1. Gymnophylla.—These singular crustaceans have long soft flexible bodies, the eyes stalked and movable, the first antennae small and filiform, the second lamellar in the female, in the male prehensile; this last character gives rise to some very fanciful developments. There are three families, two of which form companies rather severely limited. Thus the Polyartemiidae, which compensate themselves for their stumpy little tails by having nineteen instead of the normal eleven pairs of branchial feet, consist exclusively of Polyartemia forcipata (Fischer, 1851). This species from the high north of Europe and Asia carries green eggs, and above them a bright pattern in ultramarine (Sars, 1896, 1897). The Thamnocephalidae have likewise but a single species, Thamnocephalus platyurus (Packard, 1877), which justifies its title “bushy-head of the broad tail” by a singularity at each end. Forward from the head extends a long ramified appendage described as the “frontal shrub,” backward from the fourth abdominal segment of the male spreads a fin-like expansion which is unique. In the ravines of Kansas, pools supplied by torrential rains give birth to these and many other phyllopods, and in turn “millions of them perish by the drying up of the pools in July” (Packard). The remaining family, the Branchipodidae, includes eight genera. In the long familiar Branchipus, Chirocephalus and Streptocephalus the males have frontal appendages, but these are wanting in the “brine-shrimp” Artemia, and the same want helps to distinguish Branchinecta (Verrill, 1869) from the old genus Branchipus. Of Branchiopsyllus (Sars, 1897) the male is not yet known, but in his genera of the same date, the Siberian Artemiopsis and the South African Branchipodopsis (1898), there is no such appendage. Of the last genus the type species B. hodgsoni belongs to Cape Colony, but the specimens described were born and bred and observed in Norway. For the study of fresh-water Entomostraca large possibilities are now opened to the naturalist. A parcel of dried mud, coming for example from Palestine or Queensland, and after an indefinite interval of time put into water in England or elsewhere, may yield him living forms, both new and old, in the most agreeable variety. Some caution should be used against confounding accidentally introduced indigenous species with those reared from the imported eggs. Those, too, who send or bring the foreign soil should exercise a little thought in the choice of it, since dry earth that has never had any Entomostraca near it at home will not become fertile in them by the mere fact of exportation.
2. Notophylla.—In this division the body is partly covered by a broad shield, united in front with the head; the eyes are sessile, the first antennae are small, the second rudimentary or wanting; of the numerous feet, sometimes sixty-three pairs, exceeding the number of segments to which they are attached, the first pair are more or less unlike the rest, and in the female the eleventh have the epipod and exopod (flabellum and sub-apical lobe of Lankester) modified to form an ovisac. Development begins with a nauplius stage. Males are very rare. The single family Apodidae contains only two genera, Apus and its very near neighbour Lepidurus. Apus australiensis (Spencer and Hall, 1896) may rank as the largest of the Entomostraca, reaching in the male, from front of shield to end of telson, a length of 70 mm., in the female of 64 mm. In a few days, or at most a fortnight, after a rainfall numberless specimens of these sizes were found swimming about, “and as not a single one was to be found in the water-pools prior to the rain, these must have been developed from the egg.” Similarly, in Northern India Apus himalayanus was “collected from a stagnant pool in a jungle four days after a shower of rain had fallen,” following a drought of four months (Packard).
