Bousfield 1978 Phylogeny of Amphipod PDF

E . L . B O U S F I E L D , F . R . S . C* .
A Revised Classification and Phylogeny of Amphipod Crustaceans
ABSTRACT
This study analyses the probable origin and evolution of the Amphipoda, a group of small- to medium-sized crustaceans that have diversified in pelagic and benthic habitats of marine, freshwater, and terrestrial environments of the world. The study includes a revision of the suborder Gammaridea based on new concepts of superfamilies and their taxonomic relationships. The classification encompasses some 80 families and 20 superfamilies of which 13 are newly elevated and diagnosed; also formally proposed and diagnosed are the families Urothoidae, Typhlogammaridae, and Gammaroporeiidae and the genus Gamrnaropoveia. The superfamily revision is based primarily on the morphology of the reproductive male stage, as well as on the plesiomorphic-apomorphic condition of taxonomic characters of the body and appendages previously established at family and superfamily levels of significance. A scheme of probable phylogenetic relationships between amphipod suborders and gammaridean superfamily groups is proposed. The present hypothesis on the probable origin and evolution of the Amphipoda is based partly on the fossil record of amphipods and related malacostracans, on modern distributions of component superfamily groups in relation to geochronology of continental drift and plate tectonics, on pertinent evolutionary models proposed (by others) for other malacostracan groups of comparable distributions and means of dispersal, and on the stimulus to evolution of new animal taxa provided by the proliferation of new plant food resources during the Carboniferous and again in the Cretaceous period. The evidence suggests a late Palaeozoic origin for the Amphipoda, with diversification maximal during the Mesozoic and continued (in more apomorphic groups) throughout the Cenozoic to the present.
*National Museum of Natural Sciences, Ottawa.
TRANSACTIONS OF THE ROYAL SOCIETY OF CANADA l SERIES IV 1 VOLUME XV1 1 1978
344 The Royal Society of Canada
The order Amphipoda comprises a group of small- to medium-sized crustaceans that are widespread in marine and freshwater environments of the world. They constitute an important element of aquatic energy cycles by converting plant and animal protein and organic detritus into a form that is suitable as food for larger animals, especially fishes, and ultimately man. Described to date are more than 5000 species, ranging in length from less than 1 millimetre to more than 25 centimetres (lobstersized). In view of the many aquatic environments and habitats as yet unexplored for amphipods, especially in the deep ocean basins and in hypogean freshwater and terrestrial biomes, extrapolation of current rates of new species descriptions yields a probable total world amphipod fauna of more than 25,000 species. Amphipods, along with isopods, tanaids, cumaceans, mysids, and a few rare hypogean ordinal groups, are members of the malacostracan superorder Peracarida. More familiar to the layman are the larger-sized eucaridan malacostracans, to which belong the decapod and euphausiid shrimps, the lobsters, hermit crabs, and crabs. Peracaridans, however, may be considered more specialized morphologically, largely by reduction or loss of such basic crustacean features as the carapace (and thoracic respiratory chamber), biramous limbs, stalked eyes, antenna1 accessory flagella, by "cephalization" of the mouthparts in a clearly defined buccal mass (as in Insecta), and by direct development of the egg (within the female thoracic brood pouch), in contrast to the complicated free-swimming larval metamorphosis of the Eucaridans. Within the orders of Peracarida, the Amphipoda would seem of relatively recent evolution. This is inferred from the advanced morphology of mouthparts and thoracic and abdominal appendages, the numerical dominance in coastal marine and freshwater environments, and their relatively low subordinal diversity (four suborders versus ten in the Isopoda), none of which are primarily parasitic (versus three such suhorders in Isopoda). Amphipods are less numerous and less diverse than isopods in both the deep sea and on land, environments that have presumably been penetrated by primarily coastal marine organisms only with great difficulty and only over immense periods of time. Furthermore, the proven amphipod fossil record extends back only to the Eocene, whereas fossil mysids, tanaids, cumaceans, and isopods are recorded unequivocally from the Triassic, Permian, and Carboniferous (Hessler 1969; Kukalova-Peck 1973; Schram 1977). The classification of the Amphipoda, based on a natural (phyletic) system, has long posed special difficulties for crustacean taxonomists. The order has been subdivided into four suborders, the Gammaridea, Hyperiidea, Caprellidea, and Ingolfiellidea, of which the Gammaridea with more than 4500 described species is by far the largest and most diverse. Paradoxically, however, the high degree of morphological di-
BousJield: Revised ClassiJication ofAmphipod Crustaceans 345
versity and prevalence of convergent or parallel evolution within differing gammaridean family and superfamily groups, coupled with the very limited fossil record, has enormously complicated the tracing of natural phylogenies. The presence of numerous forms "intermediate" between related genera, families, superfamilies, and even suborders tends to blur the critical definition of these subjective concepts (see Stebbing 1888). The presence of so many intermediate stages probably reflects, however, the relatively recent evolution of the amphipods as a group, since the subsequent time span has presumably been insufficient for their total ecological replacement by more advanced descendants. Basic gammaridean taxonomy and classification were advanced most notably by Bate (1862), G.O. Sars (1895), and Stebbing (1888, 1906). Their systems were modified by J.L. Barnard (1958, 1969a, 1974), who proposed the family Gammaridae as the "basic" gammaridean amphipod type from which all other families and family groupings radiated, but departed from previous formats by presenting the families alphabetically rather than phyletically. Previous arrangements (especially of Sars, 1895) placed the Lysianassidae, Phoxocephalidae, and other "mysidlike" marine groups with specialized pelagic terminal males, and the Talitridae in the most primitive or plesiomorphic morphological position. Attempts to consolidate the rapidly growing and rather unwieldy family groups (some 3500 species in about 60 families in 1958) into superfamily units were commenced by Bulycheva with the Talitroidea (1957), and continued by J.L. Barnard with informal groupings of related families (1969a) and formal designation of the Corophioidea (1973). Bousfield (1973) informally grouped related families on a systematics-ecology and guidebook-utilitarian basis, but also recognized the need to break up large polyphyletic families such as the (old) "Gammaridae" into more natural and critically definable units. Reorganization of gammaroidean (gammaridan) amphipods on a family and superfamily basis was completed for oral presentation at the International Groundwater Colloquium at Schlitz in 1975, and published shortly thereafter (Bousfield 1977). J.L. Barnard and co-workers (1976, 1977), Holsinger (1977), Holsinger and Longley (1979, in press), and Stock (1977) have since confirmed the essential validity of most of the family and superfamily units and generic realignments and proposed some internal modifications to these. Despite such recent advances in amphipod systematics, most families of Gammaridea remain unevaluated with respect to generic inclusion and superfamily association. Some very large and morphologically diverse families (e.g. Lysianassidae) are greatly in need of family and subfamily subdivision and realignment. The need to group the present 80+ gammaridean family units into a relatively small number of natural superfamily units and to establish probable phyletic relationships among
them has become critical to the proper development of amphipod systematics and to the understanding of their ecology, physiology, life cycles, and general biology. By contrast, the systematics of decapod and isopod crustaceans are on a much more clearly defined and sounder basis (see Kaestner 1970). Within the Amphipoda, Bowman and Griiner (1973) have proposed a very neat and natural subdivision of the suborder Hyperiidea into infraorders, superfamilies, and families that employs reproductive morphology as a primary taxonomic tool and provides a tempting model for reclassification of the Gammaridea. Ruffo (1970) and Stock (1976) have established a workable classification of the small, relict suborder Ingolfiellidea. However, the contributions of McCain (1970), Vassilenko (1974), and Laubitz (1976a) have not yet produced a universally acceptable taxonomic system for the Caprellidea. Within this setting, the purpose of the present study is to complete the formal grouping of gammaridean families into natural superfamily units, based on taxonomic characters proposed and utilized in the gammaridan superfamily revision (Bousfield 1977), to outline probable phylogenetic relationships among the superfamily and subordinal groups, and to assess the direction of evolutionary thrust within the order Amphipoda. The principal basis for the present superfamily reclassification of the Gammaridea and the attendant concepts of phylogeny and evolution were developed during presentation of a short course on amphipod (and peracaridan) biology at the College of Marine Studies, University of Delaware, in 1977 and at the Bamfield Marine Station, British Columbia, in 1978. I am especially grateful to my colleagues Dr M.R. Carriker and Dr K.S. Price at Delaware, and Dr J.E. McInerney at Bamfield, for their foresight and stimulus in making possible the course presentation and ideas developed here, and to the many students and associates who contributed variously to the development and testing of these concepts.* I would also thank especially Dr J.R. Holsinger for access to his exciting unpublished information on hypogean amphipods from artesian wells in the southwestern United States and for commentary on the present concepts, and also Dr Eric Mills and Diana Laubitz for critical review of the manuscript.
PROPOSED SUPERFAMILY RECLASSIFICATION O F THE GAMMARIDEA
The systematic reclassification of gammaridean amphipods proposed here is based on consideration of taxonomic characters previously held to be of family and superfamily significance (see Bousfield 1977).t These
* Essential elements of this paper were also presented at the 4th International Symposium
on Gammarus and Niphargus at Blacksburg, Virginia, September 1978.
t The plesiomorphy-apomorphy of some morphological characters given in Table 11 of
BousJield: Revised Classification ofAmphipod Crustaceans 347
characters are illustrated in Figures 1 to 5 and assessed below on the basis of their plesiomorphic (primitive) or apomorphic (derived, advanced) condition. Thus, each superfamily grouping is seen to have a characteristic appearance or facies, based either on a few outstanding and diagnostic characteristics (as in Lysianassoidea, Oedicerotoidea, Talitroidea, Ampeliscoidea, and Corophioidea) or on a series or combination of taxonomic features (Melitoidea, Liljeborgioidea, Crangonyctoidea) that holds more or less consistently for all component family members. Because of infragroup variation caused by adaptive radiation on the one hand, and convergence on the other, a "best fit" (or consensus) of several major characters must be used to confirm superfamily inclusion. The prime basis for the present superfamily reclassification is the morphology of the reproductive male stage or instar. This basis reflects "reproductive morphology" in more primitive extant peracaridans such as Mysidacea and Cumacea. In most instances, especially where pairing takes place pelagically in the water column, the male dies shortly afterwards, and this stage is therefore terminal. In primarily benthic males which "amplex" with, or carry, the female prior to copulation, the male may pass through two or three adult moult phases or stages, or even reach a "post-mature" stage. Four main reproductive, or adult male, types are recognized here, in order of plesiomorphy-apomorphy: 1 Pelagic terminal male instar in which the antennae bear calceoli, and usually also brush setae, in some or most group members. 2 Pelagic male instar in which the antennae bear brush setae only and calceoli are not present in any group member. 3 Non-pelagic or benthic male instar in which antennae bear calceoli in some group member(s), rarely also brush setae. 4 Non-pelagic or benthic male stage in which all group members lack calceoli and brush setae, but may possess aesthetascs especially on antenna 1 . Type 1 males are usually strongly sexually dimorphic (Figure IA-D). In comparison with the female (which is usually benthic and infaunal, Figure lc), the antennae (especially A2) are elongate and armed with sensory structures, the eyes are larger and more heavily pigmented, the abdomen and pleopods (swimming legs) are much more powerfully developed, the tail fan (especially uropod 3) is large and foliaceous (plumose-setose), and the body is smaller and more slender. Gnathopods 1 and 2 are rarely, or only slightly, sexually dimorphic, and
that paper is reversed herewith. Thus, the plesiomorphic condition includes nonsexually dimorphic gnathopods, lack of dorsal abdominal groups of spinules, absence of baso-facial spine on peduncle of uropod 1, and apically notched telson lobes.
FIGURE
0-Q.
1. Morphological types of adult male instars, antenna1 armature, and corresponding females: types 1, A-D, F; type 2, E , G-I; type 3, J - N ; type 4,
Bous$eld: Revised ClassiJication ofAmphipod Crustaceans 349
