Principles of Corporate Finance 12th

Edition Brealey Solutions Manual

Go to download the full and correct content document:
https://testbankfan.com/product/principles-of-corporate-finance-12th-edition-brealey-s
olutions-manual/
More products digital (pdf, epub, mobi) instant
download maybe you interests ...

Principles of Corporate Finance 12th Edition Brealey

Test Bank

https://testbankfan.com/product/principles-of-corporate-
finance-12th-edition-brealey-test-bank/

Principles of Corporate Finance 11th Edition Brealey

Solutions Manual

https://testbankfan.com/product/principles-of-corporate-
finance-11th-edition-brealey-solutions-manual/

Principles of Corporate Finance 10th Edition Brealey

Solutions Manual

https://testbankfan.com/product/principles-of-corporate-
finance-10th-edition-brealey-solutions-manual/

Principles of Corporate Finance 11th Edition Brealey

Test Bank

https://testbankfan.com/product/principles-of-corporate-
finance-11th-edition-brealey-test-bank/
Principles of Corporate Finance 10th Edition Brealey
Test Bank

https://testbankfan.com/product/principles-of-corporate-
finance-10th-edition-brealey-test-bank/

Principles of Corporate Finance Concise 2nd Edition

Brealey Test Bank

https://testbankfan.com/product/principles-of-corporate-finance-
concise-2nd-edition-brealey-test-bank/

Fundamentals of Corporate Finance 7th Edition Brealey

Solutions Manual

https://testbankfan.com/product/fundamentals-of-corporate-
finance-7th-edition-brealey-solutions-manual/

Fundamentals of Corporate Finance 9th Edition Brealey

Solutions Manual

https://testbankfan.com/product/fundamentals-of-corporate-
finance-9th-edition-brealey-solutions-manual/

Fundamentals of Corporate Finance 8th Edition Brealey

Solutions Manual

https://testbankfan.com/product/fundamentals-of-corporate-
finance-8th-edition-brealey-solutions-manual/
Chapter 07 - Introduction to Risk and Return

CHAPTER 7

Introduction to Risk and Return

The values shown in the solutions may be rounded for display purposes. However, the answers were
derived using a spreadsheet without any intermediate rounding.

Answers to Problem Sets

1. Expected payoff = (.10 × $500) + (.50 × $100) + (.40 × $0)

= $100

Rates of return:

($500 – 100) / $100 = 400%

($100 – 100) / $100 = 0%
($0 – 100) / $100 = –100%

Expected rate of return = (.10 × 400%) + (.50 × 0%) + (.40 × –100%)

Expected rate of return = 0%

Variance = .10(400% – 0)2 + .50(0% – 0)2 + .40(–100% – 0)2

Variance = 20,000

Standard deviation = 20,000.5

Standard deviation = 141.42%

Est. Time: 01 – 05

2. a. Average nominal return = (.172 + .010 + .161 + .331 + .127) / 5

Average nominal return = .1602, or 16.02%

Variance = [(.172 – .1602)2 + (.010 – .1602)2 + (.161 – .1602)2 + (.331 –

.1602)2 + (.127 – .1602)2] / 5
Variance = .010595

Standard deviation = .010595.5

Standard deviation = .1029, or 10.29%

b. Average real return = {[(1.172 / 1.015) – 1] + [(1.010 / 1.030) – 1] + [(1.161

/ 1.017) – 1] + [(1.331 / 1.015) – 1] + [(1.127 /
1.008) – 1]} / 5
Average real return = .1412, or 14.12%

Est. Time: 01 – 05

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

3. Ms. Sauros:

Average return = [.249 + (–.009) + .186 + .421 + .152] / 5

Average return = .1998, or 19.98%

Variance = [(.249 – .1998)2 + (–.009 – .1998)2 + (.186 – .1998)2 + (.421 –

.1998)2 + (.152 – .1998)2] / 5
Variance = .019485

Standard deviation = .0194

Standard deviation = .1396, or 13.96%

S&P 500:

Average return = (.172 + .010 + .161 + .331 +.127) / 5

Average return = .1602, or 16.02%

Variance = [(.172 – .1602)2 + (.010 – .1602)2 + (.161 – .1602)2 + (.331 –

.1602)2 + (.127 – .1602)2
Variance = .010595

Standard deviation = .010595.5

Standard deviation = .1029, or 10.29%

Est. Time: 01 – 05

4. a. False. Investors prefer diversified portfolios because diversification

reduces variability and therefore reduces risk. However, the diversification
of an individual company does not necessarily make it less risky.

b. True. Stocks must be less than perfectly positively correlated in order to

obtain diversification benefits.

c. False. The risk eliminated by diversification is called specific risk, or the

risk surrounding an individual company or industry. Market risk will still
exist in a fully diversified portfolio.

d. False. It is true that the greatest benefit to diversification occurs when

stocks are uncorrelated. However, most stocks tend to move in the same
direction. There are still benefits to diversification any time stocks are less
than perfectly positively correlated.

e. False. The contribution to portfolio risk depends on the relationship of the

stock to the market as a whole.

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

f. True. Market risk is the risk that relates to a diversified portfolio.

g. True. The only risk inherent in a diversified portfolio is market risk.

h. False. An undiversified portfolio with a beta of 2 is twice as risky as the

market portfolio. Part of that risk will be market risk and part will be
specific risk.

Est. Time: 01 – 05

5. Perfect negative correlation does the most to reduce risks because the stocks
always move in opposite directions. When one goes up, the other goes down,
and vice versa.

Est. Time: 01 – 05

6.

Est. Time: 01 – 05

7. a. σp = 1.3 × 20% = 26%

b. σp = 0 × 20% = 0%

c. βp = 15% / 20% = .75

d. βp = Less than 1

βp = 20% / 20% = 1, This would be the beta if the portfolio were

diversified. Since the portfolio is non-diversified, the beta must be less
than 1 because part of the portfolio’s risk is specific, or unique, risk.

Est. Time: 01 – 05

8. βp = {(5 × 1.2) + [(10 – 5) × 1.4)]} / 10

βp = 1.3

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

Beta measures systematic risk which cannot be eliminated by diversification.

Est. Time: 01 – 05

9. The beta of each stock is given by the slope of the line, or the rise divided by the
run. The run is the range of the market returns while the rise is the range of the
stock returns.

