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 3 (LUMBRI- C IDAE) 2. Setal arrangement perichaetine (Fig. 2) Pheretima (MEGASCO• LECIDAE) - Setal arrangement lumbricine (Fig. 2) Diplocardia (ACANTHO• DRILIDA E) 3. Prostomium tanylobous (fig. 1), setae closely paired (Fig. 2) at least over part of the body 4 - Prostomium epilobous (fig. 1) or if tanylobous, setae widely paired or distant (Fig. 2) over the whole body 5 4. Clitellum begins on segment 24 Bimastos eiseni (Levin• sen)•• - Clitellum begins on or after segment 26 Lumbricus 5. Clitellum ends after segment 28 6 - Clitellum ends before segment 28 Eiseniella 6. Tubercula pubertatis absent, or exceptionally, present as simple thickenings of the edges of the clitellum Bimastos - Tubercula pubertatis present as ridges or isolated papillae (plate Sb) 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 (excep• tionally as separate tubercles) extending over only part of the length of the clitellum Dendrobaena •• Only one common sp ecies. 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 Octolasium 9. Spermathecal pores (Fig. 3) in line with seta 'd' or more often near the mid-dorsal line, body trapezoidal in crossection Eisenia - Spermathecal pores situated laterally between setae 'c' and 'd' or 'a' and 'b' and 'c' and 'd'. Body not trape• zoidal in crossection 10 10. Prostomium with longitudinal ridges Eophila - Prostomium without longitudinal ridges 11 11. Calciferous glands with two lateral pouches in segment Terrestrial Allolobo• phrna - Calciferous glands without lateral pouches. Amphibious 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 charac• ter 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.

MEGASCOLECIDAE Genus Pheretima Two pairs of spermathecal pores in segmental grooves 7/8 and 8/9. Pheretima 1st dorsal pore 11/12. 70- 170 mm. 10- 150 segs. Reddish-brown, californica clitellum creamy to dark grey. (Kinberg) Three pairs of spermathecal pores on the anterior edges of seg• P. hupiensis ments 7, 8 and 9. 150-220 mm. Light green/greenish buff with (Michaelsen) 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 P.posthuma and 8/9. 1st dorsal pore 12/ 13. 100- 160 mm. 120- 150 segs. Rich (Vaillant) brown. ACANTHODRILIDAE Genus Diplocardia 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 Red/brown or red/violet, Lumbricus irridescent dorsally, pale rubellus• yellow ventrally. 25-105 Hoffmeister mm. 95-120 segs. 28-33 29-33 6/7 Chestnut to violet L. castaneus • brown; brown/yellow (Savigny) ventrally, irridescen t, clitellum orange. 30-70 mm. 82-100 segs. 31, 32-37 33-36 7/8 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. 33, 34-39 34, 35, 5/6 Red-brown, lighter ven- L.festivus• 36-38 trally, irridescent dor- (Savigny) sally, prominent clitel- lum. Not found in large numbers. 48-108 mm. 100-143 segs. Genus Eiseniella 22,23-26, 23-25, 26 4/5 Male pores in 13. Dark Eiseniella 27 brown, greenish, golden tetraeda f . yellow, red. Body typica • quadrangular behind (Savigny) the clitellum. 30-60 mm. 60-90 segs. Male pore in 15. Other• E. tetraeda wise as f. typica (Savigny) fhercynia• (Michaelsen)

• British species. t for numbering see p. 12. KEY TO GENERA OF TERRESTRIAL EARTHWORMS 215

Genus Bimastos Tubercula 1st dorsal C/ite/lum pubertatis pore 20, 21, Absent Setae ab > cd. Red- Bimastos 22-29,30 brown. giese/eri 105-115 segs. 50-80 (Smith) mm.

22-29 27,28 Reddish-brown. B. tumidis Eise segs 30-6. 20-50mm 23-28 Absent 20-50 mm. 40-60 segs. B. palustris up to 75 mm. (Moore)

23, 24, 24, 25, 5/6 Reddish dorsally, yel- B. parvus 25-31, 32 26-30 lowish ventrally. 25--40 (Eisen) or absent mm. 90-110 segs. 23-32 Absent Rose-red. B. or 24-33 98-122 segs. 60-90 mm. longicinctus (Smith& Gittins)

24,25-32, Absent 5/6 Prostomium tanylobous, B. eiseni• 33 body cylindrical, reddish (Levinsen) or violet dorsally, yel- lowish ventrally, clitel- lum red. 30- 64 mm. 75- 111 segs. 25, 26, 28, 29-30, 5/6 Setae widely paired, dor- B. tenuis•t 27-30, 31, 31 or sally red-brown with (Eisen) 32, 33 absent bluish tint. Intersegmen- tal grooves and ventral side light. 20-85 mm. 90-120 segs.

27-37 Absent Pale red to chestnut B. zeteki brown, often localized (Smith & whitish banding anter- Gittins) iorly. 100-140 mm. 110- 140 segs. Genus Dendrobaena 25,26-31, 28-30 5/6 Rosy to deep red, last Dendrobaena 32 posterior segments yel• subrubicundat low. 27-90 mm. 50-100 (Eisen) segs. • British species. t Now Dendrobaena rubida (Sav.) f. tenuis (Eisen). l Now Dendrobaena rubida (Sav.) f. subrubicunda (Eisen). 216 BIOLOGY OF EARTHWORMS

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. • British species. KEY TO GENERA OF TERRESTRIAL EARTHWORMS 217

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. • British species. 218 BIOLOGY OF EARTHWORMS

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. • British species. References

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

Acanthodrilacae, 43 61, 68, 91, 92, 98, 104, 106, Acanthodrilidae, 39, 42, 43, 45, 46, 107, 108, 109, 111, 112, 113, 54, 55, 62, 212, 213 114, 122, 125, 126, 127, 128, Acanthodrilus, 38, 45, 54 129, 135, 139, 144, 171, 185, Actinomyces, 159 188, 197, 198, 199, 204, 206, Aeogaster, 47 207,208,218 Aeolosomatidae, 13, 38 culpijera, 217 aerophilus, Thominx, 195 dubiosa, 194 africanus, Hyperiodrilus, 120, 121, japonica, 116 125, 129, 131 limicola, 208 Ocnerodrilus, 194 longa, 57, 58, 59, 61, 68, 89, agglutinatus, Bacillus, 158 91, 107, 109, 111, 118, 119, agrestis, Pheretima, 88 122, 123, 125, 126, 128, 129, Aikinetocystis, 124 131, 135, 136, 137, 138, 139, Allolobophora, 52, 57, 66, 107, 196, 145, 146, 165, 166, 185, 192, 213,217 195, 198, 199, 201, 207, 208, antipae, 27, 217 217 caliginosa f. trapezoides, 23, 217 miniscula, 217 caliginosa, 1, 3, 5, 8, 53, 56, 58, nocturna, 57, 58, 68, 70, 71, 104, 59, 60, 68, 70, 71, 97, 104, 105, 105, 108, 109, 110, 111, 113, 107, 108, 109, 110, 111, 112, 118, 122, 125, 126, 135, 199, 113, 114, 115, 118, 122, 125, 201, 207, 218 126, 128, 129, 131, 132, 135, ribaucourti, 217 136, 137, 138, 139, 140, 145, robusta, 123 146, 148, 149, 151, 162, 169, rosea, 9, 10, 58, 59, 60, 66, 68, 170, 171, 173, 194, 195, 197, 105, 107, 108, 109, 110, 111, 198, 199, 206, 207 112, 113, 115, 122, 125, 126, caliginosa f. typica, 217 128, 135, 139, 185, 198, 199, chlorotica, 3, 11, 56, 58, 59, 60, 204,206,207,208,217 254 BIOLOGY OF EARTHWORMS

