A simplified key to common genera of terrestrial earthworms

A simplified key to common

genera of terrestrial earthworms

1. Clitellum commencing in front of the 15th segment (not found in the British Isles) 2

- Clitellum commencing after the 15th segment

2. Setal arrangement perichaetine (Fig. 2)

- Setal arrangement lumbricine (Fig. 2)

3. Prostomium tanylobous (fig. 1), setae closely paired (Fig. 2) at least over part of the body

- Prostomium epilobous (fig. 1) or if tanylobous, setae widely paired or distant (Fig. 2) over the whole body

4. Clitellum begins on segment 24

- Clitellum begins on or after segment 26 5. Clitellum ends after segment 28

- Clitellum ends before segment 28 6. Tubercula pubertatis absent, or exceptionally, present

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

papillae (plate Sb) 7. Setae widely-paired or distant, at least posteriorly

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

the body, tubercula pubertatis as ridges (excep

tionally as separate tubercles) extending over only part of the length of the clitellum

•• Only one common species.

3 (LUMBRI-

C ID AE)

Pheretima (MEGASCO

LECIDAE)

Diplocardia (ACANTHO

DRILID A E)

4

5 Bimastos eiseni (Levinsen)•• Lumbricus 6 Eiseniella

Bimastos

7 8 9

Dendrobaena

KEY TO GENERA OF TERRESTRIAL EARTHWORMS 213

-Setae closely-paired anteriorly (in the region of the hearts), distant posteriorly. Tubercula pubertatis as ridges as long or longer than the clitellum

9. Spermathecal pores (Fig. 3) in line with seta 'd' or more often near the mid-dorsal line, body trapezoidal in crossection

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

10. Prostomium with longitudinal ridges - Prostomium without longitudinal ridges 11. Calciferous glands with two lateral pouches in segment

Terrestrial

- Calciferous glands without lateral pouches. Amphibious

Octolasium

Eisenia

10 Eophila 11

Allolobophrna Helodrilus

Note. Internal characteristics have been used only in couplet 11. However, most Helodrilus species are amphibious, living in mud at the bottom of ditches, ponds, etc., and in wet river banks, whereas Allolobophrna species are terrestrial. Therefore the habitat can usually be used as the key character thus avoiding the necessity of dissection.

Simple key to species of terrestrial earthworms

Most common .and widespread species of earthworms can be identified using the characters in the first two or three columns of the key. If rarer species that are not included in the key are found, they may be misidentified on the basis of the characters in the first three columns, but reference to the additional descriptions given should usually be sufficient to avoid confusion.

Genus Pheretima MEGASCOLECIDAE

Two pairs of spermathecal pores in segmental grooves 7/ 8 and 8/9. 1st dorsal pore 11/12. 70- 170 mm. 10- 150 segs. Reddish-brown, clitellum creamy to dark grey. 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 in segmental grooves 5/ 6, 6/ 7, 7/8 and 8/9. 1st dorsal pore 12/ 13. 100- 160 mm. 120- 150 segs. Rich brown.

ACANTHODRILIDAE Genus Diplocardia

Pheretima californica (Kinberg) P. hupiensis (Michaelsen)

P.posthuma (Vaillant)

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

(Ude)

214 BIOLOGY OF EARTHWORMS

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

7/8 and 8/9. 180-300 mm. 125-160 segs. Anterior dorsal surface (Garman) pale flesh coloured.

Two pairs of spermathecal pores in segmental grooves 7/8 and D. riparia 8/9. 200-270 mm. 135-160 segs. Anterior dorsal surface dark (Smith) brown.

LUMBRICIDAE Genus Lumbricus

Tuberculat 1st dorsalt Clitellumt pubertatis pore

26, 27-32 28-31 7/8

28-33

31, 32-37

33, 34-39

29-33

33-36

34, 35, 36-38

6/7

7/8

5/6

Genus Eiseniella

22,23-26, 27

23-25, 26 4/5

• British species.

Red/brown or red/violet, irridescent dorsally, pale yellow ventrally. 25-105 mm. 95-120 segs.

Lumbricus rubellus• Hoffmeister

Chestnut to violet L. castaneus • brown; brown/yellow (Savigny) ventrally, irridescen t, clitellum orange. 30-70 mm. 82-100 segs.

Setae widely-paired both L. terrestris • ends of the body, Linnaeus strongly pigmented, brown-red dorsally, yel-lowish ventrally. 90-300 mm. 110-160 segs.

Red-brown, lighter ven- L.festivus• trally, irridescent dor- (Savigny) sally, prominent clitel-lum. Not found in large numbers. 48-108 mm. 100-143 segs.

Male pores in 13. Dark Eiseniella brown, greenish, golden tetraeda f . yellow, red. Body typica • quadrangular behind (Savigny) the clitellum. 30-60 mm. 60-90 segs.

Male pore in 15. Otherwise as f. typica

t for numbering see p. 12.

E. tetraeda (Savigny) fhercynia• (Michaelsen)


Genus Bimastos

Tubercula C/ite/lum

20, 21, 22-29,30

22-29

23-28

23, 24, 25-31, 32

23-32 or 24-33

24,25-32, 33

25, 26, 27-30, 31, 32, 33

27-37

pubertatis Absent

27,28

Absent

24, 25, 26-30

or absent

Absent

Absent

28, 29-30, 31 or

absent

Absent

Genus Dendrobaena 25,26-31, 28-30 32

1st dorsal pore

5/ 6

5/ 6

5/ 6

5/ 6

Setae ab > cd. Red- Bimastos brown. giese/eri 105-115 segs. 50-80 (Smith) mm.

Reddish-brown. Eise segs 30-6. 20-50mm

20-50 mm. 40-60 segs. up to 75 mm.

B. tumidis

B. palustris (Moore)

Reddish dorsally, yel- B. parvus lowish ventrally. 25--40 (Eisen) mm. 90-110 segs.

Rose-red. 98-122 segs. 60-90 mm.

B. longicinctus (Smith& Gittins)

Prostomium tanylobous, B. eiseni• body cylindrical, reddish (Levinsen) or violet dorsally, yel-lowish ventrally, clitel-lum red. 30- 64 mm. 75-111 segs.

Setae widely paired, dor- B. tenuis•t sally red-brown with (Eisen) bluish tint. Intersegmen-tal grooves and ventral side light. 20-85 mm. 90-120 segs.

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

Rosy to deep red, last posterior segments yellow. 27-90 mm. 50-100 segs.

Dendrobaena subrubicundat (Eisen)

• British species. t Now Dendrobaena rubida (Sav.) f. tenuis (Eisen). l Now Dendrobaena rubida (Sav.) f. subrubicunda (Eisen).


Tubercula 1st dorsal Clitel/um pubertatis pore

25,26-28, 25-30 5/6 or Smoky-grey with red D.platyura 29,30 sometimes pigment posteriorly. 80- (Fitzinge11)

8/ 9 180 mm. 120-160 segs.

26, 27-31, 29-30 5/6 Dark red dorsally, lighter D. rubida• 32 red ventrally. 30-60 mm. (Savigny)

50-100 segs.

24,25-26, 30 and 31 5/ 6 Violet, purple or olive D. veneta 27-32, 33 brown, dorsal pigment f. typica

bands separated by non- (Rosa) pigmented zones. Some-times uniformly unpig-men ted. 50-155 mm. 80-225 segs.

27, 28, 31- 32, 33 4/ 5 Red, violet, yellow or D. octaedra • 29-33, 34 copper. Posterior octa- (Savigny)

gonal. 17-40 mm. 79-95 segs.

28-33, 34 29,30-32 5/ 6 Dorsally pale red, 1st D. attemsi segment and ventrally (Michaelsen) and clitellum white. 20-50 mm. 100-150 segs.

31-36 33-34 4/ 5 Red-violet, slightly irri- D. mamma/is • descent. 30-65 mm. 83- (Savigny) 100 segs.

33- 37 34, 35- 36, Undetect- Red dorsally or unpig- D.pygmaea• 37 able men ted. 30- 32 mm. (Savigny)

103-180 segs.

Genus Octolasium

29-34 30-33 11/ 12 Blue-grey with (usually) Octolasium lilac-blue dorsal line. Last cyaneum 4- 5 segments yellow, (Savigny) anterior segments pink, clitellum red-orange. 50-160 mm. 100- 150 segs.

30-35 31-34 8/9, 9/ 10 White, grey, blue or O.lacteum• or 10/ 11 rose-pink, clitellum pink (Oerley)

or orange. 30-160 mm. 90-180 segs.



Tubercula 1st dorsal Clitellum pubertatis pore

Genus Eisenia

24, 25, 28-30, 31 4/ 5 Red, purple or brown; Eisenia 26-32 yellowish ventrally. Dor- foetida•

sal surface pigment al- (Savigny) temating with light inter-segmental zones. 32-130 mm. 80-110 segs.

Genus Eophila

33, 34 35, 36, Yellowish or grey. 52- Eophila 35-42,43, 37-41, 42, 140 mm. 132-170 segs. icterica • 44 43,44 (Savigny)

Genus Allolobophora

25, 26-31, 29-30, 31 4/ 5 Pale red, without pig- Allolobophora 32, 33 or 30- 32 ment, prominent and rosea•

flattened clitellum. 25- (Savigny) 85 mm. 120-150 segs.

26-32 28, f29 5/ 6 Grey, unpigmented, A. culpifera andf29, 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 men ted. 50- 90 mm. (Michaelsen) 100-130 segs.

26, 27, 31 and 33 11/12 Anterior, especially the A. caliginosa 28-34, 35 or 12/ 13 first few segments pink, f typica•

otherwise pale grey, (Savigny) yellowish 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 trapezoides connected by a narrow (Duges) bridge.

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) ISO mm. 160-200 segs.



TubeTcula 1st dorsal Clitellum pubeTtatis pore

27,28-35 32-34 Body cylindrical, colour A.ltmga• 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, eli-

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

27,28-34, 31-35 10/ 11, 55-100 mm. 152-194 A. tubeTculata 35 11/ 12 segs. Body cylindrical, Eisen

or 12/13 unpigmented, greyish.

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

28, 29-37 31 and 33 4/ 5 Light or dark green, yel- A. chkJrotico • and 35 low, grey, pink, slate- (Savigny)

blue, clitellum pink, green or grey. 30-70 mm. 80-138 segs.

Genus Helodrilus 21, 22-32 29-30 4/ 5 Setae black in fully Helodrilus

mature individuals, flesh- oculatus• coloured, body unpig- Hoffmeister mented. 35- 75 mm. 95-150 segs.


