A simplified key to common genera of terrestrial earthworms

A simplified key to common

genera of terrestrial earthworms

I. Clitellum commencing in front of the 15th segment (not found in the British Isles)

- Clitellum commencing after the 15th segment

2. Setal arrangement perichaetine (Fig. 2)

- Setal arrangement lumbricine

3. Male pores on segment 17 or in 17/18 Spermathecae posterior to segment 10

Male pores on segment 18 Spermathecae anterior to segment 10

Male pores on segment 19 (semi-aquatic)

4. Prostomium tanylobous (Fig. I), setae closely paired (Fig. 2) at least over part of the body

- Prostomium epilobous (Fig. I), or iftanylobous, setae widely paired or distant over the whole body

2

4 (LUMBRI

CIDAE)

Pheretima (MEGASCo

LECIDAE)

3

Eudrilius (EUDRI

LIDAE)

Diplocardia (ACANTHo

DRILIDAE)

Sparganophilus (SPARGANo

PHILIDAE)

Lumbricus

5

246 BIOLOGY OF EARTHWORMS

5. Clitellum ends after segment 28 6 - Clitellum ends before segment 28, body section

quadrangular Eiseniella 6. Tubercula pubertatis absent, or exceptionally, present

as simple thickenings of the edges of the c1itellum Bimastos - Tubercula pubertatis present as ridges or isolated

papillae (Plate 5b) 7 7. Setae widely-paired or distant, at least posteriorly 8

- Setae closely-paired throughout the length of the body 9 8. Setae widely paired or distant throughout the length

of the body, tubercula pubertatis as ridges (exceptionally as separate tubercles) extending over only part of the length of the c1itellum Dendrobaena

Setae closely-paired anteriorly (in region of the hearts), distant posteriorly. Tubercula pubertatis

as ridges as long or'longer than the c1itellum Octo/asion 9. Spermathecal pores (Fig. 3) in line WIth seta 'd' or

more often near the mid-dorsal line. Body trapezoidal in cross-section.

Spermathecal pores situated laterally between setae 'c' and 'd' or 'a' and 'b' and 'c' and 'd'. Body not trapezoidal in cross-section

10. Prostomium with longitudinal ridges - Prostomium without longitudinal ridges I I. Calciferous glands with two lateral pouches in seg

ment. Terrestrial

Calciferous glands without lateral pouches. Am-

Eisenia and Eisenoides

10

Eophila II

Allolobophora

phibious Helodrilus 'Note. Both Sparganophilus and Helodrilus species live in mud bordering streams, rivers and lakes, which should provide confirmation of identification if the key does not provide sufficient data.'

Simple key to species of terrestrial earthworms

'The first part of the following key describes species belonging to families other than the Lumbricidae. The great majority of the representatives of these families are to be found as indigenous and peregrine species throughout tropical and subtropical parts of Africa, South America and Asia. However, certain members of these families are to

SIMPLIFIED KEY TO COMMON GENERA 247 be found in Europe, and particularly )J"orth America, usually as peregrine species (but occasionally indigenous, such as Diplocardia spp.). The commoner species found in North America are listed.'

MEGASCOLECIDAE Genus Pheretima One pair of spermathecal pores in S/6. 1st dorsal pore in 11/12-13/14. Clitellum on 14-16. 20-S6 mm. 8S-97 segs. Colour unpigmented, white. Two pairs of spermathecal pores in segmental grooves 7/8 and 8/9. 1st dorsal pore rr/I2. 70-170 mm. IO-ISO segs. Reddishbrown, clitellum creamy to dark grey. Two pairs of spermathecal pores (small) in S/6, 6/7. 1st dorsal pore in 10/1 I. Clitellum on segments 14-16, often not fully cover.ing segments 14 and 16.40-150 mm. 75-102 segs. Colour (usually dorsal only) yellow, brownish, brownish-red, grey. Three pairs of spermathecal pores (small) in 5/6-7/8. 1st dorsal pore in 10/11. Clitellum on !I4-!I6. 25-175 mm. 75-95 segs. Colour greyish-brown with violet irridescence, reddish-brown, yellow. Three pairs of spermathecal pores on the anterior edges of segments 7, 8 and 9. 150-220 mm. Light green/greenish buff with purple green dorsal line, clitellum milky or chocolate, pale grey ventrally. Four pairs of spermathecal pores (small) in 5/6-8/9. 1st dorsal pore in rr/I2 or I2/13. Clitellum on 14-16 not quite reaching 16/17.45-145 mm. 80--100 segs. Colour reddish-brown, greyish, very dark brown to black, sometimes bluish at mid-dorsal line. Four pairs of spermathecal pores (minute) in segmental grooves 5/6-8/9. 1st dorsal pore usually in I I/I2. 49-95 mm. 80-1 IS segs. Clitellum on 14-16. Colour, dorsal surface only (except for first few segments), reddish purple, reddish brown, yellowish-brown, chocolate, greyish.

ACANTHODRILIDAE Genus Dlplocardw

I. Clitellum forms a complete ring around the body. 40--120 mm. 90-120 segs. Anterior dorsal surface pale flesh coloured.

Clitcllum not a complete ring, but saddle-shaped. 2. Three pairs of spermathecal pores in segmental grooves 6/7,

7/8 and 8/9. 180-300 mm. I25-160 segs. Anterior dorsal surface pale flesh coloured.

Two pairs of spermathecal pores in segmental grooves 7/8 and 8/9. 200-270 mm. 135-160 segs. Aflterior dorsal surface dark brown.

Pheretima mlntma (Horst) P. calt/ornica (Kinberg)

P. momsi (Beddard)

P. hawayana (Rosa)

P. hupienm (Michaelsen)

P. rodericencis (Grube)

P. dtffrmgens (Baird)

DiplocardIa singulctris (Ude)

D. communis (Garman)

D. riparza (Smith)


EUDRILIDAE Dorsal pores lacking. Intersegmental furrows visible on c1itel- Eudri/us lum. 90-185 mm. 140-21 I segs. Colour, dorsal surface only, red. eugeniae

(Kinberg)

SP ARGANOPHILIDAE Dorsal pores lacking. Prostomium zygolobous (Fig. I). Anus Spargano-dorsal. phi/us eiseni

(Smith) The species of the Family Lumbricidae tabulated here are those which are found most commonly in Britain, North America and in Europe. Amongst these species will be found those that have been transported to various other parts of the world. British and American species are indicated. Although misidentification may arise on the basis of the characters in the first three columns if species not in the key are found, reference to the additional descriptions given in the fourth column should usually be sufficient to avoid confusion.

LUMBRICIDAE Genus Lumbricus

Clitellum 26,27-32

28-33

31, 32-37

Tubercula [st dorsal pubertatis

28-3 1

pore 7/8

6/7

7/8

5/6

t North American species.

Red/brown or red/violet, irridescent dorsally, pale yellow ventrally. 25-105 mm. 95-120 segs. Chestnut to violet brown; brown/yellow ventrally, irridescent, c1itellum orange. 30-70 mm. 82-100 segs. Setae widely-paired both ends of the body, strongly pigmented, brown-red dorsally, yellowish ventrally. 90-300 mm. 110-160 segs. Red-brown, lighter ventrally, irridescent dorsally, prominent c1itellum. Not found in large numbers. 48-108 mm. 100-143 segs.

.. British species.

Lumbricus rubellus·t Hoffmeister

L. castaneus. t (Savigny)

L. terres(ris. t Linnaeus

.t L. festivus (Savigny)

SIMPLIFIED KEY TO COMMON GENERA 249

Tubercula I st dorsal Clttellum pubertatis pore

Genus Eiseniella 22, 23-26, 23-2S, 26 27

"

Genus Bimastos 20, 21, Absent 22-29,30

22-29

23-28

23,24, 2S-3 I , 32

24, 2S-32, 33

Absent

24,2S 26-30

or absent

Absent

Absent

Absent

None

4/S

"

S/6

Male pores in 13. Dark brown, greenish, golden yellow, red. Body quadrangular behind the c1itellum. 30-60 mm. 60-90 segs. Male pore in IS. Otherwise as f. typica

Setae ab > cd. Redbrown. 105-1 I S segs. s0--80 mm.

Reddish-brown, segs 30-60. 20-S0 mm.

Eiseniella tetraeda f typica* (Savigny)

E. tetraeda (Savigny) f hercynia*

(Michaelsen)

Bimastos gieselerit (Smith)

B. tumidi/ Eisen

S/6, 6/7 20-S0 mm. 40--60 segs. B. palustrist up to 7S mm. (Moore)

S/6 Reddish dorsally, yel- B. parvust

lowish ventrally. 2S-40 (Eisen)

5/6

S/6

S/6

mm. 90--110 segs.

Rose-red. 98-122 segs. 60--<)0 mm.

Prostomium tanylobous, body cylindrical, reddish or violet dorsally, yellowish ventrally, c1itel-lum red. 30-64 mm. 7S-I I I segs.

B. longicinctus t (Smith & Gittins)

B. eiseni* (Levinsen)

Pale red to chestnut B. zeteki t brown, often localized (Smith & whitish banding anter- Gittins) iorly. 100--140 mm. 110-140 segs.

Dark reddish brown to B. heim-b ·t brown setae closely urgen

paired. (Smith)


Clitellum Tubercula pubertatis

Genus Dendrobaena 25, 26-31, 28-30

32

25, 26, 27- 28, 29-30, 30, 31, 31 or 32, 33 absent

25,26-28, 29,30

26, 27-31,

32

24,25-26, 27-32, 33

28-33, 34

30 and 31

31-32, 33

I st dorsal pore

5/6 Rosy to deep red, last Dendrobaena posterior segments yel~ rubida low, 27-90 mm. 50-100 (Savigny)* segs. .r subrubi

cunda (Eisen)

Setae widely paired, dor- D. rubida sally red-brown with (Savigny)* bluish tint. Intersegmen- .r tenuis tal grooves and ventral (Eisen) side light. 20-85 mm. 90-120 segs.

5/6 or Smoky-grey with red D. platyura sometimes pigment posteriorly. 80- (Fitzinger)

8/9 180 mm. 120-160 segs.

5/6 Dark red dorsally, lighter D. rubida*t red ventrally. 30--60 mm. (Savigny)

5/6

4/5

5/6

4/5

50-100 segs.

Violet, purple or olive D. veneta brown, dorsal pigment .r typica* bands separated by non- (Rosa) pigmented zones. Some-times uniformly unpig-mented. 50-155 mm. 80--225 segs.

Red, violet, yellow or D. copper. Posterior octa- octaedra*t gonal. 17-40 mm. 79-95 (Savigny) segs.

Dorsally pale red, 1st D. attemsl segment and ventrally (Michaeland clitellum white. 20- sen) 50 mm. 100--150 segs.

Red-violet, slightly irri- D. descent. 30-65 mm. 83- mammalis*t 100 segs. (Savigny)

Undetect- Red dorsally or unpig- D. pygmaea* able mented. 30-32 mm. 103- (Savigny)

180 segs.


Clztellum Tubercula pubertatls

Genus Octolasium

Genus Eisenia 24, 25, 28~30, 31

26--J2

27, 28~32,

33,34

Genus Eisenoides

30-32

24, 25~30, 27~39

31

Genus Eophila 33, 34 35, 36,

Genus Allolobophora 25, 26~31, 29~30, 31 32, 33 or 30~32

1St dorsal pore

11/12

8/9, 9/10 or 101 II

4/5

5/6

5/6

4/5

4/5

Blue-grey with (usually) lilac-blue dorsal line. Last 4~5 segments yellow, anterior segments pink, clitellum redorange. 50~I60 mm. IOO~ ~I50 segs.

White, grey, blue or rose-pink, clitellum pink or orange. 30~I60 mm. 90~I80 segs.

Red, purple or brown; yellowish ventrally. Dorsal surface pigment alternating with light intersegmental zones. 32~I30 mm. 80~I 10 segs. Light to bright reddish, mostly on dorsal aspect.

Octolasion cyaneum*t (Savigny)

O. lacteum*t (Oerley)

Eisenia !oetida*t (Savigny)

E. hortensis* t (Michaelsen)

Reddish, body cylindri- Eisenoides cal. 95~I20 mm. I03~I58 carolinsegs. Setae closely paired. ensls t

(Michaelsen)

Pigmented brown never E. usually blood shows lonnbergit 90~I75 mm. 100~152 (Michael-segs. sen)

Yellowish or grey. 52~ Eophila 140 mm. 132~I70 segs. lcterica*

(Savigny)

Pale red, without pig- Alloloment, prominent and bophora flattened clitellum. 25~ rosea*t 85 mm. 120~150 segs. (Savigny)


Tubercula ISt dorsal Clitellum pubertatis pore

26-32 28, t29 5/6 Grey, unpigmented, A. culpifera and t29, 30 usually blood shows (Tetry)

through epidermis. 20-60 mm. 80-120 segs.

25,26-33 30 and 31 4/5 Whitish grey, unpig- A. antipae mented. 5D-90 mm. 100- (Michael-130 segs. sen)

26,27, 31 and 33 rr/12 Anterior, especially the A. caliginosa 28-34,35 or 12/13 first few segments pink, f typica*t

otherwise pale grey, yel- (Savigny) lowish clitellum. 40-100 mm. 120-150 segs.

26,27 31-33 Colour as for f. typica. A. caliginosa 28-34, 35 Tubercula pubertatis of (Savigny)

two raised tubercles, f tra-connected by a narrow d t pezOl es bridge. (Duges)

27-32,33 Absent Indistinct Grey, unpigmented. 80- A. miniscula 100 segs. 22-25 mm. (Rosa)

27, 28-35 32 and 34 Body cylindrical, pale A. terrestris grey, unpigmented. 90- (Savigny) 150 mm. 160-200 segs.

27, 28-35 32-34 Body cylindrical, colour A. longa*t as for A. terrestris. Ude 90- 150 mm. 171- 181 segs.

27, 28-35 31 and 33 Segments posterior to A. nocturna* (extending 13 divided by two Evans

into grooves into three rings segment 32) Dark reddish brown, cli-

tellum paler. 90-180 mm. 200-250 segs.

27,28-34, 31-35 10/II 55-100 mm. 152- 194 A. tubercu-35 II/I2 segs. Body cylindrical, culata* t

or 12/13 unpigmented, greyish. Eisen

28-35, 36 33 and 34 4/5 or 5/6 Unpigmented, anterior A.limicola*t pink, the rest of the body Michaelsen pinkish grey. Bulbous anterior. 40- 100 mm. 86-146 segs.


Tubercula IS! dorsal Clitellum Puber!atis hore

31 and 33 and 35

Genus Helodrilus 21, 22-32 29-30

Nomenclature

Allolobophora caliginosa

4/5

4/5

Light or dark green, yel- A. chlorlow, grey, pink, slate- otica·t

blue, c1itellum pink, (Savigny) green or grey, 30-70 mm. 80-138 segs.

Setae black in fully Helodrilus mature individuals, flesh- oculatus· coloured, body unpig- Hoffmeister mented. 35-75 mm. 95-ISO segs.

Cernosvitov and Evans (1947) described Allolobophora caliginosa as two varieties, f. typica Savigny and f. trapezoides Duges, depending upon whether the papillae of the tubercula pubertatis were distinct, or joined by a narrow translucent bridge. Cain (1955) considered that all British specimens should be called A. caliginosa Savigny, and the name A. trapezoides Duges be given to continental specimens which, apart from the narrow bridge, have other characters which separate them from A. caliginosa, such as differences in the number of segments, position of the first dorsal pore and genital markings, and extension of the glandular tumescences around the male pores. Although Omodeo (1956) called the continental worms A. caliginosa f. trapezoides, Gerard in his synopses of the British Lumbricidae (1964) agreed with Cain, and listed only A. caliginosa Savigny, with no varieties. Gates (1972) also recognized A. trapezoides as a separate species. At the same time, Gates


postulated that the specific name cafiginosa for the British species is incorrect, and on the evidence of original identification by Eisen of a group of Scandinavian earthworms, and identifications by Friend (1923) and Davies (1960) states that the correct name should be turgida. Gerard disagrees on the grounds of priority. Nevertheless, Allolobophora turgida Eisen is used frequently, especially by American authors in place of Allolobophora cafiginosa Savigny. No doubt, in the next few years, this dichotomy will be resolved.

Octolasion lacteum Oerley

Very close indeed in relationship to O. cyaneum, almost indistinguishable except for a one segment difference in position of clitellum and tubercula pubertatis, it has been suggested (Omodeo, 1959) that they are diverse forms of the same agamous species, i.e. parthenogenetic morphs. The name given to this 'species', lacteum in 1881 is in common use, but Savigny's Enterion tyrtaeum 1826 was recognized by Cernosvitov as the same species (as lacteum) although he merely left it as a foot-note instead of claiming priority for tyrtaeum over lacteum. Gates (1972) has reinvestigated the situation, and states that the name of this taxon must be Octolasion tyrtaeum Eisen. Both tyrtaeum and lacteum are currently used in literature.

Because of the many changes in taxonomic nomenclature that have taken place during recent years, many authors continue to use names which have since been superseded, or which are still well-established. A list of such synonyms is given below.

Eisenia rosea Savigny = Allolobophora rose a Savigny.

Allolobophora minima Muldal = Allolobophora muldali Omodeo.

Bimastos beddardi Michaelsen = Bimastos parvus Eisen.

Eisenia veneta. Rosa = Dendrobaena veneta f. typica Rosa.