are usually similar (e.g. Phoxocephaloidea) but may be very dissimilar . (e.g. Lysianassidae) (Figure 4 ~ ) Calceoli are small, translucent, platelike, cup-like, or barrel-shaped structures (vibration sensors?) located usually on the posterior margins of the peduncles andlor flagellar segments of antenna 1, and on the anterior margins of the peduncles and flagellar segments of antenna 2 (Figure l ~F ,, K , L, M). Brush setae are short, fine hair-like setae, close-set or clustered in short transverse rows (sexual chemoreceptors?) located on the posterior peduncular segments and on the proximal (often conjoint or fused) segments of the flagellum of antenna 1, and on the anterior margins of the distal peduncular segments of antenna 2 (Figure 1E , G). Type 2 males (Figure 1 ~ - H may be sexually dimorphic in form of the ) abdomen (Figure 2 ~B), pleopods, and tail fan, and the gnathopods may , be weakly dimorphic and weakly amplexing (Fig. 4 ~ ) The proximal . flagellar segments of antenna 1 (and occasionally the accessory flagellum) tend to be conjoint or fused (Figure IH), and the peduncle of antenna 2 longer (Figure IG), but the body size smaller than in the female (Figure 11). Type 3 males are almost invariably non-pelagic in body form, and are only slightly sexually dimorphic in the form of antennae, peraeopods, and uropods and eye size, but the body is usually larger than in the female and the gnathopods (Figure 13, N ) are moderately dissimilar and moderately to strongly amplexing and sexually dimorphic. Calceoli may be present on the flagellar segments of antenna 1 and the peduncle and flagellum of antenna 2 (Figure l ~L ,, M). In Gammaroidea, Eusiroidea, and Oedicerotoidea the calceoli are plate- or cup-(flask)-shaped (Figure IF, J , K , L) but in Crangonyctoidea calceoli are elongate, club- or barrelshaped (Figure l ~N),, somewhat as in the Phoxocephaloidea (Figure 1 ~ )In some Gammaroidea and Eusiroidea antennae of the Q may also . be calceolate. Type 4 males (Figure 10) are almost exclusively non-pelagic, benthic, fossorial, tubicolous, or hypogean, usually much larger than females (Figure IQ) in body size, and the gnathopods are usually very dissimilar, strongly amplexing, and strongly sexually dimorphic (Figure 4c, D , E). Antenna 1 may retain traces of brush setae, but more often its peduncular segment 3 and proximal flagellar segments bear aesthetascs (e.g. Liljeborgioidea), slender, rod-like, sensory structures (Figure lp). Other superfamily characters may vary as follows: The body may be smooth or carinate (Figure 2 ~c), or processiferous (Figure 2 ~ ) and the urosome may be smooth or toothed (occasionally , sexually dimorphic; Figure 2 ~ B) or bear dorsal groups of spines or , spinules (Figure 2 ~ )and some segments may be fused (Figure 2 ~ ) . , The dorsal anterior head margin may be prolonged into a strong (Figure 2 ~N), or weak (Figure 2 ~I),rostrum. The eye (usually paired,
FIGURE
2. Morphological types of body and urosome armature, A-F; rostrum and eye type, G-o; coxal plates, P-U.
Bousjield: Revised Class$cation of Amphipod Crustaceans 351
lateral) may be reniform (Figure 2 ~ ) subrectangular (Figure 21), or , rounded (Figure 2 ~ ) dorsally confluent (Figure 2 ~ ) , the rostrum ; on (Figure 2 ~or on the interantennal head lobe (Figure 20). It may be very ) large, covering most of the head (Figure 2 ~ ) small (Figure 2 ~ ) single , , , facets with cornea1 lenses (Figure 2 ~ )or lacking. The mouthparts are termed "basic" when plates and palps are more or less fully developed, fully setose and spinose, and the mandibular (Md) molar is present (Figure 3 ~ F, J , K , o , S, W); and termed , "modified" or "derived" when plates and palps are small, weakly setose and spinose, and fused (or enlarged) and mandibular molar andlor palp is reduced (Figure 3 ~M) or vestigial (Figure 3c, H , M, N , P, U , z). , The upper lip (UL)may be apically smooth (Figure 3 ~or notched, the ) , lobes often assymetric (Figure 3 ~ c). The lower lip (LL) is primitively , tall and lacks inner lobes (Figure 3 ~ F) but may be broad, with inner lobes that are strong and distinct (Figure 3 ~ G,, H) or variously fused (Figure 31). Maxilla 1 (Mx l), inner plate is basically marginally setose; the outer plate has 11 apical spine-teeth, and the palp is strong and 2-segmented (Figure 30); apomorphically, the inner plate is small, weakly setose, or lacking (Figure 3q, R), the outer plate has fewer (7-9) spine-teeth, and the palp may be l-segmented (Figure 3 ~ or vestigial (Figure 3p), or ) . greatly enlarged (Figure 3 ~ ) In basic maxilla 2 (Mx 2), the inner and outer plates are broadly developed, marginally apically setose, and the inner plate has a row of facial setae (Figure 3s), but in the derived condition, the plates are narrow or small, and setose apically only (Figure 3 ~U), or very large and setose (Figure 3v). The basic maxilliped , (Mxpd) has well-developed, setose and spinose, inner and outer plates, and a 4-segmented dactylate palp (Figure 3w). In the derived condition, the plates may be variously reduced (or fused to the opposite member) and the palp strong (Figure 3z), or the outer plate may be enlarged (Figure 3x) or the palp reduced (Figure 3 ~or lacking (Figure 3aa). ) Anterior coxal plates are usually deep and may be very enlarged r . (figure 2 ~o) small and barely separated (Figure 2 ~ ) They may overlap each other anteriorly, and their attachment corners may be overlapped by body plates (Figure 2p, S) but may fit exactly with corresponding segments as in the plesiomorphic Stegocephalidae (Figure 2u). Coxa 4 is ) usually strongly excavate proximally on the hind margin (Figure 2 ~but may be weakly (or not) excavate as in Corophioidea (Figure 2 ~ R). , Posterior coxal plates (5-7) have the posterior lobe larger and deeper , (postero-lobate (Figures 2 ~4 ,~ ) subequal in development (equilobate) (Figure 4 ~ ) or the anterior lobe larger and deeper (anterolobate) (Fi, , gures 2 ~R, 40). The corresponding peraeopods (walking limbs) may be subequal in size and form, with broadly developed bases (segment 2),
FIGURE
3. Morphological types of mouthparts: upper lip, A-c; lower lip, mandible, J-N; maxilla 1, o-R; maxilla 2, S-v; maxilliped, W-z, aa.
D-I;
Bousfield: Revised ClassiJication ofAmphipod Crustaceans 353
and posterior-distally produced angle or corner of segment 4 and all segments non-setose (homopodous condition, Figure 4 ~ ) Heteropo. dous limbs are rather unequal in size and form, with bases typically narrow or unexpanded, and 4th segment not produced behind, and the limb segments often strongly setose as well as spinose (Figure 4 ~o ,,P). Peraeopods 3 and 4 are normally directed forward, and may be raptorial ) (Figure 45). Segment 2 (Corophioidea, Figure 4 ~ or 4 (Ampeliscoidea, Figure 4 ~may house cement-secreting glands that empty through ducts ) in the slender, elongate dactyls. Fossorial (burrowing) limbs have very broadly expanded and strongly spinose and/or setose segments (Figure 4 ~ P). , Coxal gills are sac-like, leaf-like, simple (Figure 5c), pleated (Figure 5 ~ B), or lobate (Figure 5 ~ c) respiratory structures attached to the , , inner (median) surface of the coxa, outside the brood plate (in 9 ) and may occur on all peraeopods except peraeopod 1. When present, the gill on the coxa of peraeopod 7 is usually much smaller than the anterior gills. Sterna1 gills (small, finger-like, simple or furcated respiratory processes) may project from the sternum of some thoracic (peraeonal) segments in species of freshwater families (e.g. Crangonyctidae, Hyalellidae) (Figure 5 ~ ) . Brood plates ( Q only) are laminar processes attached medially to the coxae of peraeopods 2-5 (or fewer) that are typically broad and marginally multisetose (Figure 5 c , H) but may be linear or sublinear and sparsely setose (Figure 51). Marginal setae may be simple, or hooktipped (Figure 5 ~ ) . Pleopods (first three pairs of abdominal appendages), used for swimming and setting up respiratory currents, normally have unexpanded peduncles and slender equal multisegmented rami (Figure 5 ~ )but may , have very powerfully developed (muscular) or broadened peduncles and long rami in free-swimming and free-burrowing types (Figure 5 ~ or ) much reduced and vestigial rami and peduncles, especially in hypogean or terrestrial species (Figure 5 ~ )Anterior pleopods of a few species of . ) Talitroidea and Bogidielloidea are sexually dimorphic (Figure 5 ~ and presumably assist in sperm transfer in the male. The rami of uropods 1 and 2 may be lanceolate (acutely pointed) (Figure 5 ~Q) or linear and tipped with spines (Figure 50, P). Rami may , be unequal in length (Figure 50). Uropod 3 rami are usually lanceolate (Figure 5 ~S),, and foliaceous (bearing plumose marginal setae) (Figure 5 ~but may be linear and spinose (Figure 5 ~or hook-tipped (Figure 5 u , ) ) v). The outer ramus is usually longer than the inner and may be 2segmented (Figure 5 ~S), or l-segmented by fusion or loss of the small terminal segment (Figure 5 ~T). , The telson, of fundamental importance in amphipod classification, consists of two posteriorly directed lobes that are fused proximally, and
FIGURE 4. Morphological types of gnathopods 1 and 2 (pairs), A-E; gnathopod 1, F; gnathopod 2, G-I; peraeopod 3 and 4, J-L; peraepods (and coxae) 5-7, M-P.
Bousfield: Revised Classification ofAmphipod Crustaceans 355
F I G U R E 5.
<,-I;
Morphological types of coxal gills, A-E; sterna1 gills, F ; brood plates, pleopods, J-M; uropods 1 and 2, N-P; uropod 3, Q-v; telson, W-Z, aa, bb.
separated narrowly (Figure 5aa, bb) or broadly (Figure jx), or fused . entirely (Figure 5w, z) or separated entirely (Figure 5 ~ ) The apex of each lobe may be sharply notched with small inserted spine(s) (Figure 5bb), or may be truncated or rounded, with several stout spines (Figure 5x, Y). When entire, the margin may be spinose (Figure 5w - hook , spines), minutely setose, or bare (Figure 5 ~z).
SYSTEMATIC SECTION
Superfamily Phoxocephaloidea Sars 1891 (new status) Diagnosis: Plesiomorphic, smooth-bodied, rostrate, fossorial gammarideans usually having strongly dimorphic terminal pelagic male bearing sensory brush setae and calceoli on peduncular segments of antennae 1 and 2, and calceoli on flagella esp. antenna 2; accessory flagellum strongly developed; mouthparts modified, usually with weak mandibular molar and palp, weakly armed plates of maxillae 1 and 2, and small inner plates of maxilliped; lower lip, inner lobes well developed; coxal plates deep, 4th large, strongly excavate behind; coxae 5-7 posteriorly lobate; gnathopods 1 and 2 non-amplexing, subsimilar, subchelate or chelate; peraeopods 5-7 dactylate, heteropodous, often markedly so; brood plates linear; coxal gill usually present on peraeopod 7; pleopods normal to powerful; uropods lanceolate, rami of 1 and 2 subequal; uropod 3 foliaceous, outer ramus 2-segmented; telson lobes deeply separated (or fused to a notch), apices with minute notch and spine. Composition: Phoxocephalidae Sars 1891, revised Barnard and Drummond 1978; Platyischnopidae Barnard and Drummond 1979; Urothoidae new family. Family Urothoidae new family Closely allied to Phoxocephalidae, but rostrum not "shark-snouted" nor produced as cephalic hood over base of vertically inserted antennae; molar process moderately to strongly developed; gnathopods weak, subequal, variously subchelate; pelagic terminal male stage suppressed in most genera; peraeopod 5 distinctly 7-segmented; peraeopods 6 and 7 subsimilar in form and size, strongly fossorial; abdomen not sharply narrowing or flexed at urosome; uropod 3 rami subequal; telson lobes separated to base. Composition: Urothoe Dana 1852 (type genus) (type species U. irrostratus Dana 1853); Carangolia Barnard 1961 (type - Carangolia mandibularis J.L. Barnard 1961); Cunicus Griffiths 1974 (type - Cunicus
Bousfield: Revised Classification ofAmphipod Crustaceans 357 profundus Griffiths 1974; Haustoriella K.H. Barnard 1931 (type Haustoriella psammophila Barnard 1931); Phoxocephalopsis Schellenberg 1931 (type - Phoxocephalopsis zimmeri Schellenberg 1931); Urothopsis Ledoyer 1969 (type - Urothopsis spinidigitus Ledoyer 1969); Urohaustorius Sheard 1936 (type - Urohaustorius halei Sheard 1936); Urothoides Stebbing 1891 (type- Urothoides lachneesa Stebbing 1888); Zobracho Barnard 1961 (type - Zobracho canguro J.L. Barnard 1961). Some strongly fossorial members of this family (e.g. spp, of Phoxocephalopsis, Urohaustorius) remarkably and convergently resemble Haustorius (Haustoriidae). In urothoids, peraeopod 5 is 7\egmented and segments 5 and 6 are stoutly expanded, whereas in haustoriids peraeopod 5 is 6-segmented and segments 4 and 5 are expanded. Ilistribution: Marine coastal and shelf to abyssal waters of antiboreal continental margins, largely replaced by Pontoporeioidea in the holarctic. Relationships: Except for specialized mouthpart morphology and fossorial peraeopods, the general body form and reproductive male stage, as in the Lysianassoidea, is close to the presumed ancestral gammaridean type. The superfamily is superficially similar to the Pontoporeioidea but differs markedly in the more plesiomorphic condition of the urosome, gills, gnathopods, and coxal plates, and peraeopods, and in their hathymetrical (to the abyss) and geographical (largely antiboreal) ranges. The Platyischnopidae appear divisible into two main types based on mouthpart morphology and form of the pelagic terminal male. The Indo-Pacific form (e.g. P. hevdmani Walker and P. capensis K.H. Barnard) has strongly developed and marginally spinose maxilliped outer plate and strong mandibular molar, and antenna 1 ( 8 ) is elongate. The neotropical form (P. capuciatus Oliveira, P. metagracilis J.L. Barnard) has reduced mouthparts, very similar to most phoxocephalids, and antenna 2 ( 8 ) is elongate. The form of the telson, uropod 3, and peraeopods 3 and 4 is remarkably similar to that of Bathyporeia in the Pontoporeioidea. Superfamily Lysianassoidea Dana 1849 (new status) Diagnosis: Plesiomorphic, mainly smooth-bodied, weakly rostrate, often fossorial gammarideans, usually with strongly dimorphic terminal pelagic male bearing sensory brush setae on peduncular and basally conjoint flagellar segments of antenna 1, and both calceoli and brush
setae on antenna 2; antenna 1 with strongly inflated peduncular segment 1, short segments 2 and 3, and strong accessory flagellum; eyes deep reniform, bulbous below; mouthparts highly modified; upper lip notched, epistome often anteriorly produced; lower lip tall, lacking inner lobes; mandibular molar reduced, palp slender; maxilla 1 inner plate narrow, 2-setose; outer plate with 9 apical spine-teeth; maxilla 2, plates narrow, weakly setose; maxilliped outer plate often enlarged, with strong cutting margin; coxal plates large, deep, 4th strongly excavate; coxae 5-7 posteriorly lobate; gnathopods 1 and 2 non-amplexing, markedly dissimilar, 1 variously subchelate, chelate or simple, 2 with characteristic elongate segment 3 and small microchelate propod and dactyl; peraeopods 5-7 homopodous, bases expanded; brood plates sublinear; coxal gills often pleated, present on peraeopod 7; pleopods normal; uropods lanceolate; rami subequal; uropod 3 foliaceous, outer ramis 2-segmented; telson lobes usually distally separated (narrowly), apices with minute notch and spine, or fused and entire, margin not spinose. Composition: Lysianassidae Dana 1849. Distribution: Exclusively marine, mainly benthonic carnivores and scavengers or inquilines, coastal to abyssal and hadal, world-wide but mainly arctic and antarctic. Relationships: This very large, morphologically and ecologically diverse family consisting of over 100 recognized genera and several hundred species, is urgently in need of revision. Subfamilies Lysianassinae and Uristinae proposed by Hurley (1963) could justifiably be elevated to family status. Separate family andlor subfamily recognition might be given to Cyphocaris and pelagic allies, to Trischizostoma and other fish parasites, and to several highly modified species living commensally in sponges, tunicates, and coelenterates. The superfamily is allied to the Pardaliscoidea, Phoxocephaloidea, and Stegocephaloidea, near the Gammaridean root stock. Superfamily Pontoporeioidea Sars 1882 (new status) Diagnosis: Plesiomorphic, generally smooth bodied, weakly rostrate, fossorial gammarideans; dimorphic terminal pelagic male often suppressed (lacking in Haustoriidae) but, when present, bearing plate calceoli and usually brush setae on flagella of both antenna l and 2; accessory flagellum short; mouthparts basic, with strong mandibular molar and palp; plates of maxillae 1 and 2 strongly setose and spinose; maxilliped plates and palp well developed and strongly setose; lower lips with