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Indexes

Systematic index

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

Acanthodrilacae, 43 Acanthodrilidae, 39, 42, 43, 45, 46,

54, 55, 62, 212, 213 Acanthodrilus, 38, 45, 54 Actinomyces, 159 Aeogaster, 47 Aeolosomatidae, 13, 38 aerophilus, Thominx, 195 africanus, Hyperiodrilus, 120, 121,

125, 129, 131 Ocnerodrilus, 194

agglutinatus, Bacillus, 158 agrestis, Pheretima, 88 Aikinetocystis, 124 A llolobophora, 52, 57, 66, 107, 196,

213,217 antipae, 27, 217 caliginosa f. trapezoides, 23, 217 caliginosa, 1, 3, 5, 8, 53, 56, 58,

59, 60, 68, 70, 71, 97, 104, 105, 107, 108, 109, 110, 111, 112, 113, 114, 115, 118, 122, 125, 126, 128, 129, 131, 132, 135, 136, 137, 138, 139, 140, 145, 146, 148, 149, 151, 162, 169, 170, 171, 173, 194, 195, 197, 198, 199, 206, 207

caliginosa f. typica, 217 chlorotica, 3, 11, 56, 58, 59, 60,

61, 68, 91, 92, 98, 104, 106, 107, 108, 109, 111, 112, 113, 114, 122, 125, 126, 127, 128, 129, 135, 139, 144, 171, 185, 188, 197, 198, 199, 204, 206, 207,208,218

culpijera, 217 dubiosa, 194 japonica, 116 limicola, 208 longa, 57, 58, 59, 61, 68, 89,

91, 107, 109, 111, 118, 119, 122, 123, 125, 126, 128, 129, 131, 135, 136, 137, 138, 139, 145, 146, 165, 166, 185, 192, 195, 198, 199, 201, 207, 208, 217

miniscula, 217 nocturna, 57, 58, 68, 70, 71, 104,

105, 108, 109, 110, 111, 113, 118, 122, 125, 126, 135, 199, 201, 207, 218

ribaucourti, 217 robusta, 123 rosea, 9, 10, 58, 59, 60, 66, 68,

105, 107, 108, 109, 110, 111, 112, 113, 115, 122, 125, 126, 128, 135, 139, 185, 198, 199, 204,206,207,208,217


Allolobophora--{ con td.) terrestris, 123, 217 terrestris, f. longa, 60 tuberculata, 208, 218 vogesiana, 217

Alma, 50, 62, 132 Amoebotaenia cuneatus, 194

lumbrici, 194 Andiorrhinus, 49 Andioscolex, 50 annulata, Capillaria, 194 Anoplophyra, 124 Anteoides, 50 antipae, Allolobophora, 27, 217 Aptodrilus, 49 Ascaridia galli, 196 Ascaris suum, 196 Aspidodrilus, 123 attesmi, Dendrobaena, 144, 206,

216 australis, Megascolides, xi, 19, 66 Azotobacter, 147, 156

Bacillus agglutinatus, 158 cereus, 155, 158 botulinus, 124 cereus var. mycoides, 157 idosus, 158

Barogaster, 47 Bassia longifolia, 100 beddardi, Bimastos, 125 Beddardiella, 48 Benhamia, 47 Benhaminae, 43 Bimastos, 52, 212, 215 Bimastos beddardi, 125 Bettonia, 48 Bimastos eiseni, 8, 11, 107, 122, 125,

126, 133, 137, 139, 144, 207, 208,212,215

gieseleri, 215 icenorum, 208 longicinctus, 215

liinnbergi, 125, 215 muldali, 208 palustris, 215 parvus, 19,208,215 tenuis, 8, 215 tumidis, 215 zeteki, 62, 70, 72, 122, 215

Bipalium kewensis, 120, 124 Borgertia, 48 botulinus, Bacillus, 124 bronchia/is, Cyathostoma, 193, 195 Bu"ttneriodrilus, 48

cal ifornica, Pheretima, 132, 213 cali"ginosa, Allolobophora, see

Allolobophora f. trapezoides, Allolobophora, 23,

217 f. typica, Allolobophora, 217

Callidrilus, 50 Capillaria annulata, 194

causinjlata, 19 5 mucronata, 19 5 plica, 195 putorii, 19 5

carnaria, Sarcophaga, 125 castaneus, Lumbricus, see Lumbricus causinjlata, Capillaria, 195 Causus rhombeatus, 123 Celeriella, 47 cereus var. mycoides, Bacillus, 157 Cestoda, 193, 194 Chaetopoda, 13, 37 Chilota, 45 chlorotica, Allolobophora, see

Allolobophora Chuniodrilus, 48 Clostridium, 156 coeli'color, Streptomyces, 159 Comarodrilus, 44 communis, Diplocardia, 214 controversa, Tilletia, 161, 192 corethrusus, Pontoscolex, 53, 196

SYSTEMATIC INDEX 255

crassum, Porrocaecum, 195 Craterocystis, 124 Criodrilidae, 39, 51 Criodrilinae, 51 Criodrilus, 51 Criodrilus lacuum, 62, 194 culpifera, Allolobophora, 217 cuneatus, Amoebotaenia, 194 cuneata, Taenia, 124 Curgia, 44 Curgiona, 44 cyaneum, Octolasium, see

Octolasium Cyathostoma bronchialis, 193, 195

Deccania, 45 Deinodrilus, 46, 55 Dendrobaena, 13, 52, 66, 68, 212,

215 attemsi, 144, 206, 216 mammalis, 57, 58, 123, 199, 208,

216 octaedra, 106, 107, 122, 125, 126,

133, 137, 139, 144, 207, 208, 216

platyura, 215 pygmaea, 208, 216 rubida, 67, 107, 122, 125, 126,

139, 144, 199, 206, 207,215 rubida f. tenuis, 66, 208, 215 rubida f. subrubicunda, 208 subrubicunda, 58, 59, 60, 61, 67,

69, 91, 92, 126, 129, 165, 199, 201,215

veneta var. hibernica f. dendroidea, 208

veneta var. hibernica f. typica, 208

veneta var. hortensis, 208 veneta f. typica, 208, 216 veneta var. zebra, 208

Desmogaster, 41 Dicelis, 124

Dichogaster, 4{), 46, 53, 54, 55, 77 jaculatrix, 121

Didymogaster, 44 sylvaticus, 19

Digaster, 44 Dilepus undula, 1 94 Dinodriloides, 45 Dioctophyma sp., 195 Diplocardia, 46, 54, 110, 131, 212,

213 communis, 214 egglestoni, 62, 68 riparia, 214 singularis, 213

Diplotrema, 38, 46 Diporochaeta, 46 Dtprochaeta, 124 Distichopus, 124 Drawida, 41

grandis, 118, 123 Drilocrius, 50 dubiosa, Allolobophora, 194 dunguensis, Shubotziella, 65

Echinocystis, 124 egglestoni, Dtplocardia, 62, 69 Eisenia, 52, 57, 213, 216

foetida, 13, 16, 18, 60, 61 , 65, 66, 67, 69, 75, 76, 77, 80, 81, 88, 95, 96, 123, 124, 125, 128, 131, 132, 133, 145, 159, 160, 162, 172, 194, 195, 196, 197, 199, 206, 207, 208, 216

icterica, 208 metallorum, 216 rosea, 60 veneta, 194

eiseni, Bimastos, see Bimastos Eiseniella, 51, 66, 212, 214

tetraedra, 199, 207 tetraedra f. hercynia, 208, 214

Eleusine coracana, 196


elongata, Pheretima, 196 elongatus, Metastrongylus, 194 Eminoscolex, 48 Enantiodrilus, 50 Enchytraeidae, 9, 10, 39 ensicaudatum, Porrocaecum, 19 5 Entyphoeus waltoni, 65 Eodrilus, 46 Eophila, 52, 213, 217

(Allolobophora) icterica, 126, 217 obscuricola, 217

Escherichia coli, 157 Eudi'chogaster, 46 Eudrilidae, 39, 47, 53, 54 Eudrilinae, 48 Eudriloides, 48, 65 Eudrilus, 48, 72

eugeniae, 53 eugeniae, Eudrilus, 53 Eukerria, 44 Eupolygaster, 41 Eupolytoreutus, 48 Euscolex, 48 Eutoreutus, 48 Eutrigaster, 47 Eutyphoeus, 46, 77 excavatus, Pheretima, 77 Exxus, 44

f esti'vus, Lumbricus, 67, 107, 138, 199,208,214

Fimoscolex, 50 foetida, Eisenia, see Eisenia Fraxinus pennsylvanica, 170 Frideri'cia parasitica, 123 jriendi, Lumbricus, 208 Fusarium, 161, 192

galli, Ascaridia, 196 Gardullaria, 48 gieseleri, Bimastos, 215 Glossoscolecidae, xi, 9, 21, 22, 39,

49, 53, 55, 62

Glossoscolecinae, 4, 49, 54 Glossoscolex, 50 Glyphidrilus, 50 Gordiodrilus, 45 grandis, Drawida, 118, 123 Grayallia, 124 Gregarina, 124

haemorrhoidalis, Sarcophaga, 125 hali'otidea, Testacella, 123 Haplodrilus, 45 Helodrilus, 52, 212, 218

oculatus, 208, 218 herculeus, Lumbricus, 195 Heterakis, 124 Heterodera rostochiensi's, 192 hilgendorj, Pheretima, 62 Hippopera nigeriae, 154 Hirudinea, 37 Histomonas, 194 Hoplitophyra, 124 Hoplochaetella, 46 Hoplochaetina, 46 Hormogaster, 51, 54

redii f. gigantes, 54 Hormogastridae, 39, 51, 54 Hormogastrinae, 51 Holoscolex, 50 Howascolex, 46 hupeiensis, Pheretima, 62, 110, 115,

118, 121, 128, 132, 133, 213 Hyperiodrilus, 48, 66, 113

ajricanus, 120, 121, 125, 129, 131

Hystrichis tricolor, 194

icterica, Eisenia, 208

Eophila (Allolobophora), 126, 217 idosus, Bacillus, 158 inaequalis, Ventura, 161 Iridodrilus, 48

jaculatrix, Dichogaster, 121


Kaffania, 48 Kerria, 44 Kerriona, 45 kewensis, Bipalium, 120, 124 Kynotus, 50

lacteum, Octolasium, 67, 107, 139, 156, 196, 197, 199,208,216

lacuum, Criodrilus, 62, 194 Lampito, 44, 77 latus, Protoscolex, 38 Legonea,48 Lennogaster, 47 Leptodrilus, 45 Leucodrilus, 47 Libyodrilus, 48 limicola, Allolobophora, 208 longa,Allolobophora, see

Allolobophora longifolia, Bassia, 100 liinnbergi, Bimastos, 125, 215 longicinctus, Bimastos, 215 Lumbricidae, xi, 9, 10, 11, 12, 20,

39, 40, 51-3, 54, 64, 145, 212 lumbrici, Polycercus, 194 Lumbriculidae, 3, 37, 38 Lumbricus, 5, 13, 18, 23, 28, 52, 66,

67, 68, 69, 91, 118, 212, 214 castaneus, 57, 58, 59, 60, 66, 67,

98, 105, 107, 122, 126, 135, 137, 138, 145, 199, 207, 208, 214

festivus, 67, 107, 138, 199, 208, 214

friendi, 208 herculeus, 195 terrestris, 4, 5, 7, 12, 13, 16, 20,

21, 24, 60, 61, 63, 66, 67, 69, 10, 71, 72, 13, 74, 75, 76, 77, 79, 80, 84, 87, 89, 91, 92, 95, 97, 104, 105, 108, 109, 110, 113, 118, 122, 125, 126, 127, 129, 130, 131, 134, 135, 136, 140, 141, 142, 145, 146, 149,

