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Malacostraca













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MALACOSTRACA. Under this zoological title are included several groups of Crustacea, united by characters which attest their common origin, though some, and probably all of them, were already separated in distant geological ages, and some have now attained a peculiar isolation. Throughout the whole, the researches made since 1860 have not only added a great throng of new species, genera and families, but have thrown a flood of light upon questions of their phylogeny, systematic arrangement, horizontal and bathymetric distribution, organization, habits of life and economic importance. There are at least seven orders: the stalk-eyed Brachyura, Macrura, Schizopoda, Stomatopoda, and the sessile-eyed Sympoda, Isopoda, Amphipoda. An ocular segment claimed by the former division is not present or in no case demonstrable in the latter. In neither does the terminal segment or telson, whether large or obsolescent, whether articulated or coalescent, carry appendages, unless occasionally in fusion with itself. Between the eyes and the tail-piece in all the orders nineteen segments are counted, the proof of a segment's existence depending on its separateness, complete or partial, or on a sutural indication, or else on the pair of appendages known to belong to it. All these marks may fail, and then the species must be proved to be Malacostracan by other evidence than the number of its segments; but if some exceptions exhibit fewer, none of the Malacostraca exhibits more than 19 (+ 1 or + 2) segments, unless the Nebaliidae be included. Of the corresponding pairs of appendages thirteen belong to the head and trunk, two pairs of antennae, one pair of mandibles, two pairs of maxillae, followed by three which may be all maxillipeds or may help to swell the number of trunk-legs to which the next five pairs belong. The abdomen or pleon carries the remaining six pairs, of which from three to five are called pleopods and the remainder uropods. Underlying the diversity of names and functions and countless varieties of shape, there is a common standard' to which the appendages in general can be referred. In the maxillipeds and the trunk-legs it is common to find or otherwise easy to trace a seven-jointed stem, the endopod, from which may spring two branches, the epipod from the first joint, the exopod from the second.' The first antennae are exceptional in branching, if at all, at the third joint. In the mandibles and maxillae some of the terminal joints of the stem are invariably wanting. In the rest of the appendages they may either be wanting or indistinguishable. The latter obscurity results either from coalescence, to which all joints and segments are liable, or from subdivision, which occasionally affects joints even in the trunk-legs. The carapace, formerly referred only to the antennar-mandibular segments, may perhaps in fact contain elements from any number of other segments of head and trunk, Huxley, Alcock, Bouvier giving support to this opinion by the sutural or other divisional lines in Potamobius, Nephrops, Thalassina, and various fossil genera. Not all questions of classification internal to this division are yet finally settled. Between the Brachyura and Macrura some authors uphold an order Anomura, though in a much restricted sense, the labours of Huxley, Boas, Alcock and conjointly Alphonse Milne-Edwards and Bouvier, having resulted in restoring the Dromiidea and Raninidae to the Brachyura, among which de Haan long ago placed them. The French authors argue that from the macruran lobsters (Nephropsidae) anciently diverged two lines: one leading through the Dromiidea to the genuine Brachyura; or crabs, the other independently to the Anomura proper, which may conveniently be named and classed as Macrura anomala. Spence Bate maintained that the Schizopoda ought not to form a separate order, but to be ranged as a macruran tribe, "more nearly allied to the degraded forms of the Penaeidea than to those of any other group" ("Challenger" Reports, " Macrura," p. 472, 1888). According to Sars, the 1 In Huxley's terminology the first two or three joints of the stem constitute a "protopodite," from which spring the "endopodite" and "exopodite." Sympoda (or Cumaceans), in spite of their sessile eyes, have closer affinities with the stalk-eyed orders. H. J. Hansen and others form a distinct order Tanaidea for the decidedly anomalous group called by Sars Isopoda chelifera. r. Brachyura. - For the present, as of old, the true Brachyura are divided into four tribes: Cyclometopa, with arched front as in the common eatable crab; Catometopa, with front bent down as in the land-crabs and the little oyster-crab; Oxyrhyncha, with sharpened beak-like front as in the various spider-crabs; Oxystomata, including the Raninida, and named not from the character of the front but from that of the buccal frame which is usually narrowed forwards. In these tribes the bold and active habits, the striking colours, or the fantastic diversities of structure,have so long attracted remark that recent investigations, while adding a multitude of new species and supplying the specialist with an infinity of new details, have not materially altered the scientific standpoint. New light, however, has been thrown upon the "intellectual" capacity of Crustacea by the proof that the spider-crabs deliberately use changes of raiment to harmonize with their surroundings, donning and doffing various natural objects as we do our manufactured clothes. Others have the power of producing sounds, one use to which they put this faculty being apparently to signal from their burrow in the sand that they are "not at home" to an inopportune visitor. Deep-sea exploration has shown that some species have an immensely extended range, and still more, that species of the same genus, and genera of the same family, though separated by great intervals of space, may be closely allied in character. A curious effect of parasitism, well illustrated in crabs, though not confined to them, has been expounded by Professor Giard, namely, that it tends to obliterate the secondary sexual characters. Modern research has discovered no crab to surpass Macrocheira hampferi, De Haan, that can span between three and four yards with the tips of its toes, but at the other end of the scale it has yielded Collodes malabaricus, Alcock, "of which the carapace, in an adult and egg-laden female, is less than one-sixth of an inch in its greatest diameter." The most abyssal of all crabs yet known is Ethusina abyssicola, Smith, or what is perhaps only a variety of it, E. challengeri, Miers. Of the latter the "Albatross" obtained a specimen from a depth of 2232 fathoms (Faxon, 1895), of the former from 2221 fathoms, and of this S. I. Smith remarks that it has "distinctly faceted black eyes," although in them "there are only a very few visual elements at the tips of the immobile eye-stalks." The Brachyura anomala, or Dromiidea, "have preserved the external characters and probably also the organization of the Brachyura of the Secondary epoch" (Milne-Edwards and Bouvier, 1901). They agree with the true crabs in not having appendages (uropods) to the sixth segment of the pleon, the atrophy being complete in the Homolidae and Homolodromiidae, whereas in the Dromiidae and Dynomenidae a pair of small plates appear to be vestiges of these organs. In the family Homolidae stands the strange genus Latreillia, Roux, with long slender limbs and triangular carapace after the fashion of oxyrhynch spider-crabs. In Homola the carapace is quadrilateral. Between these two a very interesting link was discovered by the "Challenger" in the species Latreillopsis bispinosa, Henderson. Bouvier (1896) has shown that Palaeinachus longipes, Woodward, from the Forest Marble of Wiltshire, is in close relationship, not to the oxyrhynch Inachidae, but to the genera Homolodromia and Dicranodromia of the Homolodromiidae, and that the Jurassic crabs in general, of the family Prosoponidae (Meyer), are Dromiidea.