Allolobophora--{ con td.) liinnbergi, 125, 215 terrestris, 123, 217 muldali, 208 terrestris, f. longa, 60 palustris, 215 tuberculata, 208, 218 parvus, 19,208,215 vogesiana, 217 tenuis, 8, 215 Alma, 50, 62, 132 tumidis, 215 Amoebotaenia cuneatus, 194 zeteki, 62, 70, 72, 122, 215 lumbrici, 194 Bipalium kewensis, 120, 124 Andiorrhinus, 49 Borgertia, 48 Andioscolex, 50 botulinus, Bacillus, 124 annulata, Capillaria, 194 bronchia/is, Cyathostoma, 193, 195 Anoplophyra, 124 Bu"ttneriodrilus, 48 Anteoides, 50 antipae, Allolobophora, 27, 217 cal ifornica, Pheretima, 132, 213 Aptodrilus, 49 cali"ginosa, Allolobophora, see Ascaridia galli, 196 Allolobophora Ascaris suum, 196 f. trapezoides, Allolobophora, 23, Aspidodrilus, 123 217 attesmi, Dendrobaena, 144, 206, f. typica, Allolobophora, 217 216 Callidrilus, 50 australis, Megascolides, xi, 19, 66 Capillaria annulata, 194 Azotobacter, 147, 156 causinjlata, 19 5 mucronata, 19 5 Bacillus agglutinatus, 158 plica, 195 cereus, 155, 158 putorii, 19 5 botulinus, 124 carnaria, Sarcophaga, 125 cereus var. mycoides, 157 castaneus, Lumbricus, see Lumbricus idosus, 158 causinjlata, Capillaria, 195 Barogaster, 47 Causus rhombeatus, 123 Bassia longifolia, 100 Celeriella, 47 beddardi, Bimastos, 125 cereus var. mycoides, Bacillus, 157 Beddardiella, 48 Cestoda, 193, 194 Benhamia, 47 Chaetopoda, 13, 37 Benhaminae, 43 Chilota, 45 Bimastos, 52, 212, 215 chlorotica, Allolobophora, see Bimastos beddardi, 125 Allolobophora Bettonia, 48 Chuniodrilus, 48 Bimastos eiseni, 8, 11, 107, 122, 125, Clostridium, 156 126, 133, 137, 139, 144, 207, coeli'color, Streptomyces, 159 208,212,215 Comarodrilus, 44 gieseleri, 215 communis, Diplocardia, 214 icenorum, 208 controversa, Tilletia, 161, 192 longicinctus, 215 corethrusus, Pontoscolex, 53, 196 SYSTEMATIC INDEX 255 crassum, Porrocaecum, 195 Dichogaster, 4{), 46, 53, 54, 55, 77 Craterocystis, 124 jaculatrix, 121 Criodrilidae, 39, 51 Didymogaster, 44 Criodrilinae, 51 sylvaticus, 19 Criodrilus, 51 Digaster, 44 Criodrilus lacuum, 62, 194 Dilepus undula, 194 culpifera, Allolobophora, 217 Dinodriloides, 45 cuneatus, Amoebotaenia, 194 Dioctophyma sp., 195 cuneata, Taenia, 124 Diplocardia, 46, 54, 110, 131, 212, Curgia, 44 213 Curgiona, 44 communis, 214 cyaneum, Octolasium, see egglestoni, 62, 68 Octolasium riparia, 214 Cyathostoma bronchialis, 193, 195 singularis, 213 Diplotrema, 38, 46 Deccania, 45 Diporochaeta, 46 Deinodrilus, 46, 55 Dtprochaeta, 124 Dendrobaena, 13, 52, 66, 68, 212, Distichopus, 124 215 Drawida, 41 attemsi, 144, 206, 216 grandis, 118, 123 mammalis, 57, 58, 123, 199, 208, Drilocrius, 50 216 dubiosa, Allolobophora, 194 octaedra, 106, 107, 122, 125, 126, dunguensis, Shubotziella, 65 133, 137, 139, 144, 207, 208, 216 Echinocystis, 124 platyura, 215 egglestoni, Dtplocardia, 62, 69 pygmaea, 208, 216 Eisenia, 52, 57, 213, 216 rubida, 67, 107, 122, 125, 126, foetida, 13, 16, 18, 60, 61 , 65, 139, 144, 199, 206, 207,215 66, 67, 69, 75, 76, 77, 80, rubida f. tenuis, 66, 208, 215 81, 88, 95, 96, 123, 124, 125, rubida f. subrubicunda, 208 128, 131, 132, 133, 145, 159, subrubicunda, 58, 59, 60, 61, 67, 160, 162, 172, 194, 195, 196, 69, 91, 92, 126, 129, 165, 199, 197, 199, 206, 207, 208, 201,215 216 veneta var. hibernica f. icterica, 208 dendroidea, 208 metallorum, 216 veneta var. hibernica f. typica, rosea, 60 208 veneta, 194 veneta var. hortensis, 208 eiseni, Bimastos, see Bimastos veneta f. typica, 208, 216 Eiseniella, 51, 66, 212, 214 veneta var. zebra, 208 tetraedra, 199, 207 Desmogaster, 41 tetraedra f. hercynia, 208, 214 Dicelis, 124 Eleusine coracana, 196 256 BIOLOGY OF EARTHWORMS elongata, Pheretima, 196 Glossoscolecinae, 4, 49, 54 elongatus, Metastrongylus, 194 Glossoscolex, 50 Eminoscolex, 48 Glyphidrilus, 50 Enantiodrilus, 50 Gordiodrilus, 45 Enchytraeidae, 9, 10, 39 grandis, Drawida, 118, 123 ensicaudatum, Porrocaecum, 19 5 Grayallia, 124 Entyphoeus waltoni, 65 Gregarina, 124 Eodrilus, 46 Eophila, 52, 213, 217 haemorrhoidalis, Sarcophaga, 125 (Allolobophora) icterica, 126, 217 hali'otidea, Testacella, 123 obscuricola, 217 Haplodrilus, 45 Escherichia coli, 157 Helodrilus, 52, 212, 218 Eudi'chogaster, 46 oculatus, 208, 218 Eudrilidae, 39, 47, 53, 54 herculeus, Lumbricus, 195 Eudrilinae, 48 Heterakis, 124 Eudriloides, 48, 65 Heterodera rostochiensi's, 192 Eudrilus, 48, 72 hilgendorj, Pheretima, 62 eugeniae, 53 Hippopera nigeriae, 154 eugeniae, Eudrilus, 53 Hirudinea, 37 Eukerria, 44 Histomonas, 194 Eupolygaster, 41 Hoplitophyra, 124 Eupolytoreutus, 48 Hoplochaetella, 46 Euscolex, 48 Hoplochaetina, 46 Eutoreutus, 48 Hormogaster, 51, 54 Eutrigaster, 47 redii f. gigantes, 54 Eutyphoeus, 46, 77 Hormogastridae, 39, 51, 54 excavatus, Pheretima, 77 Hormogastrinae, 51 Exxus, 44 Holoscolex, 50 Howascolex, 46 f e sti'vus, Lumbricus, 67, 107, 138, hupeiensis, Pheretima, 62, 110, 115, 199,208,214 118, 121, 128, 132, 133, 213 Fimoscolex, 50 Hyperiodrilus, 48, 66, 113 foetida, Eisenia, see Eisenia ajricanus, 120, 121, 125, 129, Fraxinus pennsylvanica, 170 131 Frideri'cia parasitica, 123 Hystrichis tricolor, 194 jriendi, Lumbricus, 208 Fusarium, 161, 192 icterica, Eisenia, 208 Eophila (Allolobophora), 126, 217 galli, Ascaridia, 196 idosus, Bacillus, 158 Gardullaria, 48 inaequalis, Ventura, 161 gieseleri, Bimastos, 215 Iridodrilus, 48 Glossoscolecidae, xi, 9, 21, 22, 39, 49, 53, 55, 62 jaculatrix, Dichogaster, 121 SYSTEMATIC INDEX 257

Kaffania, 48 151, 152, 155, 156, 157, 158, Kerria, 44 161, 165, 169, 171, 185, 187, Kerriona, 45 189, 194, 195, 196, 197, 198, kewensis, Bipalium, 120, 124 199, 200, 201, 203, 204, 206, Kynotus, 50 201,208,209,214 rubellus, 51, 58, 59, 60, 66, 67, 89, lacteum, Octolasium, 67, 107, 139, 91, 92, 106, 107, 122, 126, 129, 156, 196, 197, 199,208,216 130, 136, 138, 144, 145, 146, lacuum, Criodrilus, 62, 194 156, 165, 169, 194, 195, 197, Lampito, 44, 77 199,206,201,208,214 latus, Protoscolex, 38 Legonea,48 Maheina, 44 Lennogaster, 47 Malabaria, 44 Leptodrilus, 45 Malodrilus, 48 Leucodrilus, 47 mamma/is, Dendrobaena, 51, 58, Libyodrilus, 48 123, 199, 208, 216 limicola, Allolobophora, 208 Maoridrilus, 45 longa,Allolobophora, see marcessens, Serratia, 157 Allolobophora maugei, Testacella, 123 longifolia, Bassia, 100 Maupasella, 124 liinnbergi, Bimastos, 125, 215 mays, Zea, 196 longicinctus, Bimastos, 215 Megachaetina, 48 Lumbricidae, xi, 9, 10, 11, 12, 20, Megadrili, xi, 39 39, 40, 51-3, 54, 64, 145, 212 Megascolecidae, 7, 9, 10, 20, 21, 22, lumbrici, Polycercus, 194 38, 39, 41, 42, 43, 53, 54, 55, Lumbriculidae, 3, 37, 38 123, 213 Lumbricus, 5, 13, 18, 23, 28, 52, 66, Megascolecinae, 43 67, 68, 69, 91, 118, 212, 214 Megascolex, vii, 27, 38, 44, 55, 125 castaneus, 57, 58, 59, 60, 66, 67, Megascolides, 46, 54 98, 105, 107, 122, 126, 135, 137, australis, xi, 19, 66 138, 145, 199, 207, 208, 214 merulae, Syngamus, 195 festivus, 67, 107, 138, 199, 208, mesomelinus, Quedius (Micro- 214 saurus), 123 friendi, 208 Metadrilus, 48 herculeus, 195 Metaradiophyra, 124 terrestris, 4, 5, 7, 12, 13, 16, 20, Metascolex, 48 21, 24, 60, 61, 63, 66, 67, 69, Metastrongylus, 124, 193 10, 71, 72, 13, 74, 75, 76, 77, elongatus, 194 79, 80, 84, 87, 89, 91, 92, 95, pudentotectus, 194 97, 104, 105, 108, 109, 110, salami, 194 113, 118, 122, 125, 126, 127, Microchaetinae, 50, 55 129, 130, 131, 134, 135, 136, Microchaetus, 50, 119 140, 141, 142, 145, 146, 149, Microdrili, xi, 39, 41 258 BIOLOGY OF EARTHWORMS

Microscolex, 45, 54 Octolasium, 52, 57, 66, 140, 146, modestus, 196 189, 212, 216 phosphoreus, 53 cyaneum, 58, 67, 107, 108, 115, millardi, Perionyx, 119 122, 126, 135, 139, 185, 199, Mil/sonia, 47 206, 207, 208, 216 miniscula, Allolobophora, 217 lacteum, 67, 107, 139, 156, 196, modestus, Microscolex, 196 197, 199,208, 216 Moniligaster, 41 Oligochaeta, xi, 6, 37, 38, 39 Moniligastridae, 38, 39, 40, 123 Onesia sepulchralis, 125 Moniligastrinae, 41 subalpina, 125 Monocystis, 19, 124 Onychochaeta, 49 Monogaster, 47 Opisthodrilus, 50 mucronata, Capillaria, 195 Myxocystis, 124 palustris, Bimastos, 215 Parabursaria, 124 Naididae, 3, 38 Parachilota, 46 Nellocystis, 124 paradoxa, Paricterotaenia, 194 Nellogaster, 44 Parascolex, 48 Nelloscolex, 44 parasitica, Fridericia, 123 Nematocystis, 124 Parendrilinae, 47 Nematoda, 124, 194 Parendrilus, 48 Nematogenia, 45 Paricterotaenia paradoxa, 194 Nemantodrilus, 48 parvus, Bimastos, 19, 208,215 Nenmanniella, 48 Paulistus, 44 Neochaeta, 44 peguana, Pheretima, 194 Neodrilacae, 43 Periodrilus, 45 Neodrilus, 45 Perionyx, 7, 44, 77 nigeriae, Hippopera, 154 millardi, 119 Nocardia polychromogens, 159 Periscolex, 50 nocturna, Allolobophora, see Allolo- Perissogaster, 44 bophora Phalaris, 139 Notoscolex, 44, 121 Pheretima, 7, 8, 18, 22, 27, 40, 44, termiticola, 123 53, 55, 61, 65, 66, 72, 81, 82, Notykus, 48 88, 91, 116, 124, 157, 194, 197, 212, 213 Ocnerodrilidae, 39, 42, 43, 44, 45, 55 agrestis, 88 Ocnerodrilinae, 43 californica, 132, 213 Ocnerodrilus, 45 elongata, 196 africanus, 194 excavatus, 77 octaedra, Dendrobaena, see Dendra• hilgendorf, 62 haena hupeiensis, 62, 110, 115, 118, 121, Octochaetidae, 39, 42, 43, 46 128, 132, 133, 213 Octochaetus, 46 peguana, 194 SYSTEMATIC INDEX 259