3. Conchophylla.—Though concealed within the bivalved shell-cover, the mouth-parts are nearly as in the Gymnophylla, but the flexing of the caudal part is in contrast, and the biramous second antennae correspond with what is only a larval character in the other phyllopods. In the male the first one or two pairs of feet are modified into grasping organs. The small ova are crowded beneath the dorsal part of the valves. The development usually begins with a nauplius stage (Sars, 1896, 1900). There are four families: (a) The Limnadiidae, with feet from 18 to 32 pairs, comprise four (or five) genera. Of these Limnadella (Girard, 1855) has a single eye. It remains rather obscure, though the type species originally “was discovered in great abundance in a roadside puddle subject to desiccation.” Limnadia (Brongniart, 1820) is supposed to consist of species exclusively parthenogenetic. But when asked to believe that males never occur among these amazons, one cannot but remember how hard it is to prove a negative. (b) The Lynceidae, with not more than twelve pairs of feet. This family is limited to the species, widely distributed, of the single genus Lynceus, established by O.F. Müller in 1776 and 1781, and first restricted by Leach in 1816 in the Encyclopaedia Britannica (art. “Annulosa,” of that edition). Leach there assigns to it the single species L. brachyurus (Müller), and as this is included in the genus Limnetis (Lovén, 1846), that genus must be a synonym of Lynceus as restricted. (c) Leptestheriidae. Estheria (Rüppell, 1837) was instituted for the species dahalacensis, which Sars includes in his genus Leptestheria (1898); but Estheria was already appropriated, and of its synonyms Cyzicus (Audouin, 1837) is lost for vagueness, while Isaura (Joly, 1842) is also appropriated, so that Leptestheria becomes the name of the typical genus, and determines the name of the family. (d) Cyclestheriidae. This family consists of the single species Cyclestheria hislopi (Baird), reported from India, Ceylon, Celebes, Australia, East Africa and Brazil. Sars (1887) having had the opportunity of raising it from dried Australian mud, found that, unlike other phyllopods, but like the Cladocera, the parent keeps its brood within the shell until their full development.
Cladocera.—In this suborder the head is more or less distinct, the rest of the body being in general laterally compressed and covered by a bivalved test. The title “branching horns” alludes to the second antennae, which are two-branched except in the females of Holopedium, with each branch setiferous, composed of only two to four joints. The mandibles are without palp. The pairs of feet are four to six. The eye is single, and in addition to the eye there is often an “eye-spot,” Monospilus being unique in having the eye-spot alone and no eye, while Leydigiopsis (Sars, 1901) has an eye with an eye-spot equal to it or larger. The heart has a pair of venous ostia, often blending into one, and an anterior arterial aorta. Respiration is conducted by the general surface, by the branchial lamina (external branch) of the feet, and the vesicular appendage (when present) at the base of this branch. The “abdomen,” behind the limbs, is usually very short, occasionally very long. The “postabdomen,” marked off by the two postabdominal setae, usually has teeth or spines, and ends in two denticulate or ciliate claws, or it may be rudimentary, as in Polyphemus. Many species have a special glandular organ at the back of the head, which Sida crystallina uses for attaching itself to various objects. The Leydigian or nuchal organ is supposed to be auditory and to contain an otolith. The female lays two kinds of eggs—“summer-eggs,” which develop without fertilization, and “winter-eggs” or resting eggs, which require to be fertilized. The latter in the Daphniidae are enclosed in a modified part of the mother’s shell, called the ephippium from its resemblance to a saddle in shape and position. In other families a less elaborate case has been observed, for which Scourfield has proposed the term protoephippium. In Leydigia he has recently found a structure almost as complex as that of the Daphniidae. In some families the resting eggs escape into the water without special covering. Only the embryos of Leptodora are known to hatch out in the nauplius stage. Penilia (Dana, 1849) is perhaps the only exclusively marine genus. The great majority of the Cladocera belong to fresh water, but their adaptability is large, since Moina rectirostris (O.F. Müller) can equally enjoy a pond at Blackheath, and near Odessa live in water twice as salt as that of the ocean. In point of size a Cladoceran of 5 mm. is spoken of as colossal.
Dr Jules Richard in his revision (1895) retains the sections proposed by Sars in 1865, Calyptomera and Gymnomera. The former, with the feet for the most part concealed by the carapace, is subdivided into two tribes, the Ctenopoda, or “comb-feet,” in which the six pairs of similar feet, all branchial and nonprehensile, are furnished with setae arranged like the teeth of a comb, and the Anomopoda, or “variety-feet,” in which the front feet differ from the rest by being more or less prehensile, without branchial laminae.