151, 152, 155, 156, 157, 158, 161, 165, 169, 171, 185, 187, 189, 194, 195, 196, 197, 198, 199, 200, 201, 203, 204, 206, 201,208,209,214

rubellus, 51, 58, 59, 60, 66, 67, 89, 91, 92, 106, 107, 122, 126, 129, 130, 136, 138, 144, 145, 146, 156, 165, 169, 194, 195, 197, 199,206,201,208,214

Maheina, 44 Malabaria, 44 Malodrilus, 48 mamma/is, Dendrobaena, 51, 58,

123, 199, 208, 216 Maoridrilus, 45 marcessens, Serratia, 157 maugei, Testacella, 123 Maupasella, 124 mays, Zea, 196 Megachaetina, 48 Megadrili, xi, 39 Megascolecidae, 7, 9, 10, 20, 21, 22,

38, 39, 41, 42, 43, 53, 54, 55, 123, 213

Megascolecinae, 43 Megascolex, vii, 27, 38, 44, 55, 125 Megascolides, 46, 54 australis, xi, 19, 66 merulae, Syngamus, 195 mesomelinus, Quedius (Micro-

saurus), 123 Metadrilus, 48 Metaradiophyra, 124 Metascolex, 48 Metastrongylus, 124, 193

elongatus, 194 pudentotectus, 194 salami, 194

Microchaetinae, 50, 55 Microchaetus, 50, 119 Microdrili, xi, 39, 41


Microscolex, 45, 54 modestus, 196 phosphoreus, 53

millardi, Perionyx, 119 Mil/sonia, 47 miniscula, Allolobophora, 217 modestus, Microscolex, 196 Moniligaster, 41 Moniligastridae, 38, 39, 40, 123 Moniligastrinae, 41 Monocystis, 19, 124 Monogaster, 47 mucronata, Capillaria, 195 Myxocystis, 124

Naididae, 3, 38 Nellocystis, 124 Nellogaster, 44 Nelloscolex, 44 Nematocystis, 124 Nematoda, 124, 194 Nematogenia, 45 Nemantodrilus, 48 Nenmanniella, 48 Neochaeta, 44 Neodrilacae, 43 Neodrilus, 45 nigeriae, Hippopera, 154 Nocardia polychromogens, 159 nocturna, Allolobophora, see Allolo-

bophora Notoscolex, 44, 121

termiticola, 123 Notykus, 48

Ocnerodrilidae, 39, 42, 43, 44, 45, 55 Ocnerodrilinae, 43 Ocnerodrilus, 45

africanus, 194 octaedra, Dendrobaena, see Dendra

haena Octochaetidae, 39, 42, 43, 46 Octochaetus, 46

Octolasium, 52, 57, 66, 140, 146, 189, 212, 216

cyaneum, 58, 67, 107, 108, 115, 122, 126, 135, 139, 185, 199, 206, 207, 208, 216

lacteum, 67, 107, 139, 156, 196, 197, 199,208, 216

Oligochaeta, xi, 6, 37, 38, 39 Onesia sepulchralis, 125

subalpina, 125 Onychochaeta, 49 Opisthodrilus, 50

palustris, Bimastos, 215 Parabursaria, 124 Parachilota, 46 paradoxa, Paricterotaenia, 194 Parascolex, 48 parasitica, Fridericia, 123 Parendrilinae, 47 Parendrilus, 48 Paricterotaenia paradoxa, 194 parvus, Bimastos, 19, 208,215 Paulistus, 44 peguana, Pheretima, 194 Periodrilus, 45 Perionyx, 7, 44, 77

millardi, 119 Periscolex, 50 Perissogaster, 44 Phalaris, 139 Pheretima, 7, 8, 18, 22, 27, 40, 44,

53, 55, 61, 65, 66, 72, 81, 82, 88, 91, 116, 124, 157, 194, 197, 212, 213

agrestis, 88 californica, 132, 213 elongata, 196 excavatus, 77 hilgendorf, 62 hupeiensis, 62, 110, 115, 118, 121,

128, 132, 133, 213 peguana, 194


posthuma, 35, 119, 213 stelleri, 8

phosphoreus, Microscolex, 53 plica, Capillaria, 195 Plagiochaeta, 45 Plagiotoma, 124 Platydrilus, 47 Platyhelminthes, 123 platyura,L>endrobaena, 215 Pleurocystis, 124 Pliconogaster, 44 Plutellus, 38, 46, 54 Pollenia rudis, 124 Polycercus, 124

lumbrici, 194 Polychaeta, 37 polychromogens, Nocardia, 159 Polytoreutus, 48 Pontodrilus, 46 Pontoscolex, 4, 50

corethrusus, 53, 196 Porrocaecum crassum, 195

ensicaudatum, 19 5 posthuma, Pheretima, 119, 213 Priodochaeta, 47 Priodoscolex, 47 Pronaidites, 38 Protoscolex, 3 7

latus, 38 Protozoa, 124, 194 pudentotectus, M etastrongylus, 194 putorii, Capillaria, 195 pygmaea, L>endrobaena, 208, 216 Pygmaeodrilus, 45, 55 Pythium, 161, 192

Quechua, 44 Quedius (Microsaurus) mesomelinus,

123 Quercus robur, 1 70

Ramiella, 46 redii f. gigantes, Hormogaster, 54

Rhabditis, 124 Rhinodrilus, 49 Rhododrilus, 45 Rhynocystis, 124 ribaucourti, Allolobophora, 217 riparia, L>iplocardia, 214 Rillogaster, 47 robur, Quercus, 170 robusta, Allolobophora, 123 Rosadrilus, 48 rosea, Allolobophora, see Allolo-

bophora Rotatoria, 124 rubellus, Lumbricus, see Lumbricus rubida, L>endrobaena, 67, 107, 122,

125, 126, 139, 144, 199, 206, 201,215

rubida f. tenuis, L>endrobaena, 66, 208,215

rudis, Pollenia, 124

salami, Metastrongylus, 194 Sarcophaga carnaria, 125

haemorrhoidalis, 12-5 striata, 125

Schubotziella, 48 dunguensis, 65

Scolioscolides, 47 scutulum, Testacella, 124 sepulchralis, Onesia, 125 Serratia marcessens, 157 singularis,L>iplocardia, 213 skrjabinomorpha, Syngamus, 124 Sparganophilinae, 50 Sparganophilus, 50 Spenceriella, 46 Sphaeractinomyzon, 124 Spirochaeta, 124 Spiroptera, 124

turdi, 195 stelleri, Pheretima, 8 Stephanurus, 124, 195 striata, Sarcophaga, 125


Streptomyces coeUcolor, 159 subalpina, Onesia, 125 subrubicunda, Dendrobaena, see

Dendrobaena suum, Ascaris, 196 sylvaticus, Didymogaster, 19 Syngamus, 124

merulae, 19 5 skrjabinomorpha, 195 trachea, 193, 195

Syngenodrilinae, 41 Syngenodrilus, 41 Synoecnema, 124

Taenia cuneata, 124 Teleudrilus, 48 tenuis, Bimastos, 8, 215 termiticola, Notoscolex, 123 terrestris, Allolobophora, 123, 217 terrestris, Lumbricus, see Lum-

bricus Testacella haliotidea, 123

maugei, 123 scutulum, 124

tetraeda, Eiseniella, 199, 207 f. hercynia, Eiseniella, 208, 214 f. typica, Eiseniella, 139, 208, 214

Thamnodrilus, 49 Thelohania, 124 Thominx aerophilus, 195

Tilletia controversa, 161, 192 Tonoscolex, 47 trachea, Syngamus, 193, 195 Travoscalides, 47 tricolor, Hystrichus, 194 Trigaster, 46, 54 Tritogenia, 50 Trocheta subviridis, 124 Tubificidae, 38 tumidis, Bimastos, 215 turdi, Spiroptera, 195

Udeina, 46 undula, Dilepus, 194

veneta, Eisenia, 194 f. typica, Dendrobaena, 208, 216

Ventura inaequalis, 161 vogesiana, Allolobophora, 217

waltoni, Entyphoeus, 65 Wegeneriella, 47 Woodwardiella, 44

Yagansia, 45

Zapotecia, 46, 54 Zea mays, 196 zeteki, Bimastos, 62, 70, 72, 122,

215

Author index

Agarwal, G. W., 196, 219 Aichberger, R. von, 157, 219 Aisyazhnyuk, A. A., 219 Alicata, J. E., 194, 246 Allee, W. C., 125, 219 Allen, R. W., 219 Anderson, H. L., 237 Anstett, M., 162, 219 Arbit, J., 76, 219 Arldt, T., 38, 219 Arlidge, G. Z., 183, 184, 234 Arnold, M. K., 183, 226, 232 Arrhenius, 0., 125, 219 Arthur, D. R., 81, 93, 179, 180, 208,

219 Arthur, J. H., 227 Ash, J. S., 224 Aspi:ick, H., 184, 236 Atlavinyte, 0., 156, 219, 220 Avel, M., xiii, 219

Bachelier, G., xiii, 20, 125, 137, 180, 220

Bahl, K. N., 18, 61, 83, 118, 220 Baker, W. L., 220 Bakhtin, P. U., 165, 166, 220 Baldwin, F. M., 74, 77, 220 Ball, D. F., 242 Ball, R. C., 220 Baluev, V. K., 168, 220 Banage, W. B., 104, 221 Barker, R. J., 183, 185, 220

Barley, K. P., 98, 103, 117, 138, 139, 140, 145, 147, 148, 149, 151, 162, 176, 177, 196, 220,221

Barrett, T. J., 221 Bassalik, K., 155, 157, 163, 165, 221 Bather, E. A., 38, 221 Bauer, K., 181, 221 Baweja, K. D., 161, 221 Baylis, H. A., 121, 123, 221 Beauge, A., 122, 221 Bejsovec, J., 196, 221 Belousova, N. K., 103, 226 Benham, W. B., 11, 26, 38, 45, 46,

50,221 Beynon, K., 227 Bharucha-Reid, R. P., 76, 221 Bhatt, J. V., 147, 157, 161, 234 Bigger, J. H., 182, 221 Blancke, E., 147, 163, 221 Blankwaardt, H. F. H., 181, 221 Block, W., 104, 221 Bocock, K. L., 222 Bodenheimer, F. S., 168, 222 Bollen, W. B., 153, 242 Bornebusch, C. H., 98, 125, 222 Bouche, M. B., 102, 222 Boyd, J. M., 107, 137, 222 Boynton, D., 133, 222 Bradley, J., 165, 168, 223 Braunig, G., 80, 88, 238 Bray, J. R., 146, 222 Bretnall, G. H ., 222


Bretscher, K., 98, 222 Breza, M., 194, 222 Brown, B. R., 140, 222 Brown, D. M., 222 Bruel, W. E. van der, 160, 222 Briisewitz, G., 157, 158, 159, 222 Buahin, G. K. A., 181, 221 Buntley, C. J., 223 Byzova, Yu B., 182, 183, 223, 230

Capstick, C. K., 222 Carter, G. S., 96, 223 Causey, D., 223 Cernosvitov, L., 52, 123, 223 Chadwick, L. C., 165, 168, 223 Chandler, R. F., 100, 226 Chapman, G., 223 Chen, C. M., 190, 223 Cockerell, T . D. A., 125, 223 Cohen, S., 148, 223, 224 Coin, C. J., 223 Combault, A., 23, 169, 223 Compton, 0. C., 133, 222 Conder, P. J., 185, 224 Cossens, G. G., 168, 170, 173, 247 Couperus, H., 79, 225 Couture, G., 189, 237 Cragg, J. B., 224 Cramp, S., 185, 224 Crompton, E., 179, 180, 224 Crossley, D. A., 140, 146, 188, 224,

