SECONDARY SEXUAL CHARACTERS IN THE LOWER CLASSES OF THE ANIMAL KINGDOM.
These characters absent in the lowest classes—Brilliant
colours—Mollusca —Annelids—Crustacea, secondary sexual characters
strongly developed; dimorphism; colour; characters not acquired before
maturity—Spiders, sexual colours of; stridulation by the
males—Myriapoda.
With animals belonging to the lower classes, the two sexes are not
rarely united in the same individual, and therefore secondary sexual
characters cannot be developed. In many cases where the sexes are
separate, both are permanently attached to some support, and the one
cannot search or struggle for the other. Moreover it is almost certain
that these animals have too imperfect senses and much too low mental
powers to appreciate each other’s beauty or other attractions, or to
feel rivalry.
Hence in these classes or sub-kingdoms, such as the Protozoa,
Coelenterata, Echinodermata, Scolecida, secondary sexual characters, of
the kind which we have to consider, do not occur: and this fact agrees
with the belief that such characters in the higher classes have been
acquired through sexual selection, which depends on the will, desire,
and choice of either sex. Nevertheless some few apparent exceptions
occur; thus, as I hear from Dr. Baird, the males of certain Entozoa, or
internal parasitic worms, differ slightly in colour from the females;
but we have no reason to suppose that such differences have been
augmented through sexual selection. Contrivances by which the male
holds the female, and which are indispensable for the propagation of
the species, are independent of sexual selection, and have been
acquired through ordinary selection.
Many of the lower animals, whether hermaphrodites or with separate
sexes, are ornamented with the most brilliant tints, or are shaded and
striped in an elegant manner; for instance, many corals and
sea-anemones (Actiniae), some jelly-fish (Medusae, Porpita, etc.), some
Planariae, many star-fishes, Echini, Ascidians, etc.; but we may
conclude from the reasons already indicated, namely, the union of the
two sexes in some of these animals, the permanently affixed condition
of others, and the low mental powers of all, that such colours do not
serve as a sexual attraction, and have not been acquired through sexual
selection. It should be borne in mind that in no case have we
sufficient evidence that colours have been thus acquired, except where
one sex is much more brilliantly or conspicuously coloured than the
other, and where there is no difference in habits between the sexes
sufficient to account for their different colours. But the evidence is
rendered as complete as it can ever be, only when the more ornamented
individuals, almost always the males, voluntarily display their
attractions before the other sex; for we cannot believe that such
display is useless, and if it be advantageous, sexual selection will
almost inevitably follow. We may, however, extend this conclusion to
both sexes, when coloured alike, if their colours are plainly analogous
to those of one sex alone in certain other species of the same group.
How, then, are we to account for the beautiful or even gorgeous colours
of many animals in the lowest classes? It appears doubtful whether such
colours often serve as a protection; but that we may easily err on this
head, will be admitted by every one who reads Mr. Wallace’s excellent
essay on this subject. It would not, for instance, at first occur to
any one that the transparency of the Medusae, or jelly-fish, is of the
highest service to them as a protection; but when we are reminded by
Haeckel that not only the Medusae, but many floating Mollusca,
crustaceans, and even small oceanic fishes partake of this same
glass-like appearance, often accompanied by prismatic colours, we can
hardly doubt that they thus escape the notice of pelagic birds and
other enemies. M. Giard is also convinced (1. ‘Archives de Zoolog.
Exper.’ Oct. 1872, p. 563.) that the bright tints of certain sponges
and ascidians serve as a protection. Conspicuous colours are likewise
beneficial to many animals as a warning to their would-be devourers
that they are distasteful, or that they possess some special means of
defence; but this subject will be discussed more conveniently
hereafter.
We can, in our ignorance of most of the lowest animals, only say that
their bright tints result either from the chemical nature or the minute
structure of their tissues, independently of any benefit thus derived.
Hardly any colour is finer than that of arterial blood; but there is no
reason to suppose that the colour of the blood is in itself any
advantage; and though it adds to the beauty of the maiden’s cheek, no
one will pretend that it has been acquired for this purpose. So again
with many animals, especially the lower ones, the bile is richly
coloured; thus, as I am informed by Mr. Hancock, the extreme beauty of
the Eolidae (naked sea-slugs) is chiefly due to the biliary glands
being seen through the translucent integuments—this beauty being
probably of no service to these animals. The tints of the decaying
leaves in an American forest are described by every one as gorgeous;
yet no one supposes that these tints are of the least advantage to the
trees. Bearing in mind how many substances closely analogous to natural
organic compounds have been recently formed by chemists, and which
exhibit the most splendid colours, it would have been a strange fact if
substances similarly coloured had not often originated, independently
of any useful end thus gained, in the complex laboratory of living
organisms.