Bousfield: Revised Classijication ofAmphipod Crustaceans 359
distinct inner lobes; coxal plates deep, 4th moderately excavate behind; coxa 5 anteriorly or equi-lobate, coxae 6 and 7 posteriorly lobate; gnathopods 1 and 2 weakly amplexing, unlike, subchelate or simple; peraeopods 5-7 adactylate (or nearly so), markedly heteropodous; brood plates sublinear to broad; coxal gill lacking on peraeopod 7; sterna1 gills occasionally present (some Pontoporeia); pleopods normal to very powerful; uropods sublinear or lanceolate, rami of 1 and 2 subequal; uropod 3 foliaceous to spinose, outer ramus usually 2segmented; telson lobes variously separated or fused, apices with spines or setae. Composition: Pontoporeiidae Sars 1882; Haustoriidae Stebbing 1906. AS currently restricted, the revised family Pontoporeiidae contains the following genera: Pontoporeia Kroyer 1842, Priscillina, Stebbing 1888, Amphiporeia Shoemaker 1929, and Bathyporeia Lindstrom 1855. The Haustoriinae (emend Bousfield 1965) (North American genera) is herewith elevated to family status.
NOTE:
Distribution: Almost exclusively limited to shallow coastal marine, brackish, and fresh waters of temperate holarctic regions, south in the Americas to Ecuador, and to South Africa. Relationships: In most characters the family Haustoriidae seems readily derivable from the Amphiporeia-Bathyporeia branch of the family Pontoporeiidae. However, the Pontoporeia-Priscillina branch differs in having an elongate antenna 1, and more apomorphic condition of the coxal plates, peraeopods, uropods, and telson, and broader brood plates. These features, combined with the weakly spinulose urosome, suggest a close relationship to family Gammaridae within the Gammaroidea. Superfamily Gammaroidea Leach 1914 (new status Bousfield 1977) Diagnosis: As defined by Bousfield (1977). Composition: *Acanthogammaridae Garjajeff 1901; Anisogammaridae Bousfield 1977, revised 1979; Gammaridae Leach 1814; Gammaroporeiidae new family; Macrohectopidae Sowinsky 1915; Mesogammaridae Bousfield 1977, revised 1979; Pontogammaridae Bousfield 1977; Typhlogammaridae new family; Behningiella-Zernovia (parasitic) complex?; Zphigenella-Pachyschesis complex.
* Family Caspicolidae Birstein
1945 is removed to Incertae Sedis by Bousfield (1977).
Family Typhlogammaridae new syn: Gammaroidean family group 2 Bousfield (1977) Diagnosis: As defined by Bousfield (1977), with component genera. Type genus: Typhlogammarus Schaferna 1906. Family Gammaroporeiidae new family syn: Gammaroidean family group 10, Bousfield (1977) p. 296 Diagnosis: As defined by Bousfield (1977) p. 296. Type genus: Gammaroporeia new genus. Diagnosis: With the characters of the family. Type species: Micruropus alaskensis Bousfield and Hubbard 1968. Distribution: Coastal cold-water estuarine and brackish waters, southeastern Alaska to west-central British Columbia (Queen Charlotte Sound). Relationships: The Gammaroidea is an essentially fresh- and brackishwater, mainly epigean superfamily group of the Northern Hemisphere distinguished from epigean crangonyctids by the more basic mouthparts, stouter and more setose antennae, longer accessory flagellum, cup- or plate-shaped (rather than club- or barrel-shaped) calceoli borne only on flagellum (rarely peduncle of antenna 2) by the non-pelagic male, by the dissimilar, more strongly amplexing gnathopods, generally heteropodous-setose peraeopods, strongly dorsally spinulose urosome, foliaceous uropod 3, and deeply separate telson lobes that are apically rounded and strongly spinose. Gammaroideans apparently diverged very early from the gammaridean ancestral stock, perhaps close to Pontoporeioidea, and radiated within cold fresh waters of the Northeqn Hemisphere. A third of the genera and about one-quarter of the species are endemic to Lake Baikal, and more than 70 per cent of the world species are exclusively freshwater. Superfamily Crangonyctoidea Bousfield 1977 Diagnosis: As given by Bousfield (1977). Composition: Neoniphargidae Bousfield 1977; Paramelitidae Bousfield 1977; Crangonyctidae Bousfield I973 (revised Holsinger 1977); Pseudocrangonyx-Sternophysinx family group (Bousfield 1977). Distribution: Holarctic and antiboreal continental fresh waters, epigean and hypogean, but lacking in South America, central Africa, Madagascar, and tropical continental regions. Sporadic in Eurasia, but dominant
Bousjield: Revised Class$cation of Amphipod Crustaceans 361
in North American fresh waters. The world distribution is essentially similar to that of the Astacura (freshwater crayfishes).
Relationships: The hypogean group of Pseudocrangonyx (Korea, Japan), Paracrangonyx (New Zealand), Procrangonyx (Japan), and Sternophysinx (South Africa), previously aligned with the Bogidielloidea (Bousfield 1977), is herewith transferred to the Crangonyctoidea on the basis of the more similar condition (especially to hypogean members of Crangonyctidae) of mouthparts, peraeopods, uropods, and telson and presence of sternal gills. Their convergently "bogidielloidean" features of elongate body, small coxal plates, enlarged gnathopod l , reduced pleopods, linear brood plates, and lack of coxal gill on peraeopod 7, however, would justify separate family recognition. The enigmatic family group represented by the New Zealand brackish and freshwater genus Phreatogammarus is tentatively reassigned to the gammaroideans. Although epigean males apparently lack calceoli (recent observation), in gross body morphology, reniform eye shape, elongate antennae, strong accessory flagellum, basic mouthparts, anterolobate coxae, heteropodous peraeopods, dorsally spinose urosome, and separated strongly spinose telson lobes, this small group is strongly similar to gammaroideans (e.g. Mesogammarus), all known members of which are confined to the Northern Hemisphere. However, the enlarged \pinose gnathopods, linear rami of uropod 3 (of which the outer is the shorter and one-segmented), and short uropods 1 and 2 are reminiscent of Bogidielloidea, whereas the broad brood plates, rounded epimeral plates, and presence of sternal gills (in one hypogean species) are more characteristic of Crangonyctoidea. Epigean members of Phreatogammarus differ in several important characters, especially those given by Hurley (1954) and the presence of a baso-facial spine on uropod 1, that in combination would justify their separate generic recognition. Crangonyctoideans appear to be a very ancient continental freshwater group, in distribution scarcely overlapping with bogidielloideans, and are perhaps in the process of replacement by gammaroideans in Eurasian surface fresh waters, but not in North America where freshwater gammaroideans are less common and little diversified. Crangonyctoideans are most readily derivable from a pontogeneioidean-like marine stock. Superfamily Niphargoidea Karaman 1962 (new status Bousfield 1977) Diagnosis: As outlined by Bousfield (1977). Composition: Niphargidae Karaman 1962; PseudoniphargusAllocrangonyx complex?; Austroniphargus complex?
362 The Royal Society of Canuda
Distribution: Essentially European and western Asiatic, south of Pleistocene glaciation, with outliers in Madagascar, Madeira, and eastcentral United States; exclusively hypogean, mostly in fresh but also in brackish water. Relationships: This very large, remarkably uniform, essentially European hypogean family, containing nearly 250 species in about 15 generic and subgeneric groupings, awaits revision by Niplzavgus taxonomists. Formal family andlor subfamily groupings are justifiable, based on such high-level characters as the degree of similarity of the gnathopods, antero- or postero-lobate condition of the coxae, heteropodous or homopodous state of the peraeopods, the degree of development and setation, andlor reduction of uropod 3, and reduction of the pleopod rami. Certain Karamaniella-type niphargids, capable of rolling into a ball, resemble salentinellids (Liljeborgioidea) but the resemblance appears convergent and superficial. In general aspect of mouthparts, gnathopods, brood plates, uropod 3, and telson, the similarity of niphargids to plesiomorphic crangonyctids (e.g. Neoniphargidae) is especially strong. However, the absence of coxal gills on peraeopod 7 and sterna1 gills on any segment, combined with the well-developed dorsal abdominal spine groups, excludes the family Niphargidae from the present diagnosis of Crangonyctoidea. Superfamily Bogidielloidea Hertzog 1936 (new status Bousfield 1977) Diagnosis: As defined by Bousfield (1977). Composition: Bogidiellidae Hertzog 1936; Spelaeogammavus complex. Distribution: Exclusively hypogean in warm temperate and tropical fresh waters of all continents and continental islands (except Antarctica); the only primary freshwater amphipods known from central Africa and Central and South America, exclusive of the epigean Hyalellidae. Relationships: These highly apomorphic, subvermiform, eyeless, exclusively hypogean gammaridean amphipods bear certain resemblances to crangonyctids and to niphargids but the form of the gnathopods, uropods, and telson suggests a primary ancestral type near to Phreatogammarus. As with crangonyctids and niphargids, a few members occur in brackish waters close to the sea, or in former marine embayments, but none are fully marine (as are some members of the suborder Ingolfiellidea with which bogidiellids frequently occur in hypogean fresh waters).
Bousjield: Revised ClassiJication ofAmphipod Crustaceans 363
Superfamily Eusiroidea Stebbing 1888 (new status) Diagnosis: Plesiomorphic, variously carinate or processiferous, rostrate, free-swimming and epibenthic amphipods, usually with moderately dimorphic terminal male stage; peduncles and flagella of antennae 1 and 2 bearing calceoli and less often brush setae; accessory flagellum small, vestigial, or lacking; eyes large, subrectangular; mouthparts more or less basic; upper lip without pronounced marginal notch; lower lip, inner lobes lacking or weakly developed; mandibular molar and palp usually strong; inner plates of maxilla 1 and 2 marginally setose, outer plate of maxilla 1 with 11 with apical spine teeth; maxilliped plates and palp well developed, setose; coxal plates medium deep, 4th excavate; anterior peraeonal segments short, abdomen (especially pleon) segments large; coxae 5-7 posteriorly lobate; gnathopods 1 and 2 non- (or weakly) amplexing, subsimilar, usually weakly subchelate (gnathopod 1 vestigial in Bateidae), occasionally large and raptorial (Eusiridae); peraeopods 5-7 basically homopodous; brood plates large, broad; coxal gills simple, present on peraeopod 7; sterna1 gills occasionally present (some Pontogeneiidae); pleopods well developed, often powerful; uropods lanceolate, rami of 1 subequal, of 2 unequal; uropod 3 , rami foliaceous, outer ramus 1-segmented; telson lobes distally separated, usually narrowly, apices with small notch and spine (seta), or entire. Composition: Eusiridae Stebbing 1888; Pontogeneiidae Stebbing 1906; 13ateidae Stebbing 1906; Gammarellidae Bousfield 1977; Calliopiidae Sars 1893; Paramphithoidae Stebbing 1906; Amathillopsidae Pirlot 1934. Ilistribution: Mainly north Pacific and antiboreal, coastal to shelf marine, brackish and fresh water (Paramoera, Pseudomoera) and hypogean (Awacaris? Paramoerella). Relationships: The Paramphithoidae appears to be a polyphyletic as\emblage in need of critical revision. The type genus and species of this family, Paramphithoe hystrix Ross 1835, is closely related to Calliopiidae in mouthpart morphology and general body characters, but is less close to Stegocephaloidea and Pleustidae. Amathillopsis is herewith reassigned to its own family. Eclysis is clearly an Astyridae. Superfamily Oedicerotoidea Lilljeborg 1865 (new status) 1)iagnosis: Moderately apomorphic, strongly rostrate, strongly fossorial rtlilrine amphipods having a moderately dimorphic terminal male stage; I ~ L I Ssetae borne on peduncles of antennae 1 and 2 or conjoint flagellum ~ of antenna 1, and occasionally calceoli on flagellum of 2; accessory tl:~gellumlacking or vestigial; eyes large, occasionally lateral, but usu-
ally fused mid-dorsally at base of rostrum; mouthparts basic but somewhat modified; upper lip, margin not distinctly incised; lower lip broad, inner lobes well developed; mandibular molar variable, usually strong, palp slender; maxilla inner plates weakly setose, apex of maxilla 1 outer plate with eight spine-teeth; maxilliped outer plate and palp strong; coxal plates medium, 4th not strongly excavate; coxae 5 and 6 deep, equi-lobate; gnathopods 1 and 2 non- (or weakly) amplexing, subsimilar, subchelate (or chelate), carpal lobes usually prominent; peraeopods 5 and 6 subsimilar, homopodous; peraeopod 7 large, dissimilar, segments elongate; brood plates linear; coxal gill usually lacking on peraeopod 7; pleopods strong; uropods narrow lanceolate, rami subequal; uropod 3 , rami non-foliaceous, outer ramus l-segmented; telson lobes fused (or nearly so), distal margin nearly bare. Composition: Oedicerotidae Lilljeborg 1865. Distribution: Coastal marine and brackish (to fresh) waters of all continents, offshore, shelf, and abyssal, especially diverse in cold waters of the northern hemisphere. Relationships: This large, rather compact family may be polyphyletic, and requires further categorization. A few plesiomorphic relict genera and species of the Southern Hemisphere (e.g. Exoediceros, Metoediceros, Patuki, Bathyporeiapus) have lateral eyes, non-pelagic males, calceolate antennae, and amplexing gnathopods, possibly secondarily derived as in corresponding members of fossorial Pontoporeiidae (Amphipoveia, etc.) in North American Atlantic waters. The antiboreal brackish and freshwater genus Paracalliope resembles oedicerotids in the short antenna 1, elongate form of peraeopod 7 and lack of coxal gill, and slender lanceolate uropods; however, its basic mouthparts, calceolate antenna 1, and unmodified homopodous peraeopods 3-6 and broad brood plates are eusiroidean (Calliopiidae). Its peculiar dimorphic amplexing gnathopods and subrotund lateral eyes, in combination with the above characters, would justify recognition in a separate family, transitional between calliopiids and oedicerotids and confirming the probable phylogeny of the Oedicerotoidea (see Barnard 1972b). Kanaloa Barnard 1970, also transitional, has unique gnathopods and elongate uropod 3. Superfamily Leucothoidea Dana 1852 (new status) Diagnosis: Apomorphic, rostrate, smooth-bodied, occasionally processiferous, crawling, clinging, or inquilinous marine gammaroideans, lacking a pelagic terminal male stage; antennae various, lacking calceoli or brush setae; peduncle of antenna 2 often strong; accessory flagellurn
Bousfield: Revised Clussijication of Amphipod Crustaceans 365