227 Curry, L. L., 220

Darwin, C., xii, 74, 88, 121, 139, 196,224

Davey, S. P., 182, 224 Davis, B. N. K., 185, 186, 224 Dawson, A. B., 32, 157, 165, 166,

224 Dawson,R. C., 224 Day, G. M., 147, 157, 158, 224 Decker, G. C., 182, 186, 221, 237

Dennis, E. B., 182, 183, 226 Devigne, J., 224 Dhawan, C. L., 224 Dhennin, L., 193, 224 Dingwall, A. R., 107, 170, 231 Doane, C. C., 182, 183, 185, 225 Dobson, R. M., 171, 225 Doeksen, J., xiii, 69, 77, 79, 99, 126,

225, 229 Doerell, E. C., 180, 225 Dotterweich, H., 127, 225 Dowdy, W. W., 110, 131, 225 Dreidax, L., 169, 226 Drift, J. van der, xii, 181, 184, 225,

226, 229 Dubash, P. J., xiii, 64, 65, 78, 96,

248 Duley, F. L., 248 Dustman, E. H., 185, 226 Dutt, A. K., 165, 166, 173, 226 Dzangaliev, A. D., 103, 226

Eaton, T . H., Jr, 100, 226 Eberhardt, A. 1., 125, 226 Edwards, C. A., 99, 103, 140- 2, 144,

150, 174, 175, 181-8, 190, 204, 221, 224, 226, 227, 232

El-Duweini, A. K., 98, 104, 126, 128, 132, 136, 226, 227

Ellenby, C., 192, 227 Ellis, D. J., 237 Empson, D. W., 226 Escherich, K., 123, 227 Escritt, J. R. , 179-81, 227 Evans, A. C., 52, 56- 60, 66, 68-71,

100, 104, 105, 110--13, 118, 119, 122, 123, 128, 129, 130, 138, 145, 163, 167, 174, 176, 197- 9

Feldkamp, J., 6, 228 Fenton, G. R., 228 Finck, A., 127, 153, 165, 228 Fleming, W. E., 183, 228

AUTHOR INDEX 263

Ford, J., 98, 228 Fox, C. J. S., 181, 228 Franz, H., 146, 228 French, M. C., 186, 224

Gansen, P. S. van, 80, 82, 228 Garner, M. R., 118, 206, 228 Gast, J., 139, 228 Gates, G. E., 39, 42-7, 51, 69, 77,

115, 121, 229 Genov, T., 194, 229 Geoghegan, M. J., 166, 229 Gerard, B. M., 56, 60, 68, 106, 108,

109, 112, 114, 115, 127-9, 229 Gersch, M., 229 Gewehr, H., 245 Ghabbour, S. 1., 98, 104, 121, 126,

128, 132, 132, 136, 160, 226, 227, 229

Ghilarov, M. S., 157, 168, 172, 182, 183, 229, 230

Giesecke, F., 147, 163, 221 Gilbert, 0., 222 Gish, C. D., 186, 187, 230 Glasgow, L. L., 187, 247 Goffart, H., 183, 184, 230 Gorham, E., 146, 222 Graff, 0., xiii, 60, 130, 132, 148,

153, 154, 174, 176, 206, 230 Grant, W. C., 62, 106, 115, 118, 127,

131, 132, 172, 230 Grassi, B., 194, 230 Gray, J., 94, 230 Greenwood, D. E., 183, 230 Griffiths, D. C., 184, 231 Grigor'eva, T. G., 182, 231 Grove, A. J., 14, 15, 17, 25, 26, 29,

34, 231, 240 Guild, W. J., MeL., 137, 138, 145,

165, 167, 174, 176, 197-9, 227, 228,231

Gunthart, E., 182, 231 Gurianova, 0. Z., 165, 231

Hadley, C. H., 183, 228 Hamblyn, C. J., 107, 170, 231 Handa, B. K., 224 Handley, W. R. C., 222 Hanel, E., 74, 231 Hanley, I. M., 228 Hardman, J. A., 185-7, 249 Harmsen, G., 161, 231 Harrison, R. B., 185, 186, 224 Hasenbein, G., 197, 231 Haswell, W. A., 124, 231 Heath, G. W., 103, 140-2, 144, 149,

179, 204, 226, 227, 231, 232, 235

Heck, L. von, 75, 76, 232 Heimburger, H . V., 232 Hensen, V., xii, 232 Herlant-Meewis, H., 87, 232 Hess, W . N., 28, 88, 232 Heungens, A., 144, 182, 184, 232 Hill, J. P., 124, 231 Hirst, J."M., 161, 232 Hobmaier, A., 194, 232 Hoeksema, K. J., 165, 232 Hoffman, J. A., 161, 192, 232 Hogben, L., 131, 232 Hollister, P. L., 219 Hopkins, A. R., 165, 174-7, 182-4,

232, 233 Hopkins, H. T., 233 Hopp, H., 98, 112, 115, 131, 165,

167, 16~ 171, 174-~ 233,246 Howell, C. D., 88, 233 Hoy, H. M., 182, 183, 233 Hub!, H., 87, 233 Hunt, L. B., 185, 186, 233 Hutchinson, S. A., 158, 161, 191,

233 Hyche, L. L., 184, 233 Hyman, L. H ., 69, 233

Inove, T ., 190, 233 Isa, A. L., 237


Jacks, G. V., 167, 233 Jacob, A., 179, 180, 233 Jacobson, G. M., 148, 153, 173, 237 Jeanson-Luusinang, C., 126, 158,

234 Jefferson, P., 100, 176, 179, 180,234 Jennings, A. C., 148, 149, 162, 220 Jevniaux, C., 224 Joachim, A. W. R., 165, 234 Johnson, M.,L., 79, 80, 234 Johnstone-Wallace, D. B., 99, 179,

234 Jongerious, A., 232 Joshi, N. V., 163, 168, 234

Kahsnitz, H. G., 169, 234 Kalmus, H., 234 Kamel, M., 158, 161, 191, 233 Kanwar, J. S., 165, 240 Karmanova, E. M., 194, 195, 234 Karpachevskii, L. 0., 144, 241 Keilin, D., 125, 234 Kelkar, B. V., 163, 168, 234 Kelly, W. A., 243 Kelsey, J. M., 183, 184, 234 Kevan, D. K. McE., xiii, 234 Khambata, S. R., 147, 157, 161, 234 King, H. G. C., 140, 142, 232, 235 Kirberger, C., 79, 235 Kirk, R. L., 131, 182- 4, 232 Kirk, V. M., 233 Kleinig, C. R., 145, 151, 221 Knop, J., 27, 235 Kobatake, M., 160, 235 Kollmannsperger, F., 235 Kollmannsperger, G., 112, 122, 130,

137,235 Kondo, K., 190, 233 Korschelt, E., 61, 235 Kozlovskaya, L. S., 156, 158, 235 Kring, J. B., 184, 235 Krivanek, J. 0., 76, 235 Kriiger, F., 78, 103, 235

Kubiena, W. L., 137, 235

Kiihnelt, W., xiii, 235 Kurcheva, G. F., 142, 235

Laan, H. van der, 184, 236 Ladell, W. R. S., 99, 235 Lahr, J. P., 219 Lakhani, K. H., 236 Lauer, A. R., 76, 236 Laverack, M. S., xiii, 16, 78, 83, 84,

89, 140, 236 Lawrence, R. D., 148, 236 Lee, K. E., 68, 42-4, 168, 197, 236 Legg, D. C., 182-4, 236 Leitenberger, L., 146, 228 Lesser, E. J., 79, 236 Lewis, H. B., 148, 223, 224 Lidgate, H. J., 183, 236 Lindquist, B., 139, 147, 236 Lipa, J. ]., 182, 236 Lissman, H. W., 94, 230 Liv, C. L., 223 Ljungstrom, P. 0., 119, 123, 197,236 Lofty, J. R., 103, 121, 131, 134, 135,

171, 174-6, 181-3, 188, 225-7, 231, 237, 243

Long, W. H., 183, 237 Love, C. W., 222 Low, A. J., 98, 165, 237 Lowe, D. G., 245 Luckman, W. H., 183, 186, 237 Lugauskas, A., 156, 220 Lukose, J., 123, 237 Lund, E. E., 194, 237 Lunt, H. A., 148, 153, 173, 237

Madge, D. S., 67, 104, 115, 121, 125, 129, 131, 146, 237

Magalhaes, P. S., 194, 237 Maldague, M., 189, 237 Mamajev, B. M., 172, 230 Mamytov, A., 165, 166, 237 Mangold, 0 ., 89, 139, 237

AUTHOR INDEX 265

Marapao, B. P., 237 Marshall, V. G., 171, 237 Martin, A. W., 84, 166, 238 Martin, J. P., 249 Mayne, D. W., 247 McCalla, T. M., 176, 248 McLeod, J. H., 123, 237 Meer, K. van der, 232 Meggitt, F. J., 194, 238 Mellanby, K., 238 Merker, E., 80, 88, 238 Meyer, L., 163, 238 Michaelsen, W., 37-40, 44-8, 50,

51,238 Michon,}., 60, 61, 69, 130, 238 Miles, H. B., 160, 238 Millar, H. R., 148, 236 Millott, N., 81, 238 Minderman, G., 225 Moment, G. B., 69, 95, 238, 239 Monnig, H. 0., 194, 239 Moore, A. R., 239 Moore, B., 52, 90, 125, 239 Morris, H. M., 99, 176, 239 Morrison, F. 0., 182, 239 Mozgovoy, A. A., 195, 239 Muldal, S., 239 Muller, G., xii, xiii, 139, 168, 239 Muller, P. E., 239 Murchie, W. R., 62, 68, 70, 72, 106,

122, 239, 240

Nakamura, Y., 115, 116, 125, 240, 248

Nathans, S., 98, 103, 243 Needham, A. E., 146, 148-50, 240 Negi, L. S., 219 Nelson, J. M., 98, 240 Newell, G. E., 14, 15, 17, 25, 26, 29,

34, 231, 240 Nicoi, H., 180, 240 Nielson, C. 0., 240 Nielson, R. L., 168, 170, 171, 240

Nijhawan, S.D., 165, 240 Nye, P. H., 122, 127, 153, 154, 240

O'Connor, F. B., 243 Ogg, W. G., 180, 240 Oldham, C., 240 Olson, H. W., 125, 127, 240 Omodeo, P., 42, 43, 241 Otanes, F. G., 190, 241

Panditesekera, D. G., 165, 234 Papedick, R. I., 223 Pare!, T. S., 241 Parker, G. H., 90, 241 Parle, J. N., 155, 156, 159, 160, 166,

