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THE LIFE OF CRUSTACEA
BY
W. T. CALMAN, D.Sc.
WITH THIRTY-TWO PLATES AND EIGHTY-FIVE FIGURES
METHUEN & CO. LTD.
36 ESSEX STREET W.C.
LONDON
First Published in 1911
PREFACE
This sketch of the Natural History of the Crustacea deals chiefly with their habits and modes of life, and attempts to provide, for readers unfamiliar with the technicalities of Zoology, an account of some of the more important scientific problems suggested by a study of the living animals in relation to their environment.
I am indebted to the Trustees of the British Museum for leave to reproduce certain figures prepared for the "Guide to the Crustacea, Arachnida, Onychophora, and Myriopoda exhibited in the Department of Zoology"; also to Sir Ray Lankester, K.C.B., F.R.S., and to Messrs. A. and C. Black for the use of a number of figures from my volume on Crustacea in the "Treatise on Zoology," edited by Sir Ray Lankester.
The source of these figures is indicated in the explanation attached to each. Of the remaining illustrations, some are reproduced from photographs of specimens in the collection of the British Museum; the others have been drawn from Nature, or copied from the original figures of various authors, by Miss Gertrude M. Woodward, to whom I am much indebted for the care and skill which she has given to their preparation.
W. T. C.
CONTENTS
LIST OF ILLUSTRATIONS IN THE TEXT
FULL-PAGE PLATES
Group of a Common Species of Acorn-shell or Rock Barnacle (Balanus balanoides). Natural Size } 42 IV. Penæus caramote, from the Mediterranean. About Half Natural Size. (From Brit. Mus. Guide) 57 V. The Common Spiny Lobster (Palinurus vulgaris). Much reduced. (From Brit. Mus. Guide) 59 VI. Munida rugosa. British. Reduced 60 VII. The Common Hermit Crab, Eupagurus bernhardus, in the Shell of a Whelk. Reduced. (From Brit. Mus. Guide) 62 VIII. The "Northern Stone Crab," Lithodes maia. Much reduced. The Last Pair of Legs are folded out of Sight in the Gill Chambers. (From Brit. Mus. Guide) 63 IX. { The Common Shore Crab (Carcinus mænas). Reduced
Dromia vulgaris, carrying on its Back a Mass of the Sponge, Clione celata. British. Reduced } 68 X. Calappa flammea. Brazil. Reduced 72 XI. The Giant Japanese Crab, Macrocheira kæmpferi. Male. The Scale of the Figure is given by a Two-foot Rule placed below the Specimen. (From Brit. Mus. Guide) 76 XII. Squilla mantis, from the Mediterranean. About One-half Natural Size. (From Brit. Mus. Guide 82 XIII. { A Swimming Crab, Portunus depurator. British. Reduced
A Spider Crab, Maia squinado, dressed in Fragments of Weeds. British. Reduced } 96 XIV. { Corystes cassivelaunus. Male (on Left) and Female (on Right). British. Reduced
Albunea symnista, One of the Hippidea. Indian Seas. Reduced } 100 XV. { Ocypode cursor. West Africa. Reduced
Gelasimus tangeri. Male Above, Female Below. West Africa. Reduced } 104 XVI. A Deep-sea Hermit Crab, Parapagurus pilosimanus, sheltered by a Colony of Epizoanthus. From Deep Water off the West of Ireland. Slightly reduced 124 XVII. A Deep-sea Prawn, Nematocarcinus undulatipes. Slightly reduced. (From Brit. Mus. Guide) 128 XVIII. Bathynomus giganteus. About One-half Natural Size. (From Lankester's "Treatise on Zoology," after Milne-Edwards and Bouvier) 131 XIX. { Latreillia elegans, One of the Dromiacea which esembles a Spider Crab. From the Mediterranean. Natural Size
The Gulf-weed Crab, Planes minutus. Slightly enlarged } 155 XX. {
The Land Crayfish, Engæus cunicularis. Tasmania. Natural Size } 177 XXI. Palæmon jamaicensis. A Large Freshwater Prawn of the Family Palæmonidæ. West Indies. Much reduced 179 XXII. Atya scabra. A Freshwater Prawn of the Family Atyidæ, West Indies. Reduced 180 XXIII. { The River Crab of Southern Europe, Potamon edule (or Telphusa fluviatilis). Reduced
Sesarma chiragra. A Freshwater Crab of the Family Grapsidæ. From Brazil. Slightly reduced } 182 XXIV. Æglea lævis. South America. Natural Size 184 XXV. The Blind Crayfish of the Mammoth Cave of Kentucky, Cambarus pellucidus. Natural Size 186 XXVI. { A West Indian Land Crab, Gecarcinus ruricola Reduced
A Land Hermit Crab, Cœnobita rugosa. Reduced } 190 XXVII. The Coconut Crab, Birgus latro. Much reduced 196 XXVIII. Group of Barnacles, Coronula diadema, on the Skin of a Whale. Japan. Reduced 209 XXIX. { Cymothoa œstrum. An Isopod Parasite of Fish. Slightly Enlarged
Sacculina carcini attached under the Abdomen of a Common Shore Crab. Reduced } 220 XXX. The "Norway Lobster," Nephrops norvegicus. About One-third Natural Size. (From Brit. Mus. Guide) 240 XXXI. The Common Edible Crab, Cancer pagurus. British. Much reduced 248 XXXII. Piece of Timber From Ryde Pier, showing Damage caused by Limnoria AND Chelura. (From Brit. Mus. Guide) 255
THE LIFE OF CRUSTACEA
CHAPTER I
INTRODUCTORY
Everyone has some acquaintance with the animals that are grouped by naturalists under the name Crustacea. The edible Crabs, Lobsters, Prawns, and Shrimps, are at least superficially familiar, either as brought to the table or as displayed in the fishmonger's, and the most unobservant of seaside visitors must have had his attention attracted by living specimens of some of the more obtrusive species, such as the common Shore Crab. Many, however, will be surprised to learn that the Barnacles coating the rocks on the seashore, the Sand-hoppers of the beach, and the Woodlice of our gardens, are members of the same class. Still less is it suspected, by those who have not given special attention to the subject, that the living species of the group number many thousands, presenting strange diversities of structure and habits, and playing important parts in the general economy of Nature.