posthuma, 35, 119, 213 Rhabditis, 124 stelleri, 8 Rhinodrilus, 49 phosphoreus, Microscolex, 53 Rhododrilus, 45 plica, Capillaria, 195 Rhynocystis, 124 Plagiochaeta, 45 ribaucourti, Allolobophora, 217 Plagiotoma, 124 riparia, L>iplocardia, 214 Platydrilus, 47 Rillogaster, 47 Platyhelminthes, 123 robur, Quercus, 170 platyura,L>endrobaena, 215 robusta, Allolobophora, 123 Pleurocystis, 124 Rosadrilus, 48 Pliconogaster, 44 rosea, Allolobophora, see Allolo- Plutellus, 38, 46, 54 bophora Pollenia rudis, 124 Rotatoria, 124 Polycercus, 124 rubellus, Lumbricus, see Lumbricus lumbrici, 194 rubida, L>endrobaena, 67, 107, 122, Polychaeta, 37 125, 126, 139, 144, 199, 206, polychromogens, Nocardia, 159 201,215 Polytoreutus, 48 rubida f. tenuis, L>endrobaena, 66, Pontodrilus, 46 208,215 Pontoscolex, 4, 50 rudis, Pollenia, 124 corethrusus, 53, 196 Porrocaecum crassum, 195 salami, Metastrongylus, 194 ensicaudatum, 19 5 Sarcophaga carnaria, 125 posthuma, Pheretima, 119, 213 haemorrhoidalis, 12-5 Priodochaeta, 47 striata, 125 Priodoscolex, 47 Schubotziella, 48 Pronaidites, 38 dunguensis, 65 Protoscolex, 3 7 Scolioscolides, 47 latus, 38 scutulum, Testacella, 124 Protozoa, 124, 194 sepulchralis, Onesia, 125 pudentotectus, M etastrongylus, 194 Serratia marcessens, 157 putorii, Capillaria, 195 singularis,L>iplocardia, 213 pygmaea, L>endrobaena, 208, 216 skrjabinomorpha, Syngamus, 124 Pygmaeodrilus, 45, 55 Sparganophilinae, 50 Pythium, 161, 192 Sparganophilus, 50 Spenceriella, 46 Quechua, 44 Sphaeractinomyzon, 124 Quedius (Microsaurus) mesomelinus, Spirochaeta, 124 123 Spiroptera, 124 Quercus robur, 170 turdi, 195 stelleri, Pheretima, 8 Ramiella, 46 Stephanurus, 124, 195 redii f. gigantes, Hormogaster, 54 striata, Sarcophaga, 125 260 BIOLOGY OF EARTHWORMS

Streptomyces coeUcolor, 159 Tilletia controversa, 161, 192 subalpina, Onesia, 125 Tonoscolex, 47 subrubicunda, Dendrobaena, see trachea, Syngamus, 193, 195 Dendrobaena Travoscalides, 47 suum, Ascaris, 196 tricolor, Hystrichus, 194 sylvaticus, Didymogaster, 19 Trigaster, 46, 54 Syngamus, 124 Tritogenia, 50 merulae, 19 5 Trocheta subviridis, 124 skrjabinomorpha, 195 Tubificidae, 38 trachea, 193, 195 tumidis, Bimastos, 215 Syngenodrilinae, 41 turdi, Spiroptera, 195 Syngenodrilus, 41 Synoecnema, 124 Udeina, 46 undula, Dilepus, 194 Taenia cuneata, 124 Teleudrilus, 48 veneta, Eisenia, 194 tenuis, Bimastos, 8, 215 f. typica, Dendrobaena, 208, 216 termiticola, Notoscolex, 123 Ventura inaequalis, 161 terrestris, Allolobophora, 123, 217 vogesiana, Allolobophora, 217 terrestris, Lumbricus, see Lum- bricus waltoni, Entyphoeus, 65 Testacella haliotidea, 123 Wegeneriella, 47 maugei, 123 Woodwardiella, 44 scutulum, 124 tetraeda, Eiseniella, 199, 207 Yagansia, 45 f. hercynia, Eiseniella, 208, 214 f. typica, Eiseniella, 139, 208, 214 Zapotecia, 46, 54 Thamnodrilus, 49 Zea mays, 196 Thelohania, 124 zeteki, Bimastos, 62, 70, 72, 122, Thominx aerophilus, 195 215 Author index

Agarwal, G. W., 196, 219 Barley, K. P., 98, 103, 117, 138, 139, Aichberger, R. von, 157, 219 140, 145, 147, 148, 149, 151, 162, Aisyazhnyuk, A. A., 219 176, 177, 196, 220,221 Alicata, J. E., 194, 246 Barrett, T. J., 221 Allee, W. C., 125, 219 Bassalik, K., 155, 157, 163, 165, 221 Allen, R. W., 219 Bather, E. A., 38, 221 Anderson, H. L., 237 Bauer, K., 181, 221 Anstett, M., 162, 219 Baweja, K. D., 161, 221 Arbit, J., 76, 219 Baylis, H. A., 121, 123, 221 Arldt, T., 38, 219 Beauge, A., 122, 221 Arlidge, G. Z., 183, 184, 234 Bejsovec, J., 196, 221 Arnold, M. K., 183, 226, 232 Belousova, N. K., 103, 226 Arrhenius, 0., 125, 219 Benham, W. B., 11, 26, 38, 45, 46, Arthur, D. R., 81, 93, 179, 180, 208, 50,221 219 Beynon, K., 227 Arthur, J. H., 227 Bharucha-Reid, R. P., 76, 221 Ash, J. S., 224 Bhatt, J. V., 147, 157, 161, 234 Aspi:ick, H., 184, 236 Bigger, J. H., 182, 221 Atlavinyte, 0., 156, 219, 220 Blancke, E., 147, 163, 221 Avel, M., xiii, 219 Blankwaardt, H. F. H., 181, 221 Block, W., 104, 221 Bachelier, G., xiii, 20, 125, 137, 180, Bocock, K. L., 222 220 Bodenheimer, F. S., 168, 222 Bahl, K. N., 18, 61, 83, 118, 220 Bollen, W. B., 153, 242 Baker, W. L., 220 Bornebusch, C. H., 98, 125, 222 Bakhtin, P. U., 165, 166, 220 Bouche, M. B., 102, 222 Baldwin, F. M., 74, 77, 220 Boyd, J. M., 107, 137, 222 Ball, D. F., 242 Boynton, D., 133, 222 Ball, R. C., 220 Bradley, J., 165, 168, 223 Baluev, V. K., 168, 220 Braunig, G., 80, 88, 238 Banage, W. B., 104, 221 Bray, J. R., 146, 222 Barker, R. J., 183, 185, 220 Bretnall, G. H., 222 262 BIOLOGY OF EARTHWORMS

Bretscher, K., 98, 222 Dennis, E. B., 182, 183, 226 Breza, M., 194, 222 Devigne, J., 224 Brown, B. R., 140, 222 Dhawan, C. L., 224 Brown, D. M., 222 Dhennin, L., 193, 224 Bruel, W. E. van der, 160, 222 Dingwall, A. R., 107, 170, 231 Briisewitz, G., 157, 158, 159, 222 Doane, C. C., 182, 183, 185, 225 Buahin, G. K. A., 181, 221 Dobson, R. M., 171, 225 Buntley, C. J., 223 Doeksen, J., xiii, 69, 77, 79, 99, 126, Byzova, Yu B., 182, 183, 223, 230 225, 229 Doerell, E. C., 180, 225 Capstick, C. K., 222 Dotterweich, H., 127, 225 Carter, G. S., 96, 223 Dowdy, W. W., 110, 131, 225 Causey, D., 223 Dreidax, L., 169, 226 Cernosvitov, L., 52, 123, 223 Drift, J. van der, xii, 181, 184, 225, Chadwick, L. C., 165, 168, 223 226, 229 Chandler, R. F., 100, 226 Dubash, P. J., xiii, 64, 65, 78, 96, Chapman, G., 223 248 Chen, C. M., 190, 223 Duley, F. L., 248 Cockerell, T . D. A., 125, 223 Dustman, E. H., 185, 226 Cohen, S., 148, 223, 224 Dutt, A. K., 165, 166, 173, 226 Coin, C. J., 223 Dzangaliev, A. D., 103, 226 Combault, A., 23, 169, 223 Compton, 0. C., 133, 222 Eaton, T . H., Jr, 100, 226 Conder, P. J., 185, 224 Eberhardt, A. 1., 125, 226 Cossens, G. G., 168, 170, 173, 247 Edwards, C. A., 99, 103, 140- 2, 144, Couperus, H., 79, 225 150, 174, 175, 181-8, 190, 204, Couture, G., 189, 237 221, 224, 226, 227, 232 Cragg, J. B., 224 El-Duweini, A. K., 98, 104, 126, 128, Cramp, S., 185, 224 132, 136, 226, 227 Crompton, E., 179, 180, 224 Ellenby, C., 192, 227 Crossley, D. A., 140, 146, 188, 224, Ellis, D. J., 237 227 Empson, D. W., 226 Curry, L. L., 220 Escherich, K., 123, 227 Escritt, J. R., 179-81, 227 Darwin, C., xii, 74, 88, 121, 139, Evans, A. C., 52, 56- 60, 66, 68-71, 196,224 100, 104, 105, 110--13, 118, 119, Davey, S. P., 182, 224 122, 123, 128, 129, 130, 138, 145, Davis, B. N. K., 185, 186, 224 163, 167, 174, 176, 197- 9 Dawson, A. B., 32, 157, 165, 166, 224 Feldkamp, J., 6, 228 Dawson,R. C., 224 Fenton, G. R., 228 Day, G. M., 147, 157, 158, 224 Finck, A., 127, 153, 165, 228 Decker, G. C., 182, 186, 221, 237 Fleming, W. E., 183, 228 AUTHOR INDEX 263

