short, scarcely guarding article 6.
Oediceros Gnathopod 1, palm transverse ............ +. . Carolobatea Gnathopod 1, palm oblique. . . . Bue iad Ra Ma colette eer 7D) Gnathopods 1-2: article 5 not deanery lobate 2 EE al eta Matas MOEN eT Gnathopods, 1—2: article 5 grossly lobate |. . 3.20: 0-5) 2... 2 28 Gnathopod’2isubchelate: 70.26) ait. thei. 2 ee ene Hie ee aes V4 Gnathopod 2 simple (fig. 1887) ... . SR aa fpuihyporeianus Eyes unpaired, coalesced; article 2 of nereonod 5 broadly lobed postero- distally ue: . . . Methalimedon Eyes paired; article 2 oF Mereonod 5 cincuaty narrowed posterodistally. Parhalimedon Back multicarinate ... . ... . . Acanthostepheia Back not multicarinate, te aaliy, mouth Bec mionally tuberculate . . . 24 Coxa 4 produced subacutely posterodistally ........2.2... 25 Coxai 4 not produced posterodistally jn) 3 3 es 28 Eyes coalesced dorsally or absent . ....... Oe ae 26 Myvesspairedvand lateral’. i) ec od l@ediceconsic (typical) Article 1 of antenna 2 swollen .... Cregeen OM
Article 1 of antenna 2 not swollen. . Oediccreides ana Paroedicertides
378 U.S. NATIONAL MUSEUM BULLETIN 271
27. Posterior lobe on article 5 of gnathopod 2 guarding article 6 (fig. 1381).
Monoculodes
Posterior lobe on article 5 of gnathopod 2 projecting at right angles. Oediceropsis 28. Pereopods 1-2: article 7 absent. ......... ... .. .. Exoediceros Pereopods) 1-2 article: 7-presemtic: five 1 coien etek har Ed Caleta eee) 29. Hyes Pomplctels coalescediaeg an) a. SETURL ath Rea} () Kyes separated by a median line or epee or = foram PeCmmemme rs teeter adn co) lb 30. Eyes forming asemicircular ring. ......... ... . Gulbarentsia Eyes forming a circle or oval . . . . . Paraperioculodes 31. Gnathopod 2: lobe of article 5, if lene! anefectine erectly at nearly right angles, not guarding article6.. . 55 a Gnathopod 2: lobe of article 5 DrOVeCtne distally at aravlle of 45° or less, guarding article6... . SAL ae Se A ERG AAS 32. Eyes paired and lateral (@heorctical Moretti) . . . . . . Oediceropsis Byes:contiguous:at midline or absentiaiy <a 4) ai fs ye cele cee 33. Maxilla 2, outer plate lacking stout spine . . PPR aA SG Es coves Maxilla 2, outer plate bearing stout spine (often pifid) . . Anoediceros
34. Basal portion of flagellum on antenna 2 swollen. Subgenus Lopiceros (see Oediceroides) Basal portion of flagellum on antenna 2 slender. ... . OEE ao 35. Coxae 3 or 4 excavate ventrally (fig. 136b), antennae pube qual in length. Aceroides (incl. subgenus Patozdes) Coxae 3 or 4 not excavate ventrally, antenna 1 very disproportionately
shorter than antenna2 ... . . . . Oediceroides (= Oediceropsoides)
36. Coxa 4 excavate ventrally, rostrum very small or obsolete (fig. 137d), eyes Absent mn mee ... . . Aceroides
Coxa 4 not excavate “capetnalli either rostaai lerees extending more than halfway along article 1 of antenna 1, oreyes present. ....... .37
37. Article 3 of antenna 1 less than half as long as article1. . . . Monoculodes Article 3 of antenna 1 as long as article 1. .... . . . .Monoculopsis
Genera of Oedicerotidae
Acanthostepheia Boeck Acanthostepheia Boeck, 1871.—Stebbing, 1906.
Type-species: Amphithonotus malmgreni Goés, 1866 (monotypy). See Stephensen, 1938.
Cutting edge of mandible projecting and toothed, molar apparently ridged; inner lobes of lower lip separate; gnathopods similar to each other, subchelate, stout, article 5 with large posterior lobe projecting at right angles; uropod 2 reaching end of uropod 3; body dorsally multicarinate (combining character). Species: 3, arctic, littoral to bathyal (552 m).
Aceroides Sars Aceroides Sars, 1895.—Stebbing, 1906.
Aceropsis Sars, 1895. (Patoides) J. L. Barnard, 1964a (subgenus).
MARINE GAMMARIDEAN AMPHIPODA 379
Type-species: Halicreion latipes Sars, 1882 (monotypy). See Sars, 1895.
Cutting edge of mandible scarcely projecting, teeth obsolescent, molar large, ridged; inner lobes of lower lip separate; gnathopods similar, subchelate, moderately stout, article 5 with sharp posterior lobe projecting distalwards; uropod 2 reaching end of uropod 3; mandibular palp article 2 straight (Arrhis). Probably Aceroides limicola K. H. Barnard, 1925, is incorrectly assigned here; its mandible apparently has cutting teeth. Species: 4, arctic-boreal, littoral to bathyal (1,110 m) and S. Atlantic (1,280 m).
Ficure 137.—Oedicerotidae: Head: a, Oediceropsis brevicornis Liljeborg (Sars, 1895, pl. 114); 5, Monoculodes longirostris (Goés) (Sars, 1895, pl. 108); c, Oediceros saginatus Krdyer (Sars, 1895, pl. 102); d, Aceroides latipes (Sars, 1895, pl. 120); e, dorsal, Oediceros; f, Monoculodes carinatus (Bate) (Sars, 1895, pl. 105); g, Arrhis phyllonyx (M. Sars) (Sars, 1895, pl. 119); h, Westwoodilla caecula (Bate) (Sars, 1895, pl. 115, as Halimedon mulleri); 1, Westwoodilla acutifrons (Sars, 1895, pl. 116); 7, Monoculodes packardi Boeck (Sars, 1895, pl. 109). Mandible: k, Westwoodilla caecula; 1, Oediceros; m, Synchelidium haplocheles (Grube) (Sars, 1895, pl. 112, as S. brevicarpum). Upper lip: n, Oediceros. Maxilliped: 0, Oediceros. Maxillae 1, 2: p,q, Oediceros. Gnathopod 1: r, Westwoodilla caecula; s, Synchelidium. Lower lip: t, Perioculodes longimanus (Bate and Westwood) (Sars, 1895, pl. 110); u, Oediceros.
285-135 O - 69 - 25
380 U.S. NATIONAL MUSEUM BULLETIN 271
Anoediceros Pirlot
Anoediceros Pirlot, 1932.
Type-species: A. hanseni Pirlot, 1932 (original designation).
Cutting edge of mandible projecting, toothed, molar medium, ridged; inner lobes of lower lip separate; outer plate of maxilla 2 bearing a stout spine (combining character); gnathopods similar, subchelate, stout, article 5 with posterior lobe projecting at right angles; uropod 2 reaching end of uropod 3. Species: 1, Indo-Pacific, bathyal-abyssal.
l m Ficure 138.—Oedicerotidae: Gnathopod 1: a, Aceroides latipes (Sars, 1895, pl. 120); 3, Paroediceros lynceus (M. Sars) (Sars, 1895, pl. 103); c, Exoediceropsis chiltoni Schellen- berg (1931); d, Oediceros saginatus Kréyer (Sars, 1895, pl. 102); e, Monoculodes tenutro- stratus Boeck (Sars, 1895, pl. 109); f, Bathyporeiapus magellanicus Schellenberg (1931). Gnathopod 2: g, Monoculodes carinatus (Bate) (Sars, 1895, pl. 105); h, Oediceros; 1, Bathyporeiapus; j, Aceroides; k, Synchelidium haplocheles (Grube) (Sars, 1895, pl. 112);
1, Monoculodes tenuirostratus; m, Westwoodilla caecula (Bate) (Sars, 1895, pl. 115, as Halimedon mulleri).
MARINE GAMMARIDEAN AMPHIPODA 381
Arrhinopsis Stappers Arrhinopsis Stappers, 1911.
Type-species: <A. longicornis Stappers, 1911 (monotypy). See Gurjanova, 1951.
Cutting edge of mandible projecting, toothed, molar bulging, setulose, but unridged; inner lobes of lower lip separate; gnathopods similar to each other, subchelate, moderately stout, lobe of article 5 long and guarding article 6; uropod 2 reaching end of uropod 3. Species: 1, arctic, littoral.
Arrhis Stebbing
Aceros Boeck, 1861 (homonym, Aves). Arrhis Stebbing, 1906 (new name).
Type-species: Leucothoe phyllonyx M. Sars, 1858 (original designa- tion and subsequent synonymy). See G. O. Sars, 1895.
Cutting edge of mandible not projecting, untoothed, molar large, ridged; inner lobes of lower lip separate; gnathopods similar, subchelate moderately stout, article 5 with sharp posterior lobe pointing distally; uropod 2 reaching end of uropod 3; mandibular palp article 2 strongly curved (Aceroides). Species: 3, arctic, littoral to abyssal.
Bathymedon Sars Bathymedon Sars, 1895.—Stebbing, 1906
Type-species: Halimedon longimanus Boeck, 1871 (original desig- nation). See Sars, 1895.
Cutting edge of mandible not projecting, untoothed, molar large, ridged; inner lobes of lower lip separate; gnathopods somewhat similar to each other, usually gnathopod 2 more slender, posterior lobe of article 5 moderately developed in gnathopod 1, becoming obsolescent occasionally in gnathopod 2; uropod 2 reaching end of uropod 3; “article 2 of mandibular palp not as curved as in Westwood- illa.”” Species: 16, cosmopolitan coldwater, littoral to abyssal.
Bathyporeiapus Schellenberg Bathyporeiapus Schellenberg, 1931.
Type-species: B. magellanicus Schellenberg, 1931 (monotypy).
Cutting edge of mandible projecting, toothed, molar large, ridged; inner lobes of lower lip separate; gnathopods somewhat dissimilar, slender, gnathopod 1 subchelate, gnathopod 2 simple; uropod 2 apparently only slightly exceeding peduncle of uropod 3. Species: 1, antiboreal, littoral.
382 U.S. NATIONAL MUSEUM BULLETIN 271
Carolobatea Stebbing Carolobatea Stebbing, 1899d.—Stebbing, 1906.
Type-species: Halimedon schneideri Stebbing, 1888 (original desig- nation).
Cutting edge of mandible projecting slightly, teeth obsolescent, molar medium, cup-shaped, dentate; inner lobes of lower lip separated; enathopods similar to each other, subchelate, small, article 5 with small posterior lobe, palm of gnathopod 1 transverse (combining charac- ter); uropod 2 reaching end of uropod 3. Species: 1, New Zealand, littoral.
Exoediceropsis Schellenberg Exoediceropsis Schellenberg, 19381.
Type-species: . chaltoni Schellenberg, 1931 (monotypy).
Cutting edge of mandible projecting, toothed, molar small, leaf- like, produced into a spine; inner lobes of lower lip separate; gnatho- pods similar to each other, subchelate, slender, article 5 not lobed; uropod 2 reaching end of uropod 3; coxa 4 produced acutely posteriorly (combining character). Species: 1, antiboreal, S.W. Atlantic, littoral.
Exoediceros Stebbing Exoediceros Stebbing, 1899d.—Stebbing, 1906.
Type-species: Oedicerus fossor Stimpson, 1855 (original designa- tion). See Haswell, 1880c (as Oedicerus arenicola).
Cutting edge of mandible projecting, toothed, molar large, ridged; {lower lip unknown]; gnathopods similar to each other, subchelate, article 5 lobed; uropod 2 apparently not reaching far along uropod 3; eyes not contiguous (Halicreion); article 7 of pereopods 1 and 2 absent. Species: 2, S.E. Australia, littoral.
Gulbarentsia Stebbing
Barentsia Stebbing, 1894 (homonym, Bryozoa). Gulbarentsia Stebbing, 1894 (new name, footnote, p. 2).
Type-species: Barentsia hoeki Stebbing, 1894 (monotypy). See Gurjanova, 1951.
Cutting edge of mandible projecting slightly, toothed; molar medium, cup-shaped, dentate; inner lobes of lower lip separate; enathopods similar to each other, subchelate, stout, lobe of article 5 projecting strongly but not especially guarding article 6; uropod 2 reaching end of uropod 3; eyes completely fused, forming a semicircular ring (Paraperioculodes). Species: 2, arctic, littoral.
MARINE GAMMARIDEAN AMPHIPODA 383
Halicreion Boeck Halicreion Boeck, 1871.—Stebbing, 1906.
Type-species: Oediceros aequicorms Norman, 1869a (monotypy and subsequent synonymy). See Sars, 1895 (as H. longicaudatus).
Cutting edge of mandible projecting slightly, poorly but distinctly toothed, molar large, ridged; inner lobes of lower lip separate; gnatho- pods similar to each other, subchelate, stout, article 5 strongly lobate, lobe on gnathopod 2 sufficiently long to appear to be guarding article 6; uropod 3 very long, uropod 2 not exceeding peduncle of uropod 3; eyes contiguous (Hxoediceros). Species: 3, bipolar, coldwater littoral to bathyal (730 m, S. Atlantic).
Methalimedon Schellenberg Methalimedon Schellenberg, 1931.
Type-species: MM. nordenskjoldi Schellenberg, 1931 (monotypy).
Cutting edge of mandible projecting, toothed, molar large, ridged; inner lobes of lower lip separate; gnathopods similar to each other, feeble, subchelate, article 5 not lobate; uropod 2 apparently reaching end of uropod 3; eyes unpaired, coalesced, base of pereopod 5 broadly lobed distally (Parhalimedon). Species: 1, subantarctic, littoral (to 310 m).
Monoculodes Stimpson
Monoculodes Stimpson, 1853.—Stebbing, 1906. Kroyera Bate, 1857a, 1858a.
Type-species: MM. demissus Stimpson, 1853 (monotypy). Species obscure.
Antenna 1 not longer than antenna 2, article 3 less than half as long as article 1 (Monoculopsis); antenna 2 neither enlarged nor elongate (Oediceroides); cutting edge of mandible projecting, toothed, molar large, ridged; gnathopods dissimilar, subchelate, gnathopod 1 usually stout, lobe of article 5 long or short, but usually not appearing to guard article 6; gnathopod 2 usually slender, lobe of article 5 usually very long and guarding article 6; both gnathopods merging to those of Oediceroides; uropod 2 reaching end of uropod 3. Species: 46, cosmo- politan in N. Hemisphere, littoral to bathyal (only 2 species in S. Hem- isphere, Tasman Sea, 610 m, and S. Georgia area).
Monoculopsis Sars Monoculopsis Sars, 1895.—Stebbing, 1906.
Type-species: Monoculodes longicornis Boeck, 1871 (monotypy).
384 U.S. NATIONAL MUSEUM BULLETIN 271
Antenna 1 longer than antenna 2, article 3 as long as article 1 (Mono- culodes); cutting edge of mandible projecting and sharp but poorly toothed, molar large, ridged; inner lobes of lower lip separate; gnathopods dissimilar or similar, subchelate, article 5 with lobe guarding article 6 on gnathopod 2, lobe not guarding as closely on gnathopod 1; uropod 2 reaching end of uropod 3. Species: 2, bisubpolar, littoral.
Oedicerina Stephensen Oedicerina Stephensen, 1931.
Type-species: 0. ingolfi Stephensen, 1931 (monotypy). See Gur- janova, 1951.
Cutting edge of mandible slightly projecting, toothed, molar large, ridged; inner lobes of lower lip separate; [gnathopods unknown]; coxa 4 with a large blunt posterior lobe. Type specimen badly damaged. Species: 1, N. Atlantic (1,802 m).
Oediceroides Stebbing
Oediceroides Stebbing, 1888; 1906. Oediceropsoides Shoemaker, 1925. (Lopiceros) J. L. Barnard, 1961 [a valid subgenus].
Type-species: Oecdiceropsis rostrata Stebbing, 1883 (selected by Pirlot, 1932).
Cutting edge of mandible projecting, toothed, molar large, ridged; inner lobes of lower lip separate; gnathopods similar to each other, stout, subchelate, posterior lobes of article 5 well developed, pro- jecting at right angles or nearly so, or not guarding article 6 (Monocu- lodes), lobes becoming obsolescent; uropod 2 reaching end of uropod 3; inner plate of maxilla 1 with three or more setae (Paroedicerordes) ; antenna 2 usually enlarged and elongate (Monoculodes). Species: 24, cosmopolitan cold-water, littoral to abyssal. Oediceroides cystrfera and O. brevirostris Schellenberg (1931) probably incorrectly assigned here.
Oediceropsis Liljeborg
Oediceropsis Liljeborg, 1865.—Stebbing, 1906.
Type-species: 0. brevicornis Liljeborg, 1865 (monotypy). See Sars, 1895.
Eyes lateral; cutting edge of mandible projecting, toothed, molar large, ridged; inner lobes of lower lip separate; gnathopods similar to each other, subchelate, stout, lobe of article 5 projecting at right angles; uropod 2 reaching end of uropod 3. Article 1 of antenna 2 swollen in type-species. Bridged through 0. prozima Chevreux (1908c) to Parodiceroides. Species: 2, boreal N. Atlantic, bathyal.
MARINE GAMMARIDEAN AMPHIPODA 385
Oediceros Kr¢éyer Oediceros Krgyer, 1842.—Stebbing, 1906.
Type-species: O. saginatus Krgyer, 1842 (monotypy). See Sars, 1895.
Cutting edge of mandible projecting and well toothed, molar lack- ing ridges; inner lobes of lower lip separate; gnathopods similar to each other, subchelate, lobes of article 5 well developed and project- ing at right angles; uropod 2 reaching end of uropod 3. Species: 3, arctic-boreal N. Atlantic, littoral.
Paraperioculodes K. H. Barnard Paraperioculodes kK. H. Barnard, 1931; 1932.
Type-species. P. brevimanus K. H. Barnard, 1931 (original des- ignation) ; 1932.
Eyes completely fused, forming a circle or oval (Gulbarentsia, Oedi- ceroides); cutting edge of mandible apparently projecting, toothed, molar ridged; inner lobes of lower lip separate; gnathopods similar to each other, moderately stout, subchelate, posterior lobes of article 5 projecting somewhat distally; uropod 2 reaching end of uropod 3; cora 4 apparently only bluntly and poorly or not produced posteriorly (Paroediceroides). Species: 3, antarctic, littoral to bathyal (569 m).
Parhalimedon Chevreux Parhalimedon Chevreux 1906a; 1906b.
Type-species: P. turqueti Chevreux, 1906a (original designation) ; 1906b.
Eyes paired; base of pereopod 5 sharply narrowed distally (Methali- medon); cutting edge of mandible projecting, toothed, molar large, ridged; inner lobes of lower lip separate; gnathopods similar to each other, feeble, subchelate, elongate, fifth articles lacking posterior lobes; uropod 2 reaching only to end of peduncle of uropod 3. Spe- cies: 2, antarctic littoral and tropical bathyal (230 m).
Paroediceroides Schellenberg Paroediceroides Schellenberg, 1931.
Type-species: P. sinuata Schellenberg, 1931 (monotypy).
Cutting edge of mandible projecting, toothed, molar large, ridged; inner lobes of lower lip separate; gnathopods similar to each other, stout, subchelate, posterior lobe of article 5 well developed, par- tially guarding article 6; uropod 2 reaching end of uropod 3; inner plate of maxilla 1 with one seta (Oediceroides) ; telson deeply emarginate in type-species. Species: 2, subantarctic littoral and E. Pacific bathyal.
386 U.S. NATIONAL MUSEUM BULLETIN 271
Paroediceros Sars Paroediceros Sars, 1895.—Stebbing, 1906.
Type-species: Oediceros lynceus M. Sars, 1858 (original designa- tion).
Cutting edge of mandible projecting, toothed, molar unridged; inner lobes of lower lip separate; gnathopods dissimilar, subchelate, article 5 of gnathopod 1 scarcely lobed, of gnathopod 2 with a long posterior lobe guarding article 6; uropod 2 reaching end of uropod 3. Species: 5, arctic-subarctic, littoral to deep bathyal.
Perioculodes Sars Perioculodes Sars, 1895.—Stebbing, 1906.
Type-species: Monoculodes longimanus Bate and Westwood, 1868 (monotypy).
Cutting edge of mandible projecting, toothed, molar very small, conical, unridged; inner lobes of lower lip coalesced; gnathopods similar to each other, subchelate, article 5 produced into a lobe guarding article 6; uropod 2 reaching end of uropod 3; telson entire (Perioculopsis). Species: 3, Ceylon, Red Sea, boreal E. Atlantic, littoral.
Perioculopsis Schellenberg Perioculopsis Schellenberg, 1925b.
Type-species: P. lophopus Schellenberg, 1925b (monotypy).
Cutting edge of mandible projecting, not distinctly toothed, molar small, unridged; inner lobes of lower lip apparently coalesced; gnatho- pods similar to each other, subchelate, article 5 produced into a lobe guarding article 6; uropod 2 reaching end of uropod 3; telson emarginate (Perioculodes). Species: 1, W. Africa, littoral.
Pontocrates Boeck Pontocrates Boeck, 1871.—Stebbing, 1906.
Type-species: Kroyera arenaria Bate, 1858a (selected as Oedicerus norvegicus Boeck, 1861, by Boeck, 1876, a species now synonymous with K. arenaria). See Sars, 1895 (as P. norvegica).
Cutting edge of mandible projecting, toothed, molar medium, ridged; inner lobes of lower lip poorly developed but separated by an incision; gnathopods dissimilar, gnathopod 1 stout, subchelate, article 5 produced into a long posterior lobe partially guarding article 6, gnathopod 2 slender, chelate, long lobe of article 5 guarding article 6; uropod 2 reaching end of uropod 38. Species: 3, subantarctic, boreal, littoral.
MARINE GAMMARIDEAN AMPHIPODA 387
Synchelidium Sars Synchelidium Sars, 1895.—Stebbing, 1906.
Type-species: Kroyeria haplocheles Grube, 1864b (selected by Chevreux and Fage, 1925). See Sars, 1895 (as S. brevcarpum).
Cutting edge of mandible projecting, toothed, molar small, conical, with apical spine, unridged; inner lobes of lower lip flat but separate; enathopods dissimilar, gnathopod 1 subchelate, stout, article 5 form- ing a lobe guarding article 6, gnathopod 2 slender, chelate, article 5 unproduced ;* uropod 2 reaching end of uropod 3. Species: 7, boreal, littoral to bathyal.
*Possibly a long lobe of article 5 has been amalgamated with the posterior margin of article 6.
Westwoodilla Bate
Westwoodia Bate, 1857a (homonym, Hymenoptera). Westwoodilla Bate, 1862 (new name).—Stebbing, 1906. Halimedon Boeck, 1871.
Type-species: Westwoodia caecula Bate, 1857a (monotypy). See Sars, 1895 (as Halimedon mullert).
Cutting edge of mandible poorly produced, poorly toothed, molar large, ridged; ‘‘article 2 of palp more strongly curved than in Bathy- medon”; inner lobes of lower lip separate; gnathopods similar to each other, feeble, subchelate, article 5 scarcely lobed; uropod 2 reaching end of uropod 3. Species: 9, arctic, warm-temperate, littoral to abyssal.
Pagetinidae FicurE 139
Diacnosis.—Accessory flagellum absent; mandibular molar eva- nescent; coxae short, all visible; outer lobes of maxillipeds poorly developed, inner obsolete; uropod 3 uniramous; telson short, entire; urosomites 2-3 coalesced. See Stenothoidae, Thaumatelsonidae, Sebidae, Phliantidae, Cressidae, Kuriidae.
Description.—Accessory flagellum absent; urosomites 2-3 coa- lesced; coxae short, all visible; upper lip probably incised; mandible with 3-articulate palp, molar evanescent; lower lip as in Pleustidae, with tilted, oval outer lobes astride coalesced inner lobes; maxilla 1 weak, with 1-articulate palp, inner lobe possibly absent, outer weakly setose! or spinose; maxillipeds slender, outer lobes small, inner lobes vestigial; gnathopods small, subchelate; uropod 3 uni- ramous, telson short, entire.
388 U.S. NATIONAL MUSEUM BULLETIN 271
RELATIONSHIP.—The Stenothoidae, Cressidae, and Thaumatel- sonidae have enlarged coxae 2-4, more elongate telsons than do Pagetinidae and coxa 1 is small and hidden by the following coxae.
The Pardaliscidae have biramous third uropods.
The Eophliantidae and Phliantidae lack mandibular palps, the former having cylindrical bodies, the latter having depressed bodies.
The Pagetinidae bear some resemblance to Sebidae in mouthparts, uropods, telson, and coxae but Sebidae have chelate gnathcopods, elongate third artcles of gnathopod 2 (with one exception) and well-formed plates of the maxillipeds.
Key to the Genera of Pagetinidae
These genera, possibly synonymous, are based on characters which may be conditions due to different methods of preservation, mounting technique, and observation.
0
Ficure 139.—Pagetinidae: a, head and gnathopod 1 of Pagetina genarum K. H. Barnard (1932). Upper lip: b, Heterocressa monodi Nicholls (1938). Mandible: c, Heterocressa. Maxilla 1: d, Pagetina, e, Heterocressa. Maxilliped: f, Pagetina; g, Heterocressa. Pereo- pods 1,3,5: h,j,k, Pagetina. Gnathopod 2: 1, Pagetina. 1, Pagetina, dorsal view of tel- son and uropod 3. Uropod 3: m, Heterocressa. Telson: n, Heterocressa. Lower lip: o, Pagetina.
MARINE GAMMARIDEAN AMPHIPODA 389
PU ToOpod ao withi2-arviculave ramus ely). 15) ee yes aes Heterocressa Wropodisawithyl-articulate ramus 4) ote he es Pagetina
Genera of Pagetinidae
Heterocressa Nicholls Heterocressa Nicholls, 1938.
Type-species: HH. monodi Nicholls, 1938 (original designation). Species: 1, Macquarie Island, littoral.
Pagetina K. H. Barnard Pagetina K. H. Barnard, 1931; 1932.
Type-species: P. genarum K. H. Barnard, 1931 (original designa- tion) ; 1932. Species: 1, S. Georgia Island, littoral.
Paramphithoidae {including Amathillopsidae Pirlot, 1934]
Figures 140-142, rroNTISPIECE
Dracenosis.—One or more of coxae 1-4 (usually all) acutely pointed midventrally, occasionally bifid, accessory flagellum vestigial or absent, at most composed of two articles. See Acanthonotozomatidae, Lepechinellidae, Pleustidae, Calliopiidae (especially Oradarea), Eu- siridae, Haustoriidae, Astyridae.
DescriptTion.—Rostrum usually well developed; body often pro- cessiferous; accessory flagellum vestigial or absent; coxae large, 1-4 usually acuminate, or coxae slightly excavate ventrally to form weak anterior-posterior points (weak bifidation); mouthparts variable, mandible always with molar (except Bathypanoploea) and 3-articulate palp, molar varying in extent and degree of trituration surface; lower lip with or without inner lobes; plates of maxillipeds well developed but palp varying from three to four articles; gnathopods feeble, either subchelate or simple; uropod 3 with elongate lanceolate rami; telson entire or partially cleft, slightly elongate or not.
Rewationsuip.—Most Acanthonotozomatidae, also with pointed coxae, differ from Paramphithoidae by the lack of a distinct mandi- bular molar and the conically grouped mouthparts. The genus Bathy- panoploea (Acanthonotozoma australis Chilton, 1912) in the Acan- thonotozomatidae does not have conically grouped mouthparts and so
390
U.S. NATIONAL MUSEUM BULLETIN 271
=F
iepperri ey, 508) Q TU ese
ves e, ro. re
=o, aad Soper
Ficure 140.—Paramphithoidae: a, Actinacanthus tricarinatus (Stebbing, 1888); b, Amathillopsis spinigera Heller (Sars, 1885).
MARINE GAMMARIDEAN AMPHIPODA 391
Ficure 141.—Paramphithoidae: a, Epimeria cornigera (Fabricius) (Sars, 1895, pl. 128); b, Paramphithoe hystrix (Ross) (Sars 1895 pl. 130. as Acanthozone cuspidata).
392 U.S. NATIONAL MUSBUM BULLETIN 271
confounds the distinctions between the two families. I include that genus also in the key of Paramphithoidae although retaining Schellen- berg’s (1931, 1939) assignment of it to the Acanthonotozomatidae.
A few paramphithoids like Amathillopsis grevei J. L. Barnard (1961) have very weak acuminations on coxae 1—4 and might be keyed into the Calliopiidae. Clerppides of the latter family may also have sub- acuminate coxae.
The Lepechinellidae differ from Paramphithoidae mainly by the slender second articles of pereopods 3-5, the possession of coalesced inner lobes on the lower lip, and apparently the fusion of pleonites 5 and 6.
The genus Amathillopleustes Pirlot (1934) described in the Amathil- lopsidae was preoccupied by Cleonardopsis K. H. Barnard (1916). Since this genus bears rounded coxae it cannot be retained in the Paramphithoidae (=Amathillopsidae); it bears an elongate 1-artic- late accessory flagellum and cleft telson, hence should be transferred to the Eusiridae. Note that the Eusiridae and Pontogeneiidae have been amalgamated.
The Pleustidae are very closely related to Paramphithoidae but have characteristic lower lips. Mesopleustes is an intermediate genus because of its weakly pleustid lower lip and tendency to coxal acumination.
Haustoriids have a well-developed accessory flagellum and fossorial appendages.
Eusiridae lack fully acuminate coxae. Rhachotropis, however, intergrades Eusiridae and Paramphithoidae.
Oradarea Walker, originally described in the Calliopiidae but transferred to Paramphithoidae by Pirlot (1934) usually lacks the pointed coxae of paramphithoids although male O. shoemaker Pirlot has slightly acuminate coxae, Oradarea should be returned to the Calliopiidae. Parepimeriella Schellenberg (1931) also has rounded, untapering coxae, bears the lower lip characteristic of Pleustidae, and should be transferred to that family.
Eelysis K. H. Barnard (1932) and Epimeriella Walker (1907) probably should be assigned to the Astyridae and the Paramphithoidae amended to include only genera with triturative molars. Helysis is particularly close to Astyra, differing from the latter in the absence of an accessory flagellum.
One species of Gammaridae, Weyprechtia heuglini (Buchholz), has coxae 4-5 forming a ventral crescentic curve like some paramphithoids but that genus may be distinguished in its multiarticulate accessory flagellum.
MARINE GAMMARIDEAN AMPHIPODA 393 Nomenclatural Changes in Paramphithoidae
Amathillopsis Heller (1875) is removed from Gammaridae and Amathillopsidae and assigned to Paramphithoidae, thus resulting in amalgamation of Amathillopsidae with Paramphithoidae.
Amathillopleustes Pirlot (1934) is a synonym of Cleonardopsis K. H. Barnard (1916) and they are removed to the Eusiridae.
Oradarea Walker (1903) is removed to the Calliopiidae.
Parepimeriella Schellenberg (1931) is removed to the Pleustidae.
Key to the Genera of Paramphithoidae
1. Body densely covered with articulate spines (see ee . Uschakoviella
Body lacking articulate’spines . °°) 2 2455. 2.4 5. 52
2. Molar of mandible absent .... St ane iathypanaploes i Molar of mandible only a setose evnerirn or cone Ge. LADD) es cre nea eS Molar of mandible well developed, large, ridged (fig. 1427)... .....4
3. Pereopod 5 as long as or longer than pereopod4...... ... . Eclysis Pereopod 5 shorter than pereopod4 ... . . . . Epimeriella
4. Maxillipedal palp bearing only three articles 2 Gre 1429) eee aeRO di WRES 5 Maxillipedal palp bearing four articles (fig. 142m). ...........6
5. Lower lip with inner lobes (fig. 1420)... .... .. . . Parepimeria ? Lower lip lacking inner lobes (fig. 142k). . . . A . . . Metepimeria
6. Coxae 4-5 together forming a crescentic curve Below, Ge ATC) ea Malis Coxae 4—5 together not forming a crescentic curve below (fig. 1406) . . .8
7. Gnathopods subchelate, dactyl slender, simple (fig. 142b) . . . Epimeria ‘4 Gnathopods simple, dactyl stout, spinose as De a ERA is ree s
8. Accessory flagellum present .. . aes ts SUSAN NE Naa oi Attn er oe USE) Accessory flagellum absent ... . i oer sir’ 10
9. Accessory flagellum long and spiniform, aathopods Bronely subche- MACE len . . . . Amathillopsis Accessory dpe Teta hor, aeolteliee or earl like, gnathopods simple even though portions occasionally stout. . . . .. . . . Parepimeria
10. Article 5 of gnathopods shorter than article6 .. . . . . Paramphithoe Article 5 of gnathopods much longer than article6 ... . Actinacanthus
1 Assigned to Acanthonotozomatidae provisionally.
2 Chevreux (1912b).
3K. H. Barnard (1932) states maxillipedal palp article 4 is present.
4 Note resemblance in coxae 4-5 of Weyprechtia (Gammaridae).
5 See also Halirages stebbingi Schellenberg (1931) (=H. huzleyanus Stebbing, 1888, not Bate), in the Callio- piidae, a species with acuminate anterior coxae; it differs from paramphithoids and other members of Hali- rages in its stout gnathopods with ovate articles.
Genera of Paramphithoidae
Actinacanthus Stebbing
Acanthechinus Stebbing, 1888 (homonym, Echinodermata). Actinacanthus Stebbing, 1906 (new name).
394 U.S. NATIONAL MUSEUM BULLETIN 271
Type-species: Acanthozone tricarinata Stebbing, 1883 (monotypy).
Accessory flagellum absent; mandibular molar large, ridged; lower lip lacking inner lobes; maxillipedal palp 4-articulate; gnathopods poorly subchelate, articles 5 and 6 slender and very elongated, article 5 longer than 6 (Paramphithoe); telson entire. Species: 1, S. Indian Ocean (282 m).
Amathillopsis Heller
Amathillopsis Heller, 1875.—Stebbing, 1906.—Pirlot, 1934—J. L. Barnard, 1964c. Acanthopleustes Holmes, 1908.
Type-species: A. spinigera Heller, 1875 (original designation). See Sars, 1885; Gurjanova, 1951.
Accessory flagellum spine-like, 2-articulate, article 2 tiny; mandib- ular molar large, ridged; lower lip apparently with coalesced inner lobes; maxillipedal palp 4-articulate; gnathopods subchelate, articles 5 and 6 ovoid, article 5 with posterior lobe; telson slightly emarginate. Species: 7, cosmopolitan, (cold littoral) bathyal and abyssal.
Eclysis K. H. Barnard Eclysis K. H. Barnard, 1932.
Type-species: FH. sumilis K. H. Barnard, 1932 (monotypy).
Accessory flagellum absent; mandibular molar a setulose conical lamina; lower lip lacking inner lobes; maxillipedal palp 4-articulate; genathopods scarcely subchelate, article 5 with small posterior lobe; pereopod & equal to or longer than pereopod 4 (Epimeriella); telson cleft halfway. Species: 1, S. Georgia Islands, littoral.
Epimeria Costa
?Vertumnus White, 1847a (nomen nudum). Epimeria Costa, 1851b.—Stebbing, 1906.
Type-species: Gammarus corniger J. C. Fabricius, 1779 (mono- typy and subsequent synonymy). See Sars, 1895.
Accessory flagellum absent; mandibular molar large, ridged; lower lip lacking inner lobes; maxillipedal palp 4-articulate; gnathopods subchelate, dactyl slender, simple (Pseudepimeria); telson emarginate or notched; cozae 4-5 together forming a crescentic curve below (com- bining character). Species: 19, cosmopolitan cold-water, littoral to abyssal.
MARINE GAMMARIDEAN AMPHIPODA 395
Epimeriella Walker Epimeriella Walker, 1906b; 1907.
Type-species: EH. macronyx Walker, 1906b; (monotypy) 1907.
Accessory flagellum absent; mandibular molar a setose conical lamina; [lower lip unknown]; maxillipedal palp 4-articulate; gnatho- pods subchelate, short article 5 with slight posterior lobe; pereopod 5 shorter than pereopod 4 (Eelysis); telson incised. Species: 3, ant- arctic, littoral to bathyal.
Ficure 142.—Paramphithoidae: Gnathopod 1: a, Parpimeria crenulata Chevreux (1912b); b, Paramphithoe hystrix (Ross) (Sars, 1895, pl. 130, as Acanthozone cuspidata); c, Eclysis similis K, H. Barnard (1932); d, Actinacanthus tricarinatus (Stebbing, 1888). Gnathopod 2: e, Pseudepimeria grandirostris Chevreux (1912b); f, Paramphithoe; g, Actinacanthus. Uropod 3: h, Paramphithoe. Mouthparts, Paramphithoe: i, upper lip; j, mandible; k, lower lip; /jm, maxillae 1, 2; 7, maxilliped. Lower lip: 0, Parepimeria. Mandible: p, Eclysis. Maxilliped: g, Parepimeria. Telson: r, Paramphithoe; s, Epimeria loricata Sars (1895, pl. 129); ¢, Eelysis.
285-135 O - 69 - 26
396 U.S. NATIONAL MUSEUM .BULLETIN 271
Metepimeria Schellenberg Metepimeria Schellenberg, 1931.
Type-species: M. acanthurus Schellenberg, 1931 (monotypy).
Accessory flagellum absent; mandibular molar large, ridged; lower lip lacking inner lobes; maxillipedal palp 3-articulate; gnatho- pods simple; telson deeply notched. Species: 1, antiboreal, littoral.
Paramphithoe Bruzelius
Acanthosoma Ross, 1835 (homonym, Hemiptera). Paramphithoe Bruzelius, 1859.—Stebbing, 1906. Acanthozone Boeck, 1871.
Type-species: Acanthosoma hystrix Ross, 1835 (selected by Bate, 1862, p. 377!; possible synonym of Oniscus cuspidatus Lepechin, 1780, fide Gurjanova, 1951; et al).
Accessory flagellum absent; mandibular molar large, ridged; lower lip lacking inner lobes; maxillipedal palp 4-articulate; gnatho- pods subchelate, article 5 shorter than article 6 (Actinacanthus) ; telson truncate. Species: 4, arctic, littoral to bathyal.
Parepimeria Chevreux Parepimeria Chevreux 1911c; 1912a; 1912b.
Type-species: P. crenulata Chevreux, 1912a; 1912b (designated by Chevreux, 1912a).
Accessory flagellum 1-articulate; mandibular molar large, ridged; lower lip with inner lobes; maxillipedal palp 3-articulate (or 4-articu- late: K. H. Barnard, 1932); gnathopods simple, articles 5 and 6 slightly elongate; telson entire. Species: 3, antarctic, littoral to bathyal.
Pseudepimeria Chevreux Pseudepimeria Chevreux, 1912a; 1912b.
Type-species: P. grandirostris Chevreux, 1912a (original designa- tion) ; 1912b.
Accessory flagellum 1-articulate; mandibular molar large, ridged; lower lip lacking inner lobes; maxillipedal palp 4-articulate; gnatho- pods simple, dactyl stout, spinose (Epimeria); telson cleft; corae 4-5 together forming a crescentic curve below (combining character). Species: 1, antarctic, bathyal.
MARINE GAMMARIDEAN AMPHIPODA 397
Uschakoviella Gurjanova Uschakoviella Gurjanova, 1955.
Type-species: U. echinophora Gurjanova, 1955 (original designa- tion). See Shoemaker, 1964.
Body covered with articulate spines; accessory flagellum absent; lower lip lacking inner lobes; mandibular molar large, ridged; maxilli- pedal palp 4-articulate; gnathopods scarcely subchelate, but palms transverse; telson cleft slightly. Species: 1, boreal N. Pacific, littoral to abyssal.
Pardaliscidae Fiaures 1438, 144
Draenosis.—Mandible lacking molar; inner plates of maxillipeds small or vestigial; coxae all short; accessory flagellum present, well developed, often sexually dimorphic. See Stilipedidae, Acantho- notozomatidae.
Derscription.—Accessory flagellum multiarticulate, often sexually dimorphic (“lacking’”’ in one genus), base of primary flagellum often conjoint in male; rostrum usually prominent, head occasionally like that of Pseudotiron in Synopiidae; upper lip incised or rounded; man- dibles flat, almost elytriform; molar of mandible absent, palp 3- articulate or absent, article 3 often shortened; inner plates of max- illipeds evanescent, outer plates occasionally small; coxae very short; enathopods powerful or feeble; rami of uropod 3 elongate, lanceolate or subfoliaceous; telson rarely elongate but often of medium length, deeply cleft or entire; mouthparts grouped quadrately (except coni- cally in Halicella).