THE SUB-KINGDOM OF THE MOLLUSCA.
Throughout this great division of the animal kingdom, as far as I can
discover, secondary sexual characters, such as we are here considering,
never occur. Nor could they be expected in the three lowest classes,
namely, in the Ascidians, Polyzoa, and Brachiopods (constituting the
Molluscoida of some authors), for most of these animals are permanently
affixed to a support or have their sexes united in the same individual.
In the Lamellibranchiata, or bivalve shells, hermaphroditism is not
rare. In the next higher class of the Gasteropoda, or univalve shells,
the sexes are either united or separate. But in the latter case the
males never possess special organs for finding, securing, or charming
the females, or for fighting with other males. As I am informed by Mr.
Gwyn Jeffreys, the sole external difference between the sexes consists
in the shell sometimes differing a little in form; for instance, the
shell of the male periwinkle (Littorina littorea) is narrower and has a
more elongated spire than that of the female. But differences of this
nature, it may be presumed, are directly connected with the act of
reproduction, or with the development of the ova.
The Gasteropoda, though capable of locomotion and furnished with
imperfect eyes, do not appear to be endowed with sufficient mental
powers for the members of the same sex to struggle together in rivalry,
and thus to acquire secondary sexual characters. Nevertheless with the
pulmoniferous gasteropods, or land-snails, the pairing is preceded by
courtship; for these animals, though hermaphrodites, are compelled by
their structure to pair together. Agassiz remarks, “Quiconque a eu
l’occasion d’observer les amours des limaçons, ne saurait mettre en
doute la séduction deployée dans les mouvements et les allures qui
préparent et accomplissent le double embrassement de ces
hermaphrodites.” (2. ‘De l’Espèce et de la Class.’ etc., 1869, p. 106.)
These animals appear also susceptible of some degree of permanent
attachment: an accurate observer, Mr. Lonsdale, informs me that he
placed a pair of land-snails, (Helix pomatia), one of which was weakly,
into a small and ill-provided garden. After a short time the strong and
healthy individual disappeared, and was traced by its track of slime
over a wall into an adjoining well-stocked garden. Mr. Lonsdale
concluded that it had deserted its sickly mate; but after an absence of
twenty-four hours it returned, and apparently communicated the result
of its successful exploration, for both then started along the same
track and disappeared over the wall.
Even in the highest class of the Mollusca, the Cephalopoda or
cuttle-fishes, in which the sexes are separate, secondary sexual
characters of the present kind do not, as far as I can discover, occur.
This is a surprising circumstance, as these animals possess
highly-developed sense-organs and have considerable mental powers, as
will be admitted by every one who has watched their artful endeavours
to escape from an enemy. (3. See, for instance, the account which I
have given in my ‘Journal of Researches,’ 1845, p. 7.) Certain
Cephalopoda, however, are characterised by one extraordinary sexual
character, namely that the male element collects within one of the arms
or tentacles, which is then cast off, and clinging by its sucking-discs
to the female, lives for a time an independent life. So completely does
the cast-off arm resemble a separate animal, that it was described by
Cuvier as a parasitic worm under the name of Hectocotyle. But this
marvellous structure may be classed as a primary rather than as a
secondary sexual character.