241 Parshley, H. M., 90, 241 Patel, H. K., 183, 190, 241 Patel, R. M., 241 Pathak, A. N., 163, 246 Pattarudriah, M., 190, 196, 242 Peachey, J. E., 241 Peredel'sky, A. A., 188, 241 Perel, T. S., 144, 241 Perrier, 44-6, 48, 4Q. Petrov, B. C., 125, 241 Phillips, E. F., 125, 241 Pickard, J. A., 183, 185, 186, 247 Polivka, J. B., 181- 3, 241, 242 Polsky, M. N., 165, 166, 220 Pomerat, G. M., 79, 242 Ponomareva, S. 1., 153, 155, 165,

166, 176, 242 Poryadkova, N. A., 241 Powers, W. L., 153, 242 Prabhoo, N. R., 242 Pratt, K. C., 69, 248 Prosser, C. L., 88, 242 Puh, P. C., 127, 153, 242 Purdy, L. H., 161, 192, 232

Radionova, L. Z., 241 Raffy, A., 79, 242


Ragg, J. M., 242 Ralph, C. L., 79, 242 Ramsay, J. A., 83, 91, 242 Rao, K. S. K., 219 Raw, F., 99-101, 103, 104, 134, 141,

142, 144, 146, 181, 184, 231, 242, 243

Reichle, D. E., 224, 227 Reinecke, A. J., 119, 123, 197, 236 Reynoldson, T. B., 98, 103, 243 Rhee, J. A. van, 98, 103, 104, 146,

169, 171, 243 Rhoades, W. C., 183, 243 Ribaudcourt, E., 169, 243 Richards, J. G., 170, 243 Richardson, H. C., 125, 134, 179-

81, 244 Richter, G., 182, 183, 244 Robertson, J.D., 80, 244 Robinson, J. S., 75, 76, 244 Rodale, R., 180, 183, 244 Roots, B. 1., 82, 90, 92, 127, 129,

244 Rovelli, G., 194, 230 Ruschmann, G., 147, 244 Russell, E. J., xii, 133, 14 7, 168, 244 Rysavy, B., 194, 195, 244 Ryzhikov, K. M., 195, 244

Sacho, R. J., 186, 233 Salisbury, E. J., xii, 125, 127, 172,

244 Saroja, K., 79, 244 Sastry, K. S. S., 190, 196, 242 Satchell, J. E., xiii, 58, 60, 61, 661

68, 69, 89, 90, 98, 100, 102, 107, 108, 125, 126, 130, 133, 135, 137, 139, 140, 142, 146, 148, 149, 151, 152, 155, 160, 176, 230, 236, 240, 245

Saussey, M., 123, 168, 245 Scharpenseal, H. W., 245 Schmid, L. A., 87, 245

Schmidt, H., SO, 76, 245 Schneider, K. C., 31, 245 Schread, J. C., 121, 182- 4, 191, 246 Schrevan, D. van, 161, 231 Schwartz, B., 194, 246 Scopes, N. E. A., 184, 249 Scott, H. E., 184, 246 Scrickhande, J. C., 163, 246 Sharma, R. L., 224 Shindo, B., 246 Shiraishi, K., 133, 246 Sims, R. W., 42, 43, 66, 246 Singh, A., 224 Sison, P. L., 190, 241 Skarbilovic, T. S., 195, 246 Slater, C. S., 167, 169, 171, 175, 180,

181, 233, 246 Smallwood, W. M., 30, 86, 246 Smith, F ., 41, 187, 247 Smith, R. D., 247 Sokolov, D. F., 241 Stafford, C. J., 227 Stephenson, J., xii, 9, 12, 13, 16, 23,

24, 31, 33, 35, 37- 40, 42- 4, 46, 49, 62, 66, 79, 91, 94, 124, 197, 247

Stickel, L. F., 185, 226, 247 Stickel, W. H., 247 Stockdill, S. M. J., 168, 170, 173,

247 Stockli, A., 98, 122, 127, 147, 153,

155,.159, 167, 247 Stokes, B. M., 247 Stolte, H. A., xiii, 124, 196, 247 St0p-Bowitz, C., 247 Storey, I. F., 232 Stough, H. B., 31, 247 Stringer, A., 183, 185, 186, 247 Sun, K. H., 69, 248 Svendsen, J. A., 98, 103, 107, 138,

248 Swaby, R. J., 165, 166, 248 Swartz, R. D., 76, 210, 248

AUTHOR INDEX 267

Takano, S., 125, 248 Tembe, V. B., xiii, 64, 65, 78, 96,

248 Tenney, F. G., 161, 248 Teotia, S. P., 156, 163, 165-7, 176,

248 Thompson, A. R., 183, 184, 186,

227,248 Tischler, W., 176, 248 Torvik, M. M., 219 Tracey, M. V., 81, 248 Trifonov, D., 190, 248 Tromba, F. G., 195, 248 Twinn, D. C., 222

Uhlen, G., 169, 180, 248 Urquhart, A. T., xii, 248

Villot, F. C. A., 194, 249 Vogel, R., 194 Volz, P., 168, 249

Waid, J. S., 222 Waite, R. H., 249 Waksman, S. A., 161, 166, 248, 249 Walton, W. R., 99, 190, 249 Ward, W. C., 232 Waters, R. A. S., 112, 117, 139, 169,

170, 176, 249 Watkin, B. R., 179: 249

Way, M. J., 184, 249 Weber, G., 182, 184, 249 Wehr, E. E., 237 Weisbach, W. W., 197, 249 Went, J. C., 157, 249 Wheatley, G. A., 185-7, 249 Wherry, E. T., 125, 126, 249 Whiting, A. E., 227 Whitney, W. K., 184, 249 Wiegland, K., 179, 180, 233 Wilke, D. E. von, 60, 61, 98, 249 Wilcox, H. G., 232 Wingerden, C. G. van, 69, 126, 225 Witkamp, M., 150, 250 Wittich, W., 89, 150, 250 Wojewodin, A. W., 181, 250 Wolf, A. V., 90, 131, 250 Wollny, E., 167, 168,250 Woodhead, A. A., 195, 250 Woodman, M. G., 222

Yegorova, S. V., 241 Yerkes, R. M., 75, 76, 250

Zarrow, M. T ., 79, 242 Zhdannikova, E. N., 156, 158 Zhinkin, L., 96, 250 Zicsi, A., 98, 102, 175, 190, 250 Zrazhevski, A. I., 156, 157, 170,

250

General index

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

Absorption zone, 81 Accessory reproductive bodies, 7 Acetylcholine, 86, 96 Acid, 89 Acid-intolerant species, 125, 126 Acid soils, 88, 125, 127, 169, 178-

81, 201 Acid-tolerant species, 125, 126 Actinomycetes, 155, 156- 8 Activity, effects of temperature

on, 130 seasonal changes in, 113

Activity cycles, 87 Adders, 123 Adolescence, 70 Adrenaline, 87 Adverse conditions, 62, 74, 88 Adverse effects, on earthworms,

196, 197 on soil, 196, 197

Adverse periods, 67 Aeration, 106, 133- 6, 167, 168

effects of, 13 3 Aestivation, 63, 67- 9, 100, 127 Affinities, 37-9 Africa, 41, 44-8, 50-5, 67, 121 Age-class composition, lOS Aggregates, 165- 7

Aggregations, 106, 107, 131, 139, 165- 7,209

Agriculture, effects on earthworms, 174-89

Alaska, 54 Albumen cells, 4, 14, 15, 36 Alder, 146 Aldicarb, 184 Aldrin, 182, 187 Alfalfa, 165, 176 Algeria, 51 Alimentary canal, 17, 19-21, 124

innervation of, 30, 31 Alimentary plexus, 23 Allantoin, 81, 148 Alluvium, population in, 136, 137 Alpine forest, 146 America, Central, 46, 49, 50, 53-5

North, 46, 49-52, 54 South, 46, 47, 49,50,52- 4

Ammonia, 81- 4, 118, 127, 148 Ammonium, 149, 162 Ammonium sulphate, 181 Amoebocytes, 18, 19, 82 Ampulla, 27 Amylase, 81, 121 Anabiosis, 91 Anaerobic conditions, 77, 96

GENERAL INDEX 269

Anaerobic respiration, 79 Andes, 45, 50 Antarctica, 55 Antarctic Islands, 45 Antibiotic, 160 Anus, 4, 19, 69, 90 Apple scab, 161 Aquatic species, 9, 10 Aquatic worms, 38, 66, 91 Arable, 99, 103, 104, 122, 123, 131,

135, 166, 174, 176, 190 Arciform muscles, 63 Arctic forest, 146 Argentina, 49, 50 Artesian irrigation, 172 Arthropods, 14 7 Ascospores, 161 Asexual reproduction, 67 Ash, 140, 149, 150, 170 Asia, 44, 50-3, 55 Association, 123 Association neurones, 86 Atrazine, 181 Auckland, 45, 46 Australasia, 55 Australia, 44-6, 53, 55, 103, 115,

151, 176 Austria, 51 Autotomy, 72 Available mineral nutrients, 152- 4 Axon,32 Azinphosmethyl, 184 Azores, 51, 52

Bacteria, 27, 80, 147, 155, 156, 158, 160, 161, 166

Bacterial gums, 166, 167 Bacteriocide, 160 Bacteroidal cells, 82 Badgers, 123 Bahamas, 41, 45 Bardsey Island, 103 Barley, 169

Basal cells, 15 Basal membrane, 16 Basalt, 163 Base exchange capacity, 153 Basement membrane, 15, 16 Bassia tree, 100 Beans, 196 Beech, 140-2, 144, 198 Beet, 140 Behaviour, 71- 7 Behavioural experiments, 210, 211 Benefactors, earthworms as, 197 Benzopyrine, 197 Bermuda, 49 BHC, 182 Biomass, xii, 102 Birch, 140, 144, 150 Birds, 123, 193-6 Black head, 194 Bladex, 181 Blood, 90, 124 Blue-green algae, 157 Body wall, 13, 14, 15- 17, 33 Bolivia, 49, 50 Borneo, 41 Bowling green, 191 Brazil, 44-6, 49, 50 Breeding, 61 Breeding period, 62 Bristle worms, 37 Brown bodies, 19 Buccal cavity, 4, 19, 21

innervation of, 30, 32 Buccal chamber, 80, 89 Buccal epithelium, 89 Bulgaria, 190 Bullock droppings, 58, 198

as food, 138 Burma, 41, 50, 55, 115, 121 Burrowing, 92, 117-22 Burrowing experiments, 204- 6 Burrows, 73, 74, 81, 99, 100, 102,

108, 174, 185, 190,204


Burrows--{ con td.) depth of, 118 permanent, 118

Bursa propulsoria, 66 Butyric acid, 156

Caesium, 189 Cages, 142 Cage for studying earthworm acti-

vity, 205 Calcareous sand, population in, 137 Calciferous gland, 20, 42-4, 127, 166 Calcium, 153, 154, 166 Calcium carbonate, 80 Calcium humate, 166 California, 45, 46

Lower,44 Cameroons, 47, 48, 50, 122 Campbell Islands, 45 Canada, 54 Canary Islands, 51, 52 Cape of Good Hope, 54 Cape Verde Islands, 45 Capillaries, 85

intra-epidermal, 24 Capillary blood vessels, 13 Capillary water, 167 Capricorn, tropic of, 54 Carabid beetles, 123 Carbaryl, 184, 191 Carbofuran, 184 Carbohydrates, soluble, 140 Carbon nitrogen ratio, 150-2 Carbon dioxide, 78, 80, 133

tension, 151 Caroline Islands, 41 Carrots, 196 Casting, 117-22

time of day of, 121 Castor meal, effect of, 134 Casts, 81, 111, 120, 148, 152, 154,