2. Macrura. - The Macrura anomala, or Anomura in restricted sense, are popularly known through the hermit-crabs alone. These only partially represent one of the three main divisions, Paguridea, Galatheidea, Hippidea. The first of these is subdivided into Pagurinea, Lithodinea, Lomisinea, each with a literature of its own. Among the Pagurinea is the Birgus latro, or robber-crab, whose expertness in climbing the coco-nut palm need no longer be doubted, since in recent years it has been noted and photographed by trustworthy naturalists in the very act. Alcock "observed one of these crabs drinking from a runnel of rain-water, by dipping the fingers of one of its chelipeds into the water and then carrying the wet fingers to its mouth." Hermits of the genus Coenobita he found feeding voraciously on nestling sea-terns. That pagurids must have the usually soft pleon or abdomen protected by the shell of a mollusc is now known to be subject to a multitude of exceptions. Birgus dispenses with a covering; Coenobita can make shift with half the shell of a coco-nut; Chlaenopagurus wraps itself up in a blanket of colonial polyps; Cancellus tanneri, Faxon, was found in a piece of dead coral rock; Xylopagurus rectus, A. Milne-Edwards, lodges in tubes of timber or bits of hollow reed. The last-named species has a straight symmetrical abdomen, with the penultimate segment expanded and strongly calcified to form a back-door to the very unconventional habitation. This it enters head-foremost from the rear, while "hermits" in general are forced to go backwards into their spiral or tapering shelters by the front. Some of the species can live in the ocean at a depth of two or three miles. Some can range inland up to a considerable height on mountains. The advantage that this group has derived from the adoption of mollusc xvrr. r5 a shells as houses or fortresses, ready built and light enough for easy transport, is obviously discounted by a twofold inconvenience. There is nothing to ensure that the supply will be equal to the demand, and Nature has not arranged that the borrowed tenement shall continue to grow with the growth of its new tenant. To meet these defects it is found that numerous species encourage or demand the companionship of various zoophytes, simple or colonial. These sometimes completely absorb the shell on which they are settled, but then act as a substitute for it, and in any case by their outgrowth they extend the limits of the dwelling, so that the inmate can grow in comfort without having to hunt or fight for a larger abode. Among the Lithodinea, or stone crabs, besides important readjustments of classification (Bouvier, 1895, 1896), should be FIG. 1. - Neolithodes grimaldii, A. Milne-Edwards and Bouvier.