Blmastos tenuls Eisen = Dendrobaena rubida Savigny f. tenuis Eisen.

Allolobophora icterica Savigny = Eophlla ieterica Hoffmeister.

Eophila oculata Hoffmeister = Helodrilus oculatus Hoffmeister.

Dendrobaena subrubicunda Eisen = Dendrobaena rubida Savigny f. Sub-rubicunda Eisen.

Genl,ls Octo/asion oerley = Octo/asium oerley.

SIMPLIFIED KEY TO COMMON GENERA 255 The following species, otherwise placed in the genus Allolobophora

Eisen, will also be found (especially in current North American publications) in the proposed new genus Aporrectodia Gates: lzmlcola, longa, moebii, rosell, trapezoides, tllberculata, tllrgida.

Glossary

Acanthodriline. Having the male pores in segment 18 and the prostatic pores in segments I7 and 19.

Aclitellate. Without a clitellum. Not necessarily confined to juvenile stages.

Amphimitic. Sexual reproduction in earthworms involving two parents. Anthropochorous. Transported by the agency of man (usually unin

tentional). Astomate. The condition in which the nephridia are closed, i.e. without

a nephrostome. Autotomy. The process of self induced loss of segments - as when a

worm is gripped by the tail. Brown bodies. Rounded bodies containing tissue debris such as setae

and corpuscles, and foreign material. Chaeta, chaetae. Alternative for seta, setae. Classical system. The classification of the Oligochaeta originated by

Michaelsen in 1900, and developed by Stephenson in 1930 in 'The o ligochaeta' .

Clitellate. Possessing a clitellum. Also used to describe the stage in a worm's growth during which it has a clitellum, when it is a more specific term than 'adult'.

Clitellum. A region of epidermal thickening, containing gland cells which secrete the cocoon material.

Copulatory chamber. An invagination of the body wall into the coelom which contains the male pore.

Copulatory pouches. An old term for spermathecae. Copulatory setae. Setae in the same segment as, and near to, the

spermathecae. Diapause. As applied to earthworms, a non-active state during which

the worm has an empty gut and stays tightly coiled in a mucus lined


cell within the soil to protect itself against adverse environmental conditions. It may be optional or obligatory.

Endemic. Indigenous. Epilobous. The condition when the continuation of the prostomium q. v.

backwards does not reach the first segmental groove. Estivation. A summer diapause.

Genital tumescences. Raised areas of the epidermis from which genital setae grow, found in the Lumbricidae.

Haemerophilic. Not adversely affected by human interference with the environment.

Haemerophobic. Adversely affected by human interference with the environment.

Heteromorphic. The condition when regeneration occurs so that a head regenerates instead of a tail, and vice versa.

Holandric. Testes restricted to segments 10 and II, or a homeotic equivalent.

Hologynous. Ovaries restricted to segments 12 and 13, or a homeotic equivalent.

Homeotic. The state in which an organ or series of organs are in a segment or segments in which they are not normally found.

Intersegmental furrow. The actual boundary between two adjacent segments.

Intersegmental groove. The visible annular depression that indicates externally the position of the boundary between two adjacent segments.

Juvenile. The term used for earthworms from the time of emergence from the cocoon to when the first indications of maturity, such as genital tumescences, seminal grooves and genital pores appear.

Lumbricine setal arrangement. Having four pairs of setae per segment.

Macroic. Large. A substitute for meganephridial. Male ducts. Male gonoducts. Sperm ducts q.v. Male funnels. Funnel shaped internal end of sperm duct. Male pores. Exterior openings of the sperm ducts. Meganephridia. Large nephridia. A term now little used, the condition

meganephridial now being encompassed by the term holonephric. Micronephridia. Small nephridia, usually numerous, often encompassed

by the term meronephric. Meronephridia. Divided nephridia, individual tubules often present in

large numbers. Can be either large - megameronephridia, or small -micro meronephridia.

GLOSSARY 259

Megascolecoid. Referring to worms placed III the classical family Megascolecidae.

Oviducts. Female gonoducts.

Parthenogenesis. Reproduction in which the ova develops without being fertilized by a spermatozoa, hence it only involves one parent.

Peregrine. Foreign, not indigenous. The agency is almost always transportation by man.

Perichaetine setal arrangement. Many setae arranged in a ring right round each segment with only a break in the mid-dorsal and midventral regions.

Peristomium. The most anterior segment of an earthworm surrounding the mouth, and which bears the prostomium.

Phylogeny, Phylogenetic. Pertaining to past evolutionary development as opposed to individual development (autogeny).

Polyandric. Having testes in more than segments 10-1 I.

Polymorphic. Pertaining to polymorphism. Polymorphs anslllg from parthenogenesis and geographical distribution are considered important in systematics.

Proandry, Proandric. Testes restricted to segment 10 or a homeotic equivalent.

Progyny, Progynous. Ovaries restricted to segment 12, or a homoeotic equivalent.

Prolobous. The condition when there is no continuation of the prostomium backwards into the peristomium.

Prostate. Glands associated with the male ducts, usually tubular, opening beside the male pores in acanthodrilid, octochaetid and ocnerodrilid worms.

Prostomium. A lobe-like appendage attached to the front of the dorsal aspect of the peristomium.

Racemose. Applied to prostates when those organs are divided into many lobes (cf. a bunch of grapes).

Seminal grooves. Permanent longitudinal grooves on the ventral surface of an earthworm associated with the male and prostatic pores which form a passage for seminal and other fluids.

Seminal vesicles. Septal pockets in which spermatozoa mature. Seta. Setae. Stiff bristle-like structures protruding from follicles through

the epidermis. Usually sigmoid in shape, except specialized setae. May be enlarged at the extreme portions of the body.

Somatic. Pertaining to any part of the body except the genital organs. Sperm ducts. Ducts carrying sperm to the bodies exterior.


Stomate. Pertaining to an 'open' nephridium, i.e. a nephridium with a funnel opening (usually) to the exterior.

Tanylobous. The condition when the continuation of the prostomium reaches backwards to the first segmental groove.

Taxon. Any unit in a system of classification. Testis sac. Closed part of the coelom containing the testes and male

funnels of a particular segment. Zygolobous. Condition when the prostomium as seen from above, is not

differentiated from the peristomium by any superficial markings.

References

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Systematic index

(Italic page numbers indicate the more important references in the text whereas bold numbers refer to illustrations.)

Acanthodrilacae, 44 Acanthodrilidae, 44, 46,51-2, 61, 65, 66,

67, 74, 75, 122, 245, 247 Acanthodrilinae, 43, 44, 45, 46 Acanthodrilini, 45 Acanthodrilus, 41, 51 Actinomyces, 187 advenltlum, Blpalium, 147 Aeolosomatidae, 13, 4 I, 42 aerophilus, Thomznx, 227 affinis, Dichogaster, 80 africanus, Hyperiodrilus, 75, 136, 142, 143,

ISO, 154, 157 afrlcanus, Ocnerodrilus, 226 Agastrodilus, 52 agglutinatus, Bacdlus, 185 agItans, Uropoda, 149 agrestis, Pheretlma, 63, 107 Aikinetocystzs, 148 alexandrt, Pheretlma, 79 Allolobophora, 59, 69, 81, 82, 127,228,246,

250, 251, 256 antipae, 30, 252 caligznosa, 26, 68, 70, 71, 73, 80, 83, 87,

98, 102, 117, 125, 126, 128, 129, 130, 13 1, 132, 133, 134, 135, 136, 138, 139, 145, 146, 149, ISO, IS I, 152, 153, 154, 157, 158, 161, 162, 163, 164, 165, 167, 172, 174, 176, 179, I!!9, 197, 19/5, 200, 201, 216, 226, 227, 229, 232, 239, 240, 241, 255

caligznosa f. tYPlca, 63, 252, 255 caligznosa f. trapezoides, 26, 63, 80, 252,

255 chlorolica, 62, 68, 70, 71, 72, 73, 80, 83,

93, I I I, II8, 125, 126, 128, 129, 130, 13 1,133,134,135,136,145,146,148, ISO, 151, 152, 153, 154, ISS, 161, 164, 170,200,216,219,229,231,232,239, 240, 241, 242, 253

culpifera, 252

dubiosa, 226 japonica, 137 limlcola, 62, 241, 252 longa, 62, 69, 70, 71, 73, 80, 84, 93, 108,

II I, 128, 130, 131, 138, 140, 145, 146, 150, 153, 154, 161, 162, 165, 172, 173, 192, 193, 199,216,224,227,231,232, 234, 240, 2 ... , 252

minzma, 256 mlnzscula, 252 muldall, 62, 256 noclurna, 69, 70,84,87,125,126, 12M, 129,

130, 131, 132, 134, 136, 13M, 140, 145, 149, ISO, 151, 161, 162,232,234,240, 241, 252

robusl a, 146 rosea, 9, 63, 70, 71, 73, 81, 83, 125, 126,

127, 128, 129, 130, 131, 132, 133, 134, 138, 145, 146, 150, 151, 152, 157, 161, 165, 216, 231, 232, 239, 240, 251, 256

terrestns, 146, 252 terrestris f. longa, 73 trapezoldes, 62, 255 tuberculata, 62, 80, 93, 241, 252 turglda, 62, 80, 255

Alluroididae, 43 Alma, 57, 75, 157, 158 Amoebotaenza cuneatus, 226

lumbri, i, 226 Andiorrhinus, 56 AndlOswlex, 56 annulata, Capil/arza, 226 anomala, Mil/soma, 74, 125, 136, 138, 152 anomala, Pheretima, 79 Anoplophyra, 148 antipae, Al/olobophora, 30, 252 apricaria, Pluvla/zs, 147 Aporrectodza, 256

/zmlcola, 256 longa, 256 moebll, 256


Aporrectodia--cont.

rosea, 256 trapezoides, 256 tuberculata, 256 turgida, 256

Aptodrilus, 56 Ascaridia gallt, 228 Ascaris suum, 228 Aspidodrtlus, 146 altemst, Dendrobaena, 170, 239, 250 austrabs, Megascolides, 19, 81 avestcula, U dema, 79 Azotobacter, 175, 183

Bactllus agglutinatus, 185 botulinus, 148 cereus, 182, 185 cereus var. mycoides, 184 tdosus, 185

Balanteodrtlus, 5 I Barogaster, 52 Bassia longifolia, 120 beatrix, Octochaetana, 80 beddardi, Btmastos, 62, 150, 256 Beddardiella, 54 Benhamia, 52 Benhaminae, 44 bermudensis, Pontodrilus, 61, 79 Bettonta, 54 btcmta, Pheretima, 63 Bimastos, 39, 59, 64, 246, 249

beddardt, 62, ISO, 256 elsem, 13, 127, 128, 145, 149, ISO, 159,

163, 165, 170, 240, 241, 245, 249 gieseleri, 62, 249 heimburgeri, 62, 249 tcenorum, 241 longicinctus, 62, 249 lonnbergi, 149, 150 muldali, 241 palustris, 62, 249 parvus, 20, 62, 93, 241, 249, 256 tenms, 8, 256 tumidts, 62, 82, 85, 249 weicht, 62 zetek!, 62, 72, 74, 75, 85, 86, 88, 89, 145,

249 Bipalium adventttum, 147

kewensts, 143, 147 bishamban, Ramiella, 80 Borgert/a, 54 botulmus, Bactllus, 148 Branchiobdellidae, 42 bolam, Dtchogaster, 63, 80 bronch/alis, Cyathostoma, 226, 227 Buttnmodrilus, 54

Caleb/ella, 52

(!Iiz/ormco, Pherer;ma, 22, 25, 1,3, 102, 154, 157, 158, 247

caligmosa, Allolobophora, 26, 68, 70, 71, 73, 80,83,87,98, 102, "7, 125, 126, 128, 129, 130, 131, 132, 133, 134, 135, 136, 138,139,145,146,149, ISO, 151, 152, 153, 154, 157, 158, 161, 162, 163, 164, 165, 167, 172, 174, 176, 179, 189, 197, 198, 200, 201, 216, 226, 227, 229, 232, 239, 240, 241, 255

caltginosa f. trapezOldes, Allolobophora, 26, 63, 80, 252, 255

caltgmosa f. ryptca, Allolobophora, 63, 252, 255

Calltdnlus, 57 campanulata, Pherettma, 79 canus, Larus, 147 Captllaria annulata, 226

causmflata, 227 mucronata, 227

plica, 227

putom, 227 carnana, Sarcophaga, 149 carolinensis, Eisenoides, 251 castaneus, Lumbricus, 38, 62, 69, 70, 71, 73,

80,82,93,98, ,,8, 126, 127, 128, 145, 146, 150, 161, 163, 164, 165, 172, 232, 241, 242, 248

causinflata, Captllana, 227 Causus rhombeatus, 147 Celenella, 52 cereus var. mycoldes, BaCIllus, 184 Cestoda, 225, 226 Chi/ota, 51 chlorot/ca, Allolobophora, 62, 68, 70, 71, 72,

7~80,83,93, III, 118, 125, 12~ 12~ 129, 130, 13 1, 133, 134, 135, 136, 145, 146, 148, 150, 151, 152, 153, 154, ISS, 161, 164, 170, 200, 216, 219, 229, 231, 232,239,24°, 241, 242, 253

ChunlOdnlus, 54 Clitellata, 40 Clostrtdium, 183 coeltcolor, Streptomyces, 187 Comarodnlus, 49 commUniS, Diplocardia, 247 commumsstma, Pherettma, 93 controversa, Tilletta, 188, 224 coracana, Eleusme, 228 corethrurus, Pontoscolex, 61, 66, 79, 228 Corvus corone, 147 crassum, Porrocaecum, 227

CraterocysttS, 148 Criodrilidae, 47,58, 61

SYSTEMATIC INDEX 30 3 Criodrilinae, 43 CrlOdrilus, 58, 114 Criodrilus lacuum, 61, 75, 226 culpifera, Allolobophora, 252 cuneatus, Amoebotaenia, 226 cuneata, Ta,nia, 148 Curgiona, 50 cyaneum, Octolasion, 62, 70, 82, 128, 129,

138,145,150,151,161,165,216,232, 239, 240, 241, 25 1, 255

Cyathostoma bronchialis, 226, 227

Decachaetus, 51 Deccania, 50 dejecta, Pheretima, 79 Deinodrilus, 67 Dendrobaena, 14, 59, 81, 84, 246, 250

attemsi, 170, 239, 250 mammalis, 62, 6cj, 70, 146, 232, 241, 250 octaedra, 62, 80, 98, 127, 128, 13 1, 145,

146, 149, 150, 159, 163, 165, 170,240, 241, 250

platyura, 250 pygmaea, 241, 250 rub,da, 62, 80, 82,127,131,145,146,149,

150, 165, 170, 232, 239, 240, 250, 256 rubida f. tenuis, 81, 241, 250, 256 rubida f. subrubicunda, 146, 241, 250, 256 subrubicunda, 19, 70, 71, 73, 82, 86, III,

150, 153, 154, 192,232, 240, 256 veneta var. hibernica f. dendroidea, 241 veneta var. hibernica f. typica, 146, 241 veneta var. hortensis, 241 veneta f. tYP,ca, 241, 250, 256 veneta var. zebra, 241

Desmogaster, 48 Diaphrodrilus, 50 Dicelis, 148 D,chogaster, 47, 52, 95

bolaui, 63 jaculatrtx, 142

sa liens, 63 Dichogastidae, 45 Didymogaster, 49

sylvatlcus, 19 dijJringens, Pheretima, 63, So, 122, 247 D,gaster, 49 Dilepus undula, 226 DtnodrtlOldes, 51 Dinodrilus, 52

Dioctophyma, sp., 227 Dionyx, 148 DIplocardia, 51, 61, 64, 13 1, 157, 245

communis, 247 egglestont, 75, 84

jiortdana, 122

mlsslssipptensis, 89, 122 rtparta, 140, 247 slngularis, 61, 247

Diplocardiinae, 43 Dzplotrema, 41, 51 Dzporochaetti, 51

Distichopus, 148 Dorgiodrilus, 50 Drawida, 48, 66

grandis, 140, 146 nagana, 8o papillifer, 80 rara, 80

Drzlocrzus, 57, 66 dubzus, Microscolex, 61, 79 dubiosa, Allolobophora, 226 dunguensis, Schubotziella, 78

Echinocystis, 148 egglestoni, Diplocardia, 75, 84 Eisenia, 63, 64, 69, 246, 251

foetida, 13, 17, 19,62,71,73,78, So, 81, 82, 85, 92, 93, 95, 99, 100, 108, 115, 116, 147, 148, 149, 153, 155, 156, 157, 158, 159, 172, 186, 188, 189, 196,200, 201,226,227, 228, 229, 232, 239, 240, 241,25 1

hortenslS, 62, 25 I lonnbergz, 251

rosea, 62, 63, 72, 73, So, 98, 140, 158, 256

veneta, 226, 256 zebra, 62

elseni, Bimastos, 13, 127, 128, 145, 149, 150, 159, 163, 165, 170,240,241,245, 249

eiseni, Lumbricus, 62 eiseni, S parganophilus, 63, 154, 248 Eiseniella, 59,81,246, 249

tetraedra, 62, 80, 165, 232, 240, 241, 249 tetraedra f. hercynza, 241, 249 tetraedra f. tYP,ca, 165, 248, 249

