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Studies in Systems, Decision and Control 182
Irina V. Gashenko
Yulia S. Zima
Armenak V. Davidyan Editors
Optimization of the
Taxation System:
Preconditions,
Tendencies, and
Perspectives
Studies in Systems, Decision and Control
Volume 182
Series editor
Janusz Kacprzyk, Polish Academy of Sciences, Warsaw, Poland
e-mail: kacprzyk@ibspan.waw.pl
The series “Studies in Systems, Decision and Control” (SSDC) covers both new
developments and advances, as well as the state of the art, in the various areas of
broadly perceived systems, decision making and control–quickly, up to date and
with a high quality. The intent is to cover the theory, applications, and perspectives
on the state of the art and future developments relevant to systems, decision
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engineering, computer science, physics, economics, social and life sciences, as well
as the paradigms and methodologies behind them. The series contains monographs,
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control spanning the areas of Cyber-Physical Systems, Autonomous Systems,
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Systems, Automation, Manufacturing, Smart Grids, Nonlinear Systems, Power
Systems, Robotics, Social Systems, Economic Systems and other. Of particular
value to both the contributors and the readership are the short publication timeframe
and the world-wide distribution and exposure which enable both a wide and rapid
dissemination of research output.
Armenak V. Davidyan
Editors
123
Editors
Irina V. Gashenko Armenak V. Davidyan
Rostov State University of Economics Rostov State University of Economics
Rostov-on-Don, Russia Rostov-on-Don, Russia
Yulia S. Zima
Rostov State University of Economics
Rostov-on-Don, Russia
This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Contents
v
vi Contents
Taxation system is one of the most important elements of the institutional envi-
ronment for conducting entrepreneurial activities in the economic system. Its
effectiveness determines the successfulness of the state budget and development of
business, which determines the importance and significance of the problem of
taxation optimization. The purpose of the book is to determine the perspectives and
to develop the system of measures for reforming the taxation system of modern
Russia for its optimization. It provides an authors’ opinion on the issue of tax
optimization in the context of development of small and medium entrepreneurship
and offer innovational mechanisms for fighting tax evasion.
Emphasis on small and medium business in the context of modern Russia is the
peculiarity of the book, which distinguishes it from other publications on this topic.
This issue was selected for research due to the fact that existing publications in the
sphere of taxation are usually devoted to studying general fiscal problems in a state,
while in the conditions of new economy, the role of small and medium business
grows, and peculiarities and perspectives of taxation optimization should be studied
by the modern scientific society.
Another peculiarity of the book, which ensures its topicality and attractiveness
for the wide audience is studying the process of optimization of the system of
taxation not in the isolated manner but in connection to the tendency of informa-
tization (digitization) of the modern economy. This ensures not only the uniqueness
of the approach to treatment of optimality of taxation system but also the originality
of the offered authors’ recommendations for achieving it in economic practice of
modern socioeconomic systems by the example of Russia.
The basic hypothesis, which became the initial point for conducting a complex
of scientific and practical research, presented in this book, consists in the idea that
the current informatization (digitization) opens new possibilities and perspectives
for optimization of the taxation system. The purpose of the book is to study the
actual tendencies and preconditions to optimize the taxation system of modern
Russia and to substantiate expedience and to develop practical recommendations
for using the possibilities of its informatization (digitization) in this process.
ix
x Introduction
1 Introduction
Importance and relevance of taxes and taxation for modern economic systems is
practically assured and stressed by all scientists and experts in the area of economic
government administration. However, the complexity and versatility of taxes and
taxation brings an ambiguity in the interpretation of their notion and essence, which
presently need clarification in the frames of each scientific research depend on its
objectives.
Heterogeneity and incoherence, as well as inconsistency of available definitions
of concept and essence of taxes and taxation undermines scientific foundations for
studying this economic phenomenon (and process) and causes an issue to systemize
a knowledge accumulated in this area and to compose a clarified integral notion and
essence of taxes and taxation.
Due to globalization, by now taxes and taxation have become a way to provide a
global competitiveness of economic systems instead of the one to solve internal
macroeconomic tasks. In this regard, the world tax systems competitiveness is
ranked, for example, in accordance with the most established one, International Tax
Competitiveness Index, annually represented by Tax Foundation on the values of
corresponding index calculated by this international organization.
Performances that determine the tax systems competitiveness of are their
attractiveness for international investors (the lower tax rates and the higher stability
and predictability of tax policy, the more its attractiveness) and its neutrality (the
lesser omissions related to the application of tax benefits, the more neutrality). At
the same time, internal needs due to successful performance of functions imposed
on them, and the peculiarities of economic systems are not taken into account.
For this reason, in this study we put forward a hypothesis that the competi-
tiveness of the tax system does not guarantee a successful performance of functions
(and vice versa). The purpose of the study is to investigate available interpretations
of the notion and the essence of taxes and taxation and to represent the compre-
hensive one, as well as to reveal the relation between competitiveness and suc-
cessful performance of the tax system.
The Notion and the Essence of Taxes … 5
3 Results
Table 1 Fundamental approaches to reveal the essence and functions of taxes and taxation
Approach Key function Essence of taxes Representatives of approach
of taxes and and taxation
taxation
Revenue Revenue A way to assure Aslanov (2015), Miller et al. (2016),
state budget Nerudová and Dvořáková (2014),
revenues Nerudová and Solilová (2017)
Regulatory Regulating A way to Musayev and Musayeva (2018), Popkova
administer an et al. (2018a, b), Shokeen et al. (2017),
economic activity Zhang et al. (2016)
Social Social A way to ensure a Dagan (2017), Dallyn (2017), Faizal et al.