Although with the Mollusca sexual selection does not seem to have come
into play; yet many univalve and bivalve shells, such as volutes,
cones, scallops, etc., are beautifully coloured and shaped. The colours
do not appear in most cases to be of any use as a protection; they are
probably the direct result, as in the lowest classes, of the nature of
the tissues; the patterns and the sculpture of the shell depending on
its manner of growth. The amount of light seems to be influential to a
certain extent; for although, as repeatedly stated by Mr. Gwyn
Jeffreys, the shells of some species living at a profound depth are
brightly coloured, yet we generally see the lower surfaces, as well as
the parts covered by the mantle, less highly-coloured than the upper
and exposed surfaces. (4. I have given (‘Geological Observations on
Volcanic Islands,’ 1844, p. 53) a curious instance of the influence of
light on the colours of a frondescent incrustation, deposited by the
surf on the coast-rocks of Ascension and formed by the solution of
triturated sea-shells.) In some cases, as with shells living amongst
corals or brightly-tinted seaweeds, the bright colours may serve as a
protection. (5. Dr. Morse has lately discussed this subject in his
paper on the ‘Adaptive Coloration of Mollusca,’ ‘Proc. Boston Soc. of
Nat. Hist.’ vol. xiv. April 1871.) But that many of the nudibranch
Mollusca, or sea-slugs, are as beautifully coloured as any shells, may
be seen in Messrs. Alder and Hancock’s magnificent work; and from
information kindly given me by Mr. Hancock, it seems extremely doubtful
whether these colours usually serve as a protection. With some species
this may be the case, as with one kind which lives on the green leaves
of algae, and is itself bright-green. But many brightly-coloured,
white, or otherwise conspicuous species, do not seek concealment;
whilst again some equally conspicuous species, as well as other
dull-coloured kinds live under stones and in dark recesses. So that
with these nudibranch molluscs, colour apparently does not stand in any
close relation to the nature of the places which they inhabit.
These naked sea-slugs are hermaphrodites, yet they pair together, as do
land-snails, many of which have extremely pretty shells. It is
conceivable that two hermaphrodites, attracted by each other’s greater
beauty, might unite and leave offspring which would inherit their
parents’ greater beauty. But with such lowly-organised creatures this
is extremely improbable. Nor is it at all obvious how the offspring
from the more beautiful pairs of hermaphrodites would have any
advantage over the offspring of the less beautiful, so as to increase
in number, unless indeed vigour and beauty generally coincided. We have
not here the case of a number of males becoming mature before the
females, with the more beautiful males selected by the more vigorous
females. If, indeed, brilliant colours were beneficial to a
hermaphrodite animal in relation to its general habits of life, the
more brightly-tinted individuals would succeed best and would increase
in number; but this would be a case of natural and not of sexual
selection.
SUB-KINGDOM OF THE VERMES: CLASS, ANNELIDA (OR SEA-WORMS).
In this class, although the sexes, when separate, sometimes differ from
each other in characters of such importance that they have been placed
under distinct genera or even families, yet the differences do not seem
of the kind which can be safely attributed to sexual selection. These
animals are often beautifully coloured, but as the sexes do not differ
in this respect, we are but little concerned with them. Even the
Nemertians, though so lowly organised, “vie in beauty and variety of
colouring with any other group in the invertebrate series”; yet Dr.
McIntosh (6. See his beautiful monograph on ‘British Annelids,’ part i.
1873, p. 3.) cannot discover that these colours are of any service. The
sedentary annelids become duller-coloured, according to M. Quatrefages
(7. See M. Perrier: ‘L’Origine de l’Homme d’après Darwin,’ ‘Revue
Scientifique’, Feb. 1873, p. 866.), after the period of reproduction;
and this I presume may be attributed to their less vigorous condition
at that time. All these worm-like animals apparently stand too low in
the scale for the individuals of either sex to exert any choice in
selecting a partner, or for the individuals of the same sex to struggle
together in rivalry.
SUB-KINGDOM OF THE ARTHROPODA: CLASS, CRUSTACEA.
In this great class we first meet with undoubted secondary sexual
characters, often developed in a remarkable manner. Unfortunately the
habits of crustaceans are very imperfectly known, and we cannot explain
the uses of many structures peculiar to one sex. With the lower
parasitic species the males are of small size, and they alone are
furnished with perfect swimming-legs, antennae and sense-organs; the
females being destitute of these organs, with their bodies often
consisting of a mere distorted mass. But these extraordinary
differences between the two sexes are no doubt related to their widely
different habits of life, and consequently do not concern us. In
various crustaceans, belonging to distinct families, the anterior
antennae are furnished with peculiar thread-like bodies, which are
believed to act as smelling-organs, and these are much more numerous in
the males than in the females. As the males, without any unusual
development of their olfactory organs, would almost certainly be able
sooner or later to find the females, the increased number of the
smelling-threads has probably been acquired through sexual selection,
by the better provided males having been the more successful in finding
partners and in producing offspring. Fritz Müller has described a
remarkable dimorphic species of Tanais, in which the male is
represented by two distinct forms, which never graduate into each
other. In the one form the male is furnished with more numerous
smelling-threads, and in the other form with more powerful and more
elongated chelae or pincers, which serve to hold the female. Fritz
Müller suggests that these differences between the two male forms of
the same species may have originated in certain individuals having
varied in the number of the smelling-threads, whilst other individuals
varied in the shape and size of their chelae; so that of the former,
those which were best able to find the female, and of the latter, those
which were best able to hold her, have left the greatest number of
progeny to inherit their respective advantages. (8. ‘Facts and
Arguments for Darwin,’ English translat., 1869, p. 20. See the previous
discussion on the olfactory threads. Sars has described a somewhat
analogous case (as quoted in ‘Nature,’ 1870, p. 455) in a Norwegian
crustacean, the Pontoporeia affinis.)