ReELationsHip.—The Stilipedidae differ from the Pardaliscidae in the complete absence of an accessory flagellum. Halicoides, assigned to Pardaliscidae, also lacks an accessory flagellum (but see Birstein and Vinogradov, 1960), bears a peculiar process on article 2 of antenna 1 but has other aspects of the Pardaliscidae. Both pairs of maxillae in stilipedids are highly foliaceous, unlike those of pardaliscids.
Halicella of the Pardaliscidae has its mouthparts formed into a conical bundle like the Acanthonotozomatidae but its coxae are short, and the inner plates of its maxilliped are absent. Acanthonotozomatids have long coxae and well-developed inner plates of the maxillipeds, as do Astyridae.
Laphystiopsidae have well-developed inner plates of the maxilliped, a distinct mandibular molar and lack an accessory flagellum.
398 U.S. NATIONAL MUSEUM BULLETIN 271
Ficure 143.—Pardaliscidae: a, Pardalisca cuspidata Krgyer (Sars, 1895, pl. 141); b, Nicippe tumida Bruzelius (Sars, 1895, pl. 144). Upper lip: c, Pardalisca; d, Nicippe. Accessory flagella, Pardalisca: e, female; f, male. Mandibles: g,h, right and left, Pardalisca; 1, Halice abyssit Boeck (Sars, 1895, pl. 145). Maxilla 2:7, Necochea pardella J. L. Barnard (1962d). Antenna 1, peduncle: k, Pardaliscoides tenellus Stebbing (1888). Telson: /, Parpano cebus J. L. Barnard (1964a).
MARINE GAMMARIDEAN AMPHIPODA 399
Revisionary Notes on Pardaliscidae
Halicoides Walker poses a problem because pleonites 4—5 of the type-specimen of the type-species were injured and never clarified. It was described as lacking an accessory flagellum but bearing a unique scale on article 2 of antenna 1. Presumably these are also injuries. If
Ficure 144.—Pardaliscidae: a, Pardisynopia tambiella J. L. Barnard (1961). Lower lip: b, Pardaliscella boecki (Malmgren) (Sars, 1895, pl. 143); c, Pardalisca cuspidata Krgyer (Sars, 1895, pl. 141). Pardalisca: d,e, maxillae 1-2; f, maxilliped. Maxilliped: g, Halicella parasitica Schellenberg (1926a). Gnathopod 1: h, Pardalisca; 1, Pardaliscella; 7, Nicippe tumida Bruzelius (Sars, 1895, pl. 144). Gnathopod 2: k, Pardalisca; l, Nicippe; m, Pardaliscella. Telson: n, Pardalisca.
400 U.S. NATIONAL MUSEUM BULLETIN 271
an accessory flagellum does exist and if pleonites 4—5 do bear large teeth then the genus would be referable to Halice. Meanwhile it is treated as distinct in the following key. Birstein and Vinogradov (1960, 1962, 1964) have referred their species, Halice nana and H. indica, to Halicoides; and Synopioides tertia Stephensen (1931) to Halice (see Birstein and Vinogradov, 1962) but suggest the latter also might be assigned to Halicoides. A different point of view seems more reasonable until some of these species, including Halice anacantha K. H. Barnard (1925) have been more thoroughly described. Birstein’s and Vinogradov’s (1962) diagnosis of Halice is reasonable, except that they include Halice tertia and H. anacantha which spoil the definition, because H. tertia does not have a conjoint article 1 of the primary flagellum of antenna 1, and both HZ. tertia and H. anacantha have obsolescent teeth on pleonites 4—5. The flagellum of H. anacantha is unknown except for K. H. Barnard’s statement that the complete antenna resembles that of Halice abyssi. If these two aberrant species were removed from Halice the diagnosis could be strongly tightened and should be so done as a provisional measure until further light can be shed by redescriptions and discovery of expected new abyssal pardaliscids. Undoubtedly H. tertia is congeneric with Pardisynoma tambiella J. L. Barnard (1961) and P. synopiae J. L. Barnard (1962b) but scarcely with Halicoides anomalus (type) if conjointing of the primary first antennal flagellum and obsolescence of pleonal teeth are diagnostic. Thus, Pardisynopia is retained as a genus distinct from Halice and Halicoides which have conjoint flagella. Halice anacantha K. H. Barnard is temporarily removed to Pardisynopia until the condition of antenna 1 is verified. Halicoides indica and H. nana also are provisionally assigned to Pardisynopia, even though uropod 3 of H. nana appears to lack article 2 on the outer ramus.
Key to the Genera of Pardaliscidae
Tes Melson centinen(oaVASh ee en Sale Mera A ute 2c eso rey Seiiee pease Parpano Telsondeeply cleft: (fig.;144n)). 6). a 2 2. Articles 4-5 of pereopods 1-2 inflated (fig. 144a) .........2.. 3 Articles 4-5 of pereopods 1-2 not inflated (fig. 143a) ......... 5
3. Antenna 1 lacking accessory flagellum (or bearing scale on article 2). Halicoides Antenna lbearing- accessory flagellum <) 20.) 2) 59k) eee 4
4, Antenna 1: basal part of flagellum inflated or narrow, but segmented (fig. 148¢), flagellar article 1 scarcely longer than article 3 of peduncle, urosomal
teeth vestiovall si. ayy 28h org a in Sia aaa aie a ae eect Pardisynopia Antenna 1: basal part of flagellum inflated, conjoint ! (fig. 143f), thus flagellar article 1 nearly as long as peduncle, urosomal teeth strong . . . Halice
1 Incompletely segmented.
MARINE GAMMARIDEAN AMPHIPODA A401
5. Mouthparts forming a cone-like bundle below head ... . . . Halicella Mouthparts not forming a cone-like bundle below head . Baines 6
6. Article 6 of both gnathopods 1 and 2 much longer hen ant fale 5, Erticle 5 with long narrow posterior lobe (fig. 1447) . . . . . . . Nicippe Article 5 of either gnathopods 1 or 2 subequal to or loner aha 6, but posterior lobe of article 5 broad and shallow if present . . year iNareesil 5c Sa
7. Antenna 1 with elongate peduncular article 2 (fig. 143h) . . Pardaliscoides Antenna 1 with short peduncular article 2 (fig. 148a) ......... 8
8. Mandible lacking palp, pereopods prehensile ..... ... . Parahalice Mandible with palp, pereopods not prehensile ............49
Ome Maxalla; 2 obsolescent Gig. 1439), 0. be. ee ~Necochea Maxilla 2 well developed (fig. 144e) . ... esata ater] 6)
10. Article 5 of gnathopods much longer than 6 Ga 144h iy states amrgetuaee dL Articles 5 and 6 of gnathopods subequal, or article 6 slightly longer than 5 (figs. 1442,m) . . . Aa Iga cun U7
11. Gnathopodal dactyls aorainallly leer shaped: nor eamece Gea 1443,)), maxilli- pedal palp more than 3 times as long as medial edge of outer plate.
Princaxelia
Gnathopodal dactyls short, spinose (figs. 144h,k), medial edge of outer plate on maxilliped nearly as long as palp (fig. 144f) . . .. . . . . Pardalisca
12. Upper lip grossly incised asymmetrically (fig. 143c) . . . . Pardaliscopsis Upper lip nearly symmetrically incised (fig. 143d). ......... 18
13. Antenna 1: basal part of flagellum narrow, segmented (fig. 143e), flagellar article 1 scarcely longer than article 3 of peduncle . . . . Pardaliscella Antenna 1: basal part of flagellum inflated, unsegmented (fig. 143f), flagellar articlemlmearlydasilongiasspeduncleynane tary ss et a er yaks:
A enGnathoOpod 2ssimple cry soi es wc a Wen ae eee ou en ene alice Gnathopod:2sulbchelate 7 os ioe 8 wiser ous oe eee Se Areulfia
Genera of Pardaliscidae
The upper lip and lower lip may have important generic characters but they have not been adequately described for most species. They are included when possible in the generic diagnoses to follow.
Arculfia J. L. Barnard
Arculfia J. L. Barnard, 1961.
Type-species: A. trago J. L. Barnard, 1961 (original designation).
Article 2 of antenna 1 shorter than article 1; upper lip rounded below; inner lobes of lower lip coalesced; gnathopod 1 simple, articles 5 and 6 subequal, article 5 not lobed, gnathopod 2 subchelate, dactyls simple; telson deeply cleft. Species: 1, Tasman Sea (610 m).
Halice Boeck
Halice Boeck, 1871.—Stebbing, 1906.—Birstein and Vinogradov, 1962. Synopiordes Stebbing, 1888; 1906.
402 U.S. NATIONAL MUSEUM BULLETIN 271
Type-species: H. abysst Boeck, 1871 (selected by Boeck, 1876). See Sars, 1895.
Article 2 of antenna 1 shorter than article 1, article 1 of primary flagellum as long as or longer than peduncular article 1 (Pardisynopia) ; [upper and lower lips not described]; gnathopods simple, article 6 slender, longer than 5, article 5 not lobed, dactyls simple or rarely half spinose (Hl. macronyz); urosomal teeth strong (Pardisynopia); telson cleft 40 percent or more; head usually normal in appearance, with projecting lateral lobes and narrow rostrum but occasionally formed into blunt anterior ‘‘shark nose” condition like Pseudotiron (Synopiidae). Species: 11, cosmopolitan, bathy-abyssopelagic.
Halicella Schellenberg Halicella Schellenberg, 1926a.
Type-species: H. parasitica Schellenberg, 1926a (monotypy).
Mouthparts forming a cone-like bundle below head; article 2 of antenna 1 shorter than article 1; [upper lip and lower lip not described]; enathopods simple, article 6 longer than 5, 5 not lobed, dactyls simple; telson deeply cleft. Species: 1, antarctic, littoral.
Halicoides Walker Halicoides Walker, 1896.—Stebbing, 1906.
Type-species: H.anomala Walker, 1896 (monotypy). See Chevreux and Fage, 1925.
Antenna 1 lacking accessory flagellum (but see Birstein and Vindo- gradov, 1960), article 2 shorter than article 1; [upper lip and lower lip not described]; gnathopods simple, articles 5 and 6 subequal, dactyls simple; telson deeply cleft. Species: ?1+, Bay of Biscay, littoral (possibly also tropical-bathypelagic). See ‘‘Revisionary Notes on Pardaliscidae.”’
Necochea J. L. Barnard Necochea J. L. Barnard, 1962d.
Type-species: WN. pardella J. L. Barnard, 1962d (original designation).
Article 2 of antenna 1 shorter than article 1; [upper and lower lip not described]; mazilla 2 obsolescent; gnathopods simple, article 6 shorter than 5, 5 not lobed, dactyls with a few setules; telson deeply cleft; coxa 5 with nearly twice as much surface area as other coxae. Species 1, subantarctic, abyssal.
MARINE GAMMARIDEAN AMPHIPODA 403
Nicippe Bruzelius Nicippe Bruzelius, 1859.—Stebbing, 1906.
Type-species: N. tumida Bruzelius, 1859 (monotypy). See Sars, 1895.
Article 2 of antenna 1 shorter than article 1; upper lip slightly bilobed; lower lip with coalesced inner lobes; gnathopods slightly subchelate, articles 5 and 6 stout, article 6 longer than 5, 5 with a large posterior lobe, dactyls simple; telson deeply cleft. Species: 2, bipolar, bathyal.
Parahalice Birstein and Vinogradov Parahalice Birstein and Vinogradov, 1962.
Type-species: P. mirabilis Birstein and Vinogradov, 1962 (orig- inal designation).
Article 2 of antenna 1 shorter than article 1; mandible lacking palp; [upper lip and lower lip not described]; gnathopods simple, article 6 slightly longer than 5, 5 not lobed, dactyls simple; all pereopods pre- hensile; telson cleft one third. Species: 1, central Pacific, bathyal or abyssopelagic.
Pardalisca Kr¢éyer Pardalisca Krgyer, 1842.—Stebbing, 1906.
Type-species: P. cuspidata Kroyer, 1842 (monotypy). See Sars, 1895.
Article 2 of antenna 1 shorter than article 1; upper lip very asym- metrically lobed; lower lip with inner lobes separate; gnathopods simple, article 6 much shorter than article 5, dactyls spinose; telson deeply cleft; medial edge of inner plate on mazilliped nearly as long as palp (Princazelia). Species: 7, bipolar, littoral to bathyal.
Pardaliscella Sars Pardaliscella Sars, 1895.—Stebbing, 1906.
Type-species: Pardalisca boeckii Malm, 1871 (monotypy).
Article 2 of antenna 1 shorter than article 1, base of flagellum seg- mented (combining character); upper lip weakly incised and slightly asymmetrical; lower lip with inner lobes coalesced; gnathopods simple, article 6 slightly shorter than 5, dactyl with one spine; telson cleft only halfway; [rostrum obsolete]. Species: 5, boreal, littoral to abyssal.
404 U.S. NATIONAL MUSEUM BULLETIN 271
Pardaliscoides Stebbing Pardaliscoides Stebbing, 1888; 1906.
Type-species: P. tenellus Stebbing, 1888 (monotypy). See Stebbing, 1897.
Article 2 of antenna 1 longer than article 1; [upper and lower lips not described]; gnathopods simple, article 6 shorter than 5, 5 not lobed, dactyls with tiny setules; telson deeply cleft. Species: 2, N. Atlantic, S. Pacific, bathyal to hadal.
Pardaliscopsis Chevreux Paradaliscopsis Chevreux, 1911b.
Type-species: P. tenuipalpa Chevreux, 1911b (original designa- tion).
Article 2 of antenna 1 shorter than article 1; upper lip very asym- metrically lobed; lower lip with coalesced inner lobes; gnathopods simple, articles 5 and 6 subequal, 5 not lobed, dactyls lacking spines; telson deeply cleft. Species: 1, N. Atlantic, abyssal.
Pardisynopia J. L. Barnard Pardisynopia J. L. Barnard, 1961.
Type-species: P. tambiella J. L. Barnard, 1961 (original designa- tion).
Article 2 of antenna 1 shorter than article 1; article 1 of primary flagellum half or less as long as peduncular article 1 (Halice and Hali- coides); upper lip scarcely incised below; lower lip with coalesced inner lobes; gnathopods simple, article 6 longer than 5 or subequal, 5 searcely lobed posteriorly on gnathopod 1, dactyls simple; pleonites 4-5 with teeth obsolescent (Halice); telson deeply cleft. Species: 6 (3 provisional), probably cosmopolitan, bathyal.
Parpano J. L. Barnard Parpano J. l. Barnard, 1964a.
Type-species: P. cebus J. L. Barnard, 1964a (original designation).
Article 2 of antenna 1 equal to or shorter than article 1; upper lip symmetrically lobed; lower lip apparently with coalesced inner lobes; enathopods simple, article 6 much longer than 5, latter obsolescent, dactyls simple; telson entire; uropod 3 much reduced in size. Species: 2, Caribbean, abyssal.
MARINE GAMMARIDEAN AMPHIPODA 405
Princaxelia Dahl Princaxelia Dahl, 1959.
Type-species: P. stephensent Dahl, 1959 (original designation).
Article 2 of antenna 1 shorter than article 1; upper lip scarcely incised, lobes slightly asymmetrical; [lower lip unknown]; mazillipedal palp more than three times as long as medial edge of outer plate (Par- dalisca); gnathopods simple, article 5 longer than 6, dactyls simple; telson deeply cleft. Species: 2, N. Atlantic, S. W. Pacific, bathyal to hadal.
Phliantidae, revised
[see J. L. Barnard, 1964c]
Ficures 145-147
Draqnosis.—Accessory flagellum absent; body dorsally depressed, at least anterior coxae splayed; mandible lacking palp, molar degraded; palp of maxilla 1 degraded or absent; gnathopods feeble; usually uropod 3 lacking inner ramus, often lacking both rami. See Prophli- antidae, Eophliantidae, Kuriidae, Dogielinotidae, Talitroidea, Dex- aminidae, Pagetinidae, Acanthonotozomatidae, Colomastigidae, Cressidae.
Description.—Phliantidae are depressed dorsally, except Phlias which is “‘compressed”’ (Stebbing, 1906), with pleon flexed below re- mainder of body, segments armored and coxae projecting laterally; accessory flagellum absent; mandible lacking palp, molar absent or represented by a smooth protuberance or spine; lower lip with or without inner lobes; maxilla 1 with degraded palp, 1-articulate or represented by a spine when present; maxilla 2 small, often with coalesced lobes; plates of maxilliped well developed, palp 2- to 4- articulate; gnathopods feeble, usually simple, but subchelate in genera Plioplateia and Ceina, article 3 of gnathopod 2 elongate; uropod 1 biramous (except Temnophlias), uropod 2 usually biramous (except Temnophlias and Pereionotus), uropod 3 uniramous in most genera or rami absent; telson usually entire (or cleft in Cena).
Some students, particularly Chevreux (1911a) have considered that uropod 2 is missing in Pereionotus and that the last biramous uropod in that genus is uropod 3. However, Pereionotus as stated by Stebbing (1906) probably has a small uropod 3 lacking rami, perhaps over- looked by Chevreux. Uropod 3 has not been clearly described in some genera (especially Ceina).
Rexatronsuip.—The Prophliantidae have biramous third uropods, a cleft telson, and often have a degraded flagellum on antenna 2.
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Ceina bridges the Phliantidae and Prophliantidae in its subchelate gnathopods and cleft telson but is retained in Phliantidae because of its nonbiramous third uropods and presence of a flagellum on antenna 2,
Figure 145.—Phliantidae: Heterophlias seclusus Shoemaker (1933a): a, dorsal view; b, lateral view.
The Podoceridae have a mandibular palp and molar. The Tali- troidea have well-developed triturating mandibular molars (except for Najna which otherwise has the talitroid facies).
The Kuriidae have a well-developed triturating molar and poorly developed outer plates of the maxilliped. The Dexaminidae, less Anatylinae, have well-developed triturating molars. The palp of max- illa 1 is normal in all dexaminids and segments 2-3 of the urosome are coalesced.
Ficure 146.—Phliantidae and Calliopiidae: a, Ceina carinata (Pirlot, 1936, as Periphlias carinatus). Upper lip: b, Pereionotus testudo (Montagu) (Chevreux and Fage, 1925). Lower lip: c, Heterophlias seclusus Shocmaker (1933a). Maxilla 2: d, Pereionotus; e, Heterophlias. Maxilla 1: f, Heterophlias; g, Plioplateia triquetra K. H. Barnard (1916). Mandible: h, Heterophlias. Body, dorsal: 1, Sancho platynotus Stebbing (1897) [Callio- piidae]; 7, Temnophlias capensis K. H. Barnard (1916); », T. hystrix K. H. Barnard (1954). Maxilliped: 7, Sancho; m, T. capensis. Telson and uropod 3: k, Sancho.
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The Acanthonotozomatidae have conically bundled mouthparts, normal uropod 3, and a mandibular palp.
Gnathopod 2 of some phliantids faintly resembles that of Lysi- anassidae because of the elongate article 2.
The Temnophlias-Sancho Problem
Gurjanova (1962) discussed at length the interrelationships of Sancho Stebbing, Chosroes Stebbing, and Temnophlias K. H. Barnard and their relationships to the Phliantidae. Sancho and Chosroes have been placed in or near the Calliopiidae despite the clear development of what Gurjanova calls a dorsal shield which is another way of denoting their extreme dorsoventral depression. They are even more depressed than typical phliantids. Gurjanova develops the Phliantidae from
Ficure 147.—Phliantidae: Gnathopod 1: a, Plioplateia triquetra K. H. Barnard (1916); b, Ceina carinata (Pirlot, 1936 as Periphlias carinatus); c, Heterophlias seclusus Shoemaker (1933a). Gnathopod 2:d, Ceina. Maxilliped: ¢, Heterophlias; f, Temnophlias capensis K. H. Barnard (1916); g, Pereionotus testudo (Montagu) (Chevreux and Fage, 1925). Pleopod 3: h, Iphinotus typicus (Thomson) (Stebbing, 1899a, as I. chiltonz); 1, Heterophlias; 7, Quasimodia capricornis Sheard (1936a); k, Iphiplateia whitelegger Stebbing (1899a); 1, Plioplateia. Pleopod 1: m, Heterophlias. Uropod 3: n, Heterophlias; o, Iphinotus; p, Temnophlias. Uropod 2: g, Temnophlias. ‘Telson: r, Iphinotus.
MARINE GAMMARIDEAN AMPHIPODA 409
organisms like Sancho and Chosroes, which unlike typical phliantids still bear triturative mandibular molars, mandibular palps, well- developed biramous third uropods, free urosomal segments, and, ex- cept for female Sancho, typical uropods 1 and 2. Sancho and Chosroes, however, apparently flex the abdomen under the thorax like phliantids and have somewhat splayed coxae. Temnophlias is linked to Sancho and Chosroes and Gurjanova suggests that all three should be joined together as an entity separate from the Phliantidae and Calliopiidae. Temnophlias has the flexed abdomen, the degradation of the urosome, its appendages and the simplification of mouthparts unlike Sancho and Chosroes, but it differs from the Phliantidae in the (presumably secondary) reduction of the coxae and the extension of the lateral pereonal margins as pleurae, presumably as a substitute for the reduced coxae. In this way Temnophlias resembles various isopods like Munna and Pleurogonium and amphipods like Podocerus. This modification is far more extreme in the unornamented type-species Temnophlias capensis K. H. Barnard (1916) than its so-called congener, 7. hystrix K. H. Barnard (1954) which has a thick, triquetral body with strong dorsal ornamentation and bifidly acuminate coxae.
I suggest that Sancho and Chosroes together should be allocated to a new family and Temnophlias to another new family in recognition of these strong differences from Calliopiidae, Phliantidae, and each other. Temnophlias can be segregated from Phliantidae by the munnid-like body which seems to be a secondary depression and lateral pleuroniza- tion of a cylindrioid body. In this respect Temnophlias may be more closely related to the cylindrioid Eophliantidae than to the Phhiantidae.
Key to the Genera of Phliantidae
1. Palp of maxilliped 2-articulate (fig. 147f) ...... . . . Temnophlias Palpiotmaxilliped: 3-articulate (ig N45 ecg ee mss ey 2 Palpvofmaxilliped 4-articulate:(fig. 147e) 7 2). Sse
Zee WO LOMOdss * DIFAMOUS A yee yk ee nei a el an ae eta Malanth nner ta cree Phlias Uropodesmot DiramMOuUsi is se os ae ee A Re baht ehhh wl Seema ise Oh 3
3. Lobes of maxilla 2 coalesced (fig. 146d), maxilla 1 lacking palp (fig. 146f).
Pereionotus Lobes of maxilla 2 separate (fig. 146e), maxilla 1 with spine representing palp. Palinnotus
4. Gnathopods subchelate or male gnathopod 2 chelate. ....... EGO
Gnathopodsisimiple aie space ey oem cia meee en eke alan) RU eae Oa Det Be 6
on
Maxilla 1 lacking palp; pleopod 3 apparently with rami subequal to each other and with about seven setose articles each; uropod 3 with one ramus. Ceina
Maxilla 1 with palp; pleopod 3 with vestigial, short, oval rami lacking setae; MULOPOCtolaACKIn GATAMNIe Ss Ve i ee Mine Aliig oc thane dois Weal Na lia e Plioplateia
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6. Uropod 3: peduncle distinct from ramus (fig. 1470) . ......... 7 Uropod 3: peduncle not distinct from ramus (fig. 147n) ....... =. 8 7. Pleopod 3: inner ramus well developed (fig.147h) ....... Iphinotus Pleopod 3: inner ramus vestigial (fig. 1477) . . ....... Quasimodia
8. Pleopod 3: inner ramus setose, half as long as outer. Pariphinotus and Heterophlias Pleopod 3: inner ramus a tiny nonsetose scale (fig. 147k) . . . Iphiplateia
Genera of Phliantidae
Ceina Della Valle
Ceina Della Valle, 1893.—Stebbing, 1906.—Nicholls, 1939. Periphlias Pirlot, 1936.
Type-species: Nicea egregia Chilton, 1883 (monotypy).
Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods subchelate; [pleopod 3 unknown, apparently biramous, rami subequal to each other]; uropod 3 with one ramus. Species: 2, Sulu Sea, New Zealand, Juan Fernandez Island, littoral.
Heterophlias Shoemaker Heterophlias Shoemaker, 1933a.
Type-species: H. seclusus Shoemaker, 1933a (monotypy).
Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods simple; inner ramus of pleopod 3 half as long as outer; uropod 3 lack- ing rami. Possibly not distinct from Pariphinotus. Species: 1, pan- America, tropical, littoral.
[phinotus Stebbing
Iphigenia Thomson, 1882 (homonym, Mollusca). Iphinotus Stebbing, 1899a; 1906.
Type-species: Iphigenia typica Thomson, 1882 (monotypy and subsequent synonymy).
Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods simple; rami of pleopod 3 well developed, equal; uropod 3 uniramous. Species: 1, New Zealand, littoral.
[phiplateia Stebbing Iphiplateia Stebbing, 1899a; 1906.
Type-species: I. whitelegger Stebbing, 1899a (monotypy).
Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods simple; inner ramus of pleopod 3 a tiny, nonsetose scale; uropod 3 lacking rami. Species: 1, E. Australia, littoral.
MARINE GAMMARIDEAN AMPHIPODA 411
Palinnotus Stebbing
Palinnotus Stebbing, 1900; 1906.
Type-species: Pereconotus thomsoni Stebbing, 1899a (original designation).
Maxilla 1 bearing a spine-like palp; lobes of maxilla 2 separate (Pereionotus) ; maxillipedal palp 3-articulate; gnathopods simple; rami of pleopod 3 equal; uropod 3 lacking rami. Species: 3, Japan Sea, S. Africa, E. Australia, littoral.
Pariphinotus Kunkel Pariphinotus Kunkel, 1910.
Type-species: P. tuckeri Kunkel, 1910 (monotypy).
Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods simple; inner ramus of pleopod 3 half as long as outer; “uropod 3 absent.”” Possibly synonymous with Heterophlias. Species: 1, Ber- muda, littoral.
Pereionotus Bate and Westwood
Pereionotus Bate and Westwood, 1863.—Stebbing, 1906. Icridium Grube, 1864a.
Type-species: Oniscus testudo Montagu, 1808 (monotypy). See Chevreux and Fage, 1925.
Maxilla 1 lacking palp; lobes of maxilla 2 coalesced (Palinnotus) ; maxillipedal palp 3-articulate; gnathopods simple; rami of pleopod 3 equal in length; uropod 3 apparently absent or not detected; uropod 2 with two rami in male, one ramus in female. Species: 1, tropical EK. Atlantic, Mediterranean, Red Sea, littoral.
Phlias Guerin Phlias Guerin, 1836.—Stebbing, 1906.
Type-species: P. serratus Guerin, 1836 (monotypy).
Obscure genus resembling Pereionotus. Maxilla 1 lacking palp; maxillipedal palp 3-articulate; gnathopods simple; pleopod 3 with rami equal in length; uropod 3 “‘biramous,’”’ but probably confused with uropod 2. Guerin’s dorsal figure of urosome shows a setose pair, of “uropods” probably representing a pair of pleopods. Species: 1, between Iles Malovines* and Port Jackson, Australia, ?littoral.
*Falkland Islands; thus, type-locality of this species obscure.
285-135 O - 69 - 27
412 U.S. NATIONAL MUSEUM BULLETIN 271
Plioplateia K. H. Barnard
Plioplateia K. H. Barnard, 1916.
Type-species: P. triquetra K. H. Barnard, 1916 (monotypy).
Maxilla 1 with palp; maxillipedal palp 4-articulate; gnathopods subchelate; pleopodal peduncles not expanded; both rami of pleopod 3 vestigial, short, oval, lacking setae; uropod 3 lacking rami. Species: 1, S. Africa, littoral.
Quasimodia Sheard Quasimodia Sheard, 1936a.
Type-species: @. womersleyi Sheard, 1936a (present selection).
Maxilla 1 lacking palp; maxillipedal palp 4-articulate; gnathopods simple; inner ramus of pleopod 3 vestigial; uropod 3 with a single ramus, occasionally with a vestigial second ramus. Species: 3, S. Australia, littoral.
Temnophlias K. H. Barnard
Temnophlias K. H. Barnard, 1916.
Type-species: 7. capensis K. H. Barnard, 1916 (monotypy).
Maxilla 1 lacking palp; maxillipedal palp 2-articulate; gnathopods simple or parachelate with chela subdistal; rami of pleopod 3 equal in length; ramus of uropod 3 completely fused to peduncle; body extraordinarily depressed and unornamented or triquetral and bearing dorsal segmental teeth, pereonal segments laterally discontiguous as in munnid isopods, coxae either small and simple or of medium size and acuminately bifid, pereopods parachelate or simple, chela sub- distal; antennal flagella of three or four or fewer articles; uropods 1-2 uniramous. Probably should be removed to a new family, see remarks in ‘‘Relationship.”’ Species: 2, S. Africa, littoral.
Phoxocephalidae
Figures 148-151
Diacenosis.—Accessory flagellum well developed and multiarticu- late; pereopod 5 shorter than pereopod 4 and structurally dissimilar to it; head elongate, with overhanging rostrum, usually large, flat, and hoodlike, occasionally reduced in size or dorsoventrally thickened, but always distinct; pereopods strongly spinose for burrowing; uropod 3 biramous; telson cleft. See Haustoriidae, Gammaridae.
Description.—Accessory flagellum well developed; peduncles of antennae very short, flagella very short except occasionally in males,
MARINE GAMMARIDEAN AMPHIPODA 413
antenna 2 peduncle usually heavily spinose; rostrum often hood- shaped and enveloping base of antenna 1 or narrowed and projecting between first antennae, usually flattened dorsoventrally ; body without carinae except on urosome; mouthparts basic, mandibular molar triturative or not, often spinose; palp of maxilla 1 uni- or biarticulate; lower lip with inner lobes; gnathopods subchelate, small or of medium
CHC TTT 4 hla:
Ah \: ‘ 1 J? | TF tors ; R / y a & y,\ sy TAT
SS
a
Figure 148.—Phoxocephalidae: a, Harpinia plumosa (Kréyer) (Sars, 1895, pl. 52); b, Leptophoxus falcatus Sars (1895, pl. 50); c, Mandibulophoxus uncirostratus (Giles) (Pillai, 1957).
414 U.S. NATIONAL MUSEUM BULLETIN 271
size (in one genus, Joubinella, gnathopod 1 larger than gnathopod 2); uropod 3 biramous, outer ramus often reduced in size; telson short, or of medium length, cleft.
ReEiatronsuip.— Most Haustortidae lack a conspicuous rostrum but intergradations occurring in genera such as Platyischnopus prevent absolute distinction; many haustoriids have pereopod 5 as long as or longer than pereopod 4 and of structure similar to each other, but genera such as Pontoporeia have pereopods 4 and 5 like those of the Phoxocephalidae. Those haustoriids having semblance of a rostrum never have pereopod 5 smaller and of different structure than 4 and are restricted arbitrarily from the Phoxocephalidae. Probably the families should be reorganized on the basis of pereopod 5 and by disregarding the condition of the rostrum.
Nomenclatural Changes in Phoxocephalidae
Phoxocephalus capuciatus Oliveira (1955) is synonymized with Platy- ischnopus gracilipes Schellenberg (1931).
Seven species of Harpinia and Harpinipsis are removed to Pseud- harpinia; they have a basal ensiform process on antenna 2 and distal spines or setae on the rami of uropod 2.
Several other species of Harpinia lacking a basal ensiform process on antenna 2 and presumably bearing elongate male antenna 2 are removed to Harpimopsis.
Key to the Genera of Phoxocephalidae
1. Article 2 of pereopod 3 more than twice as wide as article 3 (fig. 151c) . . .2 Article 2 of pereopod 3 about as wide as article 3 (fig. 151d). . . . . .10
2.) Palp/of maxilla 1 biarticulate (ig) 150))) 2.275 ss; PSE CN ie tani o> Palp of maxilla 1 uniarticulate (fig. 1507). . . . 7
3. Gnathopod 1 larger than gnathopod 2 and Bhi cpeecliiee Ge NO), 5 man- dibular molar triturative (fig. 150e) ... . ... . . .Joubinella Gnathopods 1 and 2 similar in size, or gnathopod 2 eee than 1, mandibular molar not triturative (fig. 150f) .... PBN pe)
4. Palp article 4 of maxilliped bearing two fia @istell aBe, nor dlr ire. (fig. 150m), rostrum minute (fig. 149c). . .. . : Pee fo ta)
Palp article 4 of maxilliped not bearing large distal seine or 7 bannine one nail, claw-like (fig. 150n), rostrum well developed (figs. 149a,d) ..... .6
5. Mandibular molar vestigial, a small spinose protuberance . Microphoxus Mandibular molar very large, minutely setulose (fig. 97d).
Urothoides (Haustoriidae)
6. Mandibular palp sickle-shaped, borne on a large process of the mandibular
body, Gigs) 150g) ae: . . . . .Mandibulophoxus
Mandibular palp not Bick leehaned: not Bosna on a large process of the
mandibular body, (fig: 150e).. Basi) Seoes uy a dae araphoxus
MARINE GAMMARIDEAN AMPHIPODA 415
7. Article 3 of maxillipedal palp produced (fig. 1501) . . .....2.2.2. 2.8 Article 3 of maxillipedal palp unproduced (fig. 150n) ......2.2. 2.9
8. Molar of mandible triturative, large (fig. 150e) . . . . . .Leptophoxoides Molar of mandible nontriturative, small (fig. 150f) . . . . .Leptophoxus
9. Molar of mandible triturative (fig. 150e). .... .. . . Phoxocephalus Molar of mandible nontriturative, smooth (or ponte eo 150f) . Metaphoxus ee i ae Hotel ecaibe tcstire nareiu be Kyes present .. : Sade Meena ica unless sti
11. Antenna 2 with bagel sracitionen y process Ge 1500) Stn Geet pe Bel 2 Antenna 2 lacking basal ensiform process (fig. 1506) . . . . . Harpiniopsis
12. Rami of uropod 2 with apical spines (fig. 151f) . . . . . . Pseudharpinia Rami of uropod 2 lacking apical spines (fig. 15le) ... . . . .Harpinia
18. Antenna 2 with basal ensiform process (fig. 150a) .. . . . Heterophoxus Antenna 2 lacking basal ensiform process (fig. 150b). . ...... =.14!
14. Dactyl of maxillipedal palp composed of long curved claw bearing short nail-like spine .. . . . . Mandibulophoxus Dactyl of maxillipedal elt ieompoced oF phone quned or straight body bearing elongate, curved, claw-like spine .... . . . .. .Proharpinia
1 And see Phozocephalus coralis K. H. Barnard (1932) to be relegated to a new genus.
Genera of Phoxocephalidae
Harpinia Boeck
Harpina Boeck, 1871 (homonym, Coleoptera). Harpinia Boeck, 1876 (new name).—Stebbing, 1906.
Type-species: Phoxus plumosus Kroyer, 1842 (selected by Boeck, 1876). See Sars, 1895.
Antenna 2 with basal ensiform process, male antenna 2 short (Harpin- topsis); mandibular molar a small, occasionally weakly spinose, nontriturative protuberance; palp of maxilla 1 biarticulate; maxilli- pedal palp article 3 not produced, article 4 short, with one or a few apical spines or setae; article 2 of pereopod 3 slender, scarcely wider than article 3; eyes absent; rami of uropod 2 lacking distal spines or setae (Pseudharpinia). Species: about 21, N. Atlantic, ? N. Pacific, cold-water, primarily deep sublittoral and bathyal.
Harpiniopsis Stephensen Harpiniopsis Stephensen, 1925a.
Type-species: H. similis Stephensen, 1925a (monotypy).
Antenna 2 lacking basal ensiform process, male antenna 2 as long as body (Harpinia, Pseudharpinia); mandibular molar a small, occasionally weakly spinose, nontriturative protuberance; palp of maxilla 1 biarticulate (often obscurely); maxillipedal palp article 3 not produced, article 4 short, with one or a few apical spines or setae;
416 U.S. NATIONAL MUSEUM BULLETIN 271
article 2 of pereopod 3 slender, scarcely wider than article 3; eyes absent. Species: about 19, cosmopolitan, bathyal and abyssal.
Ficure 149.—Phoxocephalidae: a, Paraphoxus tridentatus J. L. Barnard (1960); d, Joubinella traditor Pirlot (1932). Head, dorsal: c, Microphoxus minimus J. L. Barnard (1960); d, Paraphoxus oculatus Sars (1895, pl. 51).
MARINE GAMMARIDEAN AMPHIPODA 417
Heterophoxus Shoemaker
Heterophozus Shoemaker, 1925.
Type-species: Harpinia oculata Holmes, 1908 (monotypy and subsequent synonymy).
Mandibular molar a spinulose nontriturative protuberance; palp of maxilla 1 biarticulate; maxillipedal palp article 3 not produced, article 4 short, with a stout apical spine and setae; article 2 of pereo- pod 3 slender, scarcely wider than article 3; eyes present or absent; antenna 2 with basal ensiform process (Proharpima). Species: 4, E. Pacific, S. Africa, antarctic, littoral to deep bathyal.
Figure 150.—Phoxocephalidae: Antenna 2: a, Heterophoxus oculatus (Holmes) (J. L Barnard, 1960); b, Phoxocephalus holbolli (Krdyer) (Sars, 1895, pl. 49). Antenna 1: c, Phoxocephalus. Upper lip: d, Phoxocephalus. Mandibles and palps: e, Phoxocephalus; f, Leptophoxus falcatus Sars (1895, pl. 50); g, Mandibulophoxus uncirostratus (Giles) (J. L. Barnard, 1957a). Lower lip: h, Phoxocephalus. Maxilla 1: 1, Phoxocephalus; j, Harpinia plumosa (Kréyer) (Sars, 1895, pl. 52). Maxilla 2: k, Phoxocephalus. Maxil- lipeds: 1, Leptophoxus; m, Microphoxus minimus J. L. Barnard (1960); », Phoxocephalus. Gnathopod 1: 0, Joubinella strelkovi Gurjanova (1952); p, Phoxocephalus.
A18 U.S. NATIONAL MUSEUM BULLETIN 271
Joubinella Chevreux Joubinella Chevreux, 1908a.
Type-species: J. ciliata Chevreux, 1908a (original designation).
Mandibular molar well developed, ridged; palp of maxilla 1 bi- articulate; maxillipedal palp article 3 not produced, article 4 long, subobtuse or claw-like, setulose or spinulose; article 2 of pereopod 3 more than twice as wide as article 3; gnathopod 1 larger than gnatho- pod 2; gnathopods resembling those of Eusirus; flagellum of antenna 2 reduced to two or three articles. Species: 5, Atlantic-Pacific, epi- to bathypelagic.
Ficure 151.—Phoxocephalidae: Gnathopod 2: a, Phoxocephalus holbolli (Krgyer) (Sars, 1895, pl. 49); b, Joubinella strelkovi Gurjanova (1952). Pereopod 3: c, Phoxocephalus; d, Harpinia plumosa (Krgyer) (Sars, 1895, pl. 52). Uropod 2: e, Phoxocephalus ; f, Pseudharpinia dentata Schellenberg (J. L. Barnard, 1960). Uropod 3, Phoxocephalus: g, female; h, male. Telson: 1, Phoxocephalus; 7, Harpinia.
Leptophoxoides J. L. Barnard
Leptophoxoides J. L. Barnard, 1962d.
Type-species: L. molaris J. L. Barnard, 1962d (original designation).
Mandibular molar wel] developed, ridged; palp of maxilla 1 uniartic- ulate; maxillipedal palp article 3 apically produced, article 4 of medium length, subclaviform, with two apical] setae; article 2 of pereopod 3 broad, more than twice as wide as article 3; eyes absent. Species: 1, S. Atlantic, abyssal.
MARINE GAMMARIDEAN AMPHIPODA 419
Leptophoxus Sars Leptophoxus Sars, 1895.—Stebbing, 1906.
Type-species: Phozxus falcatus Sars, 1882 (monotypy).
Mandibular molar a small nontriturative protuberance; palp of maxilla 1 biarticulate; maxillipedal palp article 3 apically produced, article 4 long, claw-like; article 2 of pereopod 3 more than twice as wide as article 3; eyes absent. Species: 1, subarctic-boreal, littoral to abyssal.
Mandibulophoxus J. L. Barnard Mandibulophozus J. L. Barnard, 1957a; 1960.
Type-species: Phoxus uncirostratus Giles, 1890 (original designation and subsequent synonymy).