BetaA = (0 – 20) / (–10 – 10) = 1

BetaB = (–20 – 20) / (–10 – 10) = 2

BetaC = (–30 – 0) / (–10 – 10) = 1.5

BetaD = (15 – 15) / (–10 – 10) = 0

BetaE = (10 – (–10) / (–10 – 10) = –1

Est. Time: 01 – 05

10. a. r = [(1 + R) / (1 + h)] – 1

r1929 = {[1 + (–.145)] / (1 + .002)} – 1 = –.1467, or –14.67%

r1930 = {[1 + (–.283)] / [1 + (–.060)]} – 1 = –.2372, or –23.72%
r1931 = {[1 + (–.439)] / [1 + (–.095)]} – 1 = –.3801, or –38.01%
r1932 = {[1 + (–.099)] / [1 + (–.103)]} – 1 = .0045 or .45%
r1933 = [(1 + .573) / (1 + .005)] – 1 = .5652, or 56.52%

b. Average real return = [–.1433 + (–.2372) + (–.3801) + .0045 + .5652] / 5

Average real return = –.0382, or –3.82%

c. Risk premium1929 = –.145 – .048 = –.1930, or –19.30%

Risk premium1930 = –.283 – .024 = –.3070, or –30.70%
Risk premium1931 = –.439 – .011 = –.4500, or –45.00%
Risk premium1932 = –.099 – .010 = –.1090, or –10.90%
Risk premium1933 = .573 – .003 = .5700, or 57.00%

d. Average risk premium = [–.1930 + (–.3070) + (–.4500) + (–.1090) + .5700]

/5
Average risk premium = –0978, or –9.78%

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

e. σRisk premium = {[–.1930 – (–.0978)]2 + [–.3070 – (–.0978)]2 + [–.4500 – (–

.0978)]2 + [–.1090 – (–.0978)]2 + [.5700 – (–.0978)]2]} / 5

σRisk premium = .1246, or 12.46%

Est. Time: 06 – 10

11. a. A long-term United States government bond is always considered to be

safe in terms of the dollars received. However, the price of the bond
fluctuates as interest rates change, and the rate at which coupon
payments received can be invested also changes as interest rates
change. And, of course, the payments are all in nominal dollars, so
inflation risk must also be considered.

b. It is true that stocks offer higher long-run rates of return than do bonds, but
it is also true that stocks have a higher standard deviation of return. So,
which investment is preferable depends on the amount of risk one is
willing to tolerate. This is a complicated issue and depends on numerous
factors, one of which is the investment time horizon. If the investor has a
short time horizon, then stocks are generally not preferred.

c. Unfortunately, 10 years is not generally considered a sufficient amount of

time for estimating average rates of return. Thus, using either a 5- or 10-
year average is likely to be misleading.

Est. Time: 06 – 10

12. The risk to Hippique shareholders depends on the market risk, or beta, of the
investment in the black stallion. The information given in the problem suggests
that the horse has very high unique risk, but we have no information regarding
the horse’s market risk. So, the best estimate is that this horse has a market risk
about equal to that of other racehorses, and thus this investment is not a
particularly risky one for Hippique shareholders.

Est. Time: 01 – 05

13. In the context of a well-diversified portfolio, the only risk characteristic of a single
security that matters is the security’s contribution to the overall portfolio risk. This
contribution is measured by beta. Lonesome Gulch is the safer investment for a
diversified investor because its beta of .10 is lower than the beta of Amalgamated
Copper of .66. For a diversified investor, the standard deviations are irrelevant.

Est. Time: 01 – 05

14. a. σP2 = .602 × .102 + .402 × .202 + 2(.60 × .40 × 1 × .10 × .20)

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

σP2 = .0196

b. σP2 = .602 × .102 + .402 × .202 + 2(.60 × .40 × .50 × .10 × .20)
σP2 = .0148

c. σP2 = .602 × .102 + .402 × .202 + 2(.60 × .40 × 0 × .10 × .20)

σP2= .0100

Est. Time: 06 – 10

15. a. Refer to Figure 7.13 in the text. With 100 securities, the box is 100 by 100.
The variance terms are the diagonal terms, and thus there are 100
variance terms. The rest are the covariance terms. Because the box has
(100 × 100) terms altogether, the number of covariance terms is:
Number of covariance terms = 1002 – 100 = 9,900
Half of these terms (i.e., 4,950) are different.

b. Once again, it is easiest to think of this in terms of Figure 7.13. With 50

stocks, all with the same standard deviation (.30), the same weight in the
portfolio (.02), and all pairs having the same correlation coefficient (.40),
the portfolio variance is:

σ2 = 50(.02)2(.30)2 + [(50)2 – 50](.02)2(.40)(.30)2 = .03708

σ = .193, or 19.3%

c. For a fully diversified portfolio, portfolio variance equals the average

covariance:

σ2 = (.30)(.30)(.40) = .036
σ = .190, or 19.0%

Est. Time: 06 – 10

16. a. Refer to Figure 7.13 in the text. For each different portfolio, the relative
weight of each share is (1 / number of shares (n) in the portfolio), the
standard deviation of each share is .40, and the correlation between pairs
is .30. Thus, for each portfolio, the diagonal terms are the same, and the
off-diagonal terms are the same. There are n diagonal terms and (n2 – n)
off-diagonal terms. In general, we have:

Variance = n(1 / n)2(.4)2 + (n2 – n)(1 / n)2(.3)(.4)(.4)

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

For one share:

Variance = 1(1)2(.4)2 + 0 = .160000

For two shares:

Variance = 2(.5)2(.4)2 + 2(.5)2(.3)(.4)(.4) = .104000

The results are summarized in the second and third columns of the
table in part (c) below.

b. The underlying market risk that cannot be diversified away is the second
term in the formula for variance above:

Underlying market risk = (n2 – n)(1 / n)2(.3)(.4)(.4)

As n increases, [(n2 – n)(1 / n)2] = [(n – 1) / n] becomes close to 1, so that

the underlying market risk is: [(.3)(.4)(.4)] = .048.

c. This is the same as Part (a), except that all of the off-diagonal terms are
now equal to zero. The results are summarized in the fourth and fifth
columns of the table below.
(Part a) (Part a) (Part c) (Part c)
No. of Standard Standard
Shares Variance Deviation Variance Deviation
1 .160000 .400 .160000 .400
2 .104000 .322 .080000 .283
3 .085333 .292 .053333 .231
4 .076000 .276 .040000 .200
5 .070400 .265 .032000 .179
6 .066667 .258 .026667 .163
7 .064000 .253 .022857 .151
8 .062000 .249 .020000 .141
9 .060444 .246 .017778 .133
10 .059200 .243 .016000 .126

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

Graphs for Part (a):

Portfolio Variance Portfolio Standard Deviation

0.2 0.5

Standard D eviation
0.4
0.15
Variance

0.3
0.1
0.2
0.05
0.1

0 0
0 2 4 6 8 10 12 0 2 4 6 8 10 12
N umber of Securities Number of Securities

Graphs for Part (c):

Portfolio Variance Portfolio Standard Deviation

0.2 Standard D eviation 0.5

0.4
0.15
Variance

0.3
0.1
0.2
0.05
0.1

0 0
0 2 4 6 8 10 12 0 2 4 6 8 10 12
N umber of Securities Number of Securities

Est. Time: 16 – 20

17. The table below uses the format of Figure 7.13 in the text in order to calculate the
portfolio variance. The portfolio variance is the sum of all the entries in the
matrix. Portfolio variance is .03178.