social justice (2017), Hearson (2017)
Supervision Supervising Self-controlled Cui and Notteboom (2017), Spartà and
(tax) system Stabile (2017), Verma et al. (2017), Wu
and Lu (2018)
Source Compiled by the authors
6 I. V. Gashenko et al.
Table 2 Original data to analyze the dependence of successful revenue performance of the tax
system and its competitiveness
Country The tax system State budget Country’s credit rating,
competitiveness, points from 0 balance, % of the points from 0 to 100
to 100 (position) (x) GDP (y) (max) (y)
Estonia 100.0 (1) 0.3 (12) 76.8 (24)
New 88.7 (2) 0.6 (9) 86.3 (15)
Zealand
Switzerland 85.2 (3) −12.4 (129) 20.0 (125)
Turkey 73.7 (11) −2.3 (54) 52.6 (68)
The United 70.8 (14) −3.1 (72) 88.9 (13)
Kingdom
Norway 70.7 (15) 2.9 (4) 94.8 (2)
Canada 69.1 (17) −1.9 (48) 92.3 (8)
Japan 66.8 (22) −4.2 (93) 82.0 (19)
Germany 66.6 (23) 0.8 (8) 94.7 (3)
Mexico 62.2 (25) −2.9 (70) 71.0 (34)
USA 55.1 (30) −4.4 (95) 93.4 (4)
Italy 47.7 (34) −2.4 (61) 68.4 (37)
France 43.4 (35) −3.3 (76) 84.0 (16)
Russia 34.42 (58) −3.7 (84) 54.2 (61)
Source Compiled by the authors based on: Tax Foundation (2018), World Economic Forum
(2018)
Fig. 1 Regression curves displaying the dependence of successful revenue performance of the tax
system on its competitiveness. Source Constructed by authors
Table 3 Original data to analyze the dependence of successful regulating performance of the tax
system and its competitiveness
Country The tax system Place in a global rating Index of Economic
competitiveness, points due to a living Freedom, points from 0
from 0 to 100 standard of the to 100 (max) (position)
(max) (position) (x) population (y) (y)
Estonia 100.0 (1) 27 78.8 (7)
New 88.7 (2) 2 84.2 (3)
Zealand
Switzerland 85.2 (3) 4 55.9 (123)
Turkey 73.7 (11) 88 65.4 (58)
The United 70.8 (14) 10 78.0 (8)
Kingdom
Norway 70.7 (15) 1 74.3 (23)
Canada 69.1 (17) 8 77.7 (9)
Japan 66.8 (22) 23 72.3 (30)
Germany 66.6 (23) 11 74.2 (25)
Mexico 62.2 (25) 61 64.8 (63)
USA 55.1 (30) 18 75.7 (18)
Italy 47.7 (34) 30 62.5 (79)
France 43.4 (35) 19 63.9 (71)
Russia 34.42 (58) 101 58.2 (107)
Source Compiled by the authors based on: Tax Foundation (2018), Legatum Institute (2018),
The Heritage Foundation (2018)
Fig. 2 Regression curves displaying the dependence of successful regulating performance of the
tax system on its competitiveness. Source Constructed by authors
As can be seen from Fig. 2, with increase in the tax system competitiveness of 1
point, the living standard of the population is reduced of 0.7459 items, and the
value of the economic freedom index is increased of 0.2433 points. Moderate
values of correlation coefficients in both cases (17.36 and 25.60% respectively)
evidence a statistical insignificance of dependencies revealed.
Within a social approach, taxes and taxation are perceived as a way to ensure a
social justice with focus on their social function (when admitting all other
8 I. V. Gashenko et al.
Table 4 Original data to analyze the dependence of successful social performance of the tax
system and its competitiveness
Country The tax system competitiveness, The European index of social justice,
points from 0 to 100 (position) (x) points from 0 to 10 (position) (y)
Estonia 100.0 (1) 6.19 (12)
The United 70.8 (14) 6.22 (11)
Kingdom
Germany 66.6 (23) 6.71 (7)
Italy 47.7 (34) 4.84 (25)
France 43.4 (35) 6.29 (10)
Denmark 67.0 (21) 7.39 (1)
Sweden 81.8 (6) 7.31 (2)
Finland 68.2 (19) 7.14 (3)
Poland 54.4 (31) 5.79 (15)
Portugal 51.9 (33) 5.36 (20)
Spain 59.8 (28) 4.96 (24)
Greece 57.2 (29) 3.70 (28)
Source Compiled by the authors based on: Tax Foundation (2018), Bartelsmann Stiftung (2018)
The level of social justice
6
y = 0.0317x + 3.9595
4 R² = 0.1959
0
0 20 40 60 80 100 120
The tax system competitiveness
Fig. 3 The regression curve displaying the dependence of successful social performance of the
tax system on its competitiveness. Source Compiled by the authors
functions). The performance of social justice is the social justice index composed
by Bartelsmann Stiftung for the European countries (Table 4 and Fig. 3).
As can be seen from Fig. 3, with increase in the tax system competitiveness of 1
point, the level of social justice is increased of 0.0317 points. The moderate value of
the correlation coefficient (19.59%) evidences a statistical insignificance of the
dependency revealed.
A supervision approach stresses an importance of a supervising function of taxes
and taxation (when admitting all other functions) defining them as a self-regulatory
(tax) system. It is believed that the more complex the tax system, the more difficult
to exercise its self-supervision. The performance of the tax system complexity is a
The Notion and the Essence of Taxes … 9
Table 5 Original data to analyze the dependence of successful supervising performance of the tax
system and its competitiveness
Country The tax system complexity, points from The tax system competitiveness,
0 to 100 (position) (x) points from 0 to 100 (position) (y)
Estonia 48.70 (14) 100.0 (1)
New Zealand 34.50 (9) 88.7 (2)
Switzerland 28.80 (19) 85.2 (3)
Turkey 41.10 (88) 73.7 (11)
The United 30.70 (23) 70.8 (14)
Kingdom
Norway 37.50 (28) 70.7 (15)
Canada 20.90 (16) 69.1 (17)
Japan 47.40 (68) 66.8 (22)
Germany 48.90 (41) 66.6 (23)
Mexico 52.10 (115) 62.2 (25)
USA 43.80 (36) 55.1 (30)
Italy 48.0 (112) 47.7 (34)
France 62.20 (54) 43.4 (35)
Russia 47.50 (52) 34.42 (58)
Source Compiled by the authors based on: Tax Foundation (2018), World Bank Group,
PricewaterhouseCoopers (2018)
70
60
competitiveness
y = -0.2756x + 60.685
The tax system
50 R² = 0.2054
40
30
20
10
0
0 20 40 60 80 100 120
The tax system complexity
Fig. 4 Regression curves displaying the dependence of the tax system competitiveness on
successful supervising performance. Source Compiled by the authors
Regulating
Promoting
function
(attractiveness for investors
ta abroad)
Self-supervision xe
s
Function to provide
Tax system competitiveness
Social function
Economic entities
Fig. 5 Complex modern concept of the essence and functions of taxes and taxation Source
Compiled by the authors
4 Conclusions
Thus, hypothesis under suggestion has been proofed: multiple tests of nature and
strength of relation between the tax systems competitiveness and their successful
performances on the statistical data by sampling of different countries of the world
The Notion and the Essence of Taxes … 11
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2017-1/. Data accessed 02.06.2018.