noticed the evidence of their cosmopolitan range, and the species Neolithodes agassizii (Smith) and N. grimaldii, Milne-Edwards and Bouvier, which carry to an extreme the spinosity characteristic of the group (fig. 1). S. I. Smith's investigations on the early stages of Hippa talpoida, Say, were published in 1877.

With regard to the accessions to knowledge in the enormous group of the genuine Macrura, reference need only be made to the extensive reports in which Spence Bate, S. I. Smith, Faxon, WoodMason, Alcock, and others have made known the results of celebrated explorations. Various larval stages have been successfully investigated by Sars. Alcock (1901) describes from his own observation the newly hatched Phyllosoma larva of Thenus orientalis, Fabricius. An admirable discrimination of the larval and adult characters of the genus Sergestes has been given by H. J. Hansen (Proc. Zool. Soc., London, 1896). Singularity excites our wonder in Thaumastocheles zaleucus, v. Willemoes Suhm, which makes up for its vanished eyes by its extraordinarily elongate and dentated claws; in Psalidopus huxleyi, Wood-Mason and Alcock (1892), bristling with spikes from head to tail; in the Nematocarcinidae, with their long thread-like limbs and longer antennae; in species of Aristaeopsis reported by Chun from deep water off the east coast of Africa, bright red prawns nearly a foot long, with antennae about five times the length of the body. That certain species, particularly many from deep water, have disproportionately large eggs, is explained by the supposition that the young derive the advantage of being hatched in an advanced stage of development.

3. ScHIzoPODA. - This order of animals for the most part delicately beautiful, has for the moment five families - Lophogastridae, Eucopiidae, Euphausiidae, Mysidae and Anaspididae. In the Euphausiidae the digitiform-arborescent branchiae, as if conscious of their own extreme elegance, remain wholly uncovered. In the two preceding families they are partially covered. In theMysidae the branchiae are wanting, and some would form this family into a separate order, Mysidacea. In Anaspides, a peculiar fresh-water genus discovered in 1892 by G. M. Thomson on Mount Wellington, in Tasmania, the gills are not arborescent, and there are seven segments of the trunk free of the carapace (fig. 2). A membranaceous carapace separates the Euco piidae from the more solidly invested Lophogastridae. Among many papers that the student will find it necessary to consult may be mentioned the "Challenger" Report on Schizopoda, by Sars, 1885, dealing with the order at large; "British Schizopoda," by Norman Ann. Nat. Hist, (1892); "Decapoden and Schizopoden," PlanktonExpedition (Ortmann, 1893); "Euphausiidae," by Stebbing, Proc. Zool. Soc. (London, 1900); Mysidae of the Russian Empire, by Czernia y ski (1882-1883); and Mysidae of the Caspian, by Sars (1893-1895-1897).