Korea
BHP BP Fiat Heineken Electric Nestlé Sony Tata
BHP .0006126 .0003781 .0005062 .0000893 .0002841 -.0000090 .0002636 .0006050
BP .0003781 .0013231 .0007832 .0002051 .0003290 .0000529 .0008359 .0005157
Fiat .0005062 .0007832 .0028971 .0002063 .0003558 -.0000653 .0013274 .0008420
Heineken .0000893 .0002051 .0002063 .0005085 .0001334 .0001203 .0004677 .0002866
Korea Electric .0002841 .0003290 .0003558 .0001334 .0012102 .0000042 .0003120 .0002211
Nestlé -.0000090 .0000529 -.0000653 .0001203 .0000042 .0001470 .0001563 .0000474
Sony .0002636 .0008359 .0013274 .0004677 .0003120 .0001563 .0031416 .0005206
Tata Motors .0006050 .0005157 .0008420 .0002866 .0002211 .0000474 .0005206 .0023900

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
 Chapter 07 - Introduction to Risk and Return

Est. Time: 21 – 25

18. “Safest” means lowest risk; in a portfolio context, this means lowest variance of
return. Half of the portfolio is invested in British Petroleum stock (BP), and half of
the portfolio must be invested in one of the other securities listed. Thus, we
calculate the portfolio variance for seven different portfolios to see which is the
lowest (see table below). The safest attainable portfolio is comprised of British
Petroleum stock (BP) and Nestlé.

Company with BP: Variance Variance

BHP = .52 × .19802 + .52 × .29102 + 2 × .5 × .5 × .42 × .1980 × .2910 = .04307

Fiat Chrysler = .52 × .43062 + .52 × .29102 + 2 × .5 × .5 × .40 × .4306 × .2910 = .09259

Heineken = .52 × .18042 + .52 × .29102 + 2 × .5 × .5 × .25 × .1804 × .2910 = .03587

Korea Electric = .52 × .27832 + .52 × .29102 + 2 × .5 × .5 × .26 × .2783 × .2910 = .05106

Nestlé = .52 × .09702 + .52 × .29102 + 2 × .5 × .5 × .12 × .0970 × .2910 = .02522

Sony = .52 × .44842 + .52 × .29102 + 2 × .5 × .5 × .41 × .4484 × .2910 = .09819

Tata Motors = .52 × .39112 + .52 × .29102 + 2 × .5 × .5 × .29 × 39.11 × .2910 = .07591

Est. Time: 11 – 15

19. a-1. Change in stock’s rate of return = .05 × –.25 = –.0125, or –1.25%

a-2. Change in stock’s rate of return = –.05 × –.25 = .0125, or 1.25%

b. “Safest” implies lowest risk. Assuming the well-diversified portfolio is

invested in typical securities, the portfolio beta is approximately one. The
largest reduction in beta is achieved by investing the $20,000 in a stock
with the negative beta.

Est. Time: 06 – 10

20. E(r)P = .12 = .1wa + .15(1 – wa); wa = .60; (1 – wa) = .40

σp = (.602 × .202 + .402 × .402 + 2 × .60 × .40 × .50 × .20 × .40).5

σp = .2433, or 24.33%

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

Est. Time: 06 – 10

21. a. In general:

σP = (x12σ12 + x22σ22 + 2x1x2ρ12σ1σ2).5

Thus:

σP = (.52 × .35802 + .52 × .21002 + 2 × .5 × .5 × .30 × .3580 × .2100).5

σP = .2331, or 23.31%

b. We can think of this in terms of Figure 7.13 in the text, with three
securities. One of these securities, T-bills, has zero risk and, hence, zero
standard deviation. Thus:

σP = [(1/3)2 × .35802 + (1/3)2 × .21002 + 2 × (1/3) × (1/3) × .30 × .3580 ×

.2100].5
σP = .1554, or 15.54%

Another way to think of this portfolio is that it is comprised of one-third

T-Bills and two-thirds a portfolio which is half Bank of America and half
Starbucks. Because the risk of T-bills is zero, the portfolio standard
deviation is two-thirds of the standard deviation computed in Part (a)
above:
σP = (2/3) × 23.31% = 15.54%

c. With 50% margin, the investor invests twice as much money in the
portfolio as he had to begin with. Thus, the risk is twice that found in Part
(a) when the investor is investing only his own money:

σP = 2 × 23.31% = 46.62%

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

d. With 100 stocks, the portfolio is well diversified, and hence the portfolio
standard deviation depends almost entirely on the average covariance of
the securities in the portfolio (measured by beta) and on the standard
deviation of the market portfolio. Thus, for a portfolio made up of 100
stocks each with Bank of America’s beta of 1.57, the portfolio standard
deviation is approximately:

σP = 1.57 × 23.0% = 36.11%

For stocks like Starbucks, the approximate standard deviation is:

σP = .83 × 23.0% = 19.09%

Est. Time: 11 – 15

22. For a two-security portfolio, the formula for portfolio risk is:

σP2= x12σ12 + x22σ22 + 2x1x2ρρ12σ1σ2

If security one is Treasury bills and security two is the market portfolio, then σ1 is
zero, and σ2 is 20%. Therefore:

σP2 = x22σ22 = x22 × .202

σP = .20x2

Portfolio expected return = x1(.06) + x2(.06 + .085)

Portfolio expected return = .06x1 + .145x2

Expected Standard
Portfolio x1 x2
Return Deviation
1 1.0 .0 .060 .000
2 .8 .2 .077 .040
3 .6 .4 .094 .080
4 .4 .6 .111 .120
5 .2 .8 .128 .160
6 .0 1.0 .145 .200

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
 Chapter 07 - Introduction to Risk and Return

Portfolio Risk & Return

.16
.14
.12
Expected Return

.10
.08
.06
.04
.02
.00
.00 .05 .10 .15 .20 .25
Standard Deviation

Est. Time: 11 – 15

23. The matrix below displays the variance for each of the eight stocks along the
diagonal and each of the covariances in the off-diagonal cells:

Variances/Covariances BHP BP Fiat Heineken Korea Nestlè Sony Tata σs,mkt

BHP 0.0392040 0.0241996 0.0323983 0.0057151 0.0181841 -0.0005762 0.0168688 0.0387189 0.0218391
BP 0.0241996 0.0846810 0.0501218 0.0131241 0.0210562 0.0033872 0.0534986 0.0330049 0.0353842
Fiat 0.0323983 0.0501218 0.1854164 0.0132056 0.0227688 -0.0041768 0.0849557 0.0538905 0.0548225
Heineken 0.0057151 0.0131241 0.0132056 0.0325442 0.0085349 0.0076995 0.0299298 0.0183442 0.0161372
Korea 0.0181841 0.0210562 0.0227688 0.0085349 0.0774509 0.0002700 0.0199664 0.0141496 0.0227976
Nestlè -0.0005762 0.0033872 -0.0041768 0.0076995 0.0002700 0.0094090 0.0100038 0.0030349 0.0036314
Sony 0.0168688 0.0534986 0.0849557 0.0299298 0.0199664 0.0100038 0.2010626 0.0333202 0.0562007
Tata 0.0387189 0.0330049 0.0538905 0.0183442 0.0141496 0.0030349 0.0333202 0.1529592 0.0434278

The covariance of BP with the market portfolio (σBP, Market) is the mean of the eight
respective covariances between BP and each of the eight stocks in the portfolio.
(The covariance of BP with itself is the variance of BP.) Therefore, σBP, Market is
equal to the average of the eight covariances in the second row or, equivalently,
the average of the eight covariances in the second column. Beta for BP is equal
to the covariance divided by the market variance, which we calculated at 0.03178
in problem 17. The covariances and betas are displayed in the table below:

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Chapter 07 - Introduction to Risk and Return