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Tax System of a State: Federal, Regional
and Local Taxes and Fees
Abstract Purpose The purpose of the paper is studying the peculiarities to adjust
contradictions in the Russian tax system and drawing up a practical model to
organize tax system of the country and to implement a fiscal federalism in modern
Russia. Methodology The methodology for testing a hypothesis under suggestion
includes methods of economic statistics, namely, methods of structural and hori-
zontal analysis, as well as ones of comparative and plan-fact analysis and for-
malization (graphical representation of data). Information and analytical
background of the study is the official statistical data of the Federal Tax Service of
the Russian Federation, the Ministry of Finance of the Russian Federation, and the
Federal State Statistics Service of the Russian Federation (Rosstat) for 2008–2017.
Results We revealed deviations from preferable (target) peculiarities to adjust
contradictions in the tax system of modern Russia that lead to imbalance of local
fiscal subsystems, escalation of deficit in all level-budgets, as well as establishing
the dependence of territories on intergovernmental transfers. We have built a
practical model for organization of the tax system and for implementation of fiscal
federalism in the Russian Federation, which has not only advantages (such as
integrity, etc.), but also disadvantages (enhanced limitation of fiscal initiatives and
opportunities of territories, multiple overlapping of tax revenue flows, etc.).
Recommendations We advise an improvement of the tax system in modern Russia
through adjustment of revealed disadvantages.
Keywords Tax system of the country Federal Regional and local taxes and fees
Fiscal federalism
1 Introduction
Functions of taxes and taxation are performed through establishment, operation and
development of the tax system of the country. The fundamental principle of tax
systems in most modern countries of the world is a fiscal federalism, which
involves a simultaneous delimitation and strong interrelation of federal, regional
and local fiscal systems. Following this principle causes three inevitable contra-
dictions in the tax system:
– The collision of federal–national (more internationally-oriented) and territorial–
regional and local (more nationally-oriented) interests;
– The collision of interests in public performance (resulting in growth of expen-
ditures and a budget deficit) and in maintaining remunerativeness (with possible
surplus) of government budgets;
– The collision between national interests of balanced territorial development
(granting temporary federal support to the territories to activate their own
economic growth) and territorial interests to rely on federal support (establishing
dependence on federal support without following-up their own initiatives to
activate economic growth).
The peculiarities of adjusting these contradictions define specific nature of
practical models to organize the tax system of the country and to implement fiscal
federalism. The Concept of long-term social and economic development of the
Russian Federation until 2020 approved by the decree No. 1662-p. as of November
17, 2008 states the following preferred (target) peculiarities of adjusting these
contradictions (Government of the Russian Federation 2018a):
– prevailing of territorial interests (as evidenced by declared decentralization of
the fiscal system and its commitment to provide regional economy
development);
– prevailing of interests in public performance (as evidenced by declared
large-scale goals of social and economic development along with a refusal to
increase the total tax burden in economy);
– prevailing of national interests in balanced territorial development (as evidenced
by declared heightened commitment of the tax system to the issues of levelling
up income and enhancing spur impact of the tax system on economic
development).
Given that the above-noted concept was developed and adopted before the crisis
of the Russian economy in 2009, current hypothesis of this research consists in the
author’s suggestion that the preferred (target) peculiarities to adjust contradictions
of the tax system are not fully achieved (mispresented) in modern Russia due to
Tax System of a State … 15
crisis impact (among other). The purpose of the paper is studying the peculiarities to
adjust contradictions in the Russian tax system and drawing up a practical model to
organize tax system of the country and to implement a fiscal federalism in modern
Russia.
A review of available studies and papers on the topic allowed identifying two basic
theoretical models of the state tax system organization and the implementation of
fiscal federalism—decentralized and centralized—upon which practical models of
countries are to be built.
A decentralized model is applied in countries such as the Great Britain, the USA,
Canada, Japan, etc. It involves a high degree of financial (fiscal) independence of
territories within the state, wide opportunities to introduce their own taxes, define
performances (for example, tax rates), and to make loans, as well as the lack of
activity in taking measures aimed at levelling-up imbalances in local budgets and
overcoming unremunerativeness. This model is studied in papers by authors such as
Del Bo (2018), Haddow (2018), Kimura (2017), Nicholson-Crotty et al. (2006), and
Xing (2018).
A centralized model is much more widespread and applied in countries such as
Germany, France, Mexico, China, etc. It is related to a low degree of financial
(fiscal) independence of territories within the state, their strictly limited opportu-
nities to introduce their own taxes and define performances (for example, tax rates),
and to implement loans, as well as activity in taking measures aimed at levelling-up
imbalances in local budgets and overcoming unremunerativeness. It is considered in
papers by scientists such as Grabova et al. (2018), Gunlicks (2012), Hoyt (2017),
and Sun et al. (2017).
Modern Russia follows a centralized theoretical model enshrined in the Tax Code
of the Russian Federation (TC RF) N 146-ФЗ as of July 31, 1998. Pursuant to it, the
tax system in modern Russia is a three-tiered one with the following categories and
corresponding taxes and fees (Government of the Russian Federation 2018b):
– Federal (macro-level category): value added tax, excise duties, personal income
tax, corporate income tax, mineral extraction tax, water tax, levies for the use of
fauna and aquatic, state duty;
– Regional (meso-level category): corporate property tax, gambling tax, transport
tax;
– Local (micro-level category): land tax, individual property tax, sales tax.
The Russian practice of organizing tax system and implementing fiscal feder-
alism is discussed in publications by experts such as Dyukina (2017), Gashenko
et al. (2018), Ibragimov et al. (2014), Popkova et al. (2018a, b), Shvetsov and
Balikoev (2017).
16 I. V. Gashenko et al.
3 Results
20000.0
15000.0
8181.5
10000.0
5000.0 9162.0
0.0
Fig. 1 Structure of revenues and expenditures of the consolidated budget in the Russian
Federation in 2017, bln. rub. Source Compiled by the authors based on: Federal Tax Service of the
Russian Federation (2018), Ministry of Finance of the Russian Federation (2018), Rosstat (2018)
Tax System of a State … 17
As can be seen from Fig. 1, in the revenue structure of the consolidated budget
of the Russian Federation (RUB 17,343.5 billion) proportions of the federal budget
(RUB 9162 billion, 52.83%) and consolidated budgets of territories (entities) (RUB
8181.5 billion, 41.17%) are distributed rather equally. In the structure of expen-
ditures, ones related to implementation of national interests are prevailing (RUB
4682.9 billion, 27%), including:
– Expenditures on national issues (RUB 648.6 billion);
– Expenditures on national defense (RUB 1591.5 billion);
– Expenditures on national security and law enforcement activity (RUB
1070.2 billion);
– Expenditures on national economy (RUB 1372.7 billion).