[Fig.4. Labidocera Darwinii (from Lubbock). Labelled are: a. Part of
right anterior antenna of male, forming a prehensile organ. b.
Posterior pair of thoracic legs of male. c. Ditto of female.]
In some of the lower crustaceans, the right anterior antenna of the
male differs greatly in structure from the left, the latter resembling
in its simple tapering joints the antennae of the female. In the male
the modified antenna is either swollen in the middle or angularly bent,
or converted (Fig. 4) into an elegant, and sometimes wonderfully
complex, prehensile organ. (9. See Sir J. Lubbock in ‘Annals and Mag.
of Nat. Hist.’ vol. xi. 1853, pl. i. and x.; and vol. xii. (1853), pl.
vii. See also Lubbock in ‘Transactions, Entomological Society,’ vol.
iv. new series, 1856-1858, p. 8. With respect to the zigzagged antennae
mentioned below, see Fritz Müller, ‘Facts and Arguments for Darwin,’
1869, p. 40, foot-note.) It serves, as I hear from Sir J. Lubbock, to
hold the female, and for this same purpose one of the two posterior
legs (b) on the same side of the body is converted into a forceps. In
another family the inferior or posterior antennae are “curiously
zigzagged” in the males alone.
[Fig. 5. Anterior part of body of Callianassa (from Milne-Edwards),
showing the unequal and differently-constructed right and left-hand
chelae of the male. N.B.—The artist by mistake has reversed the
drawing, and made the left-hand chela the largest.
Fig. 6. Second leg of male Orchestia Tucuratinga (from Fritz Müller).
Fig. 7. Ditto of female.]
In the higher crustaceans the anterior legs are developed into chelae
or pincers; and these are generally larger in the male than in the
female,—so much so that the market value of the male edible crab
(Cancer pagurus), according to Mr. C. Spence Bate, is five times as
great as that of the female. In many species the chelae are of unequal
size on the opposite side of the body, the right-hand one being, as I
am informed by Mr. Bate, generally, though not invariably, the largest.
This inequality is also often much greater in the male than in the
female. The two chelae of the male often differ in structure (Figs. 5,
6, and 7), the smaller one resembling that of the female. What
advantage is gained by their inequality in size on the opposite sides
of the body, and by the inequality being much greater in the male than
in the female; and why, when they are of equal size, both are often
much larger in the male than in the female, is not known. As I hear
from Mr. Bate, the chelae are sometimes of such length and size that
they cannot possibly be used for carrying food to the mouth. In the
males of certain fresh-water prawns (Palaemon) the right leg is
actually longer than the whole body. (10. See a paper by Mr. C. Spence
Bate, with figures, in ‘Proceedings, Zoological Society,’ 1868, p. 363;
and on the nomenclature of the genus, ibid. p. 585. I am greatly
indebted to Mr. Spence Bate for nearly all the above statements with
respect to the chelae of the higher crustaceans.) The great size of the
one leg with its chelae may aid the male in fighting with his rivals;
but this will not account for their inequality in the female on the
opposite sides of the body. In Gelasimus, according to a statement
quoted by Milne Edwards (11. ‘Hist. Nat. des Crust.’ tom. ii. 1837, p.
50.), the male and the female live in the same burrow, and this shews
that they pair; the male closes the mouth of the burrow with one of its
chelae, which is enormously developed; so that here it indirectly
serves as a means of defence. Their main use, however, is probably to
seize and to secure the female, and this in some instances, as with
Gammarus, is known to be the case. The male of the hermit or soldier
crab (Pagurus) for weeks together, carries about the shell inhabited by
the female. (12. Mr. C. Spence Bate, ‘British Association, Fourth
Report on the Fauna of S. Devon.’) The sexes, however, of the common
shore-crab (Carcinus maenas), as Mr. Bate informs me, unite directly
after the female has moulted her hard shell, when she is so soft that
she would be injured if seized by the strong pincers of the male; but
as she is caught and carried about by the male before moulting, she
could then be seized with impunity.