Market
Covariance Variance Beta
BHP 0.0218391 0.0317801 0.6871943
BP 0.0353842 0.0317801 1.1134082
Fiat 0.0548225 0.0317801 1.7250607
Heineken 0.0161372 0.0317801 0.5077763
Korea 0.0227976 0.0317801 0.7173556
Nestlè 0.0036314 0.0317801 0.1142674
Sony 0.0562007 0.0317801 1.7684269
Tata 0.0434278 0.0317801 1.3665106

Est. Time: 21 – 25

Copyright © 2017 McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.
Another random document with
no related content on Scribd:
 (p. 126).
Idotheidae (p. 127).
Valvifera (p. 127)
Arcturidae (p. 127).
Asellidae (p. 128).
Asellota (p. 127) Munnopsidae
(p. 128).
Oniscoida (p. 128).
Microniscidae
(p. 130).
Cryptoniscidae
(p. 130).
Liriopsidae (p. 130).
Hemioniscidae
Cryptoniscina (pp. 129, 130) (p. 130).
Cabiropsidae
(p. 130).
Epicarida
Podasconidae
(p. 129)
(p. 130).
Asconiscidae
(p. 130).
Dajidae (p. 130).
Phryxidae (p. 130).
Bopyrina (pp. 129, 130, 132) Bopyridae (pp. 130,
133).
Entoniscidae
(pp. 130, 134).
Phreatoicidae
Phreatoicidea (p. 136)
(p. 136)
Lysianassidae
(p. 137).
Haustoriidae (p. 137).
Crevettina (p. 137) Gammaridae
Amphipoda (p. 138).
(p. 136) Talitridae (p. 139).
Corophiidae (p. 139).
Caprellidae (p. 139).
Laemodipoda (p. 139)
Cyamidae (p. 140).
Hyperina (p. 140).
Hoplocarida
Stomatopoda (p. 141) Squillidae (p. 143).
(p. 141)
Eucarida Euphausiidae
Euphausiacea (p. 144)
(p. 144) (p. 144).
Decapoda Macrura Nephropsidae
(p. 152) (p. 153) (p. 154).
Nephropsidea (p. 154)
Astacidae (p. 157).
Parastacidae (p. 157).
Eryonidea (p. 157) Eryonidae (p. 158).
Peneidae (p. 162).
Sergestidae (p. 162).
Peneidea (pp. 158, 162)
Stenopodidae
(p. 162).
Caridea (pp. 158, 163) Pasiphaeidae
(p. 163).
Acanthephyridae
(p. 163).
Atyidae (p. 163).
Alpheidae (p. 163).
 Psalidopodidae
(p.164).
Pandalidae (p. 164).
Hippolytidae
(p. 164).
Palaemonidae
(p. 164).
Glyphocrangonidae
(p. 164).
Crangonidae (p. 164).
Palinuridae (p. 167).
Loricata (p. 165)
Scyllaridae (p. 167).
Callianassidae
Thalassinidea (p. 167)
(p. 167).
Aegleidae (p. 169).
Galatheidae (p. 169).
Galatheidea (p. 168)
Porcellanidae
(p. 170).
Albuneidae (p. 171).
Hippidea (p. 170)
Hippidae (p. 171).
Pylochelidae (p. 180).
Anomura Paguridae (p. 180).
(p. 167) Eupagurinae
(p. 180).
Pagurinae (p. 180).
Paguridea (p. 171)
Coenobitidae (p. 181).
Lithodidae (p. 181).
Hapalogastorinae
(p. 181).
Lithodinae (p. 181).
Brachyura Dromiidae (p. 184).
(p. 181) Dynomenidae
Dromiacea (p. 183)
(p. 184).
Homolidae (p. 184).
Calappidae (p. 187).
Leucosiidae (p. 188).
Oxystomata (p. 185)
Dorippidae (p. 188).
Raninidae (p. 188).
Corystidae (p. 190).
Atelecyclidae
(p. 190).
Cancridae (p. 191).
Cyclometopa (p. 188) Portunidae (p. 191).
Xanthidae (p. 191).
Thelphusidae =
Potamonidae
(p. 191).
Maiidae (p. 193).
Parthenopidae
Oxyrhyncha (p. 191) (p. 193).
Hymenosomatidae
(p. 193).
Catometopa (p. 193) Carcinoplacidae
(p. 195).
Gonoplacidae
(p. 195).
Pinnotheridae
(p. 195).
 Grapsidae (p. 196).
Gecarcinidae
(p. 196).
Ocypodidae (p. 196).

TRILOBITA (p. 221).

Families
Agnostidae (p. 244).
Shumardiidae (p. 245).
Trinucleidae (p. 245).
Harpedidae (p. 245).
Paradoxidae (p. 246).
Conocephalidae = Conocoryphidae (p. 247).
Olenidae (p. 247).
Calymenidae (p. 247).
Asaphidae (p. 249).
Bronteidae (p. 249).
Phacopidae (p. 249).
Cheiruridae (p. 250).
Proëtidae (p. 251).
Encrinuridae (p. 251).
Acidaspidae (p. 251).
Lichadidae (p. 252).
 ARACHNIDA (p. 255).

DELOBRANCHIATA = MEROSTOMATA (pp. 258, 259).

Orders. Sub-Orders. Families. Sub-Families.
Xiphosurinae
Xiphosuridae (p. 276).
Xiphosura (pp. 258, 259, 276)
(p. 276) Tachypleinae
(p. 276).
Eurypterida = Gigantostraca (pp. 258, 283) Eurypteridae (p. 290).

EMBOLOBRANCHIATA (pp. 258, 297).

Buthinae (p. 306).