Mandibular molar a setulose, nontriturative protuberance; palp of maxilla 1 biarticulate; maxillipedal palp article 3 not produced, article 4 long, claw-like, tipped with short nail; article 2 of pereopod 3 more than twice as wide as article 3 (type) or not; eyes present or absent; mandibular palp set on extension of mandibular body (combining character). Species: 2, Indo-E. Pacific tropical, S. Africa, littoral.
Metaphoxus Bonnier Metaphoxus Bonnier, 1896.—Stebbing, 1906.
Type-species: MM. typicus Bonnier, 1896 (monotypy).
Mandibular molar a setulose, nontriturative protuberance; palp of maxilla 1 uniarticulate; maxillipedal palp article 3 not produced, article 4 long, claw-like; article 2 of pereopod 3 more than twice as wide as article 3; eyes present. Species: 4, boreal-warm temperate, littoral to deep bathyal.
Microphoxus J. L. Barnard Microphozus J. L. Barnard, 1960.
Type-species: MM. minimus J. L. Barnard, 1960 (original desig- nation).
Mandibular molar a setulose nontriturative protuberance; palp of maxilla 1 biarticulate; maxillipedal palp article 3 not produced, article 4 long, but with two large apical setae; article 2 of pereopod 3 more than twice as wide as article 3; eyes present. Species: 1, Pacific Costa Rica, littoral.
420 U.S. NATIONAL MUSEUM BULLETIN 271
Paraphoxus Sars
Paraphozus Sars, 1895.—Stebbing, 1906.—J. L. Barnard, 1960. Pontharpinia Stebbing, 1897.
Parharpinia Stebbing, 1899d.
Protophoxus K. H. Barnard, 1930.
Trichophozus K. H. Barnard, 1930.
Metharpinia Schellenberg, 1931.
Type-species: Phozus oculatus Sars, 1879 (monotypy).
Mandibular molar a setulose nontriturative protuberance; palp of maxilla 1 biarticulate; maxillipedal palp article 3 not produced, article 4 long, claw-like; article 2 of pereopod 3 more than twice as wide as article 3; eyes present. Species: 46, cosmopolitan, mainly littoral, occasionally bathyal in high latitudes.
Phoxocephalus Stebbing
Phorus Kr¢gyer, 1842 (homonym, Coleoptera). Spinifer Krgyer, 1842 (homonym, Mollusca). Phoxocephalus Stebbing, 1888 (new name); 1906.
Type-species: Phoxus holbolli Kréyer, 1842 (selected by Boeck, 1876). See Sars, 1895.
Mandibular molar well developed, ridged; palp of maxilla 1 uni- articulate; maxillipedal palp article 3 not produced, article 4 long, claw-like; article 2 of pereopod 3 more than twice as wide as article 3; eyes present or vestigial. Species: 7 (one dubious), cosmopolitan cold-water, littoral to bathyal.
Proharpinia Schellenberg Proharpinia Schellenberg, 1931.
Type-species: P. antipoda Schellenberg, 1931 (monotypy).
Mandibular molar a setulose nontriturative protuberance; palp of maxilla 1 biarticulate; maxillipedal palp article 3 not produced, article 4 of medium length or short, with a long apical spine and accessory setae, but appearing long and claw-like because of the long spine; article 2 of pereopod 3 slender, scarcely wider than article 3; eyes present; antenna 2 lacking basal ensiform process (Heterophorus). Species: 4, Galapagos, antiboreal, littoral.
Pseudharpinia Schellenberg Pseudharpinia Schellenberg, 1931.
Type-species: P. dentata Schellenberg, 1931 (monotypy). Antenna 2 urth basal ensiform process, male antenna 2 unknown in type and elongate in some species (Harpiniopsis); mandibular molar a
MARINE GAMMARIDEAN AMPHIPODA 421
small, occasionally weakly spinose, nontriturative protuberance; palp of maxilla 1 biarticulate (often obscurely); maxillipedal palp article 3 not produced, article 4 short, with one or a few apical spines or setae; article 2 of pereopod 3 slender, scarcely wider than article 3; eyes absent; rami of uropod 2 bearing distal spines or setae (Harpinia and Harpiniopsis). Species: about 8, antarctic littoral and probably cosmopolitan in bathyal-abyssal.
Pleustidae Fieures 152, 153
Driaenosis.—Accessory flagellum composed of a small scale or ab- sent; lower lip formed of two oval outer lobes obliquely astride nearly coalesced inner lobes; telson entire; upper lip incised. See Calliopiidae, Paramphithoidae, Liljeborgiidae, Isaeidae, Eusiridae, Ampithoidae, Amphilochidae, Ischyroceridae, Laphystiopsidae, Oedicerotidae.
Description.—Body with or without dorsal processes; rostrum long or inconspicuous; accessory flagellum vestigial; upper lip incised ; mandibles bearing a 3-articulate palp, molar triturative or nontritura- tive; lower lips characteristic (see diagnosis) ; remaining mouthparts basic, article 3 of palp of maxilliped occasionally produced; gnathopods subchelate, feeble or powerful; coxae of medium length or long; uropod 3 with rami much longer than peduncle, lanceolate, outer ramus usually shorter than inner; telson entire or apically sculptured, cleft nearly one fifth in one species of Austropleustes.
RELATIONSHIP.—One genus of Amphilochidae, Amphilochoides, pos- sesses a lower lip like those of pleustids but most amphilochids can be characterized by the small, partially hidden coxa 1, and some amphilochids have both coxae 1 and 2 concealed. Pseudamphilochus however has a normal coxa 1 and cleft telson and thus resembles one species of Austropleustes. See family key G for distinctions.
Only the shape of the lower lip separates this family from the Calliopiidae. Two genera, Harpinioides and Mesopleustes, have lower lips intermediate between the two families.
The Laphystiopsidae are very similar to the Pleustidae and further study is needed to clarify their systematic position.
Key to the Genera of Pleustidae
1. Mandibular molar with a triturating surface composed of ridges and teeth (Giese ISTE Pee a ee bls al Cl aia a ona ial ennui ranula ND muita ty eMeSetrn tac 2 Mandibular molar a simple, small, smooth hump (fig. 1538c) ...... 5 2. Maxillipedal palp article 3 bearing a distal process (fig. 15372).
Stenopleustes Maxillipedal palp article 3 lacking a distal process (fig. 153h). . . .. . 3
422 U.S. NATIONAL MUSEUM BULLETIN 271
3. Gnathopods simple, slender (fig. 1530) . . . .. =. =... . Parepimeriella Gnathopods subchelate, stout (figs. 153l,m). . . . Pee eae ye
4, Rostrum less than one fourth as long as article 1 of amnienne L. Pleusymtes, new genus
Rostrum nearly as long as article 1 of antenna 1... . . . Mesopleustes 5. Maxillipedal palp article 3 bearing a distal process (fig. 1532) . . . . . . 6 Maxillipedal palp article 3 lacking a distal process (fig. 153h) . . . . 7
6. Uropod 3 with a large peduncular process supporting the rami (fig. 153g). Austropleustes Uropod 3 lacking accessory peduncular process .. . . . . Neopleustes 7. Gnathopods with distally bent, slender, tapering fin articles, essentially simples (fig4i20) ane . . . . . Harpinioides !
Gnathopods subchelate, with arate c or solneotanrallar: unbent sixth articles. 8
8. Antennae short, gnathopods bearing lobate article 5 (fig. 153m) . Pleustes ? Antennae long, gnathopods usually lacking lobate article 5 (fig. 153p).
Parapleustes 2
1 See this genus in the Calliopiidae. 2 The differences between Pleustes and Parapleustes are subjective. See Barnard and Given (1960, p. 39) for discussion.
Genera of Pleustidae
Austropleustes K. H. Barnard Austropleustes K. H. Barnard, 1931; 1932.
Type-species: A. cuspidatus K. H. Barnard, 1931 (original desig- nation) ; 1932.
Mandibular molar a small smooth protuberance; article 3 of maxil- lipedal palp distally produced; gnathopods slender but subchelate; uropod 3 with a large peduncular process supporting the rami (Neo- pleustes). Species: 2, antarctic, deep littoral.
Mesopleustes Stebbing Mesopleustes Stebbing, 1899d; 1906.
Type-species: Pleustes abyssorum Stebbing, 1888 (original desig- nation).
Mandibular molar well developed, ridged; article 3 of maxillipedal palp not distally produced; gnathopods subchelate; rostrum nearly as long as article 1 of antenna 1 (Pleusymtes). Because lower lip is not typical of other pleustids this genus probably should be assigned to the Calliopiidae, noting that it also has affinities with Param- phithoidae because of the tendency of the coxae toward acumination. Species: 1, probably cosmopolitan, bathyal-abyssal.
MARINE GAMMARIDEAN AMPHIPODA 423
f} f} 4, t} t} t} t} i} | 1 i) ! \ \ \ ‘i i i
Figure 152.—Pleustidae: a, Stenopleustes malmgreni (Boeck) (Sars, 1895, pl. 125); b, Pleustes panopla (Kr¢yer) (Sars, 1895, pl. 121).
424 U.S. NATIONAL MUSEUM BULLETIN 271
Neopleustes Stebbing
Neopleustes Stebbing, 1906.
Type-species: Amphitoe pulchella Krgyer, 1846 (original designa- tion). See Sars, 1895 (as Paramphathoe).
Mandibular molar a small smooth protuberance; article 3 of maxil- lipedal palp produced distally; gnathopods subchelate; uropod 3 lacking a large peduncular process (Austropleustes). Species: 6, arctic- boreal, littoral to bathyal (1416 m).
Figure 153.—Pleustidae: Upper lip: a, Pleustes panopla (Kr@yer) (Sars, 1895, pl. 121). Mandible: b, Sympleustes glaber (Boeck) (Sars, 1895, pl. 126); c, Pleustes. Lower lip: d, Pleustes; e, Mesopleustes abyssorum (Stebbing, 1888). Maxillae 1, 2: fig, Pleustes. Maxillipeds: h, Pleustes; 1, Stenopleustes latipes (M. Sars) (Sars, 1895, pl. 127); 7, Meso- pleustes, palp. Gnathopod 1: k, Austropleustes cuspidatus K. H. Barnard (1932); J, Neopleustes pulchellus (Kr@yer) (Sars, 1895, pl. 122); m, Pleustes. Gnathopod 2: n, Pleustes; 0, Parepimeriella irregularis Schellenberg (1931); p, Neopleustes. Uropod 3: gq, Austropleustes, showing peduncular process [shaded]; 7, Pleustes. ‘Telson: 5, Neopleustes; t, Pleustes.
MARINE GAMMARIDEAN AMPHIPODA 425
Parapleustes Buchholz
Parapleustes Buchholz, 1874.—Sexton, 1909. Incisocalliope J. L. Barnard, 1959a.
Type-species: [P. glacilis Buchholz, 1874, =—lapsus] P. gracilis Buchholz, 1874 (monotypy). See Sars, 1895 (as Paramphithoe brevi- corns) .
Mandibular molar a small smooth protuberance; article 3 of maxil- lipedal palp not distally produced; gnathopods subchelate; antennae “Jong,” gnathopods usually lacking a lobe on article 5, rostrum reaching only one third along article 1 (Pleustes). Species: 14, arctic-boreal, littoral.
Parepimeriella Schellenberg Parepimeriella Schellenberg, 1931.
Type-species: P. irregularis Schellenberg, 1931 (monotypy). Mandibular molar large, ridged; article 3 of maxillipedal palp not distally produced; gnathopods simple, slender. Species: 1, Falkland Islands, littoral. Pleustes Bate
Pleustes Bate, 1858b.—Stebbing, 1906.
Type-species: Amphithoe panopla Krgyer, 1838 (monotypy and subsequent synonymy). See Sars, 1895.
Mandibular molar a small smooth protuberance; article 3 of maxillipedal palp not distally produced; gnathopods subchelate, stout ; antennae “‘short,”’ gnathopods bearing a strong posterior lobe on article 5, rostrum usually reaching apex of peduncle of antenna 1 in typical species [except Pleustes behningi (Gurjanova, 1938) which should be transferred to Parapleustes]. Species: 7, arctic-boreal, littoral.
Pleusymtes, new genus
Type-species: Amphithopsis glaber Boeck, 1861 (present selec- tion). See Sars, 1895.
Species included: All species listed by Barnard and Given (1960, p. 40) under the name “‘Sympleustes.”’
Mandibular molar large, ridged; article 3 of maxillipedal palp not produced; gnathopods subchelate; rostrum extending less than one quarter along article 1 of antenna 1 (Mesopleustes). Species: 17, arctic-boreal, littoral to abyssal.
Stenopleustes Sars, new synonymy
Stenopleustes Sars, 1895.—Stebbing, 1906. Sympleustes Stebbing, 1899d (type: Amphithoe [sic] latipes M. Sars, 1858).
426 U.S. NATIONAL MUSEUM BULLETIN 271
Type-species: Amphithopsis malmgreni Boeck, 1871 (selected by Chevreux and Fage, 1925).
Mandibular molar large, ridged; article 3 of maxillipedal palp apically produced; gnathopods subchelate. Species: 7, arctic-boreal, littoral to bathyal.
Podoceridae
Figures 154-156
Diaanosis.—Accessory flagellum variable, often absent; urosome markedly depressed, segment 1 more than twice as long as segment 2, often considerably longer; telson entire; pereopods not glandular. See Corophiidae, Cheluridae.
Description.—Accessory flagellum variable, multiarticulate, ves- tigial or absent; body variable, broadly depressed or slender and subcylindrical, often carinate and toothed, urosome strongly de- pressed, segment 1 more than twice as long as any following segment, occasionally segment 6 apparently fused with segment 5, segments 5 and 6 reduced in length; coxae usually small and serially discon- tiguous, occasionally splayed; mouthparts basic, except upper lip slightly bilobed; gnathopods of medium size or large, subchelate; uropod 1 normal; uropods 2 and 3: one or the other often missing, vestigial or abnormal; uropod 3 biramous only in Jcilius, generally with one or no rami, telson circular or oval, short, entire; pereonites 6-7 coalesced in Dulichia and Paradulichia.
Revationsuip.—The elongation of pleonite 4 distinguishes this family from the Corophiidae.
Podoceridae often have the subcylindrical bodies of Eophliantidae and Colomastigidae but the latter two families may be distinguished by their degraded mandibles lacking palps.
Runanga J. L.. Barnard (1961) is removed to the Corophiidae.
Key to the Genera of Podoceridae
ft Antennal lackime accessory, flagellum) salem eine il sein
Antenna 1 bearing accessory flagellum . . . ao
Pleon with only five distinct segments pr eredine ‘ihe telecon Ge 156a). Laetmatophilus !
to
Pleon with six distinct segments preceding the telson (fig. 156b) . . . .3 3. Pleonite 5 carrying well-formed uropods ...... . .. . .Cyrtophium Pleonite > lacking wropeds. 2) 3.25.4 es a Leipsuropus?
1 See Dryopoides (Aoridae or Ischyroceridae) bearing distinct uropod 3.
2 Analysis of Leipswropus may be defective; the loss of uropod 2 but the retention of uropod 3 seems il- logical; the so-called third uropod may indeed be the second and this genus would then be referable to Laetmatophilus.
MARINE GAMMARIDEAN AMPHIPODA 427
4. Three pairs of uropods present (third often minute) (fig. 156b), pereonites
G-(itreer ae oe: Bala ese)
Two pairs of sexeinosls miccent Ge 156), sonuontes 6- 7 conieccede arenes)
eNO ropod:3 biramous* (igs. lo6c9)) 2.) 2k so me urge Vole CRS Uropod 3 uniramous or lacking rami (figs. 156a 1) Hive Sts acme Sakae te iat)
6. Antenna 1 shorter than antenna 2 (fig. 154c) . . .. . . . . . Podocerus Antenna 1 as long as or longer than antenna 2 (figs. 154a,b) . .....7
“I
Coxae 1-4 tiny, separated from Gren other by a distance morethan their width (fig. 1546), pereopods 1—2 shorter than the gnathopods (fig. 1546). Neoxenodice‘ Coxae small, separated from each other by a distance of one half their width (fig. 155a), pereopods 1-2 longer than the gnathopods (fig. 155a).
Xenodice‘ Se Wropode2sbiramous lance (igalloGe)inye ten a) oe eee ee Danlichia Uropod 2 uniramous, very small (fig. 156c). . . .. . . . . Paradulichia
3 Inner ramus of uropod 3 fragile and often missing from specimens but its presence is indicated by a supporting process of the peduncle. 4 Uncinotarsus (Aoridae) keys to this position but gnathopod 1 is larger than 2.
Genera of Podoceridae
Cyrtophium Dana
Cyrtophium Dana, 1852b.—Stebbing, 1906.
Type-species: ©. orientale Dana, 1853 (selected by monotypy of Dana, 1853). See Bate, 1862.
Accessory flagellum absent; antennae subequal in length; pleon with six segments and three pairs of uropods; uropod 2 with two rami, uropod 3 lacking rami. Species: 2, Singapore, E. Australia. littoral.
Dulichia Kroyer Dulichia Kr¢éyer, 1845.—Stebbing, 1906. Dyopedos Bate, 1857a,b.
Type-species: D. spinosissima Krgyer, 1845 (monotypy). See Sars, 1895.
Accessory flagellum present; antenna 1 slightly longer than 2; pleon with only five segments and only two pairs of uropods; uropod 2 biramous; uropod 3 absent; pereonites 6-7 coalesced. Species: 17, arctic-boreal, littoral to abyssal.
Icilius Dana
Icilius Dana, 1849; 1852a.—Stebbing, 1906.
Type-species: J. ovalis Dana, 1852a (selected by monotypy of Dana, 1852a). See Stebbing, 1888 (as I. danae, I. ellipticus, and I. australis).
285-135 O - 69 - 28
428 U.S. NATIONAL MUSEUM BULLETIN 271
V ee) Nh ; ;
Figure 154.—Podoceridae: a, Dulichia spinosissima Kréyer (Sars, 1895, pl. 228); b, Neo- xenodice caprellinoides Schellenberg (J. L. Barnard, 1962d); c, Podocerus variegatus Leach (Chevreux and Fage, 1925).
MARINE GAMMARIDEAN AMPHIPODA 429
Accessory flagellum present; antenna 1 much shorter than 2; pleon with six segments and three pairs of uropods; uropod 2 biramous; uropod 3 “‘biramous,” inner ramus fragile, often missing but indicated by a supporting process on peduncle. Species: 4, Indonesia, Australia, littoral.
el i)
XK Ce ff eh
al Ree Hy It
Ficure 155.—Podoceridae: a, Xenodice frauenfeldti Boeck (Sars, 1895, pl. 227); b, Laetmato- philus tuberculatus Bruzelius (Sars, 1895, pl. 226). Upper lip: c, Laetmatophilus. Mandi- ble: d, Paradulichia typica Boeck (Sars, 1895, pl. 232); ¢, Laetmatophilus. Laetmatophilus: f, lower lip; g,h, maxillae 1, 2.
430 U.S. NATIONAL MUSEUM BULLETIN 271
Laetmatophilus Bruzelius
Laetmatophilus Bruzelius, 1859.—Stebbing, 1906.
Type-species: L. tuberculatus Bruzelius, 1859 (monotypy). See Sars, 1895.
Accessory flagellum absent; antennae subequal; pleon with only five segments and only two pairs of uropods; uropod 2 lacking rami, uropod 3 absent. Species: 7, cosmopolitan, littoral to bathyal.
Ficure 156.—Podoceridae: Urosome, uropods and telson, dorsal view: a, Laetmatophilus tuberculatus Bruzelius (Sars, 1895, pl. 226); b, Podocerus chelonophilus (Chevreux and de Guerne) (Chevreux and Fage, 1925); c, Paradulichia typica Boeck (Sars, 1895, pl. 232); d, Dulichia spinosissima Kr@yer (Sars, 1895, pl. 228); ¢, Icilius ovatus Dana (Stebbing, 1906). Maxilliped: f, Laetmatophilus. Gnathopods 1, 2: g,h, Laetmatophilus. Uropod 3: 1, Icilius danae Stebbing (1910); 7, Icilius punctatus Haswell (Stebbing, 1910); , Laetmatophilus.
Leipsuropus Stebbing Leipsuropus Stebbing, 1899d; 1906.
Type-species: Cyrtophium parasiticum Haswell, 1880b, 1885 (monotypy).
Accessory flagellum absent; antenna 1 shorter than 2; pleon with (6?) segments: “uropod 2 absent, uropod 3 short, foliaceous, ovate, lacking rami’’: these facts probably erroneous; probably uropod 3 1s absent, uropod 2 lacks rami, hence referring the genus to Laetmatophilus. Species: 1, E. Australia, littoral.
MARINE GAMMARIDEAN AMPHIPODA 431
Neoxenodice Schellenberg
Neoxenodice Schellenberg, 1926c.
Type-species: NN. caprellinoides Schellenberg, 1926c (monotypy).
Accessory flagellum present: antennae equal in length; pleon with sIx segments and three pairs of uropods; uropod 2 with two rami, uropod 3 lacking rami; corae tiny, separated by a distance more than their width, pereopods 1-2 shorter than the gnathopods (Xenodice). Species: 1, antiboreal, abyssal.
Paradulichia Boeck
Paradulichia Boeck, 1871.—Stebbing, 1906.
Type-species: P. typica Boeck, 1871 (monotypy). See Sars, 1895.
Accessory flagellum present; antenna 1 longer than 2; pleon with only five segments and two pairs of uropods; uropod 2 uniramous, very small, uropod 3 absent; pereonites 6-7 coalesced. Species: 3, arctic-boreal, N. Atlantic, littoral to bathyal.
Podocerus Leach
Podocerus Leach, 1814b.—Stebbing, 1906. Platophium Dana, 1852b. Dexiocerella Haswell, 1885.
Type-species: P. variegatus Leach, 1814b (monotypy). See Chevy- reux and Fage, 1925.
Accessory flagellum present; antenna 1 shorter than 2; pleon with six segments and three pairs of uropods; uropod 2 with two rami; uropod 3 lacking rami. Species: 21, cosmopolitan, littoral (rarely bathyal, 569 m in antarctic).
Xenodice Boeck A enodice Boeck, 1871.—Stebbing, 1906.
Type-species: X. frauenfeldti Boeck, 1871 (monotypy). See Sars, 1895.
Accessory flagellum present; antenna 1 slightly longer than 2; pleon with six segments and three pairs of uropods; uropod 2 with two rami; uropod 3 lacking rami; corae small, separated by a distance half their width, pereopods 1-2 longer than the gnathopods (Neoxrenodice). Species: 1, subarctic, N. Atlantic, littoral to bathyal.
432 U.S. NATIONAL MUSEUM BULLETIN 271
Prophliantidae, revised Fiegur#s 157, 158
Diaenosis.—Accessory flagellum absent; body laterally compressea; mandible lacking palp, molar smooth or evanescent; gnathopods feeble but clearly subchelate; telson deeply cleft; uropod 3 biramous; all of urosomal segments coalesced; antennae very short, not longer than head. See Phliantidae, Eophliantidae, Dexaminidae, Atylidae, Colomastigidae, Kuriidae.
Description.—Body laterally compressed, urosomites 1-3 coa- lesced; mandible lacking palp, molar smooth or evanescent; lower lip with inner lobes; palp of maxilla 1 uniarticulate; maxilla 2 normal; maxillipedal inner plates small, outer greatly elongate; gnathopods feeble, subchelate; coxae of medium size or large (coxa 5); antennae very short, not longer than head, antenna 2 in type-genus lacking flagellum, in second genus bearing flagellum; uropod 3 biramous; telson deeply cleft.
RELATIONSHIP.—The Eophliantidae have cylindrical bodies and reduced coxae and the gnathopods are simple or parachelate, not subchelate.
The Talitroidea have distinct urosomal segments and a well- developed mandibular molar.
The Phliantidae have uniramous or aramous third uropods. The genus Ceima in that family forms a bridge to the Prophliantidae.
The Kuriidae have a flagellum on antenna 2 and a uniramous third uropod.
The Atylidae have a mandibular palp.
The Dexaminidae (including Anatylidae) have only urosomites 2 and 3 fused.
The Prophliantidae are characterized especially by coalesced uro- somal segments, biramous third uropods, lack of mandibular palps, and cleft telsons.
Haustoriopsis Schellenberg, 1938, is removed from the Haustoriidae to this family. It resembles the type-genus in all respects except that it bears a 3-articulate antennal flagellum whereas Prophlias lacks a flagellum. The coalesced urosomal segments, large coxa 5, similar maxuliped with small inner plates and elongate outer plates, 1- articulate first maxillary palp, absence of mandibular palp, smooth and bulging mandibular molar, all point to the close relationship of these two genera. Ruffo (1959) suggested that Haustoriopsis be re- moved to the Dexaminidae but I believe that the Prophliantidae should be segregated from that family on the distinctive urosome. Presumably Prophliantidae have been derived from dexaminid-like ancestors.
MARINE GAMMARIDEAN AMPHIPODA 433 Nomenclatural Changes in Prophliantidae
The following genera are removed to the Eophliantidae: Biancolina Della Valle (1893), Bircenna Chilton (1884), Oeinina Stephensen (1933), Cylindryllioides Nicholls (1938), Hophliantis Sheard (1936a), and Wandelia Chevreux (1906a).
Cerna Della Valle (1893) is removed to the Phliantidae.
Kuria Walker and Scott (1903) is removed to the Kuriidae.
Figure 157.—Prophliantidae: a, Prophlias anomalus Nicholls (1939); b, Haustoriopsis reticulatus Schellenberg (1938); c, Prophlias, antenna 2.
434 U.S. NATIONAL MUSEUM BULLETIN 271
Key to the Genera of Prophliantidae
1. Antenna 2 with flagellum (fig. 157b) ..... =. =... .. . Haustoriopsis Antenna 2 lacking flagellum (fig. 157a) . . ... =.=... . .. . Prophlias
€
Figure 158.—Prophliantidae: Prophlias anomalus Nicholls (1939): a,b, gnathopods 1, 2; c, head; d, mandible; e, lower lip; f,g, maxillae 1, 2; A, maxilliped; 7, uropod 3; 7, telson. Haustoriopsis reticulatus Schellenberg (1938): k, mandible; /, maxilla 1; m, uropod 3.
Genera of Prophliantidae
Haustoriopsis Schellenberg
Haustoriopsis Schellenberg, 1938.
Type-species. H. reticulatus Schellenberg, 1938 (monotypy). Species: 1, Bismarck Archipelago, littoral.
Prophlias Nicholls
Prophlias Nicholls, 1939.
Type-species: P. anomalus Nicholls, 1939 (monotypy). Species: 1, W. Australia, littoral.
MARINE GAMMARIDEAN AMPHIPODA 435
Sebidae
Figure 159
Draenosis.—Accessory flagellum 2-articulate, elongate; molar of mandible obsolescent; plates of maxillipeds of medium size; gnatho- pods 1 and 2 chelate, gnathopod 1 larger than 2, article 3 of gnatho- pod 2 elongate; antenna 1 peduncle elongate; uropod 3 uniramous; telson entire; urosomites 2-3 coalesced. Monogeneric. See Lysi- anassidae, Leucothoidae, Anamixidae, Pagetinidae.
Description.—Accessory flagellum 2-articulate, elongate; body smooth or with poorly developed carinae, urosomites 2—3 coalesced; mandibular palp article 3 short, molar evanescent; remaining mouth- parts basic although plates of maxillipeds somewhat small; coxae of medium length, rounded-quadrate below; gnathopods chelate, rather
ty)
Ficure 159.—Sebidae: Seba armata Chevreux (1900): a, lateral view; b, upper lip; c, mandible; d, maxilla 1; ¢, maxilliped; f,g, gnathopods 1, 2; h, telson; 7, antenna 1; 7, uropod 3. Seba saundersi Stebbing (1888): k, lower lip.
436 U.S. NATIONAL MUSEUM BULLETIN 271
slender, first larger than second, and occasionally weakly chelate, article 3 of gnathopod 2 elongate; uropod 3 with its single ramus longer than the short peduncle; telson entire.
RELATIONSHIP.—The Leucothoidae have biramous third uropods. The Lysianassidae have a short peduncle of antenna 1 with article 2 much shorter than 1 whereas the Sebidae not only have an elongate peduncle but article 2 is much longer than 1. Few Lysianassidae have a uniramous uropod 3 and when so it is never elongate.
The Stenothoidae have subchelate gnathopods and coxa 1 is con- cealed by coxa 2, but mouthparts and uropod 3 indicate a close rela- tionship of the two groups.
Platyischnopus in the Haustoriidae resembles the Sebidae in its subchelate gnathopods but differs from sebids in its biramous third uropods, much larger outer plates of the maxilliped and cleft telson
Genera of Sebidae
Seba Bate
Seba Bate, 1862.—Stebbing, 1906.—K. H. Barnard, 1957. Teraticum Chilton, 1884.
Grimaldia Chevreux, 1889.
Paravalettia K. H. Barnard, 1916.
Type-species: S. innominata Bate, 1862 (monotypy). Species: 7, probably cosmopolitan cold-water, littoral to bathyal.
Stegocephalidae
Ficures 160, 161
DraGgnosis.—Mouthparts projecting in a conical bundle below head; mandible lacking palp and molar; accessory flagellum 1- or 2-articulate; gnathopods feeble, simple or weakly subchelate. See Amphilochidae, Lysianassidae.
DescripTion.—Peduncle of antenna 1 short; mouthparts variable; upper lip bilobed, mandible Jacking palp and molar, lower lip without inner lobes; palp of maxilla 1 large or small, with one or two articles, maxilla 2 usually with two plates, outer plate occasionally absent, often set on extended base so as to appear geniculate; maxillipedal palp with three or four articles; gnathopods feeble, simple or scarcely subchelate; coxae quadrate or rounded below, or first three often acuminate below, first four usually forming a continuous shield, coxa 1 never hidden by coxa 2; uropod 3 variable, peduncle long or short, rami long or short; telson short, or medium, entire or cleft.
MARINE GAMMARIDEAN AMPHIPODA 437
Ficure 160.—Stegocephalidae: a, Stegocephalus inflatus Krdyer (Sars, 1895, pl. 69). Head: b, Stegocephalus. Upper lip: c, Stegocephalus; d, Stegocephalina ingolf' Stephensen (1925a). Mandible: ¢, Stegocephalus; f, Andaniella pectinata Sars (1895, pl. 72); g, pair, Andaniexis abyssi (Boeck) (Sars, 1895, pl. 71). Lower lip: h, Stegocephalus. Maxilla 1: 1, Andaniexis; 7, Stegocephalus;, k, Phippsia gibbosa (Sars, 1895, pl. 71); J, Stegocephalina.
438 U.S. NATIONAL MUSEUM BULLETIN 271
RELATIONSHIP.—The loss of both mandibular palp and molar is approached only in the Phliantidae, Eophliantidae, and Pro- phliantidae but in those families the accessory flagellum is absent. The characteristic coxae and globular shape of stegocephalids plus the conical grouping of the mouthparts are unmistakable characters.
Ficure 161.—Stegocephalidae: Maxilla 2: a, Phippsia gibbosa (Sars, 1895, pl. 71); 8, Stegocephalina ingolfi Stephensen (1925a); c, Stegocephalus inflatus Krdyer (Sars, 1895, pl. 69); d, Andaniexis abyssit (Boeck) (Sars, 1895, pl. 71). Maxilliped: e, Phippsia; f, Andaniotes simplex K. H. Barnard (1930); g, Andaniella pectinata Sars (1895, pl. 72); h, Stegocephalus. Telson: 1, Stegocephalus; 7, Andaniopsis nordlandica (Boeck) (Sars, 1895, pl. 72). Pereopod 4:k, Andaniopsis; l, Stegocephalus. Pereopod 5: m, Andaniopsis; n, Stegocephalus; 0, Tetradeion crassum (Chilton, 1924). Pereopod 5: p, Stegocephalus. Uropod 3: q, Stegocephalus; r, Andaniella. Pereopod 2: 5, Parandaniexis mirabilis Schel- lenberg (1929b). Gnathopod 1: t, Stegocephalus. Gnathopod 2: u, Stegocephalus.
MARINE GAMMARIDEAN AMPHIPODA 439
Some Stenothoidae lack a mandibular palp and all have a shield- like coxa 4 but the first coxa is always very small and hidden by the second coxa and the accessory flagellum is a vestigial scale or absent.
The Amphilochidae have a small coxa 1 partially hidden by follow- ing coxae and the mouthparts project in a quadrate bundle.
The general aspect, head, coxae, body shape, mouthpart bundle, telson, uropods, pereopods, and gnathopods of Stegocephalidae suggest affinities with the Acanthonotozomatidae but the absence of mandib- ular palps and molars and the elytriform mandibular body distin- guish the Stegocephalidae.
Stegocephalidae have considerable resemblance to Lysianassidae. Some stegocephalid species have an elongate article 3 on gnathopod 2 which resembles that of those Lysianassidae having gnathopod 2 simple (not subchelate). However, most lysianassids have a mitten- shaped or minutely chelate article 6. All Stegocephalidae lack mandib- ular palps and molars, have some foliaceous portions on the maxillae, have a 1- or 2-articulate accessory flagellum and a highly characteristic configuration of coxae 1-4. This combination of characters never occurs in Lysianassidae.
Andaniotes ingens Chevreux (1906c) probably should form the type of a new genus because of its reduced first maxillary palp and narrow article 2 of pereopod 4 in combination but in other special characters it fits Huandania more than Andaniotes and so is transferred to the former. Andaniotes simplex KK. H. Barnard (1930) also should be distinguished as a new genus by its 3-, not 4-articulate maxillipedal palp, having articles 1 and 2 coalesced.
Key to the Genera of Stegocephalidae
1. Mandible bearing multitoothed (often minutely) incisor (fig. 160f) .. . 2 Mandible bearing smooth incisor (fig. mie Fe NUE aes Ad UNIO aR TOMI IH inaete td Eve 2eeehelson cleht, (fe slGlle)ac Ausra es ye RAE SpE NECROSS Nene eae hey cers Fine Melsonventuiren(fies Gl). nnn oe ete ek cere eee nae Swen cao aay Be Waxilla 2 lacking outer plater. =.) 8) eR. h i) see fie ipuchivstecorenhalie Maxilla 2 bearing outer plate... . Sita Wy oy ee isla ing 4. Palp article 2 of maxilliped produced sNsonnastiiliy (i 161e) . . Phippsia Palp article 2 of maxilliped not produced . He SR Ae chet 3) 5. Palp of maxilla 1 biarticulate (figs. 1607,h) 6 Palp of maxilla 1 uniarticulate (fig. 1607) 8 6. Article 2 of pereopod 4 broad (fig. 1611) ies i Article 2 of percopod 4 slender (fig. 1614) ..... . Gre rocentalopia 7. Outer plate of maxilla 2 gaping and geniculate (figs. 161a,b,c) . . Phippsiella Outer plate of maxilla 2 normal (fig. 161d)... . . . Pseudandaniexis ! 8. Article 2 of pereopod 4 slender (fig. 161k) PPM e Oy die He ROU aE one Auuicle Ziolpercopod4ubroad (ie Ol) sys 2. ee ene eee lO
1 Telson unknown.
440
es
10.
11.
12.
13.
14.
15.
16.
17.
U.S. NATIONAL MUSEUM BULLETIN 271
Outer plate of maxilla 2 gaping and geniculate (figs. 161a,b,c).
Stegocephaloides Outer plate of maxilla 2 normal (fig. 161d) ...... .. .. . Steleuthera Upper and lower lips and maxilla 1 not elongate (figs. 160c,h), spines of outer plate on maxilla 2 with hooks (fig. 16lc) . .... . .. . Stegocephalus Upper and lower lips and maxilla 1 elongate (fig. 160d), spines of outer plate on maxilla 2 lacking hooks (fig. 16ld) ..... .. . . Stegocephalina Pereopod! 5 wath)seven articles Gigs: U6limin)i)2) 2 ee le Pereopod 5 with three articles (fig. 16lo) .. . : . . . Tetradeion Inner plate of maxilliped reaching end of palp ariacle 1 Ges, 161h). Andaniopsis Inner plate of maxilliped scarcely reaching base of palp article 1 (figs. NGG) ae er Moegie, us} Palp of maxilla 1 cmniaratioilate, annals 2 of mercopod 4 dence Ge, 161k). Andaniella
Palp of maxilla 1 biarticulate, article 2 of pereopod 4 broad (fig. 1611). Pseudandaniexis ! Te ee Telson entire or apically slit ... . 5 co eaten Flagellar article 1 of antenna 1 eabecmal to or ahora? than sacle: pedun- cular article 5 of antenna 2 subequal to or shorter than article 4; pleonite 6 longer than peduncle of uropod 3. . . . . Andaniotes 2 Mreientanea) Flagellar article 1 of antenna 1 much longer than peduncle; peduncular article 5 of antenna 2 longer than article 4; pleonite 6 shorter than peduncle
of uropod3 . . Ba bait ane area Pereopod 2 deumetly pabenclare Ge 161s) ... .. . . Parandaniexis Pereopod 2simple .... MA AU tenn near MEAGRE rs rcs LG. Palp of maxilla 1 Particulate’: SNE TMG aint Ce AN ane XS Palp of maxilla 1 uniarticulate ........... =... Parandania
Genera of Stegocephalidae
Andaniella Sars
Andaniella Sars, 1895.—Stebbing, 1906.
Type-species: Andania pectinata Sars, 1882 (monotypy).
Mandibular incisor toothed; palp of maxilla 1 uniarticulate; outer plate of maxilla 2 not geniculate nor gaping; inner plate of maxiliped scarcely reaching base of palp article 1 (Andaniopsis), article 2 of palp not produced; article 2 of pereopods 3-4 slender, of pereopod 5 broad; telson entire. Species: 1, boreal N. Atlantic, littoral to bathyal.
Andaniexis Stebbing
Andania Boeck, 1871 (homonym, Lepidoptera). Andaniexis Stebbing, 1906 (new name).
Type-species: Amdania abyssi Boeck, 1871 (selected by Boeck, 1876). See Sars, 1895.
MARINE GAMMARIDEAN AMPHIPODA 44]
Mandibular incisor not toothed; palp of maxilla 1 biarticulate; outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of maxilliped not produced; article 2 of pereopod 3 slender, of pereopods 4-5 broad; telson entire. Species: 6, cosmopolitan, bathy-abysso- pelagic.
Andaniopsis Sars Andantopsis Sars, 1895.—Stebbing, 1906.
Type-species: Andania nordlandica Boeck, 1871 (monotypy). See Sars, 1895.
Mandibular incisor toothed; palp of maxilla 1 uniarticluate; outer plate of maxilla 2 not geniculate nor gaping; inner plate of mazxilliped reaching distal end of palp article 1 (Andaniella), palp article 2 not produced; article 2 of pereopods 3-4 slender, of article 5 broad; telson entire. Species: 1, boreal N. Atlantic, littoral.
Andaniotes Stebbing. new synonymy
Andaniotes Stebbing, 1897; 1906. Metandania Stephensen, 1925a.
Type-species: Anonyx corpulentus Thomson, 1882 (monotypy). See Stebbing, 1888 (as Andania abyssorum).
Mandibular incisor not toothed; palp of maxilla 1 uniarticulate; outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of maxilliped not produced; article 2 of pereopod 3 slender, of pereopods 4—5 broad; telson cleft one third; flagellar article 1 of antenna 1 equal to or shorter than peduncle; peduncular article 5 of antenna 2 equal to or shorter than article 4; pleonite 6 longer than peduncle of uropod 3 (Hu- andania). Species: 4, antarctic, N. Atlantic, littoral to bathyal.
Bathystegocephalus Schellenberg
Bathystegocephalus Schellenberg, 1926b.
Type-species: Stegocephalus globosus Walker, 1909 (monotypy).
Mandibular incisor toothed; palp of maxilla 1 uniarticulate; outer plate of maxilla 2 absent; palp article 2 of maxilliped not produced; article 2 of pereopods 3-4 slender, of pereopod 5 broad; telson cleft one third. Species: 1, Indo-Pacific, S. Atlantic, bathypelagic.
Euandania Stebbing Euandania Stebbing, 1899d; 1906.
Type-species: Andania gigantea Stebbing, 1888 (original designa- tion).
442 U.S. NATIONAL MUSEUM BULLETIN 271
Mandibular incisor not toothed; palp of maxilla 1 uniarticulate; outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of maxilliped not produced; article 2 of pereopod 3 slender, of pereopod 4 either broad [or slender H. ingens (Chevreux)], of pereopod 5 broad; telson cleft one third; flagellar article 1 of antenna 1 much longer than peduncle; peduncular article 5 of antenna 2 longer than article 4; pleo- mite 6 shorter than peduncle of uropod 8 (Andaniotes). Species: 2, cosmopolitan, bathypelagic.