Ford, J., 98, 228 Hadley, C. H., 183, 228 Fox, C. J. S., 181, 228 Hamblyn, C. J., 107, 170, 231 Franz, H., 146, 228 Handa, B. K., 224 French, M. C., 186, 224 Handley, W. R. C., 222 Hanel, E., 74, 231 Gansen, P. S. van, 80, 82, 228 Hanley, I. M., 228 Garner, M. R., 118, 206, 228 Hardman, J. A., 185-7, 249 Gast, J., 139, 228 Harmsen, G., 161, 231 Gates, G. E., 39, 42-7, 51, 69, 77, Harrison, R. B., 185, 186, 224 115, 121, 229 Hasenbein, G., 197, 231 Genov, T., 194, 229 Haswell, W. A., 124, 231 Geoghegan, M. J., 166, 229 Heath, G. W., 103, 140-2, 144, 149, Gerard, B. M., 56, 60, 68, 106, 108, 179, 204, 226, 227, 231, 232, 109, 112, 114, 115, 127-9, 229 235 Gersch, M., 229 Heck, L. von, 75, 76, 232 Gewehr, H., 245 Heimburger, H . V., 232 Ghabbour, S. 1., 98, 104, 121, 126, Hensen, V., xii, 232 128, 132, 132, 136, 160, 226, 227, Herlant-Meewis, H., 87, 232 229 Hess, W . N., 28, 88, 232 Ghilarov, M. S., 157, 168, 172, 182, Heungens, A., 144, 182, 184, 232 183, 229, 230 Hill, J. P., 124, 231 Giesecke, F., 147, 163, 221 Hirst, J."M., 161, 232 Gilbert, 0., 222 Hobmaier, A., 194, 232 Gish, C. D., 186, 187, 230 Hoeksema, K. J., 165, 232 Glasgow, L. L., 187, 247 Hoffman, J. A., 161, 192, 232 Goffart, H., 183, 184, 230 Hogben, L., 131, 232 Gorham, E., 146, 222 Hollister, P. L., 219 Graff, 0., xiii, 60, 130, 132, 148, Hopkins, A. R., 165, 174-7, 182-4, 153, 154, 174, 176, 206, 230 232, 233 Grant, W. C., 62, 106, 115, 118, 127, Hopkins, H. T., 233 131, 132, 172, 230 Hopp, H., 98, 112, 115, 131, 165, Grassi, B., 194, 230 167, 16~ 171, 174-~ 233,246 Gray, J., 94, 230 Howell, C. D., 88, 233 Greenwood, D. E., 183, 230 Hoy, H. M., 182, 183, 233 Griffiths, D. C., 184, 231 Hub!, H., 87, 233 Grigor'eva, T. G., 182, 231 Hunt, L. B., 185, 186, 233 Grove, A. J., 14, 15, 17, 25, 26, 29, Hutchinson, S. A., 158, 161, 191, 34, 231, 240 233 Guild, W. J., MeL., 137, 138, 145, Hyche, L. L., 184, 233 165, 167, 174, 176, 197-9, 227, Hyman, L. H ., 69, 233 228,231 Gunthart, E., 182, 231 Inove, T ., 190, 233 Gurianova, 0. Z., 165, 231 Isa, A. L., 237 264 BIOLOGY OF EARTHWORMS

Jacks, G. V., 167, 233 Kubiena, W. L., 137, 235 Jacob, A., 179, 180, 233 Kiihnelt, W., xiii, 235 Jacobson, G. M., 148, 153, 173, 237 Kurcheva, G. F., 142, 235 Jeanson-Luusinang, C., 126, 158, 234 Laan, H. van der, 184, 236 Jefferson, P., 100, 176, 179, 180,234 Ladell, W. R. S., 99, 235 Jennings, A. C., 148, 149, 162, 220 Lahr, J. P., 219 Jevniaux, C., 224 Lakhani, K. H., 236 Joachim, A. W. R., 165, 234 Lauer, A. R., 76, 236 Johnson, M.,L., 79, 80, 234 Laverack, M. S., xiii, 16, 78, 83, 84, Johnstone-Wallace, D. B., 99, 179, 89, 140, 236 234 Lawrence, R. D., 148, 236 Jongerious, A., 232 Lee, K. E., 68, 42-4, 168, 197, 236 Joshi, N. V., 163, 168, 234 Legg, D. C., 182-4, 236 Leitenberger, L., 146, 228 Kahsnitz, H. G., 169, 234 Lesser, E. J., 79, 236 Kalmus, H., 234 Lewis, H. B., 148, 223, 224 Kamel, M., 158, 161, 191, 233 Lidgate, H. J., 183, 236 Kanwar, J. S., 165, 240 Lindquist, B., 139, 147, 236 Karmanova, E. M., 194, 195, 234 Lipa, J. ]., 182, 236 Karpachevskii, L. 0., 144, 241 Lissman, H. W., 94, 230 Keilin, D., 125, 234 Liv, C. L., 223 Kelkar, B. V., 163, 168, 234 Ljungstrom, P. 0., 119, 123, 197,236 Kelly, W. A., 243 Lofty, J. R., 103, 121, 131, 134, 135, Kelsey, J. M., 183, 184, 234 171, 174-6, 181-3, 188, 225-7, Kevan, D. K. McE., xiii, 234 231, 237, 243 Khambata, S. R., 147, 157, 161, 234 Long, W. H., 183, 237 King, H. G. C., 140, 142, 232, 235 Love, C. W., 222 Kirberger, C., 79, 235 Low, A. J., 98, 165, 237 Kirk, R. L., 131, 182- 4, 232 Lowe, D. G., 245 Kirk, V. M., 233 Luckman, W. H., 183, 186, 237 Kleinig, C. R., 145, 151, 221 Lugauskas, A., 156, 220 Knop, J., 27, 235 Lukose, J., 123, 237 Kobatake, M., 160, 235 Lund, E. E., 194, 237 Kollmannsperger, F., 235 Lunt, H. A., 148, 153, 173, 237 Kollmannsperger, G., 112, 122, 130, 137,235 Madge, D. S., 67, 104, 115, 121, 125, Kondo, K., 190, 233 129, 131, 146, 237 Korschelt, E., 61, 235 Magalhaes, P. S., 194, 237 Kozlovskaya, L. S., 156, 158, 235 Maldague, M., 189, 237 Kring, J. B., 184, 235 Mamajev, B. M., 172, 230 Krivanek, J. 0., 76, 235 Mamytov, A., 165, 166, 237 Kriiger, F., 78, 103, 235 Mangold, 0 ., 89, 139, 237 AUTHOR INDEX 265

Marapao, B. P., 237 Nijhawan, S.D., 165, 240 Marshall, V. G., 171, 237 Nye, P. H., 122, 127, 153, 154, 240 Martin, A. W., 84, 166, 238 Martin, J. P., 249 O'Connor, F. B., 243 Mayne, D. W., 247 Ogg, W. G., 180, 240 McCalla, T. M., 176, 248 Oldham, C., 240 McLeod, J. H., 123, 237 Olson, H. W., 125, 127, 240 Meer, K. van der, 232 Omodeo, P., 42, 43, 241 Meggitt, F. J., 194, 238 Otanes, F. G., 190, 241 Mellanby, K., 238 Merker, E., 80, 88, 238 Panditesekera, D. G., 165, 234 Meyer, L., 163, 238 Papedick, R. I., 223 Michaelsen, W., 37-40, 44-8, 50, Pare!, T. S., 241 51,238 Parker, G. H., 90, 241 Michon,}., 60, 61, 69, 130, 238 Parle, J. N., 155, 156, 159, 160, 166, Miles, H. B., 160, 238 241 Millar, H. R., 148, 236 Parshley, H. M., 90, 241 Millott, N., 81, 238 Patel, H. K., 183, 190, 241 Minderman, G., 225 Patel, R. M., 241 Moment, G. B., 69, 95, 238, 239 Pathak, A. N., 163, 246 Monnig, H. 0., 194, 239 Pattarudriah, M., 190, 196, 242 Moore, A. R., 239 Peachey, J. E., 241 Moore, B., 52, 90, 125, 239 Peredel'sky, A. A., 188, 241 Morris, H. M., 99, 176, 239 Perel, T. S., 144, 241 Morrison, F. 0., 182, 239 Perrier, 44-6, 48, 4Q. Mozgovoy, A. A., 195, 239 Petrov, B. C., 125, 241 Muldal, S., 239 Phillips, E. F., 125, 241 Muller, G., xii, xiii, 139, 168, 239 Pickard, J. A., 183, 185, 186, 247 Muller, P. E., 239 Polivka, J. B., 181- 3, 241, 242 Murchie, W. R., 62, 68, 70, 72, 106, Polsky, M. N., 165, 166, 220 122, 239, 240 Pomerat, G. M., 79, 242 Ponomareva, S. 1., 153, 155, 165, Nakamura, Y., 115, 116, 125, 240, 166, 176, 242 248 Poryadkova, N. A., 241 Nathans, S., 98, 103, 243 Powers, W. L., 153, 242 Needham, A. E., 146, 148-50, 240 Prabhoo, N. R., 242 Negi, L. S., 219 Pratt, K. C., 69, 248 Nelson, J. M., 98, 240 Prosser, C. L., 88, 242 Newell, G. E., 14, 15, 17, 25, 26, 29, Puh, P. C., 127, 153, 242 34, 231, 240 Purdy, L. H., 161, 192, 232 Nicoi, H., 180, 240 Nielson, C. 0., 240 Radionova, L. Z., 241 Nielson, R. L., 168, 170, 171, 240 Raffy, A., 79, 242 266 BIOLOGY OF EARTHWORMS

Ragg, J. M., 242 Schmidt, H., SO, 76, 245 Ralph, C. L., 79, 242 Schneider, K. C., 31, 245 Ramsay, J. A., 83, 91, 242 Schread, J. C., 121, 182- 4, 191, 246 Rao, K. S. K., 219 Schrevan, D. van, 161, 231 Raw, F., 99-101, 103, 104, 134, 141, Schwartz, B., 194, 246 142, 144, 146, 181, 184, 231, 242, Scopes, N. E. A., 184, 249 243 Scott, H. E., 184, 246 Reichle, D. E., 224, 227 Scrickhande, J. C., 163, 246 Reinecke, A. J., 119, 123, 197, 236 Sharma, R. L., 224 Reynoldson, T. B., 98, 103, 243 Shindo, B., 246 Rhee, J. A. van, 98, 103, 104, 146, Shiraishi, K., 133, 246 169, 171, 243 Sims, R. W., 42, 43, 66, 246 Rhoades, W. C., 183, 243 Singh, A., 224 Ribaudcourt, E., 169, 243 Sison, P. L., 190, 241 Richards, J. G., 170, 243 Skarbilovic, T. S., 195, 246 Richardson, H. C., 125, 134, 179- Slater, C. S., 167, 169, 171, 175, 180, 81, 244 181, 233, 246 Richter, G., 182, 183, 244 Smallwood, W. M., 30, 86, 246 Robertson, J.D., 80, 244 Smith, F., 41, 187, 247 Robinson, J. S., 75, 76, 244 Smith, R. D., 247 Rodale, R., 180, 183, 244 Sokolov, D. F., 241 Roots, B. 1., 82, 90, 92, 127, 129, Stafford, C. J., 227 244 Stephenson, J., xii, 9, 12, 13, 16, 23, Rovelli, G., 194, 230 24, 31, 33, 35, 37- 40, 42- 4, 46, Ruschmann, G., 147, 244 49, 62, 66, 79, 91, 94, 124, 197, Russell, E. J., xii, 133, 14 7, 168, 244 247 Rysavy, B., 194, 195, 244 Stickel, L. F., 185, 226, 247 Ryzhikov, K. M., 195, 244 Stickel, W. H., 247 Stockdill, S. M. J., 168, 170, 173, Sacho, R. J., 186, 233 247 Salisbury, E. J., xii, 125, 127, 172, Stockli, A., 98, 122, 127, 147, 153, 244 155,.159, 167, 247 Saroja, K., 79, 244 Stokes, B. M., 247 Sastry, K. S. S., 190, 196, 242 Stolte, H. A., xiii, 124, 196, 247