The proportion of federal budget expenditures on national interests’ imple-
mentation is 21% (RUB 3642.2 billion), including:
– Expenses for housing and public services (RUB 66.7 billion);
– Expenses for social and cultural activities (RUB 2935.5 billion);
– Expenses for education (RUB 343.2 billion);
– Expenses for healthcare (RUB 245.4 billion);
– Expenses for environmental protection (RUB 51.6 billion).
The proportion of local budget expenditures on national interests’ implementa-
tion is 33.30% (RUB 5775.3 billion), including:
– Expenses for housing and public services (RUB 770.9 billion);
– Expenses for social and cultural activities (RUB 5004.5 billion).
To reveal the peculiarities of ratio between interests in public performance and in
maintaining remunerativeness (with possible surplus) of government budgets in
modern Russia, let’s turn to the balance dynamics in consolidated budget of the
Russian Federation in 2008–2017 (Fig. 2).
3000
2000
1000
0
-1000
trend
-2000
-3000
-4000
2008 2009 2010 2011 2012 2013 2014 2015 2016 2017
Series1 2014.2 -2448.6 -1584.7 860.8 260.4 -848.7 -845.6 -2819.5 -3142.2 -3569.1
Fig. 2 Balance dynamics in consolidated budget of the Russian Federation in 2008–2017, RUB
billion. Source Compiled by the authors based on: Federal Tax Service of the Russian Federation
(2018), Rosstat (2018)
18 I. V. Gashenko et al.
As can be seen from Fig. 2, the expenditures of the Russian Federation con-
solidated budget exceed revenues over almost all period under study, and the
balance of this budget is featured by negative dynamics and a downward trend,
which evidences an escalation of the budget deficit. In comparison with 2008, when
the budget surplus amounted to RUB 2014.2 billion, in 2017 there was a budget
deficit of RUB 3569.1 billion. This evidences the prevailing of interests in public
performance over the ones in maintaining budget remunerativeness.
To reveal the peculiarities of ratio of the national interests of balanced territorial
development (granting temporary federal support to the territories to activate their
own economic growth) and the territorial interests to rely on federal support
(establishing dependence on federal support without following-up their own ini-
tiatives to activate economic growth) in modern Russia, let’s turn to balance
dynamics in consolidated budgets of the Russian Federation territories (entities) in
2008–2017 (Fig. 3).
As can be seen from Fig. 3, over the entire period under study we observe a
negative balance in consolidated budgets of the Russian Federation territories
(entities). The highest gravity of their deficit had occurred in 2013 (641.5
RUB billion.). Despite the upward trend, in 2017 still there is a negative balance in
consolidated budgets of the Russian Federation territories (entities) in the amount of
RUB 24.5 billion. It points to the establishment of the dependence of modern
Russia territories on federal support and the prevailing of territorial interests.
Based on the results of our analysis, we built up the following practical model of
the tax system organization and the implementation of fiscal federalism in modern
Russia (Fig. 4).
As can be seen from Fig. 4, the practical model of the tax system organization
and the implementation of fiscal federalism has a complicated structure. In addition
to the advantages connected with its high degree of unification, strong interrelation
and interdependence of all-level fiscal system, and their mutual support for
achieving balance and remunerativeness, this model is featured by the following
disadvantages:
0
-100
-200
-300
-400
-500
-600 trend
-700
2008 2009 2010 2011 2012 2013 2014 2015 2016 2017
Series1 -54.3 -329.1 -99.6 -34.9 -278.7 -641.5 -447.6 -171.6 -12.6 -24.5
Fig. 3 Balance dynamics in consolidated budgets of the Russian Federation territories (entities) in
2008–2017, RUB billion. Source Compiled by the authors based on: Federal Tax Service of the
Russian Federation (2018), Rosstat (2018)
Tax System of a State … 19
Inter-government
Federal fiscal system National
transfers
Taxes (macroeconomic)
Levelling-up imbalances level
Fig. 4 Practical model of the tax system organization and the implementation of fiscal federalism
in Russia. Source Compiled by the authors
4 Conclusion
Thus, hypothesis under suggestion is proofed and deviations from the preferable
(target) peculiarities to adjust contradictions of the tax system in modern Russia
leading to imbalance in local fiscal systems, escalation of budget deficit at all levels,
and establishing the dependence of territories on inter-government transfers are
revealed.
A practical model built by us to organize the tax system and to implement fiscal
federalism in the Russian Federation has not only advantages (such as integrity,
etc.), but also disadvantages (enhanced limitation of fiscal initiatives and
20 I. V. Gashenko et al.
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Napoleon. His soldiers in drinking at pools sucked up the small leeches not thicker
than a horse's hair, whose presence in the hinder part of the mouth cavity produced
divers objectionable results, such as spitting of blood and hindered respiration.
The purely aquatic leeches swim by undulations, and also crawl by the help of the
two suckers, like a "Geometer" caterpillar. But when a land-leech is dropped into the
water it at once sinks to the bottom and crawls out; it does not swim, but can survive
immersion for a long period. In this it resembles the earthworms, which can also
survive a prolonged immersion, and even in the case of some are indifferent to the
medium, land or water, in which they live; the land-leech, however, is entirely
dependent upon damp surroundings; a dry air is fatal to it. The land-leech of Japan
leaves a slimy trail behind it as it crawls, in this respect recalling the land Planarian
Bipalium kewense.
GEPHYREA AND PHORONIS
BY
CHAPTER XV
GEPHYREA
INTRODUCTION—ANATOMY—DEVELOPMENT—SIPUNCULOIDEA—PRIAPULOIDEA—
ECHIUROIDEA—EPITHETOSOMATOIDEA—AFFINITIES OF THE GROUP.
The animals included in the above-named group were formerly associated with the
Echinodermata. Delle Chiaje[468] states that Bohadsch of Prague in 1757 was the
first to give an accurate description of Sipunculus under the name of Syrinx, but
Linnaeus, who noted that in captivity the animal always kept its anus directed
upwards, re-named it Sipunculus. Lamarck[469] placed the Gephyrea near the
Holothurians; and Cuvier[470] also assigned them a position amongst the
Echinoderms. He mentions Bonellia, Thalassema, Echiurus, Sternaspis, and three
species of Sipunculus, one of which, S. edulis, "sert de nourriture aux Chinois qui
habitent Java, et qui vont la chercher dans le sable au moyen de petits bambous
préparés."