Elsenoides, 59, 64, 246, 250 caro/inensis, 25 I

Eleuszne coracana, 228 elongata, Pheretzma, 79, 228 elongatus, Metastrongylus, ;u6 Eminoscolex, 54

emissartus, Haplotaxis, 62 Enantiodrilus, 56 Enchytraeidae, 8, II, 42, 146, 209 ensicaudatum, Porrocaecum, 227 EnterlOn tyrtaeum, 256 Eodrilus, 51, 245


Eophda, 60, 64, 66, 246, 250 (Allolobophora) icterica, 150, 241, 251,

256 Eophila oculata, 256 E phymodrdus, 55 encetorum, Turdus, 147 Erithacus rubecula, 147 Eschmchia colt, 184 Estherella, 56 Eudlchogaster, 53 Eudmodrdoides, 5 I Eudrilidae, 42, 43, 46,53-4,61,66,75,78,

79, 83, 245, 248 Eudrilinae, 43, 46,54-5 Eudriloides, 54 Eudnlus, 55, 88

eugemae, 61, 66, 142, 248 eugeniae, Eudrilus, 61, 66, 142, 248 Eukerria, 50

hortensis, 79 saltensls, 63 subandina, 79

Eupolygaster, 48 Eupolytoreutus, 55 europaea, Talpa, 147 europeus, Evinaceus, 147 Euscolex, 55 Eutoreutus, 55 Eutrigaster, 53 Eutyphoeus, 53, 95

waltoni, 78, 142 Evinaceus europeus, 147 excavatus, Penonyx, 80, 95 eXlgus, Pheretlma, 79 Enus, 45, 49

jestil'us, Lumbncus, 62, 82,127,165,232,241, 248

Fimoscolex, 56 Fletcherodrtlus, 49 flondana, Diplocardia, 122

jiJetida, Eisenia, 13, 17, 19, 62, 71, 73, 78, 80, 81, 82, 85, 92, 93, 95, 99, 100, 108, 115, 116, 147, 148, 149, 153, 155, 156, 157, 158, 159, 172, 186, 188, 189, 196, 200, 201, 226, 227, 228, 229, 232, 239, 240,241, 251

forbesi, Haplotaxls, 62

Fraxinus pennsylvanica, 198 Fndericia parasttica, 146 friendl, Lumbricus, 241 Fusarium, 188, 224

galli, Ascaridia, 228

Gardullaria, 55

Gatesla, 50 gieseleri, Bimastos, 62, 249 glabra, Pheretima, 79 Glossoscolecidae, xiv, 4, 23, 42, 43, 47,

55-6, 61, 66, 75 Glossoscolecinae, 8, 36, 43 Glossoscolex, 56 Glyphidrilus, 57 gordiodes, Haplotaxis, 62 GordlOdrilus, 50 gracilus, Pontodrilus, 61 grandls, Drawida, 140, 146 Grayallia, 148 Gregarma, 148

haemorrhoidalis, Sarcophaga, 149 Haasema, 55 haltotidea, Testacella, 147 Haplodrilus, 50 Haplotaxidae, 43, 61, 62 Haplotaxis, emiSSarlUS, 62

forbesi, 62 gordlOdes, 62 ichthyophagous, 62

Hastirogaster, 48 hawayana, Pheretlma, 63, 80, 247 heimburgerl, Bimastos, 62, 249 Helodrilus, 55, 60, 64, 66, 246

afrlcanus, 150 oculatus, 241, 253, 256

herculeus, Lumbricus, 227

Heterakis, 148 Heterodera rostochlenSls, 224

Heteroporodnlis, 49 hdgendorfi, Pheretima, 63, 72, 79 Hlppopera, 55

nzgenae, 181 Hirudinea, 40, 41 Hlstomona murchiel, 148 Histomonas, 226

H oloscolex, 56 Hoplttophyra, 148 Hoplochaetella, 53 Hoplochaetina, 53 Hormogaster, 58

redli f. gigantes, 65 Hormogastridae, 58, 65 Hormogastrinae, 43 hortensis, Elsenia, 62, 251

Eukerria, 79 houlleti, Pheretlma, 63 Howascolecini, 45 Howascolex, 53 hupelensis, Pheretlma, 24, 63, 73, 74, 75,

127, 131, 138, 140, 142, 153, 157, 158

SYSTEMATIC INDEX 305

Hypmodrilus, 55, 79 afncanus, 75, 136, 142, 143, 150, 154,

157 Hyslrlchis triwlor, 226

Icenorum, Bimastos, 241

Ichthyophagous, Haplotaxis, 62 icterica, Eophila (Allolobophora), ISO, 241,

251, 256 idosus, Bactllus, 185 Ilyogema, 50 maequalis, Ventura, 188 mfantalis, Pheretima, 79 Iridodri/us, 55

jacu/atrix, Dichogaster, 142

Japomca, Allolobophora, 137

KafJania, 55 Kerriona, 50 kewensis, Bipalium, 143, 147 Kynotus, 57

/acteum, Octo/aston, 82, 98, 128, 146, 153, 165, 183,228,229,232,241,251,255, 256

/acuum, Cnodri/us, 61, 75, 226 Lampito, 49, 80, 95

mani/ii, 80 Larus canus, 147 latus, Protoscolex, 41 Legonea,55 Lennogaster, 53 Lennoscolex, 49

pumila, 80 Leptodrtlus, 5 I, 54 Leucodrilus, 53 /evIS, Pherellma, 63 Libyodrilus, 54 /imicola, Allolobophora, 62, 241, 252 Liodrtits, 50 /onga, Allolobophora, 62, 69, 71, 73, 80, 84, 93,

108, II I, 128, 130, 131, 138, 140, 145, 146, 150, 153, 154, 161, 162, 165, 172, 173, 192, 193, 199,216,224,227,231, 232, 234, 240, 241, 252

longa, Aporrectod,a, 256 /ongicinctus, Bimastos, 62, 249 longifolta, Bassta, 120 /onnbergi, Blmastos, 149, 150 liinnbergi, Eisenoides, 62, 251 lot'endgel, Pherellma, 63 lumbnci, All%bophora, 226 /umbncl, Polycercus, 226 Lumbricidae, 8, 10, I I, 12, 13, 20, 35, 36,

42, 43, 58, 60, 61, 64, 172, 240, 241, 245, 246, 248, 255

Lumbriculidae, 2, 5, 6, 22, 30, 39, 40, 42, 47, 80, 98, 225

Lumbricus, 4, 14, 17, 19, 22, 25, 26, 30, 33, 34, 36, 37, 60, 62, 81, 83, 84, 85, 110, 139, 245, 248

castaneus, 38, 62, 69, 70, 71, 73, So, 82, 93, 98, 118, 126, 127, 128, 145, 146, 150, 161, 163, 164, 165, 172,241,242, 248

eiseni, 62 festivus, 62, 82, 127, 165, 232, 241, 248 friendi, 241

hercu/eus, 227 rubellus, 62, 69, 70, 71, 73, 80, 82, 93, 98,

108, III, 127, 128, 145, 146, 150, 154, 155, 162, 165, 170, 172, 173, 183, 192, 196, 226, 227, 229, 232, 239, 240, 241, 248

terrestris,6, 13,21,27,39,62,72,76, So, 81, 82, 83, 84, 85, 86, 88, 89, 90, 91, 92, 93, 94, 97, 98, 103, 106, 107, 108, 114, 115, 117, 125, 126, 128, 129, 130, 13 1, 132, 134, 136, 138, 139, 140, 145, 146, 148, 149, 150, 151, 152, 153, 154, 156, 157, 160, 161, 162, 166, 168, 169, 172, 173, 177, 178, 179, 182, 183, 184, 185, 188, 192, 195, 196, 198, 199, 200, 201,206,216,218, 21 9,226,227,228, 229,23°,23 1,232,234,235,236,237, 239, 240, 241, 242, 248

Mahema, 50

Malabarta, 50 Malabarini, 45 Malodrtlus, 55 mamma lis, Dendrobaena, 62, 69, 70, 146,

232, 241, 250 mamlti, Lampito, 80 Maortdrtlus, 5 I marcessens, Serratia, 184 mariana, Pieea, 198 mauget, Testacella, 147 Maupasella, 148 mays, Zea, 228

Megachaetma, 54 Megadrili, xv, 42, 43, 61, 65, 66, 79 Megascolecidae, 8, 10, 13, 19, 20, 22, 23,

25, 30, 32, 36, 38, 39, 43, 44, 45, 46, 47, 48-50 , 62, 66, 80, 146, 147, 245, 247

Megascolecinae, 7, 43, 44, 45 Megasco/ex, 7, 30, 41, 49, 149 Megascoltdes, 13, 53, 61, 199

australis, 19, 81


Meles meles, 147 meles, Meles, 147 merid,ana, Pheretima, 79 merula, Turdus, 147 merulae, Syngamus, 227 mesomeltnus, Microsaurus, 147 Metadrilus, 54 Metaradiophyra, 148 Metascolex, 55 Metastrongylus, 148, 225

elongatus, 226 pudentotectus, 226 salamI, 226

Microchaetidae, 47, 57, 66, 223 Microchaetinae, 43 Microchaetus, 57, 142 Microdrili, 42, 48, 82 Microsaurus mesomelinus, 147 MicrosCIJlex, 51

dublUS, 61, 79 modestus, 228 phosphoreus, 61

millardi, Perionyx, 140 Millsonta, 53

anomala, 74, 125, 136, 138, 152 minima, Allolobophora, 256 mtnima, Pheretima, 63, 247 mtniscula, Allolobophora, 252 mississippiensls, DiplocardIa, 89, 122 modestus, Mlcrosco/ex, 228 moebli, Aporrectodia, 256 Monil(gaster, 48 Moniligastridae, 41, 42, 43, 46, 47-8, 66,

67, 80, 146, 147 Moniligastrinae, 42, 43 Monocystls, 19, 148 Monogaster, 53 morrisl, Pheretima, 63, 80, 247 mucronata, Capillaria, 227 muldali, Allolobophora, 62, 256 muldali, Bimastos, 241 murchiei, HistlOstoma, 148 Myxocystis, 148

nagana, Drawida, 80 Naididae, 2, 41, 42 N annodrilts, 50 Nellocystis, 148 Nellogaster, 49 Nelloscolex, 49 Nemantodrilus, 54 NematocystlS, 148 Nematoda, 226 Nematogenia, 50 Neochaeta, 49 N eodrilacae, 44, 52

Neodrtlus, 52 Neogaster, 53 Neummaneona, 55 ntgerzae, Hippopera, 181 NocardIa polychromogens, 187 nocturna, Allolobophora, 69, 70, 84, 87, 125,

126, 128, 129, 130, 131, 132, 134, 136, 138, 140, 145, 149, 150, 151, 161, 162, 232, 234, 240, 241, 252

Notoscolex, 49, 142 termlticola, 147

Notykus, 54

oCCldentalts, Ocnerodrilus, 63, 80 Ocnerodrilinae, 43, 44, 45, 46 Ocnerodrilini, 45 Ocnerodrilus, 5 I

afrtcanus, 226 occidentalis, 63, 80

octaedra, Dendrobaena, 62, 80, 98, 127, 128, 13 1, 145, 146, 149, 150, 159, 163, 165, 170, 240, 241, 250

Octochaetidae, 44, 46,52-3, 63, 66, 67, 80 Octochaetinae, 43, 45, 46 Octochaetini, 45 Octochaetoides, 53 Octochaetona beatrtx, 80

surensts, 80 OClochaelus, 53 Octo/aslon, 60, 69, 166, 173, 219, 246, 251

cyaneum, 62, 70, 82, 128, 129, 138, 145, 150, 151, 161, 165, 216, 232, 239, 240, 241, 251, 255

lacteum, 82, 98, 128, 146, 153, 165, 183, 228,229,232,241,25 1,255,256

tyrtaeum, 62, 256 oculala, Eophtla, 256 oculatus, Helodrtlis, 241, 253, 256 Oligochaeta, 22, 40, 41, 42 Omodeona, 53 Onesla sepulchralis, 149

subalpina, 149 Onychochaeta, 56 Opisthodrilus, 56

palustrls, Blmastos, 62, 249 paptl/ifer, Drawida, 80 papllliJer, Plutellus, 61, 79 Parabursaria, 148 Parachilota, 52 paradoxa, Paricterotaenia, 226 Parascolex, 55 parasltlca, Frtdericla, 146 Parendrilinae, 43,54 Parendrtlus, 54 Partcterotaenia paradoxa, 226


pan'us, Blmastos, 20, 62, 93, 241, 249, 256 Paultstus, 51 peguana, Pheretima, 226 Pellogaster, 53 pennsyivanica, Fraxmus, 198 Penodrtlus, 52 Perionychini, 45 Perionyx, 7, 49, 95, 114

excavatus, 80, 95 mtllardl, 140

Penscolex, 56 Perzssogaster, 49 Phalans, 164 Pheretlma, 7, 8, 18,21,23,25,30,32,36,37,

38, 39, 47, 49, 66, 67, 73, 78, 81, 88, 100, 101, 107, 110, 137, 148, 184,226, 229, 245, 247

agrestls, 63, 107 alexandn, 79 anomala, 79 blcmta, 63 calyarn/ca, 22, 25, 63, 102, 154, 157,

158, 247 campanulata, 79 (ommuntssima, 93 defecta, 79 dlfjrmgens, 63, 80, 122, 247 e!ongata, 79, 228 eXlgus, 79 glabra, 79 hawayana, 63, 80, 247 htlgendorji, 63, 72, 79 houlletl, 63 hupelensls, 24, 63, 73, 74, 75,127, I31, 138,

140, 142, 153, 157, 158, 247 mfanttllS, 79 levIS, 63 lovertdgel, 63 mendlana, 79 minIma, 63, 247 morrisl, 63, 80, 247 peguana, 226 posthuma, 39, 63, 140

rimosa, 79, 80 rodertCencls, 247 stellert, 8 vanans, 79 Virgo, 79 voeltzkowl, 79 zoyslae, 79

Pheodrilidae, 42 phosphoreus, Mlcroscolex, 61 Picea manana, 198 pltca, Captllaria, 227 Plaglochaeta, 52 Plaglotoma, 148

Platydnlus, 54 Platyhelminthes, 148 platyura, Dendrobaena, 250 Pleuroij'stlS, 148 Pliconogaster, 50 Pliogaster, 50 Plutellus, 41, 52, 61, 65

paptlltfer, 61, 79 umbellulartae, 61, 79

Pluvlaits apricarta, 147 Pollenia rud,s, 149 Polycercus, 148

lumbrict, 226 Polychaeta, 40, 41 polychromogenes, Nocardia, 187 Polytoreutus, 55 Pontodnlus, 52

bermudenSlS, 61, 79 gracilis, 61

Pontoscolex, 56, 66 corethrurus, 61, 66, 79, 228

Porrocaecum crassum, 227 enslcaudatum, 227

posthuma, Pheretlma, 39, 63, 140 Priodochaeta, 53 PrlOdoscolex, 4, 53 Pronaldites, 41 Protoscolex, 41

latus, 41 Protozoa, 98, 99, 148, 188, 226 pudentotectus, Metastrongylus, 226 pumila, Lennoscolex, 80 putorti, Caplllana, 227 pygmaea, Dendrobaena, 241, 250 Pygmaeodrilus, 5 I Pythium, 188, 224

Quechuona, 5 I Quedius (Mlcrosaurus) mesomeiinus, 147 Quercus robur, 198

Ramiella, 53 blshambari, 80

Ramzellona, 53 rara, Drawlda, 80 redii f. gigantes, Hormogaster, 65 RhabditIs, 148 Rhmodrtlus, 56 Rhododrilus, 52 rhombeatus, Causus, 147 Rhynocystls, 148 Rillogaster, 53 nmosa, Pheretlma, 79, 80 nparta, DIplocardIa, 140, 247 robur, Quercus, 198 robusta, Allolobophora, 146


roder/eenets, Pherettma, 247 Rosadrilus, 55 rosea, Al/olobop'hora, 9, 62, 63, 70, 73, 81,

83, 125, 126, 127, 128, 129, 130, 131, 132, 165

rosea, Elsema, 62,' 63, 72, 73, 80, 98, 140, 158, 256

rostoehiensis, Heterodera, 224 rubecula, Erithacus, 147 rubel/us, Lumbricus, 62, ~, 70, 71, 73, 80,

82, 93, 98, 108, I I I, 127, 128, 145, I~I~I~I~I~I~I~I~ 173, 183, 192, 196, 226, 227, 229, 232, 239, 240, 241, 248

rublda, Dendrobaena, 62, 80, 82, 127, 131, 145, 146, 149, 150, 165, 17°,232, 239, 240, 250, 256

rubida f. subrubicunda, Dendrobaena, 146, 241, 250, 256

rubida f. tenuis, Dendrobaena, 81, 241, 250, 256

rudts, Pol/mia, 149

salami, Metastrongylus, 226 saitens, Dichogaster, 63 saltensis, Eukerria, 63 Sareophaga earnaria, 149

haemorrhoidalis, 149 strrata, 149

Sehubotztella, 55 dunguensts, 78

Seolail/us, 54 Seolioscolides, 53 scutulum, Testacella, 147 sepulchralis, Onesia, 149 Serratia'marcessens, 184 singularis, Diplocardia, 61, 247 skrJabmomorpha, Syngamus, 227 smlthii, Sparganophilus, 63 sonomae, Sparganophllus, 63 Sparganophilidae, 47,57, 63, 65, 245, 248 Sparganophilinae, 43 Sparganophilus, 57, 245, 246

elSent, 63, 154, 248 smithii, 63 sonomae, 63

S penceriel/a, 53 S phaeractinomyzon, 148 Spirochaeta, 148 Spiroptera, 148

turdi, 227 stellerr, Pheretlma, 8 Stephanurus, 148, 227 striata, Sarcophaga, 149 Streptomyces coellcolor, 187 Sturmus vulgaris, 147

subalpina, Onesra, 149 subandina, Eukerria, 79 subrubicunda, Dendrobaena, 19, 70, 71, 73,