Buthidae (p. 306)
Centrurinae (p. 306).
Diplocentrinae
(p. 307).
Urodacinae (p. 307).
Scorpionidae Scorpioninae
(p. 306) (p. 307).
Hemiscorpioninae
(p. 307).
Scorpionidea (pp. 258, 297) Ischnurinae (p. 307).
Chaerilidae (p. 307).
Megacorminae
(p. 308).
Chactidae (p. 307) Euscorpiinae
(p. 308).
Chactinae (p. 308).
Vejovidae (p. 308).
Bothriuridae (p. 308).
Thelyphonidae (p. 312).
Schizonotidae = Tartaridae (p. 312).
Pedipalpi (pp. 258, 308) Tarantulidae = Tarantulinae (p. 313).
Phrynidae Phrynichinae (p. 313).
(p. 312) Charontinae (p. 313).
Araneae (pp. 258, 314) Liphistiidae (p. 386).
Paratropidinae
(p. 387).
Actinopodinae
(p. 387).
Aviculariidae = Miginae (p. 387).
Mygalidae
Ctenizinae (p. 388).
(p. 386)
Barychelinae (p. 389).
Aviculariinae
(p. 389).
Diplurinae (p. 390).
Atypidae (p. 390).
Filistatidae (p. 391).
Oecobiidae = Urocteidae (p. 392).
Sicariidae = Scytodidae (p. 393).
Hypochilidae (p. 393).
Leptonetidae (p. 393).
Oonopidae (p. 393).
Hadrotarsidae (p. 394).
Dysderinae (p. 394).
Dysderidae (p. 394)
Segestriinae (p. 395).
Caponiidae (p. 395).
Prodidomidae (p. 395).
Drassinae (p. 396).
Clubioninae (p. 397).
Drassidae (p. 396)
Liocraninae (p. 397).
Micariinae (p. 397).
Palpimanidae (p. 398).
Eresidae (p. 398).
Dictynidae (p. 398).
Psechridae (p. 399).
Zodariidae = Enyoidae (p. 399).
Hersiliidae (p. 400).
Pholcidae (p. 401).
Argyrodinae (p. 402).
Episininae (p. 402).
Theridioninae
(p. 403).
Theridiidae (p. 401) Phoroncidiinae
(p. 404).
Erigoninae (p. 404).
Formicinae (p. 405).
Linyphiinae (p. 405).
Theridiosomatinae
(p. 407).
Tetragnathinae
(p. 407).
Argiopinae (p. 408).
Epeiridae (p. 406) Nephilinae (p. 408).
Epeirinae (p. 408).
Gasteracanthinae
(p. 409).
Poltyinae (p. 410).
Arcyinae (p. 410).
Dinopinae (p. 410).
Uloborinae (p. 410).
Uloboridae (p. 410)
Miagrammopinae
(p. 411).
Archeidae (p. 411).
Mimetidae (p. 411).
Thomisinae =
Misumeninae
(p. 412).
Philodrominae
(p. 413).
Thomisidae (p. 412) Sparassinae (p. 414).
Aphantochilinae
(p. 414).
Stephanopsinae
(p. 414).
Selenopinae (p. 414).
Zoropsidae (p. 415).
Platoridae (p. 415).
Cybaeinae (p. 415).
Ageleninae (p. 416).
Agelenidae (p. 415)
Hahniinae (p. 416).
Nicodaminae (p. 416).
 Pisauridae (p.416).
Lycosidae (p. 417).
Ctenidae (p. 418).
Senoculidae (p. 418).
Oxyopidae (p. 419).
Attidae = Salticidae (p. 419).
Palpigradi (pp. 258, 422).
Galeodidae (p. 428).
Rhagodinae (p. 429).
Solpuginae (p. 429).
Daesiinae (p. 429).
Solifugae = Solpugae (pp. 258, 423) Solpugidae (p. 429)
Eremobatinae
(p. 429).
Karshiinae (p. 429).
Hexisopodidae (p. 429).
Cheliferidae (p. 436)
Garypinae (pp. 436,
Chernetidea = Chernetes = Cheliferinae
437).
Pseudoscorpiones (pp. 258, 430) (p. 436).
Obisiinae (pp. 436,
437).
Podogona = Ricinulei (pp. 258, 439) Cryptosteinmatidae (p.440).
Cyphophthalmi
Sironidae (p. 448).
(p. 447)
Mecostethi = Phalangodidae (p. 448).
Laniatores Cosmetidae (p. 449).
(p. 448) Gonyleptidae (p. 449).
Phalangidea = Opiliones (pp. 258,
440) Sclerosomatinac
Phalangiidae (p.449).
Plagiostethi = (p. 449)
Phalangiinae (p. 450).
Palpatores
Ischyropsalidae (p. 451).
(p. 449)
Nemastomatidae (p. 451).
Trogulidae (p. 452).
Acarina = Acari = Acaridea (pp. 258, Vermiformia Eriophyidae = Phytoptidae (p. 464).
454) (p. 464) Demodicidae (p. 465).
Sarcoptinae (p. 466).
Analgesinae (p. 466).
Astigmata (p. 465) Sarcoptidae (p. 466)
Tyroglyphinae
(p. 466).
Oribatidae (p. 467).
Argasidae (p. 469).
Ixodoidea (p. 468)
Metastigmata Ixodidae (p. 469).
(p. 467) Gamasinae (p. 470).
Gamasidae (p. 470) Dermanyssinae
(p. 471).
Heterostigmata
Tarsonemidae (p. 471).
(p. 471)
Prostigmata (p. 471) Bdellidae (p. 471).
Halacaridae (p. 472).
Hydrachnidae (p. 472).
Trombidiidae Limnocharinae
(p. 472) (p. 472).
Caeculinae (p. 472).
Tetranychinae
(p. 472).
Cheyletinae (p. 473).
Erythraeinae (p. 473).
 Trombidiinae
(p. 473).
Notostigmata
Opilioacaridae (p. 473).
(p. 473)

Orders.

TARDIGRADA (pp. 258, 477).

PENTASTOMIDA (pp. 258, 488).

PYCNOGONIDA = PODOSOMATA = PANTOPODA (p. 501).

Families.
Decolopodidae (p. 531).
Colossendeidae = Pasithoidae (p. 532).
Eurycididae = Ascorhynchidae (p. 533).
Ammotheidae (p. 534).
Rhynchothoracidae (p. 535)
Nymphonidae (p. 536).
Pallenidae (p. 537).
Phoxichilidiidae (p. 538).
Phoxichilidae (p. 539).
Pycnogonidae (p. 539).
 CRUSTACEA

CHAPTERS I and III-VII

BY

GEOFFREY SMITH, M.A. (Oxon.)

Fellow of New College, Oxford

CHAPTER II
BY

The Late W. F. R. WELDON, M.A. (D.Sc. Oxon.)

Formerly Fellow of St. John’s College, Cambridge, and Linacre Professor of Human and
Comparative Anatomy, Oxford
 CHAPTER I
CRUSTACEA—GENERAL ORGANISATION