The name Gephyrea[471] was first used by Quatrefages, who regarded these
animals as bridging the gulf between the Worms and the Echinoderms. He included
in this group the genus Sternaspis (vide p. 335), now more usually classed with the
Chaetopoda.
The Gephyrea are exclusively marine. They are subcylindrical animals, which can
either retract the anterior end of their body—the introvert—carrying the mouth into
the interior; or are provided with a long flexible but non-retractile proboscis. The
latter is easily cast off. They usually bear spines or hooks of a hard chitinous
character, secreted by the epidermis or outermost layer of cells. The mouth is at the
base of the proboscis or at the end of the protractile part, the anus is at the other
end of the body or on the dorsal surface. The nervous system consists of a ring
round the mouth and of a ventral nerve-cord. A vascular system is present as a rule.
Nephridia are found which act as excretory organs, and in most cases also as ducts
for the generative cells. The Gephyrea are bisexual, and the male is sometimes
degenerate.
The group may be divided into four Orders:—(i.) Sipunculoidea; (ii.) Priapuloidea;
(iii.) Echiuroidea; (iv.) Epithetosomatoidea; of these the first is by far the largest,
both in number of genera and of species.
The introvert occupies about one-sixth or one-fifth of the total body length. It is
somewhat narrower than the trunk, and is covered by a number of small flattened
papillae, some of which lie with their free ends directed backward, overlapping one
another like tiles on a roof. In some other genera, as Phymosoma, the introvert
bears rows of horny hooks, which are apt to fall off as the animal grows old.
The trunk has from thirty to thirty-two longitudinal furrows, the elevations between
which correspond with a similar number of muscles lying in the skin. This
longitudinal marking is crossed at right angles by a circular marking of similar origin,
the elevations of which correspond with the circular muscles in the skin. These two
sets of markings thus divide the skin of the trunk into a number of small square
areas, very regularly arranged (Fig. 212).
The outline of the trunk is more or less uniform, but it is capable of considerable
change according to the state of contraction of its muscles. The circular muscles, for
instance, may be contracted at one level, thus causing a constriction at this spot.
The colour of S. nudus is a somewhat glistening greyish-white.
Fig. 211.—Right half of the anterior end of Sipunculus nudus L., seen from the inner
side and magnified. a, Funnel-shaped grooved tentacular crown leading to the
mouth; b, oesophagus; c, strands breaking up the cavity of the tentacular crown
into vascular spaces; c', heart; d, brain; e, ventral, and e', dorsal retractor
muscles; f, ventral nerve-cord; G, vascular spaces in tentacular crown.
The anterior end of the fully-expanded Sipunculus may be termed the head; here
the skin is produced into a frayed fringe which stands up in the shape of a funnel
round the mouth. This fringe is grooved on its internal surface with numerous little
gutters, all of them lined with cilia, which by their constant motion keep up a current
which sweeps food into the mouth. The fringe may be in the form of a simple ring
round the mouth, or the ring may be folded in at the dorsal side so as to take the
form of a double horse-shoe (Figs. 211 and 212).
Scattered over the surface of the body, and opening by narrow tubes which pierce
the cuticle, are a number of glandular bodies which may be either bi- or multi-
cellular. The glandular cells are apparently enlarged and modified epidermal cells;
they are arranged in a cup-shaped manner, with their apices directed towards the
orifice. They are crowded with granules, which are presumably poured out over the
cuticle, but the exact function of the secretion is entirely unknown. They have a well-
developed nerve supply.
Digestive System.—The mouth lies in the centre of the fringe, and is not provided
with any kind of jaw or biting armature; it leads directly into the thin-walled
alimentary canal, the first part of which is ciliated. The alimentary canal is not
marked out into definite regions, but passes as a thin-walled semi-transparent tube
to the posterior end of the body, and then turns forward again and opens to the
exterior by an anus situated about an inch below the junction of the introvert with the
trunk, on the median dorsal line. The descending and ascending limbs of the
alimentary canal are coiled together in a spiral, which may be more or less close in
different individuals. The whole is supported by numerous fine muscular strands,
which pass from the walls of the intestine to the skin, and by a spindle-muscle,
which runs from the extreme posterior end of the trunk up the axis of the spiral and
terminates in the skin close to the anus.
No glands open into the alimentary canal at any point of its course, but near the
anus a simple diverticulum, or pocket, of unknown function arises. The size of this
outgrowth differs enormously in different individuals. The alimentary canal near the
anus also bears two tuft-like organs, which, however, do not open into the intestine,
but probably have some function in connexion with the fluid in the body-cavity.
Along the whole course of the alimentary canal there runs a ciliated groove, into
which the food does not pass, but the cilia of which probably keep in motion a
current of water whose function may be respiratory.
Fig. 212.—Sipunculus nudus L., with introvert and head fully extended, laid open by
an incision along the right side to show the internal organs. × 2. a, Mouth; b,
ventral nerve-cord; c, heart; d, oesophagus; e, intestine; f, position of anus; g,
tuft-like organs; h, right nephridium; i, retractor muscles; j, diverticulum on
rectum. The spindle-muscle is seen overlying the rectum.
Vascular System.—On the dorsal surface of the anterior end of the alimentary
canal lies a contractile vessel, usually termed the heart. It is a tube about an inch
long, ending blindly behind, but opening in front into a ring-shaped space
surrounding the mouth and partially enveloping the brain. From this ring-like vessel
numerous branches are given off which pass into the fringe round the mouth, and
probably the chief function of the heart is by its contraction to force fluid into this
fringe, and so to extend it. The heart contains a corpusculated fluid. A similar but
shorter tube is found on the ventral surface of the anterior end of the alimentary
canal in the species in question; it also opens into the ring which surrounds the
mouth.
Sipunculus is not well provided with sense-organs, but in an animal which lives
buried in sand we should not expect to find these very highly developed. On the
introvert there are certain patches of epithelium bearing long stout cilia, which have
been regarded as tactile in function, and there is a tubular infolding reaching the
brain, which almost certainly has some sensory function. Ward[474] has termed this
"the cerebral organ." It consists of a duct lined with ciliated cells, which opens to the
exterior in the middle dorsal line outside the tentacular fringe. The duct leads down
to the brain, and expands at its lower end into a saucer-shaped space, covering that
portion of the brain where its substance is continuous with the external epithelium.