4. Stomatopoda.-ThiS order, at one time a medley of heterogeneous forms, is now confined to the singularly compact group of the Squillidae. Here the articulation of the ocular segment is unusually distinct, and here two characters quite foreign to all the preceding groups come into view. The second maxillipeds are developed into powerful prehensile organs, and the branchiae, instead of being connected with the appendages of head and trunk, are developed on the pleopods, appendages of the abdomen. At least three segments of the trunk are left uncovered by the carapace. The developing eggs are not carried about by the mother, but deposited in her subaqueous burrow, "where they are aerated by the currents of water produced by the abdominal feet of the parent." An excellent synopsis of the genera and species is provided by R. P. Bigelow (Proc. U.S. Mus. vol. xvii., 1894). For the habits and peculiarities of these and many other Crustaceans, A. E. Verrill and S. I. Smith on the Invertebrates of Vineyard Sound should be consulted (1874). The general subject has been illuminated by the labours of Claus, Miers, Brooks ("Challenger" Report, 1886), and the latest word on the relationship between the various larvae and their respective genera has been spoken by H. J. Hansen (PlanktonExpedition Report, 1895). The striking forms of Alima and Erichthus, at one time regarded as distinct genera, are now with more or less certainty affiliated to their several squillid parents.

5. SYMPoDA. - This order of sessile-eyed decapods was absolutely unknown to science till 1779. A species certainly belonging to it was described by Lepekhin in 1780, but the obscure Gammarus esca, " food Gammarus" beloved of herrings, described by J. C. Fabricius in the preceding year, may also be one of its members. Nutritious possibilities are implied in Diastylis rathkii, Kroyer, one of the largest forms, which, though slender and rarely an inch long, in its favourite Arctic waters is found "in incalculable masses, in thousands of specimens" (Stuxberg, 1880). Far on in the 19th century FIG. 3. - Pseudocuma pectinatum, Sowinsky.

eminent naturalists were still debating whether in this group there were eyes or no eyes, whether the eyes were stalked or sessile, whether the animals observed were larval or adult. The American T. Say in 1818 gave a good description of a new species and founded the premier genus Diastylis, but other investigators derived little credit from the subject till more than sixty years after its introduction by the Russian Lepekhin. Then Goodsir, Kroyer, Lilljeborg, Spence Bate and one or two others made considerable advances, and in 1865 a memorable paper by G. O. Sars led the way to the great series of researches which he has continued to the present day. The name Cumacea, however, which he uses cannot be retained, being founded on the preoccupied name Cuma (Milne-Edwards, 1828). The more recent name Sympoda (see Willey, Results, pt. v. p. 609, 1900) alludes to the huddling together of the legs, which is conspicuous in most of the species. Ten families are now distinguishedDiastylidae, Lampropidae, Platyaspidae, Pseudocumidae, all with an articulated telson; without one, the Bodotriidae (formerly called Cumidae), Vaunthompsoniidae, Leuconidae, Nannastacidae, Campylaspidae, Procampylaspidae. All the Leuconidae and Procampylaspidae are blind, and some species in most of the other families. Usually the sides of the carapace are strangely produced into a mock rostrum in front of the ocular lobe, be it oculiferous or not. The last four or five segments of the trunk are free from the carapace. The slender pleon has always six distinct segments, the sixth carrying two-branched uropods, the preceding five armed with no pleopods in the female, whereas in the male the number of pairs varies from five to none. The resemblance of these creatures to miniature Macrura is alluded to in the generic name Nannastacus, meaning dwarf-lobster. In this genus alone of the known Sympoda the eyes sometimes form a pair, in accordance with the custom of all other malacostracan orders except this and of this order itself in the embryo (Sars, 1900). The most but not the only remarkable character lies in the first maxillipeds. These, with the main stem more or less pediform, have the epipod and exopod modified for respiratory purposes. The backward-directed epipods usually carry branchial vesicles. The forward-directed exopods either act as valves or form a tube (rarely two tubes), protensile and retractile, for regulating egress of water from the branchial regions. This mechanism as a whole is unique, although, as Sars observes, the epipod of the first maxillipeds has a respiratory function also in the Lophogastridae and Mysidae and in the cheliferous isopods. As a rule armature of FIG. 2. - Anaspides tasmaniae, Thomson.

the carapace is much more developed in the comparatively sedentary female than in the usually more active male. Only in the male do the second antennae attain considerable length, with strong resemblance to what is found in some of the Amphipoda. About 150 species distributed among thirty-four genera are now known, many from shallow water and from between tide-marks, some from very great depths. H. J. Hansen concludes that "they are all typically ground animals, and as yet no species has been taken under such conditions that it could be reckoned to the pelagic plankton." As they have been found in all zones and chiefly by a very few observers, it is probable that a great many more species remain to be discovered. In recent years thirteen species, all belonging to the same genus Pseudocuma (fig. 3), have been recorded by Sars from the Caspian Sea. A bibliography of the order is given in that author's Crustacea of Norway, vol. iii. (1899-1900).