Parandania Stebbing Parandania Stebbing, 1899d; 1906.
Type-species: Andania boecki Stebbing, 1888 (original designa- tion).
Mandibular incisor not toothed; palp of maxilla 1 uniarticulate; outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of maxilliped not produced; article 2 of pereopod 3 slender, of pereopods 4—5 broad; telson entire. Species: 1, cosmopolitan, bathypelagic.
Parandaniexis Schellenberg Parandaniexis Schellenberg, 1929b. —
Type-species: P. mirabilis Schellenberg, 1929b (monotypy). Mandibular incisor not toothed; palp of maxilla 1 biarticulate; outer plate of maxilla 2 not geniculate nor gaping; palp article 2 of maxiliped not produced; article 2 of pereopods 3-4 slender, of pereopod 5 broad; telson entire; pereopod 2 subchelate. Species: 1, tropical E. Pacific, abyssal. Phippsia Stebbing
Aspidopleurus Sars, 1895 (homonym, Pisces). Phippsia Stebbing, 1906 (new name).
Type-species: Stegocephalus gibbosus Sars, 1882 (monotypy). Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped produced distomedially; article 2 of pereopods 3-4 slender, of pereopod 5 broad; telson cleft one third. Species: 2, arctic-boreal, bathy- pelagic. Phippsiella Schellenberg
Phippsiella Schellenberg, 1925a.
Type-species: Stegocephalus similis Sars, 1895 (monotypy). Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped not
MARINE GAMMARIDEAN AMPHIPODA 443
produced; article 2 of pereopod 3 slender, of pereopods 4-5 broad; telson cleft more than halfway. Species: 7, probably cosmopolitan coldwater, littoral to abyssal.
Pseudandaniexis Nicholls
Parandaniexis Nicholls, 1938 (homonym, Amphipoda). Pseudandaniexis Nicholls, 1938, Corrigenda (new name).
Type-species: Parandaniexis mixtus Nicholls, 1938 (original desig- nation).
Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer plate of maxilla 1 not geniculate nor gaping; inner plate of maxilliped reaching proximal end of palp article 1 (Andaniopsis), palp article 2 of maxilliped unproduced; article 2 of pereopod 3 slender, of pereopod 5 broad; [telson broken and not clearly analyzed]. Species: 1, antarctic, bathyal.
Stegocephalina Stephensen
Stegocephalina Stephensen, 1925a.
Type-species: SS. ingolfi Stephensen, 1925a (monotypy).
Mandibular incisor toothed; palp of maxilla 1 uniarticulate; outer plate of maxilla 2 geniculate and gaping; palp article 2 of maxilliped not produced; article 2 of pereopod 3 slender, of pereopods 4-5 broad; telson cleft halfway; upper lip, lower lip, and mazilla 1 elongate; outer plate of marilla 2 lacking hooks (Stegocephalus). Species: 1, boreal N. Atlantic, bathyal.
Stegocephaloides Sars Stegocephaloides Sars, 1895.—Stebbing, 1906.
Type-species: Stegocephalus christianiensis Boeck, 1871 (original designation).
Mandibular incisor toothed; palp of maxilla 1 uniarticulate; owter plate of maxilla 2 geniculate, gaping (Steleuthera); palp article 2 of maxilliped not produced; article 2 of pereopods 3-4 slender, of pereo- pod 5 broad; telson cleft. Species: 7, cold-water, N. and S. Atlantic, primarily bathyal.
Stegocephalopsis Schellenberg Stegocephalopsis Schellenberg, 1925a.
Type-species: Cancer ampulla Phipps, 1774 (monotypy). See Gurjanova, 1962.
285-135 O - 69 - 29
444 U.S. NATIONAL MUSEUM BULLETIN 271
Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped not produced; article 2 of pereopods 3-4 slender, of pereopod 5 broad; telson cleft halfway or more. Species: 3, arctic-boreal, littoral to bathyal.
Stegocephalus Kr¢éyer
Stegocephalus Kr@éyer, 1842.—Stebbing, 1906.
Type-species: S. inflatus Krgyer, 1842 (monotypy). See Sars, 1895.
Mandibular incisor toothed; palp of maxilla 1 uniarticulate; outer plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped not produced; article 2 of pereopod 3 slender, of pereopods 4-5 broad; telson cleft halfway; upper lip, lower lip, and maxilla 1 not elongate; spines of outer plate of mavilla 2 with hooks (Stegocephalina). Species: 2, subaretic-boreal, littoral to bathyal.
Steleuthera J. L. Barnard Steleuthera J. L. Barnard, 1964a.
Type-species: S. maremboca J. L. Barnard, 1964a (original designation).
Mandibular incisor toothed; palp of maxilla 1 untarticulate; outer plate of maxilla 2 not geniculate (Stegocephaloides); palp article 2 of maxilliped not produced; article 2 of pereopods 3-4 slender, of pereo- pod 5 broad; telson poorly cleft. Species: 1, Peru, hadal.
Tetradeion Stebbing Tetradeion Stebbing, 1899d; 1906.
Type-species: Cyproidia crassa Chilton, 1883 (original designa- tion). See Hurley, 1955.
Mandibular incisor toothed; palp of maxilla 1 biarticulate; outer plate of maxilla 2 geniculate, gaping; palp article 2 of maxilliped distomedially produced; article 2 of pereopods 3-4 slender, of pereopod 5 broad; pereopod 5 with only three articles; telson entire. Species: 1, New Zealand, littoral.
Stenothoidae
FIGURE 162
Diaanosis.—Accessory flagellum absent or composed of one or two vestigial articles; mandibular molar evanescent; coxa 1 very small, always partially covered by following coxae; coxa 4 enlarged,
MARINE GAMMARIDEAN AMPHIPODA 445
shield-like, not posterodorsally excavate; uropod 3 uniramous; outer lobes of maxilliped vestigial; telson entire; pereopod 3 with article 2 slender. See Amphilochidae, Thaumatelsonidae, Cressidae, Leucothoidae, Anamixidae, Phliantidae, Pagetinidae.
DescripTion.—Rostrum inconspicuous; accessory flagellum absent or 1- or 2-articulate, vestigial; body smooth or carinate; coxa 1 small, hidden by following coxae; coxa 4 enlarged, shield-like, not posterodorsally excavate; upper lip incised; mandible with weak, sparsely spinose molar, not triturative, palp absent and when present 1-, 2-, or 3-articulate; lower lip usually with inner lobes amalgamated, outer lobes with blunt extremities; maxilla 1 with 2- or 1-articulate palp; maxilla 2 small, stout; maxillipeds slender, with outer lobes vestigial; gnathopods usually powerful, subchelate, occasionally feeble, gnathopod 1 often simple; uropod 3 uniarticulate, ramus 2-articulate; telson of medium length, entire; pereopod 3 with slender article 2.
ReLarionsHip.—The Amphilochidae resemble stenothoids but have a biramous uropod 3 and well-developed outer lobes on the maxilliped.
The Thaumatelsonidae have some urosomal segments coalesced and the telson usually strongly thickened. Perhaps the Thaumatel- sonidae deserve only subfamily status as members of the Stenothoidae. The Cressidae differ from the Stenothoidae by the coalescence of the telson with pleonite 6 and the expanded article 2 of pereopod 3.
Classification of Genera of Stenothoidae
The genera of Stenothoidae seem to be segregated from each other by very unsatisfactory means, such as conditions of mandibular palp, maxilla 1, accessory flagellum, and to some extent breadth of article 2 on pereopods 3-5. In other familes these characters (see following key) often distinguish species-groups into natural genera but in the Stenothoidae such “pigeon-holing” throws together species with great diversity in gnathopods. Several genera thus represent grades of structure rather than clades. In these small organisms mandibular palps are very difficult to work with taxo- nomically as palps are hard to observe and may break off during dissection. There is a strong possibility that some species may indeed have palps in some populations and lack them in others. It is prudent to mount the head of the animal on a slide and look for the mandib- ular palp before dissection. Often the articulation line separating palp articles 1 and 2 of the first maxillae is difficult to resolve.
Shoemaker (1955) in his key to Stenothoidae shows the logic in reestablishing the genera Prometopa and Microstenothoe which were
MARINE GAMMARIDEAN AMPHIPODA 447
formerly synonymous with Metopa and Stenothoe, for his key points out the fact that Proboloides and Metopoides are characterized only by the presence or absence of an accessory flagellum. Because Pro- metopa and Microstenothoe bear accessory flagella, it is inconsistent to reduce them without also eliminating the genera Proboloides and Metopoides.
Nomenclatural Changes in Stenothoidae
Mesostenothoides Gurjanova (1938) is removed to Stenothordes.
Metopoides Della Valle (1893) (Stebbing, 1906) is reestablished.
Metopella nasutigenes (Stebbing, 1888) is removed to Probolisca.
Microstenothoe Pirlot (1933b) is reestablished.
Parametopella stelleri Gurjanova (1948) omitted from index by J. L. Barnard (1958a).
Parametopella minuta (Holmes), erroneously assigned to this genus, is removed to Stenothoe.
Prometopa Schellenberg (1931) is reestablished.
Stenothoe minuta Holmes, (Stebbing, 1906; Kunkel, 1918), erroneously assigned to Parametopella previously.
Stenula J. L. Barnard (1962c) was erected for species previously assigned to Mesostenothoides.
Key to the Genera of Stenothoidae (emended after Shoemaker, 1955)
1. Pereopods 4 and 5: article 2 linear (figs. 162¢,) . .. ren Key vA Pereopod 4: article 2 linear (fig. 162v); pereopod 5, mite D ended (figs. IGAS)) 6 5 i HUG 2 cheb aan IONS Pereopods 4 and 5: “onal 2 oanendad Gres 162a, a Sia De ne Maal iat] Bente KEY A
lemelalprotamaxidla taunianrticmlaten (hg. G2q)) a) 4) aes mien Gea eign) Sec eee Palp of maxilla 1 biarticulate (fig. 162h) ...... .. .. . .Probolisca 2. Mandibular palp absent (fig. 162c)......... ... . Parametopella Mandibular palp l-articulate .. . ..... . . Metopelloides Mandibular palp 2- or 3-articulate (fiz. 162d) Tae oti an ee ne Ee COpella:
ass
Figure 162.—Stenothoidae: a, Stenothoe marina (Bate) (Sars, 1895, pl. 80). Upper lip: b, Stenothoe. Mandible: c, Stenothoe; d, Proboloides gregarius (Sars, 1895, pl. 84). Lower lip: e, Stenothoe; f, Proboloides. Maxilla 1: g, Metopa alderi (Bate) (Sars, 1895, pl. 86); h, Stenothoe. Maxilla 2:1, Stenothoe. Maxilliped: 7, Stenothoe; k, Metopa; 1, Probolotdes. Gnathopod 1: m, Metopa pusilla (Sars, 1895, pl. 90); n, Metopa alderi; 0, Metopa robusta (Sars, 1895, pl. 96). Gnathopod 2: p, Metopella longimana (Boeck) (Sars, 1895, pl. 97); ¢, Metopa alderi. "Telson: r, Stenothoe. Pereopod 5:5, Mesometopa neglecta (Hansen) (Sars, 1895, pl. 97); t, Metopella; u, Stenothoe. Pereopod4:v, Metopella. Uropod 3: w, Stenothoe.
448
KEY B
1. Palp of maxilla 1 uniarticulate . Palp of maxilla 1 biarticulate. . Bilan 2. Mandibular palp l-articulate or absent . Mandibular palp 2- or 3-articulate .
KEY C
1. Palp of maxilla 1 uniarticulate . Palp of maxilla 1 biarticulate
2. Mandibular palp absent. Mandibular palp l-articulate Mandibular palp 2- or 3-articulate .
3. Accessory flagellum absent . Accessory flagellum l1-articulate
4. Mandibular palp absent Mandibular palp present
5. Accessory flagellum 1l-articulate Accessory flagellum absent
6. Mandibular palp 1-articulate Mandibular palp 2- or 3-articulate .
7. Accessory flagellum absent Accessory flagellum 1- or 2-articulate
Genera of Stenothoidae
Mesometopa Gurjanova Mesometopa Gurjanova, 1938.
Type-species:
U.S. NATIONAL MUSEUM BULLETIN
27:1
.2
. Mesoproboloides
. Stenothoides . Mesometopa
5B
Bes LA . Parametopa . Stenula
12 epee
. Metopa
. Prometopa ao Microstenothoe . Stenothoe
. Prostenothoe En rte of Proboloides Metopoides
Metopa esmarki Boeck, 1872 (original designation).
Mandibular palp 2- or 3-articulate; palp of maxilla 1 uniarticulate; article 2 of pereopod 4 linear, of article 5 expanded. Species: 4,
arctic-boreal, littoral.
Mesoproboloides Gurjanova
Mesproboloides Gurjanova, 19388.
Type-species: designation).
Metopella cornuta Schellenberg,
1926a
(original
Mandibular palp 3-articulate; palp of maxilla 1 biarticulate; arti- cle 2 of pereopod 4 linear, of pereopod 5 expanded. Species: 2,
subantarctic, littoral. Metopa Boeck Metopa Boeck, 1871.—Stebbing, 1906.
Metopina Norman, 1900b (homonym, Diptera). Sthenometopa Norman, 1902.
Type-species:
1876). See Gurjanova, 1948, 1951; Shoemaker, 1955.
Leucothoe clypeata Kroyer, 1842 (selected by Boeck,
MARINE GAMMARIDEAN AMPHIPODA A49
Mandibular palp 2- or 3-articulate; palp of maxilla 1 uniarticulate; article 2 of pereopods 4—5 expanded; accessory flagellum absent (Pro- metopa). Species: 49, subarctic-boreal, littoral to abyssal, (one species Prometopa tuberculata is subantarctic).
Metopella Sars
Metopella Sars, 1985.—Stebbing, 1906.
Type-species: Metopa longimana Boeck, 1871 (selected by Gur- janova, 1938). See Sars, 1895.
Mandibular palp 2- or 3-articulate; palp of maxilla 1 uniarticulate; article 2 of pereopods 4-5 linear. Species: 8, arctic-boreal, littoral.
Metopelloides Gurjanova Metopelloides Gurjanova, 1938, 1951.
Type-species: Metopella micropalpa Shoemaker, 1930 (original designation).
Mandibular palp 1l-articulate; palp of maxilla 1 untarticulate; article 2 of pereopods 4-5 linear. Species: 9, arctic-boreal, littoral,
Metopoides Della Valle
Metopoides Della Valle, 1893.—Stebbing, 1906.
Type-species: Metopa magellanica Stebbing, 1888 (selected by Gurjanova, 1938).
Mandibular palp 2- or 3-articulate; palp of maxilla 1 biarticulate; article 2 of pereopods 4-5 expanded; accessory flagellum 1-articulate (Proboloides). Species are combined with Proboloides; most Meto- poides are subantarctic.
Microstenothoe Pirlot Microstenothoe Pirlot, 1933b. Type-species: M. ascidiae Pirlot, 1933b (original designation). Mandibular palp absent; palp of maxilla 1 biarticulate; article
2 of pereopods 4—5 expanded; accessory flagellum 1-articulate (Steno- thoe). Species combined with Stenothoe.
Parametopa Chevreux Parametopa Chevreux, 1901b. Type-species: P. kervillei Chevreux, 1901b (original designation). See Chevreux and Fage, 1925. Mandibular palp absent; palp of maxilla 1 uniarticulate; article 2 of pereopods 4—5 expanded. Species: 3, subarctic and warm-temper- ate in Atlantic, littoral.
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Parametopella Gurjanova Parametopella Gurjanova, 1938, 1951.
Type-species: Stenothoe cypris Holmes, 1905 (original designation). See Kunkel, 1918.
Mandibular palp absent; palp of maxilla 1 uniarticulate; article 2 of pereopods 4-5 linear. Species: 3, subarctic-boreal, littoral.
Probolisca Gurjanova Probolisca Gurjanova, 1938.
Type-species: Metopa ovata Stebbing, 1888 (original designation). Mandibular palp 2- or 3-articulate; palp of maxilla 1 biarticulate; article 2 of pereopods 4-5 linear. Species: 3, antiboreal, littoral.
Proboloides Della Valle
Proboloides Della Valle, 1893.—Stebbing, 1906.—K. H. Barnard, 1932. Proboltella Walker, 1906b, 1907.
Type-species: Metopa gregaria Sars, 1882 (selected by Gurjanova, 1938). See Sars, 1895.
Mandibular palp 2- or 3-articulate; palp of maxilla 1 biarticulate; article 2 of pereopods 4-5 expanded; accessory flagellum absent (Metopoides). Species (incl. Metopoides): 34, bipolar, littoral to abyssal.
Prometopa Schellenberg
Prometopa Schellenberg, 1926a.
Type-species: P. tuberculata Schellenberg, 1926a (monotypy).
Mandibular palp 2- or 3-articulate; palp of maxilla 1 uniarticulate; article 2 of pereopods 4—5 expanded; accessory flagellum 1-articulate (Metopa). Species: 1; included with Metopa.
Prostenothoe Gurjanova Prostenothoe Gurjanova, 1938.
Type-species: P. sextonae Gurjanova, 1938 (original designation) ; 1951. Mandibular palp 1-articulate; palp of maxilla 1 biarticulate; article 2 of pereopods 4—5 expanded. Species: 1, Japan, Okhotsk Sea, littoral. Stenothoe Dana
Stenothoe Dana, 1852b.—Stebbing, 1906. Probolium Costa, 1853c, 1857.
Montagua Bate, 1857a (homonym, Decapoda). Montaguana Chilton, 1883.
MARINE GAMMARIDEAN AMPHIPODA 451
Type-species: S. validus Dana, 1853 (selected by monotypy of Dana, 1853). See Chevreux and Fage, 1925.
Mandibular palp absent; palp of maxilla 1 biarticulate; article 2 of pereopods 4-5 expanded; accessory flagellum absent (Microstenothoe). Species: 32, cosmopolitan, littoral to bathyal.
Stenothoides Chevreux
Stenothoides Chevreux, 1900!—J. L. Barnard, 1962c. Mesostenothoides Gurjanova, 1938. Type-species: S. perrieri Chevreux, 1900 (monotypy). Mandibular palp 1-articulate or absent; palp of maxilla 1 uni- articulate; article 2 of pereopod 4 linear, of article 5 expanded. Spe- cies: 6, subarctic-boreal, littoral.
Stenula J. L. Barnard Stenula J. L. Barnard, 1962c.
Type-species: Stenothoides latipes Chevreux and Fage, 1925 (ori- ginal designation).
Mandibular palp 1-articulate; palp of maxilla 1 uniarticulate; article 2 of pereopods 4-5 expanded. Species: 10, arctic-boreal, littoral to bathyal.
Stilipedidae
FicgurEs 163, 164, FRONTISPIECE
Dracnosis.—Accessory flagellum absent or vestigial, less than 3-articulate; mandible broad and lacking molar; maxillae foliaceous (figs. 163¢,d); gnathopods simple but not especially slender. See Pardaliscidae Hyperiopsidae, Laphystiopsidae, Astyridae.
DescripTion.—Peduncles of antennae very short, accessory flagel- lum vestigial or absent; upper lip asymmetrically bilobed; mandible very broad, pardaliscid in appearance, palp 3-articulate, thin, lacking molar; lower lips variable; maxilla 1 variable, some parts always foliaceous; maxilla 2 foliaceous; plates of maxilliped large, palp variable in length; gnathopods simple but not especially slender;
,coxae long; uropod 3 biramous, peduncle short, rami elongate, lanceolate; telson apically emarginate, short.
ReELATIONSHIP.— Differing from the Acanthonotozomatidae by the foliaceous maxillae (except Mazilliphimedia in that family) and the mouthparts not being arranged in a conical bundle.
The Astyridae differ from the Stilipedidae by the weaker foli- aceousness of the first maxilla but intergradation occurs in the genus
452 U.S. NATIONAL MUSEUM BULLETIN 2711
Alexandrella assigned to the Stilipedidae, for its first maxillae are not as foliaceous as those of the type-genus Stilipes. The palp is especially strongly geniculate, resembling species of Hyperiopsidae. Astyridae bear a distinct but nontriturative mandibular molar. Astyrovdes is removed from the Astyridae and synonymized with Alexandrella.
The Laphystiopsidae have normal maxillae, a mandibular molar, and small maxillipedal plates.
Ficure 163.—Stilipedidae: a, lexandrella dentata Chevreux (1912b). Maxilla 1: 3, Alexandrella; c, Stilipes sanguineus (Hurley, 1954e). Maxilla 2: d, Stilipes. Maxilliped: e, Alexandrella; f, Stilipes.
MARINE: GAMMARIDEAN AMPHIPODA 453
The absence of a mandibular molar distinguishes Stilipedidae from the Synopiidae and Hyperiopsidae. Stilipedids resemble the Par- daliscidae in the mandible but differ by the long coxae and the foliaceous second maxillae. The first maxillary palp of some pardali- scids is foliaceous but not the inner plate. Pardaliscids have a multi- articulate accessory flagellum (except Halicoides).
Ficure 164.—Stilipedidae: Mandible: a,b, Stilipes sanguineus (Hurley, 1954e). Upper lip: c, Alexandrella dentata Chevreux (1912b); d, Stilipes sanguineus. Lower lip: e, Stilipes sanguineus; f, Alexandrella. Gnathopod 2: g, Alexandrella. Gnathopods 1, 2: h,1, Stilipes sanguineus. Telson: 7, Stilipes distincta Holmes (1908). Uropod 3: k, Stzlipes distincta.
Key to the Genera of Stilipedidae
1. Palp of maxilliped greatly exceeding outer plate (fig. le upper lip very
weakly emarginate (fig. 164d) ... . . . . . Stilipes Palp of maxilliped not aes outer plat (i 163¢),. upper lip deeply inciseds (tig G40) es 3 : ... . . . Alexandrella
Genera of Stilipedidae
Alexandrella Chevreux, new synonymy
Alexandrella Chevreux, 1911c; 1912a; 1912b. Astyroides Birstein and Vinogradova, 1960.
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Type-species: A. dentata Chevreux, 1912a; 1912b (designated by Chevreux, 1912a). Species: 2, antarctic to N.W. Pacific, littoral to hadal.
Stilipes Holmes
Stilipes Holmes, 1908. Cacao K. H. Barnard, 1931; 1932. Type-species: S. disltincta Holmes, 1908 (original designation). See
Shoemaker, 1964. Species: 3, N.E. Pacific, antiboreal, littoral to bathyal.
Synopliidae, revised {including Tironidae]
Figures 165-167
Dracnosis.—Head massive,* and often galeate, produced into a downturned or deflexed rostrum; gnathopods feeble, scarcely sub- chelate, often simple; telson elongate (rarely short), but always as long as peduncle of uropod 1; eyes when present coalesced. See Astyridae, Vitjazianidae, Argissidae, Hyperiopsidae, Oedicerotidae.
Descrietion.—The characters of this family are so subtle that they are practically indefinable, yet taxonomists have little trouble in recognizing a synopiid. An inclusive diagnosis has to be inter- spersed with the words ‘‘most species” since there are so many excep- tions to any pair of character alternatives, yet the majority of species shares anyone of the following characters and all species share the majority of them.
Antenna 1 with multiarticulate accessory flagellum (except Jeddo); base of primary flagellum conjoint in male; eyes when present coal- esced dorsally; mandible with poorly developed, slender palp (except Synopia), molar well developed, often extraordinarily large, smooth and dominating mandible, article 3 usually shortened considerably (except in Bruzeliopsis); outer lobes of lower lip frequently notched; mouthparts otherwise like basic gammaridean; coxa 4 never much larger than coxa 3; coxae long (as contrasted with Pardaliscidae) ; uro- pods 1-2 with outer ramus shortened; telson cleft (except in Bruzelia, Bruzeliopsis, and some Synopia), elongate (except Synopia), as long as peduncle of uropod 1; head massive, produced into a deflexed or
*As tall as or taller than long (length from posterior margin of head to anterior margin of lateral lobe), head (except Tiron) as long as pereonites 1-3 combined.
MARINE GAMMARIDEAN AMPHIPODA 455
f ee |
Figure 165.—Synopiidae: a, Tiron spiniferum (Stimpson) (Sars, 1895, pl. 140, as T. acanthurus); b, Syrrhottes serratus Sars (1895, pl. 137). Accessory flagella: c, Tiron; d, Syrrhoe crenulata Goés (Sars, 1895, pl. 136). Mandible: e, Synopia variablis Spandl (J. L. Barnard, 1965).
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Figure 166.—Synopiidae: a, Syrrhoe crenulata Goés (Sars, 1895, pl. 136); b, Bruzelia typica Boeck (Sars, 1895, pl. 138). Upper lip: ¢, Tiron spiniferum (Stimpson) (Sars, 1895, pl. 140, as T. acanthurus). Mandible: d, Bruzelia; e, Tiron. Lower lip: f, Syrrhottes serratus Sars (1895, pl. 137); g, Tiron. Telson: h, Bruzelia tuberculata Sars (1895, pl. 139).
MARINE GAMMARIDEAN AMPHIPODA 457
downturned rostrum; pereonites 1-4 often very short; gnathopods feeble, slender, scarcely subchelate; eyes when present dorsally confluent.
RELATIONSHIP.—Synoplids differ from the basic gammaridean mainly by the massive head with large rostrum, poorly developed mandibular palp in most genera, feeble gnathopods, and elongated telson. From the Eusiridae they differ by the combination of the same characters, except for the elongate telson. They are similar to the Oedicerotidae in head, eyes, and general appearance, but all Oedicerotidae have a vestigial or no accessory flagellum, usually powerful gnathopods and short, uncleft telson.
Synopiidae differ from Pardaliscidae by the possession of a man- dibular molar but often appear similar in other ways, except that the coxae of Pardaliscidae are always very short.
The Vitjazianidae differ from Synopiidae by the small, poorly rostrate head.
The Argissidae have a normal gammaridean head, practically no rostrum, and a unique pattern in the shapes of coxae 1-4.
Astyridae resemble Synoptidae closely and differ from them in the occurrence of just one or two articles in the accessory flagellum, the setose, laminate, and not triturative mandibular molar. All synoplids, except Bruzeliopsis, have a reduced palp article 3 on the mandible, whereas astyrids have an elongated article 3. All Astyridae have short telsons and widely spaced outer lobes on the lower lip.
A revision of Synopiidae is presented herein and where generic composition has been changed, the species have been listed.
Key to the Genera of Synopiidae
1. Mandibular molar very large and smooth (minutely setulose or fuzzy oc- casionally), completely dominating mandible, body of mandible extremely bulkyacsubclobularydwartinejoalpn (igs GGG) 18 je ln ee 2
Mandible as in first triplet but palp absent . .. . PARKA edd Mandibular molar of medium size, occasionally emeoth os minutely fuzzy but usually triturative, body of mandible usually stout but not extra- ordinarily bulky or subglobular, palp relatively strong (fig. 166e). . . 5
2. Coxa 3 strongly expanded anteroposteriorly and much larger than coxa 4 (fig. 166a) (more than twice as much surface area), coxa 3 with strong posterodorsal excavation dorsal to expansion, article 1 of antenna 1 very elongate and bearing long subconical distal process, articles 2 and 3 short and subequal in length, coxa 4 very small and shaped like a comma (fig. OOO) eae Ree aii . . . . . Bruzeliopsis
Coxae 3 and 4 atineqinell | in size ‘oad Teneal coxa 3 never with more than 1.5— times as much surface area as 4, eoxa 3 weakly expanded or not expanded distally, lacking distinct posterodorsal excavation, coxa 4 adze-shaped or quadrate, its posteroventral lobe or midposterior cusp distinetly directed
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U.S. NATIONAL MUSEUM BULLETIN 271
posteriorwards and at right angles to anterior margin, posterodorsal excavation almost right-angular, articles 2 and 3 of antenna 1 even if shortened always together as long as article 1, distal tooth of latter if presenti shortzandihOokc likey men ee any oer amee n n ea MelSONVeN tires pedicels kena ea dieg ee Aalst 5 tice) Cac Oe SEEN Ta ee es a 4 Telson cleft .... . SORA ah, ae Gyrrhoites Palms of gnathopods recta: one or ro corrate spines, coxa 4 adze-shaped, tooth distinctly posteroventral . ... . .. . . . Bruzelia Palms of gnathopods with 4 nonserrate hint: spines, coxa 4 rectangular and bearing midposterior cusp . . . . . .Stephobruzelia, new genus Both telson and peduncle of aoa 3 very short and subequal in length, mandibular palp extremely stout (fig. 165e). . .... . . . .Synopia Telson elongate, twice as long as peduncle of uropod 3 even when peduncle elongate, mandibular palp slender (figs. 166d,e) .... . Pa RAY ck | cro AG) Coxa 3 strongly expanded distally, some portion of distal half at least twice as broad as proximal end, expansion directed posteriorly, posterodorsal margin strongly excavate, coxa 4 much smaller than 3, comma-shaped and usually with less than half as much surface area as coxa3d...... 7 Coxa 3 rectangular or quadrate, scarcely expanded distally, occasionally with anteroventral corner attenuated only, posterior margin nearly parallel with anterior margin, coxa 4 almost as long as 3, its surface area scarcely smaller than that of coxa 3, coxa 4 adze-shaped or chisel-shaped . . . 9 Coxa 3 rectangular or subquadrate, scarcely expanded distally, coxa 4 noticeably shorter than 3 and weakly comma-shaped . some species a Tiron Article 6 of gnathopods linear, slender, ae simple, lacking strong serrate spines... . .. . . . Pseudotiron Article 6 of gnathopods, if ample ror feat or 2 Hechenmulan either ovate and tumid or bearing distinct transverse palms, palms bearing one or two serrate spines .. . Ae OU eames Telson cleft more than ellinnee “elias of apeunapades teens . Syrrhoe Telson cleft less than 20%, palms of gnathopods oblique or indistinct. Ileraustroe, new genus Article 6 of both gnathopods linear, slender, een simple, lacking strong serrate spines .. . . . Tiron Article 6 of gnathopod 1 tumid but ail 6 of Dthoped 2 2 pertcetly linear and rectangular . . Syrrhoe fimbriatus! Stebbing and Robertson, 1891 Article 6 of both gnathopods, if simple not linear or rectangular, either ovate and tumid or bearing transverse palms, palms bearing one or two serrate
spines... Air errcanuises, wll) Palms of Ereihonodl trmavemel engin Hose 6 one very taree spine giving chelate appearance to palms. . . . . . . . . Garosyrrhoe Palms of gnathopods oblique or Gpsolescent Heating one or two serrate spines but not appearing chelate ........ .. .. . Austrosyrrhoe
' Currently assigned to Austrosyrrhoe.
Genera of Synopiidae
Austrosyrrhoe K. H. Barnard
Austrosyrrhoe K. H. Barnard, 1925.
MARINE GAMMARIDEAN AMPHIPODA 459
Type-species: A. crassipes K. H. Barnard, 1925 (monotypy). Poorly known species.
Composition: A. septentrionalis Stephensen, 1931.
Mandible of medium size, molar of medium size and not dominating mandible, apparently weakly triturative or fuzzy; coxa 3 rectangular or subquadrate, scarcely expanded distally, posterior margin nearly parallel with anterior margin and not strongly excavate; coxa 4 adze- shaped, nearly as long as coxa 8, its surface area scarcely smaller than coxa 3; article 6 of gnathopods not a perfectly linear rectangle, palms oblique or obsolescent and bearing one or two large serrate spines; telson elongate and deeply cleft. Species: 2, Atlantic, bathyal- abyssal.
Bruzelia Boeck
Bruzelia Boeck, 1871.—Stebbing, 1906.
Ficure 167.—Synopiidae: Tiron spiniferum (Stimpson) (Sars, 1895, pl. 140, as 7. acan- thurus): a,b, maxillae 1, 2; c, maxilliped; d,e, gnathopods 1, 2; f, uropod 3; g, telson. Syrrhoe crenulata Goés (Sars, 1895, pl. 136): h,i, gnathopods 1, 2. Synopta scheeleana Bruzelius (Stebbing, 1888): 7,k, gnathopods 1, 2; /, pereopod 2.
285-135 O - 69 - 30
460 U.S. NATIONAL MUSEUM BULLETIN 271
Type-species: 8B. typica Boeck, 1871 (monotypy). See Sars, 1895.
Composition: B. australis Stebbing, 1910; B. diodon K. H. Barnard, 1916; B. tuberculata Sars, 1882, 1895.
Mandible enlarged, molar enlarged, dominating mandible, smooth or fuzzy; coxa 3 rectangular, not distally expanded except for antero- ventral cusp, posterior margin parallel with anterior margin and not excavate; coxa 4 nearly as long as coxa 3, surface area subequal to coxa 3, adze-shaped; article 6 of gnathopods trapezoidal or subovate, palms oblique, bearing one or two large serrate spines; telson elongate and entire. Species: 4, probably cosmopolitan, mainly cold-water, bathyal-abyssal (B. australis is littoral),
Bruzeliopsis Chevreux
Bruzeliopsis Chevreux, 1911b.
Type-species: B. albertt Chevreux, 1911b (original designation).
Composition: B. turba J. L. Barnard, 1964a; Bruzelia cuspidata J. L. Barnard, 1962d.
Mandible enlarged, molar enlarged, dominating mandible, smooth or fuzzy; coxa 3 distally expanded, with broad quadrate posterior lobe, posterodorsal margin excavate; coxa 4 much smaller than 3, comma-shaped; article 6 of gnathopods trapezoidal or subovate, palms oblique, bearing one or two nonserrate or distally flagellate locking spines; telson elongate and entire or minutely cleft apically; article 1 of antenna 1 elongate and bearing conical distal cusp. Species: 3, Atlantic, bathyal-abyssal.
Garosyrrhoe J. L. Barnard Garosyrrhoe J. L. Barnard, 1964a.
Type-species: Syrrhoites bigarra J. L. Barnard, 1962a (original designation).
Mandible of medium size, molar of medium size and not dominating mandible, apparently weakly triturative or fuzzy; coxa 3 softly rec- tangular, posterior margin nearly parallel with anterior margin and not excavate; coxa 4 expanded midposteriorly, posterodorsal margin sloping, not concave but giving appearance of posterodorsal excavation, coxa 4 with larger surface area than coxa 3; article 6 of gnathopods trapezoidal or subovate, palms transverse and each defined by one large serrate spine lending chelate appearance; telson elongate and deeply cleft. Species: 1, California, littoral.
Ileraustroe, new genus
Type-species: Austrosyrrhoe ilergetes J. L. Barnard, 1964a (present selection).
MARINE GAMMARIDEAN AMPHIPODA 461
Composition: Austrosyrrhoe ?torpens J. L. Barnard, 1964a (not J. L. Barnard, 1962d) (=new species).
Mandible of medium size, molar of medium size or slightly enlarged and not dominating mandible [palp large], fuzzy; coxa 3 strongly expanded distally, with midposterior or posteroventral lobe, postero- dorsal margin excavate; coxa 4 much smaller than 3, comma-shaped; article 6 of gnathopods trapezoidal or subovate, palms oblique, bearing at least one large serrate spine; telson elongate but cleft only 20 percent or less. Species: 2, Mediterranean, Caribbean, abyssal.
Jeddo J. L. Barnard Jeddo J. L. Barnard, 1962d.
Type-species: J. sumplisyrrhis J. L. Barnard, 1962d (original designation).
Mandible enlarged, molar enlarged, dominating mandible, smooth or fuzzy; mandibular palp absent; coxa 3 distally expanded, with large posteroventral lobe, posterodorsal margin excavate; coxa 4 much smaller than 3, hemi-ovate; article 6 of gnathopods essentially linear and rectangular but not strongly elongate, simple, lacking distinct locking spines; telson elongate and cleft about halfway. Species: 1, S. Atlantic, deep bathyal.
Pseudotiron Chevreux
Pseudotiron Chevreux, 1895.—Stebbing, 1906.
Type-species: P. bouvieri Chevreux, 1895 (original designation).
Mandible of medium size, molar of medium size, columnar and triturative; coxa 3 strongly expanded distally, with quadrate postero- ventral lobe, posterodorsal margin forming quadrate excavation, coxa 4 much smaller than 3, comma-shaped; article 6 of gnathopods a nearly perfectly linear rectangle, simple and lacking spine(s) ; telson elongate and deeply cleft. Species: 3, Indo-Atlantic, Mediterranean, bathy- abyssopelagic.
Stephobruzelia, new genus
Type-species: Bruzelia dentata Stephensen, 1931 (present selection).
Mandible enlarged, molar enlarged, dominating mandible, smooth or fuzzy; coxa 3 rectangular, not distally expanded except for antero- ventral cusp, posterior margin parallel with anterior and not excavate; coxa 4 as long as and as large as coxa 3, rectangular, with midposterior cusp; article 6 of gnathopods trapezoidal or subovate, palms oblique, bearing four large, nonserrate, distally blunt spines; telson elongate and entire. Named for K. Stephensen. Species: 1, subarctic, bathyal.
462 U.S. NATIONAL MUSEUM BULLETIN 271
Synopia Dana Synopia Dana, 1852b.—Stebbing, 1906.
Type-species: S. ultramarina Dana, 1853 (present selection). See Stebbing, 1888 (as S. scheeleana).
Mandible of medium size, molar of medium size, triturative or fuzzy, palp extremely tumid (fig. 165e); coxa 3 strongly expanded dis- tally, with quadrate posteroventral lobe, posterodorsal margin excavate; coxa 4 much smaller than 3, very short, weakly comma- shaped; article 6 of gnathopods slender or tumid, subovate, not perfectly linear, simple, posterior margins setose but lacking distinct locking spines, dactyl of gnathopod 2 vestigial (fig. 167k); telson and peduncle of uropod 8 both short and subequal in length, telson cleft or entire, distally castellate in forms with entire telson; mazillipeds foliaceous. Species: 3, tropical, epipelagic.
Syrrhoe Goés Syrrhoe Goés, 1866.—Stebbing, 1906.
Type-species: S. crenulata Goés, 1866 (selected by Boeck, 1876). See Sars, 1895.
Mandible of medium size, molar of medium size and not dominating mandible, apparently weakly triturative or fuzzy; coxa 3 strongly expanded distally, with posteroventral lobe, posterodorsal margin excavate; coxa 4 much smaller than 3, comma- or weakly adze-shaped ; article 6 of gnathopods trapezoidal, subovate and elongate, palm nearly transverse and armed with serrate spine not giving chelate appearance; telson elongate and deeply cleft. Species: 8, probably cosmopolitan, littoral to bathyal.
Syrrhoites Sars Syrrhoites Sars, 1895.—Stebbing, 1906.
Type-species: Bruzelia serrata Sars, 1879 (original designation). See Sars, 1895.
Composition: S. anaticauda K. H. Barnard, 1930; Kinda lorida J. L. Barnard, 1962d; S. pacificus Nagata, 1965; S. pusilus Enequist, 1950; K. sorpresa J. L. Barnard, 1962d; S. tenellus K. H. Barnard, 1925; S. terceris J. L. Barnard, 1964a; S. walkeri Bonnier, 1896.
Mandible enlarged, molar enlarged, dominating mandible, smooth or fuzzy; coxa 3 rectangular, not distally expanded, posterior margin parallel with anterior margin and not excavate; coxa 4 nearly as long as coxa 3, surface area subequal to coxa 3, adze-shaped; article 6 of gnathopods trapezoidal or subovate, palms oblique, bearing one or two large, nonserrate or distally flagellate locking spines; telson
MARINE GAMMARIDEAN AMPHIPODA 463
elongate and deeply cleft. Species: 9, cosmopolitan, cold-water, deep littoral to abyssal.
Tiron Liljeborg
Tiron Liljeborg, 1865.—Stebbing, 1906.—Shoemaker, 1955. Tessarops Norman, 1868 (homonym, Arachnida).
Type-species: Lysianassa spinifera Stimpson, 1853 (monotypy and subsequent synonomy). See Sars, 1895 (as T. acanthurus).
Mandible of medium size, molar of medium size, strongly projecting, columnar and triturative; coxa 3 softly rectangular, posterior margin almost parallel with anterior margin and not strongly excavate; coxa 4 variable, typically adze-shaped and almost as long as coxa 3, surface area of coxa 4 nearly equal to coxa 3; rarely coxa 4 distinctly shorter and smaller than 3 and weakly comma-shaped; article 6 of gnathopods a nearly perfectly linear rectangle, simple and lacking distinct locking spine(s); telson elongate and deeply cleft. Species: 8, cosmopolitan, littoral (rarely to bathyal).
