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Chapter IX 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 limacons, ne saurait mettre en doute la seduction deployee dans
les mouvements et les allures qui preparent et accomplissent le double
embrassement de ces hermaphrodites." (2. 'De l'Espece 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'apres 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 Muller 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 Muller 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 Muller, '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 Muller).
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 Muller), showing the differently-
constructed chelae of the two male forms.]
Fritz Muller 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
Muller, '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 Muller, 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 Muller, 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 Muller (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.)
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