6. IsoPODA. - This vast and populous order can be traced far back in geological time. It is now represented in all seas and lands, in fresh-water lakes and streams, and even in warm springs. It adapts itself to parasitic life not only in fishes, but in its own class Crustacea, and that in species of every order, its own included. In this process changes of structure are apt to occur, and sometimes unimaginable sacrifices of the normal appearance. The order has been divided into seven tribes, of which a fuller summary than can here be given will be found in Stebbing, History of Crustacea (1893). The first tribe, called Chelifera, from the usually chelate or claw-bearing first limbs, may be regarded as Isopoda anomala, of whieh some authors would form a separate order, Tanaidea. Like the genuine isopods, they have seven pairs of trunk-legs, but instead of having seven segments of the middle body (or peraeon) FIG. 4. - Rhabdosoma piratum, Stebbing.

normally free, they have the first one or two of its segments coalesced with the head. Instead of the breathing organs being furnished by the appendages of the pleon with the heart in their vicinity, the respiration is controlled by the maxillipeds, with the heart in the peraeon (see Delage, Arch. Zool. exper. et gen., vol. ix., 1881). There are two families, Tanaidae and Apseudidae. Occasionally the ocular lobes are articulated.

The genuine Isopoda are divided among the Flabellifera, in which the terminal segment and uropods form a flabellum or swimming fan; the Epicaridea, parasitic on Crustaceans; the Valvifera, in which the uropods fold valve-like over the branchial pleopods; the Asellota, in which the first pair of pleopods of the female are usually transformed into a single opercular plate; the Phreatoicidea, a fresh-water tribe, known as yet only from subterranean waters in New Zealand and an Australian swamp nearly 6000 ft. above sea-level; and lastly, the Oniscidea, which are terrestrial. Only the last of these, under the contemptuous designation of wood-lice, has established a feeble claim to popular recognition. Few persons hear without surprise that England itself possesses more than a score of species in this airbreathing tribe. Those known from the world at large number hundreds of species, distributed among dozens of genera in six families. That a wood-louse and a land-crab are alike Malacostracans, and that they have by different paths alike become adapted to terrestrial life, are facts which even a philosopher might condescend to notice. Of the other tribes which are aquatic there is not space to give even the barest outline. Their swarming multitudes are of enormous importance in the economy of the sea. If in their relation to fish it must be admitted that many of them plague the living and devour the dead, in return the fish feed rapaciously upon them. Among the most curious of recent discoveries is that relating to some of the parasitic Cymothoidae, as to which Bullar has shown that the same individual can be developed first as a male and then as a female. Of lately discovered species the most striking is one of the deep-sea Cirolanidae, Bathynomus giganteus, A. M. Edwards (1879), which is unique in having supplementary ramified branchiae developed at the bases of the pleopods. Its eyes are said to contain nearly 4000 facets. The animal attains what in this order is the monstrous size of 9 in. by 4. A general uniformity of the trunk-limbs in Isopoda justifies the ordinal name, but the valviferous Astacillidae, and among the Asellota the Munnopsidae, offer some remarkable exceptions to this characteristic. Among many essential works on this group may be named the Monogr. Cymothoarum of Sch16dte and Meinert (1879-1883); "Challenger" Report, Beddard (1884-1886); Cirolanidae, H. J. Hansen (1890); Isopoda Terrestria, Budde-Lund (1885); Bopyridae, Bonnier (1900); Crustacea of Norway, vol. ii. (Isopoda), Sars (1896-1899), while their multitude precludes specification of important contributions by Benedict, Bovallius, Chilton, Dohrn, Dollfus, Fraisse, Giard and Bonnier, Harger, Haswell, Kossmann, Miers, M`Murrich, Norman, Harriet Richardson, Ohlin, Studer, G. M. Thomson, A. O. Walker, Max Weber and many others.