Superfamily Talitroidea [includes Talitridae, Hyalidae, and Hyalellidae] FicurREs 168, 169
Diaenosis.—Accessory flagellum absent; mandible lacking palp; uropod 3 essentially uniramous; tiny scale-like inner ramus rarely present. See Phliantidae, Dogielinotidae, Eophliantidae.
Description.—Accessory flagellum absent; body rarely with teeth or carinations; coxae of medium size; mandible lacking palp, molar large and strongly triturative (except Najna) ; lower lip without inner lobes; palp of maxilla 1 often reduced or absent; maxilliped occasion- ally with fewer than four palp articles; gnathopods usually powerful, occasionally simple or feeble; uropod 3 small, usually with one ramus, inner ramus reduced to a scale when present; telson very short, entire, apically emarginate, cleft, or appearing completely bilobate.
RELAtTIonsuip.—The following families, lacking mandibular palps, differ from the Talitroidea in their coalesced fifth and sixth pleonites: Dexaminidae, Prophliantidae, and Kuriidae.
The Dogielinotidae, which probably should be assigned to the Talitroidea, differ by their haustoriid-phoxocephalid aspect, with multispinose antennae, short spinose pereopods and shape of head, bearing a conspicuous but small rostrum. Their epistome has a nasi- form anterior lobe.
The Eophliantidae differ from talitroids by their cylindrical bodies,
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MARINE GAMMARIDEAN AMPHIPODA 465
short coxae which generally do not touch each other, and degraded mandibular molars.
The Phliantidae differ from Talitroidea by their degraded man- dibular molars and the general aspect of their depressed, massive, often processiferous bodies with laterally splayed coxae. Phliantids* never have enlarged male second gnathopods as do most talitroids, especially marine ones.
*See supplement.
Classification of Talitroidea
Bulycheva (1957) partitioned the Talitridae, emended and restricted the concept of the Talitridae and removed several genera to two new families, the Hyalidae and Hyalellidae. Morphologically there are some small conflicts in her arrangement but they detract little from what appears to be a sensible and logical arrangement, although the classificatory criteria used are to a large extent not qualitative and indeed to some extent are concerned with the ecology of the organisms.
As so outlined, in her extensive paper, the Talitridae are confined to entirely terrestrial genera, many species of which dwell on the strand but nevertheless are not aquatic. The Hyalidae are exclusively marine, being characterized by cleft telsons. The Hyalellidae inhabit both marine and fresh waters, with more genera but few species in the oceans than in freshwater, where one genus, Hyalella, has 27 species. Morphologically, it is impossible strictly and qualitatively to separate the terrestrial Talitridae from the concepts of the other two families. Bulycheva (1957) in her figure 22 has given an excellent synopsis of the telsons of the three families, showing Talitridae with generally uncleft, heavily spinose telsons; Hyalidae with cleft, poorly spinose telsons; and Hyalellidae with uncleft, poorly spinose telsons. She shows the fluvial Chiltonia mihiwaka in the Hyalellidae with uncleft telson but Hurley (1954a) shows it with partially cleft telson, al- though other chiltonias may have the telson uncleft. Chaltonia is not assignable to Talitridae because of the poorly spinose telson. Najna shows both uncleft and minutely cleft telsons in the same species; Bulycheva assigned the genus to the Hyalidae, but it should be removed to a new family because of its mandibular molar.
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Ficure 168.—Talitroidea: a, Talitrus saltator (Montagu) (Sars, 1895, pl. 9, as T. locusta); b, Hyale chevreuxi K. H. Barnard (Chevreux 1901la, as H. macrodactylus). Gnathopod 2: c, Insula antennulella Kunkel (1910); d, Chiltonta mthiwaka (Chilton) (Hurley, 1954a); e, Hyale bassargint Derjavin (Gurjanova, 1951); f, Talitrus; g, Allorchestes plumicornis (Heller) (Gurjanova, 1951, as A. ptilocerus). Gnathopod 1: h, Talitrus; i, Insula.
466 U.S. NATIONAL MUSEUM BULLETIN 271
When present the first maxillary palp is supposed to be 2-articulate in Talitridae and 1-articulate in the other two families, but the marine Parallorchestes, obviously related to Hyalidae, has a 2-articulate first maxillary palp, hence bridges the gap to the terrestrial
Talitridae.
ch [) LMT a Hi t | AQ ays i A (\ iN \ N d l n
Ficure 169.—Talitroidea: a, Najna sp. (J. L. Barnard, 1962c). Upper lip: b, Talitrus saltator (Montagu) (Sars, 1895, pl. 9, as T. locusta). Mandible: c, Talitrus; d, Najna. Lower lip: e, Talitrus. Maxilla 1: f, Parhyalella batesoni Kunkel (1910); g, Hyale nilssoni (Rathke) (Sars, 1895, pl. 11); 4, Talitrus. Maxilla 2: 1, Talitrus. Maxilliped: 7, Insula antennulella Kunkel (1910); k, Parhyale hawaiensis (Dana) (Chevreux, 190la, as Hyale brevipes); 1, Najna; m, Chiltonia mthiwaka (Chilton) (Hurley, 1954a); n, Talitrus. Uropod 3: 0, Parhyale (Shoemaker, 1956b); p, Chiltonia; q, Talitrus. Telson: r, Talitrus; s,
Hyale.
MARINE GAMMIARIDEAN AMPHIPODA 467
There are exceptions to the neat morphological partitioning of talitrid-like animals but there are definitely three or more facies and perhaps three lines of evolution as Bulycheva’s arrangement points out. She did not assign all existing genera to families, which I do in the following scheme, considering for the uses of this handbook that they are subfamilies.
Subfamily Talitrinae: Taltrus, Talorchestia, Orchestia, Orchestoidea. Subfamily MHyalellinae: Hyalella, Parhyalella, Insula, Chiltonia, Afrochiltonia, Neobule, Austrochiltonia, [Najna?] but see below. Subfamily Hyalinae: Hyale, Parhyale, Allorchestes, Parallorchestes.
?(Subfamily Najninae): Nayjna. ?(Subfamily Dogielienotinae): See as a family, p. 207.
Probably Neobule Haswell was improperly described and classified. It has not been recovered since its original description in 1880. It has the aspect of a Parhyalella but may belong to another family, such as the Kuriidae, Prophliantidae, or Lysianassidae. Its ‘“biramous posterior pleopoda” (presumably uropoda) may be an erroneous observation and the condition of the mandibular molar was not stated. It is the only talitrid presumed to have clearly biramous third uropods and so confuses what would otherwise be a clear diagnosis of the family.
Key to the Genera of Talitroidea
Neobule Haswell, a dubious genus, is not included.
1. Terminal maxillipedal palp article unguiform (claw-shaped) (figs. 1697,k) . 2
Terminal maxillipedal palp article not unguiform (figs. 1691,m,n).. . .8 ZeUropod:s: with minutesanner ramus) (fig. 1690)". 2 25 9 2. so Uropod 3 uniramous (fig. 169q) . . . . aarali( bac un pis Gee a ten
3. Palp of maxilla 1 uniarticulate (fig. 169h) . ulate . . . . Parhyale Palp of maxilla 1 biarticulate (often faintly so) (fig. 1694) . . Parallorchestes
A PEaMclSONeMtiney CIEL G OM iis jen i Milieu se itn ee ere Mauna ance vines Amen MCA a Telson cleft (fig. 169s)... . Seiarer ran Naren een Naley UR Diner tae Mee ae ARL RITA
5. Maxilla 1 lacking palp (fig. 169f) . WM yeas Cente ac a ee arnyalella Maxilla 1 bearing palp (figs. 1699,h). ... . peaaaieen tle
6. Male gnathopods 1 and 2 similar to each Other in size sand morphology (fig. 169a).... . . . .Najna and Insula
Male gnathopod 2 sail Nanece ana of difterent aneratbllores than gnathopod
Te Gan) GIO) 3 eo . . . Hyalella (freshwater)
7. Article 5 of male pantheped 2 aneceaal by ticles 4 and 6 (fig. 168e) . Hyale Article 5 of male gnathopod 2 produced between articles 4 and 6 (fig.
GSO) tie tes Sheila, ae Bo ea Ne A orchestés
8. Male gnathopod 1 Fimpic ‘Ge. ‘168h) . Sa Waele een) Clam Meccan Mea ea Cut iaen pAWiNG i Male gnathopod 1 subchelate (fig. 1681) ..... Geel)
9. Male gnathopod 2 feebly chelate (fig. 168f). . . . | Talitrus Ganexiie))
Male gnathopod 2 large and strongly subchelate (fig. 168e). Orchestoidea (terrestrial)
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10. Female gnathopod 1 simple (fig. 168h) . . . . . Talorchestia (terrestrial) Female gnathopod 1 subchelate ... . He eegeMecniny nly ay
11. Article 5 of male gnathopod 2 prodiced ipeinasen cmsniies 4 and 6 Ge NGLA) Go geo) . . . Najna
Article 5 of male paaenenadl 2 iesked peugeenl perieles 4 inl 6 Ge 168e) . 12
12. Antenna 1 shorter than peduncle of antenna 2 (fig. 168a), female gnathopod 2 mitten-shaped (fig. 168f), maxilla 1 usually with palp (even though minute), uropod 3 with ramus... .. . . .Orchestia (terrestrial) Antennae 1 and 2 subequal in length, female gnathopod 2 normally sub- chelate (fig. 168d), maxilla 1 ee palp, uropod 3 usually lacking a
TAMAS (igs OOD) eee hn nene Bair) esti IES}
13. Gnathopods 1 and 2 alike in both ; sexes, ener ole caathonod 2 not en- larcedinian .. . . .Afrochiltonia (freshwater) Gnathopod 2 of male onleweradl, mnie Ghatrom femelle iss ree
14. Malepleopodinormal. ... . . . . . . Austrochiltonia Gaestnrratien) Male pleopod 1 with inner ramus attenuated to backturned, whip-like Lash) oo eae Si aE oie um hil toniay (iEestwjau els)
Genera of Talitroidea
Afrochiltonia K. H. Barnard Afrochiltonia K. H. Barnard, 1955.
Type-species: Chiltonia capensis K. H. Barnard, 1916 (original designation).
Maxilla 1 lacking palp; article 4 of maxillipedal palp short, conical; gnathopods of both sexes subchelate, male gnathopod 2 not larger than 1, female gnathopod 2 like gnathopod 1; male pleopod 1 normal (Chiltonia); uropod 3 lacking inner ramus; telson entire. Species: 1, S. Africa, freshwater.
Allorchestes Dana
Allorchestes Dana, 1849.—Stebbing, 1906. Aspidophoreia Haswell, 1880d.
Type-species: A. compressa Dana, 1852a (selected by Chevreux and Fage, 1925). See Stebbing, 1899a.
Maxilla 1 bearing palp; article 4 of maxillipedal palp unguiform; gnathopods of both sexes subchelate, male gnathopod 2 larger than 1, article 5 produced between articles 4 and 6, female gnathopod 2 like gnathopod 1; uropod 3 lacking inner ramus; telson cleft. Species: 12, cosmopolitan, littoral. At least two kinds of telson occur in this genus, one like that of Hyale, with subconical lobes fully split from each other and set at an angle to each other forming a tent; and the other a quadratiform flat plate with partial cleft. Reorganization of this genus needs investigation.
MARINE GAMMARIDEAN AMPHIPODA 469
Austrochiltonia Hurley
Austrochiltonia Hurley, 1958.
Type-species: Hyalella australis Sayce, 1901 (original designation).
Maxilla 1 lacking palp; article 4 of maxillipedal palp short, conical; gnathopods subchelate in both sexes, male gnathopod 2 much larger than 1, article 5 not projecting between 4 and 6, female gnathopod 2 like 1; male pleopod 1 normal (Chiltonia); uropod 3 lacking inner ramus; telson entire. Species: 2, Australia, freshwater.
Chiltonia Stebbing
Chiltonia Stebbing, 1899a.—Stebbing, 1906.
Type-species: Hyalella mihiwaka Chilton, 1898 (original desig- nation). See Hurley, 1954a.
Maxilla 1 lacking palp; article 4 of maxillipedal palp short, conical; genathopods subchelate in both sexes, male gnathopod 2 much larger than 1, article 5 not projecting between articles 4 and 6, female enathopod 2 like 1; male pleopod 1 uth inner ramus attenuated to a backturned whip-like lash (Afrochiltonia and Austrochiltonia); uropod 3 lacking ramus; telson entire. Species: 3, New Zealand, freshwater.
Hyale Rathke
Hyale Rathke, 1837.—Stebbing, 1906.
Nicea Nicolet, 1849.
(Allorchestina) Brandt, 185la (valid subgenus). Type by present selection: Amphithoe [sic] prevostiz Milne Edwards, 1830.
Galanthis Bate, 1857a.
Type-species: H. pontica Rathke, 1837 (monotypy). See Sars, 1895 (as H. lubbockiana).
Maxilla 1 with 1-articulate palp (but note Sars, 1895, pl. 11); article 4 of maxillipedal palp unguiform; gnathopods subchelate in both sexes, male gnathopod 2 larger than 1, article 5 not projecting between articles 4 and 6, female gnathopod 2 like gnathopod 1; uropod 3 lacking inner ramus; telson cleft. Species: 48, cosmopolitan, especially tropics, littoral.
Hyalella Smith Hyalella Smith, 1874.—Stebbing, 1906.
Type-species: Amphitoe aztecus Saussure, 1858 (monotypy and subsequent synonymy). See Bulycheva, 1957.
Maxilla 1 with short l-articulate palp; article 4 of maxillipedal palp unguiform; gnathopods of both sexes subchelate, male gnathopod 2 larger than 1, article 5 produced between articles 4 and 6; female
470 U.S. NATIONAL MUSEUM BULLETIN 271
enathopod 2 like gnathopod 1 or minutely chelate; uropod 3 lacking inner ramus; telson entire. Species: 27, primarily S. America, freshwater.
Insula Kunkel
Insula Kunkel, 1910.
Type-species: J. antennulella Kunkel, 1910 (montypy).
Maxilla 1 bearing a palp; article ?4 (Kunkel claims 3 and figures only 3 articles) of maxillipedal palp unguiform; gnathopods subchelate in male (female unknown), relatively alike in size, rather slender (possibly juvenile), article 5 lobed between articles 4 and 6; uropod 3 lacking inner ramus; telson entire. Species: 1, Bermuda, littoral.
Najna Derzhavin
Najna Derzhavin, 1937.—Gurjanova, 1951.
Type-species: N. consiliorum Derzhavin, 1937 (monotypy).
Mandibular molar obsolete; maxilla 1 with vestigial palp; article 4 of maxillipedal palp tiny, short, not unguiform; gnathopods similar in both sexes, subchelate, gnathopod 2 not larger than 1, article 5 lobed between articles 4 and 6; uropod 3 lacking inner ramus; telson entire or slightly notched [probably requires erection of new family]. Species: 1, boreal, N. Pacific, littoral.
[Neobule Haswell]
Neobule Haswell, 1880b.—Stebbing, 1906.
Type-species: N. algicola Haswell, 1880b (monotypy).
A problematical genus, probably not a talitroid; uropod 3 biramous, rami equal in size but small; gnathopods subchelate, gnathopod 2 larger than 1, palms nearly transverse. Species: 1, Australia, littoral.
Orchestia Leach
Orchestia Leach, 1814a.—Stebbing, 1906. Scamballa White, 1847a. Parorchestia Stebbing, 1899a.
Type-species: Oniscus gammarellus Pallas, 1766 (monotypy and subsequent synonymy). See Sars, 1895 (as Orchestia littorea).
Maxilla 1 usually with palp, 2-articulate or absent; article 4 of maxillipedal palp a vestigial bud or absent; gnathopods subchelate in both sexes, male gnathopod 2 much larger than 1, female gnathopod 2 mitten-like; uropod 3 lacking inner ramus; telson entire. Species: 69, cosmopolitan, primarily tropical, terrestrial (beachhoppers).
MARINE GAMMARIDEAN AMPHIPODA A471
Orchestoidea Nicolet
Orchestoidea Nicolet, 1849.—Stebbing, 1906. Talitronus Dana, 1850, 1852a. Megalorchestia Brandt, 1851b.
Type-species: O. tuberculata Nicolet, 1849 (monotypy).
Maxilla 1 with small 2-articulate palp; palp article 4 of maxilliped a vestigial bud or absent; male gnathopod 1 subchelate but often poorly, gnathopod 2 larger than 1, subchelate, article 5 not produced between articles 4 and 6, female gnathopod 1 simple, gnathopod 2 mitten-like; uropod 3 lacking inner ramus; telson entire. Species: 12, primarily E. Pacific, terrestrial (beachhoppers).
Parallorchestes Shoemaker Parallorchestes Shoemaker, 1941a.
Type-species: Allorchestes ochotensis Brandt, 1851c (original desig- nation). See J. L. Barnard, 1962c.
Maxilla 1 with large 2-articulate palp; palp article 4 of maxilliped unguiform; gnathopods subchelate, male gnathopod 2 much larger than 1, article 5 produced between articles 4 and 6, female gnathopod 2 like 1; uropod 3 with small scale-like inner ramus; telson cleft. Species: 1, boreal N. Pacific, littoral.
Parhyale Stebbing
Parhyale Stebbing, 1897; 1906. Hyaloides Schellenberg, 1939.
Type-species: P. fasciger Stebbing, 1897 (monotypy). See Shoe- maker, 1956b.
Maxilla 1 with 1-articulate palp; article 4 of maxillipedal palp unguiform; gnathopods subchelate, male gnathopod 2 larger than 1, article 5 produced between articles 4 and 6, female gnathopod 2 like enathopod 1 and slightly larger; uropod 3 with small scale-like inner ramus; telson cleft. Species: 2, circumtropical, littoral.
Parhyalella Kunkel
Parhyalella Kunkel, 1910.—Schellenberg, 1938. Exhyalella Stebbing, 1917.
Type-species: P. batesoni Kunkel, 1910 (monotypy).
Maxilla 1 lacking palp; article 4 of maxillipedal palp unguiform; gnathopods subchelate, male gnathopod 2 larger than 1, article 1 produced between articles 4 and 6, female gnathopod 2 like male but smaller; uropod 3 uniramous; telson entire. Species: 7, pantropical, freshwater and marine.
472 U.S. NATIONAL MUSEUM BULLETIN 271
Talitrus Latreille
Talitrus Latreille, in Bosc, 1802.—Latreille, 1802.
?(Talitrorchestia) Brandt, 185la (subgenus). Type-species: Orchestza cloqueti Audouin, 1826 (obscure species) .
Talitroides Bonnier, 1898.—Stebbing, 1906. Type-species: 7. bonnieri Stebbing, 1906 (monotypy).
Talitriator Methuen, 1913.
Type-species: Cancer (Gammarus) saltator Montagu, 1808 (new name for Oniscus locusta Pallas, 1766), selected by Boeck (1876). This designation remains questionable. When Bosc (1802), in copying Latreille’s manuscript, erected Talitrus (for which the latter deserves nomenclatural credit in Bose), two species ‘‘Gammarus locusta Fab.” and “Oniscus gammarellus Pallas” were assigned to it (vol. 1 of Bose, p. 78). In vol. 2 of the same work, the assigned species were stated as Talitrus grillus, a new species later attributed to Bose, but which, as indicated in the description, should be attributed to Latreille. By strict adherence to rules, and without either original description, Bose’s second volume or Latreille’s work of a few months later in 1802, we would expect the genus Talitrus to be composed of two species, one of which is precisely the type-species of the earlier genus Gammarus, the other of which later came to be a species of Orchestia. Only by reference to volume 2 do we assume Latreille was referring earlier to the species described by Pallas and not that described by Fabricius. They become homonyms when thrown together in Cancer by other authors. Thus by elimination, now outmoded, Oniscus gammarellus Pallas could yet be selected as the type-species of Talitrus and Orchestia would fall as a synonym to Talitrus, a highly undesirable occurrence at this late day in amphipodan taxonomy. And, G. locusta Fabricius (identical with Cancer locusta Linnaeus) could also be selected. If necessary, the case should be submitted to the ICZN in the hope that they would fix Oniscus locusta Pallas as type-species, and eliminate the ambiguity of Latreille (in Bosc) having used Fabri- clus’ name as author of the species.
Maxilla 1 with small 2-articulate palp; article 3 of maxillipedal palp a vestigial bud; gnathopod 1 of both sexes simple, gnathopod 2 of both sexes small, feebly chelate, mitten-shaped; uropod 3 lacking inner ramus; telson entire or slightly notched. Species: 11, (terrestrial, circumtropical, biboreal).
Talorchestia Dana
(Talorchestia) Dana, 1852b (subgenus).—Stebbing, 1906.
Type-species: Talitrus gracilis Dana, 1852a (selected by Chevreux and Fage, 1925).
MARINE GAMMARIDEAN AMPHIPODA 473
Maxilla 1 with small 2-articulate palp; article 4 of maxillipedal palp a vestigial bud or absent; gnathopod 1 of both sexes simple or poorly subchelate, male gnathopod 2 larger than 1, subchelate, article 5 not produced between articles 4 and 6, female gnathopod 2 small, mitten- shaped; uropod 3 lacking inner ramus; telson entire or slightly notched. Species: 39, circumtropical, amphiboreal terrestrial (some beach- hoppers).
Thaumatelsonidae Fieures 170, 171
Diacnosis.—Accessory flagellum absent or vestigial; mandibular molar evanescent; coxa 1 very small or absent, always partially covered by following coxae; coxa 4 large, shield-like, not excavate posteriorly; uropod 3 uniramous; outer lobes of maxilliped vestigial; telson entire, greatly thickened dorsoventrally; urosomal segments coalesced or partially so; article 2 of pereopod 3 (and 4-5) linear. See Stenothoidae, Cressidae, Pagetinidae, Leucothoidae, Anamixidae, Amphilochidae.
Description.—Rostrum inconspicuous; accessory flagellum absent or vestigial; body smooth or pleon with dorsal carinae; coxa 1 small or absent, hidden by following coxae; coxa 4 large, shield-like, not excavate posteriorly; upper lip probably incised; mandible with evanescent molar, palp present or absent; lower lips unknown; maxilla 1 with 2-articulate palp; maxilla 2 normal; maxilliped slender, outer plates small or vestigial; gnathopods moderately powerful, subchelate or chelate; uropod 3 uniramous, the ramus 2-articulate; telson greatly thickened dorsoventrally; urosomal segments coalesced or partially so, or urosomite 1 often with large dorsal process covering remainder of urosome dorsally.
RELATIONSHIP.—T he Stenothoidae have distinct urosomal segments and unthickened telsons.
The Cressidae have the unthickened telson fused with urosomite 3.
Parathaumatelson Gurjanova (1938) is synonymized with Pseudo- thaumatelson Schellenberg (1931).
Key to the Genera of Thaumatelsonidae
1. Gnathopod 2 chelate (fig. 1717)... ..... +... . Prothaumatelson Cnathopods2;subchelate (igseplaiee yen ne in ere cine fn) eee Ui nae
2. Palp of mandible 1-articulate or absent Pseudothaumatelson (= Parathaumatelson) Palprotimandiblers-articulabe: ys 0 awit) ee ee Thaumatelson
MARINE GAMMARIDEAN AMPHIPODA 475
Genera of Thaumatelsonidae
Prothaumatelson Schellenberg Prothaumatelson Schellenberg, 1931.—Gurjanova, 1938. Type-species: Thaumatelson nasutum Chevreux, 1912a; 1912b (monotypy). Species: 2, bipolar, littoral. Pseudothaumatelson Schellenberg
Pseudothaumatelson Schellenberg, 1931.—Gurjanova, 1938. Parathaumatelson Gurjanova, 1938.
Type-species: P. patagonicum Schellenberg, 1931 (present selec- tion). : Species: 3, subantarctic, littoral.
Ficure 171.—Thaumatelsonidae: Mouthparts, Thaumatelson herdmani Walker (1907): a, mandible; b,c, maxillae 1, 2; d, maxilliped. Prothauwmatelson nasutum (Chevreux, 1912b): e, maxilliped. Gnathopod 1: f, Thawmatelson walkeri Chilton (1912b); g, Pro- thaumatelson carinatum Shoemaker (1955). Gnathopod 2: h, Thaumatelson herdmani; 1, Thaumatelson walkeri; 7, Prothaumatelson carinatum. Uropod 3: k, Prothaumatelson nasutum. Urosome and pleonite 3, urosomites coalesced, bearing 3 uropods and large telson: 1, Thaumatelson walkeri.
285-135 O - 69 - 31
476 U.S. NATIONAL MUSEUM BULLETIN 271
Thaumatelson Walker
Thaumatelson Walker, 1906b, 1907.
Type-species: T. herdmani Walker, 1906b; 1907 (monotypy). See Schellenberg, 1931. Species: 3, subantarctic, littoral.
Vitjazianidae Ficure 172
Diaanosis.—Accessory flagellum long, composed of a few long articles; base of primary flagellum conjoint; gnathopod 1 simple, enathopods feeble, anterior members intersimilar; coxae short. See Synopiidae, Astyridae, Eusiridae, Liljeborgiidae, Hyperiopsidae, Melphidippidae, Argissidae.
Description.—Accessory flagellum long, composed of only a few articles; base of primary flagellum of antenna 1 conjoint, forming an article nearly as long as peduncle; body smooth except for slight carinations on urosome; rostrum slightly developed; coxae very short to medium in length, scarcely touching; upper lip minutely incised; lower lip with or without inner lobes; mouthparts otherwise basic; enathopods feeble, gnathopod 1 simple, gnathopod 2 simple or sub- chelate; uropods biramous; telson short, cleft.
RELATIONSHIP.—The Hyperiopsidae differ from Vitjazianidae by the bent first maxillary palps, the poorly toothed primary cutting edge and small molar of the mandible and the elongate fourth articles of pereopods 1 and 2.
The Eusiridae have an elongate peduncle of antenna 1 and usually large subchelate gnathopods. Liljeborgiidae have large subchelate enathopods.
The Melphidippidae have an elongate uropod 3, and an elongate peduncle of antenna 1.
The Astyridae have a 1-articulate accessory flagellum, a reduced mandibular molar, and gaping lower lip.
The Stilipedidae lack an accessory flagellum, have foliaceous first maxillae, short gnathopods, and no mandibular molar.
—
Ficure 172.—Vitjazianidae: a, Vitjaziana gurjanovae Birstein and Vinogradov (1955); b, Vemana compressa J. L. Barnard (1964a). Lower lip: c, Vemana; d, Vitjaziana. Telson: e, Vitjaziana. Maxilla 1: f, Vitjaziana; g, Vemana. Antenna 1, basal portion: h, Vitjaziana. Maxilliped: 1, Vitjaziana. Uropod 3: 7, Vemana. Mandible: k, Vit- jaziana; 1, Vemana. Upper lip: m, Vitjaziana. Gnathopod 1: n, Vitjaziana; 0, Vemana. Gnathopod 2: p, Vitjaziana; gq, Vemana. Maxilla 2: 7, Vitjaziana.
MARINE GAMMARIDEAN AMPHIPODA
Neveeee
le
Li a
Oo Mie . ve
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A few setae of pereopods 1-5 are elongate in some vitjazianids and the term “fossorial’” might be applied to them but other fossorial families do not have a basally conjoint primary flagellum on antenna 1.
Vitjazianids have definitive similarity to Gammaridae and Eusiri- dae, from which they differ by the very short peduncle and the long conjoint base of the primary flagellum on antenna 1. By evidence of gnathopods and mandible, but not of telson and head the vitjazianids are closely related to synopiids (tironids) and demonstrate again the difficulty in defining families because of the centripetal radiation of evolutionary lines around the Gammaridae.
Key to the Genera of Vitjazianidae
1. Gnathopod 2 subchelate; coxae of medium length; mandibular palp claviform. Vemana Gnathopod 2 simple; coxae minute; mandibular palp linear. . . Vitjaziana
Genera of Vitjazianidae
Vemana J. L. Barnard Vemana J. L. Barnard, 1964a. Type-species: V. compressa J. L. Barnard, 1964a (original designation). Species: 2, Caribbean, bathy-abyssopelagic. Vitjaziana Birstein and Vinogradov Vitjaziana Birstein and Vinogradov, 1955.
Type-species: V. gurjanovae Birstein and Vinogradov, 1955 (origi- nal designation). Species: 1, N. W. Pacific, abyssopelagic.
Incertae Sedis
FiGurRE 173
Didymocheila K. H. Barnard Didymocheila K. H. Barnard, 1931; 1932.
Type-species: D. spongicola K. H. Barnard, 1931 (original desig- nation) ; 1932.
Having all characteristics of Lysianassidae except article 3 of gnathopod 2 not elongate and article 6 strongly elongate, gnathopods
MARINE GAMMARIDEAN AMPHIPODA 479
1 and 2 slender, subequal in size and weakly chelate; mouthparts from lateral view conically grouped, upper lip substyliform, entire, mandible substyliform, cutting edge slightly toothed, molar well developed, ?barrel-shaped or cup-shaped, unknown if triturative, 3-articulate palp attached level with molar; lower lip with Trischizostoma appear- ance; inner plate of maxilla 1 heavily setose medially, palp 2-articulate; maxilla 2 and maxillipeds normal; uropod 3 lacking rami, composed of a simple, ovate peduncle; telson broader than long, entire, apically concave.
With strong resemblance to Sebidae in gnathopods but article 2 of antenna 1 peduncle not elongate, upper lip elongate, mandibular molar and palp strong, uropod 3 lacking any rami, telson very short, etc. Species: 1, S. Georgia Islands, littoral.
i ¢
Figure 173.—Incertae Sedis: a, Sphaerophthalmus grobbeni Spandl (1924). Didymocheila spongicola K. H. Barnard (1932): b, lateral aspect; c, telson ard third uropods; d,e, gnatho- pods 1, 2; f, mandible; g, lower lip.
d
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Metoediceros Schellenberg
Metoediceros Schellenberg, 1931.
Type-species: MM. fuegiensis Schellenberg, 1931 (monotypy).
Head poorly rostrate, not massive; eyes lateral; antennae short, subequal in length, peduncle of antenna 1 slightly longer than head, articles 1-3 successively shorter, male antenna 2 with calceoli, ac- cessory flagellum absent; upper lip not incised; [epistome unknown]; mandible lacking palp, with large triturative molar and toothed incisor; lower lip with inner lobes separate, outer lobes with mandibular extensions; inner plate of maxilla 1 with one terminal seta, outer plate with 10 spines, palp 4-articulate, article 4 unguiform; coxae long as in normal gammaridean, coxa 4 not excavate posteriorly; gnathopods 1-2 weakly subchelate, of medium size, similar to each other, articles 5-6 subequal in length; pereopods generally like those of Oediceros saginatus (see Sars, 1895, pl. 102), thus fossorial, pereopods i-4 with vestigial dactyls; uropods 1-2 normal, apices of rami spinose; uropod 3 reduced to a small simple triangular peduncle; telson nearly circular, entire. Affinities with Dogielinotidae but epistome unknown. Species: 1, Tierra del Fuego, brackish water.
Sphaerophthalmus Spandl Sphaerophthalmus Spandl, 1923; 1924.
Type-species: S. grobbent Spandl, 1923; 1924 (monotypy).
Probably assignable to the Dexaminidae.
Head and thorax very tall, eye forming a large bead; apparently urosomal segments 2-3 coalesced, very small and flat; metasome dorsally carinate; accessory flagellum apparently absent, antenna 1 long, slender, article 2 longer than 1, flagellar articles numerous, antenna 2 “3-articulate”’; maxillipeds apparently with medium sized inner lobes, enormous subclavate outer lobes nearly exceeding the moniliform 4- (?3-) articulate palp, the terminal article of which lacks distinction; other mouthparts not described; gnathopod 1 stout, like that of stegocephalids, gnathopod 2 more slender, elongate, article 6 expanded and bearing transverse palm as in gnathopod 1; pereopods of medium length, second articles of pereopods 3-5 expanded; uropods 1-2 normal, rami interequal, apically spinose, uropod 3 with short peduncle and leaf-lke rami; telson elongate, cleft nearly to base. Species: 1, Red Sea, littoral.
MARINE) GAMMARIDEAN AMPHIPODA A81
Supplement
Recent studies by the writer in Hawai, New Zealand, and Australia suggest the following changes and additions to the classificatory scheme of Gammaridea.
The Eusiridae and Calliopiidae must be amalgamated.
The Aoridae must be amalgamated with the Isaeidae; most genera of the Corophiidae should also be placed in the greater Isaeidae but Corophium and Paracorophium might be retained within a subfamily.
Ceina is to be removed from the Phliantidae to form the type of a new family with close affinities to the Hyalidae; the family will be characterized by the loss of rami on uropod 3 and the rough skin texture; it has affinities with various freshwater chiltonias that may require removal from the Hyalellidae to form a group distinct from the American hyalellas.
Biancolina is to be removed from the Eophliantidae to form the type of a new family. The biramous uropod 3 of Biancolina is one mark of its distinction from Eophliantidae. Amphitholina should also be removed from Eophliantidae and returned to the Ampithoidae as type of a new subfamily.
The freshwater genus Paracalliope is added to the list of marine genera as one of its species, Oedicerus novizealandiae Dana (1853), is definitely intertidal in New Zealand. That species is not a member of Carolobatea as heretofore considered (Chilton, 1909). Stebbing (1906) had already suggested Dana’s species to be identical with P. fluviatilis (Thomson) but the two are definitely distinct.
Paracalliope Stebbing Paracalliope Stebbing, 1899d; 1906.
Type-species: Calliope fluviatilis Thomson, 1879 (original desig- nation).
Eusirid-calliopiid with incipient characters of Oedicerotidae, es- pecially in the disproportionately elongate pereopod 5 with elongate, terminally setose dactyl and elongate uropod 3.
Eyes paired, discontiguous; accessory flagellum vestigial; inner plates of maxillae 1-2 heavily setose medially, maxilla 2 with addi- tional submarginal row of medial setae; female gnathopods weak, those of male stout, with erect lobes on fifth articles, hands twisted mediad, palms bearing large spines; coxa 4 weakly or not postero- dorsally excavate; telson ovoid, thin, uncleft; wrosomites 2-3 coalesced. Species: 5 (2 undescribed), New Zealand, Australia, Philippine Islands, India, marine littoral, brackish and freshwaters.
Literature Cited
[*=Not seen, reference copied from Stebbing, 1888.]
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484 U.S. NATIONAL MUSEUM BULLETIN 271
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1901b. Description d’un crustacé amphipode nouveau de la famille des Stenothoidae (Parametopa Kervillei nov. gen. et sp.) capturé au moyen d’une nasse par M. Henri Gadeau de Kerville, dans la région d’Omonville-la-Rogue (Manche). Bull. Soe. Amis Sci. Nat. Rouen, 1900, pp. 231-237, pl. 3.
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MARINE GAMMARIDEAN AMPHIPODA 487
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Paracyphocaris praedator type d’un nouveau genre de Lysianassidae. Bull. Mus. Océanog., Monaco, no. 32, pp. 1-6, figs. 1-3.
Diagnoses d’amphipodes nouveaux provenant de l’expédition ant- arctique du Francais. Bull. Soc. Zool. France, vol. 30, pp. 159-165. figs. 1-3.
Description d’un amphipode (Katius obesus, nov. gen. et sp.), suivie d’une liste des amphipodes de la tribu des Gammarina remenés par le filet 4 grande ouverture pendant la derniére campagne de la Princesse-Alice en 1904. Bull. Mus. Océanog., Monaco, no. 35, pp. 1-5, figs. 1-7, list on p. 7.
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Diagnoses d’amphipodes nouveaux provenant des campagnes de la Princesse-Alice dans 1|’Atlantique nord. Bull. Inst. Océanog., Monaco, no. 121, pp. 1-15, figs. 1-8.
Diagnoses d’amphipodes nouveaux provenant des campagnes de la Princesse-Alice dans ]’Atlantique nord. Bull. Inst. Océanog., Monaco, no. 129, pp. 1-12, figs. 1-6.
Diagnoses d’amphipodes nouveaux provenant des campagnes de la Princesse-Alice dans ]’Atlantique nord. Bull. Inst. Océanog., Monaco, no. 150, pp. 1-7, figs. 1-3.
Diagnoses d’amphipodes nouveaux provenant des campagnes de la Princesse-Alice dans 1’Atlantique nord. Bull. Inst. Océanog.. Monaco, no. 156, pp. 1-4, figs. 1-2.
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Diagnoses d’amphipodes nouveaux provenant des campagnes de la Princesse-Alice dans l Atlantique nord. Bull. Inst. Océanog., Monaco, no. 204, pp. 1-3, figs. 1—6.
Sur les amphipodes des expédition antarctiques francaises. C.R. Acad. Sci. Paris, vol. 153, pp. 1166-1168.
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Amphipodes: Deuxiéme Expédition Antarctique Francaise (1908- 1910) commandée par le Dr. Jean Charcot. Sci. Nat: Doce. Sci., pp. 79-186, figs. 1-62.
488 U.S. NATIONAL MUSEUM BULLETIN 271
CuEevrevx E.—Continued 1914. Diagnoses d’amphipodes nouveaux provenant des campagnes de la Princesse-Alice dans Atlantique nord. Bull. Inst. Océanog., Monaco, no. 296, pp. 1—4, figs. 1-3. 1919. Note préliminaire sur les amphipodes recueillis par les expéditions du Travailleur et du Talisman (1880-1883). Bull. Mus. Hist. Nat., Paris, 1919, no. 7; 1920, no. 1, pp. 574-580, 7-12. 1920. Sur quelques amphipodes nouveaux ou peu connus provenant des cotes de Bretagne. Bull. Soc. Zool. France, vol. 45, pp. 75-87, figs. 1-9. 1925. Amphipodes I: Gammariens. Voyage de la Goélette Melita aux Canaries et au Sénégal (1889-1890). Bull. Soc. Zool. France, vol. 50, pp. 278-311, figs. 1-12. 1926. Diagnoses d’amphipodes nouveaux provenant des campagnes de la ‘‘Princesse-Alice’’ dans l’Atlantique et dans l’Océan Arctique. Bull. Inst. Océanog., Monaco, no. 475, pp. 1-12, figs. 1-6. 1927. Crustacés amphipodes: Expéd. Sci. ‘‘Travailleur” et du “Talisman” pendant les annés 1880, 1881, 1882, 1883. Malacostracés (suite), vol. 9, pp. 41-152, pls. 1-14. 1935. Amphipodes provenant des campagnes du Prince Albert Ie, de Monaco. Résult. Campagnes Sci. par Albert Iet, Monaco, vol. 90, pp. 1-214, pls. 1-16. CHEVREUX, E., and Bouvisr, E. -L. 1892. Perrierella crassipes, espéce et genre nouveaux d’amphipodes des cétes de France. Bull. Soc. Zool. France, vol. 17, pp. 50—54, fig. 1. CHEVREUX, E., and Faas, L. 1925. Amphipodes. Faune de France, vol. 9, pp. 1-488, figs. 1-438. CHILTON, C. 1883. Further additions to our knowledge of the New Zealand Crustacea. Trans. Proc. New Zealand Inst., vol. 15, pp. 69-86, pls. 1-38. 1884. Additions to the sessile-eyed Crustacea of New Zealand. Trans. Proc. New Zealand Inst., vol. 16, pp. 249-265, pls. 17-21. 1898. A new freshwater amphipod from New Zealand. Ann. Mag. Nat. Hist., ser. 7, vol. 1, pp. 423-427, pl. 18. 1900. A New Zealand species of the amphipodan genus Cyproidia. Ann. Mag. Nat. Hist., ser. 7, vol. 5, pp. 241-246, pl. 5. 1909. The Crustacea of the subantarctic islands of New Zealand. Sub- antarctic Islands of New Zealand, art. 26, pp. 601-671, figs. 1-19. 1912. The Amphipoda of the Scottish National Antarctic Expedition. Trans. Roy. Soc.. Edinburgh, vol. 48, pp. 455-520, pls. 1-2. 1914. A new amphipodan genus and species (family Dexaminidae) from New Zealand. Journ. Linn. Soc. London, Zool., vol. 32, pp. 331-336, pls. 26-27. 1921. Report on the Amphipoda obtained by the F.I.S. “Endeavour” in Australian seas. Biol. Res. Fish. Exper. F.I.S. ‘‘Endeavour,”’ 1909-14, vol. 5, pp. 33-92, figs. 1-16. 1924. Some New Zealand Amphipoda: No. 5. Trans. Proc. New Zealand Inst., vol. 55, pp. 631-637, figs. 1-20. Costa, A. *185la. Fauna del Regno di Napoli. *1851b. Catalogo dei crostacei Italiani e di moltri altri del Mediterraneo per Fr. Gugl. Hope. Napoli, pp. 1-48, pl. 1. *1853a. Fauna del Regno di Napoli.