In Phymosoma this cavity is produced into two finger-shaped processes, which are
sunk into the brain and are lined by cells crowded with a dense black pigment.[475]
They are probably rudimentary eyes, perhaps distinguishing only between darkness
and light. The pits appear to be absent in Sipunculus nudus, but Andrews states
they are found, although without pigment, in S. gouldii.[476]
The eggs break away from the ovary in a very undeveloped condition, but whilst
floating about in the body-cavity they increase in size and secrete a thick membrane
around them. They have a well-marked nucleus, and are oval in outline.
The mother-cells of the spermatozoa also break away in an immature condition, and
complete their development in the nutritive fluid of the body-cavity. They divide into
a number of spermatozoa, usually eight or sixteen, which remain in contact. They
each develop a tail, which projects outwards, and aids the cluster in swimming
along. These clusters of spermatozoa are about the same size as the ova of the
female, and, like them, make their way into the "brown tubes." The exact way in
which this is accomplished is not very clear, but the cilia on the funnel-shaped
internal opening of the tube seem to have some power of selecting the generative
cells when they come within their reach, and of passing them on, whilst they reject
the much smaller corpuscles of the perivisceral fluid, which are never found in the
nephridia.[477] Once inside the internal opening, the clusters break up and the
spermatozoa escape singly into the sea. Here they meet with and fertilise the eggs
which have escaped from the body of the female.
The fluid of the body-cavity contains corpuscles, which are kept in active circulation
by the constant contractions of the body-wall, and by numerous tufts of cilia which
are borne on the inner surface of the skin. The dorsal blood-vessel is one of the
latest organs to arise.
The larva swims actively about for a month, during which time it increases greatly in
size; it then undergoes a somewhat sudden metamorphosis. The ciliated ring and
the structures related to the oesophagus begin to disappear, the distinction between
the head and the rest of the body is obliterated, and the head becomes relatively
small. The mouth changes its position, and becomes terminal instead of being
somewhat ventral, and the tentacular membrane begins to appear. At the same time
the larva relinquishes its free-swimming life, and sinks to the bottom; it begins
creeping amongst the sand by protruding and retracting the anterior part of its body,
and takes on all the characters and habits of the adult.
I. Order Sipunculoidea.
Besides the genus Sipunculus, the Order Sipunculoidea includes ten other genera.
A key to these, taken for the most part from Selenka's admirable monograph, is
given on page 424.
The ventral side of each tentacle is grooved and ciliated, and the grooves are
continued into the ciliated mouth. Their dorsal surface is pigmented, and in the
hollow of the horse-shoe lies a deeply pigmented epithelium covering the brain.
A blood-vessel courses up each tentacle, and usually two channels return the blood
to the vascular ring which surrounds the mouth. In those forms which possess
tentacles on the dorsal side of the mouth only, the ventral part of the vascular ring
lies in the lower lip, which is tumid and swollen. The brain supplies a nerve to each
tentacle.
When the introvert is retracted the tentacular ring is withdrawn and to some extent
collapsed; in this condition it would be almost touching the rough external surface of
the introvert. In some species of Phymosoma the delicate appendages of the head
are guarded from the hooks on the introvert by a thin membrane or collar,[479] which
completely ensheaths the retracted head.
When the introvert is fully extended the dorsal blood-vessel contracts and sends its
blood forward into the vascular ring, and thence into the tentacles or tentacular fold,
which are thus erected. In several species of Sipunculus, as S. nudus, S.
norvegicus, S. robustus, S. tesselatus, there is a ventral blind tube as well as a
dorsal, into which the blood is withdrawn when the head is retracted. In many other
species in various genera, such as Phymosoma weldonii and Ph. asser,
Dendrostoma signifer, S. vastus, the lumen of the dorsal vessel is increased by
numerous hollow blind processes which it bears, hanging freely into the body-cavity.
Three very small genera of Sipunculids—Onchnesoma, Petalostoma, and Tylosoma
—are devoid of all trace of vascular system and of tentacles; the mouth opens in the
centre of the anterior end of the introvert. In Onchnesoma the dorsal part of the lip is
somewhat produced, so that the head has somewhat the shape of a Doge's cap,
and in Petalostoma there are two leaf-like processes of the body-wall which guard
the mouth.
The extent to which the intestine is coiled varies very much even in the same
species; the axis of the coil is often supported by a spindle-muscle, but this is
sometimes absent. The caecum, which opens into the rectum of S. nudus, is again
a very variable structure, and when it is present varies remarkably in size.
The food of Sipunculids seems to consist almost entirely of sand, and their only
nourishment must be such small microscopic organisms or particles of animal and
vegetable débris as are to be found mixed with the sand. The alimentary canal is, as
a rule, quite full of sand, and yet in spite of the tenuity of its walls they never seem
to be ruptured. If the contents of the digestive tube be washed out with a pipette, it
will be found that it requires considerable force to dislodge many of the sand-
particles lying next the wall. These are more or less embedded in crypts or pockets
of the wall, and as the sand passes along the intestine they probably serve as more
or less fixed hard points, against which the sharp edges of the sand particles are
worn off. Amongst the sand are usually to be found pieces of shell, sometimes with
a diameter equal to that of the alimentary canal; these are usually rounded, but their
angles may have been removed by attrition before they entered the mouth of the
Sipunculid.
The enormous amount of sand and mud which passes through the bodies of the
Sipunculids shows that they must take a considerable part in modifying the mineral
substances which form the bottom of the sea. Just as earthworms, as shown by
Darwin, play a considerable rôle in the formation of soil, so must these animals, in
conjunction with Echinids and Holothurians, effect considerable modifications in the
sand and mud which pass through their bodies. Mr. J. Y. Buchanan[480] is "led to
believe that the principal agent in the comminution of the mineral matter found at the
bottom of both deep and shallow seas and oceans, is the ground fauna of the sea,
which depends for its subsistence on the organic matter which it can extract from
the mud." The minerals at the bottom of the sea are exposed to a reducing process
in passing through the bodies of the animals which eat them, and subsequently to
an oxidising process due to the oxygen dissolved in the sea-water acting on the
minerals extruded from the animals' bodies.