7. Amphipoda. - As in the genuine Isopoda, the eyes of Amphipoda are always sessile, and generally paired, and, in contrast to crabs and lobsters, these two groups have only four pairs of mouthorgans instead of six, but seven pairs of trunk-legs instead of five. From the above-named isopods the present order is strongly differentiated by having heart and breathing organs not in the pleon, but in the peraeon, or middle body, the more or less simple branchial vesicles being attached to some or all of the last six pairs of trunklegs. Normally the pleon carries six pairs of two-branched appendages, of which the first three are much articulated flexible swimming feet, the last three few-jointed comparatively indurated uropods. There are three tribes, Gammaridea, Caprellidea, Hyperiidea. The middle one contains but two families, the cylindrical and often thread-like skeleton shrimps, Caprellidae, and their near cousins, the broad, flattened, so-called whale-lice, Cyamidae. This tribe has the pleon dwindled into insignificance, whereas in the other two tribes it is powerfully developed. The Hyperiidea are distinguished by having their maxillipeds never more than three-jointed. In the companion tribes these appendages have normally seven joints, and always more than three. The order thus sharply divided is united by an intimate interlacing of characters, and forms a compact whole at present defying intrusion from any other crustacean group. Since 1775, when J. C. Fabricius instituted the genus Gammarus for five species, of which only three were amphipods, while he left five other amphipods in the genus Oniscus, from this total of eight science has developed the order, at first very slowly, but of late by great leaps and bounds, so that now the Gammaridea alone comprise more than 1300 species, distributed among some 300 genera and 39 families. They burrow in the sands of every shore; they throng the weeds between tide-marks; they ascend all streams; they are found in deep wells, in caverns, in lakes; in Arctic waters they swarm in numbers beyond computation; they find lodgings on crabs, on turtles, on weed-grown buoys; they descend into depths of the ocean down to hundreds or thousands of fathoms; they are found in mountain streams as far above sea-level as some of their congeners live below it. The Talitridae, better known as sandhoppers, can forgo the briny shore and content themselves with the damp foliage of inland forests or casual humidity in the crater of an extinct volcano. Over the ocean surface, as well as at various depths, float and swim innumerable Hyperiidea - the wonderful Phronima, glass-like in its glassy barrel hollowed out of some Tunicate; the Cystisoma, 4 or 5 in. long, with its eye-covered head; the Rhabdosoma, like a thin rod of glass, with needle-like head and tail, large eyes, but limbs and mouth-organs all in miniature, and the second antennae of the male folding up like a carpenter's rule (fig. 4). On jelly-fishes are to be found species of Hyperia and their kindred, so fat and wholesome that they have been commended to shipwrecked men in open boats as an easily procurable resource against starvation. Many of the Amphipoda are extremely voracious. Some of them are even cannibals. The Cyamidae afflict the giant whale by nibbling away its skin; the Chelura terebrans is destructive to submerged timber. But, on the other hand, they largely help to clear the sea and other waters of refuse and carrion, and for fishes, seals and whales they are food desirable and often astoundingly copious. From the little flea-like species, scarcely a tenth of an inch long, up to the great and rare but cosmopolitan Eurythenes gryllus, Lichtenstein, and the still larger Alicella gigantea, Chevreux, nearly half a foot long, captured by the prince of Monaco from a depth of 2936 fathoms, not one of these ubiquitous, uncountable hordes has ever been accused of assailing man. For the naturalist they have the recommendation that many are easy to obtain, that most, apart from the very minute, are easy to handle, and that all, except as to the fleeting colours, are easy to preserve.

A nearly complete bibliography of the order down to 1888 will be found in the "Challenger" Reports, vol. xxviii., and supplementary notices in Della Valle's Monograph of the Gammarini (1893)', the scope of his work, however, not covering the Hyperiidea and Oxycephalidae of Bovallius (1889, 1890); but since these dates very numerous additions to the literature have been made by Birula, Bonnier, Norman, Walker and others, especially the Crustacea of Norway, vol. i. (Amphipoda), Sars (1890-1895), demanding attention, and the quite recent Amphipoda of the Hirondelle, Chevreux (1900), and Hyperiidea of the Plankton-Expedition, Vosseler (1901). (T. R. R. S.)



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