The Crustacea are almost exclusively aquatic animals, and they play a
part in the waters of the world closely parallel to that which insects play
on land. The majority are free-living, and gain their sustenance either as
vegetable-feeders or by preying upon other animals, but a great number
are scavengers, picking clean the carcasses and refuse that litter the
ocean, just as maggots and other insects rid the land of its dead cumber.
Similar to insects also is the great abundance of individuals which
represent many of the species, especially in the colder seas, and the
naturalist in the Arctic or Antarctic oceans has learnt to hang the
carcasses of bears and seals over the side of the boat for a few days in
order to have them picked absolutely clean by shoals of small Amphipods.
It is said that these creatures, when crowded sufficiently, will even attack
living fishes, and by sheer press of numbers impede their escape and
devour them alive. Equally surprising are the shoals of minute Copepods
which may discolour the ocean for many miles, an appearance well known
to fishermen, who take profitable toll of the fishes that follow in their
wake. Despite this massing together we look in vain for any elaborate
social economy, or for the development of complex instincts among
Crustacea, such as excite our admiration in many insects, and though
many a crab or lobster is sufficiently uncanny in appearance to suggest
unearthly wisdom, he keeps his intelligence rigidly to himself, encased in
the impenetrable reserve of his armour and vindicated by the most
powerful of pincers. It is chiefly in the variety of structure and in the
multifarious phases of life-history that the interest of the Crustacea lies.
Before entering into an examination of these matters, it will be well to
take a general survey of Crustacean organisation, to consider the plan on
which these animals are built, and the probable relation of this plan to
others met with in the animal kingdom.
The Crustacea, to begin with, are a Class of the enormous Phylum
Arthropoda, animals with metamerically segmented bodies and usually
with externally jointed limbs. Their bodies are thus composed of a series
of repeated segments, which are on the whole similar to one another,
though particular segments may be differentiated in various respects for
the performance of different functions. This segmentation is apparent
externally, the surface of a Crustacean being divided typically into a
number of hard chitinous rings, some of which may be fused rigidly
together, as in the carapace of the crabs, or else articulated loosely.
 Each segment bears typically a pair of jointed limbs, and though they
vary greatly in accordance with the special functions for which they are
employed, and may even be absent from certain segments, they may yet
be reduced to a common plan and were, no doubt, originally present on
all the segments.
Passing from the exterior to the interior of the body we find, generally
speaking, that the chief system of organs which exhibits a similar
repetition, or metameric segmentation, is the nervous system. This
system is composed ideally of a nervous ganglion situated in each
segment and giving off peripheral nerves, the several ganglia being
connected together by a longitudinal cord. This ideal arrangement,
though apparent during the embryonic development, becomes obscured
to some extent in the adult owing to the concentration or fusion of ganglia
in various parts of the body. The other internal organs do not show any
clear signs of segmentation, either in the embryo or in the adult; the
alimentary canal and its various diverticula lie in an unsegmented body-
cavity, and are bathed in the blood which courses through a system of
narrow canals and irregular spaces which surround all the organs of the
body. A single pair, or at most two pairs of kidneys are present.
The type of segmentation exhibited by the Crustacea is thus of a limited
character, concerning merely the external skin with its appendages, and
the nervous system, and not touching any of the other internal organs.[1]
In this respect the Crustacea agree with all the other Arthropods, in the
adults of which the segmentation is confined to the exterior and to the
nervous system, and does not extend to the body-cavity and its contained
organs; and for the same reason they differ essentially from all other
metamerically segmented animals, e.g. Annelids, in which the
segmentation not only affects the exterior and the nervous system, but
especially applies to the body-cavity, the musculature, the renal, and often
the generative organs. The Crustacea also resemble the other Arthropoda
in the fact that the body-cavity contains blood, and is therefore a
“haemocoel,” while in the Annelids and Vertebrates the segmented body-
cavity is distinct from the vascular system, and constitutes a true
“coelom.” To this important distinction, and to its especial application to
the Crustacea, we will return, but first we may consider more narrowly
the segmentation of the Crustacea and its main types of variation
within the group. In order to determine the number of segments which
compose any particular Crustacean we have clearly two criteria: first, the
rings or somites of which the body is composed, and to each of which a
pair of limbs must be originally ascribed; and, second, the nervous
ganglia.
 Around and behind the region of the mouth there is very little difficulty
in determining the segments of the body, if we allow embryology to assist
anatomy, but in front of the mouth the matter is not so easy.
In the Crustacea the moot point is whether we consider the paired eyes
and first pair of antennae as true appendages belonging to two true
segments, or whether they are structures sui generis, not homologous to
the other limbs. With regard to the first antennae we are probably safe in
assigning them to a true body-segment, since in some of the
Entomostraca, e.g. Apus, the nerves which supply them spring, not from
the brain as in more highly specialised forms, but from the commissures
which pass round the oesophagus to connect the dorsally lying brain to
the ventral nerve-cord. The paired eyes are always innervated from the
brain, but the brain, or at least part of it, is very probably formed of
paired trunk-ganglia which have fused into a common cerebral mass; and
the fact that under certain circumstances the stalked eye of Decapods
when excised with its peripheral ganglion[2] can regenerate in the form of
an antenna, is perhaps evidence that the lateral eyes are borne on what
were once a pair of true appendages.
Now, with regard to the segmentation of the body, the Crustacea fall
into three categories: the Entomostraca, in which the number of segments
is indefinite; the Malacostraca, in which we may count nineteen
segments, exclusive of the terminal piece or telson and omitting the
lateral eyes; and the Leptostraca, including the single recent genus
Nebalia, in which the segmentation of head and thorax agrees exactly
with that of the Malacostraca, but in the abdomen there are two
additional segments.
It has been usually held that the indefinite number of segments
characteristic of the Entomostraca, and especially the indefinitely large
number of segments characteristic of such Phyllopods as Apus, preserves
the ancestral condition from which the definite number found in the
Malacostraca has been derived; but recently it has been clearly pointed
out by Professor Carpenter[3] that the number of segments found in the
Malacostraca and Leptostraca corresponds with extraordinary exactitude
to the number determined as typical in all the other orders of Arthropoda.
This remarkable correspondence (it can hardly be coincidence) seems to
point to a common Arthropodan plan of segmentation, lying at the very
root of the phyletic tree; and if this is so, we are forced to the conclusion
that the Malacostraca have retained the primitive type of segmentation in
far greater perfection than the Entomostraca, in some of which many
segments have been added, e.g. Phyllopoda, while in others segments
have been suppressed, e.g. Cladocera, Ostracoda. It may be objected to
this view of the primitive condition of segmentation in the Crustacea that
the Trilobites, which for various reasons are regarded as related to the
ancestral Crustaceans, exhibit an indefinite and often very high number
of segments; but, as Professor Carpenter has pointed out, the oldest and
most primitive of Trilobites, such as Olenellus, possessed few segments
which increase as we pass from Cambrian to Carboniferous genera.
The following table shows the segmentation of the body in the
Malacostraca, as compared with that of Limulus (cf. p. 263), Insecta, the
primitive Myriapod Scolopendrella, and Peripatus. It will be seen that the
correspondence, though not exact, is very close, especially in the first four
columns, the number of segments in Peripatus being very variable in the
different species.
 Table showing the Segmentation of various Arthropods
Malacostraca. Limulus. Insecta. Myriapoda. Peripatus.
(Scolopendrella).
1 Eyes Median Eyes
eyes
2 1st antennae Rostrum Antennae Feelers Feelers
3 2nd antennae Chelicerae Intercalary
segment
4 Mandibles Pedipalpi Mandibles Mandibles Mandibles
5 1st maxillae 1st walking Maxillulae Maxillulae 1st jaw-claw
legs
6 2nd maxillae 2nd „ „ 1st 1st maxillae 2nd jaw-
maxillae claw
7 1st maxillipede 3rd „ „ 2nd 2nd maxillae 1st leg
maxillae
8 2nd maxillipede 4th „ „ 1st leg 1st leg 2nd „
9 3rd maxillipede Chilaria 2nd „ 2nd „ 3rd „
10 1st ambulatory Genital 3rd „ 3rd „ 4th „
operculum
11 2nd „ 1st gill- 1st 4th „ 5th „
book abdominal
12 3rd „ 2nd „ 2nd „ 6th „ 6th „
13 4th „ 3rd „ 3rd „ 6th „ 7th „
14 5th „ 4th „ 4th „ 7th „ 8th „
15 1st abdominal 5th „ 5th „ 8th „ 9th „
16 2nd „ No 6th „ 9th „ 10th „
appendages
17 3rd „ „ 7th „ 10th „ 11th „
18 4th „ „ 8th „ 11th „ 12th „
19 5th „ „ 9th „ 12th „ 13th „
20 6th „ „ 10th „ Reduced limbs 14th „
21 [4] „ Cercopods [5]