minute or lacking; eyes rounded, medium, lateral; mouthparts highly modified, apomorphic; upper lip with distinct median notch, lobes usually asymmetric; lower lip broad, inner lobes usually well developed or medially fused; mandible, molar usually small or lacking, incisor and Iacinia strong, palp slender, reduced, or lacking; maxillae small, inner plates nearly bare; maxilla 1 outer plate with 6-7 apical spine teeth; maxilliped, plates small, inner may be fused to opposite member, palp strongly dactylate; coxal plates various, usually deep, 4th often very broadly expanded, excavate behind; coxae 5-7 posteriorly lobate; gnathopods 1 and 2 usually moderately to strongly amplexing, often strongly dissimilar, subchelate, chelate, carpochelate, or simple; peraeopods 5-7 homopodous, bases variously expanded or linear; brood plates large, broad; coxal gills simple, lacking on peraeopod 7; pleopods normal; uropods lanceolate, rami of 2 and often 3 markedly unequal; uropod 3, peduncle often elongate, outer ramus l-segmented (2-segmented where inner ramus lacking), non-foliaceous; telson entire, ~ ~ s u a lwith median ventral keel, distal margin bare or nearly so. ly Composition: Pleustidae Buchholz 1874; Laphystiopsidae Stebbing 1899; Amphilochidae Boeck 1871; Leucothoidae Dana 1852; Anamixidae Stebbing 1897; Maxillipiidae Ledoyer 1973; Colomastigidae Stebbing 1899; Pagetinidae K.H. Barnard 1932; Nihotungidae .I.L. Barnard 1972; Stenothoidae (including Metopidae) Boeck 1971; Cressidae Stebbing 1899; Thaumatelsonidae Gurjanova 1938. This heterogeneous assemblage of families is difficult to classify because of their highly modified body form and mouthparts, presumably in response to member exploitation of highly diverse habitats and food resources. The group may prove to be polyphletic, with pleustids and laphystiopsid fish parasites derived from pontogeneoidean ancestors, the amphipolochid-leucothoid-anamixids from stegocephalid types, and the remainder (often with fused urosome segments) from pleustids. Distribution: World-wide coastal marine, mainly high salinity waters; especially diverse in the North Pacific and Antarctic regions; some fiimilies parasitic or commensal on fishes (Laphystiopsidae), hydroids (Stenothoidae), sponges (Leucothoidae, Anamixidae, Colomastigidae), li~nicates(Leucothoidae), or utilizing plant debris (Pleustidae, Amphilochidae). 3trpcrfamily Talitroidea Costa 1857 (elevated Bulycheva 1957) F,firrpnosis: Apomorphic mostly smooth-bodied, arostrate, benthonic f%iu ine, freshwater and terrestrial gammarideans, lacking a pelagic termini11 male stage, calceoli, or brush setae on antennae; antenna 1 usually
shorter than 2, accessory flagellum totally lacking; peduncle of 2 occasionally elongate (Talitridae); eyes various, usually rounded, subrectangular, lateral, occasionally large (Talitridae); mouthparts modified; upper lip rounded below; lower lips tall, lacking inner lobes; mandible usually with strong molar and incisor, palp lacking; maxilla 1, inner plate slender with 2 (usually) apical setae, outer plate with 9 (typically) apical spine teeth; maxilla 2 distally setose; maxilliped plates well developed, palp 4th segment occasionally reduced or lacking; coxal plates medium deep, 4th strongly excavate, 2 and 3 often with posterior marginal process; coxae 5-7 posteriorly lobate; gnathopods 1 and 2 typically strongly amplexing, variously dissimilar, subchelate, chelate, or simple; peraeopods 5-7 homopodous, secondarily heteropodous in some Talitridae; brood plates basically broad, linear or lacking (in some Talitridae), marginal setae with hooked tips; coxal gills simple, occasionally lobate, lacking on peraeopod 7 ; sterna1 gills present in some Hyalellidae; pleopods normal, modified (Phliantidae, Eophliantidae), reduced, or vestigial (especially Talitridae); urosome segments 2 and 3 very short, telescoping dorsally; uropods linear, tips spinose, rami of 1 and 2 subequal; uropod 3 small, uniramous (or with minute inner ramus), 1segmented; telson lobes separated to base or variously fused, apices spinose. Composition: Hyalidae Bulycheva 1957; Hyalellidae Bulycheva 1957; Talitridae Costa 1857, revised Bulycheva 1957; Ceinidae Barnard 1972 (Ceininae elevated); Dogielinotidae Gurjanova 1954; Najnidae Barnard 1972; Eophliantidae Sheard 1936, revised Barnard 1972; Phliantidae Stebbing 1899, revised Barnard 1969; Temnophliantidae Griffiths 1975; Kuriidae Barnard 1964. The revision of Talitroidea proposed by Barnard (1972b), while contributing useful new information and taxonomic concepts, contains an unlikely and unrealistic treatment of the first four families listed above, and is retrogressive to the concept of Bulycheva (1957). This remarkable homogeneous superfamily group contains the Talitridae, the sole amphipod family with members that have become entirely terrestrial and able to complete their life cycle remote from the sea; most members seem yet dependent on trace amounts of sodium- or chloride-ion derived from rain or wind-blown sources and unable to withstand hard-frost winters (see Bousfield 1968). Terrestrial existence is accompanied by reduction of antenna l , maxilliped palp, brood plates, and pleopods, but has led to enlargement of antenna 2, peculiar modification of gnathopod 2 (cf. Lysianassidae), lobiform gills, a shortened urosome, and saltatory (jumping) behaviour. The relatively plesiomorphic families Hyalidae and Hyalellidae and component genera need comprehensive revision.
Bousjeld: Revised ClassiJication ofAmphipod Crustaceans 367
The Talitroidea probably diverged early from a pontogeneioid-like ancestral type, but specialized in exploiting large particulate decaying plant material as food. Distribution: Mainly marine, shallow water and intertidal, fresh water, and terrestrial, mainly in tropical and temperate regions, with greatest diversity in the Southern Hemisphere. Superfamily Stegocephaloidea Dana 1852 (new status) Diagnosis: Plesiomorphic, smooth or processiferous, strongly rostrate, free-swimming or benthonic gammarideans, lacking a terminal pelagic male stage; antenna 1, peduncle 1 may be enlarged, accessory flagellum small or lacking, basal flagellar segments may be conjoint (c?); antennae I and 2 lack calceoli and usually brush setae; eyes (when present) deep-reniform; mouthparts modified, more or less arranged in cone-like buccal mass: upper lip distally notched; lower lip tall, lacking inner lobes; mandibular molar lacking or vestigial, palp slender or lacking; plates of maxillae rather strongly setose and normally spinose; coxal plates deep, often acute below, exactly matching respective peraeon segments; 4th very strongly excavate behind; coxae 5-7 lobate posteriorly; gnathopods 1 and 2 non-amplexing, somewhat dissimilar, weakly subchelate or chelate; peraeopods 5-7 homopodous, bases variously expanded; brood plates broad; coxal gills simple, present on peraeopod 7; pleopods normal; uropods lanceolate, rami slightly unequal; uropod 3 foliaceous, or lanceolate-spinose; outer ramus usually 2-segmented; telson lobes usually separated distally, apices with minute notch and spine, or lacking. Composition: Stegocephalidae Dana 1852; Acanthonotozomatidae Stebbing 1906; Ochlesidae Stebbing 1910; Lafystiidae Sars 1893. Distribution: Coastal marine and shelf waters of holarctic and antiboreal regions, mainly colder waters; Lafystiidae are external parasites of groundfishes. Relationships: The Stegocephalidae have coxal plates that join exactly with their corresponding body segments (no overlapping angles), and unlike others of the superfamily lack mandibular palps and pigmented eyes and are smooth-bodied. The swollen peduncular segment 1, con.joint flagellar segments, and accessory flagellum resemble the condition in Lysianassidae, but the gnathopods, peraeopods, uropods, and telson are not unlike the synopiid-pardaliscid complex.
Superfamily Synopioidea Dana 1853 (new status) syn: Synopioid superfamily group, Bousfield 1977 (in part) Diagnosis: Plesiomorphic, rostrate, abdominally processiferous, epibenthic and pelagic gammarideans, having dimorphic terminal male stage; conjoint flagellar segment of antenna 1 and elongate peduncle of antenna 2 bear brush setae; calceoli lacking; accessory flagellum strong; eyes subrotund, often dorsally confluent, large; mouthparts basic, upper lip with shallow median notch, lower lip broad, inner lobes variously developed; mandibular molar strong, palp usually with reduced segment 3; inner plates of maxillae setose, outer plate of maxilla 1 with 9-1 1 spine teeth; maxilliped plates and palp well developed; coxal plates medium deep, 4th excavate; coxae 5-7 posteriorly lobate; peraeon segments short, abdomen (pleon) large; gnathopods 1 and 2 non-amplexing, subsimilar, subchelate or simple; peraeopods 5-7 essentially homopodous, bases posteriorly expanded; brood plates medium broad; coxal gill on peraeopod 7; pleopods normal to powerful; uropods 1 and 2 lanceolate, apices spinose, rami unequal; uropod 3 lanceolate, foliaceous, outer ramus 2-segmented; telson large, lobes distally separated (narrowly) or fused completely, apices with small notch and spine. Composition: Synopiidae Dana 1853 (including Tironidae Stebb. 1906); Argissidae Walker 1904. Distribution: Coastal to abyssal, marine, pelagic and epibenthic on soft substrata in all oceans, mainly tropical and temperate, mainly antiboreal. Relationships: In most features synopiids are close to the predicted gammaridean ancestral type. They show close affinities with pardaliscoideans, dexaminoideans, melphidippoideans, and with benthic brackish and freshwater liljeborgiids on the one hand, and with stegocephaloideans and phoxocephalids on the other. Superfamily Pardaliscoidea Boeck 1871 (new status) syn: Pardaliscoid superfamily group, Bousfield 1977 Diagnosis: Plesiomorphic, weakly rostrate, abdominally processiferous, pelagic marine gammarideans with dimorphic terminal male stage; conjoint flagellar segment of antenna 1 and peduncle of antenna 2 bearing brush setae; accessory flagellum usually strong, occasionally vestigial or lacking; eyes (when present) deep reniform, lateral; mouthparts modified: upper lip with distal notch, lobes asymmetrical; lower lip broad, inner lobes well developed or fused; mandibular molar vestigial
Bousjeld: Revised Classijkation of Amphipod Crustaceans 369
or lacking, incisor (and lacinia) strong, palp slender, terminal segment often short; inner plates of maxillae sparsely setose, outer plate of n~axilla with 7-8 apical spine teeth, palp often greatly enlarged; maxil1 liped inner plate reduced, outer plate large, palp strong; coxal plates \hallow, 4th not strongly excavate behind; coxae 5-7 equi- or anteriorly lobate; gnathopods 1 and 2 non-amplexing, subsimilar, subchelate or \imple, occasionally enlarged and raptorial; peraeopods 3 and 4 tend to hc raptorial; peraeopods 5-7 elongate, homopodous or heteropodous, txtses not broad1y expanded; brood plates broad; coxal gill on peraeopod 7; pleopods powerful; uropods lanceolate, tips weakly spinose, rami unequal; uropod 3 lanceolate, foliaceous, outer ramus 2-segmented; tclson lobes deeply separated (apices with notch and spines(s)).
('omposition: Pardaliscidae Boeck 1871; Stilipedidae Holmes 1908; tlyperiopsidae Bovallius 1886; Astyridae Pirlot 1934; Vitjazianidae 13irstein and Vinogradov 1955. Ilistribution: Mainly open ocean abyssal and hadal, pelagic, carnivorous :imphipods, penetrating deeper coastal waters (fiords), in all oceans. Relationships: This superfamily group is closely allied to the Synopioidea, except for its mouthpart morphology, and is close to the presumed ancestral type of the sub-order Hyperiidea. Some component deep sea families are yet poorly defined; extreme modifications of mouthparts and pereopods (often bizarre) tend to obscure natural relationships. Eclysis properly belongs in Astyridae. Vemana J.L. Barnard (1964), however, is a very distinct amphipod type, bearing only the most superficial resemblance to Vitjaziana. Vemana would serve as the type of unique new familly that combines both plesiomorphic (in antennae, mouthparts, and urosome) and apomorphic (in gnathopods or peraeopods, and coxal plates) features, and might be placed near the I ,ysianassoidea andlor Pontoporeioidea. Superfamily Liljeborgioidea Stebbing 1899 (new status) syn: Synopioid superfamily group (Bousfield 1977, in part) Iliagnosis: Plesiomorphic, weakly rostrate and weakly abdominally processiferous benthonic gammarideans, lacking a pelagic terminal mule stage; antennae lacking brush setae but antenna 1 flagellum often wit h elongateaesthetascs; accessory flagellum present, occasionally very i~sge; antenna 2 peduncle stout; eyes (when present) subrotund, lateral; fnrrttthparts modified; upper lip with median notch; lower lip, broad, Inner lobes variously developed or fused; mandibular molar vestigial or $&@king,incisor strong, palp slender, terminal segment often small;
maxillae inner plates weakly setose, maxilla 1 outer plate with 7-9 apical spine teeth, palp normal; maxilliped inner plate small, outer normal, palp strong; coaxal plates medium deep, 4th excavate behind; coxae 5-7 posteriorly lobate; peraeon segments normal; abdomen large; gnathopods 1 and 2 variable, weakly to strongly amplexing, subsimilar but occasionally markedly unlike (Sebidae); peraeopods 5-7 homopodous, basis strongly expanded; brood plates linear; coxal gill lacking on peraeopod 7; pleopods normal or reduced; uropods 1 and 2 lanceolate, subapically notched, rami unequal; uropod 3 lanceolate, non-foliacous, outer ramus 2-segmented or fused, inner ramus occasionally short or lacking; telson lobes distally separated (narrowly) or fused, apices with notch and spine. Composition: Liljeborgiidae Stebbing 1899; Sebidae Walker 1908; Salentinellidae Bousfield 1977. Distribution: Coastal marine or brackish, inquilinous in polychaete tubes, sponges, corals, etc. (Liljeborgiidae, Sebidae) or freshwater hypogean (Salentinellidae), mostly in tropical and temperate regions. Relationships: Except for the lack of a pelagic terminal male stage, this assemblage is remarkably similar to both synopioids (especially Argissidae) and pardaliscoids. The genus Seborgia (Sebidae) has characteristics that link all three member families of the superfamily. Superfamil y Dexaminoidea Leach 1814 (new status) syn: Dexaminidae and related families, Bousfield 1973 (emended) Diagnosis: Plesiomorphic, rostrate and abdominally carinate, nestling and fossorial marine gammarideans, having a dimorphic pelagic terminad male stage; peduncles of antennae 1 and/or 2 bear bush setae, but lack calceoli; antenna 2 peduncle elongate; accessory flagellum lacking (occasionally vestigial); eyes reniform or rounded, lateral; mouthparts somewhat modified; upper lip, apical margin slightly incised; lower lip, inner lobes variously (usually strongly) developed; mandibular molar strong, palp normal (or lacking); inner plates of maxillae weakly setose; outer plate of maxilla 1 with 11 apical spine teeth; maxilliped plates normal, palp reduced; coxal plates medium to deep, 4th excavate; coxae 5-7 equi- or anteriorly lobate; gnathopods 1 and 2 weakly amplexing, subsimilar, subchelate; peraeopods 5-7 heteropodous, bases various; brood plates medium broad; some coxal gills pleated or foliaceous, present on peraeopod 7; pleopods normal; urosome segments 5 and 6 coalesced; uropods lanceolate, rami of 2 unequal; uropod 3 foliaceous, outer ramus l-segmented; telson lobes deeply and narrowly separated, apices with shallow notch and spine.
Bousfield: Revised Classification ofAmphipod Crustaceans 371