Satchell, J. E., xiii, 58, 60, 61, 661 St0p-Bowitz, C., 247 68, 69, 89, 90, 98, 100, 102, 107, Storey, I. F., 232 108, 125, 126, 130, 133, 135, 137, Stough, H. B., 31, 247 139, 140, 142, 146, 148, 149, 151, Stringer, A., 183, 185, 186, 247 152, 155, 160, 176, 230, 236, 240, Sun, K. H., 69, 248 245 Svendsen, J. A., 98, 103, 107, 138, Saussey, M., 123, 168, 245 248 Scharpenseal, H. W., 245 Swaby, R. J., 165, 166, 248 Schmid, L. A., 87, 245 Swartz, R. D., 76, 210, 248 AUTHOR INDEX 267

Takano, S., 125, 248 Way, M. J., 184, 249 Tembe, V. B., xiii, 64, 65, 78, 96, Weber, G., 182, 184, 249 248 Wehr, E. E., 237 Tenney, F. G., 161, 248 Weisbach, W. W., 197, 249 Teotia, S. P., 156, 163, 165-7, 176, Went, J. C., 157, 249 248 Wheatley, G. A., 185-7, 249 Thompson, A. R., 183, 184, 186, Wherry, E. T., 125, 126, 249 227,248 Whiting, A. E., 227 Tischler, W., 176, 248 Whitney, W. K., 184, 249 Torvik, M. M., 219 Wiegland, K., 179, 180, 233 Tracey, M. V., 81, 248 Wilke, D. E. von, 60, 61, 98, 249 Trifonov, D., 190, 248 Wilcox, H. G., 232 Tromba, F. G., 195, 248 Wingerden, C. G. van, 69, 126, 225 Twinn, D. C., 222 Witkamp, M., 150, 250 Wittich, W., 89, 150, 250 Uhlen, G., 169, 180, 248 Wojewodin, A. W., 181, 250 Urquhart, A. T., xii, 248 Wolf, A. V., 90, 131, 250 Wollny, E., 167, 168,250 Villot, F. C. A., 194, 249 Woodhead, A. A., 195, 250 Vogel, R., 194 Woodman, M. G., 222 Volz, P., 168, 249 Yegorova, S. V., 241 Waid, J. S., 222 Yerkes, R. M., 75, 76, 250 Waite, R. H., 249 Waksman, S. A., 161, 166, 248, 249 Zarrow, M. T ., 79, 242 Walton, W. R., 99, 190, 249 Zhdannikova, E. N., 156, 158 Ward, W. C., 232 Zhinkin, L., 96, 250 Waters, R. A. S., 112, 117, 139, 169, Zicsi, A., 98, 102, 175, 190, 250 170, 176, 249 Zrazhevski, A. I., 156, 157, 170, Watkin, B. R., 179: 249 250 General index

(Italic page numbers indicate the more important references in the text whereas bold numbers refer to illustrations.) Absorption zone, 81 Aggregations, 106, 107, 131, 139, Accessory reproductive bodies, 7 165- 7,209 Acetylcholine, 86, 96 Agriculture, effects on earthworms, Acid, 89 174-89 Acid-intolerant species, 125, 126 Alaska, 54 Acid soils, 88, 125, 127, 169, 178- Albumen cells, 4, 14, 15, 36 81, 201 Alder, 146 Acid-tolerant species, 125, 126 Aldicarb, 184 Actinomycetes, 155, 156- 8 Aldrin, 182, 187 Activity, effects of temperature Alfalfa, 165, 176 on, 130 Algeria, 51 seasonal changes in, 113 Alimentary canal, 17, 19-21, 124 Activity cycles, 87 innervation of, 30, 31 Adders, 123 Alimentary plexus, 23 Adolescence, 70 Allantoin, 81, 148 Adrenaline, 87 Alluvium, population in, 136, 137 Adverse conditions, 62, 74, 88 Alpine forest, 146 Adverse effects, on earthworms, America, Central, 46, 49, 50, 53-5 196, 197 North, 46, 49-52, 54 on soil, 196, 197 South, 46, 47, 49,50,52- 4 Adverse periods, 67 Ammonia, 81- 4, 118, 127, 148 Aeration, 106, 133- 6, 167, 168 Ammonium, 149, 162 effects of, 13 3 Ammonium sulphate, 181 Aestivation, 63, 67- 9, 100, 127 Amoebocytes, 18, 19, 82 Affinities, 37-9 Ampulla, 27 Africa, 41, 44-8, 50-5, 67, 121 Amylase, 81, 121 Age-class composition, lOS Anabiosis, 91 Aggregates, 165- 7 Anaerobic conditions, 77, 96 GENERAL INDEX 269

Anaerobic respiration, 79 Basal cells, 15 Andes, 45, 50 Basal membrane, 16 Antarctica, 55 Basalt, 163 Antarctic Islands, 45 Base exchange capacity, 153 Antibiotic, 160 Basement membrane, 15, 16 Anus, 4, 19, 69, 90 Bassia tree, 100 Apple scab, 161 Beans, 196 Aquatic species, 9, 10 Beech, 140-2, 144, 198 Aquatic worms, 38, 66, 91 Beet, 140 Arable, 99, 103, 104, 122, 123, 131, Behaviour, 71- 7 135, 166, 174, 176, 190 Behavioural experiments, 210, 211 Arciform muscles, 63 Benefactors, earthworms as, 197 Arctic forest, 146 Benzopyrine, 197 Argentina, 49, 50 Bermuda, 49 Artesian irrigation, 172 BHC, 182 Arthropods, 14 7 Biomass, xii, 102 Ascospores, 161 Birch, 140, 144, 150 Asexual reproduction, 67 Birds, 123, 193-6 Ash, 140, 149, 150, 170 Black head, 194 Asia, 44, 50-3, 55 Bladex, 181 Association, 123 Blood, 90, 124 Association neurones, 86 Blue-green algae, 157 Atrazine, 181 Body wall, 13, 14, 15- 17, 33 Auckland, 45, 46 Bolivia, 49, 50 Australasia, 55 Borneo, 41 Australia, 44-6, 53, 55, 103, 115, Bowling green, 191 151, 176 Brazil, 44-6, 49, 50 Austria, 51 Breeding, 61 Autotomy, 72 Breeding period, 62 Available mineral nutrients, 152- 4 Bristle worms, 37 Axon,32 Brown bodies, 19 Azinphosmethyl, 184 Buccal cavity, 4, 19, 21 Azores, 51, 52 innervation of, 30, 32 Buccal chamber, 80, 89 Bacteria, 27, 80, 147, 155, 156, 158, Buccal epithelium, 89 160, 161, 166 Bulgaria, 190 Bacterial gums, 166, 167 Bullock droppings, 58, 198 Bacteriocide, 160 as food, 138 Bacteroidal cells, 82 Burma, 41, 50, 55, 115, 121 Badgers, 123 Burrowing, 92, 117-22 Bahamas, 41, 45 Burrowing experiments, 204- 6 Bardsey Island, 103 Burrows, 73, 74, 81, 99, 100, 102, Barley, 169 108, 174, 185, 190,204 270 BIOLOGY OF EARTHWORMS

Burrows--{con td.) microbes in, 164 depth of, 118 pH of, 127, 172 permanent, 118 seasonal production, 112, 119 Bursa propulsoria, 66 stability of, 173 Butyric acid, 156 Cattle, 145 Caucasus, 52 Caesium, 189 Cells, albumen, 4, 14, 15, 36 Cages, 142 central, 25, 26 Cage for studying earthworm acti- chloragogen, 13, 18, 82, 83, 96 vity, 205 ciliated, 21 Calcareous sand, population in, 137 clitellar gland, 18, 65 Calciferous gland, 20, 42-4, 127, 166 cocoon-secreting, 36 Calcium, 153, 154, 166 coelomic epithelial, 17 Calcium carbonate, 80 drainpipe, 26 Calcium humate, 166 epidermal, 15, 36 California, 45, 46 epithelial, 15, 36, 81 Lower,44 epithelial brush border, 81 Cameroons, 47, 48, 50, 122 excretory, 82 Campbell Islands, 45 giant, 32 Canada, 54 glandular, 4, 15, 21, 24, 36 Canary Islands, 51, 52 goblet, 14 Cape of Good Hope, 54 light-sensitive, 88 Cape Verde Islands, 45 marginal, 25, 26 Capillaries, 85 mucous, 4, 15, 36 intra-epidermal, 24 nerve, 87 Capillary blood vessels, 13 peritoneal, 82 Capillary water, 167 photoreceptor, 4, 15, 16, 32 Capricorn, tropic of, 54 proprioreceptor, 32 Carabid beetles, 123 replacing, 15 Carbaryl, 184, 191 reproductive, 62 Carbofuran, 184 secretory, 87 Carbohydrates, soluble, 140 sensory, 14, 15, 87 Carbon nitrogen ratio, 150-2 supporting, 16 Carbon dioxide, 78, 80, 133 uric, 81, 82 tension, 151 Cellulase, 81 , 121, 155 Caroline Islands, 41 Cellulose, 150, 156 Carrots, 196 Central cells, 25, 26 Casting, 117-22 Cephalization, 3, 52 time of day of, 121 Cereals, 166, 178 Castor meal, effect of, 134 Cereal stooks, 190 Casts, 81, 111, 120, 148, 152, 154, Cerebral ganglia, 28, 32, 75, 87, 88, 156, 157, 159, 160, 163,165,167, 210 169,176,180,190,191,196, 201 Ceylon, 41, 44- 6, 55, 123 GENERAL INDEX 271