Telson Telson Telson Telson Telson

The appendages of the Crustacea exhibit a wonderful variety of
structure, but these variations can be reduced to at most two, and
possibly to one fundamental plan. In a typical Crustacean, besides the
paired eyes, which may be borne on stalks, possibly homologous to highly
modified limbs, there are present, first, two pairs of rod-like or
filamentous antennae, which in the adult are usually specialised for
sensory purposes, but frequently retain their primitive function as
locomotory limbs even in the adult, e.g. Ostracoda; while in the Nauplius
larva, found in almost all the chief subdivisions of the Crustacea, the two
pairs of antennae invariably aid in locomotion, and the base of the second
antennae is usually furnished with sharp biting spines which assist
mastication. Following the antennae is a pair of mandibles which are
fashioned for biting the food or for piercing the prey, and posterior to
these are two pairs of maxillae, biting organs more slightly built than the
mandibles, whose function it is to lacerate the food and prepare it for the
more drastic action of the mandibles. So far, with comparatively few
exceptions, the order of specialisation is invariable; but behind the
maxillae the trunk-appendages vary greatly both in structure and function
in the different groups.
As a general rule, the first or first few thoracic limbs are turned
forwards toward the mouth, and are subsidiary to mastication; they are
then called maxillipedes; this happens usually in the Malacostraca, but to
a much less extent in the Entomostraca; and in any case these
appendages immediately behind the maxillae never depart to any great
extent from a limb-like structure, and they may graduate insensibly into
the ordinary trunk-appendages. The latter show great diversity in the
different Crustacean groups, according as the animals lead a natatory,
creeping, or parasitic method of life; they may be foliaceous, as in the
Branchiopoda, or biramous, as in the swimming thoracic and abdominal
appendages of the Mysidae, or simply uniramous, as in the walking legs of
the higher Decapoda, and the clinging legs of various parasitic forms.
Without going into detailed deviations of structure, many of which will
be described under the headings of special groups, it is clear from the
foregoing description and from Fig. 1 (p. 10), that three main types of
appendage can be distinguished: first, the foliaceous or multiramous;
second, the biramous; and, third, the uniramous.
We may dismiss the uniramous type with a few words: it is obviously
secondarily derived from the biramous type; this can be proved in detail
in nearly every case. Thus, the uniramous second antennae of some adult
forms are during the Nauplius stage invariably biramous, a condition
which is retained in the adult Cladocera. Similarly the uniramous walking
legs of many Decapoda pass through a biramous stage during
development, the outer branches or exopodites of the limbs being
suppressed subsequently, while the primitively biramous condition of the
thoracic limbs is retained in the adults of the Schizopoda, which doubtless
own a common ancestry with the Decapoda. The only Crustacean limb
which appears to be constantly uniramous both in larval and adult life is
the first pair of antennae.
We are reduced, therefore, to two types—the foliaceous and biramous.
Sir E. Ray Lankester,[6] in one of his most incisive morphological essays,
has explained how these two types are really fundamentally the same. He
compares, for instance, the foliaceous first maxillipede (Fig. 1, A), or the
second maxilla (Fig. 1, B) of a Decapod, e.g. Astacus, with the foliaceous
thoracic limb of Branchipus (Fig. 1, D), and with the typically biramous
first maxillipede of a Schizopod (Fig. 1, F).
In each case there is present, on the outer edge of the limb, one or more
projections or epipodites which are generally specialised for respiratory
purposes, and may carry the gills. The 6th and 5th “endites” in the
foliaceous limb (Fig. 1, D) are compared with the exopodite and
endopodite respectively of the biramous limb, while the endites 4–1 of the
foliaceous limb are found in the basal joints of the biramous limb.
Lankester presumes that the biramous type of limb throughout has been
derived from the foliaceous type by the suppression of the endites 1–4, as
discrete rami, and the exaggerated development of the endites 5 and 6, as
above indicated.
 Fig. 1.—Appendages of Crustacea (A-G) and
Trilobita (H). A, First maxillipede of
Astacus; B, second maxilla of Astacus; C,
second walking leg of Astacus; D, thoracic
limb of Branchipus; E, first maxillipede of
Mysis; F, first maxillipede of
Gnathophausia; G, thoracic limb of
Nebalia; H, thoracic limb of Triarthrus. bp,
basipodite; br, bract; cp, carpopodite; cxp,
coxopodite; cx.s, coxopoditic setae; dp,
dactylopodite; end, endopodite; ep,
epipodite; ex, exopodite; ip, ischiopodite;
mp, meropodite; pp, propodite; 1–6, the six
endites.