82, 85, III, 150, 153, 154, 192, 232, 240, 256

subvirrd,s, Trocheta, 147 surensis, Oetoehaetona, 80 suum, Asmris, 228 sylvaticus, Didymogaster, 19 Sylvodrilus, 52 Syngamus, 148

merulae, 227 skrjabinomorpha, 227 trachea, 225, 227

Syngenodrilinae, 43 synoecnema, 148

Taenia euneata, 148 Talpa europaea, 147 Tazelaarra, 5 I Teleudrilus, 55 tenuis, Bimastos, 8, 256 termiticola, Notoscolex, 147 terrestris, Al/olobophora, 146, 252 terrestris f. longa, Al/olobophora, 73 trapezoides, Allolobophora, 62, 255

f. trapezoldes, caligmosa, Allolobophora, 26, 63, 80, 252, 255

turgida, A71olobophora, 62, 80, 255 f. (ypica, caliginosa, Allolobophora, 63,

252, 255 terrestns, Lumbricus, 6, 13, 21, 27, 39, 62,

72, 76, 80, 81, 82, 83, 84, 85, 86, 88, 89, 90, 91, 92, 93, 94, 97, 98, 103, 106, 107, 108, I II, 114, 115, 117, 125, 126, 128, 129, 130, 131, 132, 134, 136, 138, 139, 140, 145, 146, 148, 149, 150, 151, 152, 153, 154, 156, 157, 160, 161, 162, 166, 168, I~, 172, 173, 177, 178, 179, 182, 183, 184, 185, 188, 192, 195, 196, 198, 199, 200, 201, 206, 216, 218, 219,226,227,228,229,230,23 1, 232, 234, 235, 236, 237, 239, 240, 241,

242, 248 Testacella hallOlidea, 147

mauger, 147 scutulum, 147

tetraedra, Etseniella, 62, 80, 165, 232, 240, 241, 249

f. hercynia, Eiseniella, 241, 249 f. typrea, Elseniel/a, 165, 241, 249

Thamnodrrloldes, 56 Thamnodrilus, 56 Thatonta, 51 Teleuloreutus, 55 Thelohanta, 148


Thommx, aerophtlus, 227 Til/etla controversa, 188, 224 Tonoscolex, 50, 53 trachea, Syngamus, 225, 227 tncolor, Hystrichus, 226 Trtgaster, 53 Trocheta subvtrldts, 147 tuberculata, Allolobophora, 62, 80, 93, 241,

252

Tubificidae, 42 tumtdis, Bimastos, 62, 82, 85, 249 turdi, Sptroptera, 227 Turdus ericetorum, 147 Turdus merula, 147 turgida, Allolobophora, 62, 80, 255 tyrtaeum, Enterion, 256 tyrtaeum. OctolastOn, 62, 256

Udema, 52 avesicula, 79

undula, Dilepus, 226 Uropoda agitans, 149

vartans, Pheretima, 79 veneta, Eisenza, 226, 256

veneta f. typtca, Dendrobaena, 241, 250, 256 reneta var. hortensts, Dendrobaena, 241 veneta var. htberntca f. dendrotdea, Dendro-

baena, 241 veneta var. htbernica f. typlca, Dendrobaena,

146, 241 veneta var. zebra, Dendrobaena, 241 Ventura inaequalis, 188 Virgo, Pherettma, 79 voeltzkowi, Pheretima, 79 vulgaris, Sturmus, 147

waltoni, Eutyphoeus, 78, 142 Wegenerzella, 53 Wegenenona, 53 welcht, Bimastos, 62 Woodwardtella, 50

Yagansta, 52

Zapotecta, 52 Zea mays, 228 zebra, Eisenta, 62 zeteki, Bimastos, 62, 72, 74, 75, 85, 86, 88,

89, 145,249 zoyslae, Pheretima, 79

Author index

(Italic page numbers indicate the more important references in the text whereas bold numbers refer to illustrations.)

Abrahamsen, G., 123, 146, 261 Agarwal, G. W., 228, 261 Aichberger R. von., 184, 261 Aisyazhnyuk, A. A., 261 Aldag, R., 196, 261 Alicata, J. E., 294 Allee, W. c., 149, 261 Allen, R. W., 261 Almeida, A. M., 97 Anderson, H. L., 282 Anstett, M., 261 Arbit, ]., 93, 261 Aristotle, xv Arldt, T., 41, 261 Arlidge, G. Z., 214, 280 Arnold, M. K., 214, 269, 276 Arrhenius, 0., 149, 261 Arthur, D. R., 99, Il3, 209, 210, 241, 262 Arthur, J. H., 271 Ash, J. S., 266 Aspock, H., 215, 281 Atlavinyte, 0., xvii, 174, 183, 197, 230,

262 Avei, M., 262 Axeisson, B., 209, 262

Bacheiier, G., xvii, 149, 163, 210 Bahl, K. N., 18, 73, 102, 140, 262 Baker, W. L., 262 Bakhtin, P. U., 192, 193, 263 Baldwin, F. M., 90, 94, 263 Ball, D. F., 289 Ball, R. c., 263 Baluev, V. K., 196, 263 Banage, W. B., 123, 264 Barker, R. ]., 213, 215, 263 Barley, K. P., 118, 124, 138, 164, 165, 167,

172, 173, 174, 176, 179, 189, 197, 205, 206, 228, 229, 263

Barrett, T. j., xvii, 199, 263 Bassalik, K., 182, 184, 190, 192, 263

Bather, E. A., 41, 263 Bauer, K., 212, 213, 263 Baweja, K. D., 188, 224, 263 Baydonabiciene, Z., 262 Baylis, H. A., 142, 146, 263 Beauge, A., 144, 263 Bejsovec, J., 228, 264 Beiousova, N. K., 124, 26Q Bengston, S.-A., 147, 264 Benham, W. B., 13, 29, 41, 264 Beynon, K., 270 Bharucha-Reid, R. P., 93, 264 Bhatt,]. V., 174, 184, 188,224,280 Bigger, J. H., 213, 264 B1ancke, E., 175, 190, 264 Blankwaardt, H. F. H., 212, 264 Block, W., 123, 264 Bocock, K. L., 264 Bodenheimer, F. S., 149, 194, 264 Boice, R., 94, 140, 292 Bollen, W. B., 181, 289 Bornebusch, C. H., 118, 124, 149, 264 Bouche, M. B., xvii, 122, 264 Boyd, ]. M., 163, 264, 265 Boykins, E. A., 217 Boynton, D., 159, 265 Bradley, ]., 192, 196, 266 Brain, R. c., 273 Braunig, G., 98, 107, 284 Bray, J. R., 173, 265 Bretnall, G. H., 265 Bretscher, K., 118, 265 Breza, M., 226, 265 Brinkhurst, R. 0., 61, 265 Brown, B. R., 166, 265 Brown, D. M., 265 Bruel, W. E. van der, 187, 265 Briisewitz, G., 184, 185, 186, 265 Buahin, G. K., 211, 265 Budaviciene, I., 262 Buntley, C. J., 265

AUTHOR INDEX 3 I I

Byzova, Y. 8., 97, 98, 213, 265, 274

Cain, A. J., 265 Capstick, C. K., 264 Carter, G. 5., 1I6, 266 Causey, D., 226 Cernosvitov, L., 53, 60, 146, 255, 266 Chadwick, L. c., 192, 196, 266 Chandler, R. F., 120, 269 Chapman, G., 266 Chen, C. M., 222, 266 Christensen, R. E., 274 Cialdini, R. 8., 290 Clebsch, E. E. G., 290 Cockerell, T. D. A., 149, 266 Cohen, S., 103, 266 Coin, C. ]., 266 Combault, A., 26, 196, 266, 291 Compton, O. c., 159, 265 Conder, P. J., 215, 266 Cori, C. J., 227 Cossens, G. G., 194, 197, 201, 295 Couperus, H., 97, 268 Couture, G., 219, 283 Cragg,]. B., 266 Cramp,S., 215, 266 Crompton, E., 210, 267 Crossley, D. A., 166, 173,219,267, 270 Curry, L. L., 263 Czerwinski, Z., r81, 267

DaCiulyte, Y. A., 230, 262 Darwin, c., xii, xvii, 90, 108, 144, 165,

228, 267 Datta, L. G., 94, 267 Davey, S. P., 212, 267 Davies, H., 267 Davis, 8. N. K., 215, 217, 267 Dawson, A. 8., 35, 267 Dawson, R. c., 184, 192, 193, 267 Day, G. M., 174, 185, 267 Decker, G. c., 213, 217, 264, 283 Dennis, E. 8., 212, 213, 269 Devigne, J., 267 Dhawan, C. L., 267 Dhennin, L., 225, 267 Dingwall, A. R., 128, 197,276 Dixon, R. M., 194, 288 Djuffill, R. A. Jr., 297 Doane, C. c., 213, 215 Dobson, R. M., 199, 268 Doeksen, J., xvii, 84, 95, 97, 119, 151,268,

29 1

Doerell, E. c., 210, 268 Dotterweich, H., 151, 268 Douglas, D. E., xvii, 199, 272

Douhalei, N., 281 Dowdy, W. W., 131, 157, 269 Dreidax, L., 144, 196, 269 Drift, ]. van der, xvii, 212, 214, 264, 268,

269, 291 Dubash, P. J., xvi, 77, 78, I16, 296 Duley, F. L., 296 Dustman, E. H., 215, 269 Dutt, A. K., 192, 193, 201, 269 Dzangaliev, A. D., 124, 269

Eaton, T. H. Jr., 120, 269 Eberhardt, A. I., 149, 269 Edwards, C. A., 1I9, 120, 124, 151, 178,

179, 199,2°3,206,2°9, 21I, 212, 213, 214,215,217,218,222,237,265,267, 269, 270, 276

Ehlers, W., 206, 270 EI-Duweini, A. K., 22, 102, 118, 125, 150,

152, 154, 156, 157, 158, 161, 270 Ellenby, c., 224, 270 Ellis, D. ]., 283 Empson, D. W., 269 Escherich, K., 147, 271 Escrit, J. R., 209, 210, 211, 271 Evans, A. c., xvii, 60, 62, 68, 69, 71, 72,

81, 82, 84, 85, 87, 120, 125, 131, 132, 133, 134, 136, 139, 140, 141, 144, 145, 1~lnl~I~I~I~I~I~ 172, 190, 194,203,205,229,231,232, 255, 266,27 1

Feldkamp. ]., 5, 27' Fenton, G. R., 271 Finck, A., 151, 181, 192, 271 Fleming, W. E., 213, 271, 272 Ford, J., 1I8, 272 Fox, C. ]. S., 212, 272 Franz, H., 173, 272 Fraser, C. H. T., 93, 272 French, M. c., 217, 267 Fujita, 0., 93, 278

Gaddie, R. E. Sr., xvii, 199, 272 Gansen, P. S. van, 99, 100, 272 Garner, M. R., 139, 239, 272 Gast, J., 165, 272 Gates, G. E., xiii, 41, 43, 44, 46, 50, 51,

52, 57, 58, 61, 62, 63, 64, 65, 66, 79, 80, 81, 82, 85, 95, 115, 136, 142, 255, 256, 272, 273

Gavrilov, K., 62, 63, 79, 273, 279 Genov, T., 226, 273 Geoghegan, M. ]., 193, 273


Gerard, B. M., xvii, 68, 72, 84, 126, 129, 130, 133, 135, 136, 152, 153, 155,255, 273, 274

Gersch, M., 230, 274 Gewehr, H., 293 Ghabbour, S. I., 102, 118, 125, 144, 150,

152, 154, 156, 157, 158, 161, 187,27°, 274

Ghilarov, M. S., xvii, 184, 195, 200, 213, 274

Ghoca, M. L., 290 Giesecke, F., 175, 190, 264 Gilbert, 0., 264 Gish, CD., 216, 217, 218, 220, 274 Gissel-Neilson, G., 287 Glasgow, L. L., 218, 295 Goffart, H., 213, 214, 274 Gordon, W. T., 278 Gorham, E., 173, 265 Graff, 0., xvii, 72, 155, 157, 176, 181, 196,

203, 205, 239, 25 1, 274, 275 Grant, W. C, 24, 74, 127, 131, 138, 140,

152, 153, 157, 181, 275 Grassi, B., 226, 275 Gray,]., 113, 114, 275 Greenwood, D. E., 213, 275 Griffiths, D. C, 214, 275 Grigor'eva, T. G., 213, 275 Grove, A. ]., xvii, 15, 16, 18, 28, 29, 32,

37, 275 Guild, W. J., xvii, 62, 68, 69, 71, 72, 81,

82, 84, 87, 120, 125, 127, 131, 132, 133, 134, 136, 141, 144, 145, 150, 153, 154, 155, 162, 164, 165, 172, 173, 192, 194, 203, 205, 229, 231, 232, 27 1, 275

Gunthart, E., 2I)', 276 Gurianova, O. Z., 192, 276

Hadley, C H., 213, 271 Hamblyn, C J., 127, 128, 197, 276 Handa, B. K., 267 Handley, W. R. C, 271 Hanel, E., 90, 276 Hanley, I. M., 265, 272 Hardman, ]. A., 154, 215, 216, 217, 218,

298 Harman, W. ]., 276 Harmsen, G., 189, 276 Harrison, R. B., 215, 267 Hasenbein, G., 230, 276 Haswell, W. A., 148, 276 Heath, G. W., 124, 166, 168, 169, 170, 176,

209, 237, 269, 270, 276, 280 Heck, L. von, 92, 93, 276 Heimburger, H. V., 276 Hensen, V., xvi, 276

Hess, W. N., 31, 33, 35, 107, 276 Heungens, A., 170, 213, 214, 215, 277 Hill, ]. P., 148, 276 Hirst, ]. M., 188, 277 Hobmaier, A., 226, 277 Hoeksema, K. ]., 192, 277 Hoffman, ]. A., 188, 224, 277 Hogben, L., 157, 277 Hogg, T. W., 230 Hollister, P. L., 261 Hook, R. J. van, 230, 277 Hopkins, A. R., 213, 214, 277 Hopkins, H. T., 192, 203, 204, 206, 207,

212, 277 Hopp, H., !I 8, 133, 136, 156, 192, 194,

196, 199, 203, 204, 206, 207, 208, 277, 295

Howell, CD., 94, 107, 277, 278 Hoy, H. M., 213, 278 Hubl, H., 106, 277 Huhta, V., 209, 278 Hunt, L. B., 215, 217, 278 Hutchinson, S. A., 185, 224, 278 Hyche, L. L., 214, 278 Hyman, L. H., 85, 278

Inoue, T., 278 Ireland, M. P., 220, 221, 278 Isa, A. L., 282 Iwahara, S., 93, 278

Jacks, G. V., 94, 278 Jacob, A., 209, 210, 279 Jacobson, G. M., 176, 180, 181, 201, 283 Jakubczyk, H., 267 Jamieson, B. G. M., 45, 46, 61, 265, 279 Janda, V., 62, 279 Jeanson-Luusinang, C, 150, 185, 279 Jefferson, P., 120, 205, 210, 279 Jennings, A. C, 176, 189, 263 Jevniaux, C, 267 Joachim, A. W. R., 192, 279 Johnson, M. L., 98, 279 Johnstone-Wallace, D. B., 119, 208, 279 Jongerious, A., 277 Joshi, N. V., 190, 196, 279 Julin, E., 64, 279

Kahsnitz, H. G., 196, 279 Kalmus, H., 279 Kamel, M., 185, 188, 224, 278 Kanwar, ]. S., 181, '90, 192, 287 Karmanova, E. M., 226, 227, 279 Karpachevskii, L. 0., 288 Keilin, D. 280 Kelkar, B. V., 190, 196, 279

AUTHOR INDEX 313 Kelly, W. A., 291 Kelsey, j. M., 214, 280 Kevan, D. K. McE., xvii, 280 Khambata, S. R., 174, 184, 188,224, 280 King, H. G. c., 166, 169, 276, 280 Kirberger, c., 97, 280 Kirk, R. L., 157, 214, 277 Kirk, V. M., 277 Kleinig, C. R., 172, 179, 197, 263, 280, 287 Kleist, S. M., 290 Knop, J., 29, 280 Kobatake, M., 187,280 Kobayashi,S., 63, 280 Kollmannsperger, F., 133, 144, ISS, 163,

280 Kollmannsperger, G., 144, 280 Kondo, K., 222, 278 Korschelt, E., 72, 280 Kozlovskaya, L. 5., 183, 185, 280 Kring, ]. B., 214, 215, 281 Krivaneck, J. 0., 93, 280 Kriiger, F., 96, 124, 281 Kubiena, W. L., 163, 281 Kiihnelt, W., xvii, 163, 281 Kurcheva, G. F., 169, 281

Lan, H. van der, 281 Ladell, W. R. 5., 1I9, 281 Lahr, ]. P., 261 Lakhani, K. H., 121, 177, 281 Lal, R., 206, 281 Lauer, A. R., 93, 281 Lavelle, P., 74, 136, 138, 144, 153, 281 Laverack, M. S., xvii, 17,96, !O3, !O8, 166,