[Fig.8. Orchestia Darwinii (from Fritz Müller), showing the
differently-constructed chelae of the two male forms.]
Fritz Müller states that certain species of Melita are distinguished
from all other amphipods by the females having “the coxal lamellae of
the penultimate pair of feet produced into hook-like processes, of
which the males lay hold with the hands of the first pair.” The
development of these hook-like processes has probably followed from
those females which were the most securely held during the act of
reproduction, having left the largest number of offspring. Another
Brazilian amphipod (see Orchestia darwinii, Fig. 8) presents a case of
dimorphism, like that of Tanais; for there are two male forms, which
differ in the structure of their chelae. (13. Fritz Müller, ‘Facts and
Arguments for Darwin,’ 1869, pp. 25-28.) As either chela would
certainly suffice to hold the female,—for both are now used for this
purpose,—the two male forms probably originated by some having varied
in one manner and some in another; both forms having derived certain
special, but nearly equal advantages, from their differently shaped
organs.
It is not known that male crustaceans fight together for the possession
of the females, but it is probably the case; for with most animals when
the male is larger than the female, he seems to owe his greater size to
his ancestors having fought with other males during many generations.
In most of the orders, especially in the highest or the Brachyura, the
male is larger than the female; the parasitic genera, however, in which
the sexes follow different habits of life, and most of the Entomostraca
must be excepted. The chelae of many crustaceans are weapons well
adapted for fighting. Thus when a Devil-crab (Portunus puber) was seen
by a son of Mr. Bate fighting with a Carcinus maenas, the latter was
soon thrown on its back, and had every limb torn from its body. When
several males of a Brazilian Gelasimus, a species furnished with
immense pincers, were placed together in a glass vessel by Fritz
Müller, they mutilated and killed one another. Mr. Bate put a large
male Carcinus maenas into a pan of water, inhabited by a female which
was paired with a smaller male; but the latter was soon dispossessed.
Mr. Bate adds, “if they fought, the victory was a bloodless one, for I
saw no wounds.” This same naturalist separated a male sand-skipper (so
common on our sea-shores), Gammarus marinus, from its female, both of
whom were imprisoned in the same vessel with many individuals of the
same species. The female, when thus divorced, soon joined the others.
After a time the male was put again into the same vessel; and he then,
after swimming about for a time, dashed into the crowd, and without any
fighting at once took away his wife. This fact shews that in the
Amphipoda, an order low in the scale, the males and females recognise
each other, and are mutually attached.
The mental powers of the Crustacea are probably higher than at first
sight appears probable. Any one who tries to catch one of the
shore-crabs, so common on tropical coasts, will perceive how wary and
alert they are. There is a large crab (Birgus latro), found on coral
islands, which makes a thick bed of the picked fibres of the cocoa-nut,
at the bottom of a deep burrow. It feeds on the fallen fruit of this
tree by tearing off the husk, fibre by fibre; and it always begins at
that end where the three eye-like depressions are situated. It then
breaks through one of these eyes by hammering with its heavy front
pincers, and turning round, extracts the albuminous core with its
narrow posterior pincers. But these actions are probably instinctive,
so that they would be performed as well by a young animal as by an old
one. The following case, however, can hardly be so considered: a
trustworthy naturalist, Mr. Gardner (14. ‘Travels in the Interior of
Brazil,’ 1846, p. 111. I have given, in my ‘Journal of Researches,’ p.
463, an account of the habits of the Birgus.), whilst watching a
shore-crab (Gelasimus) making its burrow, threw some shells towards the
hole. One rolled in, and three other shells remained within a few
inches of the mouth. In about five minutes the crab brought out the
shell which had fallen in, and carried it away to a distance of a foot;
it then saw the three other shells lying near, and evidently thinking
that they might likewise roll in, carried them to the spot where it had
laid the first. It would, I think, be difficult to distinguish this act
from one performed by man by the aid of reason.
Mr. Bate does not know of any well-marked case of difference of colour
in the two sexes of our British crustaceans, in which respect the sexes
of the higher animals so often differ. In some cases, however, the
males and females differ slightly in tint, but Mr. Bate thinks not more
than may be accounted for by their different habits of life, such as by
the male wandering more about, and being thus more exposed to the
light. Dr. Power tried to distinguish by colour the sexes of the
several species which inhabit the Mauritius, but failed, except with
one species of Squilla, probably S. stylifera, the male of which is
described as being “of a beautiful bluish-green,” with some of the
appendages cherry-red, whilst the female is clouded with brown and
grey, “with the red about her much less vivid than in the male.” (15.