Composition: Atylidae Lilljeborg 1865; Anatylidae Bulycheva 1955; Lepechinellidae Schellenberg 1926; Dexaminidae Leach 1814; Prophliantidae Nicholls 1940, revised Barnard 1969. Distribution: Marine coastal and abyssal; world-wide but mainly Pacific and antiboreal; inquilinous with tunicates (some Dexaminidae). Relationships: This very compact superfamily group is closely related to 9ynopiids but differs in the more apomorphic condition of the mouthparts and peraeopods, and the fused urosome segments, pleated gills, etc. The Anatylidae and Lepechinellidae are sufficiently distinct from related dexaminids and atylids, in both morphology and ecology, to warrant continued recognition as separate family units. Superfamily Ampeliscoidea Bate 1861 (new status) Diagnosis: Apomorphic, arostrate, tube-building, fossorial gammarideans, having strongly dimorphic terminal male stage; peduncles of antenna 1 and 2 and basal flagellar segments of 1 bear bush setae but lack calceoli; antenna 2 peduncle elongate, antenna 1 usually short, lacking i\ccessory flagellum; eyes consisting of two (or one) pairs of facets with cornea1 lenses; mouthparts somewhat modified; upper lip with distinct median notch, lower lip broad, inner lobes well developed; mandibular molar and palp strong; maxilla 1 inner plate bare, outer with 11 apical \pine teeth; maxilla 2, inner plate moderately setose; maxilliped plates normal, palp slightly reduced. Coxal plates deep, regular, 4th strongly excavate; coxae 5-7 anteriorly lobate; gnathopods 1 and 2 non~~rnplexing, rather dissimilar, weakly subchelate or simple; peraeopods 3 and 4 glandular, dactyls simple, with ducts; peraeopods 5-7 strongly heteropodous, bases variously expanded; brood plates linear; coxal gills pleated, lacking on peraeopod 7; pleopods normal; urosome segment 1 carinate, 2 and 3 coalesced; uropods lanceolate, rami subequal; uropod 3 foliaceous, outer ramus l-segmented; telson lobes deeply and narrowly separated, apices with minute notch and spine. (lomposition: Ampeliscidae Bate 1861. 1)istribution: World-wide, marine coastal and abyssal; building sac-like tubes in soft stable sediments. Relationships: The close morphological relationship of Ampeliscidae to tlexaminoidean amphipods is obvious. However, the structurally differu n t head, types of eyes, dissimilar, non-amplexing gnathopods, glanduI:\r peraeopods, linear brood plates, and lack of coxal gill on peraeopod 7 in combination are considered distinctive at the superfamily level. Un-
like the tube-building Corophioidea, ampeliscoids have cement glands located mainly in limb segment 4 of peraeopods 3 and 4 (rather than in segment 2) and the tubes are blind sacs constructed vertically (rather than horizontally) in the sediments. Since ampeliscoids are morphologically unlike corophioids, the tube-building capability has probably arisen independently and convergently in these two groups. Superfamily Melphidippoidea Stebbing 1899 (new status Bousfield 1977) Diagnosis: Apomorphic, rostrate, abdominally processiferous marine gammaroideans, usually with pelagic terminal male stage; brush setae on elongate peduncle of antennae 1 and 2; calceoli lacking; accessory flagellum present, variable; eyes reniform, lateral; mouthparts more or less basic: upper lip slightly notched distally; lower lip, inner lobes variously developed; mandibular molar strong, palp slender; inner plates of maxillae normally setose, outer plate of maxilla 1 with 9 spine teeth; maxilliped plates strong, inner margin of outer plate with strong spine-teeth, palp somewhat reduced; coxal plates shallow (deep in fossorial forms), 4th weakly excavate; coxae 5-7 anteriorly lobate; gnathopods 1 and 2 weakly amplexing (strongly so where terminal pelagic male lacking), dissimilar, subchelate or simple; peraeopods 5-7 heteropodous, basis variously expanded; brood plates linear; coxal gills simple, lacking on peraeopod 7; pleopods normal; uropods lanceolate, rami of 2 unequal, tips spinose; uropod 3 peduncle elongate, rami weakly foliaceous, outer ramus l-segmented or minutely 2-segmented; telson lobes distally separated, apices with broad notch and spine(s). Composition: Melphidippidae Stebbing 1899; Hornellia-Cheirocratus family group Bousfield 1977;Megaluropus family group Bousfield 1977. Distribution: Marine coastal and shelf, epibenthic and pelagic, mainly in tropical and temperate continental regions. Relationships: Megaluropus, despite its superficial and convergent similarity in coxal plates and other characters to Argissa, and retention of a small coxal gill on peraeopod 7, is clearly assignable to the Melphippoidea and not to the Synopioidea. The Cheirocratus-Casco group has lost the pelagic male stage, and would seem morphologically close to the predicted ancestral type of Melitoidea. Superfamily Melitoidea Bousfield 1977 Diagnosis: As defined by Bousfield (1977). Component families: Hadziidae Karaman (1932); Melitidae Bousfield 1973; Carangoliopsidae Bousfield 1977.
Bousjeld: Revised ClassiJication ofAmphipod Crustaceans 373
Distribution: Mainly in coastal marine and estuarine (brackish) waters of tropical and temperate regions, world-wide; hypogean in warm temperate brackish sinks and former marine embayment groundwaters; a few genera and species are abyssal. Relationships: This very large and morphologically diverse superfamily group may prove to be polyphyletic. Since the original designation of family and generic members (1977), several authors, especially J.L. Barnard and colleagues (Barnard 1976; Zimmerman and Barnard 1977), have organized the genera in related groups that may require recognition at subfamily andlor family level. Especially difficult to treat are the hypogean hadzioids, weckelioids, metaniphargoids, etc., occuring in groundwaters in (and adjacent to) the former Tethys Sea region. J.R. Holsinger and colleagues are pioneering the discovery of hypogean genera and species in southwestern United States and Mexico (personal communication) and Stock (1977) has recently revised the Hadziidae. However, failing alternative designation, the genus Metacrangonyx is retained in the Hadziidae (see Barnard 1976). Weckelids are reliably distinguished from bogidiellids (with which they often CO-occur North in American groundwaters) by their prognathous buccal mass and plesiomorphic mouthparts, small gnathopod 1, expanded bases of peraeopods 6-7, more elongate uropods 1 and 2, lanceolate rami of ilropod 3 , and more deeply divided but less spinose telson lobes. However, the weckelioids converge toward the basic gammaroidean superfamily morphology but differ in loss of accessory flagellum, eyes, mandibular palp, pedunculate coxal gill on peraeopod 7, and the presence of linear brood plates and markedly dissimilar gnathopods.
Superfamily Corophioidea Dana 1849 (new status J.L. Barnard 1973) Diagnosis: Apomorphic, arostrate, cylindrical, mostly smooth-bodied, tube-building gammaridean amphipods lacking a terminal pelagic male stage; antenna1 peduncles typically strongly developed and setose, lacking brush setae and/or calceoli; accessory flagellum variously developed or lacking; eyes rounded, lateral, often on interantennal lobes; mouthparts basic; upper lip slightly notched distally; lower lip inner lobes well developed; mandibular molar strong, palp strong, occasionally weak or lacking; maxilla 1, inner plate weakly setose, outer with 10 (often only 8 or 7) spine teeth; maxilla 2, inner plate setose; maxilliped plates and palp strong; coxal plates shallow, often basally separated, 4th not excavate behind; coxae 5-7 deeply antero-lobate; gnathopods 1 and 2 strongly amplexing, dissimilar (often grossly), subchelate, carpochelate, or simple; peraeopods 3 and 4 glandular (in basis), ducting through simple dactyls; peraeopods 5-7 heteropodous; bases usually not expanded; brood plates usually broad; coxal gills simple, lacking on
peraeopod 7, occasionally also on peraeopod 2; pleopods normal, peduncles occasionally modified or reduced; rami of uropods linear, of uropod 2 unequal; uropod 3, rami short, linear, never foliaceous, outer ramus l-segmented (rarely 2) often tipped with hook spines; telson short, lobes usually totally fused distally and fleshy (except Biancolinidae), apex with cusps or hooks. Composition: Photidae Boeck 1872;Isaeidae Dana 1853; Ischyroceridae Stebbing 1899; Ampithoidae Stebbing 1899; Biancolinidae Barnard 1972; Aoridae Stebbing 1899; Cheluridae Allman 1847; Corophiidae Dana 1849; Podoceridae Stebbing 1906. Remarks: Barnard (1973)has usefully transferred Ericthonius, Cerapus, and relatives from Corophiidae to Ischyroceridae, and has fused Aoridae with Photidae and Corophiidae. Bousfield (1973) has maintained the separation of these three families, and transferred from Corophiidae to Aoridae all genera with accessory flagellum, coxal gills on peraeopod 2, and enlarged gnathopod 1. Further revision seems desirable (see Myers 1974). Distribution: Mainly coastal marine and estuarine in tropical and temperate regions. Nearly all members of this very large and diverse superfamily group (except some Podoceridae and Cheluridae) construct open-ended tubes, nests, or small "houses" of bottom debris, sediments. or algal pieces which are fixed horizontally (rarely vertically) on the substratum, or are portable. Suborder Caprellidea Leach 1814 Diagnosis: As given by Vassilenko (l974), emend Laubitz (1976a). Distribution: Marine coastal to abyssal, mainly Pacific and Antarctic, tropical to temperate; clinging to and feeding on epibionts; Cyamidae are external parasites of Cetacea. Relationships: Vassilenko (1974) has divided the suborder into four families and two subfamilies. By applying the system of Bowman and Gruner (for hyperiids), the caprellids might be separated into two infraorders (cyamids and caprellids) and the caprellids into two superfamilies, the first containing phtisicids and dodecadids, and the second containing the podocerid-like Caprogammaridae, and the Aeginellidae and Caprellidae, somewhat after the scheme proposed by McCain (1970; see Table I). Laubitz (1976b) has carefully derived the suborder from Dulichia-type podocerid gammarideans.
BousJield: Revised Class$cation of Amphipod Crustaceans 375
Suborder Hyperiidea H. ~ i l n e - ~ d w a r 1830 ds Diagnosis: As given by Bowman and Griiner (1973). Distribution: Open ocean to coastal shelf and high salinity, deep fiord systems; pelagic and abysso-pelagic; carnivorous, or commensalparasitic on coelenterates, salps, and other pelagic invertebrates. Relationships: The external morphology of the twenty-one families, six superfamilies, and two infraorders into which Bowman and Griiner (1973) have divided this suborder is superficially diverse, but fundamentally very uniform. The Hyperiidea exhibit strong affinities with the S ynopioid-pardaliscoid branch of the Gammaridea. Similarities between hyperiids and pardaliscoids (Astyridae, Hyperiopsidae, Vitjazianidae, etc.) seem more than superficial. These include highly modified carnivorous mouthparts, subsimilar, non-amplexing gnathopods, small anterior coxal plates and variably homopodous-heteropodous posterior coxae and peraeopods, broad brood plates, large abdomen with powerful pleopods, and lanceolate uropods. The distinct pelagic terminal male stage, usually bearing brush setae on conjoint flagella of antenna 1, but lacking calceoli, is a fundamental common character. However, hyperiids differ apomorphically not by additional structures but by reduction or modification of essentially pardaliscoidean characters. Thus hyperiids lack maxillipedal palps, accessory flagellum, and coxal gill on peraeopod 7, and possess carpochelate peraeopods, variously fused urosomites, and entire, unarmed telson. Suborder Ingolfiellidea Hansen 1903 Diagnosis: As defined by Ruffo (1970) and Stock (1976). Distribution: Hypogean in littoral marine, brackish and limnetic freshwater continental areas of tropical and warm temperate zones; also abyssal benthic (North Atlantic). Relationships: This small suborder (25 species) has been divided into two families, the relatively morphologically uniform Ingolfiellidae, with six genera and subgenera, and the remarkably divergent Metaingolfiellidae based on a single species from a brackish-water cavern in Italy. Noodt (1965) and others have remarked on the close superficial resemblance of component ingolfiellids to some blind, vermiform members of the hypogean gammaridean family Bogidiellidae with which they overlap considerably in continental fresh and brackish water regions. Ingolfiellids are highly apomorphic in the reduced condition of their
mouthparts, coxal plates, gills, brood plates, pleopods, and uropods, the fusion of peraeon segment 1 with the head (in Metaingolfiellidae), the presence of aesthetascs on antenna 1, and the carpochelate (or carposubchelate) gnathopods. They are plesiomorphic in the presence of minute unpigmented ocular lobes, accessory fiagellum, subsimilar gnathopods with toothed dactyls (in Ingolfiellidae), homopodous peraeopods, and lanceolate uropods. Mills (1967) has also noted the convergent similarity of the deep-sea Zngoljiella to the associated infaunal pardaliscid Epevopeus in general body form, reduced condition of coxae, pleopods, and mouthparts, and the presence of carpochelate gnathopods and lanceolate uropods. The Ingolfiellidea may have diverged very early from the ancestral amphipod root stock; its original epigean members in both marine and freshwater environments have presumably been totally replaced by the more recently evolved gammarideans (especially), and only a few highly apomorphic relict ingolfiellids yet survive in freshwater hypogean and deep-sea marine habitats. The presence of ocular lobes uniquely in Ingolfiellidea, and their occurrence in marine and freshwater biotopes, fully justify its subordinal recognition. The Caprellidea and Hyperiidae do not have unique structures and are distinctive only by reduction or modification of typically gammaridean features; these factors and their strictly marine habitus combine as basis for possible reduction to gammaridean superfamily status.
PROPOSED PHYLOGENETIC RELATIONSHIPS GAMMARIDEAN SUPERFAMILIES WITHIN
The present reclassification divides the Gammaridea into twenty superfamilies (summarized in Table 1). Hypothetical but probable phylogenetic relationships within these superfamily groupings and their relationship to other suborders of the Amphipoda are schematically presented in Figure 6. The thickness or strength of the radiating "branches" is proportional to the size of the subtended group(s), indicated by the number of described species in each superfamily or subordinal "box." The degree to which "name boxes" cross character lines is proportionate to the degree of plesiomorphy-apomorphy in those characters shown by component families and genera. In formulating this monophyletic scheme, the ancestral amphipod is assumed to be a free-swimming somewhat mysid-like animal having a strongly dimorphic terminal male instar, with subsimilar, nonamplexing gnathopods, and plesiomorphic condition in all (or most) characters illustrated in Figures 1-5 and described in the text (pp. 347-56). The morphological gap between the embryologically similar
Bousfield: Revised Class$cation of Amphipod Crustaceans 377