281, 282 Lawrence, R. D., 175, 282 Lee, K. E., xvii, 43, 46, 84, 195, 229, 282 Legg, D. c., 213, 215, 282 Leitenberger, L., 173, 272 Lesser, E. j., 98, 282 Lewis, H. B., 103, 175, 266 Lidgate, H. ]., 213, 282 Lindquist, B., 165, 175, 282 Lipa, J. ]., 213, 282 Lissman, H. W., 113, 114, 275 Liv, C. L., 222, 266 Ljungstrom, P.O., xvii, 61, 66, 79, 142,

147, 229, 282 Lofty, J. R., 120, 124, 151, 156, 160, 161,

199,203,205,206,209, 21I, 212, 214, 219, 268, 269, 270, 275, 282, 290

Lohm, u., 262 Long, W. H., 213, 282 Love, C. W., 265 Low, A. ]., 118, 192,283 Lowe, D. G., 293

Luckman, W. H., 213, 217, 283 Lugauskas, A., 183, 262 Lukose, J., 283 Lund, E. E., :zz6, 283 Lunt, H. A., 176, ISo, 181, 201, 283 Lyons, C. H., 214, 296

Madge, D. 5., xvii, 75, 83, 123, 142, 144, ISO, 154, 157, 173,283

Magalhaes, P. 5., 226, 283 Maldague, M., 219, 283 Mamajev, B. M., 200, 274 Mamytov, A., 192, 193, 283 Mangold, 0., 108, 165, 166, 283 Marapao, B. P., 106, 283 Marshall, V. G., 198, 283 Martin, A. W., 102, 283 Martin, ]. P., 193, 297 Matlu, N., 278 Mayne, D. W., 295 McCalla, T. M., 205, 296 McInnes, D. c., 297 McInroy, D., 229, 284 McLeod,]. H., 147, 284 Meer, K. van der, 277 Meggitt, F. J., 226, 284 Mellanby, K., 284 Mendes, E. G., 97, 284 Merker, E., 98, 107, 284 Meyer, L., 190, 284 Michaelsen, W., 40, 41, 42, 47, 61, 64,

284 Michon, ]., 72, 73, 74, 84, 86, 153, 155,

284 Miles, H. B., 188, 284, 285 Millar, H. R., xvi, 175, 281 Millott, N., 99, 285 Minderman, G., 268 Moment, G. B., 85, lIS, 285 Monnig, H. 0., 226, 285 Moore, A. R. 285 Moore, B., 109, 149, 285 Morris, H. M., 119, 205, 285 Morrison, F. 0., 213, 285 Mozgovoy, A. A., 227, 285 Muldal, 5., 62, 79, 285 Miiller, G., xvii, 285 Miiller, P. E., 165, 195, 285 Murchie, W. R., xvii, 61, 62, 71, 75, 84,

85, 86, 89, 127, 136, 145, 285, 286

Nakamura, Y., 137, 138, 149, 286, 296 Nathans, 5., 118, 124, 291 Needham, A. E., 103, 173, 175, 176, 177,

286 Negi, L. 5., 261


Nelson, J. M., 1I8, 286 Newell, G. E., xvii, 15, 16, 18, 28, 29, 32,

37, 1I2, 275, 286 Nicol, H., 210, 287 Nielson, C. 0., 196, 197, 199, 220, 287 Nielson, M. G., 287 Nielson, R. L., 286 Nijhawan, S. D., 181, 190, 192, 287 Nilsson, A., 264 Noble, J. c., 197, 202, 287 Nordstrom, S., 146, 156, 161,264, 287 Nowak, E., 267

O'Connor, F. B., 291 Ogg, W. G., 210, 287 Ogles, G. D., 149, 296 Oldham, c., 287 Oliver, J. H., 148, 287 Olson, H. W., 149, 152, 287 Omodeo, P., 43, 44, 46, 62, 65, 255, 287 Otanes, F. G., 222, 287

Panditesekera, D. G., 192, 279 Papedick, R. I., 265 Parker, G. H., 1I0, 288 Parle, J. N., 182, 183, 186, 187, 193, 288 Parshley, H. M., 110, 288 Patel, H. K., 213, 222, 288 Pathak, A. N., 190, 294 Peachey, J. E., 288 Pensson, T., 262 Peredel'sky, A. A., 218, 288 Perel, T. S., 170, 288 Peterson, A. E., 195, 288 Petrov, B. c., 149, 288 Phillips, E. F., 149, 288 Pickard, J. A., 213, 215, 217, 296 Piearce, T. G., 146, 151, 165, 288 Polivka, J. B., 21 I, 213, 214, 289 Polsky, M. N., 192, 193, 263 Pomerat, G. M., 97, 289 Ponomareva, S. I., 181, 182, 192, 193, 206,

289 Poryadkova, N. A., 288 Powers, W. L., 181, 289 Prabhoo, N. R., 289 Pratt, K. c., 85, 296 Prosser, C. L., 289 Puh, P. c., 151, 181, 289 Purdy, L. H., 188, 224, 277 Puttarudriah, M., 222, 228, 289

Raffy, A., 97, 289 Ragg, J. M., 289 Ralph, C. L., 97, 289 Ramsay, J. A., 101, 102, 110, I I I, 289

Rao, K. S. K., 261 Raw, F., 1I9, 120, 122, 124, 125, 159, 168,

169, 170, 173,21 1,214,221,275,289, 290

Reichle, D. E., 267, 270 Reinecke, A. ]., 142, 147, 157, 229, 282,

290 Ressler, R. H., 94, 290 Reynolds, J. W., xvii, 62, 63, 79, 122, 124,

147, 229, 290 Reynolds, W. M., 229, 290 Reynoldson, T. B., 1I8, 124, 290, 291 Rhee, J. A. van, xvii, 118,' 124, 126, 173,

196, 198, 200, 221, 290 Rhoades, W. c., 214, 291 Ribaudcourt, E., 196, 291 Richards, ]. G., 197, 291 Richardson, H. c., 150, 159,210,211,291 Richter, G., 213, 291 Robertson, J. D., 99, 291 Robinson, J. S., 92, 93, 292 Rodale, R., xvii, 210, 213, 292 Rodionova, L. Z., 288 Roots, B. I., 100, II 0, 152, 292 Rossenkoetter, ]. S., 94, 140, 292 Rovelli, G., 226, 275 Roy, S. K., 141, 144, 292 Rundgren, S., 131, 146, 156, 161, 264,

287, 292 Ruschmann, G., 175, 187, 292 Russell, E. J., xii, xvi, 159, 175, 196, 292 Rysav)', B., 226, 227, 292 Ryzhikov, K. M., 227, 292

Sacho, R. ]., 217, 278 Salisbury, E., xvi, 149, 151, 200, 292 Saroja, K., 292 Sastry, K. S. S., 222, 228, 289 Satchell, J. E., xi, xvii, 70, 72, 81, 84, 85,

86, 108, 109, 1I8, 120, 121, 123, 127, 128,129,149,150,151,155, 159,161, 163, 165, 166, 16g, 173, 175, 177, 178, 179, 180, 182, 188,205,275,281,286, 293

Saussey, M., 146, 195, 293 Scharpenseal, H. W., 293 Schmid, L. A., 106, 293 Schmidt, H., 92, 294 Schneider, K. c., 34, 294 Schread, J. c., 142, 213, 214, 223, 294 Schrevan, D. van, 189, 276 Schwartz, B., 226, 243, 294 Scopes, N. E. A., 214, 298 Scott, H. E., 214, 294 Scrickhande, J., 190, 294 Sharma, R. L., 267

AUTHOR INDEX 315

Shindo, B., 294 Shiraishi, K, 98, 159, 294 Sims, R. W., xvii, 43, 44, 45, 46, 294 Singh, A., 267 Sison, P. L., 222, 288 Skarbilovic, T. S., 227, 294 Slater, C. S., 194, 196, 199, 204, 210, 277,

294, 295 Smallwood, W. M., 33, 105, 295 Smith, F., 295 Smith, R. D., 218, 295 Sokolov, D. F., 288 Stafford, C. J., 270 Stephenson, J., xvi, xvii, 10, 12, 13, 16, 18,

26, 27, 36, 38, 40, 41, 42, 43, 44, 47, 64, 65, 81, 89, 98, IIO, II4, 148,229, 257, 295

Stickel, L. F., 215, 269, 295 Stickel, W. H., 295 Stock dill, S. M. J., 195, 197, 201, 295 Stockli, A., II8, 144, 175, 181, 182, 186,

191, 194, 295 Stokes, B. M., 295 Stolte, H. A., xvi, 148, 228, 295 Stone, P. c., 149, 296 Stop-Bowitz, c., 296 Storey, 1. F., 277 Stough, H. B., 34, 296 Stringer, A., 2 I 3, 214, 2 I 5, 217, 296 Sun, K. H., 85, 296 Svendsen, l A., II8, 122, 124, 127, 165,

296 Swaby, R. J., 192, 296 Swartz, R. D., 93, 296

Takano, S., 149, 296 Tembe, V. B., xvi, 77, 78, 117,296 Tenney, F. G., 189, 296 Teotia, S. P., 183, 190, 192, 193, 194,

205, 296 Thomson, A. J., 140, 297 Thompson, A. R., 213, 214, 215, 217, 270,

296 Tillinghast, E. K., 102, 103, 297 Tischler, W., 205, 297 Torvic, M. M., 261 Tracey, M. V., 99, 297 Trifonov, D., 222, 297 Tromba, F. G., 227, 297 Twinn, D. c., 264

Uhlen, G., 210, 297

Urquhart, A. T., xvi, 297

Vail, V. A., 82, 85, 89, 297 Villot, F. C. A., 226, 297 Vimmerstedt, l P., 202, 297 Vogel, R., 226, 297 Volz, P., 195, 297

Waid, J. S., 264 Waite, R. H., 297 Waksman, S. A., 189, 193, 296, 297 Walton, W. R., 119, 222, 298 Ward, W. c., 277 Watanabe, H., 73, 298 Waters, R. A. S., 133, 138, 139, 165, 197,

198, 205, 298 Watkin, B. R., 208, 298 Way, M. ]., 214, 298 Weber, G., 213, 214, 298 Wehr, E. E., 283 Weisbach, W. W., 229, 298 Went, J. c., 184, 185, 298 Westeringh, W. van def, 2 II, 298 Wheatley, G. A., 215, 216, 217, 218, 298 Wherry, E. T., 149, IS0, 298 White, G., xi Whiting, A. E., 270 Whitney, W. K, 214, 298 Wiegland, K., 209, 210, 279 Wilcke, D. E. von, 72, 73, 118, 298 Wilkinson, G. E., 195, 206, 298 Wilcox, H. G., 277 Wingerden, C. G. van, 84, 151, 268 Witkamp, M., 177, 299 Wittich, W., 108, 166, 177, 299 Wojewodin, A. W., 212, 299 Wolf, A. V., 110, 156,299 Wollny, E., 194, 196, 299 Woodhead, A. A., 227, 299 Woodman, M. G., 264 Wooton, R. l, 220, 278 Wright, M. A., 166,299

Yegorova, S. V., 288 Yerkes, R. M., 90, 92, 93, 299

Zajonc, 1., xvii, 124, 209, 299 Zarrow, M. T., 97, 289 Zhdannikova, E. N., 183, 185, 280 Zhinkin, L., II6, 299 Zicsi, A., xvii, 118, 122, 204, 222, 299 Zrazhevski, A.!., 183, 184, 198, 299

General index

(Italic page numbers indicate the more important references In the text whereas bold numbers refer to illustrations.)

Absorption zone, 100 Acclimation, 157 Accessory reproductive bodies, 8 Acetycholine, 106, 116 Acid intolerant species, 150

tolerant species, 149, 150 soils, 149, 150, 151, 197, 210, 234

reaction of worms to, 127, 243 Acids, 98, 108, 151, 175 Actinomycetes, 182, 183, 185, 186 Activity, 138, 145, 146, 154, 178

cycles, 93, 97, 106 effects of temperature on, 133, 155, 156 seasonal changes in, '3', 132, 134, 136

Adders, 147 Adolescence, 87 Adrenaline, 106 Adverse conditions, 70, 83, 225

effects on soil, 228-9 periods, 83

Aeration, 127, [59, [94-5, 205, 228 Aestivation, 83-4, 152 Afferent nephridial vessel, 18

typhlosolar vessel, 18 Africa, xv, 49, 50, 53, 54, 55, 56, 57, 58,

59, 60, 65, 66, 74, 125, 136, 142, 147, 152, 222, 229, 246

Age-class composition, u6 African night crawler, 66 Aggregates, 187, 190, 193, 194, 198, 201

formation of, [92-4 Agriculture, effects on earthworms, 128,

'45, 203-2[

Agricultural practice, 47 soils, 152, 228

Albumen cells, 14, 15 Alcohol, as a preservative, 233 Alder, 173 Aldicarb, 2 I 5 Aldrin, 212, 213, 214, 218 Alfalfa, 192, 206

Algeria, 58 Alimentary canal, I, 20-2, 23, 34, 37, 100,

148, 183, 184, 185 innervation of, 33, 34 plexus, 25

Alkaline soils, 234 Alkaloid substances, 108 Allantoin, 100, 175 Alluvium, population in, 162, 163 America, 55, 57, 74

Central, 5', 52, 56, 57, 65, 66, 67 North, xv, 52, 53, 59, 61, 64, 144, 205,

246, 247, 248 South, 50, 51, 52, 56, 57, 59,65,66,67,

246, 248 Ammonia, 100, 101, 102, 103, 152, 175,

176, 186 Ammonium, 176, 189

sulphate, 172, 211 Amoebocytes, 19, 101 Amphimitic reproduction, 79 Ampulla, 29 Amputation, 80 Amylase, 99, 142 Anabiosis, 110 Anaerobic conditions, 95, 116 Anaerobic respiration, 98 Anaesthesis, 113 Andes, 56 Antarctica, 67 Antibacterial extracts, 187 Antibiotic, 187 Anus, I, 20, 85, 95, 100, IIO, 248 Apertures, male and female, 9, 12

spermathecal, 12 Apple scab, 188, 2 I 5 Aquatic species, 9, I I, 43 Arable, 119, 122, 124, 133, 144, 145, 146,

156, 160, 161, 193,203,205,216,222, 233, 235

Arciform muscles, 76

GENERAL INDEX 317

Arctic forest, 173 Argentina, 51, 55, 56 Artesian irrigation, 206 Arthropods, 174, 182 Ascospores, 188 Asexual reproduction, 79, 80, 8 I Ash, 166, 177, 237 Asia, 49, 56, 59, 64, 67, 246 Asia Minor, 59, 64 Association neurones, 105 Atrazine, 2 I 2

Auckland,s I Australasia, 65, 67 Australia, 49, 50, 51, 53, 59, 60, 67, 124,

138, 159, 179, 202, 206 Austria, 58 Autotomy, 85, 89 Available mineral nutrients, 180-1

A voiding reaction, 151 Axon, 34 Azinphosmethyl, 94, 214 Azores, 59, 60

Bacteria, 19, 29, 98, 99, 148, 166, 174, 182, 183, 185, 186, 187, 188, 193

Bacterial gums, 194 Bacteriocide, 187 Bacteroidal cells, 101 Badgers, 147 Bahamas, 48, 50 Bahlia, 52 Bait, earthworms as, 229 Bardsey Island, 124 Barley, 196, 197 Basal cells, 14

membrane, 17, 33 Basalt, 190 Base exchange capacity, 180, 20 I Basement membrane, 16 Bassia tree, 120 Beans, 228, 237 Beech, 124, 166, 169, 170, 237 Beet, 166, 237 Behaviour, 88-95, 93, 94, 110 Behavioural experiments, 88, 243-4 Benomyl, 214 Benzopyrine, 230 Bermuda, 56 BHC, 213 Biocides, 2 I I

Biology xvii, 68-95 Biomass, xv, 123, 179 Birch, 166, 170, 177, 237 Birds, 147, 215, 220, 223, 225, 226, 227 Blackbirds, 147 Black Spruce, 198

Blackhead, 226 Bladder, 102, 103 Blood, 100, 101, 104, 110

plasma, 96 supply, subcutaneous, 27 system, 103 vessels, 14, 15, 18, 22, 26, 33, 116

Blue-green algae, 184 Body cavity, 9

fluids, 97, I I I

length of, 60 number of segments of, e 5 temperature of, 153 weight of, 110, 152, 173, 174

Body wall, 4, 14-17, 15, 19, 22, 33, 34, 99, 101, 103, 105, 110

nerves supplying, 105 Bolivia, 56 Borneo, 48, 66 Bowling greens, 223 Brazil, 50, 51, 52, 53, 56, 57 Breeding, 73, 199, 200

Breeding period, 12, 73, 125 Bristle worms, 40 Brown bodies, 19 Brown earth, 123 Bronchial dilation substance, 229 Buccal cavity, 17, 20, 23, 24, 33

innervation of, 35 Buccal chamber, 98, 108

epithelium, 17 Bulgaria, 222 Bullock droppings, as food, 72, 164, 231 Burma, 48, 49, 50, 57, 95, 142 Burrowing, 110, 138-45, 171, 172, 183,