Mr. Ch. Fraser, in ‘Proc. Zoolog. Soc.’ 1869, p. 3. I am indebted to
Mr. Bate for Dr. Power’s statement.) In this case, we may suspect the
agency of sexual selection. From M. Bert’s observations on Daphnia,
when placed in a vessel illuminated by a prism, we have reason to
believe that even the lowest crustaceans can distinguish colours. With
Saphirina (an oceanic genus of Entomostraca), the males are furnished
with minute shields or cell-like bodies, which exhibit beautiful
changing colours; these are absent in the females, and in both sexes of
one species. (16. Claus, ‘Die freilebenden Copepoden,’ 1863, s. 35.) It
would, however, be extremely rash to conclude that these curious organs
serve to attract the females. I am informed by Fritz Müller, that in
the female of a Brazilian species of Gelasimus, the whole body is of a
nearly uniform greyish-brown. In the male the posterior part of the
cephalo-thorax is pure white, with the anterior part of a rich green,
shading into dark brown; and it is remarkable that these colours are
liable to change in the course of a few minutes—the white becoming
dirty grey or even black, the green “losing much of its brilliancy.” It
deserves especial notice that the males do not acquire their bright
colours until they become mature. They appear to be much more numerous
than the females; they differ also in the larger size of their chelae.
In some species of the genus, probably in all, the sexes pair and
inhabit the same burrow. They are also, as we have seen, highly
intelligent animals. From these various considerations it seems
probable that the male in this species has become gaily ornamented in
order to attract or excite the female.
It has just been stated that the male Gelasimus does not acquire his
conspicuous colours until mature and nearly ready to breed. This seems
a general rule in the whole class in respect to the many remarkable
structural differences between the sexes. We shall hereafter find the
same law prevailing throughout the great sub-kingdom of the Vertebrata;
and in all cases it is eminently distinctive of characters which have
been acquired through sexual selection. Fritz Müller (17. ‘Facts and
Arguments,’ etc., p. 79.) gives some striking instances of this law;
thus the male sand-hopper (Orchestia) does not, until nearly full
grown, acquire his large claspers, which are very differently
constructed from those of the female; whilst young, his claspers
resemble those of the female.
CLASS, ARACHNIDA (SPIDERS).
The sexes do not generally differ much in colour, but the males are
often darker than the females, as may be seen in Mr. Blackwall’s
magnificent work. (18. ‘A History of the Spiders of Great Britain,’
1861-64. For the following facts, see pp. 77, 88, 102.) In some
species, however, the difference is conspicuous: thus the female of
Sparassus smaragdulus is dullish green, whilst the adult male has the
abdomen of a fine yellow, with three longitudinal stripes of rich red.
In certain species of Thomisus the sexes closely resemble each other,
in others they differ much; and analogous cases occur in many other
genera. It is often difficult to say which of the two sexes departs
most from the ordinary coloration of the genus to which the species
belong; but Mr. Blackwall thinks that, as a general rule, it is the
male; and Canestrini (19. This author has recently published a valuable
essay on the ‘Caratteri sessuali secondarii degli Arachnidi,’ in the
‘Atti della Soc. Veneto-Trentina di Sc. Nat. Padova,’ vol. i. Fasc. 3,
1873.) remarks that in certain genera the males can be specifically
distinguished with ease, but the females with great difficulty. I am
informed by Mr. Blackwall that the sexes whilst young usually resemble
each other; and both often undergo great changes in colour during their
successive moults, before arriving at maturity. In other cases the male
alone appears to change colour. Thus the male of the above
bright-coloured Sparassus at first resembles the female, and acquires
his peculiar tints only when nearly adult. Spiders are possessed of
acute senses, and exhibit much intelligence; as is well known, the
females often shew the strongest affection for their eggs, which they
carry about enveloped in a silken web. The males search eagerly for the
females, and have been seen by Canestrini and others to fight for
possession of them. This same author says that the union of the two
sexes has been observed in about twenty species; and he asserts
positively that the female rejects some of the males who court her,
threatens them with open mandibles, and at last after long hesitation
accepts the chosen one. From these several considerations, we may admit
with some confidence that the well-marked differences in colour between
the sexes of certain species are the results of sexual selection;
though we have not here the best kind of evidence,—the display by the
male of his ornaments. From the extreme variability of colour in the
male of some species, for instance of Theridion lineatum, it would
appear that these sexual characters of the males have not as yet become
well fixed. Canestrini draws the same conclusion from the fact that the
males of certain species present two forms, differing from each other
in the size and length of their jaws; and this reminds us of the above
cases of dimorphic crustaceans.