156, 157, 159, 160, 163,165,167, 169,176,180,190,191,196, 201

microbes in, 164 pH of, 127, 172 seasonal production, 112, 119 stability of, 173

Cattle, 145 Caucasus, 52 Cells, albumen, 4, 14, 15, 36

central, 25, 26 chloragogen, 13, 18, 82, 83, 96 ciliated, 21 clitellar gland, 18, 65 cocoon-secreting, 36 coelomic epithelial, 17 drainpipe, 26 epidermal, 15, 36 epithelial, 15, 36, 81 epithelial brush border, 81 excretory, 82 giant, 32 glandular, 4, 15, 21, 24, 36 goblet, 14 light-sensitive, 88 marginal, 25, 26 mucous, 4, 15, 36 nerve, 87 peritoneal, 82 photoreceptor, 4, 15, 16, 32 proprioreceptor, 32 replacing, 15 reproductive, 62 secretory, 87 sensory, 14, 15, 87 supporting, 16 uric, 81, 82

Cellulase, 81 , 121, 155 Cellulose, 150, 156 Central cells, 25, 26 Cephalization, 3, 52 Cereals, 166, 178 Cereal stooks, 190 Cerebral ganglia, 28, 32, 75, 87, 88,

210 Ceylon, 41, 44- 6, 55, 123

GENERAL INDEX 271

Chaetopoda, xi Chaetotaxy, 4-7 Chemical sampling methods, 100,

110, 111 Chemoreception, 88, 89 Chemozem, 155 Chickens, 194, 195 Chile, 45, 51, 52 China, 41, 55, 190 Chitin, 155 Chitinase, 81, 121 Chloragogen cells, 13, 18, 82, 83, 96 Chloragosomes, 18, 82 Chlordane, 182, 187, 191 Chlorfenvinphos, 184 Chloropicrin, 181 Chlorpropham, 181 Choice experiments, 75 Ciliated cells, 21 Circular muscle, 4, 13, 14, 16, 17,

24,92 innervation of, 30

Circulation, 84, 85 Circulatory system, 20, 21 Circumoesophageal nerve ring, 94 Circumpharyngeal connectives, 28,

29, 30, 88 Citric acid, 89 Clasping, 65 Clay, 108, 118, 163

population in, 136 Clay chimneys, 121 Climate, 62 Cli tellar gland cells, 18, 65 Clitellum, 9, 10, 11, 12, 32, 35, 36,

41, 49, 51, 52, 61, 63, 64, 69 Clitellum position, 9, 12 Clover, 139, 162, 169, 178 Cluster flies, 124 Cobalt, 188 Cocoon, development time of, 59, 60

formation, 10, 65 incubation time, 60

Cocoon production, 61, 64, 66, 87, 106

effects of food on, 138 effects of soil moisture on, 128 effects of temperature on, 130

Cocoon secreting cells, 36 Cocoon shape, 66 Cocoon size, 66 Cocoon wall, 66 Cocoons, 37, 56, 57, 67, 92, 107,

110, 115, 198, 199, 206, 207 seasonalproductionof,57,58, 114

Coelom, 13, 16, 17-19, 24, 28, 84, 124

Coelomic epithelial cells, 17 Coelomic fluid, 9, 13, 18, 19, 23, 72,

78, 82,83, 90-2,127 Coelomocytes, 77 Collagenous fibres, 4, 13 Colombia, 45, 49, SO Colour of earthworms, 13 Commensalism, 123 Commissural vessel, 22 Comparisons of sampling methods,

101, 102 Compost, 172 Concentration factor, 186 Condensed tannin, 140 Conductivity, 99 Congo, 47,50 Connectives, circumpharyngeal, 28,

29, 30, 88 subpharyngeal, 30

Control of earthworms, 191 Copper sulphate, 181 Copulation, 11, 39, 62, 63- 5 Copulatory pouches, 65 Corsica, 51 Costa Rica, 45, 4 7 Cotton, 187 Creatinine, 83 Cretaceous era, 38 Crop, 19, 20, 80, 157, 185


Cropping, effects of, 176-8 Crop yields, effect of earthworms

on, 168-71, 177 Cuba, 45 Cultivation, effects on earthworms,

174-6, 175 Cultivations, 182

effects of, 174-6 Culture, growth in, 60 Culture methods, 198-200 Cultures, 66, 69, 206, 207 Cuticle, 4, 13-16 Cysticercoid stage, 124 Cysticercus, 193

D- D, 181 DDE, 185, 187 DDT, 183, 185, 187 Dead roots, 112 Death, 61, 69, 70 Deciduous woodland, 146 Dehydration tropism, 90 Dendrogram, 42 Denmark, 125 Deserts, 53, 137 Desmids, 157 Desiccation, 67, 109 Development, 61, 69 Diapause, 63, 67- 9, 128, 201

facultative, 57, 68, 111 obligatory, 57, 68, 111, 207

Diatoms, 157 Diazinon, 184 Dicotyledenous plants, 38 Dieldrin, 183, 187 Diets, 206 Digestion, 80, 81 Digestive system, 20 Dimensions of worms, 208 Dimethylbenzanthrene, 197 Dipterous larvae, 124 Disking, effects of, 17 4, 17 5 Disease transmission, 191-6

Dispersive powder, 106 Dissection, 200 Distributions, 200 Disulfoton, 96 Diurnal activity, 151, 152 Diurnal cycles, 76, 77, 79 Diurnal respiratory cycles, 87 Diurnal weight changes, 90 DNOC, 181 Dog's mercury, 140 Dorsal pores, 2, 9, 19, 23, 52, 72, 90,

127 Dorsal vessel, 17, 20, 21, 22, 23, 30,

84, 85 Dorso-intestinal vessel, 17, 23 Dorsa-subneural vessel, 17, 21- 3,

85 Drainage, 167, 168 Drainpipe cells, 26 Droughts, 127 Ducks, 194 Dung, 106, 138, 139, 145, 146, 162,

169, 170, 176, 180 effects of, 134, 135

Dung-pats, 107 Dursban, 184 Dutch elm disease, 185 Dwarf bunt, 161, 192 Dyfonate, 184

Earthworm farms, 171, 197 Earthworm pies, 197 Earthworms, as benefactors, 197

as pests, 190, 191 Ecology, 98-140 Ecuador, 46, 49, 50 Eelworm cysts, 192 Eggs, 207 Egypt, 50, 52, 62, 104, 126, 138 Elderberry, 140 Electrical inhibition of growth, 95 Electrical potential, 95 Electrical sampling methods, 99

GENERAL INDEX 273

Electrical stimuli, response to, 90 Electric shock, 210 Electrode, 100 Eleocytes, 18, 19 Elm, 140, 144, 150 Endemic species, 53 Endrin, 183, 187, 191 England, 54 Environmental factors, influence of,

125- 40 Enzymes, 80, 81, 141, 155 Epidermal cells, 15, 36 Epidermal sense organ, 15 Epidermis, 4, 9, 13, 14, 16, 23, 24 Epilobous prostomium, 3 Epineurium, 31 Epithelial brush-border cells, 81 Epithelial cells, 15, 81 Epithelial sense organ, 4, 14, 32 Epithelium, 4, 19, 23, 30 Essential intermediate hosts, 193 Ethiopia, 48 Eudrilid, 63, 65 Euprostates, 47 Europe, 40, 49, 51-3, 171 Excretion, 81-5 Excretory cells, 82 Excretory organs, 9 Excretory system, 24, 25- 8, 42, 43 Experimental cages, 202 Experiments with earthworms,

198- 211 External segmental grooves, 17, 18

Facultative diapause, 57, 68, 111 Faeces, 148, 149, 155, 171

microbial content of, 159 Fallow, 103, 121, 176 Families, 39, 55 Farmyard manure, 58 Fats, 82 Fecundity, effects of food on, 138

effects of temperature on, 129

Feeding, 72, 74 Female pores, 8, 9, 10, 12, 63, 40,

41, 49, 51 Female vagina, 196 Fenestrae, 31 Fertilization, 63, 65, 66 Fertilizer, 171 Fertilizers, effects of, 178-81 Fever, 197 Flatworm, 124 Flooded soil, 92, 129 Fibre, collagenous, 4, 13

intra-epidermal nerve, 86 lateral giant, 31, 32, 86 median dorsal longitudinal giant,

85,86 motor, 86

Fibres, nerve, 15 Fibrillae, 31 Field capacity, 167 Field studies, 198-211 Filtration, 84 Fish bait, 99, 197 Fistula, 196 5 HT, 87 Fodder, as food, 138 Follicles, 4 Food, 88, 137, 174, 176 Food supply, 138-40

effect of, 138-40 Food type, effect of, 60 Foot and mouth disease, 193 Forest soil, 166 Formalin, 13, 100, 102, 103, 199-

201,209 Formalin sampling, 72, 101, 171 Formaldehyde, 89 Fossil records, 37 Fossil worms, 38 Fragmentation, 141-5 France, 46, 54 Fried earthworms, 197 Frost, effect of, 131


Fungal mycelia, 139 Fungal spores, 158, 159, 161, 191 Fungi,80, 118,156-61,166,191,192 Fungicide, 141 Fungicides, effects on worms, 181

Gambia, SO Ganglia, cerebral, 28, 32, 75, 87, 88,

210 sub-oesophageal, 28 pharyngeal, 30

Gate worms, 193 Genera, 39-55 Generator, 100 Genital openings, 7-9, 39 Genital organs, 13, 37 Genital pores, 1, 3, 5, 7, 10 Genital seta, 5 Geographical distribution, 53-5 Germany, xiii, 51, 103, 122 Ghana, 48 Giant cells, 32 Giant fibres, 31, 32, 86 Gizzard, 19, 20, 41, 49, 51, 52, 80,

81, 157, 185 Glaciation, 54 Gland, calciferous, 20

mucous, 23, 78 pharyngeal, 80 salivary, 80

Glandular cells, 4, 15, 21, 24, 36 Glandular secretions, 63 Glucose, 16, 89, 160 Glycogen, 79, 82 Goblet cells, 14 Golf courses, 181, 191 Granite particles, 163 Grape husks, 162 Grass, 162, 169, 174, 176, 180, 191 Grassland, 135, 176

Greenland, 52 Gregarines, 124 Growing zone, 69 Growth, 69, 10, 71, 129

effect on food on, 138 Growth periods, 59, 60 Guatemala, 45, 50 Guinea, 47 Gut, 94 Gut contents, 102 Guthion, 184 Guyana, 45, 49, 50

Habitat, 103 Haemochromagen, 84 Haemoglobin, 13, 78 Haiti, 46 Handsorting, 98, 99, 102, 103 Rants, 103 Harrowing, 174 Hawaii, 52 Hay, 169, 178 Hearts, 20, 21, 22, 23, 44, 45, 85 Heat extraction, 101 Health, 122 Heptachlor, 183, 187, 191 Heptachlor epoxide, 187 Herbage, 106, 112 Herbicides, effects on worms, 181,