Chaetopoda, xi Cocoon production, 61, 64, 66, 87, Chaetotaxy, 4-7 106 Chemical sampling methods, 100, effects of food on, 138 110, 111 effects of soil moisture on, 128 Chemoreception, 88, 89 effects of temperature on, 130 Chemozem, 155 Cocoon secreting cells, 36 Chickens, 194, 195 Cocoon shape, 66 Chile, 45, 51, 52 Cocoon size, 66 China, 41, 55, 190 Cocoon wall, 66 Chitin, 155 Cocoons, 37, 56, 57, 67, 92, 107, Chitinase, 81, 121 110, 115, 198, 199, 206, 207 Chloragogen cells, 13, 18, 82, 83, 96 seasonalproductionof,57,58, 114 Chloragosomes, 18, 82 Coelom, 13, 16, 17-19, 24, 28, 84, Chlordane, 182, 187, 191 124 Chlorfenvinphos, 184 Coelomic epithelial cells, 17 Chloropicrin, 181 Coelomic fluid, 9, 13, 18, 19, 23, 72, Chlorpropham, 181 78, 82,83, 90-2,127 Choice experiments, 75 Coelomocytes, 77 Ciliated cells, 21 Collagenous fibres, 4, 13 Circular muscle, 4, 13, 14, 16, 17, Colombia, 45, 49, SO 24,92 Colour of earthworms, 13 innervation of, 30 Commensalism, 123 Circulation, 84, 85 Commissural vessel, 22 Circulatory system, 20, 21 Comparisons of sampling methods, Circumoesophageal nerve ring, 94 101, 102 Circumpharyngeal connectives, 28, Compost, 172 29, 30, 88 Concentration factor, 186 Citric acid, 89 Condensed tannin, 140 Clasping, 65 Conductivity, 99 Clay, 108, 118, 163 Congo, 47,50 population in, 136 Connectives, circumpharyngeal, 28, Clay chimneys, 121 29, 30, 88 Climate, 62 subpharyngeal, 30 Cli tellar gland cells, 18, 65 Control of earthworms, 191 Clitellum, 9, 10, 11, 12, 32, 35, 36, Copper sulphate, 181 41, 49, 51, 52, 61, 63, 64, 69 Copulation, 11, 39, 62, 63- 5 Clitellum position, 9, 12 Copulatory pouches, 65 Clover, 139, 162, 169, 178 Corsica, 51 Cluster flies, 124 Costa Rica, 45, 4 7 Cobalt, 188 Cotton, 187 Cocoon, development time of, 59, 60 Creatinine, 83 formation, 10, 65 Cretaceous era, 38 incubation time, 60 Crop, 19, 20, 80, 157, 185 272 BIOLOGY OF EARTHWORMS

Cropping, effects of, 176-8 Dispersive powder, 106 Crop yields, effect of earthworms Dissection, 200 on, 168-71, 177 Distributions, 200 Cuba, 45 Disulfoton, 96 Cultivation, effects on earthworms, Diurnal activity, 151, 152 174-6, 175 Diurnal cycles, 76, 77, 79 Cultivations, 182 Diurnal respiratory cycles, 87 effects of, 174-6 Diurnal weight changes, 90 Culture, growth in, 60 DNOC, 181 Culture methods, 198-200 Dog's mercury, 140 Cultures, 66, 69, 206, 207 Dorsal pores, 2, 9, 19, 23, 52, 72, 90, Cuticle, 4, 13-16 127 Cysticercoid stage, 124 Dorsal vessel, 17, 20, 21, 22, 23, 30, Cysticercus, 193 84, 85 Dorso-intestinal vessel, 17, 23 D- D, 181 Dorsa-subneural vessel, 17, 21- 3, DDE, 185, 187 85 DDT, 183, 185, 187 Drainage, 167, 168 Dead roots, 112 Drainpipe cells, 26 Death, 61, 69, 70 Droughts, 127 Deciduous woodland, 146 Ducks, 194 Dehydration tropism, 90 Dung, 106, 138, 139, 145, 146, 162, Dendrogram, 42 169, 170, 176, 180 Denmark, 125 effects of, 134, 135 Deserts, 53, 137 Dung-pats, 107 Desmids, 157 Dursban, 184 Desiccation, 67, 109 Dutch elm disease, 185 Development, 61, 69 Dwarf bunt, 161, 192 Diapause, 63, 67- 9, 128, 201 Dyfonate, 184 facultative, 57, 68, 111 obligatory, 57, 68, 111, 207 Earthworm farms, 171, 197 Diatoms, 157 Earthworm pies, 197 Diazinon, 184 Earthworms, as benefactors, 197 Dicotyledenous plants, 38 as pests, 190, 191 Dieldrin, 183, 187 Ecology, 98-140 Diets, 206 Ecuador, 46, 49, 50 Digestion, 80, 81 Eelworm cysts, 192 Digestive system, 20 Eggs, 207 Dimensions of worms, 208 Egypt, 50, 52, 62, 104, 126, 138 Dimethylbenzanthrene, 197 Elderberry, 140 Dipterous larvae, 124 Electrical inhibition of growth, 95 Disking, effects of, 17 4, 17 5 Electrical potential, 95 Disease transmission, 191-6 Electrical sampling methods, 99 GENERAL INDEX 273

Electrical stimuli, response to, 90 Feeding, 72, 74 Electric shock, 210 Female pores, 8, 9, 10, 12, 63, 40, Electrode, 100 41, 49, 51 Eleocytes, 18, 19 Female vagina, 196 Elm, 140, 144, 150 Fenestrae, 31 Endemic species, 53 Fertilization, 63, 65, 66 Endrin, 183, 187, 191 Fertilizer, 171 England, 54 Fertilizers, effects of, 178-81 Environmental factors, influence of, Fever, 197 125- 40 Flatworm, 124 Enzymes, 80, 81, 141, 155 Flooded soil, 92, 129 Epidermal cells, 15, 36 Fibre, collagenous, 4, 13 Epidermal sense organ, 15 intra-epidermal nerve, 86 Epidermis, 4, 9, 13, 14, 16, 23, 24 lateral giant, 31, 32, 86 Epilobous prostomium, 3 median dorsal longitudinal giant, Epineurium, 31 85,86 Epithelial brush-border cells, 81 motor, 86 Epithelial cells, 15, 81 Fibres, nerve, 15 Epithelial sense organ, 4, 14, 32 Fibrillae, 31 Epithelium, 4, 19, 23, 30 Field capacity, 167 Essential intermediate hosts, 193 Field studies, 198-211 Ethiopia, 48 Filtration, 84 Eudrilid, 63, 65 Fish bait, 99, 197 Euprostates, 47 Fistula, 196 Europe, 40, 49, 51-3, 171 5 HT, 87 Excretion, 81-5 Fodder, as food, 138 Excretory cells, 82 Follicles, 4 Excretory organs, 9 Food, 88, 137, 174, 176 Excretory system, 24, 25- 8, 42, 43 Food supply, 138-40 Experimental cages, 202 effect of, 138-40 Experiments with earthworms, Food type, effect of, 60 198- 211 Foot and mouth disease, 193 External segmental grooves, 17, 18 Forest soil, 166 Formalin, 13, 100, 102, 103, 199- Facultative diapause, 57, 68, 111 201,209 Faeces, 148, 149, 155, 171 Formalin sampling, 72, 101, 171 microbial content of, 159 Formaldehyde, 89 Fallow, 103, 121, 176 Fossil records, 37 Families, 39, 55 Fossil worms, 38 Farmyard manure, 58 Fragmentation, 141-5 Fats, 82 France, 46, 54 Fecundity, effects of food on, 138 Fried earthworms, 197 effects of temperature on, 129 Frost, effect of, 131 274 BIOLOGY OF EARTHWORMS

Fungal mycelia, 139 Greenland, 52 Fungal spores, 158, 159, 161, 191 Gregarines, 124 Fungi,80, 118,156-61,166,191,192 Growing zone, 69 Fungicide, 141 Growth, 69, 10, 71, 129 Fungicides, effects on worms, 181 effect on food on, 138 Growth periods, 59, 60 Gambia, SO Guatemala, 45, 50 Ganglia, cerebral, 28, 32, 75, 87, 88, Guinea, 47 210 Gut, 94 sub-oesophageal, 28 Gut contents, 102 pharyngeal, 30 Guthion, 184 Gate worms, 193 Guyana, 45, 49, 50 Genera, 39-55 Generator, 100 Habitat, 103 Genital openings, 7-9, 39 Haemochromagen, 84 Genital organs, 13, 37 Haemoglobin, 13, 78 Genital pores, 1, 3, 5, 7, 10 Haiti, 46 Genital seta, 5 Handsorting, 98, 99, 102, 103 Geographical distribution, 53-5 Rants, 103 Germany, xiii, 51, 103, 122 Harrowing, 174 Ghana, 48 Hawaii, 52 Giant cells, 32 Hay, 169, 178 Giant fibres, 31, 32, 86 Hearts, 20, 21, 22, 23, 44, 45, 85 Gizzard, 19, 20, 41, 49, 51, 52, 80, Heat extraction, 101 81, 157, 185 Health, 122 Glaciation, 54 Heptachlor, 183, 187, 191 Gland, calciferous, 20 Heptachlor epoxide, 187 mucous, 23, 78 Herbage, 106, 112 pharyngeal, 80 Herbicides, effects on worms, 181, salivary, 80 182 Glandular cells, 4, 15, 21, 24, 36 Hermaphrodite, 7, 37, 63 Glandular secretions, 63 Herts, 14, 103 Glucose, 16, 89, 160 Heteroxanthine, 84 Glycogen, 79, 82 Himalayas, East, 41 Goblet cells, 14 Holland, 77, 103 Golf courses, 181, 191 Holonephridia, 28, 44, 45 Granite particles, 163 Honey jars, 200 Grape husks, 162 Horizontal distribution, 209 Grass, 162, 169, 174, 176, 180, 191 Horizontal migration, 107 Grassland, 135, 176 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 Kale, 140 Human welfare, 40 Kentucky, 14 Humidity, 18 Kenya, 47 Humification, 145-7 'Kilner' jars, 198 Humus, 147 Kommetjies, 119 Hungary, 51 Hydrogen sulphide, 77 Lactic acid, 79 Hydrostatic pressure, 90 Lamellae, 31 Hypotonic urine, 91 Larch, 140 Lashing movements, 72 Iceland, 52 Lateral giant fibres, 31, 32, 86 Immature worms, proportions of, Lateral-neural vessel, 17, 22, 85 104, 105 Lateral-oesophageal subneural Impotency, 197 vessel, 22, 23 Incubation period, 62 subneural vessel, 22, 23 India, 41, 44-7, 50-3, 55, 62, 115, Latex, 206 190 Latex casts, 118, 206 Indonesia, 55 Lawns, 191 Infectivity of cysts, effects of Lead arsenate, 181 worms on, 192 Leaf burial, 202, 203 Inorganic minerals, 121 effects of temperature, 130, 131 Inorganic salts, 106 Leaf disks, 142, 203, 204 Insecticide residues, 185 rate of breakdown of, 144 Insecticides, effects on worms, Leaf litter, 106 181-8, 209 Learning, 74, 75-7, 211 uptake into worms, 186 Leaves, 74 Internal salt concentration, 91 Leeches, 37, 124 Intersegmental grooves, 1, 9, 11 Legume, 178 Intestinal microflora, 147 Lespedeza, 1 7 6 Intestinal wall, innervation, 30 Lettuce, 140, 190 Intestine, 19, 20, 21, 80, 81, 85, 96, Leys, 176, 177 157 Liberia, 45, 48 Ireland, 54 Life cycles, 56-62 Iron, 189 Life history studies, 206-8 Israel, 51, 52 Life span, 60, 61 Italy, 51 Light, 63 reaction to wavelength of, 88 Japan, 41, 51, 52, 55, 115, 190, Light loam, 118 197 populations in, 136, 137 Jaundice, 197 Light reactions, 87, 88 Java, 41, 44 Light receptors, 88 Jordan, 51 Light-sensitive cells, 88 Jurassic era, 38 Lignin, 150 276 BIOLOGY OF EARTHWORMS