1 Ill l L O f
i 2
l i l a g ~ ct e r m i n a l m a l e s t a g e
I nlceolate
antennae
llon~opodous peraeapads 5-7 irllld a o s t e r o - l o b a t e coxae)
k~t~trirh~rnate number o f known s p e c i e s
ANCESTRAL AMPHIPOD
I I G U R E 6. Hypothetical phylogenetic relationships
gi~mmaridean superfamilies.
of arnphipod suborders and
hut primitive Mysidacea and apomorphic Amphipoda is immense, much greater than between members of the Cumacea-Isopoda phyletic series (Siewing 1963). Yet mysid-like vestiges, including minute unpigmented ucular lobes and "pleopod-like" uropods, persist even in the Ingolfiellidea, and presumably characterized the ancestral amphipod. The gammtiridean prototype had presumably lost the vestigial stalked eye, and the abdominal appendages had become morphologically and functionkilly separated into two distinct groups, viz., three pairs of pleopods (for ~wimmingand production of respiratory current) and three pairs of uropods (for benthic propulsion). * This single character reliably sepamtes amphipods from all other peracaridan (as well as other malacostrac m ) orders since all these have five pairs of pleopods and one pair of tiro pods.
" Macpherson and Steele (1979) have demonstrated fusion of the posterior three nerve
ganglia into a single neural ganglion in the urosome of Gammarus, a unique and r~pomorphic feature of the Amphipoda within the Peracarida.