The Ctenopoda comprise two families: (a) the Holopediidae, with a solitary species, Holopedium gibberum (Zaddach), queerly clothed in a large gelatinous involucre, and found in mountain tarns all over Europe, in large lakes of N. America, and also in shallow ponds and waters at sea-level; (b) the Sididae, with no such involucre, but with seven genera, and rather more than twice as many species. Of Diaphanosoma modiglianii Richard says that at different points of Lake Toba in Sumatra millions of specimens were obtained, among which he had not met with a single male.
The Anomopoda are arranged in four families, all but one very extensive. (a) Daphniidae. Of the seven genera, the cosmopolitan Daphnia contains about 100 species and varieties, of which Thomas Scott (1899) observes that “scarcely any of the several characters that have at one time or another been selected as affording a means for discriminating between the different forms can be relied on as satisfactory.” Though this may dishearten the systematist, Scourfield (1900) reminds us that “It was in a water-flea that Metschnikoff first saw the leucocytes (or phagocytes) trying to get rid of disease germs by swallowing them, and was so led to his epoch-making discovery of the part played by these minute amoeboid corpuscles in the animal body.” For Scapholeberis mucronata (O.F. Müller), Scourfield has shown how it is adapted for movement back downwards in the water along the underside of the surface film, which to many small crustaceans is a dangerously disabling trap. (b) Bosminidae. To Bosmina (Baird, 1845) Richard added Bosminopsis in 1895. (c) Macrotrichidae. In this family Macrothrix (Baird, 1843) is the earliest genus, among the latest being Grimaldina (Richard, 1892) and Jheringula (Sars, 1900). Dried mud and vegetable débris from S. Paulo in Brazil supplied Sars with representatives of all the three in his Norwegian aquaria, in some of which the little Macrothrix elegans “multiplied to such an extraordinary extent as at last to fill up the water with immense shoals of individuals.” “The appearance of male specimens was always contemporary with the first ephippial formation in the females.” For Streblocerus pygmaeus, grown under the same conditions, Sars observes: “This is perhaps the smallest of the Cladocera known, and is hardly more than visible to the naked eye,” the adult female scarcely exceeding 0.25 mm. Yet in the next family Alonella nana (Baird) disputes the palm and claims to be the smallest of all known Arthropoda. (d) Chydoridae. This family, so commonly called Lynceidae, contains a large number of genera, among which one may usually search in vain, and rightly so, for the genus Lynceus. The key to the riddle is to be found in the Encyclopaedia Britannica for 1816. There, as above explained, Leach began the subdivision of Müller’s too comprehensive genus, the result being that Lynceus belongs to the Phyllopoda, and Chydorus (Leach, 1816) properly gives its name to the present family, in which the doubly convoluted intestine is so remarkable. Of its many genera, Leydigia, Leydigiopsis, Monospilus have been already mentioned. Dadaya macrops (Sars, 1901), from South America and Ceylon, has a very large eye and an eye-spot fully as large, but it is a very small creature, odd in its behaviour, moving by jumps at the very surface of the water. “To the naked eye it looked like a little black atom darting about in a most wonderful manner.”
| Fig. 1.—Dolops ranarum (Stuhlmann). |
The Gymnomera, with a carapace too small to cover the feet, which are all prehensile, are divided also into two tribes, the Onychopoda, in which the four pairs of feet have a toothed maxillary process at the base, and the Haplopoda, in which there are six pairs of feet, without such a process. To the Polyphemidae, the well-known family of the former tribe, Sars in 1897 added two remarkable genera, Cercopagis, meaning “tail with a sling,” and Apagis, “without a sling,” for seven species from the Sea of Azov. The Haplopoda likewise have but a single family, the Leptodoridae, and this has but the single genus Leptodora (Lilljeborg, 1861). Dr Richard (1895, 1896) gives a Cladoceran bibliography of 601 references.