Lime, 140, 144, 1 SO, 169, 178-80 Membrane, basal, 16 Lima beans, 169 basement, 15, 16 Lipase, 81, 121 mesenteric, 18 Lithium, 96 Menazon, 184 Litter, breakdown of, 141-5 Mercuric chloride, 100 fragmentation of, 141-5 Meronephridia, 46 humification, 145-7 Mesenteric membrane, 18 incorporation of, 141 Mesh bag, 142, 143, 203 rate of burial, 142 Metabolism, effects of temperature see Leaf litter on, 130 Litter turnover, 145-7 Metal, 140 Loam, 198 Metham sodium, 181 Locomotion, 92-4 Methyl bromide, 181 Longitudinal muscle, 4, 13, 14, 16, Mexico, 45, 46, 49, SO, 54 17,24,92 Microbial cell, 159 Lucerne, 166 Microbial protein, 148 Lumbricine setae, 6, 7 Microftora, 141, 160 Luminescence, 77 Micronephridia, 28 Lung worms, 193, 194 Micro-organisms, 80, 121, 147, Lymph glands, 82 155-62, 166 Lymphocytes, 18 and earthworms, 155-62 dispersal' of, 161 Madagascar, 45, 46, SO numbers in casts, 157 Magnesium, 153, 173 numbers in gut, 156 Magnesium sulphate, 99, 199 Middle tube, 24, 26, 83, 84 Malagasy, 49, 54 Migrations, 77 Malathion, 184 Millet, 169 Malay Archipelago, 41, 46, 49, 50 Mineral elements, 154 Male pores, 8, 12, 39- 43, 47, 49, 51, Mineral nutrients, 152 52, 62- 5 Minerals, effect of, 134 Malic acid, 89 Mineral soil, 146, 147, 164 Mammals, 19 5 Mineralization of nitrogen, 147- 50 Mangolds, 135 Mites, 147 Manure, 172 Mixed woodland, 103 Maoris, 197 Moisture, 127-9, 137 Marginal cells, 25, 26 effects of, 127- 9 Mass migration, 77 reaction to, 209 Mating, 62, 63 Moisture content, 99, 106, 111, 112, Maze, 76, 96, 140 134 Mechanized soil washing, 99 Moisture equivalent, 173 Medium loam, population in, 136, Moisture gradient, 128 137 Moles, 123, 190 Meganephridia, 28 Molybdenum, 153 GENERAL INDEX 277

Moorland soil, 171 Nephridiopore, 9, 23, 24, 42, 43, 81, Mor soil, 103, 104, 122, 164 91, 148 species in, 13 7 Nephridium, 9, 24, 26, 27, 83 Mortality, 70 Nephrostome, 24, 25, 26, 27, 82-4 Motor fibres, 86 lower lip, 25 Mountain ranges, 53 peritoneum, 25 Mouth, 80, 90 upper lip, 25 Mowrah meal, 100 prostomial, 28, 29 Mozambique, 48, 50 Nerve depressant, 96 Mucocytes, 19 Nerve, segmental, 28, 29, 30, 86 Mucoproteins, 148 septal, 29 Mucous cells, 4, 15, 36 Nerve cells, 87 Mucous tubes, 65 Nerve cord, 17, 85, 96, 210 Mucus, 63, 80, 81 Nerve fibres; 15 Mud, 62 Nerve plexus, 14 Mull soil, 104, 122, 165 Nerves, electric potentials in, 88 species in, 137 Nervous System, 28, 29, 30-2, 85- Muscle, arciform, 63 90 Muscle, circular, innervation of, 30 fine structure, 31 dorsal, 16 Neural lamella, 31 fibres, 16, 17 Neurofibrils, 16 innervation, 86 Neuroglia, 31 Muscle, longitudinal, 4, 13, 14, 16, Neuropile, 31, 32 17,24,92 Neurosecretions, 87 longitudinal, innervation of, 30 New Caledonia, 44-6, 55 setal, 4 New South Wales, 44, 51, 52, 103, sphincter, 9, 18 145 subcuticular, 13 New Zealand, 44-7, 51, 53, 55, 68, ventral, 16 167, 169, 170, 173, 197 Muscoid flies, 124 Newly-emerged worms, 69 Muscular tube, 24, 26 Nigeria, 45, 48, 50, 104, 11 3, 146 Mycobacteria, 160 Nile Valley, 122 Nitrate, 147, 149, 162, 173 Napthalene, 142 Nitrate of ammonia, effects of, 134 Narrow tube, 24, 25, 26, 83, 84 Nitrate of soda, 148, 178,180 Nematodes, 19, 80, 124, 181, 193 Nitro-chalk, 179 Nephridia, 2, 20, 81, 82, 41, 47, 49 Nitrogen, 168, 178 enteronephric, 27 effect on growth, 138 exonephric, 27, 28 effect on worms, 179 holonephridia, 28 Nitrogen excretion, 151 meganephridia, 28 Nitrogen-fixing bacteria, 14 7 meronephridia, 28 Nitrogen humification, 147- 50 micronephridia, 27, 28 Nitrogen mineralization, 147- 50 278 BIOLOGY OF EARTHWORMS

Noradrenalin, 87 Oxalic acid, 89 Novocaine, 87 Oxidation-reduction potential, 133 Number of samples, 102 Oxygen, 23,78- 80,96 Nutrient transport, 82 consumption of, 159, 162 Nutrition, 58, 80 debt, 79, 151 Nylon net, 143 tension, 133, 151

Oak, 140-2, 144, 149, 150, 157, 170 Pacific Ocean Islands, 55 woodland, 103 Pakistan, 41, 51, 52, 55 Oat straw, as food, 138 Palaearctic zone, 40, 54 Oats, 169 Palatability, 89, 203 Obligatory diapause, 57, 68, 111, experiments, 203, 204 207 of leaf litter, 140 Oesophageal cavity, 20 Panama, 45, 49, 50 Oesophageal pouches, 20 Papillae, 5, 7, 10-12 Oesophagus, 19, 20, 80, 85 Paraguay, 44- 6, 49, 50 Oocytes, 63 Paraquat, 182 Oogenesis, 63 effect on worms, 182 Oogonia, 63 Parasites, 123-5 Oothecae, 56 Parathion, 96, 184 Optic organelle, 32 Parthenogenesis, 63, 66 Orchards, 101, 103- 5, 133, 141, 146, Passive agents, 196 187 Pasture, 68, 99, 103, 104, 112, 113, Ordovician period, 38 121-3, 127, 131, 139, 161, 163, Organ, excretory, 9 166, 169, 173, 176, 177 genital, 13, 37 Pasture productivity, effect of respiratory, 23, 78 worms on, 170 sense, 89 Peas, 169 Organic matter, 80, 81, 106, 121, Peat, 58, 122 165, 168, 171, 173, 174, 176, Peaty soil, population in, 137 180, 201, 204 Penes, 65 cycles, 141- 54 Peregrine species, 53 effect of, 133, 134, 135, 136 Perichaetine setae, 6, 7 Organelle, optic, 16 Peri-enteric plexus, 17, 23, 85 Organo-chlorines, 183 Peristalsis, 92, 93, 94 Organo-phosphates, 183 Pacific Islands, 44 Osmotic pressure, 91 Peristomium, 1, 3, 52 Osmotic pressure of urine, 83 Peritoneal cells, 82 Ova, 56, 63, 65, 66 Peritoneum, 4, 13, 14, 17, 23, 63 Ovarian funnels, 33 innervation of, 31 Ovaries, 32, 40, 41, 51, 63, 96 Peritrophic membrane, 81 Oviduct, 63 Persia, 52 Ovisacs, 33, 63 Peru, 44, 49 GENERAL INDEX 279

Pesticides, effects of, 181-4 Population, estimation of, 98-102 uptake in earthworms, 184-8 numbers, 102 Pests, earthworms as, 190, 191 structure, 104- 6 Petroleum ether, 160 Populations, 70, 104 pH, 89,106,125-7,168,172,201 Pores, cuticular, 13 effects of, 125-7 dorsal, 2, 9, 19, 23, 52, 72, 90, 127 reaction to, 209 female, 8, 9, 10, 12, 40, 41, 49, 51, Pharyngeal ganglion, 30 63 Pharyngeal gland, 19 genital, 1, 3, 5, 7, 10 Pharyngeal intestinal plexus, 32 prostatic, 2, 7 Pharyngeal nerve plexus, 32 sperm, 52 Pharynx, 19,20, 30,80 spermathecal, 2, 8, 12, 40, 41, 47 Phillipines, 41, 190 Porphyrins, 13 Phorate, 184, 191 Porosity, 167, 168 Phosphorescence, 77 Post-clitellar region, 7 Phosphoric acid, 89 Post-septal canal, 24 Phosphorus, 153, 173 Potassium, 153 Photoreceptor cells, 4, 15, 16, 32 Potassium cyanide, 96 Physiology, 78-97 Potassium permanganate, 89, 100-3 Pig litter, 103 Potato root eelworm, 192 Pigmentation, 13 Predation, 58 Pigs, 193, 195 Predators, 121, 123- 5 Piles, 197 Pregnancy, 197 Pine, 144 Preseptal canal, 24 woodland, 103 Preservation, 200 needles, 138 Proboscis, 1 Plant cover, 134 Prolobous prostomium, 3 Plant growth substances, 171 Propham, 181 Plant pathogens, 161 Proprioreceptor cell, 32 Ploughing, 17 4, 17 5 Prostates, 7, 36, 41- 3 Plug, 74 Prostatic fluid, 65 Polders, 171 Prostomial nerve, 28, 29 Polynesia, 46 Prostomium, 1, 2, 3, 4, 12, 28, 29, Polyphenols, 140, 142 52, 87, 89, 91, 92 reaction to, 89 innervation, 28, 32 Polysaccharide gums, 160, 166 Protease, 81 Population, biomass, 102 Protozoa, 19, 80, 160, 161 horizontal distribution, 106, 107 Protozoan parasites, 124 seasonal activity, 11~17 seasonal changes in, 71, 11~17 Quadrat, 101, 102, 203 size of, 102- 4 Queensland, 44- 6 vertical distribution, 107- 10 Quiescence, 67-9, 109-11, 128 distribution of, 106- 10 Quinine, 16, 89 280 BIOLOGY OF EARTHWORMS