TABLE l Superfamilies and component families and family groups* Superfamily Phoxocephaloidea Urothoidae new family Phoxocephalidae Sars 1891 Plan ischnooidae Barnard & Superfamily Lysianassoidea Lysianassidae Dana 1849 Superfamily Pontoporeioidea Pontoporeiidae Sars 1882 Haustoriidae Stebbing 1906 Superfamily Gammaroidea Acanthogammaridae Gariaieff 1901 ~niso~ammaridae usfieli ~o 1977 Gammaridae Leach 1814 Gammaroporeiidae new family , Macrohectopidae Sowinsky 1915 Mesogammaridae Bousfield 1977 Pontogammaridae Bousfield 1977 Typhlogammaridae new family Behnigiella-Zernouia family group ? Zphigenella-Pachyschesis family group? Phreatogammarus family group? Superfamily Crangonyctoidea Neoniphargidae Bousfield 1977 Paramelitidae Bousfield 1977 Crangonyctidae Bousfield 1973 emend 1977 Pseudocrangonyx-Sternophysinx family group Superfamily Niphargoidea Niphargidae Karaman 1962 Pseudoniphargus-Allocrangonyx family group ? Austroniphargus family group? Superfamily Bogidielloidea Bogidiellidae Hertzog 1936 Spelaeogamrnarus family group Superfamily Eusiroidea Eusiridae Stebbing 1888 Pontogeneiidae Stebbing 1906 Bateidae Stebbing 1906 Gammarellidae Bousfield 1977 Calliopiidae Sars 1893 Paramphithoidae Stebbing 1906 (revised) Amathillopsidae Pirlot 1934 Superfamily Oedicerotoidea Oedicerotidae Lilljeborg 1865 Superfamily Leucothoidea Pleustidae Buchholz 1874 Laphystiopsidae Stebbing 1899 Amohilochidae Boeck 1871 ~eucothoidae Dana 1852 Anamixidae Stebbing 1897 Maxillipiidae Ledoyer 1973 Colomastigidae Stebbing 1899 Pagetinidae K.H. Barnard 1932 Nihotungidae J.L. Barnard 1972 Stenothoidae Boeck 1871 Cressidae Stebbing 1899 Thaumatelsonidae Gurjanova 1938 Superfamily Talitroidea Hyalidae Bulycheva 1957 Hyalellidae Bulycheva 1957 Talitridae Costa 1857 emend Bulycheva 1957 Ceinidae Barnard 1972 (revised) Dogielinotidae Gurjanova 1954 Najnidae Barnard 1972 Eophliantidae Sheard 1938, revised Barnard 1972 Phliantidae Stebbing 1899, revised Barnard 1969 Temnophliantidae Griffiths 1975 Kuriidae Barnard 1964 Superfamily Stegocephaloidea Stegocephalidae Dana 1852 Acanthonotozomatidae Stebbing 1906 Ochlesidae Stebbing 1910 Lafystiidae Sars 1893 Superfamily Synopioidea Synopiidae Dana 1853 Argissidae Walker 1904 Superfamily Pardaliscoidea Pardaliscidae Boeck 1871 Stilipedidae Holmes 1908 Hyperiopsidae Bovallius 1886 Astyridae Pirlot 1934 Vitjazianidae Birstein & Vinogradov 1955 (revised) Superfamily Liljeborgioidea Liljeborgiidae Stebbing 1899 Sebidae Walker 1908 Salentinellidae Bousfield 1977 Superfamily Dexaminoidea Atylidae Lilljeborg 1865 Anatylidae Bulycheva 1955 Lepechinellidae Schellenberg 1926
BousJeld: Revised ClassiJication ofAmphipod Crustaceans 379 T A B L E 1 (Continued) Dexaminidae Leach 1814 Prophliantidae Nicholls 1940 (revised Barnard) Superfamily Ampeliscoidea Ampeliscidae Bate 1861 Superfamily Melphidippoidea Melphidippidae Stebbing 1899 Hornellia-Cheirocratus family group Megaluropus family group Superfamily Melitoidea Hadziidae Karaman 1932 Melitidae Bousfield 1973 Carangoliopsidae Bousfield 1977 Superfamily Corophioidea Photidae Boeck 1872 Isaeidae Dana 1853 Ischyroceridae Stebbing 1899 (revised) Ampithoidae Stebbing 1899 Biancolinidae Barnard 1972 Aoridae Stebbing 1899 (revised) Cheluridae Allman 1847 Corophiidae Dana 1849 (revised) Podoceridae Stebbing 1906 Suborder CAPRELLIDEA Infraorder CAPRELLIDA Superfamily Phtisicoidea Paracercopidae K. & V. 1972 Phtisicidae Vassilenko 1968 Dodecadidae Vassilenko 1968 Superfamily Caprelloidea Caprogammaridae K. & V. 1966 Aeginellidae Vassilenko 1968 Caprellidae White 1847 ~nfraorder CYAMIDA Superfamily Cyamoidea Cyamidae White 1847 Suborder INGOLFIELLIDEA Ingolfiellidae Hansen 1903 Metaingolfiellidae Ruffo 1969
* The Hyperiidea are treated by Bowman and Griiner (1973).
Within the Gammaridea, two main trunk lines diverged (Figure 6). In the larger and taxonomically more diverse line (to the right), pelagic males retained (or developed!) calceoli as well as brush setae, whereas only brush setae are present in pelagic males of the smaller trunk line to the left). The small, enigmatic, highly plesiomorphic group of stegocephaloideans, lacking a pelagic male, forms perhaps a third main type that diverged somewhere between the lysianassids and the synopiids. The (calceolate) main trunk members are almost entirely benthic or infaunal, inquilinous, or epi-parasitic types, most of which (except lysianassids, phoxocephalids, and leucothoids) have successfully penetrated brackish and primary freshwater environments, both epigean and hypogean; one group (talitroideans) penetrated the upper tidal zone and the terrestrial (leaf litter) environment. Almost all of the freshwater and terrestrial types have lost the pelagic terminal male stage (except Pontoporeia) and many evolved heteropodous limbs, anteriorly lobate coxae, spinulose urosomes, and linear, spine-tipped uropod rami. Epigean freshwater members largely retained calceoli, which are lost or lacking in almost all fully hypogean (eyeless) amphipods. The gnathopods became more or less dissimilar, and moderately to strongly
amplexing in form and%iction. The mouthparts of generalized plantand detritus-feeding (scavenging) groups remained essentially basic. The filter feeders, however, developed strong inner lobes on the lower lips, and the mouthparts of the carnivores and flesh scavengers, the solid particle feeders, and the various micro-predators, parasites, and inquilines became highly modified, largely by reduction or modification of plates, palps, setae, and spines. The other main trunk line (non-calceolate) remained essentially marine, except for penetration of brackish and coastal epigean fresh waters by some liljeborgiids and melitoids. It diverged further into two main groups. The first of these encompasses the synopioidpardaliscoidean complex that remained largely pelagic and oceanic, mainly carnivorous, and was presumably ancestral to the suborder Hyperiidea; and the small benthic, epibenthic, and commensal Liljeborgioideans, a few of which became microphreatic freshwater forms. All these retained the plesiomorphic lanceolate condition of the uropods and telson, the mainly homopodous peraeopods, the non-amplexing gnathopods, and (except for liljeborgioids) the strongly dimorphic pelagic male form. The second group became essentially benthic crawlers, clingers, nest builders or tube builders, and mostly deposit- or trypton-feeders. All developed heteropodous limbs, anteriorly lobate coxae, and apomorphic linear uropod rami. The more primitive superfamily groups of this branch (dexaminoids, ampeliscoids, and melphidippoids) largely retained the pelagic male stage, although they developed somewhat dissimilar and variously amplexing type gnathopods. The more highly apomorphic and successful corophioideans, their descendent caprellideans, and the melitoideans burst free of the pelagic male stage and developed highly dissimilar and sexually dimorphic amplexing type gnathopods that may also function in sex display, fighting, and sound production. Their bodies became cylindrical or depressed, with small, non-excavate anterior coxal plates. They also evolved very specialized ambulatory limbs for construction of protective tubes or domiciles, more elongate and setose antennae for foodgathering, and specialized uropod rami and telson. These posterior appendages bear small hook-like spines by means of which the animal clings to the entrance of the domocile during food-gathering. The pleopods were relegated mainly to the production of a respiratory current, or were lost entirely (Caprellidea). With respect to evolution of other superfamily characters, the accessory flagellum was retained most strongly in the plesiomorphic pelagic groups, especially in calceolate types, and in almost all freshwater members, but was much reduced or entirely lost in the benthic infaunal marine and terrestrial groups and in melitoidean hypogean members. Elongation of the peduncle of antenna 2 is more frequent in benthic and
Bousjield: Revised Classi~cation Amphipod Crustaceans 381 of
epibenthic groups, and in highly apomorphic groups in which it may be used in physical grasping or holding of the female during amplexus (some Corophioidea, Talitridae, possibly Caprellidea). Gnathopods are strongly raptorial in marine pelagic carnivores (Eusiridae and Pardaliscoidea), a function assumed by peraeopods 3 and 4 in some Hyperiidea. Several free-burrowing groups have enlarged, subsimilar, fossorial gnathopods (Phoxocephaloidea, Oedicerotidae, and some freshwater Crangonyctoidea and Niphargoidea). The size and marginal setation of brood plates apparently set no functionally discernible pattern, but may reflect the respiratory physiology of the developing eggs, a subject about which little is known. Coxal gills are primarily simple, plate-like or sac-like, respiratory lobes that may be attached to the coxae of all peraeopods but peraeopod (gnathopod) 1. Foliaceous or lobate development occurs in marine benthic forms tolerant of anoxia and a high carbon dioxide concentration (some Lysianassidae, Ampeliscoidea, etc.), and sterna1gills may be typical offreshwater counterparts that live in deep lake sediments (Pontoporeia) or in oxygen-deficient groundwaters (Crangonyctoidea). Loss of coxal gill on peraeopod 7 occurs mainly in benthic and most highly apomorphic groups, with specialized habits andlor respiratory current production. Development of linear uropods with apical spine groups is typical of apomorphic benthic superfamilies, presumably in response to bottom "siding" and saltation motion, and reduced need of a swimming tail fan. Development of dorsal abdominal spine clusters, especially in gammaroidean freshwater groups, presumably also provides increased purchase to bottom "sidling" species. Uropod 3 has remained lanceolate and/or foliaceous in all groups that swim extensively, including most bottom crawlers and infaunal burrowers, but has become nonfoliaceous, linear, and much reduced in tube-building Corophioidea, the terrestrial and intertidal Talitroidea, and many hypogean crangonyctids, niphargids, and bogidiellids. The most primitive form of the amphipod telson seems to have been the "partly cleft" form, with apically sharply notched lobes. In most benthic and hypogean freshwater species, the lobes have fused distally to become an entire plate, which is marginally spinose or smooth; but in some infaunal free-burrowing groups (Lysianassoidea, Phoxocephaloidea) and freshwater gammaroideans, the lobes are variously separated to the base, and in a few instances (Eohaustorius of Pontoporeioidea), the lobes are widely separated on urosome 6.
Probable origin and direction of diversij?cation in amphipod crustaceans The answer to how and when the Amphipoda diverged from its peracaridan ancestral type can only be obtained indirectly. The fossil
record of the group is extremely limited and relatively very recent. Much evidence of its past morphological radiation and of patterns of global dispersal has been eradicated by glaciation in the Southern Hemisphere during the past 20 million years. However, amphipod evolutionary chronology can be deduced, in part, from consideration of the recently established chronology of major geological events such as continental drift (plate tectonics) and of major climatic changes in the seas and on the continents, and from the much better fossil records of major plant and animal groups with which amphipods are now intimately associated. Thus, the remarkably close taxonomic relationship of the crangonyctoid genus Falklandella in eastern Falkland Islands to counterpart primary freshwater species of Pavamelita in South Africa indicates that their presumed common ancestor must have existed during the early Mesozoic, since these two continental land masses have been separated by the South Atlantic Ocean for about 150 million years (Dietz and Holden 1970). We may also look to the model provided by Schminke (1974) for intercontinental relationships in a primary freshwater hypogean malacostracan group, the Bathynellacea, which lacks a fossil record. This model seems especially applicable to CO-occurringhypogean gammaridean (Crangonyctoidea, Bogidielloidea) and ingolfiellidean amphipods whose oligohaline and marine penetration from fresh waters is believed to be secondary and whose dispersal capacity is also extremely limited. Similarly, the model provided by Brundin (1966) based on the "continental drift" dispersal of epigean dipterous insects (Chironomidae) in the Southern Hemisphere would seemingly apply well to both hypogean and epigean amphipod superfamily and family groups such as the terrestrial Talitridae (see Bousfield 1968). These cryptozoic, leaf-litter crustaceans are confined mainly to rain forests of continents and continental islands of the Southern hemisphere (not yet found in South America), and to the Indian subcontinent and to certain Caribbean continental islands (Jamaica) and mountainous areas of Central America in the Northern Hemisphere. Thus ingolfiellids, bogidielloideans, crangonyctoideans, and advanced talitroideans must have attained their present basic morphology prior to the Cretaceous (beginning of continental drift and breakup of Pangea), but have radiated morphologically very little since that time. Since these four groups are highly derived (apomorphic), we might assume that the more plesiomorphic marine amphipod groups (from which these were presumably derived, Figure 6) had attained their definitive morphology at an even earlier period. On the assumption that the initial period of evolution of bogidielloideans, crangonyctoideans, and talitroideans is at least equal to their subsequent "continental-drift" history (about 120 million years), gammaridean amphipods, as a group, probably attained their
Bousjield: Revised Classification ofAmphipod Crustaceans 383
plesiomorphic (marine) morphology during the late Palaeozic. Based on the fossil record, Schram (1977) indicates the Permian as the period of maximum dispersal of higher malacostracans, including peracaridans, into Pangean coastal and fresh waters and the Mesozoic as the main period of radiation of peracaridans. The actual evolution of the order amphipoda may have occurred towards the close of the Carboniferous Period, and during the period of rapid diversification of invertebrate orders described by Eldridge (1976). This timing would conform reasonably well with that predicted for apomorphic peracaridan groups of the mysid-tanaid-isopod evolutionary line whose fossil record extends back to the Permian and Carboniferous. Interestingly, modern amphipod-like phreatoicid isopods that occupy biomes and geographical ranges similar to the terrestrial talitroideans differ surprisingly little from their Triassic fossil precursors (see Hessler 1969). By contrast, the superfamily Gammaroidea appears to be of relatively recent development. Most families and genera have diversified in continental fresh waters of Eurasia, mainly in coastal brackish and fresh waters of the Pacific rim region (see Tzvetkova 1975), in the Pontocaspian region (see Carasau et al. 1955) and in Lake Baikal during the past 50 million years (see Bazikalova 1945). Only a few genera (in family Gammaridae, Anisogammaridae, etc.) are known from North America, and these are mostly apomorphic marine and brackish-water species, dispersed readily along holarctic continental margins (see Bousfield 1979). Pertinently, amphipod fossils from the upper Eocene (Baltic amber deposits) include only crangonyctoideans (Palaeogammarus, near Crangonyx), whereas these eyed, epigean types are almost totally excluded from that European region today and appear to have been replaced subsequently by gammaroidean counterparts. The high diversity and dominance of crangonyctoideans (Cvangonyx, Synuvella) and low diversity of gammaroideans in surface fresh waters of North America are enigmatic. Failing explanation based on differential Pleistocene glaciation, the difference may be attributed to the relatively recent evolution and diversification of gammaroideans in Eurasia, most primary freshwater types of which have been isolated from the westward-drifting North American continent. Assuming, therefore, that the Amphipoda (like other orders of Peracarida) had evolved by the late Palaeozoic, what causal basis might be proposed for the origin and diversification of this relatively apomorphic group? The writer tends to agree with the views of Siewing (1963) and Hessler and Newman (1975) that the head shield, thoracic pleura, and lack of a carapace in Amphipoda and other higher peracaridans are apomorphic derivations from a caridoid- or mysid-like ancestor, rather than with the reverse hypothesis put forth by Dahl (1976) that the
caridoid-like form may be the apomorphic or derived condition. In view of the very fragmentary nature of the fossil record, however, the subject is by no means closed. Early plesiomorphic malacostracan supraordinal groups (Phyllocarida, Syncarida) were essentially filter-feeders and have fossil records extending back variously to the Cambrian. During those early Palaeozoic periods, primary plant food sources consisted (presumably in both marine and freshwater environments) of blue-green and unicellular algae, primitive red, brown, and green thallophytic algae and bacteria and perhaps mosses, and lichens (Brown 1935). During the Devonian, the evolution of pteridophytes commenced on land, and the process of radiation and diversification of equisites, lycopods, cycads, and primitive conifers and ginkgos reached full bloom during the Carboniferous. An immense new dimension to primary food resources was thus afforded aquatic as well as terrestrial organisms of that period. In coastal marine areas, these new food sources very probably stimulated the evolution of a host of new invertebrate feeding types, both directly as consumers of plant materials and indirectly through exploitation of associated biomes and new niches, including prey-predator and hostparasite relationships. Eucaridans, especially decapod crustaceans, may have evolved mainly as scavengers and macro-carnivores in coastal marine energy systems, but were tied to shallow seas by their planktotrophic larval biology. Some primitive decapods (Astacura) did penetrate fresh waters, especially in temperate continental regions, by suppressing larval development within the egg stage, carried and protected on the female pleopods. During the Mesozoic and subsequently, these freshwater Macrura appear to have been largely replaced in tropical and warm temperate coastal freshwater areas by the more advanced Anomura (e.g. Aeglidae) and Brachyura (e.g. some Grapsidae). In these forms, larval development is very abbreviated and non-feeding, or takes place within the egg. Terrestrial decapods, however, are few (e.g. Coenobitidae, Gecarcinidae), are restricted to immediate coastal areas of tropical and warm temperature continents and islands, and have marine planktotrophic larvae. Peracaridan crustaceans, however, are characterized by direct development of eggs, protected in the thoracic brood pouch of the female. Higher peracaridans are independent of a carapace-enclosed respiratory chamber. Amphipod peracaridans, especially, are aided in propulsion and food-gathering by mechanically efficient, uniramous, and rigid thoracic and abdominal limbs, and by cephalization of the mouthparts in a buccal mass that is often prognathous. All these features would admirably facilitate the occupation of freshwater and humid terrestrial environments, and the direct exploitation of the primary new plant food resources (above). In amphipods also, the early loss of apelagic terminal
BousJield: Revised Classijication of Amphipod Crustaceans 385
male stage was prerequisite to colonization of humid terrestrial environments, and to penetration of lotic epigean habitats (by gammaroideans) and lentic interstitial and cave biotopes (by ingolfielloids, bogidielloids, niphargoids, and hadzioids. In the sea, similar advances in reproductive biology and morphology enabled amphipods to penetrate and diversify greatly within cold, dark, arctic and antarctic regions and within the deep sea. In these environments, photosynthetic activity is non-existent, or too seasonally brief to support most planktotrophic larvae or planktotrophic organisms on which predaceous larvae might feed. In the deep sea, primary food sources occur on or in the sediments, and consist (to a significant degree) of the remains of sinking plant and animal debris of terrestrial or coastal marine origin (see Wolff 1977). Perhaps Archaeozoic and early Palaeozoic deep sea bottoms were devoid of macro-life perhaps not only because of higher temperatures and more anoxic conditions, but because they predated the evolution of particulate organic matter of terrestrial origin, andlor animal converters or secondary producers that had developed life history mechanisms for living there. The earliest large marine herbivorous arthropods, the trilobites, were presumably "locked into" a primitive filter-feeding and scavenging morphology, stenotopic physiology, and primitive reproductive biology. They were thus presumably unable to exploit these new food resources and new biomes or meet the competition from more efficient crustacean types, and disappeared totally from the scene by the close of the Palaeozoic. Likewise, the most primitive filter-feeding malacostracan superorders have all but disappeared and persist only as relicts in a few essentially anoxic marine habitats (e.g. Nebaliacea) or in remote Tasmanian alpine freshwater pools (e.g. Anaspidacea) or in the freshwater hypogean (e.g. Bathynellacea). Only the carnivorous stomatopod branch of Hoplocarida persisted in the marine environment and radiated in the Mesozoic (Schram 1977). In contrast, even the most plesiomorphic marine amphipod superfamilies were able to exploit the new organic food resources, especially in shallow-water, high-energy, intertidal habitats (e.g. Phoxocephaloidea, Synopioidea, Oedicerotoidea, Melphidippoidea) or scavenge dead or dying carcasses of fishes, marine reptiles (especially during the Mesozoic), and marine mammals (since the Cretaceous; e.g. Lysianassidae), or prey upon myriad small planktotrophic crustaceans such as copepods and their larvae (e.g. Eusiridae, Pardaliscoidea, and Hyperiidea). The more apomorphic benthic marine superfamilies may have radiated contemporaneously with the evolution and development of coastal salt marsh plants, sea grasses, mangrove swamps, and their associated coral and sponge communities during the Cretaceous (see McCoy and Heck, 1976). Thus, a host of new food sources and house-
building materials would then have opened up to the essentially inquilinous Leucothoidea on the one hand, and to the nestling and tube-building Dexaminoidea, Ampeliscoidea, Melitoidea, and Corophioidea on the other. The apomorphic cyamid whale parasites presumably evolved in concert with marine cetaceans in Cenozoic times, and thereby indicate that the Caprellidae and more plesiomorphic relatives date from the Cretaceous or earlier. The sand-burrowing Pontoporeioidea have radiated in shallow, high-energy coastal marine margins of the widening North Atlantic and adjacent seas during the Cenozoic (see Bousfield 1970), but the "pelagic male" types persist in fresh waters only as phylogenetic relicts in large lacustrine bodies such as the Great Lakes and the Baltic and Caspian seas. The gammaroideans, however. have diversified immensely in all lotic and lentic epigean environments, presumably in post-Cretaceous synchrony with proliferation of angiosperms as a primary food resource. Thus the amphipods seem to have exploited successfully many of the permanent marine and freshwater niches available to organisms of their size range. They have only begun to utilize food resources available through symbiotic-parasitic relationships with other marine and freshwater invertebrates. Attendant changes in their morphology have yet barely reached the family level of recognition in most cases (e.g. in Hyperiidea, Liljeborgiidae, Colomastigidae, Lafystiidae, Lysianassidae). By contrast, parasitic morphological modifications have attained superfamily and subordinal level in the Isopoda, and ordinal and subclass level in even more plesiomorphic crustacean groups (e.g. in Branchiura, Copepoda, Cirripedia). In the marine intertidal zone, talitroideans dominate the exploitation of plant debris as a food resource, despite competitive pressure from isopods and decapods from the marine environment and insects from the terrestrial. The superior marine dispersal mechanisms of semiterrestrial talitroideans enable them to better colonize remote oceanic volcanic islands (e.g. Hawaii) and dominate the leaf litter and associated cryptozoic habitats there, even in competition with some insects, myriapods, and oniscoideans, and predation by arachnids, lizards, etc. (see Bousfield and Howarth 1976). On continental land masses, however, the primitive "terrestrial" physiology and gill respiration of amphipods ties them to limited regions with winter-mild climate, high humidity, and high chloride-ion soils. Reasons for the apparent absence of amphipods from warm, fresh surface waters of tropical continental areas, especially in the Indo-Pacific, central Africa, and to some extent South America, are not clear. Colonization of these regions may yet be accomplished by the Talitroidea, and perhaps by the Melitoidea, two eminently preadapted superfamily groups. During the late Cenozoic and Pleistocene, the Hyalellidae
BousJield: Revised Classijication of Amphipod Crustaceans 387
(Talitroidea) have been "on the march," with rapid exploitation and diversification in Lake Titicaca in South America, and wide dispersal in North America since the central American land bridge was established about one million years ago. Amphipods have not developed the resistant egg stage of primitive branchiopod crustaceans that has enabled the latter to exploit fully the food resources of temporary fresh and brackish pools. However, their physiological eurytopicity, sterna1 gill respiration, and variable capacity to live hypogeically as young, combined with their capacity for wind-blown dispersal, provide both hyalellids and crangonyctids with significant capability to colonize and exploit such environments. Likewise, the Gammaroidea have radiated remarkably (to genus and even family level) within Cenozoic Lake Baikal. Assuming long-term stability of climatological and hydrological conditions in post-Pleistocene times, one might predict a renewed penetration and diversification of gammaroideans in North America, and eventual adaptive radiation and proliferation of species flocks in the large deep lakes bordering the Precambrian Shield.
SELECTED REFERENCES
The following list embodies only references in the text, and basic references to new taxa in the table (superfamily and family). Complete references for citations of gammaridean taxa prior to 1968 are provided in Barnard (1969a) and Stebbing (1906), the two primary taxonomic references of this study, and for caprellidean taxa, in Vassilenko (1974). A complete bibliography is also available through the author. Barnard, J.L. 1958. Index to the Recent Families, Genera, and Species of the Gammaridean Amphipoda (Crustacea).Allan Hancock Found. h b l . Occas. Pap. 19. - 1964. Deep-sea Amphipoda (Crustacea) collected by the R/V"Vema" in the eastern Pacific Ocean and the Caribbean and Mediterranean seas. Bull. A m . Mus. Nut. Hist. 127 (1): 3-46. - 1969a. The Families and Genera of Marine Gammaridean Amphipoda. U.S. Natl. Mus. Bull. 271. - 1969b. Gammaridean Amphipoda of the Rocky Intertidal of California: Monterey Bay to La Jolla. U.S. Natl. Mus. Bull. 258. - 1970. Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands. Smithson. Contrib. Zool. 34. - 1972a. Gammaridean Amphipoda of Australia. Part I . Smithson. Contrib. Zool. 103. - 1972b. The Marine Fauna of New Zealand: Algae-Living Littoral Gammaridea (Crustacea, Amphipoda). N.Z. Dept. Sci. Ind. Res. Bull. 210 (N.Z. Oceanogr. Inst. Mem. 62). - 1973. Revision of Corophiidae and Related Families (Amphipoda). Smithson. Contrib. Zool. 151.