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*1853b. Descrizione di tre nuovi crostacei del Mediterraneo discoperti dal Rev. G.F. Hope. Fauna del Regno di Napoli, fasc. 83, 10 pp., 3 pls.
*1853c. Relazione sulla memoria del Dottor Achille Costa, di ricerche su’ crostacei anfipodi del regno di Napoli. Rend. Soc. Reale Borbonica, Accad. Sci., Anno 1853, pp. 166-178.
1857. Ricerche sui crostacei anfipodi del regno di Napoli. Mem. Reale Accad. Sci. Napoli, vol. 1, pp. 165-235, pls. 1-4 [two separates seen, one with color-tinted plates].
1867. Saggio della collezione de’crostacei del Mediterraneo del Museo Zoologico della Universita di Napoli inviato alla Esposizione di Parigi del 1867. [Reprint from:] Ann. Mus. Zool. Univ. Napoli, vol. 4, pp. 1-13, pls. 1-2 [see Stebbing, 1888, for underived title and pagination].
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1904. Sur un type nouveau d’amphipode Grandidierella Mahafalensis, provenant de Madagascar. Bull. Soc. Philomath., 1904, vol. 6, pp. 166-174, figs. 1-18.
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1945. Menigratopsis svennilssoni n. gen. et spec., a lysianassid amphipod from the sound. Kungl. Fysiog. Sallskapets I Lund Forhandl., vol. 15, no. 24, pp. 1-7, figs. 1-4.
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1849. Synopsis of the genera of Gammaracea. American Journ. Sci. Arts, ser. 2, vol. 8, pp. 135-140.
1850. A new genus of Orchestidae. American Journ. Sci. Arts, ser. 2, vol 9, p. 185.
1852a. Conspectus crustaceorum quae in orbis terrarum circumnavigatione, Carolo Wikles e classe reipublicae faederatae duce, lexit et descripsit Jacobus D. Dana, Pars III: [Amphipoda. No. I]. Proc. American Acad. Arts Sci., vol. 2, pp. 201-220.
1852b. On the classification of the crustacean Choristopoda or Tetradecapoda. American Journ. Sci. Arts, ser. 2, vol. 14 [appendix], pp. 297-316.
1853. Crustacea, Part II. U.S. Expl. Exped., vol. 14, pp. 689-1618, atlas of 96 pls.
DE.vLA VALLE, A.
1893. Gammarini del Golfo di Napoli. Fauna und Flora des Golfes von
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1927. The Gammaridae of the Kamchatka Expedition 1908-1909. [Russk.]. Gidrobiol. Zhurnal, vol. 6, pp. 1-13, pls. 1-4 [in Russian with English summary].
1937. Talitridae of the Soviet coast of the Japan Sea. Issledovanija Morej SSSR, fase. 23, pp. 87-112, pls. 1-6 [in Russian with English summary].
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Dysowsky, B. N.
1874. Beitrage zur naéheren Kenntniss der in dem Baikal-vorkommenden niederen Krebse aus der Gruppe der Gammariden. Herausgegeben von der russischen entomologischen Gesellschaft zu St. Petersburg. pp. 1-190, pls. 1-14. W. Besobrasoff & Comp., St. Petersburg.
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*1780. Fauna Groenlandica, systematice sistens animalia Groenlandiae occidentalis hactenus indagata, quoad nomen specificum, triviale, vernaculumque; synonyma auctorum plurium, descriptionem, lo- cum, victum, generationem, mores, usum, capturamque singuli, provt detegendi occasio fuit, maximaque parte secundum proprias observationes Othonis Fabricii. Hafniae et Lipsiae.
Gites, G. M.
1885. Natural history notes from H. M.’s Indian Marine Survey Steamer ‘Investigator,’ Commander Alfred Carpenter, R. N., Commanding, No. 2: Description of a new species of the amphipod genus Melita from the Bay of Bengal. Journ. Asiatic Soc. Bengal, vol. 54, pp. 69-71, pl. 3.
1888. Natural History notes from H. M.’s Indian Marine Survey Steamer ‘Investigator’, Commander Alfred Carpenter, R.N., D.S.O., commanding, No. 9: Further notes on the Amphipoda of Indian waters. Journ. Asiatic Soc. Bengal, vol. 57, pp. 220-255, pls. 6-12.
1890. Natural History notes from H.M.’s Indian Marine Survey Steamer ‘Investigator’, Commander Alfred Carpenter, R.N., D.S.O., commanding, No. 15: Descriptions of seven additional new Indian amphipods. Journ. Asiatic Soc. Bengal, vol. 59, pp. 63-74, pl. 2.
Gois, A.
1866. Crustacea Amphipoda maris Spetsbergiam alluentis, cum speciebus aliis arcticis enumerate. Ofversigt Kongl. Vetenskaps-Akad. Forhandl., 1865, pp. 517-536, pls. 36-41 [reprint, pp. 1-20].
GRUBB, A. E.
1861. Hin Ausflug nach Triest und dem Quarnero. Beitrige zur Kenntniss der Thierwelt dieses Gebietes. Berlin, Nicolaische Verlagsbuch- handlung, prefacial pp., pp. 1-175, pls. 1-5 [gammarideans occur on pp. 24, 135-138, 168].
1864a. Beschreibungen einiger Amphipoden der istrischen Fauna. Archiv fir Naturgeschichte, Jahrg. 30, vol. 1, pp. 195-213, pl. 5.
1864b. Die Insel Lussin und ihre Meeresfauna. Nach einem sechswéchent- lichen Aufenthalte geschildert von Dr. Adolph Eduard Grube. F. Hirt, Breslau, pp. 1-116, 1 pl., 1 chart.
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GuERIN, F.-E.
1836. Description de quelques genres nouveaux de crustacés appartenant a la famille des hypérines. Magasin de Zoologie Journal, Ann. 6, Cl. 7, pp. 1-10, + 2 unnumbered pp., pls. 17-18, 19.
GuRJANOVA, E.
1934. Neue Formen von Amphipoden des Karischen Meeres. Zool. Anzeiger, vol. 108, pp. 122-130, figs. 1-6.
1938. Amphipoda, Gammaroidea of Siaukhu Bay and Sudzukhe Bay (Japan Sea). Rep. Japan Sea Hydrobiol. Exped. of the Zool. Inst. of the Acad. Sci. USSR, 1934, pt. 1, pp. 241-404, figs. 1-59 [in Russian with English summary].
1948. Amphipoda Tixogo Okeana II: Stenothoidae dal’nevostochyx morei. Pamjati Akademika Sergeja Alekseevicha Zernova, Akad. Nauk SSSR, pp. 287-325, figs. 1-21 [in Russian].
1951. Bokoplavy morej SSSR i sopredel’nykh vod (Amphipoda-Gam- maridea). Opred. po Faune SSSR, Akad. Nauk SSSR, vol. 41, pp. 1—-1029, figs. 1-705 [in Russian].
1952. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz dal’neve- stochnykh morel. Akad. Nauk SSSR, Trudy Zool. Inst., vol. 12, pp. 171-194, figs. 1-17 [in Russian].
1953. Novye dopolnenija k dal’nevostochnoi fauna morskik bokoplavov. Trudy Zool. Inst., Akad. Nauk SSSR, vol. 13, pp. 216-241, figs. 1-19 [in Russian].
1955. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz severnoi chasti Tixogo Okeana. Trudy Zool. Inst., Akad. Nauk SSSR, vol. 18, pp. 166-218, figs. 1-23 [in Russian].
1958. K fauna amfipod (Amphipoda) Ostrova Makkuori. Biol. Sovet. Antart. Exped., vol. 3, pp. 55-56 [in Russian].
1962. Bokoplavy severnoi chasti Tixogo Okeana (Amphipoda-Gammaridea) chast’ 1. Opred. po Faune SSSR, Akad. Nauk SSSR, vol. 74, pp. 1-440, figs. 1-143 [in Russian].
Hansen, H. J.
1887. Malacostraca marina Groenlandiae occidentalis. Oversigt over det vestlige Grgnlands Fauna af malakostrake Havkrebsdyr. Vid. Medd. Nat. Foren., Kjgbenhavn, 1887, pp. 5-226, pls. 2-7.
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1964. Description d’Uncinotarsus pellucidus nov. gen., nov. sp., nouvel amphipode gammarien mésopsammique de la famille des Aoridae. Bull. Soc. Zool. France, vol. 89, pp. 125-135, figs. 1-4.
HASWELL, W. A.
1880a. Preliminary report on the Australian Amphipoda. Ann. Mag. Nat. Hist., ser. 5, vol. 5, pp. 30-34.
1880b. On Australian Amphipoda. Proc. Linn. Soe. New South Wales, vol. 4, pp. 245-279, pls. 7-12.
1880c. On some additional new genera and species of amphipodous crus- taceans. Proc. Linn. Soc. New South Wales, vol. 4, pp. 319-350 pls. 18-24.
1880d. On some new amphipods from Australia and Tasmania. Proc. Linn. Soc. New South Wales, vol. 5, pp. 97-105, pls. 5-7.
1882. Catalogue of the Australian stalk- and sessile-eyed Crustacea. Australian Mus., Sydney, pp. i-xxiv, 1-324 (plus Addenda et Corregenda), pls. 1—4.
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HasweE.ui, W. A.—Continued
1885. Notes on the Australian Amphipoda. Proc. Linn. Soc. New South
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1875. Die Crustacean, Pycnogoniden und Tunicaten der K.K. Osterr.- Ungar. Nordpol-Expedition. Denkschr. der Kaiserl. Akad. Wissensch., Math.-Nat. K1., vol. 35, pp. 25-46, pls. 1-5.
Herest, J. F. W.
1793. Garneelasseln. Onisci gammarelli. Part 6 in vol. 2 of “Versuch einer Naturgeschichte der Krabben und Krebse nebst einer system- atischen Beschreibung ihrer verschiedenen Arten’’, pp. 105-146 [pls. 34-36, not seen].
HeERpMAN, W. A.
1889. Second annual report of the Liverpool Marine Biological Station on
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*1873. Mémoire sur les crustacés rares ou nouveaux des cétes de France, (Vingt et uniéme article): Description de crustacés nouveaux appartenant 4 la légion des Edriophtalmes, de l’ordre des Amphipodes, de la famille des Piscicoles, de la tribu des Enoplopodes, nobis, du genre des Ichthyomyzoques, nobis. Ann. Sci. Nat., ser. 5, vol. 17, pp. 1-16, pl. 4.
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*1879. Carcinologisches, grésstentheils gearbeitet in der zoologischen Station der niederlindischen zoologischen Gesellschaft. Tijdschrift der Nederland. Dierkund. Vereeniging., vol. 4, pp. 97-161, pls. 5-10.
1889. Crustacea Neerlandica, II: Nieuwe lijst van tot de fauna van Neder- land behoorende Schaaldieren, met bijvoeging van enkele in de Noordzee verder van de Kust waargenomen Soorten. Tijdschrift der Nederland. Dierkund. Vereeniging., ser. 2, vol. 2, pp. 170-234, pls. 7-10.
Houmss, S. J.
1903. Synopses of North-American invertebrates. American Nat., vol. 37, pp. 267-292.
1905. The Amphipoda of southern New England. Bull. U.S. Bureau of Fisheries, vol. 24, pp. 459-529. pls. 1-13, many figs.
1908. The Amphipoda collected by the U.S. Bureau of Fisheries steamer, “‘Albatross,”’ off the west coast of North America, in 1908 and 1904, with descriptions of a new family and several new genera and species. Proc. U.S. Nat. Mus., vol. 35, pp. 489-543, figs. 1-46.
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1954a. Studies on the New Zealand amphipodan fauna No. 2: The family Talitridae: The freshwater genus Chiltonia Stebbing. Trans. Roy. Soc. New Zealand, vol. 81, pp. 563-577, figs. 1-3.
1954b. Studies on the New Zealand amphipodan fauna No. 7: The family Corophiidae, including a new species of Paracorophium. Trans. Roy Soc. New Zealand, vol. 82, pp. 431-460, figs. 1-7.
1954¢c. Studies on the New Zealand amphipodan fauna No. 9: The families Acanthonotozomatidae, Pardaliscidae and Liljeborgiidae. Trans. Roy. Soc. New Zealand, vol. 82, pp. 763-802, figs. 1-14.
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1954d. Studies on the New Zealand amphipodan fauna No. 6: Family Colomastigidae, with descriptions of two new species of Colo- mastiz. Trans. Proc. Roy. Soe. New Zealand, vol. 82, pp. 419- 429, figs. 1-3.
1954e. Studies on the New Zealand amphipodan fauna. No. 10: A new species of Cacao. Trans. Roy. Soc. New Zealand, vol. 82, pp. 803-811, figs. 1-3.
1955. Studies on the New Zealand amphipodan fauna No. 12: The marine families Stegocephalidae and Amphilochidae. Trans. Roy. Soe. New Zealand, vol. 83, pp. 195-221, figs. 1-9.
1958. Austrochiltonia, a new generic name for some Australian freshwater amphipods. Ann. Mag. Nat. Hist., ser. 138, vol. 1, pp. 765-768.
1963. Amphipoda of the family Lysianassidae from the west coast of North and Central America. Allan Hancock Found. Publ., Occ. Pap. no. 25, pp. 1-165, 49 figs.
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1838. Gr6énlands amfipoder beskrevne af Henrik Kroyer. Kongel. Danske Vidensk. Selsk. Naturvid. Math. Afhandl., vol 7, pp. 229-326, pls. 1-4.
1842. Une nordiske Slaegter og Arter af Amfipodernes Orden, henh¢grende til Familien Gammarina. (Forelgbigt Uddrag af et Stgrre Ar- bejde). Naturh. Tidsskr., vol. 4, pp. 141-166.
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1966. A new family Caprogammaridae (Amphipoda, Gammaridea) found in the north-west Pacific. Crustaceana, vol. 10, pp. 192-198, 4 figs.
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*1802. Histoire naturelle, génerale et particuliére des crustacés et des insectes. Ouvrage faisant suite aux oeuvres de Leclerc de Buffon, et partie du cours complet d’histoire naturelle redigé par C.S. Sonnini, vols. 1-4, An X, A. Paris.
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LATREILLE P. A.—Continued 1806. Genera crustaceorum et insectorum secundum ordinem naturalem in familias disposita, iconibus exemplisque plurimis explicata, vol. 1, pp. i-xviii, 1-302 + emendanda, Amand Koenig, Parisiis et Argentorati.
*1825. Encyclopedie Methodique. Histoire Naturelle. Entomologie ou histoire naturelle des crustacés, des arachnides et des insectes, par M. Latreille, vol. 10, par MM. Latreille, le peletier de Saint Fargeau, Serville et Guérin. A. Paris.
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1815. The zoological miscellany; being descriptions of new or interesting
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*1822. In M. W. Mandt, Observationes in historiam naturalem et anato- mian comparatam in itinere Groenlandico factae. Dissertatio inauguralis quam consensu et auctoritae gratiosi medicorum ordinis in universitate literaria berolinensi ut summi in medicina et chirurgia honores rite sibi concedantur die xxii. M. Julii A. MDCCCXXII. H.L.Q.S., publice defendet auctor Martinus Gulielmus Mandt Beyenburgensis.
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*1851. Norges Crustacéer, Crustacea, a clariss, M.W. v. Dueben in Nor- vegia ad Christiansund et Bergen 1843-44 collecta. Ofversigt af Kongl. Vetenskaps-Akad. Férhandl., Arg. 8, no, 1, pp. 19-25.
1855. Ofversigt af de inom Skandinavien hittills funna arterna af slagtet Gammarus Fabr. af V. Liljeborg. Kongl. Vetenskaps-Akad. Handl, for 1853, pp. 443-460.
1856 Om Hafs-Crustaceer vid Kullaberg i Skane. Ofversigt Kongl. Vetenskaps-Akad, Férhandl., vol. 12, pp. 117-1388.
1865. On the Lysianassa magellanica H. Milne Edwards and on the Crus- tacea of the suborder Amphipoda and subfamily Lysianassina found an [sic] the coast of Sweden and Norway. Nova Acta Regiae Soc. Sci. Upsaliensis [Trans. Sci. Soe. Upsala], ser. 3, vol. 6, no. 1, pp. 1-38, pls. 1-5 [reprints of 25 pp. and 38 pp. seen].
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1855. Bidrag till kannedomen om Ostersjons Invertebrat-Fauna. Ofversigt
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Mau, A. W.
1871. Om tv4 for vetenskapen nya Amfipod-species fran Bohuslan, af hvilka det ena ar typ for ett nytt genus inom Pontoporeinernas grupp. Ofversigt Kongl. Vetenskaps-Akad. Férhandl., 1870, [vol. 27], pp. 543-548, pl. 5.
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1890. Crustacea Malacostraca. Det Videnskabelige Udbytte af Kanon- baaden ‘‘Hauchs’’ Togter i de Danske Have Indenfor Skagen i Aarene 1883-86 . . . C.G. Joh. Petersen . . . Copenhagen, pp. 147-232, pls. 1-2, atlas of 14 charts.
METHUEN, P. A.
1913. Description of an amphipod belonging to the family Talitridae, from the Woodbush, Transvaal. Proc. Zool. Soc. London, 1913, pt. 1, pp. 109-112, pls. 10-11.
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1875. Descriptions of new species of Crustacea collected at Kerguelen’s Island by the Rev. A. E. Eaton. Ann. Mag. Nat. Hist., ser. 4, vol. 16, pp. 73-76.
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1830. Extrait de recherches pour servir 4 l/histoire naturelle des crustacés amphipodes. Ann. Sci. Nat., vol. 20, pp. 353-399, pls. 10-11.
1840. Histoire naturelle des crustacés, comprenant l’anatomie, la physio- logie et la classification de ces animaux. Roret, Paris, vol. 3, pp. 1-638, pls. 1-42 for all vols. [@ammaridea, pp. 11-70, pls. 29-30].
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1808. Descriptions of several marine animals found on the south coast of Devonshire. Trans. Linn. Soc. London, vol. 9, pp. 81-114, pls. 2-8.
1813. Descriptions of several new or rare animals, principally marine, discovered on the south coast of Devonshire. Trans. Linn. Soe. London, vol. 11, pp. 1-26, pls. 1-5.
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1933a. Les amphipodes de l’expédition du Siboga, Deuxiéme partie: Les amphipodes gammarides, II: Les amphipodes de la mer profonde. 1: (lLysianassidae, Stegocephalidae, Stenothoidae, Pleustidae, Lepechinellidae). Siboga-Exped., Mon. 38c, pp. 115-167, figs. 35-60.
1933b. Un nouvel amphipode ascidicole. Bull. Inst. Océanog., Monaco, no. 633, pp. 1-6, fig. 1.
1934. Les amphipodes de l’expédition du Siboga, Deuxiéme partie: Les amphipodes gammarides, II: Les amphipodes de la mer profonde. 2: Hyperiopsidae, Pardaliscidae, Astyridae nov. fam., Tironidae, Calliopiidae, Paramphithoidae, Amathillopsidae nov. fam., Eu- siridae, Gammaridae, Aoroidae, Photidae, Ampithoidae, Jassidae. Siboga-Exped., Mon. 33d, pp. 167-235, figs. 61-100.
1936. Les amphipodes de l’expédition du Siboga, Deuxiéme partie: Les amphipodes gammarides, II: Les amphipodes de la mer profonde. 3: Addendum et partie générale. III: Les amphipodes littoraux. 1: Lysianassidae, Ampeliscidae, Leucothoidae, Stenothoidae, Phliantidae, Colomastigidae, Ochlesidae, Liljeborgiidae, Oedi- cerotidae, Synopiidae, Eusiridae, Gammaridae. Siboga-Exped., Mon. 33e, pp. 237-328, figs. 102-146.
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Pirtot J. M.—Continued 1938. Les amphipodes de l’expédition du Siboga, Deuxiéme partie: Les amphipodes gammarides, III: Les amphipodes littoraux. 2: Familles des Dexaminidae, Talitridae, Aoridae, Photidae, Ampitho- idae, Corophiidae, Jassidae, Cheluridae et Podoceridae. Siboga- Exped., Mon. 33f, pp. 329-359, figs. 147-161. OMT Shy breve 1915. The fauna associated with the crinoids of a tropical coral reef: with especial reference to its colour variations. Pap. Dep. Mar. Biol., Carnegie Inst. Washington, vol. 8, pp. 71—96, figs. 1-7, 1 pl. RaTHKE, H. 1837. Zur Fauna der Krym. Ein Beitrag. Mém. Acad. Impér. Sci. Saint-Pétersbourg, vol. 3, pp. 291-454, pls. 1-10. 1843. Beitrage zur Fauna Norwegens. Verhandl. Kaiserl. Leopolinisch- Carolinischen Akad. Naturforsch., Breslau, vol. 20, pt. 1, pp. 1-264, 264b, 264c, pls. 1-12 [Amphipoda, pp. 63-98, pls. 3-4]. Reip, D. M 1940. On Gammarus (Pectenogammarus) planicrurus, subgen. et sp. n. (Crustacea, Amphipoda). Ann. Mag. Nat. Hist., ser. 11, vol. 6, pp. 287-292, fig. 1. RoBeERtTson, D. 1892. A second contribution towards a catalogue of the Amphipoda and Isopoda of the Firth of Clyde and west of Scotland. Proc. Trans. Nat. Hist. Soc. Glasgow, vol. 3, pp. 199-223. Ross, J. C. 1835. Appendix to the narrative of a second voyage in search of a North- West Passage, and of a residence in the Arctic regions during the years 1829, 1830, 1831, 1832, 1833. By Sir John Ross, C.B., K.S.A., K.C.8., &e., &e., Captain in the Royal Navy. Including the reports of Commander, now Captain, James Clark Ross, R.N., F.R.S., F.L.S., &c. and the discovery of the northern magnetic pole. A. W. Webster, London, pp. vii-c, pls. A-C, [Amphipoda, pp. 86-92, partly by R. Owen]. RuFFo, S. 1949. Amphipodes (II): Result. Voy. de la Belgica en 1897-99 . . . . Rapp Sci., Zool., pp. 1-58, figs. 1-18. 1952. Studi sui Crostacei Anfipodi XXXIV Microphotis blachei n. gen. n. sp. (Amphipoda-Photidae) delle acque del fiume Mekong (Cam- bogia). Mem. Mus. Civ. Stor. Nat. Verona, vol. 3, pp. 35-40, figs. 1-3. 1953. Studi sui Crostacei Anfipodi XXXVI Nota critica su Biancolina algicola D. Val. (Amphipoda Prophliantidae). Atti. Accad. Agric., Sci. Lett. Verona, ser. VI, vol. 3, pp. 1-9, figs. 1-2. 1958. Amphipodes terrestres et des caux continentales de Madagascar, des Comores et de la Réunion. Mem. Inst. Sci Madagascar, vol. 12, pp. 35-64, figs. 1-9. 1959. Contributo alla conoscenza degli anfipodi del Mar Rosso (1). Sea Fisheries Res. Sta., Haifa, Bull. 20, pp. 1-36, figs. 1-6. SABINE, E. *1821. An account of the animals seen by the late northern expedition whilst within the arctic circle being no. X., of the Appendix to Capt. Parry’s Voyage of Discovery. pp. 51-57. London. [pls. 1-2]. [See the 1824 edition below, which is similar.]
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1824. Invertebrate Animals. [Marine Invertebrate Animals (per title page) ]. pp. 219-239, pls. 1—2 of ‘‘A supplement to the appendix of Captain Parry’s voyage for the discovery of a North-West Passage, in the years 1819-20. Containing an account of the subjects of natural history.’’ Appendix X. Natural History. John Murray, London.
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1936b. Amphipoda from a South Australia reef, Part I. Ree. So. Australian Mus., vol. 5, pp. 445-455, figs. 1-4.
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1930. The Amphipoda of the Cheticamp Expedition of 1917. Contr. Canadian Biol. Fisheries, n. s., vol. 5, no. 10, pp. 1-141, figs. 1-54.
1933a. Two new genera and six new species of Amphipoda from Tortugas. Pap. Tortugas Lab., Carnegie Inst. Washington, vol. 28, pp. 245- 256, figs. 1-8 (Publ. 435).
1933b. Amphipoda from Florida and the West Indies. American Mus. Novitates, no. 598, pp. 1-24, figs. 1-13.
1935. The Amphipods of Porto Rico and the Virgin Islands. Sci. Survey, of Porto Rico and the Virgin Islands (N.Y. Acad. Sci.), vol. 15, pp. 229-253, figs. 1-5.
1941a. A new genus and a new species of Amphipoda from the Pacific Coast of North America. Proc. Biol. Soc. Washington, vol. 54, pp. 183-186.
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1942. Amphipod crustaceans collected on the presidential cruise of 1938. Smithson. Misc. Coll., vol. 101, no. 11, pp. 1-52, figs. 1-17.
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1945b. The amphipod genus Unciola on the east coast of America. American Midl. Nat., vol. 34, pp. 446-465, figs. 1-9.
1955. Amphipoda collected at the Arctic Laboratory, Office of Naval Research, Point Barrow, Alaska, by G. E. MacGinitie. Smith- sonian Mise. Colls., vol. 128, no. 1, pp. 1-78, figs. 1-20.
1956a. A new genus and two new species of amphipods from Dry Tortugas, Florida. Journ. Washington Acad. Sci., vol. 46, pp. 61-64, figs. 1-2.
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1883. The Challenger Amphipoda. Ann. Mag. Nat. Hist., ser. 5, vol. 11, pp. 203-207.
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1899d. Revision of Amphipoda (continued). Ann. Mag. Nat. Hist., ser. 7, vol. 4, pp. 205-211.
1900. Arctic Crustacea: Bruce collection. Ann. Mag. Nat. Hist., ser. 7, vol. 5, pp. 1-16.
1904. Biscayan plankton collected during a cruise of H.M.S. ‘Research,’ 1900, Part II: The Amphipoda and Cladocera, with notes on a larval thyrostracan. And an appendix on their distribution by G. Herbert Fowler. Trans. Linn. Soc. London, ser. 2, Zool., vol. 10, pp. 13-54, pls. 2-3.
1906. Amphipoda I: Gammaridae. Das Tierreich, vol. 21, pp. 1-806, figs. 1-127.
1907. The fauna of brackish ponds at Port Canning, Lower Bengal, Part V: Definition of a new genus of Amphipoda, and description of the typical species. Rec. Indian Mus., vol. 1, pp. 159-162, pl. 7.
1908a. On two new species of northern Amphipoda. Journ. Linn. Soc. Lon- don, Zool., vol. 30, pp. 191-197, pls. 27-28.
1908b. South African Crustacea (Part IV). Ann. So. African Mus., vol. 6, pp. 1-96, pls. 1-14.
1910. Crustacea, Part 5: Amphipoda. Sci. Res. Trawling Exped. H.M.C.S. “Thetis’’. Australian Mus., Mem. 4, vol. 2, pp. 565-658, pls. 47-60.
1914. Crustacea from the Falkland Islands collected by Mr. Rupert Val- lentin, F.L.S., Part II. Proce. Zool. Soc. London, vol. 1 (1914), pp. 341-378, pls. 1-9.
1917. South-African Talitridae. Ann. Mag. Nat. Hist., ser. 8, vol. 20, p. 435.
1918. Some Crustacea of Natal. Ann. Durban Mus., vol. 2, pp. 47-75, pls. 8-12.
STEBBING, T. R. R., and Rosertson, D.
1891. On four new British Amphipoda. Trans. Zool. Soc. London, vol. 13,
pp. 31-42, pls. 5-6. STEPHENSEN, K.
1915. Isopoda, Tanaidacea, Cumacea, Amphipoda (excl. Hyperiidea). Rep. Danish Oceanog. Expeds. 1908-10 to the Mediterranean and Adjacent Seas 2, Biol., D, vol. 1, pp. 1-53, figs. 1-33.
1923. Crustacea Malacostraca, V: (Amphipoda, I). Danish Ingolf-Exped., vol. 3, no. 8, pp. 1-100, figs. 1-22.
1925a. Crustacea Malacostraca, VI: (Amphipoda, II). Danish Ingolf- Exped., vol. 3, no. 9, pp. 101-178, figs. 23-53.
504 U.S. NATIONAL MUSEUM BULLETIN 271
STEPHENSEN K.—Continued
1925b. Danaella mimonectes (n. gen., n. sp.), a new bathypelagic gammarid (Fam. Lysianassidae) from south Greenland waters. Vid. Medd. Dansk. Nat. Foren., vol. 80, pp. 423-428, figs. 1-3.
1927. Crustacea from the Auckland and Campbell Islands. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. XL. Vid. Medd. Dansk. Nat. Foren., vol. 83, pp. 289-390, figs. 1-33.
1931. Crustacea Malacostraca, VII: (Amphipoda, III). Danish Ingolf- Exped., vol. 3, no. 11, pp. 179-290, figs. 54-81.
1935. Ceinina japonica (N. Gen., N. Sp.), a new aberrant species of the amphipodan family Talitridae from Japan. Trans. Sapporo Nat. Hist. Soc., vol. 13, pp. 63-68, figs. 1-4.
1934. Re-description of Hvyperiopsis vgringi G.-O. Sars. (Crust. Amphip.). Tromsg Mus. Arshefter Naturhist. Avd. Nr. 7, vol. 53, no. 3, pp. 112 figs) 14
1938. The Amphipoda of N. Norway and Spitsbergen with adjacent waters. Troms¢g Mus. Skrifter, vol. 3, pp. 141-278, figs. 20-31.
1940. Marine Amphipoda. Zool. Iceland, vol. 3, no. 26, pp. 1-111, figs. 1-13.
1942. The Amphipoda of N. Norway and Spitsbergen with adjacent waters. Troms¢g Mus. Skrifter, vol. 3, pp. 363-526, figs. 53-78.
1944a. Amphipoda. The Zoology of East Greenland. Medd. Grgnland, Komm. Videns. Undersggel., vol. 121, no. 14, pp. 1-165, figs. 1-18.
1944b. Crustacea Malacostraca, VIII: (Amphipoda IV). Danish Ingolf- Exped., vol. 3, no. 13, pp. 1-51, figs. 1-38, 10 charts.
1949. The Amphipoda of Tristan da Cunha. Res. Norwegian Sci. Exped. to Tristan da Cunha 1937-1938, no. 19, pp. 1-61, figs. 1-23.
STIMPson, W.
1853. Synopsis of the marine Invertebrata of Grand Manan: or the region about the mouth of the Bay of Fundy, New Brunswick. Smith- sonian Contr. Knowledge, vol. 6, pp. i-iv-+ 5-66, pls. 1-3.
1855. Description of some new marine Invertebrata. Proc. Acad. Nat.
Sci., Philadelphia, vol. 7, pp. 385-394. Stout, V. R.
1912. Studies in Laguna Amphipoda. First Ann. Rep. Laguna Mar. Lab., pp. 134-149, figs. 74-84.
1913. Studies in Laguna Amphipoda. Zool. Jahrb., Syst., vol. 34, pp. 633-659, figs. 1-3.
STUXBERG, A.
1880. LEvertebratfaunan i Sibiriens Ishaf. Férelépande Studier grundade pa de zoologiska undersékningarna under Prof. A. E. Nordenskiélds Ishafs-expedition 1878-79. Bihang K. Svenska Vet. Akad. Handl., vol. 5, no. 22, pp. 1-76, 1 chart.
TATTERSALL, W. M.
1906. The marine fauna of the coast of Ireland, Part VIII: Pelagic Amphi- poda of the Irish Atlantic slope. Fisheries, Ireland, Sci. Invest. 1905 (4), pp. 1-39, pls. 1-5.
1922. Amphipoda and Isopoda. The Percy Sladen Trust Expedition to the Abrolhos Islands (Indian Ocean). Journ. Linn. Soc. London, Zool., vol. 35, pp. 1-19, pls. 1-3.
TEMPLETON, R.
1836. Descriptions of some undescribed exotic Crustacea. Trans. Entomol.
Soc. London, vol. 1, pp. 185-198, pls. 20-21 and part of pl. 22.
MARINE GAMMARIDEAN AMPHIPODA 505
Tuomson, G. M. 1879. New Zealand Crustacea, with descriptions of new species. Trans. Proc. New Zealand Inst., vol. 11, pp. 230-248, pl. 10. 1880. New species of Crustacea from New Zealand. Ann. Mag. Nat. Hist., ser. 5, vol. 6, pp. 1-6, pl. 1. 1882. Additions to the crustacean fauna of New Zealand. Trans. Proc. New Zealand Inst., vol. 14, pp. 230-238, pls. 17-18. 1884. Descriptions of new crustaceans. Trans. Proc. New Zealand Inst., vol. 16, pp. 234-240, pls. 12-13. THORSTEINSON, EH. D. 1941. New or noteworthy amphipods from the North Pacific coast. Univ. Washington Publ. Oceanog., vol. 4, pp. 50-96, pls. 1-8. TzvetKova, N. L. 1965. Novyje rod gammarid (Amphipoda, Gammaridae) iz pribrezhnyx uchastkov Japonskogo Morja. Zool. Zhurnal, Akad. Nauk SSSR, vol. 44, pp. 1631-1636, figs. 1-2. VANHOFFEN, E. 1897. Die Fauna und Flora Grénlands. Grénland-Expedition der Gesell- schaft fiir Erkunde zu Berlin 1891-1893, vol. 2, pt. 1, pp. 1-883,
pls. 1-8. WALKER, A. O. 1891a. On Pherusa fucicola (Leach). Ann. Mag. Nat. Hist., ser. 6, vol. 7, pp. 418-422.
1891b. On Pherusa fucicola, Leach [sic]. Ann. Mag. Nat. Hist., ser. 6, vol. 8, pp. 81-83.
1895. Revision of the Amphipoda of the L.M.B.C. district. Proc. Trans. Liverpool Biol. Soc., vol. 9, pp. 287-320, pls. 18-19.
1896. On two new species of Amphipoda Gammarina. Ann. Mag. Nat. Hist., ser. 6, vol. 17, pp. 343-346, pl. 16.
1898. Crustacea collected by W. A. Herdman, F.R.S., in Puget Sound, Pacific coast of North America, September, 1897. Trans. Liver- pool Biol. Soc., vol. 12, pp. 268-287, pls. 15-16.
1903. Amphipoda of the ‘Southern Cross’ Antarctic Expedition. Journ. Linn. Soc. London, vol. 29, pp. 38-64, pls. 7-11.
1904. Report on the Amphipoda collected by Professor Herdman, at Ceylon, in 1902. Suppl. Rep., Ceylon Pearl Oyster Fisheries— 1904, no. 17, pp. 229-300, pls. 1-8.
1906a. Preliminary descriptions of new species of Amphipoda from the ‘Discovery’ Antarctic Expedition, 1902-1904. Ann. Mag. Nat. Hist., ser. 7, vol. 17, pp. 452-458.
1906b. Preliminary descriptions of new species of Amphipoda from the ‘Discovery’ Antarctic Expedition, 1902-1904. Ann. Mag. Nat. Hist., ser. 7, vol. 18, pp. 13-18.
1907. Crustacea, III: Amphipoda. Nat. Antarctic. Exped., British Mu- seum (Nat. Hist.), vol. 3, pp. 1-39, pls. 1-13.
1908. Amphipoda from the Auckland Islands. Ann. Mag. Nat. Hist., ser. 8, vol. 2, pp. 33-39, pl. 5.
1909. Amphipoda Gammaridea from the Indian Ocean, British East Africa, and the Red Sea. Trans. Linn. Soc. London, ser. 2, Zool., vol. 12, pp. 323-344, pls. 42-43.
WALKER, A. O., and Scort, A.
1903. II: Decapod and sessile-eyed crustaceans from Abd-el-Kuri. Nat.
Hist. Sokotra and Abd-el-Kuri, pp. 216-232, pls. 14A, 14B.
506 U.S. NATIONAL MUSEUM BULLETIN 271
Wuitt, A.
1847a. List of the specimens of Crustacea in the collections of the British Museum, pp. i-viii, 1-143. E. Newman, London. 1847b. Descriptions of new or little-known Crustacea in the collection at the British Museum. Ann. Mag. Nat. Hist., ser. 2, vol. 1, pp. 221-228. ZADDACH, HE. G.
1844. Synopseos crustaceorum Prussicorum prodromus. Dissertatio zoo- logica, quam scripsit et ex auctoritate amplssimi philosophorum ordinis in academia Albertina pro facultate docendi adipiscenda
die Xi. M. Decembris ...H. J. Daljowski, Regiomonti, pp. i-viii, 1-39.
Appendix I
Dissection of an Amphipod For Right-handed Operators.
In a flat-bottomed syracuse dish, the amphipod is laid on its left side with its legs projecting away from the operator, so that it appears up-side down through the oculars of the stereoscope. The body is covered to more than twice its depth with alcohol to prevent the effects of surface tension during dissection and to ameliorate the glare of reflected light from projecting legs.
A pair of jeweler’s forceps with very fine points, a fine dissecting needle such as an insect pin mounted on a stick of wood, and a coarse, standard dissecting needle are used. At least five standard glass slides, one depression slide, and six thin cover slips are needed. A small drop of glycerine is placed on two of the flat slides, a tiny drop each on three of the flat slides, and the depression-slide concavity is sparingly filled with glycerine (other media are used for permanent mounts; permanent slides have the disadvantage of restricting the manipulation of mounted parts for 3-dimensional observation; parts from glycerine slides may be stored permanently in alcohol in a tiny vial made of a bit of capillary tubing with one end closed by melting in a fire, the other end stoppered with cotton, pith, or plastic foam).
One commences removing the pereopods (legs) of the amphipod at either the fourth or fifth coxa (sideplate) depending on which of these coxae is largest or would pull away from the body without entangling other legs or coxae. The amphipod is up-side down on its left side, being held with a coarse needle in the left hand through a body segment or with forceps or a blunt stick, and the coxa is being pulled and ripped gently at its base with the fine forceps. In most cases the coxa can be pulled free of the body carrying some of its proximal musculature. Occasionally the firmness of the attachment dictates the use of a fine scalpel.
When the coxa is removed, the remainder of the leg and gill (and if a female the brood lamella) will come with it. As the legs are excised identifying marks are noted in order to record the leg sequence for positioning on the slide. Particularly confusing are coxae 3 and 4 because they are often similar in size and shape, as are the last three pereopods.
507
285-135 O - 69 - 33
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Pereopods and gnathopods are removed to one side of the dish until all seven legs have been collected. Antennae 1 and 2 are dissected at their bases (right side only). Care in removal of antenna 2 at its juncture is needed because it often breaks easily at joint 2 or 3.
The seven coxae-legs and two antennae are removed in a group from the dish of alcohol to the flat slide with the largest drop of glyc- erine. When placed in the glycerine the parts will disperse the drop, but a light breath of air will accelerate evaporation of the alcohol and the amalgamation of the puddle. The legs must be fully immersed in the glycerine to prevent drying and uptake of air bubbles. Do not put on the cover slip.
The right uropods 1, 2, and 3, both lobes of the telson, and one member of each pair of the pleopods are removed and placed on two of the flat slides with tiny drops of glycerine; the parts are manip- ulated, while the glycerine puddles coalesce, and arranged so that their respective dorsal (uropods) and anterior (pleopods) sides are up. A clean cover slip, gripped in the forceps, is lowered horizontally over the glycerine until it can be dropped smartly onto the puddle without engaging air bubbles. Glycerine is to be applied sparingly so as to prevent excessive sliding of the cover slip. If the perimeter of the cover slip lacks glycerine it may be added later by placing a small drop at the edge.
Before removing mouthparts determine whether they are grouped in a coniform or quadratiform bundle from lateral view.
Mouthparts are removed from the head, again with the amphipod head pointing away from the observer so that motion to the right with the forceps can be used to snap off the mouthparts. The maxil- lipeds, which are the most posterior mouthparts, cover all the other mouthparts and must be removed at their base first; both maxillipeds will come off together. More anteriorly, a pair of bilobed second maxillae is to be removed and then the first maxillae, each of which appears to have three lobes (inner lobe, outer lobe, and palp but in a few genera lacking a palp). The inner lobes are difficult to remove in connection with the outer unless special care is taken and caution must be exercised not to damage the lower lip. Mandibles are removed next; they are usually brittle and easily broken; they are most easily removed by rotating them to ascertain the basal muscular attach- ment and snipping this with forceps. Sclerotic connections to upper and lower lips also must be broken to avoid their damage. Usually each mandible will have a palp. After maxilla 1 and the mandibles are removed, a lower lip and an upper lip will remain; the lower lip is extensive and for removal must be grabbed deeply in its muscular and tendon attachments without separating the inner and outer lobes. After practice one may desire to remove lower lips before dissecting
MARINE GAMMARIDEAN AMPHIPODA 009
mandibles as the two mouthparts often are closely connected with tissues and the mandibles will tear the lower lip when being removed.