182 Hermaphrodite, 7, 37, 63 Herts, 14, 103 Heteroxanthine, 84 Himalayas, East, 41 Holland, 77, 103 Holonephridia, 28, 44, 45 Honey jars, 200 Horizontal distribution, 209 Horizontal migration, 107 Horizontal movement, 170, 173

Gravelly sand, population in, 136, Hormones, 87 137 Human excreta, 158

Greenhouses, 77 Human faeces, 196

GENERAL INDEX 275

Human urine, 196 Human welfare, 40 Humidity, 18 Humification, 145-7 Humus, 147 Hungary, 51 Hydrogen sulphide, 77 Hydrostatic pressure, 90 Hypotonic urine, 91

Iceland, 52 Immature worms, proportions of,

104, 105 Impotency, 197 Incubation period, 62 India, 41, 44-7, 50-3, 55, 62, 115,

190 Indonesia, 55 Infectivity of cysts, effects of

worms on, 192 Inorganic minerals, 121 Inorganic salts, 106 Insecticide residues, 185 Insecticides, effects on worms,

181-8, 209 uptake into worms, 186

Internal salt concentration, 91 Intersegmental grooves, 1, 9, 11 Intestinal microflora, 147 Intestinal wall, innervation, 30 Intestine, 19, 20, 21, 80, 81, 85, 96,

157 Ireland, 54 Iron, 189 Israel, 51, 52 Italy, 51

Japan, 41, 51, 52, 55, 115, 190, 197

Jaundice, 197 Java, 41, 44 Jordan, 51 Jurassic era, 38

Kale, 140 Kentucky, 14 Kenya, 47 'Kilner' jars, 198 Kommetjies, 119

Lactic acid, 79 Lamellae, 31 Larch, 140 Lashing movements, 72 Lateral giant fibres, 31, 32, 86 Lateral-neural vessel, 17, 22, 85 Lateral-oesophageal subneural

vessel, 22, 23 subneural vessel, 22, 23

Latex, 206 Latex casts, 118, 206 Lawns, 191 Lead arsenate, 181 Leaf burial, 202, 203

effects of temperature, 130, 131 Leaf disks, 142, 203, 204

rate of breakdown of, 144 Leaf litter, 106 Learning, 74, 75-7, 211 Leaves, 74 Leeches, 37, 124 Legume, 178 Lespedeza, 1 7 6 Lettuce, 140, 190 Leys, 176, 177 Liberia, 45, 48 Life cycles, 56-62 Life history studies, 206-8 Life span, 60, 61 Light, 63

reaction to wavelength of, 88 Light loam, 118

populations in, 136, 137 Light reactions, 87, 88 Light receptors, 88 Light-sensitive cells, 88 Lignin, 150


Lime, 140, 144, 1 SO, 169, 178-80 Lima beans, 169 Lipase, 81, 121 Lithium, 96 Litter, breakdown of, 141-5

fragmentation of, 141-5 humification, 145-7 incorporation of, 141 rate of burial, 142 see Leaf litter

Litter turnover, 145-7 Loam, 198 Locomotion, 92-4 Longitudinal muscle, 4, 13, 14, 16,

17,24,92 Lucerne, 166 Lumbricine setae, 6, 7 Luminescence, 77 Lung worms, 193, 194 Lymph glands, 82 Lymphocytes, 18

Madagascar, 45, 46, SO Magnesium, 153, 173 Magnesium sulphate, 99, 199 Malagasy, 49, 54 Malathion, 184 Malay Archipelago, 41, 46, 49, 50 Male pores, 8, 12, 39- 43, 47, 49, 51,

52, 62- 5 Malic acid, 89 Mammals, 19 5 Mangolds, 135 Manure, 172 Maoris, 197 Marginal cells, 25, 26 Mass migration, 77 Mating, 62, 63 Maze, 76, 96, 140 Mechanized soil washing, 99 Medium loam, population in, 136,

137 Meganephridia, 28

Membrane, basal, 16 basement, 15, 16 mesenteric, 18

Menazon, 184 Mercuric chloride, 100 Meronephridia, 46 Mesenteric membrane, 18 Mesh bag, 142, 143, 203 Metabolism, effects of temperature

on, 130 Metal, 140 Metham sodium, 181 Methyl bromide, 181 Mexico, 45, 46, 49, SO, 54 Microbial cell, 159 Microbial protein, 148 Microftora, 141, 160 Micronephridia, 28 Micro-organisms, 80, 121, 147,

155-62, 166 and earthworms, 155-62 dispersal' of, 161 numbers in casts, 157 numbers in gut, 156

Middle tube, 24, 26, 83, 84 Migrations, 77 Millet, 169 Mineral elements, 154 Mineral nutrients, 152 Minerals, effect of, 134 Mineral soil, 146, 147, 164 Mineralization of nitrogen, 147- 50 Mites, 147 Mixed woodland, 103 Moisture, 127-9, 137

effects of, 127- 9 reaction to, 209

Moisture content, 99, 106, 111, 112, 134

Moisture equivalent, 173 Moisture gradient, 128 Moles, 123, 190 Molybdenum, 153

GENERAL INDEX 277

Moorland soil, 171 Mor soil, 103, 104, 122, 164

species in, 13 7 Mortality, 70 Motor fibres, 86 Mountain ranges, 53 Mouth, 80, 90 Mowrah meal, 100 Mozambique, 48, 50 Mucocytes, 19 Mucoproteins, 148 Mucous cells, 4, 15, 36 Mucous tubes, 65 Mucus, 63, 80, 81 Mud, 62 Mull soil, 104, 122, 165

species in, 137 Muscle, arciform, 63 Muscle, circular, innervation of, 30

dorsal, 16 fibres, 16, 17 innervation, 86

Muscle, longitudinal, 4, 13, 14, 16, 17,24,92

longitudinal, innervation of, 30 setal, 4 sphincter, 9, 18 subcuticular, 13 ventral, 16

Muscoid flies, 124 Muscular tube, 24, 26 Mycobacteria, 160

Napthalene, 142 Narrow tube, 24, 25, 26, 83, 84 Nematodes, 19, 80, 124, 181, 193 Nephridia, 2, 20, 81, 82, 41, 47, 49

enteronephric, 27 exonephric, 27, 28 holonephridia, 28 meganephridia, 28 meronephridia, 28 micronephridia, 27, 28

Nephridiopore, 9, 23, 24, 42, 43, 81, 91, 148

Nephridium, 9, 24, 26, 27, 83 Nephrostome, 24, 25, 26, 27, 82-4

lower lip, 25 peritoneum, 25 upper lip, 25 prostomial, 28, 29

Nerve depressant, 96 Nerve, segmental, 28, 29, 30, 86

septal, 29 Nerve cells, 87 Nerve cord, 17, 85, 96, 210 Nerve fibres; 15 Nerve plexus, 14 Nerves, electric potentials in, 88 Nervous System, 28, 29, 30-2, 85-

90 fine structure, 31

Neural lamella, 31 Neurofibrils, 16 Neuroglia, 31 Neuropile, 31, 32 Neurosecretions, 87 New Caledonia, 44-6, 55 New South Wales, 44, 51, 52, 103,

145 New Zealand, 44-7, 51, 53, 55, 68,

167, 169, 170, 173, 197 Newly-emerged worms, 69 Nigeria, 45, 48, 50, 104, 11 3, 146 Nile Valley, 122 Nitrate, 147, 149, 162, 173 Nitrate of ammonia, effects of, 134 Nitrate of soda, 148, 178,180 Nitro-chalk, 179 Nitrogen, 168, 178

effect on growth, 138 effect on worms, 179

Nitrogen excretion, 151 Nitrogen-fixing bacteria, 14 7 Nitrogen humification, 147- 50 Nitrogen mineralization, 147- 50


Noradrenalin, 87 Novocaine, 87 Number of samples, 102 Nutrient transport, 82 Nutrition, 58, 80 Nylon net, 143

Oak, 140-2, 144, 149, 150, 157, 170 woodland, 103

Oat straw, as food, 138 Oats, 169 Obligatory diapause, 57, 68, 111,

207 Oesophageal cavity, 20 Oesophageal pouches, 20 Oesophagus, 19, 20, 80, 85 Oocytes, 63 Oogenesis, 63 Oogonia, 63 Oothecae, 56 Optic organelle, 32 Orchards, 101, 103- 5, 133, 141, 146,

187 Ordovician period, 38 Organ, excretory, 9

genital, 13, 37 respiratory, 23, 78 sense, 89

Organic matter, 80, 81, 106, 121, 165, 168, 171, 173, 174, 176, 180, 201, 204

cycles, 141- 54 effect of, 133, 134, 135, 136

Organelle, optic, 16 Organo-chlorines, 183 Organo-phosphates, 183 Osmotic pressure, 91 Osmotic pressure of urine, 83 Ova, 56, 63, 65, 66 Ovarian funnels, 33 Ovaries, 32, 40, 41, 51, 63, 96 Oviduct, 63 Ovisacs, 33, 63

Oxalic acid, 89 Oxidation-reduction potential, 133 Oxygen, 23,78- 80,96

consumption of, 159, 162 debt, 79, 151 tension, 133, 151

Pacific Ocean Islands, 55 Pakistan, 41, 51, 52, 55 Palaearctic zone, 40, 54 Palatability, 89, 203

experiments, 203, 204 of leaf litter, 140

Panama, 45, 49, 50 Papillae, 5, 7, 10-12 Paraguay, 44- 6, 49, 50 Paraquat, 182

effect on worms, 182 Parasites, 123-5 Parathion, 96, 184 Parthenogenesis, 63, 66 Passive agents, 196 Pasture, 68, 99, 103, 104, 112, 113,

121-3, 127, 131, 139, 161, 163, 166, 169, 173, 176, 177

Pasture productivity, effect of worms on, 170

Peas, 169 Peat, 58, 122 Peaty soil, population in, 137 Penes, 65 Peregrine species, 53 Perichaetine setae, 6, 7 Peri-enteric plexus, 17, 23, 85 Peristalsis, 92, 93, 94 Pacific Islands, 44 Peristomium, 1, 3, 52 Peritoneal cells, 82 Peritoneum, 4, 13, 14, 17, 23, 63

innervation of, 31 Peritrophic membrane, 81 Persia, 52 Peru, 44, 49

GENERAL INDEX 279

Pesticides, effects of, 181-4 uptake in earthworms, 184-8

Pests, earthworms as, 190, 191 Petroleum ether, 160 pH, 89,106,125-7,168,172,201

effects of, 125-7 reaction to, 209

Pharyngeal ganglion, 30 Pharyngeal gland, 19 Pharyngeal intestinal plexus, 32 Pharyngeal nerve plexus, 32 Pharynx, 19,20, 30,80 Phillipines, 41, 190 Phorate, 184, 191 Phosphorescence, 77 Phosphoric acid, 89 Phosphorus, 153, 173 Photoreceptor cells, 4, 15, 16, 32 Physiology, 78-97 Pig litter, 103 Pigmentation, 13 Pigs, 193, 195 Piles, 197 Pine, 144

woodland, 103 needles, 138

Plant cover, 134 Plant growth substances, 171 Plant pathogens, 161 Ploughing, 17 4, 17 5 Plug, 74 Polders, 171 Polynesia, 46 Polyphenols, 140, 142

reaction to, 89 Polysaccharide gums, 160, 166 Population, biomass, 102

horizontal distribution, 106, 107 seasonal activity, 11~17 seasonal changes in, 71, 11~17 size of, 102- 4 vertical distribution, 107- 10 distribution of, 106- 10