- 1974. Evolutionary patterns in gammaridean Amphipoda. Crustaceana 27 (2):
137-46.
- 1976. Affinities of Paraniphargus lelouparurn Monod, a blind anchialine am-
phipod (Crustacea) from the Galapagos Islands. Proc. Biol. Soc. Wash. 89 (36): 421-32. - 1978. The cavernicolous fauna of Hawaiian lava tubes 9. Amphipoda (Crustacea) from brackish lava ponds. Pac. Insects 17. Barnard, J.L., and Drummond, M. 1978. The Phoxocephalidae of Australia. Smithson. Contrib. Zool. 245. - 1979. Gammaridean Amphipoda ofAustralia. Part I V : The Platyischnopidae. Smithson. Contrib. Zool. 276. Bate, C.S. 1862. Catalogue of the Specimens ofAmphipodous Crustacea in the Collection of the British Museum, London. Bazikalova, A.Y. 1945. Amphipoda of Lake Baikal. Trud. Baikal Limnol. Sta. 11. Bousfield, E.L. 1965. The Haustoriidae of New England (Crustacea: Amphipoda). Proc. U . S . Natl. Mus. 117 (3512): 159-240. - 1968. Terrestrial adaptations in Crustacea. Sess. I . Discussion. Transition to land. A m . Zool. 8 (3): 393-8. - 1970. Adaptive radiation in sand-burrowing amphipod crustaceans. Chesapeake Sci. 11 (3): 143-54. - 1973. Shallow-Water Gammaridean Amphipoda of New England. Cornell University Press, Ithaca, N . Y . - 1977. A new look at the systematics of gammaroidean amphipods of the world. Crustaceana, Suppl. 4: 282-3 16. - 1979. The Amphipod Superfamily Gammaroidea in the Northeastern PaciJic Region:SystematicsandDistributionalEcology. Bull. Biol. Soc. Wash. 79(3). Bousfield, E.L., and Howarth, F.G. 1976. The cavernicolous fauna of Hawaiian lava tubes 8. Terrestrial Amphipoda (Talitridae), including a new genus and species with notes on its biology. Pac. Insects 17 (1): 144-54. Bousfield, E.L., and Hubbard, J.D. 1968. New Records of Gammaridean Amphipod Crustaceans from the Intertidal Zone of Prince William Sound, Alaska. Natl. Mus. Can. Nat. Hist. Pap. 40. Bowman, T.E., and Griiner, H. 1973. The Families and Genera of Hyperiidea (Crustacea: Amphipoda). Smithson. Contrib. Zool. 146. Brown, W.H. 1935. The Plant Kingdom. Ginn and Company, Boston, New York. Brundin, L. 1966. Transantarctic Relationships and Their SigniJicance, as evidenced by Chironomid Midges. Kgl. Svensk. Akad. Handl. (4) 11 (1). Bulycheva, A.I. 1957. The Sea Fleas of the Seas of the U.S.S.R. Adjacent and Waters (Amphipoda-Talitroideu) (in Russian). Keys to the Fauna of the U . S . S . R . 65. Acad. Sci. USSR. Carasau, S., Dobreanu, E., and Manolache, C. 1955.Fauna Republicii Populare Romine Crustacea. Amphipoda. Acad. Republ. Pop. Rom. 14 (4). Dahl, E. 1976. Structural plans as functional models exemplified by the Crustacea Malacostraca. 2001. Scr. 5: 163-6. Dietz, R.S., and Holden, J.C. 1970. Reconstruction of Pangaea: Breakup and dispersion of continents. Permian to present. J. Geophys. Res. 75: 4939-56. Eldridge, M. 1976. Rates of evolution revisited. Palaeogeogr. Palaeoclimatol. Palaeoecol. 2: 174-9. Griffiths, C.L. 1974. The Amphipoda of Southern Africa. Part 4. The Gam-
BousJield: Revised Classzjication ofAmphipod Crustaceans 389 maridea and Caprellidea of the Cape Province east of Cape Agulhas. Ann. S . Afr. M M . 65 (9): 251-336. - 1975. The Amphipoda of Southern Africa. Part 5. The Gammaridae and Caprellidae of the Cape Province west of Cape Agulhas. Ann. S . Afr. Mus. 67 (5): 91-181. Hessler, R.R. 1969. Peracarida. In R.C. Moore (ed.), Treatise on Invertebrate Palaeontology. Part R . Arthropoda. 4: ~ 3 6 0 - ~ 3 9 3 . Geol. Soc. America, Inc., and Univ. Kansas. Hessler, R.R., and Newman, W.A. 1975. A trilobitomorph origin for the Crustacea. Fossils and Strata 4: 437-59. Holsinger, J.R. 1977. A review of the systematics of the Holarctic amphipod family Crangonyctidae. Crustaceana, Suppl. 4: 244-81. Holsinger, J.R., and Longley, G . 1979. The Subterranean Amphipod Crustacean Fauna ofan Artesian Well in Texas. Smithson. Contrib. Zool. (in press). Hurley, D.E. 1954. Studies on the New Zealand amphipod fauna 4. The family Gammaridae including a revision of the freshwater genus Phreatogatnmarus Stebbing. Trans. Roy. Soc. N . Z . 81 (4): 601-18. - 1963. Amphipoda of the Family Lysianassidae from the West Coast of North and Central America. Allan Hancock Found. Publ. Occas. Pap. 25 Kaestner, A. 1970. Lehrbuch der Speziellen Zoologie. English translation and adaptation. Znvertebrate Biology by H.W. and L.R. Levi. Vol. III. Crustacea. Wiley, Interscience, New York, London. Kukalova-Peck, J. 1973. A Phylogenetic Tree of the Animal Kingdom (Including Orders and Higher Categories).Natl. Mus. Can. Publ. Zool. No. 8. Laubitz, D.R. 1976a. On the taxonomic status of the family Caprogammaridae Kudryaschov and Vassilenko (Amphipoda). Crustaceana 3 1 (2): 145-50. - 1976b. A revision of the genera Dulichia Krgyer and Paradulichia Boeck (Amphipoda, Podoceridae). Can. J . Zool. 55 (6): 942-82. Ledoyer, M. 1969. Amphipodes gammariens de quelques biotopes de substrat meuble de la region de Tulear (Republique Malagache). Etude systematique et ecologique. Ann. Univ. Madagascar (Sci. Nat. Math.) 6: 249-96. - 1973. Amphipodes Gammariens de la Frondaison des Enhalus de la region de Nossi-Be (Systematique et Ecologie). Tethys, Suppl. 5: 25-36. Macpherson, B.R., and Steele, V.J. 1979. The microanatomy of the central nervous system of Gammarus setosus Dementieva (Crustacea, Amphipoda). The suboesophageal ganglion and ventral ganglion chain. Proc. Znt. Symp. Groundwater Biol., Blacksburg, Va. (in press). McCain, I. 1970. Familial taxa within the Caprellidea (Crustacea, Amphipoda). Proc. Biol. Soc. Wash. 82 (65): 837-42. McCoy, E.D., and Heck, K.L. 1976. Biogeography of corals, seagrasses, and mangroves: An alternative to the centre of origin concept. Syst. ,7001. 25 (3): 201-10. Mills, E.L. 1967. Deep-sea amphipoda from the western North Atlantic Ocean. 1. Ingolfiellidea and an unusual new species in the gammaridean family Pardaliscidae. Can. J. Zool. 45: 347-55. Myers, A.A. 1974. A first record of the genus Pseudomegamphopus Myers (Crustacea, Amphipoda) from the Indo-West Pacific with a redescription of P. jassopsis (K.H. Barnard) comb. nov. Trans. Roy. Soc. S . Afr. 41 (2): 195-202. Noodt, W. 1965. Interstitielle Amphipoden der konvergenten Gattungen Zngolfiella Hansen und Pseudingolfiella n. gen. aus Siidamerika. Crustaceana 9 (l): 17-30.
Ruffo, S . 1970. ConsidCrations Apropos de la systematique et de la biogeographie des Ingolfielles (Crustacea. Amphipoda). Livre centenaire Emile G. Racovitza, 223-30. Bucarest. Sars, G.O. 1895. An Account of the Crustacea of Norway. Vol. 1. Amphipoda. Christiana and Copenhagen. Schminke, H.K. 1974. Mesozoic intercontinental relationships as evidenced by bathynellid Crustacea (Syncarida: Malacostraca). Syst. Zool. 23 (2): 157-64. Schram, F.R. 1977. Paleozoogeography of Late Paleozoic and Triassic Malacostraca. Syst. Zool. 26 (4): 367-79. Siewing, R. 1963. Studies in malacostracan morphology: results and problems. Phylogeny and Evolution of Crustacea. Special Publ. Mus. Comp. Zool. 85-103. Stebbing, T.R.R. 1888. Report on the Amphipoda collected by H.M.S. Challenger during thebyears 1873-76. In Rept. Sci. Results Voy. H.M.S. Challenger. 1873-76. Zool. 29. - 1906. Amphipoda I. Gammaridea. Das Tierreich, Vol. 21. Stock, J.H. 1976. A new member of the crustacean suborder Ingolfiellidea from Bonaire. Stud. Fauna Curaqao Other Caribb. Zsl. LV (164): 56-75. - 1977. The taxonomy and zoogeography of the hadziid Amphipoda, with emphasis on the West Indian taxa. Stud. Fauna Curaqao Other Caribb. Zsl. LV (177): 1-130. Tzvetkova, N.L. 1975. Littoral Gammaridae ofNorthern andFar-Eastern Seas of the U.S.S.R. and Adjacent Waters (in Russian). Acad. Nauk, USSR. Zool. Inst. Leningrad. Vassilenko, S.V. 1974. Caprellidea(Marine Skeleton Shrimps) of the Seas of the U.S.S.R. and Adjacent Waters (in Russian). Acad. Sci. USSR. Opred. Fauna USSR, Leningrad, No. 107. Wolff, T. 1977. Diversity and fauna1 composition of the deep-sea benthos. Nature 267 (5614): 780;s. Zimmerman, R.J., and Barnard, J.L. 1977. A new genus of primitive marine hadziid (Amphipoda) from Bimini and Puerto Rico. Proc. Biol. Soc. Wash, 89 (50): 565-80.
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E . L . B O U S F I E L D , F . R . S . C* .

A Revised Classification and Phylogeny of Amphipod Crustaceans

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llon~opodous peraeapads 5-7 irllld a o s t e r o - l o b a t e coxae)

k~t~trirh~rnate number o f known s p e c i e s

I I G U R E 6. Hypothetical phylogenetic relationships

of arnphipod suborders and

378 The Royal Society of Canada

* The Hyperiidea are treated by Bowman and Griiner (1973).

380 The Royal Society of Canada

Bousjield: Revised Classi~cation Amphipod Crustaceans 381 of

382 The Royal Society of Canada

Bousjield: Revised Classification ofAmphipod Crustaceans 383

384 The Royal Society of Canada

BousJield: Revised Classijication of Amphipod Crustaceans 385

386 The Royal Society of Canada

BousJield: Revised Classijication of Amphipod Crustaceans 387

388 The Royal Society of Canada

390 The Royal Society of Canada

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