The rate at which the sand passes through the body of Sipunculus is unfortunately
unknown, but that at any one moment a considerable quantity is contained in the
intestine is shown by the fact that the average weight of five specimens of S. nudus
from Naples, taken at random, was 19.08 grms., whilst the average weight of sand
washed out of their alimentary canal was 10.03 grms. The sand contained in five
other specimens of the same species measured respectively 6 c.c., 7 c.c., 6.5 c.c.,
7.5 c.c., and 7.5 c.c., giving an average of 6.9 c.c. for each individual.
Onchnesoma and Tylosoma have only one retractor muscle; Aspidosiphon and
Phascolion have, as a rule, two; Phymosoma and Sipunculus have four, and
perhaps this is the more usual number.
Phascolion, Tylosoma, and Onchnesoma have but one "brown tube"; in Phascolion
this is the right, in Onchnesoma it is sometimes the right and sometimes the left that
persists. Most other genera retain two, but there are many exceptions; for instance,
Phascolosoma squamatum has but one, and so has Aspidosiphon tortus, and in
both cases it is that of the left side. No Sipunculid has more than two. It has been
pointed out by Selenka that those species which have but one brown tube are, as a
rule, inhabitants of tubes or shells, and do not move actively about in the sand.
The eggs of all members of the family, with the exception of the genus Phymosoma,
are spherical, but those of the last-named genus are elliptical. They are always
surrounded by a thick membrane, the "zona radiata," pierced by numerous pores.
I. The longitudinal muscles in the body-wall divided into 17-41 distinct bundles.
Four retractor muscles.
A. Body covered with papillae. Numerous filiform tentacles which seldom (or
never?) surround the mouth, but stand above and dorsal to it in a horse-
shoe, with the opening dorsal. No rectal caecum. Hooks usually present.
Four retractors (in Ph. Rupellii only two?). Heart almost always without
caeca. Eye-spots always present. Eggs oval, flat, reddish. Almost entirely
small tropical species
1. Phymosoma
II. The longitudinal muscles in the body-wall form a continuous sheath, and are
not split up into bundles.
A. Two brown tubes. Numerous tentacles form a wreath round the mouth.
Alimentary canal forms a complete spiral, free behind except in Ph. Hanseni.
Spindle-muscle usually present. One or more ligaments present, but only on
the anterior convolutions of the intestine. Adhesive papillae always absent.
Hooks very frequently absent. Eggs spherical. Found in all seas.
3. Phascolosoma
B. Two free brown tubes. Only four or six plumed tentacles. A complete
intestinal spiral, not attached behind. Spindle-muscle always present. One or
more ligaments present, but only on the anterior convolutions of the
intestine. Hooks are present, but sometimes fall off early in life. Heart usually
bears caeca. Found only in the tropics.
4. Dendrostoma
C. Only one brown tube, that of the right side, present; it is attached to the
body-wall throughout its entire length. Numerous tentacles form a circle
round the mouth. The alimentary canal forms no spiral, or an incomplete
one. No spindle-muscle, but the intestine is attached to the body-wall
throughout its length by numerous ligaments. Adhesive papillae often
present. Not more than two retractors. Spherical eggs. Inhabits Mollusc
shells or tubes. Found in all seas
5. Phascolion
III. At both ends of the trunk a distinct horny shield, or tube-like cornification, or
a calcareous ring at the anterior end of the trunk. Hooks sometimes present.
Longitudinal muscles continuous or split up into bundles.
A. A shield at both ends of the trunk. Introvert excentric, arising from the
ventral side of the anterior shield. Tentacles small and few in number,
arranged in a horse-shoe above the mouth. A spindle-muscle, which arises
from the posterior end of the body, traverses the intestinal coil. Two
retractors only, these are the ventral; they are frequently fused together from
their point of origin.
6. Aspidosiphon
B. A calcareous ring surrounds the anterior end of the trunk, from the middle
of which the introvert is extruded. Longitudinal muscles continuous. Hooks
bifid. Tropical.
7. Cloeosiphon
C. A corneous ring, from which the introvert issues, surrounds the anterior
end of the trunk, and the posterior end of the trunk is produced into a
corneous spike. Six pinnate tentacles encircle the mouth. Four retractors.
Hooks present on the introvert. Longitudinal muscles continuous. Intestine
not coiled throughout in a spiral nor fastened posteriorly. Spindle muscle
present.
8. Golfingia
IV. No tentacles, but two leaf-like extensions of the body-wall guard the mouth.
Four retractors. Few intestinal loops, quite free. No vascular system.
9. Petalostoma
B. No introvert (?). Body cylindrical, thickly covered with papillae, which are
larger and more crowded at both ends of the trunk.
11. Tylosoma
The genus Sipunculus contains sixteen species. They are the largest and the most
conspicuous members of the group. They have a very wide distribution, some
species, as S. nudus (Fig. 212) and S. australis, being almost cosmopolitan. They
are most common in temperate and tropical seas, but S. norvegicus and S.
priapuloides are found far north, but always at considerable depths, 100 to 200
fathoms.
The following account of the habits of Sipunculus gouldii is taken from Mr.
Andrews'[482] paper on that species:—
"This Sipunculus is very abundant in certain small areas of compact, fine sand
darkened by organic matter and not laid bare at ordinary low tide. In such places,
only a few square metres in extent, they pierce the sand in all directions to a depth
of more than half a metre, making burrows with persistent lumen running from the
surface downward and then laterally, but with no regularity in direction.
"Kept in aquaria, the dependence of the animal upon the nature of the sand and its
method of locomotion may be readily observed. A vigorous individual buries itself in
a few moments in the following manner: Running out the introvert to nearly its full
extent, and applying it to the surface of the sand till some spot of less resistance is
found, the animal still further expands the introvert so that it penetrates the sand,
provided this is not too dense and firm, for then the body is merely shoved
backward. When the introvert is inserted, the contraction of the longitudinal muscles
of the body-wall brings the whole body forward somewhat, in case the introvert is
fixed in the sand. In case soft ooze was present, this fixation did not take place, and
the introvert was merely pulled out again, but when the sand was of the right
consistency the introvert was fixed by becoming much swollen at the tip, and then
constricted just posterior to this swollen area. This bulb-like area exerts lateral
pressure on the sand, as could be seen by movements of the grains. The swelling
of the anterior end of the introvert is brought about by the body-wall contracting
elsewhere, and forcing in liquid to distend that end. Owing to the curved form
assumed by the body in the normal contracted state when first removed from its
burrow, the entrance of the introvert may often be nearly vertical, and hence the
entire body is soon raised nearly upright in the water above the sand. If the body
has thus been warped forward sufficiently to become somewhat fixed in the sand,
the introvert is rolled in and again thrust forward from this new point of resistance,
and so on till the animal is entirely buried. This locomotion increases in speed as the
creature becomes more completely surrounded by sand, and is the only means of
moving from place to place.