Radiation, effect on A. chlorotica, Saccharose, 89 188 Sahara, 54 Radiocaesium, 146 St Thomas Island, 44, 46 Radioisotopes, effect on worms, Salivary gland, 80 188, 189 Sampling, 98 uptake into worms, 188, 189 Sampling tool, 98 Radishes, 196 Sand, 163 Ragi, 196 Sardinia, 46, 50, 54 Rain, reactions, to, 77 Scavenging, 71 Raw humus, 134 Scotland, 122 Reception zone, 80 Scots pine, 150 Reduction division, 63 Seasonal abundance, 106,107 Regeneration, 94-6, 209 Seasonal activity, 110-17 Regenerative phenomena, 87 Secondary sex characters, 87 Replacing cells, 1 5 Secretory cells, 87 Reproduction, 62-7, 87, 105 Seedlings, 190 Reproductive cells, 62 Segment numbering, 3 Reproductive cycle, 87 Segmental contractions, 93 Reproductive potential, 106 Segmental ganglia, 28 Reproductive system, 32, 33, 34, 35, Segmental nerves, 28, 29, 30, 86 36 Segmentation, 1-4 Resorption, 84 Segments, 3, 69 Respiration, 78-80, 150, 151 Seminal fluid, 64, 65 effects of temperature on, 130 Seminal funnel, 34 Respiratory system, 23, 24 Seminal grooves, 2, 8, 12, 63 Resting phase, 68 Seminal vesicles, 32, 34, 35, 52, 62, Retinella, 32 124, 197 Rheumatism, 197 Senescence, 69 Rhizopods, 157 Sense organs, 89 Rhodesia, 50 Sensory cells, 14, 15, 87 Rice, 190 Septa, 1, 17, 18, 26, 29, 31 Ridges, 10, 11 Septa innervation of, 30 Rigor, 91 Septal nerves, 29 Rio de Janeiro, 50 Seta of L. terrestris, 4 Rodents, 194 Setae, 4, 5, 24, 40, 41, 47, 49, Rolling, 174 51 Roots, 139, 141 closely-paired, 6 Rotary cultivation, 174 distant-paired, 6 Rotations, 176 genital, 5 Rotifers, 80 lumbricine, 6 Rowan, 150 normal, 4, 12 Russia, 51, 52 perichaetine, 6 Rye, 157, 169, 176 sexual, 49, 51, 53, 65 GENERAL INDEX 281

ventral, 12 Soil profile, 141 widely-paired, 6 with earthworms, 143 Setal muscles, 4 without earthworms, 143, 164 Setal arrangement, 6, 7 Soil structure, 196, 163-8 Setal form, 4, 5 Soil texture, 106 Sewage sludge, 58 Soil turnover, 164, 165 Sexual activity, 62 Soil type, 136, 137 Sexual development, 63 Soil washing, 99 Sexual maturity, 10, 67, 71, 161 mechanized, 99 Sexual organs, 37, 96 South Shetland Islands, 55 Seychelles, 44 Soybeans, 169, 177, 178 Sheep droppings, as food, 138 Species associations, 122, 123 Shell 19805, 181 Specific gravity, 99 Shrews, 123 Sperm funnels, 33, 35, 62 Siberia, 51, 52 Sperm pores, 52 Sicily, 51, ~4 Sperm transference, 39, 65 Sieve, 99, 101 Spermathecae, 8, 11, 32, 34, 35, Silt, 163 36, 39, 41, 47, 51, 52, 64, 65, Silurian period, 38 124 Simazine, 181 Spermathecal openings, 63 Simple key, 213 Spermathecal pores, 2, 8, 12, 40, 41, Size of populations, 102-4 47 Size of samples, 102 Spermatogenesis, 62, 197 Slime, production of, 77 Spermatogonia, 62 Slime tube, 63 Spermatozoa, 62, 65 Slugs, 123 Sphincter muscles, 9, 18 Sodium chloride, 89 Sporangia, 38 Soil, forest, 166 Springtails, 147 garden, 167, 169 Spruce, 140, 170 heavy, 118 Squirter earthworm, 19 Soil acidity, 88, 125, 127, 169, 178- Staphylinid beetles, 123 81,201 Stimuli, chemical, 72, 88, 89 Soil aeration, 165, 167, 168 electrical, 72, 75, 76, 171 Soil amelioration, 171-3 electrical response to, 90, 91 Soil consumption, 146 light, 76, 87 Soil drainage, 167, 168 photic, 88 Soil erosion, 197 sensory, 86 Soil fertility, 154, 163- 73 touch, 72, 76 Soil formation, 172 vibrations, 77 Soil indicators, 168 Stones, in the bladder, 197 Soil microflora, 134 Straw, 58, 198 Soil moisture, 56, 100, 168 Stubble-mulch farming, 176 Soil porosity, 167, 168 Stubble striping, 175 282 BIOLOGY OF EARTHWORMS

Subcuticular muscle, 13 lethal, 131-3 Sub-epidermal nerve plexus, 30, 31 seasonal changes in, 114 Sub-epithelial nerve plexus, 31 Temperature preferenda, 130, 131, Submergence, 129 132 Subneural vessel, 17, 21, 22, 85 Temperatute tolerance, 132, 133 Suboesophageal ganglion, 28 Temperatures, optimum for Subpharyngeal connectives, 30 growth,206 Subpharyngeal ganglion, 29 Termites, 123 Subtillers, 176 Testa! sacs, 34, 62 Sucrose, 16, 89 Testes, 32, 34, 40, 41, 51, 124 Sudan, SO holoandric, 33 Sulphate of ammonia, 180 meroandric, 3 3 efftl4:ts of, 134 Thigmotactic reactions, 89-90 Sumatra, 41 Timothy grass, 176 Sumithion, 184 Tissue connective, 16, 17 Summer, 62, 68 'T'-maze, 75, 76, 210 Sunda Islands, 41 Tobacco, 190 Superphosphate, 178, 180 Tobago, 54 Supra-intestinal vessel, 22 Togoland, 48, 50 Surface-dwelling species, 199 Tooth powder, 197 Survey, 186 Touch, 63, 71, 72 Survival in water, 79, 209 Transplantation, 96, 97 Swine, 194 Transverse commissure, 88 Symbiotic bacteria, 81 Transverse septa, 17 Sympathetic system, 30 Triassic era, 38 Synapses, 86 Triazine herbicides, 181 Trinidad, 54 Tactile receptors, 89 Tropical areas, 79 Tanylobous prostomium, 3 Tropical forest, 146 Tanzania, 47, 48, 50 Tropics, 122 Tapeworms, 124, 193, 194 Tube, middle, 24, 26, 83, 84 Tartaric acid, 89 mucous, 65 Tasmania, 46 muscular, 24, 26 Taxonomy, 37-55 narrow, 24, 25 , 26, 83, 84 TCA, 181 slime, 63 Teleospores, 192 wide, 24, 26, 83, 84 Temperate forest, 146 Tubercula pubertatis, 3, 10, 11, 12, Temperate zones, 68 52 T emperature, 56, 57, 60, 67, 79, 96, Tubercules, 10 100, 106, 108-12, 115, 129-33, Tumours, 197 151, 198, 211 Tunis, 51 conditioning, 133 Turnover of soil and litter, 145-7 effect on respiration, 79 T y phlosole, 21, 85 GENERAL INDEX 283

Uganda, 50, 104 lateral-oesophageal subneural, 22, Ubiquitous species, 126 23 Ultra-violet light, 77, 80, 88 nephridial, 23 Urea, 81-4, 148 subneural, 17, 21, 22, 85 Uric acid, 84, 148 supra-intestinal, 22 Uric cells, 81, 82 supra-intestino ventral, 23 Urine, 81, 91, 148, 170 typhlosolar, 23 osmotic pressure of, 83 ventral, 17, 21, 22, 23, 30 U.S.A., 38, 46, 50, 53, 54, 103, 105, ventra-intestinal, 23, 85 115, 125, 128, 175 ventra-oesophageal, 85 U.S.S.R., 103, 172, 176 ventro-parietal, 17, 23, 85 Vibrations, 72 Valerionic acid, 79 Victoria, 46 Vas deferens, 33, 34, 35, 40, 62 Viruses, 124 Vas efferens, 34, 35, 62 Voltage, 99 Vascular system, 21- 3 Vegetable crops, 190 Wales, 103 Venezuela, 49 Water conservation, 90 Ventral muscle, 16 Water loss, 68 Ventral nerve cord, 28, 29, 30, 31, Water relationships, 90-2 34, 88, 93, 96 Water-stable aggregates, 163, 165, structure of, 3 2 166-8 Ventral vessel, 17, 85 Weathering, 44-6, 49, 53, 54, 142 Ventral view, 12 Westmorland, 103 Vertical distribution, 108, 109, Wheat, 169, 176 209 effects of, 135 Vertical migration, 58, 67 Wide tube, 24, 26, 83, 84 Vessel, afferent nephridial, 17 Winter, 68 afferent typhlosolar, 17 Withdrawal reactions, 71 blood, 17, 21, 96 Woodland, 135, 151 capillary, 7 8 Worm cairns, 74 commissural, 22 Wound tissue, 95 dorsal, 17, 20, 21, 22, 23, 30, 84, Wych Elm, 140 85 Yeasts, 157, 159 dorsa-intestinal, 17, 23 Yield increases, 170 dorsa-subneural, 17, 21- 3, 85 efferent nephridial, 17 Zanzibar Island, 48 extra-oesophageal, 22 Zinophos, 184 lateral-neural, 17, 22, 85 Zurich, 122 lateral-oesophageal, 21, 22 Zygolobous prostomium, 3