190, 194 experiments, 237-9 speed of, 139

Burrows, 88, 89, 90, 91, I II, 119, 121, 122, 128, 139, 191,211,216,222,235,237, 243

depth of, 140 diameter of, 140 permanent, 129

Bursa propulsoria, 79 Butyric acid, 183

Caecae, of intestine, 22

Caecum, 24 Caesium, 219 Cage for studying earthworm activity, 238 Cages, 169 Calcareous sand, populations in, 163 Calciferous gland, 20, 22, 44, 45, 46, 50,

55, 57, 58, 59,99, 146, 151, 193,246 tissues, 20


Calcium, 99, 146, 165, 180, 220, 221 carbonate, 19, 99 humate, 193 metabolism, 146

California, Lower, 50, 51, 52, 65 Cambridge, 124 Cameroons, 53, 54, 55, 57, 144 Campbell Islands, 5 I Canada, 64, 140 Canary Islands, 59 Cape Verde Islands, 5 I Capillaries, 25, 26, 100, 104

intra-epidermal, 26 Capillary blood vessels, 14 Capillary water, 194 Carabid beetles, 147 Carbamate, fungicides, 2 I 4

insecticides, 2 I 5 Carbaryl, 215, 223 Carbofuran, 214, 215 Carbohydrates, soluble, 166 Carbon, combustion, 178, 179 Carbon nitrogen ratio, 177-80, 181, 202

dioxide, 96, 98, 159, 178 Carnivorous slugs, 147 Caroline Islands, 48 Carrots, 228 Casting, 138-45

in different habitats, 144 time of day of, 142

Castor meal, effect of, 160 Casts, 100, 131, 140, 142, 144, 145, 151,

154, 172, 176, 180, 181, 183, 184, 185, 186, 187, 188, 189, 190, 191, 192, 193, 194, 196, 201, 205, 206, 222, 223, 224, 225, 229, 234

microbes in, 184, 186 pH of, 200 minerals in, 180, 181 production of, 234-5 seasonal production of, 133, 141, 144 stability of, 192, 193, 201

Cattle, 173 Caucasus, 59, 60, 64 Cells, albumen, 14, IS, 39

central, 27, 28 chloragogen, 13, 19, 100, 101, I16 ciliated, 22, 53 clitellar gland, 79 cocoon secreting, 39

Cells, albumen, 14, 15 basal, 14 chloragogen, 13, 19, 100, 101, I16 ciliated, 22, 53 coelomic epithelial, 17 columnar, 39

drainpipe, 29 epithelial, 14, 27 epidermal, 105 epithelial brush border, 99 excretory, 100 fibrous, 34 ganglia, 34 giant, 35 gland, 14, 22, 38, 39, 79, 175 goblet, 14, 15, 39, 99 marginal, 27, 28 mucous, 14, 38, 39 nerve, 34, 106 peritoneal, 19, 101 photoreceptor, 15, 16, 107 proprioceptor, 35 reception, 114 replacing, 14 reproductive, 75 secretory, 106 sensory, 14, IS, 16, 17, 34, 106 sperm, 39 supporting, 14, 16 uric, 101

Cellulase, 99, 144 Cellulose, 177, 182 Centipedes, 147 Central cells, 27, 28 Cephalization, 4, 60 Cereal stooks, 222 Cereals, 193, 206, 207 Cerebral ganglia, 21, 31, 33, 34, 92, 106,

107, I13, 243 Ceylon, xiv, 48, 49, 50, 52, 57, 147 Chaetotaxy, 4-7 Chemical extractants, 120

sampling methods, 120, 121 stimuli, response to, 89, 122 transformation, 102 transmitter substances, 106

Chemoreception, 108-g Chernozem, 182 Chickens, 226, 227 Chile, 51, 59 China, 48 Chitin, 182 Chitinase, 99, 144 Chloragogen cells, 13, 19, 100, 101, I16 Chloragogenous tissue, 19 Chloragosomes, 19, 100 Chlordane, 213, 214, 218, 223 Chlorfenvinphos, 214 Chloropicrin, 2 I I

Chlopropham, 212 Choice experiments, I 13 Cilia, 27, 29

GENERAL INDEX 319

Ciliated cells, 22, 53 tubes, 101, 103 protozoa, 148

Circular muscle, 14, 15, 17, 18,22, 26, 31, 32, 34, 36, 38, 112, 114

innervation of, 33 Circulation, 103-4

Circulatory system, 23, 24, 85, 103 Circumoesophageal nerve ring, 114

connectives, 31, 32, 33, 107 Citric acid, 108 Clasping, 78 Clay, 129, 162, 190

chimneys, 142 population in, 162, 163

Clitellar gland cells, 79 segments, 13 tissue, 39

Clitellum, I, 10-13, 36, 38, 39, 48, 49, 55, 59, 60, 73, 76, 78, 81, 84, 86, 245, 246,247,248,249,250,251,252,253, 255

position of, I I, 13 Clover, 164, 196, 208 Cluster flies, 149 Cobalt, 168 Cocoon, development time of, 72

formation, 10, 78, 79 incubation time, 74 production, 10, 68, 69, 70, 73, 77, 106,

155, 231, 240 Cocoon production, effects of food on, 72,

164 effects of soil moisture on, 153 effects of temperature on, 68, 83, 155

Cocoons, 40, 68, 76, 77, 79, 81, 82, 85, 86, I I I, 119, 128, 138, 148, 154, 155, 164, 23 1,232,24°,242

seasonal production of, 135, 136 secretion of, I

shape, 81 size, 232 wall, 81

Coelom, 17-20, 22, 27, 148 Coelomic corpuscles, 29

cavity, I

epithelial cells, 17 fluid, 9, 13, 17, 18, 19, 26, 96, 100, 101,

110,112,15 1 tubes, 54

Coelomocytes, I, 95 Collagenous fibres, 14 Colombia, 50, 51, 56, 57 Comensalism, 146 Commercial breeding, 199

Commissural vessels, 25 Commissures, 25, 26, 107 Comparisons of sampling methods, 122

Compost, 146, 200 species, living in, 98, 200

Computer techniques, 46 Concentration factor, 218, 220 Condensed tannins, 166 Conductivity, 119 Congo, 54, 55, 57 Connective tissue, 17, 34, 38 Connectives, circumpharyngeal, 31, 32,

33, 107 subpharyngeal, 33

Consumption of soil and litter, 172-4

Contralateral nerve, 105 Contrasting soils, experiments with, 239 Control of earthworms, 223

Copper, 221 sulphate, 2 I I

Copulation,S, 39, 42, 76-83, 77, 82 Copulatory chambers, 81

pouches, 78 setae, 58

Corpuscular bodies, 19 Corsica, 58, 66 Costa Rica, 50, 53, 57 Cotton, 217 Creatinine, 101 Cretaceous period, 46 Crop, 20, 21, 22, 99, 177, 184, 215, 223 Crop growing areas, 47

plants, 109 yields, effects of earthworms on, 195-9,

198 Cropping, effects of, 194, 206-7, 207, 208 Cross-fertilization, 76, 79 Crows, 147 Crumb structure, 108, 187 Cuba, 51 Cultivation, effects on earthworms, 203---{"

2°4,206 Cultivations, 205 Culture, growth in, 85, 86

methods, 232 Cultures, 68, 70, 72, 81, 172, 194,218,219,

23 1, 232, 237, 239 Cutaneous respiration, 20 Cuticle, I, 14, 15, 16, 20, 22, 26, 27, 38, 96,

99, 110, 175 Cyanazine, 2 I 2

C ysticercoid stage, 148 Cysticercus, 225

D-D,211


DOE, 215, 218 Dead roots, 161 Deciduous woodland, 165 Dehydration, 109

tropism, 110

Dendrites, 106 Dendrogram, 46 Denmark, 124, 149 Depth zones, 128 Dermis, 106 Deserts, 6 I, 64, 66 Desmids, 184 Desiccation, 14, 20, 70, 72, 83, 154 Development, 73, 74, 86, 116, 239 Diapause, 76, 83~4, 151, 234

facultative, 70, 83, 84, 153 obligatory, 69, 83, 84, 242

Diatoms, 184 Diazinon, 214 Dicotyledenous plants, 41 Dieldrin, 213, 217, 218 Diets, 164, 165 Digestion, 22, 98~IOO Dimensions of worms, 241 Dimethylbenzanthrene, 230 Dipterous larvae, 148 Direct drilling, effects of, 206 Disking, effects of, 203 Disease transmission, 223~8 Dispersive power, 127, 128 Dissection, 233, 242 Distributions, 48, 49, J27~JI, 149, 159,

163, 233~4, 242 Disulfoton, t16, 214 Diurnal activity, 138, 178

changes, 110

respiratory cycles, 97, 106 weight changes, I II

Diverticula, of buccal cavity, 20 DNOC, 212 Dogs Mercury, 166 Dominant species, 145 Dormancy, 152 Dorsal pores, 8, 9, 10, 19, 20, 26, 55, 57,

58, 60, 88, 110, 151, 248, 249, 250, 25 1, 252, 253, 255

of A. rosea, 10 Dorsal vessel, 18,21,22,23,24,25,26,37,

104 valves of, 25

Dorso-intestinal vessel, 18, 25, 104 subneural vessel, 18, 23, 25, 104

Drainage, 194, 205, 228 Drainpipe cells, 29 Droughts, 84, 152 Ducks, 226

Dung, 95, 124, 127, 160, 161, 164, 165, 172, 173, 174, 197, 198

pats, 127 Dursban, 214 Dutch elm disease, 2 I 5 Dwarf bunt, 188, 224 Dyfonate, 214

Earthworms, as bait, I 19 as benefactors, 221)-30

as pests, 222~9 control of, 223

Earthworm farms, 229 pies, 229 to kill, 233

Ecology, xvii, J J8~67 Ecuador, 56 Eelworm cysts, 224 Egg sacs, 54 Eggs, I, 40, 224, 228, 242

dispersal of, 40 Egypt, 57, 59, 75, 125, 150, 154, 161 Elderberry, 166 Electrical inhibition of growth, I I 5

potential, 107 shock, 93, 243, 244 stimuli, response to, 89, J09

Electrode, 92, 93, 120, 243, 244 Eleocytes, 19 Elm, 166, 170, 173, 177, 237 Endemic species, 47, 61, 64, 66 Endrin, 213, 214, 218, 223 England, 41, 65, 129, 131, 140, 141, 145,

147 Environmental factors, influence of, 70, 72,

108, 141)-67

Enzymes, 99, 142, 168, 182, 225 Epidermal cells, 105

sense organ, 16 Epidermis, 10, 14, 15, 16, 17, 26, 27, 32,

34, 36, 39, 96, 106, 110, 175, 252 Epilobous prostomium, 2, 4 Epineurium, 34 Epithelial brush border cells, 99

cells, 14, 27 sense organs, 14, 35

Epithelium, 17, 20, 25, 99 Essential intermediate hosts, 225 Ethiopia, 55 Euprostates, 53 Europe, 47, 58, 59, 60, 64, 65, 144, 149,

181, 19~ 199. 205. 246 Excretion, I, 25, Ioo~3, 175. 177 Excretory canals. 30

cells, 100 ducts. 30

GENERAL INDEX 321

organs, 10, 100, 101 system, 27-]I, 46, II I

Experimental cages, 235, 236 Experiments with earthworms, 190, 231-

244 field, 233-44 palatability, 236--7 physiological, 242 pot, 197, 213 standardization of, 94

Extraction efficiency, 123 External segmental grooves, 17, 18 Extra-oesophageal vessel, 25

Facultative diapause, 70, 83, 84, 153 Faeces, 102, 140, 142, 182, 199, 228

microbial content of, 182, 186 Fallow, 123, 124, 145, 206 Families, 42---{,0

Farmyard manure, 70, 205, 239 Fats, 101 Fecundity, effects of food on, 164

effects of moisture on, 152 effects of temperature on, 154

Feeding, 82 surface, 89

Female pores, 8, I I, 45, 48, 49, 55, 57, 58, 60

Fenestrae, 34 Fertilization, 40, 76--83

internal, 79 Fertilizer, 151, 198, 199, 207, 209 Fertilizers, effects of, 207-11

mineral, 210 ni trogenous, 209 organic, 207 Fever, 229

Fibre, collagenous, 14 intra-epidermal nerve, 105 lateral giant, 34, 105 median dorsal longitudinal giant, 105 motor, 105 nerve, 16, 105, 108

Fibrillae, 34 Field experiments, 233-44

capacity, 195 studies, 174

Filtration, 102 Fish bait, I 19 Fistula, 228 Flatworm, 147 Flooded soil, 153 Fodder, as food, 164 Follicles, 4, 75 Food, 22, 68, 70, 72, 89, 90, 93, 98, 108,

113, 117, 127, 139, 146, 160, 163, 164,

165, 166, 167,205,208,229,231,235, 239, 242

adsorption, 22 earthworms, as, 229 effect of supply, 164-7 preferences, 108 supply, 68, 139, 164-7

Food material, availability of, 177 bullock droppings, 72, 164, 23 I compost, 146 dung, IZ7, 160, 164 fodder, 164 herbage, 127 horse droppings, 72, 23 I leaf, 89, 90, 98, 127, 235 manure, 93 oat straw, 164 organic matter, 127 plant, 89, 90, 98, 108, 164 roots, 133 sheep droppings, 164, 240 straw, 231, 239

Foot and mouth disease, 225 Forest soil, 123, 193, 194, 201 Formalin, 14, 120, 122, 123, 124,232,233,

234, 242 sampling, 88, 121, 232

Formaldehyde, 109 Fossil records, 41 Fossil worms, 41 Fragmentation, 168 France, 52, 66 Freshwater worms, 13 Fried earthworms, 229 Frost, effect of, 156 Fumigants, 2 I I

Fungal hyphae, 186, 193 mycelia, 165 spores, 185, 224

Fungi, 98, 183, 185, 186, 187, 188, 193, 194, 224

Fungicides, 168 copper, 211

effects on worms, 2II, 214

Gambia, 53, 57 Ganges valley, 5 I Ganglia, 21, 31, 32, 35, 92, 106, 107

cells, 35 cerebral, 21, 31, 35, 92, 106, 107, 113,

243 sub-oesophageal, 31, 106 sub-pharyngeal, 32 ventral, 106


Gape-worms, 225 Garlic, 19 Gaseous diffusion, 97 Genera, 42---{)O

Generator, 120 Genital and other apertures, 8-10 Genital organs, 36, 40

pores, 5, I I

seta, 5, 6, 60, 8 I systems, 36

Geographical distribution, 61-7, 233 Germany, 58, 65, 124, 144 Ghana, 54, 55, 144 Giant cells, 35

fibres, 34, 35, 86, 105, 106 dorsal, 105 longitudinal, 105 medial, 105

Gizzard, 20, 21, 22, 24, 48, 49, 55, 57, 58, 59, 60, 98, 99, 184

Glaciation, 64 Gland, calciferous, 20, 22, 44, 45, 46, 55,

57,58,59,99, 146, 151, 193 cell, 14, 22, 38, 39, 79, 175 lymph, 24, 101 mucous, 26, 96 papillae, 8 pharyngeal, 20, 99 prostatic, 46, 48 salivary, 99

Glandular epidermis, 39 secretions, 76

Glucose, 17, J08, 187 Glycogen, 97, 101 Goblet cells, 14, 15, 39,99 Golden plovers, 147 Golf courses, 210, 223 Gonads, I

Granite particles, 190 Granules, 13

yellow refractive, 100 Grape husks, 189 Grass, 1'22, 124, 145, 184, 189, 193, 196,

208 reseeding with, 146

Grassland, 123, 144, 145, 159, 161, 194, 205, 206, 211

earthworms, as pests of, 223

Gravelly loam, populations in, 163 Gravelly sand, populations in, 162 Great Britain, 229, 248 Green ash, 198 Greenhouses, 95 Greenland, 59 Gregarines, 148 Growing zone, 85

Growth, 85-7 effects of food on, 164 effects of temperature on, 164 electrical inhibitions of, 115 increment, 86 periods, 71, 72, 73, 155 post emergence, 85

Guatemala, 51, 57 Guinea, 53 Gulls, 147 Gut, 18, 20, 23, 25, 30, 41, 100, 104, 114,

166, 172, 173, 182, 183, 219 contents of, 165, 182, 183, 186, 187 enteronephric, 30 microflora in, 182 micro-organisms in, 183

Habitats, 123, 124, 127, 144, 233 arable land, 233 chalk downs, 233 heathland, 144, 233 heterogenity of, 127 lakes, 246 moorland, 233 numbers and weights in, 124 pastures, 233 rivers, 246 streams, 246 woodlands, 233

Haemochromagen, 103 Haemoglobin, 14, 26, 85, 96 Haemorophilic spp., 65 Haemorophobic spp., 65 Hairs, 14 Haiti, 52 Handsorting, 118-19, 122, 124, 233, 234,

242 Hants, 124 Harrowing, 203 Hawaii, 59 Hay, 196 Hearts, 23, 24, 25, 37, 50, 104

anterior loop of, 24 intestinal, 25 lateral oesophageal, 50 valves, 25, 104

Heat extraction, 121-2

Heath, Cal/una, 145 Heathland, 144, 233 Heavy metals, effects on worms, 220--1

pattern of uptake, 220 Hebrides, 163 Hedgehogs, 147 Heptachlor, 214, 218, 223 Heptachlor epoxide, 218 Herbage, 127