The upper lip and epistome are not removed from the head at this time. Their interrelationship from lateral view must be pre- served. The observer should note the condition of the ventral margin of the upper lip from anterior view (rounded, incised, truncated) before mounting the carcass on the depression slide.
Mouthparts are transferred to a tiny drop of glycerine on a flat slide, arranged in sequence and fitted with a cover slip. Preferably the mouthparts are arranged so that the following parts project upward or are on top: mandibular molars and the inner lobes of lower lip, maxillae, and maxillipeds. If the base of the maxillipeds curves upward, it may be cut off so that the cover slip will set firmly. The mandibles should be arranged with the molars projecting obliquely toward the observer or directly lateral, if the center of gravity so permits. Often mandibles are placed on a separate slide with supports for the cover slip to prevent crushing. Supports may be made of wire or sand grains.
Because a unilateral dissection has been made, the remaining amphipod carcass has a complete set of pereonal and pleonal parts remaining on one side (the left if done by a right-handed operator). Coxa 1 and any other (left) legs are removed which would obscure the head and pleon from lateral view. The carcass is mounted right side down in the glycerine of the depression slide and a cover slip firmly set. If the amphipod is so large that it will be crushed by the cover slip or lie in a tilted position, two pieces of wire of appro- priate thickness (or variously thick insect needles, pins, paper clips cut with nipping pliers) are placed on each side of the amphipod, which is covered with sufficient glycerine to fill the area between the two wires, and the cover slip set on the supports. The top glass should fit the carcass snugly so as to hold it in place but not to crush it. Glycerine has sufficient surface tension so that it will not leak _out from under the elevated coverslip as long as the slide is kept in a horizontal position.
One now returns to the first flat slide on which were placed the seven right pereopods (including gnathopods) and the two right antennae. They are arranged in order from anterior to posterior in two rows. Gills from legs 2 to 7 (or 2-6 or 2-5) are removed and placed in sequence on the fourth flat slide of glycerine. If the animal is a female, the brood lamellae are removed in sequence and placed in a row on another slide. At this stage one is working from glycerine to glycerine without the effects of a change in surface tension so that it is easy to keep the parts in order, making notes of charac- teristics that will permit proper orientation. Cover slips are set in place.
510 U.S. NATIONAL MUSEUM BULLETIN 271
If the amphipod species is dimorphic, a slide of antennae, gnath- opods, and uropod 3 of the other sex should be prepared for rapid identification; but a full dissection should be prepared for descriptive purposes.
Parts of greatly differing thickness should not be intermingled on the same slide as the thinner parts will not be properly fitted by the cover slip. Delicate parts may need artificial support of the cover slip as noted above in discussion of the mandibles. Dirt may be removed from heavily setose appendages by use of a fine camel’s- hair brush.
Eventually the student will gain sufficient experience for examination of most parts without dissection. Even mouthparts can be partially to fully examined by careful manipulation under a fine stereoscope with adequate 2-directional light sources. Mandibles often can be rotated for viewing molars without their complete removal. This protects unique specimens from unnecessary damage or loss of parts, or the need to mount parts permanently.
The taxonomist anticipating a need to illustrate the organism will leave the telson and the left first coxa attached to the carcass so that a full lateral view of the amphipod is preserved. The telson can be removed for flat mounting after the lateral view is drawn. Usually the left legs distal to their coxae are removed and mounted. The lateral in toto drawing represents a composite reconstruction of body and coxae drawn first, with legs superimposed on the drawing by use of a microprojector or camera lucida in which degree of magnification can be replicated. In this way legs are attached to the body drawing in perfectly flat but somewhat unnatural condition. One must deter- mine accurately the attachment loci of the legs to their coxae by study of the proposed slides 5 and 6 noted below. Generally, it is preferable to make slides of the following composition if illustrations are to be made.
1. Depression slide with carcass.
2. Mandible, maxilliped, lower lip, with support for cover slip.
3. Maxillae, 1, 2.
4. Antennae 1, 2, with support for cover slip to allow rotation and
examination of all surfaces by movement of cover slip.
5. Gnathopod 2, if especially thick.
6. Gnathopod 1 and pereopods 1-5.
7. Pleopods 1-3.
8. Uropods 1-3, with support for cover slip to allow rotation of parts. 9. Telson (removed from carcass after lateral illustration).
10. Upper lip and epistome (removed from carcass after lateral
illustration). 11. Left legs, except for their coxae.
MARINE GAMMARIDEAN AMPHIPODA 511
12. Gills, with support for cover slip. 13. Brood lamellae.
Very frequently, preserved amphipods have broken appendages. Sometimes the loss of uropod 3 is a consistent occurrence especially in gammarids and oedicerotids. So few Gammaridea lack a third uropod that the first assumption always should be that the part has been removed accidently and close examination should be made for sockets and musculature indicating the loss.
Antennae are often broken and such specimens should be avoided until experience is sufficient to recognize amphipods by other means. In the photid-corophiid complex, legs (except gnathopods) and antennae are frequently autotomized when the animals are preserved, and specialists usually have found other means of identification in those families.
The ecologist making a study of a single species should be prepared to take special care in preservation of his material to ensure com- pleteness of the specimens. He may find slow dilution of seawater or special anesthetics suitable to kill the organisms slowly and to prevent autotomy.
Appendix II
Amphipod Analytical Sheet A Checklist of Characters to be Examined Before Using Keys
Before the keys are utilized for the identification of an amphipod, the morphological characteristics of the organism should be deter- mined. This requires a complete dissection, mounting, and examina- tion of parts on microscope slides as discussed in Appendix I. The following checklist may be useful as a guide to characters that should be determined. This procedure is very laborious at first but pays dividends in the final result and makes familiar, more rapidly, those characters which ultimately are most important. Principal characters are italicized.
The checklist is designed to indicate the characters present by circling the pertinent items in the sentences, or adding percentages to blanks or zeros to the ends of inappropriate sentences. Descriptive phrases applied to various shapes are minimal in number; the observer may have to compose further refinements. But descriptive refine- ments can be overextended to the unnecessary wastage of time, for this checklist applies to initial analysis of the generic position and not to fine details necessary for specific identity.
One may find it useful to replicate copies of this checklist for the keeping of records.
BODY
(Pereon and Pleon [metasome and urosome))
General, segments: Abnormal disproportion or enlargement (example, Danaella). Cylindricalization (examples, Colomastigidae, Eophliantidae). Dorsal depression with or without splaying of coxae. Ornamentation: Teeth, dorsal and lateral; pereon 1, 2, 3, 4, 5, 6, 7; pleon 1, 2, 3, 4, 5, 6. Spine groups, dorsal, especially on pleon 1, 2, 3, 4, 5, 6. Elongation of metasome: example, is it as long as 5 or 6 pereonites? (for dis- tinguishing a few Gammaridae and Eusiridae). Urosome: Coalescence of segments: 1-2, 1-3, 2-3 (examples: Ampeliscidae, Kuriidae Podoceridae). Dorsal depression (examples, Corophiidae, Cheluridae). Elongation of wrosomite 1 (example, Podoceridae). Pleonal epimera: Shape and ornamentation from lateral view, especially of epimeron 3 (primarily for identification to species). 512
MARIND GAMMARIDEAN AMPHIPODA 013
COXAE Size:
Normal: like basic gammaridean; 1-4 of medium length, subquadrate, 5-7 shorter, successively smaller, 5-6 slightly lobed.
Elongation (example, Stegocephalidae).
Reduction (example, Podoceridae).
Disproportional sizes of 1-4: 1 long, 2 shorter, 3 shorter, 4 long (example, Argissidae).
Coza 1 absent or vestigial (rare examples, Bateidae, Anamizis).
Coxa 1 reduced in size and partially to fully covered by following coxae (example, Stenothoidae). (Caution: keep amphipod specimen in unflexed condition for proper observation.)
Coxae 1-2 or 1-3 reduced in size and covered by following coxae (primarily genera of Lysianassidae).
Coxa 3 larger than 4 (example, Synopiidae).
Coxa 5 as long as 4 (rare).
Position:
Serial contiguity: coxae contiguous or overlapping.
Serial discontiguity (example, Podoceridae, Eophliantidae).
Concealment of one coxa by another (see above).
Lateral splaying (Phliantidae).
Shape:
Coxae 1-4 subquadrate.
Acumination of coxae 1, 2, 3, 4.
Excavation posteriorly of coxa 4: posterior margin straight, concave or deeply excavate and/or bearing posterior lobe (contrast Stegocephalidae and Isaeidae).
Coxa 1 tapered, expanded, oval, semicircular, quadrate, conical, acutely lobed.
Ventral serrations.
Special patterns: Disproportional sizes of 1-4 (Argissidae, see above). Crescentic curve formed ventrally by coxae 4-5 together (example Epimeria).
HEAD Size:
Length as a function of one or more pereonites (1, 1-2, 1-3), head measured on horizontal axis from front of lateral lobe to perpendicular line from posterior cephalic extent.
Massive (see definition in glossary).
Shape:
Normal gammaridean: head cuboidal, with lateral lobes.
Globular: subspheroid; neck cylindrical.
Galeate (see definition in glossary).
Rostrum: Length in relation to head ( %) and to article 1 of antennal ( %). Shape: acute, spatulate; horizontal, deflexed.
Lateral lobe:
Shape and extent of projection.
Notch or ornamentation.
Ocular bulge on side of head.
Marginal details of anteroventral corner of head near insertion of antenna 2.
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EYES Composition: Presence or absence (careful examination required because eyes often lose pigment in preservative). Paired ommatidial mass below cephalic cuticle (common). Cuticular lenses in lateral pairs (Ampeliscidae almost exclusively). Occasionally lenses occupy anterior surface of head. Diffused pigment or stain. Quadrigeminous lenticular bodies (example, Argissidae). Bright pigmentary masses enveloping brain (especially Ampeliscidae). Shape: ovoid, flask-like, reniform. Position: Near lateral cephalic surface. In lateral lobes. In rostrum (especially Oedicerotidae). Dorsally confluent (especially Oedicerotidae). Accessory detached ommatidia (pattern often confused by preservational accidents). Occupying cephalic extent almost fully (Hyperiopsidae).
ANTENNA 1
Length: As percent of total body (front of head to base of telson ( %). In relation to antenna 2 and to its peduncle ( %). Flagellum: Proportion to peduncle ( %). Elongation of basal article (often conjoint, composed of incompletely seg- mented articles). Proportion to peduncular article 3, especially when latter elongated ( %). Number of articles: 0, 1, 2, 3, 4, 5, 6-10, 11-15, 16++. Peduncle: Proportion to head. Relative lengths of all three articles (value of 100 assigned to length of article 1): 1=100%; 2=—( %);3=( %). Ornamentation on any article, all sides. Distinctive spines or setal bundles. Possible geniculation between articles (example, several Haustoriidae). Accessory flagellum (attached to article 3 of peduncle, medial): Number of articles: 0, 1, 2, 3, 4, 5-10, 11+. A fused scale. Special shape. Elongation of basal article. Accessory organs: Calceoli. Aesthetases, if especially enlarged or in dense bundles.
ANTENNA 2
Length as percent of body length (including head) ( %). Peduncle: proportion to head ( %). Relative proportions of articles: 4=100%;5=( %). Tumidity of articles 3, 4, or 5; article 1 large and subspherical. Gland cone and/or ensiform process on articles 2 and 1; extreme enlargement and shape. (Ensiform process generically important primarily in Phoxo- cephalidae. Gland cone often medial and hidden from lateral view.)
MARINE GAMMARIDEAN AMPHIPODA 515
Flagellum: proportion to peduncle ( %) or article5( %). Number of articles: 0, 1, 2, 3, 4, 5, 6-10, 11-15, 16+. Ornaments: Aesthetases, calceoli on peduncle and/or flagellum. Distinctive spine groups. “Fossorial’’ condition, with long plumose setae.
MOUTHPARTS
From lateral view forming a conical bundle below head (rare) or a quadrate bundle (common). To be examined before dissection. Amalgamation of mandibles and mazillae into ventral keel (Anamixidae).
EPISTOMAL-LABRAL COMPLEX (prebuccal)
Lateral view (primarily of generic importance in Lysianassidae) :
Epistome and labrum separated by notch (common) or coalesced.
Epistome formed as lobe dominating labrum, vice versa, or produced together. Shape of lobes:
Epistome: flat, rounded, acute.
Labrum: flat, rounded, acute. Prebueccal mass inconspicuous and of normal gammaridean proportions.
UPPER LIP (anterior view)
Ventral margin: rounded, truncate, incised, lobed asymmetrically or symmetri- cally.
MANDIBLE
Shape and size of body: bulky (Synopiidae), styliform (Acanthonotozomatidae) , elytriform (Stegocephalidae, Pardaliscidae), OR normal (Gammaridae). Incisor: normal, extremely broadened; needle-like, toothed, untoothed, teeth separated by flat margin. Lacinia mobilis, if present on either right or left mandible: toothed, special shape such as vermiform. Spines proximal to lacinia mobilis: 1-2; 3-6; 7+. Molar: Absent. Size: small (fig. 1267), medium (fig. 126e), large (fig. 126g), fully dominating mandible (fig. 166d). Shape: cylindrical, cuboidal, laminate, conical, tuberous. - Texture: triturative (rasp-like), spinose (spines articulate), setulose, minutely fuzzy, striate, smooth. Accessory seta or spine on triturative molar. Palp: Number of articles: 0, 1, 2, 3. Attachment position relative to molar: over (level with), distal to, proximal to (variation from “‘level”’ primarily in Lysianassidae). Relative length of articles: 1=( %);2=100%; 3=( %). Shape: Article 8: cylindrical, falconiform, tuberculiform. Article 2: occasionally curved strongly. Article 1: rarely with distal cusp. Setation: article 3: distal only, medial. Disymmetry of right and left members (especially lacinia mobilis, incisor, spine- row).
516 U.S. NATIONAL MUSEUM BULLETIN 271
LOWER LIP
Normal gammaridean (fig. 2).
Inner lobes: weak (fig. 71d), absent.
Mandibular projection of outer lobes: pointed, obtuse, absent.
Outer lobes: distally notched; medially excavate (Ampithoidae, figs. 50c,d). Special shapes: Pleustidae (fig. 37a), Trischizostoma (fig. 125v).
MAXILLA 1 Inner plate: Size: absent (very rare), small (fig. 127f), medium (fig. 127e), as large (broad and bulky) as outer plate (rare). Setation: terminal, medial, or both. Number of setae: 1, 2, 3, 4-6, 6-12. Structure of setae: normal; sickle-shaped or strongly constricted. Outer plate: Number of spines: 1-4, 5-6, 7-8, 9-11+. Shape of spines: normally slender, extremely stout, some bifid, some serrate, in two distinct groups by position or structure. Palp: Number of articles: 0, 1, 2. Normal structure: article 1 short, article 2 long. Article 1 long, article 2 short. Modifications: strongly bent (geniculate), foliaceows, bearing scales (examples: Stilipedidae, H yperiopsidae).
MAXILLA 2
Normal gammaridean (fig. 1270).
Abnormally small, plates partially coalesced, setae very sparse. Breadth of lobes: subequal, inner broader, outer broader.
Axial divergence of lobes (fig. 127)). |
Extension of outer plate on basal article (fig. 161a).
Specialized spines (rare).
Extent of medial setation on inner plate: strong, sparse, absent.
MAXILLIPED Inner lobes (proximal): Size: vestigial (fig. 128e), normal (fig. 128a). Abnormal shape: foliaceous (rare), styliform (note: inner lobes often appear styliform if not fully depressed by cover slip). Outer lobes: Size relative to inner: usually larger, vestigial (fig. 114d), foliaceous (rare). Spination: absent, medial, distal. Palp: Extension in relation to outer plate: shorter, equal, longer. Number of articles: 0, 2, 3, 4. Medial or terminal extensions of articles, e.g., lobes, cusps: articles 1, 2, 3. Elongation of articles 1, 2, 3. © Terminal palp article (usually 4): claw-like (normal); barrel-shaped; vestigial; bearing distal nail, spine, or setae.
MARINE GAMMARIDEAN AMPHIPODA 517
GNATHOPODS
Judgment of gnathopods 1-2 as: feeble together (fig. 165a), normal (basic gam- maridean with gnathopod 2 powerful), powerful together (fig. 115a).
GNATHOPOD 1 (excluding coxa 1)
Present, vestigial, or absent (Bateidae and Paranamizis only). Size (or length) relative to gnathopod 2: smaller, equal, larger. Sexual dimorphism: similar or different in male and female. Articles: 2: Length in relation to coxa 1 (rarely important except when coxa | abnormal). 3: Length normal; or elongate (like fig. 3a). 4: Merochelation: with strong thumb-like extension (rare). 5: Length relative to article6( %). Posterior lobe: present, weak, absent. Carpochelation: with strong distoposterior tooth or teeth forming thumb or guarding article 6. 6: Breadth: relative to article 5; wider, equally wide, narrower. Shape: ovate, pyriform, quadrangular, rectangular, linearly rectangular, styliform. Palm: Present, absent, undecided. Slope: transverse, oblique: slight; moderate; extreme. Chela if present: parachelate (describe if strongly chelate). Definition of proximoposterior corner of palm: spines, protuberance, tooth, change in slope only. Ornamentation: special spines, teeth. 7: Fit of the dactyl to palm: congruent, overlapping, not fitting. Shape and ornaments: claw-like (normal); vestigial; absent (rare); with special setae or spines; hidden in setae or cirri; flagelliform. Distal articles especially scaly or with small stiff setae (Lysianassidae).
GNATHOPOD 2 Articles: 3: Length normal; elongate (fig. 3a). 4: Merochelation: with strong thumb-like extension. 5: Length relative to article6( %). Posterior lobe: present, weak, absent. Carpochelation: with strong distoposterior tooth or teeth forming thumb or guarding article 6. Seales or stiff setae (pineapple cushion of Lysianassidae). General shape: cup-like, elongate rectangle, ovate. 6: Breadth relative to article 5: wider, equal, narrower. Shape: ovate, pyriform, rectangular, linearly rectangular, mitten-like (Lysi- anassidae, Talitroidea), quadrangular. Palm: Present, absent, undecided. Slope: transverse; oblique: slight, moderate, extreme. Chela if present: parachelate (describe if strongly chelate). Definition of proximoposterior corner of palm: spines, protuberance, tooth, change in slope only. Ornamentation: special spines, teeth.
518 U.S. NATIONAL MUSEUM BULLETIN 271
7: Fit of this dactyl to palm: congruent, overlapping, not fitting. Shape and ornaments: claw-like (normal); vestigial; absent (rare); with special setae, spines, or processes.
Sexual dimorphism in female: gnathopod 2 like male but much smaller, like gnath-
opod 1 and of similar size, like gnathopod 1 but larger.
PEREOPODS 1-2
Internal glands present or absent.
Orientation of pereopod 2 like that of pereopod 3 (Hohaustorius). Chelate or prehensile.
Articles 4—5, 4-6, or 4, 5, 6 inflated strongly (rare).
Article 4 extraordinarily elongate (Ampeliscidae, Hyperiopsidae). Special spines on article 6 near claw: spines striate, hooked. Article 7 absent (Haustoriidae).
PEREOPODS 3-5
Relative lengths: pereopod3( %);4=100%;5=( %).
General stucture: All similar in structure and slightly longer successively (normal). Article 2 ecpanded: pereopod3( ),4( ),5(_). Expanded lobe of article 2 of pereopod 5 different from pereopods 3 and 4. Chelate, subchelate, or prehensile: pereopods 38, 4, 5. Fossorial setation (see glossary) present, absent. Article 7 absent (Haustoriidae, Stegocephalidae): pereopds 3, 4, 5. Pereopod 5 reduced to fewer than 6 articles.
PLEOPODS
(Rarely significant in marine Gammaridea [but see Phliantidae especially]). Relative length (size) of each pair: 1=100%;2=( %);3=( %). Length of longest ramus relative to peduncle ( %). Length of inner ramus to outer ( %) (note absence of rami or low number of articles). Shape of coupling hooks on peduncles. Lobation of peduncles. UROPODS 1 and 2
Absence (rare) or presence.
Projection along following uropods. Uropod 1 reaching ( %) along uropod 2;( %) along uropod 3. Uropod 2 reaching ( %) along uropod 3.
Relative length of rami: outer or inner shortened (occasional), inner absent or vestigial (rare).
Spination density of peduncle and rami (usually of specific value only).
Incision of inner ramus (example Anonyz).
UROPOD 3 Absence (rare) or presence. Rami absent (rare). Length relative to other uropods; extension beyond longest of other uropods ( % of its own length).
MARINE GAMMARIDEAN AMPHIPODA 519
Length of peduncle relative to wrosomal segment 3 ( %), to peduncles of other uropods ( % of peduncle of uropod 1), or to telson ( %).
Length of longest ramus relative to peduncle ( %).
Length of inner ramus to outer ( %).
Shape of rami: styliform, lanceolate, barrel-shaped, foliaceous.
Articles of outer ramus (1 or 2).
Minute ornamentation and hooks on rami (especially Ischyroceridae and Am- pithoidae).
Special peduncular processes.
TELSON
Absence (rare) or presence. Fused to urosomite 3 (rare). General shape and length (make sketch). Length in relation to urosomite 3( %) or uropod3( %). Degree of cleft between lobes ( %), emarginate only. Ornamentation: apically pointed, notched, trifid, truncate, rounded, concave. Greatly enlarged and with ventral keel (rare), forming dorsoventral plate. Dorsoventrally thickened (‘‘fleshy’’) (example, Isaeidae), bearing lateral nobs, scales, hooks. SEXUAL DIMORPHISM
(primarily for identification to species)
Especially: Antennae: Eyes: Gnathopods: Coxae: Pleonal epimera 1-3: Uropod 3: Urosomal teeth:
Appendix III Glossary
accessory flagellum. Thesecondary ramus of antenna 1, often absent or vestigial (fig. 1), and attached medially to peduncular article 3.
aesthetasc, aesthete. Sensory setae of antennae, flattened and nontapering.
article. The segment of an appendage (fig. 1).
calceolus. A small globular or helmet-shaped, articulate sense organ on the antennae; presumably a modified aesthetasc; of rare occurrence in Gammaridea and most often seen in Eusiridae (fig. 89)).
carpochelate. Immovable finger of prehensile appendage occurring on carpus (article 5); examples: Lewcothoe, Microdeutopus.
chela. Immovable finger of prehensile appendage.
chelate. Descriptive of the palm of a gnathopod protruding as an immovable finger on which the dactyl closes (fig. 129h). See parachelate, carpochelate, propodochelate, merochelate, complexly chelate.
claw, claw-like. Descriptive of a talon or simple, tapering nail. [Not descriptive of chelae as used in decapod terminology.]
compressed. Flattened from side to side.
conjoint. Describing the basal amalgamation of flagellar articles on antennae.
corneal lens. A biconvex cuticular body occurring directly in or on the chitinous cephalic surface in Ampeliscidae; contrasted with subcuticular ommatidia.
coxa, coxal plate. [Terms used synonymously herein.] Article 1 of a pereonal appendage, expanded into a lateral lamella (fig. 1). [Terms for other articles of the appendages such as basis, ischium, merus carpus, propodus, and dactyl are frequently but not universally used in Gammaridea; instead, the articles are simply numbered. ]
dactyl. Talon-like terminal article of pereopods (article 7) or maxillipeds (articles 3 or 4).
depressed. Flattened dorsoventrally.
emarginate. Descriptive of the concave posterior end of an uncleft telson (fig. 42¢e).
entire. Descriptive of an uncleft telson (fig. 38f).
epimeron. A lateral pleuron of pleonites 1-3; the ventrolateral plate-like extension of the body segment (fig. 1).
epistome. The anterior surface of the head above the labrum; this area is often extended ventrally to appear as a part of the labrum and may be anteriorly produced as a cusp or lobe (fig. 4a).
flagellum. The distal portion of either antenna 1 or 2; on antenna 1 it commences with article 4, on antenna 2 with article 6; because basal peduncular articles of antenna 2 are often difficult to resolve, the juncture may be recognized between the elongated final peduncular article and the shortened first flagellar article which is followed by similar short articles; on antenna 1, however, article 1 of the flagellum is occasionally elongate and apparently composed of non- segregated (thus conjoint) daughter articles (fig. 1).
520
MARINE GAMMARIDEAN AMPHIPODA 521
fossorial. Associated with the habit of burrowing, often referring to the exces- sively spinose or setose condition of appendages used for burrowing by Gam- maridea; especially applicable to Haustoriidae, Oedicerotidae, Phoxocephalidae with some setae of articles 4-6 of pereopods 3-5 more than half as long as those articles; and some spines in groups forming submarginal rows perpendicu- lar to margins; long setae also occur on “‘filter’’ feeders such as Ampeliscidae.
galeate. Descriptive of the helmet-shaped heads of various oedicerotids and synopiids.
gamopod. A gnathopod; referring to the use of gnathopods for grasping members of the opposite sex during amplexus.
geniculate. Permanently bent, usually in reference to the flexeda ntennae of some haustoriids, or the outer lobes of maxilla 2 in some stegocephalids in which the bend occurs between articles; or applicable to bent palps of maxilla in Hyperiop- sidae in which the bend occurs on one article.
gnathopod. One member of the first two pairs of free thoracic appendages; these appendages differ in function and usually in appearance from following pereo- pods; often called pereopods.
incisor. The apical portion of the mandible usually formed into a toothed chewing edge or untoothed chopping plate.
joint. The juncture between two articles of an appendage.
labrum. (See upper lip.) ’
lacinia mobilis. An articulated accessory plate proximal to the mandibular incisor, often absent or missing on either left or right mandibles, occasionally indistinguishable from a spine of the spine-row.
low lip (labium). A fleshy complex posterior to the mandibles, always composed of at least one pair of lobes (outer), often with a medioproximal pair of inner lobes; the lateroproximal ends of the outer lobes are often attentuated as alae and are denoted as mandibular lobes (fig. 2g).
mandible. The anterior movable appendage of the buccal group; usually composed of a body bearing a distal incisor, a lacinia mobilis, spine row, molar, and d-artculate palp (fig. 2f).
massive. A term applied to the heads of Synopiidae and Oedicerotidae; head as long as pereonites 1-3 combined and as tall as or taller than long (length not including rostrum). Heads of Ampeliscidae and Phoxocephalidae are elongate but not massive; heads of Acanthonotozomatidae are as tall as long but are not as long as pereonites 1-3 combined.
maxilla 1. A pair of cephalic appendages posterior to the lower lip; for taxonomic purposes only three portions of each member are named: the medial lobe (plate) usually bearing marginal setae, the lateral and larger lobe (plate) bearing terminal spines, and, attached to the outer lobe, a palp usually com- posed of two articles but occasionally absent (fig. 29).
maxilla 2. A pair of cephalic appendages posterior to maxilla 1; for taxonomic purposes each member recognized as a pair of lobes (plates) medial and lateral, usually strongly setose (fig. 272).
maxillipeds. The posteriormost pair of ‘‘cephalic’’ appendages, representing the primitive first thoracic segment now amalgamated with the head but in amphipod taxonomy not included in the sequential numbering of thoracic appendages; for taxonomic purposes recognized as a pair of basally amalga- mated appendages, each member composed of a proximal (inner) plate, a distal (outer) plate, and a palp of four articles, rarely reduced to 3 or 2 articles or absent (fig. 21).
merochelate. Immovable finger of prehensile appendage occurring on merus (article 4); example: gnathopod 1 of Aora.
22 U.S. NATIONAL MUSEUM BULLETIN 271
mesosome. The pereon or thorax. [Term rarely used.]
metasome. Pleonites 1-3. [Term rarely used.]
molar. A process of the mandible, located on the midmedial margin; when completely developed it is a large, massive, subcylindrical body with a surface of ridges and teeth used for grinding (triturative) (fig. 2f).
ommatidium (singular), ommatidia (plural), ommatidial (adjective). Terms applying to the parts of the subintegumentary compound eye, not to be confused with the corneal lenses of the integument of Ampeliscidae.
palm. A posterior surface or margin of article 6 of a gnathopod or pereopod on which article 7 (dactyl) closes for the purpose of prehension; usually recog- nizable because of expansion of article 6 or by occurrence of special spines or ornamentation and usually with a proximal defining limit marked by a change in marginal slope or occurrence of special spines (fig. 2e).
palp. Terminal articles of a buccal appendage, in Amphipoda occurring only on mandibles, first maxillae, and maxillipeds as the stenopodous terminal articles distal to the expanded outer plates or main body.
parachelate. A rarely used term in Amphipoda applied to propodochelate gnathopods and pereopods in which the immovable finger is distinct, but article 6 is otherwise unexpanded or nonpalmate and the dactyl strongly overlaps the apex of the immovable finger; gnathopods of various Eophliantidae are good examples but the term may also be applied to numerous other cases, such as those linear, chelate gnathopods of Sebidae, Didymocheila, and various second gnathopods of Lysianassidae.
peduncle. The basal articles of a fundamentally biramous appendage; in Amphipoda applied to antennae, pleopods, and uropods; antenna 1 with three peduncular articles, antenna 2 with five peduncular articles (but appendage not biramous); pleopods with one definitive ,-eduncular article but remnants of others occurring proximally; uropods each with one peduncular article (fig. 1).
pereon. The complex of seven free thoracic segments bearing gnathopods and pereopods, not including the maxillipeds (fig. 1).
pereonite. A segment of the pereon.
pereopod. A walking, grasping, standing, or feeding appendage attached to a pereonite; normally composed of seven articles, including coxa; in Amphipoda the first two pairs are often termed gnathopods and only the last five pairs of thoracic legs are called pereopods (fig. 1).
plate. A flattened lobe on an article of a maxilla or maxilliped.
pleopod. A biramous swimming appendage on pleonites 1-3, one pair for each pleonite (fig. 1).
pleon. The abdomen (of six free segments in Gammaridea, rarely with some segments coalesced). See metasome and urosome.
pleonite. A segment of the pleon.
prebuccal complex. The labrum and epistome together.
prehensile. Adapted for seizing or grasping; applicable to but rarely used for gammaridean gnathopods; especially useful in denoting pereopods of cyphocarids Isaea, Pleonexes, etc., which either are subchelate or chelate, or have distinct, spinose palms or nonskid surfaces indicating their use in grasping.
propodochelate. Synonymous with chelate.
propodus. ‘The sixth article of a thoracic appendage (especially used to denote the palmar article of a gnathopod).
scale, scale-like. Terms applied to the accessory flagellum when forming a small lamella immovably fused to article 3 of antenna 1; and to the inner ramus of uropod 3 when strongly reduced and plate-like.
MARINE GAMMARIDEAN AMPHIPODA 523
simple. Used in amphipod taxonomy to denote the absence of spines or setae on appendages; or the occurrence of buta single article in the ramus of a uropod; or especially to the absence of a palm ona gnathopod or pereopod. Distinction between subchelate and simple is often weak.
splayed. Descriptive of the lateral spreading of coxae, especially in Phliantidae.
subchelate. Article 6 of a gnathopod or pereopod having a distal palm against which article 7 closes; a prehensile condition in which the palm is not produced to form a finger; intermediate in condition between chelate and simple. Com- plexly subchelate or complexly chelate are terms referring to the formation of a false chela by protrusion of teeth, cusps, or lobes from articles other than the sixth and upon which article 7 impinges to form a prehensile condition; occurring especially in Aoridae, Corophiidae, and Leucothoidae (fig. 2e). See “simple.”
telson. A flap dorsal to the anus attached to pleonite 6, primitively bilobed but usually in Amphipoda with bases coalesced and often with lobes completely coalesced to form a single plate (figs. 1, 2d).
triturative. Descriptive of the rasp-like surface of a mandibular molar, composed of teeth, ridges, and cusps.
upper lip (labrum). A fleshy lobe attached to the anterior cephalic margin in front of the mandibles; occasionally the anterior surface of the labrum _ pro- trudes as a lobe or cusp; often the cephalic area to which the labrum is attached is recognizable as an “‘epistome’”’ and may also be lobed; or both labrum and epistome may be indistinguishable and produced together as a single lobe.
uropod. Oue member of the three pairs of terminal pleonal appendages, each formed of a peduncle and two rami (fig. 1) (occasionally rami of uropod 3 reduced or absent, rarely rami of uropods 1—2 absent or reduced).
urosome. The complex of pleonites 4, 5, 6, carrying uropods, and telson. Often numbered as urosomites 1, 2, 3 (fig. 1).
urosomite. A segment of the urosome.