In addition to those just mentioned, a few Crustacea are sufficiently well known to be distinguished by popular names, such, for example, as Crayfish and Hermit Crabs, but for the vast majority no names are available except those of technical zoology. In the following pages, therefore, while technical terms have been introduced as sparingly as possible, the unfamiliarity of the animals themselves makes it needful to use many unfamiliar names.
In the classification of the Animal Kingdom, the Crustacea form one of the divisions of a comprehensive group, or Phylum, known as Arthropoda. The typical members of this group have a more or less firm external skeleton, the body is divided into segments, there are jointed
It is not easy to summarize in a few words the characters common to all Crustacea, and distinguishing them from the other groups of Arthropoda. As a rough guide to classification, it is useful to remember that an Insect can generally be recognized by having three pairs of walking legs, an Arachnid by having four pairs, and a Centipede or Millipede by having a great many pairs, all nearly alike. The Crustacea, on the other hand, show great diversity in the number and arrangement of their walking or swimming legs, but they rarely show any special resemblance to those of the other large groups of Arthropoda. Thus, for example, a common species of Woodlouse, Armadillidium vulgare, is very similar at first sight to the Millipede Glomeris marginata, but it has only seven pairs of walking legs, while the Millipede has seventeen or nineteen pairs.
More precisely, it may be said of the great majority of Crustacea that they are aquatic animals, breathing by gills or by the general surface of the body, having two pairs of "feelers," or antennæ, on the front part of the head, and at least three pairs of jaws. Exceptions to each of these statements will be mentioned in later chapters in dealing with parasites and other highly modified types. In such cases, however, the larval or young stages afford indications of affinity, and comparison with less modified forms enables us to trace a connection with the typical Crustacea.
The best way to form a conception of a group of animals, however, is not to attempt in the first place to define its limits, but to begin by studying the structure of some typical and central species, and afterwards to note the divergences from this type presented by other members of the group. Speaking very generally, it may be said that these divergences are of two kinds. On the one hand there are characters that have no apparent relation to the animal's habits and mode of life, and on the other hand there are modifications of structure which are more or less plainly of use to the animal. It is to characters of the former class that we look for evidence of an animal's affinities, and it is upon them that our systems of classification are chiefly based. The characters of the second class—"adaptive" characters, as they are called—become of importance when we study the animal "as a going concern," so to speak, and endeavour to understand how its life is carried on in relation to its surroundings.
In pursuance of this plan of study, the next chapter will be devoted to a description of the Common Lobster as a type of the Crustacea. In the third chapter a survey of the classification of the group will be given; since, however, the characters on which the classification is based cannot be explained fully without entering into technical details which are beyond the scope of this work, this survey will be restricted to what is necessary for comprehension of the succeeding chapters. In the fourth chapter some account is given of the young or larval stages of Crustacea, and of the changes they undergo in the course of development.
In the next five chapters the Crustacea are classified according to their habitats, and those living in the shallow waters, the depths, and the surface of the ocean, in the fresh waters, and on land, are discussed in turn; while a separate chapter is devoted to the curious forms that live as parasites on, or as associates with, other animals. The last two chapters deal respectively with the Crustacea as they affect man, and with the past history of the group as revealed by fossil remains.
CHAPTER II
THE LOBSTER AS A TYPE OF CRUSTACEA
The most noticeable feature distinguishing the Lobster[2] (Fig. 1) at first sight from other familiar animals is the jointed shelly armour that encases its body and limbs. Over the fore part of the body this armour is continuous, forming a shield, or carapace, which projects in front, between the eyes, as a toothed beak, or rostrum; on the hinder part—the tail, or abdomen—it is divided into six segments, or somites, connected with each other by movable joints. Each of these somites carries on the under-side a pair of fin-like limbs, or swimmerets, the last pair of which (uropods) are much larger than the others, and are spread out at the sides of a middle tail-plate, or telson, forming what is known as the tail-fan. Since the fore part of the body also has a series of paired limbs, constructed, as will be shown later, on the same plan as the swimmerets, it is concluded that this part also is built up of somites, which have become soldered together. That this conclusion is correct is shown by comparison with some of the lower Crustacea in which this part of the body is divided up into eight separate somites, like those of the abdomen, each carrying, in place of the swimmerets, a pair of walking legs. In front of these eight somites, forming what is called the thorax, is the head—a part of the body which is never, in any Crustacean, broken up into distinct somites, but which, since it carries five pairs of appendages, must consist of at least five somites. The part of the body covered by the Lobster's carapace includes both the head and the thorax, and is known, therefore, as the cephalothorax. It is necessary to bear in mind that the parts of the body to which the names head, thorax, and abdomen, are applied in Crustacea are by no means exactly equivalent to those which bear the same names in Insects, for example, and that, beyond a rough similarity in position, they have no sort of relation to the parts so named in the body of a vertebrate animal.