GENERAL INDEX 323

Herbicides, effects on worms, 205, 211, 212

Herefordshire, 4 I Hermaphrodite, I, 8, 36, 40, 76 Hertfordshire, 124 Heteroxanthine, 103 Himalayas, East, 48 Holland, 95, 124 Holonephridia, 30, 3 I, 48, 57, 58 Horizontal distribution, 127-8, 242

migration, 128 Hormones, 106 Horse droppings, as food, 72, 231 5HT,106 Human excreta, 185

faeces, 228 urine, 228, 229

Humidity, 19, 242 Humification, 172-4 Humus, 159, 174, 193 Hungary, 58 Hydrogen sulphide, 95 Hydrostatic pressure, 110 Hypotonic urine, 110

Iceland, 59, 60, 147 Immature worms, proportions of, 125 Impotency, 229 Incorporation of fallen leaves, 168 Inclusions, 101

inorganic, 19 Incubation period, 73, 74, 75 India, xv, 48, 49, 50, 5', 52, 53, 57, 58, 59,

60,65,66,73, 136, '4', '42, 144, 181, 222

Indiana, 124 Indonesia, 57 Infectivity of cysts, effects of worms on,

228 Iriprganic minerals, '42

salts, 127 Insecticides, effects of, 211, 242

residues, 2 I 5 to control earthworms, 223

Inter\nediate hosts, earthworms as, 225 Internal salt concentration, I I I

Intersegmental grooves, I, 8, '3, 18, 248, 250

§Wtum,103 zones, 13, 25 I

Intestinal epithelium, 99 microflora, 174, 186 wall, 33 worms, 99

Intestine, 18, 20, 21, 22, 24, 48, 59, 98, 99,

100,104,113,114,152,168,184,187, 215, 220, 224

dorsal wall of, 22 Intra-epidermal capillaries of L. terres/rtS,

26 Introduced species, 63 Invertebrates as predators, 147 Iron, 220 Irridescence, 14, 247 Irradiation, 2 I 9 Irrigation, artesian, 206 Irritants, 109 Isobenzan, 2 I 4 Isotopes, 2 I 9 Israel, 58, 59, 60 Italy, 58, 59, 64, 66 Ivory Coast, 144

Japan, 48, 58, 59, 64, 66, 73, 137, 138, 222, 229

Jaundice, 229

Java, 48 Jordan, 59 Jurassic period, 41

Kale, 166 Kentucky, 41 Kenya, 54 Kidneys, 101 'Kilner' jars, 23 I Kommetjies, 142

Lactic acid, 97, 98 Lake District, 163 Lamellae, 34 Larch, 166 Lashing movement, 88 Lateral giant fibres, 34, 105

nerve cells, 34 neural vessel, 18, 23, 24, 104 oesophageal hearts, 25, 50 oesophageal sub-neural vessel, 25, 26 oesophageal vessels, 23, 24, 25, 104 sub-neural commissures, 25 vessel, 23, 24, 25, 104

Latex, 139 Lawns, 223 Lead, 220, 22 I Lead arsenate, 2 I I

Leaf, apple, 170 ash, 237 bean, 237 beech, 170, 237 beet, 237 birch, 170, 237


Leaf-conI. burial, 236 cabbage, 237 disks, 109, 165, 169, 170, 236 elm, 170, 173, 237 fall, 161, 168, 173, 235 kale, 237 lime, 170, 237 litter, 89, 90, 98, 108, 127, 165, 169, 170,

173 mineral content of, 165 oak, 170, 237 palatability of, 236-7 protein content of, 166 ribs, 170 sugar content of, 166 veins, 170 weathering, 169

Leaf burial, effects of temperature on, 156 Leaf disks, rate of breakdown of, 170 Leeches, 40, 41, 147 Legume, 206, 207 Lettuce, 166, 222 Leys, 176, 194, 206 Liberia, 50, 54 Life cycles, 68-75, 239

history studies, 239 span, 72, 73

Light intensity, 93 means of detecting, IS photonegative response, 107 photopositive response, 107 reactions, 106-7, 178 receptors, 106, 107 sensory. cells, 106 stimuli, 107 ultra-violet, 107

Light loam, populations in, 162, 163 Lignin, 177 Lime, 170, 177, 197, 201, 208, 210, 211,

234,237 Lipase, 99 Lithium, II6 Litter, 89, 90, 98, 108, 122, 127, 165, 168,

169, 170, 171, 172, 173, 174, 176, 177, 178, 180, 189

breakdown of, 168-72, 178, 210 CN ratio of, 177-8 consumption, 172-4 decomposition, 178, 189 feeders, 146 forests, 165 fragmentation of, 168-72 hazel, 173 humification, 172-4 incorporation of, 168

layer, 122 pine, 170 rate of burial, 169 species, 98 turnover, 172-4

Little Barrier Island, 53 Liver, 101 Loam, 129, 162, 163, 231, 237 Locomotion, 110, /[2-14 Lonbok,48 Longitudinal canals, 30

muscle, 4, 15, 17, 18, 22, 26, 34, 36 ridges, 13 vessel, 22

Lucerne, 193 Lumen, 27, 29, 30, 101 Luminescence, 95 Luminous slime, 95 Lung worms, 225, 226 Lymph glands, 24, 101 Lymphocytes, 19

Madagascar, 50, 5 I, 52, 53, 57 Magnesium, 180, 201

chloride, 113 sulphate, 119, 232

Malagasy, 56 Malathion, 214 Malay Archipelago, 48, 52, 56, 57

Peninsula, 57, 66 Malaya, 49 Malaysia, 66 Male cell, 37, 75

duct, 38 pore, 5, 8, 39, 44, 47, 49, 53, 55, 57, 58,

59, 60, 75, 76, 78, 245, 249, 255 Mammals, 147, 215, 220, 227 Manchuria, 66 Mangolds, 161 Manure, 70, 93, 188, 200, 205, 239 Maoris, 229 Marginal cells, 27, 28 Maryland, 136 Mass migration, 94, 95 Mating 75--83, 81, 106 Maze, 90, 92, 93, 95, 166, 206, 229, 243 Mechanical, damage, effects on population,

204 stimulation, 122

Mechanized soil washing, 119 Medium loam, population in, 162, 163 Meganephridia, 30, 54 Megascolecid group, xi Membrane, basal, 17, 33

basement, 16 mesenteric, 18, 22, 23

GENERAL INDEX 325

Membrane--cont. peri trophic, 100

Menazon, 214 Mercuric chloride, 120 Meronephridia, 30, 3 I Mesh bag, 169, 170, 236, 237 Metabolism, effects of temperature on, 155 Metallic residues in soil, 220 Metham sodium, 211 Methyl bromide, 2 I I Mexico, 51, 52, 53, 55, 57, 65 Michigan, 136 Microbial activity, 84, 174, 185, 187

cell, 186 protein, 175

Microflora, 159, 168, 174, 182, 186, 187 Micronephridia, 30, 54 Micro-organisms, 142, 168, 174, 182-9,

184, 185, 186, 187, 193 and earthworms, 182-9 dispersal of, 188 numbers in casts, 182, 184 numbers in gut, 183

Middle tube, 27, 29, 102, 103 Mid-dorsal line, 18, 3 I

-ventral line, 32 Migrations, 128, 131 Millet, 196 Mineral elements, 160, 181

inorganic, 142 nutrients, 180-1

particles, 22, 190 soil, 181

Minerals, effect of, 160 Mineralization of nitrogen, 174-7, 209 Mississippi, 64 Mites, 174 Mixed woodland, 124 Moisture, 83, 127, 131, 132, 152-4, 159,

163, 195, 242, 243 content of earthworms, 106, I I I, 112,

152

content of soil, 68, 109, IIO, II8, 121, 127, 131, 132, 133, 141, 152, 153, 159

equivalent, 201 gradient, 154, 243 reaction to, 242, 243

Moles, 147, 222, 223 Molybdenum, 181 Moorland soil, 123 Morphology, xvi, 1-39 Mor soil, 123, 124, 145, 163, 192

species in, 123, 163 Mortality, 86 Motor fibres, 105

nerves, 105

Mountain ranges, 61 Mouth, 2, 12, 20, 21, 31, 110 Mowrah meal, 120 Mozambique, 54, 57 Mucoproteins, 175 Mucous cells, 14, 38, 39

glands, 26, 96 membrane, 114 tubes, 78

Mucus, 14, 76, 94, 99, 100, 108, 138 Mud,75 Mulching, 205 Mull soil, 123, 145, 163, 192

species in, 123, 145, 163 Multiplication rate, 128 Muscles, arciform, 76

circular, 14, 15, 17, 18, 22, 26, 31, 32, 34, 36, 38, 112, 114

dorsal, 34 fibres, 18 innervation, 105 longitudinal, 4, IS, 17, 18, 22, 26, 34,

36 setal, 4 sphincter, 10, 18, 19, 30 subcuticular, 13 ventral, 17

Muscoid flies, 149 Muscular configurations, 113

contractions, I 13 rhythm, II3 tube, 27, 29

Mycobacteria, 187

Naphthalene, 169 Narrow tube, 27, 28, 29, 102, 103 Nematodes, 19,98, 148, 21 I, 215 Nephridia, I, 10,21,26,27,28,29,3°,31,

48, 54, 55, 57, 58, 100, 101, 103, IIO enteronephric, 30 exonephric, 29, 30 funnel, 27, 28 holonephridia, 30, 3 I, 48, 57, 58 loops, 30 macroic, 55 meganephridia, 30, 54 meronephridia, 30, 3 I micronephridia, 30 septal, 21 tubes, 27, 28 tufted, 30, 154 vessel, 26

Nephridiopore, 9, 10, 26, 27, 30, 44, 46, 57, 58, 100, 103, 110

Nephrostome, 27, 28, 29, 101, 102, 103 lower lip, 28


N ephrostome-cont. peritoneum, 28 upper lip, 28 prostomial, 31, 32

Nerve cells, 34 contralateral, 105 cord, 18,31, 32, 34, 35, 36, 37, 86, 104,

107, "3, "4, 116, 242 depressant, I 16 fibres, 16, 105, 108 impulses, 108 intra-epidermal, 105 lateral, 34 motor, 105 plexus, 15, 32, 33, 34, 35 prostomium, 32, 33 prostomial, 31, 32 ring, 32, 33, 34, 35 segmental, 31, 32 septal, 32, 33 tissue, 14

Nervous system, 31-5, 31, 32, 104, /05-9, 116

Netherlands, 128, 211 Nettle, 166 Neural lamella, 34

lateral vessel, 18, 23, 24, 104 Neurofibrils, 16, 35 Neuroglia, 34 Neurones, association, 105 Neuropile, 34, 35 N eurosecretions, 106 New Caledonia, 51, 52, 67 New Guinea, 229 New South Wales, 49, 59, 60, 172 New Zealand, xv, 49,50,51,52,53,59,67,

84, 138, 139, 181, 195, 197, 201, 229 Newly-emerged worms, 85 Nigeria, 50, 54, 55, 57, 75, 123, 136, 144,

173, 181, 195, 206 Nile Valley, 144 Nitrate, 175, 176, 180, 189, 201

of soda, 160, 175, 209, 211 Nitro-chalk, 208, 209 Nitrogen, 102, 174-7, 180, 181, 189, 201,

202, 209 available, 180 content of L. terrestYlS, 177 excretion, 27, 100, 176, 179 from earthworm corpses, 175 humification, 174-7 mineralization, 174-7, 209 non-available, 176 production, 180 turnover, 176

Noradrenalin, 106

Novocaine, 106 Number of samples, 122 Nutrients, 100, 180-1, 195 Nutrition, 70 Nylon net, 166, 169

Oak, 124, 144, 166, 168, 169, 177, 184, 198 leaf litter, 169 woodland, 124, 144

Oats, 169, 197 Obligatory diapause, 69, 83, 84, 242 Observation cage, 237, 238 Oesophageal crop, 58

hearts, 25, 50 pouches, 20 region, 18 sacs, 2I

wall,20 Oesophagus, 20, 22, 24, 37, 45, 59, 98, 99,

104, 113 Ohio, 149, 152 Oocytes, 36, 76 Oogenesis, 76 Oogonia,76 Oothecae, 68, 74 Optic organelle, 16, 35 Optimum temperature, 116, 133, 157, 239 Orchards, 109, 122, 123, 124, 168, 171,

173, 211, 214, 216 Ordovician period, 4 I Organ, bush-like, 30

excretory, 10, 100, 101 genital, 36, 40 gill-like, 26 photoreceptor, 35 reproductive, 26 respiratory, 26, 98 sense, 14, 35, 105, 108 tactile, 109

Organic matter, 93, 98, 99, 127, 142, lSI)-

161, 164, 168-81, 183, 190, 192, 194, 199, 201, 203, 205, 210, 21 1,235, 237

breakdown, 168-81, 190, 205 cycles, 168-81 decomposition, 168, 176, 183 effect of, 2 I2

incorporation of, 235 surface mat of, 159, 168, 172, 2 II

Organo-chlorines, 214, 215-18, 217 Organo-phosphates, 218 Osmotic pressure, I 10

of blood, 110 of urine, 101, 102 regulation, I I I

Ova, I, 68, 76, 79, 81

GENERAL INDEX 327

Ovarian funnels, 36 Ovaries, :n, 36, 37, 48, 49, 54, 58, 59, 76,

81, 117 Oviduct, 21, 36, 37, 54, 76, 81 Ovisacs, 36, 48, 76, 81 Oxalic acid, 108 Oxidation-reduction potential, 159 Oxygen, 26, 96, 97, 98, 116, 159, 176, 178,

179, 186, 187, 189 consumption, 98, 176, 179, 186, 187, 189 debt, 97, 178 partial pressure of, 98 tension, 159, 178

Pacific Islands, 49 Pakistan, 48, 59, 60 Palearctic zone, 47 Palatability, 161

experiments, 236-7 of leaf litter, 108, 109, 161, 166

Panama, 50, 56 Papillae, glandular, 8 Paraguay, 52, 56 Paraquat, 212 Parasites, 19, 147-9, 223, 225, 226, 227,

228 eggs of, 228 nematode, 19 protozoan, 19, 148 transmission of, 226, 227

Parathion, 116, 2 I 4 Parthenogenesis, 63, 64, 76, 79, 8 I, 242

facultative, 79 Passive agents, earthworms as, 228 Pasture, 84, 123, 124, 129, 133, 136, 138,

144, 151, 156, 159, 161, 165, 169, 172, 181, 188, 190, 191, 193, 194, 197,201, 206,207,208,217, 231, 232, 233, 234

productivity, effect of worms on, 198 Peas, 196 Peat, 72, 237 Penes, 78, 8 I Peregrine species, 49, 61, 64, 66, 246 Perichaetine setae,S, 245 Peri-enteric plexus, 18, 25, 104 Peristalsis, II 2-1 4, II3 Peristomium, 2, 60 Perithecia, 188 Peritoneal cells, 19, 101 Peritoneum, 14, 15, 17, 25, 28, 34, 76 Peritrophic membrane, 100 Persia, 59 Peru, 51 Pesticides, effects of, 2II-15

uptake in earthworms, 215-18

Pests, earthworms as, 222-30

Petroleum ether, 187 pH, 99, 108, 109, 120, 127, 14g---52, 150,

159, ISo, 181, 195, 201, 210, 234, 243 effects of, 150, 151,234 reaction to, experiments, 243

Phalaris leaves, 164 roots, 164

Pharyngeal ganglion, 33 gland, 20, 99 intestinal plexus, 35 nerve plexus, 35

Pharynx, 20, 21, 24, 31, 98 Pheromone, 140 Philippines, 48, 66, 222 Phorate, 214, 223 Phosphorescence, 95 Phosphoric acid, 108 Phosphorus, ISo, 181, 201, 210 Photoreceptor cells, 15, 16, 107 Photosensitivity, 107

drug depressants, 107 Physiological experiments, 242 Physiology, xvii, 96-II7 Pig, kidney of, 227

litter, 124 Pigment migration, 106 Pigmentation, 13-14, 84, 85 Pigs, 225 Piles, 229 Pine litter, 170

needles, 165 woodland, 124

Plant breakdown, xvi cover, 159 growth, 181, 195, 206, 228 material, 164, 207, 231 pathogens, earthworms as vectors of, 188 residues, 193

Ploughing, 145, 151, 177, 203, 204 effects on worms, 2 I2

Plugs, burrow, 89 Podsol, 123 Polyhydric phenol, 166 Polynesia, 52 Polyphenols, 108, 166, 169

water soluble, 166, 169 Polysaccharide gums, 188, 193 Population,

adolescent, 87 biomass, 206 density of, 162 distribution of, 127-31

Population, estimation of, 118-23, 233 factors limiting, 177 geographical, 6I -7, 203 horizontal distribution, 127, 128, 242


Population-con/. numbers, 162 pyramid, 74 reduction of, 147 seasonal activity, 131-8, 134 seasonal changes in, 87, IJI-8, 132, 133,

136, 137 size of, 12J-7 structure, 125-7 vertical distribution, 128-31, 129, 138,

242 Populations, 118, 127, 128, 136, 138,

147, 151, 152, 156, 160, 161, 162, 164, 177, 178, 179, 189, 195, 203, 205, 206, 207, 214

Pores, cuticular, 14 dorsal, 8, 19, 20, 26, 55, 57, 58, 60, 88,

IIO, 151,248,249,250,251,252,253, 256

female, 8, I I, 48, 49, 55, 57, 58, 60 genital,s, I I

male, 39, 44, 53, 57, 58, 59, 60 prostatic, 8, 42, 44 spermathecal, 8, 48, 53, 60, 245, 247