Population, estimation of, 98-102 numbers, 102 structure, 104- 6

Populations, 70, 104 Pores, cuticular, 13

dorsal, 2, 9, 19, 23, 52, 72, 90, 127 female, 8, 9, 10, 12, 40, 41, 49, 51,

63 genital, 1, 3, 5, 7, 10 prostatic, 2, 7 sperm, 52 spermathecal, 2, 8, 12, 40, 41, 47

Porphyrins, 13 Porosity, 167, 168 Post-clitellar region, 7 Post-septal canal, 24 Potassium, 153 Potassium cyanide, 96 Potassium permanganate, 89, 100-3 Potato root eelworm, 192 Predation, 58 Predators, 121, 123- 5 Pregnancy, 197 Preseptal canal, 24 Preservation, 200 Proboscis, 1 Prolobous prostomium, 3 Propham, 181 Proprioreceptor cell, 32 Prostates, 7, 36, 41- 3 Prostatic fluid, 65 Prostomial nerve, 28, 29 Prostomium, 1, 2, 3, 4, 12, 28, 29,

52, 87, 89, 91, 92 innervation, 28, 32

Protease, 81 Protozoa, 19, 80, 160, 161 Protozoan parasites, 124

Quadrat, 101, 102, 203 Queensland, 44- 6 Quiescence, 67-9, 109-11, 128 Quinine, 16, 89


Radiation, effect on A. chlorotica, 188

Radiocaesium, 146 Radioisotopes, effect on worms,

188, 189 uptake into worms, 188, 189

Radishes, 196 Ragi, 196 Rain, reactions, to, 77 Raw humus, 134 Reception zone, 80 Reduction division, 63 Regeneration, 94-6, 209 Regenerative phenomena, 87 Replacing cells, 1 5 Reproduction, 62-7, 87, 105 Reproductive cells, 62 Reproductive cycle, 87 Reproductive potential, 106 Reproductive system, 32, 33, 34, 35,

36 Resorption, 84 Respiration, 78-80, 150, 151

effects of temperature on, 130 Respiratory system, 23, 24

Resting phase, 68 Retinella, 32 Rheumatism, 197 Rhizopods, 157 Rhodesia, 50 Rice, 190 Ridges, 10, 11 Rigor, 91 Rio de Janeiro, 50 Rodents, 194 Rolling, 174 Roots, 139, 141 Rotary cultivation, 174 Rotations, 176 Rotifers, 80 Rowan, 150 Russia, 51, 52 Rye, 157, 169, 176

Saccharose, 89 Sahara, 54 St Thomas Island, 44, 46 Salivary gland, 80 Sampling, 98 Sampling tool, 98 Sand, 163 Sardinia, 46, 50, 54 Scavenging, 71 Scotland, 122 Scots pine, 150 Seasonal abundance, 106,107 Seasonal activity, 110-17 Secondary sex characters, 87 Secretory cells, 87 Seedlings, 190 Segment numbering, 3 Segmental contractions, 93 Segmental ganglia, 28 Segmental nerves, 28, 29, 30, 86 Segmentation, 1-4 Segments, 3, 69 Seminal fluid, 64, 65 Seminal funnel, 34 Seminal grooves, 2, 8, 12, 63 Seminal vesicles, 32, 34, 35, 52, 62,

124, 197 Senescence, 69 Sense organs, 89 Sensory cells, 14, 15, 87 Septa, 1, 17, 18, 26, 29, 31 Septa innervation of, 30 Septal nerves, 29 Seta of L. terrestris, 4 Setae, 4, 5, 24, 40, 41, 47, 49,

51 closely-paired, 6 distant-paired, 6 genital, 5 lumbricine, 6 normal, 4, 12 perichaetine, 6 sexual, 49, 51, 53, 65

GENERAL INDEX 281

ventral, 12 widely-paired, 6

Setal muscles, 4 Setal arrangement, 6, 7 Setal form, 4, 5 Sewage sludge, 58 Sexual activity, 62 Sexual development, 63 Sexual maturity, 10, 67, 71, 161 Sexual organs, 37, 96 Seychelles, 44 Sheep droppings, as food, 138 Shell 19805, 181 Shrews, 123 Siberia, 51, 52 Sicily, 51, ~4 Sieve, 99, 101 Silt, 163 Silurian period, 38 Simazine, 181 Simple key, 213 Size of populations, 102-4 Size of samples, 102 Slime, production of, 77 Slime tube, 63 Slugs, 123 Sodium chloride, 89 Soil, forest, 166

garden, 167, 169 heavy, 118

Soil acidity, 88, 125, 127, 169, 178-81,201

Soil aeration, 165, 167, 168

Soil amelioration, 171-3 Soil consumption, 146 Soil drainage, 167, 168 Soil erosion, 197 Soil fertility, 154, 163- 73 Soil formation, 172 Soil indicators, 168

Soil microflora, 134 Soil moisture, 56, 100, 168 Soil porosity, 167, 168

Soil profile, 141 with earthworms, 143 without earthworms, 143, 164

Soil structure, 196, 163-8 Soil texture, 106 Soil turnover, 164, 165 Soil type, 136, 137 Soil washing, 99

mechanized, 99 South Shetland Islands, 55 Soybeans, 169, 177, 178 Species associations, 122, 123 Specific gravity, 99 Sperm funnels, 33, 35, 62 Sperm pores, 52 Sperm transference, 39, 65 Spermathecae, 8, 11, 32, 34, 35,

36, 39, 41, 47, 51, 52, 64, 65, 124

Spermathecal openings, 63 Spermathecal pores, 2, 8, 12, 40, 41,

47 Spermatogenesis, 62, 197 Spermatogonia, 62 Spermatozoa, 62, 65 Sphincter muscles, 9, 18 Sporangia, 38 Springtails, 14 7 Spruce, 140, 170 Squirter earthworm, 19 Staphylinid beetles, 123 Stimuli, chemical, 72, 88, 89

electrical, 72, 75, 76, 171 electrical response to, 90, 91 light, 76, 87 photic, 88 sensory, 86 touch, 72, 76 vibrations, 77

Stones, in the bladder, 197 Straw, 58, 198 Stubble-mulch farming, 176 Stubble striping, 175


Subcuticular muscle, 13 Sub-epidermal nerve plexus, 30, 31 Sub-epithelial nerve plexus, 31 Submergence, 129 Subneural vessel, 17, 21, 22, 85 Suboesophageal ganglion, 28 Subpharyngeal connectives, 30 Subpharyngeal ganglion, 29 Subtillers, 176 Sucrose, 16, 89 Sudan, SO Sulphate of ammonia, 180

efftl4:ts of, 134 Sumatra, 41 Sumithion, 184 Summer, 62, 68 Sunda Islands, 41 Superphosphate, 178, 180 Supra-intestinal vessel, 22 Surface-dwelling species, 199 Survey, 186 Survival in water, 79, 209 Swine, 194 Symbiotic bacteria, 81 Sympathetic system, 30 Synapses, 86

Tactile receptors, 89 Tanylobous prostomium, 3 Tanzania, 47, 48, 50 Tapeworms, 124, 193, 194 Tartaric acid, 89 Tasmania, 46 Taxonomy, 37-55 TCA, 181 Teleospores, 192 Temperate forest, 146 Temperate zones, 68 T emperature, 56, 57, 60, 67, 79, 96,

100, 106, 108-12, 115, 129-33, 151, 198, 211

conditioning, 133 effect on respiration, 79

lethal, 131-3 seasonal changes in, 114

Temperature preferenda, 130, 131, 132

Temperatute tolerance, 132, 133 Temperatures, optimum for

growth,206 Termites, 123 Testa! sacs, 34, 62 Testes, 32, 34, 40, 41, 51, 124

holoandric, 33 meroandric, 3 3

Thigmotactic reactions, 89-90 Timothy grass, 176 Tissue connective, 16, 17 'T'-maze, 75, 76, 210 Tobacco, 190 Tobago, 54 Togoland, 48, 50 Tooth powder, 197 Touch, 63, 71, 72 Transplantation, 96, 97 Transverse commissure, 88 Transverse septa, 17 Triassic era, 38 Triazine herbicides, 181 Trinidad, 54 Tropical areas, 79 Tropical forest, 146 Tropics, 122 Tube, middle, 24, 26, 83, 84

mucous, 65 muscular, 24, 26

narrow, 24, 25, 26, 83, 84

slime, 63 wide, 24, 26, 83, 84

Tubercula pubertatis, 3, 10, 11, 12, 52

Tubercules, 10 Tumours, 197 Tunis, 51 Turnover of soil and litter, 145-7 T yphlosole, 21, 85

GENERAL INDEX 283

Uganda, 50, 104 Ubiquitous species, 126 Ultra-violet light, 77, 80, 88 Urea, 81-4, 148 Uric acid, 84, 148 Uric cells, 81, 82 Urine, 81, 91, 148, 170

osmotic pressure of, 83 U.S.A., 38, 46, 50, 53, 54, 103, 105,

115, 125, 128, 175 U.S.S.R., 103, 172, 176

Valerionic acid, 79 Vas deferens, 33, 34, 35, 40, 62 Vas efferens, 34, 35, 62 Vascular system, 21- 3 Vegetable crops, 190 Venezuela, 49 Ventral muscle, 16 Ventral nerve cord, 28, 29, 30, 31,

34, 88, 93, 96 structure of, 3 2

Ventral vessel, 17, 85 Ventral view, 12 Vertical distribution, 108, 109,

209 Vertical migration, 58, 67 Vessel, afferent nephridial, 17

afferent typhlosolar, 17 blood, 17, 21, 96 capillary, 7 8 commissural, 22 dorsal, 17, 20, 21, 22, 23, 30, 84,

85 dorsa-intestinal, 17, 23 dorsa-subneural, 17, 21- 3, 85 efferent nephridial, 17 extra-oesophageal, 22 lateral-neural, 17, 22, 85 lateral-oesophageal, 21, 22

lateral-oesophageal subneural, 22, 23

nephridial, 23 subneural, 17, 21, 22, 85 supra-intestinal, 22 supra-intestino ventral, 23 typhlosolar, 23 ventral, 17, 21, 22, 23, 30 ventra-intestinal, 23, 85 ventra-oesophageal, 85 ventro-parietal, 17, 23, 85

Vibrations, 72 Victoria, 46 Viruses, 124 Voltage, 99

Wales, 103 Water conservation, 90 Water loss, 68 Water relationships, 90-2 Water-stable aggregates, 163, 165,

166-8 Weathering, 44-6, 49, 53, 54, 142 Westmorland, 103 Wheat, 169, 176

effects of, 135 Wide tube, 24, 26, 83, 84 Winter, 68 Withdrawal reactions, 71 Woodland, 135, 151 Worm cairns, 7 4 Wound tissue, 95 Wych Elm, 140

Yeasts, 157, 159 Yield increases, 170

Zanzibar Island, 48 Zinophos, 184 Zurich, 122 Zygolobous prostomium, 3

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A simplified key to common genera of terrestrial earthworms

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