"On a smooth surface, or on one not presenting the right degree of resistance, the
Sipunculus does not change its position, but remains till death finally occurs, rolling
its introvert in and out and contracting its body-wall to no purpose.
"The essential factors in the mechanism bringing about this hydrostatic locomotion
are an elongated contractile sac filled with liquid, and some means of definitely co-
ordinating the contractions of the sac.
"In natural environment the animals are found with sometimes one, sometimes the
other end nearer the surface of the sand: in the aquaria the same was observed, but
when the water became stagnant and impure the anterior end with expanded
branchiae was often protruded somewhat above the surface of the sand."
The genus Phascolosoma contains at least twenty-five species, for the most part
small. Ph. margaritaceum, however, measures[483] 10 cm. in length, and Ph.
flagriferum, 13 cm. The latter is produced at the hinder end of its trunk into a long
whip-like process, which recalls the horny spike of Golfingia. Most species live free,
but a few inhabit the shells of dead Gasteropods or of Dentalium, or the abandoned
tubes of worms. They occur in practically all seas.
Fig. 216.—Specimens of the Coral Heteropsammia cochlea, with Aspidosiphon
heteropsammiarum or A. michelini living in a state of commensalism with them.
(From Bouvier.)
Dendrostoma contains but five species, which are all found within the tropics in the
Pacific or in the West Atlantic. They are shallow-water forms, and some are found
between tide-marks.
Phascolion is a smaller genus, containing but ten species, which may have been
derived independently from different species of Phascolosoma, and in this case the
genus should be broken up. The members of this genus live in Mollusc shells, such
as Dentalium, Turritella, Buccinum, Chenopus (Aporrhais), Nassa, Strombus, and
generally acquire the coiled shape of their host. They are usually attached to the
shell by means of certain adhesive papillae found on their posterior end. Ph. strombi
fills its shell with mud, which must be kept together by some secretion of the animal.
The body lies in a tube in this mud, and the introvert projects from the small round
opening at the end of the tube, and explores the ground in every direction. They are
found in all seas, but more especially in the colder waters.
The genus Aspidosiphon includes nineteen species, which are, with few exceptions,
exclusively confined to the Indian Ocean and neighbouring seas, including the Red
Sea. The exceptions are A. armatus from the Norwegian coast, and A. mülleri from
the Mediterranean and Adriatic. A. truncatus is also stated to occur at Panama, the
Bahamas, and at Mauritius. The remaining species almost all occur in the Malay
Archipelago and neighbouring islands, and as was the case with Phymosoma, this
part of the world seems to be the headquarters of the genus. A. mülleri lives in the
interstices of rocks and stones, and occasionally in disused Mollusc shells.
Petalostoma comprises but one species, P. minutum, which is found in the English
Channel.
Onchnesoma comprises two species, O. steenstrupii and O. sarsii, both found off
the coast of Norway at considerable depths between 200 and 300 fathoms.
Tylosoma comprises one species, T. lütkenii, also from the Norwegian coast. It is
dredged from stony ground in 50 to 80 fathoms.
Anatomy.—This Order consists of the two genera Priapulus and Halicryptus. Both
are cylindrical animals with the mouth at one end and the anus at the other. The
introvert is short, and is covered with rows of chitinous spines, which are continued
to some extent over the body.
The skin is folded in a series of rings, and the body is usually somewhat swollen
posteriorly. P. caudatus bears a curious caudal appendage, beset with a number of
hollow lobes somewhat grape-like in appearance. This is situated ventral to the
anus; its lumen is continuous with that of the body-cavity, but it can be separated
from it by the action of a sphincter muscle. Two such appendages exist in P.
bicaudatus.
There cannot be said to be any head in the Priapuloidea; they have no tentacles or
tentacular fringe, no proboscis, and no distinct brain; simply a round aperture, the
mouth, which is surrounded by a groove in the skin, at the bottom of which the
circumoesophageal nerve-cord lies. The mouth leads into a very muscular pharynx
lined with stout chitinous teeth; this passes into an intestine, which is as a rule
straight, but in P. glandifer it has a single loop.
The Priapuloidea possess no vascular system and no brown tubes. Their skin has in
the main the same structure as that of the Sipunculids, with spines, glandular
bodies, and papillae with sensory hairs which resemble similar structures on
Phymosoma varians. Retractor muscles arise from the longitudinal muscles of the
skin, and are inserted into the pharynx; they are short and not constant in number.
The nervous system has retained throughout its primitive connexion with the
epidermis. In almost all animals the nervous system is formed from the epiblast or
outermost cellular layer of the embryo; it usually, however, breaks away from this
and sinks into the body. Thus in Sipunculus it lies within the body-cavity, and has
retained its primitive connexion with the outer layers of the skin only in the region of
the brain; but in the Priapulids the nervous system, which consists of a ring round
the mouth and of a ventral cord, lies embedded in the skin, and the nerve cells are
directly continuous with the cells of the epidermis. The nerve-ring lies at the base of
a groove in the skin, which forms a kind of gutter round the mouth; the ventral
nerve-cord is visible exteriorly as a light line which marks the ventral surface of the
animal. In no place is the ring or cord differentiated in any way, and there cannot be
said to be any brain or special sense-organs. Numerous nerves are given off from
the ring to the pharynx and intestine, and from the cord to the body-wall.
The sexes are distinct, but they differ from the other Gephyrea in the nature of their
reproductive organs. In mature specimens the ovaries or testes are easily
recognisable, lying to the right and left of the alimentary canal. The reproductive
glands are continuous with ducts, which act as oviducts and vasa deferentia
respectively. Both glands and ducts are attached to the body-wall by a mesentery.
Nothing is known of the embryology of either member of this family, but both genera
appear to be sexually mature from the end of May until October.
Priapulus.—The body is continued into one or two caudal appendages, beset with
hollow papillae; these are ventral to the anus. The introvert forms ¼ to ⅓ of the total
body-length; it is covered with spines in conspicuous longitudinal rows, the rest of
the body being ringed. The retractor muscles are numerous, and are attached to the
body-wall, some anteriorly and some posteriorly.
P. caudatus Lam. (Fig. 218). Hab. Coasts of Greenland, Norway, Great Britain,
the North Sea, and the Baltic.