The essential fact that the two types of limb are built on the same plan
may be considered as established; but it may be urged that the biramous
type represents this common plan more nearly than the foliaceous. It is,
at any rate, certain that in the maxillipedes of the Decapoda we witness
the conversion of the biramous type into the foliaceous by the expansion
of the basal joints concomitantly with the assumption by the maxillipedes
of masticatory functions. Thus in the Decapoda the first maxillipede is
decidedly foliaceous owing to the expanded “gnathobases” (Fig. 1, A, bp,
cxp), and the second maxillipedes are flattened, with their basal joints
somewhat expanded and furnished with biting hairs; but in the
“Schizopoda” (e.g. Mysis) the first maxillipede is a typical biramous limb,
though the expanded gnathobases in some forms are beginning to project
(Fig. 1, E), while the limb following, which corresponds to the second
maxillipede of Decapods, is simply a biramous swimming leg. Besides this
obvious conversion of a biramous into a foliaceous limb, further evidence
of the fundamental character of the biramous type is found, first, in its
invariable occurrence in the Nauplius stage, which does not necessarily
mean that the ancestors of the Crustacea possessed this type of limb in
the adult, but which does imply that this type of limb was possessed at
some period of life by the common ancestral Crustacean; and, second, the
limbs of the Trilobita, a group which probably stands near the origin of
the Crustacea, have been shown by Beecher to conform to the biramous
type (Fig. 1, H). Furthermore, the thoracic limbs of Nebalia, an animal
which combines many of the characteristics of Entomostraca and
Malacostraca, and is therefore considered as a primitive type, despite
their flattened character, are really built upon a biramous plan (Fig. 1, G).
In conclusion, we may point out that this view of the Crustacean limb,
as essentially a biramous structure, agrees with the conclusion derived
from our consideration of the segmentation of the body, and points less to
the Branchiopoda as primitive Crustacea and more to some generalised
Malacostracan type.
So far we have shortly dealt with those systems of organs which are
clearly affected by the metameric segmentation of the body; we must now
expose the condition of the body-cavity to a similar scrutiny. If we
remove the external integument of a Crustacean, we find that the internal
organs do not lie in a spacious and discrete body-cavity, as is the case in
the Annelids and Vertebrates, but that they are packed together in an
irregular system of spaces (“haemocoel”) in communication with the
vascular system and containing blood. In the Entomostraca and smaller
forms generally, a definite vascular system hardly exists, though a central
heart and artery may serve to propel the blood through the irregular
lacunae of the body-cavity; but in the larger Malacostraca a complicated
system of arteries may be present which pour the blood into fairly
definitely arranged spaces surrounding the chief organs. These spaces
return the blood to the pericardium, and so to the heart again through the
apertures or ostia which pierce its walls.
This condition of the body-cavity or haemocoel is reproduced in the
adults of all Arthropods, but in some of them by following the
development we can trace the steps by which the true coelom is replaced
by the haemocoel. In the embryos of all Arthropods except the Crustacea,
a true closed metamerically segmented coelom is formed as a split in the
mesodermal embryonic layer of cells, distinct from the vascular system.
During the course of development the segmented coelomic spaces and
their walls give rise to the reproductive organs and to certain renal organs
in Peripatus, Myriapoda, and Arachnida (nephridia and coxal glands),
but the general body-cavity is formed as an extension of the vascular
system, which is laid down outside the coelom by a canaliculisation of the
extra-coelomic mesoderm. In the embryos of the Crustacea, however,
there is never at any time a closed segmented coelom, and in this respect
the Crustacea differ from all other Arthropods. The only clear instance in
which metamerically repeated mesodermal cavities have been seen in the
embryo Crustacean is that of Astacus; here Reichenbach[7] states that in
the abdomen segmental cavities are formed which subsequently break
down; but even in this instance no connexion has been shown to subsist
between these embryonic cavities and the reproductive and excretory
organs of the adult.
Since the connexion between the coelom and the excretory organs is
always a very close one throughout the animal kingdom, interest naturally
centres upon the renal organs in Crustacea, and it has been suggested
that these organs in Crustacea represent the sole remains, with the
possible exception of the gonads, of the coelom. Since, at any rate, a part
of the kidneys appears to be developed as a closed sac in the mesoderm,
and since they possess a possible segmental value, this suggestion is
plausible; but, on the other hand, since there are never more than two
pairs of kidneys, and since they are totally unconnected with the gonads
or with any other indication of a segmented coelom, the suggestion
remains purely hypothetical.
The renal organs of the Crustacea, excluding the Malpighian tubes
present in some Amphipods which open into the alimentary canal, and
resemble the Malpighian tubes of Insects, consist of two pairs—the
antennary gland, opening at the base of the second antenna, and the
maxillary gland, opening on the second maxilla. These two pairs of glands
rarely subsist together in the adult condition, though this is said to be the
case in Nebalia and possibly Mysis; the antennary glands are
characteristic of adult Malacostraca[8] and the larvae of the Entomostraca,
while the maxillary glands (“shell-glands”) are present in adult
Entomostraca and larval Malacostraca, that is to say, the one pair
replaces the other in the two great subdivisions of the Crustacea. The
shell-gland of the Entomostraca is a simple structure consisting of a
coiled tube opening to the exterior on the external branch of the second
maxilla, and ending blindly in a dilated vesicle, the end-sac. The
antennary gland of the Malacostraca is usually more complicated: these
complications have been studied especially by Weldon,[9] Allen, and
Marchal[10] in the Decapoda. In a number of forms we have a tube opening
to the exterior at the base of the second antenna, and expanding within to
form a spacious bladder into which the coiled tubular part of the kidney
opens, while at the extremity of this coiled portion is the vesicle called the
end-sac. This arrangement may be modified; thus in Palaemon Weldon
described the two glands as fusing together above and below the
oesophagus, the dorsal commissure expanding into a huge sac stretching
dorsally down the length of the body. This closed sac with excretory
functions thus comes to resemble a coelomic cavity, and the view that it is
really coelomic has indeed been upheld.
A modified form of this view is that of Vejdovský, who describes a
funnel-apparatus leading from the coiled tube into the end-sac of the
antennary gland of Amphipods; he regards the end-sac alone as
representing the coelom, while the funnel and coiled tube represent the
kidney opening into it.
Not very much is known of the development of these various structures.
Some authors have considered that both antennary and maxillary glands
are developed in the embryo from ectodermal inpushings, but the more
recent observations of Waite[11] on Homarus americanus indicate that the
antennary gland at any rate is a composite structure, formed by an
ectodermal ingrowth which meets a mesodermal strand, and from the
latter are produced the end-sac and perhaps the tubular excretory
portions of the gland with their derivatives.
With regard to the possible metameric repetition of the renal organs, it
is of interest to note that by feeding Mysis and Nebalia on carmine,
excretory glands of a simple character were observed by Metschnikoff
situated at the bases of the thoracic limbs.
The alimentary canal of the Crustacea is a straight tube composed of
three parts—a mid-gut derived from the endoderm of the embryo, and a
fore- and hind-gut formed by ectodermal invaginations in the embryo
which push into and fuse with the endodermal canal. The regions of the
fore- and hind-gut can be recognised in the adult by the fact of their being
lined with the chitinous investment which is continued over the external
surface of the body forming the hard exoskeleton, while the mid-gut is
naked. The chitinous lining of fore- and hind-gut is shed whenever the
animal moults. In the Malacostraca, in which a complicated “gastric mill”
may be present, the chitinous lining of this part of the gut is thrown into
ridges bearing teeth, and this stomach in the crabs and lobsters reaches a
high degree of complication and materially assists the mastication of the
food. The gut is furnished with a number of secretory and metabolic
glands; the so-called liver, which is probably a hepatopancreas, opening
into the anterior end of the mid-gut, is directed forwards in most
Entomostraca and backwards in the Malacostraca, in the Decapoda
developing into a complicated branching organ which fills a large part of
the thorax. In the Decapoda peculiar vermiform caeca of doubtful
function are present, a pair of which open into the gut anteriorly where
fore-passes into mid-gut, and a single asymmetrically placed caecum
opens posteriorly into the alimentary tract where mid- passes into hind-
gut.
The disposition of these caeca, marking as they do the morphological
position of fore-, mid-, and hind-gut, is of peculiar interest owing to the
variations exhibited. From some unpublished drawings of Mr. E. H.
Schuster, which he kindly lent me, it appears that in certain Decapods,
e.g. Callianassa subterranea, the length of the mid-gut between the
anterior and posterior caeca is very long; in Carcinus maenas it is
considerable; in Maia squinado it is greatly reduced, the caeca being
closely approximated; while in Galathea strigosa the caeca are greatly
reduced, and the mid-gut as a separate entity has almost disappeared.
The relation of these variations to the habits of the different crabs and to
their modes of development is unknown.
The reproductive organs usually make their appearance as a small
paired group of mesodermal cells in the thorax comparatively late in life;
and neither in their early development nor in the adult condition do they
show any clear signs of segmentation or any connexion with a coelomic
cavity. The sexes are usually separate, but hermaphroditism occurs
sporadically in many forms, and as a normal condition in some parasitic
groups (see pp. 105–107). The adult gonads are generally simple paired
tubes, from the walls of which the germ-cells are produced, and as these
grow and come to maturity they fill up the cavities of the tubes; special
nutrient cells are rarely differentiated, though in some cases (e.g.
Cladocera) a few ova nourish themselves by devouring their sister-cells
(see p. 44). The oviducts and vasa deferentia are formed as simple
outgrowths from the gonadial tubes, which acquire an opening to the
exterior; they are usually poorly supplied with accessory glands, the
epithelium of the canals often supplying albuminous secretions for
cementing the eggs together, while the lining of the vasa deferentia may
be instrumental in the formation of spermatophores for transferring large
packets of spermatozoa to the female. In the vast majority of Crustacea
copulation takes place, the male passing spermatophores or free
spermatozoa into special receptacles (spermathecae), or into the oviducts
of the female. The spermatophores are hollow chitinous structures in