Porphyrins, 13 Porosity, 194-5 Post-clitellar region, 6 Post-septal canal, 27 Pot experiments, 197, 213 Potassium, 180, 181, 210

cyanide, II 6 permanganate

Potato root eelworm, 224 Pouches, 18 Predation, 70 Predator, 20, 143, 147-9 Pregnancy, testing, 229 Preseptal canal, 27 Preservation, 2JJ Proboscis, 2 Prolobous prostomium, 2, 4 Propham, 212 Proprioceptor cells, 35 Prostates, 8, 21, 39, 44, 45, 49, 53 Prostatic fluid, 78

gland, 46, 48 pore, 8, 42, 44

Prostomial nerve, 31, 32 Prostomium, 2, 4, 12, 15, 17, 31, 32, 33,

57, 60, 95, 106, 108, 110, 112, 245, 248, 249

enervation, 3 I, 35 Protease, 99, 142 Protective slime, 95 Protein, 101, 102, 103, 166, 175 Protein content of earthworms, 229

Proteolytic enzymes, 99 Protozoa, 98, 99, 148, 188, 226 Protozoan parasites, 19, 148 Pseudotsuga mor, 124

QlIadrat, II8, 121, 123, 234, 235, 242 Quaternary ice caps, 64 Queensland, 49, 5 I, 52 Quiescence,8J-4, 130, 13 1, 136, 153 Quinine, 17

Radiation, 95 effect on A. chlorotlCa, 219

gamma, 219 Radioactive fallout, 2 I 8

pollution of soil by, 2 I 8 Radiocaesium, 173, 219 Radio-iron, 219 Radioisotopes, effect on worms, 2I 8-1 9

uptake into worms, 218, 219 Radishes, 228 Ragi,228 Rain, reactions to, 95 Rainfall, 235 Raw humus, 159 Reception cells, 114

zone, 99 Receptors, 17, 35 Rectum, 195 Reduction division, 76 Regeneration, 80, 85, Il4-16, 243 Regenerative phenomena, 80, 106, lIS,

II6 Rennin, 99 Replacing cells, 14 Reproduction, 75-8J, 106

asexual, 79, 80, 8 I effects of temperature on, 154

Reproductive cells, 75 cycle, 106 potential, 127 structures, 21 system, 36-9, 36, 37, 60

Reservoir, 27 Reservoir host, earthworms as, 225 Resorption, 101, 102 Response, learned, 94 Respiration, I, 26, 96-8, I 16, 178, 180

anaerobic, 98 cutaneous, 20 depression of, 98 effects of temperature on, 154, 155 rates of, 97, 98

Respiratory exchange, 187 system, 26-7

Resting phase, 84

GENERAL INDEX 329

Retinella, 35 Rheumatism, 229 Rhizopods, 184 Rhodesia, 54, 57 Rice, 222 Ridges, 12 Rigor, 110 Rio de Janeiro, 56 Rodents, 226 Rolling, effects of, 203 Roots, 161, 164, 165, 168, 193, 198, 200,

205, 222 Rotary cultivation, 203 Rotations, 206 Rothamsted, 125, 126, 129, 133, 145, 151,

160, 161, 172 Rotifers, 98 Rowan, 177 Russia, 58, 59, 60, 163 Rye, 184, 197, 206, 201)

Septal nerves, 32, 33 wall,27 vessels, 25

Septum, 22, 27, 29, 32, 36 intersegmental, 102

Setae, I, 4, 5, 6, 7, 41, 48, 53, 55, 57, 58, 60, 78, I12, 114, 175, 245, 248, 249, 250, 25 1, 253

closely-paired, 6, 7 copulatory, 58 distant-paired, 6, 7 genital,s, 6, 60, 8 I lateral, 12

lumbricine, 5 normal, of L. lerreslns, 5 penial,81 perichaetine, 5, 245 physical stimulation of, 5 ventral, 12 widely-paired, 6, 7

Setal arrangements,s, 7, 245 lumbricine, 7 perichaetine, 7

Setal muscles, 4 Setal form, 4, 5 Sewage sludge, 70 Sexual activity, 75

development, 76 Secretory zones, 99 Seedlings, 222 Segment numbering, 233 Segmental bands, 13

contractions, I 12 ganglia, 31 grooves, 17

nerves, 31, 32 Segmentation, 1-4, 239

internal, I

Segments, I, 8, 27, 105, 109, 151 Selenium, 220 Semi-aquatic species, 9, II, I II, 154 Seminal fluid, 78, 79

grooves, 8, 12, 76, 247 recepticles, 8 I vessels, 36, 37

Seminal vesicles, 21, 37, 57, 58, 60, 79, 81, 148,230

anterior, 37 mid,37 posterior, 37

Senescence, 85 Sense organs, 14, 15, 35, 105, 108

epidermal, 16 epithelial, 35 photoreceptor, 35

Sensory cells, 14, 15, 16, 17, 34, 106 hairs, 35 input, 106 light receptive role, 106

Septa, 1, 18, 20, 25, 30, 104, 112 innervation of, 34 maturity, 12, 72, 86, 87, 155, 188, 242 organs, 40

Seychelles, 50 Sheep droppings, as food, 164, 240 Siberia, 59, 60, 64 Sicily, 58, 66 Sieve, 119, 121, 232 Silurian period, 4 I Simazine, 212 Simple key, 245-53 Size of sample, 122 Slime, 95

luminous, 95 protective, 95 tube, 76

Slugs, 147 Smallpox, 229 Snakes, 147 Sodium chloride, I I 1 Soil, 98, 99, 109, 145, 149, 150, 151, 152,

154, 156, 159, 163, 168, 170, 181, 184, 185, 200, 216

acidity, 108, 149, 150, 151, 208 adverse effects on, 228-9 aeration, xvi, 127, 190, 194-5, 205, 206,

228 air space, 194 amelioration, 199-203, 229 animals, 168 climate, 68


Soil-cont. climate, fluctuations of, 68 compaction, 192 consumption, 172-4 degeneration, 2 I I

density, 202 drainage, 194-5, 205, 206, 228 erosion, 192, 228, 229 fertility, xvi, 174, 181, 190-2, 199, 205 flooded, 153 formation, xvi, 200 fumigants, 21 I horizons, 192 indicators, 195 ingestion of, 190 litter species, 98 microbial activity in, 174 microflora, 159 mineral fraction of, 191 minerals in, 18 I moisture, 68, 109, IIO, II 8, 121, 127,

13 1, 132, 13L 141, 15~ 153, 159 movement through, 14 nutrient content, 195 organic matter, 161 organisms, 174 particles, 174, 190, 193 particles, breaking down of, 190 pathogens, 21 I pathogens, control of, 2 I I

pH, 200 profile, 168, 171, 192, 199 reclamation, 199, 200 stability, 211 sterilized, 188 structure, xvi, 190-2, 194, 228 surface, 188, 201, 21 I, 223, 234 temperature, 68, 84, 97, 121, 127, 129,

13 1, 132, 133, 138, 155, 157, 179 texture, 127 throughput, rate of, 123 turnover, 172-4, 191-2, 205

Soil type, 162-3, 191-2, 205 agricultural, 152, 228 air filled, 153 alluvial, 162, 163 brown earth, 181 brown podzol, 196 clay, 162, 163 clay content, 161 clay, effect of, 161 coniferous forest, 146 coniferous mor, u4, 145 dark, 163 Egyptian, 161 fallow, 124

forest, 123, 193, 194, 201 garden, 194 gravelly loam, 163 light, 162, 163 light loam, 162, 163 light sandy, 163 medium loam, 162, 163 mor, 123, 124, 145, 163, 192 mull, 123, 145, 163, 192 open, 162 orchard, 159, 171, 21 I peaty, 145, 163 ploughed, 180, 200 sandy, 144 shallow acid peat, 163 tropical, 123 tropical, populations in, 123 water-holding capacity, 190, 195 water-logged, 109, I II, 156 water-saturated, 153 woodland mull, 145, 163

South Africa, 222, 229 South America, 50, 5 I South Shetland Islands, 67 Saccharose, 108 Salamanders, 147 Salivary glands, 99 Salt, III

internal concentration, I I I

saturated solution, 242 Sample number, 122

size, 122 Sampling methods, 118-23, 233, 234, 242

chemical, 120-1, 131, 133 digging, 232 electrical, 122

formalin, u4, 232, 233, 234, 242 handsorting, 118-19, 233, 234, 242 heat extraction, 121-2 soil cores, 118, 122 vibration, 122 vibrating rod, 122 washing, I 18, 122 wet sieving, u4

Sampling tool, 118 Sand, 190 Sardinia, 52, 58, 66 Savannah, 66, 125, 195 Scandinavia, 255 Scotland, 145, 162 Seasonal abundance, u6, 128, 139

activity, 131-8

changes, 126, 139 Secondary sex characters, 84, 106 Secretory cells, 106 Soybeans, 196, 207

GENERAL INDEX 331

Species, 42---{)0 associations, [45-7

Specific gravity, 232 Sperm, 81

cells, 39 funnels, 36, 37, 38, 48, 75 sacs, 38 transference, 42, 78

Spermathecae,8, 13,21,36,37,39,42,48, 54, 59, 60, 78, 81, 148

Spermathecal ducts, 39 openings, 76, 78, 79 pores, 8, 48, 53, 60, 245, 247

Spermatogenesis, 75, 230 Spermatogonia, 75 Spermatophores, 8 I Spermatozoa, 75, 78, 79 Sphincter muscles, 10, 18, 19, 30 Spores, 184, 186, 188

fungal, 185, 224 Sports fields, 213, 223

earthworm control on, 213 Spraying, 168 Springtails, 174 Spruce, 166 Squirter earthworm, 19 Staphylinid beetles, 147 Starlings, 147 Stems, 168 Stewart Island, 51 Stimuli, 89, 90, 106

adverse, 105 avoidance of, 92 chemical, 89, 90, 105, 108 electrical, 89, 109, 199, 243, 244 electrical response to, 89, [09 intersegmental, 112 light, 76, 93, 107 light response to, 107 photic, 107 tactile, 15, 88, 93 touch, 76, 109 I 12 vibrations, 89, 95

Stomach, birds, 226 swine, 226

Stones in the bladder, 229 Straw, 70, 174, 197, 23 1, 239, 240 Stress, 20 Stretch receptors, 35 Stubble-mulch farming, 205 Stubble stripping, 204 Sub-aquatic species, 43 Sub-Arctic Islands, 51 Subcuticular muscle, 13 Sub-epidermal nerve plexus, 33, 34, 105 Sub-epithelial nerve plexus, 34

Submergence, 154 Subneural vessels, 18, 23, 25, 26, 104 Suboesophageal ganglion, 31, 106 Subpharyngeal connectives, 33

ganglion, 32 Substrate, 109 Subsoil, 186, 190 Subtillers, 205 Sucrose, 17, 108 Suction pump, pharynx as, 20 Sudan, 57 Sulphate of ammonia, 172, 21 I

effects of, 210 Sumatra, 48, 66 Sumithion, 214 Sunda Islands, 48 Superphosphate, 208, 210 Supra-intestinal, glands, 57

ventral commissures, 25, 26 vessel, 23, 24, 25, 26

Surface casting species, 131 debris, 216, 232 dwelling species, 121, 150, 204, 232

Survival, 152 in sugar solutions, 242 in test chambers, 242 in water, 152, 242

Sweden, 131, 146, 161 Swine, 226

lungs of, 226 stomach of, 226

Switzerland, 64, 144 Symbiotic bacteria, 99, 182 Sympathetic system, 33 Synapses, 105 Syria, 58, 59, 60 Systematic affinities, 40-2

Tactile organs, 109 receptors, 109

Tannins, 166 Taste, 17 Tanylobous prostomium, 2, 4, 249 Tanzania, 54, 57 Tapeworms, 225 Tartaric acid, 108 Tasmania, 51 Taxonomy, 40---{)7 TeA, 212 Teleospores, 224 Telodrin, 214 Temperate forest, 173

zones, 65, 97, 168 Temperature, 68, 61), 70, 72, 83, 135, 141,

179, [54-8, 158, 23 1, 235, 239 acclimation, 157


Temperature--cont. conditioning, 157, 158 effect of, 244 effect on excretion, 103 effect on respiration, 97 gradient, 157 lethal, 156-}, 158 optimum, 116, 133, 157, 239 preferenda, 156, 157, 158 seasonal changes in, 158

Tension receptors, 35 Termites, 147

mounds of, 147 Testis, 37 Testes, 21, 36, 48, 49, 59, 75, 79, 81, 148

funnels, 59 holoandric, 36 meroandric, 36

Thermal death point, 157, 158 Thigmotactic reactions, [09

Thiophanate-methyl, 214 Thrushes, 147 Timothy grass, 206 Tissue, connective, 17, 34, 38

nuclei of, 38 production, 177

'T' maze, 90, 92, 93, 94 Toads, 147 Tobacco, 222 Togoland, 55, 57 Tooth powder, 229 Topsoil feeders, 146 Touch, 88 Trachea, 227

of bird and chickens, 226, 227 Transplantation, [[6-[7 Transverse commissure, 107

septa, 17 Triassic era, 41 Triazine herbicides, 212 Trichlorphon, 214 Tropical areas, 97

forest, 173 lowlands, 64 soil, 123

Tropics, 136, 144 Tube, ciliated, 101, 103

middle, 27, 29, 102, 103 mucous, 78 muscular, 27, 29 narrow, 27, 28, 29, 102, 103 slime, 76 wide, 29, 102, 103

Tubercula pubertatis, II, 12, 13, 60, 81, 245,248,249,250,25 1,252,253,255

Tubercules, 12, 252

Tubificity, 41 Tumours, 230 Tunis, 58 Turf, earthworms as pests of, 223 Turnover of soil and litter, [72-4, [9[-2,

205 Typhlosolar vessel, 25 Typhlosole, 22, 25, 57, 104

Uganda, 57, 123 Ubiquitous species, 150 Ultra-violet light, 95, 98, 107

effects of, 95, 98 Urea, 100, 101, 102, 103, 175 Uric acid, 100, 103, 175

cells, 101 Urine, 100, 101, 102, I II, 175, 176, 198,

228, 229 osmotic pressure of, 101, 102

U.S.A., 57, 64, 65, 66, 124, 127, 131, 133, 136, 138, 147, 149, 152, 156, 180, 195, 199, 204, 206, 21 6, 229

U.S.S.R., 124, 200, 206

Vacuoles, 100 Vagina, 228 Variance of species, 60

of subspecies, 60 Valerionic acid, 98 Vas deferens, 21, 36, 37, 38, 39, 48, 53, 75,

81 racemose, 39 termination of, 53 tubular, 39

Vas efferens, 37, 38, 75 Vascular system, I, 22---{)

Vesiculae seminalis, 3 I Vector diagram, 46 Vectors of animal viruses,

earthworms as, 225 Vegetable crops, 222, 228 Vegetative cells, 184 Venezuela, 53, 56 Ventral muscle, 17

nerve cord, I, 23, 31, 32, 33, 34, 35, 37, 105, 107, 114

vessel, 18, 23, 24, 25, 37, 103 view, 12

Vertebrates, 147 Vertical distribution, [28-JI, 129, 138, 242

migration, 131 Ventral ganglion, 106 Vessel, afferent nephridial, 18

afferent typhlosolar, 18 blood, 15, 18, 22, 26, 33, 38, Il6 capillary, 14

GENERAL INDEX 333

Vessel-cont. commissural, 25 dorsal, 18, 21, 22, 23, 24> 25, 26, 37, 104 dorso-intestinal, 18, 25, 104 dorso-subneural, 18, 23, 25, 104 efferent nephridial, 18 lateral-neural, 18, 23, 24, 104 lateral-oesophageal, 23, 24, 25, 104 lateral-oesophageal subneural, 25, 26 longitudinal, 22 nephridial, 26 subneural, 18, 23, 25, 26, 104 supra-intestinal, 23, 24, 25, 26 typhlosolar, 25 ventral, 18, 23, 24, 25, 37', 103 ventro-intestinal, 25, 104 ventro-oesophageal, 104 ventro-parietal, 18, 25, 26, 104

Vibrations, 89, 95, 122 Victoria, 49, 5 I Viruses, 148 Voltage, 109

Wales, 124, 151, 220 Water, 103, 152

conservation, 109, 110 infiltration rates, 195 loss, IIO, 152 relationships, 109--I 2

uptake, I10, 154 Water stable aggregates, 187, 190, 192, 193

Weathering, 169, 191 Weight, 86, 139

live, 123, 167, 173, 218 seasonal increases in, 86 wet, 218

Westmorland, 124 West Indies, 50, 51, 52, 53, 55, 56, 66, 67 Wheat, 160, 196, 206, 212 Wide tube, 29, 102, 103 Withdrawal reactions, 88 Woodland, 123, 124, 145, 156, 159, 161,

168, 169, 172, 173, 177, 179, 233 beech, 124 mixed, 124 oak, 124, 144 species, 146

Worm cairns, 89 Wound tissue, 114 Wych Elm, 166

Yeasts, 184, 186 Yield increases, 195, 196, 197 Yolk, 40, 76 Yucatan, 51

Zambesia, 57 Zanzibar Island, 54 Zinc, 220, 22 I Zinophos, 214 Zurich, 144 Zygolobous prostomium, 2, 4, 57, 248

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A simplified key to common genera of terrestrial earthworms

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