The male is generally much smaller than the female, sometimes to an
extraordinary degree (20. Aug. Vinson (‘Araneides des Iles de la
Reunion,’ pl. vi. figs. 1 and 2) gives a good instance of the small
size of the male, in Epeira nigra. In this species, as I may add, the
male is testaceous and the female black with legs banded with red.
Other even more striking cases of inequality in size between the sexes
have been recorded (‘Quarterly Journal of Science,’ July 1868, p. 429);
but I have not seen the original accounts.), and he is forced to be
extremely cautious in making his advances, as the female often carries
her coyness to a dangerous pitch. De Geer saw a male that “in the midst
of his preparatory caresses was seized by the object of his attentions,
enveloped by her in a web and then devoured, a sight which, as he adds,
filled him with horror and indignation.” (21. Kirby and Spence,
‘Introduction to Entomology,’ vol. i. 1818, p. 280.) The Rev. O.P.
Cambridge (22. ‘Proceedings, Zoological Society,’ 1871, p. 621.)
accounts in the following manner for the extreme smallness of the male
in the genus Nephila. “M. Vinson gives a graphic account of the agile
way in which the diminutive male escapes from the ferocity of the
female, by gliding about and playing hide and seek over her body and
along her gigantic limbs: in such a pursuit it is evident that the
chances of escape would be in favour of the smallest males, while the
larger ones would fall early victims; thus gradually a diminutive race
of males would be selected, until at last they would dwindle to the
smallest possible size compatible with the exercise of their generative
functions,—in fact, probably to the size we now see them, i.e., so
small as to be a sort of parasite upon the female, and either beneath
her notice, or too agile and too small for her to catch without great
difficulty.”
Westring has made the interesting discovery that the males of several
species of Theridion (23. Theridion (Asagena, Sund.) serratipes,
4-punctatum et guttatum; see Westring, in Kroyer, ‘Naturhist.
Tidskrift,’ vol. iv. 1842-1843, p. 349; and vol. ii. 1846-1849, p. 342.
See, also, for other species, ‘Araneae Suecicae,’ p. 184.) have the
power of making a stridulating sound, whilst the females are mute. The
apparatus consists of a serrated ridge at the base of the abdomen,
against which the hard hinder part of the thorax is rubbed; and of this
structure not a trace can be detected in the females. It deserves
notice that several writers, including the well-known arachnologist
Walckenaer, have declared that spiders are attracted by music. (24. Dr.
H.H. van Zouteveen, in his Dutch translation of this work (vol. i. p.
444), has collected several cases.) From the analogy of the Orthoptera
and Homoptera, to be described in the next chapter, we may feel almost
sure that the stridulation serves, as Westring also believes, to call
or to excite the female; and this is the first case known to me in the
ascending scale of the animal kingdom of sounds emitted for this
purpose. (25. Hilgendorf, however, has lately called attention to an
analogous structure in some of the higher crustaceans, which seems
adapted to produce sound; see ‘Zoological Record,’ 1869, p. 603.)
CLASS, MYRIAPODA.
In neither of the two orders in this class, the millipedes and
centipedes, can I find any well-marked instances of such sexual
differences as more particularly concern us. In Glomeris limbata,
however, and perhaps in some few other species, the males differ
slightly in colour from the females; but this Glomeris is a highly
variable species. In the males of the Diplopoda, the legs belonging
either to one of the anterior or of the posterior segments of the body
are modified into prehensile hooks which serve to secure the female. In
some species of Iulus the tarsi of the male are furnished with
membranous suckers for the same purpose. As we shall see when we treat
of Insects, it is a much more unusual circumstance, that it is the
female in Lithobius, which is furnished with prehensile appendages at
the extremity of her body for holding the male. (26. Walckenaer et P.
Gervais, ‘Hist. Nat. des Insectes: Apteres,’ tom. iv. 1847, pp. 17, 19,
68.)