285-135 O - 69 - 34
Index and Familial Assignment of Gammaridean Genera
[Valid genera in capitals; subgenera in capitals and parentheses; dubiously valid genera in brackets. See Stebbing (1888, 1906) for additional nomina dubia and nuda not assignable to valid genera. Page numbers of primary citations of genera in boldface.|
Acanthechinus, to ACTINACANTHUS, Paramphithoidae, 393 Acanthogrubia, to CYMADUSA, Ampithoidae, 144 ACANTHOHAUSTORIUS, Haustori- idae, 254 ACANTHONOTOZOMA, Acanthono- tozomatidae, 120 ACANTHONOTOZOMELLA, thonotozomatidae, 122 ACANTHONOTOZOMOIDES, Acan- thononotozomatidae, 122 Acanthonotus, to ACANTHONOTO- ZOMA, Acanthonotozomatidae, 120 Acanthopleustes, to AMATHILLOP- SIS, Paramphithoidae, 394 Acanthosoma, to PARAMPHITHOH, Paramphithoidae, 396 ACANTHOSTEPHEIA, Oedicerotidae, 378 Acanthozone, to PARAMPHITHOE, Paramphithoidae, 396 ACCEDOMOERA, Eusiridae, 219 ACEROIDES, Oedicerotidae, 378 Aceropsis, to ACEROIDES, Oedicero- tidae, 378 Aceros, to ARRHIS, Oedicerotidae, 381 ACIDOSTOMA, Lysianassidae, 317 Acidostomella, to SOCARNOIDES, Lysianassidae, 363 ACONTIOSTOMA, Lysianassidae, 317 ACTINACANTHUS, Paramphithoi- dae, 393 ACUMINODEUTOPUS, Aoridae, 150 ADELIELLA, Lysianassidae, 320 AFROCHILTONIA, Hyalellidae, Talitroidea, 468 ALEXANDRELLA, Stilipedidae, 161, 263, 453
524
Acan-
see
ALIBROTUS, dubious genus, Lysianas- sidae (not discussed, see Stebbing, 1906)
ALICELLA, Lysianassidae, 320
ALLOGAUSSIA, Lysianassidae [also assigned to ? Orchomene], 320, 353
ALLORCHESTES, Hyalidae, see Tali- troidea, 468
(Allorchestina), subgenus to H YALE, Hyalidae, see Talitroidea, 469
AMAR YLLIS, Lysianassidae, 320
Amathia, to GAMMARELLUS, Gam- maridae, 242
Amathilla, to GAMMARELLUS, Gammaridae, 242
Amathillopleustes, to CLEON A RDOP- SIS, Eusiridae, 223, 392, 393
AMATHILLOPSIS, Paramphithoidae,
394
AMBASIA, Lysianassidae, 321
AMBASIELLA, Lysianassidae, 321
AMBASIOPSIS, Lysianassidae, 321
AMPELISCA, Ampeliscidae, 8, 130
AMPELISCIPHOTIS, Isaeidae, 271
AMPHIDEUTOPUS, Aoridae, 150, 267 , 269
AMPHILOCHELLA, Amphilochidae, 136
AMPHILOCHOIDES, Amphilochidae, 136, 421
AMPHILOCHOPSIS, Amphilochidae, 136
AMPHILOCHUS, Amphilochidae, 136
AMPHIPOREIA, MHaustoriidae, 254
AMPHITHOIDES, Ampithoidae, 143
AMPHITHOLINA, Eophliantidae, 142, 211, 481
Amphithonotus, to DEXAMINE, Dex- aminidae, 203
MARINE GAMMARIDEAN AMPHIPODA
AMPHITHOPSIS, 174
AMPITHOE, Ampithoidae, 143
ANAMIXIS, Anamixidae, 146
ANATYLUS, Dexaminidae, 202
ANCHIPHIMEDIA, Acanthonotozo- matidae, 122
Andania, to ANDANIEXIS, Stego- cephalidae, 440
ANDANIELLA, Stegocephalidae, 440
ANDANIEXIS, Stegocephalidae, 440
ANDANIOPSIS, Stegocephalidae, 441
ANDANIOTES, Stegocephalidae, 441
ANELASMOPUS, Gammaridae, 217, 236
ANISOGAMMARUS, Gammaridae, 42 237
Anisopus, to PLEONEXES, Ampi- thoidae, 145
ANOEDICEROS, Ocedicerotidae,
ANONYX, Lysianassidae, 324
AORA, Aoridae, 152
AORCHO, Aoridae, 152, 184, 186, 264, 267
AOROIDES, Aoridae, 152
APHERUSA, Calliopiidae, 4, 174
Araneops, to AMPELISCA, Ampelis- cidae, 130
ARCULFIA, Pardaliscidae, 401
ARGISSA, Argissidae, 159
ARISTIAS, Lysianassidae, 324
ARISTIOPSIS, Lysianassidae, 324
AROUI, Lysianassidae, 328
ARRHINOPSIS, Oedicerotidae, 381
ARRHIS, Oedicerotidae, 381
Aruga, to LYSIANASSA, Lysianas- sidae, 348
Arugella, to LYSIANASSA, Lysianas- sidae, 348
Aspidophoreia, to ALLORCHESTES, Hyalidae, see Talitroidea, 468
Aspidopleurus, to PHIPPSIA, Stego- cephalidae, 442
ASTYRA, Astyridae, 161, 392
Astyroides, to ALEXANDRELLA, Stilipedidae, 161, 453
ATYLOELLA, Eusiridae, 222
Atyloides, to SCHRADERIA, Eusiri- dae, 230
ATYLOPSIS, Calliopiidae, 175, 218
ATYLUS, Atylidae, 163
Aucklandia, to PARAMOERA, Eusiri- dae, 227
Calliopiidae, 26,
380
y45)
Audouinia, to COROPHIUM, Coro- phiidae, 190
(AUDULLA), subgenus of GAMMA- ROPSIS, Isaeidae, 271
AUST ROCHILTONIA, Hyalellidae,see Talitroidea, 469
Austronisimus, to PARAL YSIANOP- SIS, Lysianassidae, 356
AUSTROPLEUSTES, Pleustidae, 115, ZOOM A 22) ey
AUSTROSYRRHOE, Synopiidae, 458
Autonoe, to LEMBOS, Aoridae, 154
AZOTOSTOMA, Lysianassidae, 328
Barentsia, to GULBARENTSIA, Oedi- cerotidae, 382
BATEA, Bateidae, 164
BATH YAMARYLLIS, Lysianassidae, 328
BATHCALLISOMA, Lysianassidae [also assigned to SCOPELOCHEIRUS], 328, 362
BATH YCERADOCUS, Gammaridae, 238
BATHYMEDON, Oedicerotidae, 381
BATH YPANOPLOEA, Acanthonoto- zomatidae (provisional) [seeParam- phithoidae], 122, 389
BATH YPHOTIS, Ischyroceridae, 109, 231, 267, 276
BATH YPOREIA, Haustoriidae, 254
BATH YPOREIAPUS, Oedicerotidae, 381
BATH YSCHRADERIA, Eusiridae, 222
BATH YSTEGOCEPHALUS, Stegoce- phalidae, 441
BEAUDETTIA, Beaudettiidae, 37, 167
Bellia, to HAUSTORIUS, Haustoriidae, 257
Bemlos, to LEMBOS, Aoridae, 154
BIANCOLINA, Eophliantidae, 39, 211, 433, 481
BIRCENNA. Eophliantidae, 212, 433
Boeckia, to LEPTOCHEIRUS, Aoridae 154
BOECKOSIMUS, Lysianassidae, 330
BOGENFELSIA, Ischyroceridae, 186, 276
BONNIERELLA, Ischyroceridae (and subgenus to MEGAMPHOPUS, Isaeidae), 267, 273, 276
Boscia, to MELITA, Gammaridae, 245
BOUVIERELLA, Calliopiidae, 175
13,
526
BOVALLIA, Eusiridae, 222 BRUZELIA, Synopiidae, 459 Bruzeliclla, to JASSA, Ischyroceridae, 279 BRUZELIOPSIS, Synopiidae, 82, 460 BYBLIS, Ampeliscidae, 130 BYBLISOIDES, Ampeliscidae, 132 Cacao, to STILIPES, Stilipedidae, 454 Caliniphargus, to MELITA, Gammari- dae, 245 f Callimerus, to AMPHILOCHUS, Am- philochidae, 136 Calliope, to CALLIOPIUS, Calliopiidae, 176 CALLIOPIELLA, Calliopiidae, 175 CALLIOPIUS, Calliopiidae, 4, 176 Callisoma, to SCOPELOCHEIRUS, Lysianassidae, 362 CAMACHO, Corophiidae, 188, 267 CARANGOLIA, Haustoriidae, 108, 256 CARDENIO, Haustoriidae, 256 CARINOBATEA. Bateidae, 164 ‘“Carinogammar us’’, see ‘‘Marine Car-
inate Gammarus,’’ Gammaridae, 244
CAROLOBATEA, Oedicerotidae, 382, 481
CASCO, Gammaridae, 107, 238
CEBOCARIS, Lysianassidae, 331
CEINA, Phliantidae, 39, 410, 432, 433, 481
CEININA, Eophliantidae, 213, 433
CENTROMEDON, Lysianassidae, 333
CERADOCOIDES, Gammaridae, 238
CERADOCOPSIS, Gammaridae, 239
CERADOCUS, Gammaridae, 28, 42, 239
Cerapodina, to CERAPUS, Corophiidae, 188
CERAPOPSIS, Corophiidae, 188
CERAPUS, Corophiidae, 188
Chagosia, to ASTYRA, Astyridae, 161
Charcotia, to WALDECKIA, Lysian- assidae, 368
(CHEIRIMEDEIA), subgenus PROTOMEDHEIA, Isaeidae, 274
CHEIRIMEDON. Lysianassidae, 333
CHEIRIPHOTIS, Isaeidae, 271
CHEIROCRATELLA, Gammaridae, 239
CHEIROCRATUS, Gammaridae, 32, 239
CHELURA, Cheluridae, 39, 180
of
U.S. NATIONAL MUSEUM BULLETIN 271
CHEVALIA, Isaeidae, 106, 271
CHEVREUXIELLA, Lysianassidae, 335, 373
CHEVREUXIUS, Corophiidae, 190
CHILTONIA, Hyalellidae, see Tali- troidea, 43, 469
Chimaeropsis, to ARGISSA, Argissidae, 159
Chironesimus, to ANONYX, Lysian- assidae, 324
Chloris, to PARAPHERUSA, Gammar- idae, 247
CHOSROKHES, Calliopiidae, 25, 39, 176, 217,219,408, 409
CLARENCIA, Calliopiidae, 176
CLEIPPIDES, Calliopiidae, 176, 283, 392
CLEONARDO, Eusiridae, 222
CLEONARDOPSIS, Eusiridae, 392,393
CLEPIDECRELLA, Lysianassidae, 338
COLOMASTIX, Colomastigidae, 184
CONCHOLESTES, Corophiidae, 156, 190
223,
COREMAPUS, Aoridae, 152
COROPHIUM, Corophiidae, 102, 190, 481
Cratippus, to COLOMASTIX, Colo-
mastigidae, 184
Cratophium, to JASSA, Ischyroceridae, 279
CRESSA, Cressidae, 200
CRESSINA, Cressidae, 200
CRYBELOCEPHALUS, Lysianassi- dae, 338
CRYBELOCYPHOCARIS, Lysianas- sidae, 338
Cuviera, to LEUCOTHOE, Leucothoi- dae, 289
CYCLOCARIS, Lysianassidae, 338 CYCLOTELSON, Amphilochidae, 138 CYLINDRYLLIOIDES, Eophlianti- dae, 213, 433 CYMADUSA, Ampithoidae, 144 CYPHOCARIS, Lysianassidae, 339 CYPROIDEA, Amphilochidae, 139 Cyproidia, to CYPROIDEA, Amphilo- chidae, 139 CYPSIPHIMEDIA, matidae, 123 CYRTOPHIUM, Podoceridae, 427 DANAELLA, Lysianassidae, 339, 373
Acanthonotozo-
MARINE GAMMARIDEAN AMPHIPODA
Danaia, to CRESSA, Cressidae, 200
Darwinia, to LAF YSTIUS, Lafystiidae, 283
DAUTZENBERGIA, Eusiridae, 223
Dercothoe, to ?7MICROPROTOPUS, Isaeidae, 274
DERJUGIANA, Lysianassidae, 339
Dermophilus, to ?7LAFYSTIUS, Lafy- stiidae, 283
DEXAMINE, Dexaminidae, 203
DEXAMINELLA, Dexaminidae, 203
DEXAMINOIDES, Dexaminidae, 203
DEXAMONICA, Dexaminidae, 203
Dexiocerella, to PODOCERUS, Podo- ceridae, 431
DID YMOCHEILA, see Incertae Sedis, 39, 110,294,478
DJERBOA, Eusiridae, 167, 223
DOGIELINOTUS, Dogielinotidae, 208
Dorbaneila, to LEPECHINELLA, Lepechinellidae, 286
Dryope, Driope, to UNCIOLA, Coro- phiidae, 197
DRYOPOIDES, Aoridae (2), Coro- phiidae (<7), 153, 190, 275, 426
DULICHIA, Podoceridae, 427
Dulichiella, to MELITA, Gammaridae, 245
Dyopedos, to DULICHIA, Podoceridae, 427
ECHINIPHIMEDIA, Acanthonotozo- matidae, 124
“Eechinogamma rus’’, se ANISOGAM- MARUS, Gammaridae, 238
ECLYSIS, Paramphithoidae, 115, 160, 161, 392, 394
Egidia, to UROTHOE, Haustoriidae, 260
Eiscladus, to PHOTIS, Isaeidae, 274
ELASMOPOIDES, Gammaridae, 240
ELASMOPUS, Gammaridae, 28, 37, 42, 166, 217, 240
Elimedon, to 7HIPPOMEDON, Lysi- anassidae [also assigned to PARA- CENTROMEDON], 345, 355
ENDEVOURA, Lysianassidae, 342
ENSAYARA, Lysainassidae, 342
(EOGAMMARUS), subgenus of ANI- SOGAMMARUS, Gammaridae, 237
EOHAUSTORIUS, Haustoriidae, 256
EOPHLIANTIS, Eophliantidae, 213, 433
527
Ephippiphora, to WALDECKIA, Ly- sianassidae, 368
Epidesura, to ATYLUS, Atylidae, 163
EPIMERIA, Paramphithoidae, 394
EPIMERIELLA, Paramphithoidae, 115, 160, 161, 392, 395
Epimeriopsis, void ab initio, see BATH - YPANOPLOEA, Acanthonotozo- matidae, 122
ERICTHONIUS, Corophiidae, 39, 192, 275
EKriopis, to ERIOPISA, Gammaridae, 240
ERIOPISA, Gammaridae, 240
ERIOPISELLA, Gammaridae, 241
EUANDANIA, Stegocephalidae, 441
EUCALLISOMA, Lysianassidae, 342
EUON YX, Lysianassidae, 342
EURYMERA, Eusiridae, 223
Euryporeia, to KHURYTHENES, Lysi- anassidae, 343
Eurystheus, to GAMMAROPSIS, Is- aeidae, 271
Eurytenes, to EURYTHENES, Lysi- anassidae, 343
EURYTHENES, Lysianassidae, 343
EUSIRELLA, Eusiridae, 90, 114, 225, 292
EUSIROGENES, Eusiridae, 225, 233
EUSIROIDES, Eusiridae, 173, 225
EUSIROPSIS, Eusiridae, 114, 225, 292
EUSIRUS, Eusiridae, 173, 226
Euxinia, to PONTOGAMMARUS (freshwater), Gammaridae, 248
EXAMPITHOE, Ampithoidae, 144
Exhyalella, to PARH YALELLA, Hya- lellidae, see Talitroidea, 471
EXOEDICEROPSIS, Oe¢edicerotidae, 382
EXOEDICEROS, Oedicerotidae, 382
Exungia, to COLOMASTIX, Colo- mastigidae, 184
FALKLANDELLA, Gammaridae, 97, 116, 170, 241
FIGORELLA, Lysianassidae, 343
Fimbriella, to GAMMAROPSIS, Is- aeidae, 271
GAINELLA, Lysianassidae, 343
Galanthis, to HYALE, Hyalidae, see Talitroidea, 469
Gallea, to CYPROIDEA, Amphilochi- dae, 139
528
GAMMARACANTHUS, Gammaridae, 241
GAMMARELLA, Gammaridae, 241
GAMMARELLUS, Gammaridae, 242
GAMMAROPSIS, Isaeidae, 28, 38, 271
GAMMARUS, Gammaridae, 242
GAROSYRRHOE, Synopiidae, 460
GAVIOTA, Corophiidae, 192
GITNAN, Amphilochidae, 138
GITANOGEITON, Amphilochidae, 138
GITANOPSIS, Amphilochidae, 138
Glauconeme, to UNCIOLA, Corophii- dae, 197
Glycera, to GLYCERINA, Lysianassi- dae, 345
GLYCERINA, Lysianassidae, 345
GNATHIPHIMEDIA, Acanthonoto-
_ zomatidae, 124 GOESIA, Isaeidae, 272 Gossea, to APHERUSA, Calliopiidae,
174 Gracilipes, to RHACHOTROPIS, EKusiridae, 229 GRANDIDIERELLA, — Corophiidae, 147, 155, 192
Grayia, to GAMMARELLUS, Gam- maridae, 142
Grimaldia, to SEBA, Sebidae, 436
Grubia, to CYMADUSA, Ampithoidae, 144
Guerina, to TRISCHIZOSTOMA, Lysianassidae, 365
Guerinia, to TRISCHIZOSTOMA, Lysianassidae, 365
GUERNEA, Dexaminidae, 203
GULBARENTSIA, Oedicerotidae, 382
HALICE, Pardaliscidae, 401
HALICELLA, Pardaliscidae, 114, 402
HALICOIDES, Pardaliscidae, 402, 453
HALICREION, Oedicerotidae, 383
Halimedon, to WESTWOODILLA, Oedicerotidae, 387
HALIRAGES, Calliopiidae, 108, 177, 393
HALIRAGOIDES, Calliopiidae, 177
HANSENELLA, Aoridae, 153
HAPLOCHEIRA, Isaeidae, 272
HAPLOOPS, Ampeliscidae, 132
HARCLEDO, Eusiridae, 226
Harmomia, Harmonia, to PARA- PHERUSA, Gammaridae, 247
Harpina, to HARPINIA, Phoxo- cephalidae, 415 —
U.S. NATIONAL MUSEUM BULLETIN
271
HARPINIA, Phoxocephalidae, 415
HARPINIOIDELLA, Eusiridae, 167, 226 HARPINIOIDES, Calliopiidae, 177,
218, 219, 421, 422 HARPINIOPSIS, Phoxocephalidae, 415 Haustoriella, to PHOXOCEPHALOP-
SIS, Haustoriidae, 257 HAUSTORIOIDES, Dogielinotidae, 3,
209 HAUSTORIOPSIS,
251, 434 HAUSTORIUS, Haustoriidae, 256 Hela, to NEOHELA, Corophiidae, 194 Helella, to NEOHELA, Corophiidae,
Prophliantidae,
194
Helleria, to GUERNEA, Dexaminidae, 203
Hemijassa, to JASSA, Ischyroceridae, 279
HETEROCRESSA, Pagetinidae, 389
HETEROPHILIAS, Phliantidae, 410
HETEROPHOXUS, Phoxocephalidae, 417
HIPPOMEDON, Lysianassidae, 345
HIRONDELLEA. Lysianassidae, 345
Hoplonyx, to TMETONYX, Lysian- assidae, 365
HOPLOPHEONOIDES, dae, 139
HOPLOPLEON, Amphilochidae,
HORNELLIA, Gammaridae, 242
HYALE, Hyalidae, see Talitroidea, 28, 42, 207, 469
HYALELLA, Hyalelliade, troidea, 42, 469
Hyaloides, to PARH YALE, Hyalidae, see Talitroidea, 471
HYPERIOPSIS, Hyperiopsidae,
ICHNOPUS, Lysianassidae, 346
Ichthyomyzocus, to 7LAFYSTIUS, La- fystiidae, 283
ICILIUS, Podoceridae, 427
Icridium, to PEREIONOTUS, Phli- antidae, 411
Iduna, to LILJEBORGIA, Liljebor- giidae, 294
IDUNELLA, Liljeborgiidae, 293
ILERA USTROE, Synopiidae, 460
Incisocalliope, to PARAPLEUSTES, Pleustidae, 425
INSULA, Hyalid ae, see Talitroidea, 470
Amphilochi-
140
see Tali-
263
MARINE GAMMARIDEAN AMPHIPODA
Iphigenia, to IPHINOTUS, Phliantidae, 410
IPHIMEDIA, Acanthonotozomatidae, 124
IPHIMEDIELLA, Acanthonotozoma- tidae, 124
Iphimediopsis Della Valle, to PANOP- LOEA, Acanthonotozomatidae, 126
Imphimediopsis Schellenberg, to BATH- YPANOPLOEA, Acanthomotozo- matidae, 122
IPHINOTUS, Phliantidae, 410
IPHIPLATHIA, Phliantidae, 410
ISAEA, Isaeidae, 38, 272
ISAEOPSIS, Ischyroceridae, 278
ISCH Y ROCERUS, Ischyroceridae, 279
Janassa, to PARAJASSA, Ischyroceri- dae, 279
JASSA, Ischyroceridae, 6, 116, 141, 279
JEDDO, Synopiidae, 113, 159, 461
JOUBINELLA, Phoxocephalidae, 171, 418
KAMAKA, Corophiidae, 106, 194
Katius, to KURYTHENES, Lysianas- sidae, 343
KERGUELENIA, Lysianassidae, 346
KERM YSTHEUS, Isaeidae, 272
KOROGA, Lysianassidae, 346
Kroyera, to MONOCULODES, Oedi- cerotidae, 383
KUPHOCHEIRA, Isaeidae, 273
KURIA, Kuriidae, 281, 433
KYSKA, Lysianassidae, 346
LABRIPHIMEDIA, Acanthonotozom- atidae, 125
LAETMATOPHILUS, Podoceridae, 430
LAF YSTIUS, Lafystiidae, 283
Lakota, to ANONYX, Lysianassidae, 324
Lalaria, to AORA, Aoridae, 152
Lampra, to TRITAETA, Dexaminidae,
206 LAOTHOES, Calliopiidae, 177 LAPHYSTIOPSIS, Laphystiopsidae, 86, 115, 285
LEIPSUROPUS, Podoceridae, 430 LEMBOIDES, Aoridae, 153 LEMBOPSIS, Aoridae, 153
LEMBOS, Aoridae, 28, 154
LEPECHINELLA, Lepechinellidae, 163, 286
Lepidactylis, to ?HAUSTORIUS,
Haustoriidae, 256
029
LEPIDEPECREELLA, Lysianassidae, 347
LEPIDEPECREOIDES, dae, 347
LEPIDEPECREOPSIS, Lysianassidae, 347
LEPIDEPECREUM, Lysianassiade, 348
LEPTAMPHOPUS, Calliopiidae, 178
LEPTOCHEIRUS, Aoridae, 154, 267, 269
Leptochela, to EHUON YX, Lysianassi- dae, 342
LEPTOPHOXOIDES, Phoxocephali- dae, 418
LEPTOPHOXUS, 419
Leptothoe, to MAERA, Gammaridae, 243
LEUCOTHOE, Leucothoidae, 28, 289
LEUCOTHOELLA, Leucothoidae, 290
LEUCOTHOIDES, Leucothoidae, 290
LIAGOCERADOCUS, Gammaridae, 217, 243
LILJEBORGIA, Liljeborgiidae, 294
Lilljeborgiella, to LILJEBORGIA Liljeborgiidae, 294
Linguimaera, to MAERA, Gammaridae, 243
LIOUVILLEA, Eusiridae, 226
LISTRIELLA, Liljeborgiidae, 29, 36, 294
Lonchomerus, to AORA, Aoridae, 152
(LOPICEROS), subgenus of OEDI- CEROIDES, Oedicerotidae, 384
Lusyta, to JASSA, Ischyroceridae, 279
Lycesta, to LEUCOTHOE, Leucothoi- dae, 289
LYSIANASSA, Lysianassidae, 348
Lysianassina, to LYSIANASSA, Lysi- anassidae, 348
Lysianax, to LYSIANASSA, Lysianas- sidae, 348
LYSIANELLA, Lysianassidae, 349
Lysianopsis, to LYSIANASSA, Lysi- anassidae, 348
Macleayia, to JASSA, Ischyroceridae, 279
MACROPISTHOPUS, Ampithoidae, 144
MAERA, Gammaridae, 28, 42, 243
MAERACUNHA, Gammaridae, 243
MAERELLA, Gammaridae, 243
Lysianassi-
Phoxocephalidae,
530
Maeroides, to GAMMAROPSIS, Isaei-
dae, 271
MAEROPSIS, Gammaridae, 244
MANDIBULOPHOXUS, Phoxocepha-
lidae, 419
MAORIPHIMEDIA, Acanthonotozo- matidae, 125
‘““Marine Carinate, Gammarus,” Gam- maridae, 244
(MARINOGAMMARUS), Gammari- dae, see GAMMARUS, 242
MAXILLIPHIMEDIA, Acanthonoto- zomatidae, 125, 451
Megalonoura, to MEGALUROPUS, Gammaridae, 244
Megalorchestia, to ORCHESTOIDEA,
Talitridae, see Talitroidea, 471
MEGALUROPUS, Gammaridae,
94, 107, 157, 158, 244, 369
Megamoera, to MELITA, Gammaridae,
245
MEGAMPHOPUS, Isaeidae, 273
MELITA, Gammaridae, 245
MELITOIDES, Gammaridae, 245
MELPHIDIPPA, Melphidippidae, 371
MELPHIDIPPELLA, Melphidippidae,
371
MELPHISANA, Melphidippidae, 172, 371
MENIGRATES, Lysianassidae, 349
MENIGRATOPSIS, Lysianassidae, 349
MESOCYCLOCARIS, Lysianassidae, 349
MESOCYPHOCARIS, Lysianassidae, 350
MESOGAMMARUS, Gammaridae 245
MESOMETOPA, Stenothoidae, 448
MESOPLEUSTES, Pleustidae, 88, 167,
30,
392, 422
MESOPROBOLOIDES, Stenothoidae, 448
Mesostenothoides, to STENOTHOI-
DES, Stenothoidae, 451
METACERADOCOIDES, Gammari-
dae, 245
METACERADOCUS, Gammaridae 246
METACYCLOCARIS, Lysianassidae,
350
METACYPHOCARIS, Lysianassidae, 350
U.S. NATIONAL MUSEUM
BULLETIN 271
METALEPTAMPHOPUS, piidae, 108, 178, 369 METAMBASIA, Lysianassidae, 350 Metandania, to ANDANIOTES, Ste- gocephalidae, 441 METAPHOXUS, Phoxocephalidae, 419 METEPIMERIA, Paramphithoidae
Callio-
396
METEUSIROIDES, Eusiridae, 167, 227
METHALIMEDON, Oedicerotidae 383
Metharpinia, to PARAPHOXUS, Phoxocephalidae, 420 METOEDICEROS, [Oedicerotidae], see Incertae Sedis, 37, 113, 207, 376, 377, 480 METOPA, Stenothoidae, 448 METOPELLA, Stenothoidae, 449 METOPELLOIDHES, Stenothoidae, 449 Metopina, to METOPA, Stenothoidae, 448 METOPOIDHES, ’Stenothoidae, 449 Microcheles, to IPHIMEDIA, Acantho- notozomatidae, 124 MICRODEUTOPUS, Aoridae, 154 MICROJASSA, Ischyroceridae, 279 MICROLYSIAS, Lysianassidae, 351 MICROPHOTIS, Isaeidae, 273 MICROPHOXUS, Phoxocephalidae, 419 Microplax, to LILJEBORGIA, Lilje- borgiidae, 294 MICROPROTOPUS, Isaeidae, 110, 274 MICROSTENOTHOEH, Stenothoidae, 449 MONOCULODES, Oedicerotidae, 383 MONOCULOPSIS, Oedicerotidae, 383 Montagua, to STENOTHOEH, Steno- thoidae, 450 Montaguana, to STENOTHOE, Steno- thoidae, 450 Mulleria, to MAERA, Gammaridae, 243 Naenia, to PODOCEROPSIS, Isaeidae, 274 NAJNA, see Talitroidea (provisional), 37, 59, 113, 209, 406, 470 NANNON YX, Lysianassidae, 351 NECOCHEA, Pardaliscidae, 402 NEOAMBASIA, Lysianassidae; 351 NEOBULE, Talitroidea (provisional), 470
MARINE GAMMARIDEAN AMPHIPODA
NEOCYPROIDEA, Amphilochidae, 140 Neogammaropsis, to HLASMOPUS,
Gammaridae, 240 NEOHAUSTORIUS, Haustoriidae, 257 NEOHELA, Corophiidae, 194 NEOMEGAMPHOPUS, Aoridae, 155,
267, 269 NEOMICRODEUTOPUS, Aoridae, 155, 184, 194 Neophotis, to CHEVALIA, Isaeidae,
271 NEOPLEUSTES, Pleustidae, 424 NEOXENODICEH, Podoceridae, 431 NETAMELITA, Gammaridae, 246 Nicea, to HYALE, Hyalidae, see Tali- troidea, 469 NICIPPE, Synopiidae, 403 NIPPOCHELURA, Cheluridae, 181 Normania, to NORMANION, Lysi- anassidae, 351 NORMANION, Lysianassidae, 351 Nototropis, to ATYLUS, Atylidae, 163 OCHLESIS, Ochlesidae, 373 OCOSIN GO, Lysianassidae, 352 ODIUS, Acanthonotozomatidae, 125 OEDICERINA, Oedicerotidae, 384 OEDICEROIDES, Oedicerotidae, 384 OEDICEROPSIS, Oedicerotidae, 384 Oediceropsoides, to OEDICEROIDES, Oedicerotidae, 384 OEDICEROS, Oedicerotidae, 385 ONESIMOIDES, Lysianassidae, 352 ONISIMUS, Lysianassidae, 352 “ONISIMUS” auct., to BOECKO- SIMUS, Lysianassidae, 330 Opis, to OPISA, Lysianassidae, 353 OPISA, Lysianassidae, 353 ORADAREA, Calliopiidae, 393 ORCHESTIA, Talitroidae, 470 ORCHESTOIDEA, Talitridae, Talitroidae, 12, 471 ORCHOMENT, Lysianassidae, 40, 353 Orchomenella, to ORCHOMENE. Lysianassidae, 365 “Orchomenella”’ greenlandica, Lysian- assidae, see keys, 306, 311, 313 Orchomenopsis, to ORCHOMENE, Lysianassidae, 353 Orthopalame, to MICROPROTOPUS, Isaeidae, 274
178, 392, Talitridae, see
see
531 Otus, to ODIUS, Acanthonotozo- matidae, 125 PACH YCHELIUM, Lysianassidae, 354 PACH YNUS, Lysianassidae, 354 PAGETINA, Pagetinidae, 389 PALINNOTUS, Phliantidae, 411 PANOPLOEA, Acanthonotozomatidae, 126 PANOPLOEOPSIS, matidae, 126 PARACALLIOPE (freshwater), Cal- liopiidae, see supplement, 172, 481 PARACALLISOMA, Lysianassidae, 89, 111, 354 PARACALLISOMOPSIS, sidae, 355 [PARACENT ROMDEON], sidae [also assigned to POMEDON|], 345, 355 PARACERADOCUS, Gammaridae, 246 PARACOROPHIUM, 194, 481 Paracyclocaris, to LEPIDEPECRE- ELLA, Lysianassidae, 347 PARACYPHOCARIS, Lysianassidae, 355 PARACYPROIDEA, Amphilochidae, 140 PARADEXAMINE, Dexaminidae, 206 PARADRYOPEH, Aoridae, 155, 275 PARADULICHIA, Podoceridae, 431 PARAGRUBIA, Ampithoidae, 144 PARAHALICE, Pardaliscidae, 114, 403 PARAHAUSTORIUS, Haustoriidae, 257 PARAJASSA, Ischyroceridae, 279 PARALEPECHINELLA, Lepechinel-
Acanthonotozo-
Lysianas-
Lysianas- ?HIP-
Corophiidae,
lidae, 288 PARALEPTAMPHOPUS, (fresh- water), Calliopiidae, 172 PARALEUCOTHOE, Leucothoidae, 291
PARALIBROTUS, Lysianassidae, 356 PARALICELLA, Lysianassidae, 356 PARALLORCHESTES, Hyalidae, see Talitroidae, 471 PARALYSIANOPSIS, 356 PARAMBASIA, Lysianassidae, 356 PARAMETOPA, Stenothoidae, 449 PARAMETOPELLA, Stenothoidae, 450
Lysianassidae,
032
PARAMOERA, Eusiridae, 227 PARAMPHITHOHE, Paramphithoidae,
119, 176, 396
Paranaenia, to GAMMAROPSIS, Isaeidae, 271
PARANAMIXIS, Anamixidae, 146, 164
PARANCHIPHIMEDIA, Acantho-
notozomatidae, 126 PARANDANIA, Stegocephalidae, 442 Parandaniexis Nicholls, to PPEUDAN -
DANIEXIS, Stegocephalidae, 443 PARANDANIEXIS Schellenberg,
Stegocephalidae, 442 PARAOROIDES, Aoridae, 155 PARAPANOPLOEA, Acanthonotozo-
matidae, 127 PARAPERIOCULODES,
tidae, 385 PARAPHERUSA, Gammaridae, 107,
116, 170, 247, 275 PARAPHOXUS, Phoxocephalidae, 420 PARAPLEUSTES, Pleustidae, 425 PARARGISSA, Hyperiopsidae, 32, 112,
158, 264 Pararistias, to PERRIERELLA, Ly-
sianassidae, 358 Parastyra, to ASTYRA, Astyridae, 161 Parathaumatelson, to PSE UDOTHA U-
MATELSON, Thaumatelsonidae,
475 Paratryphosites, to HIPPOMEDON,
Lysianassidae, 345 Paratylus, to ATYLUS, Atylidae, 163 Paravalettia, to SEBA, Sebidae, 436 PARAWALDECKIA, _Lysianassidae,
357 “Parawaldeckia kidderi,’’ see PARA-
WALDECKIA, Lysianassidae, 357 PARDALISCA, Pardaliscidae, 403 PARDALISCELLA, Pardaliscidae, 251,
403 PARDALISCOIDES, Pardaliscidae,404 PARDALISCOPSIS, Pardaliscidae, 404 PARDISYNOPIA, Pardaliscidae, 404 PARELASMOPUS, Gammaridae, 247,
Oedicero-
291 PAREPIMERIA, Paramphithoidae, 396 PAREPIMERIELLA, Pleustidae, 392, 393, 425
PAREUSIROGENKHES, Eusiridae, 227 PARHALIMEDON, Oedicerotidae, 385
U.S. NATIONAL MUSEUM BULLETIN 271
Parharpinia, to PARAPHOX US, Phoxocephalidae, 420
PARH YALH, Hyalidae, seeTalitroidea, 471
PARH YALELLA, Talitroidea, 471
PARIPHIMEDIA, Acanthonotozoma- tidae, 127
PARIPHIMEDIELLA, Acanthonoto- zomatidae, 127
PARIPHINOTUS, Phliantidae, 411
PAROEDICEROIDES, Oedicerotidae,
Hyalellidae, see
385 PAROEDICEROS, Oedicerotidae, 386 PARONESIMOIDES, Lysianassidae, 357
PARONESIMUS, Lysianassidae, 357
Paropisa, to SOPHROSYNE, Lysian- assidae, 363
Parorchestia, to ORCHESTIA, Tali- tridae, see Talitroidea, 470
PARPANO, Pardaliscidae, 404
PARUNCIOLA, Corophiidae, 196, 267
(PATOIDES), subgenus, to ACE RO- IDES, Oedicerotidae, 378
(PECTENOGAMMARUS), subgenus of GAMMARUS, Gammaridae, 242
PELTOCOXA, Amphilochidae, 140
PELTOPES, Amphilochidae, 141
PEREIONOTUS, Phliantidae, 411
PERIOCULODES, Oedicerotidae, 386
PERIOCULOPSIS, Oedicerotidae, 386
Periphlias, to CEINA, Phliantidae, 410
PERRIERELLA, Lysianassidae, 358
Phaedra, removed from APHERUSA, Calliopiedae, to dubious status, 174
Pherusa, to GAMMARELLA, Gam- maridae, 241
Pherusana, to GAMMARELLA, Gam- maridae, 241
PHIPPSIA, Stegocephalidae, 442
PHIPPSIELLA, Stegocephalidae, 442
PHLIAS, Phliantidae, 61, 411
PHOT IS, Isaeidae, 8, 274
PHOXOCEPHALOPSIS, Haustoriidae, 108, 257
PHOXOCEPHALUS, Phoxocephalidae, 420
PHOXOSTOMA, Lysianassidae, 358
Phoxus, to PHOXOCEPHALUS, Phococephalidae, 420
MARINE GAMMARIDEAN AMPHIPODA
Phylluropus, to MEGALUROPUS, Gammaridae, 158, 244
Platamon, to HIPPOMEDON, Lysian- assidae, 345
Platophium, to PODOCERUS, Podo- ceridae, 431
PLAT YISCHNOPUS, 258, 414, 436
PLEONEXHES, Ampithoidae, 145
PLEUSTES, Pleustidae, 425
PLEUSYMTES, Pleustidae, 425
PLIOPLATEIA, Phliantidae, 412
PODOCEROPSIS, Isaeidae, 274
PODOCERUS, Podoceridae, 22, 28, 39, 409, 431
PODOPRION, Lysianassidae, 358
PODOPRIONELLA, Lysianassidae, 358
PODOPRIONIDES, Lysianassidae, 359
POLYCHEIRA, Dexaminidae, 26, 30, 206
Pontharpinia, to PARAPHOXUS, Phoxocephalidae, 420
PONTOCRATES, Oedicerotidae, 386
PONTOGENEIA, Eusiridae, 172, 228
PONTOGENEIELLA, Eusiridae, 228
PONTOGENEOIDES, Eusiridae, 167, 173, 228
PONTOPOREIA, Haustoriidae, 258
PRACHYNELLA, Lysianassidae, 359
Prinassus, to GUERNEA, Dex- aminidae, 203
PRINCAXELIA, Pardaliscidae, 405
Priscilla, to PRISCILLINA, Haustori- idae, 258
PRISCILLINA, Haustoriidae, 258
Proboliella, to PROBOLOIDES, Sten- othoidae, 450
PROBOLISCA, Stenothoidae, 450
Probolium, to STENOTHOE, othoidae, 450
PROBOLOIDES, Stenothoidae, 450
PROCYPHOCARIS, Lysianassidae, 359
PROHARPINIA, Phoxocephalidae, 420
PROLAPHYSTIOPSIS, Laphystiopsi- dae, 115, 283, 285
PROLAPHYSTIUS, Laphystiopsidae, 86, 170, 174, 285
PROMETOPA, Stenothoidae, 450
Haustoriidae,
108,
Sten-
033
[PRONANNON YX], Lysianassidae [also assigned to 7LYSIANASSA], 348, 359
PROPHLIAS, Prophliantidae, 434
PROSTEBBINGIA, Eusiridae, 228
PROSTENOTHOE, Stenothoidae, 450
PROTHAUMATELSON, Thaumatel- sonidae, 475
PROTOHAUSTORIUS, Haustoriidae, 259
Protohyperiopsis, to PARARGISSA, Hyperiopsidae, 264
PROTOMEDEHIA, Isaeidae, 274
Protophoxus, to PARAPHOXUS, Phoxocephalidae, 420
Pseudalibrotus, to ONISIMUS, Lysi- anassidae, 352
PSEUDAMBASIA, Lysianassidae, 360
PSEUDAMPHILOCHUS, Amphilo- chidae, 36, 37, 115, 139, 217, 292, 421
PSEUDANDANIEXIS, Stegocephali- dae, 443
PSEUDEPIMERIA, Paramphithoidae, 396
PSEUDERICTHONIUS, Corophiidae, 196
(PSEUDEURYSTHEUS), subgenus of GAMMAROPSIS, Isaeidae, 271
PSEUDHARPINIA, Phoxocephalidae, 420
PSEUDIPHIMEDIELLA, Acanthono- tozomatidae, 127
Pseudiphimediopsis, to BATH YPAN- OPLOEA, Acanthonotozomatidae, 122
PSEHUDISCH YROCERUS, Ischyro- ceridae, 280
PSEUDOANON YX, Lysianassidae, 360
Pseudoceradocus, to QUADRIVISIO, Gammaridae, 247
PSEUDOHAUSTORIUS, Haustori- idae, 259
PSEUDOKOROGA, Lysianassidae, 360
PSEUDOMOERA, Eusiridae, 229
Pseudonesimus, to SCHISTURELLA, Lysianassidae, 361
Pseudophthalmus, to Ampeliscidae, 130
PSEUDOPONTOGENEIA, Eusiridae, 172, 229
AMPELISCA,
534
PSEUDORCHOMENEH, Lysianassidae, 360
PSEUDOTHAUMATELSON, Thau- matelsonidae, 475
PSEUDOTIRON, Synepiidae, 109, 397, 461
Pseudotryphosa, to ?7?URISTES, Lys- ianassidae, 367
Pterygocerus, to Haustoriidae, 256
Ptilocheirus, to LEPTOCHEIRUS, Aoridae, 154
Pyctilus, to ERICTHONIUS, Coro- phiidae, 192
QUADRIVISIO, Gammaridae, 247
QUASIMODIA, Philantidae, 412
REGALIA, Calliopiidae, 178, 283
RHACHOTROPIS, Eusiridae, 33, 173, 229, 392
RIFCUS, Lysianassidae, 361
RONCO, Eusiridae, 229
ROZINANTE, Etsiridae, 230
RUDILEMBOIDES, Aoridae, 156
RUNANGA, Corophiidae, 196, 426
SANCHO, Calliopiidae, 25, 26, 39, 108, 179, 215, 408, 409
Scamballa, to ORCHESTIA, Talitridae, see Talitroidae, 470
SCHISTURELLA, Lysianassidae, 361
SCHRADERIA, Eusiridae, 230
SCOPELOCHEIROPSIS, Lysianas-
HAUSTORIUS,
sidae, 361 SCOPELOCHEIRUS, Lysianassidae, 362 SEBA, Sebidae, 436 (SEGAMPHOPUS), subgenus of
MEGAMPHOPUS, Isaeidae, 273 Sextonia, to IDUNELLA, Liljeborgi- idae, 293 SHACKLETONIA, Lysianassidae, 362 Shoemakerella, to LYSIANASSA, Lysi- anassidae, 348 SIPHONOECETES, Corophiidae, 197 SOCARNESLLA, Lysianassidae, 362 SOCARNEI, Lysianassidae, 362 SOCARNODES, Lysianassidae, 363 SOCARNOPSIS, Lysianassidae, 363 SOPH ROSHNE, Lysianassidae, 363 SPHAEROPHTHALMUS, © see certae Sedis, 202, 480 Spinifer, to PHOXOCEPHALUS, Phoxocephalidae, 420
In-
U.S. NATIONAL MUSEUM BULLETIN
271
Stebbingia, to siridae, 227 STEGOCEPHALINA, Stegocephalidae, 443 STEGOCEPHALOIDES, Stegocepha- lidae, 443 STEGOCEPHALOPSIS, lidae, 443 STEGOCEPHALUS, Stegocephalidae, 444 STEGOPLAX, Amphilochidae, 141 STELEUTHERA, Stegocephalidae, 444 Stenia, dubious genus, Lysianassidae (not discussed, see Stebbing, 1906) STENOPLEURA, Calliopiidae, 179 STENOPLEUROIDES, Calliopiidae, 179 STENOPLEUSTHS, Pleustidae, 425 STENOTHOE, Stenothoidae, 28, 450 STENOTHOIDES, Stenothoidae, 451 STEN ULA, Stenothoidae, 451 STEPHENSENTA, Lysianassidae, 364 STEPHOBRUZELIA, Synopiidae, 461 Sthenometopa, to METOPA, Steno- thoidae, 448 STILIPES, Stilipedidae, 161, 454 Stimpsonella, to MICRODEUTOPUS, Aoridae, 154 Stimpsonia, to Aoridae, 154 STOMACONTION, Lysianassidae, 364 Suleator, to HAUSTORIUS, Haus- toriidae, 257 SUNAMPHITOE, Ampithoidae, 145 Sympleustes, to STENOPLEUSTHES, Pleustidae, 425 SYNCHELIDIUM, Oedicerotidae, 387 SYNDEXAMINE, Dexaminidae, 206 SYNOPIA, Synopiidae, 34, 107, 115,
PARAMOERA, Eu-
Stegocepha-
MICRODEUTOPUS,
113,
116, 462 Synopioides, to HALICEH, Pardalisci- dae, 401
SYRRHOEH, Synopiidae, 462
SYRRHOITES, Synopiidae, 462
Talitriator, to TALITRUS, Talitridae, see Talitroidea, 472
Talitroides, to TALITRUS, Talitridae, see Talitroidea, 472
Talitronus, to ORCHESTOIDEA, Tal- itridae, see Talitroidea, 471
MARINE GAMMARIDEAN AMPHIPODA
(Talitrorchestia), subgenus, to ?TALI- TRUS, Talitridae, see Talitroidea, 472
TALITRUS Talitridae, see Talitroidea, 472
TALORCHESTIA, Talitridae, see Tal- itroidea, 472
TEMNOPHLIAS, Phliantidae, 21, 22, 26, 412
Teraticum, to SEBA, Sebidae, 436
Tessarops, to TIRON, Synopiidae, 463
TETRADEION, Stegocefhalidae, 444
Tetromatus, to AMPELISCA, Ampel- iscidae, 130
Tetronychia, to HIRONDELLEA, Lysianassidae, 345
THAUMATELSON, Thaumatelsoni- dae, 476
Thersites, to BATHYPOREIA, Haus- toriidae, 254
Thoelaos, to LAOTHOES, Calliopiidae, 177
THORIELLA, Lysianassidae, 364, 373
TIRON, Synopiidae, 463
TMETON YX, Lysianassidae, 365
“Tmetonyx’’ auct., to TRYPHOSEL- LA, Lysianassidae, 365
Trichophoxus, to PARAPHOXUS, Phoxocephalidae, 420
TRIODOS, Ampeliscidae, 132
TRISCHIZOSTOMA, Lysianassidae, 365
TRITAETA, Dexaminidae, 206
Tritropis, to RHACHOTROPIS, Eusi- ridae, 229
TROPICHELURA, Cheluridae, 182
Tryphosa, to ORCHOMENEH, Lysian- assidae, 353, 365
O30
“Tryphosa”’ auct., to TR YPHOSELLA, Lysianassidae, 365 TRYPHOSELLA, Lysianassidae, 365 TRYPHOSITES, Lysianassidae, 366 TRYPHOSOIDES, Lysianassidae, 367 UNCINOTARSUS, Aoridae, 156, 190, 427 UNCIOLA, Corophiidae, 197 UNCIOLELLA, Corophiidae, 197 URISTES, Lysianassidae, 367
Uristoides, to URISTES, Lysianas- sidae, 367
UROHAUSTORIUS, Haustoriidae, 259
UROTHOE, Haustoriidae, 108, 260 UROTHOIDES, Haustoriidae, 108, 260 USCHAKOVIELLA, Paramphithoidae, 397 VALETTIA, Lysianassidae, 367 VALETTIOPSIS, Lysianassidae, VEMANA, Vitjazianidae, 478 Vertumnus, to 7EPIMERIA, Paramph- ithoidae, 394 VIJAYA, Lysianassidae, 368 VITJAZIANA, Vitjazianidae, 478 WALDECKIA, Lysianassidae, 368 WANDELIA, Eophliantidae, 213, 433 Westwoodia, to WESTWOODILLA, Oedicerotidae, 387 WESTWOODILLA, Oedicerotidae, 387 WEYPRECHTIA, Gammaridae, 170, 247, 392, 393 Wyvillea, to JASSA, Ischyroceridae, 279 XENOCHEIRA, Aoridae, 156 Xenoclea, to PODOCEROPSIS, Isaei- dae, 274 XENODICH, Podoceridae, 431 ZARAMILLA, Eusiridae, 230 ZOBRACHO, Haustoriidae, 260
368
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